Odenbaugh&Griffith. Philosophy of Biology
Odenbaugh&Griffith. Philosophy of Biology
Odenbaugh&Griffith. Philosophy of Biology
Philosophy of Biology
First published Fri Jul 4, 2008; substantive revision Mon Jun 1, 2020
The growth of philosophical interest in biology over the past forty years reflects the increasing prominence
of the biological sciences in the same period. There is now an extensive literature on many different
biological topics, and it would be impossible to summarise this body of work in this single entry. Instead, this
entry sets out to explain what philosophy of biology is. Why does biology matter to philosophy and vice
versa? A list of the entries in the encyclopedia which address specific topics in the philosophy of biology is
provided at the end of the entry.
Three different kinds of philosophical inquiry fall under the general heading of philosophy of biology. First,
general theses in the philosophy of science are addressed in the context of biology. Second, conceptual
problems within biology itself are subjected to philosophical analysis. Third, appeals to biology are made in
discussions of traditional philosophical questions.
Philosophy of biology can also be subdivided by the particular areas of the life sciences with which it is
concerned. Biology is an extremely diverse set of disciplines, ranging from historical sciences such as
paleontology to engineering sciences such as biotechnology. Different philosophical issues occur in each
field. The latter part of the entry discusses how philosophers have approached some of the main disciplines
within biology.
In another important early debate philosophers set out to solve a conceptual problem within biology itself.
The concept of reproductive fitness is at the heart of evolutionary theory, but its status has always been
problematic. It has proved surprisingly hard for biologists to avoid the criticism that, “[i]f we try to make
laws of evolution in the strict sense we seem to reduce to tautologies. Thus suppose we say that even in
Andromeda ‘the fittest will survive’ we say nothing, for ‘fittest’ has to be defined in terms of ‘survival’”
(Smart 1959, 366). This came to be know as the “tautology problem”. Alexander Rosenberg and Mary B.
Williams argued that fitness is an irreducible primitive which derives its meaning from its place in an
axiomatic formulation of evolutionary theory (Rosenberg 1983; Sober 1984a; Williams and Rosenberg
1985). If correct, this would solve the tautology problem since axioms are often thought of as tautologous. In
the 1970’s the new generation of philosophers of biology offered a different solution to the tautology
problem in two steps. First, they began by arguing that fitness is a supervenient property of organisms: the
fitness of each particular organism is necessarily dependent on some specific set of physical characteristics
of the organism and its particular environment, but two organisms that have the same fitness may do so in
virtue very different sets of physical characteristics (Rosenberg 1978). Second, they argued that this
supervenient property is a propensity – which is a probabilistic disposition represented by a probability
distribution over possible numbers of offspring (Mills and Beatty 1979). Although fitness is defined in terms
of reproductive success, it is not a tautology that the fittest organisms have the most offspring, any more than
it is a tautology that dice produce even numbers more often than they produce sixes. The propensities of fit
organisms to survive and of dice to fall equally often on each side both allow us to make fallible predictions
about what will happen, predictions that become more reliable as the size of the sample increases. It remains
unclear, however, whether it is possible to specify a probability distribution or set of distributions that can
play all the roles actually played by fitness in population biology (Beatty & Finsen 1989, Sober 2001; Pence
& Ramsey 2013).
The phrase ‘conceptual problems’ should be understood very broadly. The conceptual work done by
philosophers of biology in many cases merges smoothly into theoretical biology. It also sometimes leads
philosophers to examine and criticize the chains of argument constructed by biologists, and thus to enter
directly into ongoing biological debates. In the same way, the first kind of philosophy of biology we have
described – the use of biological examples to work through general issues in the philosophy of science –
sometimes feeds back into biology itself through specific recommendations for improving biological
methodology. It is a striking feature of the philosophy of biology literature that philosophers often publish in
biology journals and that biologists often contribute to philosophy of biology journals. The philosophy of
biology also has a potentially important role as a mediator between biology and society. Popular
representations of biology derive broad lessons from large swathes of experimental findings and theoretical
work. Philosophers of science have an obvious role in evaluating these interpretations of the significance of
specific biological findings (Stotz and Griffiths 2008). As two important examples, philosophers of biology
have provided a great deal of clarity regarding creationism/intelligent design (Kitcher 1982; Ruse 1982;
Pennock 2000; Sarkar 2007) and sociobiology/evolutionary psychology (Kitcher 1985; Buller 2006;
Richardson 2010; Barker 2015).
