Ofrecimientos Evolucion

Biology & Philosophy (2020) 35:30
https://doi.org/10.1007/s10539-020-09747-1
The evolutionary role of affordances: ecological

psychology, niche construction, and natural selection
Manuel Heras‑Escribano1,2
Received: 11 March 2019 / Accepted: 10 April 2020 / Published online: 22 April 2020
© Springer Nature B.V. 2020
Abstract
This paper aims to examine the evolutionary role of affordances, that is, the pos‑
sibilities for action available in our environments. There are two allegedly compet‑
ing views for explaining the evolutionary role of affordances: the first is based on
natural selection; the second is based on niche construction. According to the first,
affordances are resources that exert selection pressure. The second view claims that
affordances are ecological inheritances in the organism’s niche that are the prod‑
uct of a previous alteration of the environment. While there seems to be a mutu‑
ally exclusive definition of affordances in each of these views, I argue in this paper
that the views are not competing but, rather, complementary. In this sense, affor‑
dances play the role of either resources or ecological inheritances depending on the
temporal stage of the evolutionary process. I make this argument by analyzing how
natural selection and niche construction affect each other even when they function
independently from each other. In this light, if these two evolutionary mechanisms
exert their power in parallel but at two different stages in the evolutionary history of
a given econiche, then there is room to claim that affordances can be understood as
both resources and ecological inheritances. This dual aspect of affordances shows
their evolutionary role.
Keywords Affordances · Evolution · Niche construction · Natural selection
* Manuel Heras‑Escribano
[email protected]
1
Department of Logic and Philosophy of Science, University of the Basque Country,
Donostia‑San Sebastián, Spain
2
Embodied Cognitive Science Unit, Okinawa Institute of Science and Technology, Onna, Japan
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Vol.:(0123456789)
30 Page 2 of 27 M. Heras‑Escribano
Introduction
Affordances are the possibilities for acting in our environment: objects of a cer‑
tain size are graspable, floors are walkable, obstacles are avoidable, etc. These
affordances are the main object of study of ecological psychology, a branch of
psychology pioneered by James and Eleanor Gibson. The main aim of this paper
is to show how we should understand the evolutionary role of affordances. To that
end, some arguments are offered in favor of the compatibility of affordances with
two different evolutionary mechanisms, namely, niche construction and natural
selection.
Theorists of the ecological approach often claim that their view is linked to
evolution and biology (Gibson 1979/2015, Michaels and Carello 1981, Reed
1996, Withagen and Chemero 2009). This has been a constant of the ecological
approach, which is apparent in classical contributions like the following:
Ecological theories not only assume that organisms exist in a rich sea of
information about their environments, but also that they evolved in a rich
sea of information. Consequently, it is supposed that the structure and func‑
tion of the perceptual systems have become tailored to the available infor‑
mation. (Michaels and Carello 1981: 15)
Ecological psychologists assume that organisms evolve thanks to ecological
information and affordances, but how exactly do affordances play a role in their
evolutionary history? Here, I aim to reunite key contributions in the literature,
mainly that of Reed (1996), with a picture according to which affordances are
understood as elements of niche construction processes. This reuniting is possible
because ecological psychology and niche construction share the organismal level
of analysis and the idea that organisms are active beings that modify their environ‑
ments. However, my proposal would be incomplete if natural selection, considered
by many as the major evolutionary mechanism, were missing from this picture.
There have been previous attempts to relate niche construction and affordances
(Magnani and Bardone 2010; Withagen and van Wermeskerken 2010; Heras-
Escribano and De Pinedo-García 2018; Heras-Escribano and De Jesus 2018,
Heras-Escribano 2019). This paper intensifies this line of research and proposes
the introduction of natural selection into the picture. The main effort to relate
affordances and natural selection has been offered by Reed (1991, 1996). This
article inherits and expands some of Reed’s main ideas by relating them to niche
construction processes in order to illuminate the role of affordances within evo‑
lutionary processes. In sum, I propose an approach in which affordances acquire
their evolutionary role thanks to two different mechanisms (niche construction and
natural selection). Thus, I aim to show that, in addition to their important role in
our cognitive lives, affordances also play a role in our evolutionary history.
The combination of these two evolutionary mechanisms in a single picture
seems to be problematic for some authors, especially with respect to their differ‑
ent approaches to the relation between organism and environment. Reed claimed
that affordances should be understood as environmental resources exerting
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The evolutionary role of affordances: ecological psychology,… Page 3 of 27 30
selection pressure and that the role of organisms in this picture is to compete
for those resources (Reed 1996: 26–27). Hence, Reed was a selectionist who
claimed that there is an asymmetry between organism and environment in which
the environment is much more relevant for the organism than the organism for
the environment (ibid.). In this view, natural selection is the main driver in the
evolutionary process, and the role of the organism is minimized with respect to
that of the environment. Affordances would be purely environmental resources
that do not depend on animals for their existence, which has been criticized by
authors such as Chemero (2009: 146), who claimed that Reed’s picture fails to
do justice to the mutuality of animal and environment at the basis of the original
Gibsonian proposal (see also Withagen and van Wermeskerken 2010: 495).1 In
line with Chemero’s criticism, authors such as Walsh (2014) claim that the intro‑
duction of affordances into the evolutionary picture necessarily implies a change
in our ways of conceiving evolution and the environment, because an environ‑
ment composed of affordances “is constantly shifting with changes in organismal
form” (Walsh 2014: 223). In particular, compatible with certain views on niche
construction theory2 is the idea that affordances play a key role in the evolution
of the organism because affordances offer a picture in which the organism-envi‑
ronment reciprocity is taken as central (Withagen and van Wermeskerken 2010:
504). Withagen and van Wermeskerken share Walsh’s view on the mutual shifting
of organism and environment in evolution; they claim that Reed’s selectionism is
unsatisfactory regarding the role of affordances in evolution since “the niche con‑
struction perspective suggests alternative roles [to the one offered by Reed] for
affordances in evolutionary dynamics” (Withagen and van Wermeskerken 2010:
497, emphasis added). In this view, affordances would be aspects of niches that
have been ecologically inherited by ancestors that modified the niche (Withagen
and van Wesmerkerken 2010; Heras-Escribano and De Pinedo-García 2018). So,
authors such as Withagen and van Wesmerkerken call for a new, alternative defi‑
nition of the evolutionary role of affordances. This call shows that the definition
of affordances as resources within natural selection processes and the definition
of affordances as aspects within niche construction processes are incompatible
according to some authors.
1
A further analysis on the principles of ecological psychology will be offered in "Ecological psychology
and affordances" section.
2
Walsh proposes an affordance landscape in which the environment is shifting with changes in the
organismal form (a symmetrical view in which both aspects affect each other) as opposed to an envi‑
ronment in which the organism is a passive subject constantly affected by natural selection (an asym‑
metrical view in which only the environment affects the organism). Walsh (2014) also claims that niche
construction endorses a view of the environment that aligns with the latter view rather than with the
former. However, the idea of the affordance landscape that he offers has been proposed to be compatible
with niche construction theory, because niche construction theory holds that “organism and environment
reciprocally affect each other in their mutual development, and those affections are always determined
by the previous ones, which makes every interaction constitutive or formative of the following affec‑
tion” (Heras-Escribano and De Pinedo-García 2018: 12). For a further discussion of the compatibility of
Walsh’s affordance landscape and niche construction theory, see Heras-Escribano and De Pinedo-García
(2018).
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Thus, the role of affordances in evolution is understood in two very different

