Evolution 1

EvolutionEvolution (also known as biological or organic evolution) is the change over time in one or more inherited traitsfound in populations of organisms.[1] Inherited traits are particular distinguishing characteristics, includinganatomical, biochemical or behavioural characteristics, that result from gene–environment interactions. Evolutionmay occur when there is variation of inherited traits within a population. The major sources of such variation aremutation, genetic recombination and gene flow.[2] [3] [4] [5] This process has produced all the diversity of livingorganisms. Charles Darwin characterized the result as endless forms most beautiful and most wonderful.[6]

Two processes are generally distinguished as common causes of evolution. One is natural selection, a process inwhich there is differential survival and/or reproduction of organisms that differ in one or more inherited traits.[1]

Another cause is genetic drift, a process in which there are random changes to the proportions of two or moreinherited traits within a population.[7] [8]

In speciation, a single ancestral species splits into two or more different species. Speciation is visible in anatomical,genetic and other similarities between groups of organisms, geographical distribution of related species, the fossilrecord and the recorded genetic changes in living organisms over many generations. Speciation stretches back over3.5 billion years during which life has existed on earth.[9] [10] [11] [12] It is thought to occur in multiple ways such asslowly, steadily and gradually over time or rapidly from one long static state to another.The scientific study of evolution began in the mid-nineteenth century, when research into the fossil record and thediversity of living organisms convinced most scientists that species evolve.[13] The mechanism driving these changesremained unclear until the theory of natural selection was independently proposed by Charles Darwin and AlfredWallace in 1858. In the early 20th century, Darwinian theories of evolution were combined with genetics,palaeontology, and systematics, which culminated into a union of ideas known as the modern evolutionarysynthesis.[14] The synthesis became a major principle of biology as it provided a coherent and unifying explanationfor the history and diversity of life on Earth.[15] [16] [17]

Evolution is currently applied and studied in various areas within biology such as conservation biology,developmental biology, ecology, physiology, paleontology and medicine. Moreover, it has also made an impact ontraditionally non-biological disciplines such as agriculture, anthropology, philosophy and psychology.

History of evolutionary thought

Jean-Baptiste Lamarck

The roots of naturalistic thinking in biology can be dated to at least the 6th century BCE,with the Greek philosopher Anaximander.[18]

Early Christian Church Fathers and Medieval European scholars treated the Genesiscreation narrative as allegory and believed that natural organisms were unstable andcapricious, but the Protestant Reformation inspired Biblical literalism and a naturaltheology in which the concept of species was essentialist, static and fixed. All entitieswithin a species were seen as sharing a common essence.[19]

As emerging science explored mechanical philosophy in the 18th century,proto-evolutionary ideas were set out by a few natural philosophers such as PierreMaupertuis in 1745 and Erasmus Darwin in 1796.[20]

The word evolution was first used in relation to development of species in 1762, when Charles Bonnet used it for hisconcept of "pre-formation", in which females carried a miniature form of all future generations. The term graduallygained a more general meaning of growth or progressive development.[21] The first published modern use of theword has been attributed to the Edinburgh New Philosophical Journal in 1826, edited by Robert Jameson butarguably authored by Robert Edmond Grant.[22]

Evolution 2

The Bible-based Ussher chronology of the 1650s had calculated creation at 4004 BC, but by the 1780s geologistsassumed a much older world. Wernerians thought strata were deposits from shrinking seas, but James Huttonproposed a self-maintaining infinite cycle. Georges Cuvier's paleontological work in the 1790s established the realityof extinction, which he explained by local catastrophes, followed by repopulation of the affected areas by otherspecies. He held that species were fixed, and marginalised the ideas of the biologist Jean-Baptiste Lamarck abouttransmutation of species which were only taken up by radicals.Geologists such as Adam Sedgwick adapted Cuvier's catastrophism to show repeated worldwide annihilation andcreation of new fixed species adapted to a changed environment, identifying the most recent catastrophe as thebiblical flood. In opposition to this view, Charles Lyell adapted Hutton's concept into a stricter uniformitarianismwhich strongly influenced the young geologist Charles Darwin during the Beagle expedition. Initially Darwinfollowed Lyell's idea of repeated "centres of creation" of fixed species, but questioned Lyell's views and in 1836,near the end of the voyage, he expressed doubts that species were fixed.At this time, most natural historians conceived of "species" in terms of essences or ideals; actual individuals wereeither good or bad examples of the ideal. But natural scientists began to view regular patterns in nature statistically(that is, in terms of probabilities rather than determinism). Thomas Robert Malthus took this approach to humanpopulations in An Essay on the Principle of Population that influenced Darwin. Beginning with Darwin, "species"were conceived in statistical terms; actual individuals were expected to be different, with most diverging from theaverage form, and species were viewed as variable and intergrading units.[23] [24] [25] [26]

Around 1854 CharlesDarwin began writing out

what became On the Originof Species.

Darwin formulated his idea of natural selection in 1838 and was still developing histheory in 1858 when Alfred Russel Wallace sent him a similar theory, and both werepresented to the Linnean Society of London in separate papers.[27] At the end of 1859,Darwin's publication of On the Origin of Species explained natural selection in detail andpresented evidence leading to increasingly wide acceptance of the occurrence ofevolution. Thomas Henry Huxley applied Darwin’s ideas to humans, using paleontologyand comparative anatomy to provide strong evidence that humans and apes shared acommon ancestry. This caused an uproar around the world since it implied that thecreation myth in the Christian Bible was false, and humans did not have a special placein the universe.[28]

Debate about the mechanisms of evolution continued, and Darwin could not explain thesource of the heritable variations which would be acted on by natural selection.[29] Like Lamarck, he still thoughtthat parents passed on adaptations acquired during their lifetimes,[30] a theory which was subsequently dubbedLamarckism.[31] In the 1880s, August Weismann's experiments indicated that changes from use and disuse were notheritable, and Lamarckism gradually fell from favour.[32] [33] More significantly, Darwin could not account for howtraits were passed down from generation to generation. In 1865 Gregor Mendel found that traits were inherited in apredictable manner.[34] When Mendel's work was rediscovered in the 1900s, disagreements over the rate of evolutionpredicted by early geneticists and biometricians led to a rift between the Mendelian and Darwinian models ofevolution.

Hugo Marie de Vries was unaware of Gregor Mendel's work and rediscovered the laws of heredity in the 1890s. DeVries suggested the concept of genes as part of his mutation theory of evolution.[35] The rediscovery of GregorMendel's work provided a better understanding of how variation occurs in plant and animal traits. That variation isthe main fuel used by natural selection to shape the wide variety of adaptive traits observed in organic life. Eventhough Hugo de Vries and other early geneticists rejected gradual natural selection, their rediscovery of andsubsequent work on genetics eventually provided a solid basis on which the theory of evolution stood even moreconvincingly than when it was originally proposed.[36] At the beginning of the 20th century, Thomas Hunt Morganwas able to demonstrate that genes are carried on chromosomes and are the mechanical basis of heredity. Thesediscoveries formed the basis of the modern science of genetics.[37]

Evolution 3

The apparent contradiction between Darwin's theory of evolution by natural selection and Mendel's work wasreconciled in the 1920s and 1930s by evolutionary biologists such as J.B.S. Haldane, Sewall Wright, and particularlyRonald Fisher, who set the foundations for the establishment of the field of population genetics. The end result was acombination of evolution by natural selection and Mendelian inheritance, the modern evolutionary synthesis.[38] Inthe 1940s, the identification of DNA as the genetic material by Oswald Avery and colleagues and the subsequentpublication of the structure of DNA by James Watson and Francis Crick in 1953, demonstrated the physical basis forinheritance. Since then, genetics and molecular biology have become core parts of evolutionary biology and haverevolutionised the field of phylogenetics.[14]

In its early history, evolutionary biology primarily drew in scientists from traditional taxonomically orienteddisciplines, whose specialist training in particular organisms addressed general questions in evolution. Asevolutionary biology expanded as an academic discipline, particularly after the development of the modernevolutionary synthesis, it began to draw more widely from the biological sciences.[14] Currently the study ofevolutionary biology involves scientists from fields as diverse as biochemistry, ecology, genetics and physiology,and evolutionary concepts are used in even more distant disciplines such as psychology, medicine, philosophy andcomputer science.In the 1960s, scientists such as W. D. Hamilton[39] [40] [41] and George C. Williams[42] pioneered a gene-centeredview of evolution, with concepts such as kin selection.In 1975, E. O. Wilson's book Sociobiology established a significant place for evolutionary theory in psychology,[43]

giving rise to the field of evolutionary psychology. Critics argue that the hypotheses of evolutionary psychology aredifficult or impossible to test, for example because many current traits probably evolved to serve different functionsthan they do now.[43] While testing the hypotheses of evolutionary psychology is difficult, it is not impossible.[43]

Evolutionary Psychologists say that good evolutionary hypotheses can be corroborated or contradicted by data.[43]

The presence that evolutionary theory holds in psychology has been steadily increasing.[43]

In the 21st century, current research in evolutionary biology deals with several areas where the modern evolutionarysynthesis may need modification or extension, such as assessing the relative importance of various ideas on the unitof selection and evolvability and how to fully incorporate the findings of evolutionary developmental biology.[44] [45]

Evolution 4

Heredity

DNA structure. Bases are in the centre,surrounded by phosphate–sugar chains in a

double helix.

