Adaptation: “A Critique of Some Current Evolutionary Thought”

Author(s): Helena Cronin

Source: The Quarterly Review of Biology , Vol. 80, No. 1 (March 2005), pp. 19-26
Published by: The University of Chicago Press
Stable URL: https://www.jstor.org/stable/10.1086/431021

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Volume 80, No. 1 THE QUARTERLY REVIEW OF BIOLOGY March 2005

ADAPTATION:
“A CRITIQUE OF SOME CURRENT EVOLUTIONARY THOUGHT”

Helena Cronin
Centre for Philosophy of Natural and Social Science

London School of Economics

London WC2A 2AE England
e-mail: h.cronin@lse.ac.uk

abstract
In his classic Adaptation and Natural Selection: A Critique of Some Current Evolutionary
Thought (1966), George Williams showed definitively that our understanding of adaptation, a
central concept of evolutionary theory, must be gene-centered. The purpose of adaptations is to further
the replication of genes. Genes are machines for turning out more genes; and adaptations are the
means by which genes pluck resources from the world to promote this self-replication. Thus adaptations
transform potential resources from part of the indifferent world-at-large into tailor-made environments,
environments brimming with resources for organisms’ distinctive adaptive needs. Systematically dif-
ferent adaptive problems therefore give rise to different environments; and so different species, for
example, have different environments. Thus a gene-centered analysis of adaptations implies a gene-
centered theory of environments. Without genes to specify what constitutes an environment, environ-
ments would not exist. Rather than being separate from biology, an autonomous, independent force,
environments are themselves the products of biology. So a gene-centered view, far from depreciating the
environment, furnishes a rich and precise understanding of its importance.

tered route: “It is at the level of the gene . . .

Some Current Evolutionary
Thought—Again
A DAPTATION: the design features of liv-
ing things. This striking property of the
organic world is, as George Williams urged in
his classic work, Adaptation and Natural Selec-
tion: A Critique of Some Current Evolutionary
Thought, “a phenomenon of pervasive impor-
tance in biology” and a “key . . . concept” of
evolutionary theory (Williams 1966:5). But
the concept had become misused and mis-
understood, even to the point of impeding
scientific progress. George’s impetus for writ-
ing the book was to “help to purge biology”
(Williams 1966:4) of such errors.
George’s battle was mainly with levels of
selection. He showed unequivocally that the
path to understanding was down the gene-cen-
that we have the most fundamental and most
universally applicable understanding of adap-
tation” (Williams 1966:71). In recent years, the
concept of adaptation has again become
abused, even traduced. The battle this time is
largely about the relationship between genes
and environments. Once again, a steadily
gene-centered analysis is needed.
I shall first set out a gene-centered notion
of adaptation, then present the latest miscon-
ceptions, and finally illustrate how this per-
spective can dispel them.
What Adaptations Are For
The mathematician Paul Erdös used to
describe himself as a machine for turning cof-
fee into theorems. In much the same way,
genes are machines for turning stars into a

