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Preliminaries
Explainvs.assess

PhilosophyandEthicsof MolecularBiology Lecture3

Dr.SamuelSchindler CenterforScienceStudies DepartmentofPhysicsandAstronomy AarhusUniversity
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Queries:good! Reminderforpresenters:
Handouttotutor24hrsbeforepresentation! Printoutthehandouts SeealsoAULAondesignofhandouts
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Preliminaries
Newseminarinstructions
Inadditiontorequiredtext,readonepresentation text(assignmentinseminars) Identifyoneargument (informally)and/orcome upwithaquestion forthepresenters

Todaystopic
Thephenomenonofaltruism Units/levelsofselection R.Dawkinsview:theunitofselectionisnot theorganismbutthegene AttacksonDawkinsview

Seminarbehaviour/workethics

Recall
Thetheoryofevolution;twobasicprinciples
Variation (individualdifferences) Naturalselection(limitedfoodresources, competition,environmentalchanges,etc.)

Darwinsview
InDarwinsview,naturalselection operatesonindividuals E.g.fastantelopes
Intheancestralpopulationtherewas varietyofantelopeswithdifferentspeeds Yet,theslowoneswereweededoutby predators Onlythefastonessurvived(and procreated)
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Traditionalunitsofselection: Theindividualorganism(cf.Darwin1859/ standardview!) Groups ofindividuals(e.g.K.Lorentz) Entirespecies(S.J.Gould)

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Darwinsview
Problemwiththispicture:altruism
Someanimalssacrificethemselvesto savethemembersoftheirgroup/species Examples:bees stingingintruders(kills themselves);birds givingalarmcalls (focusingthepredatorsattentionon them)

Darwinsview
Darwinsuggestedthataltruismmakes sensefromtheevolutionaryperspective becauseitsavesgroups ofindividuals
Otherdefenders:Konrad Lorentz

Note:thisisinconsistent withhisoverall viewthatselectionworksonindividuals, ratherthangroups!

Thelatteristhestandardview

Note:iftheindividualistheunitof selection,whywouldtheindividual sacrificeitselfforother individuals?

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Infact,thestandardviewrendersaltruism evenunlikely .
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Individualismandaltruism
Supposethat
agroupconsistsofindividualswhoregularly exhibitaltruisticbehaviortowardstheirgroup peers thereareonlyfewegoisticindividualsinthose groups thealtruistic/egoistictraitisinherited

Individualismandaltruism
Theegoisticindividualshaveaselective advantageoveraltruisticonesinthatgroup:
Theycanbenefitfromthealtruisticbehavioroftheir peerswithout havingtoexposethemselvestorisks

Then
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Becauseegoisticbehaviorisadvantageousinthe survivalofthefittest,overtime,altruismwill disappear inthosegroups Hence,evenifaltruismdoesoccasionallyoccur,it isgoingtoselectedagainstbyevolution!

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Individualismandaltruism
theoptions: 1. Altruisticbehaviorisachimera
Optionadoptedbymanybiologists/theoristsof evolution

Puzzle: Howcanweaccommodatealtruism onthetheoryofevolution?

RichardDawkinsanswer:
Wecandothatbymakinggenes,not organisms,theunitofselection

2. Altruisticbehaviorisrealandneedstobe explained
whatwellfocusonhere
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RichardDawkins
Dawkins:TheSelfishGene
Published1976

Theselfishgene
Fromtheperspectiveofthegeneseye,altruism makesperfectlysense: Individualssacrificethemselvesinordertosave thegroupbecause
Thesurvivalchanceofparticulargenes (thatthe individualshareswiththegroup)isincreased!