ways: from Reed’s point of view, which is based on natural selection, they are mere
resources that preexist and exert selection pressure on the animals that take them;
from the point of view of those that support the compatibility of niche construction
and affordances, affordances are ecological inheritances that emphasize the reci‑
procity of organism and environment as a necessary starting point for understanding
evolution, which introduces a certain symmetry into the picture. Supporters of each
view take both to be incompatible. The (apparent) incompatibility of both views
is based on two main aspects: first, the role of the environment; second, the con‑
tributions of the organism in the environment. According to the selectionist view
endorsed by Reed, neither the environment nor affordances depend upon the animal
to exist. This asymmetry between organism and environment, giving priority to the
environment in this case, is key for illustrating the incompatibilities of both views.
Reed summarized his view as follows:
[P]articular instances of an affordance may be realized (literally, made real) by
an animal, as when a seagull drops shells on a rock to split them. But the affor‑
dance is a feature of the environment of all animals (in this case, all the suf‑
ficiently large and hard surfaces available to any gull) and exists independently
of the particular animal even when it is not being used. An affordance is only
a relation when an animal perceives or uses it (…) Affordances in the animal’s
niche are not relations; they are resources (…) (Reed 1996: 26, emphasis in the
original)
Against those who take affordances as relations between environmental aspects
and organismal properties (Chemero 2009), Reed claims that affordances are purely
environmental aspects that enter into a relation as such when they are perceived or
taken. This subtle difference leads to unexpected consequences, because those who
accept that affordances are relations are more inclined to relate affordances to niche
construction rather than to Reed’s view (Withagen and van Wesmerkerken 2010).
The main source of the (apparent) incompatibility between Reed’s view and the
niche construction view is that the latter view, unlike Reed’s, accepts that the organ‑
ism creates the environment. A clear antecedent of niche construction is The Dialec-
tical Biologist (Laland et al. 2016), in which Levins and Lewontin claim that “[t]he
organism influences its own evolution, by being both the object of natural selection
and the creator of the conditions of that selection” (Lewontin 1985: 106, emphasis
added). Reed totally rejected this view and, against Lewontin’s view, claims that “it
is true that birds, beavers, and humans significantly alter the landscape (…), but it
is false that we create the landscape” (Reed 1996: 27, emphasis in the original). In
this sense, Reed’s view seems totally opposed to that of Lewontin, which inspired
niche construction theory. Nevertheless, I think that this dispute is merely semantic
because, despite the use of the word “creation” or “alteration,” Lewontin and Reed
share a conceptual core.
My aim in this paper is to show that this opposition between affordances as inher‑
itances and affordances as resources is merely apparent. For this reason, I propose
that the combination of niche construction and natural selection within a single
picture does not necessarily alter the regular functioning of either mechanism. In
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my view, affordances exert selective pressure at t1, and then these same affordances
are inherited by the offspring at t2. Once these affordances are inherited, they exert
selective pressures regarding the configuration of the niche. I propose to show that
affordances are both resources and ecological inheritances, although they function
as either resources or inheritances depending on the moment of the evolutionary
process in a given niche. This picture allows for the compatibility of both definitions
of affordances (as both resources and ecological inheritances), which are accom‑
modated as two different roles of affordances within a diachronic explanation of the
evolutionary process.
The paper progresses as follows: first, in "Ecological psychology and affor‑
dances" section, I introduce ecological psychology, which is an embodied, situated,
and informational approach to cognition based on the perception of affordances, that
is, the possibilities for action in our environment. Then, in "The extended evolution‑
ary synthesis and niche construction theory" section, I introduce the main aspects
of niche construction theory. Niche construction is a process by which organisms
modify their environments, and some of these modifications facilitate their adap‑
tation. "Affordances, niche construction, and natural selection" section shows how
affordances are compatible with niche construction and natural selection. I propose
a unifying picture of how these key evolutionary mechanisms can be combined, and
I emphasize the importance of affordances in that unified picture so as to reveal that
they fulfill a different role at different stages of the evolutionary process (either as
resources or as ecological inheritances).
Ecological psychology and affordances
Ecological psychology is an approach to perception and perceptual learning pio‑

neered by Gibson and Gibson (Gibson 1979/2015, Gibson and Pick 2000). Ecologi‑
cal psychology was conceived as a third path in psychology beyond behaviorism and
cognitivism. Ecological psychology rejects the cognitivist approach (according to
which cognition is a matter of computing or manipulating representations according
to certain rules) on the basis that cognition is itself a capacity of the organism-as-
a-whole rather than the functioning of inner abstract processes. At the same time,
it also rejects the stimulus–response view of behaviorism since this view takes the
organism as a passive being that merely reacts against certain worldly impinge‑
ments. The starting point of the ecological approach is that, because the organism
is an agent (an active explorer of its environment), we cannot explain how an organ‑
ism perceives and acts if we do not appeal to the history of interactions established
between that organism and its environment. In this sense, ecological psychology is
a clear antecedent of the embodied, situated, and anti-cognitivist approaches to per‑
ception and action that are now gaining momentum in the cognitive sciences (Gib‑
son 1979/2015, Richardson et al. 2008).
The first idea that motivates the ecological approach is that perception is con‑
tinuous with action. This means that we cannot differentiate perception and action
because they are two sides of the same cyclical process: in fact, organisms explore
their surroundings and, thanks to this exploration, perceive aspects that cannot be
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perceived otherwise (Richardson et al. 2008). These aspects, in turn, guide the
behavior of the organisms, allowing them to continue exploring and starting this
process over and over again. In this sense, we perceive because we act, and our
action determines the way in which we perceive. This idea comes from the pragma‑
tist tradition in which the sensory capacities cannot be differentiated from our active
skills (Dewey 1896). Thus, organisms are taken as active agents rather than passive
beings that merely react to stimuli.
The second idea that motivates the ecological approach is situatedness: since per‑
ception and action always occur in a given place and time, the main unit of analysis
according to ecological psychology should not be restricted to the neural system; on
the contrary, the suitable level of analysis should include the organism and its envi‑
ronment, which means that explaining perception is explaining the interaction or
coupling of organism and environment. For this reason, the main unit of analysis is
the organism-environment system (O–E system). It is important to remark that Gib‑
son’s idea of environment differs from the idea of world inasmuch as the environ‑
ment is the surroundings of the organism described in relation to organisms’ capaci‑
ties, whereas the world is a description of the surroundings of the animal described
in terms of particles and forces (Gibson 1979/2015: 4–11). For this reason, the
ecological approach includes a series of terms (“O–E system,” “ecological informa‑
tion,” “specificity,” and “affordance”) that aim to do justice to how the organism
couples with the environment, this coupling being a starting point for a new means
of understanding psychology. When we explain cognition from this perspective, we
are postulating an ecological scale, that is, a methodological framework that works
as the proper level for understanding cognition from an ecological standpoint.
The ecological approach emphasizes that the organism-environment coupling
forms a system, which in turn is the main unit of analysis of psychological pro‑
cesses. But how is this reciprocity achieved? What is the organism seeking in its
exploratory activity, and what does it find to maintain its own activity? According
to ecological psychology, this reciprocity is established through ecological informa-
tion. This kind of information is the way in which we describe the energies of the
environment in relation to the bodies and capacities of the organisms that perceive
them (Heras-Escribano and De Pinedo 2018: 575). A peculiar aspect of this kind
of information is that it allows us to perceive the environment in a direct way. In
this sense, ecological psychology reacts against “indirect” approaches to cognition
like cognitivism (Gibson 1979/2015). Cognitivism states that the stimulation of sen‑
sory receptors is sufficient for achieving perception, while the ecological approach
claims otherwise. Imagine a human inside a densely fog-filled room. In this exam‑
ple, light stimulates that person’s retina, but she would be unable to perceive her
surroundings. The organism cannot perceive the surroundings of the room because
light does not reverberate; hence, it does not offer any kind of information about the
available affordances (Gibson 1979/2015: 46, Chemero 2009: 107–8). Once the fog
is removed, light can reflect and reverberate on the surfaces, thus gaining its struc‑
ture and finally showing the human the possibilities for acting (Glotzbach and Heft
1982: 111). In this case, light is informative. This is how ecological information is
formed. The person in the room detects this information because she is an agent,
that is, because she acts in order to discover the different affordances available in
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the room. As she explores her environment, she detects or picks up the ecological
variables that provide information about the room’s affordances, which amounts to
saying she discovers what she can and can’t do. Unlike cognitivism, there is no need
to postulate inner representations: behavior can be accounted for solely in terms of
the organism-environment coupling. There is no need to postulate representations
of the outer world to make sense of perception because ecological information is
a description of how the outer world is related to our capacities; it shows how the
distribution of energy arrays is disposed in relation to the agent’s movement: “[t]he
optical disturbances created by an approaching car, for example, do not resemble the
car; rather they uniquely specify it and its path of locomotion in relation to oneself”
(Gibson and Pick 2000: 18). The expression “in relation to oneself” is key for stress‑
ing that we do not represent the world as a static picture: rather, we perceive the
world from a concrete spatiotemporal location and always in relation to our bodily
constitution. For all these reasons, according to the ecological approach, “percep‑
tion is a direct—noninferential, noncomputational—process, in which information
is gathered or picked up in active exploration of the environment” (Chemero 2009:
106; see also Michaels and Carello 1981).
A typical example of an ecologically-informational variable is tau or time-to-
contact (Lee 2009). This variable consists in “the ratio of the optical size [of the
approaching object] to the rate of optical expansion [of the same object during
time]” (Jacobs and Michaels 2007: 324); in other words, when something (an object,
an organism) is approaching you, that object or organism progressively expands in
your visual field. The relation between the expansion and the approximation gives
information about the time-to-contact between the perceiver and the object or organ‑
ism that is perceived. This example shows how the environment consists in the sur‑
roundings as they relate to the organism, since time-to-contact is an agent-related
metric that helps the organism anticipate when collision will happen. This is not
expressed in neutral, agent-unrelated metrics like meters per second. Agency is key
in this picture, and this implies that the metrics of what happens in the environ‑
ment relate to the action and position of the agent-perceiver. Needless to say, time-
to-contact suffices for guiding behavior, since the optical expansion in each moment
reveals the available affordances for the perceiver: avoidance, collision, climbing,
vaulting, crawling, grasping, etc.
As we can see, ecological information shows or reveals the available affordances.
In the terminology of ecological psychology, this means that ecological information
specifies the available possibilities for each agent’s actions. Specificity is a term that
refers to the one-to-one correspondence of ecological information and affordance
perception: the detection of certain ecological variable corresponds to the perception
of affordances and vice versa (Chemero 2009: 111). As Käufer and Chemero (2015)
claim, specificity works in such a way that the presence of ecological information
guarantees the presence of affordances. This concept is the bedrock upon which the
scientific character of ecological psychology is built, because it establishes a law‑
ful correspondence and regularity between ecological information and affordances
(Jacobs and Michaels 2002: 129, Richardson et al. 2008: 177). This lawful regular‑
ity is understood in a very particular way within the ecological approach: “Ecologi‑
cal information is lawful not in the Newtonian sense of being universal in space and
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time, but in an ecological sense of being regular within an ecological context of

