text: Adaptive divergence may occur when a population becomes geographically divided: followed by an accumulation of genetic differences as they adapt to their own unique environments. Reproductive barriers do not evolve as a consequence of external forces that drive populations toward speciation. Rather, the evolution of reproductive isolation, leading to speciation, is generally thought to be an incidental by-product of genetic divergence, particularly adaptive changes that evolve through natural selection in response to different environmental conditions in separate geographic areas.

text: Adaptive divergence may occur when a population becomes geographically divided: followed by an accumulation of genetic differences as they adapt to their own unique environments. Reproductive barriers do not evolve as a consequence of external forces that drive populations toward speciation. Rather, the

entity_name: Evolution

text: evolution

text: of reproductive isolation, leading to speciation, is generally thought to be an incidental by-product of genetic divergence, particularly adaptive changes that evolve through

entity_name: Natural selection

text: natural selection

text: in response to different environmental conditions in separate geographic areas.

method: UvABottomUpChangeOrder

method: UNH-tfidf-stem

method: UNH-tfidf-ptsim

method: UNH-tfidf-lem

text: Natural selection acts on the phenotype, or the observable characteristics of an organism, but the genetic (heritable) basis of any phenotype that gives a reproductive advantage may become more common in a population. Over time, this process can result in populations that specialise for particular ecological niches (microevolution) and may eventually result in the emergence of new species (macroevolution). In other words, natural selection is a key process in the evolution of a population. Natural selection can be contrasted with artificial selection, in which humans intentionally choose specific traits, whereas in natural selection there is no intentional choice.

text: Natural selection acts on the phenotype, or the observable characteristics of an organism, but the

entity_name: Genetics

text: genetic

text: (heritable) basis of any phenotype that gives a reproductive advantage may become

entity_name: Allele frequency

text: more common in a population

text: . Over time, this process can result in populations that specialise for particular

entity_name: Ecological niche

text: ecological niche

text: s (

entity_name: Microevolution

text: microevolution

text: ) and may eventually result in the

entity_name: Speciation

text: emergence of new species

entity_name: Macroevolution

text: macroevolution

text: ). In other words, natural selection is a key process in the evolution of a population. Natural selection can be contrasted with artificial selection, in which

entity_name: Human

text: human

text: s intentionally choose specific traits, whereas in natural selection there is no intentional choice.

method: dangnt-nlp

text: Interactions between organisms can produce both conflict and cooperation. When the interaction is between pairs of species, such as a pathogen and a host, or a predator and its prey, these species can develop matched sets of adaptations. Here, the evolution of one species causes adaptations in a second species. These changes in the second species then, in turn, cause new adaptations in the first species. This cycle of selection and response is called coevolution. An example is the production of tetrodotoxin in the rough-skinned newt and the evolution of tetrodotoxin resistance in its predator, the common garter snake. In this predator-prey pair, an evolutionary arms race has produced high levels of toxin in the newt and correspondingly high levels of toxin resistance in the snake.<ref>

text: Interactions between organisms can produce both conflict and cooperation. When the interaction is between pairs of species, such as a

entity_name: Pathogen

text: pathogen

text: and a

entity_name: Host (biology)

text: host

text: , or a predator and its prey, these species can develop matched sets of adaptations. Here, the evolution of one species causes adaptations in a second species. These changes in the second species then, in turn, cause new adaptations in the first species. This cycle of selection and response is called coevolution. An example is the production of

entity_name: Tetrodotoxin

text: tetrodotoxin

text: in the

entity_name: Rough-skinned newt

text: rough-skinned newt

text: and the evolution of tetrodotoxin resistance in its predator, the

entity_name: Common garter snake

text: common garter snake

text: . In this predator-prey pair, an

entity_name: Evolutionary arms race

text: evolutionary arms race

text: has produced high levels of toxin in the newt and correspondingly high levels of toxin resistance in the snake.<ref>

text: Gene flow involves the exchange of genes between populations and between species. The presence or absence of gene flow fundamentally changes the course of evolution. Due to the complexity of organisms, any two completely isolated populations will eventually evolve genetic incompatibilities through neutral processes, as in the Bateson-Dobzhansky-Muller model, even if both populations remain essentially identical in terms of their adaptation to the environment.

text: Gene flow involves the exchange of genes between populations and between species. The presence or absence of gene flow fundamentally changes the course of evolution. Due to the complexity of organisms, any two completely isolated populations will eventually evolve genetic incompatibilities through neutral processes, as in the

entity_name: Bateson–Dobzhansky–Muller model

text: Bateson-Dobzhansky-Muller model

text: , even if both populations remain essentially identical in terms of their adaptation to the environment.

text: The continuum concept is an idea, coined by Jean Liedloff in her 1975 book The Continuum Concept, that human beings have an innate set of expectations (which Liedloff calls the continuum) that our evolution as a species has designed us to meet in order to achieve optimal physical, mental, and emotional development and adaptability. According to Liedloff, in order to achieve this level of development, young humans—especially babies—require the kind of experience to which our species adapted during the long process of our evolution by natural selection. For infants, these include, for example, that they experience:

text: The continuum concept is an idea, coined by

entity_name: Jean Liedloff

text: Jean Liedloff

text: in her 1975 book The Continuum Concept, that

text: human being

text: s have an

entity_name: Intrinsic and extrinsic properties

text: innate

text: set of expectations (which Liedloff calls the continuum) that our evolution as a species has designed us to meet in order to achieve optimal physical, mental, and emotional

entity_name: Human development (biology)

text: development and adaptability

text: . According to Liedloff, in order to achieve this level of development, young humans—especially babies—require the kind of experience to which our species adapted during the long process of

entity_name: Human evolution

text: our evolution

text: by

text: natural selection

text: . For infants, these include, for example, that they experience:

text: The definition of a "gene" has never been a trivial issue, with numerous definitions and notions proposed throughout the years since the discovery of the human genome. First, genes were conceived in the 1900s as discrete units of heredity, then it was thought as the blueprint for protein synthesis, and in more recent times, it was being defined as genetic code that is transcribed into RNA. Although the definition of a gene has evolved greatly over the last century, it has remained a challenging and controversial subject for many researchers. With the advent of the ENCODE/GENCODE project, even more problematic aspects of the definition have been uncovered, including alternative splicing (where a series of exons are separated by introns), intergenic transcriptions, and the complex patterns of dispersed regulation, together with non-genic conservation and the abundance of noncoding RNA genes. As GENCODE endeavours to build an encyclopaedia of genes and gene variants, these problems presented a mounting challenge for the GENCODE project to come up with an updated notion of a gene.

text: Watson et al. provide support for a specific specialization in language-dependent humor. Its adaptive value has both extrinsic and intrinsic components: humor facilitates social bonding if shared extrinsically, and provides pleasure if enjoyed in one's own mind. In addition, Johnson-Frey (2003) proposed a unique human specialization for tool use. According to Johnson-Frey, humans' ability to use tools is based on complex cognitive mechanisms, not just advanced sensorimotor skills. Rather than it being considered a purely physical specialization based only in motor areas of the brain, Johnson-Frey argues that tool use should be classified as a cognitive phenomenon due to its foundation in cognition. On a more philosophical level, Boyer (2003) argues that "religious thought and behavior" is a specialization that originally developed as a by-product of brain function, and its adaptive purposes led to its continued evolution by natural selection. Krueger et al. (2007) have argued that trust, which may form the foundation for helping and altruism and thus the basis of human social interaction, is also a cognitive specialization.

text: Watson et al. provide support for a specific specialization in language-dependent

entity_name: Humour

text: humor

text: . Its

entity_name: Adaptive behavior

text: adaptive

text: value has both

text: extrinsic

text: and

text: intrinsic

text: components: humor facilitates social bonding if shared extrinsically, and provides pleasure if enjoyed in one's own mind. In addition, Johnson-Frey (2003) proposed a unique human specialization for tool use. According to Johnson-Frey, humans' ability to use tools is based on complex cognitive mechanisms, not just advanced

entity_name: Motor skill

text: sensorimotor

text: skills. Rather than it being considered a purely physical specialization based only in

entity_name: Motor cortex

text: motor areas

text: of the brain, Johnson-Frey argues that tool use should be classified as a cognitive phenomenon due to its foundation in cognition. On a more

entity_name: Philosophy

text: philosophical

text: level, Boyer (2003) argues that "

entity_name: Religion

text: religious

text: thought and behavior" is a specialization that originally developed as a by-product of brain function, and its adaptive purposes led to its continued evolution by

text: natural selection

text: . Krueger et al. (2007) have argued that trust, which may form the foundation for helping and altruism and thus the basis of human social interaction, is also a cognitive specialization.

text: After his Nobel prize award, Edelman began research into the regulation of primary cellular processes, particularly the control of cell growth and the development of multi-celled organisms, focussing on cell-to-cell interactions in early embryonic development and in the formation and function of the nervous system. These studies led to the discovery of cell adhesion molecules (CAMs), which guide the fundamental processes that help an animal achieve its shape and form, and by which nervous systems are built. One of the most significant discoveries made in this research is that the precursor gene for the neural cell adhesion molecule gave rise in evolution to the entire molecular system of adaptive immunity.

text: After his Nobel prize award, Edelman began research into the regulation of primary

entity_name: Cell (biology)

entity_section: Cellular processes

text: cellular process

text: es, particularly the control of cell growth and the development of

entity_name: Multicellular organism

text: multi-celled organism

text: s, focussing on cell-to-cell interactions in early

entity_name: Embryogenesis

text: embryonic development

text: and in the formation and function of the nervous system. These studies led to the discovery of

entity_name: Cell adhesion molecule

text: cell adhesion molecules

text: (CAMs), which guide the fundamental processes that help an animal achieve its shape and form, and by which nervous systems are built. One of the most significant discoveries made in this research is that the precursor

entity_name: Gene

text: gene

text: for the neural cell adhesion molecule gave rise in evolution to the entire molecular system of

entity_name: Adaptive immune system

text: adaptive immunity

method: ICT-DRMMTKS

text: Competition can have both beneficial and detrimental effects. Many evolutionary biologists view inter-species and intra-species competition as the driving force of adaptation, and ultimately of evolution. However, some biologists disagree, citing competition as a driving force only on a small scale, and citing the larger scale drivers of evolution to be abiotic factors (termed 'Room to Roam'). Richard Dawkins, prefers to think of evolution in terms of competition between single genes, which have the welfare of the organism 'in mind' only insofar as that welfare furthers their own selfish drives for replication (termed the 'selfish gene').

text: Competition can have both beneficial and detrimental effects. Many evolutionary biologists view inter-species and intra-species competition as the driving force of

entity_name: Adaptation

text: adaptation

text: , and ultimately of

text: evolution

text: . However, some biologists disagree, citing competition as a driving force only on a small scale, and citing the larger scale drivers of evolution to be abiotic factors (termed 'Room to Roam').

entity_name: Richard Dawkins

text: Richard Dawkins

text: , prefers to think of evolution in terms of competition between single genes, which have the welfare of the organism 'in mind' only insofar as that welfare furthers their own selfish drives for replication (termed the 'selfish gene').

text: Adaptive evolution results from the propagation of advantageous mutations through positive selection. This is the modern synthesis of the process which Darwin and Wallace originally identified as the mechanism of evolution. However, in the last half century there has been considerable debate as to whether evolutionary changes at the molecular level are largely driven by natural selection or random genetic drift. Unsurprisingly, the forces which drive evolutionary changes in our own species’ lineage have been of particular interest. Quantifying adaptive evolution in the human genome gives insights into our own evolutionary history and helps to resolve this neutralist-selectionist debate. Identifying specific regions of the human genome that show evidence of adaptive evolution helps us find functionally significant genes, including genes important for human health, such as those associated with diseases.

text: Adaptive evolution

text: results from the propagation of advantageous

entity_name: Mutation

text: mutations

text: through

entity_name: Directional selection

text: positive selection

text: . This is the modern synthesis of the process which

entity_name: Charles Darwin

text: Darwin

text: and

entity_name: Alfred Russel Wallace

text: Wallace

text: originally identified as the mechanism of evolution. However, in the last half century there has been considerable debate as to whether evolutionary changes at the molecular level are largely driven by natural selection or random genetic drift. Unsurprisingly, the forces which drive evolutionary changes in our own species’ lineage have been of particular interest. Quantifying adaptive evolution in the human genome gives insights into our own evolutionary history and helps to resolve this

text: neutralist-selectionist debate

text: . Identifying specific regions of the human genome that show evidence of adaptive evolution helps us find functionally significant

text: gene

text: s, including genes important for human health, such as those associated with diseases.

text: The dictionary definition of Evolution is any process of formation, growth or development. In biological evolution the main principle behind this development is survival, we evolved to become stronger and quicker, we also evolved to become intelligent. But as we became intelligent biological evolution subsided to a new concept, cultural evolution. Cultural evolution moves at a much faster rate than biological evolution and this is one reason why it isn’t very well understood. But as survival is still the main driving force behind life and that intelligence and knowledge is currently the most important factor for that survival, we can reasonably assume that cultural evolution will progress in the direction of furthering intelligence and knowledge.

text: The dictionary definition of Evolution is any process of formation, growth or development. In

text: biological evolution

text: the main principle behind this development is survival, we evolved to become stronger and quicker, we also evolved to become intelligent. But as we became intelligent biological evolution subsided to a new concept,

entity_name: Cultural evolution

text: cultural evolution

text: . Cultural evolution moves at a much faster rate than biological evolution and this is one reason why it isn’t very well understood. But as survival is still the main driving force behind life and that intelligence and knowledge is currently the most important factor for that survival, we can reasonably assume that cultural evolution will progress in the direction of furthering intelligence and knowledge.

text: Evolution does not attempt to explain the origin of life (covered instead by abiogenesis), but it does explain how the extremely simple early lifeforms evolved into the complex ecosystem that we see today. Based on the similarities between all present-day organisms, all life on Earth originated through common descent from a last universal ancestor from which all known species have diverged through the process of evolution. All individuals have hereditary material in the form of genes that are received from their parents, then passed on to any offspring. Among offspring there are variations of genes due to the introduction of new genes via random changes called mutations or via reshuffling of existing genes during sexual reproduction. The offspring differs from the parent in minor random ways. If those differences are helpful, the offspring is more likely to survive and reproduce. This means that more offspring in the next generation will have that helpful difference and individuals will not have equal chances of reproductive success. In this way, traits that result in organisms being better adapted to their living conditions become more common in descendant populations. These differences accumulate resulting in changes within the population. This process is responsible for the many diverse life forms in the world.

text: Evolution does not attempt to explain the origin of

entity_name: Life

text: life

text: (covered instead by

entity_name: Abiogenesis

text: abiogenesis

text: ), but it does explain how the extremely simple early lifeforms evolved into the complex ecosystem that we see today. Based on the similarities between all present-day organisms, all life on

text: Earth

text: originated through

entity_name: Common descent

text: common descent

text: from a

entity_name: Last universal common ancestor

text: last universal ancestor

text: from which all known

entity_name: Species

text: species

text: have diverged through the process of evolution. All individuals have hereditary material in the form of

text: gene

text: s that are received from their parents, then passed on to any offspring. Among offspring there are variations of genes due to the introduction of new genes via random changes called mutations or via reshuffling of existing genes during

entity_name: Sexual reproduction

text: sexual reproduction

text: . The offspring differs from the parent in minor random ways. If those differences are helpful, the offspring is more likely to survive and reproduce. This means that more offspring in the next generation will have that helpful difference and individuals will not have equal chances of

entity_name: Reproduction

text: reproductive

text: success. In this way, traits that result in organisms being better

text: adapted

text: to their living conditions become more common in descendant populations. These differences accumulate resulting in changes within the population. This process is responsible for the many diverse life forms in the world.

text: DCPS focuses on two scientific leitmotifs: “Function and Diversity”and “Plant and Environment”. The research findings from bothareas are transferred to practices supporting sustainable use andthe protection of plant diversity. Here, the spectrum ranges fromthe development of genetic resources (Applied Plant Sciences)tothe management of varied eco-systems. Public relations and communicationtools are also used to present the findings in a moreaccessible form to a broader audience.Function and Diversity research aims to produce new insights intothe evolutionary development of the morphological and functionaldiversity of plants. The evolution of new traits and the mechanismsof genome evolutions are studied by combining comparative genomicswith phylogenetic, biochemical and molecular techniques.The topics addressed include the evolution and function of signaltransfer processes in plant cells, the role of the numerous plantsecondary metabolites, the uptake and utilization of nutrients aswell as the development of morphological features. This work isbased on the rapidly growing information available from genomesequencing. In the meantime this goes far beyond the model plantArabidopsis, previously the focus of much research, and opens upnumerous approaches capable of contributing to a better understandingof gene functions and the evolution of biological diversity.The close links to the newly established Berlin Consortium for Genomicsin Biodiversity Research play a significant role in researchinto genome sequence and structure.In the Plant and Environment field of research, the focus is onplant functions and adaptive reactions in the context of constantlychanging environmental conditions. This complex topic requires a transdisciplinary approach to understand the context (phylogeny,plant geography, ecology) as well as the mechanisms (physiology,biochemistry, molecular biology). Together with its regional partners,the Dahlem Centre of Plant Sciences covers the broad spectrum of expertise requiredhere. The research concentrates on plant reactions to such abioticstress factors as light intensity, extremes of temperature, lack ofwater and nutrients, as well as plant biotic interactions with otherorganisms, for example, insects or fungi. The research groupsare working on such diverse themes as, for instance, the responsesof plants to pathogens, plants’ cellular memory for environmentalstress, the role of root architecture in fungi colonisation and itssignificance for crop yields and the mechanisms of neighbouringplants to communicate with each other.

text: It seems quite reasonable to assume that organisms acquire useful adaptations during their lifetime. Such adaptations are the result of an exploratory search which samples various iterations of potential outputs in order to discern and select the most appropriate ones. Thus, it is plausible that ‘learning’, which can be described as an elaborate and iterative form of phenotypic modification that allows an organism to adjust its response to the same inputs over time based on the outcomes of previous outputs, could have a significant influence on evolution of a new species in the long run. Therefore, it would be quite wasteful to forego the advantage of the exploration performed by the organism to facilitate the evolutionary search for increased fitness if information about the acquired (learned) characteristics (new phenotypes) was not transferred to the genotype or at least retained in some (non-genomic) fashion to facilitate transgenerational inheritance. Indeed, this type of interaction between learning and evolution was independently proposed in the late 1800s by Baldwin, Osborn and Morgan and is often referred to as the ‘Baldwin effect’.  However, the Baldwin effect remained under-appreciated because of its Lamarckian connotation and consequently it was inferred by many that learning cannot guide evolution.

text: It seems quite reasonable to assume that organisms acquire useful adaptations during their lifetime. Such adaptations are the result of an exploratory search which samples various iterations of potential outputs in order to discern and select the most appropriate ones. Thus, it is plausible that ‘learning’, which can be described as an elaborate and iterative form of phenotypic modification that allows an organism to adjust its response to the same inputs over time based on the outcomes of previous outputs, could have a significant influence on evolution of a new species in the long run. Therefore, it would be quite wasteful to forego the advantage of the exploration performed by the organism to facilitate the evolutionary search for increased fitness if information about the acquired (learned) characteristics (new phenotypes) was not transferred to the genotype or at least retained in some (non-genomic) fashion to facilitate transgenerational inheritance. Indeed, this type of interaction between learning and evolution was independently proposed in the late 1800s by Baldwin, Osborn and Morgan and is often referred to as the ‘

entity_name: Baldwin effect

text: Baldwin effect

text: ’.  However, the Baldwin effect remained under-appreciated because of its Lamarckian connotation and consequently it was inferred by many that learning cannot guide evolution.

text: A special case of natural selection is sexual selection, which is selection for any trait that increases mating success by increasing the attractiveness of an organism to potential mates. Traits that evolved through sexual selection are particularly prominent among males of several animal species. Although sexually favoured, traits such as cumbersome antlers, mating calls, large body size and bright colours often attract predation, which compromises the survival of individual males. This survival disadvantage is balanced by higher reproductive success in males that show these hard-to-fake, sexually selected traits.

text: A special case of natural selection is

entity_name: Sexual selection

text: sexual selection

text: , which is selection for any trait that increases mating success by increasing the attractiveness of an organism to potential mates. Traits that evolved through sexual selection are particularly prominent among males of several animal species. Although sexually favoured, traits such as cumbersome antlers, mating calls, large body size and bright colours often attract predation, which compromises the survival of individual males. This survival disadvantage is balanced by higher reproductive success in males that show these

entity_name: Handicap principle

text: hard-to-fake

text: , sexually selected traits.

text: Among domesticated animals, yaks (Bos grunniens) are the highest dwelling animals of the world, living at . The yak is the most important domesticated animal for Tibet highlanders in Qinghai Province of China, as the primary source of milk, meat and fertilizer. Unlike other yak or cattle species, which suffer from hypoxia in the Tibetan Plateau, the Tibetan domestic yaks thrive only at high altitude, and not at lowlands. Their physiology is well-adapted to high altitudes, with proportionately larger lungs and heart than other cattle, as well as greater capacity for transporting oxygen through their blood. In yaks, hypoxia-inducible factor 1 (HIF-1) has high expression in the brain, lung, and kidney, showing that it plays an important role in the adaptation to low oxygen environment. On 1 July 2012 the complete genomic sequence and analyses of a female domestic yak was announced, providing important insights into understanding mammalian divergence and adaptation at high altitude. Distinct gene expansions related to sensory perception and energy metabolism were identified. In addition, researchers also found an enrichment of protein domains related to the extracellular environment and hypoxic stress that had undergone positive selection and rapid evolution. For example, they found three genes that may play important roles in regulating the bodyʼs response to hypoxia, and five genes that were related to the optimisation of the energy from the food scarcity in the extreme plateau. One gene in particular, ADAM-17, is known to be involved in regulating response to low oxygen levels that is also found in Tibetan highlanders.

text: Among

entity_name: List of domesticated animals

text: domesticated animals

text: , yaks (

entity_name: Domestic yak

text: Bos grunniens

text: ) are the highest dwelling animals of the world, living at . The yak is the most important domesticated animal for Tibet highlanders in

entity_name: Qinghai

text: Qinghai Province

text: of

entity_name: China

text: China

text: , as the primary source of

entity_name: Milk

text: milk

entity_name: Meat

text: meat

text: and

entity_name: Fertilizer

text: fertilizer

text: . Unlike other yak or

entity_name: Cattle

text: cattle

text: species, which suffer from hypoxia in the Tibetan Plateau, the Tibetan domestic yaks thrive only at high altitude, and not at lowlands. Their physiology is well-adapted to high altitudes, with proportionately larger lungs and heart than other cattle, as well as greater capacity for transporting oxygen through their blood. In yaks,

entity_name: Hypoxia-inducible factors

text: hypoxia-inducible factor

text: 1 (HIF-1) has high expression in the

entity_name: Brain

text: brain

entity_name: Lung

text: lung

text: , and

text: kidney

text: , showing that it plays an important role in the adaptation to low oxygen environment. On 1 July 2012 the complete genomic sequence and analyses of a female domestic yak was announced, providing important insights into understanding mammalian

text: divergence

text: and adaptation at high altitude. Distinct gene expansions related to

text: sensory perception

text: and energy metabolism were identified. In addition, researchers also found an enrichment of protein domains related to the extracellular environment and hypoxic stress that had undergone positive selection and rapid evolution. For example, they found three genes that may play important roles in regulating the bodyʼs response to hypoxia, and five genes that were related to the optimisation of the energy from the food scarcity in the extreme plateau. One gene in particular, ADAM-17, is known to be involved in regulating response to low oxygen levels that is also found in Tibetan highlanders.

text: The British biologist Alfred Russel Wallace is best known for independently proposing a theory of evolution due to natural selection that prompted Charles Darwin to publish his own theory. In his famous 1858 paper, Wallace proposed natural selection as a kind of feedback mechanism which keeps species and varieties adapted to their environment.

text: The British biologist

text: Alfred Russel Wallace

text: is best known for independently proposing a theory of

text: evolution

text: due to

text: natural selection

text: that prompted

text: Charles Darwin

text: to publish his own theory. In his famous 1858 paper, Wallace proposed natural selection as a kind of feedback mechanism which keeps species and varieties adapted to their environment.

text: During the development of the modern synthesis, Sewall Wright developed his shifting balance theory, which regarded gene flow between partially isolated populations as an important aspect of adaptive evolution. However, recently there has been substantial criticism of the importance of the shifting balance theory.

text: During the development of the modern synthesis, Sewall Wright developed his

text: shifting balance theory

text: , which regarded gene flow between partially isolated populations as an important aspect of adaptive evolution. However, recently there has been substantial criticism of the importance of the shifting balance theory.

text: In common with biological evolution through natural selection or animal husbandry, the members of a population undergoing artificial evolution modify their form or behavior over many reproductive generations in response to a selective regime.

text: In common with biological

text: evolution

text: through

text: natural selection

text: or

text: animal husbandry

text: , the members of a population undergoing artificial evolution modify their form or behavior over many reproductive generations in response to a selective regime.

text: The term is sometimes also colloquially used to refer to acceptance of the modern evolutionary synthesis, a scientific theory that describes how biological evolution occurs. In addition, the term is used in a broader sense to cover a world-view on a wide variety of topics, including chemical evolution as an alternative term for abiogenesis or for nucleosynthesis of chemical elements, galaxy formation and evolution, stellar evolution, spiritual evolution, technological evolution and universal evolution, which seeks to explain every aspect of the world in which we live.

text: The term is sometimes also colloquially used to refer to acceptance of the

text: modern evolutionary synthesis

text: , a

text: scientific theory

text: that describes how biological

text: evolution

text: occurs. In addition, the term is used in a broader sense to cover a world-view on a wide variety of topics, including chemical evolution as an alternative term for

text: abiogenesis

text: or for

text: nucleosynthesis

text: of

text: chemical element

text: s,

text: galaxy formation and evolution

text: stellar evolution

text: spiritual evolution

text: technological evolution

text: and

text: universal evolution

text: , which seeks to explain every aspect of the world in which we live.

text: The evolutionary ecologist Dr. Daniel Janzen began to explicate the idea of ecological fitting with a 1980 paper that observed that many instances of ecological interactions were inferred to be the result of coevolution when this was not necessarily the case, and encouraged ecologists to use the term coevolution more strictly. He observed that the existing defense traits of plants were likely produced by co-evolution with herbivores or parasites that no longer co-occurred with the plants, but that these traits were continuing to protect the plants against new attacks.He expanded this idea in a 1985 paper written while visiting Santa Rosa National Park in Costa Rica.  While there, he observed that almost all of the species in the park occupied large geographic ranges, and despite the heterogeneity of habitats across these ranges, individuals were mostly identical across locations, indicating that little local adaptation had taken place. He described the cyclical life history pattern he believed responsible for this pattern: a species begins as a small population occupying a small area with little genetic variation, but then over the course of a few generations grows to occupy a large area, either because of the emergence of a genotype successful over a wider range, or because of the removal of a geographic barrier.  This large interconnected population is now subject to many contradictory selection pressures and thus remains evolutionarily static until a disturbance separates populations, restarting the cycle.  This cyclic life history pattern is dependent on three premises: that the ancestral range of most species is smaller than the ones now occupied, that biological communities have porous borders and are thus subject to invasion, and that species possess robust genotypes that allow them to colonize new habitats without evolution. Thus, many biological communities may be made up of organisms that despite their complex biological interactions have very little evolutionary history with each other.

text: The evolutionary ecologist Dr.

text: Daniel Janzen

text: began to explicate the idea of ecological fitting with a 1980 paper that observed that many instances of ecological interactions were inferred to be the result of

text: coevolution

text: when this was not necessarily the case, and encouraged ecologists to use the term coevolution more strictly. He observed that the existing defense traits of plants were likely produced by co-evolution with herbivores or parasites that no longer co-occurred with the plants, but that these traits were continuing to protect the plants against new attacks.He expanded this idea in a 1985 paper written while visiting

text: Santa Rosa National Park

text: in

text: Costa Rica

text: .  While there, he observed that almost all of the species in the park occupied large geographic ranges, and despite the

text: heterogeneity

text: of habitats across these ranges, individuals were mostly identical across locations, indicating that little local adaptation had taken place. He described the cyclical life history pattern he believed responsible for this pattern: a species begins as a small population occupying a small area with little

text: genetic variation

text: , but then over the course of a few generations grows to occupy a large area, either because of the emergence of a

text: genotype

text: successful over a wider range, or because of the removal of a geographic barrier.  This large interconnected population is now subject to many contradictory selection pressures and thus remains evolutionarily static until a disturbance separates populations, restarting the cycle.  This cyclic life history pattern is dependent on three premises: that the ancestral range of most species is smaller than the ones now occupied, that biological communities have porous borders and are thus subject to invasion, and that species possess robust genotypes that allow them to colonize new habitats without evolution. Thus, many biological communities may be made up of organisms that despite their complex biological interactions have very little evolutionary history with each other.

text: Many of Darwin's supporters accepted evolution on the basis that it could be reconciled with design. In particular, Asa Gray considered natural selection to be the main mechanism of evolution and sought to reconcile it with natural theology. He proposed that natural selection could be a mechanism in which the problem of evil of suffering produced the greater good of adaptation, but conceded that this had difficulties and suggested that God might influence the variations on which natural selection acted to guide evolution. For Darwin and Thomas Henry Huxley such pervasive supernatural influence was beyond scientific investigation, and George Frederick Wright, an ordained minister who was Gray's colleague in developing theistic evolution, emphasised the need to look for secondary or known causes rather than invoking supernatural explanations: "If we cease to observe this rule there is an end to all science and all sound science."

text: Many of Darwin's supporters accepted evolution on the basis that it could be reconciled with design. In particular,

text: Asa Gray

text: considered natural selection to be the main mechanism of evolution and sought to reconcile it with natural theology. He proposed that natural selection could be a mechanism in which the

text: problem of evil

text: of suffering produced the greater good of adaptation, but conceded that this had difficulties and suggested that God might influence the variations on which natural selection acted to guide evolution. For Darwin and

text: Thomas Henry Huxley

text: such pervasive supernatural influence was beyond scientific investigation, and

text: George Frederick Wright

text: , an ordained minister who was Gray's colleague in developing theistic evolution, emphasised the need to look for secondary or known causes rather than invoking supernatural explanations: "If we cease to observe this rule there is an end to all science and all sound science."

text: Darwin's theory of evolution by natural selection is the only scientific explanation for why an animal's behaviour is usually well adapted for survival and reproduction in its environment. However, claiming that a particular mechanism is well suited to the present environment is different from claiming that this mechanism was selected for in the past due to its history of being adaptive. The literature conceptualizes the relationship between function and evolution in two ways. On the one hand, function and evolution are often presented as separate and distinct explanations of behaviour.

text: Darwin's theory of

text: evolution

text: by natural selection is the only scientific explanation for why an animal's behaviour is usually well adapted for survival and reproduction in its environment. However, claiming that a particular mechanism is well suited to the present environment is different from claiming that this mechanism was selected for in the past due to its history of being adaptive. The literature conceptualizes the relationship between function and evolution in two ways. On the one hand, function and evolution are often presented as separate and distinct explanations of behaviour.

text: This special adaptation is now recognised as an example of natural selection in action. The adaptation account of the Tibetans has become the fastest case of human evolution in the scientific record, as it is estimated to have occurred in less than 3,000 years.

text: This special

text: adaptation

text: is now recognised as an example of

text: natural selection

text: in action. The adaptation account of the Tibetans has become the fastest case of

text: human evolution

text: in the scientific record, as it is estimated to have occurred in less than 3,000 years.

text: Although biological evolution has been vocally opposed by some religious groups, many other groups accept the scientific position, sometimes with additions to allow for theological considerations.  The positions of such groups are described by terms including "theistic evolution", "theistic evolutionism" or "evolutionary creation".  Theistic evolutionists believe that there is a God, that God is the creator of the material universe and (by consequence) all life within, and that biological evolution is a natural process within that creation. Evolution, according to this view, is simply a tool that God employed to develop human life. According to the American Scientific Affiliation, a Christian organization of scientists:

text: Although biological

text: evolution

text: has been vocally opposed by some religious groups, many other groups accept the scientific position, sometimes with additions to allow for theological considerations.  The positions of such groups are described by terms including "

text: theistic evolution

text: ", "theistic evolutionism" or "evolutionary creation".  Theistic evolutionists believe that

text: there is a God

text: , that God is

text: the creator

text: of the

text: material

text: universe and (by consequence) all life within, and that biological evolution is a

text: natural process

text: within that creation. Evolution, according to this view, is simply a tool that God employed to develop

text: human life

text: . According to the

text: American Scientific Affiliation

text: , a

text: Christian

text: organization of scientists:

text: For example, barn owls in Europe exhibit a cline in reference to their plumage coloration. Their feathers range in coloration from white to reddish-brown across the geological range of the southwest to the northeast. This study sought to find if this phenotypic variation was due to selection by calculating the Qst values across the owl populations. Because high gene flow was still anticipated along this cline, selection was only expected to act upon the QTLs that incur locally adaptive phenotypic traits. This can be determined by comparing the Qst values to Fst (fixation index) values. If both of these values are similar and Fst is based on neutral markers then it can be assumed that the QTLs were based on neutral markers (markers not under selection or locally adapted) as well. However, in the case of the barn owls the Qst value was much higher than the Fst value. This means that high gene flow was present allowing the neutral markers to be similar, indicated by the low Fst value. But, local adaptation due to selection was present as well, in the form of varying plumage coloration since the Qst value was high, indicating differences in these non-neutral loci. In other words, this cline of plumage coloration has some sort of adaptive value to the birds.

text: For example, barn owls in Europe exhibit a cline in reference to their plumage coloration. Their feathers range in coloration from white to reddish-brown across the geological range of the southwest to the northeast. This study sought to find if this phenotypic variation was due to selection by calculating the Qst values across the owl populations. Because high gene flow was still anticipated along this cline, selection was only expected to act upon the QTLs that incur locally adaptive phenotypic traits. This can be determined by comparing the Qst values to Fst (

text: fixation index

text: ) values. If both of these values are similar and Fst is based on neutral markers then it can be assumed that the QTLs were based on neutral markers (markers not under selection or locally adapted) as well. However, in the case of the barn owls the Qst value was much higher than the Fst value. This means that high gene flow was present allowing the neutral markers to be similar, indicated by the low Fst value. But, local adaptation due to selection was present as well, in the form of varying plumage coloration since the Qst value was high, indicating differences in these non-neutral loci. In other words, this cline of plumage coloration has some sort of adaptive value to the birds.

text: Kin selection causes changes in gene frequency across generations, driven by interactions between related individuals. This dynamic forms the conceptual basis of the theory of social evolution. Some cases of evolution by natural selection can only be understood by considering how biological relatives influence each other's fitness. Under natural selection, a gene encoding a trait that enhances the fitness of each individual carrying it should increase in frequency within the population; and conversely, a gene that lowers the individual fitness of its carriers should be eliminated. However, a hypothetical gene that prompts behaviour which enhances the fitness of relatives but lowers that of the individual displaying the behaviour, may nonetheless increase in frequency, because relatives often carry the same gene. According to this principle, the enhanced fitness of relatives can at times more than compensate for the fitness loss incurred by the individuals displaying the behaviour, making kin selection possible. This is a special case of a more general model, "inclusive fitness". This analysis has been challenged, Wilson writing that "the foundations of the general theory of inclusive fitness based on the theory of kin selection have crumbled" and that he now relies instead on the theory of eusociality and "gene-culture co-evolution" for the underlying mechanics of sociobiology.

text: Kin selection causes changes in

text: gene

text: frequency across generations, driven by interactions between related individuals. This dynamic forms the conceptual basis of the theory of

text: social evolution

text: . Some cases of

text: evolution

text: by

text: natural selection

text: can only be understood by considering how biological relatives influence each other's

text: fitness

text: . Under natural selection, a gene encoding a

text: trait

text: that enhances the fitness of each individual carrying it should increase in frequency within the

text: population

text: ; and conversely, a gene that lowers the individual fitness of its carriers should be eliminated. However, a hypothetical gene that prompts behaviour which enhances the fitness of relatives but lowers that of the individual displaying the behaviour, may nonetheless increase in frequency, because relatives often carry the same gene. According to this principle, the enhanced fitness of relatives can at times more than compensate for the fitness loss incurred by the individuals displaying the behaviour, making kin selection possible. This is a special case of a more general model, "

text: inclusive fitness

text: ". This analysis has been challenged, Wilson writing that "the foundations of the general theory of inclusive fitness based on the theory of kin selection have crumbled" and that he now relies instead on the theory of

text: eusociality

text: and "gene-culture co-evolution" for the underlying mechanics of

text: sociobiology

text: Evolution is the process of change in all forms of life over generations, and evolutionary biology is the study of how evolution occurs. Biological populations evolve through genetic changes that correspond to changes in the organisms' observable traits. Genetic changes include mutations, which are caused by damage or replication errors in an organism's DNA. As the genetic variation of a population drifts randomly over generations, natural selection gradually leads traits to become more or less common based on the relative reproductive success of organisms with those traits.

text: Evolution is the process of change in all forms of

text: life

text: over generations, and

text: evolutionary biology

text: is the study of how

text: evolution

text: occurs. Biological

text: population

text: s evolve through

text: genetic

text: changes that correspond to changes in the

text: organism

text: s'

text: observable traits

text: . Genetic changes include

text: mutations

text: , which are caused by damage or replication errors in an organism's

text: DNA

text: . As the

text: genetic variation

text: of a population

text: drifts

text: randomly over generations,

text: natural selection

text: gradually leads traits to become more or less common based on the relative

text: reproductive success

text: of organisms with those traits.

text: The gene-centered view of evolution is a synthesis of the theory of evolution by natural selection, the particulate inheritance theory, and the non-transmission of acquired characters. It states that those alleles whose phenotypic effects successfully promote their own propagation will be favorably selected relative to their competitor alleles within the population. This process produces adaptations for the benefit of alleles that promote the reproductive success of the organism, or of other organisms containing the same allele (kin altruism and green-beard effects), or even its own propagation relative to the other genes within the same organism (intragenomic conflict).

text: The gene-centered view of evolution is a synthesis of the theory of

text: evolution

text: by

text: natural selection

text: , the

text: particulate inheritance theory

text: , and the

text: non-transmission of acquired characters

text: . It states that those

text: allele

text: s whose phenotypic effects successfully promote their own propagation will be favorably selected relative to their competitor alleles within the population. This process produces adaptations for the benefit of alleles that promote the reproductive success of the

text: organism

text: , or of other organisms containing the same allele (kin altruism and green-beard effects), or even its own propagation relative to the other

text: gene

text: s within the same organism (intragenomic conflict).

text: The current theory of evolution, the modern evolutionary synthesis (or neo-darwinism), explains that evolution of species occurs through a combination of Darwin’s mechanism of natural selection and Gregor Mendel’s theory of genetics as the basis for biological inheritance and mathematical population genetics. Essentially, the modern synthesis introduced the connection between two important discoveries; the units of evolution (genes) with the main mechanism of evolution (selection).

text: The current theory of evolution, the

text: modern evolutionary synthesis

text: (or neo-darwinism), explains that evolution of

text: species

text: occurs through a combination of Darwin’s mechanism of natural selection and

text: Gregor Mendel

text: ’s theory of genetics as the basis for biological inheritance and mathematical population genetics. Essentially, the modern synthesis introduced the connection between two important discoveries; the units of evolution (genes) with the main mechanism of evolution (selection).

text: Humanist ecology naturally favours the permanent adaptation and the best possible development of humanity, and of the human being, in an uncertain universal environment in permanent evolution, with a mind open enough to consider all the possibilities. In humanist ecological comprehension, it is vain to want to freeze an arbitrary cultural schemata and choose the apparent equilibrium and the supposed future of one moment of evolution. A permanent evolutive adaptation is necessary, as much biologically as mentally. That requires relativity and a caution in the analysis. According to this concept, for the human mind, any representation belongs to the domain of belief, considering the uncertainty of the relation of man to the universe, and the natural imperfection of his senses to represent his environment and his interactions with this environment; the reality perceived by Man being only one representation of reality, particular in mankind. Humanist ecology admits this relativized relation with belief but refutes any final and locking form, knowing that no truth can be final for the human mind without upsetting its natural evolutive necessity. This school of thought accepts belief in the present, for want of anything better, but dictates that you must take care to verify and update beliefs.

text: Humanist ecology naturally favours the permanent adaptation and the best possible development of humanity, and of the human being, in an uncertain universal environment in permanent evolution, with a mind open enough to consider all the possibilities. In humanist ecological comprehension, it is vain to want to freeze an arbitrary cultural schemata and choose the apparent equilibrium and the supposed future of one moment of evolution. A permanent evolutive adaptation is necessary, as much biologically as mentally. That requires relativity and a caution in the analysis. According to this concept, for the

text: human mind

text: , any representation belongs to the domain of belief, considering the uncertainty of the relation of man to the universe, and the natural imperfection of his senses to represent his environment and his interactions with this environment; the reality perceived by Man being only one representation of reality, particular in mankind. Humanist ecology admits this relativized relation with

text: belief

text: but refutes any final and locking form, knowing that no truth can be final for the human mind without upsetting its natural evolutive necessity. This

text: school of thought

text: accepts belief in the present, for want of anything better, but dictates that you must take care to verify and update beliefs.

text: New genes arise from several different genetic mechanisms including gene duplication, de novo origination, retrotransposition, chimeric gene formation, recruitment of non-coding sequence, and gene truncation.

text: There is an alternate strategy for identifying fit mates which does not rely on one gender having exaggerated sexual ornaments or other handicaps, but is probably generally applicable to most, if not all sexual creatures. It derives from the concept that the change in appearance and functionality caused by a non-silent mutation will generally stand out in a population. This is because that altered appearance and functionality will be unusual, peculiar, and different from the norm within that population. The norm against which these unusual features are judged is made up of fit attributes that have attained their plurality through natural selection, while less well adapted attributes will be in the minority or frankly rare. Since the overwhelming majority of mutant features are maladaptive, and it is impossible to predict evolution's future direction, sexual creatures would be expected to prefer mates with the least number of unusual or minority features. This will have the effect of a sexual population rapidly shedding peripheral phenotypic features, thereby canalizing the entire outward appearance and behavior of all of its members. They will all very quickly begin to look remarkably similar to one another in every detail, as illustrated in the accompanying photograph of the African pygmy kingfisher, Ispidina picta. Once a population has become as homogeneous in appearance as is typical of most species, its entire repertoire of behaviors will also be rendered evolutionarily stable, including any cooperative, altruistic and social interactions. Thus, in the example above of the selfish individual who hangs back from the rest of the hunting pack, but who nevertheless joins in the spoils, that individual will be recognized as being different from the norm, and will therefore find it difficult to attract a mate. Its genes will therefore have only a very small probability of being passed on to the next generation, thus evolutionarily stabilizing cooperation and social interactions at whatever level of complexity is the norm in that population.

text: There is an alternate strategy for identifying fit mates which does not rely on one gender having exaggerated sexual ornaments or other handicaps, but is probably generally applicable to most, if not all sexual creatures. It derives from the concept that the change in appearance and functionality caused by a

text: non-silent

text: mutation

text: will generally stand out in a population. This is because that altered appearance and functionality will be unusual, peculiar, and different from the norm within that population. The norm against which these unusual features are judged is made up of fit attributes that have attained their plurality through

text: natural selection

text: , while less well adapted attributes will be in the minority or frankly rare. Since the overwhelming majority of mutant features are maladaptive, and it is impossible to predict evolution's future direction, sexual creatures would be expected to

text: prefer mates with the least number of unusual or minority features

text: . This will have the effect of a sexual population rapidly shedding peripheral phenotypic features, thereby canalizing the entire outward appearance and behavior of all of its members. They will all very quickly begin to look remarkably similar to one another in every detail, as illustrated in the accompanying photograph of the

text: African pygmy kingfisher

text: , Ispidina picta. Once a population has become as homogeneous in appearance as is typical of most species, its entire repertoire of behaviors will also be rendered

text: evolutionarily stable

text: , including any cooperative, altruistic and social interactions. Thus, in the example

text: above

text: of the selfish individual who hangs back from the rest of the hunting pack, but who nevertheless joins in the spoils, that individual will be recognized as being different from the norm, and will therefore

text: find it difficult to attract a mate

text: . Its genes will therefore have only a very small probability of being passed on to the next generation, thus evolutionarily stabilizing cooperation and social interactions at whatever level of complexity is the norm in that population.

text: A core concept in geobiology is that life changes over time through evolution. The theory of evolution postulates that unique populations of organisms or species arose from genetic modifications in the ancestral population which were passed down by drift and natural selection.

text: A core concept in geobiology is that life changes over time through

text: evolution

text: . The theory of evolution postulates that unique populations of organisms or

text: species

text: arose from genetic modifications in the ancestral population which were passed down by

text: drift

text: and

text: natural selection

text: Along with standard biological evolution, life and planet co-evolve. Since the best adaptations are those that suit the ecological niche that the organism lives in, the physical and chemical characteristics of the environment drive the evolution of life by natural selection, but the opposite can also be true: with every advent of evolution, the environment changes.

text: Along with standard biological evolution, life and planet co-evolve. Since the best adaptations are those that suit the

text: ecological niche

text: that the organism lives in, the physical and chemical characteristics of the environment drive the evolution of life by natural selection, but the opposite can also be true: with every advent of evolution, the environment changes.

text: The concept of a gene continues to be refined as new phenomena are discovered. For example, regulatory regions of a gene can be far removed from its coding regions, and coding regions can be split into several exons. Some viruses store their genome in RNA instead of DNA and some gene products are functional non-coding RNAs. Therefore, a broad, modern working definition of a gene is any discrete locus of heritable, genomic sequence which affect an organism's traits by being expressed as a functional product or by regulation of gene expression.

text: The concept of a gene continues to be refined as new phenomena are discovered. For example,

text: regulatory regions

text: of a gene can be far removed from its

text: coding region

text: s, and coding regions can be split into several

text: exon

text: s. Some

text: viruses

text: store their

text: genome

text: in

text: RNA

text: instead of DNA and some gene products are functional

text: non-coding RNA

text: s. Therefore, a broad, modern working definition of a gene is any discrete locus of heritable, genomic sequence which affect an organism's traits by being

text: expressed

text: as a functional product or by

text: regulation of gene expression

text: The genome of a given organism contains thousands of genes, but not all these genes need to be active at any given moment. A gene is expressed when it is being transcribed into mRNA and there exist many cellular methods of controlling the expression of genes such that proteins are produced only when needed by the cell. Transcription factors are regulatory proteins that bind to DNA, either promoting or inhibiting the transcription of a gene. Within the genome of Escherichia coli bacteria, for example, there exists a series of genes necessary for the synthesis of the amino acid tryptophan. However, when tryptophan is already available to the cell, these genes for tryptophan synthesis are no longer needed. The presence of tryptophan directly affects the activity of the genes—tryptophan molecules bind to the tryptophan repressor (a transcription factor), changing the repressor's structure such that the repressor binds to the genes. The tryptophan repressor blocks the transcription and expression of the genes, thereby creating negative feedback regulation of the tryptophan synthesis process.

text: The genome of a given organism contains thousands of genes, but not all these genes need to be active at any given moment. A gene is

text: expressed

text: when it is being transcribed into mRNA and there exist many cellular methods of controlling the expression of genes such that proteins are produced only when needed by the cell.

text: Transcription factor

text: s are regulatory proteins that bind to DNA, either promoting or inhibiting the transcription of a gene. Within the genome of

text: Escherichia coli

text: bacteria, for example, there exists a series of genes necessary for the synthesis of the amino acid

text: tryptophan

text: . However, when tryptophan is already available to the cell, these genes for tryptophan synthesis are no longer needed. The presence of tryptophan directly affects the activity of the genes—tryptophan molecules bind to the

text: tryptophan repressor

text: (a transcription factor), changing the repressor's structure such that the repressor binds to the genes. The tryptophan repressor blocks the transcription and expression of the genes, thereby creating

text: negative feedback

text: regulation of the tryptophan synthesis process.

text: It is believed that a sub-population of Lahontan cutthroats became isolated in Silver King Creek above Silver King Canyon Gorge after erosion made the gorge impassable to trout swimming upstream, probably between 5,000 and 8,000 years ago. The upstream population then adapted to local conditions independent of the larger population below that had evolved in pluvial Lake Lahontan during the Pleistocene. Although Lahontan cutthroats are heavily spotted, the isolated sub-population lost virtually all spotting, perhaps because spots made fish more visible and susceptible to predation in the ultra-clear and shallow mountain stream. Paiute cutthroats are also notable for a purple coloration, whereas Lahontan cutthroats have bronze coloration. The Paiute strain must have adapted to a diet mainly of insects and become less migratory since juvenile fish swimming downstream in search of larger waters would have passed below downstream barriers and left the isolated gene pool.

text: It is believed that a sub-population of Lahontan cutthroats became isolated in Silver King Creek above Silver King Canyon Gorge after erosion made the gorge impassable to trout swimming upstream, probably between 5,000 and 8,000 years ago. The upstream population then adapted to local conditions independent of the larger population below that had evolved in

text: pluvial

text: Lake Lahontan

text: during the

text: Pleistocene

text: . Although Lahontan cutthroats are heavily spotted, the isolated sub-population lost virtually all spotting, perhaps because spots made fish more visible and susceptible to predation in the ultra-clear and shallow mountain stream. Paiute cutthroats are also notable for a purple coloration, whereas Lahontan cutthroats have bronze coloration. The Paiute strain must have adapted to a diet mainly of insects and become less migratory since juvenile fish swimming downstream in search of larger waters would have passed below downstream barriers and left the isolated gene pool.

text: The development and spread of antibiotic-resistant bacteria  is evidence for the process of evolution of species. Thus the appearance of vancomycin-resistant Staphylococcus aureus, and the danger it poses to hospital patients, is a direct result of evolution through natural selection. The rise of Shigella strains resistant to the synthetic antibiotic class of sulfonamides also demonstrates the generation of new information as an evolutionary process. Similarly, the appearance of DDT resistance in various forms of Anopheles mosquitoes, and the appearance of myxomatosis resistance in breeding rabbit populations in Australia, are both evidence of the existence of evolution in situations of evolutionary selection pressure in species in which generations occur rapidly.

text: The development and spread of

text: antibiotic

text: -resistant

text: bacteria

text: is evidence for the process of evolution of species. Thus the appearance of

text: vancomycin

text: -resistant

text: Staphylococcus aureus

text: , and the danger it poses to hospital patients, is a direct result of evolution through natural selection. The rise of

text: Shigella

text: strains resistant to the synthetic antibiotic class of

text: sulfonamides

text: also demonstrates the generation of new information as an evolutionary process. Similarly, the appearance of

text: DDT

text: resistance in various forms of

text: Anopheles

text: mosquitoes, and the appearance of

text: myxomatosis

text: resistance in breeding rabbit populations in Australia, are both evidence of the existence of evolution in situations of evolutionary

text: selection pressure

text: in species in which generations occur rapidly.

text: By definition, all genes in a neutral network have equivalent function, however some may exhibit promiscuous activities which could serve as starting points for adaptive evolution towards new functions. In terms of sequence space, current theories predict that if the neutral networks for two different activities overlap, a neutrally evolving population may diffuse to regions of the neutral network of the first activity that allow it to access the second. This would only be the case when the distance between activities is smaller than the distance that a neutrally evolving population can cover. The degree of interpenetration of the two networks will determine how common cryptic variation for the promiscuous activity is in sequence space.

text: By definition, all genes in a neutral network have equivalent function, however some may exhibit

text: promiscuous activities

text: which could serve as starting points for adaptive evolution towards new functions. In terms of

text: sequence space

text: , current theories predict that if the neutral networks for two different activities overlap, a neutrally evolving population may diffuse to regions of the neutral network of the first activity that allow it to access the second. This would only be the case when the distance between activities is smaller than the distance that a neutrally evolving population can cover. The degree of interpenetration of the two networks will determine how common cryptic variation for the promiscuous activity is in sequence space.

text: Combined data from coding sequence studies in C. elegans, Drosophila, Humans and Chimps show a similar pattern of molecular evolution in sex biased genes, i.e. most of the male and female biased genes when compared to genes equally expressed in both had higher Ka/Ks ratio. Male biased genes show greater divergence than female biased genes. The Ka/Ks ratio was higher for male biased genes which are expressed exclusively in reproductive tissues e.g. testis in primate lineages. In C. elegans, which is an androdioecious species (a population consisting of only hermaphrodites and males), the rate of evolution for genes expressed during spermatogenesis was higher in males than in hermaphrodites. In Drosophila, interspecies divergence was found to be higher than intraspecific polymorphism at non synonymous sites of male biased genes which elucidated the role of positive selection and showed that male biased genes undergo frequent adaptive evolution. It should be noted here that although positive evolution is associated with most of the male and female biased genes, it’s difficult to isolate genes which shown bias solely due to sexual conflict/antagonism. Nevertheless, since sexually antagonistic genes give rise to biased expression and most biased genes are under positive selection we can argue the same in favor of sexually antagonistic genes. A similar trend as seen in coding sequence evolution was seen with gene expression levels. Interspecific expression divergence was higher than intraspecific expression polymorphism. Positive selection in Acps and Frtps has also been reported previously.

text: Combined data from coding sequence studies in

text: C. elegans

text: Drosophila

text: , Humans and Chimps show a similar pattern of molecular evolution in sex biased genes, i.e. most of the male and female biased genes when compared to genes equally expressed in both had higher

text: Ka/Ks ratio

text: . Male biased genes show greater divergence than female biased genes. The

text: Ka/Ks ratio

text: was higher for male biased genes which are expressed exclusively in reproductive tissues e.g. testis in primate lineages. In

text: C. elegans

text: , which is an

text: androdioecious

text: species (a population consisting of only hermaphrodites and males), the rate of evolution for genes expressed during spermatogenesis was higher in males than in hermaphrodites. In Drosophila, interspecies divergence was found to be higher than intraspecific polymorphism at non synonymous sites of male biased genes which elucidated the role of positive selection and showed that male biased genes undergo frequent adaptive evolution. It should be noted here that although positive evolution is associated with most of the male and female biased genes, it’s difficult to isolate genes which shown bias solely due to sexual conflict/antagonism. Nevertheless, since sexually antagonistic genes give rise to biased expression and most biased genes are under positive selection we can argue the same in favor of sexually antagonistic genes. A similar trend as seen in coding sequence evolution was seen with gene expression levels. Interspecific expression divergence was higher than intraspecific expression polymorphism. Positive selection in Acps and Frtps has also been reported previously.

text: Genes can acquire mutations in their sequence, leading to different variants, known as alleles, in the population. These alleles encode slightly different versions of a protein, which cause different phenotype traits. Colloquial usage of the term "having a gene" (e.g., "good genes," "hair colour gene") typically refers to having a different allele of the gene. Genes evolve due to natural selection or survival of the fittest of the alleles.

text: Genes can acquire

text: mutation

text: s in their sequence, leading to different variants, known as

text: allele

text: s, in the

text: population

text: . These alleles encode slightly different versions of a protein, which cause different

text: phenotype

text: traits. Colloquial usage of the term "having a gene" (e.g., "good genes," "hair colour gene") typically refers to having a different allele of the gene. Genes

text: evolve

text: due to

text: natural selection

text: or

text: survival of the fittest

text: of the alleles.

text: Rapid adaptive evolution in these species leads to offspring that have higher fitness and are better suited for their environment. Intraspecific phenotypic plasticity, pre-adaptation and post-introduction evolution are all major factors in adaptive evolution. Plasticity in populations allows room for changes to better suit the individual in its environment. This is key in adaptive evolution because the main goal is how to best be suited to the ecosystem that the species has been introduced. The ability to accomplish this as quickly as possible will lead to a population with a very high fitness. Pre-adaptations and evolution after the initial introduction also play a role in the success of the introduced species. If the species has adapted to a similar ecosystem or contains traits that happen to be well suited to the area that it is introduced, it is more likely to fare better in the new environment. This, in addition to evolution that takes place after introduction, all determine if the species will be able to become established in the new ecosystem and if it will reproduce and thrive.

text: A broad operational definition is sometimes used to encompass the complexity of these diverse phenomena, where a gene is defined as a union of genomic sequences encoding a coherent set of potentially overlapping functional products. This definition categorizes genes by their functional products (proteins or RNA) rather than their specific DNA loci, with regulatory elements classified as gene-associated regions.

text: The idea of de-evolution is based at least partly on the presumption that "evolution" requires some sort of purposeful direction towards "increasing complexity". Modern evolutionary theory, beginning with Darwin at least, poses no such presumption and the concept of evolutionary change is independent of either any increase in complexity of organisms sharing a gene pool, or any decrease, such as in vestigiality or in loss of genes. Earlier views that species are subject to "cultural decay", "drives to perfection", or "devolution" are practically meaningless in terms of current (neo-)Darwinian theory.  Early scientific theories of transmutation of species such as Lamarckism and orthogenesis perceived species diversity as a result of a purposeful internal drive or tendency to form improved adaptations to the environment. In contrast, Darwinian evolution and its elaboration in the light of subsequent advances in biological research, have shown that adaptation through natural selection comes about when particular heritable attributes in a population happen to give a better chance of successful reproduction in the reigning environment than rival attributes do. By the same process less advantageous attributes are less "successful"; they decrease in frequency or are lost completely. Since Darwin's time it has been shown how these changes in the frequencies of attributes occur according to the mechanisms of genetics and the laws of inheritance originally investigated by Gregor Mendel.  Combined with Darwin's original insights, genetic advances led to what has variously been called the modern evolutionary synthesis or neo-Darwinism. In these terms evolutionary adaptation may occur most obviously through the natural selection of particular alleles. Such alleles may be long established, or they may be new mutations. Selection also might arise from more complex epigenetic or other chromosomal changes, but the fundamental requirement is that any adaptive effect must be heritable.

text: The idea of de-evolution is based at least partly on the presumption that "evolution" requires some sort of

text: purposeful direction

text: towards "

text: increasing complexity

text: ". Modern evolutionary theory, beginning with Darwin at least, poses no such presumption and the concept of evolutionary change is independent of either any increase in complexity of organisms sharing a gene pool, or any decrease, such as in

text: vestigiality

text: or in loss of genes. Earlier views that species are subject to "cultural decay", "drives to perfection", or "devolution" are practically meaningless in terms of current (neo-)Darwinian theory.  Early scientific theories of

text: transmutation of species

text: such as

text: Lamarckism

text: and

text: orthogenesis

text: perceived species diversity as a result of a purposeful internal drive or tendency to form improved adaptations to the environment. In contrast, Darwinian evolution and its elaboration in the light of subsequent advances in biological research, have shown that

text: adaptation

text: through

text: natural selection

text: comes about when particular heritable attributes in a population happen to give a better chance of successful reproduction in the reigning environment than rival attributes do. By the same process less advantageous attributes are less "successful"; they decrease in frequency or are lost completely. Since Darwin's time it has been shown how these changes in the frequencies of attributes occur according to the mechanisms of

text: genetics

text: and the laws of inheritance originally investigated by

text: Gregor Mendel

text: .  Combined with Darwin's original insights, genetic advances led to what has variously been called the

text: modern evolutionary synthesis

text: or neo-Darwinism. In these terms evolutionary adaptation may occur most obviously through the natural selection of particular

text: allele

text: s. Such alleles may be long established, or they may be new

text: mutation

text: s. Selection also might arise from more complex

text: epigenetic

text: or other

text: chromosomal

text: changes, but the fundamental requirement is that any adaptive effect must be

text: heritable

text: Though coral have large sexually-reproducing populations, their evolution can be slowed by abundant asexual reproduction. Gene flow is variable among coral species. According to the biogeography of coral species gene flow cannot be counted on as a dependable source of adaptation as they are very stationary organisms. Also, coral longevity might factor into their adaptivity.

text: Though coral have large sexually-reproducing populations, their evolution can be slowed by abundant

text: asexual reproduction

text: Gene flow

text: is variable among coral species. According to the

text: biogeography

text: of coral species gene flow cannot be counted on as a dependable source of adaptation as they are very stationary organisms. Also, coral longevity might factor into their adaptivity.

text: Natural selection most generally makes nature the measure against which individuals and individual traits, are more or less likely to survive. "Nature" in this sense refers to an ecosystem, that is, a system in which organisms interact with every other element, physical as well as biological, in their local environment. Eugene Odum, a founder of ecology, defined an ecosystem as: "Any unit that includes all of the organisms...in a given area interacting with the physical environment so that a flow of energy leads to clearly defined trophic structure, biotic diversity and material cycles (ie: exchange of materials between living and nonliving parts) within the system." Each population within an ecosystem occupies a distinct niche, or position, with distinct relationships to other parts of the system. These relationships involve the life history of the organism, its position in the food chain and its geographic range. This broad understanding of nature enables scientists to delineate specific forces which, together, comprise natural selection.

text: Natural selection most generally makes nature the measure against which individuals and individual traits, are more or less likely to survive. "Nature" in this sense refers to an

text: ecosystem

text: , that is, a system in which organisms interact with every other element,

text: physical

text: as well as

text: biological

text: , in their local environment.

text: Eugene Odum

text: , a founder of ecology, defined an ecosystem as: "Any unit that includes all of the organisms...in a given area interacting with the physical environment so that a flow of energy leads to clearly defined trophic structure, biotic diversity and material cycles (ie: exchange of materials between living and nonliving parts) within the system." Each population within an ecosystem occupies a distinct

text: niche

text: , or position, with distinct relationships to other parts of the system. These relationships involve the life history of the organism, its position in the

text: food chain

text: and its geographic range. This broad understanding of nature enables scientists to delineate specific forces which, together, comprise natural selection.

text: In general, sexual reproduction in eukaryotes has two fundamental aspects: recombination during meiosis, and outcrossing.  It has been proposed that these two aspects have two natural selective advantages respectively.  A proposed adaptive advantage of meiosis is that it facilitates recombinational repair of DNA damages that are otherwise difficult to repair (see DNA repair as the adaptive advantage of meiosis).  A proposed adaptive advantage of outcrossing is complementation, which is the masking of deleterious recessive alleles  (see hybrid vigor or heterosis).  The selective advantage of complementation may largely account for the general avoidance of inbreeding (see Kin recognition).

text: In general, sexual reproduction in eukaryotes has two fundamental aspects: recombination during meiosis, and outcrossing.  It has been proposed that these two aspects have two natural selective advantages respectively.  A proposed adaptive advantage of meiosis is that it facilitates recombinational repair of DNA damages that are otherwise difficult to repair (see

text: DNA repair as the adaptive advantage of meiosis

text: ).  A proposed adaptive advantage of outcrossing is complementation, which is the masking of deleterious recessive alleles  (see

text: hybrid vigor or heterosis

text: ).  The selective advantage of complementation may largely account for the general avoidance of inbreeding (see

text: Kin recognition

text: ). Consequently, just as a species' biological fitness does not ensure continued survival, neither does rigorous testing protect a scientific theory from refutation in the future. Yet, as it appears that the engine of biological evolution has, over many generations, produced adaptive traits equipped to deal with more and more complex problems of survival, likewise, the evolution of theories through the scientific method may, in Popper's view, reflect a certain type of progress: toward more and more interesting problems (

text: The Nup96 gene is another example of the evolution of the genes implicated in post-copulatory isolation. It regulates the production of one of the approximately 30 proteins required to form a nuclear pore. In each of the simulans groups of Drosophila the protein from this gene interacts with the protein from another, as yet undiscovered, gene on the X chromosome in order to form a functioning pore. However, in a hybrid the pore that is formed is defective and causes sterility. The differences in the sequences of Nup96 have been subject to adaptive selection, similar to the other examples of  speciation genes described above.

text: The Nup96 gene is another example of the evolution of the genes implicated in post-copulatory isolation. It regulates the production of one of the approximately 30 proteins required to form a

text: nuclear pore

text: . In each of the simulans groups of Drosophila the protein from this gene interacts with the protein from another, as yet undiscovered, gene on the X chromosome in order to form a functioning pore. However, in a hybrid the pore that is formed is defective and causes sterility. The differences in the sequences of Nup96 have been subject to adaptive selection, similar to the other examples of  speciation genes described above.

text: In the early 20th century the modern evolutionary synthesis integrated classical genetics with Darwin's theory of evolution by natural selection through the discipline of population genetics. The importance of natural selection as a cause of evolution was accepted into other branches of biology. Moreover, previously held notions about evolution, such as orthogenesis, evolutionism, and other beliefs about innate "progress" within the largest-scale trends in evolution, became obsolete scientific theories. Scientists continue to study various aspects of evolutionary biology by forming and testing hypotheses, constructing mathematical models of theoretical biology and biological theories, using observational data, and performing experiments in both the field and the laboratory.

text: In the early 20th century the

text: modern evolutionary synthesis

text: integrated

text: classical genetics

text: with Darwin's theory of evolution by natural selection through the discipline of

text: population genetics

text: . The importance of natural selection as a cause of evolution was accepted into other branches of

text: biology

text: . Moreover, previously held notions about evolution, such as

text: orthogenesis

text: evolutionism

text: , and other beliefs about innate "progress" within the

text: largest-scale trends in evolution

text: , became

text: obsolete scientific theories

text: . Scientists continue to study various aspects of

text: evolutionary biology

text: by forming and testing hypotheses, constructing

text: mathematical models of theoretical biology

text: and biological theories, using

text: observational data

text: , and performing

text: experiment

text: s in both the

text: field

text: and the laboratory.

text: A classical requirement and constraint of the model is that the number of individuals within a population that live to reach senescence must be small in number. If this is not true for a population, then the effects of old age will not be under a selection shadow and instead affect adaptation and evolution of the population as a whole. At the same time, however, this requirement has been challenged by increasing evidence of senescence being more common in wild populations than previously expected, especially among birds and mammals, while the effects of the selection shadow remain present.

