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OurHistory1PhysiologyUNIVERSITYOFTORONTOPHYSIOLOGYourhistory PHYSIOLOGYOurHistoryWrienbyProfessorsHaroldL.AtwoodandMladenVranicEditorProfessorStephenG.MahewsDesignEvaEngcopyright2012DepartmentofPhysiologyPhysiologyUNIVERSITYOFTORONTOPASTCHAIRSOFTHEDEPARTMENTOFPHYSIOLOGYR.RamsayWright1887-1892ArchibaldByronMacallum1891-1908ThomasG.Brodie1908-1916J.J.R.Macleod1918-1928CharlesH.Best1929-1965ReginaldE.Haist1965-1975JohnT.Murphy1975-1980HaroldL.Atwood1981-1991MladenVranic1991-1995JohnR.G.Challis1995-2000JohnF.MacDonald2001-2008 OurHistory1AMessagefromtheChairIamdelightedtointroducetoyouPhysiologyOurHistory.ThefollowingpagesdescribethehistoryoftheDepartmentofPhysiologyoverthepast125years.AsyouwillreadtheDepartmentofPhysiologyhasalongandillustriouspastinresearchandeducationincludingthediscoveryofinsulin.FurthertheDepartmentofPhysiologyawardedtherstPhDattheUniversityofToronto.ThiscomprehensiveaccountwaswrittenbytwoformerChairsoftheDepartmentProfessorHaroldAtwood1981-1991andProfessorMladenVranic1991-1995.IwouldliketothankthemfortheirtremendouseffortandmemoriesTodayweareoneofthelargestBasicScienceDepartmentsintheFacultyofMedicineandoneofthelargestPhysiologyDepartmentsinNorthAmerica.Wearecommittedtoexcellenceinbothourresearchandeducationprograms.Therearecurrentlyover120Facultyandalmost300GraduateStudentsandPost-DoctoralFellowsorganizedinfourmajorresearchplatforms-CardiovascularandRespiratorySciences-EndocrinologyandDiabetes-Neuroscience-ReproductionandDevelopmentIn2008wewereabletocapturealloftheChairsofPhysiologyfromthepast30yearstogetherinaphotographbackcover.AllremainmembersoftheDepartmentofPhysiologytoday.ItisagreatprivilegetobeChairofsuchanoutstandingDepartment.Ihopethatyouenjoyreadingtheaccountofourpast125years.IfyouwouldliketondoutmoreabouttheDepartmentofPhysiologyattheUniversityofTorontopleasevisitourwebsitewww.physiology.utoronto.ca.StephenG.MatthewsPhDErnestB.andLeonardB.SmithProfessorandChairDepartmentofPhysiology 2DepartmentofPhysiologyABriefHistoricalAccountoftheDepartmentofPhysiologyUniversityofTorontoHaroldL.AtwoodandMladenVranic2008TheDepartmentofPhysiologyisbestknownasthesiteofthediscoveryofinsulinanditstherapeuticvaluesin1921andshortlythereafter.Thisworkledtooneofthemostdramaticmedicaladvanceseverrealizedandtoprolongationandimprovementofthelivesofmillionsofpeoplesufferingfromdiabetes.ThediscoverywasrecognizedbytheawardoftheNobelPrizeinPhysiologyorMedicineCanadasfirsttoF.G.BantingandJ.J.R.Macleod.BothbeforeandafterthislandmarkeventtheDepartmentofPhysiologyfosteredastrongrecordinresearchandeducationwhichcontinuestothepresentday.TheDepartmentremainspositionedtocontributeinmanywaystoadvancingknowledgeandeducation.TheDepartmentscoursewassetbyitsearliestleaderswholedbyexample.Theyestablishedastrongtraditioninresearchandteachingandcreatedthephysicalandintellectualenvironmentthatmadethediscoveryofinsulinandsubsequentresearchpossible.TheirsuccessorshaveendeavouredtoremainoncoursekeepinginviewthetwomainobjectivesoftheDepartmentalmissionresearchandeducation. OurHistory3EarlyDevelopmentoftheDepartmentofPhysiology1887-1921TheearlyevolutionofboththeDepartmentofBiologyanditsoff-shoottheDepartmentofPhysiologywasguidedbyRobertRamsayWrightaScotsmanwhohadastrongOxfordinfluencesuperposeduponhisEdinburghoriginandeducation.HecametoTorontoin1874attheageoftwenty-twofromanassistantprofessorshipintheDepartmentofZoologyoftheUniversityofEdinburgh.HewasappointedProfessorofNaturalHistoryattheUniversityofTorontoatitlelaterchangedtoProfessorofBiologyandPhysiology.In1887whenthenewMedicalSchoolwasestablishedRamsayWrightdirectedPhysiologyforfiveyears.ThusitwaslargelyunderhisdirectionthattheDepartmentsofBiologyandPhysiologytookshapeattheendofthe19thcenturyandthebeginningofthe20thcentury.Heexcelledinthethreeimportantarenasofacademiclifeteachingresearchandadministration.Hewasanaccomplishedandpopularlecturerwhohadtheremarkableabilitytomaketwodifferentdrawingsontheblackboardsimultaneouslyonewithhisrighthandandtheotherwithhisleftanddidnotscorntowintheapplauseofastudentaudiencebydisplayingthisunusualaccomplishment.1HeplacedhighimportanceonthepropertrainingoffutureresearchscientistsinlaboratoryworkandtogetherwithProfessorJamesLoudonlaterPresidentheestablishedadvancedfacilitieswithintheUniversityofTorontoforthefutureofpracticalscienceinstruction.BiologicallaboratorieswereestablishedfirstintheSchoolofPracticalScienceandlaterintheBiologicalBuildingandaBiologicalMuseumwasaddedasanadjuncttopracticalteaching.ThiswasanimportantpartofhislegacyattheUniversityofToronto.RamsayWrightwasanavidproponentoforiginalresearchandstatedthattheuniversityhasahigherfunctionthantheeducationofitsownundergraduatesnamelytheadvancementaswellasthediffusionoflearning.2Hisresearchpaperswerediversecoveringaspectsofanthropologycomparativevertebrateanatomyparasitologyandprotozoologyhiscontributionstotheanatomyandbiologyoffisheswereparticularlysignificant.HesupportedtheviewsofCharlesDarwinonevolutionwhichwerenotuniversallyacceptedbythebiologistsofthatera.Inadditiontohisresearchinterestshedevotedtimetolanguagesclassicalstudiesandmusicinallofwhichheexcelled.Athumb-nailsketchofthisinterestingandinfluentialacademicpioneerhasbeenleftbyUniversityofTorontozoologistA.G.HuntsmanCoupledwithhisthoroughnessinscholarshipandhiskeennessandvisionindiscoveringnewthingswasamostunusualpowerofclearstimulatingandindeedbrilliantexpositionofhissubjectwhichattractedstudentsofeverycharacter.HislecturestostudentswhohadjustenteredtheUniversitywerewonderfulinmatterandpresentationandsuperblyillustratedwiththenewestmaterialsandmethods.Themuseumwhichhestartedinthebuildingerectedforhisdepartmentin1889developedintoperhapstheforemostinstructionalmuseumintheworldforitstime.HisinfluencewasanoutstandingfactorinthedevelopmentofthebiologicalsciencesinthiscountrythroughoutwhichandR.RamsayWright1887-1892 