19

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20 THE QUARTERLY REVIEW OF BIOLOGY
bird’s compass; carotenoids into males of daz-
zling beauty; facial muscles into signals of true
friendship; and oxygen, water, calcium, and
iron into bears, beetles, bacteria, or bluebells.
Genes have been designed by natural selec-
tion to exploit properties of the world that
promote their self-replication; for genes are
ultimately machines for turning out more
genes. The means by which genes gain access
to the world’s properties is through adapta-
tions. In shaping adaptations over evolution-
ary time, natural selection has seized on reg-
ularities, features of the world that are stable,
recurrent, and dependable, features that can
be relied on, generation after generation, so
that adaptations can do their work.
Consider the indigo bunting (Passerina
cyanea). Natural selection faced the problem
of enabling these migratory birds to navigate
from their northern home southwards to a
winter residence and then back again. Solving
a navigation problem involves locating a fixed
point. The specialized mental machinery with
which natural selection equipped the birds
for this task can be envisaged as a set of rules:
To find a fixed point, do the following.
Pay attention to the nightly voyage of the
stars. Familiarize yourself with the pat-
terns of the major constellations. And
distinguish between the slow and the fast
travelers—those that trace a tight circle,
barely moving, and those that sweep
across the sky, scurrying at speed. Pin
down the point around which they
revolve. To store where this point is, note
where it lies in relation to the now-famil-
iar nearby stars. Compass constructed;
use for finding fixed point.
Every boy scout knows that the buntings’
fixed point must be the north celestial pole,
the place towards which the earth’s axis of
rotation points, and thus the place in the sky
corresponding to north. But every boy scout
also knows that one of the nearby stars is the
pole star, conspicuously bright and within
one degree of the pole. Why, then, did natu-
ral selection not simply enable buntings to
make more direct use of that particular star?
The answer is that, relative to evolutionary
time, the close proximity of the pole star to
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Volume 80
the celestial pole is not a regularity. The
earth’s axis of rotation wobbles considerably
in a cycle of about 27,000 years; so the posi-
tion of the celestial pole among the constel-
lations changes. Thus natural selection
latched on to a deeper regularity and thereby
a far more powerful tool. It endowed bunt-
ings with the ability to study particular char-
acteristics of these constellations and then to
pinpoint the center around which they
rotate. Thus buntings that migrate today can
use the same instincts and environmental reg-
ularities as used by their long-dead ancestors,
fashioning the same precision-built instru-
ment, regardless of where the fickle earth is
situated in its never-ending cycle.
So every newly-hatched indigo bunting
becomes, in a matter of months, a proficient
star-compass navigator. As a nestling, it devotes
hours to gazing at the stars. By the time it starts
on its nocturnal migration, it can head in the
right direction upon leaving the nest; and, as
long as some of the constellations are visible,
it can use its compass to maintain its direction
along the way, confidently keeping its back to
the celestial pole in autumn and flying towards
it in spring. Fortunately, the winter quarters
are far enough north of the equator that the
circumpolar stars do not disappear over the
horizon; so the compass can be deployed
along the entire route.
Thus, the bunting’s adaptation relies not
only on its species-specific mental machinery
but also on the presence of the stellar array.
Indeed, if buntings are experimentally
immured in a starless world during the cru-
cial nestling period, they cannot orientate
themselves when migrating. Stars are as vital
a part of a bunting’s environment as is the egg
in which it develops or the water that it
drinks. Buntings without stars are destined to
be buntings without descendants.
And so it is for every one of the great pan-
oply of adaptations that natural selection has
ever favored. Resources in the environment
need to be there as reliably, in every genera-
tion, as the information in the genes. Just as
an organism requires its genes to be passed
on intact, so it requires its environment to be
unfailingly available down the generations—
carotenoids and calcium, smiles and stars.
Thus, in evolving an adaptation, natural
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March 2005 SYMPOSIUM IN HONOR OF GEORGE C WILLIAMS
selection accumulates in the species’ gene
pool an appropriate store of information
about environmental regularities. Each living
thing enters the world replete with such infor-
mation packed into its genes, information
about the stockpile of resources that natural
selection has relied on in the design of its
adaptations, information gleaned from the