text: A classical requirement and

text: constraint

text: of the model is that the number of individuals within a population that live to reach

text: senescence

text: must be small in number. If this is not true for a population, then the effects of old age will not be under a selection shadow and instead affect adaptation and evolution of the population as a whole. At the same time, however, this requirement has been challenged by increasing evidence of senescence being more common in wild populations than previously expected, especially among birds and mammals, while the effects of the selection shadow remain present.

text: While B. lusitaniae is distributed throughout countries in Europe and North Africa, it is the sole species of the Lyme borreliosis group in southern Portugal. Lizards of the family Lacertidae are believed to be important reservoir hosts of B. lusitaniae.  They were first isolated in Portugal in 1993. These lizards that include the sand lizard and common wall lizard are known to be highly structured phylogeographically.  Migration is very limited between the lizard populations from different localities and it has shed light into the evolution and epidemiology of B. lusitaniae. The pronounced population structure of B. lusitaniae over a short geographic distance (southern Portugal) by housekeeping genes indicates that the migration rates of B. lusitaniae are rather low, because the distribution of Mediterranean lizard populations is highly parapatric.

text: While B. lusitaniae is distributed throughout countries in Europe and North Africa, it is the sole species of the Lyme borreliosis group in southern

text: Portugal

text: . Lizards of the family Lacertidae are believed to be important reservoir hosts of B. lusitaniae.  They were first isolated in Portugal in 1993. These lizards that include the

text: sand lizard

text: and

text: common wall lizard

text: are known to be highly structured phylogeographically.  Migration is very limited between the lizard populations from different localities and it has shed light into the evolution and

text: epidemiology

text: of B. lusitaniae. The pronounced population structure of B. lusitaniae over a short geographic distance (southern Portugal) by

text: housekeeping genes

text: indicates that the migration rates of B. lusitaniae are rather low, because the distribution of Mediterranean lizard populations is highly

text: parapatric

text: When mutational robustness exists, many mutants will persist in a cryptic state. Mutations tend to fall into two categories, having either a very bad effect or very little effect: few mutations fall somewhere in between. Sometimes, these mutations will not be completely invisible, but still have rare effects, with very low penetrance. When this happens, natural selection weeds out the very bad mutations, while leaving the others relatively unaffected. While evolution has no "foresight" to know which environment will be encountered in the future, some mutations cause major disruption to a basic biological process, and will never be adaptive in any environment. Screening these out in advance leads to preadapted stocks of cryptic genetic variation.

text: When

text: mutational robustness

text: exists, many mutants will persist in a cryptic state. Mutations tend to fall into two categories, having either a very bad effect or very little effect: few mutations fall somewhere in between. Sometimes, these mutations will not be completely invisible, but still have rare effects, with very low

text: penetrance

text: . When this happens, natural selection weeds out the very bad mutations, while leaving the others relatively unaffected. While evolution has no "foresight" to know which environment will be encountered in the future, some mutations cause major disruption to a basic biological process, and will never be adaptive in any environment. Screening these out in advance leads to

text: preadapted

text: stocks of cryptic genetic variation.

text: By promoting a perception that evolution is the subject of wide controversy and debate within the scientific community, whereas in fact evolution is overwhelmingly supported by scientists, the list is used to lend support to other Discovery Institute campaigns promoting intelligent design, including "Teach the Controversy", "Critical Analysis of Evolution", "Free Speech on Evolution", and "Stand Up For Science". For example, in its "Teach the Controversy" campaign, the Institute claims that "evolution is a theory in crisis" and that many scientists criticize evolution and citing the list as evidence or a resource. The Discovery Institute also asserts that this information is being withheld from students in public high school science classes along with "alternatives" to evolution such as intelligent design. The Institute uses A Scientific Dissent From Darwinism as evidence to support its claim that evolution is disputed widely within the scientific community. In 2002, Stephen C. Meyer, the founder of the Discovery Institute's Center for Science and Culture, presented the list as evidence to the Ohio Board of Education to promote Teach the Controversy. He cited it as demonstrating that there was a genuine controversy over Darwinian evolution. In the 2005 Kansas evolution hearings Meyer cited the list in support of his assertion that there was "significant scientific dissent from Darwinism" that students should be informed about.

text: By promoting a perception that evolution is the subject of wide controversy and debate within the scientific community, whereas in fact evolution is overwhelmingly supported by scientists, the list is used to lend support to other

text: Discovery Institute

entity_name: Discovery Institute intelligent design campaigns

text: campaigns

text: promoting

text: intelligent design

text: , including "

text: Teach the Controversy

text: Critical Analysis of Evolution

text: Free Speech on Evolution

text: ", and "

text: Stand Up For Science

text: ". For example, in its "Teach the Controversy" campaign, the Institute claims that "evolution is a theory in crisis" and that many scientists criticize evolution and citing the list as evidence or a resource. The Discovery Institute also asserts that this information is being withheld from students in public high school science classes along with "alternatives" to evolution such as intelligent design. The Institute uses A Scientific Dissent From Darwinism as evidence to support its claim that evolution is disputed widely within the scientific community. In 2002,

text: Stephen C. Meyer

text: , the founder of the Discovery Institute's

text: Center for Science and Culture

text: , presented the list as evidence to the

text: Ohio Board of Education

text: to promote

text: Teach the Controversy

text: . He cited it as demonstrating that there was a genuine controversy over Darwinian evolution. In the 2005

text: Kansas evolution hearings

text: Meyer cited the list in support of his assertion that there was "significant scientific dissent from Darwinism" that students should be informed about.

text: The total complement of genes in an organism or cell is known as its genome, which may be stored on one or more chromosomes. A chromosome consists of a single, very long DNA helix on which thousands of genes are encoded. The region of the chromosome at which a particular gene is located is called its locus. Each locus contains one allele of a gene; however, members of a population may have different alleles at the locus, each with a slightly different gene sequence.

text: The total complement of genes in an organism or cell is known as its

text: genome

text: , which may be stored on one or more

text: chromosome

text: s. A chromosome consists of a single, very long DNA helix on which thousands of genes are encoded. The region of the chromosome at which a particular gene is located is called its

text: locus

text: . Each locus contains one

text: allele

text: of a gene; however, members of a population may have different alleles at the locus, each with a slightly different gene sequence.

text: Docofossor had a reduced number of phalanges in its fingers. The phalangeal formula was 2-2-2-2-2 instead of the ancestral 2-3-3-3-3. This led to shortened but wide digits. Furthermore, the claw-bearing sections were enlarged and the upper phalanges shortened. African golden moles possess almost exactly the same adaptation, which provides an evolutionary advantage for digging mammals. This characteristic is due to the fusion of the bone joints between the upper and middle phalanges during embryonic development – a process influenced by the genes BMP and GDF-5. Because of the many anatomical similarities, the researchers hypothesize that this genetic mechanism may have played a comparable role in early mammaliaform evolution, as in the case of Docofossor.

text: Docofossor had a reduced number of phalanges in its fingers. The phalangeal formula was 2-2-2-2-2 instead of the ancestral 2-3-3-3-3. This led to shortened but wide

text: digits

text: . Furthermore, the claw-bearing sections were enlarged and the upper phalanges shortened. African golden moles possess almost exactly the same adaptation, which provides an evolutionary advantage for digging mammals. This characteristic is due to the fusion of the

text: bone joint

text: s between the upper and middle phalanges during embryonic development – a process influenced by the

text: gene

text: s BMP and GDF-5. Because of the many

text: anatomical

text: similarities, the researchers hypothesize that this genetic mechanism may have played a comparable role in early mammaliaform evolution, as in the case of Docofossor.

text: Natural selection in an environment produces reproductive success, which benefits the species. Speciation is a process of evolution through which new biological species are formed. Dobzhansky studied the specifics of sexual, physiological, and behavioral isolating mechanisms in Drosophila pseudoobscura and Drosophila paulistrorum. Like many of his other studies, Dobzhansky's work on reproductive isolation was aimed at studying the process of evolution in action. Bringing samples from each population back into the laboratory, Dobzhansky showed that he could vary environmental conditions so as to produce the same changes in frequency of inversion patterns that were observed with changing seasons in the field. Dobzhansky concluded that such seasonal fluctuations were the result of natural selection at work, with temperature acting as the selecting agent. These masterful studies provided concrete support for the theory of natural selection, at the same time illustrating the fruitfulness of combining field and laboratory work in the study of evolution. Adaptive evolution occurs through the dominance and survival of competing genes within a species. This is caused by increasing the frequency of those alleles whose phenotypic effects selfishly promote their own reproduction. He also believed new species could not arise from single mutations and must be isolated from others of its species by time, geography, habitat, or breeding season.

text: Thus melanism in rock pocket mice is considered a fabulous example of adaptation by natural selection. Changes in the Mc1r gene sequence are not responsible for the colour difference in the mice sampled from New Mexico, however, leading the researchers to conclude that the almost identical dark coat colours developed multiple times in rock pocket mice, an example of convergent evolution.

text: Thus

text: melanism

text: in rock pocket mice is considered a fabulous example of adaptation by

text: natural selection

text: . Changes in the Mc1r gene sequence are not responsible for the colour difference in the mice sampled from New Mexico, however, leading the researchers to conclude that the almost identical dark coat colours developed multiple times in rock pocket mice, an example of

text: convergent evolution

text: Altruistic behaviour has been explained by the gene-centred view of evolution.

text: Altruistic

text: behaviour has been explained by the

text: gene-centred view of evolution

text: Biologist Paul R. Ehrlich described The Evolution of Human Sexuality as a "classic but controversial treatise on human sexual evolution", and identified Symons' study of the development of human ovulation as a landmark. Biologist Randy Thornhill and anthropologist Craig T. Palmer, writing in A Natural History of Rape (2000), identified Symons as the first author to propose that rape is "a by-product of adaptations designed for attaining sexual access to consenting partners." They wrote that Symons has falsely been accused of basing his arguments on the assumption that behavior is genetically determined, even though he explicitly rejects that assumption and criticizes it at length. Thornhill and Palmer endorsed Symons' argument about male homosexual promiscuity. Evolutionary biologist Stephen Jay Gould, writing in The Structure of Evolutionary Theory (2002), commented that the argument that the clitoris is not adaptive, put forward by Symons and subsequently by Gould himself, has been widely misunderstood as a denial of the adaptive value of the female orgasm in general, or even as a claim that female orgasms lack significance. Anthropologist Melvin Konner called Symons' book "the classic introduction to the evolutionary dimensions" of sex.

text: Biologist

text: Paul R. Ehrlich

text: described The Evolution of Human Sexuality as a "classic but controversial treatise on human sexual evolution", and identified Symons' study of the development of human ovulation as a landmark. Biologist

text: Randy Thornhill

text: and anthropologist Craig T. Palmer, writing in

text: A Natural History of Rape

text: (2000), identified Symons as the first author to propose that rape is "a by-product of adaptations designed for attaining sexual access to consenting partners." They wrote that Symons has falsely been accused of basing his arguments on the assumption that behavior is genetically determined, even though he explicitly rejects that assumption and criticizes it at length. Thornhill and Palmer endorsed Symons' argument about male homosexual promiscuity. Evolutionary biologist

text: Stephen Jay Gould

text: , writing in

text: The Structure of Evolutionary Theory

text: (2002), commented that the argument that the clitoris is not adaptive, put forward by Symons and subsequently by Gould himself, has been widely misunderstood as a denial of the adaptive value of the female orgasm in general, or even as a claim that female orgasms lack significance. Anthropologist

text: Melvin Konner

text: called Symons' book "the classic introduction to the evolutionary dimensions" of sex.

text: Individual leaders of the LDS Church have expressed a variety of opinions on biological evolution.

text: This situation led the Musée de l'Homme to a redefinition of its mission. Jean-Pierre Mohen and his team tried to arrange the mission of the Museum, without really succeeding in giving it a strong enough muséological program. We shall find in the future Museum, the Human defined through his biological evolution, through its adaptation to its environnement, through the elaboration of a culture (by the vector of the communication among others) which defines the highlights of humanity. Finally, it will be question of a conscience of human pressure on its environment as to face the consequences of the evolutions, in the present, for the future.

text: Selection can also be classified by the level or unit of selection. Individual selection acts on the individual, in the sense that adaptations are "for" the benefit of the individual, and result from selection among individuals. Gene selection acts directly at the level of the gene. In kin selection and intragenomic conflict, gene-level selection provides a more apt explanation of the underlying process. Group selection, if it occurs, acts on groups of organisms, on the assumption that groups replicate and mutate in an analogous way to genes and individuals. There is an ongoing debate over the degree to which group selection occurs in nature.

text: Selection can also be classified by the level or

text: unit of selection

text: . Individual selection acts on the individual, in the sense that adaptations are "for" the benefit of the individual, and result from selection among individuals.

text: Gene selection

text: acts directly at the level of the gene. In

text: kin selection

text: and

text: intragenomic conflict

text: , gene-level selection provides a more apt explanation of the underlying process.

text: Group selection

text: , if it occurs, acts on groups of organisms, on the assumption that groups replicate and mutate in an analogous way to genes and individuals. There is an ongoing debate over the degree to which group selection occurs in nature.

text: Mendelian inheritance is a type of biological inheritance that follows the laws originally proposed by Gregor Mendel in 1865 and 1866 and re-discovered in 1900. These laws were initially very controversial. When Mendel's theories were integrated with the Boveri–Sutton chromosome theory of inheritance by Thomas Hunt Morgan in 1915, they became the core of classical genetics. Ronald Fisher later combined these ideas with the theory of natural selection in his 1930 book The Genetical Theory of Natural Selection, putting evolution onto a mathematical footing and forming the basis for population genetics and the modern evolutionary synthesis.

text: Mendelian inheritance is a type of

text: biological

text: inheritance

text: that follows the laws originally proposed by

text: Gregor Mendel

text: in 1865 and 1866 and re-discovered in 1900. These laws were initially very controversial. When Mendel's theories were integrated with the

text: Boveri–Sutton chromosome theory

text: of inheritance by

text: Thomas Hunt Morgan

text: in 1915, they became the core of

text: classical genetics

text: Ronald Fisher

text: later combined these ideas with the theory of

text: natural selection

text: in his 1930 book

entity_name: The Genetical Theory of Natural Selection

text: The Genetical Theory of Natural Selection

text: , putting

text: evolution

text: onto a

text: mathematical

text: footing and forming the basis for

text: population genetics

text: and the

text: modern evolutionary synthesis

text: Weismann's idea was that the relationship between the hereditary material, which he called the germ plasm (German, Keimplasma), and the rest of the body (the soma) was a one-way relationship: the germ-plasm formed the body, but the body did not influence the germ-plasm, except indirectly in its participation in a population subject to natural selection. Weismann was translated into English, and though he was influential, it took many years for the full significance of his work to be appreciated. Later, after the completion of the modern synthesis, the term neo-Darwinism came to be associated with its core concept: evolution, driven by natural selection acting on variation produced by genetic mutation, and genetic recombination (chromosomal crossovers).

text: Weismann's idea was that the relationship between the hereditary material, which he called the

text: germ plasm

text: German

text: , Keimplasma), and the rest of the body (the

text: soma

text: ) was a one-way relationship: the germ-plasm formed the body, but the body did not influence the germ-plasm, except indirectly in its participation in a population subject to natural selection. Weismann was translated into English, and though he was influential, it took many years for the full significance of his work to be appreciated. Later, after the completion of the modern synthesis, the term neo-Darwinism came to be associated with its core concept: evolution, driven by natural selection acting on variation produced by

text: genetic mutation

text: , and

text: genetic recombination

text: chromosomal crossover

text: s).

text: Stebbins was passionate about teaching evolution, advocating during the 1960s and 70s the teaching of Darwinian evolution in public schools. He worked closely with the Biological Sciences Curriculum Study to develop high school curricula based on evolution as the central unifying principle in biology. He also opposed scientific creationism groups. Stebbins was active in numerous science organizations—including the International Union of Biological Sciences, the Western Society of Naturalists, the Botanical Society of America, and the Society for the Study of Evolution—and served as President of the American Society of Naturalists. He was elected to the National Academy of Sciences in 1952. Stebbins received numerous awards for his contributions to science: the National Medal of Science, the Gold Medal from the Linnean Society of London, the Addison Emery Verrill Medal from the Yale Peabody Museum of Natural History, and the John Frederick Lewis Award from the American Philosophical Society. He was elected a Fellow of the American Academy of Arts and Sciences in 1952. He was awarded the 1983 Leidy Award from the Academy of Natural Sciences of Philadelphia.

text: Stebbins was passionate about teaching evolution, advocating during the 1960s and 70s the teaching of Darwinian evolution in public schools. He worked closely with the

text: Biological Sciences Curriculum Study

text: to develop high school curricula based on evolution as the central unifying principle in biology. He also opposed scientific creationism groups. Stebbins was active in numerous science organizations—including the International Union of Biological Sciences, the Western Society of Naturalists, the Botanical Society of America, and the Society for the Study of Evolution—and served as President of the American Society of Naturalists. He was elected to the

text: National Academy of Sciences

text: in 1952. Stebbins received numerous awards for his contributions to science: the

text: National Medal of Science

text: , the Gold Medal from the

text: Linnean Society of London

text: , the Addison Emery Verrill Medal from the Yale

text: Peabody Museum of Natural History

text: , and the John Frederick Lewis Award from the

text: American Philosophical Society

text: . He was elected a Fellow of the

text: American Academy of Arts and Sciences

text: in 1952. He was awarded the 1983

text: Leidy Award

text: from the Academy of Natural Sciences of Philadelphia.

text: Order appears spontaneously in the evolution of language as individual and population behaviour interacts with biological evolution.

text: Order appears spontaneously in the

text: evolution of language

text: as individual and population behaviour interacts with biological evolution.

text: EAC stands for "Escape from Adaptive Conflict". This name first appeared in a publication by Hittinger and Carroll 2007.The evolutionary process described by the EAC model actually begins before the gene duplication event. A singleton (not duplicated) gene evolves towards two beneficial functions simultaneously. This creates an "adaptive conflict" for the gene, since it is unlikely to execute each individual function with maximum efficiency. The intermediate evolutionary result could be a multi-functional gene and after a gene duplication its sub-functions could be carried out by specialised descendants of the gene. The end result would be the same as under the DDC model, two functionally specialised genes (paralogs). In contrast to the DDC model, the EAC model puts more emphasis on the multi-functional pre-duplication state of the evolving genes and gives a slightly different explanation as to why the duplicated multi-functional genes would benefit from additional specialisation after duplication (because of the adaptive conflict of the multi-functional ancestor that needs to be resolved). Under EAC there is an assumption of a positive selection pressure driving evolution after gene duplication, whereas the DDC model only requires neutral ("undirected") evolution to take place, i.e. degeneration and complementation.

text: Anthropogeny is the study of human origins. It is not simply a synonym for human evolution by natural selection, which is only a part of the processes involved in human origins. Many other factors besides biological evolution were involved, ranging over climatic, geographic, ecological, social, and cultural ones. Anthropogenesis, meaning the process or point of becoming human, is also called hominization.

text: Anthropogeny is the study of human origins. It is not simply a synonym for

text: human evolution

text: by

text: natural selection

text: , which is only a part of the processes involved in human origins. Many other factors besides biological

text: evolution

text: were involved, ranging over climatic, geographic, ecological, social, and cultural ones.

text: Anthropogenesis

text: , meaning the process or point of becoming human, is also called

text: hominization

text: According to Williams' 1966 book Adaptation and Natural Selection,

text: According to Williams' 1966 book

text: Adaptation and Natural Selection

text: Evolutionarily stable strategies were originally considered for biological evolution, but they can apply to other contexts. In fact, there are stable states for a large class of adaptive dynamics. As a result, they can be used to explain human behaviours that lack any genetic influences.

text: Evolutionarily stable strategies were originally considered for biological evolution, but they can apply to other contexts. In fact, there are stable states for a large class of

text: adaptive dynamics

text: . As a result, they can be used to explain human behaviours that lack any genetic influences.

text: There are many issues that comprise the Evolution v.s. Creation Debate. Those issues arise from the differences in how both camps believe the universe was formed. Generally speaking, both Evolutionists and Creationists agree that natural selection is the means by which life adapts to its environment. The four areas of evolution–astronomical, geological, chemical, and biological–are what this debate is about.

text: There are many issues that comprise the Evolution v.s. Creation Debate. Those issues arise from the differences in how both camps believe the universe was formed. Generally speaking, both Evolutionists and Creationists agree that natural selection is the means by which life adapts to its environment. The four areas of evolution–

text: astronomical

text: geological

text: chemical

text: , and

text: biological

text: –are what this debate is about.

text: Understanding the changes that have occurred during organism's evolution can reveal the genes needed to construct parts of the body, genes which may be involved in human genetic disorders. For example, the Mexican tetra is an albino cavefish that lost its eyesight during evolution. Breeding together different populations of this blind fish produced some offspring with functional eyes, since different mutations had occurred in the isolated populations that had evolved in different caves. This helped identify genes required for vision and pigmentation, such as crystallins and the melanocortin 1 receptor. Similarly, comparing the genome of the Antarctic icefish, which lacks red blood cells, to close relatives such as the Antarctic rockcod revealed genes needed to make these blood cells.

text: Understanding the changes that have occurred during organism's evolution can reveal the genes needed to construct parts of the body, genes which may be involved in human

text: genetic disorder

text: s. For example, the

text: Mexican tetra

text: is an

text: albino

text: cavefish that lost its eyesight during evolution. Breeding together different populations of this blind fish produced some offspring with functional eyes, since different mutations had occurred in the isolated populations that had evolved in different caves. This helped identify genes required for vision and pigmentation, such as

text: crystallin

text: s and the

text: melanocortin 1 receptor

text: . Similarly, comparing the genome of the

text: Antarctic icefish

text: , which lacks

text: red blood cell

text: s, to close relatives such as the Antarctic rockcod revealed genes needed to make these blood cells.

text: The lecture then moves into an explanation of natural selection, which brings forth designoid objects. To explain natural selection, Dawkins first explains artificial selection by discussing the evolution of wild cabbage into broccoli, cauliflower, cabbage, red cabbage, kohlrabi, and Brussels sprouts. He continues the discussion of artificial selection by explaining the evolution of the ancestral wolf into the many varieties of modern dog. Starting with the ancestral wolf, Dawkins imagines that everyone on one side of the room is breeding for small wolves, while everyone on the other side is breeding for big wolves. By selectively breeding the smallest or largest of each litter for a number of years, you may eventually end up with something like the Chihuahua on one side of the room, and something like a Great Dane on the other side of the room.

text: The lecture then moves into an explanation of

text: natural selection

text: , which brings forth designoid objects. To explain natural selection, Dawkins first explains

text: artificial selection

text: by discussing the evolution of

text: wild cabbage

text: into

text: broccoli

text: cauliflower

text: cabbage

text: red cabbage

text: kohlrabi

text: , and

text: Brussels sprouts

text: . He continues the discussion of artificial selection by explaining the evolution of the ancestral wolf into the many varieties of modern dog. Starting with the ancestral wolf, Dawkins imagines that everyone on one side of the room is breeding for small wolves, while everyone on the other side is breeding for big wolves. By selectively breeding the smallest or largest of each litter for a number of years, you may eventually end up with something like the

text: Chihuahua

text: on one side of the room, and something like a

text: Great Dane

text: on the other side of the room.

text: Ever since 1859, when Charles Darwin published On the Origin of Species, the biological theory of evolution has been significant. The theory posits that the human species is related to all others, living and extinct, and that natural selection is the primary survival factor. This has provided a basis for new beliefs, e.g. social Darwinism, and for new technology, e.g. antibiotics.

text: Ever since 1859, when

text: Charles Darwin

text: published

text: On the Origin of Species

text: , the biological

text: theory

text: of

text: evolution

text: has been significant. The theory posits that the human

text: species

text: is related to all others, living and extinct, and that

text: natural selection

text: is the primary survival factor. This has provided a basis for new beliefs, e.g.

text: social Darwinism

text: , and for new

text: technology

text: , e.g.

text: antibiotics

text: One interesting finding in studies of invasive species has shown that introduced populations have great potential for rapid adaptation and this is used to explain how so many introduced species are able to establish and become invasive in new environments. When bottlenecks and founder effects cause a great decrease in the population size and may constrict genetic variation, the individuals begin to show additive variance as opposed to epistatic variance. This conversion can actually lead to increased variance in the founding populations which then allows for rapid adaptive evolution. Following invasion events, selection may initially act on the capacity to disperse as well as physiological tolerance to the new stressors in the environment. Adaptation then proceeds to respond to the selective pressures of the new environment. These responses would most likely be due to temperature and climate change, or the presence of native species whether it be predator or prey. Adaptations include changes in morphology, physiology, phenology, and plasticity.

text: Adaptation is not always a simple matter, where the ideal phenotype evolves for a given external environment. An organism must be viable at all stages of its development and at all stages of its evolution. This places constraints on the evolution of development, behavior and structure of organisms. The main constraint, over which there has been much debate, is the requirement that each genetic and phenotypic change during evolution should be relatively small, because developmental systems are so complex and interlinked. However, it is not clear what "relatively small" should mean, for example polyploidy in plants is a reasonably common large genetic change. The origin of eukaryotic symbiosis is a more dramatic example.

text: Adaptation is not always a simple matter, where the ideal phenotype evolves for a given external environment. An organism must be viable at all stages of its development and at all stages of its evolution. This places constraints on the evolution of development,

text: behavior

text: and structure of organisms. The main constraint, over which there has been much debate, is the requirement that each

text: genetic

text: and phenotypic change during evolution should be relatively small, because developmental systems are so complex and interlinked. However, it is not clear what "relatively small" should mean, for example

text: polyploid

text: y in plants is a reasonably common large genetic change. The origin of

text: eukaryotic

text: symbiosis

text: is a more dramatic example.

text: For his reviews, which used population genetics to illuminate such anthropological questions as race and intelligence, the biological and cultural components of language, "scientific creationism" relationships among species, and the timetable of human evolution.

text: The adaptive mechanism of Ethiopian highlanders is quite different. This is probably because their migration to the highland was relatively earlier; for example, the Amhara have inhabited altitudes above  for at least 5,000 years and altitudes around  to  for more than 70,000 years. Genomic analysis of two ethnic groups, Amhara and Oromo, revealed that gene variations associated with haemoglobin difference among Tibetans or other variants at the same gene location do not influence the adaptation in Ethiopians. Identification of specific genes further reveals that several candidate genes are involved in Ethiopians, including CBARA1, VAV3, ARNT2 and THRB. Two of these genes (THRB and ARNT2) are known to play a role in the HIF-1 pathway, a pathway implicated in previous work reported in Tibetan and Andean studies. This supports the concept that adaptation to high altitude arose independently among different highlanders as a result of convergent evolution.

text: The adaptive mechanism of Ethiopian highlanders is quite different. This is probably because their migration to the highland was relatively earlier; for example, the

text: Amhara

text: have inhabited altitudes above  for at least 5,000 years and altitudes around  to  for more than 70,000 years. Genomic analysis of two ethnic groups, Amhara and

text: Oromo

text: , revealed that gene variations associated with haemoglobin difference among Tibetans or other variants at the same

text: gene location

text: do not influence the adaptation in Ethiopians. Identification of specific genes further reveals that several candidate genes are involved in Ethiopians, including

text: and

text: THRB

text: . Two of these genes (THRB and ARNT2) are known to play a role in the

text: HIF-1 pathway

text: , a pathway implicated in previous work reported in Tibetan and Andean studies. This supports the concept that adaptation to high altitude arose independently among different highlanders as a result of

text: convergent evolution

text: Human evolution is the part of biological evolution concerning the emergence of anatomically modern humans as a distinct species.

text: Human evolution is the part of biological

text: evolution

text: concerning the emergence of

text: anatomically modern humans

text: as a distinct species.

text: Concomitant with the idea of biological evolution is that of behavioural evolution. Artifacts also evidence a rise from simple to complex. This progression had been anticipated and foreshadowed by ancient concepts of the three-age system. Innovative evolutionists such as Christian Jürgensen Thomsen in Denmark and John Lubbock, 1st Baron Avebury, produced concepts of the Stone Age, the Bronze Age, and the Iron Age, based on the major materials used by man.

text: Concomitant with the idea of biological evolution is that of behavioural evolution. Artifacts also evidence a rise from simple to complex. This progression had been anticipated and foreshadowed by ancient concepts of the

text: three-age system

text: . Innovative evolutionists such as

text: Christian Jürgensen Thomsen

text: in Denmark and

text: John Lubbock, 1st Baron Avebury

text: , produced concepts of the

text: Stone Age

text: , the

text: Bronze Age

text: , and the

text: Iron Age

text: , based on the major materials used by man.

text: In evolutionary biology, an evolutionary arms race is a struggle between competing sets of co-evolving genes, traits, or species, that develop adaptations and counter-adaptations against each other, resembling an arms race. These are often described as examples of positive feedback.  The co-evolving gene sets may be in different species, as in an evolutionary arms race between a predator species and its prey (Vermeij, 1987), or a parasite and its host.  Alternatively, the arms race may be between members of the same species, as in the manipulation/sales resistance model of communication (Dawkins & Krebs, 1979) or as in runaway evolution or Red Queen effects. One example of an evolutionary arms race is in sexual conflict between the sexes, often described with the term Fisherian runaway. Thierry Lodé emphasized the role of such antagonistic interactions in evolution leading to character displacements and antagonist coevolution.  The escalation hypothesis put forward by Geerat Vermeij speaks of more general conflicts and was originally based on his work with marine gastropod fossils.

text: In

text: evolutionary biology

text: , an evolutionary arms race is a struggle between competing sets of

text: co-evolving

text: gene

text: s, traits, or species, that develop

text: adaptation

text: s and counter-adaptations against each other, resembling an

text: arms race

text: . These are often described as examples of

text: positive feedback

text: .  The co-evolving gene sets may be in different

text: species

text: , as in an evolutionary arms race between a

text: predator

text: species and its

text: prey

text: (Vermeij, 1987), or a

text: parasite

text: and its host.  Alternatively, the arms race may be between members of the same species, as in the manipulation/sales resistance model of communication (Dawkins & Krebs, 1979) or as in

text: runaway evolution

text: or

text: Red Queen

text: effects. One example of an evolutionary arms race is in

text: sexual conflict

text: between the sexes, often described with the term

text: Fisherian runaway

text: Thierry Lodé

text: emphasized the role of such antagonistic interactions in

text: evolution

text: leading to

text: character displacement

text: s and antagonist coevolution.  The

text: escalation hypothesis

text: put forward by

text: Geerat Vermeij

text: speaks of more general conflicts and was originally based on his work with marine

text: gastropod

text: fossils.

text: Evolvability is defined as the capacity of a system for adaptive evolution. Evolvability is the ability of a population of organisms to not merely generate genetic diversity, but to generate adaptive genetic diversity, and thereby evolve through natural selection.

text: Evolvability is defined as the capacity of a system for adaptive

text: evolution

text: . Evolvability is the ability of a population of organisms to not merely generate

text: genetic diversity

text: , but to generate

text: adaptive

text: genetic diversity, and thereby evolve through

text: natural selection

text: Research documenting speciation is abundant. Biologists have documented numerous examples of speciation in nature—with evolution having produced far more species than any observer would consider necessary. For example, there are well over 350,000 described species of beetles. Great examples of speciation come from the observations of island biogeography and the process of adaptive radiation, both explained previously. Evidence of common descent can also be found through paleontological studies of speciation within geologic strata. The examples described below represent different modes of speciation and provide strong evidence for common descent. It is important to acknowledge that not all speciation research directly observes divergence from "start-to-finish". This is by virtue of research delimitation and definition ambiguity, and occasionally leads research towards historical reconstructions. In light of this, examples abound, and the following are by no means exhaustive—comprising only a small fraction of the instances observed. Once again, take note of the established fact that, "...natural selection is a ubiquitous part of speciation...", and is the primary driver of speciation, so; hereinafter, examples of speciation will often interdepend and correspond with selection.

text: While the term primarily refers to biological evolution, there are various types of chemical evolution and it is also found in economics, historical linguistics, and many other technical fields where systems develop or change gradually over time, e.g.  stellar evolution, cultural evolution,  the evolution of an idea, metaphysical evolution, spiritual evolution, etc.

text: While the term primarily refers to

text: biological

text: evolution

text: , there are various types of

text: chemical

text: evolution and it is also found in economics, historical linguistics, and many other technical fields where

text: system

text: s develop or change gradually over time, e.g.

text: stellar evolution

text: cultural evolution

text: ,  the

text: evolution of an idea

text: metaphysical evolution

text: spiritual evolution

text: , etc.

text: Abraham Bentsionovich Korol (born October 18, 1946) is a professor in the Institute of Evolution at the University of Haifa. He is a prominent Israeli geneticist and evolutionary biologist known for his work on the evolution of sex and recombination, genome mapping and the genetics of complex traits. Korol was born in Bendery city, Moldavia (now Moldova), then part of the Soviet Union, and immigrated to Israel in 1991. Before immigrating to Israel, Korol was appointed in 1981 as a senior researcher and was awarded the degree of Doctor of Science by the Presidium of Academy of Science USSR in 1988, and became a full professor in 1991. After immigrating to Israel in 1991, Korol has established and headed the Laboratory of Population Genetics and Computational Biology in the Institute of Evolution at the University of Haifa. He became full professor there in 1996 and served as the director of the Institute of Evolution between 2008-2013.Since 1994, Korol has filled many scholarly positions including member of the steering committee of Israeli Gene Bank; member of the Human Genome Organization; member of the European Society of Evolutionary Biology; a member of the Coordinating Committee of the International Wheat Genome Sequencing Consortium; member of the Infrastructure Steering Committee of the Israeli Ministry of Science; representative of Haifa University in the Kamea program steering committee (alef and bet); member of the Advisory Committee of Absorption in Science of the Israeli Ministry of Absorption.

text: Developmental systems theory is plainly radically incompatible with both neo-Darwinism and information processing theory. Whereas neo-Darwinism defines evolution in terms of changes in gene distribution, the possibility that an evolutionarily significant change may arise and be sustained without any directly corresponding change in gene frequencies is an elementary assumption of developmental systems theory, just as neo-Darwinism’s ‘explanation’ of phenomena in terms of reproductive fitness is regarded as fundamentally shallow. Even the widespread mechanistic equation of ‘gene’ with a specific DNA sequence has been thrown into question, as have the analogous interpretations of evolution and adaptation.

text: Through his experiments, Dobzhansky discovers that mutation of genes leads to evolution within a specific species. Adaptations play a large role in genetic drift, and it is known that genes and mutations influence this genetic drift in a particular environment. Mutations can result from external influences within the surrounding environment, especially if an organism inhabits an area with harsh living conditions. An organism can adapt to its environment in order to better suit its own needs. When an organism successfully adapts, it has higher survival and reproduction rates. Therefore, there is a higher chance that its genes will be passed on to its offspring. Specific genes and alleles are then passed on to future generations to continue the trend of modern evolution that Dobzhansky presents in the book.

text: In order for a biological organism to evolve by natural selection, there must be a certain minimum probability that new, heritable variants are beneficial. Random mutations, unless they occur in DNA sequences with no function, are expected to be mostly detrimental. Beneficial mutations are always rare, but if they are too rare, then adaptation cannot occur. Early failed efforts to evolve computer programs by random mutation and selection showed that evolvability is not a given, but depends on the representation of the program.  Analogously, the evolvability of organisms depends on their genotype-phenotype map. This means that biological genomes are structured in ways that make beneficial changes less unlikely than they would otherwise be. This has been taken as evidence that evolution has created not just fitter organisms, but populations of organisms that are better able to evolve.

text: In order for a biological organism to evolve by natural selection, there must be a certain minimum probability that new, heritable variants are beneficial. Random mutations, unless they occur in DNA sequences with no function, are expected to be mostly detrimental. Beneficial mutations are always rare, but if they are too rare, then

text: adaptation

text: cannot occur. Early failed efforts to evolve computer programs by random mutation and selection showed that evolvability is not a given, but depends on the representation of the program.  Analogously, the evolvability of organisms depends on their genotype-phenotype map. This means that biological genomes are structured in ways that make beneficial changes less unlikely than they would otherwise be. This has been taken as evidence that evolution has created not just fitter organisms, but populations of organisms that are better able to evolve.

text: His experimental contributions include studies of evolution of wing shape in the genus Drosophila, the evolution of the ability to evolve, and adaptation under natural and sexual selection. His theoretical contributions have included work on good genes mechanisms in sexual selection, the evolution of variance-covariance matrices, the detection of evolutionary constraints, and the use of fluctuating asymmetry as an indicator of developmental stability.

text: Bioinformatics is both a term for the body of biological gene disease studies that use computer programming as part of their methodology, as well as a reference to specific analysis pipelines that are repeatedly used, particularly in the fields of genetics and genomics. Common uses of bioinformatics include the identification of candidate genes and nucleotides, SNPs. Often, such identification is made with the aim of better understanding the genetic basis of disease, unique adaptations, desirable properties, or differences between populations. In a less formal way, bioinformatics also tries to understand the organisational principles within nucleic acid and protein sequences.

text: Bioinformatics is both a term for the body of biological gene disease studies that use computer programming as part of their methodology, as well as a reference to specific analysis pipelines that are repeatedly used, particularly in the fields of genetics and genomics. Common uses of bioinformatics include the identification of candidate genes and nucleotides,

text: SNPs

text: . Often, such identification is made with the aim of better understanding the genetic basis of disease, unique adaptations, desirable properties, or differences between populations. In a less formal way, bioinformatics also tries to understand the organisational principles within nucleic acid and protein sequences.

text: Arthur M. Young and Edward Haskell have each independently incorporated the findings of science into a larger theory of spiritual evolution, and extended the traditional human, animal, vegetable, and mineral categories with kingdoms representing photons, atoms and molecules. Arthur M. Young goes further in considering the human state as a subset of a larger kingdom of "Dominion", of which the sixth stage is represented for example by Christ and Buddha, and the seventh (final) stage an even higher level of Enlightenment or God-realisation. Moreover, both Haskell and Young present profound accounts of evolution through these kingdoms in terms of cybernetic principles. A more "mainstream" scientific presentation of this same idea is provided by Erich Jantsch in his account of how self-organising systems evolve and develop as a series of "symmetry breaks" through the sequence of matter, life, and mind. Although abiding strictly by the understanding of science, Jantsch arranges the various elements of cosmic, planetary, biological, psychological, and human evolution in a single overall framework of emergent evolution that may or may not be considered teleological.

text: Arthur M. Young

text: and

text: Edward Haskell

text: have each independently incorporated the findings of science into a larger theory of spiritual evolution, and extended the traditional human, animal, vegetable, and mineral categories with kingdoms representing

text: photon

text: s,

text: atom

text: s and

text: molecule

text: s. Arthur M. Young goes further in considering the human state as a subset of a larger kingdom of "Dominion", of which the sixth stage is represented for example by Christ and Buddha, and the seventh (final) stage an even higher level of Enlightenment or God-realisation. Moreover, both Haskell and Young present profound accounts of evolution through these kingdoms in terms of

text: cybernetic

text: principles. A more "mainstream" scientific presentation of this same idea is provided by

text: Erich Jantsch

text: in his account of how self-organising systems evolve and develop as a series of "

text: symmetry break

text: s" through the sequence of matter, life, and mind. Although abiding strictly by the understanding of science, Jantsch arranges the various elements of cosmic, planetary, biological, psychological, and human evolution in a single overall framework of

text: emergent evolution

text: that may or may not be considered teleological.

text: One of the main sources of confusion and ambiguity in the creation–evolution debate is the definition of evolution itself. In the context of biology, evolution is genetic changes in populations of organisms over successive generations. However, the word has a number of different meanings in different fields, from evolutionary computation to molecular evolution to sociocultural evolution to stellar and galactic evolution. When biological evolution is conflated with other evolutionary processes, this can cause errors such as the claim that modern evolutionary theory says anything about abiogenesis or the Big Bang.

text: One of the main sources of confusion and ambiguity in the creation–evolution debate is the definition of evolution itself. In the context of biology, evolution is genetic changes in populations of organisms over successive generations. However, the word has a number of different meanings in different fields, from

text: evolutionary computation

text: to

text: molecular evolution

text: to

text: sociocultural evolution

text: to

text: stellar

text: and

text: galactic evolution

text: . When biological evolution is conflated with other evolutionary processes, this can cause errors such as the claim that modern evolutionary theory says anything about

text: abiogenesis

text: or the

text: Big Bang

text: The term natural selection is most often defined to operate on heritable traits, because these directly participate in evolution. However, natural selection is "blind" in the sense that changes in phenotype can give a reproductive advantage regardless of whether or not the trait is heritable. Following Darwin's primary usage, the term is used to refer both to the evolutionary consequence of blind selection and to its mechanisms. It is sometimes helpful to explicitly distinguish between selection's mechanisms and its effects; when this distinction is important, scientists define "(phenotypic) natural selection" specifically as "those mechanisms that contribute to the selection of individuals that reproduce", without regard to whether the basis of the selection is heritable. Traits that cause greater reproductive success of an organism are said to be selected for, while those that reduce success are selected against.

text: It has not been proven that genetic assimilation occurs in natural evolution, but it is difficult to rule it out from having at least a minor role, and research continues into the question.  Mathematical modeling suggests that under certain circumstances, natural selection will favor the evolution of canalization that is designed to fail under extreme conditions.  If the result of such a failure is favored by natural selection, genetic assimilation will occur.  In the 1960s C. H. Waddington and J. M. Rendel argued for the importance of genetic assimilation in natural adaptation as a means of providing new and potentially beneficial variation to populations under stress.  Their contemporary Williams argued that genetic assimilation proceeds at the cost of a loss of developmental plasticity, and should be a minor mechanism.  If it occurs frequently, genetic assimilation could contribute to punctuated equilibrium in evolution, as organisms repeatedly evolve systems of canalization, then break out of them under adverse circumstances.

text: It has not been proven that genetic assimilation occurs in natural evolution, but it is difficult to rule it out from having at least a minor role, and research continues into the question.  Mathematical modeling suggests that under certain circumstances, natural selection will favor the evolution of canalization that is designed to fail under extreme conditions.  If the result of such a failure is favored by natural selection, genetic assimilation will occur.  In the 1960s

text: C. H. Waddington

text: and J. M. Rendel argued for the importance of genetic assimilation in natural adaptation as a means of providing new and potentially beneficial variation to populations under stress.  Their contemporary Williams argued that genetic assimilation proceeds at the cost of a loss of developmental plasticity, and should be a minor mechanism.  If it occurs frequently, genetic assimilation could contribute to

text: punctuated equilibrium

text: in evolution, as organisms repeatedly evolve systems of canalization, then break out of them under adverse circumstances.

text: A deme in biological evolution is conceptually related to a meme in cultural evolution.

text: A deme in biological evolution is conceptually related to a

text: meme

text: in cultural evolution.

text: 247 genes in the testes showed evidence of adaptive evolution in the study of Nielsen et al. (2005a). This could be partially due to sexual antagonism. Male-female competition could facilitate an arms race of adaptive evolution. However, in this situation you would expect to find evidence of adaptive evolution in the female sexual organs also, but there is less evidence of this. Sperm competition is another possible explanation. Sperm competition is strong, and sperm can improve their chances of fertilising the female egg in a variety of ways, including increasing their speed, stamina or response to chemoattractants (Swanson and Vacquier 2002).

text: 247 genes in the testes showed evidence of adaptive evolution in the study of Nielsen et al. (2005a). This could be partially due to sexual antagonism. Male-female competition could facilitate an arms race of adaptive evolution. However, in this situation you would expect to find evidence of adaptive evolution in the female sexual organs also, but there is less evidence of this.

text: Sperm competition

text: is another possible explanation. Sperm competition is strong, and sperm can improve their chances of fertilising the female egg in a variety of ways, including increasing their speed, stamina or response to chemoattractants (Swanson and Vacquier 2002).

text: Lenski notes that society and culture evolve through symbols, which makes this process much more rapid, deliberative, and purposeful, compared to biological evolution. However, just like in the biological survival of the fittest, in sociocultural evolution there is a process of intersocietal selection, where less fit sociocultural systems became extinct, replaced by more efficient ones. In another analogy to biological evolution, Lenski argues that the sociocultural systems that survive do so primarily on the virtue of their level of technological advancement, which otherwise is "blind and purposeless as the outcome of the biological process of natural selection and just as indifferent to humans beliefs and values". Thus, Lenski does not argue that societies that survive are morally superior to those that are defeated—simply that they were more efficient at technological progress. This, in turn, helps those societies survive and grow. Technologies can thus be compared to genes of a society, providing it with new abilities, as new technologies allow it to do things that it could not have done before.

text: Lenski notes that society and culture evolve through symbols, which makes this process much more rapid, deliberative, and purposeful, compared to

text: biological evolution

text: . However, just like in the biological

text: survival of the fittest

text: , in sociocultural evolution there is a process of intersocietal selection, where less fit sociocultural systems became extinct, replaced by more efficient ones. In another analogy to biological evolution, Lenski argues that the sociocultural systems that survive do so primarily on the virtue of their level of technological advancement, which otherwise is "blind and purposeless as the outcome of the biological process of natural selection and just as indifferent to humans beliefs and values". Thus, Lenski does not argue that societies that survive are morally superior to those that are defeated—simply that they were more efficient at

text: technological progress

text: . This, in turn, helps those societies survive and grow. Technologies can thus be compared to

text: gene

text: s of a society, providing it with new abilities, as new technologies allow it to do things that it could not have done before.

text: Natural selection is fundamental to evolutionary theory. Variants of hereditary traits which increase an organism's ability to survive and reproduce will be more greatly represented in subsequent generations, i.e., they will be "selected for". Thus, inherited behavioral mechanisms that allowed an organism a greater chance of surviving and/or reproducing in the past are more likely to survive in present organisms. That inherited adaptive behaviors are present in nonhuman animal species has been multiply demonstrated by biologists, and it has become a foundation of evolutionary biology. However, there is continued resistance by some researchers over the application of evolutionary models to humans, particularly from within the social sciences, where culture has long been assumed to be the predominant driver of behavior.

text: Natural selection

text: is fundamental to evolutionary theory. Variants of hereditary traits which increase an organism's ability to survive and reproduce will be more greatly represented in subsequent generations, i.e., they will be "selected for". Thus, inherited behavioral mechanisms that allowed an

text: organism

text: a greater chance of surviving and/or reproducing in the past are more likely to survive in present organisms. That inherited adaptive behaviors are present in nonhuman

text: animal species

text: has been multiply demonstrated by biologists, and it has become a foundation of

text: evolutionary biology

text: . However, there is continued resistance by some researchers over the application of evolutionary models to humans, particularly from within the social sciences, where culture has long been assumed to be the predominant driver of behavior.

text: The individual in which this chromosome lies will now have a selective advantage over other individuals of this species, so the allele A* will spread through the population by the normal processes of natural selection. M, due to its proximity to A*, will be dragged through into the general population. This process only works when M is very close to the allele it has mutated. A greater distance would increase the chance of recombination separating M from A*, leaving M alone with any deleterious mutations it may have caused. For this reason, evolution of mutators is generally expected to happen largely in asexual species where recombination cannot disrupt linkage disequilibrium.

text: The individual in which this chromosome lies will now have a selective advantage over other individuals of this species, so the allele A* will spread through the population by the normal processes of

text: natural selection

text: . M, due to its proximity to A*, will be dragged through into the general population. This process only works when M is very close to the allele it has mutated. A greater distance would increase the chance of recombination separating M from A*, leaving M alone with any deleterious mutations it may have caused. For this reason, evolution of mutators is generally expected to happen largely in

text: asexual

text: species where recombination cannot disrupt linkage disequilibrium.

text: By the 1850s, whether or not species evolved was a subject of intense debate, with prominent scientists arguing both sides of the issue. The publication of Charles Darwin's On the Origin of Species fundamentally transformed the discussion over biological origins. Darwin argued that his branching version of evolution explained a wealth of facts in biogeography, anatomy, embryology, and other fields of biology. He also provided the first cogent mechanism by which evolutionary change could persist: his theory of natural selection.

text: Quantum evolution was a controversial hypothesis advanced by Columbia University paleontologist George Gaylord Simpson, who was regarded by Stephen Jay Gould as "the greatest and most biologically astute paleontologist of the twentieth century." Simpson's conjecture was that according to the geological record, on very rare occasions evolution would proceed very rapidly to form entirely new families, orders, and classes of organisms. This hypothesis differs from punctuated equilibrium in several respects. First, punctuated equilibrium was more modest in scope, in that it was addressing evolution specifically at the species level. Simpson's idea was principally concerned with evolution at higher taxonomic groups. Second, Eldredge and Gould relied upon a different mechanism. Where Simpson relied upon a synergistic interaction between genetic drift and a shift in the adaptive fitness landscape, Eldredge and Gould relied upon ordinary speciation, particularly Ernst Mayr's concept of allopatric speciation. Lastly, and perhaps most significantly, quantum evolution took no position on the issue of stasis. Although Simpson acknowledged the existence of stasis in what he called the bradytelic mode, he considered it (along with rapid evolution) to be unimportant in the larger scope of evolution. In his Major Features of Evolution Simpson stated, "Evolutionary change is so nearly the universal rule that a state of motion is, figuratively, normal in evolving populations. The state of rest, as in bradytely, is the exception and it seems that some restraint or force must be required to maintain it." Despite such differences between the two models, earlier critiques—from such eminent commentators as Sewall Wright as well as Simpson himself—have argued that punctuated equilibrium is little more than quantum evolution relabeled.

text: Quantum evolution

text: was a controversial hypothesis advanced by

text: Columbia University

text: paleontologist

text: George Gaylord Simpson

text: , who was regarded by Stephen Jay Gould as "the greatest and most biologically astute paleontologist of the twentieth century." Simpson's conjecture was that according to the geological record, on very rare occasions evolution would proceed very rapidly to form entirely new

text: families

text: orders

text: , and

text: classes

text: of organisms. This hypothesis differs from punctuated equilibrium in several respects. First, punctuated equilibrium was more modest in scope, in that it was addressing evolution specifically at the

text: species

text: level. Simpson's idea was principally concerned with evolution at higher taxonomic groups. Second, Eldredge and Gould relied upon a different mechanism. Where Simpson relied upon a

text: synergistic

text: interaction between

text: genetic drift

text: and a shift in the

text: adaptive fitness landscape

text: , Eldredge and Gould relied upon ordinary speciation, particularly Ernst Mayr's concept of allopatric speciation. Lastly, and perhaps most significantly, quantum evolution took no position on the issue of stasis. Although Simpson acknowledged the existence of stasis in what he called the bradytelic mode, he considered it (along with rapid evolution) to be unimportant in the larger scope of evolution. In his Major Features of Evolution Simpson stated, "Evolutionary change is so nearly the universal rule that a state of motion is, figuratively, normal in evolving populations. The state of rest, as in bradytely, is the exception and it seems that some restraint or force must be required to maintain it." Despite such differences between the two models, earlier critiques—from such eminent commentators as

text: Sewall Wright

text: as well as Simpson himself—have argued that punctuated equilibrium is little more than quantum evolution relabeled.

text: An important contributor should not be overlooked. Conwy Lloyd Morgan was perhaps closest to understanding the so-called "Baldwin Effect". In his "Habit and Instinct" (1896) he phrased a comparable version of the theory, as he did in an address to a session of the New York Academy of Sciences (February 1896) in the presence of Baldwin. (1896/Of modification and variation. Science 4(99) (November 20):733-739). As did Henry Fairfield Osborn (1896/A mode of evolution requiring neither natural selection nor the inheritance of acquired characteristics. Transactions of the New York Academy of Science 15:141-148). The "Baldwin Effect", building in part on the principle of "organic selection" proposed by Baldwin in "Mental Development" did only receive its name from George Gaylord Simpson in 1953. (in: Evolution 7:110-117) (see: David J. Depew in "Evolution and Learning" M.I.T.2003)

text: An important contributor should not be overlooked.

text: Conwy Lloyd Morgan

text: was perhaps closest to understanding the so-called "

text: Baldwin Effect

text: ". In his "Habit and Instinct" (1896) he phrased a comparable version of the theory, as he did in an address to a session of the

text: New York Academy of Sciences

text: (February 1896) in the presence of Baldwin. (1896/Of modification and variation. Science 4(99) (November 20):733-739). As did

text: Henry Fairfield Osborn

text: (1896/A mode of evolution requiring neither natural selection nor the inheritance of acquired characteristics. Transactions of the New York Academy of Science 15:141-148). The "Baldwin Effect", building in part on the principle of "organic selection" proposed by Baldwin in "Mental Development" did only receive its name from

text: George Gaylord Simpson

text: in 1953. (in: Evolution 7:110-117) (see: David J. Depew in "Evolution and Learning" M.I.T.2003)

text: for necessity. The differential inclusion   is a type of “evolutionary engine” (called an evolutionary system associating with any initial state x a subset of evolutions starting at x. The system is said to be deterministic if this set is made of one and only one evolution and contingent otherwise. Necessity is the requirement that at each instant, the evolution is viable (remains) in the environment K described by viability constraints, a word encompassing polysemous concepts as stability, confinement, homeostasis, adaptation, etc., expressing the idea that some variables must obey some constraints (representing physical, social, biological and economic constraints, etc.) that can never be violated. So, viability theory starts as the confrontation of evolutionary systems governing evolutions and viability constraints that such evolutions must obey. They share common features:

text: for necessity. The differential inclusion   is a type of “evolutionary engine” (called an evolutionary system associating with any initial state x a subset of evolutions starting at x. The system is said to be deterministic if this set is made of one and only one evolution and contingent otherwise. Necessity is the requirement that at each instant, the evolution is viable (remains) in the environment K described by viability constraints, a word encompassing

text: polysemous

text: concepts as stability, confinement, homeostasis, adaptation, etc., expressing the idea that some variables must obey some constraints (representing physical, social, biological and economic constraints, etc.) that can never be violated. So, viability theory starts as the confrontation of evolutionary systems governing evolutions and viability constraints that such evolutions must obey. They share common features:

text: Plant evolution is an aspect of the study of biological evolution, involving predominantly evolution of plants suited to live on land, greening of various land masses by the filling of their niches with land plants, and diversification of groups of land plants.

text: Plant evolution is an aspect of the study of

text: biological evolution

text: , involving predominantly evolution of plants suited to live on land, greening of various land masses by the filling of their

text: niches

text: with land plants, and diversification of groups of land plants.

text: Further than the basic scientific theory of evolution (notably neo-darwinian, Evolutive Humanism, developed by Richard Dawkins and Stephen Jay Gould),  looks at the necessity for Man of a permanent adaptation of both his organism and his thoughts to his universal environment. And from there to the relativized relation to belief and to uncertainty, because human ideas evolve and are transformed, as are physical qualities, in a universal process of evolution and adaptive improvement in their environment. But this does not suppose determinism, genetic or cultural, it refers to the evolutive interactivity and reactivity, partly random, of the living being with his surroundings. According to Marc Carl, "Man must necessarily learn to manage the insufficient appearance and the uncertainty of information to fit and to develop in his relational interactions with his environment, as well as physically and culturally". In a universal environment still as much as 90% unknown, this evolutive cogitation therefore tries not to be locked in premature schemata and answers. It encourages humility in proposals and intuitive boldness in investigations.A subjacent humanist thought encourages human beings to deliberately take his destiny in his hands, in careful correlation with his environment, knowing that human thought is one of the most impacting manifestations of the living being, as an agent of transformation of the environment, and consequently not only of the Earth's environment. In terms of philosophy of evolution, the organized collective thought of humanity, and in general his evolutive culture, appears as the key to development of our species in its universal environment, and the key to a possibly significant interactive modification, in the long term, of this environment. It is principally a prospective step.

text: Further than the basic

text: scientific theory

text: of evolution (notably neo-

text: darwinian

text: , Evolutive Humanism, developed by

text: Richard Dawkins

text: and

text: Stephen Jay Gould

text: ),  looks at the necessity for Man of a permanent adaptation of both his organism and his thoughts to his universal

text: environment

text: . And from there to the relativized relation to belief and to uncertainty, because human ideas evolve and are transformed, as are physical qualities, in a universal process of

text: evolution

text: and adaptive improvement in their environment. But this does not suppose

text: determinism

text: , genetic or cultural, it refers to the evolutive interactivity and reactivity, partly random, of the living being with his surroundings. According to

text: Marc Carl

text: , "Man must necessarily learn to manage the insufficient appearance and the uncertainty of information to fit and to develop in his relational interactions with his environment, as well as physically and culturally". In a universal environment still as much as 90% unknown, this evolutive cogitation therefore tries not to be locked in premature schemata and answers. It encourages humility in proposals and intuitive boldness in investigations.A subjacent humanist thought encourages human beings to deliberately take his destiny in his hands, in careful correlation with his environment, knowing that human thought is one of the most impacting manifestations of the living being, as an agent of transformation of the environment, and consequently not only of the Earth's environment. In terms of philosophy of evolution, the organized collective thought of humanity, and in general his evolutive culture, appears as the key to development of our

text: species

text: in its universal environment, and the key to a possibly significant interactive modification, in the long term, of this environment. It is principally a prospective step.

text: Due to the clinical similarity between MERS-CoV and SARS-CoV, it was proposed that they may use the same cellular receptor; the exopeptidase, angiotensin converting enzyme 2 (ACE2). However, it was later discovered that neutralization of ACE2 by recombinant antibodies does not prevent MERS-CoV infection. Further research identified dipeptyl peptidase 4 (DPP4; also known as CD26) as a functional cellular receptor for MERS-CoV. Unlike other known coronavirus receptors, the enzymatic activity of DPP4 is not required for infection. As would be expected, the amino acid sequence of DPP4 is highly conserved across species and is expressed in the human bronchial epithelium and kidneys. Bat DPP4 genes appear to have been subject to a high degree of adaptive evolution as a response to coronavirus infections, so the lineage leading to MERS-CoV may have circulated in bat populations for a long period of time before being transmitted to people.

text: Due to the clinical similarity between MERS-CoV and

text: SARS-CoV

text: , it was proposed that they may use the same cellular receptor; the exopeptidase, angiotensin converting enzyme 2 (

text: ). However, it was later discovered that neutralization of ACE2 by recombinant antibodies does not prevent MERS-CoV infection. Further research identified dipeptyl peptidase 4 (

text: ; also known as

text: ) as a functional cellular receptor for MERS-CoV. Unlike other known coronavirus receptors, the

text: enzymatic activity

text: of DPP4 is not required for infection. As would be expected, the amino acid sequence of DPP4 is highly conserved across species and is expressed in the human bronchial epithelium and kidneys. Bat DPP4 genes appear to have been subject to a high degree of adaptive evolution as a response to coronavirus infections, so the lineage leading to MERS-CoV may have circulated in bat populations for a long period of time before being transmitted to people.

text: Evolution can act to simplify as well as to complicate. This raises the possibility that seemingly irreducibly complex biological features may have been achieved with a period of increasing complexity, followed by a period of simplification.

text: The patterns of genetic adaptation among the Andeans are largely distinct from those of the Tibetan, with both populations showing evidence of positive natural selection in different genes or gene regions. However, EGLN1 appears to be the principal signature of evolution, as it shows evidence of positive selection in both Tibetans and Andeans. Even then, the pattern of variation for this gene differs between the two populations. Among the Andeans, there are no significant associations between EPAS1 or EGLN1 SNP genotypes and haemoglobin concentration, which has been the characteristic of the Tibetans. The whole genome sequences of 20 Andeans (half of them having chronic mountain sickness) revealed that two genes, SENP1 (an erythropoiesis regulator) and ANP32D (an oncogene) play vital roles in their weak adaptation to hypoxia.

text: The patterns of genetic adaptation among the Andeans are largely distinct from those of the Tibetan, with both populations showing evidence of positive natural selection in different genes or gene regions. However, EGLN1 appears to be the principal signature of evolution, as it shows evidence of positive selection in both Tibetans and Andeans. Even then, the pattern of variation for this gene differs between the two populations. Among the Andeans, there are no significant associations between EPAS1 or EGLN1 SNP

text: genotype

text: s and haemoglobin concentration, which has been the characteristic of the Tibetans. The whole genome sequences of 20 Andeans (half of them having chronic mountain sickness) revealed that two genes, SENP1 (an erythropoiesis regulator) and ANP32D (an oncogene) play vital roles in their weak adaptation to hypoxia.

text: Lactase expression persistence is largely due to natural selection, which is a component of evolution by which a trait affects the chances of the survival of organisms, and consequently, the trait becomes more prevalent in the population over time.

text: Lactase expression persistence is largely due to

text: natural selection

text: , which is a component of

text: evolution

text: by which a trait affects the chances of the survival of organisms, and consequently, the trait becomes more prevalent in the population over time.

text: In the preface, Shaw speaks of the pervasive discouragement and poverty in Europe after World War I, and relates these issues to inept government. Simple primitive societies, he says, were easily governable while the civilized societies of the twentieth century are so complex that learning to govern them properly can't be accomplished within the human lifespan: People with experience enough to serve the purpose fall into senility and die. Shaw's solution is enhanced longevity: we must learn to live much longer; a centenarian should be less than middle aged. (Shaw was in his mid-60s when the plays were written).  This change, Shaw predicts, will happen through Creative Evolution (evolutionary change that occurs because it is needed or wanted—the Lamarckian view— and not as a result of natural selection—Darwinism) as influenced by the Life Force (l'élan vital).  Neither Creative Evolution nor the Life Force were Shavian inventions. Shaw says they are his names for what the churches have called Providence and scientists call Functional Adaptation and Natural Selection (among other names) and gives due credit to Henri Bergson's élan vital.   Nevertheless, he uses both terms in Man and Superman which was written nine years before Bergson's work was published. These concepts had some currency among Shaw's contemporaries, and the Methuselah plays are based on Shaw's extrapolations from the two principles. Although both ideas are out of scientific favour as the twenty-first century begins, Shaw accepted them completely (See Commentary, below.) Shaw also advocates what he calls homeopathy as a pedagogical method, arguing that society "can only be lamed and enslaved by" education. Shaw's "homeopathic" educational method consisted of lying to students, until the students were able to see through the lies and argue with the teachers.

text: In the preface, Shaw speaks of the pervasive discouragement and poverty in Europe after World War I, and relates these issues to inept government. Simple primitive societies, he says, were easily governable while the civilized societies of the twentieth century are so complex that learning to govern them properly can't be accomplished within the human lifespan: People with experience enough to serve the purpose fall into senility and die. Shaw's solution is enhanced longevity: we must learn to live much longer; a centenarian should be less than middle aged. (Shaw was in his mid-60s when the plays were written).  This change, Shaw predicts, will happen through

text: Creative Evolution

text: (evolutionary change that occurs because it is needed or wanted—the

text: Lamarckian view

text: — and not as a result of natural selection—

text: Darwinism

text: ) as influenced by the

text: Life Force

text: (l'élan vital).  Neither Creative Evolution nor the Life Force were Shavian inventions. Shaw says they are his names for what the churches have called

text: Providence

text: and scientists call Functional Adaptation and

text: Natural Selection

text: (among other names) and gives due credit to

text: Henri Bergson

text: 's élan vital.   Nevertheless, he uses both terms in

text: Man and Superman

text: which was written nine years before Bergson's work was published. These concepts had some currency among Shaw's contemporaries, and the Methuselah plays are based on Shaw's extrapolations from the two principles. Although both ideas are out of scientific favour as the twenty-first century begins, Shaw accepted them completely (See Commentary, below.) Shaw also advocates what he calls

text: homeopathy

text: as a pedagogical method, arguing that society "can only be lamed and enslaved by" education. Shaw's "homeopathic" educational method consisted of lying to students, until the students were able to see through the lies and argue with the teachers.

text: Thus, "one sharp contrast between Dawkins and Gould is on the application of science in general, and evolutionary biology in particular, to our species". (p. 162) Yet paradoxically, Dawkins' most systematic writings on human evolution explore the differences between human evolution and that of most other organisms, in which humans pass on their values through ideas and skills which Dawkins calls memes. To Dawkins, ideas are often like pathogens or parasites, replicating throughout human populations, sometimes quite virulently, with evangelical religion being a salient example. Doubts about the reliability and accuracy of idea replication suggest Dawkins' own view of cultural evolution may not work. But his general approach has gained some popularity, as illustrated by works which explore the interaction between cultural and biological evolution, such as Peter Richerson and Robert Boyd's Not By Genes Alone,. as well as Eytan Avital and Eva Jablonka's Animal Traditions. "So though Dawkins approaches human behaviour using different tools to those of standard sociobiologists and evolutionary psychologists, he is fully committed to the idea that we can understand ourselves only in an evolutionary framework." (pp. 164–165) This contrasts with Gould. While "Of course" he accepts that humans are an evolved species, "Everything that Gould does not like in contemporary evolutionary thinking comes together in human sociobiology and its descendant, evolutionary psychology. The result has been a twenty-year campaign of savage polemic against evolutionary theories of human behaviour. Gould hates sociobiology". And "It is true that some evolutionary psychology does seem simple-minded," such as Randy Thornhill's "unconvincing" attempt to argue that a tendency towards rape is an evolutionary adaptation. (p. 165) However, contemporary evolutionary psychologists, and especially biological anthropologists, have accepted the need for caution in testing adaptationist hypotheses. (p. 165) However, even the most disciplined sociobiological approaches reflect different approaches to evolution to that exemplified by Gould. They "tend not to emphasise the importance of development and history in imposing constraints on adaptation, the problems in translating microevolutionary change into species-level change, the role of contingency and mass extinction in reshaping evolving lineages, or the importance of paleobiology to evolutionary biology", (p. 166) which likely played a part in Gould's hostility. But Sterelny suspects more most of all, Gould thought "these ideas are dangerous and ill-motivated as well as wrong. They smack of hubris, of science moving beyond its proper domain, and incautiously at that". Conversely, to Dawkins, knowledge of evolutionary underpinnings to human behaviour is potentially liberating, and "might even help us to escape the poisoned chalice of religion". (p. 166)

text: Thus, "one sharp contrast between Dawkins and Gould is on the application of science in general, and evolutionary biology in particular, to our species". (p. 162) Yet paradoxically, Dawkins' most systematic writings on human evolution explore the differences between human evolution and that of most other organisms, in which humans pass on their values through ideas and skills which Dawkins calls

text: memes

text: . To Dawkins, ideas are often like pathogens or parasites, replicating throughout human populations, sometimes quite virulently, with evangelical religion being a salient example. Doubts about the reliability and accuracy of idea replication suggest Dawkins' own view of cultural evolution may not work. But his general approach has gained some popularity, as illustrated by works which explore the interaction between cultural and biological evolution, such as

text: Peter Richerson

text: and

text: Robert Boyd

text: 's Not By Genes Alone,. as well as Eytan Avital and

text: Eva Jablonka

text: In humans, the lips are important for the production of stops and fricatives, in addition to vowels. Nothing, however, suggests that the lips evolved for those reasons. During primate evolution, a shift from nocturnal to diurnal activity in tarsiers, monkeys and apes (the haplorhines) brought with it an increased reliance on vision at the expense of olfaction. As a result, the snout became reduced and the rhinarium or "wet nose" was lost. The muscles of the face and lips consequently became less constrained, enabling their co-option to serve purposes of facial expression. The lips also became thicker. "Hence", according to one major authority, "the evolution of mobile, muscular lips, so important to human speech, was the exaptive result of the evolution of diurnality and visual communication in the common ancestor of haplorhines". It is unclear whether our lips have undergone more recent adaptation to the specific requirements of speech.

