Chapter23

TheEvolutionofPopulations
LectureOutline
Overview:TheSmallestUnitofEvolution
Onecommonmisconceptionaboutevolutionisthatorganismsevolve,inaDarwinian
sense,duringtheirlifetimes.
o Naturalselectiondoesactonindividuals.Eachindividualscombinationoftraitsaffects
itssurvivalanditsreproductivesuccessrelativetootherindividualsinthepopulation.
o Theevolutionaryimpactofnaturalselectionisapparentonlyinthechangesina
populationoforganismsovertime.

Itisthepopulation,nottheindividual,thatevolves.
Considertheexampleofthemediumgroundfinch(Geospizafortis),aseedeatingbirdthat
livesontheGalpagosIslands.
o In1977,theG.spizapopulationenduredalongperiodofdrought.Of1200birds,only
180survived.
o Thesurvivingfincheshadlarger,deeperbeaksthanthefinchesthatdied.
Soft,smallseedswereinshortsupplyduringthedrought.
Large,hardseedsweremoreabundant.
Fincheswithlarge,deepbeakscouldcrackthelargeseedsandthuswereableto
survivethefoodshortageduringthedrought.
o Followingthedrought,theaveragebeaksizeinthepopulationwaslargerthanbeforethe
drought.Thefinchpopulationhadevolvedlargerbeaksbynaturalselection.
Individualfinchesdidnotevolve.Eachbirdhadabeakofaparticularsize,whichdid
notgrowlargerduringthedrought.
Theproportionofbirdswithlargebeaksinthepopulationincreasedbecausebirds
withlargebeakswerebetterabletosurvivethedroughtandreproducesuccessfully.

Microevolutionisdefinedasachangeinallelefrequenciesinapopulationovertime.

Threemechanismscancauseallelefrequenciestochange:naturalselection,geneticdrift
(chanceeventsthatalterallelefrequencies),andgeneflow(thetransferofallelesbetween
populations).
Naturalselectionistheonlymechanismofadaptiveevolution,improvingthematchbetween
organismsandtheirenvironment.

Concept23.1Mutationandsexualreproductionproducethegeneticvariationthat
makesevolutionpossible.

CharlesDarwinproposedamechanismforchangeinspeciesovertime.

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WhatwasmissingfromDarwinsexplanationwasanunderstandingofinheritancethatcould
explainhowchancevariationsariseinapopulationwhilealsoaccountingfortheprecise
transmissionofthesevariationsfromparentstooffspring.
JustafewyearsafterDarwinpublishedTheOriginofSpecies,GregorMendelproposeda
modelofinheritancethatsupportedDarwinstheory.
o Mendelsparticulatehypothesisofinheritancestatedthatparentspassondiscrete
heritableunits(genes)thatretaintheiridentitiesinoffspring.
o AlthoughMendelandDarwinwerecontemporaries,DarwinneversawMendelspaper.
o Mendelsideassetthestageforanunderstandingofthegeneticdifferencesonwhich
evolutionisbased.
Twoprocessesproducethegeneticdifferencesthatarethebasisofevolution:mutationand
sexualreproduction.

Individualvariationoccursinallspecies,butnotallphenotypicvariationisheritable.

Phenotypeistheproductofaninheritedgenotypeandenvironmentalinfluences.

Onlythegeneticcomponentofvariationhasevolutionaryconsequences.

Geneticvariationoccurswithinapopulation.

Bothquantitativeanddiscretecharacterscontributetovariationwithinapopulation.

Discretecharacters,suchasflowercolor,areusuallydeterminedbyasinglelocuswith
differentallelesthatproducedistinctphenotypes.

o
o

Quantitativecharactersvaryalongacontinuumwithinapopulation.
Forexample,plantheightinawildflowerpopulationrangesfromshorttotall.
Quantitativevariationisusuallyduetopolygenicinheritanceinwhichtheadditiveeffects
oftwoormoregenesinfluenceasinglephenotypiccharacter.

Biologistscanmeasuregeneticvariationinapopulationatthewholegenelevel(gene
variability)andatthemolecularlevelofDNA(nucleotidevariability).
Averageheterozygositymeasuresgenevariability,theaveragepercentofgenelocithat
areheterozygous.
o Inthefruitfly(Drosophila),about86%oftheir13,700genelociarehomozygous
(fixed).
o About14%(1,920genes)areheterozygous,foranaverageheterozygosityof14%.
o Averageheterozygositycanbeestimatedusingproteingelelectrophoresis,which
measuresdifferencesintheproteinproductsofgenes.Thisapproachdoesnotmeasure
silentmutationsthatdonotaltertheaminoacidsequenceofaprotein.
o PCRbasedapproachesorrestrictionfragmentanalysesdodetectsilentmutations.
Nucleotidevariabilitymeasuresthemeanlevelofdifferenceinnucleotidesequences
(basepairdifferences)amongindividualsinapopulation.
o Infruitflies,about1%ofthebasesdifferbetweentwoindividuals.
o Twoindividualsdiffer,onaverage,at1.8millionofthe180millionnucleotidesinthe
fruitflygenome.
Averageheterozygositytendstobegreaterthannucleotidediversitybecauseagenecan
consistofthousandsofbasesofDNA.Adifferenceatonlyoneofthesebasesissufficientto
maketwoallelesofthatgenedifferentandcounttowardaverageheterozygosity.
Geneticvariationoccursbetweenpopulations.
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 Geographicvariationresultsfromdifferencesinphenotypesorgenotypesbetween
populationsorbetweensubgroupsofasinglepopulationthatinhabitdifferentareas.
o Naturalselectioncontributestogeographicvariationbymodifyinggenefrequenciesin
responsetodifferencesinlocalenvironmentalfactors.
o Geneticdriftcanalsoleadtovariationamongpopulationsthroughthecumulativeeffect
ofrandomfluctuationsinallelefrequencies.
Geographicvariationintheformofgradedchangeinatraitalongageographicaxisis
calledacline.
o Clinesmayreflecttheinfluenceofnaturalselectionbasedongradationinsome
environmentalvariable.
Newgenesandnewallelesoriginateonlybymutation.

