SFIWORKSHOP:

INTEGRATINGDEVELOPMENTANDINHERITANCE13-15February2018.NoyceConferenceRoom,SFI

TitlesandAbstractsTuesday13thTobiasUllerIntegratingdevelopmentandinheritance-introductoryremarksAbstract:Anintroductiontotheworkshopanditsmainaims.(1)Todiscussthehistoricaloriginsoftheseparationofdevelopmentandinheritanceandtheirdescriptioningeneticterms,andhowthisshapedthedevelopmentofresearchprogramswithinevolutionarybiology;(2)Toexplorethepossibleimplicationsofalternativeconceptualizationsofinheritance,andthedevelopment-heredityrelationship,thatareemergingthroughrecentadvancesinthebiologicalsciences.(3)Todiscusshowempiricalworkandmathematicalmodellingbestcanproceedwithconstructiveviewsofdevelopmentandinheritance,andwhattheimplicationsmightbeforevolutionaryprocesses.ErikPetersonWaddingtondieddisappointed;or:Weren’tweintegratinginheritanceanddevelopmentinthe1930s?Abstract:Fromtheperspectiveofbiologistsworkingintheinterwarperiod,wewereonthethresholdofacomprehensivetheorycombininginheritanceanddevelopment.AmericanscionofDrosophilagenetics,ThomasHuntMorgan,authoreda1933booklayingoutjustsuchaconjunctionbetweenhisworkandembryology.Promisingdiscoveriesinbiochemistry,biophysics,andcytologyoccurredregularlyandpointedtothiskindofconjunction.Bytheearly1940s,therewasevenanameforthesubfieldthatwouldoutlinetheintegrationofdevelopmentandinheritance:“epigenetics.”But,decadeslater,theEnglishbiologistwhocoinedthatterm,C.H.Waddington,despairedthatnosuchconjunctionhadhappened.Ifanything,thefieldofbiologycollectivelyseemedlessinterestedinintegratingdevelopmentandinheritanceinthe1970sthanithadfiftyyearsearlier.Inthisessay,Itracethereasonsforoptimismaboutsuchanintegrationinthefirsthalfofthetwentiethcenturyandwhysuchanintegrationfailedtocometofruitionduringthesecondhalf.

JessicaRiskinHowthemouselostitstail,or,Lamarck'sdangerousideaAbstract:Theclockworkcosmosofearlymodernsciencewasforthemostpartapassiveandstaticthing,itsshapeimposedbyanexternaldesigner,itsmovementsoriginatingoutsideitself.Theclassicalmechanistsoftheseventeenthcenturymostlyevacuatedforceandagencyfromthecosmos,oftenevenincludingfromitslivinginhabitants,totheprovinceofasupernaturalClockmaker.Theytherebybuiltakindofsupernaturalismintotheverystructureofmodernscience.Butnoteveryoneconcurredinthisbanishment.Fromthelateseventeenthcenturyonward,atraditionofdissentersembracedtheoppositeprinciple,thatagency--acapacitytoact,tobeself-makingandself-transforming--wasessentialtonature,especiallylivingnature.Acrucialmemberofthisdissenting,active-mechanisttraditionwastheFrenchnaturalistJean-BaptisteLamarck,professorofnaturalhistoryattheMuséumnationald’histoirenaturelleinParis,andauthoroftheterm"biology"aswellasofthefirsttheoryofwhatwenowcall"evolution".Thispaperexamineshisrigorouslynaturalistapproach--whichnaturalizedratherthanoutsourcedagency--anditsexilefromthehallsofmainstreamscience.BenoitPujolAnecologistguidetodisentanglinggeneticandnon-geneticheritabilitiesinwildpopulationsAbstract:Geneticvariationforfitnessrelatedtraitsisnottheonlyheritablesourceofadaptivepotential.Itiscrucialtodisentanglenon-geneticsources(e.g.,socialinteractions,epigenetic,ecologicalnichetransmission)ofheritablevariationtorefineourestimatesofpopulationevolutionarypotential.Weprovideapracticalguideforresearchersinterestedindistinguishingbetweengeneticandnon-geneticheritabilitiesforphenotypicand/orfitnessvariation.Onethetransgenerationalmechanismisidentified,ourtutorialshowshowtoestimatesimultaneouslybothgeneticandnon-geneticsourcesofphenotypicsimilaritywithintheframeworkofthequantitativegenetic"animalmodel".Quantitativegeneticmodelscanbeusedtoquantifythegeneticvariance,andthereforetheheritability,ofcomplextraits.Thesemodelscanaccountforgeneralenvironmentalcategoricalfactorsbutouraimistogofurther.Iwilloutlinehowamatrixofsimilaritycanbeusedalongsidethegeneticmatrixofrelatednessintheanimalmodelofquantitativegeneticstocapturetheimpactoftheenvironmentalsimilarityonevolutionaryparameters.Thisisparticularlyusefulinwildpopulationswherequantifyingtheenvironmentalcomponentsoftraitscanbechallengingbecausetheenvironmentiscomposedofmanyfactors.Wewilltrialthisstatisticaltechniqueinawildpopulationofsnapdragons,Antirrhinummajus,intheSouthofFrance.Allindividualplantshavebeentagged,phenotyped,andgenotypedsince2010.Theenvironmentofeachindividualhasbeenrecorded,andtheseenvironmentalparameterswerecombinedtoformamatrixofenvironmentalsimilarity,whichwasincludedintheanimalmodelalongsidethematrixofrelatedness.WewillalsotestthisoriginalapproachonthebasisoflongtermsurveydataintheKimbeIslandwildclownfishpopulationandon

southernFrancewildroedeerpopulation.Wewelcomeanycollaborationonsimilardatasetsinordertestwhetherhabitattransmissionandothertransgenerationalnon-geneticmechanismsgeneratepopulationpotentialtorespondtodirectionalselection.RussellBondurianskyTheparentasadevelopmentaltemplateAbstract:Shiftingviewsofreproductionanddevelopmentoverthepast150yearshavedrivenprofoundchangesintheconceptofheredity.Parentswereoncethoughttobuildtheiroffspringfromtheirownsoma,providingadirectconduitfortheinheritanceofacquiredtraits.Thisconceptofreproductionwaseventuallysupplantedbytheideathatoffspringdevelopmentisguidedbyanautonomousblueprintcontaininghereditaryinformationthatcannotbealteredbyparentalenvironmentorexperience.Today,ahybridconceptofdevelopmentandheredityisemerging,withmanyresearchersrecognizingthatoffspringtraitsareshapednotonlybytheexpressionofauniquegenomebutalsobytheextendedphenotypesoftheparents.Accordingtothisview,manyparentaltraitsprovideadevelopmentalscaffoldthatshapesoffspringphenotype.Inthistalk,Iwillfocusonaparticulartypeofscaffoldingthathasintriguingimplicationsbuthasreceivedlittleattention.Intriguingevidencefromsingle-celledeukaryotesshowsthatsomecomponentsofthecellcanplayatemplate-likeroleinshapingoffspringtraits,andtherebyimpartstructuralvariationstodescendants.Iwilldiscussthepotentialforsimilarprocessestooccurinmulti-celledorganisms,andtocontributetothetransmissionoftraitsacrossgenerations.TroyDayTheroleofadaptivevariationinevolutionAbstract:TheModernSynthesis(MS)positsthatgeneticmaterialformsthemainsubstrateforevolutionaryadaptationbutthereisnowagrowingrealizationthatnongeneticformsofinformationtransmissionbetweengenerationsmightalsoplayanimportantrole.Thishasengenderedconsiderabledebateamongevolutionarybiologistsandoneofthemainpointsofcontentionisthenatureof“mutation”ingeneticvsnongeneticsystemsofinheritance.AccordingtotheMS,geneticvariationisgeneratedinanentirelyrandomfashionwhereassomeresearchershassuggestedthatnongeneticvariationmightbegeneratedinanadaptiveway.ThisclaimhasbeenparticularlyunpalatabletocriticsofnongeneticinheritancebecauseitchallengesDarwin’smostimportantinsightthatnaturalselectionisthesolemechanismofadaptiveevolution.InthistalkIwillattempttoformalizethisdebateinamoreprecisewaywiththegoaloftryingtoachieveconsensusonthisincreasinglycontroversialissue.