4DepartmentofPhysiologyintheUnitedStatesaswellhisstudentshavebeenscattered.TheinfluencewhichthisbrilliantteacherandinvestigatorexercisedinhisownDepartmentofBiologyintheUniversityofTorontoandinthecloselyrelatedFacultyofMedicinemakeshimoneofthemostmemorablefiguresinthehistoryofCanadianeducation.3InfactitwasoneofRamsayWrightsstudentsArchibaldByronMacallumwhobecamethefirstheadofaseparatedepartmentofPhysiology.RaisedonanOntariofarmheobtainedhisundergraduatedegreefromtheUniversityofTorontoandhisdoctoraldegreefromJohnsHopkinsUniversityinBaltimore.UniversityofTorontohistorianRobinHarriswritesMacallumsgreatestcontributiontotheUniversityofTorontowastheestablishmentofresearchandscholarshipasessentialfunctionsoftheUniversity.Inthe1880swhenhefirstjoinedthefacultytheUniversitywasareputableteachinginstitutionconcernedwiththepreservationanddisseminationofknowledgebutnotwithitsadvancement.HewascertainlyamongthemostdistinguishedscientiststhatCanadahasproduced.4MacallumwasinitiallyappointedbyRamsayWrightashisfirstlecturerinBiologyin1883anddesignatedlecturerinPhysiologyin1887.InduecourseaseparateDepartmentofPhysiologyemergedfromtheDepart-mentofBiologyin1891withMacallumnowProfessorasitshead.HisappointmentwastransferredtothereconstitutedandrenovatedFacultyofMedicineandthecompletionofanewMedicalBuildingin1904madeitpossiblefortheDepartmentofPhysiologytoberemovedfromitsbirth-placeintheBiologicalBuilding.Bothofthesebuildingsweredismantledinthelate1960stomakewayforthenewMedicalSciencesBuildinginwhichthepresent-dayDepartmentofPhysiologyislocated.MacallumexcelledinresearchdevelopmentofacademicinstitutionsandinmentoringfuturescientistsbutunlikeRamsayWrighthewasnotaninspiringinstructorforundergraduates.Workawayatresearchitistheonlythingthatgiveslastingsatisfactionwashismotif.Indeedhisresearchworkreceivedinternationalacclaimparticularlyhisstudiesofkidneyfunctionandbloodcompositionshowingthatanimalsandthereforehumanshadtheirevolutionaryoriginsintheocean.HereceivedhonorarydegreesfromAberdeenMcGillTorontoTrinityCollegeDublinandYaleandbecamethefirstUniversityofTorontograduatetobeelectedtotheRoyalSocietyofLondon.In1908aChairofBiochemistrywasestablishedforMacallumattheUniversityofTorontoonlythesecondofitskindtobeestablishedintheBritishEmpire.In1917hemovedtobecomeChairmanofBiochemistryTheBiologyBuildingonQueensParkCrescentociallyopenedin1889.Itwasdemolishedin1966tomakewayfortheMedicalSciencesBuilding. OurHistory5atMcGillforafurthereightyears.HewasalsothefirstChairmanoftheNationalResearchCouncilofCanada.MacallumsimpactonthedevelopmentoftheDepartmentsofPhysiologyandBiochemistrybothatTorontoandMcGillwasenormous.HefoughthardandcompletelysucceededintransformingtheUniversityofTorontosMedicalSchoolintooneoftheleadingschoolsinNorthAmerica.HewasalsolargelyresponsibleforafundamentalchangeintheviewoftheUniversityasprimarilyateachinginstitution.Macallumslegacywastheestablishmentofresearchasanindispensiblecomplementinalldepartments.Thiswaspromotedbythedevelop-mentofgraduatestudiesatthedoctorallevel.PhysiologyledthewayinthisdirectionMacallumsupervisedthefirststudenttoobtainhisPhDattheUniversityofTorontoFredrickH.ScottPhD1900.HealsoappointedthefirstwomanlecturerintheUniversityClaraBensonwholaterbecameafoundingmemberoftheFacultyofHouseholdScience.MacallumsmonumentistheMedicalSchoolofTorontostatedhisRoyalSocietyobituary.ThroughthegenerosityoftheMacallumfamilytheannualMacallumLectureshipwasestablishedintheDepartmentofPhysiologyin1976.4ThomasG.Brodie1908-1916FollowingMacallumsassumptionoftheChairofBiochemistryin1908thenotedBritishphysiologistThomasG.BrodiewasrecruitedasheadofPhysiology.HeledtheDepartmentuntilhisuntimelydeathatage50in1916.BrodiehadbeeneducatedattheUniversityofCambridgeandhadstudiedmedicineinLondon.BeforecomingtoTorontohehadheldpositionsintheDepartmentofPhysiologyinKingsCollegeLondonLondonHospitalMedicalSchoolandatSt.ThomassHospitalMedicalSchool.At36hebecameamemberoftheRoyalSocietyofLondon.HewasconsideredoneoftheleadingexperimentalphysiologistsoftheEnglish-speakingworld.InTorontohepublishedresearchonthephysiologyandpathologyofthekidney.DuringtheFirstWorldWarhecontributedstudiesonthephysiologicaleffectsofwoundsontherespiratorysystem.HiscontributionshavebeensummedupbyMacallumDr.BrodiebegantodeveloptheDepartmentalongbiophysicallineswiththeresultthatathisdeaththelaboratoryrankedamongthethreeleadinglaboratoriesofphysiologyoneithersideoftheAtlantic.Sincehistimephysiologicalbiophysicshasbeenimportantinresearchrelatedtoneurophysiologycardiovascularandrespiratoryphysiologyanddynamicsofmetabolism.4ArchibaldByronMacallum1891-1908 