Evolution in organisms occurs through changes in heritable traits –particular characteristics of an organism. In humans, for example, eyecolour is an inherited characteristic and an individual might inherit the"brown-eye trait" from one of their parents.[46] Inherited traits arecontrolled by genes and the complete set of genes within an organism'sgenome is called its genotype.[47]

The complete set of observable traits that make up the structure andbehaviour of an organism is called its phenotype. These traits come fromthe interaction of its genotype with the environment.[48] As a result, manyaspects of an organism's phenotype are not inherited. For example,suntanned skin comes from the interaction between a person's genotype andsunlight; thus, suntans are not passed on to people's children. However,some people tan more easily than others, due to differences in theirgenotype; a striking example are people with the inherited trait of albinism,who do not tan at all and are very sensitive to sunburn.[49]

Heritable traits are known to be passed from one generation to the next viaDNA, a molecule that encodes genetic information.[47] DNA is a longpolymer composed of four types of bases. The sequence of bases along aparticular DNA molecule specify the genetic information, in a mannersimilar to a sequence of letters spelling out a sentence. Before a celldivides, the DNA is copied, so that each of the resulting two cells willinherit the DNA sequence. Portions of a DNA molecule that specify asingle functional unit are called genes; different genes have different sequences of bases. Within cells, the longstrands of DNA form condensed structures called chromosomes. The specific location of a DNA sequence within achromosome is known as a locus. If the DNA sequence at a locus varies between individuals, the different forms ofthis sequence are called alleles. DNA sequences can change through mutations, producing new alleles. If a mutationoccurs within a gene, the new allele may affect the trait that the gene controls, altering the phenotype of theorganism.[1]

However, while this simple correspondence between an allele and a trait works in some cases, most traits are morecomplex and are controlled by multiple interacting genes.[50] [51] Developmental biologists suggest that complexinteractions in genetic networks and communication among cells can lead to heritable variations that may underlaysome of the mechanics in developmental plasticity and canalization.[52]

Recent findings have confirmed important examples of heritable changes that cannot be explained by direct agency of the DNA molecule. These phenomena are classed as epigenetic inheritance systems that are causally or independently evolving over genes. Research into modes and mechanisms of epigenetic inheritance is still in its scientific infancy, however, this area of research has attracted much recent activity as it broadens the scope of heritability and evolutionary biology in general.[53] 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.[54] [55] 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 effect that modifies and feeds back into the selection regime of subsequent generations. Descendants inherit genes plus environmental characteristics generated by the ecological actions of ancestors.[56] Other examples of heritability in evolution that are not under the direct control of genes include the inheritance of cultural traits, group heritability,

Evolution 5

and symbiogenesis.[57] [58] [59] These examples of heritability that operate above the gene are covered broadly underthe title of multilevel or hierarchical selection, which has been a subject of intense debate in the history ofevolutionary science.[58] [60]

VariationAn individual organism's phenotype results from both its genotype and the influence from the environment it haslived in. A substantial part of the variation in phenotypes in a population is caused by the differences between theirgenotypes.[51] The modern evolutionary synthesis defines evolution as the change over time in this genetic variation.The frequency of one particular allele will fluctuate, becoming more or less prevalent relative to other forms of thatgene. Evolutionary forces act by driving these changes in allele frequency in one direction or another. Variationdisappears when a new allele reaches the point of fixation — when it either disappears from the population orreplaces the ancestral allele entirely.[61]

Variation comes from mutations in genetic material, migration between populations (gene flow), and the reshufflingof genes through sexual reproduction. Variation also comes from exchanges of genes between different species; forexample, through horizontal gene transfer in bacteria, and hybridisation in plants.[62] Despite the constantintroduction of variation through these processes, most of the genome of a species is identical in all individuals ofthat species.[63] However, even relatively small changes in genotype can lead to dramatic changes in phenotype: forexample, chimpanzees and humans differ in only about 5% of their genomes.[64]

Mutation

Duplication of part of achromosome.

Random mutations constantly occur in the genomes of organisms; these mutations creategenetic variation. Mutations are changes in the DNA sequence of a cell's genome and arecaused by radiation, viruses, transposons and mutagenic chemicals, as well as errors thatoccur during meiosis or DNA replication.[65] [66] [67] These mutations involve severaldifferent types of change in DNA sequences; these can either have no effect, alter theproduct of a gene, or prevent the gene from functioning. Studies in the fly Drosophilamelanogaster suggest that if a mutation changes a protein produced by a gene, this willprobably be harmful, with about 70% of these mutations having damaging effects, andthe remainder being either neutral or weakly beneficial.[68]

Due to the damaging effects that mutations can have on cells, organisms have evolvedmechanisms such as DNA repair to remove mutations.[65] Therefore, the optimalmutation rate for a species is a trade-off between costs of a high mutation rate, such asdeleterious mutations, and the metabolic costs of maintaining systems to reduce themutation rate, such as DNA repair enzymes.[69] Viruses that use RNA as their geneticmaterial have rapid mutation rates,[70] which can be an advantage since these viruses willevolve constantly and rapidly, and thus evade the defensive responses of e.g. the humanimmune system.[71]

Mutations can involve large sections of a chromosome becoming duplicated (usually bygenetic recombination), which can introduce extra copies of a gene into a genome.[72]

Extra copies of genes are a major source of the raw material needed for new genes to evolve.[73] This is importantbecause most new genes evolve within gene families from pre-existing genes that share common ancestors.[74] Forexample, the human eye uses four genes to make structures that sense light: three for colour vision and one for nightvision; all four are descended from a single ancestral gene.[75]

New genes can be created from an ancestral gene when a duplicate copy mutates and acquires a new function. This process is easier once a gene has been duplicated because it increases the redundancy of the system; one gene in the

Evolution 6

pair can acquire a new function while the other copy continues to perform its original function.[76] [77] Other types ofmutation can even create entirely new genes from previously noncoding DNA.[78] [79]

The creation of new genes can also involve small parts of several genes being duplicated, with these fragments thenrecombining to form new combinations with new functions.[80] [81] When new genes are assembled from shufflingpre-existing parts, domains act as modules with simple independent functions, which can be mixed together creatingnew combinations with new and complex functions.[82] For example, polyketide synthases are large enzymes thatmake antibiotics; they contain up to one hundred independent domains that each catalyze one step in the overallprocess, like a step in an assembly line.[83]

Changes in chromosome number may involve even larger mutations, where segments of the DNA withinchromosomes break and then rearrange. For example, two chromosomes in the Homo genus fused to produce humanchromosome 2; this fusion did not occur in the lineage of the other apes, and they retain these separatechromosomes.[84] In evolution, the most important role of such chromosomal rearrangements may be to acceleratethe divergence of a population into new species by making populations less likely to interbreed, and therebypreserving genetic differences between these populations.[85]

Sequences of DNA that can move about the genome, such as transposons, make up a major fraction of the geneticmaterial of plants and animals, and may have been important in the evolution of genomes.[86] For example, morethan a million copies of the Alu sequence are present in the human genome, and these sequences have now beenrecruited to perform functions such as regulating gene expression.[87] Another effect of these mobile DNA sequencesis that when they move within a genome, they can mutate or delete existing genes and thereby produce geneticdiversity.[66]

Sex and recombination

In the first stage of sexual reproduction, "meiosis," the number ofchromosomes is reduced from a diploid number (2n) to a haploid

number (n). During "fertilization," haploid gametes come together toform a diploid zygote and the original number of chromosomes (2n)

is restored.