AmutationisachangeinthenucleotidesequenceofanorganismsDNA.

Onlymutationsincelllinesthatformgametescanbepassedontooffspring.

Infungiandplants,manydifferentcelllinescanproducegametes.

Inanimals,mostmutationsoccurinsomaticcellsandarelostwhentheindividualdies.

Apointmutationisachangeofasinglebaseinagene.

Pointmutationscanhaveasignificantimpactonphenotype,asinthecaseofsicklecell
disease.

o
o
o

Mostpointmutationsareharmless.
MuchoftheDNAineukaryoticgenomesdoesnotcodeforproteinproducts.
Becausethegeneticcodeisredundant,somepointmutationsingenesthatcodefor
proteinsmaynotaltertheproteinsaminoacidcomposition.
Evenifthereisachangeinanaminoacidasaresultofapointmutation,itmaynotaffect
theproteinsshapeandfunction.

Onrareoccasions,amutantallelemayactuallymakeitsbearerbettersuitedtothe
environment,increasingitsreproductivesuccess.
o Thisismorelikelywhentheenvironmentischanging.

o
o

Somemutationsalterthegenenumberorsequence.
Chromosomalmutationsthatdeleteorrearrangemanygenelociatoncearealmost
alwaysharmful.
Inrarecases,chromosomalrearrangementsmaybebeneficial.
Forexample,thetranslocationofpartofonechromosometoadifferentchromosome
couldlinkgenesthatacttogetherforapositiveeffect.
Geneduplicationisanimportantsourceofnewgeneticvariation.

SmallpiecesofDNAcanbeintroducedintothegenomethroughtheactivityof
transposons.
Duplicatedsegmentscanpersistovergenerationsandprovidenewlocithatmayeventually
takeonnewfunctionsbymutationandsubsequentselection.

Beneficialincreasesingenenumberappeartohaveplayedamajorroleinevolution.

Forexample,mammalianancestorscarriedasinglegenefordetectingodorsthathasbeen
duplicatedmanytimes.
Modernhumanshaveabout1,000olfactoryreceptorgenesandmicehave1,300.

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Dramaticincreasesinthenumberofolfactorygenesbenefitedearlymammals,enabling
themtodetectfaintodorsanddistinguishamongsmells.
o Becauseofmutations,60%ofthesegeneshavebeeninactivatedinhumans.
o Mice,whichrelymoreontheirsenseofsmell,havelostonly20%oftheirolfactory
receptorgenes.
Mutationratesvaryfromorganismtoorganism.
o

Ratesofmutationsthataffectphenotypeaverageabout105mutationspergenepergamete
ineachgeneration(inotherwords,aboutonemutationforevery100,000genes)inplantsand
animals.
Atthenucleotidesequencelevel,mutationratesrangefrom1011mutationsperbasepairper
generationinprokaryotestoasmanyas104perbasepairpergenerationinviruses.
Inmicroorganismsandviruseswithshortgenerationspans,mutationratesaremuch
higherandcanrapidlygenerategeneticvariation.
Forexample,HIVhasagenerationtimeoftwodays.ItalsohasanRNAgenome,which
hasahighermutationratethanaDNAgenomebecauseofthelackofRNArepair
mechanismsinhostcells.

Asaresult,itisunlikelythatasingledrugtreatmentwilleverbeeffectiveagainstHIV.
Mutantformsofthevirusthatareresistanttothedrugwillariseandproliferate.

Themosteffectivetreatmentsaredrugcocktailsbecauseitisunlikelythatmultiple
mutationswillconferresistancetoallofthedrugsinthecocktail.

Sexualreproductionproducesuniquecombinationsofalleles.
Insexuallyreproducingpopulations,mostofthegeneticvariationresultsfromtheunique
combinationsofallelesthateachindividualreceives.
Variantallelesoriginatedfrompastmutations.However,sexualreproductionshuffles
variantallelesanddealsthematrandomtoproduceuniqueindividualgenotypes.
Threemechanismscontributetotheshuffling:crossingover,independentassortmentof
chromosomes,andfertilization.
Thecombinedeffectsofthesethreemechanismsensurethatsexualreproduction
rearrangesexistingallelesintonewcombinationseachgeneration,providingthegenetic
variationthatmakesevolutionpossible.