environments of its ancestors. From this
archive, the organism can pick out the cur-
rent instantiations of those ancestral condi-
tions, the instantiations that it encounters in
its own lifetime of that long, unbroken suc-
cession of past environments. And thus it is
able to pluck from its environment what it
needs to grow and flourish and reproduce.
It is in ways such as these that adaptations
transform potential resources from part of
the indifferent world-at-large into the organ-
ism’s own tailor-made, species-specific envi-
ronment, an environment brimming with
materials and information for the organism’s
own distinctive adaptive needs.
In the beginning, natural selection created
genes that could barely exploit the heavens
or the earth; their adaptations were meager
and their environments commensurately
without form and void. But, down evolution-
ary time, genes have bootstrapped themselves
from inchoate nakedness into magnificent
dwellings, organisms of great sophistication.
They have proliferated adaptations of ever
greater complexity, inventing new technolo-
gies and strategies. They have thereby been
able to create and exploit ever richer environ-
ments in ever more ingenious ways, trans-
forming the universe of mere physics and
chemistry into a world of colors and sounds,
friendliness and beauty.
What Constitutes an Environment?
This gene-centered understanding of adap-
tation deepens our understanding of environ-
ment. What constitutes an environment? It is
those aspects of the world, and only those
aspects of the world, that genes have evolved
to harness for their use; the rest of the world
is not part of any environment. From the
potentially infinite ways of gaining access to
the world’s resources, genes use adaptations
to carve out subsets that the organism can
exploit to solve its adaptive problems. Scien-
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21
tific analyses should follow in natural selec-
tion’s wake. Environments are adaptation-
generated, adaptation-specific; environments
should therefore be parsed along systemati-
cally adaptationist lines.
Consider some examples. Take first a spe-
cies’-eye view of environments. Different spe-
cies are clusters of different adaptations; and
different adaptations pick out different envi-
ronments. So different species necessarily
have different environments. The ethologist
Jakob von Uexküll conjured up the image of
each species dwelling inside its own soap bub-
ble, “its own world, filled with the perceptions
which it alone knows“ (von Uexküll 1934:5).
In the bunting’s soap bubble, for example,
stars are a crucial element; nevertheless,
although these stars are part of the world-at-
large, a solid and enduring aspect of our
planet’s nocturnal landscape, for animals that
do not migrate or are resolutely diurnal, stars
are not part of these species’ soap bubbles.
Inside a soap bubble, there will be further
differences. Within a species, some individ-
uals face systematically different adaptive
problems from others; these differences give
rise to different environments. Soap bubbles
will differ for, say, males and females or
embryos and adults. In our species, for exam-
ple, what is salient differs for girls and boys;
even one-day-old girls prefer a human face,
boys prefer a mechanical mobile (Connellan
et al. 2000). In the parasitoid fly species Ormia
ochracea, the female, who lays her eggs on
male crickets, finds her victim by his love call;
her hearing is so finely attuned to his sere-
nade that it has converged with the auditory
world of the female cricket and diverged from
that of her own husband. Thus, shaped by
common adaptive problems, the females’
environments unite across their species’ soap
bubbles, transcending deep and ancient tax-
onomic divides; meanwhile, shaped by differ-
ent adaptive problems, the fly and her mate
dwell in disparate worlds of sound (Robert et
al. 1992, 1994).
Now consider a gene’s-eye view. For any
adaptation, its environment consists of what-
ever regularities natural selection’s design for
the adaptation relies on. For genes, these reg-
ularities are bound to include the behavior of
neighboring genes; this is because, in addi-
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22 THE QUARTERLY REVIEW OF BIOLOGY
tion to building proteins, genes influence
other genes, turning them on and off, acti-
vating them and silencing them, marshalling
them into and out of activity. And (especially
once the organism has emerged from the
confines of egg, seed, or womb) genes’ envi-
ronmental regularities are of course also
likely to include conditions in the external
world—temperatures and toxins, love songs
and warning calls.
A gene-centered view, far from depreciat-
ing the environment, furnishes a rich and
precise understanding of its importance. We
have noted, for example, that putting genes
at the center helps us to appreciate the sig-
nificance of environmental regularity, the
steady presence of the “expected” environ-
ment down the generations. As a second
example, we can now see how the relation-
ship between genes and environments cannot
