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ColinPittendrighinhisfinalyearsatPrincetonUniversity
ThemanuscriptassociatedwiththeseremarksisanabbreviatedversionofmyPh.D.dissertation,earnedin1969undertheguidanceofColinPittendrigh(Pitt)atPrincetonUniversity.Themanuscriptwasnotpublished,ormoreaccuratelywasrejected,bySciencemagazinein1976.IchosethesubjectbecauseIwasinterestedintheecologyoffliesofthegenusDrosophila,andIwantedtogenerateevidencethatnaturalenvironmentalconditionsareasourceofselectionforthewellknowncircadianeclosionrhythmoftheseanimals.AllspeciesofDrosophilatestedhaveacircadianeclosionrhythmthatcausesadultfliestoemergefromthepupalcaseonlyinthefirstfewhoursafterdawn.IselectedonespeciesofDrosophilanativetothewesternU.S.,andasecondspeciesnativetotheeasternU.S,andexposedbothtonaturalandextremesaturationdeficitregimes.Saturationdeficitisameasureoftheevaporativepoweroftheatmosphereataparticulartemperatureandrelativehumidity.IwantedtofindoutifDrosophilainnaturemoresuccessfullyecloseintheearlymorningbecausethatisthedaylighttimewhenthesaturationdeficitisusuallyataminimum.Ifoundthistobetrueforbothspecieswhenexposedtotheirnaturalenvironment.Inthelaboratory,Ifoundapositiverelationshipbetweeneclosionsuccessandlowsaturationdeficits.Pittsteadilyencouragedmeinthiswork,reviewedthemanuscriptandmadehelpfulsuggestionsbeforeIsubmitteditforpublication.IarrivedatPrincetoninthefallof1965,eagertomeetmynewscientificmentor,ColinPittendrigh.Ihaddevelopedaninterestincircadianrhythmsasanundergraduate,anddecidedhewasthepersonIwishedtoworkwith.Icloselywatchedandlearnedfromhim,themostimportantpersoninmyprofessionaldevelopment.Asithappened,PitthadjustbecomeDeanoftheGraduateSchool,amajoradministrativejobwithofficesinhistoricNassauHall,farremovedfromhislabphysicallyandpsychologically.Hislabwasverybusy,withseveralgraduatestudents,postdocs,andfull-timelaboratoryassistantsandtechnicians.Thesuiteofroomsin19thCenturyGuyotHallwasabeehiveofscientificactivity,orperhapsIshouldsayaflyswarm.Muchoftheresearchoncircadianrhythmsrevolvedaroundoneorganism,Drosophilapseudoobscura,awesternNorthAmericanspeciesoffruitfly.AmongthestudentsinthelabatthattimewerethelateArthurWinfreeoftheUniversityofArizona,amathematicalbiologistandMacArthurFellowwhowroteTheGeometryofBiologicalTime,andJerryFeldmanofSwarthmoreCollege,whohaspublishedmanypapersonmolecularcircadianclocks.Therewereoftenvisitorstoourlaboratory.Somewerecolleaguesfromotheruniversities.Onoccasion,therewouldbereportersandfilmcrewsinterviewingPittonsubjectsasdiverseasthepossibilityofextraterrestriallife,andfederalsupportofscientificresearch.WewereoncevisitedbyJürgenAschoff,whotogetherwithPittandErwinBünninglaidthegroundworkformodernstudiesofchronobiology.Aschoffmetindividuallywitheveryresearcherinthelab,andlistenedwithinterestasweeachexplainedourwork.Myfirstresearchideawasahypothesisthattheblindcavedwellingcrustacean,Orchonectespellucidus,longisolatedundergroundinMammothCave,Kentucky,wouldhaveretainedcircadianrhythms.Iplannedtobringanimalsintoconstantlaboratoryconditions,andmeasuretheircyclicalphysicalandphysiologicalfunctions.Ispentmyfirst
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yearplanningthatproject,onlytodiscoverthatgraduatestudentThomasJeglahadjustcompletedthoseveryexperimentsforhisdissertationatYale.Hepublishedhisresults,showingthatthecrayfishdoexpressfreerunningcircadianrhythmsinconstantconditions,intheJournalofExperimentalZoologyin1968.Ihadtostartoverwithanotheridea.Pittasalwayswassupportive,andwebegantodiscussresearchalternatives.IformulatedaquestionthatwoulduseD.pseudoobscura.Iwantedtomakeupforlosttime,andthischoicemeantthatIcouldtakeadvantageofequipmentandproceduresalreadyinuseinhislaboratory.Iwasoffandrunning.Myexperimentaldesignrequiredexposingflypupaetonaturalenvironmentalconditions.IdidthisnearPrincetonusingasmalllaboratorybuiltformeinsideacenturyoldfarmhousebarnonpropertyownedbytheUniversity.Thisfieldlabmusthavebeenacostlyconstructionproject.IdonotknowbecauseafterIaskedaboutit,inshortorderitwasbuiltformeaspermyspecifications