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Biology230
2010:OdysseyI
Thisisaclosedbook/notesexam;useofcalculatorsisfine.Pleasewriteininksothatwe
mayregradeuponrequest.Youwillhavetheentireperiodinwhichtocompletetheexam.
Alistofformulaefrompopulationandquantitativegeneticsisprovidedonthelastpage.
PleasewriteyournameonALLpagesoftheexam,aspagesmaybeseparatedtofacilitate
grading.Pleaseshowallyourworksothatwemaygivepartialcredit.
1.Clearly(butsuccinctly)defineandstatethesignificanceofthefollowingterms(5pts.
each).
pangenesis
heterozygosity
2.DarwinbeginsOntheOriginofSpecieswithadiscussionofvariationunder
domestication.Itistheorizedthattheoriginbeginswithdomesticationbecauseartificial
selectionissimilartonaturalselectionbutnotnearlyascontroversial.Darwinusesthis
chaptertobegintointroducesomeofthecentralpoints(orideas)inOntheOriginof
Species(notallofwhichareuniquetoDarwin).NametwoofthemajorpointsofDarwins
theorythatareaddressedinthefirstchapter.Provideanexamplefromthebookand
describehowitdemonstratesonofthemajorpointsyounamed.(5pts).
3.
Length
Course gravel
Fine
gravel
K-no predators
R-low predation
C-high predation
Describe how the results above support Endler's hypotheses that 1) predation causes selection for
crypsis and 2) sexual selection causes selection for conspicuousness? (4 pts)
4.Alepidopteristisstudyingthegeneticsofwingpatternsinbutterfliesanddiscovered
thatthethreeformsofwingspottingpatternsonthehindwing(onespot,twospots,three
spots)followedanadditivemodelofinheritanceatasinglemendelianlocus.Duringher
fieldcollectionsofbutterfliesforbreedingexperiments,shenoticedthatthesebutterflies
wereofteneatenbybirds.Uponreviewingtheliterature,shefoundthatthefamed
entomologist,I.B.Smart,hypothesizedthattheonespotphenotypewasmorecrypticand
presumablyofhigherfitnessinhighpredationenvironments.Intrigued,shedecidedtotest
thishypothesisbyobservingpredationeventsonthethreedistinctwingphenotypes.After
painstakingfieldobservationsof500predationeventsinarandomlymatingpopulation,
sheobservedthat45threespots,210twospots,and245onespotswereeatenbybirds.
Basedontheseobservations,areonespotbutterfliesmorecryptic?Wouldpredationby
birdsbepredictedtochangethefrequencyofwingspotphenotypesinthispopulation?
Clearlystateandsupportyouranswerquantitatively.(8pts.)
5.Identifyandbrieflyoutlinetwoimportantdifferencesbetweentheevolutionarytheories
ofDarwinandLamarck.(4pts).
6.Epigeneticsisarapidlyexpandingareaofgenetics;forourpurposeshere,epigenetics
referstomechanismsofheredityoutsideofthenormalMendeliantransmissionofgenes
fromparenttooffspring.Forexample,themethylationofgenesinadevelopingoffspring
bythemothercanresultingenesilencing;providedthisoccursbeforethegermlineis
sequestered,suchevents(shutdownofgeneexpressionbymethylation)areheritableand
subsequentlypassedfromparenttooffspring.Acolleagueofminerecentlycommented
thatepigeneticscouldbeconsideredLamarcksrevenge.Explain.(6pts).
6.Theconceptofeffectivepopulationsize(Ne)isintegraltotheevaluationofallele
frequencychangeovergenerationaltime.Inlecture,wediscussedhowNemayvaryfrom
speciestospecies(e.g.,largevertebratestendtobecharacterizedbylowerNeestimates
thanaresmallinvertebrates)aswellasfromgenetogene.Inasexuallyreproducing,
diploidorganism,anautosomalgenecanbeconsideredtohaveaneffectivepopulationsize
thatis4timeshigherthanamitochondrialgene(i.e.,autosomalgeneis4Ne,mitochondrial
geneisNe).Allelsebeingequal,wouldgeneticdriftbepredictedtohaveamore
pronouncedeffectonautosomalormitochondrialgenes?Wouldnaturalselectionbe
predictedtobemoreeffectiveinchangingautosomalgeneormitochondrialgeneallele
frequencies?Clearlyexplainyouranswers.(8pts).
7.Youareinterestedintheheritabilityofbodysizeininsects(no,really,youare).For
threespeciesyouhavethedatabelowonthevarianceinbodysizeinnaturalpopulations
anddataonthevarianceinbodysizeinhighlyinbred,homozygous(geneticallyidentical)
strainsofeachspeciesthatdevelopinthewildfromfertilizedeggs.Fromthesedata1)
estimateVGineachspecies.AssumethatVDandVIareequivalentacrossallspecies.
Identifythespeciesinwhichitwillbe2)easiesttoartificiallyselectforlargerorsmaller
bodysize,and3)mostdifficulttoselectforincreasedordecreasedbodysize.Clearlystate
why.(10pts.)