6DepartmentofPhysiologyJ.J.R.MacLeod1918-1928FollowingthedeathofBrodieanothereminentBritishphysiologistJ.J.R.MacleodbecameheadoftheDepartmentofPhysiology1918-1928.MacleodhadfinishedMedicalSchoolattheUniversityofAberdeenandpost-graduatestudiesinPhysiologyinLeipzigGermanyandinAberdeen.HehadheldpositionsinPhysiologyandBiochemistryattheLondonHospitalMedicalSchoolandatCaseWesternReserveUniversityinClevelandwhereheservedasheadofthePhysiologyDepartment1903-1918.WhenhecametoTorontohewasanexpertinthefieldofcarbohydratemetabolismandhaddoneextensiveworkonfactorsthatregulatebloodglucoseconcentrationincludingtheroleofthenervoussystemdietaryfactorsandtheadrenalglandinliverglycogencontrol.Healsostudiedabnormalitiesofglucoseutilizationindiabeticdogs.Ofparticularimportancein1915hepublishedapaperentitledRapidandaccurateclinicalmethodfortheestimationofsugarinsmallquantitiesofblood.EstablishmentofthismethodologyintheDepartmentofPhysiologywascrucialintheworkleadingtothediscoveryofinsulin.MacleodgainedhisinternationalreputationnotonlythroughhisoriginalpublicationsinthefieldofcarbohydratemetabolismbutalsothroughthepublicationofaclassictextbookPhysiologyandModernMedicine.WiththeexperiencegainedinleadingBritishandAmericanUniversitiesMacleodwasinapositiontoknowwhatconstitutedastate-of-the-artDepartmentofPhysiologyandheworkedtobringTorontosDepartmentofPhysiologytothelevelofotherleadingdepartmentsthroughouttheworld.4THEGREATEVENTSOF19211923ThediscoveryofinsulinintheDepartmentofPhysiologyanditsrapiddevelopmentinTorontoasatherapeuticagentfortreatmentofdiabeteswasacollaborationinitiatedbyDr.FrederickBantingamedicaldoctorpracticinginLondonOntario.HeapproachedJ.J.R.Macleodwiththeideaoftryingtoisolateanactiveagentusingdogsasexperimentalsubjects.MacleodadvisedanddirectedtheworkprovidedthefacilitiesforitsexecutiontheservicesofaresearchstudentCharlesH.BestandlatertheexpertassistanceofabiochemistJamesCollipwhofirstsuccessfullyisolatedinsulininaformthatcouldbeusedinhumanpatients.ThedetailsofthediscoveryandthesubsequentstrugglesamongstthecollaboratorsforgloryandforcontroloftheproductionanduseofinsulinhavebeenfullyexploredintwobooksbyUniversityofTorontohistorianMichaelBliss5andmorebrieflybyUniversityProfessorMartinJ.FriedlandinTheUniversityofTorontoAHistory2fromwhichabriefexcerptispresentedBantingcametoTorontointhesummerof1921andwasgivenfacilitiesonthetopfloorofthemedicalbuildinganduseofMacleodstwosummerresearchassistants.TheresearchassistantsapparentlyflippedacointoseewhowouldstartworkingwithBanting.CharlesBestwonthetoss.The21-year-oldBeststartedhis OurHistory7JamesCollipphysiologistonleavefromtheUniversityofAlbertaworkwithBantingthedayafterfinishinghislastexaminthehonoursprograminphysiologyandbiochemistry.BantingandBestworkedassiduouslyoverthesummertryingtoextractinsulinfromdogsandachievingpromisingpreliminaryresults.TheycontinuedtheirworkinthefallwithinvolvementfromMacleodandwithhelpfromaprofessorfromtheUniversityofAlbertaJamesCollip.CollipwhohadcompletedhisPhDin1916underthesupervisionofA.B.MacallumwasonleavefortheyeartoworkwithMacleod.Collipstaskwastopurifythepancreaticextractsothatitmightbequitesafeforuseintherapeutictrials.Thefirstsuccessfulinjectionofinsulinwasadministeredtoanemaciated14-year-oldcharitypatientLeonardThompsonattheTorontoGeneralHospitalonJanuary231922withspectacularresults.InOctober1923itwasannouncedthatBantingandMacleodhadbeenawardedtheNobelPrizeinMedicine.BantingwasincensedthathewastosharetheprizewithMacleodandnotwithBestandhadtobedissuadedfromturningitdown.HewasthefirstCanadiantowinaNobelprize.HesplithissharewithBestasdidMacleodwithCollip.Therehasbeenconsiderablecontroversyovertheyearsastowhoshouldhavereceivedtheprize.Bliss5rightlytakesthepositionthatallfourdeservecreditforthediscoveryInsulinemergedin1921-22astheresultofcollaborationamonganumberofresearchersdirectedbyJ.J.R.MacleodwhoexpandeduponandcarriedtotriumphantsuccessaprojectinitiatedbyBantingwiththehelpofBest.ThesinglemostimportanttechnicalachievementwasthatmadebyCollipinthepurificationoftheextract.OntheirownBantingandBestwouldprobablynothavereachedinsulin.Inhis1922annualreportPresidentFalconerwasdiplomaticallycarefultogiveallfourcreditThisremarkableandnowwidelyknowndiscoveryhasbeenduetothepersistentandimaginativeeffortsofDr.BantingtogetherwithMr.BestandDr.CollipaidedbyotherschieflyinthePhysiologicalLaboratoryinwhichDr.Macleodputhiswideknowledgeandexperienceattheirdisposal.Thediscoveryofinsulinwasofcourseofimmensebenefittothosesufferingfromdiabetes.Onepatient6-year-oldTedRyderweighedonly26poundswhenhefirstreceivedinsulininJuly1922.In1990inhisseventiesheattendedtheceremonyattheUniversityofTorontoatwhichaplaquewasunveiledatthemedicalschoolcelebratingthediscoveryofinsulin.FollowingthesuccessfulestablishmentofinsulintherapyBantingreceivedlargeamountsofmoneyforresearchbutwasnotsuccessfulinachievingothernoteworthydiscoveries.BantingsonlysuccessattheUniversityinthesepost-insulinyearsBlisspointedlynoteswasincontinuingtoblackenthereputationofJ.J.R.Macleod.By1927MacleodhadhadenoughheandBantingapparentlydidnotspeaktoeachotheranddecidedtotakeapositionasprofessorofphysiologyathisalmamatertheUniversityofAberdeen.2 