Earlycareer:zoologist ProfessorforPublicUnderstanding ofScienceatOxford(19952008) Otherpublications

Theblindwatchmaker TheGodillusion
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Afteranattackerhasbeenfoughtoffandthe groupsavedbyanindividual:
thegenefrequencyinthegenepoolisincreased(at theexpenseoftheorganism)
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Furtherreasonswhythegene shouldbetheunitofselection.
1. JunkDNAmakesnosenseonthestandard viewofevolution(w/individuals=units) 2. Onlygeneshavetherightcharacteristics requiredbythetheoryofevolution

Furtherreasons(1)
Anotherargumentagainsttheindividual organismasunitofselection
Theexistenceofnoncoding/junkDNA:98%of ourgenome!

Fromtheperspectiveoftheorganism
sincejunkDNAhasnofunctionforthe organism/speciesthereisnogoodexplanation forwhyorganismsshouldkeepinheritingjunk DNA
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Furtherreasons(1)
Ongeneeyeview(p.44/45)
ThetruepurposeofDNAisnotthebuildingof organisms(inwhichtheyreside) Rather:thesolepurposeofDNAisittosurvive! JunkDNAisparasitic;itisaharmlessbutuseless passenger,hitchingaride!

Furtherreasons(2)
Theunitofselectionmust
havelotsofcopies
Otherwise:smallchanceofsurvival

havethepotentialtosurviveforasignificant periodofevolutionarytime(>longevity)

Neitherorganisms,norspecieshavethe secondcharacteristic

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Furtherreasons(2)
Eachindividualisunique.Youcannotget evolutionbyselectingbetweenentitieswhen thereisonlyonecopyofeachentity!
Eachindividualhasveryspecificcharacteristics:e.g. onlyoneleg

Furtherreasons(2)
Individualsarenotstablethings,theyare fleeting.genesareforever(35)
Genesare(normally)notdividedwhenpassed fromonegenerationtothenext

Sexualreproductionisnot replication.(34)
Weinheritonly50%ofourgeneticmaterialtoour progeny Afterafewgenerations:hardlyany proportionin yourdescendantsofwhatmadeYOU
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Ofcourse:geneswillnotliterally lastforever
justlongenoughtofunctionasaunitof naturalselection contrarytoorganisms
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Theselfishgene
Genesarereplicators
Onlytheytrulyreplicate

Theselfishgene
Ifweweretoldthatamanhadlivedalong andprosperouslifeintheworldofChicago gangsters,wewouldbeentitledtomake someguessesastothesortofmanhewas. Wemightexpectthathewouldhave qualitiessuchastoughness,aquicktrigger finger,andtheabilitytoattractloyalfriends. Thesewouldnotbeinfallibledeductions, butyoucanmakesomeinferencesabouta man'scharacterifyouknowsomething abouttheconditions inwhichhehas survivedandprospered.
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Organismsaremerelyvehicles ofgenes
Also:interactors Theymerelyarethegenesmeans totheend ofsurvival

Genesareselfishbecausealltheycareaboutistheir ownsurvival
Theyproduceorganisms/vehiclesonly inordertosurvive

Note:thisiswhatDawkinsthinksitisreasonableto infer
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Theselfishgene
Theargumentofthisbookisthatwe,andall otheranimals,aremachinescreatedbyour genes.LikesuccessfulChicagogangsters,our geneshavesurvived,insomecasesfor millionsofyears,inahighlycompetitive world.Thisentitlesustoexpectcertain qualitiesinourgenes.Ishallarguethata predominantqualitytobeexpectedina successfulgeneisruthlessselfishness.This geneselfishnesswillusuallygiveriseto selfishnessinindividualbehaviour.
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Genesandtheprimevalsoup
ThegenesstruggleofsurvivalNOW(in organisms)isnotmuchdifferentfromwhatit usedtobeatthebeginningoflifeonEarth!
Intheprimevalsoupreplicatorsaroseby accident(15ff.),diversifiedthroughreplication errors,andcompetedagainsteachother

Replicatorswithnumerousreplications (fecundity),lasting replication(longevity)and accurate replication(fidelity)wonout

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Theselfishgene
Theselfishness ofgenesoftendirectly translatesintothebehaviour oforganisms Prayingmantis:femalebitesoffthehead ofmaleafterorevenduringcopulation
Inordertofeedherself(andinorderto increasethemalessexualperformance!)