In asexual organisms, genes are inherited together, orlinked, as they cannot mix with genes of otherorganisms during reproduction. In contrast, theoffspring of sexual organisms contain random mixturesof their parents' chromosomes that are producedthrough independent assortment. In a related processcalled homologous recombination, sexual organismsexchange DNA between two matchingchromosomes.[88] Recombination and reassortment donot alter allele frequencies, but instead change whichalleles are associated with each other, producingoffspring with new combinations of alleles.[89] Sexusually increases genetic variation and may increase therate of evolution.[90] [91] However, asexuality isadvantageous in some environments as it can evolve inpreviously sexual animals.[92] Here, asexuality mightallow the two sets of alleles in their genome to divergeand gain different functions.[93]

Recombination allows even alleles that are closetogether in a strand of DNA to be inheritedindependently. However, the rate of recombination islow (approximately two events per chromosome per

Evolution 7

generation). As a result, genes close together on a chromosome may not always be shuffled away from each other,and genes that are close together tend to be inherited together, a phenomenon known as linkage.[94] This tendency ismeasured by finding how often two alleles occur together on a single chromosome, which is called their linkagedisequilibrium. A set of alleles that is usually inherited in a group is called a haplotype. This can be important whenone allele in a particular haplotype is strongly beneficial: natural selection can drive a selective sweep that will alsocause the other alleles in the haplotype to become more common in the population; this effect is called genetichitchhiking.[95]

When alleles cannot be separated by recombination – such as in mammalian Y chromosomes, which pass intact fromfathers to sons – harmful mutations accumulate.[96] [97] By breaking up allele combinations, sexual reproductionallows the removal of harmful mutations and the retention of beneficial mutations.[98] In addition, recombination andreassortment can produce individuals with new and advantageous gene combinations. These positive effects arebalanced by the fact that sex reduces an organism's reproductive rate, can cause mutations and may separatebeneficial combinations of genes.[98] The reasons for the evolution of sexual reproduction are therefore unclear andthis question is still an active area of research in evolutionary biology,[99] [100] that has prompted ideas such as theRed Queen hypothesis.[101]

Population genetics

White peppered moth

Black morph in peppered moth evolution

From a genetic viewpoint, evolution is a generation-to-generation change in the frequencies of alleles within apopulation that shares a common gene pool.[102] A population is a localised group of individuals belonging to thesame species. For example, all of the moths of the same species living in an isolated forest represent a population. Asingle gene in this population may have several alternate forms, which account for variations between thephenotypes of the organisms. An example might be a gene for colouration in moths that has two alleles: black andwhite.A gene pool is the complete set of alleles for a gene in a single population; the allele frequency measures the fractionof the gene pool composed of a single allele (for example, what fraction of moth colouration genes are the blackallele). Evolution occurs when there are changes in the frequencies of alleles within a population of interbreedingorganisms; for example, the allele for black colour in a population of moths becoming more common.To understand the mechanisms that cause a population to evolve, it is useful to consider what conditions are requiredfor a population not to evolve. The Hardy-Weinberg principle states that the frequencies of alleles (variations in agene) in a sufficiently large population will remain constant if the only forces acting on that population are therandom reshuffling of alleles during the formation of the sperm or egg, and the random combination of the alleles inthese sex cells during fertilisation.[103] Such a population is said to be in Hardy-Weinberg equilibrium; it is notevolving.[104]

Evolution 8

Gene flowGene flow is the exchange of genes between populations, and between species.[105] Examples of gene flow within aspecies include the migration and then breeding of organisms and the exchange of pollen. Gene transfer betweenspecies includes the formation of hybrid organisms and horizontal gene transfer.Migration into or out of a population can change allele frequencies, as well as introduce genetic variation in apopulation. Immigration may add genes to the gene pool while emigration may remove genes. Gene flow is hinderedby mountain ranges, oceans and deserts or even man-made structures such as the Great Wall of China, which hashindered the flow of plant genes.[106]

Horizontal gene transfer is the transfer of genetic material from one organism to another organism that is not itsoffspring; this is most common among bacteria.[107] In medicine, this contributes to the spread of antibioticresistance, as when one bacteria acquires resistance genes it can rapidly transfer them to other species.[108]

Horizontal transfer of genes from bacteria to eukaryotes such as the yeast Saccharomyces cerevisiae and the adzukibean beetle Callosobruchus chinensis may also have occurred.[109] [110] An example of larger-scale transfers are theeukaryotic bdelloid rotifers, which have received a range of genes from bacteria, fungi, and plants.[111] Viruses canalso carry DNA between organisms, allowing transfer of genes even across biological domains.[112] Large-scale genetransfer has also occurred between the ancestors of eukaryotic cells and prokaryotes, during the acquisition ofchloroplasts and mitochondria.[113]

MechanismsThe two main mechanisms that produce evolution are natural selection and genetic drift. Natural selection is theprocess which favours genes that aid survival and reproduction. Genetic drift is the random change in the frequencyof alleles, caused by the random sampling of a generation's genes during reproduction. The relative importance ofnatural selection and genetic drift in a population varies depending on the strength of the selection and the effectivepopulation size, which is the number of individuals capable of breeding.[114] Natural selection usually predominatesin large populations, whereas genetic drift dominates in small populations. The dominance of genetic drift in smallpopulations can even lead to the fixation of slightly deleterious mutations.[115] As a result, changing population sizecan dramatically influence the course of evolution. Population bottlenecks, where the population shrinks temporarilyand therefore loses genetic variation, result in a more uniform population.[61]

Evolution 9

Natural selection

Natural selection of a population for dark colouration.

Evolution by means of natural selection isthe process by which genetic mutations thatenhance reproduction become, and remain,more common in successive generations of apopulation. It has often been called a"self-evident" mechanism because itnecessarily follows from three simple facts:

• Heritable variation exists withinpopulations of organisms.

• Organisms produce more offspring thancan survive.

• These offspring vary in their ability tosurvive and reproduce.

These conditions produce competitionbetween organisms for survival andreproduction. Consequently, organisms withtraits that give them an advantage over theircompetitors pass these advantageous traitson, while traits that do not confer an advantage are not passed on to the next generation.[116]

The central concept of natural selection is the evolutionary fitness of an organism.[117] Fitness is measured by anorganism's ability to survive and reproduce, which determines the size of its genetic contribution to the nextgeneration.[117] However, fitness is not the same as the total number of offspring: instead fitness is indicated by theproportion of subsequent generations that carry an organism's genes.[118] For example, if an organism could survivewell and reproduce rapidly, but its offspring were all too small and weak to survive, this organism would make littlegenetic contribution to future generations and would thus have low fitness.[117]

If an allele increases fitness more than the other alleles of that gene, then with each generation this allele will becomemore common within the population. These traits are said to be "selected for". Examples of traits that can increasefitness are enhanced survival, and increased fecundity. Conversely, the lower fitness caused by having a lessbeneficial or deleterious allele results in this allele becoming rarer — they are "selected against".[119] Importantly,the fitness of an allele is not a fixed characteristic; if the environment changes, previously neutral or harmful traitsmay become beneficial and previously beneficial traits become harmful.[1] However, even if the direction ofselection does reverse in this way, traits that were lost in the past may not re-evolve in an identical form (see Dollo'slaw).[120] [121]

Evolution 10

A chart showing three types of selection.1.Disruptive selection 2.Stabilizing selection

3.Directional selection

Natural selection within a population for a trait that can vary across arange of values, such as height, can be categorised into three differenttypes. The first is directional selection, which is a shift in the averagevalue of a trait over time — for example, organisms slowly gettingtaller.[122] Secondly, disruptive selection is selection for extreme traitvalues and often results in two different values becoming mostcommon, with selection against the average value. This would be wheneither short or tall organisms had an advantage, but not those ofmedium height. Finally, in stabilizing selection there is selectionagainst extreme trait values on both ends, which causes a decrease invariance around the average value and less diversity.[116] [123] Thiswould, for example, cause organisms to slowly become all the sameheight.

A special case of natural selection is sexual selection, which isselection for any trait that increases mating success by increasing theattractiveness of an organism to potential mates.[124] Traits thatevolved through sexual selection are particularly prominent in males ofsome animal species, despite traits such as cumbersome antlers, matingcalls or bright colours that attract predators, decreasing the survival of

individual males.[125] This survival disadvantage is balanced by higher reproductive success in males that show thesehard to fake, sexually selected traits.[126]

Natural selection most generally makes nature the measure against which individuals, and individual traits, are moreor less likely to survive. "Nature" in this sense refers to an ecosystem, that is, a system in which organisms interactwith every other element, physical as well as biological, in their local environment. Eugene Odum, a founder ofecology, defined an ecosystem as: "Any unit that includes all of the organisms...in a given area interacting with thephysical environment so that a flow of energy leads to clearly defined trophic structure, biotic diversity, and materialcycles (ie: exchange of materials between living and nonliving parts) within the system."[127] Each population withinan ecosystem occupies a distinct niche, or position, with distinct relationships to other parts of the system. Theserelationships involve the life history of the organism, its position in the food chain, and its geographic range. Thisbroad understanding of nature enables scientists to delineate specific forces which, together, comprise naturalselection.

