Choices by Organisms On The Role of Free

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Biological Journal of the Linnean Society, 2023, 139, 555–562. With 2 figures.

Choices by organisms: on the role of freedom in


behaviour and evolution

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KALEVI KULL*
Department of Semiotics, University of Tartu, Jakobi St. 2, 51005 Tartu, Estonia

Received 16 January 2022; revised 31 May 2022; accepted for publication 16 June 2022

Neo-Darwinian biology has demonstrated that it is possible to construct a theory of life that excludes the role of
organisms’ free choice. In a richer theory, the latter as a possibility needs to be taken into account. For that purpose,
it is necessary to introduce the biological concept of choice, analyse its structure and roles, and consider some
implications for biological theory. It is argued here that the conditions for free choice emerge together with umwelt—
the space of synchronous options. Basically, choice does not require purpose. This leads to the conclusion that freedom
is an attribute of life.

ADDITIONAL KEYWORDS: biosemiotics – free choice – habit – knowledge – possibilities – recognition – theory
of evolution – umwelt.

INTRODUCTION conditions and structure of organismic choice. The


third section presents some implications from the
External stimuli [...] approach the animal in understanding of choice for biological theory.
the form of questions (Jakob von Uexküll, 1992
[1934]: 323).
Life is not a sequence of cause and effect, but
ON THE CONCEPT OF CHOICE IN BIOLOGY
choice (Viktor von Weizsäcker, 1940: 126).
Choice is a strange concept because it fundamentally
Expressions like ‘habitat choice’, ‘choice of food’, ‘choice includes a feature, due to which it has been largely
of partner’ and ‘choice of direction of movement’ are avoided in the natural science: indeterminacy. Choice
rather common in biological discourse. The process of is the phenomenon that is situated precisely at the
choice itself, which presupposes some freedom of doing border between physics and semiotics, between the
as such in order to be identified as choice and not as a natural sciences and the sciences of mind, between
random or a deterministic process, is however seldom the study of causes and study of freedom which means
explicitly defined and analysed in biological theory. it will also allow us to connect these two areas of study,
I suspect that this situation is due to the use of a that now largely belong to separate academic cultures.
loosely defined anthropomorphic concept of choice, In biology, the problem of choice is fundamentally
the relevance of which for other species is unclear. related to the theory of evolution, and is at the heart
Introducing a non-anthropomorphic understanding of of a contemporary remarkable paradigm change in
free choice, its widespread existence in the living world the understanding of evolution. Namely, the contrast
can be observed. This is the basic hypothesis of this between the neo-Darwinian and the post-Darwinian
study. If true, the implications for biological theory will paradigms includes some oppositions which concern
be discernible. the role of agentive activity: (1) either organisms are
This article is organized as follows. The first section replication-devices, or they are interpretive agents; (2)
briefly describes the situation in the use of the concept either environment selects, or organisms choose; (3)
of choice in biological theories. The second section either the main factor of evolution is a passive natural
introduces the key definitions, and describes the selection, or an active organic fitting.
These are not necessarily alternatives. Every
*E-mail: [email protected] experienced biologist certainly understands that

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All rights reserved. For permissions, please e-mail: [email protected]
556 K. KULL

evolution is based on both. But this does not resolve principle of parsimony, was later used to reduce
the problem, because then the question will be ‘in the enthusiasm to apply psychological concepts in
what relationship?’ In order to answer this, the precise biology [see also a comment on Morgan in Ginsburg
definitions of terms (including choice) will be required. & Jablonka (2019: 197)]. Behaviourism refused to use
A focus on the concept of choice in an evolutionary psychological concepts, and the ethological tradition
context appeared at the end of the 19 th century. could mostly do without a direct reference to the
George Romanes, in the first chapter of his Mental subjective experience of animals. Accordingly, the
evolution in animals (‘The criterion of mind’) writes: study of human choice behaviour has developed along

