Evolution - Wikipedia

6/23/2017 EvolutionWikipedia
Evolution
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Evolutionischangeintheheritablecharacteristicsofbiologicalpopulationsoversuccessivegenerations.[1][2]
Evolutionaryprocessesgiverisetobiodiversityateverylevelofbiologicalorganisation,includingthelevelsof
species,individualorganisms,andmolecules.[3]
AlllifeonEarthsharesacommonancestorknownasthelastuniversalcommonancestor(LUCA),[4][5][6]which
livedapproximately3.53.8billionyearsago,[7]althoughastudyin2015found"remainsofbioticlife"from4.1
billionyearsagoinancientrocksinWesternAustralia.[8][9]InJuly2016,scientistsreportedidentifyingasetof
355genesfromtheLUCAofallorganismslivingonEarth.[10]
Repeatedformationofnewspecies(speciation),changewithinspecies(anagenesis),andlossofspecies
(extinction)throughouttheevolutionaryhistoryoflifeonEartharedemonstratedbysharedsetsofmorphological
andbiochemicaltraits,includingsharedDNAsequences.[11]Thesesharedtraitsaremoresimilaramongspecies
thatshareamorerecentcommonancestor,andcanbeusedtoreconstructabiological"treeoflife"basedon
evolutionaryrelationships(phylogenetics),usingbothexistingspeciesandfossils.Thefossilrecordincludesa
progressionfromearlybiogenicgraphite,[12]tomicrobialmatfossils,[13][14][15]tofossilisedmulticellular
organisms.Existingpatternsofbiodiversityhavebeenshapedbothbyspeciationandbyextinction.[16]Morethan
99percentofallspeciesthateverlivedonEarthareestimatedtobeextinct.[17][18]EstimatesofEarth'scurrent
speciesrangefrom10to14million,[19][20]ofwhichabout1.9millionareestimatedtohavebeennamed[21]and1.6
milliondocumentedinacentraldatabasetodate.[22]Morerecently,inMay2016,scientistsreportedthat1trillion
speciesareestimatedtobeonEarthcurrentlywithonlyonethousandthofonepercentdescribed.[23]
Inthemid19thcentury,CharlesDarwinformulatedthescientifictheoryofevolutionbynaturalselection,
publishedinhisbookOntheOriginofSpecies(1859).Evolutionbynaturalselectionisaprocessdemonstratedby
theobservationthatmoreoffspringareproducedthancanpossiblysurvive,alongwiththreefactsabout
populations:1)traitsvaryamongindividualswithrespecttomorphology,physiology,andbehaviour(phenotypic
variation),2)differenttraitsconferdifferentratesofsurvivalandreproduction(differentialfitness),and3)traits
canbepassedfromgenerationtogeneration(heritabilityoffitness).[24]Thus,insuccessivegenerationsmembers
ofapopulationarereplacedbyprogenyofparentsbetteradaptedtosurviveandreproduceinthebiophysical
environmentinwhichnaturalselectiontakesplace.Thisteleonomyisthequalitywherebytheprocessofnatural
selectioncreatesandpreservestraitsthatareseeminglyfittedforthefunctionalrolestheyperform.[25]The
processesbywhichthechangesoccur,fromonegenerationtoanother,arecalledevolutionaryprocessesor
mechanisms.[26]Thefourmostwidelyrecognisedevolutionaryprocessesarenaturalselection(includingsexual
selection),geneticdrift,mutationandgenemigrationduetogeneticadmixture.[26]Naturalselectionandgenetic
driftsortvariationmutationandgenemigrationcreatevariation.[26]
Consequencesofselectioncanincludemeioticdrive[27](unequaltransmissionofcertainalleles),nonrandom
mating[28]andgenetichitchhiking.Intheearly20thcenturythemodernevolutionarysynthesisintegratedclassical
geneticswithDarwin'stheoryofevolutionbynaturalselectionthroughthedisciplineofpopulationgenetics.The
importanceofnaturalselectionasacauseofevolutionwasacceptedintootherbranchesofbiology.Moreover,
previouslyheldnotionsaboutevolution,suchasorthogenesis,evolutionism,andotherbeliefsaboutinnate
https://en.wikipedia.org/wiki/Evolution 1/51
"progress"withinthelargestscaletrendsinevolution,becameobsolete.[29]Scientistscontinuetostudyvarious
aspectsofevolutionarybiologybyformingandtestinghypotheses,constructingmathematicalmodelsof
theoreticalbiologyandbiologicaltheories,usingobservationaldata,andperformingexperimentsinboththefield
andthelaboratory.
Intermsofpracticalapplication,anunderstandingofevolutionhasbeeninstrumentaltodevelopmentsin
numerousscientificandindustrialfields,includingagriculture,humanandveterinarymedicine,andthelife
sciencesingeneral.[30][31][32]Discoveriesinevolutionarybiologyhavemadeasignificantimpactnotjustinthe
traditionalbranchesofbiologybutalsoinotheracademicdisciplines,includingbiologicalanthropology,and
evolutionarypsychology.[33][34]Evolutionarycomputation,asubfieldofartificialintelligence,involvesthe
applicationofDarwinianprinciplestoproblemsincomputerscience.
Contents
1 Historyofevolutionarythought
1.1 Classicaltimes
1.2 Medieval
1.3 PreDarwinian
1.4 Darwinianrevolution
1.5 Pangenesis
1.6 The'modernsynthesis'
1.7 Furthersyntheses
2 Heredity
3 Variation
3.1 Mutation
3.2 Sexandrecombination
3.3 Geneflow
4 Mechanisms
4.1 Naturalselection
4.2 Biasedmutation
4.3 Geneticdrift
4.4 Genetichitchhiking
4.5 Geneflow
5 Outcomes
5.1 Adaptation
5.2 Coevolution
5.3 Cooperation
5.4 Speciation
5.5 Extinction
6 Evolutionaryhistoryoflife
6.1 Originoflife
6.2 Commondescent
6.3 Evolutionoflife
7 Applications
8 Socialandculturalresponses
9 Seealso
10 References
11 Bibliography
12 Furtherreading
13 Externallinks
Historyofevolutionarythought
Classicaltimes
Theproposalthatonetypeoforganismcoulddescendfromanothertypegoesbackto
someofthefirstpreSocraticGreekphilosophers,suchasAnaximanderand
Empedocles.[36]SuchproposalssurvivedintoRomantimes.Thepoetandphilosopher
LucretiusfollowedEmpedoclesinhismasterworkDererumnatura(OntheNatureof
Things).[37][38]
Medieval
Incontrasttothesematerialisticviews,Aristotelianismconsideredallnaturalthingsas
actualisationsoffixednaturalpossibilities,knownasforms.[39][40]Thiswaspartofa Lucretius
medievalteleologicalunderstandingofnatureinwhichallthingshaveanintended
roletoplayinadivinecosmicorder.Variationsofthisideabecamethestandard
understandingoftheMiddleAgesandwereintegratedintoChristianlearning,but
Aristotledidnotdemandthatrealtypesoforganismsalwayscorrespondoneforone
withexactmetaphysicalformsandspecificallygaveexamplesofhownewtypesof
livingthingscouldcometobe.[41]
PreDarwinian
Inthe17thcentury,thenewmethodofmodernsciencerejectedtheAristotelian
approach.Itsoughtexplanationsofnaturalphenomenaintermsofphysicallawsthat
werethesameforallvisiblethingsandthatdidnotrequiretheexistenceofanyfixed
naturalcategoriesordivinecosmicorder.However,thisnewapproachwasslowto
takerootinthebiologicalsciences,thelastbastionoftheconceptoffixednatural
AlfredRusselWallace
types.JohnRayappliedoneofthepreviouslymoregeneraltermsforfixednatural
types,"species,"toplantandanimaltypes,buthestrictlyidentifiedeachtypeofliving
thingasaspeciesandproposedthateachspeciescouldbedefinedbythefeaturesthat
perpetuatedthemselvesgenerationaftergeneration.[42]Thebiologicalclassification
introducedbyCarlLinnaeusin1735explicitlyrecognisedthehierarchicalnatureof
speciesrelationships,butstillviewedspeciesasfixedaccordingtoadivineplan.[43]
Othernaturalistsofthistimespeculatedontheevolutionarychangeofspeciesover
timeaccordingtonaturallaws.In1751,PierreLouisMaupertuiswroteofnatural
modificationsoccurringduringreproductionandaccumulatingovermanygenerations
toproducenewspecies.[44]GeorgesLouisLeclerc,ComtedeBuffonsuggestedthat
speciescoulddegenerateintodifferentorganisms,andErasmusDarwinproposedthat
allwarmbloodedanimalscouldhavedescendedfromasinglemicroorganism(or ThomasRobertMalthus
"filament").[45]ThefirstfullfledgedevolutionaryschemewasJeanBaptiste
Lamarck's"transmutation"theoryof1809,[46]whichenvisagedspontaneousgenerationcontinuallyproducing
simpleformsoflifethatdevelopedgreatercomplexityinparallellineageswithaninherentprogressivetendency,
andpostulatedthatonalocalleveltheselineagesadaptedtotheenvironmentbyinheritingchangescausedbytheir
useordisuseinparents.[47][48](ThelatterprocesswaslatercalledLamarckism.)[47][49][50][51]Theseideaswere
condemnedbyestablishednaturalistsasspeculationlackingempiricalsupport.Inparticular,GeorgesCuvier
insistedthatspecieswereunrelatedandfixed,theirsimilaritiesreflectingdivinedesignforfunctionalneeds.Inthe
meantime,Ray'sideasofbenevolentdesignhadbeendevelopedbyWilliamPaleyinto
theNaturalTheologyorEvidencesoftheExistenceandAttributesoftheDeity(1802),
whichproposedcomplexadaptationsasevidenceofdivinedesignandwhichwas
admiredbyCharlesDarwin.[52][53][54]
Darwinianrevolution
Thecrucialbreakfromtheconceptofconstanttypologicalclassesortypesinbiology
camewiththetheoryofevolutionthroughnaturalselection,whichwasformulatedby
CharlesDarwinintermsofvariablepopulations.PartlyinfluencedbyAnEssayonthe
PrincipleofPopulation(1798)byThomasRobertMalthus,Darwinnotedthat
StatisticianRonaldFisher