A third form of philosophy of biology occurs when philosophers appeal to biology to support positions on
traditional philosophical topics, such as ethics or epistemology. The extensive literature on biological
teleology is a case in point. After a brief flurry of interest in the wake of the “modern synthesis”, during
which the term ‘teleonomy’ was introduced to denote the specifically evolutionary interpretation of
teleological language (Pittendrigh 1958), the ideas of function and goal directedness came to be regarded as
relatively unproblematic by evolutionary biologists. In the 1970s, however, philosophers started to look to
biology to provide a solid, scientific basis for normative concepts, such as illness, disorder, or malfunction
(Wimsatt 1972; Wright 1973; Boorse 1976). Eventually, the philosophical debate produced an analysis of
teleological language fundamentally similar to the view associated with modern synthesis (Millikan 1984;
Neander 1991; Godfrey-Smith 1994). According to the “selected effects” theory of function, the functions of
a trait are those activities in virtue of which the trait was selected. The idea of proper function has become
part of the conceptual toolkit of philosophy in general and of the philosophy of language and the philosophy
of mind in particular (Dretske 1991, 1997; Millikan 1995, 2004, 2005; Papineau 1987, 1993; Neander 2017;
Garson 2019).
The intense philosophical interest in evolutionary theory in the 1980’s can partly be explained by two
controversies (Segerstråle 2000). First, there was a controversy over “sociobiology” that was provoked by
the publications of E.O. Wilson’s eponymous textbook (Wilson 1975) and his Pulitzer prize winning popular
book (Wilson 1978). One extremely important critique of this application of evolutionary biology to human
social behavior came from Stephen Jay Gould and Richard Lewontin (Gould and Lewontin 1979). The
debates over adaptationism turned out to involve a diffuse set of worries about whether evolution produces
adaptations, the role of optimality models, and the methodology of evolutionary theory (Amundson 1994;
Orzack and Sober 1994; Brandon and Rausher 1996; Godfrey-Smith 2001; Millstein 2007; Forber 2009;
Potochnik 2009; Lloyd 2015). Philosophical work has helped to distinguish these strands in the debate and
reduce the confusion seen in the heated and polemical biological literature for and against adaptationism
(Orzack and Sober 2001). Second, there was the appearance of George Williams’ Adaptation and Natural
Selection (Williams 1966) and Richard Dawkins’ The Selfish Gene (Dawkins 1976). They claim that the unit
of selection is the individual Mendelian allele rather than the organism, group of organisms, or species. This
created an explosion of early philosophical work on the “units of selection” question (Brandon and Burian
1984). The early debates concerned whether there was a determinate unit of selection and what criteria
should be used for determining what it is in a given case (Hull 1980; Wimsatt 1980a, 1980b; Brandon 1982;
Sober & Lewontin 1984; Lloyd 1988; Sterelny & Kitcher 1988). As multi-level selection models and the
Price equation appeared, close attention turned to how those models should be interpreted (Godfrey-Smith &
Lewontin 1993; Godfrey-Smith & Kerr 2002; Lloyd 2005; Waters 2005; Okasha 2006; Birch 2017). For
example, for any group selection model, is there an empirically equivalent individual selection model
(similarly for genotypic and genic models)? If so, do they represent the very same causal structure? Is kin
selection a form of group selection? Arguably, philosophers made a significant contribution to the
rehabilitation of some forms of “group selection” within evolutionary biology in the 1990s, following two
decades of neglect or contempt (Sober and Wilson 1998).
The biologist Michael Ghiselin piqued the interest of philosophers when he suggested that systematics was
fundamentally mistaken about the ontological status of biological species (Ghiselin 1974). Species were not
natural kinds in the way that chemical elements are. Instead, they are historical particulars like nations or
galaxies. They have a beginning through speciation, they have parts that are integrated over time by
biological relations, and they cease to exist by going extinct (Mishler & Brandon 1987). Additionally,
individual organisms are not instances of species, as a wedding ring is an instance of gold. Instead, they are
parts of species, as one is a part of a family. As Smart had earlier noticed, this has the implication that there
can be no “laws of nature” about biological species per se, at least in the traditional sense of ‘laws of nature’
(Smart 1959). David Hull further developed and argued for the “species as individuals” thesis. He explored
its implications for a variety of topics including species names, laws of nature, and human nature (Hull 1976,
1978, 1986).
However, the view that species are individuals leaves other important questions about species unsolved and
raises new problems of its own (Kitcher 1984, 1989). For example, suppose you are a population biologist
and you need to census the individuals in a population of species. How do you decide what entities to count?
Philosophers and biologists have provided a variety of criteria for answering questions like this including
reproduction, life-cycles, genetics, sex, developmental bottlenecks, germ-soma separation, policing
mechanisms, spatial boundaries or contiguity, immune response, fitness maximization, cooperation and/or
conflict, codispersal, adaptations, metabolic autonomy, and functional integration (Clarke 2013). So, the
question of what a biological individual is indeed a pressing one (Wilson 2005; Okasha 2006; Clarke 2011;
Pradeu 2012; Bouchard and Huneman 2013; Clarke 2013; Godfrey-Smith 2013; Wilson and Barker 2013).
