I n s t r u c t o r ’s M a n u a l

Matthew E. Gruwell

INSTRUCTOR’SMANUAL

CarlT.BergstromandLeeAlanDugatkin’s

EvolutionSECONDEDITIONInstructor’sManualbyMatthewE.GruwellPENNSTATEERIE,THEBEHRENDCOLLEGE W•W•Norton&Company•NewYork•London

08Fall

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TableofContentsPartI:FoundationsofEvolutionaryBiologyChapter1:AnOverviewofEvolutionaryBiology p1Chapter2:EarlyEvolutionaryIdeasandDarwin’sInsight p5Chapter3:NaturalSelection p12Chapter4:PhylogenyandEvolutionaryHistory p19Chapter5:InferringPhylogeny p26PartII:EvolutionaryGeneticsChapter6:TransmissionGeneticsandtheSourcesofGeneticVariation p34Chapter7:TheGeneticsofPopulations p41Chapter8:EvolutioninFinitePopulations p50Chapter9:EvolutionatMultipleLoci p58Chapter10:GenomeEvolution p65PartIII:TheHistoryofLifeChapter11:TheOriginandEvolutionofEarlyLife p71Chapter12:MajorTransitions p77Chapter13:EvolutionandDevelopment p83Chapter14:SpeciesandSpeciation p88Chapter15:ExtinctionandEvolutionaryTrends p93PartIV:EvolutionaryInteractionsChapter16:SexandSexualSelection p100Chapter17:TheEvolutionofSociality p107Chapter18:Coevolution p113Chapter19:HumanEvolution p118Chapter20:EvolutionandMedicine p125

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Chapter1

AnOverviewofEvolutionaryBiology

ChapterSummaryandDiscussionPointsIntroductionCharlesDarwinevolvedbiologywhenhetheorizedthatallorganismsshareacommonancestryandthatanorganism’sfitnesswithinitsenvironmentistheresultofnaturalselection.Darwin’sevolutionarytheoryledtoanaturalexplanationforthediversityoflifeand,thus,aparadigmshiftinbiologicalthoughtthatnowreacheseverysubdisciplineofbiology.Forexample,whenanatomystudentsnowstudythestructureofbonesinthehumanwrist,itispossibletoextrapolatetheirunderstandingofskeletalstructurestoothertetrapodsandfindbonesofthesameoriginbeingusedinsimilarorverydifferentways.Section1.1:ABriefIntroductiontoEvolution,NaturalSelection,andPhylogeneticsAsaprimertotheensuingchapters,thissectionexplainstheideaofdescentwithmodificationasbeingthewayinwhichallorganismschangeovertimeduetonaturalorartificialselection,leadingtothepremisethatalllifeissubjecttotheprinciplesofevolution.Forexample,totrulyunderstandaspeciessuchasHomosapiens,onemuststudyandunderstandprimates,theirclosestrelatives.Thissectionoffersstudentsarefresheronsomeofthebasicsofgenetics,whichsetsthestagefordefiningselection.Artificialselectioniscoveredusingmultipleexamplesofcrops,includingadetailedfigureofhumanselectionofmanydifferentfoodcropsoverthousandsofyears,aswellasamorespecificexampleinvolvingstrawberries.Theseexamplesarethencontrastedwithanexampleofnaturalselectioninvolvingpesticideresistanceininsects.Antibioticresistancealsoisexploredasaproblemthatwillbesolvedonlybyusingtheprinciplesofevolutionarybiology.Lastly,tohelpstudentsseethatevolutionaryprinciplesaidinthestudyofconservationbiology,theauthorsintroducetheconceptoftreethinkingusingthephylogenetictreeoflifeandextinction.ThissectionconcludeswiththeKeyConceptQuestion(seeKeyConceptQuestion1.1):Canyouthinkofotherwaysthatevolutionarythinkingmightaffectstudiesinconservationbiology?DiscussionPoints:

• WhattwothingsdidDarwinoriginallynoticeandhowdidthatleadtohistheory?WhataresomeexamplesfromthenaturalworldthatconfirmDarwin’sidea?

• Howdoartificialandnaturalselectiondiffer?Howdoesartificialselectiondemonstratetherealityofevolution?Whenareexamplesofhumaninfluence

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causingchangeinasystemstillconsiderednaturalselection?• Whyisantibioticresistanceaseriousthreattomodernmedicine?Howwill

theprinciplesofevolutionarybiologyhelpcontrolthisproblem?Section1.2:EmpiricalandTheoreticalApproachestotheStudyofEvolution Thissectiondividesthetypesofresearchconductedinevolutionarybiologyintotwomajormethods:empiricalandtheoretical.Empiricalresearchisfurtherdividedintoobservationormanipulationofanaturalsystem.Observationstudiesinvolvegatheringdatafromasystemwithoutmanipulatingitandincludestudiessuchasresearchonthefossilrecord,inferringphylogenyfromgeneticsequences,orobservingbehavior.Thisisdemonstratedbycomparingsimilaritiesbetweenchimpsandhumansinanatomyandgenomicstoshowcommonancestryanddeterminewhatseparatesthemgenetically.Manipulationofnaturalsystemsisexplainedbyasecondexampleofempiricalstudy,whichinvolvesbreedingsystemsandtestessizein33speciesofprimates.TheoreticalbiologyisintroducedasamajordisciplineinevolutionarybiologythroughtheworkofRonaldA.FisherandWilliamD.Hamilton,inwhichtheyutilizedmathematicalmodelstopredictandunderstandsexratios.ThisisfollowedbyanactualexampleofchangingsexratiosinthebluemoonbutterflyontheSamoanislands.DiscussionPoints:

• Ofthetworesearchmethodsdiscussed,whichwouldbemostusefulintestingspecifichypotheses?

