What Affordances and Niche

The Role of Affordances in the
Evolutionary Process Reconsidered

A Niche Construction Perspective
Rob Withagen and Margot van Wermeskerken

UNIVERSITY MEDICAL CENTER GRONINGEN AND UNIVERSITY OF GRONINGEN
ABSTRACT. Gibson asserted that affordances are the primary objects of

perception. Although this assertion is especially attractive when considered
in the context of evolutionary theory, the role that affordances play in the
evolution of animals’ perceptual and action systems is still unclear. Trying
to combine the insights of both Gibson and Darwin, Reed developed a
selectionist view in which affordances are conceived as resources that exert
selection pressures, giving rise to animals equipped with action systems.
Reed’s advocacy of selectionism, however, has been criticized on several
grounds, among which is an inconsistency with recent trends in evolutionary
thinking. Current developments in evolutionary biology indeed ask for a
reconsideration of the role of affordances in the evolution of perceptual and
action systems. Adopting a niche construction perspective, we reexamine
the role of affordances in the evolutionary process. It is argued that affor-
dances and their utilization, destruction, and creation are central elements
in evolutionary dynamics. The implications for ecological psychology and
evolutionary theory are explored.
KEY WORDS: affordances, ecological psychology, evolution, niche construction,
selectionism
The concept of affordance is arguably one of the most important contributions

of the ecological movement in psychology. It was introduced in the 1960s by
James Gibson, the founder of this movement, to refer to the action possibili-
ties that the environment offers the animal. For example, for humans the
ground is walk-on-able, chairs are sit-on-able, water affords drinking, and so
on. Gibson’s assertion that the animal’s environment consists of affordances
was radical at his time (see, e.g., Costall, 2003; Withagen & Michaels, 2005).
Indeed, with this statement, Gibson took aim at the dominant view that the
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490 THEORY & PSYCHOLOGY 20(4)
meaningful world as one experiences it is a product of the mind. Ever since

the mechanization of the worldview, philosophers and later psychologists
have generally regarded the environment as meaningless, consisting merely
of matter and motion. The experienced meaningful world, then, is thought to
emerge in the perceptual processes. Asserting that the animal’s environment
consists of affordances, Gibson placed meaning in the environment. “The
meaning or value of a thing consists of what it affords” (Gibson, 1982,
p. 407). And these meaningful action possibilities can, according to Gibson,
be directly perceived. In fact, they are the primary objects of perception.
Although, as Gibson (1979/1986, pp. 138–140) himself acknowledged, the
concept of affordance is not without its antecedents, it has been regarded as
one of the most central and original concepts of the ecological approach. Over
the years it has inspired many ecological psychologists (e.g., Costall, 1995;
Fajen, 2005; Heft, 1993; Turvey, 1992; Warren, 1984) as well as researchers
in the fields of human factors (e.g., Flach, Hancock, Caird, & Vicente, 1995),
embodied and situated cognition (e.g., Clark, 1997), anthropology (e.g.,
Ingold, 2000), and archeology (Knappett, 2005).
The concept of affordance fits in nicely with Gibson’s evolutionarily inspired
approach to perception (see, e.g., Gibson, 1966, chap. 9; see also Heft, 2007;
Lombardo, 1987; Looren de Jong, 1995, 1997; Withagen & Chemero, 2009).
Indeed, the idea that animals perceive their environments primarily in terms of
affordances is very attractive from an evolutionary perspective. After all, in
order to survive and reproduce it is of primary importance that animals perceive
what affords eating, locomotion, danger, and so on. Hence, it is quite likely that
animals should evolve so as to perceive the action possibilities in their envi-
ronments. And although it is hard to demonstrate empirically that affordances
are the primary objects of perception, recent studies have revealed that humans
are fairly accurate in perceiving what behavior the environment affords
(e.g., Heft, 1993; Mark, 1987; Oudejans, Michaels, Bakker, & Dolne, 1996;
Oudejans, Michaels, van Dort, & Frissen, 1996; van der Kamp, Savelsbergh,
& Davis, 1998; Warren, 1984; Warren & Whang, 1987).
Although the idea that animals perceive their environments in terms of
affordances is attractive from an evolutionary perspective, the role that affor-
dances play in the evolution of perceptual and action systems is still unclear.
In fact, Reed (1982a, 1985, 1996) was one of the few authors who made
explicit statements about the role of affordances in this process. Being
inspired by both Gibson and Darwin, Reed put the concept of affordance in
the context of evolutionary theory. More precisely, adopting a selectionist
perspective, he developed the view that affordances are resources that exert
selection pressures, giving rise to animals equipped with action systems. Over
the last decade, however, Reed’s evolutionary construal of affordances has
been criticized for being based on “a very conventional (and highly contested)
line of selectionist theorizing within ecological and evolutionary biology”
(Costall, 1999, p. 413; see also Chemero, 2003). However, the critics of Reed
WITHAGEN & VAN WERMESKERKEN: AFFORDANCES IN EVOLUTION 491
have not provided an in-depth analysis of his selectionist view of affordances.

Given that Reed’s ecological perspective is arguably one of the most important
theoretical elaborations of Gibson’s original framework, a thorough exami-
nation of his selectionist view is in order. Furthermore, if the critique is right,
the role of affordances in evolutionary dynamics needs to be reanalyzed. The
present paper aims at doing so.
After a brief exposition of Reed’s ideas, we evaluate his selectionist view of
affordances. It is argued that this view indeed runs counter to current develop-
ments in evolutionary thinking. Among other things, the selectionist view is
criticized for underestimating the evolutionary consequences of animals’ mod-
ification of their environments. Hence, to reexamine the role of affordances in
the evolutionary process, we adopt a niche construction perspective instead.
This view of the evolutionary process emphasizes the animal–environment
coupling in the evolution of both animals and their environments. We will
argue that affordances and their utilization, construction, and destruction
play key roles in evolutionary dynamics. We end our paper by exploring the
implications for ecological psychology and evolutionary theory.
Reed’s Selectionist View of Affordances
The academic work of Reed was influenced by several eminent theorists.