constraint” (Warren 2005: 342–343). This lawful regularity shows why ecological
information such as time-to-contact specifies or guarantees the presence of certain
affordances, and these affordances change depending on the spatiotemporal coordi‑
nates in which they are perceived: for example, given a particular rate of expansion
in our visual field we can perceive not only the possibility of avoidance but also the
possibility of grasping or collision depending on the moment and the circumstances.
Now we can fully understand the nature of the main objects of perception for eco‑
logical psychology: affordances.
Affordances are possibilities for action that are available in the environment for a
particular kind of organism. Affordances are, then, aspects of the environment that
are considered in relation to the bodily constitution and capacities of the agents that
perceive them. The perception of the available opportunities for action is dependent
on the detection or picking-up of ecological information: objects are graspable for
agents with posable thumbs, steps are climbable for those who can raise their legs
at a certain length, etc. These objects of perception are quite intuitive, therefore,
and we are constantly surrounded by affordances in our daily lives. Nevertheless,
their philosophical status is still in dispute, and this is because they are challeng‑
ing for traditional philosophical categories. On the one hand, affordances are not
mind-independent aspects of the environment; on the other hand, they are not purely
subjective aspects. They escape the objective-subjective dichotomy because they are
environmental aspects relative to the bodily constitution and abilities of their per‑
ceivers (Gibson 1979/2015: 120–1). The coupling or reciprocity of organism and
environment is challenging for standard dichotomies in philosophy, and this is why
affordances and the ecological standpoint that takes the O–E system as a main unit
of analysis are innovative conceptual frameworks that allow us to understand cogni‑
tion form an innovative perspective.
This coupling between organism and environment is the key aspect that connects
ecological psychology and niche construction.3 As we have seen, affordances play
the role of illustrating the reciprocity of organism and environment at the cognitive
level, but the ecological study of affordances can also play the role of illuminating
how we can understand evolution better under the principles of niche construction.
Affordances are aspects of the environment related to bodily features or traits of the
organism that the organism encounters when exploring the environment; these affor‑
dances exert a feedback function to the organism that guides its behavior, and this
function can be described in dynamical systems. This influence of environmental
information working as a feedback mechanism for exploring the environment is key
for our developmental processes and perceptual learning, as it has been proved with
empirical evidence by ecological psychology (Gibson and Pick 2000; Jacobs and
Michaels 2007). This is the very idea that underlies niche construction: modifica‑
tions in the environment lead to an alteration in the evolutionary dynamics of the
population that cause it. In this sense, the dynamics established by the interaction
with the affordances of the environment can exert an influence in the evolutionary
3
I thank an anonymous reviewer for urging me to comment upon this idea.
13
dynamics as well: if the processes of niche construction lead to a certain modifica‑

tion of the environment and the creation of new affordances, these affordances play
a role in both the cognitive and the evolutionary history of the population that will
benefit from perceiving and using them. Thus, this well-established psychological
study of perceptual learning can serve as an illustration to emphasize the importance
of these feedback relations between organism and environment. For these reasons,
affordances are an example of how important niche construction processes are for
making sense of evolution beyond the typical examples at a genetic level. In order to
understand this in more detail, we have to describe the principles of niche construc‑
tion and how it fits within the extended evolutionary synthesis.
The extended evolutionary synthesis and niche construction theory
Niche construction theory is often presented as one of the main theories that con‑
forms to what has been called the extended evolutionary synthesis (EES), a view
that challenges the traditional approach to evolution, also known as the Modern
Synthesis.
The Modern Synthesis (MS) is considered the mainstream approach in evolu‑
tion; it consists in the combination of Darwinian natural selection with Mendelian
genetics (Huxley 1942). The viewpoint of the MS focuses on the role of genes and
genetic inheritance, and this insistence on genetic inheritance as a cornerstone of
evolution has often been called the gene’s view or the gene-centered view (Fisher
1930; Williams 1966; Maynard Smith 1998; Dawkins 1976). According to this
view, the explanation of evolutionary change and adaptation relies on the role and
functioning of genes.4 From this perspective, adaptation is taken as a process guided
by natural selection, which is the only (external) force that drives evolution by act‑
ing like a filter by which populations with the suitable genetic pool fit their par‑
ticular environmental circumstances. Random genetic mutations of some organisms
are essential for increasing their chance to reproduce against hostile environmental
conditions and, if successful, those organisms transmit their genes to their offspring
(Maynard Smith 1998: 10). In this sense, only genes transmit the essential elements
that provide the adaptation to the environment. The main reason to choose genes
instead of other aspects as the main units, or to choose the organisms’ genotype (the
unobservable genetic aspects that determine the observable features) rather than the
organisms’ phenotype (the observable features or traits), is that phenotypes, unlike
genotypes, are extremely contingent and temporary combinations of how genes are
expressed and how the environment is disposed (Williams 1966: 22–23). This allows
for dramatic variations in short periods of time that do now allow one to quantify
4
The gene-centered view is understood here as centered on individual genes as opposed to genomes
or whole organism phenotypes. This view is compatible with Dawkins’s view of organisms as survival
machines of genes, as well as with orthodox views in the Modern Synthesis cited above. To show the
intricacies of the idea and its relation to phenotypic variation, I offer a comparison of niche construction
within the Extended Synthesis and Dawkins’s extended phenotype within the Modern Synthesis at the
end of this section. I thank an anonymous reviewer for urging me to clarify this idea here.
13
and analyze the real impact of adaptation and evolution. This is an asymmetric pro‑
cess in which an external force acts upon a particular population and in which we
know whether or not adaptation is achieved by merely looking at the genetic level.
This is because “the environment is largely autonomous with respect to the organ‑
isms,” and for this reason it works “according to its own intrinsic dynamics” (God‑
frey-Smith 2001: 254). There is no coupling or significant mutual affection between
organism and environment. The environment exerts selection pressure and does so
basically via natural selection.
A this point we can see how the MS usually adopts a gene-centered view, and this
general picture offers three main consequences regarding evolution and adaptation:
first, the main (and maybe only) driving force in evolution is natural selection; sec‑
ond, genetic inheritance is the main (and maybe only) kind of inheritance that mat‑
ters for evolution and adaptation; third, organisms are receptacles in which genes are
transmitted generation by generation. This latter idea leads Dawkins (1976) to define
organisms as “survival machines” of genes.
Despite the influence of the MS and the gene-centered view, a new and emerg‑
ing approach to evolution has been gaining momentum in recent years: the extended
evolutionary synthesis (EES). It is a new view that aims to expand the claims of
the MS by incorporating different evolutionary mechanisms neglected or not suf‑
ficiently emphasized in the previous years. In particular, the EES rescues the impor‑
tance of developmental processes for evolution, as well as the reciprocal causation
and co-determination of organism and environment (Oyama 1984; Lewontin 2000;
Jablonka and Lamb 2014; Pigliucci and Muller 2010; Laland et al. 2015). Propo‑
nents of the EES argue against a restrictive view of the MS by claiming that there
are means of inheriting beyond the genetic level, and they question the primacy of
natural selection as the sole force driving evolution. In this sense, the EES aims to
gather some processes, phenomena, and mechanisms beyond natural selection (such
as developmental processes, epigenetics, niche construction, or ecological inherit‑
ances) so as to enrich and expand the main ideas of the MS. In this sense, these
often neglected aspects or mechanisms play the role of a bias in the drift of selec‑
tion. According to the picture offered by the EES, natural selection still works as a
major force in evolution, but it is not the only one. In the same vein, new forms of
inheritance beyond genetic inheritance are also considered relevant for making sense
of evolutionary drift.
According to these authors, a key example of a neglected evolutionary mechanism
is niche construction (Odling-Smee et al. 2003). Niche construction is an evolution‑
ary mechanism by which organisms alter or modify their environments in a way that
is beneficial to them and their offspring since those alterations increase their chance
of reproduction and survival (Lewontin 1985; Odling-Smee et al. 2003; Laland et al.
2015). Niche construction analyzes the ways in which the environment is shaped by
the organisms that inhabit it. In this sense, changes in the environment can be divided
in two main kinds: first, inceptive perturbations, that is, the changes in the organism’s
behavior that physically alter environmental aspects (such as the emission of detritus,
13
dairy farming, or the construction of dams5); second, inceptive relocations or the

