5.a. Teleology (History of The Concept Naturalization Naturalized Organism) (Walsh 2008)

Teleology
Teleology
D.M. Walsh
The Oxford Handbook of Philosophy of Biology
Edited by Michael Ruse
Print Publication Date: Jul 2008 Subject: Philosophy, Philosophy of Science, Metaphysics
Online Publication Date: Sep 2009 DOI: 10.1093/oxfordhb/9780195182057.003.0006
Abstract and Keywords
Teleology is a mode of explanation in which the presence, occurrence, or nature of some

phenomenon is explained by the end to which it contributes. The model of explanation is
“pure mechanism” which holds that there is a single kind of stuff in the world-”matter”
that exhibits a single kind of change, motion. It falls into three classes: the argument
from nonactuality, the argument from intentionality, and the argument from normativity.
These objections are because of early modern thinking about the natural world. These ar
guments rely on the Platonic model of transcendent teleology. Aristotelian teleology com
plements our current understanding of goal-directed, self-organizing, adaptive systems.
The success of development can be explained by plasticity which is a goal-directed capac
ity of organisms to produce and maintain a stable, well-functioning living thing. The un
derstanding of how evolution can be adaptive requires us to incorporate teleology.
Keywords: model, mechanism, argument, teleology, evolution, plasticity
Introduction
Teleology is a mode of explanation in which the presence, occurrence, or nature of some
phenomenon is explained by appeal to the goal or end to which it contributes. As an ex
planatory strategy, teleology has fallen from favor in modern times. While it retained
some currency within some branches of comparative biology, teleological thinking was
purged from the most fundamental of sciences—physics—during the scientific revolution.
Much of twentieth‐century philosophy of science, guided by reductionism and the unity of
science hypothesis (Oppenheim and Putnam 1958), took it on faith that a mode of expla
nation deemed illegitimate in physics should not find a home in any of the special sci
ences.
Yet, the role and status of teleology in biology has continued to attract considerable atten
tion. There are two general questions to be addressed with respect to biological teleolo
gy: whether teleological explanation might constitute a legitimate part of a scientist's
conceptual tool kit, and whether evolutionary biology is committed to unreduced teleolog
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Teleology
ical explanation. Received opinion answers ‘no’ to both; teleology has no legitimate place
in the natural sciences, and that goes as much for biology as for physics. But I think that
received opinion is wrong. Unreduced teleology is a legitimate, wholly natural explanato
ry mode; moreover, it is not only available to evolutionary biology, it is indispensable. Or
thodox opinion is mistaken, I believe, both in its characterization of teleological explana
tion and in its account of teleology's application to biology. Teleological explanation, I
shall argue, does not have the antinaturalistic implications commonly imputed to it. It is
naturalistically acceptable, metaphysically unimpeachable, and explanatorily au
tonomous. Be that as it may, the naturalistic credentials of teleological explanations
(p. 114) do not oblige biologists to deploy them. The case for biological teleology rests up
on the demonstration that it plays some ineliminable role in evolutionary biology. There is
general agreement that it doesn't. I claim that biologists and philosophers of biology have
failed to find any unreduced teleological commitment in biology because they have been
looking in the wrong place. Recent developmental biology, I suggest, illuminates the ine
liminable role for unreduced teleology in evolutionary explanation.
1. Barren Virgins and Mistresses