8DepartmentofPhysiologyCharlesH.BestasHeadofPhysiology1929-1965CharlesBestcompletedamedicaldegreeatTorontoandwenttotheUniversityofLondonforadoctorateandthenreturnedtoteachandconductresearchatToronto.AfterMacleodleftforScotlandin1928Bestwasappointedtothechairofphysiologywherehecontinuedtodogoodwork.OneofhisearlysuccesseswasinpurifyingtheanticoagulantheparinwhichwasfirstusedinclinicalpracticebythesurgeonGordonMurrayattheTorontoGeneralHospitalin1935.Heparinmadepossiblesuchlaterproceduresaskidneydialysisandopen-heartsurgery.2Healsoconductedgoodresearchonhistaminecholineasadietaryfactoranddevelopmentofinfusionmediaforwarcasualtiesandhecontinuedtopromoteresearchoninsulinappointinganumberoffacultymembersinendocrinologyandattractingoutstandingpost-doctoralresearchersfromabroadsomeofwhombecamefacultymembersinduecourse.InadditiontoresearchCharlesBestcontributedgreatlytomedicaleducationputtingtheDepartmentofPhysiologyinthevanguardwithhisoutstandingtext-booksTheHumanBodyandItsFunctionsBestandTaylor1932andThePhysiologicalBasisofMedicalPracticeBestandTaylor1937.Thelatterbookwentthroughmanyeditionsuptothepresentandhasbeentranslatedintomanylanguages.BestreceivedmanyhonoursforhiscontributionsincludingelectiontotheRoyalSocietyofLondonForeignAssociateoftheNationalAcademyFellowoftheCanadianandBritishRoyalSocietiesCompanionoftheOrderofCanadaand25honorarydegrees.OnBantingsdeathin1941attheageof49BestwasappointedtothechairofmedicalresearchattheUniversity.AseparateresearchinstitutehadbeenestablishedandBestbecameitsdirectoraswellasbeingheadofPhysiology.In1950theBestInstitutewasconstructedbesideandconnectedtotheBantingInstituteonCollegeStreetandforsomeyearstheDepartmentofPhysiologyhaditsresearchfacilitiesinthatbuilding.FrederickBanngrightandCharlesBestontheroofofthemedicalbuildingwithoneofthediabecdogsusedintheirexperi-mentswithinsulin.Picturethoughttobefromthesummerof1921butdatedApril1922byBannginhisscrapbook.BanngandBestslaboratoryonthesecondoorofthemedicalbuildingCharlesH.Best1929-1965 OurHistory9DuringtheSecondWorldWartheDepartmentofPhysiologycontributedresearchinseveralrelevantareasincludingtreatmentofwarinjurieshypothermiaandnightvision.Somemembersservedoverseas.OneofthesewasDr.JacobMarkowitzwhoattainedhisMDin1923andPhDinPhysiologyattheUniversityofTorontoandhadworkedonexperimentalsurgeryunderthedirectionofDr.CharlesBest.HewasafacultymemberoftheDepartmentofPhysiologyfrom1947tohisretirementin1967andProfessorEmeritusuntilhisdeathin1969.DuringtheWarhewasamemberoftheRAMCRoyalArmyMedicalCorpswastakenprisonerbytheJapanesearmyduringthefallofSingaporeandwassenttotheisolatedPOWcampatChungkai60milesnorthwestofBangkok.Theremortalityrateswerehighwith165000bothcaptivesandJapanesesoldiersdyingfromthelabourdiseaseandmalnutritioninoneyear.TheRAMCcaptiveshadthedifficulttaskofcaringforthesickundertheseconditions.Theycontendedwithcholeramalnutritionandwithtropicalulcersandinfectionswhichoftenrequiredlimbamputationstosavealife.Amputationsweredoneusingboiledhacksawstocutthroughthebonetincansformuscleretractorsandcottonthreadforstitchingwounds.Onlysmallquantitiesofdisinfectantsandanaestheticswereavailablesometimesobtainedfromguardsforabribe.Alegamputationwasdonein12to20minutes.Onehundredandfifteenthighamputationswereperformedonprisonersandabout50ofthemsurvivedtotheendofthewar.Dr.Markowitzandhiscolleaguesarecreditedwithsavingover2000lives.InlateryearshewashonouredwiththeOrderoftheBritishEmpireforhisefforts.ClemsonUniversityU.S.A.institutedtheJacobMarkowitzAwardforoutstandingresearchinsurgery.6ReginaldR.HaistChairman1965-1975ReginaldHaistwasaseniormemberoftheDepartmentwhobecameChairin1965whenBeststeppeddown.HisareaofexpertisewasendocrinologyhehadworkedandpublishedwithCharlesBestandheappointedseveralfacultymembersintheareaofendocrinologyincludingMladenVranicwhowouldlaterinturnbecomeChairoftheDepartment.Theearly1960ssawthepost-SputnikexpansionoffacultythroughouttheUniversityandHaistwasabletoappointfacultymembersintheareaofneurophysiologyinfacthebecameconvincedthatthiswasanareainwhichtheDepartmentshouldbestrengthened.ThisdecisionhadamajoreffectonsubsequentdevelopmentoftheDepartment.DuringHaistschairmanshiptheBantingandBestDepartmentofMedicalResearchwassetupasaseparateDepartmentwithitsownChairDr.IrvingFritzwhorecruitedsomeoutstandingresearchers.ThePhysiologyDepartmentmovedtoitsnewlocationinthepresent-dayMedicalSciencesBuilding.Modernlaboratorieseachtailoredtotheresearchfieldoftheinitialoccupantwereavailablein1969-70andhavebeenusedcontinuouslysincethenwithsomegrumblingaboutlackofwindowsbyoccupantsofcertainwindowlesslaboratories.HaistalsopresidedoverimportantchangesinUniversityregulationsguidingappointmentsandtenure.TherevisedregulationsbecameknownastheHaistRulesimportantbuildingblocksintheemergingre-definitionoftherelationshipbetweentheUniversityanditsfacultymembers.ReginaldR.Haist1965-1975 