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‘What activities of an organism are to be taken as very different lines, compared to the study of choice
indicative of consciousness? The answer that comes in animals (see Staddon, 2016: 270). In the studies of
most readily is, — All activities that are indicative of animal choice, what has mainly been meant by choice
Choice; wherever we see a living organism apparently is preference. This is common in the studies of animal
exerting intentional choice, we may infer that it is learning, which focus on the changes of preference
conscious choice, and, therefore, that the organism under various conditions (e.g. reinforcement) and in
has a mind’ (Romanes, 1883: 17). And he concludes: Edward Thorndike’s law of effect.
‘the distinctive element of mind is consciousness, the In her critique of Darwinism, Lynn Margulis
test of consciousness is the presence of choice, and emphasized that: ‘organisms choose’ (Margulis &
the evidence of choice is the antecedent uncertainty Sagan, 1995: 222). Sagan (2021: 6) adds:
of adjustive action between two or more alternatives’
The prototactical associations and living
[(Romanes, 1883: 18); see also comments on Romanes
choices made by organisms, always members of
in Maher (2021) and Ginsburg & Jablonka (2019: 195)].
communities in identifiable ecosystems, may lead
Charles Darwin (1871), when famously introducing
directly to evolutionary consequences: origins of
the term ‘sexual selection’, not only speaks also about
new species. [...] But if we accept Darwin’s sexual
‘sexual choice’ (indeed referring to choice in a direct
selection, and most biologists have, why then do
sense), but sees choice as the main factor of sexual
we not also consider other kinds of organism-
selection: ‘Hence in these classes, such as the Protozoa,
level choice-based selection? Prototactical living
Cœlenterata, Echinodermata, Scolecida, true secondary
beings actively decide with which other life
sexual characters do not occur; and this fact agrees
forms to associate. Which ought they try to eat?
with the belief that such characters in the higher
With whom ought they band for protection? Who
classes have been acquired through sexual selection,
might they inhabit for shelter? These “artificial
which depends on the will, desires, and choice of either
selection” decisions of the living have evolutionary
sex’ (Darwin, 1871: 321, emphasis added).
consequences that are not to be confused with the
The roots of the current post-Darwinian (as different
“mechanical” – such as the inanimate interactions
from the neo-Darwinian) paradigm go back at least to
in random or deterministic behaviours of billiard
the concept of organic selection, as defined by James
balls, moving electrons, other elementary particles
Mark Baldwin (1896: 444): ‘Whatever the method of
or solid bodies.
doing this may be, we may simply, at this point, claim
the law of use and disuse, as applicable in ontogenetic The concept of prototaxis was introduced by Ivan
development, and apply the phrase “Organic Wallin, to denote an association more general than
Selection”, to the organism’s behavior in acquiring new symbiosis, involving any innate tendency of any
modes or modifications of adaptive function with its particular species, organism or cell to respond in a
influence of structure’. Baldwin does not use here the specific way (associative or dissociative) to any other
word ‘choice’, but his meaning of the ‘organic selection’ sort of species, organism or cell (Wallin, 1923). This
that is due to agent activity is close enough to it. As concurs with Janzen’s (1985) observation that species
Piaget (1971: 299) commented: ‘When Baldwin talked coexistence in ecosystems is not due to slow, long-term
of “organic selection”, it was still only a word and a co-evolution, but due to their ecological fitting, which
rather ambiguous word at that, for although external is a quick, real-time process. More generally, ecological
selection may proceed by eliminations and survival of fitting may include any relationship an organism
the fittest, any organic or internal selection is much establishes as a result of search and choice. Its general
more like a “choice” of a more or less active kind, which role in ecosystems has been rather well demonstrated
means that i[t] is, properly speaking, a regulation.’ (see, e.g. Araujo et al., 2015).
Conwy Lloyd Morgan had similar ideas, and Likewise, for instance, Williams (1994: 84) writes,
provided a criterion for choice as ‘an alteration or with a reference to Herrnstein (1970): ‘all behavior
modification of response in the light of individual is choice, in the sense that there are always
experience’ (Morgan, 1896: 265). However, his own alternatives other than the response measured by the
other principle, formulated as Morgan’s Canon or experimenter. Thus, the animal is always “deciding”

© 2022 The Linnean Society of London, Biological Journal of the Linnean Society, 2023, 139, 555–562
ORGANISMS’ CHOICE AND FREEDOM 557

which response to perform.’ In ethology, ‘free-choice literature, for example, it is most often discussed in the
conditions’ is used as one of the experimental settings context of neurobiology (e.g. Ansermet & Magistretti,
(Graham et al., 2018). 2007; Lee et al., 2012).
Thus, the idea that organisms make choices is not My own attention has been on the role of choice
new. Young (1987: 148) mentions: ‘The realization that in the processes of semiosis and interpretation, from
choice is a property of all living things gives us great a biosemiotic perspective. Our analysis of meaning-
help in understanding the world and our place in it’. making or semiosis has shown that choice is its
However, it is important to notice that animal choice is necessary component (Kull, 2015, 2018a, b).