The model of explanation that has come to dominate the natural sciences since the ad
vent of the scientific revolution is pure mechanism.1 As C. D. Broad (1925) characterizes
it, “pure mechanism” holds that there is a single kind of stuff in the world, matter, that
exhibits a single kind of change, motion. Changes in motion are regulated and governed
by a single, unified set of fundamental laws that govern all mechanical interactions. There
is, moreover, “a simple principle of composition, according to which the behaviour of any
aggregate of particles, or the influence of any one aggregate on any other, follows in a
uniform way from the mutual influences of the constituent particles taken by
pairs” (Broad 1925, 45). To explain a natural phenomenon—a change in motion—in a sys
tem, one simply adverts to the causal, mechanical relations that hold among the parts of
the system. These are sufficient to necessitate any observed change, and hence to explain
it.
On the mechanical world view, teleological explanations aren't merely otiose; they are at
once fatuous and steeped in an occult metaphysics of ends and goals. Scientific revolu
tion and Enlightenment thinkers took great pains to expose teleology as an intellectual
imposter. Molière satirizes the teleology of his day as engaged in “dormitive virtue” expla
nations. Francis Bacon derides final causes as “barren virgins dedicated to God.” Spinoza
(Ethics, appendix 1) complains that “this doctrine concerning the end turns Nature com
pletely upside down. For what is really a cause it considers as an effect, and conversely,
[what is an effect it considers as a cause]” (quoted in Curley 1994, 112). Spinoza's princi
pal influence in this regard is Descartes (Manning 2006). According to Descartes, pure
mechanism suffices for biology because nonhuman organisms are bête machines.
The number and the orderly arrangements of the nerves, veins, bones, and other
parts of an animal do not show that nature is insufficient to form them, provided
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Teleology
you suppose that in everything nature acts in accordance with the laws of mechan
ics. (Letter to Mersenne, 1639)2
Despite the influence of mechanistic philosophy, and despite its open antipathy toward
teleology, teleological thinking was not entirely extirpated from early (p. 115) modern sci
ence. It seems to have held a tenacious grip on biology that no amount of derision or oblo
quy could loosen. There is good reason for this. Uniquely, biology is concerned with a do
main of entities—organisms—for which teleological explanation seems perfectly suited.
Buffon's characterization of organisms is most apt in this respect. Organisms, according
to Buffon, are distinguished by three capacities: (i) the capacity to self‐organize, i.e., to
make adaptive responses to the conditions of their existence, (ii) to self‐reproduce, and
(iii) to self‐nourish. The last of these incorporates the capacity that we now recognize as
metabolism, to synthesize the very material out of which organisms themselves are con
structed (McLaughlin 2000). Because organisms are both the agents and the conse
quences of these reflexive, goal‐directed processes, their activities cannot be accounted
for by pure mechanism alone. Teleology is needed to account for the distinctive charac
teristics of organisms.
Kant's Third Critique vividly articulates the tension between mechanistic and teleological
thinking in biology. According to Kant ([1793]2000), organisms must be, and yet cannot
be, judged to be the results of mechanical causes alone. Organisms must be the results of
mechanical processes, according to Kant, because they are natural entities, and natural
entities are wholly susceptible of mechanical explanation. We explain the properties of
complex entities by appeal to the causal powers of their parts and the mechanical laws of
nature. Yet Kant believes biology cannot do without teleology because we must acknowl
edge that organisms are “natural purposes.” They are self‐organizing, self‐nourishing en
tities. Each part of an organism is manufactured by the organism itself and has the prop
erties it has precisely because of the goals of the organism that it subserves. Organisms
are such functionally integrated entities that the nature of each part is explained by the
overall functioning of the organism, and the function of the organism is realized by and
explained by the parts. Organisms, Kant tells us, are both causes and effects of their
parts (Zammito 1991). Mechanical explanation accounts only for organisms as effects of
their parts. Teleological explanation is needed to account for how organisms build, modi
fy, and regulate their parts. Alas, Kant repines, teleological explanation is not applicable
to the natural world. Kant's discomfiture derives from his very modern conviction that
causal/mechanical explanation is the only naturalistically acceptable mode of explanation.
It is tempting to suppose that Darwin's theory of evolution should have alleviated any lin
gering anxiety about biological teleology. Darwin's theory identifies a wholly mechanical
process, natural selection, that has as a simple causal consequence that organisms have
the fantastic features that so impressed Buffon and perplexed Kant. Mayr remarks that
“Darwin had solved Kant's great puzzle” (Mayr 1988, 58). Michael Ghiselin (1993) called
teleology “a metaphysical delusion” for which Darwin's theory of natural selection is the
cure. Von Helmholtz ([1869]1971) praised Darwin for bringing the study of biological
form under the ambit mechanism. Considerations such as these led David Hull to declare:
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Teleology
“From the point of view of contemporary biology, both vitalism and teleology are stone‐
cold dead” (1969, 249).3 But even committed Darwinians continue to feel the Kantian ten
sion (p. 116) between the requirement of teleology and its unavailability. J. B. S. Haldane is
said to have quipped that teleology, to a biologist, is like a mistress. He can't be without
her, but he is ashamed to be seen with her in public. The claim is as apt as it is inappro
priate. Biology seems to be inexorably drawn to teleology, and yet its credentials as a sci
ence in good standing appear to require that either it renounce its dalliance with teleolo
gy or make a legitimate partner of it. Biology, it seems, can't live with teleology, and it
can't live without it.
2. Teleological Antinaturalism
A battery of standard objections suggests that teleology will never attain naturalistic re
spectability. They fall into three classes: the argument from nonactuality, the argument
from intentionality, and the argument from normativity.
The argument from nonactuality
Teleological explanations advert to goals. If goals explain, and the measure of a success
ful teleological explanation is that the presence of the item to be explained is accounted
for by showing how it conduces to—i.e., causes—the fulfillment of the goal, then the pres
ence of the item must be explained by some unactualized state of affairs. Means precede
their ends, but in teleology, ends explain their means. Teleology requires the positing of
backward causation or some occult form of causation by nonactual states of affairs. This
is the thought that gives impetus to Spinoza's “upside‐down” complaint.
The argument from intentionality
Unactualized goals may not be able to explain, but representations of unactualized goals
can. One way to avoid the commitment to nonactualia is to suppose that teleological ex
planation requires intentionality, the representation of goals. Our paradigm of successful
teleological explanation, after all, is psychological. The occurrences of actions are ex
plained by the intentions of agents, and in the same way, the features of artifacts are ex
plained by the intentions of artisans. The conviction that all teleological explanation re
quires intentionality is evident in thinkers as diverse as Aquinas and Kant and informs
modern‐day intelligent design theory. Aquinas (2006) says:
We see that things which lack intelligence, such as natural bodies, act for an end,
and this is evident from their acting always, or nearly always, in the same way,
so as to obtain the best result. Hence it is plain that not fortuitously, but
(p. 117)
designedly, do they achieve their end. Now whatever lacks intelligence cannot
move towards an end, unless it be directed by some being endowed with knowl
edge and intelligence; as the arrow is shot to its mark by the archer. Therefore
some intelligent being exists by whom all natural things are directed to their end.4
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Teleology
Kant, in reference to the apparent purposiveness of organisms, tells us:
We cannot conceive of the purposiveness which must be made the basis of our
cognition of the internal possibility of many things in nature and make it compre
hensible except by representing them and the world in general as a product of an
intelligent cause. ([1793]2000, 5:401–5)5
Of course, from the naturalist's point of view, the world is not the product of anyone's in
tentions. The natural world has no designer and it is retrograde to think otherwise. Teleo
logical explanation has no place in the natural sciences.
The argument from normativity
While the argument from nonactuality and the argument from intentionality suggest that
no natural system could fulfill the conditions required for offering a teleological explana
tion, a third argument, the argument from normativity, suggests that no natural state of
affairs could underwrite the presumed consequences of teleological explanations. Teleo
logical explanation appears to have normative import. Here again, the paradigm of inten
tional explanation is illustrative (Taylor 1963). There is a rationality constraint upon inten
tional explanation (Davidson 1963). To explain an action as the consequence of an agent's
intention is to demonstrate that the agent was rationally required (or at minimum ratio
nally permitted) to commit the act, given her goals. Generalizing the model, a teleological
explanation must explain that, given the system's pursuit of γ as a goal, and given that φ
is a means to the attainment of γ, the system ought to have produced φ (Stout 1996).
There are thought to be two consequences of this model of teleological explanation: (i) it
is the status of the goal state qua goal that explains, and (ii) teleological explanations are
normative by their very nature. Marc Bedau (1991, 1992) has argued that because of the
normativity of teleological explanation, goals can only play their presumed explanatory
role in teleology if goals themselves are intrinsically normatively evaluable: φ construed
as the means toward the attainment of some goal γ could only be something that the sys
tem ought to produce if γ is a state that the system ought to attain, but γ could not be a
state that the system ought to attain unless γ were good. Typically, naturalists are loath to
admit evaluative states of affairs into the natural world, so genuine teleological explana
tion is proscribed.6
(p. 118) 3. Transcendence and Immanence

Taken together, this battery of standard arguments mounts a redoubtable challenge to
the naturalistic status of teleological explanation. Nevertheless, by observing a distinc
tion between two formative conceptions of teleology, represented, respectively, by Plato
and Aristotle, these challenges can be deflected.
The battery of standard arguments issues from a presupposition of the Platonic concep
tion of teleology. Plato's extended discussion of teleology in the Timaeus is the locus clas
sicus of what we might call “transcendent” or “extrinsic” teleology. In the Timaeus, Plato
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Teleology
argues that teleology is the mark of craftsmanship. The magnificent order of the world re
quires the work of the demiurge. Mere material necessity (the “wandering cause”) is suf
ficient by itself only to produce chaos. Order must be imposed on recalcitrant nature. The
order of nature is a reflection of the design of the demiurge, in just the way that the func
tion of artifacts is a reflection of the design of an artisan. The distinguishing feature of
Platonic teleology is that the goals that explain the features of the explanandum are al
ways extrinsic to the explanandum itself, and they are intentional.
The Platonic conception of teleology lends support to standard antiteleological objections.

Platonic teleology is committed to “occult,” unactualized ends the intentionality of teleol
ogy, and the normativity of teleological explanation. The regularities of the world are re
flections of nonnatural, perfect forms. They are the products of the demiurge's represen
tations of these forms. The world is the way it ought to be given the intentions of the
demiurge (and the constraints of recalcitrant matter).7
Extrinsic or transcendental teleology of this sort exerts a significant hold on our imagina
tions and for good reason. It is probably an indispensable explanatory device. It is the
mode of explanation by which we account for the occurrences of actions and the features
of artifacts. To engage in action explanation, or design explanation, is to commit to a form
of extrinsic teleology. Though it is indispensable, extrinsic teleology is by no means oblig
atory. It doesn't follow from the fact that some phenomena call for an extrinsic teleologi
cal explanation that all genuine teleological phenomena do. The other great model of tele
ology from antiquity demonstrates this.
Where Plato's teleology is transcendent and supernatural, Aristotle's is immanent and

natural. By “immanent” is meant that the goals that explain the parts and activities of a
system are goals pursued by the system itself. The import of immanent teleology is most
clearly seen in Aristotle's biological works. Aristotle (1995) identifies the nature of a sub
stance with its form, a principle of change or stasis. He tells us (in De Anima) that the na
ture, form, of an organism is soul (psuche):
We may think of the form of an organism as a set of organizing principles, or a set