Concept23.2TheHardyWeinbergequationcanbeusedtotestwhethera
populationisevolving.
Forapopulationtoevolve,individualsmustdiffergeneticallyandoneofthefactorsthat
causesevolutionmustbeatwork.
Apopulationsgenepoolisdefinedbyitsallelefrequencies.
Apopulationisagroupofindividualsthatbelongtothesamespecies,liveinthesame
area,andinterbreedtoproducefertileoffspring.
Onedefinitionofaspeciesisagroupofnaturalpopulationswhoseindividualshavethe
potentialtointerbreedandproducefertileoffspring.
Populationsofaspeciesmaybeisolatedfromeachotherandrarelyexchangegenetic
material.
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 Membersofapopulationarefarmorelikelytobreedwithmembersofthesame
populationthanwithmembersofotherpopulations.
o Individualsnearthepopulationscenterare,onaverage,morecloselyrelatedtoone
anotherthantomembersofotherpopulations.
Thetotalaggregateofalltheallelesforallofthelociforalloftheindividualsina
populationiscalledthepopulationsgenepool.
o Ifonlyonealleleexistsataparticularlocusinapopulation,thatalleleissaidtobefixed
inthegenepool,andallindividualswillbehomozygousforthatgene.
o Iftherearetwoormoreallelesataparticularlocus,thenindividualscanbeeither
homozygousorheterozygousforthatgene.

Eachallelehasafrequencyinthepopulationsgenepool.

Forexample,imagineapopulationof500wildflowerplantswithtwoalleles(C RandCW)
atalocusthatcodesforflowerpigment.
o Supposethatintheimaginarypopulationof500plants,20(4%)arehomozygousforthe
CWallele(CWCW)andhavewhiteflowers.
o Oftheremainingplants,320(64%)arehomozygousfortheC Rallele(CRCR)andhavered
flowers.
o Theseallelesshowincompletedominance,so160(32%)oftheplantsareheterozygous
(CRCW)andproducepinkflowers.
Becausetheseplantsarediploid,thepopulationof500plantshas1,000copiesofthe
geneforflowercolor.
o Thedominantallele(CR)accountsfor800copies(3202forCRCR+1601forCRCW).
o ThefrequencyoftheCRalleleinthegenepoolofthispopulationis800/1,000=0.8,or
80%.
o TheCWallelemusthaveafrequencyof1.00.8=0.2,or20%.
Whentherearetwoallelesatalocus,theconventionistouseptorepresentthe
frequencyofonealleleandqtorepresentthefrequencyoftheother.

Thusp,thefrequencyoftheCRalleleinthispopulation,is0.8.

ThefrequencyoftheCWallele,representedbyq,is0.2.

Alleleandgenotypefrequenciescanbeusedtotestwhetherevolutionisoccurringina
population.
TheHardyWeinbergprincipledescribesanonevolvingpopulation.
Populationgeneticistsdeterminewhatthegeneticmakeupofapopulationwouldbeifit
werenotevolving.

Wecanthencomparedatafromarealpopulationtowhatwewouldexpecttoseeifthe
populationwerenotevolving.

Ifwefinddifferences,wecanconcludethatthepopulationisevolving,andthentrytofigure
outwhy.
TheHardyWeinbergprincipledescribesthegenepoolofapopulationthatisnot
evolving.
TheHardyWeinbergprinciplestatesthatthefrequenciesofallelesandgenotypesina
populationsgenepoolwillremainconstantovergenerationsunlessacteduponbyagents
otherthanMendeliansegregationandrecombinationofalleles.

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o
o

Theshufflingofallelesbymeiosisandrandomfertilizationhasnoeffectontheoverall
genepoolofapopulation.
SuchagenepoolissaidtobeinHardyWeinbergequilibrium.

Inourimaginarywildflowerpopulationof500plants,80%(0.8)oftheflowercolor
allelesareCRand20%(0.2)areCW.
Howwillmeiosisandsexualreproductionaffectthefrequenciesofthetwoallelesinthe
nextgeneration?
Becauseeachgametehasonlyonealleleforflowercolor,weexpectthatagametedrawn
fromthegenepoolatrandomhasa0.8chanceofbearingaC Ralleleanda0.2chanceof
bearingaCWallele.
Supposethattheindividualsinapopulationnotonlydonategametestothenext
generationatrandombutalsomateatrandom.Inotherwords,allmalefemalematingsare
equallylikely.
o Theallelefrequenciesinthispopulationwillnotchangefromonegenerationtothenext.
Itsgenotypefrequencies,whichcanbepredictedfromtheallelefrequencies,willalso
remainunchanged.
Fortheflowercolorlocus,thepopulationsgeneticstructureisinastateofHardy
Weinbergequilibrium.
o Usingtheruleofmultiplicationforprobabilities,wecandeterminethefrequenciesofthe
threepossiblegenotypesinthenextgeneration.
o TheprobabilityofpickingtwoCRalleles(toobtainaCRCRgenotype)is0.80.8=0.64,
or64%.
o TheprobabilityofpickingtwoCWalleles(toobtainaCWCWgenotype)is0.20.2=
0.04,or4%.
o HeterozygousindividualsareeitherCRCWorCWCR,dependingonwhethertheCRallele
arrivedviaspermoregg.
o Theprobabilityofbeingheterozygous(withaC RCWgenotype)is0.80.2=0.16for
CRCW,0.20.8=0.16forCWCR,andthus0.16+0.16=0.32,or32%,forCRCW+CWCR.
Asyoucansee,theprocessesofmeiosisandrandomfertilizationhavemaintainedthe
samealleleandgenotypefrequenciesthatexistedinthepreviousgeneration.
TheHardyWeinbergprinciplestatesthattherepeatedshufflingofapopulationsgene
poolovergenerationsdoesnotincreasethefrequencyofonealleleoveranother.
o Theoretically,theallelefrequenciesinourflowerpopulationshouldremain0.8forC R
and0.2forCWforever.
Togeneralizetheexample,inapopulationthathastwoalleleswithfrequenciespandq,
thecombinedfrequenciesmustaddto1,or100%.
o Thereforep+q=1.
o Ifp+q=1,thenp=1qandq=1p.
Inthewildflowerexample,pisthefrequencyofredalleles(CR)andqisthefrequencyof
whitealleles(CW).
o TheprobabilityofaCRCRoffspringisp2(anapplicationoftheruleofmultiplication).
Inourexample,p=0.8andp2=0.64.
o TheprobabilityofaCWCWoffspringisq2.
Inourexample,q=0.2andq2=0.04.
o TheprobabilityofaCRCWoffspringis2pq.
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Inourexample,20.80.2=0.32.