10DepartmentofPhysiologyJohnT.MurphyChairman1975-1980JohnMurphyMDPhDarelativelyyoungfacultymemberwhohadtrainedearlierwithNobelLaureateSirJohnEcclesassumedthechairmanshipin1975.Hehaddevelopedanoutstandingresearchprograminneurophysiologyfocusedonmotorcontrolinhighermammalsincludingprimatesseveralexcellentgraduatestudentsandpost-doctoralfellowsweretrainedinhislaboratory.Hepromotedtheareaofneurosciencebutrecognizedtheimportanceofotherareasofphysiology.UnderhisdirectiontheDepartmentwasorganizedintothreeResearchGroupsNeurophysiologyEndocrinologyandIntegrativePhysiologywhichincludedrespiratorycardiacautonomicandkidneyfunctions.EachGrouphelditsownperiodicmeetingsandinsomecasessocialevents.TheGroupsalsotookturnsselectingthelecturerfortheannualArchibaldByronMacallumLecturewhichwasinitiatedin1978.TheGroupspromotedinteractionswithandguidanceofgraduatestudents.InmodifiedformtheGroupconcepthascontinuedtothepresentday.InadditiontoitsresearchcommitmentstheDepartmentmaintainedastrongpresenceintheteachingprogramoftheMedicalSchoolespeciallyinthefirsttwoyears.AlsoastrongundergraduatetrainingprogramintheFacultyofArtsandSciencealreadyinexistenceintheearlypartofthe20thcenturywassustained.AdditionalteachingcommitmentsintheFacultiesofNursingPharmacyDentistryandPhysicalandHealthEducationaddedtotheburdenofinstructionsupportedbytheDepartment.InmitigationsomenewandorjointappointmentsaccruedtotheDepartmentduringtheregimesofHaistandMurphy.Neverthelesstheburdenofteachingwasconsiderable.Concernsdevelopedaroundthelossofresearchfundingduetoheavyteachingcommitments.GiventhewidescopeofPhysiologyandtheproblemofhavingknowledgeableexpertsinallitsbranchestheDepartmentincreasinglyaddedcross-appointedmembersfromotherDepartmentsandfromaffiliatedhospitalstobolsterareasofPhysiologyinwhichexpertiseamongprimaryappointeeswasthoughttobeinsufficientandtolightentheteachingloadforprimaryappointees.UneasyrelationshipswithemergingHospitalResearchInstitutesbegantodevelop.SeveraloftheHospitalsinparticulartheTorontoWesternHospitaltheMountSinaiHospitalandtheTorontoGeneralHospitalbegantoestablishresearchinstitutesalongthelinesoftheonethathadbeensetupearlierattheHospitalforSickChildren.Theseinstitutesbecamestaffedwithbasicscientistsmanyofwhomhadoutstandingresearchcredentials.Cross-appointmentsweresoughtbytheseindividualsintheestablishedBasicScienceDepartmentsoftheFacultyofMedicinesothattheycouldrecruitandtraingraduatestudents.Resolutionoftherequirementsforcross-appointmentandthedutiesdemandedofcross-appointeesbecameamajorpreoccupationforallsubsequentChairsofPhysiology.JohnMurphyrelinquishedthechairmanshipin1980inordertopursuespecialisttraininginneurology.AfterafewyearshegaveuphisresearchprogramandhisaffiliationwithPhysiologyandleftToronto.FollowinghisresignationasChairtherewasalengthysearchforanewChair.DuringthisperiodProfessorDonaldClarkeaseniormemberoftheDepartmentwithoutresearchresponsibilitiesbecameActingChair.HeservedagainasActingChairin1987-88duringanadministrativeleavetakenbythenextChairandhealsoservedasAssociateDeanforBasicSciencesintheearly1980s.JohnT.Murphy1975-1980 OurHistory11HaroldL.AtwoodChairman1981-1991Althoughtherewereseveralwell-qualifiedapplicantsfortheChairspositionwithintheDepartmentofPhysiologyandothersfromoutsidetheSearchCommitteewasunabletoagreeonanyofthemandbecamedeadlocked.FinallyoneoftheSearchCommitteememberspresentedanewcandidatefromtheDepartmentofZoologywhowasrecruitedasthenextChairofPhysiology.HaroldL.AtwoodwasanunusualchoicesincehehadnoformalconnectionwithtraditionalDepartmentsofPhysiologyandhismostsignificantresearchwasoninvertebrateorganisms.YouarethefirstinvertebratetoimpressthemammalswroteacolleaguefromBristolonlearningoftheappointment.AcomparativephysiologisthehadstudiedBiologyandComparativeNeurophysiologyattheUniversitiesofTorontoCaliforniaGlasgowOregonandtheCaliforniaInstituteofTechnology.IntheDepartmentofZoologyhehaddevelopedaninternationallyrecognizedresearchprogramonsynaptictransmissionofinvertebrateswithasecondaryfocusonmammalianneuromuscularphysiology.HehadalsoinitiatedadvancedcoursesinneurophysiologywhichattractedundergraduateandgraduatestudentsfrombothZoologyandPhysiologyDepartments.HisresearchwasrecognizedbyelectiontotheRoyalSocietyofCanada.IntheDepartmentofPhysiologyAtwoodcontinuedtheThreeResearchGrouptraditioninheritedfromJohnMurphy.InparticularheelectedtostrengthentheEndocrinologyGroupwithnewappointmentsratherthanlettingitfadeawaythroughimpendingretirements.Healsoappointedmorewomentofacultypositionsthanhadpreviouslybeendonethreeofhisfirstfourinitialappointeeswerewomen.OneofthemValerieWattwasthefirsttobringmodernresearchapproachesbaseduponmoleculargeneticstotheDepartment.AnothernewrecruitMiltonCharltonbroughtrecentlydevelopedapproachestophysiologybasedonopticalindicatorsforobservingintracellularcalciumdynamics.AsawholetheDepartmentsintakeofresearchgrantsandthesophisticationofresearchapproachesincreasedsteadilyduringthe1980s.AtwoodcontinuedandenlargedtwoDepartmentaladministrativeproceduresinitiatedbyJohnMurphyaSeniorAdvisoryCommitteewhichmetperiodicallytoadvisetheChaironimportantdecisionsaffectingtheDepartmentandanannualRetreatusuallyoff-campusatwhichgeneralissuescouldbethoroughlyairedanddiscussedbytheentireDepartment.PolicydirectionsweresometimesdecidedattheseRetreats.BothtraditionshavebeencontinuedbysubsequentChairs.AmongthemanyresearchcontributionsthatemanatedfromtheDepartmentofPhysiologyduringthisperiodthemostwidelyrecognizedwasthediscoveryofatrialnatriureticfactorANF.AcollaborationbetweenHaraldSonnenbergandA.DeBoldQueensUniversityshowedthatthispeptideissynthesizedandreleasedfromthecardiacatriaandcontributestohomeostasisofbloodpressure.ThisworkledtoaGairdnerAwardforthecollaboratorsandstimulatedagreatdealofresearchworld-wide.HaroldL.Atwood1981-1991 