text: In humans, the lips are important for the production of

text: stops

text: and

text: fricatives

text: , in addition to

text: vowel

text: s. Nothing, however, suggests that the lips evolved for those reasons. During primate evolution, a shift from nocturnal to diurnal activity in

text: tarsier

text: s, monkeys and apes (the

text: haplorhines

text: ) brought with it an increased reliance on vision at the expense of olfaction. As a result, the snout became reduced and the

text: rhinarium

text: or "wet nose" was lost. The muscles of the face and lips consequently became less constrained, enabling their co-option to serve purposes of facial expression. The lips also became thicker. "Hence", according to one major authority, "the evolution of mobile, muscular lips, so important to human speech, was the exaptive result of the evolution of diurnality and visual communication in the common ancestor of haplorhines". It is unclear whether our lips have undergone more recent adaptation to the specific requirements of speech.

text: Phenotypic variation (due to underlying heritable genetic variation) is a fundamental prerequisite for evolution by natural selection. It is the living organism as a whole that contributes (or not) to the next generation, so natural selection affects the genetic structure of a population indirectly via the contribution of phenotypes. Without phenotypic variation, there would be no evolution by natural selection.

text: Phenotypic variation (due to underlying heritable

text: genetic variation

text: ) is a fundamental prerequisite for

text: evolution

text: by

text: natural selection

text: . It is the living organism as a whole that contributes (or not) to the next generation, so natural selection affects the genetic structure of a population indirectly via the contribution of phenotypes. Without phenotypic variation, there would be no evolution by natural selection.

text: Other gene types showing considerable evidence of adaptive evolution (but generally less evidence than the types discussed) include: genes on the X chromosome, nervous system genes, genes involved in apoptosis, genes coding for skeletal traits, and possibly genes associated with speech (Nielsen et al. 2005a, Williamson et al. 2007, Voight et al. 2006, Krause et al. 2007).

text: At this point, Dawkins switches from explaining artificial selection to explaining natural selection. To demonstrate natural selection in a computer program, Dawkins uses a program written by Peter Fuchs to simulate the evolution of the spiderweb. The program builds "genetic" variations of a parent web, as if the web was actually being built by a child spider. For each generation, a simulation is run which randomly generates flies - some of which will hit the web, and others that will miss it. The child web that is able to capture the highest number of flies is selected as the parent for the next generation of webs. Dawkins shows the audience the "fossil record" that the program recorded after simulating a large number of generations overnight. The web starts off very simple and inefficient, but by the end it has evolved into a web that is highly efficient and highly complex. This is the same process that has led to the existence of all designoid objects.

text: At this point, Dawkins switches from explaining artificial selection to explaining natural selection. To demonstrate natural selection in a computer program, Dawkins uses a program written by Peter Fuchs to simulate the evolution of the

text: spiderweb

text: . The program builds "genetic" variations of a parent web, as if the web was actually being built by a child spider. For each generation, a simulation is run which randomly generates flies - some of which will hit the web, and others that will miss it. The child web that is able to capture the highest number of flies is selected as the parent for the next generation of webs. Dawkins shows the audience the "fossil record" that the program recorded after simulating a large number of generations overnight. The web starts off very simple and inefficient, but by the end it has evolved into a web that is highly efficient and highly complex. This is the same process that has led to the existence of all designoid objects.

text: This cultural trend seems to have emerged because the concept of progressive permanent adaptation, which is no longer appreciated only by its scientific aspects, also took a metaphysical dimension (in a sense of research of the essence of the human), which consequently encourages analysis of human evolution, an agent which potentially modifies our environment, in philosophical terms. The possibilities and the risks of this evolution, non-deterministic because of its integration into a dynamic complex living system, gives the human existence and destiny a new sense. And to be involved in such a cogitation opens a philosophical way in which curious minds could not miss advancing sooner or later, a way which rests on a regenerated metaphysics, encouraging us to take back in an evolutive way Aristoteles original concept of physis and its substance, and to search with modern conceptual tools the essence and the sense of our life.

text: This cultural trend seems to have emerged because the concept of progressive permanent adaptation, which is no longer appreciated only by its scientific aspects, also took a metaphysical dimension (in a sense of research of the essence of the human), which consequently encourages analysis of

text: human evolution

text: , an agent which potentially modifies our environment, in philosophical terms. The possibilities and the risks of this evolution,

text: non-deterministic

text: because of its integration into a dynamic complex living system, gives the human existence and destiny a new sense. And to be involved in such a cogitation opens a philosophical way in which curious minds could not miss advancing sooner or later, a way which rests on a regenerated

text: metaphysics

text: , encouraging us to take back in an evolutive way Aristoteles original concept of physis and its substance, and to search with modern conceptual tools the essence and the sense of our life.

text: A gene is a sequence of DNA that contains genetic information and can influence the phenotype of an organism. Within a gene, the sequence of bases along a DNA strand defines a messenger RNA sequence, which then defines one or more protein sequences. The relationship between the nucleotide sequences of genes and the amino-acid sequences of proteins is determined by the rules of translation, known collectively as the genetic code. The genetic code consists of three-letter 'words' called codons formed from a sequence of three nucleotides (e.g. ACT, CAG, TTT).

text: A gene is a sequence of DNA that contains genetic information and can influence the

text: phenotype

text: of an organism. Within a gene, the sequence of bases along a DNA strand defines a

text: messenger RNA

text: sequence, which then defines one or more protein sequences. The relationship between the nucleotide sequences of genes and the

text: amino-acid

text: sequences of proteins is determined by the rules of

text: translation

text: , known collectively as the

text: genetic code

text: . The genetic code consists of three-letter 'words' called codons formed from a sequence of three nucleotides (e.g. ACT, CAG, TTT).

text: Gene-culture coevolution allows humans to develop highly distinct adaptations to the local pressures and environments more quickly than with genetic evolution alone. Robert Boyd and Peter J. Richerson, two strong proponents of cultural evolution, postulate that the act of social learning, or learning in a group as done in group selection, allows human populations to accrue information over many generations. This leads to cultural evolution of behaviors and technology alongside genetic evolution. Boyd and Richerson believe that the ability to collaborate evolved during the Middle Pleistocene, a million years ago, in response to a rapidly changing climate.

text: Gene-culture coevolution allows humans to develop highly distinct adaptations to the local pressures and environments more quickly than with genetic evolution alone.

text: Robert Boyd

text: and

text: Peter J. Richerson

text: , two strong proponents of cultural evolution, postulate that the act of social learning, or learning in a group as done in group selection, allows human populations to accrue information over many generations. This leads to cultural evolution of behaviors and technology alongside genetic evolution. Boyd and Richerson believe that the ability to collaborate evolved during the Middle

text: Pleistocene

text: , a million years ago, in response to a rapidly changing climate.

text: Microbial population biology can include aspects of molecular evolution or phylogenetics. Strictly, however, these emphases should be employed toward understanding issues of microbial evolution and ecology rather than as a means of understanding more universal truths applicable to both microscopic and macroscopic organisms. The microorganisms in such endeavors consequently should be recognized as organisms rather than simply as molecular or evolutionary reductionist model systems. Thus, the study of RNA  in vitro evolution is not microbial population biology and nor is the in silico generation of phylogenies of otherwise non-microbial sequences, even if aspects of either may in some (especially unintentional) manner be analogous to evolution in actual microbial populations.

text: Microbial population biology can include aspects of

text: molecular evolution

text: or

text: phylogenetics

text: . Strictly, however, these emphases should be employed toward understanding issues of microbial evolution and ecology rather than as a means of understanding more universal truths applicable to both microscopic and macroscopic organisms. The microorganisms in such endeavors consequently should be recognized as organisms rather than simply as molecular or evolutionary

text: reductionist

text: model systems. Thus, the study of RNA

text: in vitro

text: evolution is not microbial population biology and nor is the

text: in silico

text: generation of phylogenies of otherwise non-microbial sequences, even if aspects of either may in some (especially unintentional) manner be analogous to evolution in actual microbial populations.

text: The American biologist Sewall Wright, who had a background in animal breeding experiments, focused on combinations of interacting genes, and the effects of inbreeding on small, relatively isolated populations that exhibited genetic drift. In 1932, Wright introduced the concept of an adaptive landscape and argued that genetic drift and inbreeding could drive a small, isolated sub-population away from an adaptive peak, allowing natural selection to drive it towards different adaptive peaks. The work of Fisher, Haldane and Wright founded the discipline of population genetics. This integrated natural selection with Mendelian genetics, which was the critical first step in developing a unified theory of how evolution worked.

text: The American biologist Sewall Wright, who had a background in

text: animal breeding

text: experiments, focused on combinations of interacting genes, and the effects of inbreeding on small, relatively isolated populations that exhibited genetic drift. In 1932, Wright introduced the concept of an

text: adaptive landscape

text: and argued that genetic drift and inbreeding could drive a small, isolated sub-population away from an adaptive peak, allowing natural selection to drive it towards different adaptive peaks. The work of Fisher, Haldane and Wright founded the discipline of population genetics. This integrated natural selection with Mendelian genetics, which was the critical first step in developing a unified theory of how evolution worked.

text: Recurrent evolution is a broad term, but is usually used to describe recurring regimes of selection within or across lineages. While most commonly used to describe recurring patterns of selection, it can also be used to describe recurring patterns of mutation, for example transitions are more common than transversions. It encompasses both convergent evolution and parallel evolution and can be used to describe the observation of similar repeating changes through directional selection as well as the observation of highly conserved phenotypes or genotypes across lineages through continuous purifying selection over large periods of evolutionary time.  The changes can be observed at the phenotype level or the genotype level. At the phenotype level recurrent evolution can be observed across a continuum of levels, which for simplicity can be broken down into molecular phenotype, cellular phenotype, and organismal phenotype. At the genotype level recurrent evolution can only be detected using DNA sequencing data. The same or similar changes in the genomes of different lineages indicates recurrent genomic evolution may have taken place. Recurrent genomic evolution can also occur within a lineage. An example of this would include some types of phase variation that involve highly directed changes at the DNA sequence level. The evolution of different forms of phase variation in separate lineages represent convergent and recurrent evolution toward increased evolvability. In organisms with longer generation times, any potential recurrent genomic evolution within a lineage would be difficult to detect. Recurrent evolution has been studied most extensively at the organismic level but with cheaper and faster sequencing technologies more attention is being paid to recurrent genomic evolution. Recurrent evolution can also be described as recurring or repeated evolution.

text: Recurrent evolution is a broad term, but is usually used to describe recurring regimes of

text: selection

text: within or across

text: lineages

text: . While most commonly used to describe recurring patterns of selection, it can also be used to describe recurring patterns of

text: mutation

text: , for example

text: transitions

text: are more common than

text: transversions

text: . It encompasses both

text: convergent evolution

text: and

text: parallel evolution

text: and can be used to describe the observation of similar repeating changes through

text: directional selection

text: as well as the observation of highly conserved phenotypes or genotypes across lineages through continuous

text: purifying selection

text: over large periods of evolutionary time.  The changes can be observed at the

text: phenotype

text: level or the

text: genotype

text: level. At the phenotype level recurrent evolution can be observed across a continuum of levels, which for simplicity can be broken down into molecular phenotype, cellular phenotype, and organismal phenotype. At the genotype level recurrent evolution can only be detected using

text: DNA sequencing

text: data. The same or similar changes in the genomes of different lineages indicates recurrent genomic evolution may have taken place. Recurrent genomic evolution can also occur within a lineage. An example of this would include some types of

text: phase variation

text: that involve highly directed changes at the DNA sequence level. The evolution of different forms of phase variation in separate lineages represent convergent and recurrent evolution toward increased

text: evolvability

text: . In organisms with longer generation times, any potential recurrent genomic evolution within a lineage would be difficult to detect. Recurrent evolution has been studied most extensively at the organismic level but with cheaper and faster sequencing technologies more attention is being paid to recurrent genomic evolution. Recurrent evolution can also be described as recurring or repeated evolution.

text: The principles of natural selection have inspired a variety of computational techniques, such as "soft" artificial life, that simulate selective processes and can be highly efficient in 'adapting' entities to an environment defined by a specified fitness function. For example, a class of heuristic optimisation algorithms known as genetic algorithms, pioneered by John Henry Holland in the 1970s and expanded upon by David E. Goldberg, identify optimal solutions by simulated reproduction and mutation of a population of solutions defined by an initial probability distribution. Such algorithms are particularly useful when applied to problems whose energy landscape is very rough or has many local minima.

text: The principles of natural selection have inspired a variety of computational techniques, such as "soft"

text: artificial life

text: , that simulate selective processes and can be highly efficient in 'adapting' entities to an environment defined by a specified

text: fitness function

text: . For example, a class of

text: heuristic

text: optimisation

text: algorithm

text: s known as

text: genetic algorithm

text: s, pioneered by

text: John Henry Holland

text: in the 1970s and expanded upon by

text: David E. Goldberg

text: , identify optimal solutions by simulated reproduction and mutation of a population of solutions defined by an initial

text: probability distribution

text: . Such algorithms are particularly useful when applied to problems whose

text: energy landscape

text: is very rough or has many local minima.

text: According to Simpson (1944), quantum evolution relied heavily upon Sewall Wright's theory of random genetic drift. Simpson believed that major evolutionary transitions would arise when small populations—isolated and limited from gene flow—would fixate upon unusual gene combinations. This "inadaptive phase" (by genetic drift) would then (by natural selection) drive a deme population from one stable adaptive peak to another on the adaptive fitness landscape. However, in his Major Features of Evolution (1953) Simpson wrote that it was still controversial:

text: According to Simpson (1944), quantum evolution relied heavily upon

text: Sewall Wright

text: 's theory of

text: random genetic drift

text: . Simpson believed that major evolutionary transitions would arise when small populations—isolated and limited from gene flow—would fixate upon unusual gene combinations. This "inadaptive phase" (by genetic drift) would then (by natural selection) drive a

text: deme population

text: from one stable adaptive peak to another on the

text: adaptive fitness landscape

text: . However, in his Major Features of Evolution (1953) Simpson wrote that it was still controversial:

text: Bateson's work published before 1900 systematically studied the structural variation displayed by living organisms and the light this might shed on the mechanism of biological evolution, and was strongly influenced by both Charles Darwin's approach to the collection of comprehensive examples, and Francis Galton's quantitative ("biometric") methods. In his first significant contribution, he shows that some biological characteristics (such as the length of forceps in earwigs) are not distributed continuously, with a normal distribution, but discontinuously (or "dimorphically"). He saw the persistence of two forms in one population as a challenge to the then current conceptions of the mechanism of heredity, and says "The question may be asked, does the dimorphism of which cases have now been given represent the beginning of a division into two species?”

text: Bateson's work published before 1900 systematically studied the structural variation displayed by living organisms and the light this might shed on the mechanism of biological evolution, and was strongly influenced by both

text: Charles Darwin

text: 's approach to the collection of comprehensive examples, and

text: Francis Galton

text: 's quantitative ("biometric") methods. In his first significant contribution, he shows that some biological characteristics (such as the length of forceps in earwigs) are not distributed continuously, with a normal distribution, but discontinuously (or "

text: dimorphically

text: "). He saw the persistence of two forms in one population as a challenge to the then current conceptions of the mechanism of heredity, and says "The question may be asked, does the dimorphism of which cases have now been given represent the beginning of a division into two species?”

text: Evolution – change in heritable traits of biological organisms over generations due to natural selection, mutation, gene flow, and genetic drift.  Also known as descent with modification. Over time these evolutionary processes lead to formation of new species (speciation), changes within lineages (anagenesis), and loss of species (extinction). "Evolution" is also another name for evolutionary biology, the subfield of biology concerned with studying evolutionary processes that produced the diversity of life on Earth.

text: Evolution

text: – change in

text: heritable trait

text: s of

text: biological organisms

text: over generations due to

text: natural selection

text: mutation

text: gene flow

text: , and

text: genetic drift

text: .  Also known as descent with modification. Over time these evolutionary processes lead to formation of new species (

text: speciation

text: ), changes within lineages (

text: anagenesis

text: ), and loss of species (

text: extinction

text: ). "Evolution" is also another name for

text: evolutionary biology

text: , the subfield of

text: biology

text: concerned with studying

text: evolutionary processes

text: that produced the

text: diversity of life

text: on Earth.

text: With pleiotropy, some genes control multiple traits, so that adaptation of one trait is impeded by effects on other traits that are not necessarily adaptive. Selection that influences epistasis is a case where the regulation or expression of one gene, depends on one or several others. This is true for a good number of genes though to differing extents. The reason why this leads to muddied responses is that selection for a trait that is epistatically based can mean that an allele for a gene that is epistatic when selected would happen to affect others. This leads to the coregulation of others for a reason other than there is an adaptive quality to each of those traits. Like with pleiotropy, traits could reach fixation in a population as a by-product of selection for another.

text: With

text: pleiotropy

text: , some genes control multiple traits, so that adaptation of one trait is impeded by effects on other traits that are not necessarily adaptive. Selection that influences

text: epistasis

text: is a case where the regulation or expression of one gene, depends on one or several others. This is true for a good number of genes though to differing extents. The reason why this leads to muddied responses is that selection for a trait that is epistatically based can mean that an

text: allele

text: for a gene that is epistatic when selected would happen to affect others. This leads to the

text: coregulation

text: of others for a reason other than there is an adaptive quality to each of those traits. Like with pleiotropy, traits could reach fixation in a population as a by-product of selection for another.

text: Though not directly related to molecular evolution, the mid-1960s also saw the rise of the gene-centered view of evolution, spurred by George C. Williams's Adaptation and Natural Selection (1966). Debate over units of selection, particularly the controversy over group selection, led to increased focus on individual genes (rather than whole organisms or populations) as the theoretical basis for evolution. However, the increased focus on genes did not mean a focus on molecular evolution; in fact, the adaptationism promoted by Williams and other evolutionary theories further marginalized the apparently non-adaptive changes studied by molecular evolutionists.

text: Though not directly related to molecular evolution, the mid-1960s also saw the rise of the

text: gene-centered view of evolution

text: , spurred by

text: George C. Williams

text: 's

text: Adaptation and Natural Selection

text: (1966). Debate over

text: units of selection

text: , particularly the controversy over

text: group selection

text: , led to increased focus on individual genes (rather than whole organisms or populations) as the theoretical basis for evolution. However, the increased focus on genes did not mean a focus on molecular evolution; in fact, the

text: adaptationism

text: promoted by Williams and other evolutionary theories further marginalized the apparently non-adaptive changes studied by molecular evolutionists.

text: The aquatic adaptations of the baiji and other cetaceans have happened slowly and can be linked to PSGs and/or other functional changes. Comparative genopic analyses have uncovered that the baiji have a slow molecular clock and molecular adaptations to their aquatic environment. This information leads scientists to conclude that a bottleneck must have occurred near the end of the last deglaciation, a time that coincided with rapid temperature decrease and a rise in eustatic sea level. Scientists have also looked into positively selected genes (PSGs) in the baiji genome which are used for DNA repair and response to DNA stimulus. These PSGs have not been found previously in any other mammal species. Pathways being used for DNA repair have been known to have a major impact on brain development and have been implicated in diseases including microcephaly. The slow down of the substitution rate among cetaceans may have been affected by the evolution of DNA damage pathways. Over time, river dolphins, including the baiji, have had a reduction in the size of their eyes and the acuity of their vision. This probably stems from poor visibility in fluvial and estuarine environments. When analyzing the baiji genome, scientists have found that there are four genes that have lost their function due to a frameshift mutation or premature stop codons. The baiji has the lowest single nucleotide polymorphism (SNP) frequency reported thus far among mammals. This low frequency could be related to the relatively low rate of molecular evolution in cetaceans; however, considering that the decrease in the rate of molecular evolution in the baiji was not as great as the decrease in heterozygosity rate, it is likely that much if the low genetic diversity observed was caused by the precipitous decline in the total baiji population in recent decades and the associated breedings.

text: The aquatic adaptations of the baiji and other cetaceans have happened slowly and can be linked to PSGs and/or other functional changes. Comparative genopic analyses have uncovered that the baiji have a slow molecular clock and molecular adaptations to their aquatic environment. This information leads scientists to conclude that a bottleneck must have occurred near the end of the last deglaciation, a time that coincided with rapid temperature decrease and a rise in eustatic sea level. Scientists have also looked into positively selected genes (PSGs) in the baiji genome which are used for DNA repair and response to DNA stimulus. These PSGs have not been found previously in any other mammal species. Pathways being used for DNA repair have been known to have a major impact on brain development and have been implicated in diseases including

text: microcephaly

text: . The slow down of the substitution rate among cetaceans may have been affected by the evolution of DNA damage pathways. Over time, river dolphins, including the baiji, have had a reduction in the size of their eyes and the acuity of their vision. This probably stems from poor visibility in fluvial and estuarine environments. When analyzing the baiji genome, scientists have found that there are four genes that have lost their function due to a frameshift mutation or premature stop codons. The baiji has the lowest single nucleotide polymorphism (SNP) frequency reported thus far among mammals. This low frequency could be related to the relatively low rate of molecular evolution in cetaceans; however, considering that the decrease in the rate of molecular evolution in the baiji was not as great as the decrease in heterozygosity rate, it is likely that much if the low genetic diversity observed was caused by the precipitous decline in the total baiji population in recent decades and the associated breedings.

text: She has also previously studied the evolution of the color of mice coats, and its significance for adaptation. In 2013 her team published a paper in the journal Science, describing how coat color in mice was controlled by nine separate mutations within a single gene, named "agouti." Speaking about this discovery, Hoekstra said "The question has always been whether evolution is dominated by these big leaps or smaller steps. When we first implicated the agouti gene, we could have stopped there and concluded that evolution takes these big steps as only one major gene was involved, but that would have been wrong. When we looked more closely, within this gene, we found that even within this single locus, there are, in fact, many small steps." Her work supports the hypothesis that Evolution can occur through incremental changes.

text: She has also previously studied the evolution of the color of mice coats, and its significance for

text: adaptation

text: . In 2013 her team published a paper in the journal

text: Science

text: , describing how coat color in mice was controlled by nine separate mutations within a single gene, named "agouti." Speaking about this discovery, Hoekstra said "The question has always been whether evolution is dominated by these big leaps or smaller steps. When we first implicated the agouti gene, we could have stopped there and concluded that evolution takes these big steps as only one major gene was involved, but that would have been wrong. When we looked more closely, within this gene, we found that even within this single locus, there are, in fact, many small steps." Her work supports the hypothesis that

text: Evolution

text: can occur through incremental changes.

text: Experimental evolution is the use of experiments or controlled field manipulations to explore evolutionary dynamics. Evolution may be observed in the laboratory as populations adapt to new environmental conditions and/or change by such stochastic processes as random genetic drift. With modern molecular tools, it is possible to pinpoint the mutations that selection acts upon, what brought about the adaptations, and to find out how exactly these mutations work. Because of the large number of generations required for adaptation to occur, evolution experiments are typically carried out with microorganisms such as bacteria, yeast or viruses, or other organisms with rapid generation times. However, laboratory studies with foxes and with rodents (see below) have shown that notable adaptations can occur within as few as 10-20 generations and experiments with wild guppies have observed adaptations within comparable numbers of generations. More recently, experimentally evolved individuals or populations are often analyzed using whole genome sequencing, an approach known as Evolve and Resequence (E&R).

text: Experimental evolution is the use of experiments or controlled field manipulations to explore evolutionary dynamics. Evolution may be observed in the laboratory as populations adapt to new environmental conditions and/or change by such

text: stochastic

text: processes as random

text: genetic drift

text: . With modern

text: molecular

text: tools, it is possible to pinpoint the

text: mutation

text: s that

text: selection

text: acts upon, what brought about the

text: adaptation

text: s, and to find out how exactly these mutations work. Because of the large number of

text: generation

text: s required for adaptation to occur, evolution experiments are typically carried out with

text: microorganism

text: s such as

text: bacteria

text: yeast

text: or

text: virus

text: es, or other organisms with rapid generation times. However, laboratory studies with foxes and with

text: rodents

text: (see below) have shown that notable adaptations can occur within as few as 10-20 generations and experiments with wild

text: guppies

text: have observed adaptations within comparable numbers of generations. More recently, experimentally evolved individuals or populations are often analyzed using

text: whole genome sequencing

text: , an approach known as Evolve and Resequence (E&R).

text: Group Selection is a 1971 book edited by George C. Williams, containing papers written by biologists arguing against the view of group selection as a major force in evolution.  The group of biologists writing on a single unified (if somewhat broad) theme contrasts with Williams' earlier seminal 1966 book Adaptation and Natural Selection, which Williams suspected such views to be his alone.  In particular it contains a reprint, with an erratum, of W.D. Hamilton's classic 1964 paper on inclusive fitness, "The Genetical Evolution of Social Behavior" plus a paper by John Maynard Smith entitled "The Origin and Maintenance of Sex" (pp 163–175), containing ideas on evolution of sex later developed by Maynard Smith; see especially his 1978 book The Evolution of Sex.

text: Group Selection is a

text: book edited by

text: George C. Williams

text: , containing papers written by biologists arguing against the view of

text: group selection

text: as a major force in

text: evolution

text: .  The group of biologists writing on a single unified (if somewhat broad) theme contrasts with Williams' earlier seminal 1966 book

text: Adaptation and Natural Selection

text: , which Williams suspected such views to be his alone.  In particular it contains a reprint, with an erratum, of

text: W.D. Hamilton

text: 's classic 1964 paper on

text: inclusive fitness

entity_name: The Genetical Evolution of Social Behaviour

text: The Genetical Evolution of Social Behavior

text: " plus a paper by

text: John Maynard Smith

text: entitled "The Origin and Maintenance of Sex" (pp 163–175), containing ideas on

text: evolution of sex

text: later developed by Maynard Smith; see especially his 1978 book

text: The Evolution of Sex

text: As shown above, the genes of gene expression programming have all the same size. However, these fixed length strings code for expression trees of different sizes. This means that the size of the coding regions varies from gene to gene, allowing for adaptation and evolution to occur smoothly.

text: As shown

text: above

text: , the genes of gene expression programming have all the same size. However, these fixed length strings code for

text: expression tree

text: s of different sizes. This means that the size of the coding regions varies from gene to gene, allowing for adaptation and evolution to occur smoothly.

text: A gene family is a set of several similar genes, formed by duplication of a single original gene, and generally with similar biochemical functions. One such family are the genes for human hemoglobin subunits; the ten genes are in two clusters on different chromosomes, called the α-globin and β-globin loci.  These two gene clusters are thought to have arisen as a result of a precursor gene being duplicated approximately 500 million years ago.

text: A gene family is a set of several similar genes, formed by duplication of a single original gene, and generally with similar biochemical functions. One such family are the genes for human

text: hemoglobin

text: subunits; the ten genes are in two clusters on different chromosomes, called the

text: α-globin

text: and

text: β-globin

text: loci.  These two gene clusters are thought to have arisen as a result of a precursor gene being duplicated approximately 500 million years ago.

text: The study of orphan genes emerged largely after the turn of the century. In 2003, a study of Caenorhabditis briggsae and related species compared over 2000 genes. They proposed that these genes must be evolving too quickly to be detected and are consequently sites of very rapid evolution.  In 2005, Wilson examined 122 bacterial species to try to examine whether the large number of orphan genes in many species was legitimate. The study found that it was legitimate and played a role in bacterial adaptation. The definition of taxonomically-restricted genes was introduced into the literature to make orphan genes seem less "mysterious."

text: Galápagos Islands – gene – gene-centric view of evolution – gene duplication – gene flow – gene pool – genetic drift – genetic hitchhiking – genetic recombination – genetic variation – genotype – genotype-environment correlation – genotype-environment interaction – genotype-phenotype distinction – Stephen Jay Gould – gradualism – Peter and Rosemary Grant – group selection

text: Galápagos Islands

text: gene

text: gene-centric view of evolution

text: gene duplication

text: gene flow

text: gene pool

text: genetic drift

text: genetic hitchhiking

text: genetic recombination

text: genetic variation

text: genotype

text: genotype-environment correlation

text: genotype-environment interaction

text: genotype-phenotype distinction

text: Stephen Jay Gould

text: gradualism

text: Peter and Rosemary Grant

text: group selection

text: Wheat powdery mildew is an obligate biotroph with a poorly understood evolutionary history. Sequencing its genome in 2013, many aspects of the evolution of its parasitism were unveiled. Obligate biotrophy has appeared multiple times in evolution in both Ascomycetes like B. graminis and Basidiomycetes, thus different selective pressure must have acted in the different organisms through time. It has been seen that B. graminis f.sp. triticis genome is a mosaic of haplogroups with different divergence times, which explains its unique pathogen adaptability. Haplogroup H (diverged 40-80 mya) allows for the infection of wild tetraploid wheat and H (diverged 2-10 mya) allows for the infection of both domesticated hexaploid wheat.It is hypothesized that this mosaicisms has been maintained through clonal propagation in populations with small effective size or through quasi-clonal propagation in populations with large effective size.Additionally, it has been seen that there is a positive selective pressure acting on genes that code for candidate secretor proteins and non-secreted candidate secretor proteins, indicating that these might participate in the gene-for-gene relationship of plant disease resistance.

text: Wheat powdery mildew is an obligate biotroph with a poorly understood evolutionary history. Sequencing its genome in 2013, many aspects of the evolution of its parasitism were unveiled. Obligate biotrophy has appeared multiple times in evolution in both

text: Ascomycetes

text: like B. graminis and

text: Basidiomycetes

text: , thus different selective pressure must have acted in the different organisms through time. It has been seen that B. graminis f.sp. triticis genome is a mosaic of haplogroups with different divergence times, which explains its unique pathogen adaptability. Haplogroup H (diverged 40-80 mya) allows for the infection of wild tetraploid wheat and H (diverged 2-10 mya) allows for the infection of both domesticated hexaploid wheat.It is hypothesized that this mosaicisms has been maintained through clonal propagation in populations with small effective size or through quasi-clonal propagation in populations with large effective size.Additionally, it has been seen that there is a positive selective pressure acting on genes that code for candidate secretor proteins and non-secreted candidate secretor proteins, indicating that these might participate in the

text: gene-for-gene relationship

text: of plant disease resistance.

text: Animals that exhibit ecotonal variations allow for research concerning the mechanisms that maintain population differentiation. A wealth of information about natural selection, genotypic, and phenotypic variation; adaptation and ecomorphology; and social signaling has been acquired from the studies of three species of lizards located in the White Sands desert of New Mexico. Holbrookia maculata, Aspidoscelis inornata, and Sceloporus undulatus exhibit ecotonal populations that match both the dark soils and the white sands in the region. Research conducted on these species has found significant phenotypic and genotypic differences between the dark and light populations due to strong selection pressures. For example, H. maculata exhibits the strongest phenotypic difference (matches best with the substrate) of the light colored population coinciding with the least amount of gene flow between the populations and the highest genetic differences when compared to the other two lizard species.

text: Animals that exhibit

text: ecotonal

text: variations allow for research concerning the mechanisms that maintain population differentiation. A wealth of information about natural selection, genotypic, and phenotypic variation; adaptation and

text: ecomorphology

text: ; and social signaling has been acquired from the studies of three species of lizards located in the White Sands desert of New Mexico.

text: Holbrookia maculata

text: Aspidoscelis inornata

text: , and

text: Sceloporus undulatus

text: exhibit ecotonal populations that match both the dark soils and the white sands in the region. Research conducted on these species has found significant phenotypic and genotypic differences between the dark and light populations due to strong selection pressures. For example, H. maculata exhibits the strongest phenotypic difference (matches best with the substrate) of the light colored population coinciding with the least amount of gene flow between the populations and the highest genetic differences when compared to the other two lizard species.

text: In the discussions of genetic diseases in humans, pseudogene mediated gene conversions that introduce pathogenic mutations into functional genes is a well known mechanism of mutation. In contrast, it is possible that pseudogenes could serve as templates. During the course of evolution, functional source genes which are potentially advantageous have been derived from multiple copies in their single source gene. The pseudogene-templated changes might eventually become fixed as long as they did not possess deleterious effects. So, in fact, pseudogenes can act as sources of sequence variants which can be transferred to functional genes in novel combinations and can be acted upon by selection. Lectin 11 (SIGLEC11), a human immunoglobulin that binds to sialic acid, can be considered an example of such a gene conversion event which has played a significant role in evolution. While comparing the homologous genes of human SIGLEC11 and its pseudogene in the chimpanzee, bonobo, gorilla and orangutan, it appears that there was gene conversion of the sequence of 5’ upstream regions and the exons that encode the sialic acid recognition domain, approximately 2kbp from the closely flanking hSIGLECP16 pseudogene (Hayakawa et al., 2005). The three pieces of evidence concerning this event have together suggested this as an adaptive change which is very evolutionarily important in genus Homo. Those includes that only in human lineage this gene conversion happened, the brain cortex has acquired an important expression of SIGLEC11 specifically in human lineage and the exhibition of  a change in substrate binding in human lineage when compared to that of its counterpart in chimpanzees. Of course the frequency of the contribution of this pseudogene-mediated gene conversion mechanism to functional and adaptive changes in evolution of human is still unknown and so far it has been scarcely explored (Chen et al., 2007). In spite of that, the introduction of positively selective genetic changes by such mechanism can be put forward for consideration by the example of SIGLEC11. Sometimes due to interference of transposable elements in to some members of a gene family, it causes a variation among them and finally it may also cease the rate of gene conversion due to lack of sequence similarity which leads to divergent evolution.

text: In the discussions of

text: genetic diseases

text: in humans,

text: pseudogene

text: mediated gene conversions that introduce pathogenic

text: mutations

text: into functional genes is a well known mechanism of mutation. In contrast, it is possible that pseudogenes could serve as templates. During the course of evolution, functional source genes which are potentially advantageous have been derived from multiple copies in their single source gene. The pseudogene-templated changes might eventually become fixed as long as they did not possess deleterious effects. So, in fact, pseudogenes can act as sources of sequence variants which can be transferred to functional genes in novel combinations and can be acted upon by

text: selection

text: Lectin

text: 11 (

text: SIGLEC

text: 11), a human immunoglobulin that binds to sialic acid, can be considered an example of such a gene conversion event which has played a significant role in evolution. While comparing the homologous genes of human SIGLEC11 and its pseudogene in the chimpanzee, bonobo, gorilla and orangutan, it appears that there was gene conversion of the sequence of 5’ upstream regions and the exons that encode the sialic acid recognition domain, approximately 2kbp from the closely flanking hSIGLECP16 pseudogene (Hayakawa et al., 2005). The three pieces of evidence concerning this event have together suggested this as an adaptive change which is very evolutionarily important in

text: genus

text: Homo

text: . Those includes that only in human lineage this gene conversion happened, the brain cortex has acquired an important expression of SIGLEC11 specifically in human lineage and the exhibition of  a change in substrate binding in human lineage when compared to that of its counterpart in chimpanzees. Of course the frequency of the contribution of this pseudogene-mediated gene conversion mechanism to functional and adaptive changes in evolution of human is still unknown and so far it has been scarcely explored (Chen et al., 2007). In spite of that, the introduction of positively selective genetic changes by such mechanism can be put forward for consideration by the example of SIGLEC11. Sometimes due to interference of

text: transposable elements

text: in to some members of a gene family, it causes a variation among them and finally it may also cease the rate of gene conversion due to lack of sequence similarity which leads to

text: divergent evolution

text: Since the experiment's inception, Lenski and his colleagues have reported a wide array of genetic changes; some evolutionary adaptations have occurred in all 12 populations, while others have only appeared in one or a few populations. One particularly striking adaptation was the evolution of a strain of E. coli that was able to use citrate as a carbon source in an aerobic environment.

text: Since the experiment's inception, Lenski and his colleagues have reported a wide array of genetic changes; some evolutionary

text: adaptation

text: s have occurred in all 12 populations, while others have only appeared in one or a few populations. One particularly striking adaptation was the evolution of a strain of E. coli that was able to use

text: citrate

text: as a carbon source in an aerobic environment.

text: In natural populations, selective attachment in parenting appears to emerge as a result of the combination of the mobility of the offspring and the social structure of the species. Natural selection is assumed to favor the evolution of mechanisms ensuring that the correct offspring receive the (often costly) parental investment involved in rearing. Sheep are fully mobile shortly after birth and young are likely to mix with unrelated offspring at an early age due to the flock social structure, so the ability to identify and preferentially interact with one’s own offspring is crucial for the correct allocation of maternal resources, principally milk. This is commonly seen in other ungulates and also in seals and primates, all species where the chance of misdirecting parental care is substantial.

text: In natural populations, selective attachment in parenting appears to emerge as a result of the combination of the mobility of the offspring and the social structure of the species.

text: Natural selection

text: is assumed to favor the evolution of mechanisms ensuring that the correct offspring receive the (often costly)

text: parental investment

text: involved in rearing. Sheep are fully mobile shortly after birth and young are likely to mix with unrelated offspring at an early age due to the flock social structure, so the ability to identify and preferentially interact with one’s own offspring is crucial for the correct allocation of maternal resources, principally milk. This is commonly seen in other

text: ungulate

text: s and also in

text: seals

text: and

text: primate

text: s, all species where the chance of misdirecting parental care is substantial.

text: Because of the migration to the Americas and the new introduction of peoples, the different genetic traits of people throughout Latin America show the relation within the people of this region today. Latin Americans today carry the blood of three different populations. These populations include Africans, Europeans, as well as Native people or Native Americans. There has been little research on the study of genetic admixtures because there is uncertainty in the evolution of the different ancestral populations that is. happening between individuals from Latin America. The genes influencing a person's physical appearance is also a result of the contribution of admixture between the three populations. Although the details of admixture of Africans, Europeans, and Native Americans are still unclear, exploration of the genetics of the Latin American population is a still ongoing subject. There have been studies that show how peoples of different countries in Latin America show relative characteristics regarding disease risk, for example.

text: Because of the migration to the Americas and the new introduction of peoples, the different

text: genetic traits

text: of people throughout Latin America show the relation within the people of this region today. Latin Americans today carry the blood of three different

text: populations

text: . These populations include Africans, Europeans, as well as Native people or Native Americans. There has been little research on the study of

text: genetic

text: admixtures

text: because there is

text: uncertainty

text: in the

text: evolution

text: of the different

text: ancestral populations

text: that is. happening between individuals from Latin America. The

text: genes

text: influencing a person's physical appearance is also a result of the contribution of admixture between the three populations. Although the details of admixture of Africans, Europeans, and Native Americans are still unclear, exploration of the genetics of the Latin American population is a still ongoing subject. There have been studies that show how peoples of different countries in Latin America show relative characteristics regarding disease risk, for example.

text: Pigmentation genes show particularly strong evidence of adaptive evolution in non-African populations (Williamson et al. 2007). This is likely to be because those humans that left Africa approximately 50,000 years ago, entered less sunny climates, and so were under new selection pressures to obtain enough Vitamin D from the weakened sunlight.

text: Cochran and Harpending put forward the idea that the development of agriculture has caused an enormous increase in the rate of human evolution, including numerous evolutionary adaptations to the different challenges and lifestyles that resulted. Moreover, they argue that these adaptations have varied across different human populations, depending on factors such as when the various groups developed agriculture, and the extent to which they mixed genetically with other population groups.

text: Cochran and Harpending put forward the idea that the

text: development of agriculture

text: has caused an enormous increase in the rate of

text: human evolution

text: , including numerous evolutionary

text: adaptations

text: to the different challenges and lifestyles that resulted. Moreover, they argue that these adaptations have varied across different human

text: populations

text: , depending on factors such as when the various groups developed agriculture, and the extent to which they mixed genetically with other population groups.

text: Populations located in different environments may be faced with different biotic and abiotic pressures, consequently natural selection may drive the evolution of these populations in different directions. This divergent natural selection can lead to differences in trait values among populations for those traits that are heritable and impact organism fitness. Local adaptation of a variety of traits has been demonstrated in numerous, phylogenetically disparate organisms.

text: Identifying genes on each chromosome is an active area of genetic research. Because researchers use different approaches to predict the number of genes on each chromosome, the estimated number of genes varies. The X chromosome contains about 2000 genes compared to the Y chromosome containing 78 genes, out of the estimated 20,000 to 25,000 total genes in the human genome. Genetic disorders that are due to mutations in genes on the X chromosome are described as X linked.

text: Identifying

text: genes

text: on each chromosome is an active area of genetic research. Because researchers use different approaches to predict the number of genes on each chromosome, the estimated number of genes varies. The X chromosome contains about 2000 genes compared to the Y chromosome containing 78 genes, out of the estimated 20,000 to 25,000 total genes in the human genome.

text: Genetic disorder

text: s that are due to

text: mutation

text: s in genes on the X chromosome are described as X linked.

text: Variations in the toolkit may have produced a large part of the morphological evolution of animals. The toolkit can drive evolution in two ways. A toolkit gene can be expressed in a different pattern, as when the beak of Darwin's large ground-finch was enlarged by the BMP gene, or when snakes lost their legs as distal-less became under-expressed or not expressed at all in the places where other reptiles continued to form their limbs. Or, a toolkit gene can acquire a new function, as seen in the many functions of that same gene, distal-less, which controls such diverse structures as the mandible in vertebrates, legs and antennae in the fruit fly, and eyespot pattern in butterfly wings. Given that small changes in toolbox genes can cause significant changes in body structures, they have often enabled the same function convergently or in parallel. distal-less generates wing patterns in the butterflies Heliconius erato and Heliconius melpomene, which are Müllerian mimics. In so-called facilitated variation, their wing patterns arose in different evolutionary events, but are controlled by the same genes. Developmental changes can contribute directly to speciation.

text: Variations in the toolkit may have produced a large part of the morphological evolution of animals. The toolkit can drive evolution in two ways. A toolkit gene can be expressed in a different pattern, as when the beak of Darwin's

text: large ground-finch

text: was enlarged by the

text: BMP

text: gene, or when snakes lost their legs as distal-less became under-expressed or not expressed at all in the places where other reptiles continued to form their limbs. Or, a toolkit gene can acquire a new function, as seen in the many functions of that same gene, distal-less, which controls such diverse structures as the mandible in vertebrates, legs and antennae in the fruit fly, and

text: eyespot pattern

text: in

text: butterfly

text: wing

text: s. Given that small changes in toolbox genes can cause significant changes in body structures, they have often enabled the same function

text: convergently

text: or

text: in parallel

text: . distal-less generates wing patterns in the butterflies

text: Heliconius erato

text: and

text: Heliconius melpomene

text: , which are

text: Müllerian mimics

text: . In so-called

text: facilitated variation

text: , their wing patterns arose in different evolutionary events, but are controlled by the same genes. Developmental changes can contribute directly to

text: speciation

text: He was one of the three major figures to develop the mathematical theory of population genetics, along with Ronald Fisher and Sewall Wright. Thusly he played an important role in the modern evolutionary synthesis, which is popularly called "neo-Darwinism", as in Richard Dawkins' 1976 work titled The Selfish Gene. He re-established natural selection as the premier mechanism of evolution by explaining it in terms of the mathematical consequences of Mendelian inheritance. He wrote a series of ten papers called A Mathematical Theory of Natural and Artificial Selection, on the numerical formalism underpinning natural selection. It showed that gene frequencies have direction and rates of change; and he pioneered the interaction of natural selection with mutation and animal migration. Haldane's book, The Causes of Evolution (1932), summarised these results, especially in its extensive appendix.

text: He was one of the three major figures to develop the mathematical theory of

text: population genetics

text: , along with

text: Ronald Fisher

text: and

text: Sewall Wright

text: . Thusly he played an important role in the

text: modern evolutionary synthesis

text: , which is popularly called "neo-Darwinism", as in

text: Richard Dawkins

text: ' 1976 work titled

text: The Selfish Gene

text: . He re-established

text: natural selection

text: as the premier mechanism of

text: evolution

text: by explaining it in terms of the mathematical consequences of

text: Mendelian inheritance

text: . He wrote a series of ten papers called

entity_name: A Mathematical Theory of Natural and Artificial Selection

text: A Mathematical Theory of Natural and Artificial Selection

text: , on the numerical formalism underpinning natural selection. It showed that

text: gene frequencies

text: have direction and rates of change; and he pioneered the interaction of natural selection with

text: mutation

text: and

text: animal migration

text: . Haldane's book,

text: The Causes of Evolution

text: (1932), summarised these results, especially in its extensive

text: appendix

text: Neutral theory does not deny the occurrence of natural selection. Hughes writes: "Evolutionary biologists typically distinguish two main types of natural selection: purifying selection, which acts to eliminate deleterious mutations; and positive (Darwinian) selection, which favors advantageous mutations. Positive selection can, in turn, be further subdivided into directional selection, which tends toward fixation of an advantageous allele, and balancing selection, which maintains a polymorphism. The neutral theory of molecular evolution predicts that purifying selection is ubiquitous, but that both forms of positive selection are rare, whereas not denying the importance of positive selection in the origin of adaptations." In another essay, Hughes writes: "Purifying selection is the norm in the evolution of protein coding genes. Positive selection is a relative rarity — but of great interest, precisely because it represents a departure from the norm." A more general and more recent view of molecular evolution is presented by Nei.

text: Neutral theory does not deny the occurrence of natural selection. Hughes writes: "Evolutionary biologists typically distinguish two main types of natural selection:

text: purifying selection

text: , which acts to eliminate deleterious mutations; and positive (Darwinian) selection, which favors advantageous mutations. Positive selection can, in turn, be further subdivided into

text: directional selection

text: , which tends toward fixation of an advantageous allele, and

text: balancing selection

text: , which maintains a

text: polymorphism

text: . The neutral theory of molecular evolution predicts that purifying selection is ubiquitous, but that both forms of positive selection are rare, whereas not denying the importance of positive selection in the origin of adaptations." In another essay, Hughes writes: "Purifying selection is the norm in the evolution of protein coding genes. Positive selection is a relative rarity — but of great interest, precisely because it represents a departure from the norm." A more general and more recent view of molecular evolution is presented by

text: Nei

text: Lester Frank Ward (1841–1913), sometimes referred to as the "father" of American sociology, rejected many of Spencer's theories regarding the evolution of societies. Ward, who was also a botanist and a paleontologist, believed that the law of evolution functioned much differently in human societies than it did in the plant and animal kingdoms, and theorized that the "law of nature" had been superseded by the "law of the mind". He stressed that humans, driven by emotions, create goals for themselves and strive to realize them (most effectively with the modern scientific method) whereas there is no such intelligence and awareness guiding the non-human world. Plants and animals adapt to nature; man shapes nature. While Spencer believed that competition and "survival of the fittest" benefited human society and socio-cultural evolution, Ward regarded competition as a destructive force, pointing out that all human institutions, traditions and laws were tools invented by the mind of man and that that mind designed them, like all tools, to "meet and checkmate" the unrestrained competition of natural forces. Ward agreed with Spencer that authoritarian governments repress the talents of the individual, but he believed that modern democratic societies, which minimized the role of religion and maximized that of science, could effectively support the individual in his or her attempt to fully utilize their talents and achieve happiness. He believed that the evolutionary processes have four stages:

text: Lester Frank Ward (1841–1913), sometimes referred to as the "father" of American sociology, rejected many of Spencer's theories regarding the evolution of societies. Ward, who was also a botanist and a paleontologist, believed that the law of evolution functioned much differently in human societies than it did in the plant and animal kingdoms, and theorized that the "law of nature" had been superseded by the "law of the mind". He stressed that humans, driven by emotions, create goals for themselves and strive to realize them (most effectively with the modern

text: scientific method

text: ) whereas there is no such intelligence and awareness guiding the non-human world. Plants and animals adapt to nature; man shapes nature. While Spencer believed that competition and "survival of the fittest" benefited human society and socio-cultural evolution, Ward regarded competition as a destructive force, pointing out that all human institutions, traditions and laws were tools invented by the mind of man and that that mind designed them, like all tools, to "meet and checkmate" the unrestrained competition of natural forces. Ward agreed with Spencer that authoritarian governments repress the talents of the individual, but he believed that modern democratic societies, which minimized the role of religion and maximized that of science, could effectively support the individual in his or her attempt to fully utilize their talents and achieve happiness. He believed that the evolutionary processes have four stages:

text: Evolutionary psychologists say that natural selection has provided humans with many psychological adaptations, in much the same way that it generated humans' anatomical and physiological adaptations. As with adaptations in general, psychological adaptations are said to be specialized for the environment in which an organism evolved, the environment of evolutionary adaptedness, or EEA. Sexual selection provides organisms with adaptations related to mating. For male mammals, which have a relatively fast reproduction rate, sexual selection leads to adaptations that help them compete for females. For female mammals, with a relatively slow reproduction rate, sexual selection leads to choosiness, which helps females select higher quality mates. Charles Darwin described both natural selection and sexual selection, but he relied on group selection to explain the evolution of self-sacrificing behavior. Group selection is a weak explanation because in any group the less self-sacrificing animals will be more likely to survive and the group will become less self-sacrificing.

text: Evolutionary psychologists say that

text: natural selection

text: has provided humans with many

text: psychological adaptation

text: s, in much the same way that it generated humans' anatomical and physiological adaptations. As with adaptations in general, psychological adaptations are said to be specialized for the environment in which an organism evolved, the environment of evolutionary adaptedness, or EEA.

text: Sexual selection

text: provides organisms with adaptations related to mating. For

text: male mammals

text: , which have a relatively fast reproduction rate, sexual selection leads to adaptations that help them compete for females. For

text: female mammals

text: , with a relatively slow reproduction rate, sexual selection leads to choosiness, which helps females select higher quality mates. Charles Darwin described both natural selection and sexual selection, but he relied on group selection to explain the evolution of self-sacrificing behavior. Group selection is a weak explanation because in any group the less self-sacrificing animals will be more likely to survive and the group will become less self-sacrificing.

text: Metabolism can be considered an important part of exaptation. As one of the oldest biological systems and being central to life on the Earth, studies have shown that metabolism may be able to use exaptation in order to be fit, given some new set of conditions or environment. Studies have shown that up to 44 carbon sources are viable for metabolism to successfully take place and that any one adaptation in these specific metabolic systems is due to multiple exaptations. Taking this perspective, exaptations are important in the origination of adaptations in general. A recent example comes from Richard Lenski's E. coli long-term evolution experiment, in which aerobic growth on citrate arose in one of twelve populations after 31,000 generations of evolution. Genomic analysis by Blount and colleagues showed that this novel trait was due to a gene duplication that caused oxic expression of a citrate transporter gene that is normally only expressed under anoxic conditions, thus exapting it for aerobic use. Metabolic systems have the potential to innovate without adaptive origins.

text: Metabolism can be considered an important part of exaptation. As one of the oldest biological systems and being central to life on the Earth, studies have shown that metabolism may be able to use exaptation in order to be fit, given some new set of conditions or environment. Studies have shown that up to 44 carbon sources are viable for metabolism to successfully take place and that any one adaptation in these specific metabolic systems is due to multiple exaptations. Taking this perspective, exaptations are important in the origination of adaptations in general. A recent example comes from

text: Richard Lenski

text: 's

text: E. coli long-term evolution experiment

text: , in which aerobic growth on

text: citrate

text: arose in one of twelve populations after 31,000 generations of evolution. Genomic analysis by

text: Blount

text: and colleagues showed that this novel trait was due to a gene duplication that caused oxic expression of a citrate transporter gene that is normally only expressed under anoxic conditions, thus exapting it for aerobic use. Metabolic systems have the potential to innovate without adaptive origins.

text: Coevolution can occur at many biological levels: it can be as microscopic as correlated mutations between amino acids in a protein or as macroscopic as covarying traits between different species in an environment. Each party in a coevolutionary relationship exerts selective pressures on the other, thereby affecting each other's evolution. Coevolution of different species includes the evolution of a host species and its parasites (host–parasite coevolution), and examples of mutualism evolving through time. Evolution in response to abiotic factors, such as climate change, is not biological coevolution (since climate is not alive and does not undergo biological evolution).

text: Coevolution can occur at many biological levels: it can be as microscopic as correlated mutations between amino acids in a protein or as macroscopic as covarying traits between different species in an environment. Each party in a coevolutionary relationship exerts

text: selective pressures

text: on the other, thereby affecting each other's evolution. Coevolution of different species includes the evolution of a

text: host

text: species and its

text: parasite

text: s (

text: host–parasite coevolution

text: ), and examples of

text: mutualism

text: evolving through time. Evolution in response to

text: abiotic factor

text: s, such as

text: climate change

text: , is not biological coevolution (since climate is not alive and does not undergo biological evolution).

text: The 1859 publication of Charles Darwin's theory of natural selection put forward an explanation for complexity and adaptation, which reflects scientific consensus on the origins of biological diversity, and provides a counter-argument to the watchmaker analogy. Evolutionary biologist Richard Dawkins also referred to the analogy in his 1986 book The Blind Watchmaker giving his explanation of evolution. In the 19th century, deists, who championed the watchmaker analogy, held that Darwin's theory fit with "the principle of uniformitarianism—the idea that all processes in the world occur now as they have in the past" and that deistic evolution "provided an explanatory framework for understanding species variation in a mechanical universe."

text: The 1859 publication of

text: Charles Darwin

text: 's

text: theory

text: of

text: natural selection

text: put forward an explanation for complexity and

text: adaptation

text: , which reflects scientific consensus on the origins of biological diversity, and provides a counter-argument to the watchmaker analogy. Evolutionary biologist

text: Richard Dawkins

text: also referred to the analogy in his 1986 book

text: The Blind Watchmaker

text: giving his explanation of

text: evolution

text: . In the 19th century,

text: deists

text: , who championed the watchmaker analogy, held that Darwin's theory fit with "the principle of

text: uniformitarianism

text: —the idea that all processes in the world occur now as they have in the past" and that

text: deistic evolution

text: "provided an explanatory framework for understanding species variation in a mechanical universe."

text: Genes involved in lactose metabolism show particularly strong evidence of adaptive evolution amongst the genes involved in nutrition. A mutation linked to lactase persistence shows very strong evidence of adaptive evolution in European and American populations (Williamson et al. 2007), populations where pastoral farming for milk has been historically important.

text: Genes are the nodes and the edges are directed. A gene serves as the source of a direct regulatory edge to a target gene by producing an RNA or protein molecule that functions as a transcriptional activator or inhibitor of the target gene. If the gene is an activator, then it is the source of a positive regulatory connection; if an inhibitor, then it is the source of a negative regulatory connection. Computational algorithms take as primary input data measurements of mRNA expression levels of the genes under consideration for inclusion in the network, returning an estimate of the network topology. Such algorithms are typically based on linearity, independence or normality assumptions, which must be verified on a case-by-case basis. Clustering or some form of statistical classification is typically employed to perform an initial organization of the high-throughput mRNA expression values derived from microarray experiments, in particular to select sets of genes as candidates for network nodes. The question then arises: how can the clustering or classification results be connected to the underlying biology? Such results can be useful for pattern classification – for example, to classify subtypes of cancer, or to predict differential responses to a drug (pharmacogenomics). But to understand the relationships between the genes, that is, to more precisely define the influence of each gene on the others, the scientist typically attempts to reconstruct the transcriptional regulatory network. This can be done by data integration in dynamic models supported by background literature, or information in public databases, combined with the clustering results. The modelling can be done by a Boolean network, by Ordinary differential equations or Linear regression models, e.g. Least-angle regression, by Bayesian network or based on Information theory approaches. For instance it can be done by the application of a correlation-based inference algorithm, as will be discussed below, an approach which is having increased success as the size of the available microarray sets keeps increasing

text: Genes are the nodes and the edges are directed. A gene serves as the source of a direct regulatory edge to a target gene by producing an

text: RNA

text: or protein molecule that functions as a transcriptional activator or inhibitor of the target gene. If the gene is an activator, then it is the source of a positive regulatory connection; if an inhibitor, then it is the source of a negative regulatory connection. Computational algorithms take as primary input data measurements of

text: mRNA

text: expression levels of the genes under consideration for inclusion in the network, returning an estimate of the network

text: topology

text: . Such algorithms are typically based on linearity, independence or normality assumptions, which must be verified on a case-by-case basis. Clustering or some form of statistical classification is typically employed to perform an initial organization of the high-throughput mRNA expression values derived from microarray experiments, in particular to select sets of genes as candidates for network nodes. The question then arises: how can the clustering or classification results be connected to the underlying biology? Such results can be useful for pattern classification – for example, to classify subtypes of

text: cancer

text: , or to predict differential responses to a

text: drug

text: (pharmacogenomics). But to understand the relationships between the genes, that is, to more precisely define the influence of each gene on the others, the scientist typically attempts to reconstruct the transcriptional regulatory network. This can be done by data integration in dynamic models supported by background literature, or information in public

text: database

text: s, combined with the clustering results. The modelling can be done by a

text: Boolean network

text: , by

text: Ordinary differential equation

text: s or

text: Linear regression

text: models, e.g.

text: Least-angle regression

text: , by

text: Bayesian network

text: or based on

text: Information theory

text: approaches. For instance it can be done by the application of a correlation-based inference algorithm, as will be discussed below, an approach which is having increased success as the size of the available microarray sets keeps increasing

text: Natural selection will only cause evolution if there is enough genetic variation in a population. Before the discovery of Mendelian genetics, one common hypothesis was blending inheritance. But with blending inheritance, genetic variance would be rapidly lost, making evolution by natural selection implausible. The Hardy–Weinberg principle provides the solution to how variation is maintained in a population with Mendelian inheritance. The frequencies of alleles (variations in a gene) will remain constant in the absence of selection, mutation, migration and genetic drift.

text: Natural selection will only cause evolution if there is enough

text: genetic variation

text: in a population. Before the discovery of Mendelian genetics, one common hypothesis was

text: blending inheritance

text: . But with blending inheritance, genetic variance would be rapidly lost, making evolution by natural selection implausible. The

text: Hardy–Weinberg principle

text: provides the solution to how variation is maintained in a population with Mendelian inheritance. The frequencies of alleles (variations in a gene) will remain constant in the absence of selection, mutation, migration and genetic drift.

text: Integration can be found at the genetic level due to gene linkage. Gene linkage involves multiple genes being inherited together during meiosis because they are close to each other on the same chromosome.  Alleles at different loci can be inherited together if they are tightly linked. Large genetic correlations can only be upheld if the loci that influence different characters are tightly linked, or if high levels of inbreeding in the population occur.  Even if selection favors the correlations, it will not be maintained unless those conditions are met.  Selection will favor tight linkage because it is maintained better. Poorly linked genetic correlations will not last.  Transposition allows the loci at different locations on the chromosome to move so that they can become close to each other and be inherited together.  This is significant to understanding the relationship between phenotypic integration and evolution because it is one of the mechanisms of how multiple traits that are connected to each other to evolve and change together.  For instance, the Papilio dardanus butterflies come in three different forms, each mimicking a different distasteful butterfly species.  Multiple loci contribute to these different forms, and a butterfly with alleles for form A at one locus and B at another locus would have poor fitness.  However, the multiple loci are tightly linked, so they are inherited together as a single allele.  Through transposition, these multiple loci ended up close to each other.  Mutations among these linked genes are the nonadaptive fuel which can create evolution.  Evolution may also occur because the integration may have an adaptive advantage in a particular environment for an organism.  It is also important to recognize that not only can the traits be inherited together, but inherited separately and selected together.  Another important example of phenotypic integration evolving over time is the relationship between the neurocranium and the brain.  Over the last 150 million years the number of bones in the brain has decreased while the size of the brain in mammals has changed.  Integration between the brain and the skull has evolved over this time period to reduce the number of bones in the cranium, while increasing the size of the brain.  This relationship between correlated traits has played an important role in the evolution of mammalian cranium structure and brain size. Finally, development is another crucial cause of phenotypic integration that has evolved over time.  Cell-signaling pathways which utilize integration in the form of complex interactions among specific cells in the pathway are crucial to proper development in many organisms.  The interactions among the cells in the pathway, and the interaction of the pathways with other pathways have evolved over time to create complex structures.

text: In 1924, J.B.S. Haldane calculated, using a simple general selection model, the selective advantage necessary for the recorded natural evolution of peppered moths, based on the assumption that in 1848 the frequency of dark-coloured moths was 2%, and by 1895 it was 95%. The dark-coloured, or melanic, form would have had to be 50% more fit than the typical, light-coloured form. Even taking into consideration possible errors in the model, this reasonably excluded the stochastic process of genetic drift, because the changes were too fast. Haldane's statistical analysis of selection for the melanic variant in peppered moths became a well known part of his effort to demonstrate that mathematical models that combined natural selection with Mendelian genetics could explain evolution — an effort that played a key role in the foundation of the discipline of population genetics, and the beginnings of the modern evolutionary synthesis.

text: In 1924,

text: J.B.S. Haldane

text: calculated, using a simple

text: general selection model

text: , the selective advantage necessary for the recorded natural evolution of peppered moths, based on the assumption that in 1848 the frequency of dark-coloured moths was 2%, and by 1895 it was 95%. The dark-coloured, or melanic, form would have had to be 50% more fit than the typical, light-coloured form. Even taking into consideration possible errors in the model, this reasonably excluded the

text: stochastic

text: process of genetic drift, because the changes were too fast. Haldane's statistical analysis of selection for the melanic variant in peppered moths became a well known part of his effort to demonstrate that mathematical models that combined natural selection with Mendelian genetics could explain evolution — an effort that played a key role in the foundation of the discipline of

text: population genetics

text: , and the beginnings of the

text: modern evolutionary synthesis

text: Natural selection is the evolutionary process by which heritable traits that increase an individual's fitness become more common, and heritable traits that decrease an individual's fitness become less common.

text: Natural selection is the evolutionary process by which

text: heritable

text: traits that increase an individual's fitness become more common, and heritable traits that decrease an individual's fitness become less common.

text: The concept of devolution on the other hand, requires that there be a preferred hierarchy of structure and function, and that evolution must mean "progress" to "more advanced" organisms. For example, it could be said that "feet are better than hooves" or "lungs are better than gills", so their development is "evolutionary" whereas change to an inferior or "less advanced" structure would be called "devolution". In reality an evolutionary biologist defines all heritable changes to relative frequencies of the genes or indeed to epigenetic states in the gene pool as evolution. All gene pool changes that lead to increased fitness in terms of appropriate aspects of reproduction are seen as (neo-)Darwinian adaptation because, for the organisms possessing the changed structures, each is a useful adaptation to their circumstances. For example, hooves have advantages for running quickly on plains, which benefits horses, and feet offer advantages in climbing trees, which some ancestors of humans did.

text: The concept of devolution on the other hand, requires that there be a preferred hierarchy of structure and function, and that evolution must mean "progress" to "more advanced" organisms. For example, it could be said that "feet are better than

text: hooves

text: " or "

text: lung

text: s are better than

text: gill

text: s", so their development is "evolutionary" whereas change to an inferior or "less advanced" structure would be called "devolution". In reality an evolutionary biologist defines all heritable changes to relative frequencies of the genes or indeed to

text: epigenetic

text: states in the gene pool as evolution. All gene pool changes that lead to increased fitness in terms of appropriate aspects of reproduction are seen as (neo-)Darwinian adaptation because, for the organisms possessing the changed structures, each is a useful adaptation to their circumstances. For example, hooves have advantages for running quickly on plains, which benefits horses, and feet offer advantages in climbing trees, which some ancestors of humans did.

text: High-altitude adaptations are an example of convergent evolution, with adaptations occurring simultaneously on three continents. Tibetan humans and Tibetan domestic dogs found the genetic mutation in both species, EPAS1. This mutation has not been seen in Andean humans, showing the effect of a shared environment on evolution

text: High-altitude adaptations are an example of

text: convergent evolution

text: , with adaptations occurring simultaneously on three continents. Tibetan humans and Tibetan domestic dogs found the genetic mutation in both species, EPAS1. This mutation has not been seen in Andean humans, showing the effect of a shared environment on evolution

text: When Morgan took the professorship in experimental zoology, he became increasingly focused on the mechanisms of heredity and evolution. He had published Evolution and Adaptation (1903); like many biologists at the time, he saw evidence for biological evolution (as in the common descent of similar species) but rejected Darwin's proposed mechanism of natural selection acting on small, constantly produced variations.