o Thegenotypefrequenciesmustaddto1.0:p2+2pq+q2=1.0.

Forthewildflowers,0.64+0.32+0.04=1.0.

ThisgeneralformulaistheHardyWeinbergequation.

Usingthisformula,wecancalculatethefrequenciesofallelesinagenepoolifweknow
thefrequenciesofgenotypes,orwecancalculatethefrequenciesofgenotypesifweknowthe
frequenciesofalleles.
FiveconditionsmustbemetforapopulationtoremaininHardyWeinbergequilibrium.
TheHardyWeinbergprincipledescribesahypotheticpopulationthatisnotevolving.
Realpopulationsdoevolve,however,andtheiralleleandgenotypefrequenciesdochange
overtime.
Populationsevolvebecausethefiveconditionsfornonevolvingpopulationsarerarely
metforlonginnature.
ApopulationmustsatisfyfiveconditionsifitistoremaininHardyWeinberg
equilibrium:
1. Nomutations.Alterationofallelesordeletionorduplicationofgeneswillmodifythe
genepool.
2. Randommating.Ifindividualspickmateswithcertaingenotypes,orifinbreedingis
common,themixingofgameteswillnotberandomandgenotypefrequencieswill
change.
3. Nonaturalselection.Differentialsurvivalorreproductivesuccessamonggenotypeswill
alterallelefrequencies.
4. Extremelylargepopulation.Insmallpopulations,chancefluctuationsinthegenepool
willcauseallelefrequenciestochangeovertime.Theserandomchangesarecalled
geneticdrift.
5. Nogeneflow.Geneflow,thetransferofallelesduetothemigrationofindividualsor
gametesbetweenpopulations,willchangethefrequenciesofalleles.
Departurefromanyoftheseconditionsresultsinevolutionarychange,whichiscommon
innaturalpopulations.

ItisalsocommonfornaturalpopulationstobeinHardyWeinbergequilibriumfor
specificgenes.

Apopulationcanbeevolvingatsomeloci,yetsimultaneouslybeinHardyWeinberg
equilibriumatotherloci.

Therateofevolutionarychangeinmanypopulationsissoslowthattheyappeartobe
closetoequilibrium.
o Insuchcases,wecanusetheHardyWeinbergequationtoestimategenotypeandallele
frequencies.
WecanapplytheHardyWeinbergprincipletoahumanpopulation.

WecanusetheHardyWeinbergprincipletoestimatethepercentofthehuman
populationthatcarriestheallelefortheinheriteddiseasephenylketonuria(PKU).
Aboutonein10,000babiesbornintheUnitedStatesisbornwithPKU,ametabolic
conditionthatresultsinmentalretardationandotherproblemsifleftuntreated.
o Thediseaseiscausedbyarecessiveallele.

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 NewbornsaretestedforPKU.Iftheyarediagnosedwiththedisease,theirsymptomsare
lessenedwithaphenylalaninefreediet.

IstheU.S.populationinHardyWeinbergequilibriumwithrespecttothePKUgene?

ThemutationrateforthePKUgeneisverylow.(condition1)

PeopledonotchoosetheirpartnersbasedonwhethertheycarrythePKUallele,and
inbreeding(marriagetocloserelatives)israreintheUnitedStates.(condition2)

SelectionagainstPKUactsonlyagainsttherareheterozygousrecessiveindividuals,and
thenonlyifthedietaryrestrictionsareignored.Asaresult,theeffectsofdifferential
survivalandreproductivesuccessamongPKUgenotypescanbeignored.(condition3)

TheU.S.populationisverylarge.(condition4)

PopulationsoutsidetheUnitedStateshavePKUallelefrequenciessimilartothoseseen
intheUnitedStates,sogeneflowdoesnotalterallelefrequenciessignificantly.
(condition5)

Becausetheconditionsaremet,thepopulationisinHardyWeinbergequilibrium.
Fromtheepidemiologicdata,weknowthatthefrequencyofhomozygousrecessive
individuals(q2intheHardyWeinbergprinciple)isonein10,000,or0.0001.
o Thefrequencyoftherecessiveallele(q)isthesquarerootof0.0001=0.01.
o Thefrequencyofthedominantallele(p)isp=1q,or10.01=0.99.
o Thefrequencyofcarriers(heterozygousindividuals)is2pq=20.990.01=0.0198,
orabout2%.