migrations of organisms to new habitats in which those organisms encounter new
environmental aspects or variables (Odling-Smee et al. 2003: 47). In these inceptive
perturbations and relocations, the organisms are the drivers of environmental change,
and they act in an exploratory way, but there are also counteractive approaches to
perturbations and relocations: counteractive perturbations are those actions in which
organisms respond to or compensate a drastic environmental change by modifying
their environments (like the thermo-regulation of bird nests against a drastic change
in temperature), and counteractive relocations provoke migrations to other places due
to environmental changes (like the seasonal migrations of some birds). In sum, organ‑
isms find certain cues that guide their exploration and allow them to expand their
behavioral repertoire in order to survive, reproduce, and thrive, compensating distur‑
bances in the environment and finding new habitats with resources that will benefit
them. They seek these cues in order to succeed in their goals. For this reason, accord‑
ing to both defenders and opponents of this view, “niche construction is guided by
(genetic or acquired) information” (Scott-Phillips et al. 2014: 1234).
From the point of view of niche construction, inceptive perturbations play the
role of a bias in the direction of selection, which means that apparently innocuous
alterations of the environment might generate a changing output on a larger scale
that modifies the spatiotemporal dynamics of the niche. If we analyze some of the
examples provided above, dams alter the environment not only in a way that is ben‑
eficial for their builders (humans, beavers); they also affect different aspects of the
environment that were not taken into account by their builders. For example, dras‑
tic changes in the current of rivers may help the reproduction of microorganisms
that could not reproduce otherwise, leading to new diseases or parasitic relations
between species in that habitat. In the same vein, stopping the current of a river with
a dam might create a habitat suitable for birds to stop during their migration process,
and this could also attract certain predators. In this sense, altering the environment
not only serves the purpose of benefiting the organisms that make those alterations;
they also trigger unexpected changes in the dynamics of the niche. Hence, if the
activity of the organisms produces alterations in habitats that benefit their fitness
and that also provoke unexpected changes in their niche that bias their evolutionary
drift, they are not simply receptacles of genes subject to changes in their habitats.
There are physical consequences that are produced by these alterations. And
sometimes those physical consequences are objects that become part of the habi‑
tat, like the previously mentioned example of the dam. When these objects are
5
It should be highlighted that not all alterations in the environment produced by the organism are goal-
directed and adaptive. Dairy farming in humans is clearly a goal-directed action, but the emission of
detritus in populations of squirrels is not necessarily a goal-directed action. Despite this difference, the
alterations of both inceptive perturbations can produce an equally significant alteration in their particular
ecosystems. The same goes for the adaptive role: not all such alterations are beneficial in adaptive terms,
although they equally change the evolutionary dynamics even without an adaptive impact. In this sense, I
focus here on affordance perception and affordance taking as an example of goal-directed action, but not
all aspects of niche construction processes involve this kind of aspect. I thank an anonymous reviewer for
inviting me to highlight this point.
13
inherited by the offspring of those who altered their environment, they are catego‑
rized as ecological inheritances. Ecological inheritances are defined as the physical
or material consequences of niche construction processes (Odling-Smee et al. 2003).
There are multiple kinds of ecological inheritances that range from cultural practices
like dairy farming to artifacts, buildings, tools, etc. Biological inheritances expand
beyond genetic inheritance, since those material consequences determine the devel‑
opment of organisms and their habitats. This counters the claim that genetic inherit‑
ance is the only kind of biological inheritance. Within the EES there are non-genetic
mechanisms that contribute to inheritance, such as parent–offspring transmission
of objects that alter the environment in a way that benefits their fitness. Ecologi‑
cal inheritances also alter the genetic expression of phenotypes in the development
of organisms (Donohue 2005). In this sense, the material consequences of previous
alterations of the environment are inherited, which means the benefit obtained by
the alteration is transmitted and modifies the evolutionary history of populations in
ways that were unexpected before the alteration.
In sum, the EES expands the main ideas and claims of the MS regarding evolu‑
tion and adaptation, offering a richer view of the phenomena that take into account
processes and mechanisms usually neglected. At the same time, the introduction of
these mechanisms and processes supposes a challenge to the gene-centered view
that includes certain theoretical changes in our way of understanding adaptation,
evolution, and biology in general.
The main theoretical consequences of MS and the gene-centered view are the fol‑
lowing: organisms are receptacles of genes, and the only force that drives evolution
is natural selection, which means the only kind of inheritance is genetic inheritance.
At the same time, defenders of niche construction claim that organism-environment
coupling establishes certain dynamics that bias the direction of evolution. These
alterations favor their survival and reproduction, and the material consequences of
those changes are inherited by their offspring. These material consequences show
that there is inheritance beyond genetic inheritance. All these aspects in combina‑
tion reveal that adaptation is not conceived within the EES as an asymmetric process
by which biological features evolve in response to environmental selective pressures.
In this view, adaptation is partially constituted by the (voluntary or involuntary)
activity of organisms, because they bias their evolutionary drift at the same time that
they are affected by other selective mechanisms. This mutual affection of organism
and environment emphasizes the importance of the organismal level for explaining
the causal role of these kinds of process, moving from the asymmetrical picture of
the MS to a symmetrical picture (Walsh 2014: 216).
One might think that the MS’s emphasis on the role of genes leaves aside the
importance of organisms’ active capacities and that, in turn, these capacities are
taken into account within the EES view (particularly in niche construction). Never‑
theless, a more cautious treatment of how the MS understands the activity of organ‑
isms reveals that this picture is not accurate. In particular, a supporter of the MS
view would claim that gene distributions only have selective consequences through
the activity of organisms. In this sense, the role of organismal activity is rather
essential in the traditional view. Thus, the differences between both views do not
rely on the active powers of organisms but, rather, on other aspects.
13
An example is the significantly different ways in which the MS and the EES
understand development and the conceptual relation among variation, differential
fitness, and heredity (Uller and Helanterä 2019). Variation, differential fitness, and
heredity are taken as autonomous concepts and quasi-independent processes within
the MS (Walsh 2015). Variation (random mutation) is explained independently.
At the same time, the variable rates of survival among individuals determine the
features that will be present in the offspring. Nevertheless, these processes do not
directly affect the process of inheritance, which is explained by independent Men‑
delian rules. Each element provides the materials for the next step, but every step
is explained by appealing to its own dynamics or rules (Uller and Helanterä 2019).
It is important to note that development has a reduced role in this view, because the
only evolutionary causes related to development are those that pass through the fil‑
ter of natural selection and are inherited (Walsh 2015). Development, then, is sepa‑
rate from the mechanisms that explain the content transmitted trans-generationally
(the genes), and it only affects the environmental context in which the content is
expressed (Uller and Helanterä 2019). However, there are authors who claim that
this quasi-independence should not be taken literally, since the developmental pro‑
cesses that produce recurrent phenotypic expressions are closely related to processes
that produce recurrent selection insofar as the way organisms respond to certain
disturbances during their ontogenetic development shapes their environment and
their adaptation to it; furthermore, these responses will affect the way in which their
offspring will treat the already modified environment, favoring survival and repro‑
duction (Uller and Helanterä 2019; Sultan 2015). In this sense, the EES does not
take biological processes as quasi-independent. The emphasis on development and
the intertwinement of development and fitness is another clear difference between
the MS and the EES, which aggravates the differences between both syntheses and
helps to offer a different picture of the role of the organismal level in adaptation.
These differences establish different views regarding the status of certain evo‑
lutionary processes in evolutionary biology, a discussion carried out between sup‑
porters of niche construction theory and defenders of the MS. These discussions
emphasize the role of niche construction in evolution. There are biologists who
claim that the MS can accommodate the main contributions of niche construction,
mainly because they think that phenomena such as niche construction do not neces‑
sitate rethinking evolutionary theory as it is conceived within the MS (Laland et al.
2014). Niche construction processes in this view, according to Laland et al. (2014),
are reduced to mere feedback relations between organism and environment, and
these feedback relations, although they play a part in evolution by altering it, are not
essential for it (the only essential aspects are—according to these authors—natural
selection, drift, mutation, recombination, and gene flow). Some authors might claim
that niche construction resembles well-established ideas in the literature that do not
need to appeal to mechanisms beyond natural selection, such as Dawkins’s extended
phenotype. Although overlaps exist between the idea of extended phenotype and
niche construction, there are differences: first, the extended phenotype only focuses
on selective feedback from environmental modification to the genes responsible for
those modifications, but niche construction takes into account other environmental
aspects that are potentially unrelated to those genes; second, unlike the extended
13
phenotype, niche construction theory emphasizes the legacy of modifications that