text: When Morgan took the professorship in experimental zoology, he became increasingly focused on the mechanisms of heredity and evolution. He had published Evolution and Adaptation (1903); like many biologists at the time, he saw evidence for biological evolution (as in the

text: common descent

text: of similar species) but rejected Darwin's proposed mechanism of

text: natural selection

text: acting on small, constantly produced variations.

text: These genes were first shown to be arranged into a physically-linked chromosomal cluster in amphioxus, an invertebrate with a single member of each of the three gene families. All the ParaHox genes in the amphioxus genome are therefore in the ParaHox gene cluster. In contrast, the human genome has six ParaHox genes (GSX1, GSX2, PDX1, CDX1, CDX2, CDX4), of which three genes (GSX1, PDX1 (=IPF1), CDX2) are physically linked to form a human ParaHox gene cluster on chromosome 13. Mouse has a homologous ParaHox gene cluster on chromosome 5. The other three human ParaHox genes are remnants from duplicated ParaHox gene clusters that were generated in the 2R genome duplications at the base of vertebrate evolution. Some vertebrates, notably chondrichthyan fish and coelacanths, have retained an additional ParaHox gene (PDX2).

text: These genes were first shown to be arranged into a physically-linked chromosomal cluster in

text: amphioxus

text: , an

text: invertebrate

text: with a single member of each of the three gene families. All the ParaHox genes in the amphioxus genome are therefore in the ParaHox gene cluster. In contrast, the human genome has six ParaHox genes (

text: ), of which three genes (

text: ) are physically linked to form a human ParaHox gene cluster on

text: chromosome 13

text: . Mouse has a homologous ParaHox gene cluster on chromosome 5. The other three human ParaHox genes are remnants from duplicated ParaHox gene clusters that were generated in the

text: genome duplications

text: at the base of vertebrate evolution. Some vertebrates, notably

text: chondrichthyan

text: fish and

text: coelacanths

text: , have retained an additional ParaHox gene (PDX2).

text: Natural selection can act at different levels of organisation, such as genes, cells, individual organisms, groups of organisms and species. Selection can act at multiple levels simultaneously. An example of selection occurring below the level of the individual organism are genes called transposons, which can replicate and spread throughout a genome. Selection at a level above the individual, such as group selection, may allow the evolution of cooperation, as discussed below.

text: Natural selection can act at

text: different levels of organisation

text: , such as genes, cells, individual organisms, groups of organisms and species. Selection can act at multiple levels simultaneously. An example of selection occurring below the level of the individual organism are genes called

text: transposon

text: s, which can replicate and spread throughout a genome. Selection at a level above the individual, such as

text: group selection

text: , may allow the evolution of cooperation, as discussed below.

text: The most widely accepted theory of evolution states that organisms will diversify based on natural selection where changes caused by mutations improve the organism's chance of survival. Adaptive mutation states that rather than mutations and evolution being random, they are in response to specific stresses. In other words, the mutations that occur are more beneficial and specific to the given stress, instead of random and not a response to anything in particular. It should be noted that the term stress refers to any change in environment, such as temperature, nutrients, population size, etc. Tests with microorganisms have found that for adaptive mutation, more of the mutations observed after a given stress were effective at dealing with the stress than chance alone would suggest is possible. This theory of adaptive mutation was first brought to academic attention in the 1980s by John Cairns.

text: Selective breeding to enhance the most beneficial traits is the structure upon which agriculture is built.  Genetic manipulation expedited the evolution process to make crop plants and animals larger and more disease resistant. Research into wild-type mutations has allowed the creation of genetically modified crops that are more efficient food producers, which is a major reason behind world hunger being at all-time lows .  Genetic alteration of plants leads to not only larger crop production, but also more nutritious products, allowing isolated populations to receive vital vitamins and mineral that would otherwise be unavailable to them.  Utilization of these wild-type mutations has also led to plants capable of growing in extremely arid environments, making more of the planet habitable than ever before.  As more is understood about these genes, agriculture will continue to become a more efficient process, which will be relied upon to sustain a continually growing population. Amplification of advantageous genes allows the best traits in a population to be present at much higher percentages than normal, although this practice has been the subject of some ethical debate.  These changes have also been the reason behind certain plants and animals being almost unrecognizable when compared to their ancestral lines.

text: Critics of Dawkins's biological approach suggest that taking the gene as the unit of selection (a single event in which an individual either succeeds or fails to reproduce) is misleading; the gene could be better described, they say, as a unit of evolution (the long-term changes in allele frequencies in a population). In The Selfish Gene, Dawkins explains that he is using George C. Williams's definition of the gene as "that which segregates and recombines with appreciable frequency." Another common objection is that a gene cannot survive alone, but must cooperate with other genes to build an individual, and therefore a gene cannot be an independent "unit". In The Extended Phenotype, Dawkins suggests that from an individual gene's viewpoint, all other genes are part of the environment to which it is adapted.

text: Critics of Dawkins's biological approach suggest that taking the

text: gene

text: as the unit of selection (a single event in which an individual either succeeds or fails to reproduce) is misleading; the gene could be better described, they say, as a unit of evolution (the long-term changes in

text: allele

text: frequencies in a population). In The Selfish Gene, Dawkins explains that he is using

text: George C. Williams

text: 's definition of the gene as "that which segregates and recombines with appreciable frequency." Another common objection is that a gene cannot survive alone, but must cooperate with other genes to build an individual, and therefore a gene cannot be an independent "unit". In The Extended Phenotype, Dawkins suggests that from an individual gene's viewpoint, all other genes are part of the environment to which it is adapted.

text: In 1859, Charles Darwin set out his theory of evolution by natural selection as an explanation for adaptation and speciation. He defined natural selection as the "principle by which each slight variation [of a trait], if useful, is preserved." The concept was simple but powerful: individuals best adapted to their environments are more likely to survive and reproduce. As long as there is some variation between them and that variation is heritable, there will be an inevitable selection of individuals with the most advantageous variations. If the variations are heritable, then differential reproductive success leads to a progressive evolution of particular populations of a species, and populations that evolve to be sufficiently different eventually become different species.

text: In 1859, Charles Darwin set out his theory of evolution by natural selection as an explanation for

text: adaptation

text: and speciation. He defined natural selection as the "principle by which each slight variation [of a trait], if useful, is preserved." The concept was simple but powerful: individuals best adapted to their environments are more likely to survive and reproduce. As long as there is some variation between them and that variation is

text: heritable

text: , there will be an inevitable selection of individuals with the most advantageous variations. If the variations are heritable, then differential reproductive success leads to a progressive evolution of particular populations of a species, and populations that evolve to be sufficiently different eventually become different species.

text: Sports are the most extensively studied non-instrumental aspect of human culture. A sport, in this article, will be defined as a "subset of games that require phsical skill" .  therefore excluding games of chance. Sports are both ornamental and competitive aspects of culture, as winning is a central component of games, and ornamental because there is no direct evolutionary function or use of sport . Sports are responsible for developing moral codes, collaborative networks, social capital, and social status .  Sports also help construct collective cultural identities . The evolution of sport as a cultural invention can be linked to specific evolutionary processes. Applying cultural evolution theory, framing culture and the development of sports and games as a system analogous to biological adaptations and evolution, as well as understanding how biologically evolved mechanisms of the brain have created conditions for culture and culturally expressive activities such as sport, we can understand how both sexes have evolved to compete in sport. These theories are cultural evolution theory and evolutionary psychology respectively . The third and most compelling perspective, dual-inheritance theory, a fragment of cultural evolution theory, frames the human ability to create culture as depending on both biologically shaped learning mechanisms, innate mating needs and goals, and domain biases that lead to cultural transmission... In other words, culture is an adaptation.These sections will illuminate the different hypotheses for the driving force behind sport evolution.

text: Sports are the most extensively studied non-instrumental aspect of human culture. A sport, in this article, will be defined as a "subset of games that require phsical skill" .  therefore excluding games of chance. Sports are both ornamental and competitive aspects of culture, as winning is a central component of games, and ornamental because there is no direct evolutionary function or use of sport . Sports are responsible for developing moral codes, collaborative networks,

text: social capital

text: , and social status .  Sports also help construct collective cultural identities . The evolution of sport as a cultural invention can be linked to specific evolutionary processes. Applying

text: cultural evolution theory

text: , framing culture and the development of sports and games as a system analogous to biological adaptations and evolution, as well as understanding how biologically evolved mechanisms of the brain have created conditions for culture and culturally expressive activities such as sport, we can understand how both sexes have evolved to compete in sport. These theories are cultural evolution theory and

text: evolutionary psychology

text: respectively . The third and most compelling perspective,

text: dual-inheritance theory

text: , a fragment of cultural evolution theory, frames the human ability to create culture as depending on both biologically shaped learning mechanisms, innate mating needs and goals, and domain biases that lead to

text: cultural transmission

text: ... In other words, culture is an adaptation.These sections will illuminate the different hypotheses for the driving force behind sport evolution.

text: Natural selection acts on an organism's phenotype, or physical characteristics. Phenotype is determined by an organism's genetic make-up (genotype) and the environment in which the organism lives. When different organisms in a population possess different versions of a gene for a certain trait, each of these versions is known as an allele. It is this genetic variation that underlies differences in phenotype. An example is the ABO blood type antigens in humans, where three alleles govern the phenotype.

text: Natural selection acts on an organism's phenotype, or physical characteristics. Phenotype is determined by an organism's genetic make-up (genotype) and the environment in which the organism lives. When different organisms in a population possess different versions of a gene for a certain trait, each of these versions is known as an

text: allele

text: . It is this genetic variation that underlies differences in phenotype. An example is the

text: ABO

text: blood type

text: antigen

text: s in humans, where three alleles govern the phenotype.

text: 417 genes involved in the immune system showed strong evidence of adaptive evolution in the study of Nielsen et al. (2005a). This is probably because the immune genes may become involved in a evolutionary arms race with bacteria and viruses. These pathogens evolve very rapidly, so selection pressures change quickly, giving more opportunity for adaptive evolution.

text: 417 genes involved in the immune system showed strong evidence of adaptive evolution in the study of Nielsen et al. (2005a). This is probably because the immune genes may become involved in a

text: evolutionary arms race

text: with bacteria and viruses. These pathogens evolve very rapidly, so selection pressures change quickly, giving more opportunity for adaptive evolution.

text: The McDonald–Kreitman test is a statistical test often used by evolution and population biologists to detect and measure the amount of adaptive evolution within a species by determining whether adaptive evolution has occurred, and the proportion of substitutions that resulted from positive selection (also known as directional selection). To do this, the McDonald–Kreitman test compares the amount of variation within a species (polymorphism) to the divergence between species (substitutions) at two types of sites, neutral and nonneutral. A substitution refers to a nucleotide that is fixed within one species, but a different nucleotide is fixed within a second species at the same base pair  of homologous DNA sequences. A site is nonneutral if it is either advantageous or deleterious. The two types of sites can be either synonymous or nonsynonymous within a protein-coding region. In a protein-coding sequence of DNA, a site is synonymous if a point mutation at that site would not change the amino acid, also known as a silent mutation. Because the mutation did not result in a change in the amino acid that was originally coded for by the protein-coding sequence, the phenotype, or the observable trait, of the organism is generally unchanged by the silent mutation. A site in a protein-coding sequence of DNA is nonsynonymous if a point mutation at that site results in a change in the amino acid, resulting in a change in the organism's phenotype. Typically, silent mutations in protein-coding regions are used as the "control" in the McDonald–Kreitman test.

text: The McDonald–Kreitman test is a statistical test often used by evolution and population biologists to detect and measure the amount of

text: adaptive evolution

text: within a species by determining whether adaptive evolution has occurred, and the proportion of substitutions that resulted from positive selection (also known as

text: directional selection

text: ). To do this, the McDonald–Kreitman test compares the amount of variation within a species (

text: polymorphism

text: ) to the divergence between species (substitutions) at two types of sites, neutral and nonneutral. A substitution refers to a nucleotide that is fixed within one species, but a different nucleotide is fixed within a second species at the same base pair  of homologous DNA sequences. A site is nonneutral if it is either advantageous or deleterious. The two types of sites can be either synonymous or nonsynonymous within a protein-coding region. In a protein-coding sequence of DNA, a site is synonymous if a

text: point mutation

text: at that site would not change the amino acid, also known as a

text: silent mutation

text: . Because the mutation did not result in a change in the amino acid that was originally coded for by the protein-coding sequence, the phenotype, or the observable trait, of the organism is generally unchanged by the silent mutation. A site in a protein-coding sequence of DNA is nonsynonymous if a point mutation at that site results in a change in the amino acid, resulting in a change in the organism's phenotype. Typically, silent mutations in protein-coding regions are used as the "control" in the McDonald–Kreitman test.

text: Indeed, a recently proposed mathematical model to compare the adaptive value of different schemes of inheritance by Rivoire and Leibler does appear to lend credence to the MRK model. The authors considered three biological phenomena that are often considered to be either irrelevant to evolution, or absent because “forbidden” namely, i) role of phenotypic plasticity and developmental canalization, ii) reverse flow of information from phenotype to genotype (or its absence), and iii) conditions under which direct integration of information into the transmitted genotype is logically excluded as a consequence of natural selection. Their model allows for variations to be inherited, randomly produced, or environmentally induced, and, irrespectively, to be either transmitted or not during reproduction. The adaptation of the different schemes for processing variations is quantified for a range of fluctuating environments, following an approach that links quantitative genetics with stochastic control theory. Interestingly, when the authors conducted a Gedankenexperiment (German for a thought-experiment) that allowed them to compare the Darwinian and Lamarckian “modalities,” and test the conjecture that each of them is tuned to a different type of selective pressure, they found that the main controlling parameter in the model appears to be the correlation a of the environmental fluctuations, with the Lamarckian modality systematically becoming more favorable when this correlation is large, in line with the intuition that transmitting acquired information is beneficial when the selective pressure experienced by the offspring is sufficiently similar to that experienced by the parents.

text: In modern times, the term evolution is widely used, but the terms evolutionism and evolutionist are seldom used in the scientific community to refer to the biological discipline as the term is considered both redundant and anachronistic, though it has been used by creationists in discussing the creation-evolution controversy.

text: In modern times, the term

text: evolution

text: is widely used, but the terms evolutionism and evolutionist are seldom used in the

text: scientific community

text: to refer to the biological discipline as the term is considered both redundant and anachronistic, though it has been used by creationists in discussing the

text: creation-evolution controversy

text: Recent findings have confirmed important examples of heritable changes that cannot be explained by changes to the sequence of nucleotides in the DNA. These phenomena are classed as epigenetic inheritance systems. DNA methylation marking chromatin, self-sustaining metabolic loops, gene silencing by RNA interference and the three-dimensional conformation of proteins (such as prions) are areas where epigenetic inheritance systems have been discovered at the organismic level. Developmental biologists suggest that complex interactions in genetic networks and communication among cells can lead to heritable variations that may underlay some of the mechanics in developmental plasticity and canalisation. Heritability may also occur at even larger scales. For example, ecological inheritance through the process of niche construction is defined by the regular and repeated activities of organisms in their environment. This generates a legacy of effects that modify and feed back into the selection regime of subsequent generations. Descendants inherit genes plus environmental characteristics generated by the ecological actions of ancestors. Other examples of heritability in evolution that are not under the direct control of genes include the inheritance of cultural traits and symbiogenesis.

text: Recent findings have confirmed important examples of heritable changes that cannot be explained by changes to the sequence of

text: nucleotide

text: s in the DNA. These phenomena are classed as

text: epigenetic

text: inheritance systems.

text: DNA methylation

text: marking

text: chromatin

text: , self-sustaining metabolic loops, gene silencing by

text: RNA interference

text: and the three-dimensional

text: conformation

text: of

text: protein

text: s (such as

text: prion

text: s) are areas where epigenetic inheritance systems have been discovered at the organismic level. Developmental biologists suggest that complex interactions in

text: genetic networks

text: and communication among cells can lead to heritable variations that may underlay some of the mechanics in

text: developmental plasticity

text: and

text: canalisation

text: . Heritability may also occur at even larger scales. For example, ecological inheritance through the process of

text: niche construction

text: is defined by the regular and repeated activities of organisms in their environment. This generates a legacy of effects that modify and feed back into the selection regime of subsequent generations. Descendants inherit genes plus environmental characteristics generated by the ecological actions of ancestors. Other examples of heritability in evolution that are not under the direct control of genes include the inheritance of

text: cultural traits

text: and

text: symbiogenesis

text: In discussing Gould's perspective, Sterelny begins with two fundamental distinctions that Gould saw between his viewpoint and that of the Dawkins camp. Firstly, Gould thought that gene selectionists misrepresent the role of genes in microevolution, ascribing a causal role in evolution, rather than by-product record of evolutionary change. Moreover, evolutionary biologists have often neglected non-selective possibilities when formulating hypotheses about microevolutionary change. For example, contemporary sex differences in human males and females need not be adaptations, but could be evolutionary vestiges of a greater sexual dimorphism in ancestral species.

text: In discussing Gould's perspective, Sterelny begins with two fundamental distinctions that Gould saw between his viewpoint and that of the Dawkins camp. Firstly, Gould thought that gene selectionists misrepresent the role of genes in

text: microevolution

text: , ascribing a causal role in evolution, rather than by-product record of evolutionary change. Moreover, evolutionary biologists have often neglected non-selective possibilities when formulating hypotheses about microevolutionary change. For example, contemporary sex differences in human males and females need not be adaptations, but could be evolutionary vestiges of a greater sexual dimorphism in ancestral species.

text: Evolutionary psychology primarily uses the theories of natural selection, sexual selection, and inclusive fitness to explain the evolution of psychological adaptations.

text: In order to properly evaluate the Evolution/Creation debate, one must first understand what is meant by 'creation' and what is meant by 'evolution'. 'Creation' can be defined as the act of God in which the universe, and everything in it, was created out of nothing. While there are various points of view within the Creationist camp, the Young Earth Creationists seem to have the greatest number of differences from Evolutionists. When the word 'creation' or 'Creationist' are used in this article, that is what is meant by it. 'Evolution' can be defined as the naturalistic process by which life arose from chemicals, and, broadly speaking, how the universe arose from the Big Bang. This is also called 'macro-evolution'. 'Micro-evolution' generally relates to the changes to species that result in specification and adaptation. The term 'evolution' in this article refers to 'macro-evolution'. There should also be a clarification concerning the word 'theory'. In the context of science, a theory is an explanation for a natural phenomena that has been tested time and time again to be shown with near certainty to be true.

text: In Russia, the idea of "mutual aid" was used to explain evolution rather than the struggle for existence. Peter Kropotkin wrote the book Mutual Aid: A Factor of Evolution in response to the idea of evolution by natural selection. Due to the vast landscape and lack of population in proportion to land, many people of Russia did not see the struggle for existence and could not relate to Malthus's ideas on population. Thus, it was concluded that cooperation, which is more successful in battling the abiotic environment, rather than competition is a driving factor in natural selection. Rather than a struggle for existence, a mutual struggle and mutual aid drives natural selection. Kropotkin believed that Wallace and Darwin saw the struggle for existence because of their coastal location and overpopulated areas of study.

text: In

text: Russia

text: , the idea of "mutual aid" was used to explain evolution rather than the struggle for existence.

text: Peter Kropotkin

text: wrote the book

text: Mutual Aid: A Factor of Evolution

text: in response to the idea of evolution by natural selection. Due to the vast landscape and lack of population in proportion to land, many people of Russia did not see the struggle for existence and could not relate to Malthus's ideas on population. Thus, it was concluded that cooperation, which is more successful in battling the abiotic environment, rather than competition is a driving factor in natural selection. Rather than a struggle for existence, a mutual struggle and mutual aid drives natural selection. Kropotkin believed that Wallace and Darwin saw the struggle for existence because of their coastal location and overpopulated areas of study.

text: Evolutionary algorithms form a subset of evolutionary computation in that they generally only involve techniques implementing mechanisms inspired by biological evolution such as reproduction, mutation, recombination, natural selection and survival of the fittest. Candidate solutions to the optimization problem play the role of individuals in a population, and the cost function determines the environment within which the solutions "live" (see also fitness function). Evolution of the population then takes place after the repeated application of the above operators.

text: Evolutionary algorithms

text: form a subset of evolutionary computation in that they generally only involve techniques implementing mechanisms inspired by

text: biological evolution

text: such as

text: reproduction

text: mutation

text: recombination

text: natural selection

text: and

text: survival of the fittest

text: Candidate solutions

text: to the optimization problem play the role of individuals in a population, and the

text: cost function

text: determines the environment within which the solutions "live" (see also

text: fitness function

text: Evolution

text: of the population then takes place after the repeated application of the above operators.

text: If only Genic variance (σ) is used in the numerator, the heritability may be called "narrow sense" (h). It quantifies the extent to which phenotypic variance is determined by Fisher's substitution expectations variance.   \begin{align} h^2 & = \frac {\sigma ^2_A}{\sigma ^2_P} \\ & = \frac {\sigma ^2_a + \sigma ^2_d + cov_{ad}}{\sigma ^2_P} \end{align} Fisher proposed that this narrow-sense heritability might be appropriate in considering the results of natural selection, focusing as it does on change-ability, that is upon "adaptation". He proposed it with regard to quantifying Darwinian evolution.

text: Darwin had convinced most scientists that evolution as descent with modification was correct, and he was regarded as a great scientist who had revolutionised ideas. Though few agreed with his view that "natural selection has been the main but not the exclusive means of modification", he was honoured in June 1909 by more than 400 officials and scientists from across the world who met in Cambridge to commemorate his centenary and the fiftieth anniversary of On the Origin of Species. During this period, which has been called "the eclipse of Darwinism", scientists proposed various alternative evolutionary mechanisms which eventually proved untenable. Ronald Fisher, an English statistician finally united Mendelian genetics with natural selection between 1918 and his 1930 book The Genetical Theory of Natural Selection, giving the theory a mathematical footing and bringing broad scientific consensus that it was the basic mechanism of evolution, founding the basis for population genetics and the modern evolutionary synthesis, with J.B.S. Haldane and Sewall Wright, which set the frame of reference for modern debates and refinements of the theory.

text: Darwin had convinced most scientists that

text: evolution

text: as

text: descent with modification

text: was correct, and he was regarded as a great scientist who had revolutionised ideas. Though few agreed with his view that "natural selection has been the main but not the exclusive means of modification", he was honoured in June 1909 by more than 400 officials and scientists from across the world who met in

text: Cambridge

text: to

text: commemorate his centenary

text: and the fiftieth anniversary of

text: On the Origin of Species

text: . During this period, which has been called "

text: the eclipse of Darwinism

text: ", scientists proposed various alternative evolutionary mechanisms which eventually proved untenable.

text: Ronald Fisher

text: , an English

text: statistician

text: finally united

text: Mendelian genetics

text: with

text: natural selection

text: between 1918 and his 1930 book

text: , giving the theory a

text: mathematical

text: footing and bringing broad scientific consensus that it was the basic mechanism of evolution, founding the basis for

text: population genetics

text: and the

text: modern evolutionary synthesis

text: , with

text: J.B.S. Haldane

text: and

text: Sewall Wright

text: , which set the frame of reference for modern debates and refinements of the theory.

text: Examining patterns of Linkage Disequilibrium (LD) can locate signatures of adaptive evolution (Hawks et al. 2007, Voight et al. 2006). LD tests work on the basic principle that, assuming equal recombination rates, LD will rise with increasing natural selection. These genomic methods can also be applied to search for adaptive evolution in non-coding DNA, where putatively neutral sites are hard to identify (Ponting and Lunter 2006).

text: It is widely regarded as unprofessional for a biologist to say something like "A wing is for flying," although that is their normal function. A biologist would be conscious that sometime in the remote past feathers on a small dinosaur had the function of retaining heat, and that later many wings were not used for flying (e.g. penguins, ostriches). So, the biologist would rather say that the wings on a bird or an insect usually had the function of aiding flight. That would carry the connotation of being an adaptation with a history of evolution by natural selection.

text: It is widely regarded as unprofessional for a biologist to say something like "A wing is for flying," although that is their normal function. A biologist would be conscious that sometime in the remote past feathers on a small dinosaur had the function of retaining heat, and that later many wings were not used for flying (e.g. penguins,

text: ostrich

text: es). So, the biologist would rather say that the wings on a bird or an insect usually had the function of aiding flight. That would carry the connotation of being an adaptation with a history of evolution by natural selection.

text: Moreover, any system must be adapted to its environment, which implies that it is able to react adequately to changes in that environment. This is the origin of mind or intelligence, as the system should be able to select the right actions for the given conditions. These "condition-action" relations are the basis of knowledge. As systems evolve, their adaptiveness tends to increase, and therefore also their knowledge or intelligence. Thus, the general trend of evolution is self-organization, or a spontaneous increase in intelligent organization.

text: Moreover, any system must be adapted to its environment, which implies that it is able to react adequately to changes in that environment. This is the origin of mind or intelligence, as the system should be able to select the right actions for the given conditions. These "condition-action" relations are the basis of knowledge. As systems evolve, their adaptiveness tends to increase, and therefore also their knowledge or intelligence. Thus, the general trend of evolution is

text: self-organization

text: , or a spontaneous increase in intelligent organization.

text: Leary uses the eight circuits along with recapitulation theory to explain the evolution the human species, the personal development of an individual, and the biological evolution of all life.

text: Leary uses the eight circuits along with

text: recapitulation theory

text: to explain the evolution the human species, the personal development of an individual, and the

text: biological evolution

text: of all life.

text: In fact, as the diagram illustrates schematically, genes are much more complicated and elusive concepts. A reasonable modern definition of a gene is "a locatable region of genomic sequence, corresponding to a unit of inheritance, which is associated with regulatory regions, transcribed regions and/or other functional sequence regions." One of the major complicating factors is that the exons which code for the proteins are often separated by many introns, which used to be called "junk DNA" but appear to have various as-yet-ill-understood purposes. The exons can be combined in different orders (splice variants) to produce different proteins. For example the gene called Dscam in Drosophila has 110 introns and therefore tens of thousands of possible splice variants.

text: In fact, as the diagram illustrates schematically, genes are much more complicated and elusive concepts. A reasonable modern definition of a gene is "a locatable region of genomic sequence, corresponding to a unit of inheritance, which is associated with regulatory regions, transcribed regions and/or other functional sequence regions." One of the major complicating factors is that the

text: exons

text: which

text: code

text: for the proteins are often separated by many

text: introns

text: , which used to be called "

text: junk DNA

text: " but appear to have various as-yet-ill-understood purposes. The exons can be combined in different orders (

text: splice variant

text: s) to produce different proteins. For example the gene called

text: Dscam

text: in

text: Drosophila

text: has 110 introns and therefore tens of thousands of possible splice variants.

text: Evolution of the adaptive immune system occurred in an ancestor of the jawed vertebrates. Many of the classical molecules of the adaptive immune system (e.g., immunoglobulins and T cell receptors) exist only in jawed vertebrates. However, a distinct lymphocyte-derived molecule has been discovered in primitive jawless vertebrates, such as the lamprey and hagfish. These animals possess a large array of molecules called Variable lymphocyte receptors (VLRs) that, like the antigen receptors of jawed vertebrates, are produced from only a small number (one or two) of genes. These molecules are believed to bind pathogenic antigens in a similar way to antibodies, and with the same degree of specificity.

text: Evolution of the adaptive immune system

text: occurred in an ancestor of the jawed vertebrates. Many of the classical molecules of the adaptive immune system (e.g.,

text: immunoglobulin

text: s and

text: T cell receptor

text: s) exist only in jawed vertebrates. However, a distinct

text: lymphocyte

text: -derived molecule has been discovered in primitive

text: jawless vertebrates

text: , such as the

text: lamprey

text: and

text: hagfish

text: . These animals possess a large array of molecules called

text: Variable lymphocyte receptor

text: s (VLRs) that, like the antigen receptors of jawed vertebrates, are produced from only a small number (one or two) of

text: gene

text: s. These molecules are believed to bind pathogenic

text: antigen

text: s in a similar way to antibodies, and with the same degree of specificity.

text: Behe calculates the "edge of evolution" - the point at which Darwinian evolution is no longer an efficacious agent of creative biological change - by taking into account the number of mutations required to "travel" from one genetic state to another, as well as population size for the organism in question.  He concludes that purposeful design plays a major role in the development of biological complexity, through the mechanism of producing "non-random mutations", which are then subjected to the sculpting hand of natural selection.

text: The Descent of Man, and Selection in Relation to Sex is a book by English naturalist Charles Darwin, first published in 1871, which applies evolutionary theory to human evolution, and details his theory of sexual selection, a form of biological adaptation distinct from, yet interconnected with, natural selection.  The book discusses many related issues, including evolutionary psychology, evolutionary ethics, differences between human races, differences between sexes, the dominant role of women in mate choice, and the relevance of the evolutionary theory to society.

text: The Descent of Man, and Selection in Relation to Sex is a book by English

text: naturalist

text: Charles Darwin

text: , first published in 1871, which applies

text: evolution

text: ary theory to

text: human evolution

text: , and details his theory of

text: sexual selection

text: , a form of biological adaptation distinct from, yet interconnected with, natural selection.  The book discusses many related issues, including

text: evolutionary psychology

text: evolutionary ethics

text: , differences between human

text: races

text: , differences between sexes, the dominant role of women in

text: mate choice

text: , and the relevance of the

text: evolutionary theory

text: to society.

text: Strawberry poison frog, Oophaga pumilio, shows extreme variation in color and pattern between populations that have been geographically isolated for more than 10,000 years. When populations are separated by geographic distances and landscape barriers, they frequently experience restricted gene flow, which can enable phenotypic divergence between populations through selection or drift. Their variety in warning coloration is used for their visibility, toxicity and resistance to predators. When divergent phenotypes are mostly restricted to separate islands, the biogeography of color polymorphism suggests a major role for neutral process. However, Summers et al. (1997) provide evidences that neutral divergence alone is unlikely to have caused the variation in color patterns. As shown by Lande, rapid evolution in sexually selecting species is led by the interaction of random genetic drift with natural and sexual selection such as random genetic drift in female mating preferences. Color is known to play a role in male–female signaling, mate attraction, and male–male signaling in anurans. Based on Tazzyman and Iwasa’s study that involved collections of samples from main islands in the Bocas del Toro archipelago, its results proved that female preference on male calls led to call divergence and therefore divergence was driven by sexual selection. Mate choice plays a critical role in generating and maintaining biodiversity. Furthermore, spatial variation in predators or habitat features could exert divergent natural selection on coloration in response to its subjection to predator selection. It is still unclear to what extent sexual selection has driven the evolution of color morphs rather than reinforcing the reproductive isolation of morphs. In an aposematic organism such as Oophaga pumilio, we cannot attribute a phylogenetic signal of selection to female mate choice alone but  is quite possible that genetic drift would interact with female color preferences to trigger divergence

text: Strawberry poison frog, Oophaga pumilio, shows extreme variation in color and pattern between populations that have been geographically isolated for more than 10,000 years. When populations are separated by geographic distances and landscape barriers, they frequently experience restricted gene flow, which can enable phenotypic divergence between populations through selection or drift. Their variety in warning coloration is used for their visibility, toxicity and resistance to predators. When divergent phenotypes are mostly restricted to separate islands, the biogeography of color polymorphism suggests a major role for neutral process. However, Summers et al. (1997) provide evidences that neutral divergence alone is unlikely to have caused the variation in color patterns. As shown by

text: Lande

text: , rapid evolution in sexually selecting species is led by the interaction of random genetic drift with natural and sexual selection such as random genetic drift in female mating preferences. Color is known to play a role in male–female signaling, mate attraction, and male–male signaling in anurans. Based on Tazzyman and Iwasa’s study that involved collections of samples from main islands in the Bocas del Toro archipelago, its results proved that female preference on male calls led to call divergence and therefore divergence was driven by sexual selection. Mate choice plays a critical role in generating and maintaining biodiversity. Furthermore, spatial variation in predators or habitat features could exert divergent natural selection on coloration in response to its subjection to predator selection. It is still unclear to what extent sexual selection has driven the evolution of color morphs rather than reinforcing the reproductive isolation of morphs. In an

text: aposematic

text: organism such as Oophaga pumilio, we cannot attribute a phylogenetic signal of selection to female mate choice alone but  is quite possible that genetic drift would interact with female color preferences to trigger divergence

text: Darwinism: An Exposition of the Theory of Natural Selection with Some of Its Applications is an 1889 book on biological evolution by Alfred Russel Wallace, the co-discoverer of evolution by natural selection together with Charles Darwin.  This was a book Wallace wrote as a defensive response to the scientific critics of natural selection. Of all Wallace's books, it is cited by scholarly publications the most.

text: Darwinism: An Exposition of the Theory of Natural Selection with Some of Its Applications is an 1889 book on

text: biological evolution

text: by

text: Alfred Russel Wallace

text: , the co-discoverer of evolution by

text: natural selection

text: together with

text: Charles Darwin

text: .  This was a book Wallace wrote as a defensive response to the scientific critics of natural selection. Of all Wallace's books, it is cited by scholarly publications the most.

text: The similarity in all these unrelated families involves convergent evolution of the saber-like canines as a hunting adaptation.  Meehan et al. note that it took around 8 million years for a new type of saber-toothed cat to fill the niche of an extinct predecessor in a similar ecological role; this has happened at least four times with different families of animals developing this adaptation. Although the adaptation of the saber-like canines made these creatures successful, it seems that the shift to obligate carnivorism, along with co-evolution with large prey animals, led the saber-toothed cats of each time period to extinction. As per Van Valkenburgh, the adaptations that made saber-toothed cats successful also made the creatures vulnerable to extinction. In her example, trends toward an increase in size, along with greater specialization, acted as a "macro-evolutionary ratchet": when large prey became scarce or extinct, these creatures would be unable to adapt to smaller prey or consume other sources of food, and would be unable to reduce their size so as to need less food.

text: The similarity in all these unrelated families involves convergent evolution of the saber-like canines as a hunting adaptation.  Meehan et al. note that it took around 8 million years for a new type of saber-toothed cat to fill the niche of an extinct predecessor in a similar ecological role; this has happened at least four times with different families of animals developing this adaptation. Although the adaptation of the saber-like canines made these creatures successful, it seems that the shift to

text: obligate carnivorism

text: , along with

text: co-evolution

text: with large prey animals, led the saber-toothed cats of each time period to extinction. As per Van Valkenburgh, the adaptations that made saber-toothed cats successful also made the creatures vulnerable to extinction. In her example, trends toward an increase in size, along with greater specialization, acted as a "macro-evolutionary ratchet": when large prey became scarce or extinct, these creatures would be unable to adapt to smaller prey or consume other sources of food, and would be unable to reduce their size so as to need less food.

text: In the second half of the century the ideas of population genetics began to be applied in the new discipline of the genetics of behavior, sociobiology, and, especially in humans, evolutionary psychology. In the 1960s W.D. Hamilton and others developed game theory approaches to explain altruism from an evolutionary perspective through kin selection. The possible origin of higher organisms through endosymbiosis, and contrasting approaches to molecular evolution in the gene-centered view (which held selection as the predominant cause of evolution) and the neutral theory (which made genetic drift a key factor) spawned perennial debates over the proper balance of adaptationism and contingency in evolutionary theory.

text: In the second half of the century the ideas of population genetics began to be applied in the new discipline of the genetics of behavior,

text: sociobiology

text: , and, especially in humans,

text: evolutionary psychology

text: . In the 1960s

text: W.D. Hamilton

text: and others developed

text: game theory

text: approaches to explain

text: altruism

text: from an evolutionary perspective through

text: kin selection

text: . The possible origin of higher organisms through

text: endosymbiosis

text: , and contrasting approaches to molecular evolution in the

text: gene-centered view

text: (which held selection as the predominant cause of evolution) and the

text: neutral theory

text: (which made

text: genetic drift

text: a key factor) spawned perennial debates over the proper balance of

text: adaptationism

text: and contingency in evolutionary theory.

text: For example, Darwinian evolution points to the competition of a variety of species, each with heterogeneous members, within a given environment. This leads to changing species and even to new species arising. A dialectical biologist fully accepts this picture then looks for ways in which the competing creatures (which serve as the internal conflicts in the environment) lead to changes. The changes manifest in the creatures themselves, through the creatures embracing biological adaptations that provide them with advantages, and in the environment itself, as when the action of microbes encourages the erosion of rocks. Further, each species is part of the "environment" of all the others.

text: For example,

text: Darwinian evolution

text: points to the competition of a variety of species, each with heterogeneous members, within a given environment. This leads to changing species and even to new species arising. A dialectical biologist fully accepts this picture then looks for ways in which the competing creatures (which serve as the internal conflicts in the environment) lead to changes. The changes manifest in the creatures themselves, through the creatures embracing biological adaptations that provide them with advantages, and in the environment itself, as when the action of microbes encourages the erosion of rocks. Further, each species is part of the "environment" of all the others.

text: While the stocks of those species that survived are in many cases recovering, the lake ecosystem has changed irrevocably. The entire trophic web has been observed to be upset. But still, evolution runs its course: those Lake Victoria Haplochromis species that still exist are in many cases adapting to new food sources, and in time, speciation is likely to set in and produce a new adaptive radiation of these fishes. Until then, however, the ecological balance of the lake is still on the brink, and many of the cichlids that survived the peak population of Lates are still critically endangered and close to extinction.

text: While the stocks of those species that survived are in many cases recovering, the lake ecosystem has changed irrevocably. The entire

text: trophic web

text: has been observed to be upset. But still,

text: evolution

text: runs its course: those Lake Victoria Haplochromis species that still exist are in many cases

text: adapting

text: to new food sources, and in time,

text: speciation

text: is likely to set in and produce a new

text: adaptive radiation

text: of these fishes. Until then, however, the ecological balance of the lake is still on the brink, and many of the cichlids that survived the peak population of Lates are still

text: critically endangered

text: and close to extinction.

text: Pascoe accepted the fact of evolution but was an opponent to natural selection.

text: Pascoe accepted the fact of

text: evolution

text: but was an opponent to

text: natural selection

text: These outcomes of evolution are distinguished based on time scale as macroevolution versus microevolution. Macroevolution refers to evolution that occurs at or above the level of species, in particular speciation and extinction; whereas microevolution refers to smaller evolutionary changes within a species or population, in particular shifts in gene frequency and adaptation. In general, macroevolution is regarded as the outcome of long periods of microevolution. Thus, the distinction between micro- and macroevolution is not a fundamental one—the difference is simply the time involved. However, in macroevolution, the traits of the entire species may be important. For instance, a large amount of variation among individuals allows a species to rapidly adapt to new habitats, lessening the chance of it going extinct, while a wide geographic range increases the chance of speciation, by making it more likely that part of the population will become isolated. In this sense, microevolution and macroevolution might involve selection at different levels—with microevolution acting on genes and organisms, versus macroevolutionary processes such as species selection acting on entire species and affecting their rates of speciation and extinction.

text: These outcomes of evolution are distinguished based on time scale as

text: macroevolution

text: versus microevolution. Macroevolution refers to evolution that occurs at or above the level of species, in particular speciation and extinction; whereas microevolution refers to smaller evolutionary changes within a species or population, in particular shifts in

text: gene frequency

text: and adaptation. In general, macroevolution is regarded as the outcome of long periods of microevolution. Thus, the distinction between micro- and macroevolution is not a fundamental one—the difference is simply the time involved. However, in macroevolution, the traits of the entire species may be important. For instance, a large amount of variation among individuals allows a species to rapidly adapt to new

text: habitat

text: s, lessening the chance of it going extinct, while a wide geographic range increases the chance of speciation, by making it more likely that part of the population will become isolated. In this sense, microevolution and macroevolution might involve selection at different levels—with microevolution acting on genes and organisms, versus macroevolutionary processes such as

entity_section: Species selection and selection at higher taxonomic levels

text: species selection

text: acting on entire species and affecting their rates of speciation and extinction.

text: In 1987, PCR-amplification of mtDNA was first used to determine sequences. In 1991 Vigilante et al. published the seminal work on mtDNA phylogeny implicating sub-saharan Africa as the place of humans most recent common ancestors for all mtDNAs. The war between out-of-Africa and multiregionalism, already simmering with the critiques of Allan Templeton, soon escalated with the paleoanthropologist, like Milford Wolpoff, getting involved.In 1995, F. Ayala published his critical Science article "The Myth about Eve", which relied on HLA-DR sequence. At the time, however Ayala was not aware of rapid evolution of HLA loci via recombinatory process. In 1996, Parham and Ohta published their finds on the rapid evolution of HLA by short-distance recombination ('gene conversion' or 'abortive recombination'), weakening Ayala's claim (Parham had actually written a review a year earlier, but this had gone unnoticed).  A stream of papers would follow from both sides, many with highly flawed methods and sampling. One of the more interesting was Harris and Hey, 1998 which showed that the TMCRA (time to most recent common ancestor) for the PDHA1 gene was well in excess of 1 million years. Given a ploidy at this locus of 1.5 (3 fold higher than mtDNA) the TMRCA was more than double the expectation. While this falls into the 'fixation curve' of 1.5 ploidy (averaging 2 female and 1 male) the suggested age of 1.8 my is close a significantly deviant p-value for the population size, possibly indicating that the human population shrank or split off of another population.  Oddly, the next X-linked loci they examined, Factor  IX, showed a TMRCA of less than 300,000 years.

text: In 1987, PCR-amplification of mtDNA was first used to determine sequences. In 1991 Vigilante et al. published the seminal work on mtDNA phylogeny implicating sub-saharan Africa as the place of humans most recent common ancestors for all mtDNAs. The war between out-of-Africa and multiregionalism, already simmering with the critiques of Allan Templeton, soon escalated with the paleoanthropologist, like Milford Wolpoff, getting involved.In 1995, F. Ayala published his critical Science article "The Myth about Eve", which relied on

text: HLA-DR

text: sequence. At the time, however Ayala was not aware of rapid evolution of HLA loci via recombinatory process. In 1996, Parham and Ohta published their finds on the rapid evolution of HLA by short-distance recombination ('gene conversion' or 'abortive recombination'), weakening Ayala's claim (Parham had actually written a review a year earlier, but this had gone unnoticed).  A stream of papers would follow from both sides, many with highly flawed methods and sampling. One of the more interesting was Harris and Hey, 1998 which showed that the TMCRA (time to most recent common ancestor) for the PDHA1 gene was well in excess of 1 million years. Given a

text: ploidy

text: at this locus of 1.5 (3 fold higher than mtDNA) the TMRCA was more than double the expectation. While this falls into the 'fixation curve' of 1.5 ploidy (averaging 2 female and 1 male) the suggested age of 1.8 my is close a significantly deviant p-value for the population size, possibly indicating that the human population shrank or split off of another population.  Oddly, the next X-linked loci they examined, Factor  IX, showed a TMRCA of less than 300,000 years.

text: Polymorphism was crucial to research in ecological genetics by E. B. Ford and his co-workers from the mid-1920s to the 1970s (similar work continues today, especially on mimicry). The results had a considerable effect on the mid-century evolutionary synthesis, and on present evolutionary theory. The work started at a time when natural selection was largely discounted as the leading mechanism for evolution, continued through the middle period when Sewall Wright's ideas on drift were prominent, to the last quarter of the 20th century when ideas such as Kimura's neutral theory of molecular evolution was given much attention. The significance of the work on ecological genetics is that it has shown how important selection is in the evolution of natural populations, and that selection is a much stronger force than was envisaged even by those population geneticists who believed in its importance, such as Haldane and Fisher.

text: Polymorphism was crucial to research in

text: ecological genetics

text: by E. B. Ford and his co-workers from the mid-1920s to the 1970s (similar work continues today, especially on

text: mimicry

text: ). The results had a considerable effect on the mid-century

text: evolutionary synthesis

text: , and on present

text: evolutionary theory

text: . The work started at a time when

text: natural selection

text: was largely discounted as the leading mechanism for evolution, continued through the middle period when

text: Sewall Wright

text: 's ideas on

text: drift

text: were prominent, to the last quarter of the 20th century when ideas such as

text: Kimura

text: 's

text: neutral theory of molecular evolution

text: was given much attention. The significance of the work on ecological genetics is that it has shown how important selection is in the evolution of natural populations, and that selection is a much stronger force than was envisaged even by those population geneticists who believed in its importance, such as

text: Haldane

text: and

text: Fisher

text: The adaptive market hypothesis, as proposed by Andrew Lo, is an attempt to reconcile economic theories based on the efficient market hypothesis (which implies that markets are efficient) with behavioral economics, by applying the principles of evolution to financial interactions: competition, adaptation and natural selection.

text: The adaptive market hypothesis, as proposed by

text: Andrew Lo

text: , is an attempt to reconcile economic theories based on the

text: efficient market hypothesis

text: (which implies that

text: market

text: s are

text: efficient

text: ) with

text: behavioral economics

text: , by applying the principles of

text: evolution

text: to financial interactions:

text: competition

text: adaptation

text: and

text: natural selection

text: The rate of evolution is a variable of considerable interest in evolutionary biology. It concerns the limits of adaptation to natural environments as well as the limits of artificial selection.

text: The rate of evolution is a variable of considerable interest in

text: evolutionary biology

text: . It concerns the limits of adaptation to natural environments as well as the limits of artificial selection.

text: For example, the National Association of Biology Teachers in a statement endorsing evolution as noncontroversial quoted Theodosius Dobzhansky "nothing in biology makes sense except in the light of evolution" and went on to state that the quote "accurately reflects the central, unifying role of evolution in biology. The theory of evolution provides a framework that explains both the history of life and the ongoing adaptation of organisms to environmental challenges and changes." They emphasized that "Scientists have firmly established evolution as an important natural process" and that "The selection of topics covered in a biology curriculum should accurately reflect the principles of biological science. Teaching biology in an effective and scientifically honest manner requires that evolution be taught in a standards-based instructional framework with effective classroom discussions and laboratory experiences."

text: For example, the

entity_name: National Association of Biology Teachers

text: National Association of Biology Teachers

text: in a statement endorsing evolution as noncontroversial quoted

text: Theodosius Dobzhansky

entity_name: Nothing in Biology Makes Sense Except in the Light of Evolution

text: nothing in biology makes sense except in the light of evolution

text: " and went on to state that the quote "accurately reflects the central, unifying role of evolution in biology. The theory of evolution provides a framework that explains both the history of life and the ongoing adaptation of organisms to environmental challenges and changes." They emphasized that "Scientists have firmly established evolution as an important natural process" and that "The selection of topics covered in a biology curriculum should accurately reflect the principles of biological science. Teaching biology in an effective and scientifically honest manner requires that evolution be taught in a standards-based instructional framework with effective classroom discussions and laboratory experiences."

text: One of the original aspects of the noosphere concept deals with evolution. Henri Bergson, with his L'évolution créatrice  (1907), was one of the first to propose evolution is "creative" and cannot necessarily be explained solely by Darwinian natural selection. L'évolution créatrice is upheld, according to Bergson, by a constant vital force which animates life and fundamentally connects mind and body, an idea opposing the dualism of René Descartes. In 1923, C. Lloyd Morgan took this work further, elaborating on an "emergent evolution" which could explain increasing complexity (including the evolution of mind). Morgan found many of the most interesting changes in living things have been largely discontinuous with past evolution. Therefore, these living things did not necessarily evolve through a gradual process of natural selection. Rather, he posited, the process of evolution experiences jumps in complexity (such as the emergence of a self-reflective universe, or noosphere). Finally, the complexification of human cultures, particularly language, facilitated a quickening of evolution in which cultural evolution occurs more rapidly than biological evolution. Recent understanding of human ecosystems and of human impact on the biosphere have led to a link between the notion of sustainability with the "co-evolution" and harmonization of cultural and biological evolution.

text: One of the original aspects of the noosphere concept deals with

text: evolution

text: Henri Bergson

text: , with his

text: L'évolution créatrice

text: (1907), was one of the first to propose evolution is "creative" and cannot necessarily be explained solely by Darwinian

text: natural selection

text: . L'évolution créatrice is upheld, according to Bergson, by a constant

text: vital force

text: which animates life and fundamentally connects mind and body, an idea opposing the dualism of

text: René Descartes

text: . In 1923,

text: C. Lloyd Morgan

text: took this work further, elaborating on an "

text: emergent evolution

text: " which could explain increasing

text: complexity

text: (including the evolution of mind). Morgan found many of the most interesting changes in living things have been largely discontinuous with past evolution. Therefore, these living things did not necessarily evolve through a gradual process of natural selection. Rather, he posited, the process of evolution experiences jumps in complexity (such as the emergence of a self-reflective universe, or noosphere). Finally, the complexification of human cultures, particularly language, facilitated a quickening of evolution in which cultural evolution occurs more rapidly than biological evolution. Recent understanding of human ecosystems and of human impact on the biosphere have led to a link between the notion of

text: sustainability

text: with the "co-evolution" and harmonization of cultural and biological evolution.

text: The Out-of-Africa model proposes that modern humans originated in Africa and migrated north to populate Europe and Asia. These migrants most likely had a functional MC1R variant and, accordingly, dark hair and skin as displayed by indigenous Africans today. As humans migrated north, the absence of high levels of solar radiation in northern Europe and Asia relaxed the selective pressure on active MC1R, allowing the gene to mutate into dysfunctional variants without reproductive penalty, then propagate by genetic drift. Studies show the MC1R Arg163Gln allele has a high frequency in East Asia and may be part of the evolution of light skin in East Asian populations. No evidence is known for positive selection of MC1R alleles in Europe and there is no evidence of an association between MC1R and the evolution of light skin in European populations.

text: The

text: Out-of-Africa

text: model proposes that modern humans originated in Africa and migrated north to populate Europe and Asia. These migrants most likely had a functional MC1R variant and, accordingly, dark hair and skin as displayed by indigenous Africans today. As humans migrated north, the absence of high levels of

text: solar radiation

text: in northern Europe and Asia relaxed the

text: selective pressure

text: on active MC1R, allowing the gene to mutate into dysfunctional variants without reproductive penalty, then propagate by

text: genetic drift

text: . Studies show the MC1R Arg163Gln allele has a high frequency in East Asia and may be part of the evolution of light skin in East Asian populations. No evidence is known for

text: positive selection

text: of MC1R alleles in Europe and there is no evidence of an association between MC1R and the evolution of light skin in European populations.

text: 'The assumption that only the genes evolved reflects a widespread misconception about the way natural selection acts. Genes are the so-called units of selection, which are inherited, selected, or eliminated, and so they are indeed something that evolves. But every time one gene is selected over another, one design for a developmental program is selected over another as well; by virtue of its structure, this developmental program interacts with some aspects of the environment rather than others, rendering certain environmental features causally relevant to development. So, step by step, as natural selection constructs the species' gene set (chosen from the available mutations), it constructs in tandem the species' developmentally relevant environment (selected from the set of all properties of the world). Thus, both the genes and the developmentally relevant environment are the product of evolution (p. 84).

text: A Mathematical Theory of Natural and Artificial Selection is the title of a series of scientific papers by the British population geneticist J.B.S. Haldane, published between 1924 and 1934.  Haldane outlines the first mathematical models for many cases of evolution due to selection, an important concept in the modern evolutionary synthesis.

text: A Mathematical Theory of Natural and Artificial Selection is the title of a series of

text: scientific paper

text: s by the

text: British

text: population geneticist

text: J.B.S. Haldane

text: , published between 1924 and 1934.  Haldane outlines the first mathematical models for many cases of

text: evolution

text: due to

text: selection

text: , an important concept in the

text: modern evolutionary synthesis

text: Evolutionary psychologists consider Charles Darwin's theory of natural selection to be important to an understanding of psychology. Natural selection occurs because individual organisms who are genetically better suited to the current environment leave more descendants, and their genes spread through the population, thus explaining why organisms fit their environments so closely. This process is slow and cumulative, with new traits layered over older traits. The advantages created by natural selection are known as adaptations. Evolutionary psychologists say that animals, just as they evolve physical adaptations, evolve psychological adaptations.

text: Natural selection is observed in contemporary human populations, with recent findings demonstrating that the population at risk of the severe debilitating disease kuru has significant over-representation of an immune variant of the prion protein gene G127V versus non-immune alleles. Scientists postulate one of the reasons for the rapid selection of this genetic variant is the lethality of the disease in non-immune persons. Other reported evolutionary trends in other populations include a lengthening of the reproductive period, reduction in cholesterol levels, blood glucose and blood pressure.

text: Natural selection is observed in contemporary human populations, with recent findings demonstrating that the population at risk of the severe debilitating disease

text: kuru

text: has significant over-representation of an immune variant of the

text: prion protein

text: gene G127V versus non-immune alleles. Scientists postulate one of the reasons for the rapid selection of this

text: genetic variant

text: is the lethality of the disease in non-immune persons. Other reported evolutionary trends in other populations include a lengthening of the reproductive period, reduction in cholesterol levels, blood glucose and blood pressure.

text: The evolution of interactome complexity is delineated in a study published in Nature. In this study it is first noted that the boundaries between prokaryotes, unicellular eukaryotes and multicellular eukaryotes are accompanied by orders-of-magnitude reductions in effective population size, with concurrent amplifications of the effects of random genetic drift. The resultant decline in the efficiency of selection seems to be sufficient to influence a wide range of attributes at the genomic level in a nonadaptive manner. The Nature study shows that the variation in the power of random genetic drift is also capable of influencing phylogenetic diversity at the subcellular and cellular levels. Thus, population size would have to be considered as a potential determinant of the mechanistic pathways underlying long-term phenotypic evolution. In the study it is further shown that a phylogenetically broad inverse relation exists between the power of drift and the structural integrity of protein subunits. Thus, the accumulation of mildly deleterious mutations in populations of small size induces secondary selection for protein–protein interactions that stabilize key gene functions, mitigating the structural degradation promoted by inefficient selection. By this means, the complex protein architectures and interactions essential to the genesis of phenotypic diversity may initially emerge by non-adaptive mechanisms.

text: The evolution of interactome complexity is delineated in a study published in Nature. In this study it is first noted that the boundaries between

text: prokaryotes

text: , unicellular

text: eukaryotes

text: and multicellular eukaryotes are accompanied by orders-of-magnitude reductions in effective population size, with concurrent amplifications of the effects of

text: random genetic drift

text: . The resultant decline in the efficiency of selection seems to be sufficient to influence a wide range of attributes at the genomic level in a nonadaptive manner. The Nature study shows that the variation in the power of random genetic drift is also capable of influencing phylogenetic diversity at the subcellular and cellular levels. Thus, population size would have to be considered as a potential determinant of the mechanistic pathways underlying long-term phenotypic evolution. In the study it is further shown that a phylogenetically broad inverse relation exists between the power of drift and the structural integrity of

text: protein

text: subunits. Thus, the accumulation of mildly deleterious mutations in populations of small size induces secondary selection for

text: protein–protein interactions

text: that stabilize key gene functions, mitigating the structural degradation promoted by inefficient selection. By this means, the complex protein architectures and interactions essential to the genesis of phenotypic diversity may initially emerge by non-adaptive mechanisms.

text: There is an alternate strategy for identifying fit mates which does not rely on one gender having exaggerated sexual ornaments or other handicaps, but is generally applicable to most, if not all sexual creatures. It derives from the concept that the change in appearance and functionality caused by a non-silent mutation will generally stand out in a population. This is because that altered appearance and functionality will be unusual, peculiar, and different from the norm within that population. The norm against which these unusual features are judged is made up of fit attributes that have attained their plurality through natural selection, while less adaptive attributes will be in the minority or frankly rare.  Since the overwhelming majority of mutant features are maladaptive, and it is impossible to predict evolution's future direction, sexual creatures would be expected to prefer mates with the least number of unusual or minority features. This will have the effect of a sexual population rapidly shedding peripheral phenotypic features and canalizing the entire outward appearance and behaviour so that all the members of that population will begin to look remarkably similar in every detail, as illustrated in the accompanying photograph of  the African pygmy kingfisher, Ispidina picta. Once a population has become as homogeneous in appearance as is typical of most species, its entire repertoire of behaviours will also be rendered evolutionarily stable, including any altruistic, cooperative and social characteristics. Thus, in the example of the selfish individual who hangs back from the rest of the hunting pack, but who nevertheless joins in the spoils, that individual will be recognized as being different from the norm, and will therefore find it difficult to attract a mate. Its genes will therefore have only a very small probability of being passed on to the next generation, thus evolutionarily stabilizing cooperation and social interactions at whatever level of complexity is the norm in that population.

text: There is an alternate strategy for identifying fit mates which does not rely on one gender having exaggerated sexual ornaments or other handicaps, but is generally applicable to most, if not all sexual creatures. It derives from the concept that the change in appearance and functionality caused by a

text: non-silent

text: mutation

text: natural selection

text: , while less adaptive attributes will be in the minority or frankly rare.  Since the overwhelming majority of mutant features are maladaptive, and it is impossible to predict evolution's future direction, sexual creatures would be expected to

text: . This will have the effect of a sexual population rapidly shedding peripheral phenotypic features and canalizing the entire outward appearance and behaviour so that all the members of that population will begin to look remarkably similar in every detail, as illustrated in the accompanying photograph of  the

text: African pygmy kingfisher

text: , Ispidina picta. Once a population has become as homogeneous in appearance as is typical of most species, its entire repertoire of behaviours will also be rendered

text: evolutionarily stable

text: , including any altruistic, cooperative and social characteristics. Thus, in the example of the selfish individual who hangs back from the rest of the hunting pack, but who nevertheless joins in the spoils, that individual will be recognized as being different from the norm, and will therefore find it difficult to attract a mate. Its genes will therefore have only a very small probability of being passed on to the next generation, thus evolutionarily stabilizing cooperation and social interactions at whatever level of complexity is the norm in that population.

text: Agnostic evolution is the position of acceptance of biological evolution, combined with the belief that it is not important whether God is, was, or will have been involved.

text: His academic research focuses on the evolution of gene regulation.

text: His academic research focuses on the

text: evolution

text: of

text: gene regulation

text: His book The Origins of Adaptations (1963) was an early text that discussed the main themes of the modern evolutionary synthesis such as genetic drift, modes of speciation, natural selection  and population genetics. However, Grant did not describe these mechanisms of evolution as "Neo-Darwinism" or the synthetic theory, instead he referred to these mechanisms as the "causal theory." The book was awarded the 1964 Phi Beta Kappa Award in Science.

text: His book The Origins of Adaptations (1963) was an early text that discussed the main themes of the

text: modern evolutionary synthesis

text: such as

text: genetic drift

text: modes of speciation

text: natural selection

text: and

text: population genetics

text: . However, Grant did not describe these mechanisms of evolution as "

text: Neo-Darwinism

text: " or the synthetic theory, instead he referred to these mechanisms as the "causal theory." The book was awarded the 1964

text: Phi Beta Kappa Award in Science

text: What exactly constitutes an "evolutionary landscape" is confused in the literature. The term 'evolutionary landscape' is often used interchangeably with 'adaptive landscape' and 'fitness landscape', although some authors distinguish between these terms. As discussed below, different authors have different definitions of adaptive and fitness landscapes. Additionally, there is a large disagreement whether evolutionary landscape should be used as a visual metaphor disconnected from the underlying math, a tool for evaluating models of evolution, or a model in and of itself used to generate hypotheses and predictions. Clearly, the field of biology, specifically evolutionary biology and population genetics, needs to come to a consensus about what an evolutionary landscape is and how it should be used.

text: Evolution influences every aspect of the form and behaviour of organisms. Most prominent are the specific behavioural and physical adaptations that are the outcome of natural selection. These adaptations increase fitness by aiding activities such as finding food, avoiding predators or attracting mates. Organisms can also respond to selection by cooperating with each other, usually by aiding their relatives or engaging in mutually beneficial symbiosis. In the longer term, evolution produces new species through splitting ancestral populations of organisms into new groups that cannot or will not interbreed.

text: Evolution influences every aspect of the form and behaviour of organisms. Most prominent are the specific behavioural and physical

text: adaptation

text: s that are the outcome of natural selection. These adaptations increase fitness by aiding activities such as finding food, avoiding

text: predator

text: s or attracting mates. Organisms can also respond to selection by

text: cooperating

text: with each other, usually by aiding their relatives or engaging in mutually beneficial

text: symbiosis

text: . In the longer term, evolution produces new species through splitting ancestral populations of organisms into new groups that cannot or will not interbreed.

text: Evolution has caused sexually selected adaptations to occur in penis and testis size in order to maximise reproductive success and minimise sperm competition.

text: Evolution has caused sexually selected adaptations to occur in

text: penis

text: and testis size in order to maximise

text: reproductive success

text: and minimise

text: sperm competition

text: The ancestral condition of the anterior dentition on the lower jaw, based on Eocene primate fossils, suggests that earliest primates had lacked a differentiated toothcomb. Most fossil strepsirrhines lacked the stereotypic lemuriform toothcomb.  Collectively, early strepsirrhine primates are known as adapiforms. Adapiforms are considered to be a paraphyletic group (containing many but not all of the descendants of the last common ancestor of the group's members) because the lemuriforms are assumed to have evolved from one of several groups of adapiforms. In terms of ecology, the evolution of the toothcomb is assumed to have required a folivorous (leaf-eating) diet among the ancestral adapiform population, since that would select for reduced incisors, which would serve as an exaptation (a trait with adaptive value for something other than what it was originally selected for), which could then be used for personal or social grooming. However, the inclusion of the canines into the toothcomb must have required exceptional conditions, since large lemuriforms have secondarily modified caniniform premolars to substitute for the loss.

text: The ancestral condition of the anterior dentition on the lower jaw, based on Eocene primate fossils, suggests that earliest primates had lacked a differentiated toothcomb. Most fossil strepsirrhines lacked the stereotypic lemuriform toothcomb.  Collectively, early strepsirrhine primates are known as

text: adapiforms

text: . Adapiforms are considered to be a

text: paraphyletic

text: group (containing many but not all of the descendants of the last common ancestor of the group's members) because the lemuriforms are assumed to have evolved from one of several groups of adapiforms. In terms of ecology, the evolution of the toothcomb is assumed to have required a

text: folivorous

text: (leaf-eating) diet among the ancestral adapiform population, since that would select for reduced incisors, which would serve as an

text: exaptation

text: (a trait with adaptive value for something other than what it was originally selected for), which could then be used for personal or

text: social grooming

text: . However, the inclusion of the canines into the toothcomb must have required exceptional conditions, since large lemuriforms have secondarily modified caniniform premolars to substitute for the loss.

text: Several alternative hypotheses to natural selection as the driving force of evolution were proposed during the 1920s and 1930s.  Random mutation, migration or genetic drift were also seen as major forces of evolution. However, the magnitude of the changes observed can only be accounted for by natural selection. It can be seen from population genetics that a non-differential change will not cause evolution. If the allele frequencies are denoted by the algebraic terms p and q, and (say) p = 0.6 and q = 0.4, then a non-differential reduction in population size from say 2000 to 100 individuals, will still produce the same values of (approximately) p = 0.6 and q = 0.4.

text: Several alternative hypotheses to natural selection as the driving force of evolution were proposed during the 1920s and 1930s.

text: Random mutation

text: migration

text: or

text: genetic drift

text: were also seen as major forces of evolution. However, the magnitude of the changes observed can only be accounted for by natural selection. It can be seen from

text: population genetics

text: that a non-differential change will not cause evolution. If the allele frequencies are denoted by the

text: algebra

text: ic terms p and q, and (say) p = 0.6 and q = 0.4, then a non-differential reduction in population size from say 2000 to 100 individuals, will still produce the same values of (approximately) p = 0.6 and q = 0.4.

text: The early years of the twentieth century and particularly the First World War saw biological concepts and metaphors shunned by most social sciences. Even uttering the word evolution carried [Ref] [Ref] a "serious risk to one’s intellectual reputation". Darwinian ideas were also in decline following the rediscovery of Mendelian genetics but were revived, especially by Fisher, Haldane and Wright who developed the first population genetic models and as it became known the modern synthesis.Cultural evolutionary concepts, or even metaphors, revived more slowly. If there was one influential individual in the revival it was probably Donald T Campbell. In 1960 he drew on Wright to draw a parallel between genetic evolution and the "blind variation and selective retention" of creative ideas; work that was developed into a full theory of "socio-cultural evolution" in 1965 (a work that includes references to other works in the then current revival of interest in the field. Campbell (1965 26) was clear that he understood cultural evolution not as an analogy "from organic evolution per se, but rather from a general model for quasiteleological processes for which organic evolution is but one instance".

text: The early years of the twentieth century and particularly the First World War saw biological concepts and metaphors shunned by most social sciences. Even uttering the word evolution carried [Ref] [Ref] a "serious risk to one’s intellectual reputation". Darwinian ideas were also in decline following the rediscovery of Mendelian genetics but were revived, especially by

text: Fisher

text: Haldane

text: and

text: Wright

text: who developed the first population genetic models and as it became known the modern synthesis.Cultural evolutionary concepts, or even metaphors, revived more slowly. If there was one influential individual in the revival it was probably

text: Donald T Campbell

text: . In 1960 he drew on Wright to draw a parallel between genetic evolution and the "blind variation and selective retention" of creative ideas; work that was developed into a full theory of "socio-cultural evolution" in 1965 (a work that includes references to other works in the then current revival of interest in the field. Campbell (1965 26) was clear that he understood cultural evolution not as an analogy "from organic evolution per se, but rather from a general model for quasiteleological processes for which organic evolution is but one instance".

text: Among the Andeans, there are no significant associations between EPAS1 or EGLN1 and haemoglobin concentration, indicating variation in the pattern of molecular adaptation. However, EGLN1 appears to be the principal signature of evolution, as it shows evidence of positive selection in both Tibetans and Andeans. Adaptive mechanism is different among the Ethiopian highlanders. Genomic analysis of two ethnic groups, Amhara and Oromo, revealed that gene variations associated with haemoglobin difference among Tibetans or other variants at the same gene location do not influence the adaptation in Ethiopians. Instead, several genes appear to be involved in Ethiopians, including CBARA1, VAV3, ARNT2 and THRB, which are known to play a role in HIF genetic functions.