EvaJablonkaTheevolutionaryimplicationsofextrageneticinheritanceAbstract:Therearemanyextra-geneticinheritancetransmissionpathwaysandextra-geneticvariations,formepigeneticvariationsthatcanbedescribedinmolecularterms,totransmissiblevariationinsociallytransmittednon-symbolicalandsymbolicalbehaviorandvariationsinthereconstructionofecologicallegacies.WhattheyallhaveincommonisthatarebasedonvariationsininDNAbasesequence.Idiscusssomeoftheevolutionaryimplicationsofthese.Wednesday14thAlanLoveDevelopmentalbiologyandtheModernSynthesis:perspectivesandprospectsforintegratingdevelopmentandevolutionAbstract:Thehistoryofhowdevelopmentandinheritancepartedwaysintheearly20thcenturyiscrucialtounderstandingbothhowtheModernSynthesisemergedinevolutionarybiologyandwhydevelopmentwasnotconstitutiveofit.Althoughthegeneticformulationofinheritancehelpsinaccountingfortheformer,itislessrelevanttounderstandingthelatter.TounderstandwhydevelopmentwasnotcentrallyinvolvedintheModernSynthesis,weneedtounderstandthetrajectoryofresearchprogramsthatcharacterizedevelopmentalbiology.ThesetrajectoriesclusteraroundresearchquestionsunrelatedtoevolutionarychangeandcanberecoveredinpartthroughanexaminationofthetopicsandthemesfoundinfortyyearsoftheAnnualSymposiafortheSocietyforDevelopmentalBiology.Developmentalbiologistsconcentratedonunderstandingrecurringsimilaritiesinmodelorganismsthatweredomesticated¾bothintermsoftheirgeneticconstitutionandenvironmentalhomogeneity¾tominimizevariation.Thisanalysisshowsthatamajorbarrierbetweenevolutionarybiologyanddevelopmentalbiologythroughthe20thcenturywasdivergentinvestigativepracticesarisingfromalackofsharedproblems,whichcoalescedduringtheirdistinctiveperiodsofdisciplinaryprofessionalization.Additionally,muchofevolutionarydevelopmentalbiologyhasbeenshapedbythepracticesandproblemsofdevelopmentalbiology,reframedcomparatively,andignoresaspectsofdevelopmentandhereditygermanetoevolutionarychange.Toachieveamoreintegratedtheoreticalperspective,theproblemsandpracticesofprofessionalizedevolutionaryanddevelopmentalbiologymustbebridgedbyanintentionalarticulationoflinkagesbetweentheirdisparateresearchquestionsandmethods.Isuggestonefruitfulstrategyfordoingsoliesincoordinatingempiricalinquiryaroundtemporalstagesofalifehistorywherequestionsaboutthedevelopmentaloriginofvariationanditsevolutionaryrelevanceconverge.

