Agencies

Mathilde Tahar

► To cite this version:

Mathilde Tahar. Agencies. 2024. hal-04911458v1

HAL Id: hal-04911458

https://hal.science/hal-04911458v1

Preprint submitted on 24 Jan 2025 (v1), last revised 19 Nov 2025 (v2)

HAL is a multi-disciplinary open access
archive for the deposit and dissemination of sci-
entific research documents, whether they are pub-
lished or not. The documents may come from
teaching and research institutions in France or
abroad, or from public or private research centers.

L'archive ouverte pluridisciplinaire HAL, est
destinée au dépôt et à la diffusion de documents
scientifiques de niveau recherche, publiés ou non,
émanant des établissements d'enseignement et de
recherche français ou étrangers, des laboratoires
publics ou privés.

<figure><img src="image_1.png" /></figure>

Distributed under a Creative Commons Attribution 4.0 International License
“Individual playful memory”: making
sense of biological agency

**Abstract.** In recent decades, agency has become a prominent concept for conceiving of biological phenomena, both in philosophy and in biology. The concept aims to capture the spontaneity of organisms and illuminate processes overlooked by the previously dominant reductionist framework. However, its definition varies widely across and within disciplines, whether it is employed to understand development, organisation, or evolution, or applied to genes, cells, organisms, or even natural selection. This ambiguity hinders distinguishing its appropriate from inappropriate uses and establishing a cohesive theoretical basis for its various manifestations. This paper reviews the existing literature on agency from philosophy, evolutionary theory, developmental biology, and animal studies. It identifies three core capacities that unify the concept of agency: Individuality, Playful flexibility, and Memory (IPM). These capacities interact in diverse ways, with their influence varying depending on the process or context. These quantitative differences give rise to qualitatively distinct forms of agency: autonomous organisation, goal-directed choice, and inventiveness. Drawing on the IPM framework, this paper explores the relationships between these forms of agency, and argues that this approach can rationalise and formalise our understanding of agency while accommodating its diverse expressions.

**Keywords:** agency; organisation; goal-directedness; inventiveness; evolution; behavioural ecology; animal studies
“Individual playful memory”: making
sense of biological agency

Over the past decades, agency has emerged as a central concept in philosophy of biology, and in biological theory. This concept aims to capture the spontaneity of living beings and shed light on some processes underdetermined by the once-prevailing reductionist perspective. Nonetheless, the definition of biological agency is far from uniform across and within disciplines and varies according to its context and application: whether it is used in discussions on development, organisation, evolution, or in reference to genes, cells, organisms, or even natural selection. Additionally, it is shared by researchers across a broad spectrum of disciplines and serves varied purposes. In some cases, the concept is merely metaphorical, while in others it plays an operational role, being incorporated into biological theories or even acquiring ontological significance. Such a multiplicity of definitions, usually opportunistically produced to suit the needs of the argument at hand, seems to indicate the existence of multiple concepts of agency linked only by their shared terminology. Each concept has a narrow scope and a precise application; conflating its different uses can lead to confusion and risks the concept becoming widely applicable yet operationally meaningless.

However, the lack of a unified synthesis presents several issues. First, despite the use of the same word indicating a commonality of meaning, this meaning remains globally undefined. As a result, the term is often divorced from its general intuitive signification. Second, without establishing common criteria for agency, it becomes challenging to differentiate between the erroneous and justified uses of the concept. Thus, its operational value is diminished. Finally, in the absence of a common definition, agency functions as an umbrella term shared by several disciplines, but whose equivocality impedes the integration of knowledge from different fields. This paper seeks to address these issues by synthesising the disparate definitions of agency in order to identify the unifying criteria behind this diversity. Through a review of the literature, three main categories of agency are identified: organisation, choice, and invention. It is then argued that these distinct types of agency share a set of core characteristics: Individuality, Playful flexibility, and Memory (IPM). Finally, this paper demonstrates how these characteristics help to address key challenges, such as elucidating the commonalities among diverse forms of agency, enhancing the understanding of their distinctions, and clarifying their conceptual and biological articulation.

Agency or agencies?

Agency in biology

The term agency has traditionally been used in the social sciences and humanities to reflect on the rationality or irrationality, moral responsibility or irresponsibility of agents, typically human (Shapiro 2005; Emirbayer and Mische 1998). However, over the past decades, agency or at
least a form of “agential thinking” (Godfrey-Smith 2009) has increasingly permeated the field of biology. It is true that biologists do not frequently engage with the concept of agency itself. And, when they do, it seldom receives in-depth theoretical scrutiny. Nevertheless, even if agency is not a primary object of biological study, it is still present in three notable ways. First, it is used as a rhetorical device to communicate core biological concepts, especially within evolutionary studies. In this case, agency serves as a metaphor, but also reveals an actual agential mode of thinking at work in the theory. Second, ethologists invoke animals’ agency to reflect on scientific practices. Here, agency is explicitly advanced as a concept to be incorporated into biological methodologies. Finally, it is sometimes used as a concept that could actually help rebuild or correct theoretical models. The definition of agency is then generally borrowed from philosophical discourse or jointly developed in collaboration with philosophers.

## Agential thinking in evolution

An agential perspective is often used by evolutionists to convey their findings. In these instances, agency rarely becomes the subject of deep theorisation. More often, it assumes a metaphorical role: it is *as if* the subject of study were acting as an agent. Here, agency is understood teleologically: one speaks as if the object under study were an agent because it seems to be goal-directed. This approach to agency has been applied to a wide range of biological processes and entities. For instance, the action of natural selection is often described in agent-like terms, as if nature itself were striving to optimise for the most suitable traits (Dawkins 1989; Dennett 1995). Similarly, genes are sometimes depicted as metaphorical agent, pursuing their own interests, most famously in Dawkins’ *Selfish gene* (Dawkins 1989). Aligning with a more intuitive understanding of agency, the concept is also widely applied to organisms, which are often portrayed as intentional, even rational, agents. For example, it is said that the lion kills cubs that are not its own, because it *realises* that the mother won’t have any sexual activity while lactating (and actually does not ovulate), and it *wants* to increase its chances of reproduction (infanticide interrupts lactation in mothers and allows ovulation to resume).
In all these cases, agency functions as a practical, if anthropomorphic, metaphor that makes biological discourse more accessible. It offers a pedagogical framework for discussing biological functions. Agency here is merely expressing the adaptationism prevalent in evolutionary explanations, and according to which all traits and behaviours are contributing to the overall fitness of the whole (Okasha 2018). Yet, this form of agential thinking has been strongly criticised as being, in most situations, a misapplication of the concept, especially when applied to natural selection, and genes, and often even to organisms. While it seems useful in select scenarios (especially when the agents are high-cognitive organisms), it usually leads to more confusion than clarity (Okasha 2018; Tahar 2023).

## Agency for biological practice

In contrast, there are cases where agency is adopted as a concept with true operational implications in scientific practices, shedding light on genuine attributes of biological entities. This is particularly evident in behavioural ecology (also labelled ethology), where agency is employed to critically assess the scientific validity and ethical considerations in experiments. Agency then expresses animals’ subjectivity and spontaneity, prompting us to reconsider the ethical implications of our scientific practices. A particularly illustrative example of these operational implications is the controversy that arose following the publication of a widely mediatised study where researchers engaged in a game of hide-and-seek with rats (Reinhold et al. 2019). Their experiment demonstrated the rats’ capacity for learning, active participation, and even initiating new behaviours. However, despite these demonstrations, the rats were ultimately euthanised to analyse the neural mechanisms underlying their behaviours. This
decision sparked considerable controversy, with several researchers advocating for the recognition of the rats' agency as a basis for ethical reconsideration, a recognition made necessary by the very study that killed them (Webb, Woodford, and Huchard 2020).