possibly be—as is commonly assumed—zero-
sum, as if the more that environments do the
less there is for genes to do. A gene-centered
view emphasizes that, far from biology and
environment competing for causal power,
adaptations connect them in such a way that
the richer and more complex the adapta-
tions, the richer and more complex the envi-
ronments they rely on. As a final example,
recall that what constitutes an environment
cuts a swathe along a causal chain that invar-
iably includes both genes and aspects of the
outside world. This overturns the naı̈ve view
that “biology” stops where “environment”
starts, such that genes are never part of the
environment, and environment consists
exclusively of the world beyond the organism.
Without genes to specify what constitutes
an environment, environments would not
exist. Thus environments, far from being
separate from biology, an autonomous and
independent force, are themselves the prod-
ucts of biology. Environment is therefore—
necessarily—a biological issue, an adaptation-
ist issue, a gene-centered issue.
“Why Privilege Genes?”
George has given us not only a gene-cen-
tered understanding of adaptations but also
a gene-centered theory of environment. How-
ever, recent discussions of environment,
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Volume 80
rather than building on this insight, have, on
the contrary, attempted to oust genes from
center stage in the misguided belief that this
is how to give environment its due. In place
of George’s analysis, there has been much con-
fusion, involving a rash of environmental nex-
uses, developmental matrices, ontogenetic
niches, dynamic systems, interactive complex-
ities, holistic epigenetics, constructivist dialec-
tics, ontological reversals, phenotypic accom-
modations, decontextualized problem-solving
routines . . . and more. The confusions are
epitomized by a favorite rhetorical question:
“Why privilege genes?” Let us see how a gene-
centered analysis deals with some of the stan-
dard positions. (For various versions of these
arguments see e.g., Oyama 1985, 2000; Grif-
fiths and Grey 1994; Thelen and Smith 1994;
West-Eberhard 2003; for discussions see e.g.,
Dennett 1995:115 n. 10, 196–198; Sterelny
1995:162–165; Sterelny and Griffiths 1999:94–
111; for background see e.g., Popper 1972,
1990:27–51, 1999:45–56; Tooby and Cosmides
1990a:19–22, 1990b:388–389, 408–410, 1992:
69–73, 82–87; Cosmides and Tooby 1994:98–
103; Dawkins 1998; Pinker 2004).
• We are all interactionists now. Everyone agrees
that natural selection acts on the relationship
between genes and environments. So we should no
longer single out genes for privileged status. We
should accord parity to genes and environment.
Well, here is a parity test. Try specifying
“an” environment without first specifying
whether it is the environment of a bunting or
a human, a male or a female, an adaptation
for navigation or for language. The task is of
course impossible; the specification must start
in an adaptationist, and therefore gene-cen-
tered, way.
And here is another challenge to parity.
Genes use environments for a purpose—self-
replication. Environments, however, have no
purposes; so they do not use genes. Thus
genes are machines for converting stars into
more genes; but stars are not machines for
converting genes into more stars.
• Gene-environment interaction is so enmeshed
that the influences of genes and environments—in
both evolution and development—are inextricable.
Thus all attempts to disentangle them are spurious.
Inextricable? On the contrary; just ask the
right questions. It is misconceived to ask what
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percentage of the bunting’s compass or a
human language is the result of genes, and
what percentage is the result of environment.
But it does make sense to ask about the
design features of an adaptation. For bunt-
ings, at what ages do the windows of oppor-
tunity for compass-building open and close?
How important is the pole star; and how
many constellations must a nestling study?
Similarly, for humans, what exactly are the
instincts and what exactly are the environ-
ments that we need to acquire language? For
example, what is specified in the universal
grammar? What does a child require from an
environment of language users, and when?
It also makes sense to ask about the causes
of individual differences. How much of the
variation in a particular characteristic among
individuals within a population can be attrib-
uted to genetic, and how much to environ-
mental, differences? For example, what per-
centage of the differences in efficacy of
compasses in a group of buntings or linguistic
proficiency in a group of humans is attribut-
able to differences in genes, what percentage
to differences in environments? At present,
these questions (the realm of quantitative
genetics, particularly behavioral genetics)
tend not to analyze phenotypes or, conse-
quently, environments from an adaptationist
perspective. But the more that they come to
do so, the more they will shed light on the
detailed workings of both evolution and
