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ORIGINAL RESEARCH
Erkenntnis
https://doi.org/10.1007/s10670-023-00695-2
Abstract
Selected-eects theories provide the most popular account of biological teleology.
According to these theories, the purpose of a trait is to do whatever it was selected
for. The vast majority of selected-eects theories consider biological teleology to
be introduced by natural selection. We want to argue, however, that natural selec-
tion is not the only relevant selective process in biology. In particular, our proposal
is that biological regulation is a form of biological selection. So, those who ac-
cept selected-eects theories should recognize biological regulation as a distinctive
source of biological teleology. The purposes derived from biological regulation are
of special interest for explaining and predicting the behavior of organisms, given
that regulatory mechanisms directly modulate the behavior of the systems they
regulate. This explanatory power, added to the fact that regulation is widespread in
the biological world, makes the idea that regulation gives rise to its own form of
teleology a substantial contribution to the debate on biological teleology.
Received: 24 May 2022 / Accepted: 6 May 2023
© The Author(s) 2023
Biological Purposes Beyond Natural Selection: Self-
Regulation as a Source of Teleology1
JavierGonzález de Prado1· CristianSaborido1
1 The authors are listed alphabetically. Both of them have contributed equally to this work. We are
grateful to audiences at the University of Bielefeld, Complutense University (Madrid), the University
of Helsinki, the National University of Colombia (Bogotá), Nova University of Lisbon, the University
of Oslo, the University of Prague, UAM (Madrid), the University of the West of England Bristol, and
UNED (Madrid). Special thanks to Leonardo Bich, Matteo Mossio and several anonymous reviewers.
Thanks as well to Megan J. Watkins for the linguistic revision of the article. This work has been
supported by the Spanish Government research projects APID2021-128835NB-I00 and PID2021-
123938NB-I00.
Javier González de Prado
jgonzalezdeprado@fsof.uned.es
Cristian Saborido
cristian.saborido@fsof.uned.es
1 Department of Logic, History and Philosophy of Science, UNED, Madrid, Spain
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J. González de Prado, C. Saborido
Selected-eects theories provide the most popular account of biological teleology.
According to these theories, the purpose of a trait is to do whatever it was selected
for (Millikan 1984, 1989; Neander, 1991; Griths, 1993; Godfrey-Smith, 1994). The
vast majority of selected-eects theories consider biological teleology to be intro-
duced by natural selection (an exception is Garson (2012, (2017, 2019b)). We want
to argue, however, that natural selection is not the only relevant selective process in
biology. In particular, our proposal is that biological regulation is a form of biologi-
cal selection. So, those who accept selected-eects theories should also recognize
biological regulation as a distinctive source of biological teleology. The purposes
derived from biological regulation are of special interest for explaining and predict-
ing the behavior of organisms, given that regulatory mechanisms directly modulate
the behavior of the systems they regulate. This explanatory power, added to the fact
that regulation is ubiquitous in the biological world, makes the idea that regulation
gives rise to its own form of teleology a substantial contribution to the debate on
biological teleology.
We intend for our proposal to have a conciliatory, pluralistic spirit. An important
upshot of the paper is that evolutionary accounts of teleology do not need to be in
competition with approaches that, like ours, focus on the current organization of bio-
logical systems. As we will show, in both cases teleology can be seen as underlain
by selective mechanisms. However, this does not mean that these approaches are
equivalent. Given that they involve very dierent types of selective processes (natu-
ral selection and regulation, respectively), the resulting attributions of purposes and
teleological explanations also tend to be signicantly dierent. Nonetheless, these
teleological explanations can be compatible and complementary. What we want to
stress is that selected-eects theories do not commit one to considering natural selec-
tion to be the only source of genuine biological teleology. Regulation provides a clear
example of a biological selective process that operates at the level of the dynamics
of individual organisms, and that generates a distinctive form of biological teleology.
This is the plan for the paper. In the rst section we examine standard etiological
accounts of biological teleology, which rely on natural selection. These etiological
accounts are typically underpinned by selected-eects theories of teleology. In the
second section, we oer some general motivation for such selected-eects theories.
After that, we characterize the phenomenon of biological regulation. Finally, we
argue that biological regulation is a form of selection, and that, therefore, accord-
ing to selected-eects theories, it should be regarded as a source of teleology. We
also show how our proposal avoids counterexamples faced by other selected-eects
theories.
1 Etiological-evolutionary Accounts of Biological Teleology
Teleological notions such as purpose, goal, or function are widespread in biological
discourse. Such teleological notions are associated with standards of success. Pur-
posive behavior is successful if it reaches its purpose or goal, and failed otherwise.
In virtue of this link with success and failure, teleology has an evaluative dimen-
sion—understanding evaluative normativity as having to do with what is good or bad
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Biological Purposes Beyond Natural Selection: Self-Regulation as a…
(benecial or detrimental) in some way (McLaughlin, 2009). Standards of success
are evaluative, insofar as successful performances count as good qua instances of
purposive behavior. Saying that some purposive behavior has been successful is a
way of evaluating it (positively), with respect to the purpose of that behavior.
We will assume here that biological teleology is observer-independent, rather
than being due to projections of purposes by external observers (or being a form of
metaphorical or loose talk). According to this assumption, there are some entities in
the biological world such that being those entities entails having certain purposes,
and therefore being evaluable in terms of success, independently of projections from
external observers. We will also assume that biological organisms are (sophisticated)
complex physical systems. The question that arises is how certain physical systems
(i.e. biological organisms) become subject to teleological standards.
A popular answer to this question appeals to the etiology of the relevant biological
trait, more specically to its evolutionary history (Millikan, 1984, 1989; Neander,
1991; Griths, 1993; Godfrey-Smith, 1994).1 Broadly, the idea is that the purpose
of a biological trait is doing what it was naturally selected for doing. So, a token of
a biological trait has as its purpose producing some eect if producing that type of
eect is what explains the proliferation of the ancestors of that token in the face of the
pressures of natural selection. We will call this kind of approach an etiological-evo-
lutionary account of teleology. To repeat a classic example, the purpose of hearts is
to pump blood, because pumping blood explains why hearts managed to proliferate.