Biologists have been, and are still, deeply divided over the species category (Ereshefsky 1992b; Wheeler and
Meier 2000; Coyne and Orr 2004; Wilkins 2009). Consider Ernst Mayr’s famous biological species concept
(BSC). He writes, “Species are groups of interbreeding natural populations that are reproductively isolated
from other such groups” (Mayr 1963, 89). There are lots of concerns regarding the BSC (Ehrlich and Raven
1969; Sokal and Crovello 1970; Van Valen 1976; Wiley 1978). Asexual organisms do not interbreed. Thus,
on the BSC, there are no species of asexual organisms. Many species exhibit some introgression, and thus
cannot be distinct species. Last, the BSC is extremely difficult to apply to species in the fossil record since
sex organs do not normally fossilize, and reproductive behavior is difficult to corroborate. In light of the
problems with the BSC (and for other reasons), biologists have put forward other species concepts. Species
pluralism is the claim that there is no single correct species concept that classifies organisms exactly the
same (Ereshefsky 1992a); rather, there are several correct species concepts. That is, for some organisms,
different species concepts will correctly place them in distinct species. Species monism is the claim that there
is a single correct species concept. Some allege species pluralism is temporary because we will eventually
find the single best concept (Hull 1999). The debate between pluralists and monists rages on (Wilson 1999).
Biological species are often given as one of the classic examples of a “natural kind”. The philosophy of
systematics has had a major influence on recent work on classification and natural kinds (Dupré 1993, 2002;
Wilson et. al. 1997; Boyd 1999; Griffiths 1999; Wilson 1999; Okasha 2001; Walsh 2006). For example, can
there be a notion of natural kind or essentialism consistent with the “population thinking” found in biology
(Mayr 1975, Sober 1980; Ariew 2008)?
Another important topic in the philosophy of molecular biology has been the concept of the gene (Beurton,
Falk and Rheinberger 2000; Waters 2000, 2004; Griffiths and Stotz 2007). Philosophers have also written
extensively on the concept of genetic information, the general tenor of the literature being that it is difficult
to reconstruct this idea precisely in a way that does justice to the apparent weight placed on it by molecular
biologists (Sarkar 1996; Maynard Smith 2000; Godfrey-Smith 2001; Griffiths 2001; Jablonka 2002;
Rosenberg 2006). For example, does DNA actually carry semantic information (Shea 2007)? If it does not, is
this a useful fiction, or is it hindering molecular biology (Levy 2001)?
Philosophers have started to remedy the neglect of ecology and a number of major books have appeared
(Cooper 2003, Ginzburg and Colyvan 2004, Sarkar 2005, MacLaurin and Sterelny 2008). The discussions
have focused on several topics including the complicated and sometimes troubled relationship between
mathematical models and empirical data in ecology (Shrader-Frechette & McCoy 1993; Ginzburg and
Colyvan 2004; Odenbaugh 2005; Weisberg 2012), whether there are distinctive ecological laws (Cooper
2003; Mikkelson 2003; Lange 2005; Eliot 2011b, Linquist et. al. 2016), the nature and reality of ecological
communities and ecosystems (Sterelny 2006; Odenbaugh 2007; Eliot 2011), the “robustness“ of ecological
principles (Odenbaugh 2003; Weisberg & Reisman 2008; Justus 2012), the idea of ecological stability and
the “balance of nature” (Odenbaugh 2001; Cooper 2003; Mikkelson 2001; Justus 2008), the definition of
biodiversity (Sarkar 2005; MacLaurin and Sterelny 2008; Santana 2014), and the relationship between
ecology and conservation biology (Linquist 2008). Recently, there has been interesting work done on
functions in ecology as well (Jax, 2005; Odenbaugh 2010; Nunes-Neto et. al. 2014; Dussault & Bouchard
2017). Most ecologists and evolutionary biologists do not think communities or ecosystems are units of
selection, and thus the selected-effects account of functions does not readily apply. So, philosophers of
ecology have been exploring alternatives.
Even the distinction between the questions of biology and those of philosophy of biology is not absolutely
clear. As noted above, philosophers of biology address three types of questions: general questions about the
nature of science, conceptual problems within biology, and traditional philosophical questions that seem
open to illumination from the biosciences. When addressing the second sort of question, there is no clear
distinction between philosophy of biology and theoretical biology. But while this can lead to the accusation
that philosophers of biology have abandoned their calling for ‘science reporting’ it can equally well be said
that a book like The Selfish Gene (Dawkins 1976) is primarily a contribution to philosophical discussion of
biology. Certainly, the professional skills of the philosopher are as relevant to these internal conceptual
puzzles as they are to the other two types of question. All three types of question can be related to the
specific findings of the biological sciences only by complex chains of argument.
There is a great deal of new work being in done in the philosophy of biology. As examples, there is a rich
philosophical literature emerging around cancer (Plutynski 2018), cultural evolution (Sterelny 2012; Lewens
2015; O’Connor 2019), human nature (Machery 2008; Ramsey 2013; Kronfeldner 2018), microbiology
(O’Malley 2014), and paleobiology (Turner 2011; Currie 2018). Philosophy of biology still remains one of
the most dynamic and interesting areas of the philosophy of science and philosophy more generally.
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