KarenKovakaInteractinginheritancechannelsAbstract:Thisoffersanaccountofwhatitisforaninheritanceprocesstobeevolutionarilypowerful.Attheheartoftheaccountistheclaimthatweshouldtakeaninteractiveapproachtoassessingtheevolutionarypowerofdifferentinheritanceprocesses.Otherphilosophers(Sterelny2001;seediscussioninGriesemeretal.2005,Godfrey-Smith2011,Sterelny2011)haveproposedthreecriteria—internalpolicing,stabletransmission,andgenerationofvariation—thatanevolutionarilypowerfulinheritanceprocessmustmeet.Imodifythisproposalbyshowingthataninheritanceprocessmaymeetthesecriteriaeitherindependently,orininteractionwithotherinheritanceprocesses.Thisisanimportantdifference:theinteractiveapproachopensupspaceforustoviewextra-geneticinheritanceasevolutionarilypowerfulwhilealsoclarifyinghowandwhythereappeartobesuchdifferencesinpowerbetweengeneticandextra-geneticinheritance.LynnChiuHolobiontsasunitsofevolutionaryprocessesAbstract:Aholobiontisaspecificcollectionoflivingentities:asinglemacroorganism(usuallyamulticellulareukaryotichost)andthestably-persistentmicroorganismsthatresideinoronit(includingviruses,protists,bacteria,fungi,andsometimes,helminthes).Macroorganismshaveclearlyevolvedinthecontextofmicroorganisms,yetitishighlycontroversialwhetherholobiontshaveaspecificevolutionarystatusandthusatheoreticalroleinevolutionarytheory.ProponentsofHologenomeTheoryarguethatholobiontsarebonafideunitsofselection.Yetmanyarguethatholobiontsdonotsatisfythestringentcriteriaofclassicalunitsofselection.Inthispaper,Iarguethatholobiontsareimportantunitsofevolutioneveniftheyarenotunitsofselection.Theunitofselectiondebateaskswhetherholobiontsareevolutionarytransitions,thatis,productsofevolutionobjecttoselectiveforces.Iproposearevisednotionofevolutionaryunitsthatreframesholobiontsnotasobjectsofevolutionbutanexusofinteractingevolutionarycausesandprocessesoforganismalorigin.Inrecentdecades,thescopeofevolutionarycauseshasexpandedtoincludetheevolvingorganismsandotherorganisms,whichinfluenceevolutionthroughecologicaldevelopmentandnicheconstruction.Thedynamicholobiontissuchprocesseswritlarge.Iwillexamineaccountsofevolutionaryunitsthatalreadyintegratedevelopmentalprocesses,inparticular,JamesGriesemer’sprocessualaccountofreproducersanddevelopmentallyscaffolded“hybrids”andanalyzehowaddingnicheconstructionimpliesanevolutionaryunitofprocesses.ThenIwillarguethatholobiontsareexemplarsofsuchunits.Suchanaccountspellsouttheevolutionarysignificanceofholobionts,thussatisfyingthetheoreticalgoalsofHologenomesupporters,withoutcommittingtothemorerestrictiveviewthatholobiontsareunitsofselection.

RoseThorogoodCanculturehaveevolutionaryeffectsacrossspeciesboundaries?Abstract:Thebehaviourofotherscomprisesanimportantpartofananimal’secology-observinghowindividualsinteractwiththeirownabioticandbioticenvironmentsprovidesinformationandfacilitatesbehaviouralplasticity.Ifsocialinformationisusedindecision-making,itshouldthenhavepotentialtoinfluenceselectionacrossspeciesboundaries.However,beyondhumanculture,therehasbeenlittleconsiderationofhowsocially-transmittedbehaviourswithinspeciesmayleadtodifferentevolutionaryoutcomesinco-evolvingspecies.HereI’llpresenttwoexamplesfrommygroup’sworkwhereweareexploringtheemergentpropertiesofsocialtransmissionforevolutionaryarmsracesbetweenpredatorsandprey(usingParidtits),andparasitesandhosts(usingbroodparasiticcuckoosasourstudysystem).HalWhiteheadThereachofgene-culturecoevolutioninanimalsAbstract:Culture,sharedbehaviouracquiredbysociallearning,ispresentindiverseanimalspecies.Itiswellestablishedthatcultureaffectshumangeneticevolution.Herewereviewtheincreasingevidencethatthisalsooccursinnon-humansthroughseverallinkedprocesses.Culturemayhavedrivenearlyphasesofspeciationinbothbirdsandkillerwhales.Thereisincreasingevidencethatculturecanleadtolowdiversityofneutralgenesthroughculturalhitchhikingorculturallymediatedmigration(inmatrilinealwhalesandhumans).Culturecandrivetheadaptiveevolutionoffunctionalgenes(inkillerwhalesandbirds).Additionally,inculturalspeciesselectionshouldfavourtraitswithageneticbasisthatsupportorenhancesociallearning,includinglongerlives,lengthenedjuvenileperiods,menopause,andenhancedcognitiveabilities.Conceptionsofadaptiveevolutionshouldincludehoworganisms’culturesfacilitateadjustmenttoenvironmentalchallengesornewopportunities,dragginggeneticchangeintheirwake.KarthikPanchanathanTheevolutionofsensitiveperiodsinamodelofincrementaldevelopmentAbstract:Sensitiveperiods,inwhichexperienceshapesphenotypicdevelopmenttoalargerextentthanotherperiods,arewidespreadinnature.Despitearecentfocusonneural-physiologicalexplanation,fewformalmodelshaveexaminedtheevolutionaryselectionpressuresthatresultindevelopmentalmechanismsthatproducesensitiveperiods.I’llpresentanevolutionarymodelofdevelopmentI’vebeenworkingonwithcolleagues.Wemodeldevelopmentasaspecializationprocessduringwhichindividualsincrementallyadapttolocalenvironmentalconditions,whilereceivingaconstantstreamofcost-free,imperfectcuestotheenvironmentalstate.Wecomputeoptimaldevelopmentalprogrammesacrossarangeofecologicalconditionsandusetheseprogramstosimulatedevelopmentaltrajectoriesand

obtaindistributionsofmaturephenotypes.I’lldiscussboththemodellingframeworkandsomeoftheresultsofthemodel.Thursday15thSeanRiceEvolutionaryconsequencesofinteractionsbetweendevelopment,inheritance,andselectionAbstract:Inheritanceisthekeyfactormakingbiologicalevolutionpossible.Despitethiscentralrole,transmissionisoftenbundledintothesimplifyingassumptionsofevolutionarymodels,makingitdifficulttoseehowchangesinthepatternsoftransmissioninfluenceevolutionarydynamics.Iwilldiscussanewmathematicalmethodthatallowsustoderivepatternsofinheritancefromdevelopmentalprocesses,andtoincorporatetheresultsintomodelsofevolution.Iwillillustratetheapproachwithanexampleinwhichanenvironmentalfactorinfluencesbothinheritance(throughitsimpactondevelopment)andfitness.Ishowthattheresultingcorrelationbetweeninheritanceandselectioncanhaveapronouncedimpactonevolutionarydynamics.MichaelLachmannThethriftywatchmakerAbstract:InmytalkIwillexplorethetypesofvariationthatevolutiongetstotinkerwith.Iwillre-examinepointsmadeaboutplasticitybefore,byWolff,West-Eberhardandothers.Iwill,however,claimthatgenesandenvironmentshouldnotbeseenasactingonanevenfooting.InsteadIwillarguethatthegenotypeshouldbeseenasaninterpreteroftheenvironment,andthatmostoftheinformation,andevenmostofthefunctionalinformationinthephenotypecomesfromtheenvironment.Neuronallearningguidingbehaviorshouldnotbeseenasanoutlierbutinsteadasthenormofdevelopmentalprocesses.MarcFeldmanOntheevolutionofculturaltransmissionandfitnessoptimizationAbstract:Westudycompetitionbetweenverticalandobliqueculturaltransmissionofadichotomousphenotypeunderconstant,periodicallycycling,andrandomlyfluctuatingselection.Conditionsarederivedfortheexistenceofastablepolymorphisminaperiodicallycyclingselectionregime.Undersuchaselectionregime,thefateofageneticmodifieroftherateofverticaltransmissiondependsonthelengthofthecycleandthestrengthofselection.Theevolutionarilystablerateofverticaltransmissiondiffersmarkedlyfromtheratethatmaximizesthegeometricmeanfitnessofthepopulation.