An initial virtue of this approach is that it seems to capture the circular dependence
between eects and causes distinctive of teleology. According to Wright’s analysis
of teleology (1976: 39), the presence of a purposive trait is explained by its tendency
to produce certain eects, which constitute its purpose. Similarly, Walsh (2008: 113)
claims that “teleology is a mode of explanation in which the presence, occurrence,
or nature of some phenomenon is explained by the end to which it contributes.” In
the case of purposive biological traits, the existence of a current token of the trait
would be explained by the fact that past members of its lineage or reproductive chain
produced certain eects, which are therefore the eects that the current token of the
trait is supposed to have.
However, even if it is granted that the continued presence of a biological trait is
explained by its tendency to produce certain eects, it is not immediately clear why
this, on its own, would entail that the trait has a purpose and is subject to standards
of success. One should not conate necessary conditions for perpetuation with pur-
poses. Existing because of the tendency to produce some eect is not the same as
existing in order to produce that eect.
We think that the best way to vindicate etiological-evolutionary accounts is to
focus on the role of selective processes in Darwinian evolution. Most advocates
of etiological-evolutionary approaches appeal, explicitly or implicitly, to selected-
eects theories of teleology, according to which selective processes give rise to
purposes (Millikan, 1984, 1989; Neander, 1991; Griths, 1993; Kitcher, 1993;
1 Alternative accounts of biological teleology, which we will not discuss here, are provided, among others,
by systemic or dispositional theories (Cummins, 1975; Craven 2001; Boorse 2002) and by organizational
approaches (Christensen & Bickhard, 2002; Mossio et al., 2009).
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J. González de Prado, C. Saborido
Godfrey-Smith, 1994). In Griths’s words, “where there is selection there is teleol-
ogy” (1993: 420). From the perspective of these theories of teleology, purposes are
selected eects: the purpose of some entity is to produce certain eects if its tendency
to produce those eects explains why the entity was preserved through a selective
process.
Selected-eects theories of teleology oer direct support to the view that biologi-
cal purposes are introduced by natural selection, provided that it is assumed that natu-
ral selection is a genuine selective process. So, biological teleology would result not
merely from the circular dependence between eects and causes in biological traits,
but from the fact that such dependence is mediated by natural selection.
We will not oppose here the view that natural selection is a source of biological
teleology. Our goal is rather to show that natural selection is not the only relevant
form of selection in biology. In particular, biological regulation should be counted as
a selective process too, at least inasmuch as natural selection can be so counted. What
we want to argue is that those who are happy to appeal to selected-eects theories
in relation to natural selection should also be willing to do the same in relation to
biological regulation, and therefore to recognize it as a distinctive source of biologi-
cal teleology. So, for our purposes in this paper, we could just take selected-eects
theories for granted. Nevertheless, in the next section we oer further clarication of
such theories and provide some motivation for them. We will argue that the idea that
selection introduces purposes is particularly plausible once we consider the type of
(thin) evaluative normativity characteristic of teleology.
2 Selected-eects Theories of Teleology
Selection is closely related to evaluation. Indeed, selection can be seen as classica-
tion plus evaluative valence. Selective processes involve sorting items into groups
with positive and negative valences—items selected for are positively evaluated,
while items selected against receive negative evaluations, relative to the evaluative
standard associated with the relevant selective process.
Of course, if the notion of selection is taken to be constitutively linked to evalua-
tive standards, then the claim that there is evaluation wherever there is selection will
be trivially true. However, we are interested in using selected-eects theories to oer
a naturalistic account of the emergence of evaluative standards in biology. In order
to do so, we need to characterize selective process without presupposing evaluative
standards. Otherwise, we would have not explained how evaluative standards arise
in biology. The aim, therefore, is to describe, in non-evaluative terms, biological pro-
cesses that can be counted as forms of selection, and then argue that these processes
give rise to evaluative standards.2
Campbell (1960) considers that any selection process involves blind variation and
selective retention. An attractive option is to generalize this idea, thinking of selec-
tion in terms of dierential reinforcement over a set of items featuring variability
2 For abstract characterizations of selective processes, see Darden and Cain (1989) and Hull, Langman
and Glenn (2001).
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Biological Purposes Beyond Natural Selection: Self-Regulation as a…
in their properties. In selective processes, items with certain features are retained,
reproduced or promoted, which can be seen as a form of positive reinforcement,
whereas other items are inhibited or rejected, which is a form of negative reinforce-
ment. The features or eects selected for are those that explain why the relevant items
are positively reinforced. Imagine, as an example, a gardener tending their garden by
preserving owers and cutting weeds. This gardener will be positively reinforcing
the presence of owers, while inhibiting the presence of weeds. It is natural to think
of the gardener as selecting owers, but not weeds, as a satisfactory contribution to
their garden.
In general, processes driven by dierential positive and negative reinforcement
are intuitively seen as instances of selection. It makes sense to consider natural selec-
tion a selective process precisely because it involves dierential reinforcement, in the
form of dierential reproductive rates. Learning by trial and error is another example
of selection underlain by reinforcement (see, among others, Millikan 1984; Garson,
2017). In this case, the learner develops dispositions to repeat some behaviors and not
others depending on whether they are observed to produce certain outcomes reliably.
It should be stressed that dierential reproduction is only one of the possible forms
of reinforcement that can underlie selection. Given the prominence of Darwinian
evolution in biological research, it is understandable that biological selection tends
to be identied with natural selection, and therefore with dierential reproduction.
However, biological selection can take place via other forms of reinforcement. For
instance, Garson (2011, (2012, (2017, 2019b) describes neural selection as a case of
biological selection involving dierential retention without dierential reproduction.
And, in principle, there could be further types of selective reinforcement, for example
dierential activation of certain processes or dierential recruitment of some mecha-
nisms (that is, situations in which several mechanisms are available for a given task
and some of them are recruited more frequently than others, even if all those mecha-
nisms are retained). It is wise to remain fairly liberal about the types of reinforcement
potentially involved in biological selection, given the variety of shapes that reinforce-
ment can take in ordinary cases of selection (for example, selecting one’s clothes for
an interview, selecting candidates for a job, selecting a lm to watch, crop selection,
or animal breeding).