Reed (1993) himself once enumerated the following scholars as his main
sources of inspiration: Darwin, Gibson, Piaget, Marx, and Vygotsky. Although
traces of the theories of all these authors can be found throughout Reed’s
articles and books, the influence of Darwin and Gibson on his thinking is
perhaps dominant and clear from the very beginning of his career. Indeed, in
his first papers, Reed (1978, 1981, 1982b; Reed & Jones, 1977) devoted
considerable attention to Darwin, addressing issues like the lawfulness of
natural selection, selectionism, population thinking, and Darwin’s studies of
earthworms. In several of these papers, Reed (e.g., Reed, 1982b; Reed &
Jones, 1977) already explored the connection between evolutionary thinking
and Gibsonian psychology, a topic that he worked on for the rest of his
career (Mace, 1997).
When it comes to an understanding of animal perception and behavior,
Reed was convinced that evolutionary theory might benefit from an ecological
perspective, and vice versa. An ecological perspective is needed to under-
stand the evolution of organisms. Reed (1982b, 1996) argued that Darwin,
given his emphasis upon the importance of the behavioral ecology, had
already developed such a perspective. In fact, in his studies on earthworms
(Darwin, 1881), the father of modern biology already came close to the
development of the ecological concepts of information and affordances
(Reed, 1982b, 1986). And these concepts, Reed asserted, are crucial in the
understanding of the evolution of psychological functions. Furthermore,
ecological psychology, in turn, might benefit from evolutionary thinking.

In fact, Reed put considerable effort in incorporating evolutionary theory
in his development of Gibson’s (1966, 1979/1986) ecological approach to
psychology.
The ecological approach that Reed developed was relentlessly selectionist.
Following several biologists (e.g., Ghiselin, 1969; see also Darden & Cain,
1989), including the neuroscientist Edelman (1987), Reed argued that selec-
tionist thinking applies at all levels of biological reality, from the immune
system and the selection of neural groups to the selection of animal behavior
and groups of animals. The theory of selectionism that Reed defended was
based on two ideas.1 The first idea is population thinking, which implies,
among other things, that biological entities vary. This idea originated in the
work of Darwin and has been conceived as one of the most important
contributions of his theory (Mayr, 1975/1994, 2001/2002; Sober, 1980).
Population thinking, Reed (1989, 1991) stressed, applies at all levels of
biological organization, from neural cells to individual organisms to groups
of animals. That is, there is variation among all biological entities; they are
unique with respect to each other. The second element of Reed’s selectionist
view is competition. Among all biological entities there is always competition
for resources. It is this omnipresent competition that implies a ubiquitous
selection process. After all, because there is variation among the competitors,
and the resources are not conserved and sometimes scarcely available, the
biological entities that are more successful in exploiting the resources are
selected over others. Especially when the availability is limited, resources
can exert severe selection pressures. What the resources are, however, depends
on the level of biological organization. For instance, cells compete for
inorganic macromolecules, and at the level of behavior and perception there
is competition for affordances and information, respectively (Reed, 1985).
The idea that affordances and information can be conceived as resources,
and that these resources exert selection pressures, is perhaps the most impor-
tant claim of Reed’s ecological approach. As Reed (1996) asserted in his last
portrayal of his ecological perspective:
The fundamental hypothesis of ecological psychology and of this book is
that affordances and only the relative availability (or nonavailability) of
affordances create selection pressure on the behavior of individual organisms;
hence behavior is regulated with respect to the affordances of the environment
for a given animal. (p. 18)
This claim figured prominently in Reed’s work. He used this “selective
retention theory” to explain social development (Reed, 1993). Combining it
with Edelman’s theory of Neural Darwinism, Reed (1989, 1990, 1996) used
it to explain neural development and brain functioning.2 And Reed (1982a,
1988, 1996) relied heavily on selectionist thinking in his theory of action
systems (see Costall, 1999; Withagen & Michaels, 2005). In Reed’s view, the
relative persistence of some affordances in the animal’s environment has
given rise to the evolution of several distinct action systems, enabling the
animal to take advantage of these affordances. Among these systems are
the basic orienting system, the locomotion system, the appetitive system, and
the performatory system, each of which has evolved to take advantage of
different types of resources.
Setting out his ideas about affordances and the evolution of action systems,
Reed (1982a, 1996) backed up his theory with some empirical findings. The
most important evidence that he presented is the phenomenon of convergent
evolution: that is, the fact that animals descended from different lineages
nevertheless evolve similar behavioral patterns and action systems. Reed
(1996), for instance, presented the example of flying behavior, which many
species have evolved independently.
There are ... flying (really gliding) squirrels, frogs, lizards, fish, squid, and
more. Although the morphological adaptations for this form of locomotion
vary widely, the behavioral pattern of jump and glide (often via a sail-like
flap of skin) are strikingly convergent. (p. 41)
Apparently, the persistence of “the gliding potential of air” (p. 41) has given
rise to animals with similar action systems. According to Reed, such conver-
gent evolution is a “powerful proof of the efficacy of affordance in evolution”
(p. 43). It suggests that evolution gives rise to animals that are capable of
taking advantage of the relatively persistent affordances in the environment.
In fact, Reed (1982a, 1996) asserted that affordances exerting selection
pressures is a dominant force in evolution by natural selection, and he quoted
Ricklefs (1990), who argued that, “Wherever one looks, one finds conver-
gence, and this reinforces our belief that community organization depends on
local conditions of the environment more than it does on the evolutionary
origins of the species that comprise the community” (p. 672).
An Evaluation of Reed’s Selectionist View of Affordances
Reed’s work is an important attempt at developing an ecological psychology