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sometimes understood as being strictly computational What can be concluded from this brief review is that
(e.g. Real, 1991), which would exclude free choice. the concept of choice has a place in biological theory;
More recently, the natural history of freedom has however, it is not well explicated and accordingly the
received attention in biology (Ho, 1996; Heilinger, content of this concept varies. In the next section
2007; Hoffmeyer, 2010). Yudanin (2020) provides a I attempt to clarify the meaning of the ‘choice process’.
rich review of the studies of animal choice. One of the
conditions for choice he points out is self-determination
(Yudanin, 2020: 64).
THE WORKING OF CHOICE
Among the attempts to describe the mechanism of
choice, an account by Noble & Noble (2018) builds it Here the aim is to formulate the necessary and sufficient
to the harnessing of stochasticity. Their description conditions for a process to be qualified as choice—by
divides the process of choice into five stages (Noble & which is meant an indeterminate non-random action
Noble, 2018: 3): (Kull, in press). The latter is emphasized because
the term ‘choice’ is used by some authors for certain
1. A challenge has occurred—as a puzzle analogous to
algorithms of determinate actions. A little scheme
the form of a template for which a match is needed.
might be of some help in which a random process is
2. The organism searches amongst existing stored
compared with choice (Figs 1, 2).
possible fits to the problem template.
A random process can be modelled as a landscape
3. The organism activates stochastic processes within
with branching channels, upon which a ball is rolling
itself to generate further possible new solutions.
downwards (similarly to the classical picture of
4. The organism returns to direct control at this
an epigenetic landscape as a representation of an
stage, which is to compare what is thrown up by
aspect in developmental differentiation, provided by
the stochastic process with the problem template to
Waddington, Fig. 1). The pathway which the ball takes
determine what fits.
at any given branching point is random in this situation,
5. Implementation of the discovered action to solve
if no additional conditions are applied. Although, for
the problem.
the ball, there may be a short-term unstable moment
Delafield-Butt (2021: 80) points out the anticipatory at the branching point of the channel, the ball does
aspect of choice: ‘Anticipatory motor control with its not ‘see’ both available channels. The two pathways
sense of possible futures affords the organism choice. are not ‘options’ for the ball, these are options only for
It is the pivot on which sits immediate experience and an external observer. The ball simply follows one of
agency. In mammals, this pivot rests on a tripartite of these pathways by chance due to micro-determination.
information integrated from the exteroceptive senses
of the outside world (i.e. sight, sound, touch, taste),
interoceptive senses of visceral and vital physiological
need (e.g. hunger, thirst, thermoregulation), altogether
with proprioceptive senses of the body-in-motion. [ … ]
Agent action of this kind is common to all vertebrates,
and the basic system of “sense, evaluate, choose”
common to all organisms.’
Since consumer choices are an important focus
in economic theories, some work which links the
biological and economic models of choice was carried
out (Akçay, 2015). If related to humans, choice is seen
as dependent on free will. Literature on free will is vast,
but free will is seldom seen as a biological problem to
be studied and solved by biologists—which it certainly
is, particularly as the study of capacity to choose (cf. Figure 1. A simple model, in which the selection of
Brembs, 2011). In the contemporary philosophical pathway is random (modified from Waddington, 1957: 29).

© 2022 The Linnean Society of London, Biological Journal of the Linnean Society, 2023, 139, 555–562
558 K. KULL

the finite Now. This is a tiny period in which what


is presented can be synchronous, can be seen or felt
together, without determined sequence. Simultaneity
of options is an elementary and fundamental condition
of freedom in the context of choice. Collecting
information about options can be sequential, while for
the options to work as possibilities free to choose from,
these should be represented together. Choice itself