Dierential reinforcement does not presuppose evaluative standards, so we can
use the characterization of selection given above to account for the emergence of bio-
logical teleology. Let us call the causal powers responsible for the relevant patterns
of reinforcement “selective pressures.” In the selected-eects theory we are explor-
ing, selective pressures introduce teleological evaluative standards. The reason why
these standards can be said to be teleological is that they satisfy the teleological loop
postulated by Wright (1976): the reinforcement of the selected items is explained by
their tendency to produce those eects that got them selected. Notice that this type
of teleological explanation can be applied to instances of selection regardless of the
form of reinforcement involved. So, when reinforcement takes place as dierential
reproduction, we would explain such dierential reproduction by appeal to certain
eects of the items that managed to reproduce (this is what happens in explanations
of natural selection). In a similar way, when selection happens via dierential reten-
tion, we would point to relevant eects of the items selected in order to explain why
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J. González de Prado, C. Saborido
they have persisted. The teleological nature of selection does not depend, therefore,
on the involvement of dierential reproduction (see Garson 2017, 2019b).
We are not claiming, to be clear, that all instances of dierential reinforcement will
automatically count as cases of selection. Fuzzy, borderline cases are to be expected.
However, we think that when the relevant dierential reinforcement is suciently
complex and exible, the resulting processes can share enough features of paradig-
matic forms of selection to deserve being classed as such. At least, we think, this is
something that defenders of selected-eects theories should concede. In particular,
as we will argue below, biological regulation can legitimately be seen as a type of
selective process.
According to the view we are exploring, selective pressures do not presuppose
evaluative standards, but rather institute them. Thus, selected-eects theories would
be an instance of a more general metanormative approach in which evaluative stan-
dards are taken to be generated by responses or pressures that can be characterized
in non-evaluative terms (in this case, in terms of reinforcement). This metanorma-
tive commitment may seem contentious, at least when applied to normative issues
generally. However, selected-eects theories become far less controversial once we
observe that they only address the type of thin evaluative normativity underlying
teleology. The evaluations linked with ascriptions of success involve what is usually
known as attributive goodness, rather than predicative goodness (for this distinc-
tion, see Geach 1956; Thomson, 2008). One makes attributive evaluations when one
says that something is good as an instance of a certain kind (for instance, that some
footballer is good as a goalkeeper). By contrast, in evaluations involving predicative
goodness, one treats something as good simpliciter, and not just as an instance of a
kind. An example is saying that peace is good and war is bad.
When we treat a performance as successful we are assessing it as good only with
respect to the purpose of the relevant behavior (that is, good as purposive behavior).
It does not follow from this attributive evaluation that that successful performance
is good simpliciter (good in a predicative sense), or good when not assessed as an
instance of the relevant purposive behavior but as an instance of a dierent kind.
It may well be that things that are good as instances of a kind are not, all things
considered, desirable, admirable, or worth promoting. So, we are not claiming that
successful behaviors are always good in a way that makes them desirable or worth
promoting. In many cases, rather the opposite is true; for example, successful mur-
dering behaviors are to be stopped and prevented.
It is also important to distinguish evaluative normativity, which concerns what is
good and bad, from prescriptive normativity, which concerns obligations, permissions
and prohibitions. Attributive evaluations, like those associated with assessments of
success, do not need to have immediate prescriptive implications. There may be no
obligation whatsoever to engage successfully in a certain purposive behavior.
Thus, the type of normativity that we are associating with teleology is quite
lightweight, in that it does not involve prescriptions or what is good simpliciter. We
can remain neutral about whether more substantial normative standards (like those
associated with morality, or with goodness simpliciter) are generated by reinforcing
responses. As far as selected-eects theories are concerned, we only need to endorse
the plausible view that reinforcing responses can determine what counts as good with
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Biological Purposes Beyond Natural Selection: Self-Regulation as a…
respect to the standards governing activities shaped by those responses (regardless
of whether it is good simpliciter to engage in such activities). More specically, the
pressures giving form to a certain selective process would determine what counts as
good according to the standards governing that selective process. Let us insist that
this is only attributive goodness, internal to the standards of a process of selection. In
fact, it may be that there is nothing good simpliciter in the existence of a certain selec-
tive process, and it can even be the case that there are strong reasons to prevent some
kinds of selection from happening (think of employee selection processes based on
race or gender rather than merit).
It is instructive to consider at this point the examples of games and social norms
(see Bicchieri 2006). Arguably, if the participants in a game treat certain moves as
good or successful while playing the game, those performances count as good or
successful as moves in that rule-governed game. There is no further, external stan-
dard that determines whether a certain move in the game is successful, beyond the
responses and attitudes of the players of the game (at least if we are considering
non-institutionalized games that are created from scratch by the players). Something
analogous happens with many social norms, for instance norms of etiquette. Whether
certain behaviors count as good table manners in a given social dining practice
depends on whether the participants in the practice treat them as such. Indeed, what
is good manners according to a certain dining practice can be bad manners in a dif-
ferent cultural context, where participants reinforce and sanction dierent behaviors.
Of course, that some behaviors count as good manners according to a dining practice
does not mean that such behaviors are good simpliciter or should be promoted. After
all, the relevant dining practice could be sexist, classist, or bad in other respects.
In the same way that the normative standards internal to games and social prac-
tices are established by the attitudes and sanctioning dispositions of participants, the
standards internal to selective processes would be generated by the pressures con-
stitutive of the process. Given that selective processes are driven by the pressures
introducing the relevant standards, these standards can be said to actively govern
such selective processes.
3 Biological Selection Beyond Evolution
We are aware that the considerations in the previous section may not be universally
convincing. For instance, some could insist, perhaps appealing to Moorean open-
question arguments, that a purely causal process like dierential reinforcement can-
not introduce the evaluative standards underlying selection (see Bedau 1991 for such
an anti-reductionist view). While we are not moved by these strong anti-reductionist
intuitions, in particular when dealing with the lightweight normativity characteristic
of teleology, we leave that debate for another occasion. Our immediate target here
is those who accept selected-eects theories, and in particular those who are will-
ing to apply this type of theory to natural selection. So, in what follows we will
take selected-eects theories for granted, as one of our assumptions, understanding
selection in terms of dierential reinforcement. We want to extend selected-eects
theories beyond Darwinian evolution, arguing that natural selection is not the only
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J. González de Prado, C. Saborido
plausible selective process in the biological world (see Garson (2017, 2019b) for a
project in the same direction). More specically, our claim is that biological regula-
tion constitutes a form of selection, and therefore it deserves to be counted as a dis-
tinctive source of biological teleology.3
Thus, the argument put forward in this paper is conditional. If selected-eects theo-
ries are accepted, then biological regulation should be taken to introduce teleological
standards of success. To be clear, we do not intend to attack etiological-evolutionary
theories of biological teleology. The main contribution of this paper is constructive,
not critical. Our goal is to draw attention to the possibility of considering biological
regulation, in addition to natural selection, as a legitimate source of teleology. This
is, we think, a substantial contribution, given how widespread biological regulation
is in nature. Although the argumentative scope of the paper is restricted as specied
above, the conclusions we will draw are still far-reaching and worth exploring. Argu-
ably, selected-eects theories oer the dominant naturalistic account of biological
teleology, so it is interesting to note that they allow us to consider biological regula-
tion as introducing teleology in the natural world.