An active area of research is the unit of selection, with natural selection being proposed to work at the level of genes,cells, individual organisms, groups of organisms and species.[128] [129] None of these are mutually exclusive andselection can act on multiple levels simultaneously.[130] An example of selection occurring below the level of theindividual organism are genes called transposons, which can replicate and spread throughout a genome.[131]

Selection at a level above the individual, such as group selection, may allow the evolution of co-operation, asdiscussed below.[132]

Evolution 11

Genetic drift

Simulation of genetic drift of 20 unlinked allelesin populations of 10 (top) and 100 (bottom). Drift

to fixation is more rapid in the smallerpopulation.

Genetic drift is the change in allele frequency from one generation tothe next that occurs because of the role that chance plays indetermining whether a given individual will survive and reproduce. Inmathematical terms, alleles are subject to sampling error. As a result,when selective forces are absent or relatively weak, allele frequenciestend to "drift" upward or downward randomly (in a random walk). Thisdrift halts when an allele eventually becomes fixed, either bydisappearing from the population, or replacing the other allelesentirely. Genetic drift may therefore eliminate some alleles from apopulation due to chance alone. Even in the absence of selectiveforces, genetic drift can cause two separate populations that began withthe same genetic structure to drift apart into two divergent populationswith different sets of alleles.[133]

The time for an allele to become fixed by genetic drift depends onpopulation size, with fixation occurring more rapidly in smallerpopulations.[134] The precise measure of population that is important iscalled the effective population size. The effective population is always smaller than the total population since it takesinto account factors such as the level of inbreeding, the number of animals that are too old or young to breed, and thelower probability of animals that live far apart managing to mate with each other.[135]

An example of when genetic drift is probably of central importance in determining a trait is the loss of pigmentsfrom animals that live in caves, a change that produces no obvious advantage or disadvantage in completedarkness.[136] However, it is usually difficult to measure the relative importance of selection and drift,[137] so thecomparative importance of these two forces in driving evolutionary change is an area of current research.[138] Theseinvestigations were prompted by the neutral theory of molecular evolution, which proposed that most evolutionarychanges are the result of the fixation of neutral mutations that do not have any immediate effects on the fitness of anorganism.[139] Hence, in this model, most genetic changes in a population are the result of constant mutationpressure and genetic drift.[140] This form of the neutral theory is now largely abandoned, since it does not seem to fitthe genetic variation seen in nature.[141] [142] However, a more recent and better-supported version of this model isthe nearly neutral theory, where most mutations only have small effects on fitness.[116]

OutcomesEvolution influences every aspect of the form and behaviour of organisms. Most prominent are the specificbehavioural and physical adaptations that are the outcome of natural selection. These adaptations increase fitness byaiding activities such as finding food, avoiding predators or attracting mates. Organisms can also respond toselection by co-operating with each other, usually by aiding their relatives or engaging in mutually beneficialsymbiosis. In the longer term, evolution produces new species through splitting ancestral populations of organismsinto new groups that cannot or will not interbreed.These outcomes of evolution are sometimes divided into macroevolution, which is evolution that occurs at or above the level of species, such as extinction and speciation, and microevolution, which is smaller evolutionary changes, such as adaptations, within a species or population.[143] In general, macroevolution is regarded as the outcome of long periods of microevolution.[144] Thus, the distinction between micro- and macroevolution is not a fundamental one – the difference is simply the time involved.[145] 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,

Evolution 12

by making it more likely that part of the population will become isolated. In this sense, microevolution andmacroevolution 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 ofspeciation and extinction.[146] [147] [148]

A common misconception is that evolution has goals or long-term plans; realistically however, evolution has nolong-term goal and does not necessarily produce greater complexity.[149] [150] Although complex species haveevolved, they occur as a side effect of the overall number of organisms increasing, and simple forms of life stillremain more common in the biosphere.[151] For example, the overwhelming majority of species are microscopicprokaryotes, which form about half the world's biomass despite their small size,[152] and constitute the vast majorityof Earth's biodiversity.[153] Simple organisms have therefore been the dominant form of life on Earth throughout itshistory and continue to be the main form of life up to the present day, with complex life only appearing more diversebecause it is more noticeable.[154] Indeed, the evolution of microorganisms is particularly important to modernevolutionary research, since their rapid reproduction allows the study of experimental evolution and the observationof evolution and adaptation in real time.[155] [156]

AdaptationAdaptation is the process that makes organisms better suited to their habitat.[157] [158] Also, the term adaptation mayrefer to a trait that is important for an organism's survival. For example, the adaptation of horses' teeth to the grindingof grass. By using the term adaptation for the evolutionary process, and adaptive trait for the product (the bodilypart or function), the two senses of the word may be distinguished. Adaptations are produced by naturalselection.[159] The following definitions are due to Theodosius Dobzhansky.

1. Adaptation is the evolutionary process whereby an organism becomes better able to live in its habitat orhabitats.[160]

2. Adaptedness is the state of being adapted: the degree to which an organism is able to live and reproduce in agiven set of habitats.[161]

3. An adaptive trait is an aspect of the developmental pattern of the organism which enables or enhances theprobability of that organism surviving and reproducing.[162]

Adaptation may cause either the gain of a new feature, or the loss of an ancestral feature. An example that showsboth types of change is bacterial adaptation to antibiotic selection, with genetic changes causing antibiotic resistanceby both modifying the target of the drug, or increasing the activity of transporters that pump the drug out of thecell.[163] Other striking examples are the bacteria Escherichia coli evolving the ability to use citric acid as a nutrientin a long-term laboratory experiment,[164] Flavobacterium evolving a novel enzyme that allows these bacteria togrow on the by-products of nylon manufacturing,[165] [166] and the soil bacterium Sphingobium evolving an entirelynew metabolic pathway that degrades the synthetic pesticide pentachlorophenol.[167] [168] An interesting but stillcontroversial idea is that some adaptations might increase the ability of organisms to generate genetic diversity andadapt by natural selection (increasing organisms' evolvability).[169] [170]

A baleen whale skeleton, a and b label flipper bones, which were adapted from front legbones: while c indicates vestigial leg bones, suggesting an adaptation from land to

sea.[171]

Adaptation occurs through the gradualmodification of existing structures.Consequently, structures with similarinternal organisation may havedifferent functions in relatedorganisms. This is the result of a singleancestral structure being adapted tofunction in different ways. The bones

Evolution 13

within bat wings, for example, are very similar to those in mice feet and primate hands, due to the descent of allthese structures from a common mammalian ancestor.[172] However, since all living organisms are related to someextent,[173] even organs that appear to have little or no structural similarity, such as arthropod, squid and vertebrateeyes, or the limbs and wings of arthropods and vertebrates, can depend on a common set of homologous genes thatcontrol their assembly and function; this is called deep homology.[174] [175]

During adaptation, some structures may lose their original function and become vestigial structures.[176] Suchstructures may have little or no function in a current species, yet have a clear function in ancestral species, or otherclosely related species. Examples include pseudogenes,[177] the non-functional remains of eyes in blindcave-dwelling fish,[178] wings in flightless birds,[179] and the presence of hip bones in whales and snakes.[171]

Examples of vestigial structures in humans include wisdom teeth,[180] the coccyx,[176] the vermiform appendix,[176]

and other behavioural vestiges such as goose bumps[181] and primitive reflexes.[182] [183] [184] [185]

However, many traits that appear to be simple adaptations are in fact exaptations: structures originally adapted forone function, but which coincidentally became somewhat useful for some other function in the process.[186] Oneexample is the African lizard Holaspis guentheri, which developed an extremely flat head for hiding in crevices, ascan be seen by looking at its near relatives. However, in this species, the head has become so flattened that it assistsin gliding from tree to tree—an exaptation.[186] Within cells, molecular machines such as the bacterial flagella[187]

and protein sorting machinery[188] evolved by the recruitment of several pre-existing proteins that previously haddifferent functions.[143] Another example is the recruitment of enzymes from glycolysis and xenobiotic metabolismto serve as structural proteins called crystallins within the lenses of organisms' eyes.[189] [190]

A critical principle of ecology is that of competitive exclusion: no two species can occupy the same niche in thesame environment for a long time.[191] Consequently, natural selection will tend to force species to adapt to differentecological niches. This may mean that, for example, two species of cichlid fish adapt to live in different habitats,which will minimise the competition between them for food.[192]

An area of current investigation in evolutionary developmental biology is the developmental basis of adaptations andexaptations.[193] This research addresses the origin and evolution of embryonic development and how modificationsof development and developmental processes produce novel features.[194] These studies have shown that evolutioncan alter development to create new structures, such as embryonic bone structures that develop into the jaw in otheranimals instead forming part of the middle ear in mammals.[195] It is also possible for structures that have been lostin evolution to reappear due to changes in developmental genes, such as a mutation in chickens causing embryos togrow teeth similar to those of crocodiles.[196] It is now becoming clear that most alterations in the form of organismsare due to changes in a small set of conserved genes.[197]

Co-evolution

Evolution 14

Common garter snake (Thamnophis sirtalissirtalis) which has evolved resistance to

tetrodotoxin in its amphibian prey.