that is rooted in Darwinian thinking. It is conceptually interesting in that it
tries to connect Gibson’s and Darwin’s perspectives via the central ecologi-
cal concept of affordance. In addition, Reed presented evidence for his
theory of the role of affordances in the process of evolution. However, as
mentioned in the introduction, Reed’s perspective has been criticized for
being based on a contested line of thinking within evolutionary theory
(Chemero, 2003; Costall, 1999). In this section, we evaluate Reed’s selec-
tionism in the light of recent developments in evolutionary biology. We first
briefly sketch some (recent) arguments against the selectionist theory of
evolution and evaluate whether they apply to Reed’s view. Then, we address
the question of whether convergent evolution does support Reed’s selectionist
view of affordances.
Animal–Environment Mutuality
Over recent decades, the selectionist theory of evolution has been criticized
on several grounds (see, e.g., Godfrey-Smith, 1996; Ingold, 2000, 2004,
2007; Jablonka & Lamb, 2005; Lewontin, 1983, 2001; Odling-Smee, Laland,
& Feldman, 2003; Oyama, Griffiths, & Gray, 2001; H. Rose & S. Rose, 2000;
S. Rose, 1998; Ruse, 2006). However, many of the recent arguments have
been anticipated by Lewontin (1983) in his now classic book chapter “Gene,
Organism, and Environment.” Hence, we rely heavily on Lewontin’s per-
spective in our analysis of the selectionist view.
It is important to note that the above critics of the selectionist view do not
take aim at the idea of variation and selection. In fact, they fully embrace this
central tenet of evolutionary theory. What they criticize is the lack of rela-
tional thinking in the traditional selectionist view. Generally speaking, this
view treats the animal and the environment as two separate entities. The envi-
ronment is said to exist prior to and independent of the animal. And through
the process of evolution by natural selection, organisms evolve that fit in to
this preexisting environment. This view of the evolutionary process is
perhaps best captured in the metaphor of adaptation (Lewontin, 1983, 2001).
This metaphor implies that the environment “poses the problems,” and the
animal “posits solutions.” As Lewontin (1983) put it, “To make the metaphor
of adaptation work, environments or ecological niches must exist before the
organisms that fill them” (p. 280). But, as Lewontin (1983, 2001) argued, this
view of the evolutionary process and the allied conception of the environment
is seriously flawed. There is no environment independently of organisms;
there is no environment that has to be filled. After all, an environment implies
an organism. Like Gibson (1979/1986), Lewontin (2001) made a distinction
between the physical world and the environment. The physical world exists
independently of organisms—it will not cease to exist in the absence of
species. The environment, on the other hand, is determined by the organisms:
An environment is something that surrounds or encircles, but for there to
be a surrounding there must be something at the center to be surrounded.
The environment of an organism is the penumbra of external conditions that
are relevant to it because it has effective interactions with those aspects of
the outer world. (Lewontin, 2001, pp. 48–49)
Earlier critics of Reed have asserted that this point of critique applies to his
selectionist view of affordances (Chemero, 2003; Costall, 1999). Indeed,
Reed has been claimed to conceive affordances as resources that exist prior
to the animals (e.g., Chemero, 2003). However, to our minds, this critique
needs more nuance. By and large, ecological psychologists have argued for a
relational conception of the environment that at first blush is consistent with
current trends in evolutionary thinking. They, too, argue that the properties of
an animal determine what constitutes the environment. Indeed, this idea is
central in the concept of affordance. As Gibson (1979/1986) pointed out, “I
mean by [affordance] something that refers to both the environment and the
animal in a way that no existing term does. It implies the complementarity of
the animal and the environment” (p. 127). In other words, an affordance does
not exist independently of an animal; rather, it implies a fit between the
animal and environment. For example, it is the properties of a cup (i.e., its
size and form) relative to the action capabilities of my body that make it
graspable. Although the ontological status of affordances is still a highly
debated topic, the vast majority of ecological psychologists take this mutuality
as central in their conceptualization of action possibilities and, thus, the envi-
ronment (e.g., Chemero, 2003; Costall, 1995; Heft, 2001; Stoffregen, 2003;
Turvey, 1992). And Reed was no exception. Although he openly criticized
some mutualists (see, e.g., Reed, 1996, p. 26) and claimed that affordances
exist independently of animals, his ideas on affordances and resources also
had a mutualist flavor. For example, after complaining about the absence of
an explicit definition of the concept of resource in biology, Reed (1985)
offered a tentative one, stressing that “although resources exist independently
of the processes they promote, what makes something a resource is its relation
to such a process, its causal power to further that process, or type of process”
(p. 360; see also Reed, 1993, p. 54). According to Reed, the existence of a
biological process is a necessary prerequisite for something to be a resource.
Indeed, in cases where a biological process has not yet evolved, he talked
about a “potential resource” (Reed, 1993, p. 54). The complementarity of
animal and environment was also central in Reed’s (1996) last portrayal of
affordances, where he defined affordances relative to populations. Hence, in
defining affordances, Reed did not claim that affordances preexist animals;
instead he defined them relationally.
However, the earlier critics (Chemero, 2003; Costall, 1999) of Reed seem
right when it comes to his evolutionary analyses of affordances and action
systems. Indeed, there seems to be a tension between Reed’s definition of
affordances and his evolutionary examinations. In the latter, he often did not
bear witness to a relational conception of the environment. In fact, the view
that he espoused at certain points is the classic selectionist view of evolution
in which species are molded by natural selection to fit preexisting environ-
ments. To reiterate an example, Reed (1996) suggested that “the gliding
potential of air” (p. 41) has given rise to the evolution of a particular flying
behavior. Thus, the environmental property (the gliding potential of air) exists
prior to the evolution of flying behavior, and animals evolve so as to exploit
this particular resource. Reed adopted here the classic selectionist line of
thinking at which authors like Lewontin (1983, 2001) took aim. Hence,
although Reed argued that affordances should be defined relative to a popu-
lation, his evolutionary account of action systems suggests at certain points
that affordances preexist a population. But if one takes mutualism seriously,
affordances do not preexist animals; they do not exert selection pressures,
explaining the origin of action systems. Rather, affordances can only provide
the context of selection after the animal has evolved (Costall, 1999). After
all, as just mentioned, Gibson’s concept of affordance implies a fit between
animal and environment. As an example, the coming into being of the affor-
dance “walk-on-able” depends on the evolution of a certain locomotion system
(see also Chemero, 2003).
Animals Construct Their Environments