It is worth stressing that we are not claiming that selective processes introduce
teleological standards to the extent that they resemble natural selection. Rather,
according to the selected-eects theory we are putting forward, it is the other way
around: natural selection is teleological insofar as it counts as a form of selection.
There are selective processes associated with teleological standards despite the fact
that they do not meet some of the conditions characteristic of natural selection, for
instance processes in which selective pressures do not operate over past tokens of
a type, but over the current performances of some item. Think, as an example, of a
recruitment process in which candidates are selected according to their performance
in an exam. Here, the relevant selective reinforcement targets the candidates’ cur-
rent performance in the exam, and not their past behavior, or the behavior of past
individuals.
Are we stretching the notion of selection unduly? There is no reason to think so.
Many central, paradigmatic types of selection do not resemble natural selection. If
anything, it is natural selection that is a heterodox form of selection. In particular,
stereotypical cases of selection involve identiable systems or mechanisms that exert
the relevant reinforcing pressures (that is, a selector). As we will see, in biological
regulation there are regulatory sub-systems, integrated into the organism, that play
this selecting role. By contrast, in natural selection the source of reinforcement is
environmental forces, which usually do not constitute individuated mechanisms or
systems. In this respect at least, biological regulation is closer to stereotypical selec-
tive processes than natural selection.
Of course, we are not the rst to generalize selected-eects theories beyond natu-
ral selection. This generalization has already been suggested by Wimsatt (1972: 13),
who claims that “the operation of selection processes is not only not special to biol-
3 Schroeder (2014) has defended the view that natural regulation generates functions, relying on the anal-
ogy with the way in which everyday forms of regulation confer functions to objects. We follow an alterna-
tive path here: we aim to integrate biological regulation into a general selected-eects theory of teleology,
arguing that regulation constitutes a selective process.
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Biological Purposes Beyond Natural Selection: Self-Regulation as a…
ogy, but appears to be at the core of teleology and purposeful activity wherever they
occur.” Similarly, Neander (1991) notes that selected-eects theories could in prin-
ciple be applied to any selective process, not just natural selection, although she does
not develop this line of thought. Several authors, working from the perspective of
selected-eects theories, have pointed out that trial and error learning can introduce
purposes (for instance, Wimsatt 1972, 2002; Millikan, 1984; Griths, 1993).
In this way, our proposal should not be seen as conicting with standard selected-
eects approaches, but rather as continuous with them. As we have just seen, many
authors consider selected-eects theories to oer a general framework for account-
ing for teleology, not only in biology but also in relation to purposeful behavior in
other domains. However, when applied to biology, selected-eects theories tend to
focus overwhelmingly on natural selection. We want to resist this dominant tendency,
and extend selected-eects approaches to other biological processes, in particular
regulation. In this respect, Garson’s work (2011, 2012, 2017, 2019b) is especially
relevant for our purposes. Garson’s claim is that selected-eects theories should not
be restricted to processes driven by dierential reproduction, such as natural selec-
tion, but should also allow for selective processes involving dierential retention, in
particular neural selection. In this way, Garson argues that selected-eects theories
should be seen as disjunctive theories, covering both cases of dierential reproduc-
tion and of dierential retention.
We agree with the spirit of Garson’s proposal, even if we prefer to think of the
generalization of selected-eects theories as a unied view, rather than a disjunctive
one, as Garson presents it. The unifying element in this generalized theory would
be the idea of selection, characterized in terms of the notion of reinforcement. As
we have pointed out, reinforcement can take dierent shapes, including dieren-
tial reproduction and dierential retention, but this does not mean that selection is a
disjunctive kind. A more substantial dierence between our proposal and Garson’s
is that our characterization of teleological selection is wider than his. In particular,
the phenomenon we are going to focus on, biological regulation, does not typically
meet the conditions for selective processes set by Garson’s theory. According to Gar-
son (2017), the relevant forms of selection involve either dierential reproduction or
retention in a population of tokens of a type engaged in tness-relevant (competitive
or cooperative) interactions. This is not what happens in most cases of biological
regulation. Blood sugar regulation does not involve interactions within a population
of dierent levels of blood sugar. But, again, there are many paradigmatic examples
of teleological selection in which this condition is absent. Selection does not nec-
essarily require interactions among the items undergoing the selective process. For
instance, we can select which students pass a course, and which fail, by means of
an individual exam, with no interactions among the students. Indeed, there can be
selection with only one candidate. Think of the process of selecting candidates for an
award by means of an exam. This process can take place even with a single candidate:
if the candidate passes the exam, they will be selected for the award, otherwise they
will not (none will get the award). Despite not involving a population of items with
alternative traits, this is clearly a process of selection (even if not of selection over
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J. González de Prado, C. Saborido
existing alternative traits).4 So, although Garson surely discusses an interesting type
of biological selection, processes that do not t his account can nonetheless be close
to paradigmatic forms of selection.
Garson introduces the conditions we have just discussed to avoid counterexam-
ples that threaten to make selected-eects theories too liberal. We will show below
that the type of selection associated with biological regulation is not aected by at
least the most glaring of these counterexamples. We will also argue that allowing
for regulatory teleology has explanatory and theoretical benets analogous to those
vindicating selected-eects theories in relation to natural selection (and to Garson’s
neural selection). However, before that, in the next section, we present the notion of
biological regulation in more detail.
4 Biological Regulation
Biological organisms are able to regulate their behavior in the face of perturbations
and to adapt it to environmental changes, so as to remain in congurations that allow
the organism to keep existing (Rosen, 1970; Di Paolo, 2005; Bich et al., 2016).
Think, for instance, of a bacterium that modulates its movement searching for nutri-
ents in its environment, or a sick human that compensates for a feverish state through
perspiration.
