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Alfred Mirsky NATIONAL ACADEMY OF SCIENCES A L F R E D E Z R A M IRSKY 1900—1974 A Biographical Memoir by S E Y M O U R S . CO HEN Any opinions expressed in this memoir are those of the author(s) and do not necessarily reflect the views of the National Academy of Sciences. Biographical Memoir COPYRIGHT 1998 NATIONAL ACADEMIES PRESS WASHINGTON D.C. CourtesyoftheRockefellerUniversityArchives,NewYork,NewYork ALFRED EZRA MIRSKY October 17, 1900—June 19, 1974 BY SEYMOUR S. COHEN LFRED EZRA MIRSKY, SON OF Michael David Mirsky and Frieda AIttelson Mirsky, graduated from the Ethical Culture School in New York City and from Harvard College, obtain- ing a B.A. degree in 1922. He studied at the College of Physicians and Surgeons of Columbia University for two years. On receipt of a fellowship from the National Re- search Council in 1924, he worked at Cambridge University under Joseph Barcroft during the academic year 1924-1925, and completed his graduate studies under Lawrence J. Henderson at Harvard. He wrote a dissertation titled “The Haemoglobin Molecule” and received a Ph.D. from Cam- bridge in 1926. The molecularity of haemoglobin and the molecular weight of the protein were established by Theodor Svedberg and Gilbert Adair in 1925. Their results demonstrated that pro- teins are rigorously definable species of large molecules, and were important in showing that proteins, despite their size, should be described in the molecular terms of the chemist. The initial postulates of protoplasmic components as being essentially undefinable, dispersible, and colloidal aggregates were eventually replaced by the view that the Reprinted with permission from Dictionary of Scientific Biography, vol. 18, suppl. II, pp. 633-36. New York: Chas. Scribner’s Sons, 1990. 3 4 BIOGRAPHICALMEMOIRS majorconstituentsofprotoplasmcontainproteinmolecules whoseshape,charge,andstateofaggregationaremarkedly affectedbytheresponseoftheirionizablegroupstothe hydrogenionconcentration(pH).Mirsky’searlypapers demonstratethathehadadoptedarigorouslychemicalori- entationfromthebeginningofhiscareer. In1924and1925Mirskypublishedeightpapersonhe- moglobinwithMortimerL.Anson,ofwhichthefirsttwo hadBarcroftascoauthor.Ansonwasafellowstudentin Barcroft’slaboratory,aswellasalatercolleagueatthe BiophysicsLaboratoryoftheCancerCommissionatHarvard. ThecollaborationofMirskyandAnsononhemoglobin continueduntil1935.Aspresentedinpreliminaryformin Mirsky’sdissertation,thedisruptionofproteinorganiza- tionandprecipitation(denaturation)hadbeenobserved tobereversible.Themechanismsofthesephenomenawere themajorproblemsstudiedbytheseyounginvestigators forthenextdecade. On25May1926MirskymarriedRebaPaeff;theyhada daughterandason.In1927Mirskywasappointedanas- sistantinthelaboratoryofAlfredF.Cohnatthehospital oftheRockefellerInstituteforMedicalResearch.Anson wasappointedtothelaboratoryofJohnHowardNorthrop atthePrincetonbranchoftheinstitute.Cohnwasen- gagedinthequantitationofactivitiesofthehumanheart. MirskybeganhisworkinCohn’slaboratorywithstudies ofpHinthebloodofdevelopingchicks,usingaglass electrodedevelopedwithAnson.TheincreaseinpHsimu- latedthecurveofdecreaseinoxygenconsumptionduring