Table Of ContentExplaining Photosynthesis
History, Philosophy and Theory
of the Life Sciences
Volume 8
Editors:
CharlesT.Wolfe,GhentUniversity,Belgium
PhilippeHuneman,IHPST(CNRS/UniversitéParisIPanthéon-Sorbonne),France
ThomasA.C.Reydon,LeibnizUniversitätHannover,Germany
EditorialBoard:
MarshallAbrams(UniversityofAlabamaatBirmingham)
AndreAriew(Missouri)
MinusvanBaalen(UPMC,Paris)
DomenicoBertoloniMeli(Indiana)
RichardBurian(VirginiaTech)
PietroCorsi(EHESS,Paris)
FrançoisDuchesneau(UniversitédeMontréal)
JohnDupré(Exeter)
PaulFarber(OregonState)
LisaGannett(SaintMary’sUniversity,Halifax)
AndyGardner(Oxford)
PaulGriffiths(Sydney)
JeanGayon(IHPST,Paris)
GuidoGiglioni(WarburgInstitute,London)
ThomasHeams(INRA,AgroParisTech,Paris)
JamesLennox(Pittsburgh)
AnnickLesne(CNRS,UPMC,Paris)
TimLewens(Cambridge)
EdouardMachery(Pittsburgh)
AlexandreMétraux(ArchivesPoincaré,Nancy)
HansMetz(Leiden)
RobertaMillstein(Davis)
StaffanMüller-Wille(Exeter)
DominicMurphy(Sydney)
FrançoisMunoz(UniversitéMontpellier2)
StuartNewman(NewYorkMedicalCollege)
FrederikNijhout(Duke)
SamirOkasha(Bristol)
SusanOyama(CUNY)
KevinPadian(Berkeley)
DavidQueller(WashingtonUniversity,StLouis)
StéphaneSchmitt(SPHERE,CNRS,Paris)
PhillipSloan(NotreDame)
JacquelineSullivan(WesternUniversity,London,ON)
GiuseppeTesta(IFOM-IEA,Milano)
J.ScottTurner(Syracuse)
DenisWalsh(Toronto)
MarcelWeber(Geneva)
Forfurthervolumes:
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Kärin Nickelsen
Explaining Photosynthesis
Models of Biochemical Mechanisms,
1840-1960
2123
KärinNickelsen
HistoryofScience
LMUMunich
Munich
Germany
ISSN2211-1948 ISSN2211-1956(electronic)
History,PhilosophyandTheoryoftheLifeSciences
ISBN978-94-017-9581-4 ISBN978-94-017-9582-1(eBook)
DOI10.1007/978-94-017-9582-1
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Preface
Thisbookisastudyof‘lightanddarkness’—aswasthetitleofthehabilitationthesis
onwhichitisbased: metaphoricallyspeaking, intermsofdarkfailuresandbright
successes in the middle of conceptual fog; and in a very literal sense, referring to
thechemicaleffectsoflightanddarknessonthegreenpartsofplants.Itdealswith
the elucidation of the photosynthetic mechanism, which has not been the subject
ofabook-lengthstudybefore.Thismeansthatalotofprimarymaterialhadtobe
analysed,processedandbroughtintoasensiblearrangement.
Atthesametime,thebookaimstocontributetothelongstandingphilosophical
questionofhowknowledgeisgeneratedinscience. Ithasrepeatedlybeenargued,
byhistoriansofscience,byphilosophersofscienceandinparticularbythosewho
consider themselves affiliated to the ‘History and Philosophy of Science’ (HPS)
that this question can only be answered by scrutinising actual cases and trying to
understand from these some characteristic features of scientific research.1 This is
whatIamattemptingtodointhisstudy.Theresultisricherinhistoricaldetailthan
manyphilosophersofsciencemightwishtoengagewith;atthesametime,itscore
questionsaremorephilosophicallyorientedthanmanyhistoriansofsciencemight
expect. The main issue is of a methodological nature: How and for what reasons
do scientists do what they do, while they are investigating a problem, such as the
mechanismofphotosynthesis?Howdotheysettheirprioritieswhenpreferringone
optiontoanother?Accordingtowhichconventions,habitsandexpectationsdothey
organise and (re-)direct their work? It is, to sum it up, a study in the heuristics of
scientificresearch.