12DepartmentofPhysiologyAnotherareaofresearchthatbecameprominentwastheinvestigationofAlzheimersdiseasevigorouslypursuedbyDonaldMcLachlanUmbertoDeBoniandcollaborators.Inadditiontoresearchtheseinvestigatorspromotedawarenessoftheincreasingprevalenceofthisneurodegenerativediseaseinaginghumanpopulations.Inthemid-1980sadonationfromtheTanzfamilyledtoestablishmentoftheCentreforResearchinNeurodegenerativeDiseaseintherenovatedBotanyBuildingnexttotheMedicalSciencesBuildingwithDonaldMcLachlanasitsfirstDirector.AfterhisretirementoneofhisformerpostdoctoralfellowsPeterSt.GeorgeHyslopbecametheDirectorandtheCentreachievedgreatsuccessusingmoderngeneticapproachesinthestudyofseveralneurodegenerativediseases.Duringthe1980sthegraduatestudentsoftheDepartmentincreasedinnumberandprominenceandstimu-latedalotofdebateabouttheproceduresoftheDepartmentstrainingprogramwhichbecameprogressivelymodifiedinconsequence.Ledbyastudentwithorganizingambitionsin1981thegraduatestudentsinitiatedtheannualresearchdayFrontiersinPhysiologyFIPwiththeChairssupport.InduecoursetheannualMacallumLecturecametobeheldinconjunctionwithFIPenhancingthevisibilityofbothevents.ThegraduatestudentsalsoformedtheirownassociationGraduateAssociationofStudentsinPhysiologyorGASPwhichisrepresentedinvariousdecision-makingmeetingsandcommitteeswithintheDepartment.Thetrendtowardsincreasednumbersofcross-appointmentscontinuedasthehospitalresearchinstitutesgainedmomentum.ThePlayfairInstituteattheTorontoWesternHospitalwasespeciallycloselylinkedtoPhysiologybecauseitssecondDirectorWilliamTattonhadobtainedhisPhDintheDepartmentandrecruitedseveralmemberswithinterestsinneurosciencewhoweregivenappointmentsinPhysiology.RelationsbetweentheDepartmentandthePlayfairInstitutewereattimesfractiousbutthelinkwasmaintained.OneofthePlayfairresearchersJohnMacDonaldmovedtothePhysiologyDepartmentandeventuallybecameitsChairtherebyillustratingthevalueofcross-fertilizationwithintheTorontoMedicalSchoolcomplex.AfterservingtwotermsasChairAtwoodledasuccessfulgroupapplicationtotheMedicalResearchCouncilofCanadaandformedtheMedicalResearchCouncilGroupinNerveCellsandSynapses.TheGroupincludedmembersfromtheDepartmentsofPhysiologyandMedicineandtheFacultyofPharmacy.NewapproachesinresearchsuchasconfocalmicroscopyincreasedcollaborationamongGroupmembersandinvolvementintheNetworksofCentresofExcellenceprogramwithsupportforresearchfacilitiesandtrainingprogramswereamongthefeaturesfosteredbythisresearch-basedassociation.TheGroupcontinuedwithchangingmembershipintothenextcenturyJohnMacDonaldbecameitsnewDirectorin2000.AfterAtwoodsdeparturefromtheChairsOfficeanActingChairelectedbytheSeniorAdvisoryCommitteepresidedovertheDepartmentforthenextninemonths.Dr.UweAckermannacardiovascularphysiologistandaneloquentteacherkepttheDepartmentrunningsmoothlywhiletheDeanofMedicineJohnDirksattemptedmajorchangesintheorganizationoftheFaculty.Ackermanninstitutedamorerefinedandsystematicmethodforassessingthecontributionsoffacultymembersandtheirannualmerit-basedsalaryincreasesandhissystemhasbeenretainedwithmodificationsbysubsequentChairs.MeanwhileaSearchCommitteeengagedinselectingthenextChair. OurHistory13MladenVranicChairman1991-1995WithabackgroundinmedicineandphysiologyacquiredinCroatiaMladenVranichadcometoCanadaasthelastpost-doctoralfellowofCharlesBestandwasappointedtothefacultyin1965byReginaldHaist.HehadthenestablishedaproductiveandinternationallyrecognizedresearchlaboratoryfocusedonendocrinologyanddiabeteswhichproducedtraineeswhobecamesuccessfulscientistsatuniversitiesinCanadaandinothercountries.LinksbetweentheDepartmentsofPhysiologyandMedicinewerestrengthenedduringVranicschairmanship.AnagreementwasreachedwiththeChairmanoftheDepartmentofMedicineDr.ArnieAbermanwhosoonthereafterbecamethenewDeanofMedicinetoaddafacultymemberintheareaofdiabetesresearchwithspaceintheDepartmentofPhysiologyandsalaryfromtheDepartmentofMedicine.Otherpositionsindiabetes-relatedresearchwereobtainedwithpartialsupportfromtheBantingandBestDiabetesCentre.Strengtheningofresearchrelatedtoendocrinologyanddiabetesarosealsofromaninternationalconnection.TheDepartmentofPhysiologyandtheBantingandBestDiabetesCentreorganizedanumberofsymposiainitiallybetweenTorontoandtheKarolinskaInstituteinStockholmandlaterbetweenTorontoStockholmandZagreb.TheseinitiativeswerestronglysupportedbytheDirectoroftheBantingandBestDiabetesCentrewhereDr.VranicservedastheheadofInternationalRelations.TheDepartmentofPhysiologyandtheBantingandBestDiabetesCentrealsoco-organizedthecelebrationin1996ofthe75thanniversaryofthediscoveryofinsulinintheDepartmentofPhysiology.Dr.Vranicchairedtheorganizingcommittee.AnendowedlectureshipBestLecturewasestablishedbyDr.JackDavidsonwhohadobtainedhisPhDdegreeunderthesupervisionofCharlesBest.TheseactivitieshelpedtobolstertheDepartmentsacademicstrengthintheareasofendocrinologyanddiabetes.AdministrativelytheDepartmentcontinuedtooperatewithaSeniorAdvisoryCommitteeandwithtwoAssociateChairsoneforgraduateeducationandtheotherforundergraduateeducation.TheChairmanwithadvicefromtheSeniorAdvisoryCommitteeconsideredmajorissuesdepartmentalresponsesandinitiatives.TheannualDepartmentalRetreatwasmaintainedatthiseventimportantissuesarisingduringthepreviousyearweredebatedandreportsofDepartmentalcommitteespresented.AtoneoftheRetreatsmodificationstotheDepartmentsGroupstructurewereconfirmedinplaceoftheoriginalthreeverybroadgroupsmorespecializedonesweresetup.Theseincludedneurophysiologyendocrinologycardiovascularandrenalphysiologyrespiratoryphysiologyandtheoreticalphysiology.Amajorgoalofthegroupsarrangementwastobridgeinthewidestpossiblesensemolecularcellularorganandintegrativeapproaches.WithtimemolecularandcellularaspectsofphysiologywereincreasinglyemphasizedinTorontowhileotheruniversitiesQueensUniversityofMontrealUniversityofAlbertaemphasizedintegrativephysiologyespeciallyintheneurosciences.Thenumberofgroupsexpandedto8duringthenextchairmanshipbutnotallprovedviableinthelongtermandconsequentlythenumberwaseventuallyreducedto4.Thegroupshadresponsibilitiesforseminarsaspectsofgraduatestudenttrainingandexaminationandfosteringexchangeofscientificinformation.MladenVranic1991-1995 14DepartmentofPhysiologyWithincreasingnumbersofcross-appointedmemberstheDepartmenthadabout70facultymembersduringthisperiod.Onlyabout20ofthesewereprimaryappointmentslocatedintheMedicalSciencesBuildingtherestwerelocatedinotherDepartmentsbothBasicScienceandClinicalandinhospitalresearchinstitutes.TheDepartmentwasabletomaintainitscoreofprimaryappointmentsthroughexternalsalaryawardsandcreativearrangementswithotherdepartmentsdespitefinancialdifficultiesbroughtonbyyearlybudgetcutsfromtheUniversity.TheoverallratingsofthedepartmenthavebeenexcellentasindicatedintheOCGSreviewsofthegraduateprogramandintheLCMEreviewsofthemedicalprogram.VranicsresearchwasrecognizedbyelectiontotheRoyalSocietyofCanadaCanadianAcademyofHealthSciencesandtheCroatianAcademyofArtsandSciences.HeisalaureateoftheCanadianMedicalHallofFamehewasappointedOfficeroftheOrderofCanadaandtheOrderofOntarioandreceivedtheQueenElizabethIIDiamondJubileeMedal.JohnChallisChairman1995-2000WhenMladenVranichadtoleaveofficeduetotheUniversitysrequirementforofficialretirementatage65DeanAbermanlaunchedasearchforanewChair.TheresultwasrecruitmentofDr.JohnChallisfromtheUniversityofWesternOntarioUWO.EducatedinEnglandPhDCambridgeUniversityhehadmovedtoCanadaanddevelopedaremarkablysuccessfulandprolificresearchprogramintheareaofreproductiveanddevelopmentalphysiologynotprevious-lywellrepresentedinthecoreofthePhysiologyDepartmentinToronto.Hisresearchisfoundationaltoourcurrentunderstandingofendocrinedevelopmentandcontrolofthebirthprocess.HisresearchwasrecognizedbyelectiontotheRoyalSocietyofCanadatogetherwithnumerousotherawards.HehadattainedmajoradministrativepositionsatUWOVice-PresidentResearchSt.JosephsHealthCentreScientificDirectorLawsonResearchInstituteDirectorMedicalResearchCouncilGroupinFetalandNeonatalHealthandDevelopment.AsDeanAbermanremarkedafterreadinghiscredentialsHeyThisguyisaheavyhitter.JohnChallisinheritedadepartmentwitharichtraditioninexcellenceparticularlyinneuroscienceandinendocrinologydiabetesandahistoryofdiscoveryininsulin.Buthewaswillingtoexploremethodsthatcouldfurthermodernizethetraditionalphysiologicalmode.Heundertookdevelopmentofanewstrategicplanexaminingwaystoenhancerelationshipswithcolleaguesandcross-appointedmembersinthehospitalsandhospital-basedresearchinstitutes.ThushepromotedcloserrelationshipswithclinicaldepartmentsparticularlyJohnR.G.Challis1995-2000Dr.C.BestMrs.BestDr.M.Vranic OurHistory15ObstetricsandGynecologyandwithhospital-basedresearchinstitutes.NewfacultyappointmentsinthecoreDepartmentstrengthenedneuro-endocrinologydevelopmentalphysiologyandintegrativesensory-motorphysiology.Coreandnon-corefacultymemberswereequallyappreciatedandparticipatedmoreinthelifeoftheDepartment.DepartmentalcohesionwasalsoenhancedbyinstitutionofaweeklynewsletterPhysioLINKdistributedbye-mail.DuetoclosecollaborationwiththehospitalsthefacultycomplementforPhysiologygrewfrom75toabout100withallprimaryandcross-appointedmembershavingequalvotingrights.Inparallelwiththeincreaseoffacultymembershipthenumberofgraduatestudentsgrewconsiderably.Administrativeinnovationsincludedmoreformalandrigorousscrutinyofapplicationsforresearchgrantsbeforesubmission.AResearchCommitteechairedinitiallybyHaroldAtwoodregulatedtheinternalreviewprocess.TheSeniorAdvisoryGroupbecametheChairsAdvisoryGroupcomprisingSeniorAssociateChairheadsoftheUndergraduateandGraduateProgramsChairoftheResearchCommitteeahospitalrepresentativeandtwoothermembers.Meetingswereheldweekly.AtraininginitiativeindevelopmentalphysiologyledtoanagreementwithKarolinskaInstituteinStockholmtoholdannualmeetingsalternatingbetweenTorontoandStockholmforgraduatestudentspost-doctoralfellowsandclinicalfellows.ThisinitiativespearheadedbyDr.StephenMatthewscurrentChairoftheDepartmentcontinuestodaywith500alumniandrecentlyhasledtootherinitiativesincludingtheMatsSundinFellowshipinDevelopmentalHealth.Amuch-neededboostfortheDepartmentsfinancialwell-beingmaterializedintheformofanendowmentfortheChairOctober1996.InrecognitionofagifttotheUniversitybyErnestSmiththeChairofPhysiologywasnamedTheErnestB.andLeonardB.SmithChairinPhysiology.AttheturnofthecenturywhenJohnChallisleftofficetobecometheDirectorofanewlyformedInstituteoftheCanadianInstitutesforHealthResearchtheDepartmentofPhysiologyhadbecomelargerandmorediversifiedthanitscounterpartsatotherCanadianuniversitiesandcontinuedtomaintainexcellenceinresearchandeducation.Dr.J.ChallisandDr.A.AbermanDeanofMedicine1992-99 