Interactions between organisms can produce both conflict andco-operation. When the interaction is between pairs of species, such asa pathogen and a host, or a predator and its prey, these species candevelop matched sets of adaptations. Here, the evolution of one speciescauses adaptations in a second species. These changes in the secondspecies then, in turn, cause new adaptations in the first species. Thiscycle of selection and response is called co-evolution.[198] An exampleis the production of tetrodotoxin in the rough-skinned newt and theevolution of tetrodotoxin resistance in its predator, the common gartersnake. In this predator-prey pair, an evolutionary arms race hasproduced high levels of toxin in the newt and correspondingly highlevels of toxin resistance in the snake.[199]

Co-operationHowever, not all interactions between species involve conflict.[200] Many cases of mutually beneficial interactionshave evolved. For instance, an extreme cooperation exists between plants and the mycorrhizal fungi that grow ontheir roots and aid the plant in absorbing nutrients from the soil.[201] This is a reciprocal relationship as the plantsprovide the fungi with sugars from photosynthesis. Here, the fungi actually grow inside plant cells, allowing them toexchange nutrients with their hosts, while sending signals that suppress the plant immune system.[202]

Coalitions between organisms of the same species have also evolved. An extreme case is the eusociality found insocial insects, such as bees, termites and ants, where sterile insects feed and guard the small number of organisms ina colony that are able to reproduce. On an even smaller scale, the somatic cells that make up the body of an animallimit their reproduction so they can maintain a stable organism, which then supports a small number of the animal'sgerm cells to produce offspring. Here, somatic cells respond to specific signals that instruct them whether to grow,remain as they are, or die. If cells ignore these signals and multiply inappropriately, their uncontrolled growth causescancer.[65]

Such cooperation within species may have evolved through the process of kin selection, which is where oneorganism acts to help raise a relative's offspring.[203] This activity is selected for because if the helping individualcontains alleles which promote the helping activity, it is likely that its kin will also contain these alleles and thusthose alleles will be passed on.[204] Other processes that may promote cooperation include group selection, wherecooperation provides benefits to a group of organisms.[205]

Evolution 15

Speciation

The four mechanisms of speciation.

Speciation is the process where aspecies diverges into two or moredescendant species.[206]

There are multiple ways to defining thespecies concept. The choice of whichconcept to use is dependent on theparticularities of the speciesconcerned.[207] For example, somespecies concepts apply more readilytoward sexually reproducing organismswhile others lend themselves bettertoward asexual organisms. Despite thediversity of various species concepts,these various concepts can be placedinto one of three broad philosophicalapproaches: interbreeding, ecological,and phylogenetic.[208] The biologicalspecies concept (BSC) is a classicexample of the interbreeding approach.Defined by Ernst Mayr in 1942, the

BSC states that "species are groups of actually or potentially interbreeding natural populations, which arereproductively isolated from other such groups"[209] :120. Despite its wide and long-term use, the BSC like others isnot without controversy, particularly in prokaryotes,[210] and this is called the species problem.[207] Some researchershave attempted a unifying monistic definition of species, while others adopt a pluralistic approach and suggest thatthere may be a different ways to logically interpret the definition of a species.[207] [208] "

Barriers to reproduction between two diverging populations are required for the populations to become new species.Gene flow may slow this process by spreading the new genetic variants also to the other populations. Depending onhow far two species have diverged since their most recent common ancestor, it may still be possible for them toproduce offspring, as with horses and donkeys mating to produce mules.[211] Such hybrids are generally infertile. Inthis case, closely related species may regularly interbreed, but hybrids will be selected against and the species willremain distinct. However, viable hybrids are occasionally formed and these new species can either have propertiesintermediate between their parent species, or possess a totally new phenotype.[212] The importance of hybridisationin creating new species of animals is unclear, although cases have been seen in many types of animals,[213] with thegray tree frog being a particularly well-studied example.[214]

Speciation has been observed multiple times under both controlled laboratory conditions and in nature.[215] Insexually reproducing organisms, speciation results from reproductive isolation followed by genealogical divergence.There are four mechanisms for speciation. The most common in animals is allopatric speciation, which occurs inpopulations initially isolated geographically, such as by habitat fragmentation or migration. Selection under theseconditions can produce very rapid changes in the appearance and behaviour of organisms.[216] [217] As selection anddrift act independently on populations isolated from the rest of their species, separation may eventually produceorganisms that cannot interbreed.[218]

The second mechanism of speciation is peripatric speciation, which occurs when small populations of organisms become isolated in a new environment. This differs from allopatric speciation in that the isolated populations are numerically much smaller than the parental population. Here, the founder effect causes rapid speciation through both

Evolution 16

rapid genetic drift and selection on a small gene pool.[219]

The third mechanism of speciation is parapatric speciation. This is similar to peripatric speciation in that a smallpopulation enters a new habitat, but differs in that there is no physical separation between these two populations.Instead, speciation results from the evolution of mechanisms that reduce gene flow between the two populations.[206]

Generally this occurs when there has been a drastic change in the environment within the parental species' habitat.One example is the grass Anthoxanthum odoratum, which can undergo parapatric speciation in response to localisedmetal pollution from mines.[220] Here, plants evolve that have resistance to high levels of metals in the soil. Selectionagainst interbreeding with the metal-sensitive parental population produced a gradual change in the flowering time ofthe metal-resistant plants, which eventually produced complete reproductive isolation. Selection against hybridsbetween the two populations may cause reinforcement, which is the evolution of traits that promote mating within aspecies, as well as character displacement, which is when two species become more distinct in appearance.[221]

Geographical isolation of finches on theGalápagos Islands produced over a dozen new

species.

Finally, in sympatric speciation species diverge without geographicisolation or changes in habitat. This form is rare since even a smallamount of gene flow may remove genetic differences between parts ofa population.[222] Generally, sympatric speciation in animals requiresthe evolution of both genetic differences and non-random mating, toallow reproductive isolation to evolve.[223]

One type of sympatric speciation involves cross-breeding of tworelated species to produce a new hybrid species. This is not common inanimals as animal hybrids are usually sterile. This is because duringmeiosis the homologous chromosomes from each parent are fromdifferent species and cannot successfully pair. However, it is morecommon in plants because plants often double their number ofchromosomes, to form polyploids.[224] This allows the chromosomesfrom each parental species to form matching pairs during meiosis,since each parent's chromosomes are represented by a pair already.[225] An example of such a speciation event iswhen the plant species Arabidopsis thaliana and Arabidopsis arenosa cross-bred to give the new species Arabidopsissuecica.[226] This happened about 20,000 years ago,[227] and the speciation process has been repeated in thelaboratory, which allows the study of the genetic mechanisms involved in this process.[228] Indeed, chromosomedoubling within a species may be a common cause of reproductive isolation, as half the doubled chromosomes willbe unmatched when breeding with undoubled organisms.[229]

Speciation events are important in the theory of punctuated equilibrium, which accounts for the pattern in the fossilrecord of short "bursts" of evolution interspersed with relatively long periods of stasis, where species remainrelatively unchanged.[230] In this theory, speciation and rapid evolution are linked, with natural selection and geneticdrift acting most strongly on organisms undergoing speciation in novel habitats or small populations. As a result, theperiods of stasis in the fossil record correspond to the parental population, and the organisms undergoing speciationand rapid evolution are found in small populations or geographically restricted habitats, and therefore rarely beingpreserved as fossils.[231]

Evolution 17

Extinction

Tyrannosaurus rex. Non-avian dinosaurs died outin the Cretaceous–Tertiary extinction event at the

end of the Cretaceous period.