remain beyond the modifiers as inheritances; third, again unlike the extended phe‑
notype, niche construction does not require traits to have a strong genetic basis or a
strong association with fitness since niche construction processes include or acquire
characters that generate selective feedback even without being extended phenotypes
(Laland et al. 2016). Thus, the idea of extended phenotype is a narrower concept
compared to niche construction, which includes phenomena that go beyond well-
established ideas in the MS and that play a relevant role in the alteration of evolu‑
tionary dynamics. Niche construction theorists are among the advocates of the EES
who claim that MS is insufficient and should be enriched, but the relation between
the EES and the MS varies depending on each author. For example, while some
authors claim that evolutionary theory should be expanded (Kauffman 1993) or
urgently rethought (Laland et al. 2014) on the basis of new evidence so as to create
a new synthesis, other authors think that the MS should be replaced because it no
longer offers a satisfactory framework for evolutionary biology (Jablonka and Lamb
2008). The proposal offered here aligns with the views of authors such as Odling-
Smee et al. (2003), who emphasize the role of certain environmental aspects in
altering the evolutionary dynamics of a particular population, and I hold that affor‑
dances contribute significantly to these alterations.
Affordances, niche construction, and natural selection
Affordances and niche construction
There are various similarities and complementarities between niche construction

and ecological psychology. In particular, the ideas of agency and information are
key in both approaches. Furthermore, the way both concepts are depicted in each
theory shows that there is room for a certain compatibility inasmuch as we accept
that cognitive or psychological processes play a certain role in biological aspects
such as development and adaptation.
First of all, both theories emphasize the role of organismal activity. As we have
seen, although some examples of NC involve active “psychological” agents behav‑
ing in ways that are aimed at modifying the environment, other examples imply that
the modifications on the environment are fortuitous. Thus, many cases of niche con‑
struction do not involve any behavior or agency at all (e.g., chemical changes in the
soil produced by bacterial detritus). Given that I aim to reconcile niche construction
and ecological psychology, and given that ecological psychology implies volition
and agency, the niche construction processes to which I am appealing that can be
explained by the principles of ecological psychology and ecological information are
very few. Nevertheless, it could be claimed that, in these specific cases, organisms
are active explorers and modifiers of their habitats and that some of those changes
are produced in their favor, increasing their chances of survival and reproductive
success. The offspring of that population will find those previous alterations in their
environment from their very beginning, increasing as well their chances of survival
and reproduction. In this sense, the action of organisms is essential for their survival
13
and fitness from a niche construction perspective, since their capacity for altering
the environment has a direct effect on these aspects. By contrast, organisms are con‑
sidered active explorers within the ecological approach because the organism acts in
order to perceive the available affordances of the environment. Action is so crucial
for the ecological approach that Gibson’s main book claims that motion or action is
the criterion for dividing the cognitive from the non-cognitive, the animate from the
inanimate (Gibson 1979/2015: 3). Thus, niche construction and ecological psychol‑
ogy focus on the idea of organism as an active agent that explores the environment
in search of opportunities for action (affordances) that play the role of resources for
adapting to the environment.
Another source for similarities between the ecological approach and NC is
the emphasis on information. This is what guides niche construction and percep‑
tion–action processes. As argued by both niche construction theorists and their
opponents, “niche construction is guided by (genetic or acquired) information”
(Scott-Phillips et al. 2014: 1234). Leaving aside genetic information, if we ana‑
lyze the kind of information we can find at the organismal level, it seems that niche
construction endorses the idea that organismal behavior is guided by some kind of
acquired information that results from the coupling of organism and environment,
which is sometimes produced by the organism’s alteration of its own habitat. In
the same sense, ecological psychology claims that the organism-environment cou‑
pling is achieved thanks to ecological information, which is formed by ecological
or higher-order variables that result from the interaction between energy arrays and
the action of agents. For this reason, some authors point to the idea that ecological
information can be conceived as one kind of acquired information that guides niche
construction processes, proposing a continuity of cognitive and evolutionary pro‑
cesses from an organismal and informational perspective (Heras-Escribano and De
Pinedo-García 2018; Heras-Escribano and De Jesus 2018).
The final aspect shared by both the extended synthesis and ecological psychology
is the emphasis on development or the ontogenetic history of organisms adapting to
their environments. As already shown, developmental processes play a crucial role
within the EES since the experience of the organism during its own development is
crucial for compensating for the disturbances of the environment. In this sense, the
responses developed by the organism to facilitate its own adaptation to the environ‑
ment and survival will have a direct impact on its offspring, because the offspring
will enter into an environment or habitat that has been already modified for increas‑
ing the probability of its own survival and adaptation (Uller and Helanterä, 2019;
Sultan, 2015). One of the major branches of the theory since its birth, development
is also very important within ecological psychology. E. J. Gibson focused on the
importance of ontogenetic processes and perceptual learning, thereby developing a
line of research parallel to that of J. J. Gibson focused on perception (Gibson 1969;
Gibson and Pick 2000; Rader 2018; Read and Szokolszky 2018). The main innova‑
tion of this approach is Jacobs and Michaels’s (2007) direct learning theory, which
proves that participants evolve from novice to expert performers of a certain task
because they tend to pick up the specific ecological variables over the non-specific
ones. The ecological approach to perceptual learning shows how organisms make
use of specific ecological information available in their environments: organisms
13
attune to their environments when they seek the most convenient information in
order to perform a task, and for this reason they need to calibrate or adjust their
behavior to the already-found informational variable (Lobo et al. 2018a).
These similarities show that there are sufficient conceptual compatibilities
between niche construction and ecological psychology so as to consider affordances
as aspects in the niche construction process6 (at least as aspects that play a role for
niche construction in cases when niche construction implies active agency). As a
result, the material consequences of niche construction processes are not restricted
to the physical objects that are inherited: the social function and the affordances of
those objects are also inherited (Heras-Escribano and De Pinedo-García 2018). For
example, any kind of device that we use in our society (a pair of glasses, typewrit‑
ers, keys, chairs, etc.) are inherited as physical objects, but we inherit them along
with the ways in which we should deal with them. These ways of dealing with the
objects are twofold: first of all, there is the way in which we deal with them from an
individual point of view, that is, as an individual organism dealing with the environ‑
ment and aiming to satisfy its purposes and needs by perceiving the affordances of
objects; secondly, we deal with the environment as social beings, that is, as members
of a community of shared practices in which we are instructed and trained, and as
social beings we aim to follow the norms and behavioral patterns because they help
us modulate how we take the affordances of objects in different ways7(Heras-Escrib‑
ano and De Pinedo 2018: 486–587). In this sense, our niche contains not only the
physical structures of those objects as such but also their affordances (which is the
way in which those objects relate to our bodily skills and capacities) and the behav‑
ioral patterns that are socially enacted and serve to deal with those objects in an
organized manner benefitting the community. This implies that human nature at an
organismal level establishes a double dialectics: the reciprocal interplay of organism
and environment on the one hand and, on the other, that of the individual and the
community of sociocultural practices (Reed 1993, 1996; Heft 2007, 2018; Ingold
2001/2011).
6
As suggested by an anonymous reviewer, it is worth highlighting that niche construction theory applies
to all organisms, including those that are not traditionally conceived as perceivers (though they do detect
things), behavers (in most senses of the term), or fast (animated) movers. This is important to avoid
equating the words “organism” and “cognizer” and giving the false impression that niche construction
processes are only triggered by cognitive organisms. In the case of chordates, action capacities are well-
developed so as to act in a fast way and develop a certain plasticity and flexibility that increase their
cognitive capacities (Settleworth 2010), and niche construction processes are triggered thanks to these
capacities. In this sense, organisms that do not possess these capacities are also capable of producing
alterations in their environments, which trigger niche construction processes (Odling-Smee et al. 2003).
Nevertheless, there are also organisms that are usually categorized as non-cognitive but, as recent empiri‑
cal studies prove, show cognitive capacities in ecological terms, as happens in the field of plant cogni‑
tion, which is gaining momentum in the cognitive sciences (see, for example, Heras-Escribano and Calvo
2019).
7
Based on how it is written, it might seem that a greater capacity for social sharing of information
is always advantageous. However, sometimes it is not, since such capacities also carry costs that vary
according to the ecological situation of the organism. I thank an anonymous reviewer for asking me to
point this idea out.
13
The affordances of our human environment are aspects of our environment

