Table Of ContentHistory, Philosophy and Theory of the Life Sciences
Pierre-Alain Braillard
Christophe Malaterre
Explanation
in Biology
An Enquiry into the Diversity of
Explanatory Patterns in the Life Sciences
History, Philosophy and Theory of the Life
Sciences
Volume 11
Editors
CharlesT.Wolfe,GhentUniversity,Belgium
PhilippeHuneman,IHPST(CNRS/UniversitéParisIPanthéon-Sorbonne),France
ThomasA.C.Reydon,LeibnizUniversitätHannover,Germany
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Moreinformationaboutthisseriesathttp://www.springer.com/series/8916
Pierre-Alain Braillard (cid:129) Christophe Malaterre
Explanation in Biology
An Enquiry into the Diversity of Explanatory
Patterns in the Life Sciences
123
Pierre-AlainBraillard ChristopheMalaterre
IndependantScholar Départementdephilosophie&CIRST
Peyregrand,Drulhe Chairederecherche UQAMenPhilosophie
France dessciences
UniversitéduQuébecàMontréal(UQAM)
Montréal,Québec
Canada
ISSN2211-1948 ISSN2211-1956 (electronic)
History,PhilosophyandTheoryoftheLifeSciences
ISBN978-94-017-9821-1 ISBN978-94-017-9822-8 (eBook)
DOI10.1007/978-94-017-9822-8
LibraryofCongressControlNumber:2015939272
SpringerDordrechtHeidelbergNewYorkLondon
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Contents
1 ExplanationinBiology:AnIntroduction................................ 1
Pierre-AlainBraillardandChristopheMalaterre
PartI ExploringExplanatoryPluralisminBiology
2 Is There an Explanation for::: the Diversity
ofExplanationsinBiologicalStudies? ................................... 31
MichelMorange
3 ExplanationinSystemsBiology:IsItAllAboutMechanisms?....... 47
ConstantinosMekios
4 Historical Contingency and the Explanation
ofEvolutionaryTrends..................................................... 73
DerekTurner
5 Developmental Noise: Explaining the Specific
HeterogeneityofIndividualOrganisms.................................. 91
FrancescaMerlin
PartII Mechanistic Explanation: Applications
andEmendations
6 ExplaininginContemporaryMolecularBiology:Beyond
Mechanisms ................................................................. 113
FrédériqueThéry
7 EvolutionaryDevelopmentalBiologyand the Limits
ofPhilosophicalAccountsofMechanisticExplanation ................ 135
IngoBrigandt
8 TheRelevanceofIrrelevance:ExplanationinSystemsBiology....... 175
FridolinGross
v
vi Contents
9 Generalizing Mechanistic Explanations Using
Graph-TheoreticRepresentations........................................ 199
WilliamBechtel
PartIII TheRoleofMathematicsinBiologicalExplanations
10 MathematicalExplanationinBiology.................................... 229
AlanBaker
11 ExplanationandOrganizingPrinciplesinSystemsBiology........... 249
TobiasBreidenmoserandOlafWolkenhauer
12 AreDynamicMechanisticExplanationsStillMechanistic?........... 265
TarikIssadandChristopheMalaterre
PartIV TheRoleofHeuristicsinBiologicalExplanations
13 Heuristics,Descriptions,andtheScopeofMechanisticExplanation. 295
CarlosZednik
14 ProspectandLimitsofExplainingBiologicalSystems
inEngineeringTerms ...................................................... 319
Pierre-AlainBraillard
15 FromMechanismstoMathematicalModelsandBack
toMechanisms:QuantitativeMechanisticExplanations .............. 345
TudorM.Baetu
PartV New Theories of Explanation
inBiologyandElsewhere
16 BiologicalExplanationsasCursoryCoveringLawExplanations..... 367
JoelPress
17 ExplanatoryInterdependence:The CaseofStemCell
Reprogramming ............................................................ 387
MelindaBonnieFagan
18 ExplainingCausalSelectionwithExplanatoryCausal
Economy:BiologyandBeyond............................................ 413
LauraR.Franklin-Hall
Chapter 1
Explanation in Biology: An Introduction
Pierre-AlainBraillardandChristopheMalaterre
Abstract Explanation in biology has long been characterized as being different
from explanation in other scientific disciplines, in particular from explanation in
physics.One ofthereasonswas theexistencein biologyof explanationtypesthat
wereunheardofinthephysicalsciences:teleologicalandfunctionalexplanations,
historical and evolutionary explanations. More recently, owing in part to the rise
of molecular biology, biological explanationshave been depicted as mechanisms.