BeforeIexplainthesethoughtsinmoredetail,Ishallspendtherestofthispref-
aceonsomehighlyappropriatewordsofacknowledgementandgratitude.Firstand
foremost,IwouldliketothankGerdGraßhoffandallthemembersandstudentsof
theHistoryandPhilosophyofScienceDivisionoftheUniversityofBern’sInstitute
ofPhilosophy.Theparticularresearchprofileofthisdivision—which,unfortunately,
1Cf.,e.g.,SchickoreandSteinle(2006);argumentsalongtheselinescanalsobefoundinMauskopf
andSchmaltz(2011),whichistheoutcomeofoneoftherecentlyinstitutedconferenceson&HPS,
thatis,‘IntegratedHistoryandPhilosophyofScience’.
v
vi Preface
hasbeenabolishedsince—stronglyinfluencedmywork.Theprojectnotonlyben-
efited from stimulating discussions with Gerd Graßhoff and others but also from
thesabbaticalleavethatIwasgenerouslypermittedtotakeintheautumnof2008.
I spent this leave at the Max Planck Institute for the History of Science in Berlin,
andIamparticularlygratefultoLorraineDastonforhertremendoushospitalitydur-
ingmystaythere.Thisstudyalsoprofitedenormouslyfrommyacquaintancewith
Govindjee(nofirstname)oftheUniversityofIllinoisatUrbana–Champaign,whoso
generouslysharedhisimmenseknowledgeof,andhiscontagiousenthusiasmfor,the
historyofphotosynthesisresearchwithme,andwhometiculouslywentthroughthe
detailsofthisbook’smanuscript.MyextendedresearchstayatUrbana–Champaign
proved exceedingly fruitful and pleasurable, thanks largely to the hospitality and
practical help given to me by Govindjee and his wife, Rajni Govindjee. I am also
grateful to have participated in the DFG-funded network ‘Philosophy of the Life
Sciences’, organised by Maria Kronfeldner, which has provided a valuable source
ofinspirationoverthepastyears.
Furthermore, I am grateful to a large number of friends and colleagues for dis-
cussions at various occasions, useful hints and valuable material; (in alphabetical
order)ChristinaBrandt,AngelaCreager,PetraGentz-Werner,MathiasGrote,Ekke-
hardHöxtermann,JeremiahJames,ChristianJoas,FabianKrämer,GiannaPomata,
TilmanSauer,RaphaelScholl,PhillipSloan,RichardStaley,FriedrichSteinle,Ger-
hardWagenitz, MarcelWeber,VolkerWissemannandAdrianWüthrich. Thesame
holdstruefortheenormouslyhelpfulstaffatthevariousarchivesthatIconsulted,
namely the archive departments of: the ETH Zurich; the Max Planck Society; the
Berlin-BrandenburgAcademyofSciencesandHumanities;theBerlinStateLibrary;
the University of Illinois at Urbana–Champaign; the University of Chicago; the
University of Cambridge, UK; and the Braunschweig/Berlin-based Physikalisch-
TechnischeBundesanstalt.BesidestheUniversityofBern’sInstituteofPhilosophy,I
wouldalsoliketothankthefollowinginstitutionsfortheirgenerousfinancialsupport:
TheYoungAcademyattheBerlin-BrandenburgAcademyofSciencesandHuman-
ities and the GermanAcademy of Sciences Leopoldina; the Hochschulstiftung of
theBurgergemeindeBern;andtheMittelbaufondsoftheUniversityofBern.Ialso
wouldliketothankMargaretaSimons,whocarefullyeditedthebook;BasilMarti,
whomanagedthearchivaldatabases, straightenedoutthechemistryandgavethe
habilitationafinalread;JosephineMusil-GutschandClausSpenninger,whowereof
enormouspracticalhelp;and,inparticular,CaterinaSchürch,whogreatlyimproved
thegraphs,wentthroughthebibliographyandproofreadthewholemanuscripttoits
enormousbenefit.