shaped by our activity, but they are also at the same time environmental resources
that modulate our activity and, in addition, exert selection pressure on the behav‑
ior of organisms (Reed 1996: 18). For this reason, the affordances that exert selec‑
tion pressure and that remain as ecological inheritances in our niches (because they
increase our and our offspring’s chances of survival and reproduction) contribute to
the systematic bias exerted by our alterations, thereby conditioning our evolutionary
drift in a direction that could not have been achieved otherwise. This is because the
behavioral patterns acquired by social interaction teach us how to deal with affor‑
dances in a way that is beneficial for our adaptation and that of our offspring.
Some authors could claim that affordances do not offer substantial contributions
to the explanation of evolution in terms of niche construction.8 As shown earlier,
supporters of niche construction do not need the concept of affordance to make their
central point, although this per se does not preclude affordance from entering into
the picture of their central point. Yet, can affordances make a genuine contribution
to niche construction? As we have seen, genetic or acquired information guides
niche construction processes (Scott-Phillips et al. 2014). If so, then ecological infor‑
mation is one of the kinds of acquired information that help us guide niche con‑
struction processes in the realm of perception and action (Heras-Escribano and De
Pinedo-García 2018; Heras-Escribano and De Jesus 2018). According to the eco‑
logical approach, the direct perception of affordances amounts to the detection of
ecological information, and perceiving affordances helps us navigate the environ‑
ment. This idea is canonical and lies at the very core of the ecological approach
(Gibson 1979; Michaels and Carello 1981; Turvey et al. 1981; Reed 1996; Chemero
2009; Heft 2001). In this sense, ecological information is a special kind of infor‑
mation of which we are aware in our navigation of the environment; this informa‑
tion is useful for us since it guides and constraints our range of behaviors when we
explore our surroundings. We can also modify the environment and alter the avail‑
able affordances while retaining others as we please. In this sense, perceiving and
taking affordances via the detection of ecological information can be taken as a key
process in our cognitive lives, since we are constantly detecting ecological informa‑
tion that allows us to explore the environment in an efficient way. The acceptance of
ecological information as a kind of acquired information that plays a decisive role in
niche construction processes implies the acceptance of affordances as a key aspect
not only of our cognitive lives but also of our evolutionary process.
In conclusion, it can be argued that affordances and niche construction share the
same organismal level of analysis; they share the ideas that organisms are agents,
that development is crucial for perceptual learning and adaptation, and that our deal‑
ings with the environment are guided by information of all kinds, including eco‑
logical information for affordances. In this sense, the ecological approach to percep‑
tion and action can be understood as an approach that shares the main commitments
and premises of niche construction. This shows that there are cognitive processes of
8
Thanks to an anonymous reviewer for making me aware of this idea.
13
affordance perception and taking that can be inserted within such evolutionary pro‑
cesses as niche construction located at an organismal level.
At the end of "Ecological psychology and affordances" section, I emphasized the
importance of ecological psychology for niche construction theory, and some words
must be now added to emphasize the importance of niche construction to ecological
psychology.9 As shown in the introductory section, most ecological psychologists
agree that affordances are tightly related to our evolutionary processes, although few
systematic proposals have been offered to date. Niche construction can illuminate
the evolutionary impact of affordances and also various other aspects within the eco‑
logical approach like effectivities. This is because niche construction can explain
how affordances can modify phenotypes: according to the Connecticut School in
ecological psychology (Lobo et al. 2018b), the complementary aspects of affor‑
dances are animals’ effectivities, that is, properties of the organism related to affor‑
dances. Although a well-established aspect in this tradition, not much has been said
about effectivities and its evolutionary role, but niche construction can illuminate
how effectivities can change and how they can serve to create new affordances in
the environment. This is because, if we understand affordances as key environmental
aspects within niche construction processes, there is a tradeoff between phenotypic
plasticity and niche construction in which the environment induces changes in the
phenotype and, at the same time, these changes in the phenotype may involve adap‑
tive specializations that might in turn lead to changes in the environment, which is
why plasticity is a major source of niche construction that could trigger changes
within species (Laland et al. 2016: 196). For this reason, understanding affordances
from a niche construction perspective and trying to combine niche construction and
ecological psychology are quite helpful gestures for understanding not only the evo‑
lutionary role of affordances but also different aspects of the ecological approach
(such as effectivities) from an evolutionary perspective.
Introducing natural selection
The previous subsection shows that cognitive processes of perceiving and taking
affordances are compatible with a general scheme of evolution as portrayed by niche
construction processes. In this subsection, I aim to integrate natural selection—the
most influential evolutionary mechanism to date—into the picture offered above.
As a consequence, I offer here a rich view relating two evolutionary mechanisms
to affordances and highlighting the evolutionary role of these objects of perception.
Natural selection is an evolutionary mechanism that causally connects heritable
variation to differential ability to survive and reproduce in a given population. Spon‑
taneous mutations of genes and their phenotypic expression can be either benefi‑
cial or punitive depending on the external circumstances. If the organisms with that
genetic material and its correspondently developed phenotypic expression survive
9
Thanks to an anonymous reviewer for urging me to comment this idea.
13
the external circumstances, then they will pass their genes to their offspring. This
means that those individual differences are beneficial for survival and, hence,
preserved and transmitted to future generations. Those traits and genes are then
selected, and the organisms that possess them are said to be adapted to the environ‑
ment. As Darwin wrote, “[t]his preservation of favorable individual differences and
variations, and the destruction of those which are injurious, I have called Natural
Selection, or the Survival of the Fittest” (Darwin 1876/2009: 63). In this sense, nat‑
ural selection is the combination of three different principles that correspond to the
three quasi-autonomous processes presented in "Ecological psychology and affor‑
dances" section above. Each process produces the materials that are available for the
functioning of the rest of the processes. They are summarized as follows:
1. There is variation in morphological, physiological, and behavioral traits among

members of a species (the principle of variation).
2. The variation is in part heritable, so that individuals resemble their relations more
than they resemble unrelated individuals and, in particular, offspring resemble
their parents (the principle of heredity).
3. Different variants leave different numbers of offspring either in immediate or
remote generations (the principle of differential fitness).
(…) All three conditions are necessary as well as sufficient conditions for
evolution by natural selection (Lewontin 1985: 70).
As such, natural selection is a mechanism that works at a population level since