Thus,about2%oftheU.S.populationcarriesthePKUallele.

Concept23.3Naturalselection,geneticdrift,andgeneflowcanalterallele
frequenciesinapopulation.
Althoughnewmutationscanmodifyallelefrequencies,thechangefromgenerationto
generationisverysmall.

Thechangeissmallbecausemutationsarerare.

Mutationcanultimatelyhavealargeeffectonallelefrequencieswhenitproducesnew
allelesthatstronglyinfluencefitnessinapositiveornegativeway.

Nonrandommatingcanaffectthefrequenciesofhomozygousandheterozygous
genotypes,butitusuallyhasnoeffectonallelefrequenciesinthegenepool.
Thethreemechanismsthatdirectlyalterallelefrequenciestobringaboutevolutionary
changearenaturalselection,geneticdrift,andgeneflow.
Naturalselectionisbasedondifferentialreproductivesuccess.

Individualsinapopulationvaryintheirheritabletraits.

Individualswithvariationsbettersuitedtotheenvironmenttendtoproducemore
offspringthanthosewithvariationsthatarelesswellsuited.
Asaresultofselection,allelesarepassedontothenextgenerationinfrequencies
differentfromtheirrelativefrequenciesinthepresentpopulation.
Forexample,thefruitflyDrosophilamelanogasterhasanallelethatconfersresistanceto
severalinsecticides,includingDDT.
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Thisallelehasafrequencyof0%inlaboratorystrainsofDrosophilacollectedinthe
1930s,beforeDDTwasused.

Instrainsestablishedafter1960,following20yearsofDDTuse,theallelefrequencyhad
increasedto37%.

Themutationeitherarosebymutationbetween1930and1960orwaspresentinthe
population,butveryrare,priorto1930.

DDTwasastrongselectiveforce,favoringallelesthatconferredresistance.

Geneticdriftresultsfromchancefluctuationsinallelefrequenciesinsmallpopulations.
Geneticdriftoccurswhenchangesingenefrequenciesfromonegenerationtoanother
occurbecauseofchanceevents(samplingerrors)thatoccurinsmallpopulations.
o Forexample,youwouldnotbetoosurprisedifatossedcoinproducedsevenheadsand
threetailsintentosses,butyouwouldbesurprisedifyousaw700headsand300tailsin
1,000tossesyouwouldexpectcloseto500ofeach.
o Thesmallerthesample,thegreaterthechanceofdeviationfromtheexpectedresult.
Inalargepopulation,allelefrequenciesdonotchangefromgenerationtogenerationby
chancealone.
o Inasmallwildflowerpopulationwithastablesizeofonlytenplants,however,genetic
driftcancompletelyeliminatesomealleles.
Geneticdriftinsmallpopulationsmayoccurasaresultoftwosituations:thebottleneck
effectorthefoundereffect.
Thefoundereffectoccurswhenanewpopulationisstartedbyasmallnumberof
individualswhodonotrepresentthegenepoolofthelargersourcepopulation.
o Atanextreme,apopulationcouldbestartedbyasinglepregnantfemaleorasingleseed
withonlyatinyfractionofthegeneticvariationofthesourcepopulation.

Geneticdriftcontinuesfromgenerationtogenerationuntilthepopulationgrowslarge
enoughforsamplingerrorstobeminimal.
Foundereffectshavebeendemonstratedinhumanpopulationsthatstartedfromasmallgroup
ofcolonists.
Thebottleneckeffectoccurswhenthenumbersofindividualsinalargepopulationare
drasticallyreducedbyadisaster.
o Bychance,someallelesmaybeoverrepresentedandothersunderrepresentedamongthe
survivors.
o Someallelesmaybeeliminatedaltogether.
Geneticdriftcontinuestochangethegenepooluntilthepopulationislargeenoughto
eliminatetheeffectofchancefluctuations.
Thebottleneckeffectisanimportantconceptinconservationbiologyofendangered
species.
Geneticdrifthasplayedanimportantroleintheconservationofthegreaterprairie
chicken(Tympanuchuscupido).

MillionsofgreaterprairiechickensoncelivedontheprairiesofIllinois.

By1993,onlytwopopulationsofbirdsremainedinIllinois,togethertotalingfewerthan
50birds.

Thefewsurvivingbirdshadlowlevelsofgeneticvariation.Lessthan50%oftheeggs
hatched,comparedtohigherhatchingratesoflargerpopulationsinotherstates.

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 Thesedatasuggestthatgeneticdriftduringthebottleneckmayhaveledtoalossof
geneticvariationandanincreaseinthefrequencyofdeleteriousalleles.
JuanBouzatandhiscolleaguesextractedDNAfrom165museumspecimensofIllinois
greaterprairiechickens.