development.TheresultingpapermarkedtheendofMirsky’s associationwithCohn’sresearchprogram,sinceCohnhad becomemoreinterestedinhumanisticstudiesthaninthose oflaboratoryscience. ALFREDEZRAMIRSKY 5 MirskyandAnsonresumedthestudyofhemoglobinand itsdenaturationandrenaturation.Inasecondseriesof paperswrittenbetween1929and1935,theyshowedthat proteincoagulationtakesplaceintwosteps,inwhichun- foldingcanbeseparatedfromprecipitation.Horsehemo- globin,coagulatedbyvariousmethods,includingheator acid,canbesolubilizedanditsunfoldedstateconvertedby acyanidesolutiontoastateindistinguishablefromnative hemoglobin;thisproductcanthenbecrystallizedtoa denaturablehemoglobin.Thedenaturableportionofthe hemoglobinwasshowntobetheglobin;itwasfoundthat otherdenaturableproteins,suchasserumalbumin,canbe renatured.Theseresultswereextendedtotheformationof anactivetrypsinfromaninactivedenaturedenzyme. Freesulfhydrylgroupsappearedinthedenaturationof eggalbuminandserumalbumin.Althoughotherresearch- ershadproposedthatsuchgroupsaregeneratedfrom disulfidebondsduringdenaturation,anotherhypothesis wasformulatedin1936,duringMirsky’ssabbaticalyearat theCaliforniaInstituteofTechnology.MirskyandLinus Paulingthenproposedthatnativeproteinsarecoiledin specificconfigurationswhosepartsarestabilizedlargely byhydrogenbonds,andthatunfoldinganddenaturation revealgroupspreviouslyobscuredandprotectedbythe originallyfoldedchains.Thispaperwasanimportantearly statementofnow-currentviewsofproteinstructureandof themechanismofdenaturationandrenaturation.Both viewsoftheappearanceofsulfhydrylgroupsarenowbe- lievedtobecorrect,sinceinmanyproteinsnativestruc- tureisalsomaintainedbydisulfideswhosereductiongen- eratessulfhydrylsandopensthestructure. In1937Mirskystudiedchangesinmuscleproteinsinan attempttocorrelatethemwithfunctionalalterationsin themuscleasaresultofelevatedtemperature.Anirrevers- 6 BIOGRAPHICALMEMOIRS iblemuscleshortening,knownasthermalrigor,wasassoci- atedwiththeappearanceofsulfhydrylgroups,andhecon- cludedthatthisphenomenonwasduetoadenaturation. Healsobelievedthatdenaturationwaspartofmusclecon- tractiongenerally,althoughherealizedthatnotenough wasknownaboutmuscleproteinsatthetime. AninterestinstructuralproteinsledMirskytoattempt theisolationofaproteincomplexthathadbeendescribed in1938asderivedfromthecytoplasm.Thefibrousmate- rialwasfoundtocontainlargeamountsofdeoxyribonucleic acid(DNA)andwasclearlynuclearinorigin.Hethen begananewlineofinvestigation,attemptingtounder- standthestructuralandfunctionalsignificanceofnuclear nucleoproteins.Inthemid1930’sthediscoverythatthe plantvirusesareribonucleoproteinsandthatribonucleic acid(RNA)ispresentincytoplasmicparticulateshadbe- gunthegrowthofbiochemicalinterestinthenucleicac- ids,culminatinginthe1944discoverybyO.T.Averyand hiscolleaguesthatDNAisthepneumococcaltransform- ingagent.By1941enoughwasknownoftherelationof genestochromosomestoposetheproblemofthechemi- calnatureofthehereditarydeterminants.Inapproaching thisquestion,Mirsky,whoin1940hadbecomeanassoci- atememberoftheRockefellerInstitute,establishedacol- laborationwiththecytochemistArthusW.Pollister,ofthe departmentofzoologyatColumbiaUniversity.Forthe nextdecadeMirsky’slaboratoryatRockefellerInstitute wasaleadingcenterforstructuralandfunctionalstudies ofcellnuclei. Mirsky’sfirstformalpaperwithPollisterdescribesthe extractionofnucleoproteinsfromawidevarietyofanimal cells.Theirmajorapproachwasthedifferentialuseof neutralsodiumchloridesolutionsofvaryingconcentra- tions:physiologicalsalineremovedproteinandcytoplas- ALFREDEZRAMIRSKY 7 micconstituents;concentratedsaline(1Mto2M)extracted DNAandotherproteinsthatwereprecipitatedinphysi- ologicalsaline.