development and the respective roles of
genes, environments, and even that elusive
player, chance.
• Genes respond to the organism’s experience of
the world. Experience regulates which genes are
expressed and when; the ambient temperature or a
smiling face can trigger reactions in a great cascade
of genes, switching some on and others off. So genes
are vulnerable to experience; they are at the mercy
of events in the outside world.
It is fundamental to the logic of an adap-
tationist approach that it must start from
genes, then go to adaptations, and only then
to environments, organisms, and experience.
But, critics object, this does not give expe-
rience its due. Experience can influence genes
as well as genes influencing experience; the
causal arrows go not only out from genes to
experience but also back again to genes.
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23
However, this criticism confuses the logic of
the approach with a claim about the direction
of causal arrows. In order to understand the
causal pathways between genes and their envi-
ronments, we must start from genes; but this
most certainly does not imply that the path-
ways go only one way.
The objection is proposed as an alternative
to the gene-centered view. But it is not. The
role played by experience is an essential part
of this view. How could it not be, given that
the entire raison d’être of adaptations is to
enable genes to use the world’s resources?
Therefore what constitutes experience for an
organism, and how it can exploit that expe-
rience, are part of the design features of its
adaptations, and consequently part of the
design specifications of its genes. So experi-
ence will of course be in the causal chains that
turn genes on and off.
Indigo buntings provide an illuminating
example of experience altering physiology,
and consequently, it seems, genes. How do
buntings decide when to migrate and in which
direction? The timing is triggered by changes
in an environmental factor, day length. The
birds’ experience of this change alters their
physiology; their fat deposits increase, for
instance, and they feel migratory restlessness.
In this state, new aspects of their environment
become salient; their physiology alters what
constitutes experience. They become sensitive,
for example, to slight changes in barometric
pressure; this enables them to sense when
pressure patterns augur winds favorable to
their odyssey. Similarly, the trigger for whether
to orientate themselves northwards (to home)
or southwards (to their winter dwelling) is the
changing seasons. But, again, the trigger acts
by causing physiological changes. Indeed,
once those changes have occurred, they use
their own internal state rather than their exter-
nal environment as their cue. Buntings that
were experimentally altered in spring to the
autumn body-state (by manipulating the ratio
of light and dark periods) orientated them-
selves towards the south, blithely oblivious to
the approaching season and the call of their
natal home.
Thus a gene-centered understanding not
only fully incorporates the role of experience
but also sharpens and enriches the concept,
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24 THE QUARTERLY REVIEW OF BIOLOGY
making scientific sense of what is otherwise
vague and elusive.
• Evolutionary change does not have to wait for
genetic mutations; environments and phenotypes
working together can take the lead. For example, by
influencing development, an environmental change
can bring about phenotypic change; and if the new
phenotype is more viable than the old, it will spread
rapidly through the entire population; what is more,
these new phenotypes will persist down the genera-
tions as long as those environmental conditions per-
sist. So the outcome will be adaptive change without
genetic mutation. Thus, contrary to orthodox Dar-
winian thinking, environmentally induced pheno-
typic novelty can initiate and sustain adaptive evo-
lution even more readily than can genetic novelty.
George subjected arguments such as these
to adaptationist scrutiny when they were first
in vogue; he found them wanting (Williams
1966:72–81). The current revivals fare no bet-
ter. Now, as then, three questions immedi-
ately present themselves.
First, is the new phenotype within the design
of an adaptation? This would be the most likely
case given that it is so well-suited to the new
conditions. After all, adaptations are designed
to be sensitive to differences in environmental
conditions; and adaptations involved in devel-
opment are designed to build different phe-
notypes under different conditions. It could
be that neither the environmental conditions
nor the phenotype is really “new.” The new
phenotype is merely a designed response to
these environmental conditions. Indeed, if
the phenotype “shows the unique biological
property of adaptive organization” (Williams
1966:76), that is most likely to be what is hap-
pening. Such phenotypes are hardly in the
vanguard of evolutionary change.
Second, could it be that the new pheno-
types, rather than reflecting adaptations, are