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creates the conditions for choice.
From this description, one can notice that in such
cases of free choice, there still can be some additional
limitations, restrictions, preferences, motivations and
intentions to choose, for instance, the ‘left path’ instead
of ‘right path’, without such motivation being the
Figure 2. Agent’s choice between options as an elementary strict determinator of the resulting selection. Such
interpretation. preferences or motivations work as memory traces
from earlier behaviour, making some paths easier to
The branches of the pathways that were not taken by use. This means that preference or motivation does not
the rolling ball do not exist for the ball, nor have they remove the condition of the choice being free, so long as
ever. Which means its turns are not choices, no matter the preference or motivation are not algorithms that
how many ‘options’ may appear, to the observer, to be must by necessity be followed.
present in the landscape. Accordingly motivation or preference is not the
Thus, if rolling downwards, then at the branching opposite of arbitrarity or free choice, but one of its
points, at bifurcations, there is not a choice—it is rather important features. An arbitrary choice, or free choice,
a case of pure chance. But change your perspective may include more motivation or less motivation in
and assume that the closer end here is the higher one some direction, but so long as the motivation is not a
(Fig. 2). If climbing upwards, i.e. when doing work, necessity, and so long as there is still the capacity to
as any agent does, the situation could be different behave against the motivation, against an acquired
than in the earlier picture. Still, if the climber has no preference, the choice is free, and it is still a choice.
representation of possibilities, then the situation is Thus, we arrive at a very important and fundamental
not much different from the previous one—the path point for biology—the non-anthropomorphic
taken will be random. But if the climber is not ‘blind’, description of the structure of free choice (which is
which means if it has receptors which can provide a also the key to the structure of semiosis or meaning-
sign in advance about the two or more possibilities, making in the pre-linguistic realm).
then the choice can be made. All that is needed is that The necessary and sufficient conditions for free
both paths are represented simultaneously. Obviously, choice include: work, simultaneous availability
at least two receptors are required for creating such a (i.e. ‘sensing’ the existence) of alternative options
representation. (possibilities) and indeterminacy, i.e. the absence of
In the case of choice, in place of the passively rolling a controlling force or algorithm which would make
ball, we now have an active agent that can move one of the options necessary to pick [see also a similar
itself, and that can somehow register or ‘sense’ the approach in Laskey (2018)].
different pathways that are now available before it. First: the condition of the need for work. In cases
The existence of different pathways upon which to where work is being applied, the situation of bifurcating
go—i.e. the possibilities, the options—being ‘sensed’ by pathways is different than it is when small stochastic
the agent, now have to be actively chosen. Such choice, fluctuations alone determine the continuation of the
in order to be a choice (which means not determined process (i.e. the path taken) at the bifurcation point.
by a randomizer or any other determinator), means, by In the cases of ‘choice’, the energy that is directed
definition, that there is no force or pre-given algorithm towards pushing the behaviour is larger than the
or rule that has to be necessarily followed. Such non- energy of fluctuations, and is therefore decisive for
algorithmicity is necessary in order for the event to the selection of the pathway. The randomness of
be a choice, and indeed makes it a free choice. This fluctuations, therefore, is not what plays the principal
meaning of ‘free’ here, however, requires some further role in path-selection when an agent’s work is applied
discussion. to the process.
Freedom emerges from an additional feature Second: optionality in the sense of the simultaneous
that is implied—from simultaneity of represented availability of possibilities, which can be described as
possibilities, which means a brief moment of present, simultaneously representing the existence of more

© 2022 The Linnean Society of London, Biological Journal of the Linnean Society, 2023, 139, 555–562
ORGANISMS’ CHOICE AND FREEDOM 559

than one pathway, and which requires the co-existence between these two types is not strict, and both
of more than one elementary receptor in the same provide knowledge (sensu lato) to organisms via their
agent. Assuming that one elementary receptor can consequences.
detect the existence of something only once at a time,
at least two receptors are required in order to detect
two simultaneously existing elements at the same
IMPLICATIONS
time, i.e. to create non-sequentionality necessary for
indeterminacy. If the two are detectable only one by Choice, according to the definition exemplified in