of goal‐directed dispositions, to organize its matter in such a way that the organ
ism is capable of performing particular soul functions (in the particular way) dis
tinctive of its kind. (Lennox 2001, 183)
On Aristotle's view, the nature or, as we might say, “essence” of an organism consists in
(p. 119)
an “interactive unity” between the matter of which it is composed and its goal‐directed form
(Charles 2000).8 The matter of which an organism is constituted is organized by the form. At the
same time, the form of an organism is realized in, and constrained by, its matter. In biology, as in
any other natural science, one must know both the form and the matter.
The important point for our purposes is that both the form—the goal‐directed capacity of
an organism to organize matter—and the matter itself inhere in the nature of an organ
ism. According to Aristotle, the distinctive feature of any organism is its capacity to enlist
the powers of its constituent matter in the attainment of its goals of producing and main
taining a well‐functioning organism typical of its kind. The presence and nature of an
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Teleology
organism's salient features, and indeed the distinctive capacities of the organism itself,
are explained by appeal to its pursuit of its own goals. The goal‐directedness of organisms
confers on them the capacity to produce those structures and processes required for their
vital functions. According to Aristotle, goal‐directedness explains the salient features of
organisms.
The Aristotelian conception of immanent teleology doesn't fall prey to the standard bat
tery of antinaturalist objections.
The argument from nonactuality
It is goal‐directedness, not unactualized goals, that explains the presence of traits in an

organism. Goal‐directedness is an intrinsic property of a system manifested in its behav
ior. It is an occurrent, dispositional, empirically observable property of things. Teleologi
cal explanation requires no appeal to occult nonactualia.
The argument from intentionality
It is true that the paradigm of successful teleological explanation appeals to the inten
tions of an agent. Intentions are representations of an agent's goals. They are also mani
fested as an agent's goal‐directed activity. It is underdetermined, though, which of these
aspects of intentions—the intentionality or the goal‐directedness—does the explanatory
work. The Aristotelian conception of teleology suggests that it is the goal‐directedness
that takes explanatory priority. The reason is that teleological explanation, Aristotle as
serts, applies just as well to nonhuman organisms as it does to rational agents. Nonhu
man organisms are not intentional agents, but are goal‐directed nonetheless. As Aristotle
tells us, we do not need to observe intentionality to apply a teleological explanation: “It is
absurd to suppose that purpose is not present because we do not observe the agent
deliberating” (Aristotle 1970, Physics II.8).
The extrinsic teleology of action and artifact explanations is merely a special case. The
goal‐directedness of a system can be employed to explain features of the (p. 120) system
itself (intrinsic teleology) or of some other thing that it produces (extrinsic teleology).
Again, the only condition required for the proper use of teleological explanation is goal‐di
rectedness.
The argument from normativity
The argument from normativity incorporates two distinct claims. In a genuinely teleologi
cal explanation, (a) a goal's being a goal must enter into the explanation; and (b) the eval
uative status of the goal (i.e., its being good) explains why the system ought to act in a
way that brings about the goal. Aristotelian teleology suggests a naturalistic response to
each of these. Against (a), as we have seen, an Aristotelian teleological explanation does
not explain the presence of a feature by appeal to an unactualized goal, but rather to oc
current, actualized, goal‐directed activity. As for (b), what it is for a state of affairs to be a
goal is neither normative nor evaluative. A goal is simply the state to which a goal‐direct
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Teleology
ed process tends. Teleology does not require a category of value‐bearing goal states; it
only requires goal‐directedness.
But how does the goal‐directedness of a system explain what the system ought to do? The
quick answer is that it doesn't. On the Aristotelian conception, teleological explanation
per se has no normative implications. I realize that this dismissal of the argument from
normativity is a little facile. The whole issue of normativity arises, after all, from the fact
that our paradigm cases of teleological explanation clearly do have normative implica
tions; action explanations are rationalizing. It is a constraint on a good psychological ex
planation that it demonstrate to us that the agent ought to have done what she did, given
her goals (Davidson 1963). If so, the appeal to the goal‐directedness of action, at least,
must tell us what an agent ought to do. If rationalizing explanation is the paradigm of
teleological explanations, how can rationalizing explanation be normative if teleological
explanations aren't? The key to the answer lies in the distinction between two senses of
“ought.”
There are two distinct, but rarely differentiated, senses of “ought”. After John Broome
(1999), I shall call them “ought” and “normative requirement.” There are things we ought
to do simpliciter—we ought (simpliciter) to keep our promises—and there are things we
are normatively required to do. We are normatively required to bring about the means to
the attainment of our goals. This is a constraint on rational action, and it holds whether
or not the goals are goals we ought to have. For example, there is no fact of the matter, I
take it, whether one ought to prefer to watch FC Barcelona over Real Madrid, but given
an intention to watch Barcelona rather than Real, one ought to go to Camp Nou rather
than Bernabéu. That is to say, one is normatively required by the goal of watching
Barcelona to go to Camp Nou, whether or not one ought (simpliciter) to go.
(p. 121)
The ought/normative requirement distinction has two important implications. The first is
that even in the paradigm cases of teleological explanation where we accept that the tele
ology has normative import—i.e., the explanation of actions—it does not follow that the
normativity depends upon there being an intrinsically evaluable goal (or for that matter
an intrinsically evaluable explanans of any kind). Arguably, an agent ought to adopt x as a
goal only if x is intrinsically good. But whether an agent is normatively required to per
form an action can be assessed quite independently of the evaluative status of the goal: If
agent a has goal γ, and φ is a (or the only) means to the attainment of γ, then the agent is
normatively required to do φ, whether or not she ought to bring about γ. When we offer a
rationalizing, teleological explanation of an agent's actions, we demonstrate only that the
action was normatively required, given the agent's goals, and not that the goal is intrinsi
cally normatively evaluable (i.e., “good”).
The second implication of the ought/normative requirement distinction is that it allows us

consistently to hold that teleological explanations of an agent's actions do have normative
implications while teleological explanations per se do not. The relation of normative re
quirement that holds between an agent's goals and the means toward their attainment is
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Teleology
a special case of a general relation between means and goals that Aristotle called “hypo
thetical necessity” (Cooper 1987). On Aristotle's conception of teleological explanation,
we explain the presence of some phenomenon teleologically if it is (i) a consequence of
goal‐directed activity, and (ii) hypothetically necessary for the attainment of the goal: φ is
hypothetically necessary for γ just in case γ is a goal and φ is necessary for the attainment
of γ under the circumstances. The relation of normative requirement, by which we gauge
the rationality of actions, has the same form: Agent a is normatively required to do φ just
if γ is a goal (i.e., she intends to bring about γ) and φ is necessary for the attainment of γ.
Just as we can judge the appropriateness of a biological trait by how well it conduces to
the fulfillment of a biological goal, we can judge the appropriateness of an action accord
ing to whether it conduces to the fulfillment of an intentional goal. In either case, these
judgments are simply judgments about the hypothetical necessity of the explanandum.
Normative requirement is just the relation of hypothetical necessity that holds between
an agent's intended goals and the means to their fulfillment. Hypothetical necessity per
se has no normative implications; it introduces normativity only when it is applied to ra
tional action. Teleological explanations of action, then, are genuinely normative whereas
teleological explanation per se is not. The supposition that all teleological explanations
are irreducibly normative is simply an artifact of an overgeneralization of the intentional
model.
The Aristotelian model of immanent teleology allows the naturalist to resist the standard
battery of antiteleological arguments. Because teleological explanation is explanation that
appeals to goal‐directedness, all the Aristotelian model needs is an account of goal‐direct
edness that is both natural and sufficiently strong to demonstrate that a phenomenon's
being hypothetically necessary for the fulfillment of a goal can explain its occurrence.
4. Explanatory Sufficiency, Explanatory Au