Tenofthebirdswerecollectedinthe1930s,whentheIllinoispopulationwas25,000.

Fivewerecollectedinthe1960s,whenthepopulationwas1,000.

Theresearcherssurveyedsixlociandfoundthatthesmall1993populationhadlostnine
allelespresentinthemuseumspecimens.

Tocounterthenegativeeffectsofgeneticdrift,theresearchersbroughtin271birdsfrom
neighboringstates.

NewallelesenteredtheIllinoispopulation,andtheegghatchingrateimprovedtogreater
than90%.

Thisexamplehighlightsfourkeypointsaboutgeneticdrift.

1. Geneticdriftissignificantinsmallpopulations.
2. Geneticdriftcausesallelefrequenciestochangeatrandom.
3. Geneticdriftleadstoalossofallelicvariationwithinpopulations.
4. Geneticdriftcancauseharmfulallelestobecomefixed.
Apopulationmayloseorgainallelesbygeneflow.
Geneflowisthetransferofallelesamongpopulationsduetothemigrationoffertile
individualsorgametes.
o Forexample,ifanearbywildflowerpopulationconsistedentirelyofwhiteflowers,its
pollen(CWallelesonly)couldbecarriedintoourhypotheticalpopulation.
o TheresultwouldbeanincreasethefrequencyofC Wallelesinthehypotheticalpopulation
inthenextgeneration.

o
o

Geneflowtendstoreducedifferencesbetweenpopulations.
Ifextensiveenough,geneflowcanamalgamateneighboringpopulationsintoasingle
populationwithacommongenepool.
Humanstodaymigratemuchmorefreelythaninthepast,andgeneflowhasbecomean
importantagentofevolutionarychangeinhumanpopulationsthatwerepreviously
isolated.

Whenneighboringpopulationsliveindifferentenvironments,allelesmaybetransferred
thatlimittheadaptationofoneorbothallelestotheirlocalenvironment.

Forexample,bentgrass(Agrostistenuis)populationsgrownearcoppermines,where
theyareexposedtohighconcentrationsoftoxiccopper.

Ifallelesforcoppertolerancearepresentinthebentgrasspopulation,increasedcopper
tolerancerapidlyspreadswithinthelocalpopulation.

Onnearbysoilsnotcontaminatedwithcopper,coppertolerantplantsareata
disadvantageandreproducepoorly.

Becausebentgrassiswindpollinated,coppertoleranceallelesaretransferredtonon
minesoils,andcopperintolerantallelesaretransferredtominesoils.

Beneficialallelescanalsobetransferredbetweenpopulations.

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Forexample,geneflowhasresultedintheworldwidespreadofinsecticideresistance
allelesinthemosquitoCulexpipiens,avectorofWestNilevirus.

Geneflow,likemutation,canintroducenewallelesintoapopulation.

Becausegeneflowcanoccuratahigherratethanmutation,itcanhaveagreatereffecton
allelefrequencies.

Oncegeneflowormutationintroducesanewalleletoapopulation,naturalselectionmay
causethenewalleletoincrease(asintheinsecticideresistanceallelesinmosquitoes)or
decrease(asinthecoppertoleranceallelesinbentgrass).

Concept23.4Naturalselectionistheonlymechanismthatconsistentlycauses
adaptiveevolution.
Ofallthefactorsthatcanchangeagenepool,onlynaturalselectionleadstothe
adaptationofanorganismtoitsenvironment.
Thetermsstruggleforexistenceandsurvivalofthefittestaremisleadingbecausethey
suggestthatindividualscompetedirectlyincontests.

Insomeanimalspecies,malesdocompetedirectlyformates.

Reproductivesuccessisgenerallysubtler,however,anddependsonfactorsotherthanthe
battleformates.
o Forexample,abarnaclemayproducemoreeggsthanitsneighborsbecauseitismore
efficientatfilteringfoodfromthewater.
o Amothmayhavemoreoffspringthanitscompetitorsbecauseitsbodycolorsconcealit
frompredators,allowingittosurvivelongenoughtoreproduce.
Theseexamplesillustratehowadaptiveadvantagecanleadtogreaterrelativefitness:the
contributionanindividualmakestothegenepoolofthenextgeneration,relativetothe
contributionsofotherindividuals.
Althoughwemayrefertotherelativefitnessofagenotype,theentitythatissubjectedto
naturalselectionisthewholeorganism,nottheunderlyinggenotype.

Naturalselectionactsonthegenotypeindirectly,viahowthegenotypeaffectsthe
phenotype.

Therelativefitnessconferredbyaparticularalleledependsonthegeneticand
environmentalcontextsinwhichitisexpressed.

Forexample,aslightlydisadvantageousallelemaybenefitfrombeingclosetoa
favorablealleleonanadjacentlocus.

Therearethreemodesofselection:directional,disruptive,andstabilizing.
Naturalselectioncanalterthefrequencydistributionofheritabletraitsinthreeways,
dependingonwhichphenotypesinapopulationarefavored.

Thethreemodesofselectionarecalleddirectional,disruptive,andstabilizingselection.