Typically, animal behaviours' studies will use the concept but not provide a detailed theory (e.g. Warkentin 2009; Webb, Woodford, and Huchard 2020). But some do outline criteria for identifying agentive behaviours in experiments or observations, encompassing the animals' natural behaviours, social interactions, and their responses to experimental conditions (McFarland and Hediger 2009; Špinka 2019; Sueur, Zanaz, and Pelé 2023). Agency then refers to the animals' ability to control an action (Couchman 2012; Kaneko and Tomonaga 2011), resist to some experimental settings (Sueur, Zanaz, and Pelé 2023), or even manifest creativity (McFarland and Hediger 2009; Špinka and Wemelsfelder 2011).

Importantly, recognising animal's agency has implications beyond ethical considerations. It is not merely a matter of acknowledging agency to respect animals; it also has significant scientific implications. For valid research, it is crucial to limit interference with the subjects under observation, while allowing their agency to express as freely as possible (Sueur, Zanaz, and Pelé 2023; Webb, Woodford, and Huchard 2019). Therefore, experimental designs should allow animals to express their agency, avoiding undue stress or overly restrictive conditions.

## Agency for theory

The concept of agency has also recently been used in theoretical biology to challenge the dominant reductionist framework. In this context, agency is addressed not merely as a pedagogical tool or an instrument for practical and ethical purposes, but as a fully-fledged concept to be integrated in theoretical models. Its application in biological theory is particularly prominent in two main areas: the study of biological systems where it highlights the specificity of biological organisations (Nadolski and Moczek 2023; Newman 2023; Watson 2023), and in evolutionary theory (Kauffman and Clayton 2006; Laland, Odling-Smee, and Feldman 2019). In the latter, agency designates the active role played by organisms in ecological dynamics and selective pressures. As such, it is employed within the framework of the extended evolutionary synthesis, which challenges the exclusively selectionist narrative by introducing new processes, such as niche construction (Kylafis and Loreau 2008).

In theoretical biology, a great deal of discussion is devoted to the definition of agency, usually based on philosophical research. These definitions tend to closely align with those found in the philosophy of biology, with biologists either directly adopting these philosophical definitions (Nadolski and Moczek 2023; Newman 2023) or collaborating with philosophers to develop them (Kauffman and Clayton 2006; Kylafis and Loreau 2008). Hence, the distinction between biological and philosophical concepts of agency becomes somewhat irrelevant, especially when the concept is used to structure theoretical models.

## Agency in philosophy of biology

Mirroring the distinctions made in theoretical biology, philosophy also recognises two main notions of agency: one rooted in organisational theory and the other in evolutionary theory.

## Organisation

The concept of agency is frequently invoked to capture the specificity of organisms and of their relationships with the environment as opposed to other (non-living) systems. This specificity is mostly evident in the coordination and orientation of their activities towards internal goals, such
as maintenance, survival, and reproduction, but it encompasses several other key characteristics of organisms: their individuality, their ability to self-initiate actions, the normativity of their activities, and their adaptability to both internal and external changes (Barandiaran, Di Paolo, and Rohde 2009; Moreno and Mossio 2015; Virenque and Mossio 2024).

While this concept of agency primarily applies to organisms, it can be extended to lower-level entities such as cells, when their activities exhibit characteristics typically associated with agency (Skewes and Hooker 2009; A. Soto, 2024, personal communication).

Although this concept of agency is primarily aimed at understanding the organisation of biological systems, the inclusion of adaptability bridges the gap between the organisational and evolutionary approaches (Virenque and Mossio 2024).

## Evolution

In recent years, the role of organisms in shaping evolutionary processes has garnered increased recognition, particularly in terms of how they influence selective pressures through processes like niche construction. This shift in perspective has led researchers to import the concept of agency into evolutionary theory (Walsh 2015). In this context, for a biological entity to be considered an agent, it must influence its ecology or environment through its activities (e.g. Bateson 2004; Corning 2014). While this entity is typically an organism, it can also extend to include groups or populations. Here, the focus shifts from the inherent attributes of the agent to the active role it assumes in the observed dynamics.

## Choice

Denis Walsh (2015) made the first substantial contribution to defining agency from an evolutionary perspective. According to Walsh, all organisms can be considered agents, because they experience ecological conditions as affordances to which they respond based on their adaptive repertoire and with respect to their internal goals. The active role played by organisms is rooted in their capacity to select responses in alignment with their goals. Consequently, agency is closely linked to this ability to make choices, even when those are deemed unintentional – an idea further developed by several other researchers (see Corning et al. 2023; Desmond 2023).

## Inventiveness

A distinct form of agency emerges in the phenomenon of invention. An invention is a novel behaviour, that neither the organism nor any member of its group has previously exhibited. This type of agency differs from the choice model, in that it does not necessarily require goal-directedness but rather inventiveness — the ability to self-initiate a new behaviour whose novelty can be attributed to the organism itself (i.e., it is neither genetically determined, environmentally induced, nor socially learned) (Tahar 2023). Such agency, commonplace in humans, is increasingly studied in non-human animals due to its potential impact on evolution. Indeed, when an invention becomes an innovation, that is when it spreads in the population and becomes part of the culture, it can significantly influence ecological dynamics and, consequently, selective pressures (Sol 2003). Moreover, it can initiate a “Baldwin effect”: where a new useful behaviour acquired through individual inventiveness becomes a catalyst for natural selection, inducing genetic evolution (Baldwin 1896).

Although the definitions of agency in biology are varied, this does not preclude the existence of underlying commonalities, which justify the use of the same term for these diverse concepts. The challenge now is to synthesise these approaches to identify the principles that unify them. In the following section, I explore what these definitions share, beginning with the everyday meaning of agency.
# What do the different definitions of biological agency share?

The persistence of the term “agency” across diverse contexts and definitions suggests that scholars from different fields share a vague, intuitive understanding of agency, and do not tend to devise a sui generis concept, divorced from the general, common, idea of agency (Okasha 2024).

## Intuitive meaning of agency

In everyday language, agency refers to the capacity of an entity or individual to exert control or influence over something else. This concept addresses both the agent, which possesses the ability to (α) self-initiate an action, and the consequences of this action: for agency to be acknowledged, the action (β) must yield tangible effects (Merriam-Webster 2025; Oxford English Dictionary 2025). Refining this definition further, an entity is considered an agent only if (γ) it has some control over the action it initiates and if it serves as both the cause and the *raison d'être* of that action. In other words, the action would be different, or might not occur at all, if the agent were different. This means that (δ) the action depends on the agent's singularity and flexibility. This definition implies that agents are not agents all the time, in all situations. For instance, while humans in full command of their faculties are typically recognised as agents, we do not consider someone experiencing uncontrolled actions, such as hiccups or stumbling, to be exercising agency in those moments.

This initial definition already helps to eliminate certain inappropriate uses of the concept. For example, natural selection itself does not initiate or influence anything: it is merely the result of the individuals' struggle for survival and their consequent differential reproduction. Therefore, it is not a good candidate for agency. Similarly, while genes, as carriers of information, exert influence, they do not self-initiate actions; their influence is contingent upon their so-called “interpretation”, which disqualifies them as agents. Conversely, organisms could be considered agents, though not in all situations. Consider cows that release methane. The cow initiates the belch that releases methane, impacting climate change, but this process is not under the cow's control, nor does it vary significantly between individuals. Hence, in this scenario, the cow is not an agent.

While this preliminary definition is broad and does not teach us anything about the processes enabling agency or the characteristics of agentive actions, it does help to eliminate inappropriate uses of the term. One could argue that the various uses of “agency” in biology share only this vague definition and might therefore coexist without further integration. However, a more compelling approach is to hypothesise that a deeper unity underlies these definitions, potentially forming the foundation for a cohesive concept of biological agency. This paper adopts the latter perspective, aiming to explore the potential for a unified understanding of agency across different biological contexts.
# Synthesising the scientific definitions

Before unifying the conceptualisation of agency, it was necessary to compile the various definitions of the term. To achieve this, I conducted a literature review, drawing from theoretical biology, philosophy of biology, and behavioural ecology. Relevant articles were identified through data mining using the keyword 'agency' in combination with 'biology', 'evolution', 'ecology' 'organisms', or 'animals'. These diverse sources led me to distinguish three primary categories of phenomena to which agency is applied: behaviour, the organisation of biological systems, and the influence of organisms on ecological and evolutionary dynamics.