populationgrowthwouldleadtoa"struggleforexistence"inwhichfavorable
variationsprevailedasothersperished.Ineachgeneration,manyoffspringfailto
survivetoanageofreproductionbecauseoflimitedresources.Thiscouldexplainthe
diversityofplantsandanimalsfromacommonancestrythroughtheworkingof
naturallawsinthesamewayforalltypesoforganism.[55][56][57][58]Darwindeveloped
histheoryof"naturalselection"from1838onwardsandwaswritinguphis"bigbook"
onthesubjectwhenAlfredRusselWallacesenthimaversionofvirtuallythesame
theoryin1858.Theirseparatepaperswerepresentedtogetherata1858meetingofthe
LinneanSocietyofLondon.[59]Attheendof1859,Darwin'spublicationofhis
"abstract"asOntheOriginofSpeciesexplainednaturalselectionindetailandina
waythatledtoanincreasinglywideacceptanceofDarwin'sconceptsofevolutionat
theexpenseofalternativetheories.ThomasHenryHuxleyappliedDarwin'sideasto
humans,usingpaleontologyandcomparativeanatomytoprovidestrongevidencethat In1842,CharlesDarwin
humansandapessharedacommonancestry.Someweredisturbedbythissinceit pennedhisfirstsketchof
impliedthathumansdidnothaveaspecialplaceintheuniverse.[60] OntheOriginof
Species. [35]
Pangenesis
Themechanismsofreproductiveheritabilityandtheoriginofnewtraitsremainedamystery.Towardsthisend,
Darwindevelopedhisprovisionaltheoryofpangenesis.[61]In1865,GregorMendelreportedthattraitswere
inheritedinapredictablemannerthroughtheindependentassortmentandsegregationofelements(laterknownas
genes).Mendel'slawsofinheritanceeventuallysupplantedmostofDarwin'spangenesistheory.[62]August
Weismannmadetheimportantdistinctionbetweengermcellsthatgiverisetogametes(suchasspermandegg
cells)andthesomaticcellsofthebody,demonstratingthathereditypassesthroughthegermlineonly.Hugode
VriesconnectedDarwin'spangenesistheorytoWeismann'sgerm/somacelldistinctionandproposedthatDarwin's
pangeneswereconcentratedinthecellnucleusandwhenexpressedtheycouldmoveintothecytoplasmtochange
thecellsstructure.DeVrieswasalsooneoftheresearcherswhomadeMendel'sworkwellknown,believingthat
Mendeliantraitscorrespondedtothetransferofheritablevariationsalongthegermline.[63]Toexplainhownew
variantsoriginate,deVriesdevelopedamutationtheorythatledtoatemporaryriftbetweenthosewhoaccepted
DarwinianevolutionandbiometricianswhoalliedwithdeVries.[48][64][65]Inthe1930s,pioneersinthefieldof
populationgenetics,suchasRonaldFisher,SewallWrightandJ.B.S.Haldanesetthefoundationsofevolution
ontoarobuststatisticalphilosophy.ThefalsecontradictionbetweenDarwin'stheory,geneticmutations,and
Mendelianinheritancewasthusreconciled.[66]
The'modernsynthesis'
Inthe1920sand1930sthesocalledmodernsynthesisconnectednaturalselection,mutationtheory,and
Mendelianinheritanceintoaunifiedtheorythatappliedgenerallytoanybranchofbiology.Themodernsynthesis
explainedpatternsobservedacrossspeciesinpopulations,throughfossiltransitionsinpalaeontology,andcomplex
cellularmechanismsindevelopmentalbiology.[48][67]ThepublicationofthestructureofDNAbyJamesWatson
andFrancisCrickin1953demonstratedaphysicalmechanismforinheritance.[68]Molecularbiologyimprovedour
understandingoftherelationshipbetweengenotypeandphenotype.Advancementswerealsomadeinphylogenetic
systematics,mappingthetransitionoftraitsintoacomparativeandtestableframeworkthroughthepublicationand
useofevolutionarytrees.[69][70]In1973,evolutionarybiologistTheodosiusDobzhanskypennedthat"nothingin
biologymakessenseexceptinthelightofevolution,"becauseithasbroughttolighttherelationsofwhatfirst
seemeddisjointedfactsinnaturalhistoryintoacoherentexplanatorybodyofknowledgethatdescribesand
predictsmanyobservablefactsaboutlifeonthisplanet.[71]
Furthersyntheses
Sincethen,themodernsynthesishasbeenfurtherextendedtoexplainbiologicalphenomenaacrossthefulland
integrativescaleofthebiologicalhierarchy,fromgenestospecies.Thisextension,knownasevolutionary
developmentalbiologyandinformallycalled"evodevo,"emphasiseshowchangesbetweengenerations
(evolution)actsonpatternsofchangewithinindividualorganisms(development).[72][73][74]Sincethebeginningof
the21stcenturyandinlightofdiscoveriesmadeinrecentdecades,somebiologistshavearguedforanextended
evolutionarysynthesis,whichwouldaccountfortheeffectsofnongeneticinheritancemodes,suchasepigenetics,
parentaleffects,ecologicalandculturalinheritance,andevolvability.[75][76]
Heredity
Evolutioninorganismsoccursthroughchangesinheritabletraitstheinherited
characteristicsofanorganism.Inhumans,forexample,eyecolourisaninherited
characteristicandanindividualmightinheritthe"browneyetrait"fromoneoftheir
parents.[77]Inheritedtraitsarecontrolledbygenesandthecompletesetofgeneswithin
anorganism'sgenome(geneticmaterial)iscalleditsgenotype.[78]
Thecompletesetofobservabletraitsthatmakeupthestructureandbehaviourofan
organismiscalleditsphenotype.Thesetraitscomefromtheinteractionofitsgenotype
withtheenvironment.[79]Asaresult,manyaspectsofanorganism'sphenotypearenot
inherited.Forexample,suntannedskincomesfromtheinteractionbetweenaperson's
genotypeandsunlightthus,suntansarenotpassedontopeople'schildren.However,
somepeopletanmoreeasilythanothers,duetodifferencesingenotypicvariationa
strikingexamplearepeoplewiththeinheritedtraitofalbinism,whodonottanatalland
areverysensitivetosunburn.[80]
DNAstructure.Bases
areinthecentre,
HeritabletraitsarepassedfromonegenerationtothenextviaDNA,amoleculethat
surroundedby
encodesgeneticinformation.[78]DNAisalongbiopolymercomposedoffourtypesof phosphatesugar
bases.ThesequenceofbasesalongaparticularDNAmoleculespecifythegenetic chainsinadouble
information,inamannersimilartoasequenceoflettersspellingoutasentence.Beforea helix.
celldivides,theDNAiscopied,sothateachoftheresultingtwocellswillinheritthe
DNAsequence.PortionsofaDNAmoleculethatspecifyasinglefunctionalunitare
calledgenesdifferentgeneshavedifferentsequencesofbases.Withincells,thelongstrandsofDNAform
condensedstructurescalledchromosomes.ThespecificlocationofaDNAsequencewithinachromosomeis
knownasalocus.IftheDNAsequenceatalocusvariesbetweenindividuals,thedifferentformsofthissequence
arecalledalleles.DNAsequencescanchangethroughmutations,producingnewalleles.Ifamutationoccurs
withinagene,thenewallelemayaffectthetraitthatthegenecontrols,alteringthephenotypeoftheorganism.[81]
However,whilethissimplecorrespondencebetweenanalleleandatraitworksinsomecases,mosttraitsaremore
complexandarecontrolledbyquantitativetraitloci(multipleinteractinggenes).[82][83]
Recentfindingshaveconfirmedimportantexamplesofheritablechangesthatcannotbeexplainedbychangesto
thesequenceofnucleotidesintheDNA.Thesephenomenaareclassedasepigeneticinheritancesystems.[84]DNA
methylationmarkingchromatin,selfsustainingmetabolicloops,genesilencingbyRNAinterferenceandthe
threedimensionalconformationofproteins(suchasprions)areareaswhereepigeneticinheritancesystemshave
beendiscoveredattheorganismiclevel.[85][86]Developmentalbiologistssuggestthatcomplexinteractionsin
geneticnetworksandcommunicationamongcellscanleadtoheritablevariationsthatmayunderlaysomeofthe
mechanicsindevelopmentalplasticityandcanalisation.[87]Heritabilitymayalsooccuratevenlargerscales.For
example,ecologicalinheritancethroughtheprocessofnicheconstructionisdefinedbytheregularandrepeated
activitiesoforganismsintheirenvironment.Thisgeneratesalegacyofeffectsthatmodifyandfeedbackintothe
selectionregimeofsubsequentgenerations.Descendantsinheritgenesplusenvironmentalcharacteristics
generatedbytheecologicalactionsofancestors.[88]Otherexamplesofheritabilityinevolutionthatarenotunder
thedirectcontrolofgenesincludetheinheritanceofculturaltraitsandsymbiogenesis.[89][90]
Variation
Anindividualorganism'sphenotyperesultsfrombothitsgenotypeandthe
influencefromtheenvironmentithaslivedin.Asubstantialpartofthe
phenotypicvariationinapopulationiscausedbygenotypicvariation.[83]The
modernevolutionarysynthesisdefinesevolutionasthechangeovertimein
thisgeneticvariation.Thefrequencyofoneparticularallelewillbecomemore
orlessprevalentrelativetootherformsofthatgene.Variationdisappears
whenanewallelereachesthepointoffixationwheniteitherdisappears
fromthepopulationorreplacestheancestralalleleentirely.[91] Whitepepperedmoth
Naturalselectionwillonlycauseevolutionifthereisenoughgeneticvariation
inapopulation.BeforethediscoveryofMendeliangenetics,onecommon
hypothesiswasblendinginheritance.Butwithblendinginheritance,genetic
variancewouldberapidlylost,makingevolutionbynaturalselection
implausible.TheHardyWeinbergprincipleprovidesthesolutiontohow
Blackmorphinpepperedmoth
variationismaintainedinapopulationwithMendelianinheritance.The
evolution
frequenciesofalleles(variationsinagene)willremainconstantinthe