Thisprofusionofexplanatorypatternsistypicalofbiology.Theaimofthepresent
volume Explanation in Biology. An Enquiry into the Diversity of Explanatory
PatternsintheLifeSciencesistoshedsomenewlightonthediversityofexplanation
modelsinbiology.Inthisintroductorychapter,werecallthegeneralphilosophical
contextof scientific explanationas it has unfolded in the past seven decades, and
highlightthe specific issues that models of explanationhave faced in biology.We
thenshowhowthedifferentessaysgatheredinthiscollectivevolumetackleaspects
ofthisimportantdebate.
Keywords Scientific explanation (cid:129) Biologicalexplanation (cid:129) Nomologicalexpla-
nation (cid:129) Causal explanation (cid:129) Mathematic explanation (cid:129) Mechanistic explana-
tion (cid:129) Explanatorypluralism
1 Introduction
Among the different achievements of science – such as prediction, control or
simply description – explanation is generally understood as occupying a very
centralandspeciallocationontheepistemicchessboard.Yet,explanationhasalso
P.-A.Braillard
IndependentScholar,Peyregrand,12350Drulhe,France
e-mail:[email protected]
C.Malaterre((cid:2))
Départementdephilosophie&CIRST,ChairederechercheUQAMenPhilosophiedessciences,
UniversitéduQuébecàMontréal(UQAM),Casepostale8888,SuccursaleCentre-Ville,
Montréal,QC,H3C3P8,Canada
e-mail:[email protected]
©SpringerScience+BusinessMediaDordrecht2015 1
P.-A.Braillard,C.Malaterre,ExplanationinBiology,History,Philosophy
andTheoryoftheLifeSciences11,DOI10.1007/978-94-017-9822-8_1
2 P.-A.BraillardandC.Malaterre
raised numerous questions and controversies among scientists and philosophers,
especially since the development of the deductive-nomological model (Hempel
and Oppenheim 1948; Hempel 1965). Though not everybody agrees, it is largely
assumed that it is part of the task of science to explain natural phenomena, and
that its methods enable it to do so incomparably better than common sense. But
then what is a scientific explanation? What distinguishes an explanation from
something that is not an explanation? And what distinguishes an explanation that
isscientificfromonethatisnot?Understandingwhatscientificexplanationsconsist
of has been one of the most central issues in the philosophy of science as it
has developed throughout the twentieth century, and more recently as analyzed
in the realm of biology. Part of the agenda has been to identify necessary and
sufficient criteria that would enable distinguishing genuine scientific explanations
from pseudo-explanations. Hence the issue is not only a matter of descriptive
adequacy(accountingforthepracticeofscience),butalsohasnormativeambitions.
Anotherstronglydebatedtopic is whetherthereexists a single modelof scientific
explanation or several different ones, possibly depending on specific explanatory
practices across disciplines. If what is accepted as an explanation varies across
disciplines,ifwhatissoughtafterinelaboratinganexplanationalsovaries,aswell
asthereasonswhyonepursuessuchanexplanatoryquest,thenwouldn’tallofthis
begoodreasonstobeskepticalabouttheexistenceofasinglemodelofexplanation?
Thisdebatedissue hasproventobe ofmuchrelevancein biology,bothinlightof
thespecificitiesofthismultifaceteddisciplinecomparedtophysics–thelatterbeing
oftentakenasadisciplineofreferenceinphilosophyofscience–andinlightofthe
heterogeneityofexplanatorypracticeswithinbiologyitself.Inthisseventhdecade
sinceHempelandOppenheim’s“StudiesintheLogicofExplanation”,thepresent
volume Explanation in Biology. An Enquiry into the Diversity of Explanatory
Patterns in the Life Sciences proposes a collection of essays that discuss some of
the most recent philosophicalperspectives of scientific explanationin light of the
specificitiesofmodernbiology,includingsucha broadrangeofsub-disciplinesas
molecularbiology,systemsbiology,evolutionarybiologyordevelopmentalbiology.