## Agency in understanding behaviour: the ethological approach

The concept of agency as applied to the behaviour of living organisms, primarily animals but also plants (Gilroy and Trewavas 2023; Sandford 2024; Sultan, Moczek, and Walsh 2022), encompasses several key characteristics across different studies:

(1) The agent is considered an individual (Sueur, Zanaz, and Pelé 2023; Wemelsfelder 1997). However, this individuality is not isolated; the agent exists in relation to its surroundings, and its agency is contingent upon these surroundings (Carter and Charles 2013; Read and Szokolszky 2024).

(2) These surroundings are experienced by the agent as affordances. In other words agents are not merely objective but subjective entities: they have an internal experience, and an individual memory (Steward 2009; Sueur, Zanaz, and Pelé 2023; Warkentin 2009).

(3) The relationship with the environment can be asymmetrical, i.e., the agent can self-initiate some actions (Read and Szokolszky 2024; Steward 2009; Wemelsfelder 1997).

(4) Agents exercise a certain level of control over at least some of their actions. For instance, they can follow orders or not (Scott 2009), and possess the ability to resist undesirable conditions, (e.g., they can attempt to escape in captivity) (Couchman 2012; Kaneko and Tomonaga 2011; Warkentin 2009).

(5) Agency is recognised through volition, i.e., preferences or choices of resources, habitat, or behaviours (Blattner, Donaldson, and Wilcox 2020). For instance, the controversial 2019 study (Reinhold et al. 2019) was interpreted as demonstrating the rat's agency because it revealed their ability to engage in hide-and-seek without direct rewards, indicating their intrinsic motivation. In other words, through their behaviours, agents exhibit internal goals or desires (Špinka 2019; Sueur, Zanaz, and Pelé 2023; Warkentin 2009).

(6) Agency is also manifested when organisms demonstrate flexibility in their actions, meaning their behaviour does not appear automatic but is the result of a dynamic interaction with the environment. (Gilroy and Trewavas 2023; Wemelsfelder 1997; Sultan, Moczek, and Walsh 2022).

(7) In some cases, the agents exhibit 'surprising' behaviour, that seems at odds with the usual repertoire: thereby manifesting creativity (McFarland and Hediger 2009; Špinka and Wemelsfelder 2011).

(8) Agents have a tangible impact, whether on the conditions of experiments, their surroundings, or even their own destiny (Carter and Charles 2013).

## Agency in understanding biological systems: the organisational approach

The organisational approach, as shared by biologists and philosophers, reveals distinct characteristics. Through an analysis of the literature, I identified the following key features:

(A) The agent is an individual (Barandiaran, Di Paolo, and Rohde 2009; Gontier 2023).
(B) This individual is a self-organising system. In other words:

a. It exhibits a form of closure (Moreno and Mossio 2015)...

b. derived from its self-generated structure, i.e., the system is autopoietic (Moreno and Mossio 2015; Skewes and Hooker 2009).

(C) Agents are capable of initiating activities, actions, and interactions (Moreno and Mossio 2015; Newman 2023; Skewes and Hooker 2009).

(D) Agency is characterised by autonomy. In other words:

a. Agents produce their own internal norms (Barandiaran, Di Paolo, and Rohde 2009; Moreno and Mossio 2015; Skewes and Hooker 2009)...

b. which define internal goals towards which their activities are oriented (Moreno and Mossio 2015; Newman 2023; Watson 2023).

(E) Agents are adaptative, flexible systems (Skewes and Hooker 2009; Virenque and Mossio 2024).

(F) They are adaptive because they retain experiences and have memory, which gives their organisation a historical dimension (Moreno and Mossio 2015; Watson 2023).

(G) Agents' activities have causal effects both on themselves and on their surround (Moreno and Mossio 2015; Newman 2023).

It is important to note that sometimes the same word is used across the different approaches but with non-identical meanings. This is the case with goals. In the context of behaviour, goal-directedness refers to organisms actively pursuing their *desires* (5). However, in the organisational approach, goal-directedness means that the agent's activities are oriented towards *internal goals* (such as maintenance, or reproduction) (D).

*Agency in understanding the role of organisms in evolution: the evolutionary approach*

Finally, agency is used to explore the influence of organisms' activities on evolutionary dynamics. In this context, agency is defined by the following characteristics:

(i) The agent is an individual (Tahar 2023; Walsh 2015).

(ii) Agents experience their conditions as affordances, which necessitates having an internal experience and an individual memory (Tahar 2023; Walsh 2015).

(iii) Consistent with other definitions, agents are able of self-initiating actions (Aaby and Desmond 2021; Tahar 2023).

(iv) They possess some degree of control over their actions (Aaby and Desmond 2021; Trewavas 2023).

(v) Many scholars posit that agency requires goal-directedness: agents are individuals acting according to their own goals (Corning 2023; Desmond 2023; Walsh 2015). Agency is then defined by the ability to choose one's activities, according to these goals (Desmond 2023; Kauffman and Clayton 2006; Walsh 2015).

(vi) Agents can act in their ever-changing surrounds through adaptive responses, demonstrating their flexibility (Tahar 2023; Walsh 2015).

(vii) In some cases, agency is expressed through inventions that can become innovations (Tahar 2023). In other words, an agent can sometimes exceed the limits of its repertoire, which is primarily genetically, but also culturally constituted, and invent a new behaviour, which may then spread within the group.

(viii) Agents' self-initiated actions can have significant impacts on ecology or evolution (Aaby and Desmond 2021; Kauffman and Clayton 2006; Laland, Odling-Smee, and Feldman 2019; Trewavas 2023).
The next step involves organising these various definitions to identify their shared elements and distinctions, enabling a unified conceptual framework of biological agency.

# Organising the definitions

Some elements in the definitions of agency relate less to the inherent properties of the agent and more to the actions they perform and the relationship between the agent and those actions.

## Agentive actions

Regarding the nature of an agent's actions – how agents enact their agency – there is a clear consensus (Figure 1):

*Agentive actions are initiated by the agent itself. (α; 3; C; iii)*

*Agentive actions are actions with consequences. (β; 8, G, viii)*

Additionally, while not a *sine qua non* condition for organisational agency, it is often the case that:

*These actions are within the agent's control (not only does the agent trigger the action, but it also has power over its execution). (γ; 4; iv)*

While most definitions describe agentive actions as goal-directed, the ambiguity surrounding the term “goal-directedness” across different definitions warrants setting this aspect aside for the moment.

## Biological agent

The more complex issue lies in identifying the properties that enable agents to perform these agentive actions. To address this, I have drawn upon the remaining points from the three perspectives discussed. Some characteristics are consistently present across all definitions of biological agency (Figure 2).

One of the most significant is individuality:

*An agent is an individual (which might be a collective entity, as long as it performs actions as a unified whole). (1; A; i)*

Another shared characteristic is flexibility. Regardless of the approach to agency, the agent is regarded as capable of a range of behaviours or activities, making it a flexible individual. This flexibility is foundational for the volition observed by ethologists, the adaptability crucial in organisational theory, the ability of agents to choose actions from an adaptive repertoire, and even the capacity for invention. Without this flexibility, agents could not exhibit creative or adaptive behaviours. This flexibility, sometimes referred to as “adaptive improvisation” (Soen, Knafo, and Elgart 2015) or even “play” (Kortmulder 1998; G. Longo 2023, personal communication; Tahar 2023; Wemelsfelder 1997) does not necessarily imply goal-directedness. Flexibility does not equate to directionality.

*An agent is flexible in its activities and behaviours: it can play with its internal and external conditions. (6; E; vi)*

However, for flexibility to result in controlled actions rather than random behaviours, it must be channelled. This channelling arises from the agent's internal experience, which is shaped by its unique embodied perspective and individual memory. It is important to note that memory does not always require a brain. For example, worms trained to traverse rough surfaces in exchange for a reward retained the learned behaviour even after being decapitated and regenerated from their tails, suggesting that the cells in the tails recorded past experiences
(Shomrat and Levin 2013). Similarly, brainless slime moulds have been shown to overcome their aversion to caffeine to reach oatmeal they previously enjoyed, demonstrating memory-related behaviour (Boisseau, Vogel, and Dussutour 2016).