absenceofselection,mutation,migrationandgeneticdrift.[92]
Variationcomesfrommutationsinthegenome,reshufflingofgenesthroughsexualreproductionandmigration
betweenpopulations(geneflow).Despitetheconstantintroductionofnewvariationthroughmutationandgene
flow,mostofthegenomeofaspeciesisidenticalinallindividualsofthatspecies.[93]However,evenrelatively
smalldifferencesingenotypecanleadtodramaticdifferencesinphenotype:forexample,chimpanzeesand
humansdifferinonlyabout5%oftheirgenomes.[94]
Mutation
MutationsarechangesintheDNAsequenceofacell'sgenome.Whenmutationsoccur,they
mayaltertheproductofagene,orpreventthegenefromfunctioning,orhavenoeffect.Based
onstudiesintheflyDrosophilamelanogaster,ithasbeensuggestedthatifamutation
changesaproteinproducedbyagene,thiswillprobablybeharmful,withabout70%ofthese
mutationshavingdamagingeffects,andtheremainderbeingeitherneutralorweakly
beneficial.[95]
Mutationscaninvolvelargesectionsofachromosomebecomingduplicated(usuallyby
geneticrecombination),whichcanintroduceextracopiesofageneintoagenome.[96]Extra
copiesofgenesareamajorsourceoftherawmaterialneededfornewgenestoevolve.[97]
Thisisimportantbecausemostnewgenesevolvewithingenefamiliesfrompreexisting
genesthatsharecommonancestors.[98]Forexample,thehumaneyeusesfourgenestomake
structuresthatsenselight:threeforcolourvisionandonefornightvisionallfourare
descendedfromasingleancestralgene.[99]
Duplicationof
Newgenescanbegeneratedfromanancestralgenewhenaduplicatecopymutatesand
partofa
acquiresanewfunction.Thisprocessiseasieronceagenehasbeenduplicatedbecauseit
chromosome
increasestheredundancyofthesystemonegeneinthepaircanacquireanewfunctionwhile
theothercopycontinuestoperformitsoriginalfunction.[100][101]Othertypesofmutationscan
evengenerateentirelynewgenesfrompreviouslynoncodingDNA.[102][103]
Thegenerationofnewgenescanalsoinvolvesmallpartsofseveralgenesbeingduplicated,withthesefragments
thenrecombiningtoformnewcombinationswithnewfunctions.[104][105]Whennewgenesareassembledfrom
shufflingpreexistingparts,domainsactasmoduleswithsimpleindependentfunctions,whichcanbemixed
togethertoproducenewcombinationswithnewandcomplexfunctions.[106]Forexample,polyketidesynthasesare
largeenzymesthatmakeantibioticstheycontainuptoonehundredindependentdomainsthateachcatalyseone
stepintheoverallprocess,likeastepinanassemblyline.[107]
Sexandrecombination
Inasexualorganisms,genesareinheritedtogether,orlinked,astheycannotmixwithgenesofotherorganisms
duringreproduction.Incontrast,theoffspringofsexualorganismscontainrandommixturesoftheirparents'
chromosomesthatareproducedthroughindependentassortment.Inarelatedprocesscalledhomologous
recombination,sexualorganismsexchangeDNAbetweentwomatchingchromosomes.[108]Recombinationand
reassortmentdonotalterallelefrequencies,butinsteadchangewhichallelesareassociatedwitheachother,
producingoffspringwithnewcombinationsofalleles.[109]Sexusuallyincreasesgeneticvariationandmay
increasetherateofevolution.[110][111]
ThetwofoldcostofsexwasfirstdescribedbyJohnMaynardSmith.[112]Thefirstcostisthatinsexually
dimorphicspeciesonlyoneofthetwosexescanbearyoung.(Thiscostdoesnotapplytohermaphroditicspecies,
likemostplantsandmanyinvertebrates.)Thesecondcostisthatanyindividualwhoreproducessexuallycanonly
passon50%ofitsgenestoanyindividualoffspring,withevenlesspassedonaseachnewgenerationpasses.[113]
Yetsexualreproductionisthemorecommonmeansofreproductionamongeukaryotesandmulticellular
organisms.TheRedQueenhypothesishasbeenusedtoexplainthesignificanceofsexualreproductionasameans
toenablecontinualevolutionandadaptationinresponsetocoevolutionwithotherspeciesinaneverchanging
environment.[113][114][115][116]
Geneflow
Geneflowistheexchangeofgenesbetweenpopulationsandbetween
species.[117]Itcanthereforebeasourceofvariationthatisnewtoa
populationortoaspecies.Geneflowcanbecausedbythemovement
ofindividualsbetweenseparatepopulationsoforganisms,asmightbe
causedbythemovementofmicebetweeninlandandcoastal
populations,orthemovementofpollenbetweenheavymetaltolerant
andheavymetalsensitivepopulationsofgrasses.
Genetransferbetweenspeciesincludestheformationofhybrid Thisdiagramillustratesthetwofoldcostof
organismsandhorizontalgenetransfer.Horizontalgenetransferisthe sex.Ifeachindividualweretocontribute
transferofgeneticmaterialfromoneorganismtoanotherorganism tothesamenumberofoffspring(two),(a)
thatisnotitsoffspringthisismostcommonamongbacteria. [118]In thesexualpopulationremainsthesamesize
medicine,thiscontributestothespreadofantibioticresistance,as eachgeneration,wherethe(b)Asexual
whenonebacteriaacquiresresistancegenesitcanrapidlytransfer reproductionpopulationdoublesinsize
themtootherspecies. [119]Horizontaltransferofgenesfrombacteriato eachgeneration.
eukaryotessuchastheyeastSaccharomycescerevisiaeandtheadzuki
beanweevilCallosobruchuschinensishasoccurred.[120][121]Anexampleoflargerscaletransfersarethe
eukaryoticbdelloidrotifers,whichhavereceivedarangeofgenesfrombacteria,fungiandplants.[122]Virusescan
alsocarryDNAbetweenorganisms,allowingtransferofgenesevenacrossbiologicaldomains.[123]
Largescalegenetransferhasalsooccurredbetweentheancestorsofeukaryoticcellsandbacteria,duringthe
acquisitionofchloroplastsandmitochondria.Itispossiblethateukaryotesthemselvesoriginatedfromhorizontal
genetransfersbetweenbacteriaandarchaea.[124]
Mechanisms
FromaNeoDarwinianperspective,evolutionoccurswhen
therearechangesinthefrequenciesofalleleswithina
populationofinterbreedingorganisms.[92]Forexample,the
alleleforblackcolourinapopulationofmothsbecomingmore
common.Mechanismsthatcanleadtochangesinallele
frequenciesincludenaturalselection,geneticdrift,genetic
hitchhiking,mutationandgeneflow.
Naturalselection
Evolutionbymeansofnaturalselectionistheprocessby
whichtraitsthatenhancesurvivalandreproductionbecome
morecommoninsuccessivegenerationsofapopulation.Ithas
oftenbeencalleda"selfevident"mechanismbecauseit
necessarilyfollowsfromthreesimplefacts:[24]
Mutationfollowedbynaturalselectionresultsina
Variationexistswithinpopulationsoforganismswith populationwithdarkercolouration.
respecttomorphology,physiology,andbehaviour
(phenotypicvariation).
Differenttraitsconferdifferentratesofsurvivalandreproduction(differentialfitness).
Thesetraitscanbepassedfromgenerationtogeneration(heritabilityoffitness).
Moreoffspringareproducedthancanpossiblysurvive,andtheseconditionsproducecompetitionbetween
organismsforsurvivalandreproduction.Consequently,organismswithtraitsthatgivethemanadvantageover
theircompetitorsaremorelikelytopassontheirtraitstothenextgenerationthanthosewithtraitsthatdonot
conferanadvantage.[125]
Thecentralconceptofnaturalselectionistheevolutionaryfitnessofanorganism.[126]Fitnessismeasuredbyan
organism'sabilitytosurviveandreproduce,whichdeterminesthesizeofitsgeneticcontributiontothenext
generation.[126]However,fitnessisnotthesameasthetotalnumberofoffspring:insteadfitnessisindicatedbythe
proportionofsubsequentgenerationsthatcarryanorganism'sgenes.[127]Forexample,ifanorganismcould
survivewellandreproducerapidly,butitsoffspringwerealltoosmallandweaktosurvive,thisorganismwould
makelittlegeneticcontributiontofuturegenerationsandwouldthushavelowfitness.[126]
Ifanalleleincreasesfitnessmorethantheotherallelesofthatgene,thenwitheachgenerationthisallelewill
becomemorecommonwithinthepopulation.Thesetraitsaresaidtobe"selectedfor."Examplesoftraitsthatcan
increasefitnessareenhancedsurvivalandincreasedfecundity.Conversely,thelowerfitnesscausedbyhavinga
lessbeneficialordeleteriousalleleresultsinthisallelebecomingrarertheyare"selectedagainst."[128]
Importantly,thefitnessofanalleleisnotafixedcharacteristiciftheenvironmentchanges,previouslyneutralor
harmfultraitsmaybecomebeneficialandpreviouslybeneficialtraitsbecomeharmful.[81]However,evenifthe
directionofselectiondoesreverseinthisway,traitsthatwerelostinthepastmaynotreevolveinanidentical
form(seeDollo'slaw).[129][130]
Naturalselectionwithinapopulationforatraitthatcanvaryacrossarangeofvalues,suchasheight,canbe
categorisedintothreedifferenttypes.Thefirstisdirectionalselection,whichisashiftintheaveragevalueofa
traitovertimeforexample,organismsslowlygettingtaller.[131]Secondly,disruptiveselectionisselectionfor
extremetraitvaluesandoftenresultsintwodifferentvaluesbecomingmostcommon,withselectionagainstthe
averagevalue.Thiswouldbewheneithershortortallorganismshadanadvantage,butnotthoseofmedium
height.Finally,instabilisingselectionthereisselectionagainstextremetraitvaluesonbothends,whichcausesa
decreaseinvariancearoundtheaveragevalueandlessdiversity.[125][132]Thiswould,forexample,causeorganisms
toslowlybecomeallthesameheight.