Theselfishgene
Butnote:theselfishnessofthegene doesnotnecessarily implythat organismsbeselfish Geneselfishnessisverymuch compatiblewithaltruistic behaviour
Helpingotherorganismsofthesame speciesisameanstotheendofreplicating genesofthesamekind

Emperorpenguins:sometimespushtheir peersintothesea
Inordertotestwhethertherearesealsinthe water(whomightkillthem)
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Howselfisharegenes?
Thegene isthebasicunitofselfishness
Infact:Genesarecompetingdirectlywiththeir allelesforsurvival,sincetheirallelesinthegene poolarerivalsfortheirslotonthechromosomeof futuregenerations.

Howselfisharegenes?
Trulyruthlesslyselfishgenes:outlaw genes
Noregardforwhethertheirreplicationis ofanybenefittotheorganismornot

atthegenelevel,altruismmustbebadand selfishnessgood(36)

Examples:
JunkDNA Drivinggenesthatdistortmeiosisintheir ownfavour,sothattheyhaveagreater chanceofmakingitintogametes
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Howselfisharegenes?
Important:Dawkinsconceptionofselfish genesdoesnot imply
thatgenesnevercooperate

Howselfisharegenes?
Yet,singlegenesmayverywellmakea differencetothephenotype
E.g.thelength oflegs

Genesmayservetheirowninterests bestbycooperatingwithothergenes(p. 37)

Hence:selfishnessthroughcooperation

Analogy:plantfertilizer
Otherthingsbeingequal,plantfertilizermake plantsgrow Otherimportantfactors:seed,soil,sun,water, etc./onlyiftheyarepresent,doesthefertilizer makeadifferencetoplantgrowth
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E.g.:buildingalegcannotbedonebya single gene

Butasinglegenewillprofitfromorganisms thatrunfast!

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Howselfisharegenes?
Dawkinscomparesgeneselectionism tothe coachsselectionofoarsmen(i.e.,rowers)inboat racing Thecoach hastopickanidealteam
Assemblesthreecrews andshufflesthemembers randomlyandletsthemraceagainsteachother Itturnsoutthat,onaverage,therearesome individualsthattendtobeinthewinningboats,and somewhotendtobeintheloosingones

Howselfisharegenes?
Analogytogeneselection
Oarsmen =genes Rivaloarsmenforaseatintheboat=allele Rowingfast=buildingavehiclewhichissuccessfulat surviving Coach=naturalselection

Note:anygene(goodandbadones)must cooperate withothersinordertorowfast

Cooperationisintheirselfinterest

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TasksandQuestions
Explainwhyaltruism isproblematiconthe standardviewofevolution ExplainDawkinsargumentsforwhythegene shouldbeviewedastheunit ofselection (ratherthantheorganism);mention
Altruism JunkDNA Characteristicsrequiredbyevolution Inparticular:longevity
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WhoagreeswithDawkins?
Handsup!

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Whatsageneacc.toDawkins?
Evolutionarygeneconception(fromWilliams)

Argumentsagainstthegenes eyeviewofevolution
1. Dawkinsgeneconcept 2. Acrucialambiguityundermining Dawkinsaccount? 3. Developmentalsystems
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Ageneisdefinedasany portionofchromosomal materialthatpotentially lastsforenough generationstoserveasaunitofnaturalselection (p.28)

Note:veryloose definitionindeed!
Nomentionofanymolecularmechanisms(such astranslation/transcriptionetc.) 'thegenemayvaryinlength
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Theevolutionarygeneconcept
Theshorterthegeneticunit,themore likelyitistosurvive(p.29) Consider
Ageneticunitofhalfthesizeofa chromosome:thereisa50%chancethatit willbesplitaftereachmeiosis Ageneticunitof1%ofthesizeofa chromosome:1%chanceofsurvivalafter eachmeiosis

Theevolutionarygeneconcept
Note:theevolutionarygeneconceptis compatiblewithnucleotidesbeinggenes!
Theyaretheshortest portionofchromosomal materialthatpotentiallylastsforgenerations Plus:theycanfunctionasunitsofselection

Butthatsofcourseabsurd;nucleotidesare notgenes
Yet,itfollowsfromDawkinsdefinition!