expressing hitherto unexpressed genetic vari-
ation that is present in the population? This
genetic variability would not be part of any
adaptive design but would just happen to get
expressed phenotypically when a change of
conditions disrupts the organism’s environ-
ment and changes the course of develop-
ment. But, if this were how a phenotype came
about, it would be vanishingly unlikely to be
viable at all, let alone more viable than the
existing range of phenotypes. And a mecha-
nism that is vanishingly improbable must also
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Volume 80
be vanishingly unimportant in the course of
evolution.
Third, either way—whether the new phe-
notype is part of the adaptive design or an
extraordinarily improbable response—will
the phenotypic novelty ever get further than
the existing genetic variability happens to
allow? The answer is no. For any “really novel
adaptations” (Williams 1966:78) to enter the
scene, there must be genetic mutation. There
must be a genetic change that makes the
organism responsive to new aspects of the
world-at-large and in such a way as to offer
new solutions to adaptive problems.
Thus, to envisage these phenotypic changes
as evolutionary change is myopic. Indeed, all
such notions should be obliged to take an eye
test. Could that paradigmatic case of adaptive
complexity, the eye, have come about by a
“phenotype-led” process? A process that fails
this test cannot be a major protagonist. Or, as
George succinctly put it, “I question its value
as a model of adaptive evolution” (Williams
1966:75).
• Under some environmental conditions, some
responses that are thought to be instinctive fail to
work normally or to come on stream at all. Think of
Harlow’s experiments with rhesus macaques; daugh-
ters deprived of maternal care were unable, as adults,
to show maternal feelings to their own offspring. This
demonstrates that the standard maternal response is
not an instinct but is learned, a socially or culturally
acquired skill. Thus even maternal care, paradig-
matically an unconditional biological response,
depends crucially on the environment. So genes must
be relatively inconsequential.
What is behind this argument? It is the idea
that a gene-centered view sees instinctive
behavior as surfacing full-blown from innate,
endogenous forces, regardless of the state of
the world and without need for learning. To
correct this putative mistake, the argument
emphasizes how crucial a role environment
plays in instinctive behavior.
Admittedly, this notion of instinct did lurk
within certain strands of ethology in its early
days; and, although it should long have been
laid to rest, it has proved tenacious.
But this concept of instinct has no natural
place in the gene-centered view. On the con-
trary, it is because adaptations rely on regu-
larities in the environment as well as on
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design specifications in the genes that a
“mutation” in the environment can disrupt
normal functioning just as a genetic mutation
can. Think of buntings deprived of stars or
feral children deprived of language speakers.
Similarly, the macaque experiments raise the
questions: What precisely must a female
macaque glean from the presence of her
mother to equip herself for motherhood?
And what are the innate mechanisms that
enable her to do this? These experiments
were too blunt an instrument for tackling
such intricate questions. But there are cases,
such as tracking down a serendipitous “muta-
tion” in the environment of the laboratory
rat, that have nicely demonstrated how to
analyze mothering adaptively (Daly and Wil-
son 1995:1274–1277).
Where does this leave the claim that mater-
nal care is not an instinct but is learned—and,
what is more, learned through social or cul-
tural means? To unravel the confusions here,
we need to parse “learning” adaptively. First, is
the learning an adaptation? Think of humans
learning a mother tongue or buntings learn-
ing to construct a star compass. Alternatively,
is the learning not an adaptation but a skill
that uses aspects of adaptations for purposes
for which they were not designed? Think of
acquiring the skills of reading and writing.
Whichever is the case—adaptation or borrow-
ing—learning from the environment is not an
alternative to instinct; on the contrary, innate
mechanisms are what make learning possible.
Second, if the learning is an adaptation, what
are the innate mechanisms and what are the
environmental resources that it relies on? For
learning language, humans rely crucially on a
social resource, a community of language
speakers; for building compasses, buntings
REFERENCES
Connellan J, Baron-Cohen S, Wheelwright S, Batki A,
Ahluwalia J. 2000. Sex differences in human neo-
natal social perception. Infant Behavior and Devel-
opment 23:113–118.
Cosmides L, Tooby J. 1994. Origins of domain speci-
ficity: the evolution of functional organization.
Pages 85–116 in Mapping the Mind: Domain Specific-
ity in Cognition and Culture, edited by L A Hirschfeld
and S A Gelman. Cambridge: Cambridge Univer-