Living organisms are complex self-maintaining systems (see Moreno & Mossio
2015). This means that the activity of an organism contributes to the preservation of
the conditions in which its existence is viable. In this way, organisms sustain their
own existence and identity. Among other things, this involves repairing and regener-
ating their constituents, obtaining enough nutrients, disposing of waste, and avoiding
toxic substances.5
Organisms are often threatened by external and internal perturbations that push
them away from the conditions favorable to their existence (for instance, changes in
the temperature of the environment, scarcity of nutrients, or the presence of preda-
tors). The self-maintenance of organisms will, therefore, be fragile unless they are
able to react to these perturbations in ways that tend to keep the organism within its
range of viability conditions. At rst pass, biological regulation can be understood
as the capacity of organisms to modulate their own behavior in response to perturba-
4 What is essential for selection is that it presents a suitable modal prole: traits are selected because they
produce certain eects, and would not have been selected if they did not produce them. This modal prole
may require the existence of a population with alternative traits, for instance in cases in which the competi-
tion among such alternative traits explains the dierential reinforcement of traits with the relevant eects.
But, as the example of the lone award candidate shows, there are selective processes with a suitable modal
dimension despite not involving a population of traits. In this way, the lone candidate in the example above
will not be selected for the award if they do not pass the exam, regardless of the absence of competitors.
5 We will esh out the notion of regulation within an organizational framework, following, among others,
Bich et al. (2016). However, the type of biological teleology we want to explore diers from that studied
by organizational authors like Mossio, Saborido and Moreno (2009), who focus on purposes associated
with the contribution of traits to the self-maintenance of organisms. We will examine instead the purposes
that, according to a selected-eects approach, are introduced by self-regulation. We consider these two
approaches to be complementary.
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Biological Purposes Beyond Natural Selection: Self-Regulation as a…
tions that endanger their maintenance. Of course, regulation is not infallible. Organ-
isms will not always manage to counteract the eects of perturbations. Yet regulatory
responses will at least have the tendency to prevent perturbations from undermining
the organism’s viability conditions.
It is important to note, however, that a notion of regulation that includes all com-
pensatory reactions to perturbations would be too broad for our purposes (see Bich
et al., 2016; Bich, 2018; Bich et al., 2020). For instance, reversible chemical reac-
tions show a certain degree of robustness against perturbations, as encapsulated in
Le Chatelier’s principle. In general, physical systems near stable equilibrium states
tend to oppose perturbations that try to take the system away from the equilibrium
(think, for example, of a marble at the bottom of a bowl). We do not want to conate
the behavior of these simple physical systems near equilibrium with the more sophis-
ticated forms of adaptation to perturbations characteristic of biological organisms.
Thus, we will work with a restricted, regimented notion of regulation that does not
count every system near stable equilibriums as self-regulated.6
More specically, we reserve the (regimented) notion of regulation for complex
systems in which there are specialized mechanisms that modulate the behavior of the
system in response to the relevant perturbations (see Bich et al., 2016). In self-reg-
ulated organisms, therefore, there is a hierarchical distinction between higher-level
regulating mechanisms and the lower-level mechanisms and processes regulated by
them. This does not happen, for example, in the case of a marble moving around
the bottom of a bowl. Higher-level regulatory sub-systems are also absent in other
cases of simple dierential retention—for instance, in the example of the beach
rocks eroded dierentially by the sea, discussed by Garson (2017). Thus, our account
avoids potential charges of overgeneralization faced by proposals that associate tele-
ology with equilibrium systems or mere dierential retention.
To be clear, regulatory sub-systems are an integrated part of the organism, insofar
as such sub-systems are sustained by the activity of the rest of the organism. Yet it
is possible to dierentiate these regulatory sub-systems from the other parts of the
organism whose behavior they modulate. Bich et al. (2016) propose delineating this
distinction in terms of what they call dynamical decoupling: this type of decoupling
occurs when the relevant sub-systems work at “dierent intrinsic rates”, so that there
is a certain degree of independence between their activities (Bich et al., 2016: 254).
The conception of regulation that we will adopt here is nicely summarized by
Bich, Mossio and Soto:
regulation consists of the capacity to selectively modulate the rst-order self-
maintaining regime in response to specic variations of the internal and external
environment, due to the action of a dynamically decoupled dedicated control
subsystem that is sensitive to these variations. (Bich et al., 2020: 9).
6 Bedau (1992) argues against the idea that equilibrium systems can generally be taken to be goal-directed,
objecting that it leads to overgeneralization. As we explain above, we do not consider systems near equi-
librium as self-regulated. Indeed, self-regulated biological organisms are far-from-equilibrium systems
(see Moreno & Mossio 2015).
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J. González de Prado, C. Saborido
Thus, biological regulation, as we understand it, involves specialized or dedicated
sub-systems that modulate the behavior of other parts of the organism in response
to internal or external perturbations. It should be stressed that this talk of dedica-
tion does not presuppose teleological notions. In particular, we are not assuming that
dedicated regulatory mechanisms have the purpose of modulating the behavior of the
system. Self-regulation involves dedicated mechanisms in the sense that there are
sub-systems that contribute causally to the dynamics of the system by producing the
pressures that generate the relevant dierential reinforcement. We can characterize
this contribution in purely causal terms, as one would do in Cummins’s causal-role
approach to functional explanations (1975).
Regulatory modulation may amount to modifying the rate or intensity at which
some mechanism operates. For example, one way in which the secretion of insu-
lin contributes to blood sugar regulation is by stimulating the uptake of glucose in
muscle and fat tissue cells. However, as Bich et al. (2016) emphasize, in other cases
the relevant modulation is a matter of switching the regime under which the regulated
sub-system works—so that the regulated sub-system changes its mode of operation.
For instance, in bacterial chemotaxis, bacteria control the direction of their move-
ment by alternating between two modes of rotation of their agella (clockwise and
counter-clockwise). Another example is the regulation of gene expression in the lac
operon, which allows bacteria to switch from metabolizing glucose to metabolizing
lactose (for details see Jacob & Monod 1961; Müller-Hill, 1996; Bich et al., 2016).
The lac operon is a clear illustration of a regulatory mechanism that shifts between
dierent metabolic regimes of the regulated sub-system.