A second, and related, point of critique is that the selectionist view of evolution
inadequately characterizes the causal relation between animal and environment
in the evolutionary process. The selectionist view conceives this relation as a
one-way causation. Through natural selection, the environment is said to
mold the animal, but the animal does not significantly shape the environment.
However, as argued by several authors (Laland, Odling-Smee, & Feldman,
1996, 2000, 2004; Lewontin, 1983, 2001; Odling-Smee et al., 2003), this
view is mistaken. In the evolutionary process, there is no one-way causation
between animal and environment. Animals are not only molded by the envi-
ronment, they also alter their environment, and do so in a non-random way.
Indeed, animals and their environments evolve together. Hence, to capture the
relation between animal and environment, Lewontin (1983, 2001) suggested
that the metaphor of adaptation needs to be replaced by the metaphor of
construction. Animals not only determine what constitutes the environment,
they also literally construct it. As Lewontin (1983) put it in an often-cited
quote, “Organisms do not adapt to their environments; they construct them
out of the bits and pieces of the external world” (p. 280).
Lewontin’s (1983, 2001) idea that animals construct their environments has
a long history. In fact, and as illustrated below, it has a significant place in
Darwin’s original work.3 And also later authors like Morgan and Waddington
stressed the animal’s modification of the environment (see, e.g., Costall, 1993;
Odling-Smee et al., 2003). However, over the last decade the idea of animals’
alteration of their environments has been elaborated significantly under the
rubric of “niche construction” (see, e.g., Laland et al., 1996, 2000, 2004;
Odling-Smee et al., 2003). It is emphasized that animals’ modification of the
environment is not primarily a human affair, as some authors had suggested
(e.g., Dobzhansky, 1955). In fact, niche construction is ubiquitous in the animal
kingdom—even the lowest animals are capable of it. Although it remains
underappreciated by the vast majority of ecological psychologists (see Costall,
1995; Heft, 2007; and below), the idea that animals modify their environment
is by no means new in the ecological literature. Costall (1995, 2004), for
instance, had argued that many affordances are created by humans. In a similar
vein, Heft (2007) and Ingold (2000) brought the process of niche construction
to attention when arguing that the human environment is largely a product of
social processes. And also Reed stressed animals’ modification of their
environments at several points (see, e.g., Reed, 1985, 1996).
What is important for present purposes, however, is that this process

of niche construction has proven to have consequences for evolutionary
dynamics (Laland et al., 1996, 2000, 2004; Odling-Smee et al., 2003). As we
will discuss below, the process of niche construction (a) can change the
selection pressures; (b) implies an inheritance not only of genes but also of
environments; and, (c) consequently, has proven to be capable of creating
new evolutionary equilibria and trajectories. Hence, animals do not evolve
so as to fit in a preexisting environment. Rather, animals and their environ-
ments evolve together, and the animals’ alteration of the environment has a
constitutive role in this co-evolution.
Although Reed (1996, p. 27) was openly critical of Lewontin’s metaphor of
construction, he often stressed that animals can modify their environments.
According to Reed’s perspective, the animal’s environment is far from rigid.
In fact, he expressed serious concerns about the fact that humans ruin their
environment. At certain points, Reed (see, e.g., 1985, p. 369) even hinted that
animals’ modification of their environments might change the selection
pressures. However, the idea of niche construction and its evolutionary con-
sequences has gathered momentum primarily over the last decade, that is, after
Reed’s death in 1997. Hence, it is of no surprise that this idea was not central
in his evolutionary analyses of affordances and action systems. However, as
we will argue below, the niche construction perspective suggests alternative
roles for affordances in evolutionary dynamics. Hence, to bring the evolutionary
analysis of affordances up to date with current trends in biology, animals’
modification of the environment needs to be incorporated in the analysis.
The Issue of Convergent Evolution