(p. 122)
tonomy
To naturalize a phenomenon, one needs to show its place in the causal order of the world.
Naturalized immanent teleology, then, must demonstrate how goal‐directedness is a
causal consequence of unproblematic, natural phenomena. The commitment to natural
ized teleology, however, is more than merely a commitment to the existence in the world
of goal‐directed processes. It also involves a commitment to an irreducible explanatory
role for goal‐directedness. Here is a potential pitfall for naturalized teleology. There is a
tension between showing how some phenomenon is caused or causally realized, on the
one hand, and demonstrating its explanatory autonomy, on the other. It is tempting to
suppose that everything explained by some phenomenon x is wholly explained by the
causes of x. 9 Teleological explanation would be rendered otiose if everything explained by
goal‐directedness could also be explained by the causes of goal‐directedness. Naturaliz
ing teleology, then, treads a fine line between identifying the causes of goal‐directedness
and at the same time showing that goal‐directedness explains phenomena that its causes
cannot.
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The first task is quite straightforward; goal‐directedness is now no mystery. The mecha
nisms generating goal‐directed activity have been the subject of a considerable amount of
empirical research. An initial wave of interest in goal‐directedness originated in the cy
bernetics research of the 1940s–1960s. Later research into complex, adaptive, self‐orga
nizing systems (Kauffman 1995) has supplemented this understanding. Such systems are
capable of attaining and maintaining robustly persistent states by the implementation of
complex, adaptive, compensatory changes (Rosenbleuth, Wiener, and Bigelow 1943; Som
merhof 1950). Complex adaptive systems are capable of “supple adaptation”, the capacity
to pursue a goal state, and can sustain that state despite perturbations (Kitano 2004).10
Typically, an adaptive system is capable of implementing changes to its component
processes in a way that corrects for the effects of perturbations that might tend to sway
the system from its goal, or stable end‐state.
The distinguishing features of goal‐directed systems are architectural. Such systems typi
cally comprise arrangements of modules. Modules are clusters of causally integrated
processes that are decoupled from other such modules. Modules exhibit a capacity to pro
duce and maintain their integrated activities across a range of perturbations and influ
ences; they are robust (Kitano 2004). Each module exerts regulatory influence, in the
form of positive or negative feedback, over a small number of other modules. Complex,
self‐organizing systems are usually arranged as hierarchies of modules. Kauffman (1993)
identifies a general architectural constraint that must be met by such systems; “each part
must impinge on rather few other parts” (67). The consequence of this architectural
structure is that complex adaptive systems are buffered; they are capable of maintaining
their (p. 123) functional integrity across a range of perturbations. They also exhibit plas
ticity, the capacity to maintain stability in the face of perturbations by implementing nov
el, stable, adaptive responses. Robustness and the capacity to accommodate in novel
ways are the hallmarks of complex adaptive systems. Organisms, of course, are the very
paradigms of complex, adaptive, self‐organizing systems. Unsurprisingly, the principal ar
chitectural feature that confers this property on organisms is the modularity of their de
velopment (Schlosser 2004).
Goal‐directedness is an unproblematic causal consequence of the architecture of an adap

tive system. It consists in the capacity of a system as a whole to enlist the causal capaci
ties of the parts and direct them toward a stable homeostatic end‐point. Teleology's re
liance upon goal‐directedness, then, should offer no impediment to naturalism. Ludwig
von Bertalanffy presses the case:
[T]eleological behaviour directed toward a characteristic final state or goal is not

something off limits for a natural science and an anthropomorphic misconception
of processes which, in themselves, are undirected and accidental. Rather it is a
form of behaviour which can be well defined in scientific terms and for which the
necessary conditions and possible mechanisms can be indicated. (von Bertalanffy
1969, 46)11
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However, naturalized teleology requires more than mere goal‐directedness. As men

tioned, the goal‐directed activity of a system must have explanatory consequences that
are autonomous from the explanatory consequences of the mechanical interactions of the
system's parts. Teleology would be redundant if every phenomenon explained by appeal
to the goal‐directedness of a system could also be explained by appeal to the mechanical
interactions among its parts.
But this worry is misplaced. There is an independently compelling approach to explana

tion that has the added merits of demonstrating the explanatory autonomy of goal‐direct
edness over the mechanisms that cause goal‐directedness. The general idea is that we ex
plain a phenomenon e by citing some other phenomenon f such that (i) f is change relat
ing, and (ii) e counterfactually depends upon f (Woodward 2000); were f to occur, so would
e, and were f not to occur, e wouldn't either.12 When this relation holds, we say that f is a
difference maker for e. To explain a phenomenon e is to identify a difference maker—some
factor f that makes the difference between e's occurrence and its nonoccurrence. This ap
proach to explanation is often employed in defence of mechanism, but it can also be used
to demonstrate that the goal‐directedness of a system has explanatory consequences that
the mechanical interactions of its parts do not (Walsh 2006b).
Suppose we have a goal‐directed system that tends, by whatever available means, to at
tain stable end‐point R. Suppose further that, on some particular occasion, our system at
tains R by producing an outcome e, whose mechanical causes are c 1, … c n. There are two
relevant phenomena to be explained—e's occurrence, and the end‐point's instantiation of
R—and there are two distinct difference makers. The set of mechanical causes c 1, … c n is
a difference maker for e; were these to occur in relevantly different counterfactual situa
tions, e would also. So (p. 124) c 1, … cn explains the occurrence of e. But c 1, … c n does not
explain why the end‐point of this system almost invariably instantiates R. The reason is
that, even if c 1, … c n had not occurred, some sequence of mechanical causes would have
occurred that eventuated in R. The difference maker for the regularity of R is the goal‐di
rected capacity of the system as a whole. If R were not a goal for the system in question,
it would not be the case that had c 1, … c n not occurred, some other causal sequence suffi
cient to realize R would have.
The goal‐directedness of a system consists in its capacity to marshall the causal powers of
its mechanisms in a way that realizes R across a range of possible circumstances. Goal‐di
rectedness thus explains what the mere mechanical properties of the parts cannot. In this
sense, goal‐directedness has explanatory autonomy over the mechanisms that produce
it.13
Indeed, the goal‐directedness of the system as a whole figures in explanations in just the
way that Aristotle's model of teleology suggests. The goal‐directedness of a self‐organiz
ing system explains the occurrence of its particular component processes. Typically, a
self‐organizing, adaptive system will have at its disposal a broad repertoire of possible
trajectories and end‐states. Each of the component parts (or processes) itself has a reper
toire of possible effects. The regulatory control imparted by the system as a whole on the
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Teleology
activities of the parts explains why each part behaves in the way it does. After a perturba
tion, the system will effect changes among the component processes that restore the sta
ble end‐point. The changes occur precisely because they are required—in the sense of hy
pothetically necessary—for the attainment of the end‐point.
There are further Aristotelian resonances in the explanation of self‐organizing, adaptive

systems. Aristotle (1970, Physics II.8) criticizes the mechanistic explanations of the atom
ists (“materialists”) as insufficient. Mechanistic explanation is sufficient to account for a
particular outcome, say, the development of the incisors at the front of the mouth. Me
chanical causes can demonstrate how, in some sense, the effect to be explained was nec
essary. But they cannot distinguish those events that are the outcome of pure chance
from those that occur “always or for the most part” (ibid.). Matthen (2007) elucidates
Aristotle's position:
Aristotle's claim is that when an event O, brought about by a complex sequence of