In this introductorychapter,our objective is to set the contextwithin which these
latestdevelopmentsonphilosophicalmodelsofexplanationhavetakenplace.The
chapter is organized as follows. We first provide a brief overview of the major
modelsof scientific explanationthat have been developedin the past six decades,
and their “universalist” aspirations (Sect. 2). We then review the key issues that
these models generally face in biology (Sect. 3). These issues are often taken as
good reasons for developing philosophical models of explanation that are more
narrowly tailored to the claimed specificity of biology and of its sub-disciplines;
incidentally,theyare also taken as goodreasonsagainstanyuniversalistmodelof
explanation.Theythereforeshapethedebate–whichisnotablysalientinbiology–
between“universalists”whoargueinfavorofaunifiedaccountofexplanation,and
“pluralists”whoargueinfavorofaprofusionofexplanatorypatterns.Wethenshow
(Sect.4)howthedifferentcontributionsofthevolumeaddressfacetsofthisimpor-
tant debate, be it by documenting a profusion of explanatory patterns, analyzing
the specific heuristics at work in biology, or critically assessing mechanistic-type
explanationsandexploringalternativeapproaches.
1 ExplanationinBiology:AnIntroduction 3
2 SixDecades ofScientificExplanation
Elaborating a model of scientific explanation is a long-lasting aim of philosophy
of science. However, accounting for what a scientific explanation really is has
turned out to be much more difficult than one might have initially expected. As
reviewedbySalmoninhisFourDecadesofScientificExplanation(Salmon1989),
the literature on the topic is vast, triggered in one way or another by Hempel and
Oppenheim’s(1948)essay.Twodecadeslater,scientificexplanationisstillamatter
ofmuchdebate.SinceHempel’sDN model,thegeneralquestionofcharacterizing
scientific explanation has branched out to address three types of problems that
have structured the debate in the past six decades, and still structure it today: (1)
Are there uniquecharacteristics to scientific explanations?Can we define a set of
necessary and sufficient conditions for scientific explanations? (2) Is causation a
primitive notion for scientific explanation? (or should causation be construed on
the basis of explanation?)And if yes, which account of causation is suitable? (3)
Whichroledoescontextplayinexplanation,ifany?Thedebatealsoconcernsthe
ways models can or cannot address canonical counter-examples that range from
flagpoleshadows(asan illustrationof explanatoryasymmetry)to hexedtable salt
(explanatoryrelevance),syphiliticmayors(lowprobabilityexplanations)andstorms
andbarometers(correlations).Thecurrentdebateaboutexplanationinbiologyhas
inherited from all of these questions. Our aim here is to map the general context
within which this debate arose, and to survey some of the most salient models of
scientificexplanation.
Typicallyaddressingproblemsof type(1)above,the modelofscientific expla-
nation that has been the most discussed and that has influenced nearly almost all
subsequent work is the deductive-nomological(DN) model, mainly developed by
CarlHempel(HempelandOppenheim1948;Hempel1965).Asiswellknown,the
DNmodelconstruesexplanationasconsistingofadeductiveargument,thepremises
of which must contain at least one law-like generalization. According to this
model, scientific explanation involves two epistemic elements: an explanandum–
thephenomenontobeexplained–thatappearsastheconclusionoftheargument,
andanexplanansthatconsistsofthesetofpremisesthatdotheexplaining.Several
criteria constrainthe explanandumand the explanans.In particular,the explanans
must deductively entail the explanandum, and the deduction must make essential
useofgenerallaws;theexplanansmustalsohaveempiricalcontent.Thedecisionto
defineexplanationasalogicalargumentinvolvinglaws(orlaw-likegeneralizations)
originatedin partfrom empiricist worriesaboutother conceptssuch as causation,
foundtobemetaphysicallytooloaded.Butofcourse,thewholeapproachdepends
on the ability to properlydefine what a law is, and this has provento be far from
straightforward(e.g.,Ayer1956;Dretske1977;Cartwright1980).Wewillseehow
this question is especially problematic in biology. Because many cases involving
statistical generalizations are not covered by the DN model (and most special
sciences are not based on deterministic laws), Hempel added an account of what
he called inductive-statistical(IS) explanations(Hempel1965).Similarly to a DN