Extinction is the disappearance of an entire species. Extinction is notan unusual event, as species regularly appear through speciation, anddisappear through extinction.[232] Nearly all animal and plant speciesthat have lived on Earth are now extinct,[233] and extinction appears tobe the ultimate fate of all species.[234] These extinctions have happenedcontinuously throughout the history of life, although the rate ofextinction spikes in occasional mass extinction events.[235] TheCretaceous–Tertiary extinction event, during which the non-aviandinosaurs went extinct, is the most well-known, but the earlierPermian–Triassic extinction event was even more severe, withapproximately 96% of species driven to extinction.[235] The Holocene

extinction event is an ongoing mass extinction associated with humanity's expansion across the globe over the pastfew thousand years. Present-day extinction rates are 100–1000 times greater than the background rate, and up to30% of species may be extinct by the mid 21st century.[236] Human activities are now the primary cause of theongoing extinction event;[237] global warming may further accelerate it in the future.[238]

The role of extinction in evolution is not very well understood and may depend on which type of extinction isconsidered.[235] The causes of the continuous "low-level" extinction events, which form the majority of extinctions,may be the result of competition between species for limited resources (competitive exclusion).[14] If one species canout-compete another, this could produce species selection, with the fitter species surviving and the other speciesbeing driven to extinction.[128] The intermittent mass extinctions are also important, but instead of acting as aselective force, they drastically reduce diversity in a nonspecific manner and promote bursts of rapid evolution andspeciation in survivors.[239]

Evolutionary history of life

Origin of lifeHighly energetic chemistry is believed to have produced a self-replicating molecule around 4 billion years ago andhalf a billion years later the last common ancestor of all life existed.[240] The current scientific consensus is that thecomplex biochemistry that makes up life came from simpler chemical reactions.[241] The beginning of life may haveincluded self-replicating molecules such as RNA,[242] and the assembly of simple cells.[243]

Common descent

Evolution 18

The hominoids are descendants of a common ancestor.

All organisms on Earth are descendedfrom a common ancestor or ancestralgene pool.[173] [244] Current species area stage in the process of evolution, withtheir diversity the product of a longseries of speciation and extinctionevents.[245] The common descent oforganisms was first deduced from foursimple facts about organisms: First,they have geographic distributions thatcannot be explained by local adaptation.Second, the diversity of life is not a setof completely unique organisms, butorganisms that share morphological similarities. Third, vestigial traits with no clear purpose resemble functionalancestral traits, and finally, that organisms can be classified using these similarities into a hierarchy of nestedgroups – similar to a family tree.[246] However, modern research has suggested that, due to horizontal gene transfer,this "tree of life" may be more complicated than a simple branching tree since some genes have spread independentlybetween distantly related species.[247] [248]

Past species have also left records of their evolutionary history. Fossils, along with the comparative anatomy ofpresent-day organisms, constitute the morphological, or anatomical, record.[249] By comparing the anatomies of bothmodern and extinct species, paleontologists can infer the lineages of those species. However, this approach is mostsuccessful for organisms that had hard body parts, such as shells, bones or teeth. Further, as prokaryotes such asbacteria and archaea share a limited set of common morphologies, their fossils do not provide information on theirancestry.More recently, evidence for common descent has come from the study of biochemical similarities betweenorganisms. For example, all living cells use the same basic set of nucleotides and amino acids.[250] The developmentof molecular genetics has revealed the record of evolution left in organisms' genomes: dating when species divergedthrough the molecular clock produced by mutations.[251] For example, these DNA sequence comparisons haverevealed that humans and chimpanzees share 96% of their genomes and analyzing the few areas where they differhelps shed light on when the common ancestor of these species existed.[252]

Evolution 19

Evolution of life

Evolutionary tree showing the divergence of modern species from their common ancestorin the centre.Ciccarelli FD, Doerks T, von Mering C, Creevey CJ, Snel B, Bork P (2006)."Toward automatic reconstruction of a highly resolved tree of life". Science 311 (5765):

1283–87. doi:10.1126/science.1123061. PMID 16513982.  The three domains arecoloured, with bacteria blue, archaea green, and eukaryotes red.

Prokaryotes inhabited the Earth fromapproximately 3–4 billion yearsago.[253] [254] No obvious changes inmorphology or cellular organisationoccurred in these organisms over thenext few billion years.[255]

The eukaryotic cells emerged between1.6 – 2.7 billion years ago. The nextmajor change in cell structure camewhen bacteria were engulfed byeukaryotic cells, in a cooperativeassociation called endosymbiosis.[113]

[256] The engulfed bacteria and the hostcell then underwent co-evolution, withthe bacteria evolving into eithermitochondria or hydrogenosomes.[257]

Another engulfment ofcyanobacterial-like organisms led tothe formation of chloroplasts in algae and plants.[258]

The history of life was that of the unicellular eukaryotes, prokaryotes, and archaea until about 610 million years agowhen multicellular organisms began to appear in the oceans in the Ediacaran period.[253] [259] The evolution ofmulticellularity occurred in multiple independent events, in organisms as diverse as sponges, brown algae,cyanobacteria, slime moulds and myxobacteria.[260]

Soon after the emergence of these first multicellular organisms, a remarkable amount of biological diversityappeared over approximately 10 million years, in an event called the Cambrian explosion. Here, the majority oftypes of modern animals appeared in the fossil record, as well as unique lineages that subsequently becameextinct.[261] Various triggers for the Cambrian explosion have been proposed, including the accumulation of oxygenin the atmosphere from photosynthesis.[262]

About 500 million years ago, plants and fungi colonised the land, and were soon followed by arthropods and otheranimals.[263] Insects were particularly successful and even today make up the majority of animal species.[264]

Amphibians first appeared around 300 million years ago, followed by early amniotes, then mammals around 200million years ago and birds around 100 million years ago (both from "reptile"-like lineages). However, despite theevolution of these large animals, smaller organisms similar to the types that evolved early in this process continue tobe highly successful and dominate the Earth, with the majority of both biomass and species being prokaryotes.[153]

ApplicationsConcepts and models used in evolutionary biology, in particular natural selection, have many applications.[265]

Artificial selection is the intentional selection of traits in a population of organisms. This has been used for thousandsof years in the domestication of plants and animals.[266] More recently, such selection has become a vital part ofgenetic engineering, with selectable markers such as antibiotic resistance genes being used to manipulate DNA. Inrepeated rounds of mutation and selection proteins with valuable properties have evolved, for example modifiedenzymes and new antibodies, in a process called directed evolution.[267]

Evolution 20

Understanding the changes that have occurred during organism's evolution can reveal the genes needed to constructparts of the body, genes which may be involved in human genetic disorders.[268] For example, the mexican tetra is analbino cavefish that lost its eyesight during evolution. Breeding together different populations of this blind fishproduced some offspring with functional eyes, since different mutations had occurred in the isolated populations thathad evolved in different caves.[269] This helped identify genes required for vision and pigmentation.[270]

In computer science, simulations of evolution using evolutionary algorithms and artificial life started in the 1960s,and was extended with simulation of artificial selection.[271] Artificial evolution became a widely recognisedoptimisation method as a result of the work of Ingo Rechenberg in the 1960s. He used evolution strategies to solvecomplex engineering problems.[272] Genetic algorithms in particular became popular through the writing of JohnHolland.[273] Practical applications also include automatic evolution of computer programs.[274] Evolutionaryalgorithms are now used to solve multi-dimensional problems more efficiently than software produced by humandesigners, and also to optimise the design of systems.[275]

Social and cultural responses

As evolution became widely accepted inthe 1870s, caricatures of Charles Darwinwith an ape or monkey body symbolised

evolution.[276]

In the 19th century, particularly after the publication of On the Origin ofSpecies in 1859, the idea that life had evolved was an active source ofacademic debate centred on the philosophical, social and religiousimplications of evolution. Nowadays, the modern evolutionary synthesis iswidely accepted by scientists.[14] However, evolution remains a contentiousconcept for some theists.[277]

While various religions and denominations have reconciled their beliefs withevolution through concepts such as theistic evolution, there are creationistswho believe that evolution is contradicted by the creation myths found in theirrespective religions and who raise various objections to evolution.[143] [278]

[279] As had been demonstrated by responses to the publication of Vestiges ofthe Natural History of Creation in 1844, the most controversial aspect ofevolutionary biology is the implication of human evolution that humans sharecommon ancestry with apes, and that the mental and moral faculties ofhumanity have the same types of natural causes as other inherited traits inanimals.[13] In some countries, notably the United States, these tensionsbetween science and religion have fuelled the current creation-evolutioncontroversy, a religious conflict focusing on politics and public education.[280] While other scientific fields such ascosmology[281] and Earth science[282] also conflict with literal interpretations of many religious texts, evolutionarybiology experiences significantly more opposition from religious literalists.