constituent events occurs regularly, there must be an overarching cause C
responsible for the material/efficient causes of O. C is responsible for O occurring
regularly because it ensures that the materialist causes sufficient for O occur reg
ularly. The materialist explanation that cites only E [efficient causes] is deficient,
then, not because it gives causally insufficient conditions, but because it does not
specify all the causes of O. (Matthen forthcoming)
Aristotle's case is overstated, of course. Nowadays, we think that some things happen regularly
precisely because they are governed by mechanical law. But complex systems dynamics demon
strates that not all regularities are like that. Some regularities occur because they are the conse
quences of the activities of goal‐directed systems. In these systems, goal‐directedness (and not
mechanical law) plays the role of the Aristotelian “overarching cause.”
(p. 125)
The upshot of these considerations is that teleology should be seen as a naturalistically

acceptable, autonomous mode of explanation in the natural sciences. It is applicable to
self‐organizing, goal‐directed systems. According to the Aristotelian model, all that teleo
logical explanation requires is that there are natural goal‐directed systems and that goal‐
directedness explains the regulation of the component processes of such systems. It may
well be that this mode of explanation has no currency in the fundamental sciences.
Physics probably has no need of the concept of goal‐directedness. But it doesn't follow
from this that natural teleological explanation plays no role in any of the natural sciences.
It remains to be shown whether it plays an irreducible role in modern evolutionary biolo
gy.
5. Function Ascription in Evolutionary Biology

The literature on teleology and related issues in modern biology is vast. Practically all of
it in recent years has been concerned with the use of function ascriptions and adaptive
explanations.14 The motivating thought is that, if teleological explanation is to have some
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Teleology
role in biology, it is to be found in the practice of ascribing functions to traits. There is, af
ter all, at least a passing resemblance between teleological explanation and the explana
tory uses of function ascription in biology. Ascribing a function to a biological trait ap
pears to offer an explanation of the presence of the trait in question by identifying its con
tribution to an organism's vital processes. There are, evidently, “normative” distinctions
in play in biological function talk. We can distinguish functions from malfunctions and
functions from accidents (Millikan 1989; Neander 1991). The function of a trait, we are
told, is some effect that the trait ought to have in propitious conditions; a trait's failure to
perform its function counts as a malfunction. The effect of a trait that explains its pres
ence is its function; other effects are accidents.
There are two general strategies for understanding biological function talk. Both operate
under the presupposition that an unreduced teleology of goals and goal‐directedness is
unavailable to evolutionary biology.15 One strategy seeks to preserve as much as possible
of the apparently teleological tenor of function ascription without taking on the commit
ment to unreduced explanation by goals. The other eschews any form of ersatz teleology
and seeks to show that these putative explanatory consequences of function ascription in
biology are illusory (Walsh 2007).
Among those strategies that seek to emulate the teleological explanations, Larry Wright's
(1973) and Ruth Garrett Millikan's (1984, 1989) are the most influential. According to this
general approach, the biological function of a trait is some effect for which the trait has
been selected in the past. It holds that “the function of x is to f,” means “traits of x's kind
are prevalent in the population (p. 126) because they have been promoted by selection be
cause of their capacity to do f in the past.” This “selected effects' approach has become
the orthodox position on the concept of function since the early 1990s (Buller 1999).
There are two issues relevant to the assessment of the selected effects approach to func
tion. The first is its adequacy. Does it capture the full range of uses to which function as
criptions are put by biologists? The second issue is how accurately it represents the ex
planatory practices of biologists.
It has often been noted that the selected effects account cannot adequately encompass
the full range of uses to which function explanations are put in biology. In particular, it
cannot countenance novel functions. Biologists frequently ascribe novel functions to traits
or functions to novel traits (Bigelow and Pargetter 1987; Amundson and Lauder 1994;
Walsh 1996).16 There is a straightforward curative to the selected effects account that al
lows it to accommodate all of the uses of the function concept in evolutionary biology. The
effect that a trait has had in the past by dint of which it has been selected is simply its
typical contribution to the fitness of the individuals that have possessed it. Contributions
to fitness may be historical or novel. So the selected effects account may be amended in
the following fashion: The function of a trait is its typical contribution to an organism's fit
ness (Walsh 1996; Walsh and Ariew 1996).17
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Teleology
The contribution to fitness accounts preserve the presumed advantages of the historical
approach. They support the function/malfunction and function/accident distinctions. The
contribution to fitness that is typical for a trait type (in a selective regime) is the trait's
function. Tokens of the type that are incapable of making this contribution to fitness are
malfunctioning. Other effects of a trait that do not significantly affect the fitness of an in
dividual are mere accidents. Furthermore, the contribution to fitness approach enjoys sig
nificant advantages over selected effects accounts. It allows, for instance, that novel
traits may have biological functions and that a trait may have a biological function even
when it is being (or has been) selected against (Walsh and Ariew 1996; Buller 1999; Walsh
2007).
The second consideration in assessing these approaches is how accurately they charac
terize the role of functional explanations in biology. It is often taken to be a mark of the
success of this family of function theories that it emulates genuine teleological explana
tion, without incurring the commitment to goals. For example, the claim that selected ef
fect function ascriptions closely approximate teleological explanations (or indeed are gen
uinely teleological) is commonplace. For example, Neander says:
Teleological explanations of the functional variety are like purposive explanations,

in that they explicitly refer to a future effect of a trait for which that trait was se
lected. In doing so they explain the trait by implicitly referring to the causally effi
cacious selection process from which it resulted. (1991, 463)18
On this general approach, functional explanations in biology count as genuinely teleologi

cal only because teleological explanations are recast as a mere species of efficient cause
explanation (Nagel 1977). Function ascriptions may well figure in (p. 127) teleological ex
planations in this etiolated sense of teleology. But this is teleology in name only.19 The use
of function ascriptions in evolutionary biology seems not to require any unreduced appeal
to explanation by goal‐directedness. A function ascription does not explain the presence
of a trait token in an individual organism by showing it to be hypothetically necessary for
the attainment of the organism's goals. Rather, it explains the presence, prevalence, or
persistence of trait types in a population, by articulating some biologically significant ef
fect: their contributions to fitness. None of these roles requires function attributions to be
teleological.
Christopher Boorse (2002) has sought to reaffirm the full‐blooded teleological commit
ments of function ascriptions. The concept of biological function, he argues, requires us
to acknowledge a category of biological goals. He argues that the fitness of an individual
organism is simply its capacity to survive and reproduce: Survival and reproduction are
goals of organisms. So, to ascribe a function to a trait is to cite its contribution to a goal.
That may well be, but that alone is not sufficient to make function ascriptions in biology
genuinely teleological. Survival and reproduction are indeed goals, but they do not figure
in functional explanations in biology as goals; they appear simply as effects. All in all,
there is little reason to suppose that biological function ascriptions figure in any irre
ducible teleological explanations. There is little reason, then, to suppose that the extent
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Teleology
to which our account of biological function emulates genuine teleological explanation