So:sizematters!
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Theevolutionarygeneconcept
Dawkinsreaction:
Itwouldbeabsurdtosaythatthedifferenttypes ofnucleotidescompete witheachother andthatnucleotidesofthesametypeco operatewitheachother If[nucleotides]arecompetitorsatall,theyare competitorsforeachlocusseparately.Theyare indifferenttothefateoftheirexactreplicasat otherloci.(1982,p.91)
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Theevolutionarygeneconcept
CriticismbyGriffithsandNeumann Held(1999)
Dawkinsonallelesofthesametype:no competition Dawkinsonnucleotidesofthesametype: competition Hence:Dawkinsisinconsistent!
Whyistherecompetitionb/wnucleotides ofthesametypebutnocompetitionb/w allelesofthesametype?
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Theevolutionarygeneconcept
Notealso:cooperationbetweensametypesof genes/alleles isrequiredforexplaining altruism!
Onlyifthereiscooperationatadistance betweengenescanweexplainfromthegenes eyeviewwhydifferentorganisms(withsimilar genes)cooperate

Anothercriticism:Dawkinsview isunderminedbyanambiguity

Again:whatsthejustificationforthis?
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Animportantambiguity
Soberpointsoutthatthereisanimportant distinctiontobemadebetween Selectionfor X
Xs survivebecause evolutionselectsforthem

Illustrativeexample
Selectionfor ballsize
i.e.causalselection

Selectionof ballcolour
i.e.,correlativeselection

Selectionof X
Xs survivenot becauseevolutionselectedfor them,butbecauseforsomeotherreason

E.SobercriticizesDawkinsfor notdistinguishingbetween selectionfor genesandthe selectionof genes

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Animportantambiguity
Selectionfor genes
Organismssurvivebecause theyhavecertain genes Sober:notcorrect!Rather:b/corganismshave certainproperties

Acounterexample?
Selectionfor genes=false Counterexample:sicklecellanemia andMalaria
Heterozygotes withasicklecellallele andanormalallelearemorelikelyto surviveMalariathanhomozygotes withtwonormalalleles Anemiasymptoms:minimal

Selectionof genes
Organismssurvivebecausetheyhavecertain features whichareadaptedtotheenvironment and thosefeaturesarecausedbygenes Sober:triviallytrue/widelyaccepted!
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Acounterexample?
Sober:whatsbeingselectedforis thegenotype(heterozygote),not forparticulargenes(i.e.,alleles)
Howeverthereisofcourseselection of alleles

Acounterexample?
Note:selectionof genesiscompatiblewith selectionfor individuals
Itisthevehicles/interactors thatfeeltheforceof theenvironment(i.e.,naturalselection),notthe genes Itisthevehiclesthatare,orarenot,adapted totheir environments,notthegenes Itistruethat,ultimately,evolutionischangeofgene frequenciesinthegenepoolovertime
Butthissaysnothingaboutwhy certaingenesare selected
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ReplytoSober
Sterelny andKitcher (1988)
Theyargue:ifconstruedcorrectly,Sobers alleged counterexamplecanbeturnedintoaconfirmation ofDawkinsview! Crux:theexampleturnsonthefrequency ofthe respectiveallelesinthegenepool Inotherwords:geneselectionisfrequency dependent

ReplytoSober
Asicklecellallelewillbefitter incontexts in whichthenormalalleleforhemoglobinoccurs frequently
Because:insuchcontextsthesicklecell heterozygoteismorelikelythanthesicklecell homozygote And,organismswithasicklecellalleleand a normalallelewillhaveabetterchancetosurvive Malaria Hence:thesicklecellallelewillbeselectedfor
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ReplytoSober
Yetincontexts inwhichthesicklecellalleleis frequent,thesicklecellallelewillbeunfit
Thentheorganismisatahighrisktodieofsickle cellanemia Hence:normalallelewillbeselectedfor