Before we proceed to the niche construction perspective, there is one final
argument that we need to consider. As mentioned above, Reed (1996) pre-
sented evidence for his selectionist view of affordances. According to Reed,
the convergent evolution of similar action systems provides “powerful
proof for the efficacy of affordances in evolution” (p. 43). What is the status
of this argument? Does convergent evolution cast doubt on the metaphor of
construction?
Over the years, the phenomenon of convergent evolution has often been
used as an argument for the selectionist theory of evolution. The fact that
animals from different lineages nevertheless evolve similar behavioral patterns
has been treated as evidence for the view that animals evolve so as to fit a
preexisting niche (e.g., Grinnell, 1924; for a brief overview see Harmon,
Kolbe, Cheverud, & Losos, 2005). It is important to note that Reed (1996)
used convergent evolution primarily as an argument for the effectiveness of
affordances in the evolutionary process. However, as mentioned above, at
certain points Reed also presented convergent evolution as an argument for
the selectionist view of evolution. Once again, he did argue that “the gliding
potential of air” (p. 41) has given rise to the evolution of animals with similar
action systems. However, as already argued by Lewontin (1983), convergent
evolution does not provide irrefutable evidence for the selectionist theory of
evolution. Indeed, it is fully compatible with the metaphor of construction.
What Lewontin emphasized is that there are constraints on the co-evolution
of animal and environment. “The coupled equations of coevolution of organ-
ism and environment are not unconstrained. Some pathways through the
organism–environment space are more probable than others, precisely because
there are real physical relations in the external world that constrain change”
(Lewontin, 1983, p. 283). Hence, contrary to Reed, Lewontin would not
conceive the “gliding potential of air” as a resource that exerts selection pres-
sures, leading to a convergent evolution of flying behavior. Rather, Lewontin
would conceive of this physical property as a constraint on the evolution of
the animal–environment system. “When there is strong convergence ... this
should be taken as evidence about the nature of constraints on development
and physical relations, rather than as evidence for pre-existing niches” (p. 283).
How to decide between the constructionist and the selectionist explanation
for convergent evolution is a difficult issue that seems to be largely concep-
tual in nature.4 What is important for present purposes, however, is that both
the selectionist view and the constructionist view can deal with the phenome-
non of convergent evolution. Hence, it does not provide conclusive proof
for Reed’s selectionist view. It is also in keeping with the constructionist
construal of affordances that we defend below.
A Niche Construction Perspective
Reed’s ecological perspective is an interesting attempt at making a connec-

tion between Gibsonian psychology and Darwinian theory via the concept
of affordances. Yet, as we have seen, his selectionist account of affordances
cannot stand up to some current trends in evolutionary biology. The fact that
the environment (i.e., the affordances) exists by virtue of animals seems to
seriously downplay the role that affordances play in evolutionary dynamics.
After all, it means that affordances cannot account for the origin of action
systems, as Reed suggested. In the portrayal of his theory of affordances,
Chemero (2003) made some brief comments about an alternative relation
between affordances and the evolutionary process. He argued that affor-
dances are “tied to evolution” (p. 190), but surmised that they do not have a
causal role in the evolution of perceptual and action systems.
Affordances, which are the glue that holds the animal and the environment
together, exist only in virtue of selection pressures exerted on animals
by the normal physical environment [emphasis added]. They arise along
with the abilities of animals to perceive and take advantage of them.
(Chemero, 2003, p. 190)
We agree that affordances arise along with the evolution of action systems,
but believe that affordances do have a constitutive role in evolutionary
dynamics. Indeed, as we will argue below, some current thinking on evolu-
tion opens up new ways to consider the role of affordances in the evolutionary
process. In this section we sketch the niche construction perspective on
evolution, a contemporary evolutionary theory that takes animals’ modification
of the environment to be central. Our sketch and the presented examples are
based largely on the work of Odling-Smee, Laland, and Feldman (2003;
Laland et al., 1996, 2000, 2004). After a portrayal of their theory, we reex-
amine the role of affordances in the evolutionary process. It is argued not only
that affordances constitute the context of selection, but also that animals’
destruction and construction of affordances change this context, demonstrating
the key roles affordances play in evolutionary dynamics.
Outline of the Theory and Some Examples

The fact that animals modify their environments is easily observable. The
vast majority of ants and birds build nests, beavers build dams, spiders
construct nests, and humans create houses. As mentioned above, the idea of niche
construction has a relatively long history. Indeed, and as described below, it
can be traced back to Darwin. However, the renewed interest in niche con-
struction has revealed that this process has “hidden complexities” (Odling-
Smee et al., 2003, p. xi), especially for evolutionary dynamics. Indeed, recent
research has spelled out specific evolutionary consequences of the process of
niche construction (Laland et al., 1996; Odling-Smee et al., 2003). That niche
construction is evolutionarily consequential, however, is not hard to see. After
all, it has long been acknowledged that environmental changes often have
consequences for selection pressures (Endler, 1986). And if animals can bring
about these changes by constructing niches, it is likely that this process
affects the selection pressures to which they and other animals are exposed.
An early observation of this process of niche construction and its conse-
quences was provided by Charles Darwin. In his last book, Darwin (1881)
wrote about the actions and mental powers of earthworms. A large part of this
book is devoted to demonstrating that worms, through the creation of veg-
etable mold, are responsible for changing the structure and chemistry of the
topsoil. And this has profound effects on the ecosystem. As every gardener
knows, vegetable mold provides a rich structural basis from which to grow
plants. However, the change in the topsoil has implications also for the worms
themselves. Initially, worms were structurally not very well adapted to the
topsoil. Indeed, their epidermis is very sensitive and needs to be kept warm
and wet. They are better suited to live in water than in the soil. However, by
changing the chemical composition of the soil, digging burrows and dragging
leaves in, earthworms created an environment that better suits their physio-
logical make-up. This demonstrates that a fit between an environment and an
FIGURE 1. Schematic conception of the niche construction perspective

on evolution. Animals modify their environment, which can change
the selection pressures to which they are exposed. Furthermore, through
the construction of niches, animals also create an ecological inheritance
(redrawn from Laland et al., 2000).
animal can be established in basically two ways. First, as conventional