should be a criterion of that account's success.
This antiteleological approach to function has been strenuously pursued by Robert Cum
mins (1975, 2002). Cummins argues that to ascribe a function to a part of a system is sim
ply to identify the contribution that part makes to some overall activity of the system of
interest. Walsh and Ariew (1996) point out that the contribution to fitness conception of
function constitutes a special case of Cummins function, where the system of interest is
the organism and the effects of interest are typical contributions to survival and repro
duction.
The Cummins approach to function has been the subject of withering criticism, particu
larly for its alleged inability to accommodate the so‐called normative distinctions. But the
Cummins account can underwrite the function/malfunction and function/accident distinc
tions by appeal to the pragmatics of explanation. Given a system and analysis of interest,
the function of a trait is the causal contribution it makes to the characteristic activity—
i.e., the activity we wish to explain; other effects are accidents. Where the activity of in
terest is survival (and/or reproduction), then a trait token that does not contribute to fit
ness in the way other tokens of its type do can be said to be malfunctioning (Walsh 2007).
Furthermore, where the effect of interest is survival (and/or reproduction), then the typi
cal contribution that traits of the type make to this effect explains the persistence of the
trait type. Some Cummins functions explain the presence of a trait in a population as well
as ersatz teleological functions.
Functional explanation in biology requires no appeal to goals, goal‐ directedness, or

norms. It appears, then, that the practice of ascribing functions to biological traits re
quires no teleology—ersatz or genuine (Matthen 1997). If there is (p. 128) any vestige of
unreduced teleology in biology, function ascriptions are not the place to look for it.
6. Teleology in Organismal Biology

There is, I believe, a better place to look for unreduced teleological explanation in biolo
gy, one that has gone strangely ignored. It is to be found in the adaptiveness of organ
isms. Organisms are the very paradigm of goal‐directed, adaptive systems: “you cannot
even think of an organism without taking into account what variously and rather loosely
is called adaptiveness, purposiveness, goal seeking and the like” (von Bertalanffy 1969,
45). Jacques Monod, in a similar vein, identifies “one of the fundamental characteristics
common to all living things: that of being objects endowed with a purpose or project.
Rather than reject this idea … it is indispensable to recognize that it is essential to the
very definition of living things” (1973, 9). The purposive adaptiveness of organisms clear
ly has some explanatory consequences: Metabolic rates vary as a consequence of environ
mental temperature; immune responses restore well‐being in the face of disease; circula
tory and respiratory systems acclimate to changes in altitude. There must, then, be at
least a limited role for teleological explanation to play (Ayala 1970). This recognition of an
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Teleology
explanatory role for the adaptiveness of organisms is a very minor concession to teleolo
gy, one that the naturalist may make with equanimity.
Once this concession is made, however, biologists should immediately feel impelled to
make a much more momentous one. Teleology is indispensable to the central project of
evolutionary biology, viz. explaining the adaptiveness of life.
“Adaptiveness” has two radically different senses. Applied to organisms, “adaptiveness”

refers to the goal‐directed capacity to make compensatory changes to form or physiology
during one's lifetime, changes of the sort we see in acclimation or in the immune re
sponse. Applied to the process of evolution, it designates the tendency of lineages under
going selection to become increasingly well suited to the conditions of their existence
over evolutionary time. In this second sense, adaptiveness has no teleological implica
tions. It is commonly, and rightly, supposed that the adaptiveness of organisms is a simple
consequence of the adaptiveness of evolution (Ruse 1973; Hull 1973). So whereas physiol
ogy may need to avail itself of teleology, the process of evolution can be wholly explained
without it.
However, advances in the understanding of organismal development suggest that the

adaptiveness of organisms has a more profound and pervasive role to play in evolutionary
biology than has heretofore been recognized. It has been noticed that the Darwinian
process of iterated mutation and selection is insufficient, by itself, to produce adaptive
evolution. Wagner and Altenberg (1996) demonstrate (p. 129) this with the case of evolu
tionary algorithms. It is impossible, they say, to improve the workings of most computer
programs (for example, Microsoft Word) simply by introducing random perturbations
(mutations) and then selecting the outcomes. Invariably, such programs are adversely af
fected by this process. Algorithms can only evolve under selection and mutation if they
possess a certain kind of architectural structure. This suggests an analogous question for
organisms; what properties do they have that allows populations of them to undergo
adaptive improvement in evolution? The question is posed elegantly by Stuart Kauffman:
How can such wonderful systems emerge merely through random mutation and
selection? For if Darwin told us that adaptation occurs through the gradual accu
mulation of useful mutations, he has not yet told us what kinds of systems are ca
pable of accumulating useful mutations. (Kauffman 1993,173)
The question of how organisms are subject to adaptive improvement through the Darwin
ian process has become one of the most important in evolutionary biology. It goes under
the rubric of “evolvability.” Evolvability requires organisms to meet the competing de
mands of robustness and flexibility (Wagner 2005).
By evolvability, we mean the capacity to generate heritable, selectable phenotypic

variation. This capacity may have two components: (i) to reduce the potential
lethality of mutations and (ii) to reduce the number of mutations needed to pro
duce phenotypically novel traits. (Kirschner and Gerhart 1998, 8420)
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Teleology
Current developmental biology suggests that a single feature, plasticity, confers both
robustness and flexibility upon organisms. Plasticity is the capacity of an organism “to re
act to an internal or external environmental input with a change in form, state, move
ment, or rate of activity” (West‐Eberhard 2003, 33). Plasticity involves both the mainte
nance of constant structure and function in the face of perturbations and the capacity to
make novel adaptive changes in order to preserve stability. The plasticity of organisms is
simply a manifestation of their self‐organizing goal‐directedness (Walsh 2006a).
The significance of phenotypic plasticity for adaptive evolution is receiving a considerable

degree of attention (West‐Eberhard 2003, 2005a, 2005b; Kirschner and Gerhart 2005).
West‐Eberhard conjectures that the adaptiveness of adaptive evolution is driven by the
phenotypic plasticity of organisms. She proposes that adaptive evolution comprises three
stages: phenotypic accommodation, developmental recombination, and genetic accommo
dation:
Phenotypic accommodation: “Phenotypic accommodation due to phenotypic plas

ticity is the immediate adaptive adjustment of phenotype to the production of a
novel trait or trait combination. … Phenotypic accommodation reduces the amount
of functional disruption occasioned by developmental novelty” (West‐Eberhard
2003, 147). “Adaptive phenotypic adjustments to potentially disruptive effects of
the novel input exaggerate and accommodate the phenotypic change without ge
netic change” (West‐Eberhard 2005a, 613).
(p. 130)
Developmental recombination: “Developmental recombination occurs in a popula

tion of individuals because of a new, or recurrent input. A new input … causes a re
organization of the phenotype, or ‘developmental recombination.’ Given the vari
able developmental plasticity of different individuals, this process produces a pop
ulation of novel variable phenotypes, providing material for selection” (West‐Eber
hard 2005b, 6544).
Genetic accommodation: “If the phenotypic variation is associated with variation

in reproductive success, natural selection results; and to the degree that the vari
ants acted upon by selection are genetically variable, selection will produce genet
ic accommodation” (West‐Eberhard 2005a, 612).
On this account of evolution, adaptive novelties initially appear in a population not pri
marily through mutation, but through adaptive changes in phenotype made during devel
opment, facilitated by the plasticity of organisms. They are maintained in a population not
by gene selection but by the capacity of organisms reliably to produce viable phenotypes
despite genetic and environmental uncertainty. Plasticity is pivotal in both of these
processes. The significance of plasticity underscores the ineliminable role that organisms
—construed as self‐organizing, goal‐directed, adaptive systems—play in the explanation
of adaptive evolution (Walsh 2006a).
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Teleology
A crucial concept in this understanding of adaptive evolution as driven by the plasticity of