Developmentalsystems

Thus,itisnottruethat,asSoberclaims,the genotype,butnotgenesarebeingselectedfor
Onthecontrary,genes/allelesarebeingselected for,justasDawkinsclaims
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Geneticdeterminism?
Criticism:Dawkinsviewpresupposes geneticdeterminism
Geneticdeterminism:istheviewthatthe phenotypeisfully determinedbythe genotype

Geneticdeterminism?
NumberofeyesinDrosophila(wildtype)
Normally:between700and1100 (temperaturedependent!)

Mutations
UltrabarandInfrabar:bothconsistentlyless thanthewildtype However:theanswertothequestionwhich mutationhasmoreeyes,UltrabarorInfrabar iscontextdependent
Temperaturetoodetermineseyenumber(not justgenes)
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R.Gray,P.Griffiths,andothers:
geneticdeterminismiswrong! Environmentalconditionsarejustas importantasgenesintheexpressionofthe phenotype!

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Geneticdeterminism?
ChallengetoDawkins:
Genesdonotdeterminethephenotype Thereareotherimportantfactors(e.g. environment) Butifgenesdonotdeterminethe phenotype,andthereareotherimportant factors,then
.wecannotreallysaythatgenesarebeing selectedfor whencertainorganismssurvive andprocreate(andtherebyreplicategenes)
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Geneticdeterminism?
Question:doesDawkinsreallysubscribe togeneticdeterminism?Whatsmore, doeshisviewreallyrequireit? Dawkins:
Development oforganisms:Genetic causesandenvironmentalcausesarein principlenodifferent. Evolution:Geneticfactorsreplicate themselvesbutnongeneticfactorsdo not
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Geneticdeterminism?
Itsnottruethatonly genesare replicatedthroughoutgenerations Exampleforextragenetic inheritance: aphidBuchneraendosymbiosis
Aphidswithout Buchnera:stuntedgrowth, reproductivelysterile,dieprematurely MaternalBuchneramassisbeingpassedon Aphid=bladlus toeggsorembryo Remarkablyancientsymbioticrelationship
160 280millionyears!
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Geneticdeterminism?
AphidBuchneraexamplessatisfies Dawkinsthreecriteria
Longevity (sincewehaveinheritanceonan evolutionaryscale) Fecundity,andfidelity (numerousand accuratereplication)

Note:buchnera arenotpartofthegenetic programme ofaphids! Aphid=bladlus

ContrarytoDawkins:thewhole developmentalsystemisbeingreplicated indefinitely
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Geneticdeterminism?
Epigeneticinheritancethroughmethylation
Methylgroupsblockthetranscriptionofanygenes theyareattachedto Normallyremovedduringzygoteformationand reestablishedduringcelldivisions Methylationlevelisthoughttobeinfluencedby externalfactorslikediet

Developmentalsystems
GriffithsandGraysconclusion:
Theunitofselectionisnotjustthegenebut ratherentiredevelopmentalsystemswhich arebeingreplicated(onanevolutionary scale) Thegene isjustoneamongmanyother factorsdeterminingthephenotype
Others:environment

Hence:
Changeofgeneexpressionthroughchangeof externalfactors
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Hence,thegeneisnottheunitofselection
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TasksandQuestions
ExplainGriffithsandNeumannHelds challengeabouttheevolutionarygene ExplainSobers sicklecellcounterexampleto Dawkinsgeneeyeview ExplainSterelnyandKitchers responseto Sober Explaintheideaofdevelopmentalsystems withregardtotheunitofselectiondebate
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Lecturequery
ExplainDawkinsideaofthegenebeingthe unitofselection
Mentiontheideaofthelongevityofreplicators

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