Far from being a novel concept, or a mere speculative construct, regulation is a
well-studied phenomenon in the biological sciences (e.g. Heinrich & Schuster 1996;
Fell, 1997; Tsokolov, 2010). Homeostasis, the ability of organisms to maintain sta-
bility in their internal variables, is generally the result of the activity of regulatory
sub-systems. This is what happens, for instance, in thermoregulation, or in glycemia
regulation. In this latter case, the pancreas acts as a regulatory mechanism that keeps
blood sugar levels stable, via the production of insulin and glucagon (see Bich et al.,
2020). Insulin, which is released by beta cells in the pancreas when they detect an
increase of blood sugar, contributes to reducing the levels of glucose by inducing the
liver to transform glucose into glycogen, and by promoting cellular intake of glucose
in muscle and adipose tissues. By contrast, glucagon, released by pancreatic alpha
cells, has the opposite eect of increasing the levels of blood glucose.
Glycemia regulation is just an illustrative example of biological regulation, among
many possible others (think, for instance, of the regulatory roles of the thyroid, or of
adrenaline). It should be stressed that biological regulation is a widespread phenom-
enon, which takes place in virtually every biological system, and on all levels of bio-
logical organization. Most aspects of the behavior of organisms and their interactions
with their environment are shaped to some extent by regulatory mechanisms. Thus,
we can nd regulation at the basic level of gene expression, as in the lac operon. The
movement of organisms in their environment is also usually controlled by regulatory
mechanisms, even in simple cases like bacterial chemotaxis. Bacteria like E. Coli can
direct their movements toward higher concentrations of some chemical substances
(attractants, e.g., nutrients), and away from others (repellents, e.g., toxic substances),
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Biological Purposes Beyond Natural Selection: Self-Regulation as a…
despite the fact that they only have two ways of moving: swimming in a straight
line (when rotating their agella counter-clockwise) and tumbling (when rotating
the agella clockwise). This can be achieved thanks to regulatory mechanisms that
detect variations in the environmental concentration of the relevant substances. When
a decrease in the concentration of attractants is sensed, the regulatory mechanism
induces a switch to clockwise rotation of the agella, as a result of which the bacte-
rium tumbles and changes its direction of movement at random. By contrast, when
the concentration of attractants increases, the bacterium keeps swimming in a straight
direction, which tends to get it closer to the source of attractants. The key components
in this regulatory system are a group of proteins called Che proteins. These proteins
are sensitive to the activation of the transmembrane receptors in charge of detect-
ing attractants/repellents, and also control the rotation of the agella (for details see
Eisenbach 2004; Wadhams & Armitage, 2004; Bich et al., 2016). This simple regula-
tory mechanism allows bacteria to eectively navigate chemical gradients in their
environment.
The behaviors of more sophisticated organisms are also typically subject to regu-
latory controls. Consider, for instance, a predator following the trajectory of its prey
(say, a cat hunting a mouse). In order to be able to track and replicate the changes
in direction of the prey, the movements of the predator will be guided by complex
regulatory mechanisms, involving its sensory organs and central nervous system.
In general, exible behaviors that show a high degree of adaptability require the
engagement of regulatory systems.
5 Biological Regulation as a Selective Process
Our proposal is that biological regulation constitutes a selective process. Regulatory
sub-systems tend to promote, retain, or reproduce certain behaviors in organisms,
and inhibit others. The activity of regulatory sub-systems results, therefore, in the
dierential reinforcement of the behavior of the organism regulated. This ts the
characterization of selection in terms of dierential reinforcement put forward above.
Thus, it makes sense to talk of regulatory selection. In this type of selection, the rel-
evant selective pressures are generated by the constraints imposed by the regulatory
sub-system on the behavior of the regulated system.
Remember that natural selection can be counted as a selective process because it
is driven by dierential reproduction, which is a type of dierential reinforcement.
Given the involvement of dierential reinforcement in biological regulation, it should
also be considered a form of selection, at least to the same extent that Darwinian nat-
ural selection is. Indeed, many examples of biological regulation can be described in
terms of dierential reproduction or repetition of certain behaviors. Let us go back to
bacterial chemotaxis. In this case, the regulatory sub-system promotes the repetition
of rotations of the agella that direct the bacterium towards high concentrations of an
attractant (e.g., nutrients), while movements that lead away from attractants tend to
be interrupted. In virtue of this dierential reinforcement of agella rotation, it makes
sense to think of chemotaxis as selective control of the movement of bacteria.
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J. González de Prado, C. Saborido
Other examples of biological regulation are perhaps better accounted for by
appealing to forms of selective reinforcement beyond dierential reproduction or
repetition. For instance, in glycemia regulation, the release of insulin intensies or
fosters the absorption of glucose into muscle, adipose and liver cells, whereas it
inhibits the production of glucose via glycogenolysis. In this example, rather than
dierential reproduction, we nd dierential stimulation and inhibition of certain cel-
lular processes. However, there is no reason to think that these further forms of rein-
forcement cannot give rise to selective processes—a point already made by Garson
(2017) when discussing selection via dierential retention. Nothing in the notion of
selection requires reproduction, as shown by paradigmatic examples like employee
selection (which, obviously, can take place without the candidate employees repro-
ducing). What is important is that there are selective pressures that lead to dierential
reinforcement, allowing for the distinction between items selected for and against.
Our claim is that biological regulation can be counted as a selective process, insofar
as it involves relevant forms of dierential reinforcement (even if this reinforcement
does not always take the shape of dierential reproduction). Indeed, the language of
selection is dicult to avoid when describing biological regulatory mechanisms. Just
to present one example, Bich, Mossio and Soto talk of regulation as the “capacity to
selectively modulate the rst-order self-maintaining regime” (2020: 9, our emphasis).
Once it is granted that biological regulation is a selective process, it follows from
selected-eects theories that regulation introduces teleological standards. As far as
regulatory teleology is concerned, the behaviors of organisms have as their purpose
producing those eects that explain their positive selection in some relevant regula-
tory regime. That is, purposive behavior is produced by traits that operate under the
causal control of regulatory sub-systems. The (positive or negative) reinforcement of
the regulated trait is explained causally by the pressures exerted by the relevant regu-
latory mechanism. Thus, the (positive and negative) reinforcing constraints generated
by the regulatory sub-system dene the teleological evaluative standards to which the
behavior of the regulated traits is subject (of course, there could be further purposes
introduced by other selective processes, for instance natural selection). In this way,
behaviors that tend to be inhibited by a regulatory mechanism (that is, behaviors that
tend to be regulated against) count as unsuccessful or inappropriate with respect to
the standards set by that regulatory mechanism. Our proposal, in sum, is that a trait
has the purpose of Φ if it has been reinforced by some regulatory mechanism because
of its tendency to bring about Φ (in the case of positive reinforcement), or because of
its tendency not to bring about Φ (in the case of negative reinforcement).