sity Press.
152.118.24.31 on Mon, 19 Sep 2022 13:45:41 UTC
All use subject to https://about.jstor.org/terms
25
rely crucially on a physical resource, the stars.
So environmental regularities that are social or
cultural do not make the learning less innate
. . . whatever that might mean. Alternatively, if
the learning is not an adaptation but an
acquired skill, what are the adaptations that it
taps into? Reading and writing parasitise mul-
tiple adaptations, greedily and eclectically,
from our language instinct to our opposable
thumb. It is along such lines—across species,
genera and phyla—that motherhood should
be parsed. From the resulting soap bubbles
would emerge both enduring similarities and
intricate differences, reflecting the ingenuity
with which natural selection has solved the
multifarious problems of being a mother.
Conclusion
At the end of his magisterial analysis,
George stated his conviction that the modern
theory of natural selection is “the light and
the way” (Williams 1966:273). That is an apt
description of George’s own profound con-
tribution to evolutionary thinking—not least,
to the concept of adaptation. Only through
adaptations were environments constructed,
and only through understanding adaptations
can we reconstruct them. George’s gene-cen-
tered understanding of adaptations and envi-
ronments shows us still how to clarify some
current evolutionary thought.
acknowledgments
I am grateful to everyone at Stony Brook who orga-
nized the George Williams Symposium and who kindly
invited me to participate. For comments on this paper,
my thanks to Rosalind Arden, Martin Daly, Thomas E
Dickins, David Haig, Isaac Marks, Steven Pinker, Rich-
ard Webb, Margo Wilson, and particularly to Aubrey
Sheiham and Oliver Curry. Above all, my thanks to
George for the difference that he has made.
Daly M, Wilson M. 1995. Discriminative parental solic-
itude and the relevance of evolutionary models to
the analysis of motivational systems. Pages 1269–
1286 in The Cognitive Neurosciences, edited by M S
Gazzaniga. Cambridge (MA): MIT Press.
Dawkins R. 1998. The ‘information challenge.’ The
Skeptic 18(4):21–25.
Dennett D C. 1995. Darwin’s Dangerous Idea: Evolution
and the Meanings of Life. New York: Simon and
Schuster.
This content downloaded from
26 THE QUARTERLY REVIEW OF BIOLOGY
Griffiths P E, Gray R D. 1994. Developmental systems
and evolutionary explanation. Journal of Philosophy
91(6):277–304.
Oyama S. 1985. The Ontogeny of Information: Develop-
mental Systems and Evolution. Cambridge: Cam-
bridge University Press; 2000. Second Edition.
Durham (NC): Duke University Press.
Pinker S. 2004. Why nature and nurture won’t go away.
Daedalus 133:5–17.
Popper K R. 1972. Objective Knowledge: An Evolutionary
Approach. Oxford: Clarendon Press.
Popper K R. 1990. A World of Propensities. Bristol:
Thoemmes.
Popper K R. 1999. All Life is Problem Solving. London:
Routledge.
Robert D, Amoroso J, Hoy R R. 1992. The evolutionary
convergence of hearing in a parasitoid fly and its
cricket host. Science 258:1135–1137.
Robert D, Read M P, Hoy R R. 1994. The tympanal
hearing organ of the parasitoid fly Ormia ochracea
(Diptera, Tachinidae, Ormiini). Cell and Tissue
Research 275:63–78.
Sterelny K. 1995. Understanding life: recent work in
philosophy of biology. British Journal for the Philos-
ophy of Science 46(2):155–183.
Sterelny K, Griffiths P E. 1999. Sex and Death: An Intro-
duction to Philosophy of Biology. Chicago (IL): Uni-
versity of Chicago Press.
152.118.24.31 on Mon, 19 Sep 2022 13:45:41 UTC
All use subject to https://about.jstor.org/terms
Volume 80
Thelen E, Smith L B. 1994. A Dynamic Systems Approach
to the Development of Cognition and Action. Cambridge
(MA): MIT Press.
Tooby J, Cosmides L. 1990a. On the universality of
human nature and the uniqueness of the individ-
ual: the role of genetics and adaptation. Journal of
Personality 58:17–67.
Tooby J, Cosmides L. 1990b. The past explains the
present: emotional adaptations and the structure
of ancestral environments. Ethology and Sociobiology
11:375–424.
Tooby J, Cosmides L. 1992. The psychological foun-
dations of culture. Pages 19–136 in The Adapted
Mind: Evolutionary Psychology and the Generation of
Culture, edited by J H Barkow, L Cosmides, and J
Tooby. New York: Oxford University Press.
von Uexküll J. 1934. Streifzüge durch die Umwelten
von Tieren und Menschen. Berlin (Germany):
Springer. [Reprint. 1957. A stroll through the
worlds of animals and men: a picture book of invis-
ible worlds. Pages 5–80 in Instinctive Behavior: The
Development of a Modern Concept, edited by C H Schil-
ler. New York: International Universities Press.]
West-Eberhard M J. 2003. Developmental Plasticity and
Evolution. Oxford: Oxford University Press.
Williams G C. 1966. Adaptation and Natural Selection: A
Critique of Some Current Evolutionary Thought. Prince-
ton (NJ): Princeton University Press.
This content downloaded from
March 2005 SYMPOSIUM IN HONOR OF GEORGE C WILLIAMS 27

This content downloaded from

152.118.24.31 on Mon, 19 Sep 2022 13:45:41 UTC
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