*An agent is a subjective entity, registering its experiences in an individual memory.* (2; F; ii) I synthesise these three criteria – Individuality, Playful flexibility, and Memory (IPM) – as the foundational elements of biological agency:

*An agent is an individual characterised by a playful (non-directional) flexibility, and a memory that channels and directs this flexibility.*

Individuality, Playful flexibility, and Memory are the elements without which biological agency cannot be identified: they represent both the necessary conditions and the criteria by which agency can be recognised. Whether they are sufficient conditions for biological agency remains an open question. Nevertheless, this question is likely merely theoretical, as most biological entities that meet these three conditions also exhibit other characteristics posited by the organisational, ethological, or evolutionary approaches. It is also worth noting that, in line with the principle of Occam's razor, I have excluded goal-directedness from these foundational criteria. I argue that the directionality associated with agency is sufficiently accounted for by the channelling of flexibility through memory, rendering explicit references to goals unnecessary. However, as we will see, goal-directedness remains important for explaining certain properties of specific forms of agency.

What about the other criteria that were not incorporated into this synthesis? These criteria delineate distinct forms of agency, which will be examined in the following section.

# The different forms of agency

## Autonomous organisation, goal-directed choice, inventiveness

Having established the foundational elements of biological agency (IPM) we can now explore the other characteristics put forward by the different definitions, which constitute more specific forms of agency (Figure 7).

### Autonomous organisation

The organisational concept of agency introduces several additional features to IPM, forming a distinctive form of agency. These features are organised into two categories: (B) self-organisation, which encompasses closure and autopoiesis, and (D) autonomy, which includes normativity and internal goals. All of these are in fact interrelated, as self-organisation leads to the creation of internal norms that guide the agent's activities. Together with the foundational elements of agency (IPM), these characteristics define a first form of agency: *agency as autonomous organisation*.
## Goal-directed choice

Across the different approaches of agency, certain features appear to overlap. This is the case of volition and choice: the distinction between these concepts seems to lie less in their meanings and more in the roles they play within specific arguments. Both volition as discussed in ethology, and choice as framed in evolutionary biology, pertain to goal-directed actions. However, volition is often used in animal studies not only for scientific purposes but also to advocate for ethical considerations in experimental contexts (Sueur, Zanaz, and Pelé 2023; Webb, Woodford, and Huchard 2019), or in everyday interactions (Blattner, Donaldson, and Wilcox 2020; Carter and Charles 2013). This focus on volition seeks to acknowledge the moral agency (McFarland and Hediger 2009), or even citizenship of animals (Kymlicka and Donaldson 2014). In contrast, evolutionary biology tends to adopt a more theoretical stance (although the boundary between theory and ethics remains porous), likely explaining the preference for the concept of choice over volition or will. The shift away from volition, exemplified by Walsh's notion of “unintentional choice” (2015), likely reflects an effort to avoid anthropomorphism and maintain scientific rigour¹. Regardless of the terminology, both volition and choice refer to an agent's capacity to select actions based on internal goals or desires. This capacity forms a second form of agency, which I refer to as *goal-directed choice*.

## Inventiveness

The notions of creativity in the ethological approach and of inventiveness in the evolutionary one also appear to be synonymous. While ethology emphasises the element of surprise for the observer, the evolutionary perspective highlights how the invented behaviour deviates from the known repertoire. However, in both cases, they are referring to the agents' ability to produce new actions, behaviours, and relationships with the environment, often unexpected by observers. Inventiveness is distinct from choice-agency because it does not necessarily require goal-directedness. Although inventiveness can occur in goal-directed activities, such as problem-solving, it may also emerge without a clear purpose, especially during play (Bateson 2014; Burghardt 2015). This capacity for innovation constitutes a third form of agency: *agency as inventiveness*.

Despite their distinct properties, the three forms of agency share the foundational elements of agency (IPM). However, Individuality, Playful flexibility, and Memory manifest in varying degrees and hold different levels of importance across the three forms. Similarly, while all agentive actions involve initiation, control, and consequences, the expression of these elements differs between autonomous organisation, goal-directed choice, and inventiveness.

In the following section, I will explore these distinctions in greater detail.

## Comparison: the different types of agentive actions

The three forms of agency involve different types of agentive actions, distinguished by their main target and by the degree of control the agent exerts over them (Chart 1).

### Target of the action

In autonomous organisation, agentive activities are primarily oriented toward the agent itself. This explains the organisational theory's emphasis on *self-organisation*, *self-maintenance*, and *self-constitution*. The impact on the agent is direct, while any effect on the relationship with the environment or on ecological and evolutionary dynamics is indirect.
In the case of goal-directed choices, actions predominantly influence the relationship between the organism and its environment, whether through changes in habitat, resources, or social interactions. In some instances, the environment itself may become the direct target of these actions. For example, the construction of dams by beavers is a behaviour primarily aimed at transforming the environment. However, except in the case of human beings, such actions only indirectly affect the agent, and evolutionary dynamics.

Finally, invention influences both the agent and its relationship with the environment. The agent expands its behavioural repertoire, while new interactions with the environment become possible. For instance, in Jerusalem, rats invented a technique for opening pinecones, which spread and became an innovation (Zohar and Terkel 1991). This invention affects both the rat's behavioural repertoire, widening it, and its interaction with the environment, enabling the consumption of pinecones and the invasion of pine forests. As with goal-directed choice, it is conceivable that inventions could affect the environment directly, although empirical evidence for this remains limited. Except when the agent is human, it seems that the influence of invention on ecological and evolutionary dynamics remains mostly indirect.

## Control over the action

The degree of control an agent exerts over its actions varies across different types of agency. In autonomous organisation, control originates from the *agent's internal conditions*, which the agent contributes to shape, even though they are also heavily constrained by genetics and the environment.

In goal-directed choice, the agent exerts control by *selecting* one behaviour from a range of possible actions, each of which is equally likely. Although the execution of the behaviour is determined by this choice, its structure is largely predefined by the existing behavioural repertoire, shaped partly by the organism's individual experience (Memory) but mostly by past selection. Thus, while the agent controls the specific execution of the action, it does not fully control its overall structure.

In contrast, invention involves both the performance and the structure of the action being determined by the agent, as the action is not part of the initial behavioural repertoire. The agent *invents* the behaviour: in other words, it functions as both the triggering and the structuring cause of its behaviour (for this distinction, see Dretske 1997).

These differences in the control of action reflect deeper distinctions in the nature of the agent itself.

## Comparison: the different types of agents

While the three forms of agency—autonomous organisation, goal-directed choice, and inventiveness—share the foundational characteristics of individuality, playful flexibility, and memory (IPM), these elements are expressed differently across each form.

### IPM in the three forms of agency

The role of individual experience and memory remains consistent across the three forms of agency (depending on the specific action being performed). However, the significance of individuality and flexibility varies between these forms (Chart 2).

For example, high individuality is crucial in autonomous organisation, where high-level of cohesion is required for self-organisation. In contrast, goal-directed choice and inventiveness necessitate less unity, and can be performed by less cohesive groups.
Playful flexibility is especially important in inventiveness. Although this flexibility is channelled by individual experience, and sometimes contextual factors (especially in problem-solving), inventiveness demands the greatest freedom from genetic and environmental constraints. Choice, while also involving playful flexibility, operates under tighter constraints; it is channelled not only by the agent's individual experience but also by its goals and typical behavioural repertoire. In autonomous organisation, this flexibility is further narrowed, directed by the unity of the cohesive, self-oriented goal – the agent itself.

Each form of agency also possesses distinct characteristics beyond IPM, constituting their specificity. An important question is whether these forms of agency are interconnected and, if so, how they relate to one another.