ObservedVariances
InbredStrains
NaturalPopulations
Species1
0.40
0.80
Species2
0.04
0.40
Species3
0.40
0.50
8.In1965KnudSickstudiedahemoglobinpolymorphismintheAtlanticcod,Gadus
morhua.Heobtainedgenotypedatafor1000fishfromthreedifferentpopulations;the
geographicproximityofthethreepopulationsisgiveninthefigurebelow.Previouslyit
hadbeenhypothesizedthatpopulation3wasreallyjustahybridgroupcomprisedof
individualsfrompopulation1andpopulation2.Thesedatawerecollectedtotestthis
hypothesis.Recallthatmigrationissimplythemovementofindividuals,geneflowisthe
actualinterbreedingofindividualsfromdifferentpopulations,andthatonegenerationof
randommatingrestorestheHardyWeinbergequilibrium.
Region
AA
Aa
aa
Pop.1
40
320
640
Pop 1
Pop 2
Pop.2
640
320
40
Pop.3
340
320
340
Pop 3
A.Whatcanyouinferaboutthepatternsofmatingwithineachofthethreeregions?
B.Howdothedataallowyoutodiscriminatebetweenmigrationandgeneflowamongthe
threeregions?
C.ProvideanexplanationforthepatternofgenotypefrequenciesinPopulation3.
Clearlyjustifyyouranswersqualitativelyandquantitatively.(11points)
9.WhileonsabbaticalattheTjrnMarineBiologicalLaboratoryinSweden,yousample
populationsoftherockweedFucusradicansfromtheBalticSeaandtheAtlanticOcean.You
observethatBalticpopulationsarecharacterizedbyanincreasedlevelofmuco
polysaccharidesthatcoatthealgae(slime).Thisincreasedlevelofmucushasbeen
hypothesizedasaselectiveresponsetotheextremeandbrackishconditionsexperienced
intheBaltic.Yoursabbaticalhost,phycologistPhil,hasrecentlyidentifiedagene
(daycare1)thatdeterminesthelevelofmucopolysaccharidesproduced.Youscreenthis
geneinyoursamples,observeseveralallelessegregatinginthepopulations,and
subsequentlycalculateFstas0.20.ProvidetwoexplanationsforahighFstvalueobserved
betweenBalticandAtlanticpopulationsofF.radicans.(6pts.).
10.Acolleagueofyourshasanextensivedatasetonmorphologicaltraitsandlifehistories
inapopulationofredwingedblackbirdsinMarshfield,Wisconsin.Inthispopulation,she
hasmeasuredbodysize,reproductivesuccess,eggsize,eggnumber,andlifespanofeach
birdinthepopulationoveraperiodof12years.Bloodwasalsoobtainedfromeach
individual,allowingaccuratedeterminationofpaternity.Giventhecomprehensivenature
ofthedatasetandthelargenumberofbirdsexamined,youareabletoperforma
quantitativegeneticdescriptionofthesetraitsinthispopulation.Youestimateheritability
foreachtraitandalsogeneratethevariance/covariancematrixamongtraits,fromwhich
youcalculategeneticcorrelations.Youfocusonfemalelifespanandreproductiveeffort
(whichincludesbotheggsizeandnumberofeggsproduced).Wouldyoupredictthe
geneticcorrelationbetweenthesetraitstobenegative,effectivelyzero,orpositive?Why?
(8points)
11.Inclass,wediscussedhow,accordingtothepopulationgeneticmodelsofR.A.Fisher,
selectionwillalwayscauseanincreaseintheaveragefitnessofapopulation.Wethenused
theexampleofsicklecellanemiainhumanstodemonstratehowthisshortsightedness
canconstraintheadaptivepotentialofpopulations(i.e.,becauseselectioncausesa
populationtogoupalocalfitnesspeak,thepopulationcannotcrossafitnessvalleyto
climbahigherpeak).Brieflyoutlineanalternativehypothesisforevolutionarychange,
proposedbyS.Wright,thatcouldallowpopulationstocrossfitnessvalleys.Whatwas
perceivedasthebiggestproblemwithWrightsmodel?(10pts).
12.Inourdiscussionsofbothpopulationandquantitativegenetics,weemphasized
mechanismsbywhichgeneticvariationcanbemaintainedinapopulation.Whyisthe
maintenanceofgeneticvariationsoimportantinevolutionarybiology?Providetwoways
inwhichvariationcanbemaintainedat1)thelevelofasinglegene(i.e.,allelicvariation)
AND2)atthelevelofthephenotype(i.e.,foraquantitative/continuous/polygenictrait).
(10pts).
Formulas
PopulationGenetics
p2+2pq+q2=1
p'=p(1u)+q(v)
p*=v/(u+v)
q*=u/(u+v)
p*=m/s(dominant)
q*=(m/s)1/2(recessive)
p*=t/(s+t)
q*=s/(s+t)
p=m(pxtpyt)
ppopulation1t+1=ppopulation1(1m)+p
population2(m)
m(qxtqyt)=sqx2(1qx)
FSR=(HRHS)/HR
FRT=(HTHR)/HT
FST=(HTHS)/HT
(1FSR)(1FRT)=1FST
FST1/(4Nem+1)
Wbar=wAAp2+wAa2pq+waaq2
pt+1=(wAAp2+wAapq)/(wAAp2+
wAa2pq+waaq2)
QuantitativeGenetics
P=G+E
G=A+D+I
VP=VG+VE
VG=VA+VD+VI
h2=VA/VP(narrow),h2=VG/VP
(broad)
R=h2S