text: Among the Andeans, there are no significant associations between EPAS1 or EGLN1 and haemoglobin concentration, indicating variation in the pattern of molecular adaptation. However, EGLN1 appears to be the principal signature of evolution, as it shows evidence of positive selection in both Tibetans and Andeans. Adaptive mechanism is different among the Ethiopian highlanders. Genomic analysis of two ethnic groups,

text: Amhara

text: and

text: Oromo

text: gene location

text: do not influence the adaptation in Ethiopians. Instead, several genes appear to be involved in Ethiopians, including

text: and

text: THRB

text: , which are known to play a role in

text: HIF

text: genetic functions.

text: In addition to their native reaction, many enzymes perform side reactions. Similarly, binding proteins may spend some proportion of their time bound to off-target proteins. These reactions or interactions may be of no consequence to current fitness but under altered conditions, may provide the starting point for adaptive evolution. For example, several mutations in the antibiotic resistance gene B-lactamase introduce cefotaxime resistance but do not affect ampicillin resistance. In populations exposed only to ampicillin, such mutations may be present in a minority of members since there is not fitness cost (i.e. are within the neutral network). This represents cryptic genetic variation since if the population is newly exposed to cefotaxime, the minority members will exhibit some resistance.

text: In addition to their native reaction, many

text: enzymes

text: perform side reactions. Similarly, binding proteins may spend some proportion of their time bound to off-target proteins. These reactions or interactions may be of no consequence to current

text: fitness

text: but under altered conditions, may provide the starting point for adaptive evolution. For example, several mutations in the antibiotic resistance gene

text: B-lactamase

text: introduce

text: cefotaxime

text: resistance but do not affect

text: ampicillin

text: resistance. In populations exposed only to ampicillin, such mutations may be present in a minority of members since there is not fitness cost (i.e. are within the

text: neutral network

text: ). This represents cryptic genetic variation since if the population is newly exposed to cefotaxime, the minority members will exhibit some resistance.

text: Natural selection is one of the cornerstones of modern biology. The term was introduced by Darwin in his groundbreaking 1859 book On the Origin of Species, in which natural selection was described by analogy to artificial selection, a process by which animals and plants with traits considered desirable by human breeders are systematically favored for reproduction. The concept of natural selection was originally developed in the absence of a valid theory of heredity; at the time of Darwin's writing, nothing was known of modern genetics. The union of traditional Darwinian evolution with subsequent discoveries in classical and molecular genetics is termed the modern evolutionary synthesis. Natural selection remains the primary explanation for adaptive evolution.

text: Natural selection is one of the cornerstones of modern

text: biology

text: . The term was introduced by Darwin in his groundbreaking 1859 book

text: On the Origin of Species

text: , in which natural selection was described by analogy to

text: artificial selection

text: , a process by which animals and plants with traits considered desirable by human breeders are systematically favored for reproduction. The concept of natural selection was originally developed in the absence of a valid theory of

text: heredity

text: ; at the time of Darwin's writing, nothing was known of modern genetics. The union of traditional

text: Darwinian evolution

text: with subsequent discoveries in

text: classical

text: and

text: molecular genetics

text: is termed the

text: modern evolutionary synthesis

text: . Natural selection remains the primary explanation for

text: adaptive evolution

text: The Selfish Gene is a 1976 book on evolution by Richard Dawkins, in which Dawkins builds upon the principal theory of George C. Williams's Adaptation and Natural Selection (1966). Dawkins uses the term "selfish gene" as a way of expressing the gene-centred view of evolution as opposed to the views focused on the organism and the group, popularising ideas developed during the 1960s by W. D. Hamilton and others. From the gene-centred view, it follows that the more two individuals are genetically related, the more sense (at the level of the genes) it makes for them to behave selflessly with each other.

text: The Selfish Gene is a 1976 book on

text: evolution

text: by

text: Richard Dawkins

text: , in which Dawkins builds upon the principal theory of

text: George C. Williams

text: 's

text: Adaptation and Natural Selection

text: (1966). Dawkins uses the term "selfish gene" as a way of expressing the

text: gene-centred view of evolution

text: as opposed to the views focused on the organism and the

text: group

text: , popularising ideas developed during the 1960s by

text: W. D. Hamilton

text: and others. From the gene-centred view, it follows that the more two individuals are genetically related, the more sense (at the level of the genes) it makes for them to behave selflessly with each other.

text: In terms of structure, the genes are very similar, suggesting that they arose through gene duplication of an ancestral gene. The most highly related genes are the α- and γ-genes, utilizing two promoters and differing only with the presence of the unique 2a exon in the α-gene. Although substantial differences between alternative exons from the same gene have been revealed by sequence comparison (1a and 1b, 6a and 6b, and the exon 9s), most exons are, however, highly conserved between the different genes. For example, exon 1a and 1b from the α-gene vary considerably in sequence, however the sequence from exon 1a from the α-, β-, γ-, and δ-genes is highly conserved.

text: The genome sequence of Tibetans in 2010 provided the first clue to the molecular evolution of high-altitude adaptation. Genes such as EPAS1, PPARA and EGLN1 are found to have significant molecular changes among the Tibetans, and the genes are involved in haemoglobin production. These genes function in concert with another gene named hypoxia inducible factors (HIF), which in turn is a principal regulator of red blood cell production in response to oxygen metabolism. Further, the Tibetans are enriched for genes in the disease class of human reproduction (such as genes from the DAZ, BPY2, CDY, and HLA-DQ and HLA-DR gene clusters) and biological process categories of response to DNA damage stimulus and DNA repair (such as RAD51, RAD52, and MRE11A), which are related to the adaptive traits of high infant birth weight and darker skin tone and, are most likely due to recent local adaptation.

text: The

text: genome sequence

text: of Tibetans in 2010 provided the first clue to the

text: molecular evolution

text: of high-altitude adaptation. Genes such as

entity_name: Peroxisome proliferator-activated receptor alpha

text: PPARA

text: and

text: are found to have significant

text: molecular changes

text: among the Tibetans, and the genes are involved in

text: haemoglobin production

text: . These genes function in concert with another gene named

text: hypoxia inducible factors

text: (HIF), which in turn is a principal regulator of

text: red blood cell production

text: in response to oxygen metabolism. Further, the Tibetans are enriched for genes in the disease class of human reproduction (such as genes from the

text: DAZ

text: CDY

text: , and

text: HLA-DQ

text: and

text: HLA-DR

text: gene clusters) and biological process categories of response to

text: DNA damage

text: stimulus and

text: DNA repair

text: (such as

text: , and

text: ), which are related to the adaptive traits of high infant birth weight and

text: darker skin tone

text: and, are most likely due to recent local adaptation.

text: Genetic diversity plays an important role in the survival and adaptability of a species. When a population's habitat changes, the population may have to adapt to survive; the ability of the population to adapt to the changing environment will determine their ability to cope with an environmental challenge. Variation in the population's gene pool provides variable traits among the individuals of that population. These variable traits can be selected for, via natural selection, ultimately leading to an adaptive change in the population, allowing it to survive in the changed environment. If a population of a species has a very diverse gene pool then there will be more variety in the traits of individuals of that population and consequently more traits for natural selection to act upon to select the fittest individuals to survive.

text: Genetic diversity plays an important role in the survival and adaptability of a species. When a population's habitat changes, the population may have to adapt to survive; the ability of the population to adapt to the changing environment will determine their ability to cope with an environmental challenge. Variation in the population's

text: gene pool

text: provides variable traits among the individuals of that population. These variable traits can be selected for, via

text: natural selection

text: , ultimately leading to an adaptive change in the population, allowing it to survive in the changed environment. If a population of a species has a very diverse gene pool then there will be more variety in the traits of individuals of that population and consequently more traits for natural selection to act upon to select the

text: fittest

text: individuals to survive.

text: Dawkins has consistently been sceptical about non-adaptive processes in evolution (such as spandrels, described by Gould and Lewontin) and about selection at levels "above" that of the gene. He is particularly sceptical about the practical possibility or importance of group selection as a basis for understanding altruism.This behaviour appears at first to be an evolutionary paradox, since helping others costs precious resources and decreases one's own fitness. Previously, many had interpreted this as an aspect of group selection: individuals are doing what is best for the survival of the population or species as a whole. British evolutionary biologist W. D. Hamilton used gene-frequency analysis in his inclusive fitness theory to show how hereditary altruistic traits can evolve if there is sufficient genetic similarity between actors and recipients of such altruism (including close relatives). Hamilton's inclusive fitness has since been successfully applied to a wide range of organisms, including humans. Similarly, Robert Trivers, thinking in terms of the gene-centred model, developed the theory of reciprocal altruism, whereby one organism provides a benefit to another in the expectation of future reciprocation. Dawkins popularised these ideas in The Selfish Gene, and developed them in his own work. In June 2012 Dawkins was highly critical of fellow biologist E.O. Wilson's 2012 book The Social Conquest of Earth as misunderstanding Hamilton's theory of kin selection. Dawkins has also been strongly critical of the Gaia hypothesis of the independent scientist James Lovelock.

text: Dawkins has consistently been sceptical about non-adaptive processes in evolution (such as

text: spandrels

text: , described by

text: Gould

text: and

text: Lewontin

text: ) and about selection at levels "above" that of the gene. He is particularly sceptical about the practical possibility or importance of

text: group selection

text: as a basis for understanding

text: altruism

text: .This behaviour appears at first to be an evolutionary paradox, since helping others costs precious resources and decreases one's own

text: fitness

text: . Previously, many had interpreted this as an aspect of group selection: individuals are doing what is best for the survival of the population or species as a whole. British evolutionary biologist

text: W. D. Hamilton

text: used gene-frequency analysis in his

text: inclusive fitness

text: theory to show how hereditary altruistic traits can evolve if there is sufficient genetic similarity between actors and recipients of such altruism (including close relatives). Hamilton's inclusive fitness has since been successfully applied to a wide range of organisms, including

text: humans

text: . Similarly,

text: Robert Trivers

text: , thinking in terms of the gene-centred model, developed the theory of

text: reciprocal altruism

text: , whereby one organism provides a benefit to another in the expectation of future reciprocation. Dawkins popularised these ideas in The Selfish Gene, and developed them in his own work. In June 2012 Dawkins was highly critical of fellow biologist

text: E.O. Wilson

text: 's 2012 book

text: The Social Conquest of Earth

text: as misunderstanding Hamilton's theory of kin selection. Dawkins has also been strongly critical of the

text: Gaia hypothesis

text: of the independent scientist

text: James Lovelock

text: For the future, some means have to be found for MNT design evolution at the nanoscale which mimics the process of biological evolution at the molecular scale. Biological evolution proceeds by random variation in ensemble averages of organisms combined with culling of the less-successful variants and reproduction of the more-successful variants, and macroscale engineering design also proceeds by a process of design evolution from simplicity to complexity as set forth somewhat satirically by John Gall: "A complex system that works is invariably found to have evolved from a simple system that worked.  . . . A complex system designed from scratch never works and can not be patched up to make it work. You have to start over, beginning with a system that works."  A breakthrough in MNT is needed which proceeds from the simple atomic ensembles which can be built with, e.g., an STM to complex MNT systems via a process of design evolution. A handicap in this process is the difficulty of seeing and manipulation at the nanoscale compared to the macroscale which makes deterministic selection of successful trials difficult; in contrast biological evolution proceeds via action of what Richard Dawkins has called the "blind watchmaker" comprising random molecular variation and deterministic reproduction/extinction.

text: For the future, some means have to be found for MNT design evolution at the nanoscale which mimics the process of biological evolution at the molecular scale. Biological evolution proceeds by random variation in ensemble averages of organisms combined with culling of the less-successful variants and reproduction of the more-successful variants, and macroscale engineering design also proceeds by a process of design evolution from simplicity to complexity as set forth somewhat satirically by

text: John Gall

text: : "A complex system that works is invariably found to have evolved from a simple system that worked.  . . . A complex system designed from scratch never works and can not be patched up to make it work. You have to start over, beginning with a system that works."  A breakthrough in MNT is needed which proceeds from the simple atomic ensembles which can be built with, e.g., an STM to complex MNT systems via a process of design evolution. A handicap in this process is the difficulty of seeing and manipulation at the nanoscale compared to the macroscale which makes deterministic selection of successful trials difficult; in contrast biological evolution proceeds via action of what Richard Dawkins has called the "blind watchmaker" comprising random molecular variation and deterministic reproduction/extinction.

text: "W.L.S.", reviewing Cott in The Geographical Journal in 1940, begins with "In this large and well-illustrated volume the author discusses at length reason or reasons for the various colour patterns found in the animal kingdom." The reviewer goes on "He has presented us with a vast number of facts and observations which are somewhat difficult to analyse." However "W.L.S." admits that disruptive coloration "is discussed at considerable length by Mr. Cott and many remarkable instances of it are considered in detail". The review ends by mentioning that while biologists (of the 1930s) usually "reject the influence of Natural Selection in evolution, the facts of adaptive coloration as given in Mr. Cott's work are a strong argument in its favour, and must be given due weight. This is what Mr. Cott claims to have accomplished in a volume which will certainly take its place as a most valuable contribution to zoological literature."

text: "W.L.S.", reviewing Cott in The

text: Geographical Journal

text: in 1940, begins with "In this large and well-illustrated volume the author discusses at length reason or reasons for the various colour patterns found in the animal kingdom." The reviewer goes on "He has presented us with a vast number of facts and observations which are somewhat difficult to analyse." However "W.L.S." admits that disruptive coloration "is discussed at considerable length by Mr. Cott and many remarkable instances of it are considered in detail". The review ends by mentioning that while biologists (of the 1930s) usually "reject the influence of Natural Selection in evolution, the facts of adaptive coloration as given in Mr. Cott's work are a strong argument in its favour, and must be given due weight. This is what Mr. Cott claims to have accomplished in a volume which will certainly take its place as a most valuable contribution to zoological literature."

text: Gene families, part of a hierarchy of information storage in a genome, play a large role in the evolution and diversity of multicellular organisms. Gene families are large units of information and genetic variability. Over evolutionary time, gene families have expanded and contracted with new gene families being formed and some gene families being lost. In several evolutionary lineages, genes are gained and lost at relatively same rates. Adaptive expansion of gene families occurs when natural selection would favour additional gene copies. This is the case when an environmental stressor acts on a species. Gene amplification is more common in bacteria and is a reversible process. Adaptive contraction of gene families commonly results from accumulation of loss of function mutations. A nonsense mutation which prematurely halts gene transcription becomes fixed in the population, leading to the loss of genes. This process occurs when changes in the environment render a gene redundant.

text: The technique may be applied to gene frequency in a population of organisms, and to evolution of language.

text: The technique may be applied to

text: gene frequency

text: in a population of organisms, and to

text: evolution of language

text: Because of the mechanism of evolution by natural selection, the environment (“nature”) determines (“selects”) which traits will persist in a population. Therefore, there will be a gradual weeding out of disadvantageous traits over several generations as the population becomes more adapted to its environment. Any significant change in a population’s traits that cannot be attributed to other factors (such as genetic drift and mutation) will be responsive to a change in that population’s environment; in other words, natural selection is inherently reactive. Shortly following an environmental change, traits that evolved in the previous environment, whether they were advantageous or neutral, are persistent for several generations in the new environment. Because evolution is gradual and environmental changes often occur very quickly on a geological scale, there is always a period of “catching-up” as the population evolves to become adapted to the environment. It is this temporary period of “disequilibrium” that is referred to as mismatch. Mismatched traits are ultimately addressed in one of several possible ways: the organism may evolve such that the maladaptive trait is no longer expressed, the organism may decline and/or become extinct as a result of the disadvantageous trait, or the environment may change such that the trait is no longer selected against.

text: Because of the mechanism of

text: evolution

text: by natural selection, the environment (“nature”) determines (“selects”) which traits will persist in a population. Therefore, there will be a gradual weeding out of disadvantageous traits over several generations as the population becomes more adapted to its environment. Any significant change in a population’s traits that cannot be attributed to other factors (such as genetic drift and mutation) will be responsive to a change in that population’s environment; in other words, natural selection is inherently reactive. Shortly following an environmental change, traits that evolved in the previous environment, whether they were advantageous or neutral, are persistent for several generations in the new environment. Because

text: evolution is gradual

text: and environmental changes often occur very quickly on a geological scale, there is always a period of “catching-up” as the population evolves to become adapted to the environment. It is this temporary period of “disequilibrium” that is referred to as mismatch. Mismatched traits are ultimately addressed in one of several possible ways: the organism may evolve such that the maladaptive trait is no longer expressed, the organism may decline and/or become extinct as a result of the disadvantageous trait, or the environment may change such that the trait is no longer selected against.

text: A neutral network is a set of genes all related by point mutations that have equivalent function or fitness. Each node represents a gene sequence and each line represents the mutation connecting two sequences. Neutral networks can be thought of as high, flat plateaus in a fitness landscape. During neutral evolution, genes can randomly move through neutral networks and traverse regions of sequence space which may have consequences for robustness and evolvability.

text: A neutral network is a set of

text: gene

text: s all related by

text: point mutation

text: s that have equivalent function or

text: fitness

text: . Each node represents a gene sequence and each line represents the mutation connecting two sequences. Neutral networks can be thought of as high, flat plateaus in a

text: fitness landscape

text: . During

text: neutral evolution

text: , genes can randomly move through neutral networks and traverse regions of

text: sequence space

text: which may have consequences for

text: robustness

text: and

text: evolvability

text: Although Delbrück's Rockefeller Foundation fellowship expired in 1939, the Foundation matched him up with Vanderbilt University in Nashville, Tennessee, where from 1940 to 1947 he taught physics, yet had his laboratory in the biology department.  In 1941, Delbrück met Salvador Luria of Indiana University who began visiting Vanderbilt.  In 1942, Delbrück and Luria published on bacterial resistance to virus infection mediated by random mutation.  Alfred Hershey of Washington University began visiting in 1943. The Luria–Delbrück experiment, also called the Fluctuation Test, demonstrated that Darwin's theory of natural selection acting on random mutations applies to bacteria as well as to more complex organisms. The 1969 Nobel Prize in Physiology or Medicine was awarded to both scientists in part for this work. To put this work in its historical perspective, Lamarck in 1801 first presented his theory of Inheritance of Acquired Characteristics, which stated that if an organism changes during life in order to adapt to its environment (for example stretches its neck to reach for tall trees), those changes are passed on to its offspring. He also said that evolution happens according to a predetermined plan. Darwin published his theory of evolution in his 1859 book On the Origin of Species with compelling evidence contradicting Lamarck. Darwin said that evolution is not predetermined but that there are inherent variations in all organisms, and that those variations that confer increased fitness are selected by the environment and passed on to the offspring. In the feud between Lamarck and Darwin, Darwin talked of pre-existing changes, but the nature of these changes was not known and had to await the science of genetics by Gregor Mendel’s experiments on pea plants published in 1866. Support for Darwin’s theory was provided when Thomas Morgan Hunt discovered that a mutated white-eyed fruit fly among red-eyed flies was able to reproduce true white-eyed offspring. The most elegant and convincing support for Darwin’s ideas, however, was provided by the Luria-Delbruck experiment, which showed that mutations conferring resistance of the bacterium E. coli to T1 bacteriophage (virus) existed in the population prior to exposure to T1 and were not induced by adding T1. In other words, mutations are random events that occur whether or not they prove to be useful, while selection (for T1 resistance upon challenge with T1 in this case) provides the direction in evolution by retaining those mutations that are advantageous, discarding those that are harmful (T1 sensitivity in this case). This experiment dealt a blow to Lamarckian inheritance and set the stage for tremendous advances in genetics and molecular biology, launching a tsunami of research that eventually led to the discovery of DNA as the hereditary material and to cracking the genetic code.

text: Although Delbrück's

text: Rockefeller Foundation

text: fellowship expired in 1939, the Foundation matched him up with

text: Vanderbilt University

text: in

text: Nashville, Tennessee

text: , where from 1940 to 1947 he taught

text: physics

text: , yet had his laboratory in the biology department.  In 1941, Delbrück met

text: Salvador Luria

text: of

text: Indiana University

text: who began visiting Vanderbilt.  In 1942,

text: Delbrück and Luria published

text: on

text: bacteria

text: l resistance to

text: virus

text: infection mediated by random

text: mutation

text: Alfred Hershey

text: of

text: Washington University

text: The 5-speed automatic gearbox had what Mitsubishi called "fuzzy logic", which meant that the car would learn what the driver's driving characteristics were like and would adapt the gear change timings and kick down reactions accordingly. The gears could be manually selected as with most Tiptronics via steering wheel + and – buttons (a pair both sides) or via selecting the tiptronic gate with the gear lever. Power was down a little from the standard manual cars with . The GT-A gearbox did not appear again in the Evolution VIII but has been installed in the estate version of the Evolution IX Wagon. It was replaced by the Twin Clutch SST gearbox since the introduction of Evolution X.

text: The 5-speed automatic gearbox had what Mitsubishi called "fuzzy logic", which meant that the car would learn what the driver's driving characteristics were like and would adapt the gear change timings and kick down reactions accordingly. The gears could be manually selected as with most Tiptronics via steering wheel + and – buttons (a pair both sides) or via selecting the tiptronic gate with the gear lever. Power was down a little from the standard manual cars with . The GT-A gearbox did not appear again in the Evolution VIII but has been installed in the estate version of the Evolution IX Wagon. It was replaced by the

text: Twin Clutch SST

text: gearbox since the introduction of Evolution X.

text: Professor of biology Jerry Coyne sums up biological evolution succinctly:

text: Professor of biology

text: Jerry Coyne

text: sums up biological evolution succinctly:

text: Natural selection within a population for a trait that can vary across a range of values, such as height, can be categorised into three different types. The first is directional selection, which is a shift in the average value of a trait over time—for example, organisms slowly getting taller. Secondly, disruptive selection is selection for extreme trait values and often results in two different values becoming most common, with selection against the average value. This would be when either short or tall organisms had an advantage, but not those of medium height. Finally, in stabilising selection there is selection against extreme trait values on both ends, which causes a decrease in variance around the average value and less diversity. This would, for example, cause organisms to slowly become all the same height.

text: Natural selection within a population for a trait that can vary across a range of values, such as height, can be categorised into three different types. The first is

text: directional selection

text: , which is a shift in the average value of a trait over time—for example, organisms slowly getting taller. Secondly,

text: disruptive selection

text: is selection for extreme trait values and often results in

text: two different values

text: becoming most common, with selection against the average value. This would be when either short or tall organisms had an advantage, but not those of medium height. Finally, in

text: stabilising selection

text: there is selection against extreme trait values on both ends, which causes a decrease in

text: variance

text: around the average value and less diversity. This would, for example, cause organisms to slowly become all the same height.

text: Epigenetic inheritance may only affect fitness if it predictably alters a trait under selection.  Evidence has been forwarded that environmental stimuli are important agents in the alteration of epigenes.  Ironically, Darwinian evolution may act on these neo-Lamarckian acquired characteristics as well as the cellular mechanisms producing them (e.g. methyltransferase genes).Epigenetic inheritance may confer a fitness benefit to organisms that deal with environmental changes at intermediate timescales.   Short-cycling changes are likely to have DNA-encoded regulatory processes, as the probability of the offspring needing to respond to changes multiple times during their lifespans is high.  On the other end, natural selection will act on populations experiencing changes on longer-cycling environmental changes.  In these cases, if epigenetic priming of the next generation is deleterious to fitness over most of the interval (e.g. misinformation about the environment), these genotypes and epigenotypes will be lost.  For intermediate time cycles, the probability of the offspring encountering a similar environment is sufficiently high without substantial selective pressure on individuals lacking a genetic architecture capable of responding to the environment.  Naturally, the absolute lengths of short, intermediate, and long environmental cycles will depend on the trait, the length of epigenetic memory, and the generation time of the organism.Much of the interpretation of epigenetic fitness effects centers on the hypothesis that epigenes are important contributors to phenotypes, which remains to be resolved.

text: Epigenetic inheritance may only affect

text: fitness

text: Genes generally express their functional effect through the production of proteins, which are complex molecules responsible for most functions in the cell. Proteins are made up of one or more polypeptide chains, each of which is composed of a sequence of amino acids, and the DNA sequence of a gene (through an RNA intermediate) is used to produce a specific amino acid sequence. This process begins with the production of an RNA molecule with a sequence matching the gene's DNA sequence, a process called transcription.

text: Genes generally

text: express

text: their functional effect through the production of

text: protein

text: s, which are complex molecules responsible for most functions in the cell. Proteins are made up of one or more polypeptide chains, each of which is composed of a sequence of

text: amino acid

text: s, and the DNA sequence of a gene (through an RNA intermediate) is used to produce a specific

text: amino acid sequence

text: . This process begins with the production of an

text: RNA

text: molecule with a sequence matching the gene's DNA sequence, a process called

text: transcription

text: Three critical assumptions underlie the study of Mhc evolution: First, the Mhc is absent in all non-vertebrates. Second, jawless vertebrates (Agnatha) are monophyletic and are a sister group of jawed vertebrates (Gnathostoma). And third, jawless vertebrates lack the Mhc, which is present in all the jawed vertebrates. Klein’s group contributed significantly to the current general acceptance of these suppositions. The absence of the Mhc in non-vertebrates became apparent when scrutiny of non-vertebrate genomes failed to identify homologs of Mhc genes. Klein’s group provided strong support for agnathan monophyly by cloning, sequencing, and analyzing long DNA stretches of the representative agnathan and gnathostome species. And they isolated, in collaboration with Max Cooper’s group, lymphocyte-like cells, cloned, sequenced, and analyzed genes expressed in these cells, and found no evidence for expressed Mhc gene homologs. They did find, however, evidence for gradual evolution of the adaptive immune system. They could demonstrate the presence in jawless vertebrates of several auxiliary components and pathways, which the AIS co-opted when the three central receptors (Mhc, Tcr, and Bcr) emerged in the jawed vertebrates. They also contributed evidence for the omnipresence of Mhc genes in jawed vertebrates by identifying such genes in a wide range of species from bony fishes [zebrafish (Danio rerio), cichlid Aulonocara hansbaenschi, tilapia (Oreochromis niloticus), carp (Cyprinus carpio), guppy (Poecilia reticulata), threespine stickleback (Gasterosteus aculeatus), swordtail (Xiphophorus)]; through coelacanth (Latimeria chalumnae), African lungfish (Protopterus aethiopicus); birds [Bengalese finch (Lonchura striata), Darwin’s finches and their South American relatives]; to metatherian mammals [red-necked wallaby (Macropus rufogriseus) and eutherian mammal [rodents such as the mole rat (Spalax ehrenbergi)] and a variety of primates including prosimians, New World monkeys (NWM, Platyrrhini), Old World monkeys (OWM, Catarrhini) and apes]. In several of these species they worked out also the organization of the Mhcs, most notably in the zebrafish.

text: The Genetical Theory of Natural Selection is a book by Ronald Fisher which combines Mendelian genetics with Charles Darwin's theory of natural selection, with Fisher being the first to argue that "Mendelism therefore validates Darwinism" and stating with regard to mutations that "The vast majority of large mutations are deleterious; small mutations are both far more frequent and more likely to be useful", thus refuting orthogenesis. First published in 1930 by The Clarendon Press, it is one of the most important books of the modern evolutionary synthesis, and helped define population genetics. It is commonly cited in biology books, outlining many concepts that are still considered important such as Fisherian runaway, Fisher's principle, reproductive value, Fisher's fundamental theorem of natural selection, Fisher's geometric model, the sexy son hypothesis, mimicry and the evolution of dominance. It was dictated to his wife in the evenings as he worked at Rothamsted Research in the day.

text: The Genetical Theory of Natural Selection is a book by

text: Ronald Fisher

text: which combines

text: Mendelian genetics

text: with

text: Charles Darwin

text: 's theory of

text: natural selection

text: , with Fisher being the first to argue that "Mendelism therefore validates Darwinism" and stating with regard to

text: mutation

text: s that "The vast majority of large mutations are deleterious; small mutations are both far more frequent and more likely to be useful", thus refuting

text: orthogenesis

text: . First published in 1930 by

text: The Clarendon Press

text: , it is one of the most important books of the

text: modern evolutionary synthesis

text: , and helped define

text: population genetics

text: . It is commonly cited in biology books, outlining many concepts that are still considered important such as

text: Fisherian runaway

text: Fisher's principle

entity_name: Reproductive value (population genetics)

text: reproductive value

entity_name: Fisher's fundamental theorem of natural selection

text: Fisher's fundamental theorem of natural selection

text: Fisher's geometric model

text: , the

text: sexy son hypothesis

text: mimicry

text: and the

text: evolution of dominance

text: . It was dictated to his wife in the evenings as he worked at

text: Rothamsted Research

text: in the day.

text: CMA-ES stands for Covariance Matrix Adaptation Evolution Strategy. Evolution strategies (ES) are stochastic, derivative-free methods for numerical optimization  of non-linear or non-convex continuous optimization problems. They belong to the class of evolutionary algorithms and evolutionary computation.  An evolutionary algorithm is broadly based on the principle of biological evolution, namely the repeated interplay of variation (via recombination and mutation) and selection: in each generation (iteration) new individuals (candidate solutions, denoted as

text: CMA-ES stands for Covariance Matrix Adaptation Evolution Strategy.

text: Evolution strategies

text: (ES) are

text: stochastic

text: derivative-free methods

text: for

text: numerical optimization

text: of non-

text: linear

text: or non-

text: convex

text: continuous optimization

text: problems. They belong to the class of

text: evolutionary algorithms

text: and

text: evolutionary computation

text: .  An

text: evolutionary algorithm

text: is broadly based on the principle of

text: biological evolution

text: , namely the repeated interplay of variation (via recombination and mutation) and selection: in each generation (iteration) new individuals (candidate solutions, denoted as

text: Natural selection acts on the phenotype, or the observable characteristics of an organism, but the genetic (heritable) basis of any phenotype which gives a reproductive advantage will become more common in a population (see allele frequency). Over time, this process can result in adaptations that specialize organisms for particular ecological niches and may eventually result in the speciation (the emergence of new species).

text: Natural selection acts on the

text: phenotype

text: , or the observable characteristics of an organism, but the

text: genetic

text: (heritable) basis of any phenotype which gives a reproductive advantage will become more common in a population (see

text: allele frequency

text: ). Over time, this process can result in

text: adaptation

text: s that specialize organisms for particular

text: ecological niche

text: s and may eventually result in the speciation (the emergence of new species).

text: Natural selection, introduced in 1859 as the central mechanism of evolution by Charles Darwin, is the differential survival and reproduction of individuals due to differences in phenotype. The mechanism directly implies evolution, a change in heritable traits of a population over time.

text: Natural selection

text: , introduced in 1859 as the central mechanism of

text: evolution

text: by

text: Charles Darwin

text: , is the differential survival and reproduction of individuals due to differences in

text: phenotype

text: . The mechanism directly implies evolution, a change in

text: heritable

text: traits

text: of a

text: population

text: over time.

text: There is some evidence of adaptive evolution in genes linked to brain development, but some of these genes are often associated with diseases, e.g. microcephaly (see Table 2). However, there is a particular interest in the search for adaptive evolution in brain genes, despite the ethical issues surrounding such research. If more adaptive evolution was discovered in brain genes in one human population than another, then this information could be interpreted as showing greater intelligence in the more adaptively evolved population.

text: Substantial evidence in Tibetan highlanders suggests that variation in haemoglobin and blood-oxygen levels are adaptive as Darwinian fitness. It has been documented that Tibetan women with a high likelihood of possessing one to two alleles for high blood-oxygen content (which is odd for normal women) had more surviving children; the higher the oxygen capacity, the lower the infant mortality. In 2010, for the first time, the genes responsible for the unique adaptive traits were identified following genome sequences of 50 Tibetans and 40 Han Chinese from Beijing. Initially, the strongest signal of natural selection detected was a transcription factor involved in response to hypoxia, called endothelial Per-Arnt-Sim (PAS) domain protein 1 (EPAS1). It was found that one single-nucleotide polymorphism (SNP) at EPAS1 shows a 78% frequency difference between Tibetan and mainland Chinese samples, representing the fastest genetic change observed in any human gene to date. Hence, Tibetan adaptation to high altitude becomes the fastest process of phenotypically observable evolution in humans, which is estimated to occur in less than 3,000 years ago, when the Tibetans split up from the mainland Chinese population. Mutations in EPAS1, at higher frequency in Tibetans than their Han neighbours, correlate with decreased haemoglobin concentrations among the Tibetans, which is the hallmark of their adaptation to hypoxia. Simultaneously, two genes, egl nine homolog 1 (EGLN1) (which inhibits haemoglobin production under high oxygen concentration) and peroxisome proliferator-activated receptor alpha (PPARA), were also identified to be positively selected in relation to decreased haemoglobin nature in the Tibetans. Similarly, the Sherpas, known for their Himalayan explorations, exhibit similar patterns in the EPAS1 gene, which further fortifies that the gene is under selection for adaptation to the high-altitude life of Tibetans. A study in 2014 indicates that the mutant EPAS1 gene could have been inherited from archaic hominins, the Denisovans. EPAS1 and EGLN1 are definitely the major genes for unique adaptive traits when compared with those of the Chinese and Japanese. Comparative genome analysis in 2014 revealed that the Tibetans inherited an equal mixture of genomes from the Nepalese-Sherpas and Hans, and they acquired the adaptive genes from the sherpa-lineage. Further, the population split was estimated to occur around 20,000 to 40,000 years ago, a range of which support archaeological, mitochondria DNA and Y chromosome evidence for an initial colonisation of the Tibetan plateau around 30,000 years ago. The genes (EPAS1, EGLN1, and PPARA) function in concert with another gene named hypoxia inducible factors (HIF), which in turn is a principal regulator of red blood cell production (erythropoiesis) in response to oxygen metabolism. The genes  are associated not only with decreased haemoglobin levels, but also in regulating energy metabolism. EPAS1 is significantly associated with increased lactate concentration (the product of anaerobic glycolysis), and PPARA is correlated with decrease in the activity of fatty acid oxidation. EGLN1 codes for an enzyme, prolyl hydroxylase 2 (PHD2), involved in erythropoiesis. Among the Tibetans, mutation in EGLN1 (specifically at position 12, where cytosine is replaced with guanine; and at 380, where G is replaced with C) results in mutant PHD2 (aspartic acid at position 4 becomes glutamine, and cysteine at 127 becomes serine) and this mutation inhibits erythropoiesis. The mutation is estimated to occur about 8,000 years ago. Further, the Tibetans are enriched for genes in the disease class of human reproduction (such as genes from the DAZ, BPY2, CDY, and HLA-DQ and HLA-DR gene clusters) and biological process categories of response to DNA damage stimulus and DNA repair (such as RAD51, RAD52, and MRE11A), which are related to the adaptive traits of high infant birth weight and darker skin tone and are most likely due to recent local adaptation.

text: Substantial evidence in Tibetan highlanders suggests that variation in haemoglobin and

text: blood-oxygen

text: levels are adaptive as

text: Darwinian fitness

text: . It has been documented that Tibetan women with a high likelihood of possessing one to two

text: allele

text: s for high blood-oxygen content (which is odd for normal women) had more surviving children; the higher the oxygen capacity, the lower the infant mortality. In 2010, for the first time, the genes responsible for the unique adaptive traits were identified following genome sequences of 50 Tibetans and 40

text: Han Chinese

text: from

text: Beijing

text: . Initially, the strongest signal of natural selection detected was a transcription factor involved in response to hypoxia, called endothelial Per-Arnt-Sim (PAS) domain protein 1 (

text: ). It was found that one

text: single-nucleotide polymorphism

text: (SNP) at EPAS1 shows a 78% frequency difference between Tibetan and mainland Chinese samples, representing the fastest genetic change observed in any human gene to date. Hence, Tibetan adaptation to high altitude becomes the fastest process of

text: phenotypically observable

text: evolution in humans, which is estimated to occur in less than 3,000 years ago, when the Tibetans split up from the mainland Chinese population.

text: Mutations

text: in EPAS1, at higher frequency in Tibetans than their Han neighbours, correlate with decreased haemoglobin concentrations among the Tibetans, which is the hallmark of their adaptation to hypoxia. Simultaneously, two genes, egl nine homolog 1 (

text: ) (which inhibits haemoglobin production under high oxygen concentration) and peroxisome proliferator-activated receptor alpha (

text: PPARA

text: ), were also identified to be positively selected in relation to decreased haemoglobin nature in the Tibetans. Similarly, the

text: Sherpas

text: , known for their Himalayan explorations, exhibit similar patterns in the EPAS1 gene, which further fortifies that the gene is under

text: selection

text: for adaptation to the high-altitude life of Tibetans. A study in 2014 indicates that the mutant

text: gene could have been inherited from archaic hominins, the

text: Denisovans

text: . EPAS1 and EGLN1 are definitely the major genes for unique adaptive traits when compared with those of the Chinese and Japanese. Comparative genome analysis in 2014 revealed that the Tibetans inherited an equal mixture of genomes from the Nepalese-Sherpas and Hans, and they acquired the adaptive genes from the sherpa-lineage. Further, the population split was estimated to occur around 20,000 to 40,000 years ago, a range of which support archaeological, mitochondria DNA and Y chromosome evidence for an initial colonisation of the Tibetan plateau around 30,000 years ago. The genes (EPAS1, EGLN1, and PPARA) function in concert with another gene named

text: hypoxia inducible factors

text: (HIF), which in turn is a principal regulator of red blood cell production (

text: erythropoiesis

text: ) in response to oxygen metabolism. The genes  are associated not only with decreased haemoglobin levels, but also in regulating energy metabolism. EPAS1 is significantly associated with increased lactate concentration (the product of

text: anaerobic glycolysis

text: ), and PPARA is correlated with decrease in the activity of

text: fatty acid oxidation

text: . EGLN1 codes for an enzyme, prolyl hydroxylase 2 (PHD2), involved in erythropoiesis. Among the Tibetans, mutation in EGLN1 (specifically at position 12, where cytosine is replaced with guanine; and at 380, where G is replaced with C) results in mutant PHD2 (aspartic acid at position 4 becomes glutamine, and cysteine at 127 becomes serine) and this mutation inhibits erythropoiesis. The mutation is estimated to occur about 8,000 years ago. Further, the Tibetans are enriched for genes in the disease class of human reproduction (such as genes from the

text: DAZ

text: CDY

text: , and

text: HLA-DQ

text: and

text: HLA-DR

text: DNA damage

text: stimulus and

text: DNA repair

text: (such as

text: , and

text: darker skin tone

text: and are most likely due to recent local adaptation.

text: Kin selection (commonly referred to as altruism, even though it is just a sub-type) is an example of an adaptive behavior that directly influences the genetic composition of a population. It involves evolutionary strategies that favour the persistence of an organism's relatives, often at the cost of their own survival and reproduction. This will result in population gene frequency variation over successive generations, based on the interactions between related individuals. The probability of altruism increases when the cost is low for the donor, or when there is a high level of gain for the beneficiary. In addition, individuals often display altruistic behaviors when the relative is genetically similar to them. This means offspring or siblings are more likely to benefit from altruism than someone more distantly related, such as a cousin, aunt, or uncle.Kin selection has played a large role in the evolution of social and adaptive behaviors in chimpanzees. Closely related chimpanzees will form a kin group that cooperates to protect a territory, thereby increasing their access to females and resources. By working together with close relatives, they can ensure that their genes will persist in the next generation, even if circumstances make them unable to reproduce themselves. This behavioral adaptation coincides with chimpanzee's ability to distinguish kin from non-kin (referred to as visual kin selection). This allows chimpanzees to formulate large, complex societies, where they use altruistic methods to ensure their genes persist in future generations.  A wide variety of species have displayed kin selection as an adaptive behavior. Examples include lions, honeybees, and insects.

text: Kin selection

text: (commonly referred to as

text: altruism

text: , even though it is just a sub-type) is an example of an adaptive behavior that directly influences the genetic composition of a population. It involves evolutionary strategies that favour the persistence of an organism's relatives, often at the cost of their own survival and

text: reproduction

text: . This will result in population gene

text: frequency

text: variation over successive generations, based on the interactions between related individuals. The probability of altruism increases when the

text: cost

text: is low for the donor, or when there is a high level of gain for the beneficiary. In addition, individuals often display altruistic behaviors when the relative is genetically similar to them. This means offspring or siblings are more likely to benefit from altruism than someone more distantly related, such as a cousin, aunt, or uncle.Kin selection has played a large role in the evolution of social and adaptive behaviors in chimpanzees. Closely related chimpanzees will form a kin group that cooperates to protect a territory, thereby increasing their access to females and resources. By working together with close relatives, they can ensure that their genes will persist in the next generation, even if circumstances make them unable to reproduce themselves. This behavioral adaptation coincides with chimpanzee's ability to distinguish kin from non-kin (referred to as visual kin selection). This allows chimpanzees to formulate large, complex societies, where they use altruistic methods to ensure their genes persist in future generations.  A wide variety of species have displayed kin selection as an adaptive behavior. Examples include lions, honeybees, and insects.

text: Rasmussen was a biological anthropologist who specialized in both paleontology and behavioral ecology. He studied primate evolution, utilizing his knowledge of both living and fossil primates. His primary research interests were the adaptive radiation of prosimian primates, particularly their life history and evolution, as well as evolutionary origins of both simians (anthropoids) and primates in general.

text: Rasmussen was a

text: biological anthropologist

text: who specialized in both

text: paleontology

text: and

text: behavioral ecology

text: . He studied

text: primate evolution

text: , utilizing his knowledge of both living and

text: fossil primates

text: . His primary research interests were the

text: adaptive radiation

text: of

text: prosimian

text: primates, particularly their life history and evolution, as well as evolutionary origins of both

text: simian

text: s (anthropoids) and primates in general.

text: Scientists have observed and documented a multitude of events where natural selection is in action. The most well known examples are antibiotic resistance in the medical field along with better-known laboratory experiments documenting evolution's occurrence. Natural selection is tantamount to common descent in that long-term occurrence and selection pressures can lead to the diversity of life on earth as found today. All adaptations—documented and undocumented changes concerned—are caused by natural selection (and a few other minor processes). It is well established that, "...natural selection is a ubiquitous part of speciation...", and is the primary driver of speciation; therefore, the following examples of natural selection and speciation will often interdepend or correspond with one another. The examples below are only a small fraction of the actual experiments and observations.

text: Structural variation can be seen as an avenue of genome modification for adaptation by evolution. A study was conducted on ancestral diet and the evolution of the human amylase gene copy number. The consumption of starch became a huge component of the human diet with the development of agricultural societies. Amylase is the enzyme that breaks down starch and its copy number varies. These observations led to the question of whether or not the differences in starch consumption between different populations created natural selection pressures on the enzyme amylase. They tested for the differences in the amylase protein expression in saliva from different populations and compared their expression to their copy number in their respective genomes. Then they compared the starch consumption of different populations to their copy number of the amylase gene. They found that there was more amylase protein expression in saliva from people that had higher amylase copy number in their genome and there was also an association between groups of people with high starch diets and a larger amylase gene copy number. This study brought exciting results as structural variation proved an involvement in the evolution of the human population by increasing its amylase copy number over time.

text: Selection favors different traits in captive populations than it does in wild populations, so this may result in adaptations that are beneficial in captivity but are deleterious in the wild. This reduces the success of re-introductions, so it is important to manage captive populations in order to reduce adaptations to captivity.  Adaptations to captivity can be reduced by minimizing the number of generations in captivity and by maximizing the number of migrants from wild populations.  Minimizing selection on captive populations by creating an environment that is similar to their natural environment is another method of reducing adaptations to captivity, but it is important to find a balance between an environment that minimizes adaptation to captivity and an environment that permits adequate reproduction. Adaptations to captivity can also be reduced by managing the captive population as a series of population fragments.  In this management strategy, the captive population is split into several sub-populations or fragments which are maintained separately.  Smaller populations have lower adaptive potentials, so the population fragments are less likely to accumulate adaptations associated with captivity.  The fragments are maintained separately until inbreeding becomes a concern.  Immigrants are then exchanged between the fragments to reduce inbreeding, and then the fragments are managed separately again.

text: Selection

text: favors different traits in captive populations than it does in wild populations, so this may result in adaptations that are beneficial in captivity but are deleterious in the wild. This reduces the success of re-introductions, so it is important to manage captive populations in order to reduce adaptations to captivity.  Adaptations to captivity can be reduced by minimizing the number of generations in captivity and by maximizing the number of migrants from wild populations.  Minimizing selection on captive populations by creating an environment that is similar to their natural environment is another method of reducing adaptations to captivity, but it is important to find a balance between an environment that minimizes adaptation to captivity and an environment that permits adequate reproduction. Adaptations to captivity can also be reduced by managing the captive population as a series of population fragments.  In this management strategy, the captive population is split into several sub-populations or fragments which are maintained separately.  Smaller populations have lower adaptive potentials, so the population fragments are less likely to accumulate adaptations associated with captivity.  The fragments are maintained separately until

text: inbreeding

text: becomes a concern.  Immigrants are then exchanged between the fragments to reduce inbreeding, and then the fragments are managed separately again.

text: If the directional evolution model is valid then the evolution must have been slow and the timing of events would have overlapped considerably. A valid end member interpretation is that the crater population still represents a population emplaced on a mostly inactive planet, but the final throes of a global emplacement of volcanic plains has filled most of the craters with a few hundred meters of volcanic flows. If this is true, then post-tessera plains emplacement must have dragged on for most of the visible surface history of the planet and  the cessation of tessera deformation must have overlapped considerably with emplacement of plains. Thus, while a tessera/plains/rifts evolution is a valid hypothesis, that evolution could not have occurred as a "catastrophe". The highly varying levels of post-impact volcanism and deformation that the craters have experienced are consistent with a steady state model of Venus resurfacing. The craters are in a variety of stages of removal but display the same processes that have operated throughout the visible surface history. It remains a powerful constraint that the distribution of geologic features on the planet (plains, volcanoes, rifts, etc.) is decidedly more nonuniform than the crater population. This means that while the nature of resurfacing on Venus may vary regionally in the uniformitarian hypothesis, the rates must be similar.

text: A considerable number of studies have used genomic methods to identify specific human genes that show evidence of adaptive evolution. Table 2 gives selected examples of such genes for each gene type discussed, but provides nowhere near an exhaustive list of the human genes showing evidence of adaptive evolution. Below are listed some of the types of gene which show strong evidence of adaptive evolution in the human genome.

text: Tautz's research covers a broad interest area in the fields of molecular evolution and developmental genetics. His thesis dealt with the first generic description of simple sequences (now often called microsatellites).This work became later the basis for his development of the genetic fingerprinting technology that is nowadays used in paternity testing, forensics and population genetics and for which he obtained a broad patent. For his second postdoc phase in Tübingen, he switched to developmental genetics joining the groups around Herbert Jäckle and Christiane Nüsslein-Volhard. His work there focused on the molecular characterization of segmentation genes in Drosophila. He cloned and characterized the segmentation gene ‘’hunchback’’ and he developed the "whole mount" in situ hybridization technology that is nowadays used as the general standard technology for the spatial and temporal analysis of gene expression in embryos. Based on the work on segmentation processes in Drosophila, he developed a general interest in studying the evolution of segmentation processes, especially in the model system Tribolium and he was involved in generating the first full genome sequencing for this beetle species. He is considered to be one of the founders of the new field of evolution of developmental processes (often called Evo-Devo) and he was the first editor in chief of the journal Development Genes and Evolution. As professor for Zoology in Munich he expanded his interests into the fields of population genetics, speciation research and molecular phylogeny reconstruction. He studied the mechanisms of sympatric speciation of cichlids in crater lakes, hybrid speciation of sculpins in the Rhine system and was the first to show a sister group relationship between crustaceans and insects in the phylogeny of arthropods. With his move to the University of Cologne he started projects in comparative genomics, with a special focus on the evolution of new genes, which resulted in the discovery of de novo evolution of genes from non-coding sequences. Through studying a segmentation gene in insects, he found the first eukaryotic poly-cistronic peptide-coding gene. In Cologne he started also his current projects on the genetic basis of adaptations to new environmental conditions. He uses natural populations and subspecies of the house mouse (Mus musculus) as a model system for this  work, which he analyses with methods from population genetics, comparative genomics and quantitative genetics.

text: Tautz's research covers a broad interest area in the fields of

text: molecular evolution

text: and

text: developmental genetics

text: . His thesis dealt with the first generic description of simple sequences (now often called

text: microsatellite

text: s).This work became later the basis for his development of the

text: genetic fingerprinting

text: technology that is nowadays used in paternity testing, forensics and population genetics and for which he obtained a broad patent. For his second postdoc phase in Tübingen, he switched to developmental genetics joining the groups around Herbert Jäckle and

text: Christiane Nüsslein-Volhard

text: . His work there focused on the molecular characterization of segmentation genes in

text: Drosophila

text: . He cloned and characterized the segmentation gene ‘’hunchback’’ and he developed the "whole mount"

text: in situ hybridization

text: technology that is nowadays used as the general standard technology for the spatial and temporal analysis of gene expression in embryos. Based on the work on

text: segmentation processes

text: in Drosophila, he developed a general interest in studying the evolution of segmentation processes, especially in the model system Tribolium and he was involved in generating the first full genome sequencing for this beetle species. He is considered to be one of the founders of the new field of evolution of developmental processes (often called

text: Evo-Devo

text: ) and he was the first editor in chief of the journal Development Genes and Evolution. As professor for Zoology in Munich he expanded his interests into the fields of

text: population genetics

text: , speciation research and molecular phylogeny reconstruction. He studied the mechanisms of

text: sympatric speciation

text: of

text: cichlids

text: in crater lakes, hybrid speciation of

text: sculpin

text: s in the Rhine system and was the first to show a sister group relationship between crustaceans and insects in the phylogeny of arthropods. With his move to the University of Cologne he started projects in

text: comparative genomics

text: , with a special focus on the evolution of new genes, which resulted in the discovery of de novo evolution of genes from

text: non-coding sequences

text: . Through studying a segmentation gene in insects, he found the first eukaryotic poly-cistronic peptide-coding gene. In Cologne he started also his current projects on the genetic basis of adaptations to new environmental conditions. He uses natural populations and

text: subspecies

text: of the

text: house mouse

text: (Mus musculus) as a model system for this  work, which he analyses with methods from

text: population genetics

text: , comparative genomics and

text: quantitative genetics

text: Stabilizing selection (not the same thing as negative selection) is a type of natural selection in which genetic diversity decreases and the population mean stabilizes on a particular trait value. This is thought to be the most common mechanism of action for natural selection because most traits do not appear to change drastically over time. Stabilizing selection commonly uses negative selection (a.k.a. purifying selection) to select against extreme values of the character. Stabilizing selection is the opposite of disruptive selection. Instead of favoring individuals with extreme phenotypes, it favors the intermediate variants. It reduces phenotypic variation and maintains the status quo. Natural selection tends to remove the more severe phenotypes, resulting in the reproductive success of the norm or average phenotypes.   Because most traits change little over time, stabilizing selection is thought to be the most common type of selection in most populations.  However, a meta-analysis of studies that measured selection in the wild failed to find an overall trend for stabilizing selection.  The reason can be that methods for detecting stabilizing selection are complex. They can involve studying the changes that causes natural selection in the mean and variance of the trait, or measuring fitness for a range of different phenotypes under natural conditions and examining the relationship between these fitness measurements and the trait value, but analysis and interpretation of the results is not straightforward.

text: Stabilizing selection (not the same thing as

text: negative selection

text: ) is a type of

text: natural selection

text: in which

text: genetic diversity

text: decreases and the

text: population

text: mean stabilizes on a particular

text: trait

text: value. This is thought to be the most common mechanism of action for natural selection because most traits do not appear to change drastically over time. Stabilizing selection commonly uses negative selection (a.k.a. purifying selection) to select against extreme values of the character. Stabilizing selection is the opposite of

text: disruptive selection

text: . Instead of favoring individuals with extreme phenotypes, it favors the intermediate variants. It reduces phenotypic variation and maintains the status quo. Natural selection tends to remove the more severe phenotypes, resulting in the reproductive success of the norm or average phenotypes.   Because most traits change little over time, stabilizing selection is thought to be the most common type of selection in most populations.  However, a meta-analysis of studies that measured selection in the wild failed to find an overall trend for stabilizing selection.  The reason can be that methods for detecting stabilizing selection are complex. They can involve studying the changes that causes natural selection in the mean and variance of the trait, or measuring fitness for a range of different

text: phenotype

text: s under natural conditions and examining the relationship between these fitness measurements and the trait value, but analysis and interpretation of the results is not straightforward.

text: A small body's function in the environment would then be adaptive, but it wouldn't become an adaptation until enough generations had passed to in which small bodies were advantageous to reproduction for small bodies to selected for. Given this, it is best to understand that presently functional traits might not all have been produced by natural selection. The term "function" is preferable to "adaptation", because adaptation is often construed as implying that it was selected for due to past function.

text: The comparison of complete gene sets for a group of organisms allow the identification of events in gene evolution such as gene duplication or gene deletion. The evolutionary relevance of this events is really big. For example, multiple duplications of genes encoding degradative enzymes of certain families is a common adaptation in microbes to new nutrient sources. On the contrary, loss of genes is important in reductive evolution, such as in intracellular parasites or symbionts. Whole genome duplication events, which potentially duplicate all the genes in a genome at once, are drastic evolutive events with great relevance in the evolution of many clades, and whose signal can be traced with phylogenomic methods.

text: The comparison of complete gene sets for a group of organisms allow the identification of events in gene evolution such as

text: gene duplication

text: or

text: gene deletion

text: . The evolutionary relevance of this events is really big. For example, multiple duplications of genes encoding degradative enzymes of certain families is a common adaptation in microbes to new nutrient sources. On the contrary, loss of genes is important in

text: reductive evolution

text: , such as in intracellular parasites or symbionts.

text: Whole genome duplication

text: events, which potentially duplicate all the genes in a genome at once, are drastic evolutive events with great relevance in the evolution of many clades, and whose signal can be traced with phylogenomic methods.

text: Cdc14 is widely distributed, being found in most eukaryote kingdoms.  However, it is not found in all species based on searches of sequenced genomes.  One or more Cdc14 genes are found in alveolates, animals, fungi, trypanosomes, and lower plants.  However, Cdc14 genes have apparently been lost in some lineages, including higher plants, rhodophytes, and slime molds.  There is a fairly tight positive correlation between the presence of Cdc14 in a species and whether that species makes flagella or cilia. This may be related to the ancestral role of Cdc14.  Whether flagella-anchoring basal bodies or centrioles involved in mitosis appeared first during evolution has been debated, but one theory is that flagella evolved first as a motility and sensory organelle, and the basal body was later co-opted into a mitotic role.  The function of Cdc14 may have adapted to different functions during the evolution of those organelles.

text: Cdc14 is widely distributed, being found in most eukaryote kingdoms.  However, it is not found in all species based on searches of sequenced genomes.  One or more Cdc14 genes are found in alveolates, animals, fungi, trypanosomes, and lower plants.  However, Cdc14 genes have apparently been lost in some lineages, including higher plants, rhodophytes, and slime molds.  There is a fairly tight positive correlation between the presence of Cdc14 in a species and whether that species makes

text: flagella

text: or

text: cilia

text: . This may be related to the ancestral role of Cdc14.  Whether flagella-anchoring basal bodies or centrioles involved in mitosis appeared first during evolution has been debated, but one theory is that flagella evolved first as a motility and sensory organelle, and the basal body was later co-opted into a mitotic role.  The function of Cdc14 may have adapted to different functions during the evolution of those organelles.

text: The lancelets (from "lancet") — also known as amphioxi (singular, amphioxus) — comprise some 32 species of fish-like marine chordates in the order Amphioxiformes, with a global distribution in shallow temperate (as far north as Scotland) and tropical seas, usually found half-buried in sand. They are the modern representatives of the subphylum Cephalochordata, formerly thought to be the sister group of the craniates. In Asia, they are harvested commercially as food for humans and domesticated animals. They are an important object of study in zoology as they provide indications about the evolutionary origins of the vertebrates. Lancelets serve as an intriguing comparison point for tracing how vertebrates have evolved and adapted. Although lancelets split from vertebrates more than 520 million years ago, their genomes hold clues about evolution, particularly how vertebrates have employed old genes for new functions. They are regarded as similar to the archetypal vertebrate form.

text: The lancelets (from "lancet") — also known as amphioxi (singular, amphioxus) — comprise some 32 species of fish-like marine

text: chordates

text: in the order Amphioxiformes, with a global distribution in shallow

text: temperate

text: (as far north as

text: Scotland

text: ) and

text: tropical

text: sea

text: s, usually found half-buried in sand. They are the modern representatives of the subphylum

text: Cephalochordata

text: , formerly thought to be the sister group of the

text: craniate

text: s. In

text: Asia

text: , they are harvested commercially as

text: food

text: for

text: human

text: s and

text: domesticated animal

text: s. They are an important object of study in

text: zoology

text: as they provide indications about the evolutionary origins of the

text: vertebrate

text: s. Lancelets serve as an intriguing comparison point for tracing how vertebrates have evolved and adapted. Although lancelets split from vertebrates more than 520 million years ago, their genomes hold clues about evolution, particularly how vertebrates have employed old genes for new functions. They are regarded as similar to the archetypal vertebrate form.

text: The modern evolutionary synthesis is based on the concept that populations of organisms have significant genetic variation caused by mutation and by the recombination of genes during sexual reproduction. It defines evolution as the change in allelic frequencies within a population caused by genetic drift, gene flow between sub populations, and natural selection. Natural selection is emphasised as the most important mechanism of evolution; large changes are the result of the gradual accumulation of small changes over long periods of time.

text: Sewall Green Wright (December 21, 1889March 3, 1988) was an American geneticist known for his influential work on evolutionary theory and also for his work on path analysis. He was a founder of population genetics alongside Ronald Fisher and J.B.S. Haldane, which was a major step in the development of the modern evolutionary synthesis combining genetics with evolution. He discovered the inbreeding coefficient and methods of computing it in pedigree animals. He extended this work to populations, computing the amount of inbreeding between members of populations as a result of random genetic drift, and along with Fisher he pioneered methods for computing the distribution of gene frequencies among  populations as a result of the interaction of natural selection, mutation, migration and genetic drift.  Wright also made major contributions to mammalian and biochemical genetics.

text: Sewall Green Wright (December 21, 1889March 3, 1988) was an American

text: geneticist

text: known for his influential work on

text: evolutionary theory

text: and also for his work on

text: path analysis

text: . He was a founder of

text: population genetics

text: alongside

text: Ronald Fisher

text: and

text: J.B.S. Haldane

text: , which was a major step in the development of the

text: modern evolutionary synthesis

text: combining

text: genetics

text: with

text: evolution

text: . He discovered the

text: inbreeding coefficient

text: and methods of computing it in

text: pedigree animals

text: . He extended this work to

text: population

text: s, computing the amount of

text: inbreeding

text: between members of populations as a result of random

text: genetic drift

text: , and along with Fisher he pioneered methods for computing the distribution of

text: gene frequencies

text: among  populations as a result of the interaction of

text: natural selection

text: mutation

text: , migration and genetic drift.  Wright also made major contributions to

text: mammal

text: ian and biochemical genetics.

text: Phenotypes are often Gaussian distributed in a large population and a necessary condition for the natural evolution to be able to fulfill the theorem of Gaussian adaptation, with respect to all Gaussian quantitative characters, is that it may push the centre of gravity of the Gaussian to the centre of gravity of the selected individuals. This may be accomplished by the Hardy–Weinberg law. This is possible because the theorem of Gaussian adaptation is valid for any region of acceptability independent of the structure (Kjellström, 1996).

text: Phenotypes are often Gaussian distributed in a large population and a necessary condition for the natural evolution to be able to fulfill the theorem of Gaussian adaptation, with respect to all Gaussian quantitative characters, is that it may push the centre of gravity of the Gaussian to the centre of gravity of the selected individuals. This may be accomplished by the

text: Hardy–Weinberg law

text: . This is possible because the theorem of Gaussian adaptation is valid for any region of acceptability independent of the structure (Kjellström, 1996).

text: John H. Gillespie is an evolutionary biologist interested in theoretical population genetics and molecular evolution. In molecular evolution, he emphasized the importance of advantageous mutations and balancing selection. For that reason, Gillespie is well known for his selectionist stance in the neutralist-selectionist debate. He is widely considered the main proponent of natural selection in molecular evolution.  He had a well-known feud with the father of the neutral theory of molecular evolution, Motoo Kimura, initiated by a review in Science of Kimura's book in which Gillespie criticized Kimura for "using the book as a vehicle to establish for himself a niche in the history of science." Gillespie had only four PhD students during his career, Richard Hudson, James N. McNair, David Cutler, and Andrew Kern. Gillespie was a professor in the College of Biological Sciences at the University of California, Davis until his retirement in 2005.

text: John H. Gillespie is an

text: evolutionary biologist

text: interested in

text: theoretical population genetics

text: and

text: molecular evolution

text: . In molecular evolution, he emphasized the importance of advantageous mutations and

text: balancing selection

text: . For that reason, Gillespie is well known for his

text: selectionist

text: stance in the

text: neutralist-selectionist debate

text: . He is widely considered the main proponent of

text: natural selection

text: in

text: molecular evolution

text: .  He had a well-known feud with the father of the

text: neutral theory of molecular evolution

text: Motoo Kimura

text: , initiated by a review in

text: Science

text: of

text: Kimura

text: 's book in which Gillespie criticized Kimura for "using the book as a vehicle to establish for himself a niche in the history of science." Gillespie had only four PhD students during his career, Richard Hudson, James N. McNair, David Cutler, and Andrew Kern. Gillespie was a professor in the College of Biological Sciences at the

text: University of California, Davis

text: until his retirement in 2005.

text: While sociocultural evolutionists agree that an evolution-like process leads to social progress, classical social evolutionists have developed many different theories, known as theories of unilineal evolution. Sociocultural evolutionism became the prevailing theory of early sociocultural anthropology and social commentary, and is associated with scholars like Auguste Comte, Edward Burnett Tylor, Lewis Henry Morgan, Benjamin Kidd, L. T. Hobhouse and Herbert Spencer. Sociocultural evolutionism attempted to formalise social thinking along scientific lines, with the added influence from the biological theory of evolution. If organisms could develop over time according to discernible, deterministic laws, then it seemed reasonable that societies could as well. Human society was compared to a biological organism, and social science equivalents of concepts like  variation, natural selection, and inheritance were introduced as factors resulting in the progress of societies. The idea of progress led to that of a fixed "stages" through which human societies progress, usually numbering threesavagery, barbarism, and civilizationbut sometimes many more. As early as the late 18th century, the Marquis de Condorcet (1743–1794) listed ten stages, or "epochs", each advancing the rights of man and perfecting the human race. At that time, anthropology was rising as a new scientific discipline, separating from the traditional views of "primitive" cultures that was usually based on religious views.

text: While sociocultural evolutionists agree that an evolution-like process leads to social progress, classical social evolutionists have developed many different theories, known as theories of unilineal evolution. Sociocultural evolutionism became the prevailing theory of early sociocultural anthropology and

text: social commentary

text: , and is associated with scholars like

text: Auguste Comte

text: Edward Burnett Tylor

text: Lewis Henry Morgan

text: Benjamin Kidd

text: L. T. Hobhouse

text: and

text: Herbert Spencer

text: . Sociocultural evolutionism attempted to formalise social thinking along scientific lines, with the added influence from the biological theory of

text: evolution

text: . If organisms could develop over time according to discernible, deterministic laws, then it seemed reasonable that societies could as well. Human society was compared to a biological organism, and social science equivalents of concepts like

text: variation

text: natural selection

text: , and

text: inheritance

text: were introduced as factors resulting in the progress of societies. The idea of progress led to that of a fixed "stages" through which human societies progress, usually numbering threesavagery, barbarism, and civilizationbut sometimes many more. As early as the late 18th century, the

text: Marquis de Condorcet

text: (1743–1794) listed ten stages, or "epochs", each advancing the rights of man and perfecting the human race. At that time, anthropology was rising as a new scientific discipline, separating from the traditional views of "primitive" cultures that was usually based on religious views.

text: The shifting balance theory is a theory of evolution proposed in 1932 by Sewall Wright, suggesting that adaptive evolution may proceed most quickly when a population divides into subpopulations with restricted gene flow.  The name of the theory is borrowed from Wright's metaphor of fitness landscapes (evolutionary landscapes), attempting to explain how a population may move across an adaptive valley to a higher adaptive peak.  According to the theory, this movement occurs in three steps:

text: The shifting balance theory is a theory of

text: evolution

text: proposed in 1932 by

text: Sewall Wright

text: , suggesting that adaptive evolution may proceed most quickly when a

text: population

text: divides into

text: subpopulation

text: s with restricted

text: gene flow

text: .  The name of the theory is borrowed from Wright's metaphor of

text: fitness landscapes

text: evolutionary landscape

text: s), attempting to explain how a population may move across an

text: adaptive valley

text: to a higher

text: adaptive peak

text: .  According to the theory, this movement occurs in three steps:

text: Gojobori has worked extensively on the rates of synonymous and non-synonymous substitutions, positive selection, horizontal gene transfer, viral evolution, genome evolution, and comparative gene expression. In recent years, he has focused on the evolution of the brain and Central nervous system.