16DepartmentofPhysiologyJohnF.MacDonaldChairman2001-2008WhileasearchcommitteesetupbytheDeanofMedicineDavidNaylorconsideredaselectionofanewChairofPhysiologyDr.PatriciaBrubakerwastheActingChair2000-2001andtheDepartmentmaintainedastabletrajectory.ThesearchcommitteerecommendedJohnF.MacDonaldasthenextChairofPhysiology.HewastrainedinPhysiologyandNeuroscienceattheUniversityofBritishColumbiaPhDtheUniversityofSt.AndrewsMcGillUniversityandtheNationalInstitutesofHealthinBethesdaMaryland.InitiallyappointedattheUniversityofTorontoasafacultymemberintheDepartmentofPharmacologywithhislaboratoryattheTorontoWesternHospitalhemovedtotheDepartmentofPhysiologyin1990tobecomeafoundingmemberoftheMRCGroupinNerveCellsandSynapses.BycoincidencehisselectionasChairofPhysiologyoccurredalmostexactlyacenturyafteranotherCanadian-bornscientistofScottishancestryArchibaldByronMacallumhadbecomethefirstChairofthenewlyindependentDepartment.JohnMacDonaldsimpressivecredentialsinresearchasanindependentfacultymemberattheUniversityofTorontoincludediscoveryin1980-81ofvoltage-dependenceofanewclassofreceptorsfortheneurotransmitterglutamateinthemammalianbrain.ThesereceptorseventuallytermedNMDAreceptorswerelaterfoundtoberequiredforlong-termsynapticmodificationslong-termpotentiationthoughttounderliesomeformsoflearning.Aftermanysuccessfulstudiesofglutamatereceptorsheandco-workersuncoveredaroleforcalcium-permeablenon-selectivecationchannelsindelayedcelldeathofnervecellsinthebrainaffectedbystroke.Thesestudiespointthewaytodevelopmentofnewprotectiveagentsthatwillbeeffectiveformanyhoursfollowingthestrokeitself.JohnMacDonaldslaboratoryhastrainedanumberofoutstandinggraduatestudentsandpost-doctoralfellows.HisresearchcontributionshavebeenrecognizedthroughelectiontotheRoyalSocietyofCanada.JohnMacDonaldhadservedasGraduateCoordinatoroftheDepartmentduringtheChairmanshipofJohnChallis.OnceChairitsoonbecameevidentthathewasasuperblong-rangestrategicplannerfortheDepartment.MorethananypreviousChairheinvestedagreatdealoftimeinpredictingtheworst-casescenariosfortheDepartmentsfinancesbroughtonbybudgetcutsimposedbytheUniversityandforalikelydiminishedcomplementoffacultymembersandthenlookedforcreativesolutionstowardoffimpendingdoom.BywayofexplanationofhistendencytolookfortheworstoutcomescoupledwithanunusuallykeenfinancialeyehewouldcommentIamaScotchPresbyterianByanticipatingtheworstJohnMacDonaldachievedthebest.DuringhisChairmanshiptheDepartmentrecruitednewfacultymembersthroughexternalawardsandpartnershipswithresearchinstitutesandclinicaldepartmentsintheFacultyofMedicinethuscontinuingandenhancingtheapproachesinitiatedbypreviousChairs.TheDepartmentasawholeobtained11CanadaResearchChairs3CIHRNewInvestigatorawards1HeartandStrokeInvestigatorAwardandthehighlyprestigiousMichaelSmithChairinneurosciencesandJohnF.MacDonald2001-2008 OurHistory17mentalhealthheldbyDr.MinZhuo.Thisunprecedentedgrowthandlevelofsupportoccurredatatimewhenseveralotherbasicsciencedepartmentswereclosedormergedinanefforttodealwithongoingfiscalconstraints.Theareaofcardiovascularphysiologywasgreatlystrengthenedbynewappointments.TheResearchGroupsintheDepartmentnowcompriseEndocrinologyandDiabetesNeurosciencesB.R.A.I.N.GroupReproductiveandDevelopmentalPhysiologyandCardiovascularPhysiology.TheGraduateProgramintheDepartmentincreasedfrom100toabout150studentswhilethetrainingprogramwasalteredtoincludemoreparticipation-basedseminarstylegraduatecourses.TheUndergraduateprogramsweresustainedandsignificantlyanewagreementbetweentheFacultyofMedicineandtheFacultyofArtsandSciencehasledtofinancialcompensationfortheDepartmentduetothelargeteachingloadintheundergraduateprograminArtsandScience.TheundergraduateArtsandSciencestudentshavetheirownassociationUndergraduatePhysiologyStudentsAssociationUPSAsupportedbytheDepartment.TheDepartmentofPhysiologysachievementinresearchgrantsinsupportofresearchawardspublicationsandeducationalprogramshaveassureditsratingasoneofthebestoftheBasicScienceDepartmentsoftheFacultyofMedicine.IthasbecomethelargestandmostcomprehensivePhysiologyDepartmentinCanada.References1.E.H.Craigie1965.AHistoryoftheDepartmentofZoologyoftheUniversityofTorontoupto1962.TorontoUniversityofTorontoPress.2.M.L.Friedland2002.TheUniversityofTorontoAHistory.TorontoUniversityofTorontoPress.excerptsmodiedbyH.L.Atwood3.A.G.Huntsman1934.ProceedingsoftheRoyalSocietyofCanadaSer.328IV-VI.QuotedbyE.H.CraigieinReference1.4.M.Vranic1996.ThePhysiologyDepartmentUniversityofToronto1887-1921.pp.9-12.InTheDiscoveryofInsulinattheUniversityofTorontoAnexhibioncommemorangthe75thanniversary.TorontoInternaonalResearchSymposiumonDiabetes.excerptsmodiedbyH.L.Atwood5.M.Bliss1982.TheDiscoveryofInsulin.TorontoUniversityofTorontoPress.M.Bliss1992.BanngABiography.TorontoUniversityofTorontoPress.6.P.Wysong1989.HeroesandLegendsDr.JacobMarkowitz.InAdvancesinPhysiology.TorontoDepartmentofPhysiologyUniversityofToronto.editedbyH.L.Atwood 18DepartmentofPhysiologyChairs-DepartmentofPhysiology-from1981topresentBackrowJ.F.MacDonaldJ.R.G.ChallisP.L.BrubakerS.G.MahewscurrentSeatedH.L.AtwoodM.Vranic