evolutionary theory states, a species can be molded to fit an environment
through the process of natural selection. Second, animals can also establish a
fit by modifying their environment (Odling-Smee et al., 2003). And this, of
course, can change the selection pressures to which they are exposed.
Another important corollary of the niche construction perspective is an
alternative system of inheritance. If the environment is modified in a rela-
tively persistent way, then what is “passed on” to the offspring are not only
the genes but also the environment (see Figure 1). Hence, the theory of
niche construction suggests an “ecological inheritance,” an inheritance of
niches (Odling-Smee et al., 2003). As an example, the burrowing activities
of a myriad of earthworms change the structure of the topsoil in a relatively
persistent way. Hence, the worms’ offspring will inherit not only genes, as
standard evolutionary theory dictates, but also an environment that better
suits their epidermis.5 Thus, the change in the selection pressures resulting
from the burrowing activity is also relatively persistent.
There are three points that are worth emphasizing at this stage. First, it is
important to note that the theory of niche construction recognizes the
conventional idea that the environment is the context of selection. Indeed, as
depicted in Figure 1, it also states that the environment can mold the species.
The theory is fully consistent with the central tenet of evolutionary theory: the
mechanism of variation and selection. Hence, there is no strong distinction
between the niche construction perspective and the idea of the environment
as the context of selection. Second, niche construction theory also recognizes
that geological and hydrological processes can change the selection pressures
to which a population is exposed (see, e.g., Griffiths & Gray, 2001). Indeed,
a volcanic eruption can change the context of selection of many species in a
certain habitat. However, what the theory of niche construction adds to the
conventional view is that the animals’ modifications of their environments can
also be evolutionarily consequential. Third, although these modifications
can determine evolutionary dynamics, they do not always do so. Indeed, the
environmental changes that animals bring about do not always change the
context of selection. Over the last decade or so, Laland, Odling-Smee, and
Feldman have significantly developed their niche construction perspective,
conceptually, empirically, and mathematically (see, e.g., Odling-Smee et al.,
2003). We will limit ourselves here to the conceptual development and some
examples of the process.
In their book, Odling-Smee et al. (2003) defined a niche of a population as
“the sum of all the natural selection pressures to which the population is
exposed” (p. 40). They emphasized that this definition implies what we call
animal–environment mutualism—“the selection pressures are only selection
pressures relative to specific organisms” (p. 40). To develop a definition of
niche construction, Odling-Smee et al. adopted Bock’s (1980) classic scheme
that decomposes animals into features and environments into factors. In this
scheme, evolution is conceptualized as a process that promotes a match of
factors and features, what Bock termed a synerg. As an example, there is a
match between the outer shape of the fish (the feature) and its fluid environ-
ment (the factor). And matches between animal and environment, Odling-
Smee et al. asserted, can be established by both natural selection and niche
construction. Niche construction, then, is said to occur
when an organism modifies the feature–factor relationship between itself
and its environment by changing one or more factors in its environment,
either by physically perturbing factors at its current location in space and
time, or by relocating to a different space-time address, thereby exposing
itself to different factors. (Odling-Smee et al., 2003, p. 41)6
Odling-Smee, Laland, and Feldman (Laland et al., 1996, 2000, 2004;
Odling-Smee et al., 2003) used their concept of niche construction to refer to
almost any modification that the organism brings about in its environment.
The examples they present range from photosynthesis to the building of
houses. However, because the present paper is about the role of affordances
in the evolution of action systems, we focus here on the behavioral level. At
this level, numerous anatomical and behavioral adaptations to prior niche
construction activity have now been listed (see Odling-Smee et al., 2003,
for an extensive overview). For example, many animals have evolved certain
features to construct niches with more efficiency. Animals that dig burrows,
for instance, have evolved special limbs and claws that make the burrowing
activity easier. In addition, the construction of niches has also led to the
evolution of new features. For example, the burrowing activities of certain
animals have given rise to selection pressures for place learning. Similarly, in
response to the web life of their ancestors, some web spiders have evolved
responses to predators on their web; others have evolved an extra claw on
each leg that facilitates movement on the web (Preston-Mafham & Preston-
Mafham, 1996). This indicates that animals evolve anatomical and behavioral
adaptations in response to the niche construction activities of their ancestors.
Hence, the construction of niches affects the evolution of perceptual and
action systems.
The Role of Affordances from a Niche Construction Perspective

Although evolutionary biologists have not used the concept of affordance to
elucidate the process of niche construction, affordances clearly have a role in
this process. There are basically two aspects to the niche construction per-
spective on evolution. First, as mentioned earlier, this perspective recognizes
the conventional idea that through natural selection, the environment molds
the species. At the behavioral level, affordances are surely involved in this
process. Indeed, they often, if not always, form the context of selection,
although not in the sense as Reed (1982a, 1996) sometimes suggested that
affordances preexist animals and give rise to a convergent evolution of action
systems that are capable of utilizing them. Rather, affordances arise along
with the evolution of action systems and provide the context of selection after
they have evolved (Costall, 1999). Indeed, animals compete for things that
are edible, for places that afford shelter, for mates that afford reproduction,
and so on. In their struggle for survival and reproduction, animals compete
for affordances, and the animals that are better in exploiting the affordances
than their competitors have a greater chance of producing offspring.
The second, and more important, aspect of the niche construction perspective
is animals’ alteration of the environment. There is a “world-in-formation”
(Ingold, 2006, p. 12) and animals are key players in this process (Costall,
1995; Heft, 2007). This aspect of the niche construction perspective suggests
important roles for affordances in the evolution of action systems that, as far
as we know, have hitherto not been recognized. First, the process of niche
construction requires an environment consisting of affordances. After all, in
order for niche construction to occur, the environment needs to afford this
process. As examples, for a spider a place might afford building a web; and
for a beaver a certain spot in a creek might afford building a dam. Second,
and related to this, the niche construction perspective attributes an important
role to action in evolutionary dynamics. Ecological psychologists often make
a distinction between performatory and exploratory action. Performatory
activity basically consists of the utilization of affordances. Exploratory action, by
contrast, is aimed at obtaining information, and is thus central in perceiving
the environment. Both types of action play important roles in evolutionary