text: Gojobori has worked extensively on the rates of synonymous and non-

text: synonymous substitutions

text: positive selection

text: horizontal gene transfer

text: viral evolution

text: genome evolution

text: , and comparative

text: gene expression

text: . In recent years, he has focused on the evolution of the brain and

text: Central nervous system

text: The underlying molecular evolution of high-altitude adaptation has been explored and understood fairly recently. Depending on the geographical and environmental pressures, high-altitude adaptation involves different genetic patterns. At the turn of the 21st century, it was reported that the genetic make-up of the respiratory components of the Tibetan and the Ethiopian populations are significantly different.

text: The underlying

text: molecular evolution

text: of high-altitude adaptation has been explored and understood fairly recently. Depending on the geographical and environmental pressures, high-altitude adaptation involves different genetic patterns. At the turn of the 21st century, it was reported that the genetic make-up of the respiratory components of the Tibetan and the Ethiopian populations are significantly different.

text: The mediocrity principle is the philosophical notion that "if an item is drawn at random from one of several sets or categories, it's likelier to come from the most numerous category than from any one of the less numerous categories". The principle has been taken to suggest that there is nothing very unusual about the evolution of the Solar System, Earth's history, the evolution of biological complexity, human evolution, or any one nation. It is a heuristic in the vein of the Copernican principle, and is sometimes used as a philosophical statement about the place of humanity. The idea is to assume mediocrity, rather than starting with the assumption that a phenomenon is special, privileged, exceptional, or even superior.

text: The mediocrity principle is the

text: philosophical

text: notion that "if an item is drawn at random from one of several sets or categories, it's likelier to come from the most numerous category than from any one of the less numerous categories". The principle has been taken to suggest that there is nothing very unusual about the

entity_name: Formation and evolution of the Solar System

text: evolution of the Solar System

text: Earth's history

text: , the

text: evolution of biological complexity

text: human evolution

text: , or any one

text: nation

text: . It is a

text: heuristic

text: in the vein of the

text: Copernican principle

text: , and is sometimes used as a philosophical statement about the place of humanity. The idea is to assume mediocrity, rather than starting with the assumption that a phenomenon is special, privileged, exceptional, or even

text: superior

text: Understanding the changes that have occurred during an organism's evolution can reveal the genes needed to construct parts of the body, genes which may be involved in human genetic disorders. For example, the Mexican tetra is an albino cavefish that lost its eyesight during evolution. Breeding together different populations of this blind fish produced some offspring with functional eyes, since different mutations had occurred in the isolated populations that had evolved in different caves. This helped identify genes required for vision and pigmentation.

text: Understanding the changes that have occurred during an organism's evolution can reveal the genes needed to construct parts of the body, genes which may be involved in human

text: genetic disorder

text: s. For example, the

text: Mexican tetra

text: is an

text: albino

text: cavefish that lost its eyesight during evolution. Breeding together different populations of this blind fish produced some offspring with functional eyes, since different mutations had occurred in the isolated populations that had evolved in different caves. This helped identify genes required for vision and pigmentation.

text: P (peptidyl) site --P element --P53 --Palindrome --Panmictic --Paralogous genes --Paramecin --Parameter --Parental --Parental ditype --Parental generation --Parental imprinting --Parkinson's disease --Parthenogenesis --Partial digest --Partial diploid --Partial dominance --Pascal's triangle --Patent --Path diagram --Pathogen --Pattern formation --PCR --Pedigree --Penetrance --Peptide --Peptide bond --Peptidyl transferase --Permanent translocation heterozygosity --Permissive condition --PEST hypothesis --Petite mutation --Phage --Pharmacogenomics --Pharming (genetics) --Phe --Phenocopy --Phenotype --Phenotypic variance --Phenylketonuria --Pheromone --Phosphodiester bond --Photoreactivation --Phyletic evolution --Phyletic gradualism --Phylogenetic tree --Phylogenetics --Physical map --Piebald --Pilus --PKU --Plant breeding --Plasmid --Plastid --Pleiotropic mutation --Pleiotropy --Pluripotency --Point mutation --Poisson distribution --Poky mutation --Polar body --Polar granules --Polar mutation --Polarity --Polarity gene --Pollen grain --Poly(A) tail --Polyacrylamide --Polycistronic --Polycistronic mRNA --Polydactyly --Polyadenylation --Polygene --Polygenic disorder --Polymerase --Polymerize --Polymorphism --Polypeptide --Polyploid --Polyribosome --Polysaccharide --Polysome --Polytene chromosome --Population --Population genetics --Position effect --Positional cloning --Positive control --Pre-mRNA --Precancerous polyp --Predisposition --Preemptor stem --Prenatal diagnosis --Pribnow box --Primary oocyte --Primary spermatocyte --Primary structure --Primary transcript --Primase --Primer --Primosome --Prion --Probability --Probability theory --Processivity --Product of meiosis --Product rule --Proflavin --Prognosis --Prokaryote --Prokaryotic cell --Promoter --Pronucleus --Proofreader genes --Prophage --Prophase --Proplastid --Propositus --Prostate cancer --Protease --Protein --Protein engineering --Protein product --Proteome --Proteomics --Proteus syndrome --Proto-oncogene --Protoplast --Prototroph --Provirus --Pseudoallele --Pseudoautosomal gene --Pseudodominance --Pseudogene --Public policy --Punnett square --Purine --Purines --Pyrimidine --

text: P (peptidyl) site

text: P element

text: Palindrome

text: Panmictic

text: Paralogous genes

text: Paramecin

text: Parameter

text: Parental

text: Parental ditype

text: Parental generation

text: Parental imprinting

text: Parkinson's disease

text: Parthenogenesis

text: Partial digest

text: Partial diploid

text: Partial dominance

text: Pascal's triangle

text: Patent

text: Path diagram

text: Pathogen

text: Pattern formation

text: PCR

text: Pedigree

text: Penetrance

text: Peptide

text: Peptide bond

text: Peptidyl transferase

text: Permanent translocation heterozygosity

text: Permissive condition

text: PEST hypothesis

text: Petite mutation

text: Phage

text: Pharmacogenomics

text: Pharming (genetics)

text: Phe

text: Phenocopy

text: Phenotype

text: Phenotypic variance

text: Phenylketonuria

text: Pheromone

text: Phosphodiester bond

text: Photoreactivation

text: Phyletic evolution

text: Phyletic gradualism

text: Phylogenetic tree

text: Phylogenetics

text: Physical map

text: Piebald

text: Pilus

text: PKU

text: Plant breeding

text: Plasmid

text: Plastid

text: Pleiotropic mutation

text: Pleiotropy

text: Pluripotency

text: Point mutation

text: Poisson distribution

text: Poky mutation

text: Polar body

text: Polar granules

text: Polar mutation

text: Polarity

text: Polarity gene

text: Pollen grain

text: Poly(A) tail

text: Polyacrylamide

text: Polycistronic

text: Polycistronic mRNA

text: Polydactyly

text: Polyadenylation

text: Polygene

text: Polygenic disorder

text: Polymerase

text: Polymerize

text: Polymorphism

text: Polypeptide

text: Polyploid

text: Polyribosome

text: Polysaccharide

text: Polysome

text: Polytene chromosome

text: Population

text: Population genetics

text: Position effect

text: Positional cloning

text: Positive control

text: Pre-mRNA

text: Precancerous polyp

text: Predisposition

text: Preemptor stem

text: Prenatal diagnosis

text: Pribnow box

text: Primary oocyte

text: Primary spermatocyte

text: Primary structure

text: Primary transcript

text: Primase

text: Primer

text: Primosome

text: Prion

text: Probability

text: Probability theory

text: Processivity

text: Product of meiosis

text: Product rule

text: Proflavin

text: Prognosis

text: Prokaryote

text: Prokaryotic cell

text: Promoter

text: Pronucleus

text: Proofreader genes

text: Prophage

text: Prophase

text: Proplastid

text: Propositus

text: Prostate cancer

text: Protease

text: Protein

text: Protein engineering

text: Protein product

text: Proteome

text: Proteomics

text: Proteus syndrome

text: Proto-oncogene

text: Protoplast

text: Prototroph

text: Provirus

text: Pseudoallele

text: Pseudoautosomal gene

text: Pseudodominance

text: Pseudogene

text: Public policy

text: Punnett square

text: Purine

text: Purines

text: Pyrimidine

text: A common claim of creationists is that evolution has never been observed. Challenges to such objections often come down to debates over how evolution is defined (see the Defining evolution section above). Under the conventional biological definition of evolution, it is a simple matter to observe evolution occurring. Evolutionary processes, in the form of populations changing their genetic composition from generation to generation, have been observed in different scientific contexts, including the evolution of fruit flies, mice, and bacteria in the laboratory, and of tilapia in the field. Such studies on experimental evolution, particularly those using microorganisms, are now providing important insights into how evolution occurs, especially in the case of antibiotic resistance.

text: A common claim of creationists is that evolution has never been observed. Challenges to such objections often come down to debates over how evolution is defined (see the

text: Defining evolution

text: section above). Under the conventional biological definition of evolution, it is a simple matter to observe evolution occurring. Evolutionary processes, in the form of populations changing their genetic composition from generation to generation, have been observed in different scientific contexts, including the evolution of

text: fruit flies

text: mice

text: , and

text: bacteria

text: in the laboratory, and of

text: tilapia

text: in the field. Such studies on

text: experimental evolution

text: , particularly those using

text: microorganism

text: s, are now providing important insights into how evolution occurs, especially in the case of

text: antibiotic resistance

text: His first paper in 1915 demonstrated genetic linkage in mammals while subsequent works helped to create population genetics, thus establishing a unification of Mendelian genetics and Darwinian evolution by natural selection whilst laying the groundwork for modern evolutionary synthesis. His article on abiogenesis in 1929 introduced the "Primordial Soup Theory", and it became the foundation to build physical models for the chemical origin of life. Haldane established human gene maps for haemophilia and colour blindness on the X chromosome, and codified Haldane's rule on sterility in the heterogametic sex of hybrids in species. He correctly proposed that sickle-cell disease confers some immunity to malaria. He was the first to suggest the central idea of in vitro fertilisation, as well as concepts such as: the hydrogen economy, cis and trans-acting regulation, coupling reaction, molecular repulsion, darwin (as a unit of evolution) and organismal cloning. In 1957 he articulated Haldane's dilemma, a limit on the speed of beneficial evolution which subsequently proved incorrect. He willed his body for medical studies, as he wanted to remain useful even in death.

text: His first paper in 1915 demonstrated

text: genetic linkage

text: in

text: mammals

text: while subsequent works helped to create

text: population genetics

text: , thus establishing a unification of

text: Mendelian genetics

text: and Darwinian evolution by

text: natural selection

text: whilst laying the groundwork for

text: modern evolutionary synthesis

text: . His article on

text: abiogenesis

text: in 1929 introduced the "Primordial Soup Theory", and it became the foundation to build physical models for the

text: chemical origin of life

text: . Haldane established human

text: gene map

text: s for

text: haemophilia

text: and

text: colour blindness

text: on the

text: X chromosome

text: , and codified

text: Haldane's rule

text: on

text: sterility

text: in the

text: heterogametic sex

text: of

text: hybrids

text: in

text: species

text: . He correctly proposed that

text: sickle-cell disease

text: confers some immunity to

text: malaria

text: . He was the first to suggest the central idea of

text: in vitro fertilisation

text: , as well as concepts such as: the

text: hydrogen economy

text: , cis and

text: trans-acting

text: regulation,

text: coupling reaction

text: molecular repulsion

text: darwin

text: (as a unit of evolution) and

text: organismal cloning

text: . In 1957 he articulated

text: Haldane's dilemma

text: , a limit on the speed of beneficial

text: evolution

text: which subsequently proved incorrect. He willed his body for medical studies, as he wanted to remain useful even in death.

text: In a 2008 study, its authors stated that "There is considerable evidence that human sexual orientation is genetically influenced, so it is not known how homosexuality, which tends to lower reproductive success, is maintained in the population at a relatively high frequency." They hypothesized that "while genes predisposing to homosexuality reduce homosexuals' reproductive success, they may confer some advantage in heterosexuals who carry them," and their results suggested that "genes predisposing to homosexuality may confer a mating advantage in heterosexuals, which could help explain the evolution and maintenance of homosexuality in the population." However, in the same study, the authors noted that "nongenetic alternative explanations cannot be ruled out" as a reason for the heterosexual in the homosexual-heterosexual twin pair having more partners, specifically citing "socialpressure on the other twin to act in a more heterosexual way" (and thus seek out a greater number of sexual partners) as an example of one alternative explanation. Also, the authors of the study acknowledge that a large number of sexual partners may not lead to greater reproductive success, specifically noting there is an "absence of evidence relating the number of sexual partners and actual reproductive success, either in the present or in our evolutionary past."

text: Wen-Hsiung Li (Traditional Chinese:李文雄, 1942-) is a Taiwanese American scientist working in the fields of molecular evolution, population genetics, and genomics.  He is currently the James Watson Professor of Ecology and Evolution at the University of Chicago and a Principal Investigator at the Institute of Information Science and Genomics Research Center, Academia Sinica, Taiwan.

text: Wen-Hsiung Li (

text: Traditional Chinese

text: :李文雄, 1942-) is a

text: Taiwanese American

text: scientist working in the fields of

text: molecular evolution

text: population genetics

text: , and

text: genomics

text: .  He is currently the

text: James Watson

text: Professor of Ecology and Evolution at the

text: University of Chicago

text: and a Principal Investigator at the Institute of Information Science and Genomics Research Center,

text: Academia Sinica

text: Taiwan

text: On the other hand, the common definition of adaptation, a central concept in evolution, is a trait that was functional to the reproductive success of the organism and that is thus now present due to being selected for; that is, function and evolution are inseparable. However a trait can have a current function that is adaptive without being an adaptation in this sense, if for instance the environment has changed. Imagine an environment in which having a small body suddenly conferred benefit on an organism when previously body size had had no effect on survival.

text: One level of genome organization is the grouping of genes into several gene families. Gene families are groups of related genes that share a common ancestor. Members of gene families may be paralogs or orthologs. Gene paralogs are genes with similar sequences from within the same species while gene orthologs are genes with similar sequences in different species. Gene families are highly variable in size, sequence diversity, and arrangement. Depending on the diversity and functions of the genes within the family, families can be classified as a multigene families or superfamilies.

text: Estimates of the amount of adaptive evolution in non-coding DNA are generally very low, although fewer studies have been done on non-coding DNA. As with the coding DNA however, the methods currently used are relatively weak. Ponting and Lunter (2006) speculate that underestimates may be even more severe in non-coding DNA, because non-coding DNA may undergo periods of functionality (and adaptive evolution), followed by periods of neutrality. If this is true, current methods for detecting adaptive evolution are inadequate to account for such patterns. Additionally, even if low estimates of the amount of adaptive evolution are correct, this can still equate to a large amount of adaptively evolving non-coding DNA, since non-coding DNA makes up approximately 98% of the DNA in the human genome. For example, Ponting and Lunter (2006) detect a modest 0.03% of non-coding DNA showing evidence of adaptive evolution, but this still equates to approximately 1 Mb of adaptively evolving DNA. Where there is evidence of adaptive evolution (which implies functionality) in non-coding DNA, these regions are generally thought to be involved in the regulation of protein coding sequences.As with humans, fewer studies have searched for adaptive evolution in non-coding regions of other organisms. However, where research has been done on Drosophila, there appears to be large amounts of adaptively evolving non-coding DNA. Andolfatto (2005) estimated that adaptive evolution has occurred in 60% of untranslated mature portions of mRNAs, and in 20% of intronic and intergenic regions. If this is true, this would imply that much non-coding DNA could be of more functional importance than coding DNA, dramatically altering the consensus view. However, this would still leave unanswered what function all this non-coding DNA performs, as the regulatory activity observed thus far is in just a tiny proportion of the total amount of non-coding DNA. Ultimately, significantly more evidence needs to be gathered to substantiate this viewpoint.

text: Estimates of the amount of adaptive evolution in

text: non-coding DNA

text: Regarding social evolution, it has been suggested that social anagenesis/aromorphosis be viewed as  universal or widely diffused social innovation that raises social systems' complexity, adaptability, integrity, and interconnectedness.

text: Regarding

text: social evolution

text: , it has been suggested that social anagenesis/aromorphosis be viewed as  universal or widely diffused social innovation that raises social systems' complexity, adaptability, integrity, and interconnectedness.

text: The word evolution in a broad sense refers to processes of change, from stellar evolution to changes in language. In biology, the meaning is more specific: heritable changes which accumulate over generations of a population. Individual organisms do not evolve in their lifetimes, but variations in the genes they inherit can become more or less common in the population of organisms. Any changes during the lifetime of organisms which are not inherited by their offspring are not part of biological evolution.

text: The genome of an organism contains anywhere from a few hundred to thousands of different genes, all encoding a singular product or more. For numerous reasons, including organizational maintenance, energy conservation, and generating phenotypic variance, it is important that genes are only expressed when they are needed. The most efficient way for an organism to regulate genetic expression is at the transcriptional level. CREs function to control transcription by acting nearby or within a gene. The most well characterized types of CREs are enhancers and promoters. Both of these sequence elements are structural regions of DNA that serve as transcriptional regulators.

text: The

text: genome

text: of an organism contains anywhere from a few hundred to thousands of different genes, all encoding a singular product or more. For numerous reasons, including organizational maintenance, energy conservation, and generating

text: phenotypic

text: variance, it is important that genes are only expressed when they are needed. The most efficient way for an organism to

text: regulate genetic expression

text: is at the transcriptional level. CREs function to control transcription by acting nearby or within a gene. The most well characterized types of CREs are

text: enhancer

text: s and

text: promoters

text: . Both of these sequence elements are structural regions of DNA that serve as

text: transcriptional regulators

text: New beneficial mutations create fitter genotypes than those previously present in the population. When load is calculated as the difference between the fittest genotype present and the average, this creates a substitutional load. The difference between the theoretical maximum (which may not actually be present) and the average is known as the "lag load". Motoo Kimura's original argument for the neutral theory of molecular evolution was that if most differences between species were adaptive, this would exceed the speed limit to adaptation set by the substitutional load. However, Kimura's argument confused the lag load with the substitutional load, using the former when it is the latter that in fact sets the maximal rate of evolution by natural selection.

text: New beneficial mutations create fitter genotypes than those previously present in the population. When load is calculated as the difference between the fittest genotype present and the average, this creates a

text: substitutional load

text: . The difference between the theoretical maximum (which may not actually be present) and the average is known as the "lag load".

text: Motoo Kimura

text: 's original argument for the

text: neutral theory of molecular evolution

text: was that if most differences between species were adaptive, this would exceed the speed limit to adaptation set by the substitutional load. However, Kimura's argument confused the lag load with the substitutional load, using the former when it is the latter that in fact sets the maximal rate of evolution by natural selection.

text: The evolution of the number of inhabitants is known through the population censuses carried out in the town since 1793 and the research study of Louis-Edmond Marissal, Clerk of the Peace of the city, published in 1844. From the 21st century, communes with more than 10,000 population have sample surveys held every year, unlike other municipalities that have a real census every five years.<center>Evolution of the Roubaisian population in the 19th century

text: The evolution of the number of inhabitants is known through the population censuses carried out in the town since 1793 and the research study of Louis-Edmond Marissal, Clerk of the Peace of the city, published in 1844. From the 21st century, communes with more than 10,000 population have

text: sample surveys

text: held every year, unlike other municipalities that have a real census every five years.<center>Evolution of the Roubaisian population in the 19th century

text: There were serious scientific objections to the process of natural selection as the key mechanism of evolution, including Karl von Nägeli's insistence that a trivial characteristic with no adaptive advantage could not be developed by selection. Darwin conceded that these could be linked to adaptive characteristics. His estimate that the age of the Earth allowed gradual evolution was disputed by William Thomson (later awarded the title Lord Kelvin), who calculated that it had cooled in less than 100 million years. Darwin accepted blending inheritance, but Fleeming Jenkin calculated that as it mixed traits, natural selection could not accumulate useful traits. Darwin tried to meet these objections in the 5th edition. Mivart supported directed evolution, and compiled scientific and religious objections to natural selection. In response, Darwin made considerable changes to the sixth edition. The problems of the age of the Earth and heredity were only resolved in the 20th century.

text: There were serious scientific objections to the process of

text: natural selection

text: as the key mechanism of evolution, including

text: Karl von Nägeli

text: 's insistence that a trivial characteristic with no adaptive advantage could not be developed by selection. Darwin conceded that these could be linked to adaptive characteristics. His estimate that the

text: age of the Earth

text: allowed gradual evolution was disputed by

text: William Thomson (later awarded the title Lord Kelvin)

text: , who calculated that it had cooled in less than 100 million years. Darwin accepted

text: blending inheritance

text: , but

text: Fleeming Jenkin

text: calculated that as it mixed traits, natural selection could not accumulate useful traits. Darwin tried to meet these objections in the 5th edition.

text: Mivart

text: supported directed evolution, and compiled scientific and religious objections to natural selection. In response, Darwin made considerable changes to the sixth edition. The problems of the age of the Earth and heredity were only resolved in the 20th century.

text: The evolution of biological complexity is one important outcome of the process of evolution. Evolution has produced some remarkably complex organisms - although the actual level of complexity is very hard to define or measure accurately in biology, with properties such as gene content, the number of cell types or morphology all being used to assess an organism's complexity.

text: The evolution of biological complexity is one important outcome of the process of

text: evolution

text: . Evolution has produced some remarkably complex organisms - although the actual level of complexity is very hard to define or measure accurately in biology, with properties such as gene content, the number of

text: cell type

text: s or

text: morphology

text: all being used to assess an organism's complexity.

text: The "Continuity Hypothesis" (CH) provides an ontological basis for evolutionary economics.  Witt’s concept is influenced by Darwinian theories insofar as it stresses the importance of cognitive and social dispositions acquired by early humans during natural, biological evolution.  Innate dispositions of this kind are seen as substantially influencing human behavior in an economic context. However, CH places emphasis on the fact that evolutionary processes driving the economy rely on mechanisms different from those underlying biological evolution. The form of evolution relevant for economics is cultural evolution, which, in Witt’s view, is Non-Darwinian and cannot be explained by abstract “Darwinian principles”. Cultural evolution is based on humans’ cognitive apparatus as an outcome of biological evolution, but is governed by distinct and idiosyncratic mechanisms. Therefore, Witt is skeptical about the use of biological analogies in explaining the evolution of the economy. Important Non-Darwinian mechanisms include individual and social learning, the diffusion of knowledge, and the creation of variance through human creativity and imagination. These are the driving forces of socio-economic dynamics that manifest themselves in a process of self-organization. In Witt’s opinion, the role of natural selection in cultural evolution becomes negligible due to the immense increase in pace of evolutionary change and the huge reproductive success of the human species - supported by the evolution of cultural institutions. Thus, the “connection between natural and man-made evolution resolves into an ... indirect one” (Witt 2003, p. 17). In the light of CH, long term economic change is no longer mainly an optimization problem of economizing on scarce resources. Rather, economic evolution today is driven by cumulative knowledge generation and problem solving, guided by learning, insight and intentionality as well as the successive emergence of new problems. In this respect, Witt sees his CH in the tradition of Thorstein Veblen, while it is “somewhat different from, but not incompatible with, the neo-Schumpeterian approach” (Witt 2003, p. 18).

text: The "Continuity Hypothesis" (CH) provides an ontological basis for evolutionary economics.  Witt’s concept is influenced by

text: Darwinian

text: theories insofar as it stresses the importance of cognitive and social dispositions acquired by early humans during natural, biological

text: evolution

text: .  Innate dispositions of this kind are seen as substantially influencing human behavior in an economic context. However, CH places emphasis on the fact that evolutionary processes driving the economy rely on mechanisms different from those underlying biological evolution. The form of evolution relevant for economics is

text: cultural evolution

text: , which, in Witt’s view, is Non-Darwinian and cannot be explained by abstract “Darwinian principles”. Cultural evolution is based on humans’ cognitive apparatus as an outcome of biological evolution, but is governed by distinct and idiosyncratic mechanisms. Therefore, Witt is skeptical about the use of biological analogies in explaining the evolution of the economy. Important Non-Darwinian mechanisms include individual and social learning, the diffusion of knowledge, and the creation of variance through human creativity and imagination. These are the driving forces of socio-economic dynamics that manifest themselves in a process of self-organization. In Witt’s opinion, the role of natural selection in cultural evolution becomes negligible due to the immense increase in pace of evolutionary change and the huge reproductive success of the human species - supported by the evolution of cultural institutions. Thus, the “connection between natural and man-made evolution resolves into an ... indirect one” (Witt 2003, p. 17). In the light of CH, long term economic change is no longer mainly an optimization problem of economizing on scarce resources. Rather, economic evolution today is driven by cumulative knowledge generation and problem solving, guided by learning, insight and intentionality as well as the successive emergence of new problems. In this respect, Witt sees his CH in the tradition of

text: Thorstein Veblen

text: , while it is “somewhat different from, but not incompatible with, the neo-Schumpeterian approach” (Witt 2003, p. 18).

text: Studies of the genetic influences on geographic range limits tend to focus on genetic effects that restrict peripheral populations from expanding beyond their range edge. Many hypotheses have been proposed, however there has not been a general consensus on one main cause. Most hypotheses stem from the fact that peripheral populations are expected to be small, due to less optimal conditions at the edge. One hypothesis predicts that due to smaller population size at the edge, genetic drift will have a larger effect and reduce genetic variation of edge populations. With less genetic variation, populations at the edge may have difficulty adapting to changing conditions. Another hypothesis predicts that high selection pressure at the peripheral edge will only allow for a few genotypes to survive, decreasing genetic variation and future adaptation. Reduced gene flow between central and peripheral populations has also been hypothesized as a limitation for species expansion. Without introduction of new alleles from central to edge populations, genetic variation will remain low and peripheral populations will have difficulty adapting to new conditions. Contrary to this, some hypothesize that gene flow between central and peripheral populations can prevent range expansion, where gene flow causes contamination of the edge gene pool. Conditions at the center of the range differ from those at the periphery; therefore adapted alleles at the center may not benefit those at the edge who experience different conditions. The Allee effect has also been proposed to reduce range expansion, where small population sizes lead to reduced cooperation and survival. There are many potential causes of the allee effect, including mate-finding failure or inbreeding. At low densities, the challenge for males and females to locate each other is much larger, leading to decreased reproduction and survival. Small population sizes may also cause species to mate with their relatives, increasing chances of lethal alleles within the population and decreasing survival.

text: Studies of the

text: genetic

text: influences on geographic range limits tend to focus on genetic effects that restrict peripheral populations from expanding beyond their range edge. Many

text: hypotheses

text: have been proposed, however there has not been a general consensus on one main cause. Most hypotheses stem from the fact that peripheral populations are expected to be small, due to less optimal conditions at the edge. One hypothesis predicts that due to smaller

text: population size

text: at the edge,

text: genetic drift

text: will have a larger effect and reduce

text: genetic variation

text: of edge populations. With less genetic variation, populations at the edge may have difficulty adapting to changing conditions. Another hypothesis predicts that high

text: selection pressure

text: at the peripheral edge will only allow for a few

text: genotypes

text: to survive, decreasing genetic variation and future adaptation. Reduced

text: gene flow

text: between central and peripheral populations has also been hypothesized as a limitation for species expansion. Without introduction of new

text: alleles

text: from central to edge populations, genetic variation will remain low and peripheral populations will have difficulty adapting to new conditions. Contrary to this, some hypothesize that gene flow between central and peripheral populations can prevent range expansion, where gene flow causes contamination of the edge

text: gene pool

text: . Conditions at the center of the range differ from those at the periphery; therefore adapted alleles at the center may not benefit those at the edge who experience different conditions. The

text: Allee effect

text: has also been proposed to reduce range expansion, where small population sizes lead to reduced

text: cooperation

text: and survival. There are many potential causes of the allee effect, including mate-finding failure or

text: inbreeding

text: . At low densities, the challenge for

text: males

text: and

text: females

text: to locate each other is much larger, leading to decreased reproduction and survival. Small population sizes may also cause species to mate with their relatives, increasing chances of

text: lethal alleles

text: within the population and decreasing survival.

text: Different organisms house different numbers of genes within their genomes as well as different patterns in the distribution of genes throughout the genome.  Some organisms, such as most bacteria, Drosophila, and Arabidopsis have particularly compact genomes with little repetitive content or non-coding DNA.  Other organisms, like mammals or maize, have large amounts of repetitive DNA, long introns, and substantial spacing between different genes.  The content and distribution of genes within the genome can influence the rate at which certain types of mutations occur and can influence the subsequent evolution of different species.  Genes with longer introns are more likely to recombine due to increased physical distance over the coding sequence.  As such, long introns may facilitate ectopic recombination, and result in higher rates of new gene formation.

text: Different organisms house different numbers of genes within their genomes as well as different patterns in the distribution of genes throughout the genome.  Some organisms, such as most bacteria, Drosophila, and Arabidopsis have particularly compact genomes with little repetitive content or non-coding DNA.  Other organisms, like mammals or maize, have large amounts of repetitive DNA, long introns, and substantial spacing between different genes.  The content and distribution of genes within the genome can influence the rate at which certain types of mutations occur and can influence the subsequent evolution of different species.  Genes with longer

text: introns

text: are more likely to recombine due to increased physical distance over the coding sequence.  As such, long introns may facilitate

text: ectopic recombination

text: , and result in higher rates of new gene formation.

text: Genes are the primary units of inheritance in all organisms. A gene is a unit of heredity and corresponds to a region of DNA that influences the form or function of an organism in specific ways. All organisms, from bacteria to animals, share the same basic machinery that copies and translates DNA into proteins. Cells transcribe a DNA gene into an RNA version of the gene, and a ribosome then translates the RNA into a protein, a sequence of amino acids. The translation code from RNA codon to amino acid is the same for most organisms, but slightly different for some. For example, a sequence of DNA that codes for insulin in humans also codes for insulin when inserted into other organisms, such as plants.

text: Gene

text: s are the primary units of inheritance in all organisms. A gene is a unit of

text: heredity

text: and corresponds to a region of

text: DNA

text: that influences the form or function of an organism in specific ways. All organisms, from bacteria to animals, share the same basic machinery that copies and translates DNA into

text: protein

text: s. Cells

text: transcribe

text: a DNA gene into an

text: RNA

text: version of the gene, and a

text: ribosome

text: then

text: translates

text: the RNA into a protein, a sequence of

text: amino acid

text: s. The

text: translation code

text: from RNA codon to amino acid is the same for most organisms, but slightly different for some. For example, a sequence of DNA that codes for

text: insulin

text: in humans also codes for insulin when inserted into other organisms, such as plants.

text: He was the commentator for the first time in the Pro Evolution Soccer series from Pro Evolution Soccer 2008 to Pro Evolution Soccer 2015. He worked alongside with analyst Mark Lawrenson from Pro Evolution Soccer 2008 to Pro Evolution Soccer 2010. From Pro Evolution Soccer 2011 to Pro Evolution Soccer 2015, he worked together with analyst from ITV, Jim Beglin. He was then replaced by Peter Drury as the commentator for Pro Evolution Soccer 2016.

text: He was the commentator for the first time in the

text: Pro Evolution Soccer

text: series from

text: Pro Evolution Soccer 2008

text: to

text: Pro Evolution Soccer 2015

text: . He worked alongside with analyst

text: Mark Lawrenson

text: from

text: Pro Evolution Soccer 2008

text: to

text: Pro Evolution Soccer 2010

text: . From

text: Pro Evolution Soccer 2011

text: to

text: Pro Evolution Soccer 2015

text: , he worked together with analyst from ITV,

text: Jim Beglin

text: . He was then replaced by

text: Peter Drury

text: as the commentator for

text: Pro Evolution Soccer 2016

text: Following the establishment of evolutionary biology, studies of mutation and genetic diversity in natural populations, combined with biogeography and systematics, led to sophisticated mathematical and causal models of evolution. Paleontology and comparative anatomy allowed more detailed reconstructions of the evolutionary history of life. After the rise of molecular genetics in the 1950s, the field of molecular evolution developed, based on protein sequences and immunological tests, and later incorporating RNA and DNA studies. The gene-centered view of evolution rose to prominence in the 1960s, followed by the neutral theory of molecular evolution, sparking debates over adaptationism, the unit of selection, and the relative importance of genetic drift versus natural selection as causes of evolution. In the late 20th-century, DNA sequencing led to molecular phylogenetics and the reorganization of the tree of life into the three-domain system by Carl Woese. In addition, the newly recognized factors of symbiogenesis and horizontal gene transfer introduced yet more complexity into evolutionary theory. Discoveries in evolutionary biology have made a significant impact not just within the traditional branches of biology, but also in other academic disciplines (for example: anthropology and psychology) and on society at large.<ref name="Futuyma">

text: Following the establishment of

text: evolutionary biology

text: , studies of mutation and

text: genetic diversity

text: in natural populations, combined with biogeography and

text: systematics

text: , led to sophisticated mathematical and causal models of evolution. Paleontology and

text: comparative anatomy

text: allowed more detailed reconstructions of the

text: evolutionary history of life

text: . After the rise of

text: molecular genetics

text: in the 1950s, the field of

text: molecular evolution

text: developed, based on

text: protein sequences

text: and immunological tests, and later incorporating

text: RNA

text: and

text: DNA

text: studies. The

text: gene-centered view of evolution

text: rose to prominence in the 1960s, followed by the

text: neutral theory of molecular evolution

text: , sparking debates over

text: adaptationism

text: , the

text: unit of selection

text: , and the relative importance of

text: genetic drift

text: versus natural selection as causes of evolution. In the late 20th-century, DNA sequencing led to

text: molecular phylogenetics

text: and the reorganization of the tree of life into the

text: three-domain system

text: by

text: Carl Woese

text: . In addition, the newly recognized factors of

text: symbiogenesis

text: and

text: horizontal gene transfer

text: introduced yet more complexity into evolutionary theory. Discoveries in evolutionary biology have made a significant impact not just within the traditional branches of biology, but also in other academic disciplines (for example:

text: anthropology

text: and

text: psychology

text: ) and on society at large.<ref name="Futuyma">

text: The general theory of adaptive biological systems, named practopoiesis (meaning creation of actions), has been used to derive a theory that explains mental operations as an adaptive process. Much like species adapt through evolution and an organism adapts through development, the theory of anapoiesis (meaning re-creation) proposes that a thought is a process of adaptation to the immediate environment. This is performed by fast physiological machinery that can operate within a few 100s of milliseconds and relies on the mechanisms of neural adaptation. A key difference between anapoietic approach and the functional approach is that for anapoietic process much of the information needed for the mental operations is located outside the organism. If mental operations are an adaptive process, they do not juggle symbols internally (like a computer) but make guesses of what changes should be made to the nervous system and then test them against the environment.

text: The general theory of adaptive biological systems, named practopoiesis (meaning creation of actions), has been used to derive a theory that explains mental operations as an adaptive process. Much like species adapt through evolution and an organism adapts through development, the theory of anapoiesis (meaning re-creation) proposes that a thought is a process of adaptation to the immediate environment. This is performed by fast physiological machinery that can operate within a few 100s of milliseconds and relies on the mechanisms of

text: neural adaptation

text: . A key difference between anapoietic approach and the functional approach is that for anapoietic process much of the information needed for the mental operations is located outside the organism. If mental operations are an adaptive process, they do not juggle symbols internally (like a computer) but make guesses of what changes should be made to the nervous system and then test them against the environment.

text: Recent theoretical thought has developed a modified definition of cultural evolution, one that claims that the processes that give rise to cultural stability and change are parallel to those of biological evolution (Shennan 2008).   Culture evolution in this respect refers to the “changing distributions of cultural attributes in populations resulting from natural selection as well as by other process that are not related to genetic evolution” (Shennan, 2008).  In other words, it is characterized as a “Darwinian process comprising the selective retention of favorable culturally transmitted variants, as well as a variety of non-selective processes, such as drift” (Boyd & Richerson 1985; Cavalli-Sforza & Feldman 1981). Mesoudi and colleagues have explained cultural entities and processes using original Darwinian formulations (Mesoudi et al, 2006) However, critics debate to the extent to which cultural process can be explained by evolutionary models.

text: In 1944, Simpson expanded Wright's landscape to include phenotypes. in Simpson's model, the landscape was a means of visualizing the "relationship between selection, structure, and adaptation."  Unlike Wright, Simpson used the landscape to represent both natural selection and genetic drift. Uphill movements are due to positive selection, and downhill movements are due to negative selection. The size and shape of a peak indicated the relative specificity of selection; i.e. a sharp and high peak indicates highly specific selection. Another difference between Simpson's and Wright's landscapes is the level at which evolution is acting. For Wright, a population geneticist, only populations of a species were shown. In Simpson's figures, the circles drawn represent all of Equidae. The most important difference is that in Simpson's model, the landscape could vary through time, while in Wright's model, the landscape was static. It is interesting to note that Wright reviewed Simpson's work (Tempo and Mode in Evolution) and did not object to Simpson's use of evolutionary landscapes. In later writings, Simpson referred to peaks as adaptive zones. In a series of papers Russell Lande developed a mathematical model for Simpson's phenotypic landscape. Lande reconciled Wright's population level view with Simpson's use of higher taxonomic levels. Lande considers fitness peaks to be determined by the environment and thus represent ecological niches or adaptive zones for a population. Clusters of peaks inhabited by phenotypically similar populations can be viewed as higher taxonomic levels.

text: In 1944, Simpson expanded Wright's landscape to include phenotypes. in Simpson's model, the landscape was a means of visualizing the "relationship between selection, structure, and adaptation."  Unlike Wright, Simpson used the landscape to represent both natural selection and genetic drift. Uphill movements are due to

text: positive selection

text: , and downhill movements are due to

text: negative selection

text: . The size and shape of a peak indicated the relative specificity of selection; i.e. a sharp and high peak indicates highly specific selection. Another difference between Simpson's and Wright's landscapes is the level at which evolution is acting. For Wright, a

text: population geneticist

text: , only populations of a species were shown. In Simpson's figures, the circles drawn represent all of Equidae. The most important difference is that in Simpson's model, the landscape could vary through time, while in Wright's model, the landscape was static. It is interesting to note that Wright reviewed Simpson's work (Tempo and Mode in Evolution) and did not object to Simpson's use of evolutionary landscapes. In later writings, Simpson referred to peaks as adaptive zones. In a series of papers

text: Russell Lande

text: developed a mathematical model for Simpson's phenotypic landscape. Lande reconciled Wright's population level view with Simpson's use of higher taxonomic levels. Lande considers fitness peaks to be determined by the environment and thus represent ecological niches or adaptive zones for a population. Clusters of peaks inhabited by phenotypically similar populations can be viewed as higher taxonomic levels.

text: In population genetics and evolution, genetic memory represents the recorded history of adaptive changes in a species. Selection of organisms carrying genes coding for the best adapted proteins results in the evolution of species. An example for such a genetic memory is the innate immune response that represents a recording of the history of common microbial and viral pathogens encountered throughout the evolutionary history of the species. In contrast to the somatic memory of the adaptive immune response, the innate immune response is present at birth and does not require the immune system to learn to recognize antigens.

text: In

text: population genetics

text: and

text: evolution

text: , genetic memory represents the recorded history of adaptive changes in a species. Selection of organisms carrying genes coding for the best adapted proteins results in the

text: evolution

text: of species. An example for such a genetic memory is the

text: innate immune response

text: that represents a recording of the history of common microbial and viral pathogens encountered throughout the evolutionary history of the species. In contrast to the somatic memory of the adaptive immune response, the innate immune response is present at birth and does not require the immune system to learn to recognize antigens.

text: According to the modern evolutionary synthesis, adaptations are produced incrementally through natural selection, so major genetic changes will usually spread within populations only if they do not decrease the fitness of individuals. Although neutral changes (ones which provide no benefit) can spread through genetic drift, and detrimental changes can spread under some circumstances, large changes that require multiple steps will occur only if the intermediate stages increase fitness. Richard Dawkins describes the matter: "The wheel may be one of those cases where the engineering solution can be seen in plain view, yet be unattainable in evolution because it lies [on] the other side of a deep valley, cutting unbridgeably across the massif of Mount Improbable." In such a fitness landscape, wheels might sit on a highly favorable "peak", but the valley around that peak may be too deep or wide for the gene pool to migrate across by genetic drift or natural selection. Stephen Jay Gould notes that biological adaptation is limited to working with available components, commenting that "wheels work well, but animals are debarred from building them by structural constraints inherited as an evolutionary legacy."

text: According to the

text: modern evolutionary synthesis

text: , adaptations are produced incrementally through

text: natural selection

text: , so major genetic changes will usually spread within populations only if they do not decrease the

text: fitness

text: of individuals. Although

text: neutral changes

text: (ones which provide no benefit) can spread through

text: genetic drift

text: , and detrimental changes can spread under some circumstances, large changes that require multiple steps will occur only if the intermediate stages increase fitness.

text: Richard Dawkins

text: describes the matter: "The wheel may be one of those cases where the engineering solution can be seen in plain view, yet be unattainable in evolution because it lies [on] the other side of a deep valley, cutting unbridgeably across the massif of Mount Improbable." In such a

text: fitness landscape

text: , wheels might sit on a highly favorable "peak", but the valley around that peak may be too deep or wide for the

text: gene pool

text: to migrate across by genetic drift or natural selection.

text: Stephen Jay Gould

text: notes that biological adaptation is limited to working with available components, commenting that "wheels work well, but animals are debarred from building them by structural constraints inherited as an evolutionary legacy."

text: Natural selection in the evolution of a trait can be divided into three classes. Directional or positive selection refers to a situation where a certain allele has a greater fitness than other alleles, consequently increasing its population frequency (e.g. antibiotic resistance of bacteria). In contrast, stabilizing or negative selection (also known as purifying selection) lowers the frequency or even removes alleles from a population due to disadvantages associated with it with respect to other alleles. Finally, a number of forms of balancing selection exist; those increase genetic variation within a species by being overdominant (heterozygous individuals are fitter than homozygous individuals, e.g. G6PD, a gene that is involved in both sickle cell anaemia and malaria resistance) or can vary spatially within a species that inhabits different niches, thus favouring different alleles. Some genomic differences may not affect fitness.  Neutral variation, previously thought to be “junk” DNA, is unaffected by natural selection resulting in higher genetic variation at such sites when compared to sites where variation does influence fitness.

text: Natural selection

text: in the

text: evolution

text: of a trait can be divided into three classes. Directional or

text: positive selection

text: refers to a situation where a certain allele has a greater fitness than other

text: alleles

text: , consequently increasing its population frequency (e.g.

text: antibiotic resistance

text: of bacteria). In contrast, stabilizing or

text: negative selection

text: (also known as purifying selection) lowers the frequency or even removes alleles from a population due to disadvantages associated with it with respect to other alleles. Finally, a number of forms of

text: balancing selection

text: exist; those increase genetic variation within a species by being overdominant (

text: heterozygous

text: individuals are fitter than

text: homozygous

text: individuals, e.g.

text: , a gene that is involved in both

text: sickle cell anaemia

text: and

text: malaria

text: resistance) or can vary spatially within a species that inhabits different niches, thus favouring different alleles. Some genomic differences may not affect fitness.  Neutral variation, previously thought to be “junk” DNA, is unaffected by natural selection resulting in higher genetic variation at such sites when compared to sites where variation does influence fitness.

text: In natural populations natural selection (adaptation mechanism), gene flow, and mutation combine to change allele frequencies across generations. Genetic drift causes changes in allele frequency from random sampling due to offspring number variance in a finite population size, with small populations experiencing larger per generation fluctuations in frequency than large populations. There is also a theory that second adaptation mechanism exists - niche construction According to Extended Evolutionary Synthesis adaptation occur due to natural selection, environmental induction, non-genetic inheritance, learning and cultural transmission. An allele at a particular locus may also confer some fitness effect for an individual carrying that allele, on which natural selection acts. Beneficial alleles tend to increase in frequency, while deleterious alleles tend to decrease in frequency. Even when an allele is selectively neutral, selection acting on nearby genes may also change its allele frequency through hitchhiking or background selection.

text: In natural populations

text: natural selection

text: adaptation

text: mechanism),

text: gene flow

text: , and

text: mutation

text: combine to change allele frequencies across generations.

text: Genetic drift

text: causes changes in allele frequency from random sampling due to offspring number variance in a finite population size, with small populations experiencing larger per generation fluctuations in frequency than large populations. There is also a theory that second adaptation mechanism exists -

text: niche construction

text: According to

text: Extended Evolutionary Synthesis

text: adaptation occur due to natural selection, environmental induction, non-genetic inheritance, learning and cultural transmission. An allele at a particular locus may also confer some fitness effect for an individual carrying that allele, on which natural selection acts. Beneficial alleles tend to increase in frequency, while deleterious alleles tend to decrease in frequency. Even when an allele is selectively neutral, selection acting on nearby genes may also change its allele frequency through

text: hitchhiking

text: or

text: background selection

text: These experiments address various important questions in evolutionary biology: does evolution proceed in small steps or in large steps; is evolution reversible; how does complexity evolve. It has been shown that slight mutations in the amino acid sequence of hormone receptors determine an important change in their preferences for hormones. These changes mean huge steps in the evolution of the endocrine system. Thus very small changes at the molecular level may have enormous consequences. The Thornton lab has also been able to show that evolution is irreversible studying the glucocorticoid receptor. This receptor was changed by seven mutations in a cortisol receptor, but reversing these mutations didn't give the original receptor back. Indicating that epistasis plays a major role in protein evolution - an observation that in combination with the observations of several examples of parallel evolution, support the neutral network model mentioned above. Other earlier neutral mutations acted as a ratchet and made the changes to the receptor irreversible. These different experiments on receptors show that, during their evolution, proteins are greatly differentiated and this explains how complexity may evolve. A closer look at the different ancestral hormone receptors and the various hormones shows that at the level of interaction between single amino acid residues and chemical groups of the hormones arise by very small but specific changes. Knowledge about these changes may for example lead to the synthesis of hormonal equivalents capable of mimicking or inhibiting the action of a hormone, which might open possibilities for new therapies.

text: These experiments address various important questions in evolutionary biology: does

text: evolution

text: proceed in small steps or in large steps; is evolution reversible; how does

text: complexity

text: evolve. It has been shown that slight

text: mutations

text: in the amino acid sequence of hormone receptors determine an important change in their preferences for hormones. These changes mean huge steps in the evolution of the

text: endocrine system

text: . Thus very small changes at the molecular level may have enormous consequences. The Thornton lab has also been able to show that evolution is irreversible studying the

text: glucocorticoid receptor

text: . This receptor was changed by seven mutations in a cortisol receptor, but reversing these mutations didn't give the original receptor back. Indicating that

text: epistasis

text: plays a major role in protein evolution - an observation that in combination with the observations of several examples of parallel evolution, support the neutral network model mentioned above. Other earlier

text: neutral mutation

text: s acted as a

text: ratchet

text: and made the changes to the receptor irreversible. These different experiments on receptors show that, during their evolution, proteins are greatly differentiated and this explains how complexity may evolve. A closer look at the different ancestral hormone receptors and the various

text: hormone

text: s shows that at the level of interaction between single amino acid residues and chemical groups of the hormones arise by very small but specific changes. Knowledge about these changes may for example lead to the synthesis of hormonal equivalents capable of mimicking or inhibiting the action of a hormone, which might open possibilities for new therapies.

text: abiogenesis – adaptation – adaptive mutation - adaptive radiation – allele – allele frequency – allopatric speciation – altruism – : anagenesis – anti-predator adaptation – Applications of evolution - aposematism – Archaeopteryx – aquatic adaptation – artificial selection – atavism

text: abiogenesis

text: adaptation

text: adaptive mutation

text: adaptive radiation

text: allele

text: allele frequency

text: allopatric speciation

text: altruism

text: : anagenesis

text: anti-predator adaptation

text: Applications of evolution

text: aposematism

text: Archaeopteryx

text: aquatic adaptation

text: artificial selection

text: atavism

text: Natural selection is known to be one of the most important biological processes behind evolution. There are many variations of traits, and some cause greater or lesser reproductive success of the individual. Selection’s purpose is to promote certain alleles, traits, and individuals that have a higher chance to survive and reproduce in their specific environment. Since the environment has a carrying capacity, nature acts on this mode of selection on individuals to let only the most fit offspring survive and reproduce to their full potential. The more advantageous the trait is the more common it will become in the population. Disruptive selection is a specific type of natural selection that actively selects against the intermediate in a population, favoring both extremes of the spectrum.

text: Evolution in colloquial contexts can refer to any sort of progressive development or gradual improvement, and a process that results in greater quality or complexity. When misapplied to biological evolution this common meaning leads to frequent misunderstandings. For example, the idea of devolution ("backwards" evolution) is a result of erroneously assuming that evolution is directional or has a specific goal in mind (cf. orthogenesis). In reality, the evolution of an organism has no "objective" and is only showing increasing ability of successive generations to survive and reproduce in its environment; and increased suitability is only defined in relation to this environment. Biologists do not consider any one species, such as humans, to be more highly evolved or advanced than another. Certain sources have been criticized for indicating otherwise due to a tendency to evaluate nonhuman organisms according to anthropocentric standards rather than more objective ones.

text: Evolution in colloquial contexts can refer to any sort of progressive development or gradual improvement, and a process that results in greater quality or complexity. When misapplied to biological evolution this common meaning leads to frequent misunderstandings. For example, the idea of

text: devolution

text: ("backwards" evolution) is a result of erroneously assuming that evolution is directional or has a specific goal in mind (cf. orthogenesis). In reality, the evolution of an organism has no "objective" and is only showing increasing ability of successive generations to survive and reproduce in its environment; and increased suitability is only defined in relation to this environment. Biologists do not consider any one species, such as humans, to be more highly evolved or advanced than another. Certain sources have been criticized for indicating otherwise due to a tendency to evaluate nonhuman organisms according to

text: anthropocentric

text: standards rather than more objective ones.

text: Short-term EvolutionIn the short-term evolution of influenza A virus, a 2006 study found that stochastic, or random, processes are key factors.  Influenza A virus HA antigenic evolution appears to be characterized more by punctuated, sporadic jumps as opposed to a constant rate of antigenic change.  Using phylogenetic analysis of 413 complete genomes of human influenza A viruses that were collected throughout the state of New York, the authors of Nelson et al. 2006 were able to show that genetic diversity, and not antigenic drift, shaped the short-term evolution of influenza A via random migration and reassortment.  The evolution of these viruses is dominated more by the random importation of genetically different viral strains from other geographic locations and less by natural selection.  Within a given season, adaptive evolution is infrequent and had an overall weak effect as evidenced from the data gathered from the 413 genomes.  Phylogenetic analysis revealed the different strains were derived from newly imported genetic material as opposed to isolates that had been circulating in New York in previous seasons.  Therefore, the gene flow in and out of this population, and not natural selection, was more important in the short term.

text: In contrast to learning, natural selection is solely a biological concept. It is the biological and genetic component that allows an adaptive behavior to be inherited with no connection to the environment. This form of adaptive behavior is most commonly considered in ecological studies, and therefore natural selection is often used to explain ecological adaptive behavior in organisms.

text: In contrast to learning, natural selection is solely a biological concept. It is the biological and genetic component that allows an adaptive behavior to be inherited with no connection to the environment. This form of adaptive behavior is most commonly considered in

text: ecological studies

text: , and therefore natural selection is often used to explain ecological adaptive behavior in organisms.

text: George C. Williams in his influential book Adaptation and Natural Selection was one of the first to present a gene-centered view of evolution with the gene as the unit of selection, arguing that a unit of selection should exhibit a high degree of permanence.

text: George C. Williams

text: in his influential book

text: Adaptation and Natural Selection

text: was one of the first to present a

text: gene-centered view of evolution

text: with the gene as the unit of selection, arguing that a unit of selection should exhibit a high degree of permanence.

text: From this set of considerations, Boyd concluded that to maintain an accurate or effective grasp of reality one must undergo a continuous cycle of interaction with the environment geared to assessing its constant changes. Boyd, though he was hardly the first to do so, then expanded Darwin's theory of evolution, suggesting that natural selection applies not only in biological but also in social contexts (such as the survival of nations during war or businesses in free market competition). Integrating these two concepts, he stated that the decision cycle was the central mechanism of adaptation (in a social context) and that increasing one's own rate and accuracy of assessment vis-a-vis one's counterpart's rate and accuracy of assessment provides a substantial advantage in war or other forms of competition. The key to survival and autonomy is the ability to adapt to change, not perfect adaptation to existing circumstances. Indeed, Boyd noted that radical uncertainty is a necessary precondition of physical and mental vitality: all new opportunities and ideas spring from some mismatch between reality and ideas about it, as examples from the history of science, engineering and business illustrate.

text: From this set of considerations, Boyd concluded that to maintain an accurate or effective grasp of reality one must undergo a continuous cycle of interaction with the environment geared to assessing its constant changes. Boyd, though he was hardly the first to do so, then expanded

text: Darwin

text: 's

text: theory of evolution

text: , suggesting that natural selection applies not only in biological but also in social contexts (such as the survival of nations during war or businesses in free market competition). Integrating these two concepts, he stated that the decision cycle was the central mechanism of adaptation (in a social context) and that increasing one's own rate and accuracy of assessment vis-a-vis one's counterpart's rate and accuracy of assessment provides a substantial advantage in war or other forms of competition. The key to survival and autonomy is the ability to adapt to change, not perfect adaptation to existing circumstances. Indeed, Boyd noted that radical uncertainty is a necessary precondition of physical and mental vitality: all new opportunities and ideas spring from some mismatch between reality and ideas about it, as examples from the history of science, engineering and business illustrate.

text: Conserved gene clusters, such as Hox and the human β-globin gene cluster, may be formed as a result of the process of gene duplication and divergence. A gene is duplicated during cell division, so that its descendants have two end-to-end copies of the gene where it had one copy, initially coding for the same protein or otherwise having the same function. In the course of subsequent evolution, they diverge, so that the products they code for have different but related functions, with the genes still being adjacent on the chromosome. Ohno theorized that the origin of new genes during evolution was dependent on gene duplication. If only a single copy of a gene existed in the genome of a species, the proteins transcribed from this gene would be essential to their survival. Because there was only a single copy of the gene, they could not undergo mutations which would potentially result in new genes; however, gene duplication allows essential genes to undergo mutations in the duplicated copy, which would ultimately give rise to new genes over the course of evolution.  Mutations in the duplicated copy were tolerated because the original copy contained genetic information for the essential gene's function. Species who have gene clusters have a selective evolutionary advantage because natural selection must keep the genes together.  Over a short span of time, the new genetic information exhibited by the duplicated copy of the essential gene would not serve a practical advantage; however, over a long, evolutionary time period, the genetic information in the duplicated copy may undergo additional and drastic mutations in which the proteins of the duplicated gene served a different role than those of the original essential gene. Over the long, evolutionary time period, the two similar genes would diverge so the proteins of each gene were unique in their functions. Hox gene clusters, ranging in various sizes, are found among several phyla.

text: Conserved gene clusters, such as Hox and the human β-globin gene cluster, may be formed as a result of the process of gene duplication and divergence. A gene is duplicated during

text: cell division

text: , so that its descendants have two end-to-end copies of the gene where it had one copy, initially coding for the same protein or otherwise having the same function. In the course of subsequent evolution, they diverge, so that the products they code for have different but related functions, with the genes still being adjacent on the chromosome. Ohno theorized that the origin of new genes during evolution was dependent on gene duplication. If only a single copy of a gene existed in the genome of a species, the proteins transcribed from this gene would be essential to their survival. Because there was only a single copy of the gene, they could not undergo mutations which would potentially result in new genes; however, gene duplication allows essential genes to undergo mutations in the duplicated copy, which would ultimately give rise to new genes over the course of evolution.  Mutations in the duplicated copy were tolerated because the original copy contained genetic information for the essential gene's function. Species who have gene clusters have a selective evolutionary advantage because

text: natural selection

text: must keep the genes together.  Over a short span of time, the new genetic information exhibited by the duplicated copy of the essential gene would not serve a practical advantage; however, over a long, evolutionary time period, the genetic information in the duplicated copy may undergo additional and drastic mutations in which the proteins of the duplicated gene served a different role than those of the original essential gene. Over the long, evolutionary time period, the two similar genes would diverge so the proteins of each gene were unique in their functions. Hox gene clusters, ranging in various sizes, are found among several

text: phyla

text: Co-Evolution of Genes with Culture Hypothesis. The planet Earth is rich in a variety of geographical zones, all of them differing from one another in climate and living conditions they allow for. Across generations individuals populating a certain area learn to adopt and pass on to the next generations the cultural traits that promote survival and flourishing within the environment of their locality. As a result, the genes supporting the survival-relevant traits are passed on, while others fade away. In the long run, it becomes the case that it is for the surviving genes to set conditions for the cultural practices to be used and even to create the environment to which the members adapt. The process that changes the frequency of application of cultural traits is influenced by the same forces that determine the remolding of the combination of genetic variants. These forces are natural selection, mutation, drift, and migration. There is however one more force – 'a decision-making force' – in cultural evolution.  Since cultural traits are transmitted in the context of interpersonal communication, the cultural variants its participants adopt are influenced by the behavioral choices the 'communicator' and the 'learner' make.

text: Co-Evolution of Genes with Culture Hypothesis. The planet Earth is rich in a variety of geographical zones, all of them differing from one another in climate and living conditions they allow for. Across generations individuals populating a certain area learn to adopt and pass on to the next generations the cultural traits that promote survival and flourishing within the environment of their locality. As a result, the genes supporting the survival-relevant traits are passed on, while others fade away. In the long run, it becomes the case that it is for the surviving genes to set conditions for the cultural practices to be used and even to create the environment to which the members adapt. The process that changes the frequency of application of cultural traits is influenced by the same forces that determine the remolding of the combination of genetic variants. These forces are

text: natural selection, mutation, drift, and migration

text: . There is however one more force – 'a decision-making force' – in

text: cultural evolution

text: .  Since cultural traits are transmitted in the context of interpersonal communication, the cultural variants its participants adopt are influenced by the behavioral choices the 'communicator' and the 'learner' make.

text: One of the most recently proposed drivers of the evolution of skin pigmentation in humans is based on research that shows a superior barrier function in darkly pigmented skin. Most protective functions of the skin, including the permeability barrier and the antimicrobial barrier, reside in the stratum corneum (SC) and the researchers surmise that the SC has undergone the most genetic change since the loss of human body hair. Natural selection would have favored mutations that protect this essential barrier; one such protective adaptation is the pigmentation of interfollicular epidermis, because it improves barrier function as compared to non-pigmented skin. In lush rainforests, however, where UV-B radiation and xeric stress were not in excess, light pigmentation would not have been nearly as detrimental. This explains the side-by-side residence of lightly pigmented and darkly pigmented peoples.

text: One of the most recently proposed drivers of the evolution of skin pigmentation in humans is based on research that shows a superior barrier function in darkly pigmented skin. Most protective functions of the skin, including the permeability barrier and the antimicrobial barrier, reside in the

text: stratum corneum

text: (SC) and the researchers surmise that the SC has undergone the most genetic change since the loss of human body hair.

text: Natural selection

text: would have favored mutations that protect this essential barrier; one such protective adaptation is the pigmentation of

text: interfollicular

text: epidermis

text: , because it improves barrier function as compared to non-pigmented skin. In lush rainforests, however, where UV-B radiation and xeric stress were not in excess, light pigmentation would not have been nearly as detrimental. This explains the side-by-side residence of lightly pigmented and darkly pigmented peoples.

text: A comparative analysis of CpG methylation patterns between humans and primates found that there were more than 800 genes that varied in their methylation patterns among orangutans, gorillas, chimpanzees, and bonobos. Despite these apes having the same genes, methylation differences are what account for their phenotypic variation. The genes in question are involved in development. It is not the protein sequences that account for the differences in physical characteristics between humans and apes; rather, it is the epigenetic changes to the genes. Since humans and the great apes share 99% of their DNA, it is thought that the differences in methylation patterns account for their distinction. So far, there are known to be 171 genes that are uniquely methylated in humans, 101 genes that are uniquely methylated in chimpanzees and bonobos, 101 genes that are uniquely methylated in gorillas, and 450 genes that are uniquely methylated in orangutans. For example, genes involved in blood pressure regulation and the development of the inner ear’s semicircular canal are highly methylated in humans, but not in the apes. There are also 184 genes that are conserved at the protein level between human and chimpanzees, but have epigenetic differences. Enrichments in multiple independent gene categories show that regulatory changes to these genes have given humans their specific traits. This research shows that epigenetics plays an important role in evolution in primates.It has also been shown that cis-regulatory elements changes affect the transcription start sites (TSS) of genes. 471 DNA sequences are found to be enriched or depleted in regards to histone trimethylation at the H3K4 histone in chimpanzee, human, and macaque prefrontal cortexes. Among these sequences, 33 are selectively methylated in neuronal chromatin from children and adults, but not from non-neuronal chromatin. One locus that was selectively methylated was DPP10, a regulatory sequence that showed evidence of hominid adaptation, such as higher nucleotide substitution rates and certain regulatory sequences that were missing in other primates. Epigenetic regulation of TSS chromatin has been identified as an important development in the evolution of gene expression networks in the human brain. These networks are thought to play a role in cognitive processes and neurological disorders.An analysis of methylation profiles of humans and primate sperm cells reveals epigenetic regulation plays an important role here as well. Since mammalian cells undergo reprogramming of DNA methylation patterns during germ cell development, the methylomes of human and chimp sperm can be compared to methylation in embryonic stem cells (ESCs). There were many hypomethylated regions in both sperms cells and ESCs that showed structural differences. Also, many of the promoters in human and chimp sperm cells had different amounts of methylation. In essence, DNA methylation patterns differ between germ cells and somatic cells as well as between the human and chimpanzee sperm cells. Meaning, differences in promoter methylation could possible account for the phenotypic differences between humans and primates.

text: A comparative analysis of

text: CpG

text: methylation patterns between humans and primates found that there were more than 800 genes that varied in their methylation patterns among orangutans, gorillas, chimpanzees, and bonobos. Despite these apes having the same genes, methylation differences are what account for their

text: phenotypic variation

text: . The genes in question are involved in development. It is not the protein sequences that account for the differences in physical characteristics between humans and apes; rather, it is the epigenetic changes to the genes. Since humans and the great apes share 99% of their DNA, it is thought that the differences in methylation patterns account for their distinction. So far, there are known to be 171 genes that are uniquely methylated in humans, 101 genes that are uniquely methylated in chimpanzees and bonobos, 101 genes that are uniquely methylated in gorillas, and 450 genes that are uniquely methylated in orangutans. For example, genes involved in blood pressure regulation and the development of the inner ear’s semicircular canal are highly methylated in humans, but not in the apes. There are also 184 genes that are conserved at the protein level between human and chimpanzees, but have epigenetic differences. Enrichments in multiple independent gene categories show that regulatory changes to these genes have given humans their specific traits. This research shows that epigenetics plays an important role in evolution in primates.It has also been shown that cis-regulatory elements changes affect the transcription start sites (TSS) of genes. 471 DNA sequences are found to be enriched or depleted in regards to histone trimethylation at the H3K4 histone in chimpanzee, human, and macaque prefrontal cortexes. Among these sequences, 33 are selectively methylated in neuronal

text: chromatin

text: from children and adults, but not from non-neuronal chromatin. One

text: locus

text: that was selectively methylated was

text: , a

text: regulatory sequence

text: that showed evidence of hominid adaptation, such as higher

text: nucleotide substitution

text: rates and certain regulatory sequences that were missing in other primates. Epigenetic regulation of TSS chromatin has been identified as an important development in the evolution of gene expression networks in the human brain. These networks are thought to play a role in

text: cognitive processes

text: and

text: neurological disorders

text: .An analysis of methylation profiles of humans and primate

text: sperm cells

text: reveals epigenetic regulation plays an important role here as well. Since mammalian cells undergo reprogramming of DNA methylation patterns during

text: germ cell

text: development, the methylomes of human and chimp sperm can be compared to methylation in

text: embryonic stem cells

text: (ESCs). There were many hypomethylated regions in both sperms cells and ESCs that showed structural differences. Also, many of the

text: promoter

text: s in human and chimp sperm cells had different amounts of methylation. In essence, DNA methylation patterns differ between germ cells and

text: somatic cells

text: as well as between the human and chimpanzee sperm cells. Meaning, differences in promoter methylation could possible account for the phenotypic differences between humans and primates.

text: The handicap principle gains further support by providing interpretations for behaviours that fit into a single unifying gene-centered view of evolution and making earlier explanations based on group selection obsolete. A classic example is that of stotting in gazelles. This behaviour consists in the gazelle initially running slowly and jumping high when threatened by a predator such as a lion or cheetah. The explanation based on group selection was that such behaviour might be adapted to alerting other gazelle to a cheetah's presence or might be part of a collective behaviour pattern of the group of gazelle to confuse the cheetah. Instead, Zahavi proposed that each gazelle was communicating  that it was a fitter individual than its fellows. This may communicate to the cheetah to avoid chasing this fitter individual, or the message may be oriented toward the other gazelles, which would indicate its fitness to potential mates.

text: The handicap principle gains further support by providing interpretations for behaviours that fit into a single unifying

text: gene-centered view of evolution

text: and making earlier explanations based on

text: group selection

text: obsolete. A classic example is that of

text: stotting

text: in

text: gazelle

text: s. This behaviour consists in the gazelle initially running slowly and jumping high when threatened by a

text: predator

text: such as a

text: lion

text: or

text: cheetah

text: . The explanation based on group selection was that such behaviour might be adapted to alerting other gazelle to a cheetah's presence or might be part of a collective behaviour pattern of the group of gazelle to confuse the cheetah. Instead, Zahavi proposed that each gazelle was communicating  that it was a fitter individual than its fellows. This may communicate to the cheetah to avoid chasing this fitter individual, or the message may be oriented toward the other gazelles, which would indicate its fitness to potential mates.

text: This suggests, once again, that polymorphism is a common and important aspect of adaptive evolution in natural populations.