Take the example of chemotaxis. Movements that drive the bacterium away from
high concentrations of an attractant tend to be interrupted by the regulatory mecha-
nism, while movements towards high concentrations of the attractant tend to be con-
tinued. From the perspective of the regulatory mechanism governing chemotaxis,
therefore, the purpose of the bacterium’s movement is to go towards high concentra-
tions of attractants.
It should be stressed that although we are developing our characterization of regu-
lation in an organizational setting (following especially Bich et al., 2016), our pro-
posal is intended to be an instance of selected-eects theories. Other organizational
approaches account for purposes as contributions to the self-maintenance of organ-
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Biological Purposes Beyond Natural Selection: Self-Regulation as a…
isms (for instance, Mossio et al., 2009). In this type of view, regulation is not in itself
a source of teleology, but rather a way in which organisms become sensitive to the
teleological standards set by self-maintenance. In this paper, our approach is dierent
(even if compatible). Adopting the perspective of selective eects-theories, we argue
that regulation introduces purposes by virtue of being a selective process. At any
rate, the application of selected-eects theories to biological regulation vindicates
the claim by organizational theorists (see Mossio & Saborido 2016) that etiological
accounts of teleology do not need to appeal to the evolutionary history of traits.7
There can be etiological accounts that focus instead on the behavior of the current
organism, be it on the causal contribution of a trait to self-maintenance, as in standard
organizational proposals (Mossio Saborido and Moreno 2009), or on the eects that
explain the preservation or reinforcement of a trait under a regulatory regime.
The view that regulation is a source of teleology has all the explanatory virtues that
support selected-eects theories in general, and etiological-evolutionary accounts of
biological teleology in particular. First, it allows us to distinguish purposes from non-
purposive eects. The latter would be eects that are not under the control of regula-
tory mechanisms (to be sure, these eects could count as purposive with respect to
other, non-regulatory, selective processes).
Second, like other selected-eects theories, the view we are presenting accounts
for the normative dimension of teleology. More precisely, regulation, being a selec-
tive process, establishes evaluative standards, so that behaviors can be classied as
appropriate or inappropriate with respect to the standards introduced by the relevant
regulatory mechanism (remember, though, that this is just an internal evaluation rela-
tive to this regulatory process). In particular, it is perfectly possible for a trait to fail
to full the purpose conferred on it by some regulatory mechanism. This will hap-
pen when the trait behaves in ways that tend to be counteracted by the regulatory
mechanism (note that the regulatory mechanism may fail to counteract eectively
deviations in the regulated system).
Finally, the standards instituted by regulatory processes are teleological, because
they ground explanations of the presence of behaviors by appeal to their eects.
The fact that some behavior produces eects that tend to be promoted, rather than
inhibited, by a regulatory mechanism provides a relevant causal explanation for the
continued presence of the behavior. If the behavior had dierent eects, it would
probably have been inhibited by the regulatory mechanism controlling it. Therefore,
in order to explain why the behavior is being preserved (or somehow reinforced)
rather than inhibited by the (present) action of the relevant regulatory mechanism,
we have to appeal to the eects of that behavior. This is the characteristic structure of
teleological explanations. Note that teleological explanations do not always explain
7 Authors such as Garson (2019a) or Artiga and Martínez (2016) have argued that the organizational
approach is actually a version of etiological accounts. This is not disputed by defenders of organiza-
tional views, who argue that these views are perfectly compatible with the etiological characterization of
teleological explanations proposed by Wright (Saborido, 2014). Organizational approaches to functions
have not been developed as alternatives to etiological accounts in general, but to etiological-evolutionary
approaches in particular. The main dierence between these two approaches is not that one is etiological
and the other is not (both are), but that the organizational framework does not focus on evolutionary his-
tory (Mossio & Saborido, 2016).
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J. González de Prado, C. Saborido
the proliferation of a trait. Explanations that account for the preservation or continued
presence of a trait can be considered to be teleological, insofar as they do so by appeal
to the eects of that trait (see Mossio & Saborido 2016). A way of explaining the per-
sistence of a trait is to explain how it avoids being inhibited by a selective, regulatory
mechanisms to which the trait is subject.
Garson (2017) argues that, since the satisfaction of these three explanatory desid-
erata constitutes the best argument for etiological-evolutionary theories of teleol-
ogy, other selected-eects theories that satisfy those desiderata should, by parity, be
considered equally justied. He resorts to this argumentative strategy to vindicate
the idea that neural selection gives rise to functions; however, as we have seen, the
same strategy can be used to defend the view that biological regulation is a source
of teleology.
Moreover, our account captures two distinctive features of teleological behav-
iors: their persistence and plasticity—these features are highlighted, among others,
by Nagel (1979) and McShea (2012). The behavior of regulated systems is persis-
tent in the sense that deviations from the relevant behavioral trajectories tend to be
counteracted by the regulatory mechanisms. Additionally, regulatory mechanisms are
typically able to take the system back to the target behavioral trajectories from dier-
ent deviations (in the face of dierent perturbations), which confers plasticity to the
system’s behavior.
Thanks to the richness of regulatory behavior, our account is less prone to accu-
sations of liberality and overgeneralization than other selected-eects approaches.
Regulatory selection is not exposed to prominent counterexamples to selected-eects
theories discussed in the literature. For instance, Bedau (1991) considers a collec-
tion of clay crystals reproducing from crystal-seeds at dierent rates. Bedau argues
that these crystals meet the conditions for natural selection, even if we would not
attribute teleology to them. Garson (2017) discusses another example, proposed by
Kingsbury (2008), in which a group of beach rocks are eroded by the sea at dierent
rates depending on their hardness. This process can be seen as a form of dierential
retention, yet it does not involve teleology.8
None of these counterexamples aect our proposal, because they do not count as
instances of regulation. More specically, they do not feature a higher-order sub-
system exercising the relevant reinforcing pressures on a regulated system. Biologi-
cal self-regulation imposes quite stringent conditions, since it requires the presence
of higher-order regulatory mechanisms integrated in the self-maintaining organiza-
tion of the system. While this sophisticated form of organization is characteristic
of biological organisms, it is dicult to think of examples elsewhere in the natural
world, for instance in collections of rocks or clay crystals. Thus, counterexamples to
regulatory teleology are hard to come by. Even if regulation can be a phenomenon
with fuzzy limits, and dicult borderline cases are to be expected, a selected-eects
theory in terms of self-regulation does not seem to overgeneralize in problematic
ways. This is not surprising, given that the presence of a selector (in this case, in the
8 Garson’s (2017: 536–549) response to these purported counterexamples is to require that the relevant
forms of selection operate over a population of items engaged in tness-relevant interactions. The collec-
tion of beach rocks does not satisfy this condition.