Finally, I am indebted to the History Department of the Ludwig Maximilians
University Munich for granting me a sabbatical leave very soon after I started my
tenured position there in 2011. This gave me the indispensable freedom to focus
onfinalisingthisbookforpublication.And,ofcourse,therearetheusualsuspects
amongfriendsandfamilywhoknowverywell,Ihope,howmuchtheirsupporthas
beencherished.
Contents
1 Introduction.................................................. 1
1.1 ModelsofMechanisms..................................... 2
1.2 HeuristicsandStrategiesinPhotosynthesisResearch............ 6
1.3 TheIndividualandtheCommunity........................... 8
1.4 SurveyoftheBook ........................................ 12
2 InPursuitofaPathway(1843–1918) ............................ 15
2.1 TheNineteenth-CenturyConceptionofPhotosynthesis .......... 16
2.2 FindingtheChemicalMechanism............................ 18
2.2.1 JustusLiebigandtheOrganicAcidModel ............. 19
2.2.2 AdolfvonBaeyerandtheFormaldehydeModel......... 22
2.2.3 TestingandModification............................ 25
2.2.4 TheFormaldehydeProblem ......................... 28
2.3 ReconcilingtheFirstApproaches ............................ 30
2.3.1 TheFormicAcidModel............................. 31
2.3.2 TheOrganicAcid–FormaldehydeHypothesis........... 32
2.3.3 TheWaterCleavageModel .......................... 32
2.3.4 TheChlorophyllComplexModel..................... 34
2.4 FeaturesCommontotheModels............................. 36
2.4.1 SimplicityandSimplification ........................ 36
2.4.2 IncompleteEmpiricalSupport ....................... 39
2.4.3 SelectedFocusandModules......................... 40
2.5 Model-BuildingHeuristics.................................. 41
2.5.1 ExtendingaPrototype .............................. 41
2.5.2 TheTransferofCausalKnowledge ................... 42
2.5.3 TheBuildingBlockStrategy......................... 44
2.5.4 ThePrincipleofPlurality............................ 46
2.6 CollectiveVersusIndividualGoals ........................... 48
2.6.1 PhotosynthesisasaSideIssue ....................... 48
2.6.2 ConstructiveResearchOpportunism .................. 51
vii
viii Contents
3 OttoWarburgandtheTurntoManometry(1912–1925) ........... 55
3.1 NewMaterialsandMethods ................................ 56
3.1.1 OttoWarburg(1883–1970) .......................... 56
3.1.2 Manometry ....................................... 58
3.1.3 Chlorella ......................................... 60
3.1.4 BufferSolution .................................... 63
3.1.5 Bolometry ........................................ 63
3.1.6 RotatingSectors:TheFlashingLightTechnique ........ 64
3.1.7 Inhibitors......................................... 64
3.2 Warburg’sEarlyPhotosynthesisModel ....................... 65
3.2.1 ExperimentalFindingsandtheInterpretationsthereof.... 66
3.2.2 PhotosynthesisFramedasPhotolysis.................. 73
3.3 TheEfficiencyoftheProcess................................ 75
3.4 Father,SonandPhotosynthesis .............................. 78
3.4.1 EmilWarburgandPhotochemistry.................... 78
3.4.2 InfluencesonOttoWarburg.......................... 80
3.5 StudiesinCellRespiration.................................. 83
3.6 ComparisonwiththeChlorophyll-ComplexModel ............. 87
3.7 OttoWarburg’sBuildingBlocks ............................. 89
3.8 Warburg’sImpactonPhotosynthesisResearch ................. 90
4 StrugglingwiththeStandardModel(1930–1941) ................. 93
4.1 FluorescenceandtheStandardModel......................... 94
4.1.1 TheKautskyEffect................................. 94
4.1.2 JamesFranckandPhotosynthesis..................... 96
4.1.3 TheNewStandardModel ........................... 99