The teaching of evolution in American secondary school biology classes was uncommon in most of the first half ofthe 20th century. The Scopes Trial decision of 1925 caused the subject to become very rare in American secondarybiology textbooks for a generation, but it was gradually re-introduced about a generation later and legally protectedwith the 1968 Epperson v. Arkansas decision. Since then, the competing religious belief of creationism was legallydisallowed in secondary school curricula in various decisions in the 1970s and 1980s, but it returned in the form ofintelligent design, to be excluded once again in the 2005 Kitzmiller v. Dover Area School District case.[283]

Social Darwinism is ideas about "survival of the fittest" taken out of their biological context and applied tocommerce and human societies as a whole, and misused to justify social inequality, sexism, racism andimperialism.[284] The 19th century Malthusian theory developed by Whig philosopher Herbert Spencer is also seenas belonging to the social Darwinism movement.

Evolution 21

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Evolution 22

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Further readingIntroductory reading• Carroll, S. (2005). Endless Forms Most Beautiful. New York: W.W. Norton. ISBN 0-393-06016-0.• Charlesworth, C.B. and Charlesworth, D. (2003). Evolution. Oxfordshire: Oxford University Press.

ISBN 0-19-280251-8.• Dawkins, R. (2006). The Selfish Gene: 30th Anniversary Edition. Oxford University Press. ISBN 0-19-929115-2.• Gould, S.J. (1989). Wonderful Life: The Burgess Shale and the Nature of History. New York: W.W. Norton.

ISBN 0-393-30700-X.• Jones, S. (2001). Almost Like a Whale: The Origin of Species Updated. (American title: Darwin's Ghost). New

York: Ballantine Books. ISBN 0-345-42277-5.• Mader, Sylvia S. (2007). Biology. Murray P. Pendarvis (9th ed.). McGraw Hill. ISBN 978-0-07-325839-3.• Maynard Smith, J. (1993). The Theory of Evolution: Canto Edition. Cambridge University Press.

ISBN 0-521-45128-0.• Pallen, M.J. (2009). The Rough Guide to Evolution. Rough Guides. ISBN 978-1-85828-946-5.• Smith, C.B. and Sullivan, C. (2007). The Top 10 Myths about Evolution. Prometheus Books.

ISBN 978-1-59102-479-8.History of evolutionary thought• Darwin, Charles (1859). On the Origin of Species (http:/ / darwin-online. org. uk/ content/

frameset?itemID=F373& viewtype=text& pageseq=1) (1st ed.). London: John Murray. ISBN 0-8014-1319-2.• Larson, E.J. (2004). Evolution: The Remarkable History of a Scientific Theory. New York: Modern Library.

ISBN 0-679-64288-9.• Zimmer, C. (2001). Evolution: The Triumph of an Idea. London: HarperCollins. ISBN 0-06-019906-7.Advanced reading• Barton, N.H., Briggs, D.E.G., Eisen, J.A., Goldstein, D.B. and Patel, N.H. (2007). Evolution. Cold Spring Harbor

Laboratory Press. ISBN 0-87969-684-2.• Coyne, J.A. and Orr, H.A. (2004). Speciation. Sunderland: Sinauer Associates. ISBN 0-87893-089-2.• Futuyma, D.J. (2005). Evolution. Sunderland: Sinauer Associates. ISBN 0-87893-187-2.• Gould, S.J. (2002). The Structure of Evolutionary Theory. Cambridge: Belknap Press (Harvard University Press).

ISBN 0-674-00613-5.• Maynard Smith, J. and Szathmáry, E. (1997). The Major Transitions in Evolution. Oxfordshire: Oxford

University Press. ISBN 0-19-850294-X.• Mayr, E. (2001). What Evolution Is. New York: Basic Books. ISBN 0-465-04426-3.• Olson, Wendy; Hall, Brian Keith (2003). Keywords and concepts in evolutionary developmental biology.

Cambridge: Harvard University Press. ISBN 0-674-02240-8.• Genome Evolution

Evolution 32

External linksGeneral information• Evolution (http:/ / www. bbc. co. uk/ programmes/ p00545gl) on In Our Time at the BBC. ( listen now (http:/ /

www. bbc. co. uk/ iplayer/ console/ p00545gl/ In_Our_Time_Evolution))• Everything you wanted to know about evolution by New Scientist (http:/ / www. newscientist. com/ topic/

evolution)• Howstuffworks.com — How Evolution Works (http:/ / science. howstuffworks. com/ evolution/ evolution. htm)• National Academies Evolution Resources (http:/ / nationalacademies. org/ evolution/ )• Synthetic Theory Of Evolution: An Introduction to Modern Evolutionary Concepts and Theories (http:/ / anthro.

palomar. edu/ synthetic/ )• Understanding Evolution from University of California, Berkeley (http:/ / evolution. berkeley. edu/ )• Evolution of Evolution – 150 Years of Darwin's "On the Origin of Species" (http:/ / www. nsf. gov/ news/

special_reports/ darwin/ textonly/ index. jsp)History of evolutionary thought• The Complete Work of Charles Darwin Online (http:/ / darwin-online. org. uk/ )• Understanding Evolution: History, Theory, Evidence, and Implications (http:/ / www. rationalrevolution. net/

articles/ understanding_evolution. htm)On-line lectures• The Making of the Fittest (http:/ / www. molbio. wisc. edu/ carroll/ Fittest. html) – lecture by Sean B. Carroll