organisms is phenotypic repertoire. Each organism has the capacity to produce any of a
very large array of phenotypes. Organisms have their broad phenotypic repertoires be
cause development is modular. Each developmental module itself has the capacity to con
tribute to the production of a wide array of adaptive phenotypes. Each, in turn, has the
capacity to influence other modules in a wide variety of ways: “In principle, a modular
system allows the generation of diverse phenotypes by the rearrangement of its internal
module connections and [by the] relatively independent evolution of each
module” (Kitano 2004, 830).
Most phenotypic novelties are the product of the reorganization of the regulatory rela
tions among modules (Kirschner and Gerhard 2005): “A very large body of evidence …
shows that phenotypic novelty is largely reorganizational rather than a product of innova
tive genes” (West‐Eberhard 2005b, 6547). One consequence of this is that novel pheno
types tend to be less deleterious and more likely to be adaptively advantageous than the
traditional model of genetic mutation would suggest. “Variation … is both less lethal and
more appropriate to selective conditions than would be variation from random change.
Evolutionary change is thereby facilitated” (Kirschner and Gerhard 2005, 221).
One pressing question raised by phenotypic repertoire is: What determines which of the
array of outputs that a module might produce it actually does produce? Another is: What
determines which of the vast array of total phenotypes an organism might produce it ac
tually adopts? The answers lie in the self‐organizing, goal‐directed, adaptive capacities of
the organism as a whole.
The plasticity of organisms, as we have seen, consists in their capacity to regulate the
causal powers of their component parts (modules) and processes (p. 131) during develop
ment in a way that directs the organism toward the attainment of a stable, viable end‐
state. Phenotypic plasticity is the manifestation of the goal‐directedness of organisms. De
velopmental biologists increasingly appeal to this capacity in order to explain the occur
rence of particular traits during development and the constraints upon variation. Rudy
Raff argues that the robust adaptive capacities of organisms during development
provide internal, non‐random evolutionary variation. Such variation is sorted by

the internal requirement to maintain integrated function and by external selection
on resultant phenotypes. Because of the internal requirement that modified onto
genies be functional, only a subset of all theoretically possible phenotypes will be
generated. (Raff 1996, 325, emphasis added)
The explanatory role of phenotypic plasticity has all of the hallmarks of genuine, unre
duced teleology (Walsh 2006b). Plasticity is a prime example of Aristotle's “overarching
cause” of development. It explains why organisms of a particular kind resemble one an
other to the extent they do, despite the enormous amount of genetic variation within a
population and despite the occurrence of perturbations. Organisms of the same species
resemble one another, when they do, not because of similarities among their mechanical,
efficient causes, but in spite of the differences between them (Gibson 2002). Organismal
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Teleology
development occurs “always or for the most part” in the way it does because of the
buffering of development secured by the plasticity of organisms.
Furthermore, the plasticity of organisms illuminates an explanatory role for hypothetical

necessity in organismal development. Plasticity consists in the capacity of organisms to
effect compensatory changes in response to developmental perturbations. These pertur
bations are made in response to the “internal requirement that modified ontogenies be
functional” (Raff 1996: 325). That is to say, plasticity ensures that alterations to develop
ment can be made for the very reason that they are hypothetically necessary for the con
tinued proper functioning of the organism. Again, plasticity as the overarching cause ex
plains not just the regularity of development, but also the occurrence of particular com
ponents of an individual's development. This is a distinctive hallmark of Aristotelian teleo
logical explanation.
The position is this. Plasticity explains (i) the regularity of development, (ii) the nonran
domness of phenotypic novelties, (iii) the maintenance of phenotypic novelties in a popu
lation, and (iv) the capacity of organismal development to produce a viable organism typi
cal of its kind, despite perturbations and genetic and environmental vagaries. Plasticity is
a manifestation of the goal‐directedness of organisms. It is the capacity of an organism to
maintain stability by effecting adaptive, compensatory changes to its component develop
mental processes. So, the goal‐directedness of organisms explains the regularity of organ
ismal development. Moreover, it does so in a way that an appeal to the mere mechanical
processes of an organism's development cannot. The goal‐directedness of an organism
regulates the processes of its own development. So, goal‐directedness explains why the
(p. 132) particular processes that occur within an organism's development actually do oc
cur. If plasticity genuinely explains these developmental phenomena, then the explanation
of development is irreducibly teleological.
The role of plasticity in organismal development highlights the Kantian tension between
mechanistic and teleological explanations in biology (Walsh 2006b). Organisms are clear
ly susceptible of mechanical explanation. An organism's phenotype is the consequence of
the set of causal interactions among its component parts and processes. At the same
time, however, given the vast repertoire of an organism's developmental system, given
the vagaries of the environment and the enormous amount of genetic perturbation, mech
anism alone cannot explain why an organism adopts one total phenotype rather than an
other. Nor can pure mechanism alone explain the resemblances among organisms. The vi
able development of an organism, the high fidelity of phenotypic inheritance, and the re
semblances among organisms of the same kind are all dependent upon the goal‐directed
regulation of development. Goal‐directedness explains why the specific causal interac
tions that produce a particular organism actually occur. It also explains why these
processes occur as regularly as they do—“always or for the most part”—given the enor
mous amount of genetic, developmental, and environmental variation faced by a popula
tion of organisms. The goal‐directedness of organisms is the difference maker for the reg
ularity of development. Uniquely, then, on account of their self‐synthesizing, self‐repro
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Teleology
ducing, self‐regulating capacities, organisms require both mechanical and teleological