dynamics. After all, the construction of a niche requires both the perception
and utilization of affordances, that is, exploratory and performatory behavior.
And if this construction is evolutionarily consequential, then the perception
and utilization of the affordances that led to the niche are not merely a result
of an evolutionary process; rather, they (partly) determine this process. Third,
niche construction not only requires the utilization of affordances, it also
consists in a change in affordance layout. Recall that Odling-Smee et al.
(2003) defined niche construction as a process by which animals modify “the
feature–factor relationship between itself and its environment” (p. 41). Hence,
a change of the environment only counts as niche construction if it is relevant
to the animal that brings about this change. And this includes the creation and
destruction of affordances. Indeed, animals create tools, contrive places that
afford shelter, destroy things and organisms that afford harm, grow food, and
so on (Costall, 1995; Heft, 2007; Ingold, 2000). Hence, animals often create
and destroy affordances and pass the modified environment on to their
offspring. Thus, the ecological inheritance encompasses an inheritance
of affordances. It is important to emphasize, however, that changes in the
affordance layout are not exclusively the result of animal activity. Indeed, as
mentioned above, geological and hydrological processes can also alter the
affordances in an animals’ habitat. And although this does not count as
niche construction, it is recognized by the theory. Furthermore, it is impor-
tant to reiterate that creating a niche is of course not always evolutionarily
consequential–it does not have to change the context of selection. However,
as we have seen, niche construction can alter the evolutionary trajectory.
Indeed, animals have evolved specific action systems in response to the niche
construction activities of their ancestors. Hence, affordances not only
form the context of selection that animals encounter; by creating and destroying
affordances, animals also construct this context and thereby affect the
evolutionary dynamics.
Implications for Ecological Psychology and Evolutionary Theory
This paper has been about the role of affordances in evolutionary dynamics.
Although Gibson’s (1966, 1979/1986) idea that animals perceive their environ-
ments primarily in terms of affordances is very attractive from an evolutionary
perspective, the role that affordances play in the evolution of animals was still
unclear. Reed (1982a, 1985, 1996) sketched a selectionist view in which affor-
dances exert selection pressures, giving rise to the evolution of action systems.
To his mind, this selectionist approach to evolution is consistent with the eco-
logical approach to perception and action. As he put it, “because ecological psy-
chology begins with the study of how animals encounter their environment,
selectionism is a natural ally of the ecological approach” (Reed, 1996, p. 187).
However, current developments in evolutionary biology have seriously

questioned the selectionist view of evolution that Reed espoused. Indeed,
animals do not so much encounter the environment as construct it. And this
construction, current thinking in biology indicates, can be evolutionarily
consequential. We reexamined the role of affordances in the evolutionary
dynamics from the niche construction approach to evolution, arguing that
affordances play key roles in the evolutionary process. We end our paper by
exploring some implications of our constructionist construal of affordances for
ecological psychology and evolutionary theory.
Let us start by stating that we agree with Reed that ecological psychology
might benefit from evolutionary theory and vice versa. Animals’ perceptual
and action systems have evolved and, thus, theories of their functioning
should be consistent with evolutionary considerations (see also Withagen,
2004; Withagen & Chemero, 2009; Withagen & van Wermeskerken, 2009).
Indeed, an evolutionary approach can provide the “background conditions”
(Heft, 2007, p. 87) for an ecological perspective on perception and action.
The present constructionist construal of affordances suggests that in under-
standing the evolution and working of action systems, the animal–environment
reciprocity should be taken as central. After all, as niche construction theory
demonstrates, the evolution of animals is partly determined by the modifica-
tions that animals bring about in their environments. This implies a shift in
focus on the animal–environment system within ecological psychology. As
mentioned above, ecological psychologists generally take animal–environment
mutuality as central in their description of the environment. However, up to
this point, ecologically motivated studies of affordances have focused almost
exclusively on the perception and utilization of them. What remains under-
appreciated is the fact that many affordances are created and destroyed by
animals (Costall, 1995; Heft, 2007). The present constructionist construal of
affordances brings this aspect to the fore. Indeed, it states that the animal’s
alteration to the affordance layout needs to be taken into account not only to
understand the true reciprocity of animal and environment, but also to under-
stand the evolution and functioning of the animal’s perception–action systems
(Heft, 2007). This brings us to the implications of our constructionist construal
of affordances for evolutionary theory.
As mentioned above, evolutionary biologists have, as far as we know, not
used the concept of affordances to elucidate the process of evolution in gen-
eral and that of niche construction in particular. However, as mentioned in note
6, there is a serious struggle with clarifying the process of niche construction.
To our reading, the same holds true for conceptualizing the environment.
Presumably, the concept of affordance might be of help here. As Lewontin
(1983) stated,
Organisms determine what is relevant. While stones are part of a thrush’s
environment, tree bark is part of a woodpecker’s, and the undersides of
leaves part of the warbler’s. It is the life activities of these birds that
determine which parts of the world, physically accessible to all of them,