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one, then they are not true options that must now be the previous section, requires the simultaneous
chosen from. representation of something more than one, something
A receptor can be defined functionally as an organ multiple (characterized as possibilities—habits that can
or organelle that is connected to an actor via habit. If be executed). This implies that the existence of choice is
an agent has multiple receptors, linking to multiple coextensive with subjective time and space—i.e. umwelt,
habits, then the situation of incompatibility may as defined by von Uexküll (1928). Consequently, all
occur. The state of incompatibility is the state of an organisms who can make a choice have an umwelt. And
agent in which two (or more) habitual connections since umwelt is the space of possibilities, precisely those
(i.e. habits) are simultaneously excited yet cannot be who have umwelt can choose.
simultaneously executed. The existence of umwelt and choice in particular
What happens in the coexistence of incompatible organisms should be established by detailed research.
habits is that a new dimension becomes created. This A liminal example often mentioned in this respect is
new dimension is the perceptual simultaneity of more plant heliotropism. If heliotropism is not an acquired
than one—i.e. the creation of a map, the model of space, feature, then it cannot be a plant’s choice. If heliotropic
the subjective space (if space is defined as something movement has been acquired, but works completely
that consists of more than one point). The realization deterministically, then it also cannot be a choice.
of, and access to, this ‘space’ or ‘representation’, is the However, the hypothesis that the choice conditions are
prerequisite of choice. met during the period of acquisition of such a reaction
This new dimension is relational, it is logical, not to sunlight, is quite plausible.
physical. The incompatibility that characterizes The situation of choice requires the simultaneity
options cannot be based on physical necessity, which is (synchronicity) of options. An organism can only have
why it requires habits, since habits are the carriers of the freedom to make a decision if several possibilities
acquired relations (which by definition are not based are presented and available at the same time. From the
on physical necessity). physicalist point of view, this may seem impossible—
Choice requires simultaneity of options. If options time is continuous and there is at least a microscopic
are not represented simultaneously, they are not difference between events, thus everything is
options. Simultaneity is possible—as there exists the sequential. However, from an organism’s point of view,
specious present, the subjective now. The existence perceived time has a certain finite interval which is
of subjective present in animals has been confirmed interpreted as present. From the physiological point of
[see references in Kull (2018b)]. Such simultaneity is view, the specious present (Varela, 1999) appears due to
illusionary in the same manner as logical relations are the finite relaxation times of coupled functional cycles.
illusionary, or as subjectivity is illusionary. In other In this case, before a functional cycle can culminate in
words, choice creates subjective reality. action, there is another functional cycle that would lead
At this point a slight further generalization is to an alternative action, and if the actor is the same,
possible. ‘Choice’ can be defined as what happens in then there is an incompatibility between the operations,
the situation where there are possibilities present hence there occurs a true situation of indeterminate
(this being equivalent to the condition of semiosis, or choice. Moreover, the moment of choice is related to
interpretation). Possibility, by definition, cannot be an organism’s meaning making. This can be seen as
single. The field of possibilities provides the condition the fundamental point that connects phenomenology,
of choices; moreover, it makes the choices inevitable. semiotics and physiology, where these three converge.
And there are two main types of choices—strongly A structure that can support the conditions for choice
motivated [for instance recognitions, readings or is paired receptor organs. In particular, this is the case
measurements in the broad sense of Pattee (2007)] in organisms with bilateral symmetry—as in the clade
and weakly motivated (commonly called ‘free’) Bilateria among Eumetazoa—which, of course, does
choices. A choice is called strongly motivated if bias not exclude the existence of necessary integration
or preference towards one of the possibilities is strong. between receptors in the organisms of other forms and
A choice is called weakly motivated if bias towards taxa, in principle even including unicellular organisms
any of the possibilities is not strong. The boundary with membrane receptors.

© 2022 The Linnean Society of London, Biological Journal of the Linnean Society, 2023, 139, 555–562
560 K. KULL

According to our conceptualization here, possibilities and there may be several alternative habits which fit
are habits. An appropriate definition of habit can be one the same situation. Thus, there is no ultimate purpose
from Gardner (2015: 277): ‘habit is a process by which like survival or anything else that would determine
a stimulus generates an impulse to act as a result of a the choices.
learned stimulus-response association’ while reading Work alone is insufficient for choice or for
it as compatible with Peirce’s concept of habit (West intentionality. For example, various engines (including
& Anderson, 2016). As this definition says, habit is chemical ones) do work without any aboutness
what mediates [‘generates an impulse towards action’ or purpose. An additional necessary condition is