• Whatistheusefulnessoftheoreticalstudiesinevolutionarybiology?AnswerstoReviewQuestions

1. Paradigm shifts represent fundamental changes in thewaywe think aboutandstudynaturefromascientificperspective.

2. (1)All species aredescended fromoneor a fewancestral life-forms. (2)Aprocess Darwin dubbed natural selection explains the fit of organism toenvironment.

3. Thisevidence includes,but isnot limited to,moleculargenetic, anatomical,

physiological,behavioral,developmental,andhormonaldata.Suchdatacanbeamassedfromfossilsorcontemporaryorganisms.

4. Artificial selection is the human-driven analog to the process of naturalselection. In artificial selection,we selectively breed individualswith traitsthatarebeneficialtous.

5. Bacteria reproduce extremely rapidly—as fast as hourly or even faster.Bacterialpopulationsizesareenormous,providingavast supplyofgeneticvariation. Naturalselectionimposedbyantibioticresistanceisverystrong.

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For all of these reasons mutations conferring antibiotic resistance canquicklyariseandrapidlyspreadthroughbacterialpopulations.

6. Phylogeneticdiversityisameasureofdiversitythattakesintoaccounthowmuchoftheevolutionaryhistoryofthegroupbeingstudiedispreserved.

7. The two basic ways to gather empirical evidence to test hypotheses areobservationalandexperimentalstudies.

8. Neutralmutationsaremutationsthatdonotaffectfitness.

9. Sexratiomeasurestheratiooffemalestomalesinagivenpopulation.

10. Regardless of whether empirical work precedes theoretical work or viceversa, each of these informs the other, and this (ideally) leads to thegenerationofnew,testablehypotheses.

AnswerstoKeyConceptApplicationQuestions

11. Wecanapplytheexactsamereasoningweusedforthecaseoftheevolutionofantibioticresistanceinbacteriatothecasesofantiviralresistanceevolutionbyvirusesandantifungalresistanceevolutionbyfungi.Similarargumentsalsoapplyinagriculturetotheevolutionofherbicide-resistantweedsandpesticide-resistantinsects.

12. Answerswillvary,butthetheoryofcontinentaldriftandplatetectonicsrepresentsonesuchparadigmshift.Thetheoryofcontinentaldriftproposedthatcontinentsaredriftinglandmassesthatmoveovergeologicaltimeandthereforeeventhelarge-scalegeographyoftheearthisdynamicallychanging.Thetheoryofplatetectonicsprovideduswithamechanisticunderstandingofhowcontinentaldrifttakesplace.Together,thesetheoriesprovideanexplanationofwhysurfaceofourplanetlooksthewaythatitdoesnow,howlargelandmassesarecreatedanddestroyed,andsomuchmore.

13. Thestudyofdescentwithmodificationistheconceptualgluethatunitesallofthelifesciences.Withoutadoptinganevolutionaryapproachinthebiologicalsciences,wehavemanypotentiallyimportant,disparatefacts,butnocommontheoreticalperspectivetounitethem.

14. Byselectingfortraitsthatareeitheraestheticallypleasingorinsomesensepractical,humanshaveshapedeverythingfromthesize(thinkminiaturepoodletoGreatDane)tothebehavior(herding,hunting,retrieving)ofdomesticateddogbreeds.

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15. Nowthatthemoleculargenetictechniquesassociatedwithstudyinggeneexpressionarewidelyavailable,evolutionarybiologistscanstudynotjusthowdifferencesbetweenspeciesinprotein-codingDNAsequencesareassociatedwithevolutionarychange,butalsohowdifferencesinwhengenesareturnedonandoffareassociatedwithevolutionarychange.

16. Aswenotedinthechapter,a1femaleto1malesexratioissocommonthatitishardtoimagineanyothersexratio,butwhatwewanttounderstandiswhya1:1sexratioissoubiquitousinthefirstplace.Todothat,weneedtoconsiderotherpossiblesexratios,andthenexaminewhichsexratioisfavoredbynaturalselectionandunderwhatconditions.Oncewemakeclearourassumptions,mathematicalmodelsgiveusthepowertodoexactlythat.

SuggestedReadingsThislistofsuggestedreadingsisprintedattheendofthischapterinthestudenttextbookandreprintedhereforyourconvenienceandplanning.Birkhead,T.R.,andT.Pizzari.2002.Postcopulatorysexualselection.NatureReviews

Genetics3:262–273.•Thispaperwillgiveyouabetterunderstandingofthespermcompetitionandspermallocationworkwediscussedinthischapter.

Engelstädter,J.,andG.D.D.Hurst.2009.Theecologyandevolutionofmicrobesthat

manipulatehostreproduction.AnnualReviewofEcology,EvolutionandSystematics40:127–149.

•AdetailedreviewonissueswediscussedintheWolbachia/bluemoonbutterflysexratioexample.

Huxley,T.H.1863.EvidenceofMan’sPlaceinNature.D.Appleton,NewYork.•Huxley—Darwin’scolleague—presentedevidenceforhumanevolutioninthisbook.

Kuhn,T.1962.TheStructureofScientificRevolutions.UniversityofChicagoPress,Chicago.

•Inthisvolume,aclassicinthephilosophyofscience,Kuhnoutlinestheideaofaparadigmshift.

Varki,A.,D.H.Geschwind,andE.E.Eichler.2008.Explaininghumanuniqueness:genomeinteractionswithenvironment,behavior,andculture.NatureReviewsGenetics9:749–763.

•Aninterestingdiscussionofhowtounderstandwhatmoleculargeneticcomparisonstellus(anddon’ttellus)aboutsimilaritiesanddifferencesbetweenhumansandotherprimates.