evolution and adaptation focuses on which genes are successfully transmitted to the
next generation and thereby preserve their presence in a given population, as seen
in "Ecological psychology and affordances" section above. The distinction between
an organismal and a sub-organismal level with their own evolutionary mechanisms
implies that niche construction and natural selection are not incompatible processes.
In this sense, certain genes that allow for developing particular behavioral strate‑
gies for detecting ecological information can be inherited, provoking certain differ‑
ential fitness in the offspring. This is the way in which natural selection is related to
ecological information and affordances as environmental resources that exert selec‑
tive pressure, and this is the way in which Reed’s approach (1991, 1996) actualizes
Michaels and Carello’s claim according to which “it is supposed that the structure
and function of the perceptual systems have become tailored to the available infor‑
mation” (Michaels and Carello 1981: 15).
Reed (1991, 1996) proposed a means to relate the ecological approach and evo‑
lution by natural selection by emphasizing the role of ecological information. As
Reed claimed, organisms that are differentially sensitive to ecological information
13
compete in an environment where that information is a relevant resource10 that

improves the possibilities of survival and reproduction (Reed 1991: 193). In this
sense, as Reed claims, all forms of behavior are modes of competition for the use
of some resource and have therefore evolved thanks to natural selection (Reed 1985:
359). Thus, the behavioral, physiological, and genetic variation allows for a certain
degree of sensitivity to ecological information, and this variability is what allows
for competition among organisms. These informational resources work in the same
sense as other resources in nature: Organisms with greater sensitivity to that kind
of information outcompete those with lesser sensitivity. This is why organisms that
develop more efficient behaviors for exploiting those resources are more likely to
survive and reproduce than those that do not. This greater sensitivity is partially
achieved by the organism thanks to different environmental influences during its
development. And this is when a niche construction process enters the scene.
Unifying natural selection, niche construction, and affordances in a single picture
In the picture I offer, there are two evolutionary mechanisms working at the same
time. First of all, processes at the sub-organismal level such as spontaneous muta‑
tions in the genotype lead to certain (morphological, physiological, and behavioral)
variation, which is expressed in the phenotype. This process goes in a bottom-up
direction, as the expression of the genotype is also affected by developmental pro‑
cesses and the configuration of the (ecological and social) environment at the same
time. If those traits are beneficial for survival (that is, if those traits can deal with
the available affordances of the environment), they are transmitted to the following
generation, and this is why natural selection preserves that genetic pool. Meanwhile,
organisms alter their environments in beneficial ways through niche construction
processes. As such, the offspring of those organisms will benefit from those envi‑
ronmental alterations, as we have seen in "Ecological psychology and affordances"
section. Thus, we have two mechanisms working in parallel.
At this point a pertinent question arises: are these two mechanisms totally iso‑
lated from each other? I do not think so. In the case of ecological information, the
“greater sensitivity” to which Reed refers may be either the product of a spontaneous
10
An anonymous reviewer urges me to explain Reed’s emphasis on the idea of resources in his approach
to affordances. Reed defines affordances as resources for behavior in his book Encountering the World.
The idea of resource plays a key role in the definition of affordance, and it is defined as an environmental
aspect that exerts selective pressure and that gives an evolutionary advantage to the organisms that take
them. The author does not refer to a particular tradition or author in the literature on evolutionary biology
from which he takes the concept. As the anonymous reviewer claims, a justification this paper’s recourse
to Reed’s idiosyncratic way of thinking must be provided. The reason is primarily that Reed’s contribu‑
tion provides the first attempt to relate evolutionary biology and ecological psychology systematically,
thereby offering a milestone in the history of ecological psychology and pioneering new paths for under‑
standing the connection between non-representational, non-cognitivist psychology and evolutionary biol‑
ogy, a research line that has been deepened in the latest years (see, for example, Stotz 2014). In this
sense, Reed’s understanding may be rather idiosyncratic, as the reviewer claims, but his contributions
are sufficient to include his views as one key approach to take into account when discussing the relation
between evolutionary biology and affordances.
13
mutation that can be taken as favorable for detecting that information, or the prod‑
uct of training to detect it during the ontogenetic development of different agents.
In the latter case, elements like the presence of ecological inheritances or the edu‑
cation of attention taught by mates in a particular niche of a given organism dur‑
ing its development critically improve the possibility of developing a special sen‑
sitivity to detect ecological information, as happens in the case of humans (Gibson
1950: 155; Costall 1995: 477; Ingold 2001/2011: 36). Hence, the configuration of
the niche in general and the importance of the education of attention as a decisive
factor for detecting ecological information in particular have been emphasized by
theories of perceptual learning from an ecological perspective (see "Affordances and
niche construction" section above). Consequently, those agents that inhabit a niche
filled with social mediation and instruction are favored when they compete against
other organisms for the same resources (in this sense, ecological information is a
resource for perception according to Reed). This is the way in which niche construc‑
tion processes and natural selection relate with regard to the evolutionary role of
affordances. In fact, supporters of niche construction appeal to these two mecha‑
nisms as working continuously and affecting each other. This is why they claim that,
“[w]hen such modifications [of the niche] alter natural selection pressures, evolution
by niche construction is a possible outcome” (Laland et al. 2016: 192). In this sense,
modifications occur and give rise to alterations, but these new alterations also exert
selective pressure as well.
As these two evolutionary mechanisms exert their power in parallel, it would not
be strange to assume that the modifications of the environment produced in niche
construction processes make a difference in the way in which natural selection pre‑
serves a certain genetic pool, as we can expect in the case in which humans educate
their offspring to detect ecological information as a resource that they can find in
their environments.
As I recalled earlier, there are authors that hold the environment to be autono‑
mous with respect to organisms, which means that it has its own intrinsic dynam‑
ics (Godfrey-Smith 2001: 254). Yet, if we conflate natural selection and niche con‑
struction in the same picture, the alterations of the environment made by agents at
the organismal level drastically change the alleged “intrinsic dynamics” of the so-
called “autonomous” environment, and consequently the effects of niche construc‑
tion processes at the organismal level have an impact on the very functioning of
natural selection, because those environmental alterations increase the chances of
survival and reproduction of agents, which has an indirect impact at the sub-organis‑
mal level because it facilitates the preservation of the genotype of organisms. In this
picture, the environment has no fully autonomous dynamics, at least if “autonomous
dynamics” is understood in the minimal sense that the action of organisms does
not affect how environmental pressures will be exerted. For example, an environ‑
ment full of negative affordances (those that are injurious—see Gibson 1979/2015:
128–129) that minimize the probability of survival and reproduction can be restruc‑
tured by agents (willingly or unwillingly) so as to become beneficial for them and
their offspring. In this sense, it seems far from obvious that the environment pos‑
sesses dynamics isolated from the effect of the organism. This is also reflected, for
example, in aspects that involve development. In this sense, while it is true that what
13
happens at the sub-organismal, genetic level is partially autonomous from what hap‑
pens at the phenotypic, organismal level (because the principles of variation, fitness,
and heredity imply their own functioning), if we accept that developmental pro‑
cesses and environmental conditions affect the expression of genes (as they usually
do—see Reed 1985: 366; Hunter 2005; Lobo 2008; Ralston and Shaw 2008) and
that alterations made at an organismal level may facilitate the work of preserving
a certain genetic pool at a sub-organismal level, then we should not conceive the
environment as having intrinsic dynamics independent of the dynamics of organ‑
isms, as supporters of the gene-centered view claim. If the previous claim is on the
right track, some symmetry should be included in the evolutionary picture (Walsh
2015) once we aim to start offering a view that combines niche construction and
natural selection together (which seems the most plausible and reasonable approach,
given the scientific evidence in favor of both mechanisms). In this view, symmetry
is achieved because organisms constantly modify their environment, thereby altering
their pre-existing affordances so as to make a more habitable and protective habitat,
which in turn affects organisms back not only at the organismal level (via niche con‑
struction) but also at a sub-organismal level (via natural selection).
This previous reflection leads us to the main tension introduced in Sect. 1.
Namely, focusing on different mechanisms leads to different understandings of the
role of organisms, environments, and affordances in evolution. As seen previously,
both evolutionary mechanisms are related, which invites one to think that it is not
entirely clear that environments and organisms are totally independent, as some
authors suggest. At the same time, the allegedly contrasting views of affordances do
not seem to be irreconcilable. There is a tight connection between niche construc‑