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(Gardner, 2015: 280)]. What can be added here is that perception of possibilities together with choice making,
habit, in order to be a habit, should not be completely but even this is insufficient for purposefulness. Simple
automatic or deterministic; a principal feature of habit playful behaviour includes choices with no purpose
is at least a minimal freedom of it not to be followed— as such.
only then can a habit be a possibility. Together with the subjective world (umwelt), the
Habits—the possibilities with strong preferences— existence of choice implies the existence of meaning
as acquired and as the results of learning, carry making, i.e. semiosis. This is because umwelt consists
in themselves an anticipation about what may of the relations in which the organism is a part; umwelt
happen. This is because the main types of learning— is the organism’s interpretative world. Interpretation,
imprinting, conditioning and imitating—that create which is semiosis by definition, presupposes the
(correspondingly) either iconic, indexical or emonic potential of alternative interpretations—which
relations. Symbolic relations, as acquired by convention, is choice.
are an exception, as they may not correspond to any One of Karl Popper’s writings was entitled as ‘A world
earlier regularity [for terminology, see Kull (2020: 15)]. without natural selection but with problem solving’
These relations, other than symbolic, acquire (we can (Popper, 2014). The ‘problem solving’, if not just a
also say ‘they model’) the relationships or regularities metaphor for certain deterministic processes, assumes
that exist around the organism. This is why Peirce can indeterminacy, motivation and choice, referring
say that ‘knowledge is habit’ (Peirce, 1906: CP 4.531). to some fundamental freedom in the behaviour of
Since possibilities are habits, and habits embed in organisms. A problem, or a problem situation, in the
themselves the local regularities, every choice includes general sense, is a situation in which the behaviour
an anticipation as based on these regularities. has to be indeterminate, and a choice making has to
Choice (as described) does not require purpose. be possible. This occurs if coupled functional systems
Choice happens because organisms face alternatives face mutual incompatibility. For instance, perceptions
and have the capacity to take one. Consequently, from two sense organs order the opposite actions of the
purpose itself is free. Why organisms choose is not same effector. Or, if a perception orders two effectors
because of some purpose, but because of umwelt, their that lead to opposite actions. This is a situation in
now, which is the field of alternative possibilities, of which behaviour is not fully determined by any rule,
options; that is what behaviour is—to do this or that. i.e. when an organism is a little bit confused. This
Accordingly, the formation of purpose is secondary in kind of indeterminacy appears precisely together
relation to choice. with umwelt.
In the context of biological debate between teleology Finally, only a process based on choice and
and teleonomy, the understanding described above can learning, i.e. on semiosis or interpretation, provides
be seen as a ‘third way’. This is neither teleology nor an adaptiveness profoundly independent of natural
teleonomy but semiotics. There is free choice but not selection. In a more detailed analysis, six principal
necessarily any purpose. Purpose as such presupposes types of transformations in living systems can be
freedom, because the concept of purpose is defined as distinguished (Table 1).
not applicable to deterministic or random processes. Thus, there are three independent sources of
The directionality in behaviour can be a consequence innovation: mutation, environmental influence and
of preferences to which choices contribute. The choice. Mutation initiates genetic modification,
preferences in choice making are the features of habits, environmental influence modifies physiological

Table 1. The principal types of transformations in living systems (see text)

Mutational Plastic Interpretative, meaningful

Neutral Random drift Self-organizational shift Weakly motivated choice


Adaptive Natural selection Homeostatic adjustment Strongly motivated choice

© 2022 The Linnean Society of London, Biological Journal of the Linnean Society, 2023, 139, 555–562
ORGANISMS’ CHOICE AND FREEDOM 561

processes, and choice modifies habits. One should behaviour is organized by preferences—i.e. by the
notice that self-organization and self-assembly as possibilities which are biased and habit based.
the processes largely responsible for plasticity but Innovative adaptivity is greatest where there is
also quite common in non-living systems, may not freedom.
be teleonomically accommodative or functionally Freedom is not a quality of being alive, it is what
adaptive. There are also three independent test defines being alive. Life is choice making, choice
processes for congruence or functionality, i.e. processes making is freedom. Freedom is an attribute of life. It is
that can turn the modification adaptive: natural here that the main problem of biology for our era lies.

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selection, homeostatic feedback and problem solving
via choice [regarding the last, cf. Gregory (1980) on
perception as hypothesis]. These are very different
ACKNOWLEDGEMENTS
processes, but all may result in adaptive behaviour.
Since all these can to some extent be heritable, either This article is a contribution to a special issue on
genetically, epigenetically or ecologically, it should be Teleonomy in Living Systems, guest edited by Richard
concluded that there exists at least three independent I.  Vane-Wright and Peter A.  Corning, based on a
processes of evolutionary adaptation. Linnean Society meeting held on 28–29 June 2021.
Whether the indeterminacy based on I thank Dick and Peter for initiating and commenting
incompatibility of behavioural habits, and its solution on this paper, and also Don Favareau and anonymous
via choice, exists only in animals with a nervous reviewers for suggestions that improved this work.
system, or can be identified also in other organisms, I also thank the Estonian Research Council grant
including at least some types of cells, remains to PRG314 for support.
be carefully studied. Where it exists, it provides a
process for end-directed changes—given that choices
are based on alternative habits. This is a process of DATA AVAILABILITY
internal teleology, as referred to by Woodger [who This article is based on information available in the
added that ‘it would doubtless be desirable in biology published literature.
to avoid the term “teleology” if a suitable substitute
could be found’ (Woodger, 1929: 453)], and can be
identified with a kind of agency. Plasticity, excitable
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