Directionalselectionismostcommonduringperiodsofenvironmentalchangeorwhen
membersofapopulationmigratetoanewhabitatwithdifferentenvironmentalconditions.
Directionalselectionshiftsthefrequencycurveforaphenotypiccharacterinone
directionbyfavoringindividualswhodeviatefromtheaverage.
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Forexample,fossilevidenceindicatesthattheaveragesizeofblackbearsinEurope
increasedduringeachglacialperiod,onlytodecreaseagainduringthewarmer
interglacialperiods.
Largebearshaveasmallersurfacetovolumeratioandarebetteratconservingbodyheat
duringperiodsofextremecold.

Disruptiveselectionoccurswhenenvironmentalconditionsfavorindividualsatboth
extremesofthephenotypicrangeoverthosewithintermediatephenotypes.
o Forexample,twodistinctbilltypesarepresentinCameroonsblackbellied
seedcrackers.
o Largerbilledbirdsaremoreefficientinfeedingonhardseeds,andsmallerbilledbirds
aremoreefficientinfeedingonsoftseeds.
o Birdswithintermediatebillsarerelativelyinefficientatcrackingbothtypesofseedsand
thushavelowerrelativefitness.

Stabilizingselectionfavorsintermediatevariantsandactsagainstextremephenotypes.

o
o

Stabilizingselectionreducesvariationandmaintainsthestatusquoforatrait.
Humanbirthweightissubjecttostabilizingselection.
Babiesmuchlargerorsmallerthan34kghavehigherinfantmortalitythanaverage
sizedbabies.

Naturalselectionplaysakeyroleinadaptiveevolution.

Theadaptationsoforganismsareessentialtotheirsurvivalandreproductivesuccess.

Naturalselectionincreasesthefrequenciesofallelesthatenhancesurvivaland
reproduction.

Astheproportionofindividualsthathavefavorablecharacteristicsincreasesovertime,
thematchbetweenaspeciesanditsenvironmentimproves,andadaptiveevolution
occurs.

Becausethephysicalandbiologicalcomponentsofanorganismsenvironmentmay
change,adaptiveevolutionisacontinuous,dynamicprocess.
Naturalselectionistheonlyevolutionarymechanismthatconsistentlyleadstoadaptive
evolution.
Sexualselectionmayleadtopronouncedsecondarydifferencesbetweenthesexes.
CharlesDarwinwasthefirstscientisttoinvestigatesexualselection,whichisnatural
selectionformatingsuccess.
Sexualselectionresultsinsexualdimorphism,markeddifferencesbetweenthesexesin
secondarysexualcharacteristicsnotdirectlyassociatedwithreproduction.
o Malesandfemalesmaydifferinsize,coloration,andornamentation.
o Invertebrates,malesareusuallythelargerandshowiersex.

Itisimportanttodistinguishbetweenintrasexualandintersexualselection.

Intrasexualselectionisdirectcompetitionamongindividualsofonesexformatesofthe
oppositesex.
o Competitionmaytaketheformofdirectphysicalbattlesbetweenindividuals,usually
males.
Forexample,asinglemalemaypatrolagroupoffemalesandpreventothermales
frommatingwiththem.
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Morecommonly,ritualizeddisplaysdiscouragelessercompetitorsanddetermine
dominance.
Evidenceisgrowingthatintrasexualselectioncantakeplacebetweenfemalesaswell.
Inringtailedlemurs,femalesdominatemalesandalsoestablishdominance
hierarchies.

Intersexualselection,ormatechoice,occurswhenmembersofonesex(usually
females)arechoosyinselectingtheirmatesfromindividualsoftheothersex.
o Becausefemalesinvestmoreineggsandparentalcare,theyarechoosierabouttheir
matesthanmales.
o Afemaletriestoselectamatethatwillconferafitnessadvantageontheirmutual
offspring.
o Inmanycases,thefemalechoosesamalebasedonhisshowyappearanceorbehavior.
o Somemaleshowinessdoesnotseemtobeadaptiveexceptinattractingmatesandmay
evenputthemaleatconsiderablerisk.
Forexample,brightplumagemaymakemalebirdsmorevisibletopredators.
Eveniftheseextravagantfeatureshavesomecosts,anindividualthatpossessthem
willhaveenhancedfitnessiftheyhelpthatindividualgainamate.
Everytimeafemalechoosesamatebasedonappearanceorbehavior,she
perpetuatestheallelesthatcausedhertomakethatchoice.
Shealsoenablesamalewiththatparticularphenotypetoperpetuatehisalleles.
Howdofemalepreferencesforcertainmalecharacteristicsevolve?Aretherefitness
benefitstoshowytraits?
o Severalresearchersaretestingthehypothesisthatfemalesusemalesexual
advertisementstomeasurethemalesoverallhealthandimmunefunction.
o Maleswithseriousparasiticinfectionsmayhavedull,disheveledplumage.
Theseindividualsareunlikelytowinmanyfemales.
o Ifafemalechoosesashowymate,shemaybechoosingahealthyone,andherbenefitis
agreaterprobabilityofhavinghealthyoffspring.
Diploidyandbalancingselectionpreservegeneticvariation.
Whatpreventsnaturalselectionfromreducinggeneticvariationbycullingall
unfavorablegenotypes?
Thetendencyfornaturalselectiontoreducevariationiscounteredbymechanismsthat
preserveorrestorevariation,includingdiploidyandbalancedpolymorphisms.
Diploidyineukaryotespreventstheeliminationofrecessiveallelesviaselectionbecause
recessiveallelesdonotaffectthephenotypeinheterozygotes.
o Evenrecessiveallelesthatareunfavorablecanpersistinapopulationthroughtheir
propagationbyheterozygousindividuals.
Recessiveallelesareexposedtoselectiononlywhenbothparentscarrythesame
recessivealleleandcombinetworecessiveallelesinonezygote.
o Thishappensonlyrarelywhenthefrequencyoftherecessivealleleisverylow.
o Therarertherecessiveallele,thegreaterthedegreeofprotectionithasfromnatural
selection.
Heterozygoteprotectionmaintainsahugepoolofallelesthatmaynotbesuitableunder
thepresentconditionsbutmaybecomebeneficialwhentheenvironmentchanges.
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Naturalselectionitselfpreservesvariationatsomegeneloci.