Article Sources and Contributors 33

Article Sources and ContributorsEvolution  Source: http://en.wikipedia.org/w/index.php?oldid=417575551  Contributors: !!!niloivenutfotuO, .:Ajvol:., 0nlyth3truth, 10.175, 100110100, 119, 12 Cove, 123heyho, 162.129.26.xxx, 168..., 1nic, 210.50.54.xxx, 216.126.89.xxx, 2357, 24.108.14.xxx, 3 Lane, 4 Gnon, 4 trin, 40 tune, 489thCorsica, 6 Four, 6 y go, 65.68.87.xxx, 7 Fej, 8r13n, 9 noon, 99DBSIMLR, A Softer Answer, A bit iffy, A455bcd9, A8UDI, AC+79 3888, AH9, ASDFGHJKL, AVengel, Aaarrrggh, Aarnu, Aaron Schulz, AaronFX, AaronY, Aartp, Abce2, Abdullais4u, Abi Don, Abrech, Abtract, Abuzin, Academic Challenger, Acalamari, Achoo5000, Acroterion, Acsparkman, AdamRetchless, Adambiswanger1, Adambro, Adamsiepel, Adashiel, Adenosine, Adi, Adraeus, Adrian, Adriansrfr, Adz71, Aecis, Afasmit, Africangenesis, Ag545, Agathman, Agentmoose, Aggn, Agrabiec4, Ahoerstemeier, Aircorn, Aitias, Ajf99, Ajgisme, Aks818guy, Alan Peakall, AlanD, AlanHarmony, Aldaron, Aldenrw, Ale jrb, Alex.tan, Alexf, AlexiusHoratius, Alexwebb2, Alfio, Algebraist, Algorithms, Ali, Alias Flood, AlienHook, Alienus, Alpha166, AlphaEta, Alphachimp, Alphazeta33, Altenmann, Amaltheus, Amarilloarmadillo, Amcaja, AmiDaniel, Amitch, Amused Usher, Ancheta Wis, Ande B., Andrevan, Andrew Lancaster, Andrew c, AndrewTJ31, AndrewWTaylor, Andrewa, Andrewlp1991, Andrewpmk, Andrewrost3241981, Andriesb, Android79, Andromeda321, Andymarczak, Angelo, Anilocra, Animum, Another sockpuppet of Outoftuneviolin, Antandrus, Anthony, AnthonyQBachler, Apokryltaros, Apostlealex, Apostrophe, Arch dude, ArcticFrog, Ardric47, ArielGold, Arjayay, Arjun01, Arkatox, Arker, Arm, Armchair info guy, Arnoutf, Arof, Aron-Ra, Art LaPella, ArthurWeasley, Artichoker, Asbestos, Ascánder, Asemoasyourmom, Ashenai, Ashmoo, Astrobayes, Athenean, Athf1234, AuburnPilot, Audacity, Aude, Aunt Entropy, Aurush kazemini, Autonova, Avb, AvicAWB, Avicennasis, Awon, Axa4975, Axel147, AxelBoldt, Axeman89, Axxel Sel, Az1568, AzaToth, Azcolvin429, Azra99, AzureCitizen, B, B89smith, BG, Back23, Badanedwa, Badreligion, Baegis, Bakasuprman, Ballin789, Ballista, Bam2014, Banes, Barbary lion, Baristarim, Barnaby dawson, Barny Fife, Barras, Bart133, Base tonne, Basil45, Bassbreaker, BatteryIncluded, Bauxzaux, Bbatsell, Bcasterline, Bdaay, Beezhive, Beltho, BenB4, Bender235, Bendzh, Benhocking, Benzocane, BerndH, Besidesamiracle, Betacommand, Betbs, Betterusername, Bevo, Bharatveer, Bhawani Gautam Rhk, BigGoose2006, Biglonstud, Bikeable, Bill37212, Billare, BillySharps, Billystut, Binabik80, Binarypascal, Bio-queen, BirgitteSB, Bkl3x, Black Kite, BlackMaria, Blainster, Blanchardb, BlankVerse, Blue401, BlueFireIce, BlueGoose, BlueNight, BlytheG, Bobby Mason, Bobo192, Bodnotbod, Boe syl, Boffey, Bogdangiusca, Bogey97, Bollyx, Boo wall, Bookandcoffee, Boomcoach, Boothy443, Boring bus, Borisblue, Bornhj, Bow bowl, Bowslayer, BradBeattie, Brainstormer1980, Brazucs, Breathe ing, Breed3011, Brent0270, Brian0918, BrianG5, Brighterorange, Briguy, Broabey, Broalka, BrokenSegue, Bryan Derksen, Bth, Bubba73, Bucketsofg, Bull Market, Burmiester, Burtonsarpa, Busy ironing, Butros, Butwhatdoiknow, Bwatb, Bwhack, Byrll, C.Fred, CJLL Wright, CMacMillan, CUSENZA Mario, Caasiopia68, Caesura, Calamine Lotion, Calaschysm, CalebNoble, Caltechdoc, CambridgeBayWeather, Camembert, Can't sleep, clown will eat me, CanDo, Canadian-Bacon, Candorwien, CanisRufus, Canterbury Tail, CapitalR, CardinalDan, Cards44izzy, Cassan, Caulde, Causa sui, Celador, Cen Upp, Centrx, CerealKiller, ChadThomson, Chairboy, Chameleon, Chanting Fox, Chaojoker, Chaos, Chardon, CharlesGillingham, Charlesdrakew, Chas zzz brown, CheeseDreams, Cherhillsnow, Cheshire Boy, ChicXulub, Chicken Soup, Chris Melton, Chris the speller, Chris55, Chrislk02, Christian List, Christian41691, Christopher Parham, Christopherlin, Christpower79, Chun-hian, Chunky Rice, Ciar, Cinnamon, Ck lostsword, ClarenceCM3, Clayoquot, Clean Sweep, ClockworkSoul, Cloud silver, Cmayy, Coffee2theorems, Cold Sandwhich, Colin Keigher, Comder, Cometstyles, CommonJoe, CommonsDelinker, Compute14, ComputerPlace, ConfuciusOrnis, Connelly, ConservativeChristian, Conspiracyfactory, Conversion script, Coopercmu, Copper canned, Coppit, Cosmic Latte, Cougarkid, Country050, Cpl Syx, Crab or, Craigmac41, Craigy144, Crazytail2, Crazytail3, Crazytales, Creationlaw, Cretog8, Crust, CryptoDerk, Curps, CuteWombat, Cutsman, Cwilldagangsta, Cybercobra, Cyde, Cynical, Cynicism addict, Cyp, Cyr egi, DDek, DEFRT, DGG, DJ Clayworth, DKqwerty, DLH, DLR, DSG2, DVD R W, DVdm, Da Gingerbread Man, Daa89563, Dabs, Dachshundboy25, Daelin, Daeoque, Dafergu3, Dan ald, DanTheMan2, Dana boomer, Dancingspring, Danerz34, Dangee63, Daniel5127, DanielCD, Danielkueh, Danielparker, Danielrcote, Danimoth, Danny, Dante Alighieri, Dar book, Dark Shikari, Darthgriz98, Dash92, Dashmast3r, Dave souza, David D., David Fuchs, David Kernow, David Merrill, David Schaich, David Z. 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Image Sources, Licenses and ContributorsImage:Jean-Baptiste Lamarck.jpg  Source: http://en.wikipedia.org/w/index.php?title=File:Jean-Baptiste_Lamarck.jpg  License: Public Domain  Contributors: User:DuesentriebFile:Charles Darwin aged 51 crop.jpg  Source: http://en.wikipedia.org/w/index.php?title=File:Charles_Darwin_aged_51_crop.jpg  License: Public Domain  Contributors: User:Dave souzaFile:ADN static.png  Source: http://en.wikipedia.org/w/index.php?title=File:ADN_static.png  License: Public Domain  Contributors: User:Brian0918File:Gene-duplication.svg  Source: http://en.wikipedia.org/w/index.php?title=File:Gene-duplication.svg  License: Public Domain  Contributors: User:K. AainsqatsiImage:Sexual cycle.svg  Source: http://en.wikipedia.org/w/index.php?title=File:Sexual_cycle.svg  License: GNU Free Documentation License  Contributors: User:StanneredImage:Biston.betularia.7200.jpg  Source: http://en.wikipedia.org/w/index.php?title=File:Biston.betularia.7200.jpg  License: Creative Commons Attribution-Sharealike 2.5  Contributors:Kilom691, OleiImage:Biston.betularia.f.carbonaria.7209.jpg  Source: http://en.wikipedia.org/w/index.php?title=File:Biston.betularia.f.carbonaria.7209.jpg  License: Creative Commons Attribution-Sharealike2.5  Contributors: Kilom691, OleiFile:Mutation and selection diagram.svg  Source: http://en.wikipedia.org/w/index.php?title=File:Mutation_and_selection_diagram.svg  License: GNU Free Documentation License Contributors: User:ElembisFile:Selection Types Chart.png  Source: http://en.wikipedia.org/w/index.php?title=File:Selection_Types_Chart.png  License: Creative Commons Attribution-Sharealike 3.0  Contributors:User:Azcolvin429File:Allele-frequency.png  Source: http://en.wikipedia.org/w/index.php?title=File:Allele-frequency.png  License: GNU Free Documentation License  Contributors: Original uploader was Esurnirat en.wikipediaFile:Whale skeleton.png  Source: http://en.wikipedia.org/w/index.php?title=File:Whale_skeleton.png  License: Public Domain  Contributors: Meyers KonversionlexikonFile:Thamnophis sirtalis sirtalis Wooster.jpg  Source: http://en.wikipedia.org/w/index.php?title=File:Thamnophis_sirtalis_sirtalis_Wooster.jpg  License: Public Domain  Contributors:User:Wilson44691File:Speciation modes edit.svg  Source: http://en.wikipedia.org/w/index.php?title=File:Speciation_modes_edit.svg  License: Public Domain  Contributors: User:Ilmari KaronenFile:Darwin's finches.jpeg  Source: http://en.wikipedia.org/w/index.php?title=File:Darwin's_finches.jpeg  License: Public Domain  Contributors: John Gould (14.Sep.1804 - 3.Feb.1881)File:Palais de la Decouverte Tyrannosaurus rex p1050042.jpg  Source: http://en.wikipedia.org/w/index.php?title=File:Palais_de_la_Decouverte_Tyrannosaurus_rex_p1050042.jpg  License:GNU Free Documentation License  Contributors: User:David.MonniauxFile:Ape skeletons.png  Source: http://en.wikipedia.org/w/index.php?title=File:Ape_skeletons.png  License: Public Domain  Contributors: Original uploader was TimVickers at en.wikipediaImage:Collapsed_tree_labels_simplified.png  Source: http://en.wikipedia.org/w/index.php?title=File:Collapsed_tree_labels_simplified.png  License: Public Domain  Contributors: Originaluploader was TimVickers at en.wikipediaFile:Editorial cartoon depicting Charles Darwin as an ape (1871).jpg  Source:http://en.wikipedia.org/w/index.php?title=File:Editorial_cartoon_depicting_Charles_Darwin_as_an_ape_(1871).jpg  License: Public Domain  Contributors: Unknown, The Hornet is no longer inpublication and it is very likely for a 20-year-old artist in 1871 to have died before 1939

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