modes of explanation.
But, the goal‐directedness of development is required for the explanation of more than
just developmental regularity. Goal‐directedness is required to explain why adaptive evo
lution is adaptive. West‐Eberhard's account of plasticity suggests that the answer to
Kauffman's question “what kinds of lineages are capable of adaptive evolution?” is simply
“those comprising entities with the right kind of phenotypic plasticity.” Adaptive evolution
of lineages is possible because of the phenotypic plasticity of their constituent organisms.
If recent developmental biology is correct, plasticity has an ineliminable role to play in
the explanation of adaptive evolution. Plasticity, of course, is just the manifestation of the
adaptive goal‐directedness of organisms. If this is so, then the adaptiveness of evolution
is a consequence of the adaptiveness of organisms. If this is the case, then the explana
tion of adaptive evolution requires an appeal to unreduced teleology.
Conclusion
Teleology is a naturalistically acceptable, indeed indispensable, mode of explanation. I
have argued for this conclusion on two grounds. First, teleology, construed as an explana
tory strategy that explains the features of a system by appeal to its (p. 133) goal‐directed
capacities, is available to the thoroughgoing naturalist. It does not succumb to the bat
tery of standard antiteleology objections from nonactuality, intentionality, and normativi
ty. These objections are merely holdovers from the teleophobia prevalent in early modern
thinking about the natural world. Moreover, these arguments implicitly rely on the Platon
ic model of extrinsic, transcendent teleology. The Aristotelian model of teleology is more
congenial to the present‐day naturalist. According to Aristotelian teleology, the goal‐di
rectedness of a system explains the presence and nature of the system's component parts
and processes. Aristotelian teleology complements our current understanding of goal‐di
rected, self‐organizing, adaptive systems. Second, recent advances in evolutionary biolo
gy suggest that teleological explanation is indispensable. We can explain neither the fi
delity of inheritance nor the success of development without appeal to the plasticity of or
ganismal development. Plasticity itself is a goal‐ directed capacity of organisms to pro
duce and maintain a stable, well‐functioning living thing in the face of the vagaries of en
vironment and genetics. More important, the adaptive plasticity of organisms is required
to explain how a population of organisms undergoing natural selection can manifest adap
tive evolution. Phenotypic plasticity issues in genuine, unreduced teleological explana
tions. If plasticity is required for the explanation of adaptive evolution, then so is the
mode of explanation that deploys it—teleology.
The “Aristotelian purge” was seen as a pivotal achievement of early modern science.20 As
a consequence of the scientific revolution, the natural sciences learned to live without
teleology. Current evolutionary biology, I contend, demonstrates that quite the opposite
lesson needs now to be learned. The understanding of how evolution can be adaptive
requires us to incorporate teleology—issuing from the goal‐directed, adaptive plasticity of
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Teleology
organisms—as a legitimate scientific form of explanation. The natural sciences must, once
again, learn to live with teleology.
Acknowledgments
I would like to thank André Ariew, Thomas Johansen, Mohan Matthen, Marcel Quarfood,
and Joan Steigerwald for valuable help in my understanding various historical concep
tions of teleology. Various parts of this paper are drawn from talks given in London, Paris,
and Montreal. I thank the audiences for helpful discussions. I would especially like to
thank Michael Ruse for his patience and keen editorial eye.
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Notes:
(1.) The term “pure mechanism” comes from Broad (1925).
(2.) Quoted in Grene and Depew (2004, 37).
(3.) Of course, the perception of the relation of Darwin's theory to teleology is much more
complicated. Beatty (1990) and Cornell (1986) offer sophisticated and complementary
discussions.
(4.) Aquinas, Summa Theologica (2006), question 2, article 3.
(5.) Small wonder then that teleology is thought to support intelligent design and that
those who oppose intelligent design should seek to rid biology of its involvement with
teleology. See Ariew (2007).
(6.) Bedau is an exception to the usual naturalist scruples about countenancing a natural
world imbued with value.
(7.) I am indebted to Thomas Johansen's account of Plato's teleology found in his Plato's
Naturalism (2004).
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(8.) The term “interactive unity” comes from David Charles.
(9.) It is this simple thought that gave rise to the “supervenient causation” industry.
(10.) The term “supple adaptation” originates with Bedau (1992).
(11.) See Ayala (1970) for an articulation of the claim that teleology is an empirically ob
servable feature of the biological world.
(12.) This is highly oversimplified. A sophisticated and detailed account can be found in
Woodward (2003).
(13.) In this sense, goal‐directedness is an emergent property (see Broad 1925).
(14.) Allen, Bekoff, and Lauder (1999) and Buller (1999) are valuable compendia of essays
on function and teleology in biology.
(15.) Boorse (2002) is a notable exception.
(16.) Griffiths (1993) and Godfrey‐Smith (1994) attempt to provide a quick fix by distin
guishing recently selected effects from more distantly historical effects.
(17.) Ruse (1973) and Wimsatt (1972) develop early versions of the contribution‐to‐fitness
approach.
(18.) See also Lennox (1993), who suggests that Darwin's theory of natural selection
gives us a new theoretical definition of teleology.
(19.) Mayr (1988) offers a taxonomy of faux teleologies.
(20.) The term “Aristotelian purge” comes from Taylor (1989).
D.M. Walsh
D.M. Walsh is Canada Research Chair in Philosophy of Biology in the department of

Philosophy and the Institute for the History and Philosophy of Science and Technolo
gy, University of Toronto.
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Teleology

Abstract and Keywords

Teleology is a mode of explanation in which the presence, occurrence, or nature of some

Keywords: model, mechanism, argument, teleology, evolution, plasticity

Subscriber: Universitats- und Stadtbibliothek Köln; date: 12 March 2022

1. Barren Virgins and Mistresses

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The argument from nonactuality

The argument from intentionality

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Kant, in reference to the apparent purposiveness of organisms, tells us:

The argument from normativity

(p. 118) 3. Transcendence and Immanence

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The Platonic conception of teleology lends support to standard antiteleological objections.

Where Plato's teleology is transcendent and supernatural, Aristotle's is immanent and

We may think of the form of an organism as a set of organizing principles, or a set

Subscriber: Universitats- und Stadtbibliothek Köln; date: 12 March 2022

The argument from nonactuality

It is goal‐directedness, not unactualized goals, that explains the presence of traits in an

The argument from intentionality

The argument from normativity

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The second implication of the ought/normative requirement distinction is that it allows us

Subscriber: Universitats- und Stadtbibliothek Köln; date: 12 March 2022

4. Explanatory Sufficiency, Explanatory Au­

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Goal‐directedness is an unproblematic causal consequence of the architecture of an adap­

[T]eleological behaviour directed toward a characteristic final state or goal is not

Subscriber: Universitats- und Stadtbibliothek Köln; date: 12 March 2022

However, naturalized teleology requires more than mere goal‐directedness. As men­

But this worry is misplaced. There is an independently compelling approach to explana­

Subscriber: Universitats- und Stadtbibliothek Köln; date: 12 March 2022

There are further Aristotelian resonances in the explanation of self‐organizing, adaptive

Aristotle's claim is that when an event O, brought about by a complex sequence of

The upshot of these considerations is that teleology should be seen as a naturalistically

5. Function Ascription in Evolutionary Biology

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Subscriber: Universitats- und Stadtbibliothek Köln; date: 12 March 2022

Teleological explanations of the functional variety are like purposive explanations,

On this general approach, functional explanations in biology count as genuinely teleologi­

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to which our account of biological function emulates genuine teleological explanation

Functional explanation in biology requires no appeal to goals, goal‐ directedness, or

6. Teleology in Organismal Biology

Subscriber: Universitats- und Stadtbibliothek Köln; date: 12 March 2022

“Adaptiveness” has two radically different senses. Applied to organisms, “adaptiveness”

However, advances in the understanding of organismal development suggest that the

By evolvability, we mean the capacity to generate heritable, selectable phenotypic

Subscriber: Universitats- und Stadtbibliothek Köln; date: 12 March 2022

The significance of phenotypic plasticity for adaptive evolution is receiving a considerable

Phenotypic accommodation: “Phenotypic accommodation due to phenotypic plas­

Developmental recombination: “Developmental recombination occurs in a popula­

Genetic accommodation: “If the phenotypic variation is associated with variation

Subscriber: Universitats- und Stadtbibliothek Köln; date: 12 March 2022

A crucial concept in this understanding of adaptive evolution as driven by the plasticity of

provide internal, non‐random evolutionary variation. Such variation is sorted by

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Furthermore, the plasticity of organisms illuminates an explanatory role for hypothetical

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ducing, self‐regulating capacities, organisms require both mechanical and teleological

Subscriber: Universitats- und Stadtbibliothek Köln; date: 12 March 2022

4. Explanatory Sufficiency, Explanatory Au

Goal‐directedness is an unproblematic causal consequence of the architecture of an adap

However, naturalized teleology requires more than mere goal‐directedness. As men

But this worry is misplaced. There is an independently compelling approach to explana

On this general approach, functional explanations in biology count as genuinely teleologi

Phenotypic accommodation: “Phenotypic accommodation due to phenotypic plas

Developmental recombination: “Developmental recombination occurs in a popula

West‐Eberhard, M. J. 2003. Developmental Plasticity and Phenotypic Evolution. Cam

West‐Eberhard, M. J. 2005a. Phenotypic Accommodation: Adaptive Innovation due to De

West‐Eberhard, M. J. 2005b. Developmental Plasticity and the Origin of Species Differ