## Connections between the three forms of agency

The characteristics proper to autonomous organisation (self-organisation and normativity) also serve as a foundation for enabling both goal-directed choice and invention, albeit required to a lesser degree. For example, while groups may meet the criteria for goal-directed choice and inventiveness, their usual lack of closure and self-organisation makes them less suitable (although not theoretically disqualified) as candidates for autonomous organisation. Conversely, the characteristics of goal-directed choice and invention do not overlap. Goal-directed choice is not a prerequisite for autonomous organisation or invention, nor is invention necessary for autonomous organisation or choice. These forms represent *diverging* types of agency: invention involves creating new behaviours, whereas choice concerns selecting from pre-existing options. However, both forms share some characteristics. Like goal-directed choice, invention requires the agent to transcend automatic responses to the environment. Additionally, in some cases, invention can be goal-oriented, especially in problem-solving.

## Agents

A final important question is: which agents are concerned by these forms of agency?

In theory, any biological entity can perform behaviours corresponding to autonomous organisation, goal-directed choice, or inventiveness. However, some entities are more prone than others to exhibit certain forms of agency.

Autonomous organisation, with its emphasis on individuality and cohesion, is most likely to manifest in organisms, especially multicellular organisms, which exhibit higher levels of self-organisation.

Goal-directed choice is evident when organisms could have acted differently, i.e., alternative actions were conceivable by the observer. While this form of agency is more easily shown in organisms, it can also be exhibited by cells or groups, under certain conditions.

Invention can theoretically occur across a broad range of across diverse entities too, but it is predominantly observable in individuals whose behaviour can be studied over extended periods. While invention has primarily been documented in animals, recent studies on plants may lead us to reconsider this exclusivity.

This analysis demonstrates that there is no hierarchy among the three forms of agency, as they differ in how they express IPM and introduce distinct characteristics that define each form. However, this does not mean that the different forms of agency cannot be rigorously compared. Indeed, Charts 1 and 2 can be employed to identify and quantify agentive actions or to specify the agency of a biological individual. Moreover, this discussion highlights that the different types of agency, despite their distinct processes, are not mutually exclusive and may coexist within the same entities, particularly in organisms. The next step is to examine how the processes underpinning these varied forms of agency interact with one another.
# How do these forms of agency interact?

To understand how different processes contribute to each form of agency and how the three forms of agency interact, the most effective case study is the organism, as it is where these different forms of agency are most likely to coexist.

## Agency in the organism

### Autonomous organisation

The organism itself constitutes an autonomous organisation, which forms and sustains itself over time through self-directed processes of development and maintenance (Figure 3). These processes are enabled by IPM, but also by certain characteristics of the system: it is autopoietic (self-generating with structural closure) and it is autonomous (establishing internal norms and defining its own goals). The individual is not isolated: its agentive activities are influenced by its environment, and by other constraints inherited from history (such as past natural selection, the successive organisational stages it underwent during development, and prior agentive processes) (Montévil 2022; Montévil and Mossio 2015). This form of agency may or may not influence selective pressures and evolutionary processes (see next section).

The question now arises: how does this form of agency interact with others? Along with IPM that operates across all levels of agency, organisational processes contribute to shaping the behavioural repertoire of the agent, though this repertoire is also influenced by external constraints such as genetic and environmental pressures, as well as social and cultural contexts. Organisational processes also contribute to shaping the organism's goals; however, these goals are largely imposed by environmental factors, particularly those related to survival and reproduction. Combined with environmental pressures, the behavioural repertoire and natural goals, when limited, can constrain the agent's actions to the extent that it may never exercise its capacity of choice, i.e., of deliberation (Desmond 2023). Thus, autonomous organisation does not necessarily coexist with other forms of agency (Figure 3b). While this form of agency may enable goal-directed choice or even inventiveness, it doesn't guarantee their exercise. In many cases, organisms are "just" autonomous organisations, their actions being otherwise fully determined.

But sometimes, autonomous organisations *do make choices*.

### Goal-directed choice

Certain organisms possess the ability to select behaviours from their behavioural repertoire, and according to their goals (Figure 4). When organisms display this capacity, they go beyond being merely autonomous organisations and exhibit a distinct form of agency: goal-directed choice. At this level, IPM intervenes both in the definition of goalsii, in the definition of the repertoire and in the choice itself. While autonomous organisation enables this form of agency, it may not be enacted if the organism consistently responds automatically to its environment. The expression of goal-directed choice depends not only on the individual agent but also on environmental conditions, as it is less likely to manifest in stressful situations that demand quick, risk-free actions.
However, it is not only autonomous organisation that shapes agency as choice; choices themselves feed back into and influence the organisation. For instance, food choices can affect immunity or even gene expression; partnership choices can sometimes significantly affect an individuals' phenotype, especially during early developmentiii. In most cases, choices impact the organism's relationship with the environment, although not necessarily at an evolutionary scale.

## Inventive agency

Some organisms manifest inventive agency (Figure 5): in certain circumstances they invent new behaviours, outside of their existing behavioural repertoire. This inventiveness is constrained by organisational processes but not by choice processes. However, in some cases, the inventive process is channelled by goals, sharing this goal-directedness with choice agency. Like choice, invention retroactively influences the organism's overall organisation, but also directly transforms the behavioural repertoire, by introducing new patterns. This, in turn, may or may not affect evolutionary dynamics.

### Why do organisms invent rather than select from the existing repertoire?

Invention likely occurs in scenarios where no immediate, functional behaviour is required, allowing the organism to take risks and experiment. This aligns with the observed link between play and inventiveness: many non-human inventions are first observed during play. Research indicates that play promotes inventiveness and potentially adaptability across species (Bateson 2014; Burghardt 2015; Špinka, Newberry, and Bekoff 2001). The lack of a specific goal seems to free flexibility from the demands of efficiency, enabling true playfulness. Conversely, it could be argued that “necessity is the mother of invention” (Reader and Laland 2003): in emergencies, or unexpected situations, where typical behaviours fail, organisms are compelled to invent. Both perspectives may be valid. But, to this day, determining when and why individuals deviate from routine to experiment with new behaviours remains a challenge.

While Figure 3 primarily serves the organisational approach, Figures 4 and 5 cater to ethological and ecological perspectives. However, these three models offer limited insight into the long-term evolutionary implications of agency. This is the focus of the next section.

## Agency in evolution

What makes a biological agent an *evolutionary* agent? For an individual to act as an evolutionary agent, it must not only be the full cause of its action, but that action must also exert a tangible effect on the environment and selective pressures, impacting long-term evolutionary dynamics. How can each of the three forms of agency affect ecology and evolution?

To begin with, it should be stressed that these three forms of agency do not necessarily influence the evolution of species. Some organisms may be agents, because they are autonomous organisations, because they make choices, or even because they invent behaviours, but their agentive activities may nevertheless prove to have no notable consequences on the longer-term dynamics of evolution.

However, in some cases, these activities can be decisive.
## Autonomous organisation and evolution

When autonomous organisation does not significantly affect selective pressures, we observe a near-replication of forms generation after generation. However, in some instances, organisational activities have consequences on the agent-environment interaction, potentially sparking evolutionary shifts or tempering the effects of selective pressures (Figure 6). It is known, for example, that phenotypic plasticity can prevent a species from extinction or allow a population to pursue new evolutionary directions (West-Eberhard 2003). In these cases, autonomous organisation has a causal influence on evolutionary dynamics, making the organism an evolutionary agent.