Aspecialcaseofnaturalselectionissexualselection,whichisselectionforanytraitthatincreasesmatingsuccess
byincreasingtheattractivenessofanorganismtopotentialmates.[133]Traitsthatevolvedthroughsexualselection
areparticularlyprominentamongmalesofseveralanimalspecies.Althoughsexuallyfavoured,traitssuchas
cumbersomeantlers,matingcalls,largebodysizeandbrightcoloursoftenattractpredation,whichcompromises
thesurvivalofindividualmales.[134][135]Thissurvivaldisadvantageisbalancedbyhigherreproductivesuccessin
malesthatshowthesehardtofake,sexuallyselectedtraits.[136]
Naturalselectionmostgenerallymakesnaturethemeasureagainstwhichindividualsandindividualtraits,are
moreorlesslikelytosurvive."Nature"inthissensereferstoanecosystem,thatis,asysteminwhichorganisms
interactwitheveryotherelement,physicalaswellasbiological,intheirlocalenvironment.EugeneOdum,a
founderofecology,definedanecosystemas:"Anyunitthatincludesalloftheorganisms...inagivenarea
interactingwiththephysicalenvironmentsothataflowofenergyleadstoclearlydefinedtrophicstructure,biotic
diversityandmaterialcycles(ie:exchangeofmaterialsbetweenlivingandnonlivingparts)withinthesystem."[137]
Eachpopulationwithinanecosystemoccupiesadistinctniche,orposition,withdistinctrelationshipstootherparts
ofthesystem.Theserelationshipsinvolvethelifehistoryoftheorganism,itspositioninthefoodchainandits
geographicrange.Thisbroadunderstandingofnatureenablesscientiststodelineatespecificforceswhich,
together,comprisenaturalselection.
Naturalselectioncanactatdifferentlevelsoforganisation,
suchasgenes,cells,individualorganisms,groupsof
organismsandspecies.[138][139][140]Selectioncanactat
multiplelevelssimultaneously.[141]Anexampleofselection
occurringbelowtheleveloftheindividualorganismare
genescalledtransposons,whichcanreplicateandspread
throughoutagenome.[142]Selectionatalevelabovethe
individual,suchasgroupselection,mayallowtheevolution
ofcooperation,asdiscussedbelow.[143]
Biasedmutation
Inadditiontobeingamajorsourceofvariation,mutation
mayalsofunctionasamechanismofevolutionwhenthere
aredifferentprobabilitiesatthemolecularlevelfordifferent
mutationstooccur,aprocessknownasmutationbias.[144]If
Thesechartsdepictthedifferenttypesofgenetic twogenotypes,forexampleonewiththenucleotideGand
selection.Oneachgraph,thexaxisvariableisthe anotherwiththenucleotideAinthesameposition,havethe
typeofphenotypictraitandtheyaxisvariableisthe samefitness,butmutationfromGtoAhappensmoreoften
numberoforganisms.GroupAistheoriginal thanmutationfromAtoG,thengenotypeswithAwilltend
populationandGroupBisthepopulationafter toevolve.[145]Differentinsertionvs.deletionmutation
selection. biasesindifferenttaxacanleadtotheevolutionofdifferent
Graph1showsdirectionalselection,inwhicha genomesizes.[146][147]Developmentalormutationalbiases
singleextremephenotypeisfavored. havealsobeenobservedinmorphologicalevolution.[148][149]
Graph2depictsstabilizingselection,wherethe Forexample,accordingtothephenotypefirsttheoryof
intermediatephenotypeisfavoredovertheextreme evolution,mutationscaneventuallycausethegenetic
traits. assimilationoftraitsthatwerepreviouslyinducedbythe
Graph3showsdisruptiveselection,inwhichthe
extremephenotypesarefavoredovertheintermediate.
environment.[150][151][152]
Mutationbiaseffectsaresuperimposedonotherprocesses.
Ifselectionwouldfavoreitheroneoutoftwomutations,butthereisnoextraadvantagetohavingboth,thenthe
mutationthatoccursthemostfrequentlyistheonethatismostlikelytobecomefixedinapopulation.[153][154]
Mutationsleadingtothelossoffunctionofagenearemuchmorecommonthanmutationsthatproduceanew,
fullyfunctionalgene.Mostlossoffunctionmutationsareselectedagainst.Butwhenselectionisweak,mutation
biastowardslossoffunctioncanaffectevolution.[155]Forexample,pigmentsarenolongerusefulwhenanimals
liveinthedarknessofcaves,andtendtobelost.[156]Thiskindoflossoffunctioncanoccurbecauseofmutation
bias,and/orbecausethefunctionhadacost,andoncethebenefitofthefunctiondisappeared,naturalselection
leadstotheloss.LossofsporulationabilityinBacillussubtilisduringlaboratoryevolutionappearstohavebeen
causedbymutationbias,ratherthannaturalselectionagainstthecostofmaintainingsporulationability.[157]When
thereisnoselectionforlossoffunction,thespeedatwhichlossevolvesdependsmoreonthemutationratethanit
doesontheeffectivepopulationsize,[158]indicatingthatitisdrivenmorebymutationbiasthanbygeneticdrift.In
parasiticorganisms,mutationbiasleadstoselectionpressuresasseeninEhrlichia.Mutationsarebiasedtowards
antigenicvariantsinoutermembraneproteins.
Geneticdrift
Geneticdriftisthechangeinallelefrequencyfromonegenerationtothe
nextthatoccursbecauseallelesaresubjecttosamplingerror.[159]Asa
result,whenselectiveforcesareabsentorrelativelyweak,allele
frequenciestendto"drift"upwardordownwardrandomly(inarandom
walk).Thisdrifthaltswhenanalleleeventuallybecomesfixed,eitherby
disappearingfromthepopulation,orreplacingtheotherallelesentirely.
Geneticdriftmaythereforeeliminatesomeallelesfromapopulationdueto
chancealone.Evenintheabsenceofselectiveforces,geneticdriftcan
causetwoseparatepopulationsthatbeganwiththesamegeneticstructure
todriftapartintotwodivergentpopulationswithdifferentsetsof
alleles.[160]
Itisusuallydifficulttomeasuretherelativeimportanceofselectionand
neutralprocesses,includingdrift.[161]Thecomparativeimportanceof Simulationofgeneticdriftof20
adaptiveandnonadaptiveforcesindrivingevolutionarychangeisanarea unlinkedallelesinpopulationsof10
ofcurrentresearch.[162] (top)and100(bottom).Driftto
fixationismorerapidinthesmaller
Theneutraltheoryofmolecularevolutionproposedthatmostevolutionary population.
changesaretheresultofthefixationofneutralmutationsbygenetic
drift.[163]Hence,inthismodel,mostgeneticchangesinapopulationarethe
resultofconstantmutationpressureandgeneticdrift.[164]Thisformoftheneutraltheoryisnowlargely
abandoned,sinceitdoesnotseemtofitthegeneticvariationseeninnature.[165][166]However,amorerecentand
bettersupportedversionofthismodelisthenearlyneutraltheory,whereamutationthatwouldbeeffectively
neutralinasmallpopulationisnotnecessarilyneutralinalargepopulation.[125]Otheralternativetheoriespropose
thatgeneticdriftisdwarfedbyotherstochasticforcesinevolution,suchasgenetichitchhiking,alsoknownas
geneticdraft.[159][167][168]
Thetimeforaneutralalleletobecomefixedbygeneticdriftdependsonpopulationsize,withfixationoccurring
morerapidlyinsmallerpopulations.[169]Thenumberofindividualsinapopulationisnotcritical,butinsteada
measureknownastheeffectivepopulationsize.[170]Theeffectivepopulationisusuallysmallerthanthetotal
populationsinceittakesintoaccountfactorssuchasthelevelofinbreedingandthestageofthelifecycleinwhich
thepopulationisthesmallest.[170]Theeffectivepopulationsizemaynotbethesameforeverygeneinthesame
population.[171]
Genetichitchhiking
RecombinationallowsallelesonthesamestrandofDNAtobecomeseparated.However,therateofrecombination
islow(approximatelytwoeventsperchromosomepergeneration).Asaresult,genesclosetogetherona
chromosomemaynotalwaysbeshuffledawayfromeachotherandgenesthatareclosetogethertendtobe
inheritedtogether,aphenomenonknownaslinkage.[172]Thistendencyismeasuredbyfindinghowoftentwo
allelesoccurtogetheronasinglechromosomecomparedtoexpectations,whichiscalledtheirlinkage
disequilibrium.Asetofallelesthatisusuallyinheritedinagroupiscalledahaplotype.Thiscanbeimportant
whenonealleleinaparticularhaplotypeisstronglybeneficial:naturalselectioncandriveaselectivesweepthat
willalsocausetheotherallelesinthehaplotypetobecomemorecommoninthepopulationthiseffectiscalled
genetichitchhikingorgeneticdraft.[173]Geneticdraftcausedbythefactthatsomeneutralgenesaregenetically
linkedtoothersthatareunderselectioncanbepartiallycapturedbyanappropriateeffectivepopulationsize.[167]
Geneflow
Geneflowinvolvestheexchangeofgenesbetweenpopulationsandbetweenspecies.[117]Thepresenceorabsence
ofgeneflowfundamentallychangesthecourseofevolution.Duetothecomplexityoforganisms,anytwo
completelyisolatedpopulationswilleventuallyevolvegeneticincompatibilitiesthroughneutralprocesses,asin
theBatesonDobzhanskyMullermodel,evenifbothpopulationsremainessentiallyidenticalintermsoftheir
adaptationtotheenvironment.
Ifgeneticdifferentiationbetweenpopulationsdevelops,geneflowbetweenpopulationscanintroducetraitsor
alleleswhicharedisadvantageousinthelocalpopulationandthismayleadtoorganismswithinthesepopulations
evolvingmechanismsthatpreventmatingwithgeneticallydistantpopulations,eventuallyresultinginthe
appearanceofnewspecies.Thus,exchangeofgeneticinformationbetweenindividualsisfundamentallyimportant
forthedevelopmentofthebiologicalspeciesconcept.
Duringthedevelopmentofthemodernsynthesis,SewallWrightdevelopedhisshiftingbalancetheory,which
regardedgeneflowbetweenpartiallyisolatedpopulationsasanimportantaspectofadaptiveevolution.[174]
However,recentlytherehasbeensubstantialcriticismoftheimportanceoftheshiftingbalancetheory.[175]
Outcomes
Evolutioninfluenceseveryaspectoftheformand
behaviouroforganisms.Mostprominentarethespecific
behaviouralandphysicaladaptationsthataretheoutcome
ofnaturalselection.Theseadaptationsincreasefitnessby
aidingactivitiessuchasfindingfood,avoidingpredators
orattractingmates.Organismscanalsorespondto
selectionbycooperatingwitheachother,usuallyby
aidingtheirrelativesorengaginginmutuallybeneficial
symbiosis.Inthelongerterm,evolutionproducesnew
speciesthroughsplittingancestralpopulationsof
organismsintonewgroupsthatcannotorwillnot
Avisualdemonstrationofrapidantibioticresistance
interbreed.
evolutionbyE.coligrowingacrossaplatewith
Theseoutcomesofevolutionaredistinguishedbasedon increasingconcentrationsoftrimethoprim. [176]
timescaleasmacroevolutionversusmicroevolution.