.ItwasPitt,ofcourse,whoarrangedandpaidforthat.Henevertoldmehow.ThefollowingyearIneededtotakemyexperimentalfliestoawesternU.S.climate,wheretheynaturallyoccur,tofindouthowpreciselythewellstudiedcircadianeclosionrhythmexhibitedinourPrincetonlaboratoryisentrainedbynaturaltemperatureandlightcycles.PittsentmetotheRockefellerUniversityinNewYorkCitytoconferwithTheodosiusDobzhansky,atthetimethegrandoldmanofevolutionarygenetics.PittpointedouttomethatDobzhanskywasmyintellectualgrandfather,sincehehadbeenPitt’smajorprofessoratColumbia.Pittreferredtohimas“theoldboy.”DobzhanskyhadbeenworkingformanydecadeswithDrosophila,andhaddonesointheAmericanWest.Thefirstthinghesaidtomewas“Sohereistheyoungmanwhodoesnotsignhisletters.”Ihadtypedmyletterstohim,andinmycallowyearsIdidnotrealizeitwasgoodformtoincludeasignature.Hemadethisremarkwithakindlysmileonhisface,becauseDr.Dobzhanskywasamarkedlycourteous,indeedcourtly,personwhogreetedmeasacolleague.HelistenedwithinterestasIexplainedmyresearch,andsoughthisadvice.InhispronouncedRussianaccent,headdressedmeas“MeesterQuinn.”Heshowedmearoundhislab,whereanumberofresearchers,whowerefamiliartomeonlyfromtheirpublications,explainedtheirresearchprograms.Dobzhanskytheninvitedmetopresentaseminaronmyresearchtohisassembledlaboratorycolleaguesandstaff.Iaskedhimwhen,expectingalaterdate,andhereplied“thisafternoonatfour,”twohourslater.HowcouldIsayno,eventhoughIwascompletelyunpreparedandquitenervousaboutanaudiencewhohadbeenfamouslystudyingDrosophilafordecades?Isomehowgatheredmywitsandmadeacoherentpresentation,witharmwavinginplaceofslides.AfterwardsDr.DobzhanskyinvitedmetodinnerwithhimandhispostdocsattheRockefellerUniversity,aplacewheremenwererequiredtodressfordinnerinacoatandtie.Icamethatdaydressedasahumblegraduatestudent.Thegooddoctorarrangedforoneofhispost-docs,atallman,toloanmethepropergarb,soIateandconversedwiththesleevesofmycoatjacketrolledup.Veryelegant.Dr.DobzhanskyrecommendedthatIdomyfieldworkattheSouthwesternResearchStation,afacilityinArizonaoperatedthenandnowbytheAmericanMuseumofNaturalHistory.Hehadworkedthere,wherehewasrememberedfortravellingaroundtohisfieldsitesonhorseback.TheBiologyDepartmentpaidformytravelexpenses,butnotforahorse,andPitthadtheexperimentalmachineryIneededbuiltandshippedtoArizona.
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ColinPittendrighhadoneofthemostremarkablescientificmindsthatIhaveeverencountered.Hegraspedcomplexideasimmediately,andoftenforesawtheirimplicationsforresearchjustasquickly.Ialwayscarefullypreparedformymeetingswithhiminhisdistantadministrativeoffices,becauseIknewthathecouldprocessandanalyzewhateverIsaidasquicklyasIcouldexplainmyself.Afterourmeetings,Iwouldsitdownandwriteouteverythinghehadsaid,whileitwasfreshinmymind.Hisglitteringverbalandanalyticalabilitieswereequallymatched.Whenhespokeformallyhewaselegant,andheknewhowtotailorhisremarkstohisaudience.Heenjoyedexplainingevolutioninthegeneralsense.IsawhimdothatatapubliclectureinthePrincetonPublicLibrary.Iwastakenbyhislecturebecausehestatedhisideassoclearlyforaverygeneralaudience.Onanotheroccasion,Isawhimexplainevolutionaryprinciples,inonehour,toaverylargeundergraduateclass.Hisremarksweresoengagingthatwhenheconcludedtheentireclassstoodandgavehimastandingovation.ThatwastheonlytimeIsawthathappeninanyPrincetonundergraduateclass.AsIwaspushingtowardthecompletionofmydissertationinthespringof1969,IlearnedthatPittwasleavingPrinceton,aswasI.HemovedtoStanfordUniversitywherehecontinuedhisresearch,andassumedanewsetofimportantadministrativeduties.Heofcoursecontinuedhisscientificresearch.Heoncetoldmethathewasconcernedthathewaslivingonhis“scientificcapital,”implyingthathisadministrativeresponsibilitieswerecausinghimtoneglectscientificresearch.IdidnotseethathappenduringhisfinalPrincetonyears.IfeelextremelyfortunatetohavestudiedunderPitt.Hewassupportive,andbyhisexamplehehelpedmelearntothinkandworkasascientist.RonaldD.QuinnPh.D.PrincetonUniversity,1969Dr.QuinnisProfessorEmeritusfromtheDepartmentofBiologicalSciencesatCalPolyPomona(akaCaliforniaStatePolytechnicUniversity,Pomona)