4.2 TheCrucialExperimentsof1932 ............................ 108
4.2.1 RobertEmerson:Harvard,Berlin,CaltechandStanford .. 108
4.2.2 Emerson,Arnoldand2500MoleculesofChlorophyll.... 110
4.3 TheGeneralisedEquationforPhotosynthesis .................. 113
4.3.1 CornelisB.VanNielandGeneralMicrobiology......... 113
4.3.2 BacterialPhotosynthesisandtheConsequences......... 116
4.4 ThePhotosyntheticUnit.................................... 120
4.4.1 ContextandScopeofthe1936Paper.................. 122
4.4.2 CritiqueoftheStandardModel....................... 124
4.4.3 TheUnitasExplanatoryAlternative................... 125
4.5 Franck’sConservativeAlternatives ........................... 126
4.5.1 FranckandHerzfeld’sProposal ...................... 126
4.5.2 Franck’sFurtherAttempts........................... 129
4.6 IsolatedChloroplastsandWaterSplitting...................... 132
4.6.1 Robert(Robin)HillandtheChloroplastReaction ....... 132
4.6.2 ImplicationsoftheFindings ......................... 137
4.7 OntheVergeofNewPerspectives............................ 138
4.7.1 BiologicalStudiesGeneralised....................... 138
4.7.2 TheMainLinesofThought.......................... 140
4.7.3 ANewConceptionofPhotosynthesis ................. 146
Contents ix
5 TheMaximumQuantumYieldControversy(1937–1955) .......... 149
5.1 PhotochemicalQuantumYieldMeasurements.................. 150
5.2 FirstOpponentsoftheWarburg–NegeleinValue................ 151
5.3 EmersonandLewis’sChallenge ............................. 153
5.3.1 TheCarbonDioxideBurst(1939–1941) ............... 153
5.3.2 TheRedDrop(1943)............................... 156
5.4 Wartime ................................................. 157
5.5 AttemptstoFindaSolution ................................. 161
5.5.1 ThePhotosynthesisProjectatUrbana ................. 161
5.5.2 WarburgComestotheUnitedStates .................. 161
5.5.3 TheTimeSpentatUrbana........................... 164
5.5.4 TheRediscoveryoftheMaximumQuantumYield....... 167
5.5.5 Franck’sAttempttoFindaCompromise ............... 171
5.5.6 FurtherExchangesofBlows ......................... 172
5.5.7 ControversialThemesAround1950................... 174
5.6 AHardeningoftheFronts .................................. 178
5.6.1 TheOne-QuantumMechanism....................... 178
5.6.2 TheGatlinburgConferenceonPhotosynthesis,1952..... 181
5.6.3 EmersonStrikesBack(1955) ........................ 187
5.7 TheAftermath ............................................ 190
5.7.1 TheEnhancementEffect ............................ 190
5.7.2 Emerson’sDeathandBeyond........................ 193
5.8 HowControversiesEnd .................................... 194
6 ThePathofCarboninPhotosynthesis(1937–1954)................ 201
6.1 EarlyRadiotracerExperiments .............................. 202
6.1.1 KamenMeetstheCyclotron ......................... 202
6.1.2 KamenMeetsRuben ............................... 204
6.1.3 TheFirstMetabolismExperiments.................... 207
6.1.4 Carbon-14andtheEndofaCollaboration ............. 212
6.2 TheBio-OrganicChemistryGroup........................... 214
6.3 TheFirstCyclicModel..................................... 220
6.4 NewMethodsandaNewModel ............................. 226
6.4.1 PaperChromatography ............................. 226
6.5 TheSecondCyclicModel(1950) ............................ 229
6.6 InNeedofRe-Orientation .................................. 232
6.6.1 TwoUnexpectedCompounds(1952).................. 232
6.6.2 TheDegradationStudies ............................ 234
6.7 TheSaturationExperiments................................. 236
6.8 ThePathXXIPaper ....................................... 240
6.8.1 TheFraction1Protein .............................. 243
6.9 TheHeuristicsoftheBerkeleyGroup......................... 246