Macroevolutionreferstoevolutionthatoccursatorabove
thelevelofspecies,inparticularspeciationandextinctionwhereasmicroevolutionreferstosmallerevolutionary
changeswithinaspeciesorpopulation,inparticularshiftsingenefrequencyandadaptation.[177]Ingeneral,
macroevolutionisregardedastheoutcomeoflongperiodsofmicroevolution.[178]Thus,thedistinctionbetween
microandmacroevolutionisnotafundamentalonethedifferenceissimplythetimeinvolved.[179]However,in
macroevolution,thetraitsoftheentirespeciesmaybeimportant.Forinstance,alargeamountofvariationamong
individualsallowsaspeciestorapidlyadapttonewhabitats,lesseningthechanceofitgoingextinct,whileawide
geographicrangeincreasesthechanceofspeciation,bymakingitmorelikelythatpartofthepopulationwill
becomeisolated.Inthissense,microevolutionandmacroevolutionmightinvolveselectionatdifferentlevels
withmicroevolutionactingongenesandorganisms,versusmacroevolutionaryprocessessuchasspeciesselection
actingonentirespeciesandaffectingtheirratesofspeciationandextinction.[180][181][182]
Acommonmisconceptionisthatevolutionhasgoals,longtermplans,oraninnatetendencyfor"progress,"as
expressedinbeliefssuchasorthogenesisandevolutionismrealisticallyhowever,evolutionhasnolongtermgoal
anddoesnotnecessarilyproducegreatercomplexity.[183][184][185]Althoughcomplexspecieshaveevolved,they
occurasasideeffectoftheoverallnumberoforganismsincreasingandsimpleformsoflifestillremainmore
commoninthebiosphere.[186]Forexample,theoverwhelmingmajorityofspeciesaremicroscopicprokaryotes,
whichformabouthalftheworld'sbiomassdespitetheirsmallsize,[187]andconstitutethevastmajorityofEarth's
biodiversity.[188]SimpleorganismshavethereforebeenthedominantformoflifeonEarththroughoutitshistory
andcontinuetobethemainformoflifeuptothepresentday,withcomplexlifeonlyappearingmorediverse
becauseitismorenoticeable.[189]Indeed,theevolutionofmicroorganismsisparticularlyimportanttomodern
evolutionaryresearch,sincetheirrapidreproductionallowsthestudyofexperimentalevolutionandthe
observationofevolutionandadaptationinrealtime.[190][191]
Adaptation
Adaptationistheprocessthatmakesorganismsbettersuitedto
theirhabitat.[192][193]Also,thetermadaptationmayrefertoa
traitthatisimportantforanorganism'ssurvival.Forexample,
theadaptationofhorses'teethtothegrindingofgrass.By
usingthetermadaptationfortheevolutionaryprocessand
adaptivetraitfortheproduct(thebodilypartorfunction),the
twosensesofthewordmaybedistinguished.Adaptationsare
producedbynaturalselection.[194]Thefollowingdefinitions
areduetoTheodosiusDobzhansky:
1.Adaptationistheevolutionaryprocesswherebyan
organismbecomesbetterabletoliveinitshabitator
habitats.[195] Homologousbonesinthelimbsoftetrapods.The
2.Adaptednessisthestateofbeingadapted:thedegreeto bonesoftheseanimalshavethesamebasic
whichanorganismisabletoliveandreproduceina structure,buthavebeenadaptedforspecificuses.
givensetofhabitats.[196]
3.Anadaptivetraitisanaspectofthedevelopmental
patternoftheorganismwhichenablesorenhancestheprobabilityofthatorganismsurvivingand
reproducing.[197]
Adaptationmaycauseeitherthegainofanewfeature,orthelossofanancestralfeature.Anexamplethatshows
bothtypesofchangeisbacterialadaptationtoantibioticselection,withgeneticchangescausingantibiotic
resistancebybothmodifyingthetargetofthedrug,orincreasingtheactivityoftransportersthatpumpthedrugout
ofthecell.[198]OtherstrikingexamplesarethebacteriaEscherichiacolievolvingtheabilitytousecitricacidasa
nutrientinalongtermlaboratoryexperiment,[199]Flavobacteriumevolvinganovelenzymethatallowsthese
bacteriatogrowonthebyproductsofnylonmanufacturing,[200][201]andthesoilbacteriumSphingobiumevolving
anentirelynewmetabolicpathwaythatdegradesthesyntheticpesticidepentachlorophenol.[202][203]Aninteresting
butstillcontroversialideaisthatsomeadaptationsmightincreasetheabilityoforganismstogenerategenetic
diversityandadaptbynaturalselection(increasingorganisms'evolvability).[204][205][206][207][208]
Adaptationoccursthroughthegradualmodificationofexistingstructures.Consequently,structureswithsimilar
internalorganisationmayhavedifferentfunctionsinrelatedorganisms.Thisistheresultofasingleancestral
structurebeingadaptedtofunctionindifferentways.Theboneswithinbatwings,forexample,areverysimilarto
thoseinmicefeetandprimatehands,duetothedescentofallthesestructuresfromacommonmammalian
ancestor.[210]However,sincealllivingorganismsarerelatedtosomeextent,[211]evenorgansthatappeartohave
littleornostructuralsimilarity,suchasarthropod,squidandvertebrateeyes,orthelimbsandwingsofarthropods
andvertebrates,candependonacommonsetofhomologousgenesthatcontroltheirassemblyandfunctionthisis
calleddeephomology.[212][213]
Duringevolution,somestructuresmaylosetheiroriginal
functionandbecomevestigialstructures.[214]Suchstructures
mayhavelittleornofunctioninacurrentspecies,yethavea
clearfunctioninancestralspecies,orothercloselyrelated
species.Examplesincludepseudogenes,[215]thenonfunctional
Abaleenwhaleskeleton,aandblabelflipper remainsofeyesinblindcavedwellingfish,[216]wingsin
bones,whichwereadaptedfromfrontlegbones:
flightlessbirds,[217]thepresenceofhipbonesinwhalesand
whilecindicatesvestigiallegbones,suggestingan
snakes,[209]andsexualtraitsinorganismsthatreproducevia
adaptationfromlandtosea. [209]
asexualreproduction.[218]Examplesofvestigialstructuresin
humansincludewisdomteeth,[219]thecoccyx,[214]the
vermiformappendix,[214]andotherbehaviouralvestigessuchasgoosebumps[220][221]andprimitive
reflexes.[222][223][224]
However,manytraitsthatappeartobesimpleadaptationsareinfactexaptations:structuresoriginallyadaptedfor
onefunction,butwhichcoincidentallybecamesomewhatusefulforsomeotherfunctionintheprocess.[225]One
exampleistheAfricanlizardHolaspisguentheri,whichdevelopedanextremelyflatheadforhidingincrevices,as
canbeseenbylookingatitsnearrelatives.However,inthisspecies,theheadhasbecomesoflattenedthatit
assistsinglidingfromtreetotreeanexaptation.[225]Withincells,molecularmachinessuchasthebacterial
flagella[226]andproteinsortingmachinery[227]evolvedbytherecruitmentofseveralpreexistingproteinsthat
previouslyhaddifferentfunctions.[177]Anotherexampleistherecruitmentofenzymesfromglycolysisand
xenobioticmetabolismtoserveasstructuralproteinscalledcrystallinswithinthelensesoforganisms'
eyes.[228][229]
Anareaofcurrentinvestigationinevolutionarydevelopmentalbiologyisthedevelopmentalbasisofadaptations
andexaptations.[230]Thisresearchaddressestheoriginandevolutionofembryonicdevelopmentandhow
modificationsofdevelopmentanddevelopmentalprocessesproducenovelfeatures.[231]Thesestudieshaveshown
thatevolutioncanalterdevelopmenttoproducenewstructures,suchasembryonicbonestructuresthatdevelop
intothejawinotheranimalsinsteadformingpartofthemiddleearinmammals.[232]Itisalsopossiblefor
structuresthathavebeenlostinevolutiontoreappearduetochangesindevelopmentalgenes,suchasamutation
inchickenscausingembryostogrowteethsimilartothoseofcrocodiles.[233]Itisnowbecomingclearthatmost
alterationsintheformoforganismsareduetochangesinasmallsetofconservedgenes.[234]
Coevolution
Interactionsbetweenorganismscanproducebothconflictandcooperation.Whentheinteractionisbetweenpairs
ofspecies,suchasapathogenandahost,orapredatoranditsprey,thesespeciescandevelopmatchedsetsof
adaptations.Here,theevolutionofonespeciescausesadaptationsinasecondspecies.Thesechangesinthesecond
speciesthen,inturn,causenewadaptationsinthefirstspecies.Thiscycleofselectionandresponseiscalled
coevolution.[235]Anexampleistheproductionoftetrodotoxinintheroughskinnednewtandtheevolutionof
tetrodotoxinresistanceinitspredator,thecommongartersnake.Inthispredatorpreypair,anevolutionaryarms
racehasproducedhighlevelsoftoxininthenewtandcorrespondinglyhighlevelsoftoxinresistanceinthe
snake.[236]
Cooperation
Notallcoevolvedinteractionsbetweenspeciesinvolveconflict.[237]Many
casesofmutuallybeneficialinteractionshaveevolved.Forinstance,an
extremecooperationexistsbetweenplantsandthemycorrhizalfungithat
growontheirrootsandaidtheplantinabsorbingnutrientsfromthe
soil.[238]Thisisareciprocalrelationshipastheplantsprovidethefungi
withsugarsfromphotosynthesis.Here,thefungiactuallygrowinsideplant
cells,allowingthemtoexchangenutrientswiththeirhosts,whilesending
signalsthatsuppresstheplantimmunesystem.[239]
Coalitionsbetweenorganismsofthesamespecieshavealsoevolved.An
extremecaseistheeusocialityfoundinsocialinsects,suchasbees, Commongartersnake(Thamnophis
termitesandants,wheresterileinsectsfeedandguardthesmallnumberof sirtalissirtalis)hasevolvedresistance
organismsinacolonythatareabletoreproduce.Onanevensmallerscale, tothedefensivesubstance
thesomaticcellsthatmakeupthebodyofananimallimittheir tetrodotoxininitsamphibianprey.
reproductionsotheycanmaintainastableorganism,whichthensupportsa
smallnumberoftheanimal'sgermcellstoproduceoffspring.Here,
somaticcellsrespondtospecificsignalsthatinstructthemwhethertogrow,remainastheyare,ordie.Ifcells
ignorethesesignalsandmultiplyinappropriately,theiruncontrolledgrowthcausescancer.[240]
Suchcooperationwithinspeciesmayhaveevolvedthroughtheprocessofkinselection,whichiswhereone
organismactstohelpraisearelative'soffspring.[241]Thisactivityisselectedforbecauseifthehelpingindividual
containsalleleswhichpromotethehelpingactivity,itislikelythatitskinwillalsocontaintheseallelesandthus
thosealleleswillbepassedon.[242]Otherprocessesthatmaypromotecooperationincludegroupselection,where
cooperationprovidesbenefitstoagroupoforganisms.[243]
Speciation
Speciationistheprocesswhereaspeciesdivergesinto
twoormoredescendantspecies.[244]
Therearemultiplewaystodefinetheconceptof
"species."Thechoiceofdefinitionisdependentonthe
particularitiesofthespeciesconcerned.[245]For
example,somespeciesconceptsapplymorereadily
towardsexuallyreproducingorganismswhileothers
lendthemselvesbettertowardasexualorganisms.