text: l theory of human evolution had far more reaching consequences, than “just poor science”.  This theory not only justified rankings of human culture based on superficial and materialist accomplishments, but correlated it with an intrinsic biological component such as evolution. It justified cultural superiority and accepted the flawed notion of ‘advanced’ societies are better than ‘primitive’ societies due to “scientific evidence”.  This approach was further expanded into the idea of biological determinism and ideology where certain cultural groups are inferior or superior based on their genetic makeup. These were the evolutionary methods and ideological notions occupied by anthropologist in the early 19 century through late 20 century. This mentality favored and justified colonization efforts by many European powers. As a result, “the social and political implications of early cultural evolutionary thinking, such as eugenics and other atrocities, understandably promoted a reaction, and the social sciences turned towards a thoroughly anti-biological stance: The Standard Social Science Model (Barkow et al, 1992). Therefore, a strong resistance among scientist rose arguing for limited biological restraints on human behavior. This reaction led to the acceptance of social influences and culture affecting and changing human behavior. The Standard Social Science Model, assumes that “culture is selected by free agents making active, unconstrained choices, and there has been a tendency to stress the vast plethora of different cultural practices rather than to look for cultural universals” (Workman and Reader, 2004).

text: l theory of human evolution had far more reaching consequences, than “just poor science”.  This theory not only justified rankings of human culture based on superficial and materialist accomplishments, but correlated it with an intrinsic biological component such as evolution. It justified cultural superiority and accepted the flawed notion of ‘advanced’ societies are better than ‘primitive’ societies due to “scientific evidence”.  This approach was further expanded into the idea of

text: biological determinism

text: and ideology where certain cultural groups are inferior or superior based on their genetic makeup. These were the evolutionary methods and ideological notions occupied by anthropologist in the early 19 century through late 20 century. This mentality favored and justified colonization efforts by many European powers. As a result, “the social and political implications of early cultural evolutionary thinking, such as eugenics and other atrocities, understandably promoted a reaction, and the social sciences turned towards a thoroughly anti-biological stance: The Standard Social Science Model (Barkow et al, 1992). Therefore, a strong resistance among scientist rose arguing for limited biological restraints on human behavior. This reaction led to the acceptance of social influences and culture affecting and changing human behavior. The Standard Social Science Model, assumes that “culture is selected by free agents making active, unconstrained choices, and there has been a tendency to stress the vast plethora of different cultural practices rather than to look for cultural universals” (Workman and Reader, 2004).

text: In Tylor's time, the major themes of evolution were rapidly replacing earlier views based on Biblical scholarship. Biological evolution , proposed by Darwin and Wallace, hypothesized a gradual modification of species to form new species. By the methods of comparative anatomy it concluded to a general progression from simpler and generally less capable forms to more complex and more capable forms. At the top of the progression were the mammals, led by man. The projected starting point must be an Earth of no life forms at all, which was supported by the contemporaneous views of the birth and death of heavenly bodies.  A chemical pre-adaptation on early Earth was required for a starting point, an abiogenesis, or spontaneous generation of replicating molecules.

text: In Tylor's time, the major themes of evolution were rapidly replacing earlier views based on Biblical scholarship. Biological

text: evolution

text: , proposed by Darwin and Wallace, hypothesized a gradual modification of species to form new species. By the methods of

text: comparative anatomy

text: it concluded to a general progression from simpler and generally less capable forms to more complex and more capable forms. At the top of the progression were the mammals, led by man. The projected starting point must be an Earth of no life forms at all, which was supported by the contemporaneous views of the birth and death of heavenly bodies.  A chemical pre-adaptation on early Earth was required for a starting point, an

text: abiogenesis

text: , or spontaneous generation of replicating molecules.

text: While natural selection has a direction, guiding evolution towards heritable adaptations to the current environment, genetic drift has no direction and is guided only by the mathematics of chance. As a result, drift acts upon the genotypic frequencies within a population without regard to their phenotypic effects. In contrast, selection favors the spread of alleles whose phenotypic effects increase survival and/or reproduction of their carriers, lowers the frequencies of alleles that cause unfavorable traits, and ignores those that are neutral.

text: While natural selection has a direction, guiding evolution towards heritable

text: adaptation

text: s to the current environment, genetic drift has no direction and is guided only by the

text: mathematics of chance

text: . As a result, drift acts upon the

text: genotypic frequencies

text: within a population without regard to their phenotypic effects. In contrast, selection favors the spread of alleles whose phenotypic effects increase survival and/or reproduction of their carriers, lowers the frequencies of alleles that cause unfavorable traits, and ignores those that are neutral.

text: "Evolutionary epistemology" can refer to a branch of epistemology that applies the concepts of biological evolution to the growth of animal and human cognition.  It argues that the mind is in part genetically determined and that its structure and function reflect adaptation, a nonteleological process of interaction between the organism and its environment. A cognitive trait tending to increase inclusive fitness in a given population should therefore grow more common over time, and a trait tending to prevent its carriers from passing on their genes should show up less and less frequently.

text: "Evolutionary epistemology" can refer to a branch of

text: epistemology

text: that applies the concepts of biological

text: evolution

text: to the growth of animal and human cognition.  It argues that the mind is in part genetically determined and that its structure and function reflect

text: adaptation

text: , a

text: nonteleological

text: process of interaction between the organism and its environment. A cognitive trait tending to increase

text: inclusive fitness

text: in a given population should therefore grow more common over time, and a trait tending to prevent its carriers from passing on their genes should show up less and less frequently.

text: Evolution is change in the heritable characteristics of biological populations over successive generations. Evolutionary processes give rise to biodiversity at every level of biological organisation, including the levels of species, individual organisms, and  molecules.

text: Evolution is change in the

text: heritable

text: characteristics of

text: biological

text: population

text: s over successive

text: generation

text: s. Evolutionary processes give rise to

text: biodiversity

text: at every level of

text: biological organisation

text: , including the levels of

text: species

text: , individual

text: organism

text: s, and

text: molecules

text: The difference between cultural behaviors acquired through selective transmission mechanisms and behaviors resulting from psychological adaptations has resulted in some researchers differentiating between evoked and transmitted culture (Tooby and Cosmides, 1992). Evoked cultural behaviors are those that are the outputs of shared psychological mechanisms in response to local environmental cues Fessler et al., 2015). Transmitted cultural behaviors are those behaviors which are learned from one’s social group, regardless of environment. As such, investigating evoked culture may well rest in the domain of evolutionary psychology and transmitted culture studied by culture evolution, social psychology, and other disciplines. Differentiating between the two is more difficult than it may appear, however. For example, given the level of cross-cultural variation in human societies, variation between any two cultures may result from separate histories and cultural evolutionary pathways or different evoked cultures, based on different local environmental cues, while uniformity across cultures could result from convergent cultural evolution or similarly functioning psychological adaptations (Fessler et al., 2015). Specifically, if two cultures exhibit similar behaviors to avoid pathogens, can one differentiate between either the two groups independently inventing similar behaviors or the groups displaying the same underlying psychological adaptation for pathogen avoidance? As well, the issue of behavioral variation and transmitted culture may be seen as a major point of contention between evolutionary psychology and other disciplines, especially gene-culture coevolution. Evolutionary psychology and human behavioral ecology both admit to cultural transmission as a source of behavioral variation but do not see it as an evolutionary process or its potential to have significantly affected human evolution. Gene-culture coevolution would argue that culture traits may have altered genetic change and selection pressures, ultimately affecting cognition. However, as Brown et al. (2011) states:"Whether human behavioural ecologists and evolutionary psychologists are willing to accommodate the idea that some portion of human behavioural diversity could result from genetic differences that have arisen via selection pressures imposed by socially transmitted behaviour remains to be seen." 	While this may difficult, some evolutionary psychologists, such as Gangestad, Haselton, and Buss (2006) have argued that the future of the discipline rests on unifying transmitted and evoked culture. In order to do so, the authors suggest that a more comprehensive definition of culture needs to be developed since different cultural phenomena need different kinds of analysis to investigate their effects. As they state, “Culture, however, is not a "thing" with singularity; it's an umbrella concept subsuming a collection of extraordinarily varied phenomena, each of which requires scientific analysis” (Gangestad et al., 2006). Ultimately, the issues of behavioral variation through evoked or transmitted culture must rest on empirical evidence.

text: adaptive dynamics effectively combines game theory and population dynamics. As such, it can be very useful in investigating how evolution affects the dynamics of populations. One interesting finding to come out of this is that individual-level adaptation can sometimes result in the extinction of the whole population/species, a phenomenon known as evolutionary suicide.

text: adaptive dynamics effectively combines

text: game theory

text: and

text: population dynamics

text: . As such, it can be very useful in investigating how

text: evolution

text: affects the

text: dynamics of populations

text: . One interesting finding to come out of this is that individual-level

text: adaptation

text: can sometimes result in the

text: extinction

text: of the whole

text: population

text: species

text: , a phenomenon known as

text: evolutionary suicide

text: Natural selection is one of the cornerstones of modern biology. The concept, published by Darwin and Alfred Russel Wallace in a joint presentation of papers in 1858, was elaborated in Darwin's influential 1859 book On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life, which described natural selection as analogous to artificial selection, a process by which animals and plants with traits considered desirable by human breeders are systematically favoured for reproduction. The concept of natural selection originally developed in the absence of a valid theory of heredity; at the time of Darwin's writing, science had yet to develop modern theories of genetics. The union of traditional Darwinian evolution with subsequent discoveries in classical genetics formed the modern evolutionary synthesis of the mid-20th century. The addition of molecular genetics has led to evolutionary developmental biology, which explains evolution at the molecular level. While genotypes can slowly change by random genetic drift, natural selection remains the primary explanation for adaptive evolution.

text: biology

text: . The concept, published by Darwin and

text: Alfred Russel Wallace

text: in a

entity_name: On the Tendency of Species to form Varieties; and on the Perpetuation of Varieties and Species by Natural Means of Selection

text: joint presentation of papers in 1858

text: , was elaborated in Darwin's influential 1859 book

text: On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life

text: , which described natural selection as analogous to artificial selection, a process by which animals and plants with traits considered desirable by human breeders are systematically favoured for reproduction. The concept of natural selection originally developed in the absence of a valid theory of heredity; at the time of Darwin's writing, science had yet to develop modern theories of genetics. The union of traditional

text: Darwinian evolution

text: with subsequent discoveries in

text: classical genetics

text: formed the

text: modern evolutionary synthesis

text: of the mid-20th century. The addition of

text: molecular genetics

text: has led to

text: evolutionary developmental biology

text: , which explains evolution at the molecular level. While genotypes can slowly change by random

text: genetic drift

text: , natural selection remains the primary explanation for

text: adaptive evolution

text: As noted previously, many of the tests used to detect adaptive evolution have very large degrees of uncertainty surrounding their estimates. It is beyond the purview of this article to look at all the modifications applied to individual tests to overcome the associated problems. However, it will briefly discuss in general terms two of what may be the most important confounding variables that may hinder accurate detection of adaptive evolution. Demographic changes are particularly problematic and may severely bias estimates of adaptive evolution. The human lineage has undergone both rapid population size contractions and expansions over its evolutionary history, and these events will change many of the signatures thought to be characteristic of adaptive evolution (Nielsen et al. 2007). Some genomic methods have been shown through simulations to be relatively robust to demographic changes (e.g. Willamson et al. 2007). However, no tests are completely robust to demographic changes, and new genetic phenomena linked to demographic changes have recently been discovered. This includes the concept of “surfing mutations”, where new mutations can be propagated with a population expansion (Klopfstein et al. 2006).A phenomenon which could severely alter the way we look for signatures of adaptive evolution is biased gene conversion (BGC) (Galtier and Duret 2007). Meiotic recombination between homologous chromosomes that are heterozygous at a particular locus can produce a DNA mismatch. DNA repair mechanisms are biased towards repairing a mismatch to the CG base pair. This will lead allele frequencies to change, leaving a signature of non-neutral evolution (Galtier et al. 2001). The excess of AT to GC mutations in human genomic regions with high substitution rates (human accelerated regions, HARs) implies that BGC has occurred frequently in the human genome (Pollard et al. 2006, Galtier and Duret 2007). Initially, it was postulated that BGC could have been adaptive (Galtier et al. 2001), but more recent observations have made this seem unlikely. Firstly, some HARs show no substantial signs of selective sweeps around them. Secondly, HARs tend to be present in regions with high recombination rates (Pollard et al. 2006). In fact, BGC could lead to HARs containing a high frequency of deleterious mutations (Galtier and Duret 2007). However, it is unlikely that HARs are generally maladaptive, because DNA repair mechanisms themselves would be subject to strong selection if they propagated deleterious mutations. Either way, BGC should be further investigated, because it may force radical alteration of the methods which test for the presence of adaptive evolution.

text: As noted previously, many of the tests used to detect adaptive evolution have very large degrees of uncertainty surrounding their estimates. It is beyond the purview of this article to look at all the modifications applied to individual tests to overcome the associated problems. However, it will briefly discuss in general terms two of what may be the most important confounding variables that may hinder accurate detection of adaptive evolution. Demographic changes are particularly problematic and may severely bias estimates of adaptive evolution. The human lineage has undergone both rapid population size contractions and expansions over its evolutionary history, and these events will change many of the signatures thought to be characteristic of adaptive evolution (Nielsen et al. 2007). Some genomic methods have been shown through simulations to be relatively robust to demographic changes (e.g. Willamson et al. 2007). However, no tests are completely robust to demographic changes, and new genetic phenomena linked to demographic changes have recently been discovered. This includes the concept of “surfing mutations”, where new mutations can be propagated with a population expansion (Klopfstein et al. 2006).A phenomenon which could severely alter the way we look for signatures of adaptive evolution is

text: biased gene conversion

text: (BGC) (Galtier and Duret 2007). Meiotic recombination between homologous chromosomes that are heterozygous at a particular locus can produce a DNA mismatch. DNA repair mechanisms are biased towards repairing a mismatch to the CG base pair. This will lead allele frequencies to change, leaving a signature of non-neutral evolution (Galtier et al. 2001). The excess of AT to GC mutations in human genomic regions with high substitution rates (human accelerated regions, HARs) implies that BGC has occurred frequently in the human genome (Pollard et al. 2006, Galtier and Duret 2007). Initially, it was postulated that BGC could have been adaptive (Galtier et al. 2001), but more recent observations have made this seem unlikely. Firstly, some HARs show no substantial signs of selective sweeps around them. Secondly, HARs tend to be present in regions with high recombination rates (Pollard et al. 2006). In fact, BGC could lead to HARs containing a high frequency of deleterious mutations (Galtier and Duret 2007). However, it is unlikely that HARs are generally maladaptive, because DNA repair mechanisms themselves would be subject to strong selection if they propagated deleterious mutations. Either way, BGC should be further investigated, because it may force radical alteration of the methods which test for the presence of adaptive evolution.

text: The annual vs. perennial trait has been empirically associated with differing subsequent rates of molecular evolution within multiple plant lineages. The perennial trait is generally associated with a slower rate of evolution than annual species when looking at both non-coding and coding DNA. Generation time is often implicated as one of the major factors contributing to this disparity, with perennials having longer generation times and likewise an overall slower mutation and adaptation rate. This may result in higher genetic diversity in annual lineages.

text: The annual vs. perennial trait has been

text: empirically

text: associated with differing subsequent rates of

text: molecular evolution

text: within multiple plant lineages. The perennial trait is generally associated with a slower rate of evolution than annual species when looking at both

text: non-coding

text: and

text: coding DNA

text: Generation time

text: is often implicated as one of the major factors contributing to this disparity, with perennials having longer generation times and likewise an overall slower

text: mutation

text: and

text: adaptation

text: rate. This may result in higher

text: genetic diversity

text: in annual lineages.

text: Segmental duplications are also important for their role in the evolution of new genes as it is one of the primary mechanisms by which new genes are created. The most common method by which segmental duplication operates in creating new genes is by duplication of the entire gene, whether it is in tandem or in an interspersed configuration. Using whole-genome shotgun sequencing, it has been found that gene density is the greatest factor in showing correlation with segmental duplication density.  However, this does not mean that all segmental duplications are enriched for genes and that all duplication regions are sites for the formation of new genes. Rather, it has been seen that segmental duplications which contain interstitial interchromosomal duplications are regions with the most enrichment of genes. Comparing the subtelomeric and pericentromeric regions, it is seen that the subtelomeric regions contains more gene content and transcriptional activity. In addition, genes found within segmental duplications also share properties. Firstly, strong signatures of positive selection are common in segmentally duplicated genes. Secondly, these genes are five to ten times more likely than their counterparts to show interspecies and intraspecies structural and copy-number variation. Finally, immune response and xenobiotic recognition are some functions that are enriched within these genes. Together, this suggests an important role for segmental duplication within human and primate adaptive evolution.

text: The game was created by Charlie "Flayra" Cleveland, who later founded the company Unknown Worlds Entertainment. Natural Selection v1 was first publicly released on Halloween 2002, and is now at version 3.2. Natural Selection 2 was released in late 2012.

text: The game was created by Charlie "Flayra" Cleveland, who later founded the company

text: Unknown Worlds Entertainment

text: . Natural Selection v1 was first publicly released on

text: Halloween

text: 2002, and is now at version 3.2.

text: Natural Selection 2

text: was released in late 2012.

text: Studies of biological evolution generally concern the evolution of the human form. Cultural evolution involves the study of cultural change over time and space and frequently incorporate cultural transmission models. Note that cultural evolution is not the same as biological evolution, and that human culture involves the transmission of cultural information, which behaves in ways quite distinct from human biology and genetics. The study of cultural change is increasingly performed through cladistics and genetic models.

text: Studies of biological evolution generally concern the evolution of the human form. Cultural evolution involves the study of cultural change over time and space and frequently incorporate

text: cultural transmission

text: models. Note that cultural evolution is not the same as biological evolution, and that human culture involves the transmission of cultural information, which behaves in ways quite distinct from

text: human biology

text: and

text: genetics

text: . The study of cultural change is increasingly performed through

text: cladistics

text: and genetic models.

text: Fisherian runaway is a hypothesized genetic inter-sexual selection mechanism for the paradoxical evolution of elaborate and exaggerated male ornaments that neither play a role in intra-sexual selection nor serve to increase fitness in an obvious way. Fisherian runaway is an attempt to resolve this paradoxical evolution within the context of evolutionary theory. Originally hypothesized during the beginning of the modern evolutionary synthesis, Fisherian runaway suggests that male ornamentation is the result of a positive feedback ("runaway") mechanism that is driven by the "intersexual" preference in mate choice by females for males that express the most exaggerated ornaments. A strong female choice for the expression alone, as opposed to the function, of a male ornament can oppose and undermine the forces of natural selection and result in the runaway sexual selection that leads to the further exaggeration of the ornament (as well as the preference) until the costs (incurred by natural selection) of the expression become greater than the benefit (bestowed by sexual selection).

text: Fisherian runaway is a hypothesized genetic inter-sexual selection mechanism for the paradoxical evolution of elaborate and exaggerated male ornaments that neither play a role in intra-sexual selection nor serve to increase fitness in an obvious way. Fisherian runaway is an attempt to resolve this paradoxical evolution within the context of

text: evolutionary theory

text: . Originally hypothesized during the beginning of the modern evolutionary synthesis, Fisherian runaway suggests that male ornamentation is the result of a positive feedback ("runaway") mechanism that is driven by the "intersexual" preference in

text: mate choice

text: by females for males that express the most exaggerated ornaments. A strong female choice for the expression alone, as opposed to the function, of a male ornament can oppose and undermine the forces of natural selection and result in the runaway sexual selection that leads to the further exaggeration of the ornament (as well as the preference) until the costs (incurred by natural selection) of the expression become greater than the benefit (bestowed by sexual selection).

text: In general, the McDonald-Kreitman test is often thought to be unreliable because of how significantly the test tends to underestimate the degree of adaptive evolution in the presence of slightly deleterious mutations. A slightly deleterious mutation can be defined as a mutation that negative selection acts on only very weakly so that its fate is determined by both selection and random genetic drift. If slightly deleterious mutations are segregating in the population, then it becomes difficult to detect positive selection and the degree of positive selection is underestimated. Weakly deleterious mutations have a larger chance of contributing to polymorphism than strongly deleterious mutations, but still have low probabilities of fixation. This creates bias in the McDonald–Kreitman test's estimate of the degree of adaptive evolution, resulting in a dramatically lower estimate of α. By contrast, since strongly deleterious mutations contribute to neither polymorphism nor divergence, strongly deleterious mutations do not bias estimates of α. The presence of slightly deleterious mutations is strongly linked to genes that  have experienced the greatest reduction in effective population size. This means that soon after a recent reduction in effective population size in a species has occurred, such as a bottleneck, we observe a larger presence of slightly deleterious mutations in the protein-coding regions. We can make a direct connection with the increase in number of slightly deleterious mutations and the recent decrease in effective population size. For more information on why population size affects the tendency of slightly deleterious mutations to increase in frequency, refer to the article Nearly neutral theory of molecular evolution.

entity_name: Nearly neutral theory of molecular evolution

text: Nearly neutral theory of molecular evolution

text: Several recent studies have compared the amounts of adaptive evolution occurring between different populations within the human species. Williamson et al. (2007) found more evidence of adaptive evolution in European and Asian populations than African American populations. Assuming African Americans are representative of Africans, these results makes sense intuitively, because humans spread out of Africa approximately 50,000 years ago  (according to the consensus Out-of-Africa hypothesis of human origins (Klein 2009)), and these humans would have adapted to the new environments they encountered. By contrast, African populations remained in a similar environment for the following tens of thousands of years, and were therefore probably nearer their adaptive peak for the environment. However, Voight et al. (2006) found evidence of more adaptive evolution in Africans, than in Non-Africans (East Asian and European populations examined), and Boyko et al. (2008) found no significant difference in the amount of adaptive evolution occurring between different human populations. Therefore, the evidence obtained so far is inconclusive as to what extent different human populations have undergone different amounts of adaptive evolution.

text: Subsequent chapters consider the origin of life on Earth, and the physical extremes to which life as adapted. In astrobiology, it pays to think “outside the box” and imagine how strange life might be or whether post-biological evolution is possible, where the basis is mechanical or computational. A chapter on evolution shows how it is affected by the cosmic environment.

text: Subsequent chapters consider the

text: origin of life on Earth

text: , and the physical extremes to which life as adapted. In

text: astrobiology

text: , it pays to think “outside the box” and imagine how strange life might be or whether post-

text: biological evolution

text: is possible, where the basis is mechanical or computational. A chapter on evolution shows how it is affected by the cosmic environment.

text: In fact, evolution is being put to practical use in industry and widely used on a daily basis by researchers in medicine, biochemistry, molecular biology, and genetics to both formulate hypotheses about biological systems for the purposes of experimental design, as well as to rationalise observed data and prepare applications. As of January 2016 there are 428,827 scientific papers in PubMed that mention 'evolution'. Pharmaceutical companies utilize biological evolution in their development of new products, and also use these medicines to combat evolving bacteria and viruses.

text: In fact, evolution is being put to practical use in industry and widely used on a daily basis by researchers in

text: medicine

text: biochemistry

text: molecular biology

text: , and

text: genetics

text: to both formulate hypotheses about biological systems for the purposes of experimental design, as well as to rationalise observed data and prepare applications. As of January 2016 there are 428,827 scientific papers in

text: PubMed

text: that mention 'evolution'. Pharmaceutical companies utilize biological evolution in their development of new products, and also use these medicines to combat evolving bacteria and viruses.

text: In 1979, influenced by Seilacher among others, the paleontologist Stephen J. Gould and the population geneticist Richard Lewontin wrote what Wagner called "the most influential structuralist manifesto", "The Spandrels of San Marco and the Panglossian Paradigm". They pointed out that biological features (like architectural spandrels) did not necessarily have adaptation as their direct cause. Instead, architects couldn't help creating small triangular areas between arches and pillars, as arches need (evolve) to be curved, and pillars need to be vertical. The resulting spandrels are exaptations, consequences of other evolutionary changes. Evolution, they argued, did not select for a protruding human chin: instead, reducing the length of the tooth row left the jaw protruding.

text: In 1979, influenced by Seilacher among others, the

text: paleontologist

text: Stephen J. Gould

text: and the

text: population geneticist

text: Richard Lewontin

text: wrote what Wagner called "the most influential structuralist manifesto", "The

text: Spandrel

text: s of San Marco and the

text: Panglossian

text: Paradigm". They pointed out that biological features (like

text: architectural spandrels

text: ) did not necessarily have

text: adaptation

text: as their direct cause. Instead, architects couldn't help creating small triangular areas between

text: arch

text: es and

text: pillar

text: s, as arches need (evolve) to be curved, and pillars need to be vertical. The resulting spandrels are

text: exaptation

text: s, consequences of other evolutionary changes. Evolution, they argued, did not select for a protruding human chin: instead, reducing the length of the tooth row left the jaw protruding.

text: Darwin’s Cathedral: Evolution, Religion, and the Nature of Society (ISBN 9780226901350) is a 2002 book by David Sloan Wilson which proposes that religion is a multi-level adaptation—i.e., a product of cultural evolution developed through multi-level selection.

text: Darwin’s Cathedral: Evolution, Religion, and the Nature of Society (ISBN 9780226901350) is a 2002 book by

text: David Sloan Wilson

text: which proposes that

text: religion

text: is a multi-level adaptation—i.e., a product of

text: cultural evolution

text: developed through

text: multi-level selection

text: From the above definitions, it is clear that there is a relationship between adaptedness and fitness (a key population genetics concept). Differences in fitness between genotypes predict the rate of evolution by natural selection. Natural selection changes the relative frequencies of alternative phenotypes, insofar as they are heritable. Although the two are connected, the one does not imply the other: a phenotype with high adaptedness may not have high fitness. Dobzhansky mentioned the example of the Californian redwood, which is highly adapted, but a relict species in danger of extinction. Elliott Sober commented that adaptation was a retrospective concept since it implied something about the history of a trait, whereas fitness predicts a trait's future.

text: From the above definitions, it is clear that there is a relationship between adaptedness and fitness (a key

text: population genetics

text: concept). Differences in fitness between

text: genotype

text: s predict the rate of evolution by natural selection. Natural selection changes the relative frequencies of alternative phenotypes, insofar as they are

text: heritable

text: . Although the two are connected, the one does not imply the other: a phenotype with high adaptedness may not have high fitness. Dobzhansky mentioned the example of the

text: Californian redwood

text: , which is highly adapted, but a

text: relict

text: species in danger of

text: extinction

text: Elliott Sober

text: commented that adaptation was a retrospective concept since it implied something about the history of a trait, whereas fitness predicts a trait's future.

text: Natural selection can act on any heritable phenotypic trait, and selective pressure can be produced by any aspect of the environment, including sexual selection and competition with members of the same or other species. However, this does not imply that natural selection is always directional and results in adaptive evolution; natural selection often results in the maintenance of the status quo by eliminating less fit variants.

text: Natural selection can act on any heritable

text: phenotypic trait

text: , and selective pressure can be produced by any aspect of the environment, including sexual selection and

text: competition

text: with members of the same or other species. However, this does not imply that natural selection is always directional and results in adaptive evolution; natural selection often results in the maintenance of the status quo by eliminating less fit variants.

text: Evolution generally refers to biological evolution, but here it means a process in which the whole universe is a progression of interrelated phenomena, a gradual process in which something changes into a different and usually more complex form (emergence). It should not be 'biologized’ as it includes many areas of science. In addition, outside of the scientific community, the term evolution is frequently used differently from scientists' usage. Even respected dictionaries may present questionable definitions that are not scientific or only address genetic changes. Unfortunately it is common for the general public to enter into a discussion about evolution with a wrong definition in mind. This often leads to misunderstanding since scientists are viewing evolution from a different perspective. The same applies to the use of the term theory as used in the theory of evolution (see references for Evolution as theory and fact).

text: Evolution generally refers to biological

text: evolution

text: , but here it means a

text: process

text: in which the whole universe is a progression of interrelated phenomena, a gradual process in which something changes into a different and usually more complex form (

text: emergence

text: ). It should not be 'biologized’ as it includes many areas of science. In addition, outside of the scientific community, the term evolution is frequently used differently from scientists' usage. Even respected dictionaries may present questionable definitions that are not scientific or only address genetic changes. Unfortunately it is common for the general public to enter into a discussion about evolution with a wrong definition in mind. This often leads to misunderstanding since scientists are viewing evolution from a different perspective. The same applies to the use of the term theory as used in the theory of evolution (see references for

text: Evolution as theory and fact

text: Evolutionary psychologists say that natural selection has provided humans with many psychological adaptations, in much the same way that it generated humans' anatomical and physiological adaptations. As with adaptations in general, psychological adaptations are said to be specialized for the environment in which an organism evolved, the environment of evolutionary adaptedness, or EEA. Sexual selection provides organisms with adaptations related to mating. For male mammals, which have a relatively high maximal potential reproduction rate, sexual selection leads to adaptations that help them compete for females. For female mammals, with a relatively low maximal potential reproduction rate, sexual selection leads to choosiness, which helps females select higher quality mates. Charles Darwin described both natural selection and sexual selection, and he relied on group selection to explain the evolution of altruistic (self-sacrificing) behavior. But group selection was considered a weak explanation, because in any group the less altruistic individuals will be more likely to survive, and the group will become less self-sacrificing as a whole.

text: Evolutionary psychologists say that natural selection has provided humans with many psychological adaptations, in much the same way that it generated humans' anatomical and physiological adaptations. As with adaptations in general, psychological adaptations are said to be specialized for the environment in which an organism evolved, the environment of evolutionary adaptedness, or EEA. Sexual selection provides organisms with adaptations related to mating. For male

text: mammal

text: s, which have a relatively high maximal potential reproduction rate, sexual selection leads to adaptations that help them compete for females. For female mammals, with a relatively low maximal potential reproduction rate, sexual selection leads to choosiness, which helps females select higher quality mates. Charles Darwin described both natural selection and sexual selection, and he relied on group selection to explain the evolution of

text: altruistic

text: (self-sacrificing) behavior. But group selection was considered a weak explanation, because in any group the less altruistic individuals will be more likely to survive, and the group will become less self-sacrificing as a whole.

text: A variant of group selection theory rooted in the idea of population viscosity (limited offspring dispersal), first proposed by Hamilton (1964), is widely present in natural populations. This population structure builds a continuum between individual selection, kin selection, kin group selection and group selection without a clear boundary for each level. However, early theoretical models by D.S. Wilson et al. (1992) and Taylor (1992) showed that pure population viscosity cannot lead to cooperation and altruism. This is because any benefit generated by kin cooperation is exactly cancelled out by kin competition; additional offspring  from cooperation are eliminated by local competition. However, this exact cancelling out also suggests that any additional benefit of local cooperation would be sufficient to allow cooperation to evolve. Mitteldorf and D.S. Wilson (2000) later showed that if the population is allowed to fluctuate, then local populations can temporarily store the benefit of local cooperation and promote the evolution of cooperation and altruism. By assuming individual differences in adaptations, Yang (2013) further showed that the benefit of local altruism can be stored in the form of offspring quality and thus promote the evolution of altruism even if the population does not fluctuate. This is because local competition among more individuals resulting from local altruism increases the average local fitness of the individuals that survive.

text: E. coli, a Gram-negative proteobacterium, is a common model in biological research, for which comprehensive data on various aspects of its life-history is available. It has been used extensively for evolution experiments, including those related to coevolution with phages. These studies revealed - among others - that coevolutionary adaptation may be influenced by pleiotropic effects of the involved genes. In particular, binding of the bacteriophage to E. coli surface receptor is the crucial step in the virus infection cycle. A mutation in the receptor's binding site may cause resistance. Such mutations often show pleiotropic effects and may cause a cost of resistance. In the presence of phages, such pleiotropy may lead to polymorphisms in the bacterial population and thus enhance biodiversity in the community.Another model system consists of the plant- and animal-colonizing bacterium Pseudomonas and its bacteriophages. This system provided new insights into the dynamics of coevolutionary change. It demonstrated that coevolution may proceed via recurrent selective sweeps, favouring generalists for both antagonists. Furthermore, coevolution with phages may promote allopatric diversity, potentially enhancing biodiversity and possibly speciation.<ref>

text: E. coli

text: , a

text: Gram-negative

text: proteobacterium

text: , is a common model in biological research, for which comprehensive data on various aspects of its life-history is available. It has been used extensively for

text: evolution

text: experiments, including those related to

text: coevolution

text: with phages. These studies revealed - among others - that coevolutionary

text: adaptation

text: may be influenced by

text: pleiotropic

text: effects of the involved

text: genes

text: . In particular, binding of the

text: bacteriophage

text: to E. coli surface

text: receptor

text: is the crucial step in the

text: virus

text: infection

text: cycle. A

text: mutation

text: in the receptor's binding site may cause

text: resistance

text: . Such mutations often show pleiotropic effects and may cause a cost of resistance. In the presence of phages, such pleiotropy may lead to

text: polymorphisms

text: in the bacterial population and thus enhance

text: biodiversity

text: in the community.Another model system consists of the plant- and animal-colonizing bacterium

text: Pseudomonas

text: and its

text: bacteriophages

text: . This system provided new insights into the dynamics of coevolutionary change. It demonstrated that coevolution may proceed via recurrent

text: selective sweeps

text: , favouring

text: generalists

text: for both antagonists. Furthermore, coevolution with phages may promote

text: allopatric

text: diversity, potentially enhancing biodiversity and possibly

text: speciation

text: .<ref>

text: Genes are made of DNA. DNA is  a long molecule made up of individual molecules called nucleotides. Genetic information is encoded in the sequence of nucleotides, that make up the DNA, just as the sequence of the letters in words carries information on a page. The genes are like short instructions built up of the "letters" of the DNA alphabet. Put together, the entire set of these genes gives enough information to serve as an "instruction manual" of how to build and run an organism. The instructions spelled out by this DNA alphabet can be changed, however, by mutations, and this may alter the instructions carried within the genes. Within the cell, the genes are carried in chromosomes, which are packages for carrying the DNA. It is the reshuffling of the chromosomes that results in unique combinations of genes in offspring. Since genes interact with one another during the development of an organism, novel combinations of genes produced by sexual reproduction can increase the genetic variability of the population even without new mutations. The genetic variability of a population can also increase when members of that population interbreed with individuals from a different population causing gene flow between the populations. This can introduce genes into a population that were not present before.

text: Genes are made of

text: DNA

text: . DNA is  a long

text: molecule

text: made up of individual molecules called

text: nucleotide

text: s. Genetic information is encoded in the sequence of nucleotides, that make up the DNA, just as the sequence of the letters in words carries information on a page. The genes are like short instructions built up of the "letters" of the DNA alphabet. Put together, the entire set of these genes gives enough information to serve as an "instruction manual" of how to build and run an organism. The instructions spelled out by this DNA alphabet can be changed, however, by mutations, and this may alter the instructions carried within the genes. Within the

text: cell

text: , the genes are carried in

text: chromosome

text: s, which are packages for carrying the DNA. It is the reshuffling of the chromosomes that results in unique combinations of genes in offspring. Since genes interact with one another during the development of an organism, novel combinations of genes produced by sexual reproduction can increase the genetic variability of the population even without new mutations. The genetic variability of a population can also increase when members of that population

text: interbreed

text: with individuals from a different population causing

text: gene flow

text: between the populations. This can introduce genes into a population that were not present before.

text: Memetics, which comes from the meme idea described in Dawkins's The Selfish Gene, is similar to DIT in that it treats culture as an evolutionary process that is distinct from genetic transmission.  However, there are some philosophical differences between memetics and DIT.   One difference is that memetics' focus is on the selection potential of discrete replicators (memes), where DIT allows for transmission of both non-replicators and non-discrete cultural variants.  DIT does not assume that replicators are necessary for cumulative adaptive evolution. DIT also more strongly emphasizes the role of genetic inheritance in shaping the capacity for cultural evolution.  But perhaps the biggest difference is a difference in academic lineage. Memetics as a label is more influential in popular culture than in academia. Critics of memetics argue that it is lacking in empirical support or is conceptually ill-founded, and question whether there is hope for the memetic research program succeeding. Proponents point out that many cultural traits are discrete, and that many existing models of cultural inheritance assume discrete cultural units, and hence involve memes.

text: Memetics

text: , which comes from the

text: meme

text: idea described in

text: Dawkins's

text: The Selfish Gene

text: , is similar to DIT in that it treats culture as an evolutionary process that is distinct from genetic transmission.  However, there are some philosophical differences between memetics and DIT.   One difference is that memetics' focus is on the selection potential of discrete replicators (memes), where DIT allows for transmission of both non-replicators and non-discrete cultural variants.  DIT does not assume that replicators are necessary for cumulative adaptive evolution. DIT also more strongly emphasizes the role of genetic inheritance in shaping the capacity for cultural evolution.  But perhaps the biggest difference is a difference in academic lineage.

text: Memetics

text: as a label is more influential in popular culture than in academia. Critics of memetics argue that it is lacking in empirical support or is conceptually ill-founded, and question whether there is hope for the memetic research program succeeding. Proponents point out that many cultural traits are discrete, and that many existing models of cultural inheritance assume discrete cultural units, and hence involve memes.

text: Because she is such a big member of the human evolution debate, a lot of her research centers around human evolution and modern human behavior. Modern human behavior is generally thought of as beginning in Europe with the end of the Last Glacial Maximum, but Brooks thinks differently. She and Sally McBrearty feel that there is a fatal flaw with this direction of thinking. By thinking that modern human behavior evolved anywhere other than in Africa totally erases a fundamental part of African history from the archaeological record and also from the history of every human alive today. All the categories to divide time up have come from Europeans, but these frameworks often leave out key events in African history that cannot be fit into these categories. The first Europeans were Africans after the second Out of Africa movement. Homo sapiens are present in Africa 130,000 years ago (190,000 pending investigation). Homo sapiens do not appear in Europe or Asia until 40,000 years ago (the absolute earliest in Levant may be 80-120,000 years ago) and not until 62,000 years ago were then found in Australia. It is also shown that the area of inhabitable land in Africa that could have fostered human evolution is much larger than that of Europe or Asia. Their theory is that the adaption of modern human behavior was not a sudden change that was adapted by all Homo sapiens at the same time, but rather was transferred from human to human over time through Africa and out to Europe and Asia. She argues that modern human behavior was visible in Africa in the Middle Stone Age, way before it was visible in Europe. Brooks and McBrearty explain to us what it means to have modern human behavior and the evidence that has been found in Africa that dates back to the Middle Stone Age. In their paper, Brooks and McBrearty show that the evolution of humans was not a European revolution that suddenly overtook the world and replaced everything else but rather a gradual shift from Africa and out.

text: Because she is such a big member of the

text: human evolution

text: debate, a lot of her research centers around

text: human evolution

text: and

text: modern human behavior

text: Modern human behavior

text: is generally thought of as beginning in

text: Europe

text: with the end of the

text: Last Glacial Maximum

text: , but Brooks thinks differently. She and

text: Sally McBrearty

text: feel that there is a fatal flaw with this direction of thinking. By thinking that

text: modern human behavior

text: evolved

text: anywhere other than in

text: Africa

text: totally erases a fundamental part of

text: African history

text: from the

text: archaeological record

text: and also from the history of every human alive today. All the categories to divide time up have come from

text: Europeans

text: , but these frameworks often leave out key events in

text: African history

text: that cannot be fit into these categories. The first

text: Europeans

text: were

text: Africans

text: after the second

text: Out of Africa

text: movement.

text: Homo sapiens

text: are present in

text: Africa

text: 130,000 years ago (190,000 pending investigation).

text: Homo sapiens

text: do not appear in

text: Europe

text: or

text: Asia

text: until 40,000 years ago (the absolute earliest in Levant may be 80-120,000 years ago) and not until 62,000 years ago were then found in

text: Australia

text: . It is also shown that the area of inhabitable land in

text: Africa

text: that could have fostered

text: human evolution

text: is much larger than that of

text: Europe

text: or

text: Asia

text: . Their theory is that the adaption of

text: modern human behavior

text: was not a sudden change that was adapted by all

text: Homo sapiens

text: at the same time, but rather was transferred from

text: human

text: to

text: human

text: over time through

text: Africa

text: and out to

text: Europe

text: and

text: Asia

text: . She argues that

text: modern human behavior

text: was visible in

text: Africa

text: in the

text: Middle Stone Age

text: , way before it was visible in

text: Europe

text: . Brooks and

text: McBrearty

text: explain to us what it means to have

text: modern human behavior

text: and the evidence that has been found in

text: Africa

text: that dates back to the

text: Middle Stone Age

text: . In their paper, Brooks and

text: McBrearty

text: show that the

text: evolution of humans

text: was not a

text: European

text: revolution

text: that suddenly overtook the world and replaced everything else but rather a gradual shift from

text: Africa

text: and out.

text: While various religions and denominations have reconciled their beliefs with evolution through concepts such as theistic evolution, there are creationists who believe that evolution is contradicted by the creation myths found in their religions and who raise various objections to evolution. As had been demonstrated by responses to the publication of Vestiges of the Natural History of Creation in 1844, the most controversial aspect of evolutionary biology is the implication of human evolution that humans share common ancestry with apes and that the mental and moral faculties of humanity have the same types of natural causes as other inherited traits in animals. In some countries, notably the United States, these tensions between science and religion have fuelled the current creation–evolution controversy, a religious conflict focusing on politics and public education. While other scientific fields such as cosmology and Earth science also conflict with literal interpretations of many religious texts, evolutionary biology experiences significantly more opposition from religious literalists.

text: While

entity_section: Support for evolution by religious bodies

text: various religions and denominations

text: have reconciled their beliefs with evolution through concepts such as

text: theistic evolution

text: , there are

text: creationists

text: who believe that evolution is contradicted by the

text: creation myth

text: s found in their

text: religion

text: s and who raise various

text: objections to evolution

text: . As had been demonstrated by responses to the publication of

entity_name: Vestiges of the Natural History of Creation

text: Vestiges of the Natural History of Creation

text: in 1844, the most controversial aspect of evolutionary biology is the implication of

text: human evolution

text: that humans share common ancestry with apes and that the mental and

text: moral faculties

text: of humanity have the same types of natural causes as other inherited traits in animals. In some countries, notably the United States, these tensions between science and religion have fuelled the current creation–evolution controversy, a religious conflict focusing on

text: politics

text: and

entity_name: Creation and evolution in public education

text: public education

text: . While other scientific fields such as

text: cosmology

text: and

text: Earth science

text: also conflict with literal interpretations of many

text: religious text

text: s, evolutionary biology experiences significantly more opposition from religious literalists.

text: Modern descriptions of biological evolution will typically elaborate on major contributing factors to evolution such as the formation of local micro-environments, mutational robustness, molecular degeneracy, and cryptic genetic variation.  Many of these contributing factors in evolution have been isolated and described for cancer.

text: Modern descriptions of biological evolution will typically elaborate on major contributing factors to evolution such as the formation of local micro-environments, mutational robustness, molecular

text: degeneracy

text: , and cryptic genetic variation.  Many of these contributing factors in evolution have been isolated and described for cancer.

text: Speciation events are important in the theory of punctuated equilibrium, which accounts for the pattern in the fossil record of short "bursts" of evolution interspersed with relatively long periods of stasis, where species remain relatively unchanged. In this theory, speciation and rapid evolution are linked, with natural selection and genetic drift acting most strongly on organisms undergoing speciation in novel habitats or small populations. As a result, the periods of stasis in the fossil record correspond to the parental population and the organisms undergoing speciation and rapid evolution are found in small populations or geographically restricted habitats and therefore rarely being preserved as fossils.

text: Speciation events are important in the theory of

text: punctuated equilibrium

text: , which accounts for the pattern in the fossil record of short "bursts" of evolution interspersed with relatively long periods of stasis, where species remain relatively unchanged. In this theory, speciation and rapid evolution are linked, with natural selection and genetic drift acting most strongly on organisms undergoing speciation in novel habitats or small populations. As a result, the periods of stasis in the fossil record correspond to the parental population and the organisms undergoing speciation and rapid evolution are found in small populations or geographically restricted habitats and therefore rarely being preserved as

text: fossil

text: s.

text: There is currently no consensus about the extent to which capacitance might contribute to evolution in natural populations.

text: There is currently no

text: consensus

text: about the extent to which capacitance might contribute to evolution in natural populations.

text: Wagner showed that robustness can accelerate innovation in biological evolution, because it helps organisms tolerate otherwise deleterious mutations that can help create new and useful traits. In this way, robust transcription factor binding sites, for example, can facilitate the evolution of new gene expression. An additional consequence of robustness is that evolving populations of organisms can accumulate cryptic genetic variation, inconsequential variation that may provide benefits in some environments.  Wagner’s laboratory showed experimentally that such cryptic variation can indeed accelerate the evolution of an RNA enzyme to react with a new substrate molecule. Wagner has argued that robustness can also help resolve the long-standing neutralism-selectionism controversy, which revolves around the question whether frequent neutral mutations – a consequence of robustness – are important for Darwinian evolution. The reason is that neutral mutations are important stepping stones to later evolutionary adaptations and innovations. Robust systems can also bring forth useful traits – potential exaptations – that arise as mere by-products of other, adaptive traits, which can help explain the great abundance of exaptations in life’s evolution.

text: Wagner showed that robustness can accelerate

text: innovation

text: in biological evolution, because it helps organisms tolerate otherwise deleterious mutations that can help create new and useful traits. In this way, robust

text: transcription factor binding site

text: s, for example, can facilitate the evolution of new

text: gene expression

text: . An additional consequence of robustness is that evolving populations of organisms can accumulate

text: cryptic genetic variation

text: , inconsequential variation that may provide benefits in some environments.  Wagner’s laboratory showed experimentally that such cryptic variation can indeed accelerate the evolution of an

text: RNA enzyme

text: to react with a new substrate molecule. Wagner has argued that robustness can also help resolve the long-standing neutralism-selectionism controversy, which revolves around the question whether frequent

text: neutral mutation

text: s – a consequence of robustness – are important for Darwinian evolution. The reason is that neutral mutations are important stepping stones to later

text: evolutionary adaptations

text: and innovations. Robust systems can also bring forth useful traits – potential

text: exaptation

text: s – that arise as mere by-products of other, adaptive traits, which can help explain the great abundance of exaptations in life’s evolution.

text: That mutations are unpredictable, faithfully replicated, and that natural selection operates only upon the products of chance is repeated at the start of chapter seven entitled "Evolution". Monod states that the decisive factor in natural selection is not the "struggle for life" but is the differential rate of reproduction and the only mutations "acceptable" to an organism are those that "do not lessen the coherence of the teleonomic apparatus, but rather, further strengthen it in its already assumed orientation" (Monod, 119). Monod explains that the teleonomic performance is judged through natural selection and this system retains only a very small fraction of mutations that will perfect and enrich the teleonomic apparatus. Monod gives the example of antibody development to show how chance combinations can give a well defined solution. He states that the source of information for the antibodies associative structure is not the antigen itself but is instead the result of many random recombinations of part of the antibody gene. The antibody that is able to bind to the antigen is multiplied. This remarkable example shows chance as the basis for one of the most precise adaptation phenomena. Monod makes the point that selection of a mutation is due to the environmental surroundings of the organism and the teleonomic performances. He then gives some examples to show the interconnection of specific performances/behaviors and anatomical adaptations. The author spends the rest of the chapter discussing linguistic and physical human evolutionary development. Language is an utterly different from the various auditory, tactile, and visual forms of communication in that it allows the communication of an original personal association to another individual. Monod hypothesizes that language was not merely the product but one of the driving forces for the evolution of our central nervous system. He believes that rudimentary symbolic communication appeared early on and created a new selective pressure that favored development of linguistic ability and hence the brain. He then talks about the evolution of our ancestors including the development of upright posture which allowed them to become hunters. Monod lastly points out the evidence to suggest the development of the cognitive function of language in children depends upon postnatal growth of the cortex.

text: Culture & Social Cognition is the relationship between human culture and human cognitive capabilities.  Cultural cognitive evolution proposes that humans’ unique cognitive capacities are not solely due to biological inheritance, but are in fact due in large part to cultural transmission and evolution (Tomasello, 1999). Modern humans and great apes are separated evolutionarily by about six million years. Proponents of cultural evolution argue that this would not have been enough time for humans to develop the advanced cognitive capabilities required to create tools, language, and build societies through biological evolution. Biological evolution could not have individually produced each of these cognitive capabilities within that period of time. Instead, humans must have evolved the capacity to learn through cultural transmission (Tomasello, 1999). This provides a more plausible explanation that would fit within the given time frame. Instead of having to biologically account for each cognitive mechanism that distinguishes modern humans from previous relatives, one would only have to account for one significant biological adaptation for cultural learning. According to this view, the ability to learn through cultural transmission is what distinguishes humans from other primates (Tomasello, 1999). Cultural learning allows humans to build on existing knowledge and make collective advancements, also known as the “ratchet effect”. The ratchet effect simply refers to the way in which humans continuously add on to existing knowledge through modifications and improvements.  This unique ability distinguishes humans from related primates, who do not seem to build collaborative knowledge over time. Instead, primates seem to build individual knowledge, in which the expertise of one animal is not built on by others, and does not progress across time.

text: Culture & Social Cognition is the relationship between human culture and human

text: cognitive

text: capabilities.  Cultural cognitive

text: evolution

text: proposes that humans’ unique cognitive capacities are not solely due to biological inheritance, but are in fact due in large part to cultural transmission and evolution (Tomasello, 1999). Modern humans and

text: great apes

text: are separated evolutionarily by about six million years. Proponents of cultural evolution argue that this would not have been enough time for humans to develop the advanced cognitive capabilities required to create tools, language, and build societies through biological evolution. Biological evolution could not have individually produced each of these cognitive capabilities within that period of time. Instead, humans must have evolved the capacity to learn through cultural transmission (Tomasello, 1999). This provides a more plausible explanation that would fit within the given time frame. Instead of having to biologically account for each cognitive mechanism that distinguishes modern humans from previous relatives, one would only have to account for one significant biological adaptation for cultural learning. According to this view, the ability to learn through cultural transmission is what distinguishes humans from other primates (Tomasello, 1999). Cultural learning allows humans to build on existing knowledge and make collective advancements, also known as the “

text: ratchet effect

text: ”. The ratchet effect simply refers to the way in which humans continuously add on to existing knowledge through modifications and improvements.  This unique ability distinguishes humans from related primates, who do not seem to build collaborative knowledge over time. Instead, primates seem to build individual knowledge, in which the expertise of one animal is not built on by others, and does not progress across time.

text: The term evolution (from its literal meaning of "unfolding" of something into its true or explicit form) carries a connotation of gradual improvement or directionality from a beginning to an end point. This contrasts with the more general development, which can indicate change in any direction, or revolution, which implies recurring, periodic change. The term  biological devolution is coined as an antonym to evolution, indicating such degeneration or decrease in quality or complexity.

text: The term evolution (from its literal meaning of "unfolding" of something into its true or explicit form) carries a connotation of gradual improvement or directionality from a beginning to an end point. This contrasts with the more general

text: development

text: , which can indicate change in any direction, or

text: revolution

text: , which implies recurring, periodic change. The term

text: biological devolution

text: is coined as an antonym to evolution, indicating such degeneration or decrease in quality or complexity.

text: Wagner's gene network model is a computational model of artificial gene networks, which explicitly modeled the developmental and evolutionary process of genetic regulatory networks. A population with multiple organisms can be created and evolved from generation to generation. It was first developed by Andreas Wagner in 1996 and has been investigated by other groups to study the evolution of gene networks, gene expression, robustness, plasticity and epistasis.

text: Wagner's gene network model is a computational model of artificial gene networks, which explicitly modeled the developmental and evolutionary process of

text: genetic regulatory networks

text: . A

text: population

text: with multiple organisms can be created and evolved from generation to generation. It was first developed by

text: Andreas Wagner

text: in 1996 and has been investigated by other groups to study the evolution of

text: gene network

text: s,

text: gene expression

text: robustness

text: plasticity

text: and

text: epistasis

text: The rate of adaptive evolution in the human genome has often assumed to be constant over time. For example, the 35% estimate for α calculated by Fay et al. (2001) led them to conclude that there was one adaptive substitution in the human lineage every 200 years since human divergence from old-world monkeys. However, even if the original value of α is accurate for a particular time period, this extrapolation is still invalid. This is because there has been a large acceleration in the amount of positive selection in the human lineage over the last 40,000 years, in terms of the number of genes that have undergone adaptive evolution (Hawks et al. 2007). This agrees with simple theoretical predictions, because the human population size has expanded massively in the last 40,000 years, and with more people, there should be more adaptive substitutions. Hawks et al. (2007) argue that demographic changes (particularly population expansion) may greatly facilitate adaptive evolution, an argument that corroborates somewhat with the positive correlation inferred between population size and amount of adaptive evolution occurring mentioned previously.It has been suggested that cultural evolution may have replaced genetic evolution, and hence slowed the rate of adaptive evolution over the past 10,000 years. However, it is possible that cultural evolution could actually increase genetic adaption. Cultural evolution has vastly increased communication and contact between different populations, and this provides much greater opportunities for genetic admixture between the different populations (Hawks et al. 2007). However, recent cultural phenomena, such as modern medicine and the smaller variation in modern family sizes, may reduce genetic adaption as natural selection is relaxed, overriding the increased potential for adaptation due to greater genetic admixture.

text: If all genes are in linkage equilibrium, the effect of an allele at one locus can be averaged across the gene pool at other loci. In reality, one allele is frequently found in linkage disequilibrium with genes at other loci, especially with genes located nearby on the same chromosome. Recombination breaks up this linkage disequilibrium too slowly to avoid genetic hitchhiking, where an allele at one locus rises to high frequency because it is linked to an allele under selection at a nearby locus. Linkage also slows down the rate of adaptation, even in sexual populations. The effect of linkage disequilibrium in slowing down the rate of adaptive evolution arises from a combination of the Hill-Robertson effect (delays in bringing beneficial mutations together) and background selection (delays in separating beneficial mutations from deleterious hitchhikers).

text: If all genes are in

text: linkage equilibrium

text: , the effect of an allele at one locus can be averaged across the

text: gene pool

text: at other loci. In reality, one allele is frequently found in

text: linkage disequilibrium

text: with genes at other loci, especially with genes located nearby on the same chromosome.

text: Recombination

text: breaks up this linkage disequilibrium too slowly to avoid

text: genetic hitchhiking

text: , where an allele at one locus rises to high frequency because it is

text: linked

text: to an allele under selection at a nearby locus. Linkage also slows down the rate of adaptation, even in sexual populations. The effect of linkage disequilibrium in slowing down the rate of adaptive evolution arises from a combination of the

text: Hill-Robertson effect

text: (delays in bringing beneficial mutations together) and

text: background selection

text: (delays in separating beneficial mutations from deleterious

text: hitchhikers

text: Farming, which, the authors note, produces 10 to 100 times more calories per acre than foraging, carried this trend further. Over the period from 10,000 BC to AD 1, the world population increased about a hundredfold - estimates range from 40 to 170 times. An accelerated rate of evolution is a direct result of the larger human population. More people will have more mutations, thereby increasing opportunity for evolutionary change under natural selection. The spread of rapidly expanding populations eventually outpaced the spread of favourable mutations under selection in those populations, so for the ﬁrst time in human history favourable mutations could not fully disperse throughout the human species. In addition, of course, selection pressures changed once farming was adopted, favouring distinctive adaptations in different geographic areas.

text: Farming, which, the authors note, produces 10 to 100 times more calories per acre than foraging, carried this trend further. Over the period from 10,000 BC to AD 1, the world population increased about a hundredfold - estimates range from 40 to 170 times. An accelerated rate of evolution is a direct result of the larger human population. More people will have more

text: mutation

text: s, thereby increasing opportunity for evolutionary change under natural selection. The spread of rapidly expanding populations eventually outpaced the spread of favourable mutations under selection in those populations, so for the ﬁrst time in human history favourable mutations could not fully disperse throughout the human species. In addition, of course, selection pressures changed once farming was adopted, favouring distinctive adaptations in different geographic areas.

text: Dog evolution (from the Latin evolutio: "unrolling") is the biological descent with modification that led to the domestic dog. This process encompasses small-scale evolution (changes in gene frequency in a population from one generation to the next) and large-scale evolution (the descent of different species from a common ancestor over many generations).

text: On some occasions, DNA damage is not repaired, or is repaired by an error-prone mechanism that results in a change from the original sequence. When this occurs, mutations may propagate into the genomes of the cell's progeny. Should such an event occur in a germ line cell that will eventually produce a gamete, the mutation has the potential to be passed on to the organism's offspring. The rate of evolution in a particular species (or, in a particular gene) is a function of the rate of mutation. As a consequence, the rate and accuracy of DNA repair mechanisms have an influence over the process of evolutionary change. Since the normal adaptation of populations of organisms to changing circumstances (for instance the adaptation of the beaks of a population of finches to the changing presence of hard seeds or insects) proceeds by gene regulation and the recombination and selection of gene variations  – alleles  – and not by passing on irreparable DNA damages to the offspring, DNA damage protection and repair does not influence the rate of adaptation by gene regulation and by recombination and selection of alleles.  On the other hand, DNA damage repair and protection does influence the rate of accumulation of irreparable, advantageous, code expanding, inheritable mutations, and slows down the evolutionary mechanism for expansion of  the genome of organisms with new functionalities. The tension between evolvability and mutation repair and protection needs further investigation.

text: On some occasions, DNA damage is not repaired, or is repaired by an error-prone mechanism that results in a change from the original sequence. When this occurs,

text: mutation

text: s may propagate into the genomes of the cell's progeny. Should such an event occur in a

text: germ line

text: cell that will eventually produce a

text: gamete

text: , the mutation has the potential to be passed on to the organism's offspring. The rate of

text: evolution

text: in a particular species (or, in a particular gene) is a function of the rate of mutation. As a consequence, the rate and accuracy of DNA repair mechanisms have an influence over the process of evolutionary change. Since the normal adaptation of populations of organisms to changing circumstances (for instance the adaptation of the beaks of a population of finches to the changing presence of hard seeds or insects) proceeds by gene regulation and the recombination and selection of gene variations  – alleles  – and not by passing on irreparable DNA damages to the offspring, DNA damage protection and repair does not influence the rate of adaptation by gene regulation and by recombination and selection of alleles.  On the other hand, DNA damage repair and protection does influence the rate of accumulation of irreparable, advantageous, code expanding, inheritable mutations, and slows down the evolutionary mechanism for expansion of  the genome of organisms with new functionalities. The tension between evolvability and mutation repair and protection needs further investigation.

text: Cambrian explosion – camouflage – Sean B. Carroll – catagenesis – gene-centered view of evolution – cephalization – Sergei Chetverikov – chronobiology – chronospecies – clade – cladistics – climatic adaptation – coalescent theory – co-evolution – co-operation – coefficient of relationship – common descent – convergent evolution – creation-evolution controversy – cultivar – conspecific song preference

text: Cambrian explosion

text: camouflage

text: Sean B. Carroll

text: catagenesis

text: gene-centered view of evolution

text: cephalization

text: Sergei Chetverikov

text: chronobiology

text: chronospecies

text: clade

text: cladistics

text: climatic adaptation

text: coalescent theory

text: co-evolution

text: co-operation

text: coefficient of relationship

text: common descent

text: convergent evolution

text: creation-evolution controversy

text: cultivar

text: conspecific song preference

text: The evolution of the MADS-box family has been extensively studied. These genes are present even in pteridophytes, but the spread and diversity is many times higher in angiosperms. There appears to be quite a bit of pattern into how this family has evolved. Consider the evolution of the C-region gene AGAMOUS (AG). It is expressed in today's flowers in the stamens, and the carpel, which are reproductive organs. It's ancestor in gymnosperms also has the same expression pattern. Here, it is expressed in the strobili, an organ that produces pollens or ovules. Similarly, the B-genes' (AP3 and PI) ancestors are expressed only in the male organs in gymnosperms. Their descendants in the modern angiosperms also are expressed only in the stamens, the male reproductive organ. Thus, the same, then-existing components were used by the plants in a novel manner to generate the first flower. This is a recurring pattern in evolution.

text: The evolution of the

text: MADS-box

text: family has been extensively studied. These genes are present even in

text: pteridophytes

text: , but the spread and diversity is many times higher in

text: angiosperms

text: . There appears to be quite a bit of pattern into how this family has evolved. Consider the evolution of the C-region gene

text: AGAMOUS

text: (AG). It is expressed in today's flowers in the

text: stamens

text: , and the

text: carpel

text: , which are reproductive organs. It's ancestor in

text: gymnosperms

text: also has the same expression pattern. Here, it is expressed in the

text: strobili

text: , an organ that produces

text: pollens

text: or ovules. Similarly, the B-genes' (AP3 and PI) ancestors are expressed only in the male organs in

text: gymnosperms

text: . Their descendants in the modern angiosperms also are expressed only in the

text: stamens

text: , the male reproductive organ. Thus, the same, then-existing components were used by the plants in a novel manner to generate the first flower. This is a recurring pattern in

text: evolution

text: Spencer's attempt to explain the evolution of complexity was radically different from that to be found in Darwin's Origin of Species which was published two years later. Spencer is often, quite erroneously, believed to have merely appropriated and generalised Darwin's work on natural selection. But although after reading Darwin's work he coined the phrase 'survival of the fittest' as his own term for Darwin's concept, and is often misrepresented as a thinker who merely applied the Darwinian theory to society, he only grudgingly incorporated natural selection into his preexisting overall system. The primary mechanism of species transformation that he recognised was Lamarckian use-inheritance which posited that organs are developed or are diminished by use or disuse and that the resulting changes may be transmitted to future generations. Spencer believed that this evolutionary mechanism was also necessary to explain 'higher' evolution, especially the social development of humanity. Moreover, in contrast to Darwin, he held that evolution had a direction and an end-point, the attainment of a final state of equilibrium. He tried to apply the theory of biological evolution to sociology. He proposed that society was the product of change from lower to higher forms, just as in the theory of biological evolution, the lowest forms of life are said to be evolving into higher forms. Spencer claimed that man's mind had evolved in the same way from the simple automatic responses of lower animals to the process of reasoning in the thinking man. Spencer believed in two kinds of knowledge: knowledge gained by the individual and knowledge gained by the race. Intuition, or knowledge learned unconsciously, was the inherited experience of the race.

text: Spencer's attempt to explain the

text: evolution of complexity

text: was radically different from that to be found in Darwin's

text: Origin of Species

text: which was published two years later. Spencer is often, quite erroneously, believed to have merely appropriated and generalised Darwin's work on

text: natural selection

text: . But although after reading Darwin's work he coined the phrase '

text: survival of the fittest

text: ' as his own term for Darwin's concept, and is often misrepresented as a thinker who merely applied the Darwinian theory to society, he only grudgingly incorporated natural selection into his preexisting overall system. The primary mechanism of species transformation that he recognised was

text: Lamarckian

text: use-inheritance which posited that organs are developed or are diminished by use or disuse and that the resulting changes may be transmitted to future generations. Spencer believed that this evolutionary mechanism was also necessary to explain 'higher' evolution, especially the social development of humanity. Moreover, in contrast to Darwin, he held that evolution had a direction and an end-point, the attainment of a final state of equilibrium. He tried to apply the theory of biological evolution to sociology. He proposed that society was the product of change from lower to higher forms, just as in the theory of biological evolution, the lowest forms of life are said to be evolving into higher forms. Spencer claimed that man's mind had evolved in the same way from the simple automatic responses of lower animals to the process of reasoning in the thinking man. Spencer believed in two kinds of knowledge: knowledge gained by the individual and knowledge gained by the race. Intuition, or knowledge learned unconsciously, was the inherited experience of the race.

text: Although the number of base-pairs of DNA in the human genome has been known since the 1960s, the estimated number of genes has changed over time as definitions of genes, and methods of detecting them have been refined. Initial theoretical predictions of the number of human genes were as high as 2,000,000. Early experimental measures indicated there to be 50,000–100,000 transcribed genes (expressed sequence tags). Subsequently, the sequencing in the Human Genome Project indicated that many of these transcripts were alternative variants of the same genes, and the total number of protein-coding genes was revised down to ~20,000 with 13 genes encoded on the mitochondrial genome. Of the human genome, only 1–2% consists of protein-coding genes, with the remainder being 'noncoding' DNA such as introns, retrotransposons, and noncoding RNAs. Every multicellular organism has all its genes in each cell of its body but not every gene functions in every cell .

text: Although the number of base-pairs of DNA in the human genome has been known since the 1960s, the estimated number of genes has changed over time as definitions of genes, and methods of detecting them have been refined. Initial theoretical predictions of the number of human genes were as high as 2,000,000. Early experimental measures indicated there to be 50,000–100,000 transcribed genes (

text: expressed sequence tag

text: s). Subsequently, the sequencing in the

text: Human Genome Project

text: indicated that many of these transcripts were

text: alternative variants

text: of the same genes, and the total number of protein-coding genes was revised down to ~20,000 with 13 genes encoded on the

text: mitochondria

text: l genome. Of the human genome, only 1–2% consists of protein-coding genes, with the remainder being

text: 'noncoding' DNA

text: such as

text: intron

text: s,

text: retrotransposon

text: s, and

text: noncoding RNA

text: s. Every multicellular organism has all its genes in each cell of its body but not every gene functions in every cell .

text: This section contains a list of works on evolution, the change across successive generations in the heritable characteristics of biological populations.

text: This section contains a list of works on

text: evolution

text: , the change across successive

text: generation

text: s in the

text: heritable

text: characteristics

text: of

text: biological

text: population

text: s.