(Itwasshownlaterthanconcentratedsa- linedissociatedDNAandproteins.)Thesesolublecompo- nentsreassociatedandreprecipitatedatlowersalt concentrations.Hence,themildprocedure,oftensuitable fortheisolationofDNA,neverthelessintroducedrestruc- turingoftheoriginalcellularcomplex.Somethirtyyears lateritwasshownthatchromosomalsubunitsofDNAand variousproteins,knownasnucleosomes,areisolablewith- outdissociation,andhencerepresentabetterapproachto theisolationofamorenativeDNA-proteincomplex.Nev- ertheless,theDNA-proteincomplexesisolatedin1941were showntocomefromcellnucleiandwerebelievedtobe componentsofchromosomes.Mirsky’sreviewofthesta- tusofthisfieldin1943isausefulsummaryofthisearly periodofthebiochemistryofgeneticmaterial.Mirskydid notspeculateastothespecificchemicalnatureofthe gene;hesuggestedthatthenewlyisolatedDNAproteins wereeither“thegenesthemselvesorwereintimatelyre- latedtogenes.” MirskyandPollisterthenattemptedtoisolatechromatin fromthenucleiofcertaintypesofanimalcellsandthought thattheyhadobtainedthreadsofthismaterial,whichmight evenhavebeenintactchromosomes.From1946to1951 theseeffortswereextendedwiththecollaborationofHans Ris.Threadspossessingthemaincytologicalfeaturesof chromosomeswereisolatedfromcalfthymuslymphocytes andanalyzed.Over90percentwerefoundtoconsistof nucleohistonecontainingDNA.Aninsolubleresiduecon- tainedproteindeterminingtheformofthe“chromosome” aswellassomeRNAandDNA.Thesestudieswereex- tendedtotheisolationofsimilar“chromosomes”from manykindsofcells,includingmorevoluminousstructures 8 BIOGRAPHICALMEMOIRS frominterphasenuclei.Inhislastpapers,however,Mirsky becamemorecircumspectaboutproblemsofisolatingac- tivechromatin. In1948MirskybecameamemberoftheRockefeller Institute;occupyingnewlaboratories,heenlargedhisgroup, whichinadditiontoRisincludedVincentAllfrey,Marie Daly,andHerbertStern.ForeignvisitorssuchasAlberto Monroybegantoworkonproblemsofchemicalembryol- ogy.WithRis,Mirskyshowedthatdiploidsomaticcellsof anorganismcontainidenticalamountsofDNA,twicethat ofhaploidgermcells.In1950HermannJ.Mullercon- gratulatedMirskyandreferredtothe“granddiscovery”of DNAconstancy,whichsupportedtheconceptofDNAas thehereditarymaterial. In1950,inasymposiumcommemoratingthefiftieth anniversaryoftherediscoveryofMendel’swork,Mirsky presentedapapertitled“SomeChemicalAspectsofthe CellNucleus.”NotingtheconstancyofDNA,incontrast withthevariabilityofRNA,heconcludedthatDNAis partofthegenesubstance.Nevertheless,somesixyears afterAvery’sdiscovery,hewasstillunconvincedthatDNA itselfwasthesolegeneticmaterial,pointingouttheinsen- sitivityoftheassayanddifficultyofassuringthatminute quantitiesofproteinarenotattachedtotheDNA.Asnoted byNormanW.Pirie,whohadsimilarreservationsinlater yearsconcerningtheinfectivityofviralRNA,“Scepticism andobjectivityarenearneighbors.”Thechemicalevidence
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