Despitethediversityofvariousspeciesconcepts,these
variousconceptscanbeplacedintooneofthreebroad
philosophicalapproaches:interbreeding,ecological
andphylogenetic.[246]TheBiologicalSpeciesConcept
(BSC)isaclassicexampleoftheinterbreeding
approach.DefinedbyErnstMayrin1942,theBSC
statesthat"speciesaregroupsofactuallyorpotentially
interbreedingnaturalpopulations,whichare
Thefourmechanismsofspeciation reproductivelyisolatedfromothersuchgroups."[247]
Despiteitswideandlongtermuse,theBSClikeothers
isnotwithoutcontroversy,forexamplebecausethese
conceptscannotbeappliedtoprokaryotes,[248]andthisiscalledthespeciesproblem.[245]Someresearchershave
attemptedaunifyingmonisticdefinitionofspecies,whileothersadoptapluralisticapproachandsuggestthatthere
maybedifferentwaystologicallyinterpretthedefinitionofaspecies.[245][246]
Barrierstoreproductionbetweentwodivergingsexualpopulationsarerequiredforthepopulationstobecomenew
species.Geneflowmayslowthisprocessbyspreadingthenewgeneticvariantsalsototheotherpopulations.
Dependingonhowfartwospecieshavedivergedsincetheirmostrecentcommonancestor,itmaystillbepossible
forthemtoproduceoffspring,aswithhorsesanddonkeysmatingtoproducemules.[249]Suchhybridsare
generallyinfertile.Inthiscase,closelyrelatedspeciesmayregularlyinterbreed,buthybridswillbeselected
againstandthespecieswillremaindistinct.However,viablehybridsareoccasionallyformedandthesenew
speciescaneitherhavepropertiesintermediatebetweentheirparentspecies,orpossessatotallynew
phenotype.[250]Theimportanceofhybridisationinproducingnewspeciesofanimalsisunclear,althoughcases
havebeenseeninmanytypesofanimals,[251]withthegraytreefrogbeingaparticularlywellstudiedexample.[252]
Speciationhasbeenobservedmultipletimesunderbothcontrolledlaboratoryconditionsandinnature.[253]In
sexuallyreproducingorganisms,speciationresultsfromreproductiveisolationfollowedbygenealogical
divergence.Therearefourmechanismsforspeciation.Themostcommoninanimalsisallopatricspeciation,which
occursinpopulationsinitiallyisolatedgeographically,suchasbyhabitatfragmentationormigration.Selection
undertheseconditionscanproduceveryrapidchangesintheappearanceandbehaviouroforganisms.[254][255]As
selectionanddriftactindependentlyonpopulationsisolatedfromtherestoftheirspecies,separationmay
eventuallyproduceorganismsthatcannotinterbreed.[256]
Thesecondmechanismofspeciationisperipatricspeciation,whichoccurswhensmallpopulationsoforganisms
becomeisolatedinanewenvironment.Thisdiffersfromallopatricspeciationinthattheisolatedpopulationsare
numericallymuchsmallerthantheparentalpopulation.Here,thefoundereffectcausesrapidspeciationafteran
increaseininbreedingincreasesselectiononhomozygotes,leadingtorapidgeneticchange.[257]
Thethirdmechanismofspeciationisparapatricspeciation.Thisissimilartoperipatricspeciationinthatasmall
populationentersanewhabitat,butdiffersinthatthereisnophysicalseparationbetweenthesetwopopulations.
Instead,speciationresultsfromtheevolutionofmechanismsthatreducegeneflowbetweenthetwo
populations.[244]Generallythisoccurswhentherehasbeenadrasticchangeintheenvironmentwithintheparental
species'habitat.OneexampleisthegrassAnthoxanthumodoratum,whichcanundergoparapatricspeciationin
responsetolocalisedmetalpollutionfrommines.[258]Here,plantsevolvethathaveresistancetohighlevelsof
metalsinthesoil.Selectionagainstinterbreedingwiththemetalsensitiveparentalpopulationproducedagradual
changeinthefloweringtimeofthemetalresistantplants,whicheventuallyproducedcompletereproductive
isolation.Selectionagainsthybridsbetweenthetwopopulationsmaycausereinforcement,whichistheevolution
oftraitsthatpromotematingwithinaspecies,aswellascharacterdisplacement,whichiswhentwospecies
becomemoredistinctinappearance.[259]
Finally,insympatricspeciationspeciesdivergewithoutgeographicisolationorchangesinhabitat.Thisformis
raresinceevenasmallamountofgeneflowmayremovegeneticdifferencesbetweenpartsofapopulation.[260]
Generally,sympatricspeciationinanimalsrequirestheevolutionofbothgeneticdifferencesandnonrandom
mating,toallowreproductiveisolationtoevolve.[261]
Onetypeofsympatricspeciationinvolvescrossbreedingoftworelatedspeciestoproduceanewhybridspecies.
Thisisnotcommoninanimalsasanimalhybridsareusuallysterile.Thisisbecauseduringmeiosisthe
homologouschromosomesfromeachparentarefromdifferentspeciesandcannotsuccessfullypair.However,itis
morecommoninplantsbecauseplantsoftendoubletheirnumberofchromosomes,toformpolyploids.[262]This
allowsthechromosomesfromeachparentalspeciestoformmatchingpairsduringmeiosis,sinceeachparent's
chromosomesarerepresentedbyapairalready.[263]Anexampleof
suchaspeciationeventiswhentheplantspeciesArabidopsisthaliana
andArabidopsisarenosacrossbredtogivethenewspecies
Arabidopsissuecica.[264]Thishappenedabout20,000yearsago,[265]
andthespeciationprocesshasbeenrepeatedinthelaboratory,which
allowsthestudyofthegeneticmechanismsinvolvedinthis
process.[266]Indeed,chromosomedoublingwithinaspeciesmaybea
commoncauseofreproductiveisolation,ashalfthedoubled
chromosomeswillbeunmatchedwhenbreedingwithundoubled
organisms.[267]
Speciationeventsareimportantinthetheoryofpunctuated
equilibrium,whichaccountsforthepatterninthefossilrecordofshort
"bursts"ofevolutioninterspersedwithrelativelylongperiodsofstasis, Geographicalisolationoffinchesonthe
GalpagosIslandsproducedoveradozen
wherespeciesremainrelativelyunchanged.[268]Inthistheory,
newspecies.
speciationandrapidevolutionarelinked,withnaturalselectionand
geneticdriftactingmoststronglyonorganismsundergoingspeciation
innovelhabitatsorsmallpopulations.Asaresult,theperiodsofstasisinthefossilrecordcorrespondtothe
parentalpopulationandtheorganismsundergoingspeciationandrapidevolutionarefoundinsmallpopulationsor
geographicallyrestrictedhabitatsandthereforerarelybeingpreservedasfossils.[181]
Extinction
Extinctionisthedisappearanceofanentirespecies.Extinctionisnotan
unusualevent,asspeciesregularlyappearthroughspeciationanddisappear
throughextinction.[269]Nearlyallanimalandplantspeciesthathavelived
onEartharenowextinct,[270]andextinctionappearstobetheultimatefate
ofallspecies.[271]Theseextinctionshavehappenedcontinuously
throughoutthehistoryoflife,althoughtherateofextinctionspikesin
occasionalmassextinctionevents.[272]TheCretaceousPaleogene
extinctionevent,duringwhichthenonaviandinosaursbecameextinct,is
Tyrannosaurusrex.Nonavian
themostwellknown,buttheearlierPermianTriassicextinctioneventwas
dinosaursdiedoutintheCretaceous
evenmoresevere,withapproximately96%ofallmarinespeciesdrivento
Paleogeneextinctioneventattheend
oftheCretaceousperiod. extinction.[272]TheHoloceneextinctioneventisanongoingmass
extinctionassociatedwithhumanity'sexpansionacrosstheglobeoverthe
pastfewthousandyears.Presentdayextinctionratesare1001000times
greaterthanthebackgroundrateandupto30%ofcurrentspeciesmaybeextinctbythemid21stcentury.[273]
Humanactivitiesarenowtheprimarycauseoftheongoingextinctionevent[274]globalwarmingmayfurther
accelerateitinthefuture.[275]
Theroleofextinctioninevolutionisnotverywellunderstoodandmaydependonwhichtypeofextinctionis
considered.[272]Thecausesofthecontinuous"lowlevel"extinctionevents,whichformthemajorityofextinctions,
maybetheresultofcompetitionbetweenspeciesforlimitedresources(thecompetitiveexclusionprinciple).[72]If
onespeciescanoutcompeteanother,thiscouldproducespeciesselection,withthefitterspeciessurvivingandthe
otherspeciesbeingdriventoextinction.[139]Theintermittentmassextinctionsarealsoimportant,butinsteadof
actingasaselectiveforce,theydrasticallyreducediversityinanonspecificmannerandpromoteburstsofrapid
evolutionandspeciationinsurvivors.[276]
Evolutionaryhistoryoflife
Originoflife
Lifetimeline
TheEarthisabout4.54billionyearsold.[277][278][279] viewdiscuss
TheearliestundisputedevidenceoflifeonEarthdates 0 Earliesthumans
Quaternary P Flowers
fromatleast3.5billionyearsago,[7][280]duringthe h Mammals
a Dinosaurs
EoarcheanEraafterageologicalcruststartedto Karoo
n Landlife
Andean e
solidifyfollowingtheearliermoltenHadeanEon. 500 r Cambrianexplosion
Microbialmatfossilshavebeenfoundin3.48billion Cryogenian
o
z
Ediacarabiota

yearoldsandstoneinWesternAustralia.[13][14][15]
o
i Multicellular
Otherearlyphysicalevidenceofabiogenicsubstance 1000
c
life

isgraphitein3.7billionyearoldmetasedimentary
Earliestsexual
rocksdiscoveredinWesternGreenland[12]aswellas reproduction
P
"remainsofbioticlife"foundin4.1billionyearold r
1500 o
rocksinWesternAustralia.[8][9]Accordingtooneof t
e
theresearchers,"Iflifearoserelativelyquicklyon r Eukaryotes
o
Earththenitcouldbecommonintheuniverse."[8] 2000 z
o
i
Morethan99percentofallspecies,amountingtoover c
Huronian
Oxygencrisis
fivebillionspecies,[281]thateverlivedonEarthare
2500 Atmosphericoxygen
estimatedtobeextinct.[17][18]Estimatesonthe
numberofEarth'scurrentspeciesrangefrom10

millionto14million,[19][20]ofwhichabout1.9
Pongola
photosynthesis
3000 A
millionareestimatedtohavebeennamed[21]and1.6 r
c
milliondocumentedinacentraldatabasetodate,[22] h
e
leavingatleast80percentnotyetdescribed. a
3500 n Earliestoxygen
Highlyenergeticchemistryisthoughttohave
producedaselfreplicatingmoleculearound4billion Singlecelled
life LHBmeteorites
yearsago,andhalfabillionyearslaterthelast 4000
H
commonancestorofalllifeexisted.[5]Thecurrent a
d Earliestlife
scientificconsensusisthatthecomplexbiochemistry e water
a Earliestwater
thatmakesuplifecamefromsimplerchemical 4500 n EarliestEarth
reactions.[282]Thebeginningoflifemayhave Axisscale:millionsofyears.