## Goal-directed choice and evolution

The choices organisms make regarding habitats, resources, and partners contribute to niche construction, and thus affects the way they interact with their environment (Aaby and Desmond 2021). These choices can also influence offspring numbers and may influence the ecology, even if the environment is not the primary target of the action (Kylafis and Loreau 2008). For instance, the foraging paths chosen by ants have consequences for the entire ecosystem, as ants drop part of their harvest along the way. In this case, the choice likely has long-term effects, thereby contributing to shaping selective pressures. However, not all choices have such long-term consequences – this is especially true of choices made by individual agents. For example, if an individual or a small group migrates on its own and fails to survive, their quick death will prevent their choice from having lasting effects on the environment, the fate of the population, or selective pressures. In such cases, they are agents, but not evolutionary ones. Conversely, if they survive and reproduce, their choice may significantly shift the evolutionary trajectory of the entire species (Zelnik, Solomon, and Yaari 2015). Indeed, for the choice to be evolutionarily significant, its consequences must persist across generations (Aaby and Desmond 2021; Laland, Odling-Smee, and Feldman 2019). Through its choice, the agent (or more often the agents) initiates a new evolutionary dynamic, and can thus be qualified an evolutionary agent (Figure 7).

## Inventiveness and evolution

Inventions are rare and difficult to observer. However, the emergence of an inventor suggests conditions that foster inventiveness, whether in the environment, the population, or the species itself. This implies that when one invention occurs, others may arise across several generations within the same population. Nonetheless, like other forms of agentive action, inventions do not necessarily have an evolutionary impact, as they rarely spread widely enough to cause large-scale effects. The inventor is then an agent, but it is not an evolutionary agent. However, when inventions do spread, become innovations and change the way things are done, they can have long-term effects, on both organisms, their interaction with the environment, and the broader ecology (Bateson 2004; Sol 2003) (Figure 8). A prime example is the orcas in the Aleutian Islands, which innovated by including otters in their diet. This shift affected the entire ecosystem: with otters preying on urchins, their decline caused urchin populations to surge, devastating the kelp forests crucial for local biodiversity (Estes et al. 1998). Even if the orcas were to revert to their previous diet, this dietary innovation will likely have profound evolutionary consequences.

The long-term effects of innovations are even more pronounced when they are consistently transmitted through successive generations. In such cases, both the inventor and those spreading and adopting the invention can be considered evolutionary agents, as they alter selective pressures on the long-term.
To capture the evolutionary significance of these three forms of agency, it is crucial to
acknowledge the roles of individual memory, as well as the historical continuity and progression
of agentive processes. Biological agents shape a continuous history, generation after generation,
through which they can exert influence on evolutionary processes.

Remaining challenges

One remaining challenge is determining where agency ends. Should agency encompass only the direct target of the action, what the agent controls? If so, organisms would not be considered agents of the consequences of their actions, which complicates the idea that they can be agents of evolutionary change. For instance, an ant selecting its foraging path has no control over the seeds fall, yet it is the fall that impacts the environment and potentially has an evolutionary effect. Is the ant *not* an agent of this evolutionary effect? Likewise, a cow belching is not regarded as the agent of increased methane levels. But if cows were to innovate by changing their diet, thereby reducing methane emissions, this would influence their ecosystem and perhaps alter selective forces; would the cows *not* be agents of this evolutionary change? On one hand, if agency includes the actions' consequences, by a sort of "butterfly effect", organisms risk being considered agents behind nearly all evolutionary events—rendering agency a non-operational concept in evolutionary biology. On the other hand, if organisms are only agents of what they directly control, then even humans would rarely qualify as evolutionary agents. One potential solution is to differentiate between more or less direct consequences, associating these with varying degrees of agency. For example, if cows change their diet and start competing with a new species, they could be seen as "more" agents of this shift in selective pressures than of the secondary ecological consequences resulting from the changes in methane emissions due to their new diet.

Overall, our understanding of agency remains limited. It is especially true of non-human inventiveness: the conditions that facilitate invention, the supporting biological processes, and how these differ from those underlying choice are still largely unknown.

More broadly, we know very little about the physiological, organisational and cognitive
processes underlying Playful flexibility and Memory. Understanding how these processes
interplay to enable agency^iv, and how they drive different forms of agency, remains an open
question.

Concluding remarks

Recognising the role of agency prompts a shift from a deterministic view to an agent-based one that acknowledges individual specificity. While this perspective is gaining traction in organisational studies, and ecological studies, it remains undervalued in evolutionary theory. The dominant view still regards the dynamics of biological systems as irrelevant to the study of evolution, despite their role in long-term evolutionary processes (Duckworth 2021). However, considering organisms as potential agents in evolution, and thus integrating organisational and behavioural studies, could improve our understanding of evolutionary dynamics, bridging the explanatory gaps between different biological processes. A unified, integrative approach to agency could thus be critical for making agency an operational concept
in evolutionary biology. This paper represents an initial attempt to connect the different concepts of agency, and the different fields of research that study them.

## Literature cited.

Aaby, Bendik Hellem, and Hugh Desmond. 2021. ‘Niche Construction and Teleology: Organisms as Agents and Contributors in Ecology, Development, and Evolution’. *Biology & Philosophy* 36 (5): 47. https://doi.org/10.1007/s10539-021-09821-2.

‘Agency, n. Meanings, Etymology and More’, Oxford English Dictionary, 2025. Accessed 24 January 2025. https://www.oed.com/dictionary/agency_n.

Arbilly, Michal, and Arnon Lotem. 2017. ‘Constructive Anthropomorphism: A Functional Evolutionary Approach to the Study of Human-like Cognitive Mechanisms in Animals’. *Proceedings of the Royal Society B: Biological Sciences* 284 (1865): 20171616.

https://doi.org/10.1098/rspb.2017.1616.

Baldwin, James Mark. 1896. ‘A New Factor in Evolution’. *American Naturalist* 30:441-451;536-553.

Barandiaran, Xabier E., Ezequiel Di Paolo, and Marieke Rohde. 2009. ‘Defining Agency: Individuality, Normativity, Asymmetry, and Spatio-Temporality in Action’. *Adaptive Behavior* 17 (5): 367–86. https://doi.org/10.1177/1059712309343819.

Bateson, Patrick. 2004. ‘The Active Role of Behaviour in Evolution’. *Biology & Philosophy* 19 (2): 283–98. https://doi.org/10.1023/B:BIPH.0000024468.12161.83.

———. 2014. ‘Play, Playfulness, Creativity and Innovation’. *Animal Behavior and Cognition* 1 (2): 99–112. https://doi.org/10.12966/abc.05.02.2014.

Blattner, Charlotte E, Sue Donaldson, and Ryan Wilcox. 2020. ‘Animal Agency in Community’ 6.

Boisseau, Romain P., David Vogel, and Audrey Dussutour. 2016. ‘Habituation in Non-Neural Organisms: Evidence from Slime Moulds’. *Proceedings of the Royal Society B: Biological Sciences* 283 (1829): 20160446. https://doi.org/10.1098/rspb.2016.0446.

Burghardt, Gordon M. 2006. ‘Critical Anthropomorphism, Uncritical Anthropocentrism, and Naïve Nominalism’. *Comparative Cognition & Behavior Reviews* 2.

https://doi.org/10.3819/ccbr.2008.20009.

———. 2015. ‘Creativity, Play, and the Pace of Evolution’. In *Animal Creativity and Innovation*, 129–61. Elsevier. https://doi.org/10.1016/B978-0-12-800648-1.00005-X.

Carter, Bob, and Nickie Charles. 2013. ‘Animals, Agency and Resistance’. *Journal for the Theory of Social Behaviour* 43 (3): 322–40. https://doi.org/10.1111/jtsb.12019.

Corning, Peter A. 2014. ‘Evolution “on Purpose”: How Behaviour Has Shaped the Evolutionary Process: Behaviour and the Evolutionary Process’. *Biological Journal of the Linnean Society* 112 (2): 242–60. https://doi.org/10.1111/bij.12061.

Corning, Peter A. 2023. ‘Teleonomy in Evolution: “The Ghost in the Machine”’. In *Evolution ‘On Purpose’: Teleonomy in Living Systems*, edited by Peter A. Corning, Stuart A. Kauffman, Denis Noble, James A. Shapiro, Richard I. Vane-Wright, and Addy Pross, 11–31. The MIT Press. https://doi.org/10.7551/mitpress/14642.001.0001.