are actually parts of their environments. (pp. 280–281)
Like Darwin (see Reed, 1982b, 1996), Lewontin came close to the develop-
ment of the concept of affordances here. However, it seems that this ecologi-
cal concept can further specify his already mutualist conception of the
environment. Indeed, describing the action possibilities of the environment for
a particular animal, an affordance-analysis explains not only that stones are
relevant to thrushes (and are thus part of their environment), but also why. It
describes what an environmental property means to an animal, what its rele-
vance is. Given that at a behavioral level, animals compete for meaningful
properties in the struggle for existence and reproduction (e.g., items that are
edible, places that afford shelter, mates that afford reproduction), a concept
that captures this meaning seems useful. In addition, the concept of affordance
might also help to clarify the process of niche construction. In fact, reconcep-
tualizing the niche construction process in terms of affordances makes it more
concrete. Recall that Odling-Smee et al. (2003) defined this process in rather
abstract terms as the organism’s modification of “the feature–factor relation-
ship between itself and its environment” (p. 41). As argued in the present
paper, what this process actually consists in at a behavioral level is the
perception, utilization, destruction, and creation of affordances. Indeed, in
constructing a niche, animals perceive and utilize affordances and by so doing
create and destroy other action possibilities. Hence, an affordance-analysis
cannot only help in clarifying how the niche is constructed, it also helps in
understanding what the modification of a niche consists in. Indeed, animals
bring about changes in the affordance layout and this modified layout is
passed on to the offspring. Thus, as argued above, the “ecological inheritance”
encompasses an inheritance of affordances. Hence, not only can evolutionary
theory serve as the foundation for ecological psychology, but ecological
psychology, and the concept of affordances in particular, might also contribute
to the understanding of the evolutionary process.
Notes
1. For an overview of theories of selectionism, see Darden & Cain (1989).
2. It is interesting to note that Reed (1989, 1996) criticized Edelman for his “limited
psychology.” According to Reed, Edelman’s theory of Neural Darwinism neglected
the meaningful environmental properties that are involved in the neural selection
process (i.e., affordances). In Reed’s view, however, competition for affordances
should be incorporated in a theory of the development of the brain: “it is not the
animal’s brain that organizes its world, but the evolutionary ecology of the animal
that organizes its brain” (Reed, 1996, p. 69).
3. Lewontin (1983, 2001) suggested that Darwin made a strong distinction between
animal and environment. However, this seems to be a misrepresentation of
Darwin’s view. Indeed, Darwin emphasized the mutuality of animal and environ-
ment at several points (see, e.g., Costall, 2004).
4. As mentioned above, Lewontin (1983) argued that the main problem with the
selectionist view is how to define the preexisting environment.
More paradoxical is the necessity of defining environment without organisms.

... But what laws of the physical universe can be used to pick out the possible
environment waiting to be filled? In fact, we only recognize an “environment”
when we see the organism whose environment it is. (p. 280)
5. There has been quite some work on inheritance systems lately, demonstrating that
what animals inherit includes much more than genes (e.g., Jablonka & Lamb,
2005; Oyama et al., 2001). However, many of these inheritance systems are not
relevant to the issue of affordances in the evolutionary process. Hence, we limit
ourselves here to the ecological inheritance.
6. It is important to note that there has been quite some discussion on the definition
of niche construction (e.g., Godfrey-Smith, 2000; Laland et al., 2000; Reed, 1996;
van der Steen, 2000). Some authors found the choice of the term “construction”
unfortunate; others have argued that relocating to a different place does not count
as niche construction. The definition we adopt here has been formulated partly in
response to severe criticism on an earlier definition (see Laland et al., 2000) and
has been used by its founders ever since (Laland, Odling-Smee, & Gilbert, 2008;
Odling-Smee et al., 2003). However, we agree with other authors (e.g., Godfrey-
Smith, 2000) that relocating to a different place is not really an instance of niche
construction. Hence, we limit ourselves here to the modifications that the animal
brings about in its environment.
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ACKNOWLEDGEMENTS. We are grateful to Alan Costall, two anonymous

reviewers, and the editor for very helpful advice and comments on earlier
drafts of this paper.
ROB WITHAGEN is Assistant Professor at the Center for Human Movement

Sciences at the University Medical Center Groningen and the University
of Groningen. His research is concerned with the naturalization of per-
ception and action. ADDRESS: Center for Human Movement Science,
University Medical Center Groningen, PO Box 196, 9700 AD Groningen,
The Netherlands. [email: R.G.Withagen@med.umcg.nl]
MARGOT VAN WERMESKERKEN is a Ph.D. student at the Research Institute

MOVE, Faculty of Human Movement Sciences at the VU University
Amsterdam. Her research focuses on developmental and learning aspects of
action and perception. ADDRESS: Faculty of Human Movement Sciences, VU
University Amsterdam, Van der Boechorststraat 9, 1081 BT Amsterdam,
The Netherlands. [email: M.vanWermeskerken@fbw.vu.nl]

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The Role of Affordances in the

Evolutionary Process Reconsidered

Rob Withagen and Margot van Wermeskerken

ABSTRACT. Gibson asserted that affordances are the primary objects of

The concept of affordance is arguably one of the most important contributions

THEORY & PSYCHOLOGY VOL. 20 (4): 489–510

meaningful world as one experiences it is a product of the mind. Ever since

have not provided an in-depth analysis of his selectionist view of affordances.

Reed’s Selectionist View of Affordances

The academic work of Reed was influenced by several eminent theorists.

ecological psychology, in turn, might benefit from evolutionary thinking.

An Evaluation of Reed’s Selectionist View of Affordances

Reed’s work is an important attempt at developing an ecological psychology

Animals Construct Their Environments

What is important for present purposes, however, is that this process

The Issue of Convergent Evolution

A Niche Construction Perspective

Reed’s ecological perspective is an interesting attempt at making a connec-

Outline of the Theory and Some Examples

FIGURE 1. Schematic conception of the niche construction perspective

animal can be established in basically two ways. First, as conventional

The Role of Affordances from a Niche Construction Perspective

the environment. Both types of action play important roles in evolutionary

Implications for Ecological Psychology and Evolutionary Theory

However, current developments in evolutionary biology have seriously

determine which parts of the world, physically accessible to all of them,

More paradoxical is the necessity of defining environment without organisms.

Reed, E.S. (1985). An ecological approach to the evolution of behavior. In T. Johnston

Withagen, R., & Chemero, A. (2009). Naturalizing perception: Developing the

ACKNOWLEDGEMENTS. We are grateful to Alan Costall, two anonymous

ROB WITHAGEN is Assistant Professor at the Center for Human Movement

MARGOT VAN WERMESKERKEN is a Ph.D. student at the Research Institute

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