(4540)
Orangelabels:knowniceages.
includedselfreplicatingmoleculessuchasRNA[283]
Alsosee:HumantimelineandNaturetimeline
andtheassemblyofsimplecells.[284]
Commondescent
AllorganismsonEartharedescendedfromacommonancestororancestralgenepool.[211][285]Currentspeciesare
astageintheprocessofevolution,withtheirdiversitytheproductofalongseriesofspeciationandextinction
events.[286]Thecommondescentoforganismswasfirstdeducedfromfoursimplefactsaboutorganisms:First,
theyhavegeographicdistributionsthatcannotbeexplainedbylocaladaptation.Second,thediversityoflifeisnot
asetofcompletelyuniqueorganisms,butorganismsthatsharemorphologicalsimilarities.Third,vestigialtraits
withnoclearpurposeresemblefunctionalancestraltraitsandfinally,thatorganismscanbeclassifiedusingthese
similaritiesintoahierarchyofnestedgroupssimilartoafamilytree.[287]However,modernresearchhas
suggestedthat,duetohorizontalgenetransfer,this"treeoflife"maybemorecomplicatedthanasimplebranching
treesincesomegeneshavespreadindependentlybetweendistantlyrelatedspecies.[288][289]
Pastspecieshavealsoleftrecordsoftheirevolutionary
history.Fossils,alongwiththecomparativeanatomyof
presentdayorganisms,constitutethemorphological,or
anatomical,record.[290]Bycomparingtheanatomiesof
bothmodernandextinctspecies,paleontologistscaninfer
thelineagesofthosespecies.However,thisapproachis
mostsuccessfulfororganismsthathadhardbodyparts,
suchasshells,bonesorteeth.Further,asprokaryotessuch
asbacteriaandarchaeasharealimitedsetofcommon
morphologies,theirfossilsdonotprovideinformationon
Thehominoidsaredescendantsofacommonancestor. theirancestry.
Morerecently,evidenceforcommondescenthascome
fromthestudyofbiochemicalsimilaritiesbetweenorganisms.Forexample,alllivingcellsusethesamebasicset
ofnucleotidesandaminoacids.[291]Thedevelopmentofmoleculargeneticshasrevealedtherecordofevolution
leftinorganisms'genomes:datingwhenspeciesdivergedthroughthemolecularclockproducedbymutations.[292]
Forexample,theseDNAsequencecomparisonshaverevealedthathumansandchimpanzeesshare98%oftheir
genomesandanalysingthefewareaswheretheydifferhelpsshedlightonwhenthecommonancestorofthese
speciesexisted.[293]
Evolutionoflife
ProkaryotesinhabitedtheEarthfromapproximately34
billionyearsago.[295][296]Noobviouschangesin
morphologyorcellularorganisationoccurredinthese
organismsoverthenextfewbillionyears.[297]The
eukaryoticcellsemergedbetween1.62.7billionyearsago.
Thenextmajorchangeincellstructurecamewhenbacteria
wereengulfedbyeukaryoticcells,inacooperative
associationcalledendosymbiosis.[298][299]Theengulfed
bacteriaandthehostcellthenunderwentcoevolution,with
thebacteriaevolvingintoeithermitochondriaor
hydrogenosomes.[300]Anotherengulfmentof
cyanobacteriallikeorganismsledtotheformationof Evolutionarytreeshowingthedivergenceofmodern
chloroplastsinalgaeandplants.[301] speciesfromtheircommonancestorinthecentre. [294]
Thethreedomainsarecoloured,withbacteriablue,
Thehistoryoflifewasthatoftheunicellulareukaryotes, archaeagreenandeukaryotesred.
prokaryotesandarchaeauntilabout610millionyearsago
whenmulticellularorganismsbegantoappearintheoceans
intheEdiacaranperiod.[295][302]Theevolutionofmulticellularityoccurredinmultipleindependentevents,in
organismsasdiverseassponges,brownalgae,cyanobacteria,slimemouldsandmyxobacteria.[303]InJanuary
2016,scientistsreportedthat,about800millionyearsago,aminorgeneticchangeinasinglemoleculecalledGK
PIDmayhaveallowedorganismstogofromasinglecellorganismtooneofmanycells.[304]
Soonaftertheemergenceofthesefirstmulticellularorganisms,aremarkableamountofbiologicaldiversity
appearedoverapproximately10millionyears,inaneventcalledtheCambrianexplosion.Here,themajorityof
typesofmodernanimalsappearedinthefossilrecord,aswellasuniquelineagesthatsubsequentlybecame
extinct.[305]VarioustriggersfortheCambrianexplosionhavebeenproposed,includingtheaccumulationof
oxygenintheatmospherefromphotosynthesis.[306]
About500millionyearsago,plantsandfungicolonisedthelandandweresoonfollowedbyarthropodsandother
animals.[307]Insectswereparticularlysuccessfulandeventodaymakeupthemajorityofanimalspecies.[308]
Amphibiansfirstappearedaround364millionyearsago,followedbyearlyamniotesandbirdsaround155million
yearsago(bothfrom"reptile"likelineages),mammalsaround129millionyearsago,homininaearound10million
yearsagoandmodernhumansaround250,000yearsago.[309][310][311]However,despitetheevolutionoftheselarge
animals,smallerorganismssimilartothetypesthatevolvedearlyinthisprocesscontinuetobehighlysuccessful
anddominatetheEarth,withthemajorityofbothbiomassandspeciesbeingprokaryotes.[188]
Applications
Conceptsandmodelsusedinevolutionarybiology,suchasnaturalselection,havemanyapplications.[312]
Artificialselectionistheintentionalselectionoftraitsinapopulationoforganisms.Thishasbeenusedfor
thousandsofyearsinthedomesticationofplantsandanimals.[313]Morerecently,suchselectionhasbecomeavital
partofgeneticengineering,withselectablemarkerssuchasantibioticresistancegenesbeingusedtomanipulate
DNA.Proteinswithvaluablepropertieshaveevolvedbyrepeatedroundsofmutationandselection(forexample
modifiedenzymesandnewantibodies)inaprocesscalleddirectedevolution.[314]
Understandingthechangesthathaveoccurredduringanorganism'sevolutioncanrevealthegenesneededto
constructpartsofthebody,geneswhichmaybeinvolvedinhumangeneticdisorders.[315]Forexample,the
Mexicantetraisanalbinocavefishthatlostitseyesightduringevolution.Breedingtogetherdifferentpopulations
ofthisblindfishproducedsomeoffspringwithfunctionaleyes,sincedifferentmutationshadoccurredinthe
isolatedpopulationsthathadevolvedindifferentcaves.[316]Thishelpedidentifygenesrequiredforvisionand
pigmentation.[317]
Manyhumandiseasesarenotstaticphenomena,butcapableofevolution.Viruses,bacteria,fungiandcancers
evolvetoberesistanttohostimmunedefences,aswellaspharmaceuticaldrugs.[318][319][320]Thesesameproblems
occurinagriculturewithpesticide[321]andherbicide[322]resistance.Itispossiblethatwearefacingtheendofthe
effectivelifeofmostofavailableantibiotics[323]andpredictingtheevolutionandevolvability[324]ofourpathogens
anddevisingstrategiestosloworcircumventitisrequiringdeeperknowledgeofthecomplexforcesdriving
evolutionatthemolecularlevel.[325]
Incomputerscience,simulationsofevolutionusingevolutionaryalgorithmsandartificiallifestartedinthe1960s
andwereextendedwithsimulationofartificialselection.[326]Artificialevolutionbecameawidelyrecognised
optimisationmethodasaresultoftheworkofIngoRechenberginthe1960s.Heusedevolutionstrategiestosolve
complexengineeringproblems.[327]GeneticalgorithmsinparticularbecamepopularthroughthewritingofJohn
HenryHolland.[328]Practicalapplicationsalsoincludeautomaticevolutionofcomputerprogrammes.[329]
Evolutionaryalgorithmsarenowusedtosolvemultidimensionalproblemsmoreefficientlythansoftware
producedbyhumandesignersandalsotooptimisethedesignofsystems.[330]
Socialandculturalresponses
Inthe19thcentury,particularlyafterthepublicationofOntheOriginofSpeciesin
1859,theideathatlifehadevolvedwasanactivesourceofacademicdebate
centredonthephilosophical,socialandreligiousimplicationsofevolution.Today,
themodernevolutionarysynthesisisacceptedbyavastmajorityofscientists.[72]
However,evolutionremainsacontentiousconceptforsometheists.[332]
Whilevariousreligionsanddenominationshavereconciledtheirbeliefswith
evolutionthroughconceptssuchastheisticevolution,therearecreationistswho
believethatevolutioniscontradictedbythecreationmythsfoundintheirreligions
andwhoraisevariousobjectionstoevolution.[177][333][334]Ashadbeen
demonstratedbyresponsestothepublicationofVestigesoftheNaturalHistoryof
Creationin1844,themostcontroversialaspectofevolutionarybiologyisthe
implicationofhumanevolutionthathumanssharecommonancestrywithapesand
thatthementalandmoralfacultiesofhumanityhavethesametypesofnatural Asevolutionbecamewidely
causesasotherinheritedtraitsinanimals.[335]Insomecountries,notablythe acceptedinthe1870s,
UnitedStates,thesetensionsbetweenscienceandreligionhavefuelledthecurrent caricaturesofCharles
creationevolutioncontroversy,areligiousconflictfocusingonpoliticsandpublic Darwinwithanapeor
education.[336]Whileotherscientificfieldssuchascosmology[337]andEarth monkeybodysymbolised
science[338]alsoconflictwithliteralinterpretationsofmanyreligioustexts, evolution. [331]
evolutionarybiologyexperiencessignificantlymoreoppositionfromreligious
literalists.