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Biological Purposes Beyond Natural Selection: Self-Regulation as a…
shape of a regulatory subsystem) is one of the distinctive features of stereotypical
selective processes, which sets them apart from more basic forms of sorting or near
equilibrium behavior.
It is worth noting that, in addition to sharing the central explanatory virtues of
etiological-evolutionary theories, the view we are putting forward avoids some of
their problematic features. According to etiological-evolutionary theories, in order
to identify the purposes of some trait, we have to examine the evolutionary history
of its ancestors (we have to gure out what its ancestors were naturally selected for).
However, researchers are often interested in how functional traits contribute to the
organization of current organisms. That is, when attributing purposes, researchers
often want to study the actual organization and behavior of present-day organisms,
regardless of their evolutionary history. It can be argued, therefore, that in many cases
etiological-evolutionary theories do not t well with the explanatory interests that
underly the practice of ascribing purposes to organisms in biological research. This
is sometimes regarded as a problem for etiological theories (Amundson & Lauder,
1994; Christensen & Bickhard, 2002), and an advantage of alternative systemic
accounts of teleology, in which purposes are seen as current contributions of a trait to
the organisms’ organization (for instance, Cummins 1975; Craven 2001; Christensen
& Bickhard 2002; Mossio et al., 2009).
The view of biological teleology presented here does not face this problem.
Although, as a selected-eects theory, it is an etiological view, the focus is not on
the evolutionary history of the ancestors of current organisms, but rather on how
behaviors are produced under a certain regulatory regime in the actual organisms
studied. In self-regulation, the relevant selective processes operate on the very same
organisms to which the resulting purposes are attributed. Regulatory mechanisms
shape the behavior of the system they regulate. In this way, the selective pressures
created by regulatory mechanisms will be directly reected by the behavior of the
regulated system. Thus, the attribution of purposes arising from biological regulation
will be immediately relevant for explaining the current behavior of organisms. Note
as well that regulation is not a marginal phenomenon, aecting only a narrow range
of behaviors. Quite the opposite, regulation is pervasive in biology. Indeed, regula-
tory processes are crucial to explain phenomena as important as functional integra-
tion, the emergence of the complexity of organic bauplans, and even the origins of
cognition (Bich, 2018). The idea that regulation is a source of biological teleology
has, therefore, great explanatory potential.
All we have said is compatible with granting that the purposes introduced by natu-
ral selection can play a signicant role in scientic explanations (for instance, if
one wants to investigate the origin and evolution of some trait). The considerations
in this paper lead naturally to a pluralistic selected-eects theory in which dierent
teleological standards are instituted by dierent selective processes, including natural
selection but also neural selection and, as we have argued, biological regulation (and
we have also not ruled out the possibility that purposes may be established in further
ways that do not involve selection). Depending on the interests of researchers on each
occasion, some of these selective processes will be more relevant than others.
It is to be expected that, in many cases, the purposes introduced by biological
regulation will align with those instituted by natural selection, since, in general, regu-
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J. González de Prado, C. Saborido
latory mechanisms have themselves been naturally selected. However, natural selec-
tion and regulation constitute distinct sources of teleology. The purposes generated
by biological regulation are independent of the evolutionary history of the organism
exhibiting the relevant regulatory mechanisms. Indeed, conicts and divergences
between these two sources of teleology are, in principle, possible. Imagine, as a
hypothetical example, bacteria whose chemoreceptors treat as attractants substances
that were not present in the environment where they evolved, and which are not ben-
ecial for their tness. In this case, bacterial chemotaxis would have a purpose (navi-
gating towards the non-benecial attractant) that clashes with evolutionary goals.
When these misalignments occur, the purposes associated with regulation remain
particularly useful for explaining the actual behavior of organisms and the way in
which they are expected to react to perturbations.9
Before concluding, let us mention a possible objection to our account. According
to the view we have presented, regulatory sub-systems set teleological standards for
the systems they regulate, but not for their own regulatory behavior (see Schroeder
2014: 122). It seems, therefore, that regulatory mechanisms are not, themselves, sub-
ject to teleological standards. So, it would not be possible to attribute purposes to
regulatory mechanisms, and to identify cases of regulatory malfunction (at least, if
we are focusing just on teleological standards derived from regulation). This is cer-
tainly an implication of our view when dealing with systems controlled by a single
regulatory mechanism. However, in suciently sophisticated organisms, regulatory
processes may be themselves regulated by further regulatory mechanisms, creating
a complex hierarchical network of regulatory sub-systems (Bich et al., 2020: 10).
This makes room for the possibility that a regulatory mechanism malfunctions with
respect to the standards instituted by another regulatory mechanism that controls its
behavior. Although the purposes arising from regulation are ultimately established
by the regulatory activity of the organism, particular regulatory mechanisms can be
subject to teleological standards set by other parts of the regulatory network. In any
case, we leave the study of how dierent regulatory mechanisms are integrated hier-
archically in organisms for another occasion.
Funding Open Access funding provided thanks to the CRUE-CSIC agreement with Springer Nature. This
work has been supported by the Spanish Government research projects APID2021-128835NB-I00 and
PID2021-123938NBI00.
Open Access This article is licensed under a Creative Commons Attribution 4.0 International License,
which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long
as you give appropriate credit to the original author(s) and the source, provide a link to the Creative
Commons licence, and indicate if changes were made. The images or other third party material in this
article are included in the article’s Creative Commons licence, unless indicated otherwise in a credit line
to the material. If material is not included in the article’s Creative Commons licence and your intended use
is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission
directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/
licenses/by/4.0/.
9 Fagerberg (2022) criticises Garson’s generalised theory, arguing that it counts some traits as both func-
tional and dysfunctional. We do not nd this to be problematic, but rather an interesting implication of the
view. The important point is that the dierent, sometimes conicting, purposes are introduced by dierent
selective processes. As far as each selective process is concerned, the purpose of the trait is well dened.
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Biological Purposes Beyond Natural Selection: Self-Regulation as a…
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