text: A lineage is expected to evolve to maximise its inclusive fitness—the number of copies of its genes passed on globally (rather than by a particular individual). As a result, populations will tend towards an evolutionarily stable strategy. The book also coins the term meme for a unit of human cultural evolution analogous to the gene, suggesting that such "selfish" replication may also model human culture, in a different sense. Memetics has become the subject of many studies since the publication of the book.  In raising awareness of the ideas of W. D. Hamilton, as well as making its own valuable contributions to the field, the book has also stimulated research on human inclusive fitness.

text: A lineage is expected to evolve to maximise its

text: inclusive fitness

text: —the number of copies of its genes passed on globally (rather than by a particular individual). As a result, populations will tend towards an

text: evolutionarily stable strategy

text: . The book also coins the term

text: meme

text: for a unit of human

text: cultural evolution

text: analogous to the gene, suggesting that such "selfish" replication may also model human culture, in a different sense.

text: Memetics

text: has become the subject of many studies since the publication of the book.  In raising awareness of the ideas of

text: W. D. Hamilton

text: , as well as making its own valuable contributions to the field, the book has also stimulated research on

text: human inclusive fitness

text: Brook’s elucidation of the concept of art, as what he at first called ‘experimental modelling of actual and possible worlds,’ is first comprehensively spelled out in his paper ‘A new theory of art’. This rather complex theoretical structure underwent continuous modification in numerous publications as his memetic account of cultural evolution developed. He embraced Dawkins’ concept of the meme as the potent factor in cultural evolution, analogous to the function of the gene in biological evolution. However, he differed significantly from Dawkins and other meme-theorists in his way of understanding and exemplifying the meme. Instead of offering such examples as the popular song or catchphrase he insisted that these things are items of a cultural kind analogous to the items of biological kinds such as a horse, or a cabbage. Just as a horse is the product of appropriately activated genes, he argues that a popular song or a catchphrase is not itself a meme. It is an item of a predictable cultural kind that is generated by exercising memes. Memes are purposefully directed actions that can be exercised in concerted ways, as contrasted with mere behaviours. This is first unambiguously elucidated in his paper ‘Art history?’.

text: Brook’s elucidation of the concept of art, as what he at first called ‘experimental modelling of actual and possible worlds,’ is first comprehensively spelled out in his paper ‘A new theory of art’. This rather complex theoretical structure underwent continuous modification in numerous publications as his

text: memetic

text: account of cultural evolution developed. He embraced

text: Dawkins’

text: concept of the

text: meme

text: as the potent factor in cultural evolution, analogous to the function of the

text: gene

text: in biological evolution. However, he differed significantly from Dawkins and other meme-theorists in his way of understanding and exemplifying the meme. Instead of offering such examples as the popular song or catchphrase he insisted that these things are items of a cultural kind analogous to the items of biological kinds such as a horse, or a cabbage. Just as a horse is the product of appropriately activated genes, he argues that a popular song or a catchphrase is not itself a meme. It is an item of a predictable cultural kind that is generated by exercising memes. Memes are purposefully directed actions that can be exercised in concerted ways, as contrasted with mere behaviours. This is first unambiguously elucidated in his paper ‘Art history?’.

text: However, intermixing two different populations may give rise to unfit polygenic traits in outbreeding depression, yielding offspring which lack the genetic adaptations to specific environmental conditions. These, then, will have a lowered fitness than pure-bred individuals e.g. of a particular subspecies that has adapted to its local environment.

text: However, intermixing two different populations may give rise to unfit polygenic traits in

text: outbreeding depression

text: , yielding offspring which lack the genetic

text: adaptation

text: s to specific environmental conditions. These, then, will have a lowered fitness than pure-bred individuals e.g. of a particular

text: subspecies

text: that has adapted to its local environment.

text: In 2003, a drought as bad as the 1977 drought plagued the island. The major difference this time was that large ground finch had become the predominate-size birds. The large ground finch had to compete with the larger-beaked medium ground finch. The large ground finch trumped the medium finch; 90% of the medium ground finch population died due to starvation. Peter Grant said in one paper that the advantage had shifted strongly to the small-beaked members of the medium ground finch population and that evolution by natural selection had occurred again.

text: Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in heritable traits of a population over time. Charles Darwin popularised the term "natural selection", and compared it with  artificial selection.

text: Natural selection is the differential survival and reproduction of individuals due to differences in

text: phenotype

text: . It is a key mechanism of

text: evolution

text: , the change in

text: heritable

text: traits

text: of a

text: population

text: over time.

text: Charles Darwin

text: popularised the term "natural selection", and compared it with

text: artificial selection

text: Wright's explanation for stasis was that organisms come to occupy adaptive peaks. In order to evolve to another, higher peak, the species would first have to pass through a valley of maladaptive intermediate stages. This could happen by genetic drift if the population is small enough. If a species was divided into small populations, some could find higher peaks. If there was some gene flow between the populations, these adaptations could spread to the rest of the species. This was Wright's shifting balance theory of evolution. There has been much skepticism among evolutionary biologists as to whether these rather delicate conditions hold often in natural populations. Wright had a long-standing and bitter debate about this with R. A. Fisher, who felt that most populations in nature were too large for these effects of genetic drift to be important.

text: Wright's explanation for

text: stasis

text: was that organisms come to occupy

text: adaptive peak

text: s. In order to evolve to another, higher peak, the species would first have to pass through a valley of maladaptive intermediate stages. This could happen by

text: genetic drift

text: if the population is small enough. If a species was divided into small populations, some could find higher peaks. If there was some

text: gene flow

text: between the populations, these adaptations could spread to the rest of the species. This was Wright's

text: shifting balance theory

text: of evolution. There has been much skepticism among evolutionary biologists as to whether these rather delicate conditions hold often in natural populations. Wright had a long-standing and bitter debate about this with

text: R. A. Fisher

text: , who felt that most populations in nature were too large for these effects of genetic drift to be important.

text: Hugo de Vries tried to link the new genetics with evolution; building on his work with heredity and hybridization, he proposed a theory of mutationism, which was widely accepted in the early 20th century. Lamarckism also had many adherents. Darwinism was seen as incompatible with the continuously variable traits studied by biometricians, which seemed only partially heritable. In the 1920s and 1930s—following the acceptance of the Mendelian-chromosome theory— the emergence of the discipline of population genetics, with the work of R.A. Fisher, J.B.S. Haldane and Sewall Wright, unified the idea of evolution by natural selection with Mendelian genetics, producing the modern synthesis. The inheritance of acquired characters was rejected, while mutationism gave way as genetic theories matured.

text: Hugo de Vries tried to link the new genetics with evolution; building on his work with heredity and

text: hybridization

text: , he proposed a theory of

text: mutationism

text: , which was widely accepted in the early 20th century.

text: Lamarckism

text: also had many adherents.

text: Darwinism

text: was seen as incompatible with the continuously variable traits studied by

text: biometricians

text: , which seemed only partially heritable. In the 1920s and 1930s—following the acceptance of the Mendelian-chromosome theory— the emergence of the discipline of

text: population genetics

text: , with the work of

text: R.A. Fisher

text: J.B.S. Haldane

text: and

text: Sewall Wright

text: , unified the idea of evolution by

text: natural selection

text: with

text: Mendelian genetics

text: , producing the

text: modern synthesis

text: . The

text: inheritance of acquired characters

text: was rejected, while mutationism gave way as genetic theories matured.

text: Science writer Matt Ridley popularized the term in connection with sexual selection in his 1993 book The Red Queen, in which he discussed the debate in theoretical biology over the adaptive benefit of sexual reproduction to those species in which it appears. The connection of the Red Queen to this debate arises from the fact that the traditionally accepted Vicar of Bray hypothesis only showed adaptive benefit at the level of the species or group, not at the level of the gene (although the protean "Vicar of Bray" adaptation is very useful to some species that belong to the lower levels of the food chain). By contrast, a Red-Queen-type thesis that organisms are running  arms races with their parasites can explain the utility of sexual reproduction at the level of the gene by positing that the role of sex is to preserve genes that are currently disadvantageous, but that will become advantageous against the background of a likely future population of parasites.

text: Science writer

text: Matt Ridley

text: popularized the term in connection with

text: sexual selection

text: in his 1993 book

entity_name: The Red Queen: Sex and the Evolution of Human Nature

text: The Red Queen

text: , in which he discussed the debate in theoretical biology over the adaptive benefit of

text: sexual reproduction

text: to those species in which it appears. The connection of the Red Queen to this debate arises from the fact that the traditionally accepted

text: Vicar of Bray hypothesis

text: only showed adaptive benefit at the level of the species or group, not at the level of the

text: gene

text: (although the protean "Vicar of Bray" adaptation is very useful to some species that belong to the lower levels of the

text: food chain

text: ). By contrast, a Red-Queen-type thesis that organisms are running  arms races with their

text: parasite

text: s can explain the utility of sexual reproduction at the level of the gene by positing that the role of sex is to preserve genes that are currently disadvantageous, but that will become advantageous against the background of a likely future population of parasites.

text: Convergent evolution is a biological concept whereby unrelated species acquire similar traits because they adapted to a similar environment and/or played similar roles in their ecosystems. In fiction, the concept is extended whereby similar planets will result in races with similar cultures and/or histories.

text: Convergent evolution

text: is a biological concept whereby unrelated species acquire similar traits because they adapted to a similar environment and/or played similar roles in their ecosystems. In fiction, the concept is extended whereby similar planets will result in races with similar cultures and/or histories.

text: Thus, and because the adaptation to the peculiar conditions on Socorro may even have accelerated not only morphological but also molecular evolution – see also founder effect -, it cannot be said with any certainty whether or not among Mimus, the Socorro species is a quite recent island offshoot of either of the mainland species. In any case, the three taxa are very close relatives. This serves to show that evolution does not move on at a constant speed; certainly not on a morphological level, and often neither on a molecular level either. Rather, the rate of evolutionary change varies, sometimes considerably, depending on the circumstances and the strength with which natural selection acts upon a founding population.

text: Thus, and because the adaptation to the peculiar conditions on Socorro may even have accelerated not only

text: morphological

text: but also molecular evolution – see also

text: founder effect

text: -, it cannot be said with any certainty whether or not among Mimus, the Socorro species is a quite recent island offshoot of either of the mainland species. In any case, the three

text: taxa

text: are very close relatives. This serves to show that evolution does not move on at a constant speed; certainly not on a morphological level, and often neither on a molecular level either. Rather, the rate of evolutionary change varies, sometimes considerably, depending on the circumstances and the strength with which natural selection acts upon a founding population.

text: The decades before the start of the 20th century, and the first decades of that century, have been described as the eclipse of Darwinism. Darwin's work had quickly established scientific consensus that evolution occurred, but there was considerable disagreement about the mechanisms involved, and few gave as much significance to natural selection as Darwin himself. Evolution itself was assumed, but the mechanism of how it happened was in considerable debate, and none had anything near to a consensus. Among these theories were neo-Lamarckism (which merged certain aspects of Lamarck's theory of acquired characteristics with certain aspects of Darwinian evolution), orthogenesis ("straight-line" evolution, which talked about evolution towards a specific goal by forces within the organism), and the discontinuous variation of Mendelism and Hugo De Vries' mutation theory. Some of these alternative theories, in particular neo-Lamarckism and orthogenesis, allowed more easily for an interpretation of the intervention of God, which appealed to many scientists at the time. The term Darwinism had covered a wide range of ideas, many of which differed from Darwin's views, but it became associated with the minority view of August Weismann who went further than Darwin by rejecting inheritance of acquired characters and attributing all evolution to natural selection, a view also called neo-Darwinism. By the first decades of the 20th century, the debate had become generally one between continuous-variation biometricians and discontinuous-variety Mendelians. In the 1930s and 1940s, though, they were combined with natural selection into the modern evolutionary synthesis, which soon became the dominant model in the scientific community. This model has also been called Darwinism and neo-Darwinism.

text: The decades before the start of the 20th century, and the first decades of that century, have been described as

text: the eclipse of Darwinism

text: . Darwin's work had quickly established scientific consensus that evolution occurred, but there was considerable disagreement about the mechanisms involved, and few gave as much significance to natural selection as Darwin himself. Evolution itself was assumed, but the mechanism of how it happened was in considerable debate, and none had anything near to a consensus. Among these theories were

text: neo-Lamarckism

text: (which merged certain aspects of Lamarck's theory of acquired characteristics with certain aspects of Darwinian evolution),

text: orthogenesis

text: ("straight-line" evolution, which talked about evolution towards a specific goal by forces within the organism), and the discontinuous variation of

text: Mendelism

text: and

text: Hugo De Vries

text: mutation theory

text: . Some of these alternative theories, in particular neo-Lamarckism and orthogenesis, allowed more easily for an interpretation of the intervention of God, which appealed to many scientists at the time. The term

text: Darwinism

text: had covered a wide range of ideas, many of which differed from Darwin's views, but it became associated with the minority view of

text: August Weismann

text: who went further than Darwin by rejecting

text: inheritance of acquired characters

text: and attributing all evolution to natural selection, a view also called

text: neo-Darwinism

text: . By the first decades of the 20th century, the debate had become generally one between continuous-variation biometricians and discontinuous-variety Mendelians. In the 1930s and 1940s, though, they were combined with natural selection into the

text: modern evolutionary synthesis

text: , which soon became the dominant model in the scientific community. This model has also been called Darwinism and neo-Darwinism.

text: Biological evolution can be considered as a form of trial and error. Random mutations and sexual genetic variations can be viewed as trials and poor reproductive fitness, or lack of improved fitness, as the error. Thus after a long time 'knowledge' of well-adapted genomes accumulates simply by virtue of them being able to reproduce.

text: Biological evolution

text: can be considered as a form of trial and error. Random mutations and sexual genetic variations can be viewed as trials and poor reproductive fitness, or lack of improved fitness, as the error. Thus after a long time 'knowledge' of well-adapted genomes accumulates simply by virtue of them being able to reproduce.

text: In a number of works Klejn discusses the idea of the contradiction between modern culture and the nature of man.  This treatment is far from the Rousseau’s mood and is built on the basis of socio-biology. Developing the ideas of Lorenz and Desmond Morris, Klejn’s idea is built upon the fact that in every stage of man’s biological evolution man has been formed by adaptation to conditions not only of the natural environment but also of the socio-cultural milieu. However tempos of socio-cultural evolution are much faster than those of biological evolution: socio-cultural evolution has no need to wait for generations to change. While biological evolution has not terminated even a single stage of homo sapiens, socio-cultural evolution has gone through the Upper Palaeolithic, Mesolithic, Eneolithic, Neolithic, Bronze and Iron Ages, and all epochs of civilization, and has now entered the Atomic and Computer Age. Yet our psycho-physiological characteristics remain the same as in the Stone Age – they were, and are, adapted to those conditions. This accords with Freud’s observation: we are discontent with modern culture because we are not by our nature adapted to it. Hence there are a number of specific deceases and psychic problems that are developing into social ones. Yet in culture a number of compensatory mechanisms exist to diminish this tension. However, when in some milieus there is a lack of culture, from man’s inside there bursts a savage, and thus society, when left to self-organization (as in Lord of the Flies), restores (mutatis mutandis) primordial forms of life. This can be seen in the example of hard labour camps, bullying (by age) among army troops, etc..

text: Nevo received a M.Sc and  PhD (1964)   from Hebrew University His Ph.D thesiss was "Population studies of Anurans from the lower Cretaceous of Makhtesh Ramon, Israel"    He founded the Institute of Evolution at Haifa in 1973. Nevo works with many  evolutionary subjects, involving understanding and tracking of speciation processes, modifications of highly evolved traits (vision), climatic and geographic effects in sympatric speciation in insects, bacteria, fungi, mammals and crops. He is a proponent of evolutionary models where environmental stressors   acting positively to shape genetic polymorphisms. The model of "Evolution Canyon" is a microscale open environment where two opposite slopes with contrasting climatic conditions have also finer biotic adaptations  that have been researched at different biological scales, supporting the evolution in action generating evidence of sympatric speciation, and of particular impact and interest are the works in Drosophila  and in wild barley

text: Nevo received a M.Sc and  PhD (1964)   from

text: Hebrew University

text: His Ph.D thesiss was "Population studies of Anurans from the lower Cretaceous of Makhtesh Ramon, Israel"    He founded the Institute of Evolution at Haifa in 1973. Nevo works with many  evolutionary subjects, involving understanding and tracking of speciation processes, modifications of highly evolved traits (vision), climatic and geographic effects in sympatric speciation in insects, bacteria, fungi, mammals and crops. He is a proponent of evolutionary models where environmental stressors   acting positively to shape genetic polymorphisms. The model of "Evolution Canyon" is a microscale open environment where two opposite slopes with contrasting climatic conditions have also finer biotic adaptations  that have been researched at different biological scales, supporting the evolution in action generating evidence of sympatric speciation, and of particular impact and interest are the works in Drosophila  and in wild barley

text: Evolutionary psychology explores the evolutionary roots of mental and behavioral patterns, and posits that common patterns may have emerged because they were highly adaptive for humans in the environments of their evolutionary past—even if some of these patterns are maladaptive in today's environments. Fields closely related to evolutionary psychology are animal behavioral ecology, human behavioral ecology, dual inheritance theory, and sociobiology. Evolutionary psychology is distinct from, although related to, behavioral genetics. Memetics, founded by Richard Dawkins, is a related but competing field that proposes that cultural evolution can occur in a Darwinian sense but independently of Mendelian mechanisms; it therefore examines the ways in which thoughts, or memes, may evolve independently of genes.

text: Evolutionary psychology

text: explores the evolutionary roots of mental and behavioral patterns, and posits that common patterns may have emerged because they were highly

text: adaptive

text: for humans in the environments of their evolutionary past—even if some of these patterns are maladaptive in today's environments. Fields closely related to evolutionary psychology are animal

text: behavioral ecology

text: human behavioral ecology

text: dual inheritance theory

text: , and

text: sociobiology

text: . Evolutionary psychology is distinct from, although related to,

text: behavioral genetics

text: Memetics

text: , founded by

text: Richard Dawkins

text: , is a related but competing field that proposes that

text: cultural evolution

text: can occur in a

text: Darwinian

text: sense but independently of

text: Mendelian

text: mechanisms; it therefore examines the ways in which thoughts, or

text: meme

text: s, may evolve independently of genes.

text: The difference in gene frequencies between the original population and colony may also trigger the two groups to diverge significantly over the course of many generations. As the difference, or genetic distance, increases, the two separated populations may become distinct, both genetically and phenetically, although not only genetic drift but also natural selection, gene flow, and mutation contribute to this divergence. This potential for relatively rapid changes in the colony's gene frequency led most scientists to consider the founder effect (and by extension, genetic drift) a significant driving force in the evolution of new species. Sewall Wright was the first to attach this significance to random drift and small, newly isolated populations with his shifting balance theory of speciation. Following after Wright, Ernst Mayr created many persuasive models to show that the decline in genetic variation and small population size following the founder effect were critically important for new species to develop. However, there is much less support for this view today since the hypothesis has been tested repeatedly through experimental research and the results have been equivocal at best.

text: The difference in gene frequencies between the original population and colony may also trigger the two groups to

text: diverge

text: significantly over the course of many generations. As the difference, or

text: genetic distance

text: , increases, the two separated populations may become distinct, both genetically and

text: phenetically

text: , although not only genetic drift but also natural selection, gene flow, and mutation contribute to this divergence. This potential for relatively rapid changes in the colony's gene frequency led most scientists to consider the founder effect (and by extension, genetic drift) a significant driving force in the evolution of

text: new species

text: . Sewall Wright was the first to attach this significance to random drift and small, newly isolated populations with his

text: shifting balance theory

text: of speciation. Following after Wright,

text: Ernst Mayr

text: created many persuasive models to show that the decline in genetic variation and small population size following the founder effect were critically important for new species to develop. However, there is much less support for this view today since the hypothesis has been tested repeatedly through experimental research and the results have been equivocal at best.

text: Essynth  is a mathematical model for interactive sound synthesis based on the evolutionary computation which uses genetic operators and fitness functions to create sound. In Essynth, digitally sampled sounds (waveforms) are represented as individuals within a population set that describes an evolutionary path along time in generation steps.  Here the evolution is given by the population’s adaptation to another set of individuals, called here as the Target Set, which conveys all psychoacoustic features that steers the population evolution. This process is somehow similar to the “evolution pressure” that dictates the adaptation of populations in biological systems. The “best individual” within the population is the one most similar to the Target set. Unlike others deterministic sound synthesis methods such as the linear methods (i.e. additive synthesis), the non-linear (i.e. frequency modulation synthesis) and the sample-based methods (i.e. sample-based synthesis) Essynth is a non-deterministic method of sound synthesis once that its resulting sound is not fixed for the same parametric values but continuously evolves along time.

text: Essynth  is a

text: mathematical model

text: for interactive

text: sound

text: synthesis based on the

text: evolutionary computation

text: which uses

text: genetic

text: operators and fitness functions to create sound. In Essynth, digitally sampled sounds (

text: waveforms

text: ) are represented as individuals within a population set that describes an

text: evolutionary

text: path along time in generation steps.  Here the evolution is given by the population’s adaptation to another set of individuals, called here as the Target Set, which conveys all

text: psychoacoustic

text: features that steers the population evolution. This process is somehow similar to the “evolution pressure” that dictates the adaptation of populations in biological systems. The “best individual” within the population is the one most similar to the Target set. Unlike others deterministic sound synthesis methods such as the linear methods (i.e.

text: additive synthesis

text: ), the non-linear (i.e.

text: frequency modulation synthesis

text: ) and the sample-based methods (i.e.

text: sample-based synthesis

text: ) Essynth is a non-deterministic method of sound synthesis once that its resulting sound is not fixed for the same parametric values but continuously evolves along time.

text: The study of evolution is the central unifying concept in biology. Biology can be divided in various ways. One way is by the level of biological organisation, from  molecular to cell, organism to population. Another way is by taxonomic group, with fields such as zoology (all animals), ornithology (birds), and herpetology (reptiles and amphibians).A third way is by approach, such as field biology, theoretical biology, experimental evolution, and palaeontology. These alternative ways of dividing up the subject can be combined with evolutionary biology to create subfields like evolutionary ecology and evolutionary developmental biology.

text: The study of

text: evolution

text: is the central unifying concept in biology. Biology can be divided in various ways. One way is by the level of

text: biological organisation

text: , from

text: molecular

text: to

text: cell

text: , organism to

text: population

text: . Another way is by

text: taxonomic group

text: , with fields such as

text: zoology

text: (all animals),

text: ornithology

text: (birds), and

text: herpetology

text: (reptiles and amphibians).A third way is by approach, such as field biology,

text: theoretical biology

text: experimental evolution

text: , and

text: palaeontology

text: . These alternative ways of dividing up the subject can be combined with evolutionary biology to create subfields like

text: evolutionary ecology

text: and

text: evolutionary developmental biology

text: In biology, an error is said to occur when perfect fidelity is lost in the copying of information. For example, in an asexually reproducing species, an error (or mutation) has occurred for each DNA nucleotide that differs between the child and the parent. Many of these mutations can be harmful, but unlike other types of errors, some are neutral or even beneficial. Mutations are an important force driving evolution. Mutations that make organisms more adapted to their environment increase in the population through natural selection as organisms with favorable mutations have more offspring.

text: In

text: biology

text: , an error is said to occur when perfect fidelity is lost in the copying of

text: information

text: . For example, in an asexually reproducing species, an error (or mutation) has occurred for each

text: DNA

text: nucleotide

text: that differs between the

text: child

text: and the

text: parent

text: . Many of these mutations can be harmful, but unlike other types of errors, some are neutral or even beneficial. Mutations are an important force driving

text: evolution

text: . Mutations that make organisms more adapted to their

text: environment

text: increase in the population through

text: natural selection

text: as organisms with favorable mutations have more

text: offspring

text: Chimeric genes are important players in the evolution of genetic novelty.  Much like gene duplications, they provide a source of new genes, which can allow organisms to develop new phenotypes  and adapt to their environment.  Unlike duplicate genes, chimeric proteins are immediately distinct from their parental genes, and therefore are more likely to produce entirely new functions.

text: Chimeric genes are important players in the evolution of genetic novelty.  Much like

text: gene duplication

text: s, they provide a source of new genes, which can allow organisms to develop new

text: phenotype

text: s  and

text: adapt

text: to their environment.  Unlike duplicate genes, chimeric proteins are immediately distinct from their parental genes, and therefore are more likely to produce entirely new functions.

text: While the technological singularity is usually seen as a sudden event, some scholars argue the current speed of change already fits this description. In addition, some argue that we are already in the midst of a major evolutionary transition that merges technology, biology, and society. Digital technology has infiltrated the fabric of human society to a degree of indisputable and often life-sustaining dependence. A 2016 article in Trends in Ecology & Evolution argues that "humans already embrace fusions of biology and technology. We spend most of our waking time communicating through digitally mediated channels... we trust artificial intelligence with our lives through antilock braking in cars and autopilots in planes... With one in three marriages in America beginning online, digital algorithms are also taking a role in human pair bonding and reproduction". The article argues that from the perspective of the evolution, several previous Major Transitions in Evolution have transformed life through innovations in information storage and replication (RNA, DNA, multicellularity, and culture and language). In the current stage of life's evolution, the carbon-based biosphere has generated a cognitive system (humans) capable of creating technology that will result in a comparable evolutionary transition. The digital information created by humans has reached a similar magnitude to biological information in the biosphere. Since the 1980s, "the quantity of digital information stored has doubled about every 2.5 years, reaching about 5 zettabytes in 2014 (5x10^21 bytes). In biological terms, there are 7.2 billion humans on the planet, each having a genome of 6.2 billion nucleotides. Since one byte can encode four nucleotide pairs, the individual genomes of every human on the planet could be encoded by approximately 1x10^19 bytes. The digital realm stored 500 times more information than this in 2014 (...see Figure)... The total amount of DNA contained in all of the cells on Earth is estimated to be about 5.3x10^37 base pairs, equivalent to 1.325x10^37 bytes of information. If growth in digital storage continues at its current rate of 30–38% compound annual growth per year, it will rival the total information content contained in all of the DNA in all of the cells on Earth in about 110 years. This would represent a doubling of the amount of information stored in the biosphere across a total time period of just 150 years".

text: While the technological singularity is usually seen as a sudden event, some scholars argue the current speed of change already fits this description. In addition, some argue that we are already in the midst of a

text: major evolutionary transition

text: that merges technology, biology, and society. Digital technology has infiltrated the fabric of human society to a degree of indisputable and often life-sustaining dependence. A 2016 article in

text: Trends in Ecology & Evolution

text: argues that "humans already embrace fusions of biology and technology. We spend most of our waking time communicating through digitally mediated channels... we trust

text: artificial intelligence

text: with our lives through

text: antilock braking in cars

text: and

text: autopilot

text: s in planes... With one in three marriages in America beginning online, digital algorithms are also taking a role in human pair bonding and reproduction". The article argues that from the perspective of the

text: evolution

text: , several previous

text: Major Transitions in Evolution

text: have transformed life through innovations in information storage and replication (

text: RNA

text: DNA

text: multicellularity

text: , and

text: culture

text: and

text: language

text: ). In the current stage of life's evolution, the carbon-based biosphere has generated a

text: cognitive system

text: (humans) capable of creating technology that will result in a comparable

text: evolutionary transition

text: . The digital information created by humans has reached a similar magnitude to biological information in the biosphere. Since the 1980s, "the quantity of digital information stored has doubled about every 2.5 years, reaching about 5 zettabytes in 2014 (5x10^21 bytes). In biological terms, there are 7.2 billion humans on the planet, each having a genome of 6.2 billion nucleotides. Since one byte can encode four nucleotide pairs, the individual genomes of every human on the planet could be encoded by approximately 1x10^19 bytes. The digital realm stored 500 times more information than this in 2014 (...see Figure)... The total amount of DNA contained in all of the cells on Earth is estimated to be about 5.3x10^37 base pairs, equivalent to 1.325x10^37 bytes of information. If growth in digital storage continues at its current rate of 30–38% compound annual growth per year, it will rival the total information content contained in all of the DNA in all of the cells on Earth in about 110 years. This would represent a doubling of the amount of information stored in the biosphere across a total time period of just 150 years".

text: In the context of anthropology and other sciences which study the past, historicism has a different meaning. Anthropological historicism is associated with the work of Franz Boas. His theory used the diffusionist concept that there were a few "cradles of civilization" which grew outwards, and merged it with the idea that societies would adapt to their circumstances, which is called historical particularism. The school of historicism grew in response to unilinear theories that social development represented adaptive fitness, and therefore existed on a continuum. While these theories were espoused by Charles Darwin and many of his students, their application as applied in social Darwinism and general evolution characterized in the theories of Herbert Spencer and Leslie White, historicism was neither anti-selection, nor anti-evolution, as Darwin never attempted nor offered an explanation for cultural evolution. However, it attacked the notion that there was one normative spectrum of development, instead emphasizing how local conditions would create adaptations to the local environment. Julian Steward refuted the viability of globally and universally applicable adaptive standards proposing that culture was honed adaptively in response to the idiosyncrasies of the local environment, the cultural ecology, by specific evolution. What was adaptive for one region might not be so for another. This conclusion has likewise been adopted by modern forms of biological evolutionary theory.

text: In the context of

text: anthropology

text: and other sciences which study the past, historicism has a different meaning. Anthropological historicism is associated with the work of Franz Boas. His theory used the

text: diffusionist

text: concept that there were a few "cradles of civilization" which grew outwards, and merged it with the idea that societies would adapt to their circumstances, which is called

text: historical particularism

text: . The school of historicism grew in response to

text: unilinear theories

text: that social development represented adaptive fitness, and therefore existed on a continuum. While these theories were espoused by

text: Charles Darwin

text: and many of his students, their application as applied in

text: social Darwinism

text: and

text: general evolution

text: characterized in the theories of

text: Herbert Spencer

text: and

text: Leslie White

text: , historicism was neither anti-selection, nor anti-evolution, as Darwin never attempted nor offered an explanation for cultural evolution. However, it attacked the notion that there was one normative spectrum of development, instead emphasizing how local conditions would create adaptations to the local environment.

text: Julian Steward

text: refuted the viability of globally and universally applicable adaptive standards proposing that culture was honed adaptively in response to the idiosyncrasies of the local environment, the

text: cultural ecology

text: , by

text: specific evolution

text: . What was adaptive for one region might not be so for another. This conclusion has likewise been adopted by modern forms of biological evolutionary theory.

text: Many different studies have attempted to quantify the amount of adaptive evolution in the human genome, the vast majority using the comparative approaches outlined above. Although there are discrepancies between studies, generally there is relatively little evidence of adaptive evolution in protein coding DNA, with estimates of adaptive evolution often near 0% (see Table 1). The most obvious exception to this is the 35% estimate of α (Fay et al. 2001). This comparatively early study used relatively few loci (fewer than 200) for their estimate, and the polymorphism and divergence data used was obtained from different genes, both of which may have led to an overestimate of α. The next highest estimate is the 20% value of α (Zhang and Li 2005). However, the MK test used in this study was sufficiently weak that the authors state that this value of α is not statistically significantly different from 0%. Nielsen et al. (2005a)’s estimate that 9.8% of genes have undergone adaptive evolution also has a large margin of error associated with it, and their estimate shrinks dramatically to 0.4% when they stipulate that the degree of certainty that there has been adaptive evolution must be 95% or more. This raises an important issue, which is that many of these tests for adaptive evolution are very weak. Therefore the fact that many estimates are at (or very near to) 0% does not rule out the occurrence of any adaptive evolution in the human genome, but simply shows that positive selection is not frequent enough to be detected by the tests. In fact, the most recent study I mention states that confounding variables, such as demographic changes, means that the true value of α may be as high as 40% (Eyre-Walker and Keightley 2009). Another recent study, which uses a relatively robust methodology, estimates α at 10-20%  Boyko et al. (2008). I will comment on weaknesses in the methods in a subsequent section, but it is clear that the debate over the amount of adaptive evolution occurring in human coding DNA is not yet resolved.Even if low estimates of α are accurate, a small proportion of substitutions evolving adaptively can still equate to a considerable amount of coding DNA. Many authors, whose studies have small estimates of the amount of adaptive evolution in coding DNA, nevertheless accept that there has been some adaptive evolution in this DNA, because these studies identify specific regions within the human genome which have been evolving adaptively (e.g. Bakewell et al. (2007)). More genes underwent positive selection in chimpanzee evolution than in human), something I will examine later.The generally low estimates of adaptive evolution in human coding DNA can be contrasted with other species. Bakewell et al. (2007) found more evidence of adaptive evolution in chimpanzees than humans, with 1.7% of chimpanzee genes showing evidence of adaptive evolution (compared with the 1.1% estimate for humans; see Table 1). Comparing humans with more distantly related animals, an early estimate for α in Drosophila species was  45% (Smith and Eyre-Walker 2002), and later estimates largely agree with this (Eyre-Walker 2006). Bacteria and viruses generally show even more evidence of adaptive evolution; research shows values of α in a range of 50-85%, depending on the species examined (Eyre-Walker 2006). Generally, there does appear to be a positive correlation between (effective) population size of the species, and amount of adaptive evolution occurring in the coding DNA regions. This may be because random genetic drift becomes less powerful at altering allele frequencies, compared to natural selection, as population size increases.

text: Many different studies have attempted to quantify the amount of adaptive evolution in the human genome, the vast majority using the comparative approaches outlined above. Although there are discrepancies between studies, generally there is relatively little evidence of adaptive evolution in protein

text: coding DNA

text: population size

text: of the species, and amount of adaptive evolution occurring in the coding DNA regions. This may be because random

text: genetic drift

text: becomes less powerful at altering allele frequencies, compared to natural selection, as population size increases.

text: Debate over Darwin's work led to the rapid acceptance of the general concept of evolution, but the specific mechanism he proposed, natural selection, was not widely accepted until it was revived by developments in biology that occurred during the 1920s through the 1940s. Before that time most biologists regarded other factors as responsible for evolution. Alternatives to natural selection suggested during "the eclipse of Darwinism" (circa 1880 to 1920) included inheritance of acquired characteristics (neo-Lamarckism), an innate drive for change (orthogenesis), and sudden large mutations (saltationism). Mendelian genetics, a series of 19th Century experiments with pea plant variations rediscovered in 1900, was integrated with natural selection by Ronald Fisher, J. B. S. Haldane, and Sewall Wright during the 1910s to 1930s, and resulted in the founding of the new discipline of population genetics. During the 1930s and 1940s population genetics became integrated with other biological fields, resulting in a widely applicable theory of evolution that encompassed much of biology—the modern evolutionary synthesis.

text: Debate over Darwin's work led to the rapid acceptance of the general concept of evolution, but the specific mechanism he proposed, natural selection, was not widely accepted until it was revived by developments in

text: biology

text: that occurred during the 1920s through the 1940s. Before that time most

text: biologist

text: s regarded other factors as responsible for evolution. Alternatives to natural selection suggested during "

text: the eclipse of Darwinism

text: " (circa 1880 to 1920) included

text: inheritance of acquired characteristics

text: neo-Lamarckism

text: ), an innate drive for change (

text: orthogenesis

text: ), and sudden large

text: mutation

text: s (

text: saltationism

text: Mendelian genetics

text: , a series of 19th Century experiments with

text: pea

text: plant variations rediscovered in 1900, was integrated with natural selection by

text: Ronald Fisher

text: J. B. S. Haldane

text: , and

text: Sewall Wright

text: during the 1910s to 1930s, and resulted in the founding of the new discipline of

text: population genetics

text: . During the 1930s and 1940s population genetics became integrated with other biological fields, resulting in a widely applicable theory of evolution that encompassed much of biology—the

text: modern evolutionary synthesis

text: The Price equation can describe any system that changes over time but is most often applied in evolutionary biology. The evolution of sight provides an example of simple directional selection. The evolution of sickle cell anemia shows how a [heterozygote] advantage can affect trait evolution. The Price equation can also be applied to population context dependent traits such as the evolution of sex ratios. Additionally, the Price equation is flexible enough to model second order traits such as the evolution of mutability. The Price equation also provides an extension to Founder effect which shows change in population traits in different settlements

text: The Price equation can describe any system that changes over time but is most often applied in evolutionary biology. The

text: evolution of sight

text: provides an example of simple directional selection. The

entity_section: Example: evolution of sickle cell anemia

text: evolution of sickle cell anemia

text: shows how a [heterozygote] advantage can affect trait evolution. The Price equation can also be applied to population context dependent traits such as the evolution of

text: sex ratios

text: . Additionally, the Price equation is flexible enough to model second order traits such as the

text: evolution of mutability

text: . The Price equation also provides an extension to Founder effect which shows change in population traits in different settlements

text: In the mid-19th century, Charles Darwin formulated the scientific theory of evolution by natural selection, published in his book On the Origin of Species (1859). Evolution by natural selection is a process demonstrated by the observation that more offspring are produced than can possibly survive, along with three facts about populations: 1) traits vary among individuals with respect to morphology, physiology, and behaviour (phenotypic variation), 2) different traits confer different rates of survival and reproduction (differential fitness), and 3) traits can be passed from generation to generation (heritability of fitness). Thus, in successive generations members of a population are replaced by progeny of parents better adapted to survive and reproduce in the biophysical environment in which natural selection takes place. This teleonomy is the quality whereby the process of natural selection creates and preserves traits that are seemingly fitted for the functional roles they perform. Natural selection, including sexual selection, is the only known cause of adaptation but not the only known cause of evolution. Other, nonadaptive evolutionary processes include mutation, genetic drift and gene migration.

text: In the mid-19th century,

text: Charles Darwin

text: formulated the

text: scientific theory

text: of evolution by

text: natural selection

text: , published in his book

text: On the Origin of Species

text: (1859). Evolution by natural selection is a process demonstrated by the observation that more offspring are produced than can possibly survive, along with three

text: fact

text: s about populations: 1) traits vary among individuals with respect to morphology, physiology, and behaviour (

text: phenotypic variation

text: ), 2) different traits confer different rates of survival and

text: reproduction

text: (differential

text: fitness

text: ), and 3) traits can be passed from generation to generation (

text: heritability

text: of fitness). Thus, in successive generations members of a population are replaced by

text: progeny

text: of parents better

text: adapted

text: to survive and reproduce in the

text: biophysical environment

text: in which natural selection takes place. This

text: teleonomy

text: is the quality whereby the process of natural selection creates and preserves traits that are

text: seemingly fitted

text: for the

text: functional

text: roles they perform. Natural selection, including

text: sexual selection

text: , is the only known cause of adaptation but not the only known cause of evolution. Other, nonadaptive evolutionary processes include

text: mutation

text: genetic drift

text: and

text: gene migration

text: Evolution captures both the history of an organism via its phylogeny, and the history of natural selection working on function to produce adaptations.  There are several reasons why natural selection may fail to achieve optimal design (Mayr 2001:140–143; Buss et al. 1998).  One entails random processes such as mutation and environmental events acting on small populations.  Another entails the constraints resulting from early evolutionary development. Each organism harbors traits, both anatomical and behavioural, of previous phylogenetic stages, since many traits are retained as species evolve.

text: Evolution

text: captures both the history of an organism via its phylogeny, and the history of natural selection working on function to produce adaptations.  There are several reasons why

text: natural selection

text: may fail to achieve optimal design (Mayr 2001:140–143; Buss et al. 1998).  One entails random processes such as

text: mutation

text: and environmental events acting on small

text: population

text: s.  Another entails the constraints resulting from early evolutionary development. Each

text: organism

text: harbors

text: traits

text: , both

text: anatomical

text: and

text: behavioural

text: , of previous phylogenetic stages, since many traits are retained as

text: species

text: evolve.

text: Speciation, the divergence of one species into two or more descendant species, occurs when a part of the species is separated from the main population by a reproductive barrier. In allopatric and peripatric speciation, the reproductive barrier is the result of a physical barrier (e.g. flood waters, mountain range, deserts). Once separated, the species begins to adapt to their new environment via genetic drift and natural selection. After many generations and continual evolution, the populations become unable to interbreed with one another.

text: Speciation

text: , the divergence of one species into two or more descendant species, occurs when a part of the species is separated from the main population by a reproductive barrier. In allopatric and peripatric speciation, the reproductive barrier is the result of a physical barrier (e.g. flood waters, mountain range, deserts). Once separated, the species begins to adapt to their new environment via

text: genetic drift

text: and

text: natural selection

text: . After many generations and continual evolution, the populations become unable to interbreed with one another.

text: The gene-centered view of evolution, gene's eye view, gene selection theory, or selfish gene theory holds that adaptive evolution occurs through the differential survival of competing genes, increasing the allele frequency of those alleles whose phenotypic trait effects successfully promote their own propagation, with gene defined as "not just one single physical bit of DNA [but] all replicas of a particular bit of DNA distributed throughout the world". The proponents of this viewpoint argue that, since heritable information is passed from generation to generation almost exclusively by DNA, natural selection and evolution are best considered from the perspective of genes.

text: The gene-centered view of evolution, gene's eye view, gene selection theory, or selfish gene theory holds that adaptive

text: evolution

text: occurs through the differential survival of competing

text: gene

text: s, increasing the

text: allele frequency

text: of those

text: allele

text: s whose

text: phenotypic trait

text: effects successfully promote their own propagation, with gene defined as "not just one single physical bit of DNA [but] all replicas of a particular bit of DNA distributed throughout the world". The proponents of this viewpoint argue that, since

text: heritable

text: information is passed from generation to generation almost exclusively by

text: DNA

text: natural selection

text: and evolution are best considered from the perspective of genes.

text: Over many generations, the genomes of organisms can change significantly, resulting in evolution. In the process called adaptation, selection for beneficial mutations can cause a species to evolve into forms better able to survive in their environment. New species are formed through the process of speciation, often caused by geographical separations that prevent populations from exchanging genes with each other. The application of genetic principles to the study of population biology and evolution is known as the "modern evolutionary synthesis."

text: Over many generations, the genomes of organisms can change significantly, resulting in

text: evolution

text: . In the process called

text: adaptation

text: , selection for beneficial mutations can cause a species to evolve into forms better able to survive in their environment. New species are formed through the process of

text: speciation

text: , often caused by geographical separations that prevent populations from exchanging genes with each other. The application of genetic principles to the study of population biology and evolution is known as the "

text: modern evolutionary synthesis

text: "Let's Go All the Way" is a song by American group Sly Fox.  Released as a single in December 1985 from their debut studio album of the same name, the record entered the Billboard chart on December 28 and went Top 10 in both the U.S. and the UK. Despite receiving considerable commercial and critical success, the group failed to match expectations with their later singles, and are sometimes referred to as a one-hit wonder.

text: "Let's Go All the Way" is a song by American group

text: Sly Fox

text: .  Released as a

text: single

text: in December 1985 from their

text: debut studio album of the same name

text: , the record entered the Billboard chart on December 28 and went Top 10 in both the U.S. and the UK. Despite receiving considerable commercial and critical success, the group failed to match expectations with their later singles, and are sometimes referred to as a

text: one-hit wonder

text: Doxey's influence is such in Mormon thought that fifteen years after his death he was quoted in a First Presidency Message.

text: Doxey's influence is such in Mormon thought that fifteen years after his death he was quoted in a

text: First Presidency

text: Message.

text: The Moses Kent House is a historic house on River Road in Lyme, New Hampshire.  The  story wood frame house was built in 1811, and follows a typical Georgian floor plan with Federal styling.  Its interior and exterior finish details are in good condition.  The house is most notable the murals by Rufus Porter that are drawn on its walls.  Porter, at the time an itinerant painter, probably executed them between 1825 and 1830, and they are the only known example of his work in the area.  The murals depict countryside scenes and views of harbors.

text: The Moses Kent House is a historic house on River Road in

text: Lyme, New Hampshire

text: .  The  story wood frame house was built in 1811, and follows a typical Georgian floor plan with Federal styling.  Its interior and exterior finish details are in good condition.  The house is most notable the murals by

text: Rufus Porter

text: that are drawn on its walls.  Porter, at the time an itinerant painter, probably executed them between 1825 and 1830, and they are the only known example of his work in the area.  The murals depict countryside scenes and views of harbors.

text: Governor Arnold Schwarzenegger "directed inmate firefighters and staff from the California Department of Corrections and Rehabilitation", including those from CCI, to help fight the October 2007 California wildfires.

text: Governor

text: Arnold Schwarzenegger

text: "directed inmate firefighters and staff from the

entity_name: California Department of Corrections and Rehabilitation

text: California Department of Corrections and Rehabilitation

text: ", including those from CCI, to help fight the

text: October 2007 California wildfires

text: Both adaptation and random events can cause birth-and-death evolution. The probability of gene duplication is dictated by chance events and primarily occurs through unequal crossover; this excludes the rare event of whole genome duplication. Alternatively, the Fixation of duplicate genes can be influenced by natural selection or can occur randomly. Within the mammalian phylogenetic tree, a large number of gene gains and losses are observed for almost all branches, suggesting that a significant fraction of gene number changes were caused by inactivation events and random gene duplication. This process is known as genomic drift, or “random genetic drift of gene frequencies” in population genetics. Pseudogenes are also subject to genomic drift, since they are rendered as non-functional and are believed to evolve in a neutral manner.

text: Both adaptation and random events can cause birth-and-death evolution. The probability of gene duplication is dictated by chance events and primarily occurs through unequal crossover; this excludes the rare event of whole genome duplication. Alternatively, the

text: Fixation

text: of duplicate genes can be influenced by

text: natural selection

text: or can occur randomly. Within the mammalian phylogenetic tree, a large number of gene gains and losses are observed for almost all branches, suggesting that a significant fraction of gene number changes were caused by inactivation events and random gene duplication. This process is known as

text: genomic drift

text: , or “random genetic drift of gene frequencies” in population genetics.

text: Pseudogene

text: s are also subject to genomic drift, since they are rendered as non-functional and are believed to evolve in a neutral manner.

text: Edge of Sanity was a Swedish death metal band that, alongside Opeth, is commonly regarded as being the first to fuse extreme metal styles like death and black metal with progressive rock. The group was founded by Dan Swanö in 1989 and ended in 2003.

text: Edge of Sanity was a Swedish

text: death metal

text: band that, alongside

text: Opeth

text: , is commonly regarded as being the first to fuse extreme metal styles like

text: death

text: and

text: black metal

text: with

text: progressive rock

text: . The group was founded by

text: Dan Swanö

text: in 1989 and ended in 2003.

text: Yue Yang laid siege to the capital of Zhongshan for three years, during which time Duke Wu of Zongshan captured Yue Yang's son Yue Shu, killed him then cooked the body and sent the minced remains to Yue Yang. Sitting under his tent, the general ate an entire bowl full. On seeing this Marquess Wen said: "Because of me General Yang has had to eat the flesh of his own son. If he can do that, whose body would he not eat?"

text: 2008   Croydonisation, Museum of Croydon, London

text: 6. S. Musteaţă, (Ed.), Two Decades of Development in Post-Soviet States: Successes and Failures, Colecția Academica, 232, Iași, Institutul European, 2014, ISBN 978-606-24-0073-6

text: Crippen has done play by play work on the Water Channel's American Powerboat Television, ESPN's coverage of Unlimited Hydroplane Circuit, as well as NASCAR's Motor Racing Network.

text: Crippen has done play by play work on the

text: Water Channel

text: 's American Powerboat Television,

text: ESPN

text: 's coverage of Unlimited Hydroplane Circuit, as well as

text: NASCAR

text: 's Motor Racing Network.

text: He received a scholarship from UCLA, where he played at nose guard and defensive tackle. He transferred at the end of his sophomore season (1978) to Brigham Young University. He was named the starter at left defensive tackle as a junior, before moving to defensive end, where he received All-WAC and Honorable Mention All-American honors.

text: He received a scholarship from

text: UCLA

text: , where he played at

text: nose guard

text: and

text: defensive tackle

text: . He transferred at the end of his sophomore season (1978) to

text: Brigham Young University

text: . He was named the starter at

text: left defensive tackle

text: as a junior, before moving to

text: defensive end

text: , where he received All-

text: WAC

text: and Honorable Mention

text: All-America

text: n honors.

text: Krynickillus melanocephalus is a species of air-breathing land slug, a shell-less terrestrial gastropod mollusk in the family Agriolimacidae.

text: Krynickillus melanocephalus is a species of air-breathing

text: land slug

text: , a shell-less terrestrial

text: gastropod

text: mollusk

text: in the family

text: Agriolimacidae

text: The supplements and annual updates were given the same name as the publications

text: CIPFA and the Society of Management Accountants of Canada signed a mutual recognition agreement in February 2004.

text: CIPFA and the

entity_name: Certified Management Accountants of Canada

text: Society of Management Accountants of Canada

text: signed a

text: mutual recognition agreement

text: in February 2004.

text: The Railway code of Dharampur Station in "DMPR".

text: The 1994 national census reported a total population for this woreda of 228,772 in 47,812 households, of whom 118,696 were men and 110,076 were women; 3,552 or 1.55% of its population were urban dwellers. The largest ethnic group reported in Farta was the Amhara (99.95%), Amharic was spoken as a first language by 99.96%, and 99.57% of the population practiced Ethiopian Orthodox Christianity.

text: The 1994 national census reported a total population for this woreda of 228,772 in 47,812 households, of whom 118,696 were men and 110,076 were women; 3,552 or 1.55% of its population were urban dwellers. The largest ethnic group reported in Farta was the

text: Amhara

text: Amharic

text: was spoken as a first language by 99.96%, and 99.57% of the population practiced

text: Ethiopian Orthodox Christianity

text: Barlow's lark (Calendulauda barlowi) is a species of lark in the Alaudidae family. It is found in Namibia and South Africa where its natural habitat is subtropical or tropical dry shrubland. It is threatened by habitat loss.

text: Barlow's lark (Calendulauda barlowi) is a species of

text: lark

text: in the

text: Alaudidae

text: family. It is found in

text: Namibia

text: and

text: South Africa

text: where its natural

text: habitat

text: is subtropical or tropical dry

text: shrubland

text: . It is threatened by

text: habitat loss

text: Biological evolution gives considerable emphasis to the ability of organisms to adapt to unpredictable change—their survival value. In his words,

text: 8. e5 Nd7

text: 8. e5 Nd7

text: In some cases the terms of indenture were mentioned in the advertisement as well.

text: Charles Darwin is the founder of the theory of the evolution of the species. The evolutionary approach to personality psychology is based on this theory. This theory examines how individual personality differences are based on natural selection. Through natural selection organisms change over time through adaptation and selection. Traits are developed and certain genes come into expression based on an organism's environment and how these traits aid in an organism's survival and reproduction.

text: Charles Darwin

text: is the founder of the

text: theory of the evolution of the species

text: . The evolutionary approach to personality psychology is based on this theory. This theory examines how individual personality differences are based on

text: natural selection

text: . Through natural selection organisms change over time through adaptation and selection. Traits are developed and certain genes come into expression based on an organism's environment and how these traits aid in an organism's survival and reproduction.

text: The early Kanun-name (literally: "book of law") were related to financial and fiscal matters, and based on custom (örf), they tried to reconcile previously existing practices with the priorities and needs of the Ottoman state. Kanun-names were also granted to individual provinces following their conquest; these provincial books of law would typically maintain most of the taxes and dues existing under the previous rule, and simply adapt them to an Ottoman standard.

text: The early Kanun-name (literally: "book of law") were related to financial and fiscal matters, and based on custom (

text: örf

text: ), they tried to reconcile previously existing practices with the priorities and needs of the Ottoman state. Kanun-names were also granted to individual provinces following their conquest; these provincial books of law would typically maintain most of the taxes and dues existing under the previous rule, and simply adapt them to an Ottoman standard.

text: These types of TTM illustrate that an organization’s TTM goals should be aligned with its business strategy rather than pursuing speed blindly.

text: Group selection is often criticized to be too slow to happen in real biology. However, Jiang-Nan Yang recently showed with an individual-based model that the evolution of altruistic ageing occurs under fairly general conditions by kin/group selection. Group selection can be based on population viscosity (limited offspring dispersal, first proposed by Hamilton (1964) for kin selection) that is widely present in natural populations. This population structure builds a continuum between individual selection, kin selection, kin group selection and group selection without a clear boundary for each level. Although early theoretical models by D.S. Wilson et al. (1992) and Taylor (1992) showed that pure population viscosity cannot lead to cooperation/altruism because of the exact cancelling out of the benefit of kin cooperation and the cost of kin competition, this exact cancelling out also suggests that any additional benefit of local cooperation would be sufficient for the evolution of cooperation. Mitteldorf and D.S. Wilson (2000) later showed that if the population is allowed to fluctuate, then local populations can temporarily store the benefit of local cooperation and promote the evolution of altruism. By assuming individual differences in adaptations, Yang (2013) further showed that the benefit of local altruism can be stored in the form of offspring quality and thus promote the evolution of altruistic ageing even if the population does not fluctuate, this is because local competition among the young will result in an increased average local inherited fitness of survived progenies after the elimination of the less adapted by natural selection, since the young do not have strong age-associated abilities and have to depend more on inherited abilities to compete. In Yang (2013)'s model, altruistic ageing is stabilized by higher-level selection instead of just kin selection.

text: Group selection is often criticized to be too slow to happen in real biology. However, Jiang-Nan Yang recently showed with an

text: individual-based model

text: The Wii and Nintendo DS versions of Tron: Evolution.

text: The

text: Wii

text: and

text: Nintendo DS

text: versions of Tron: Evolution.

text: In 1867, when he came back from the world exhibition in Paris and brought the first chromic acid cell with him, he also met the astronomer Wilhelm Klinkerfues. Klinkerfues had developed a bifilar-hygrometer which was widely used but did not prove in laymen hands. After that Lambrecht build a hair hygrometer „Model Klinkerfues”. In 1873, after separation from Klinkerfues, he started to build new meteorological instruments such as polymeters, dew point monitors, aspiration psychrometers and so on. His weather telegraphs, weather columns and combinations of different meteorological instruments where in use at several bigger cities and foreign health resorts before 1st World War. Additionally Lambrecht engaged in construction and building of medical thermometers. At the same time he developed the so-called minimum thermometer with a narrowing of the lumen of the capillary which is placed above the mercury container. The precision instruments of Lambrecht are well-known all over the world.

text: In 1867, when he came back from the world exhibition in Paris and brought the first

text: chromic acid cell

text: with him, he also met the astronomer

text: Wilhelm Klinkerfues

text: . Klinkerfues had developed a bifilar-hygrometer which was widely used but did not prove in laymen hands. After that Lambrecht build a hair

text: hygrometer

text: „Model Klinkerfues”. In 1873, after separation from Klinkerfues, he started to build new meteorological instruments such as polymeters, dew point monitors, aspiration

text: psychrometer

text: s and so on. His weather telegraphs, weather columns and combinations of different meteorological instruments where in use at several bigger cities and foreign health resorts before 1st World War. Additionally Lambrecht engaged in construction and building of medical thermometers. At the same time he developed the so-called minimum thermometer with a narrowing of the lumen of the capillary which is placed above the mercury container. The precision instruments of Lambrecht are well-known all over the world.

text: Les Chaises Volantes, located in Walibi Sud-Ouest is a Zierer model from 1987 which was re-localised to the park in 1992.

text: Les Chaises Volantes, located in

text: Walibi Sud-Ouest

text: is a Zierer model from 1987 which was re-localised to the park in 1992.

text: In the 1990s, biological systematics used not only morphology and fossils to classify organisms, but also molecular biology. Molecular biology involves sequencing an organism's DNA and RNA and comparing the sequence with that of other living beings – the more similar they are, the more closely they are related. Comparison of even-toed ungulate and cetaceans genetic material has shown that the closest living relatives of whales and hippopotamuses is the paraphyletic group Artiodactyla.

text: In the 1990s,

text: biological systematics

text: used not only morphology and fossils to classify organisms, but also

text: molecular biology

text: . Molecular biology involves sequencing an organism's DNA and RNA and comparing the sequence with that of other living beings – the more similar they are, the more closely they are related. Comparison of even-toed ungulate and cetaceans genetic material has shown that the closest living relatives of

text: whale

text: s and

text: hippopotamus

text: es is the paraphyletic group Artiodactyla.

text: Gymnopilus aurantiobrunneus is a species of mushroom in the Cortinariaceae family.

text: Gymnopilus aurantiobrunneus is a species of

text: mushroom

text: in the

text: Cortinariaceae

text: family.

text: Scotland would have been eligible to enter the contest had Scotland gained independence as a result of the Scottish independence referendum, 2014, as Scotland would therefore have been a separate country.

text: Scotland would have been eligible to enter the contest had Scotland gained independence as a result of the

text: Scottish independence referendum, 2014

text: , as Scotland would therefore have been a separate country.

text: Lagarde is a commune in the Ariège department in southwestern France.

text: Lagarde is a

text: commune

text: in the

text: Ariège

text: department

text: in southwestern France.

text: Or similarly,

text: Or similarly,

text: C photosynthetic pathway evolution in Alloteropsis also represents an example of adaptive evolution via horizontal gene transfer in eukaryotes. It is the first such example of gene transfer between plant species that are not in direct physical contact (as in a host-parasite relationship). In each case, genes adapted for an important function in C photosynthesis have been transferred from grass lineages that diverged from Alloteropsis more than 20 million years ago, and independently evolved C photosynthesis. Horizontally inherited genes encode the photosynthesis enzymes PEPC and PEPCK. All other genes expressed in the mature C leaf of A. s. semialata were vertically inherited from a common ancestor with the C taxon A. s. eckloniana.

text: C photosynthetic pathway evolution

text: in Alloteropsis also represents an example of

text: adaptive evolution

text: via

text: horizontal gene transfer

text: in

text: eukaryote

text: s. It is the first such example of gene transfer between plant species that are not in direct physical contact (as in a host-parasite relationship). In each case, genes adapted for an important function in C photosynthesis have been transferred from grass lineages that diverged from Alloteropsis more than 20 million years ago, and independently evolved C photosynthesis.

text: Horizontally inherited genes

text: encode the photosynthesis

text: enzyme

text: s PEPC and PEPCK. All other genes expressed in the mature C leaf of A. s. semialata were vertically inherited from a

text: common ancestor

text: with the C taxon A. s. eckloniana.

text: Seven new facilities were added to the existing boardwalk area:

text: By now, she had become fed up with Dracula. The final insult came from his name being raised in conversation with a "bimbo" at the Usher club. She teleported to his castle only to find everyone dead. Lured by the only survivor, she was trapped in a furnace and almost incinerated, but managed to escape by taking the blood of the bait and teleporting out.

text: The Neolithic Revolution brought about significant evolutionary changes in humans; namely the transition from a hunter-gatherer lifestyle, in which humans foraged for food, to an agricultural lifestyle. This change occurred approximately 10,000-12,000 years ago. Humans began to domesticate both plants and animals, allowing for the maintenance of constant food resources. This transition quickly and dramatically changed the way that humans interact with the environment, with societies taking up practices of farming and animal husbandry. However, human bodies had evolved to be adapted to their previous foraging lifestyle. The slow pace of evolution in comparison with the very fast pace of human advancement allowed for the persistence of these adaptations in an environment where they are no longer necessary. In human societies that now function in a vastly different way from the hunter-gatherer lifestyle, these outdated adaptations now lead to the presence of maladaptive, or mismatched, traits.

text: The

text: Neolithic Revolution

text: brought about significant evolutionary changes in humans; namely the transition from a

text: hunter-gatherer

text: lifestyle, in which humans foraged for food, to an

text: agricultural

text: lifestyle. This change occurred approximately 10,000-12,000 years ago. Humans began to domesticate both plants and animals, allowing for the maintenance of constant food resources. This transition quickly and dramatically changed the way that humans interact with the environment, with societies taking up practices of farming and animal husbandry. However, human bodies had evolved to be adapted to their previous foraging lifestyle. The slow pace of evolution in comparison with the very fast pace of human advancement allowed for the persistence of these adaptations in an environment where they are no longer necessary. In human societies that now function in a vastly different way from the hunter-gatherer lifestyle, these outdated adaptations now lead to the presence of maladaptive, or mismatched, traits.

text: The Marlowe Theatre's pantomime is produced with Evolution Pantomimes, who produce number of pantomimes across the country. Past stars in The Marlowe Theatre pantomimes have included Natalie Imbruglia, Martine McCutcheon, Danniella Westbrook, Shelia Ferguson, Shaun Williamson, Daniel MacPherson, Robert Powell, Lewis Collins, Emma Barton, Adrian Edmonson, John Thompson, John Partridge, Toyah Willcox, Samantha Womack, Rita Simons and Gareth Gates.

text: The Marlowe Theatre's pantomime is produced with Evolution Pantomimes, who produce number of pantomimes across the country. Past stars in The Marlowe Theatre pantomimes have included

text: Natalie Imbruglia

text: Martine McCutcheon

text: Danniella Westbrook

text: Shelia Ferguson

text: Shaun Williamson

text: Daniel MacPherson

text: Robert Powell

text: Lewis Collins

text: Emma Barton

text: Adrian Edmonson

text: John Thompson

text: John Partridge

text: Toyah Willcox

text: Samantha Womack

text: Rita Simons

text: and

text: Gareth Gates

text: Though modern medical professional organizations share Freud's opinion that attempts to change sexual orientation are not helpful, they also do not find that a sort of arrested development explains homosexuality or that interpersonal relationships are what determine sexual orientation.  More scientific research has since found other biological factors that influence sexual attraction, with environmental factors weighing more on sexual behavior than sexual orientation.

text: Though modern medical professional organizations share Freud's opinion that attempts to change sexual orientation are not helpful, they also do not find that a sort of arrested development explains homosexuality or that interpersonal relationships are what determine sexual orientation.  More scientific research has since found other

text: biological factors

text: that influence sexual attraction, with

text: environmental factors

text: weighing more on sexual behavior than sexual orientation.

text: The next key step was the work of the British biologist and statistician Ronald Fisher. In a series of papers starting in 1918 and culminating in his 1930 book The Genetical Theory of Natural Selection, Fisher showed that the continuous variation measured by the biometricians could be produced by the combined action of many discrete genes, and that natural selection could change allele frequencies in a population, resulting in evolution. In a series of papers beginning in 1924, another British geneticist, J.B.S. Haldane worked out the mathematics of allele frequency change at a single gene locus under a broad range of conditions. Haldane also applied statistical analysis to real-world examples of natural selection, such as the Peppered moth evolution and industrial melanism, and showed that selection coefficients could be larger than Fisher assumed, leading to more rapid adaptive evolution as a camouflage strategy following increased pollution.

text: The next key step was the work of the British biologist and statistician Ronald Fisher. In a series of papers starting in 1918 and culminating in his 1930 book

text: , Fisher showed that the continuous variation measured by the biometricians could be produced by the combined action of many discrete genes, and that natural selection could change allele frequencies in a population, resulting in evolution. In a series of papers beginning in 1924, another British geneticist, J.B.S. Haldane worked out the mathematics of allele frequency change at a single gene

text: locus

text: under a broad range of conditions. Haldane also applied statistical analysis to real-world examples of natural selection, such as the

text: Peppered moth evolution

text: and

text: industrial melanism

text: , and showed that

text: selection coefficient

text: s could be larger than Fisher assumed, leading to more rapid adaptive evolution as a camouflage strategy following increased pollution.

text: Besides actively participating in the current discussions on evolutionary theory and genetics, Mosterín has also tackled issues like the definition of life itself or the ontology of biological organisms and species. Following in Aristotle’s and Schrödinger’s footsteps, he has been asking the simple question: what is life? He has analyzed the main proposed definitions, based on metabolism, reproduction, thermodynamics, complexity and evolution, and found all of them wanting. It is true that all organisms on Earth share many characteristics, from the encoding of genetic information in DNA to the storage of energy in ATP, but these common features merely reflect the inheritance from a common ancestor that possibly acquired them in a random way. From that point of view, our biology is the parochial science of life on Earth, rather than a universal science of life in general. Such a general biology seems impossible, as long as we do not detect and come to know other forms of life in the galaxy (in case they exist). Concerning the ontological thesis of Michael Ghiselin and David Hull on the individuality of biological species, Mosterín shows that they are neither classes nor individuals in the usual meaning of these words. He tries to extend and make more precise the available conceptual framework of the discussion. Specifically, he shows the formal equivalence of the set-theoretical and the mereological (or part and individual) approach, so that everything that can be said about the classes can be translated into the jargon of individuals, and the other way around. All these concerns converge in his recent philosophy of animality, which combines the ontology of animals as paradigmatic individuals with the insights and results of biological research. This general theory of animals provides a solid foundation for the development of anthropology, conceived as the science of human primates.

text: Besides actively participating in the current discussions on evolutionary theory and genetics, Mosterín has also tackled issues like the definition of life itself or the ontology of biological organisms and species. Following in

text: Aristotle

text: ’s and

text: Schrödinger

text: ’s footsteps, he has been asking the simple question: what is life? He has analyzed the main proposed definitions, based on metabolism, reproduction, thermodynamics, complexity and evolution, and found all of them wanting. It is true that all organisms on Earth share many characteristics, from the encoding of genetic information in DNA to the storage of energy in ATP, but these common features merely reflect the inheritance from a common ancestor that possibly acquired them in a random way. From that point of view, our biology is the parochial science of life on Earth, rather than a universal science of life in general. Such a general biology seems impossible, as long as we do not detect and come to know other forms of life in the galaxy (in case they exist). Concerning the ontological thesis of

text: Michael Ghiselin

text: and

text: David Hull

text: on the individuality of biological species, Mosterín shows that they are neither classes nor individuals in the usual meaning of these words. He tries to extend and make more precise the available conceptual framework of the discussion. Specifically, he shows the formal equivalence of the set-theoretical and the mereological (or part and individual) approach, so that everything that can be said about the classes can be translated into the jargon of individuals, and the other way around. All these concerns converge in his recent philosophy of animality, which combines the ontology of animals as paradigmatic individuals with the insights and results of biological research. This general theory of animals provides a solid foundation for the development of anthropology, conceived as the science of human primates.