Balancingselectionoccurswhennaturalselectionmaintainsstablefrequenciesoftwoor
morephenotypesinapopulation.

o
o

Onemechanismthatproducesbalancedpolymorphismisheterozygoteadvantage.
Insomesituations,individualswhoareheterozygousataparticularlocushavegreater
fitnessthanhomozygotes.
Inthesecases,naturalselectionwillmaintainmultipleallelesatthatlocus.

Heterozygoteadvantageisdefinedintermsofthegenotype,notthephenotype.

Whetherheterozygoteadvantagerepresentsstabilizingordirectionalselectiondepends
ontherelationshipbetweengenotypeandphenotype.

Ifthephenotypeofaheterozygoteisintermediatetothephenotypesofboth
homozygotes,thenheterozygoteadvantageisaformofstabilizingselection.

Heterozygoteadvantagemaintainsgeneticdiversityatthehumangeneforthe
polypeptidesubunitchainofhemoglobin.
o Arecessivealleleatthatlocuscausessicklecelldisease.
o Homozygousrecessiveindividualssufferfromsicklecelldisease.
o Homozygousdominantindividualsarevulnerabletomalaria.
o Heterozygousindividualsareresistanttomalaria.
Thefrequencyofthesicklecellalleleishighestinareaswherethemalarialparasiteis
common.
o InsomeAfricantribes,thesicklecellalleleaccountsfor20%ofthegenepool,avery
highfrequencyforsuchaharmfulallele.
o Evenatthishighfrequency,only4%ofthepopulationsuffersfromsicklecelldisease(q2
=0.20.2=0.04),while32%ofthepopulationisresistanttomalaria(2pq=20.8
0.2=0.32).
o Theaggregatebenefitofthesicklecellalleleinthepopulationbalancesitsaggregate
harm.
Asecondmechanismthatpromotesbalancedpolymorphismisfrequencydependent
selection.
Frequencydependentselectionoccurswhenthefitnessofanyonemorphdeclinesifit
becomestoocommoninthepopulation.
o Frequencydependentselectionhasbeenobservedinanumberofpredatorprey
interactionsinthewild.
Somegeneticvariations,neutralvariations,havenegligibleimpactonfitness,andthus
naturalselectiondoesnotaffectthesealleles.
o Manyofthenucleotidedifferencesinnoncodingsequenceswithinthehumangenome
appeartoconfernoselectiveadvantageordisadvantage.
Naturalselectioncannotfashionperfectorganisms.
Thereareatleastfourreasonsnaturalselectioncannotproduceperfection.
1. Selectioncanactonlyonexistingvariations.
Naturalselectionfavorsonlythefittestphenotypesamongthosecurrentlyinthe
population,whichmaynotbetheidealtraits.
Newadvantageousallelesdonotariseondemand.
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2. Evolutionislimitedbyhistoricalconstraints.
Evolutiondoesnotscrapancestralfeaturesandbuildnewcomplexstructuresor
behaviorfromscratch.
Evolutioncooptsexistingfeaturesandadaptsthemtonewsituations.
Forexample,birdsmightbenefitfromhavingwingsplusfourlegs.However,birds
descendedfromreptilesthathadonlytwopairsoflimbs.Cooptingtheforelimbsfor
flightleftonlytwohindlimbsformovementontheground.
3. Adaptationsareoftencompromises.
Eachorganismmustdomanydifferentthings.
Becausetheflippersofasealmustallowittowalkonlandandalsoswimefficiently,
theirdesignisacompromisebetweentheseenvironments.
Humanlimbsareflexibleandallowversatilemovementsbutarepronetoinjuries,
suchassprains,tornligaments,anddislocations.
Betterstructuralreinforcementofhumanlimbswouldcompromisetheiragility.
4. Chance,naturalselection,andtheenvironmentinteract.
Chanceeventsaffectthesubsequentevolutionaryhistoryofpopulations.
Forexample,thefoundersofnewpopulationsmaynotnecessarilybetheindividuals
bestsuitedtothenewenvironment,butratherthoseindividualsthatwerecarried
therebychance.

Withtheseconstraints,evolutioncannotcraftperfectorganisms.

Infact,theevidenceforevolutioncanbeseeninthemanyimperfectionsofthe
organismsitproduces.

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