tion and natural selection because, as seen earlier, modifications in the niche lead to
alterations in natural selection pressures, which may result in niche construction as a
possible outcome (Laland et al. 2016: 192). Nevertheless, even when those modifi‑
cations alter the already existing pressures, there will be others that result from this
alteration of the niche, so the alteration of the niche does not inhibit the function‑
ing of natural selection. This shows that there are not really two isolated mecha‑
nisms working in parallel; rather, there are two mechanisms that are continuous or
connected, but they function in parallel at two different times or moments. This is
quite illuminating for understanding the role of affordances, because this shows that
the allegedly competing views are not really competing. As one will recall, Reed’s
selectionist view defines affordances as resources for regulating the organism’s rela‑
tion to the environment (Reed 1996:17), whereas supporters of niche construction
understand affordances as ecological inheritances that maximize the offspring’s
chances of survival (Withagen and van Wesmerkerken 2010: 505; Heras-Escribano
and De Pinedo-García 2018: 10–11). This leads some supporters of the latter view
to claim that “the niche construction perspective suggests alternative roles [to the
one offered by Reed] for affordances in evolutionary dynamics” (Withagen and van
Wesmerkerken 2010: 497). Whence the alleged incompatibility. However, if one
pays attention to the point made earlier concerning the relation between niche con‑
struction and natural selection (Laland et al. 2016: 192), then one can claim that
there affordances play a double role in evolution: first, they work as a resource that,
in Reed’s sense, apply selective pressure; second, after seizing the affordance and
13
Fig. 1 Adaptation from Figure 1.3 in Odling-Smee et al. (2003: 14). While a is the standard evolutionary
perspective of the MS in which organisms transmit their genes from generation t to generation t + 1, b
represents the EES view in which niche construction processes are included in the picture, where organ‑
isms modify their environments by taking or giving rise to affordances in their environment (A). Each
generation ecologically inherits from their ancestors the affordances that were previously perceived,
taken, or that emerged thanks to their ancestors’ modifications of the environment within niche construc‑
tion processes
after proof of its selective advantage, affordances can work as ecological inherit‑
ances from one generation to the next (offering the same selective pressure), and
so on. In this light, Withagen and van Wesmerkerken’s (2010: 497) point should
be understood while taking into account that the alternative roles for which they
call are still compatible with the view offered by Reed (1996). Thus, affordances
can be understood as ecological inheritances of an already modified environment
that, at a different time, played the role of resources exerting selection pressure in
the new environment produced by the alteration effected by organisms in the niche
construction process: first, they exert selective pressure at stage t1, and then the
same affordances work as inheritances for the offspring at stage t2. In this sense, the
13
evolutionary role of affordances is dual: they are both resources and inheritances.
Adapted from Odling-Smee et al. (2003), the following Fig. 1 diagram exemplifies
this idea:
The reader may note that the adaptation of the diagram originally designed by
Odling-Smee, Laland, and Feldman is not fortuitous. As we have seen in previous
sections, there are different ways to understand the extended synthesis and, given
this variety, affordances are either accepted or rejected as key aspects in the niche
construction processes. In this paper, I argue in favor of the view propounded by
Odling-Smee, Laland, and Feldman and according to which the metabolism and
action of organisms in their ecosystem lead to unexpected consequences in the
direction, rate, and dynamics of the evolutionary process. For ecological informa‑
tion is pervasive in our everyday lives, and affordances are common objects of
perception for every human being. Because they are so common, they must have a
role in our evolutionary history, and I propose that they are related to the two main
evolutionary mechanisms: natural selection and niche construction. Nevertheless,
although natural selection and niche construction are two different mechanisms,
they do not have to be taken as completely discrete and isolated from each other.11
This point is emphasized in the idea that, according to these authors, the alteration
of selection pressures leads to niche construction processes as a possible outcome
(Laland et al. 2016: 192) and in the idea that niche construction processes of ances‑
tors modifies the selection of offspring (Odling-Smee et al. 2003: 11), which shows
that these two mechanisms are not as isolated as one might at first think. For this
reason, the idea that natural selection and niche construction are isolated from each
other makes no sense in light of the approach to niche construction endorsed by
authors such as Odling-Smee and Laland. The discreetness of both mechanisms is
also criticized by other supporters of the extended synthesis (Uller and Helanterä
2019). The idea of affordances casts considerable doubt on the discreetness of both
mechanisms, in addition, because they help emphasize the intertwinement of both
mechanisms in different moments of evolutionary history, playing one role at one
moment (as pressure) and another at a different moment (inheritance). Regarding
this last point, affordances may serve a greater purpose than showing the intimate
connection between natural selection and niche construction. As such, affordances
are aspects that, given their “dual” role in the evolutionary process (as pressures and
as inheritances), have the power to make more visible the impact of niche construc‑
tion process and the potential to re-configure niche construction theory in the life
sciences completely.12
Thus, affordances are key aspects of the niche construction process (Heras-
Escribano and De Pinedo-García 2018), but they are also environmental aspects that
exert selection pressure (Reed 1991, 1996). At this point it becomes clear that affor‑
dances possess an evolutionary role that cannot be avoided and, at the same time,
that this biological aspect of affordances opens the door to conceive a more organis‑
mal-centered, agential, and informational approach to evolution.
11
Thanks to an anonymous reviewer for making me aware of the importance of this idea.
12
Thanks to an anonymous reviewer for inviting me to highlight this consequence.
13
Conclusion
In this paper, I have offered a general attempt to explain the evolutionary role of
affordances. For this reason, I have delved into former views on the evolutionary role
of affordances (the selectionist view and the niche construction view), and I have
analyzed the consequences of understanding affordances in each. According to the
selectionist view (Reed 1996), affordances are mere resources that exert selection
pressure, whereas they are ecological inheritances in the niche construction process
according to the niche construction view (Withagen and van Wesmerkerken 2010;
Heras-Escribano and De Pinedo-García 2018). According to some authors (With‑
agen and van Wesmerkerken 2010), there seems to be a tension in which two dif‑
ferent views on evolution lead to two different views on affordances. Nevertheless,
I have shown how the allegedly incompatible views on affordances reconcile. These
views on affordances are complementary because niche construction and natural
selection are two different mechanisms that are connected, although one does not
alter the functioning of the other (Laland et al. 2016). The alterations of the envi‑
ronment in the niche construction process alters the selective pressure of different
aspects of the environment, but even these alterations lead to a new configuration of
the environment the elements of which will also exert selective pressure: in this sce‑
nario, this means that affordances (if they remain without alteration) exert selective
pressure at stage t1, and then these affordances are inherited by the offspring as eco‑
logical inheritances at stage t2, thereby exerting selective pressures as well depend‑
ing on the configuration of the niche. In this sense, affordances are both resources
and ecological inheritances, although they function as one or the other depending on
the moment in the evolutionary process. This dual nature of affordances shows their
evolutionary role.
Acknowledgements Thanks to Ezequiel Di Paolo for fruitful suggestions to an earlier version of this
paper and also to Cristian Saborido, Javier González de Prado, Lorena Lobo, Víctor Luque, Donald
Cross, and David Teira for their comments.
Funding This paper has been written thanks to a 2018 Leonardo Grant for Researchers and Cultural
Creators, BBVA Foundation (The Foundation accepts no responsibility for the opinions, statements
and contents included in the project and/or the results thereof, which are entirely the responsibility of
the authors), the Project FFI2016-80088-P funded by the Spanish Ministry of Science, and the FiloLab
Group of Excellence funded by the University of Granada, Spain.
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Biology & Philosophy (2020) 35:30

The evolutionary role of affordances: ecological

Keywords Affordances · Evolution · Niche construction · Natural selection

Thus, the role of affordances in evolution is understood in two very different

Ecological psychology and affordances

Ecological psychology is an approach to perception and perceptual learning pio‑

time, but in an ecological sense of being regular within an ecological context of

dynamics as well: if the processes of niche construction lead to a certain modifica‑

The extended evolutionary synthesis and niche construction theory

dairy farming, or the construction of dams5); second, inceptive relocations or the

phenotype, niche construction theory emphasizes the legacy of modifications that

Affordances, niche construction, and natural selection

Affordances and niche construction

There are various similarities and complementarities between niche construction

The affordances of our human environment are aspects of our environment

Introducing natural selection

1. There is variation in morphological, physiological, and behavioral traits among

As such, natural selection is a mechanism that works at a population level since

compete in an environment where that information is a relevant resource10 that

Unifying natural selection, niche construction, and affordances in a single picture

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