Corning, Peter A., Stuart A. Kauffman, Denis Noble, James A. Shapiro, Richard I. Vane-Wright, and Addy Pross, eds. 2023. *Evolution ‘On Purpose’: Teleonomy in Living Systems*. The MIT Press. https://doi.org/10.7551/mitpress/14642.001.0001.

Couchman, Justin J. 2012. ‘Self-Agency in Rhesus Monkeys’. *Biology Letters* 8 (1): 39–41.
https://doi.org/10.1098/rsbl.2011.0536.

Dawkins, R. 1989. *The Selfish Gene*. Business Book Summary. Oxford University Press.

https://books.google.fr/books?id=WkHO9HI7koEC.

‘Definition of AGENCY’, Merriam-Webster. 2025. Accessed 20 January 2025.

https://www.merriam-webster.com/dictionary/agency.

Dennett, D.C. 1995. *Darwin's Dangerous Idea: Evolution and the Meanings of Life*. A Touchstone Book. New York: Simon & Schuster.

https://books.google.fr/books?id=FvRqtnpVotwC.

Desmond, Hugh. 2023. ‘The Deliberation Model of Organismic Agency’.

https://philarchive.org/rec/DESTDM-6.

Dretske, Fred I. 1997. *Explaining Behavior: Reasons in a World of Causes*. 5th print. A Bradford Book. Cambridge, Mass.: MIT Press.

Duckworth, Kathryn C. Chenard, Renée A. 2021. ‘The Special Case of Behavioral Plasticity?’

In *Phenotypic Plasticity & Evolution*, edited by David W. Pfennig. Boca Raton: CRC Press.

Emirbayer, Mustafa, and Ann Mische. 1998. ‘What Is Agency?’ *American Journal of Sociology* 103 (4): 962–1023. https://doi.org/10.1086/231294.

Estes, J. A., M. T. Tinker, T. M. Williams, and D. F. Doak. 1998. ‘Killer Whale Predation on Sea Otters Linking Oceanic and Nearshore Ecosystems’. *Science* 282 (5388): 473–76.

https://doi.org/10.1126/science.282.5388.473.

Gilroy, Simon, and Tony Trewavas. 2023. ‘Agency, Teleonomy and Signal Transduction in Plant Systems’. *Biological Journal of the Linnean Society* 139 (4): 514–29.

https://doi.org/10.1093/biolinnean/blac021.

Godfrey-Smith, P. 2009. *Darwinian Populations and Natural Selection*. OUP Oxford.

https://books.google.fr/books?id=QXex0rDN_ UC.

Gontier, Nathalie. 2023. ‘Teleonomy as a Problem of Self-Causation’. *Biological Journal of the Linnean Society* 139 (4): 388–414. https://doi.org/10.1093/biolinnean/blac111.

Horowitz, Alexandra, and Marc Bekoff. 2007. ‘Naturalizing Anthropomorphism: Behavioral Prompts to Our Humanizing of Animals.’ *Anthrozoös* 20 (1): 23–35.

Kaneko, Takaaki, and Masaki Tomonaga. 2011. ‘The Perception of Self-Agency in Chimpanzees (Pan Troglodytes)’. *Proceedings of the Royal Society B: Biological Sciences* 278 (1725): 3694–3702. https://doi.org/10.1098/rspb.2011.0611.

Kauffman, Stuart, and Philip Clayton. 2006. ‘On Emergence, Agency, and Organization’.
*Biology & Philosophy* 21 (4): 501–21. https://doi.org/10.1007/s10539-005-9003-9.

Kortmulder, Koenraad. 1998. *Play and Evolution: Second Thoughts on the Behaviour of Animals*. Utrecht, the Netherlands: International Books.

Kylafis, Grigoris, and Michel Loreau. 2008. ‘Ecological and Evolutionary Consequences of Niche Construction for Its Agent’. *Ecology Letters* 11 (10): 1072–81.

https://doi.org/10.1111/j.1461-0248.2008.01220.x.

Kymlicka, W., and S. Donaldson. 2014. ‘Animals and the Frontiers of Citizenship’. *Oxford Journal of Legal Studies* 34 (2): 201–19. https://doi.org/10.1093/ojls/gqu001.

Laland, Kevin N., John Odling-Smee, and Marcus W. Feldman. 2019. ‘Understanding Niche Construction as an Evolutionary Process’. In *Evolutionary Causation*, edited by Tobias Uller and Kevin N. Lala, 127–52. The MIT Press. https://doi.org/10.7551/mitpress/11693.003.0008.

McFarland, Sarah E., and Ryan Hediger. 2009. ‘Approaching the Agency of Other Animals: An Introduction’. In *Animals and Agency: An Interdisciplinary Exploration*, edited by Sarah E. McFarland and Ryan Hediger, 1–22. Human Animal Studies, v. 8. Leiden; Boston: Brill.

Montévil, Maël. 2022. ‘Historicity at the Heart of Biology’. *Theory in Biosciences* 141 (2): 165–73. https://doi.org/10.1007/s12064-020-00320-8.

Montévil, Maël, and Matteo Mossio. 2015. ‘Biological Organisation as Closure of Constraints’. *Journal of Theoretical Biology* 372 (May):179–91.
https://doi.org/10.1016/j.jtbi.2015.02.029.

Moreno, Alvaro, and Matteo Mossio. 2015. *Biological Autonomy: A Philosophical and Theoretical Enquiry*. Cham: Springer.

Nadolski, Erica M., and Armin P. Moczek. 2023. ‘Promises and Limits of an Agency Perspective in Evolutionary Developmental Biology’. *Evolution & Development* 25 (6): 371–92. https://doi.org/10.1111/ede.12432.

Newman, Stuart A. 2023. ‘Inherency and Agency in the Origin and Evolution of Biological Functions’. *Biological Journal of the Linnean Society* 139 (4): 487–502. https://doi.org/10.1093/biolinnean/blac109.

Okasha, Samir. 2018. *Agents and Goals in Evolution*. First edition. Oxford: Oxford University Press.

———. 2024. ‘The Concept of Agent in Biology: Motivations and Meanings’. *Biological Theory* 19 (1): 6–10. https://doi.org/10.1007/s13752-023-00439-z.

Pradeu, Thomas. 2011. ‘A Mixed Self: The Role of Symbiosis in Development’. *Biological Theory* 6 (1): 80–88. https://doi.org/10.1007/s13752-011-0011-5.

Read, Catherine, and Agnes Szokolszky. 2024. ‘What Animals Can Do: Agency, Mutuality, and Adaptation’. *Biological Theory* 19 (3): 198–208. https://doi.org/10.1007/s13752-024-00463-7.

Reader, Simon M., and Kevin N. Laland. 2003. ‘Animal Innovation: An Introduction’. In *Animal Innovation*, edited by Simon M. Reader and Kevin N. Laland, 3–36. Oxford University Press. https://doi.org/10.1093/acprof:oso/9780198526223.003.0001.

Reinhold, Annika Stefanie, Juan Ignacio Sanguinetti-Scheck, Konstantin Hartmann, and Michael Brecht. 2019. ‘Behavioral and Neural Correlates of Hide-and-Seek in Rats’. *Science* 365 (6458): 1180–83. https://doi.org/10.1126/science.aax4705.

Sandford, Stella. 2024. ‘What Plant Philosophy Says about Plant Agency and Intelligence | Aeon Essays’. Aeon. 2024. https://aeon.co/essays/what-plant-philosophy-says-about-plant-agency-and-intelligence.

Scott, Shelly R. 2009. ‘The Racehorse as Protagonist: Agency, Independence, and Improvisation’. In *Animals and Agency: An Interdisciplinary Exploration*, edited by Sarah E. McFarland and Ryan Hediger, 45–66. Human Animal Studies, v. 8. Leiden/Boston: Brill. Shapiro, Susan P. 2005. ‘Agency Theory’. *Annual Review of Sociology* 31 (1): 263–84. https://doi.org/10.1146/annurev.soc.31.041304.122159.

Shomrat, Tal, and Michael Levin. 2013. ‘An Automated Training Paradigm Reveals Long-Term Memory in Planaria and Its Persistence through Head Regeneration’. *Journal of Experimental Biology*, no. 216 (pt 20) (January), 3799–3810. https://doi.org/10.1242/jeb.087809.