TheteachingofevolutioninAmericansecondaryschoolbiologyclasseswasuncommoninmostofthefirsthalfof
the20thcentury.TheScopesTrialdecisionof1925causedthesubjecttobecomeveryrareinAmericansecondary
biologytextbooksforageneration,butitwasgraduallyreintroducedlaterandbecamelegallyprotectedwiththe
1968Eppersonv.Arkansasdecision.Sincethen,thecompetingreligiousbeliefofcreationismwaslegally
disallowedinsecondaryschoolcurriculainvariousdecisionsinthe1970sand1980s,butitreturnedin
pseudoscientificformasintelligentdesign(ID),tobeexcludedonceagaininthe2005Kitzmillerv.DoverArea
SchoolDistrictcase.[339]
Seealso
Argumentfrompoordesign Evolutionaryneuroscience
Bioculturalevolution Evolutionofbiologicalcomplexity
Biologicalclassification Evolutionofplants
Evidenceofcommondescent Timelineoftheevolutionaryhistoryoflife
Evolutionaryanthropology Unintelligentdesign
Evolutionaryecology UniversalDarwinism
Evolutionaryepistemology
Evolutionaryneuroscience
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talog.php?record_id=11876).Washington,D.C.:NationalAcademyPress.ISBN9780309105866.LCCN2007015904
(https://lccn.loc.gov/2007015904).OCLC123539346(https://www.worldcat.org/oclc/123539346).Retrieved20141122.
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andstressproteinsaslenscrystallins".InJansson,BengtJrnvall,HansRydberg,Ulfetal.TowardaMolecularBasis
ofAlcoholUseandAbuse.Experientia.71.BaselBoston:BirkhuserVerlag.ISBN3764329408.ISSN1023294X(h
ttps://www.worldcat.org/issn/1023294X).LCCN94010167(https://lccn.loc.gov/94010167).OCLC30030941(https://w
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020835).OCLC18380658(https://www.worldcat.org/oclc/18380658)."ThisbookistheresultoftheSpringSystematics
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77).
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Ray,John(1686).HistoriaPlantarum[HistoryofPlants].I.Londini:TypisMariClark.LCCNagr11000774(https://l
ccn.loc.gov/agr11000774).OCLC2126030(https://www.worldcat.org/oclc/2126030).
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Evolution(PhDthesis).Problemata(inGerman).15.AfterwordbyManfredEigen.StuttgartBadCannstatt:Frommann
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WestEberhard,MaryJane(2003).DevelopmentalPlasticityandEvolution.OxfordNewYork:OxfordUniversityPress.
ISBN0195122356.LCCN2001055164(https://lccn.loc.gov/2001055164).OCLC48398911(https://www.worldcat.or
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OCLC741259265(https://www.worldcat.org/oclc/741259265).doi:10.1002/9781118017883(https://doi.org/10.1002%2F
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Illinois:UniversityofChicagoPress.ISBN0226910385.LCCN67025533(https://lccn.loc.gov/67025533).
Furtherreading
Introductoryreading
Barrett,PaulH.Weinshank,DonaldJ.Gottleber,TimothyT.,eds.(1981).AConcordancetoDarwin'sOriginof
Species,FirstEdition.Ithaca,NY:CornellUniversityPress.ISBN0801413192.LCCN80066893(https://lccn.loc.go
v/80066893).OCLC610057960(https://www.worldcat.org/oclc/610057960).
Carroll,SeanB.(2005).EndlessFormsMostBeautiful:TheNewScienceofEvoDevoandtheMakingoftheAnimal
Kingdom.illustrationsbyJamieW.Carroll,JoshP.Klaiss,LeanneM.Olds(1sted.).NewYork:W.W.Norton&
Company.ISBN0393060160.LCCN2004029388(https://lccn.loc.gov/2004029388).OCLC57316841(https://www.
worldcat.org/oclc/57316841).
Charlesworth,BrianCharlesworth,Deborah(2003).Evolution:AVeryShortIntroduction.VeryShortIntroductions.
OxfordNewYork:OxfordUniversityPress.ISBN0192802518.LCCN2003272247(https://lccn.loc.gov/200327224
Gould,StephenJay(1989).WonderfulLife:TheBurgessShaleandtheNatureofHistory(1sted.).NewYork:W.W.
Norton&Company.ISBN0393027058.LCCN88037469(https://lccn.loc.gov/88037469).OCLC18983518(https://
www.worldcat.org/oclc/18983518).
Jones,Steve(1999).AlmostLikeaWhale:TheOriginofSpeciesUpdated.LondonNewYork:Doubleday.ISBN0385
44).
(2000).Darwin'sGhost:TheOriginofSpeciesUpdated(1sted.).NewYork:RandomHouse.ISBN0375
0131).Americanversion.
Mader,SylviaS.(2007).Biology.SignificantcontributionsbyMurrayP.Pendarvis(9thed.).Boston,MA:McGrawHill
HigherEducation.ISBN9780072464634.LCCN2005027781(https://lccn.loc.gov/2005027781).OCLC61748307(h
ttps://www.worldcat.org/oclc/61748307).
MaynardSmith,John(1993).TheTheoryofEvolution(Cantoed.).CambridgeNewYork:CambridgeUniversityPress.
ISBN0521451280.LCCN93020358(https://lccn.loc.gov/93020358).OCLC27676642(https://www.worldcat.org/ocl
c/27676642).
Pallen,MarkJ.(2009).TheRoughGuidetoEvolution.RoughGuidesReferenceGuides.LondonNewYork:Rough
Guides.ISBN9781858289465.LCCN2009288090(https://lccn.loc.gov/2009288090).OCLC233547316(https://ww
w.worldcat.org/oclc/233547316).
Advancedreading
Barton,NicholasH.Briggs,DerekE.G.Eisen,JonathanA.etal.(2007).Evolution.ColdSpringHarbor,NY:Cold
SpringHarborLaboratoryPress.ISBN9780879696849.LCCN2007010767(https://lccn.loc.gov/2007010767).
Coyne,JerryA.Orr,H.Allen(2004).Speciation.Sunderland,MA:SinauerAssociates.ISBN0878930892.
Bergstrom,CarlT.Dugatkin,LeeAlan(2012).Evolution(1sted.).NewYork:W.W.Norton&Company.ISBN9780
29341924).
Gould,StephenJay(2002).TheStructureofEvolutionaryTheory.Cambridge,MA:BelknapPressofHarvardUniversity
Press.ISBN0674006135.LCCN2001043556(https://lccn.loc.gov/2001043556).OCLC47869352(https://www.worl
dcat.org/oclc/47869352).
Hall,BrianK.Olson,Wendy,eds.(2003).KeywordsandConceptsinEvolutionaryDevelopmentalBiology.Cambridge,
MA:HarvardUniversityPress.ISBN0674009045.LCCN2002192201(https://lccn.loc.gov/2002192201).
Kauffman,StuartA.(1993).TheOriginsofOrder:SelforganizationandSelectioninEvolution(https://books.google.co
m/books?id=lZcSpRJz0dgC&printsec=frontcover&dq=evolution+biological+selforganization&hl=en&sa=X&ved=0ahU
KEwiY7enKqOLQAhUG6IMKHaoAAkQ6AEIIzAA#v=onepage&q&f=false).NewYork,NY:OxfordUniversityPress.
MaynardSmith,JohnSzathmry,Ers(1995).TheMajorTransitionsinEvolution.OxfordNewYork:W.H.Freeman
Spektrum.ISBN0716745259.LCCN94026965(https://lccn.loc.gov/94026965).OCLC30894392(https://www.world
cat.org/oclc/30894392).
Mayr,Ernst(2001).WhatEvolutionIs.NewYork:BasicBooks.ISBN0465044263.LCCN2001036562(https://lccn.
loc.gov/2001036562).OCLC47443814(https://www.worldcat.org/oclc/47443814).
Minelli,Alessandro(2009).FormsofBecoming:TheEvolutionaryBiologyofDevelopment.TranslationbyMark
Epstein.Princeton,NJOxford:PrincetonUniversityPress.ISBN9780691135687.LCCN2008028825(https://lccn.l
oc.gov/2008028825).OCLC233030259(https://www.worldcat.org/oclc/233030259).
Externallinks
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Evolution(http://www.bbc.co.uk/programmes/p00545gl)onInOurTimeattheBBC.
"Evolution"(http://www.newscientist.com/topic/evolution).NewScientist.Retrieved20110530.
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D.C.:NationalAcademyofSciences.Retrieved20110530.
"UnderstandingEvolution:youronestopresourceforinformationonevolution"(http://evolution.berkeley.e
du/).UniversityofCalifornia,Berkeley.Retrieved20110530.
"EvolutionofEvolution150YearsofDarwin's'OntheOriginofSpecies' "(http://www.nsf.gov/news/speci
al_reports/darwin/textonly/index.jsp).ArlingtonCounty,VA:NationalScienceFoundation.Retrieved
20110530.
HumanTimeline(Interactive)(http://humanorigins.si.edu/evidence/humanevolutiontimelineinteractive)
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