Skewes, J. C., and C. A. Hooker. 2009. ‘Bio-Agency and the Problem of Action’. *Biology & Philosophy* 24 (3): 283–300. https://doi.org/10.1007/s10539-008-9135-9.

Soen, Yoav, Maor Knafo, and Michael Elgart. 2015. ‘A Principle of Organization Which Facilitates Broad Lamarckian-like Adaptations by Improvisation’. *Biology Direct* 10 (1): 68. https://doi.org/10.1186/s13062-015-0097-y.

Sol, Daniel. 2003. ‘Behavioural Innovation: A Neglected Issue in the Ecological and Evolutionary Literature?’ In *Animal Innovation*, edited by Simon M. Reader and Kevin N. Laland, 63–82. Oxford University Press. https://doi.org/10.1093/acprof:oso/9780198526223.003.0003.

Špinka, Marek. 2019. ‘Animal Agency, Animal Awareness and Animal Welfare’. *Animal Welfare* 28 (1): 11–20. https://doi.org/10.7120/09627286.28.1.011.

Špinka, Marek, Ruth C. Newberry, and Marc Bekoff. 2001. ‘Mammalian Play: Training for the Unexpected’. *The Quarterly Review of Biology* 76 (2): 141–68. https://doi.org/10.1086/393866.
Špinka, Marek, and Françoise Wemelsfelder. 2011. 'Environmental Challenge and Animal Agency'. In *Animal Welfare*, by Michael C. Appleby, Joy A. Mench, Barry O. Hughes, and I. Anna S. Olsson, 27–44. Wallingford, UK: CAB International.
Steward, Helen. 2009. 'Animal Agency'. *Inquiry* 52 (3): 217–31.
https://doi.org/10.1080/00201740902917119.

Sueur, Cédric, Sarah Zanaz, and Marie Pelé. 2023. 'Incorporating Animal Agency into Research Design Could Improve Behavioral and Neuroscience Research.' *Journal of Comparative Psychology* 137 (2): 129–43. https://doi.org/10.1037/com0000335.

Sultan, Sonia E., Armin P. Moczek, and Denis Walsh. 2022. 'Bridging the Explanatory Gaps: What Can We Learn from a Biological Agency Perspective?' *BioEssays* 44 (1): 2100185.
https://doi.org/10.1002/bies.202100185.

Tahar, Mathilde. 2023. 'Agency, Inventiveness, and Animal Play: Novel Insights into the Active Role of Organisms in Evolution. Special'. *Spontaneous Generations* 11 (1).
https://doi.org/10.4245/spongen.v11i1.1996.

Toates, Frederick. 2004. 'Cognition, Motivation, Emotion and Action: A Dynamic and Vulnerable Interdependence'. *Applied Animal Behaviour Science* 86 (3–4): 173–204.
https://doi.org/10.1016/j.applanim.2004.02.010.

Trewavas, Anthony. 2023. 'Agency, Teleonomy, Purpose, and Evolutionary Change in Plant Systems'. In *Evolution 'On Purpose': Teleonomy in Living Systems*, edited by Peter A. Corning, Stuart A. Kauffman, Denis Noble, James A. Shapiro, Richard I. Vane-Wright, and Addy Pross, 299–323. The MIT Press. https://doi.org/10.7551/mitpress/14642.001.0001.
Virenque, Louis, and Matteo Mossio. 2024. 'What Is Agency? A View from Autonomy Theory'. *Biological Theory* 19 (1): 11–15. https://doi.org/10.1007/s13752-023-00441-5.
Walsh, D. M. 2015. *Organisms, Agency, and Evolution*. 1st ed. Cambridge UK: Cambridge University Press. https://doi.org/10.1017/CBO9781316402719.

Warkentin, Traci. 2009. 'Whale Agency: Affordances and Acts of Resistance in Captive Environments'. In *Animals and Agency: An Interdisciplinary Exploration*, edited by Sarah E. McFarland and Ryan Hediger, 23–43. Leiden/Boston: Brill.

Watson, Richard. 2023. 'Agency, Goal-Directed Behavior, and Part-Whole Relationships in Biological Systems'. *Biological Theory*, November. https://doi.org/10.1007/s13752-023-00447-z.

Webb, Christine E, Peter Woodford, and Elise Huchard. 2019. 'Animal Ethics and Behavioral Science: An Overdue Discussion'. *BioScience* 69 (10): 778–88.
https://doi.org/10.1093/biosci/biz082.

Webb, Christine E., Peter Woodford, and Elise Huchard. 2020. 'The Study That Made Rats Jump for Joy, and Then Killed Them: The Gap between Knowledge and Practice Widens When Scientists Fail to Engage with the Ethical Implications of Their Own Work'. *BioEssays* 42 (6): 2000030. https://doi.org/10.1002/bies.202000030.

Wemelsfelder, Françoise. 1997. 'The Scientific Validity of Subjective Concepts in Models of Animal Welfare'. *Applied Animal Behaviour Science* 53 (1–2): 75–88.
https://doi.org/10.1016/S0168-1591(96)01152-5.

West-Eberhard, Mary-Jane. 2003. *Phenotypic Plasticity and Evolution*. Oxford: Oxford University Press.

Zelnik, Yuval R., Sorin Solomon, and Gur Yaari. 2015. 'Species Survival Emerge from Rare Events of Individual Migration'. *Scientific Reports* 5 (1): 7877.
https://doi.org/10.1038/srep07877.

Zohar, Opher, and Joseph Terkel. 1991. 'Acquisition of Pine Cone Stripping Behaviour in Black Rats (Rattus rattus)'. *International Journal of Comparative Psychology* 5 (1): 1–6.
https://doi.org/10.46867/C4KW2H.
i Interestingly, anthropomorphism is frequently debated in ethology (see among many others: Arbilly and Lotem 2017; Burghardt 2006; Horowitz and Bekoff 2007) but often treated as a caricatured scarecrow in philosophy.

ii It is true that IPM influences are moderate within the goals, as the survival and reproduction imperatives determine the general goal-direction. Nevertheless, IPM plays a role in shaping specific goals, reflecting individual differences.

iii For instance, the symbiosis between the squid and the bacterium *Vibrio fischeri* enables the squid to develop a luminescent organ it cannot develop otherwise (Pradeu 2011).

iv Nevertheless, there is a corpus of research investigating the cognitive processes, particularly those pertaining to memory, that underpin the phenomenon of agency in non-human animals (e.g. Toates 2004).
FIGURES

<figure><img src="image_2.png" /><figcaption>Figure 1. Properties of the agentive actions</figcaption></figure>

<figure><img src="image_3.png" /><figcaption>Figure 2. Core characteristics of biological agency (IPM)</figcaption></figure>
<figure><img src="image_4.png" /><figcaption>Figure 3. Autonomous organisation</figcaption></figure>

<figure><img src="image_5.png" /></figure>

<figure><img src="image_6.png" /><figcaption>Figure 3-b. Autonomous organisation without any other forms of agency</figcaption></figure>

<figure><img src="image_7.png" /></figure>
<figure><img src="image_8.png" /><figcaption>Figure 4. Goal-directed choice</figcaption></figure>

<figure><img src="image_9.png" /></figure>

<figure><img src="image_10.png" /><figcaption>Figure 5. Inventiveness</figcaption></figure>

<figure><img src="image_11.png" /></figure>
<figure><img src="image_12.png" /><figcaption>Figure 6. Autonomous organisation and evolution</figcaption></figure>

<figure><img src="image_13.png" /></figure>

<figure><img src="image_14.png" /><figcaption>Figure 7. Goal-directed choice and evolution</figcaption></figure>

<figure><img src="image_15.png" /></figure>
<figure><img src="image_16.png" /><figcaption>Figure 8. Inventiveness and evolution</figcaption></figure>

<figure><img src="image_17.png" /></figure>
CHARTS

<table><caption>Chart 1. Performance of the agentive action</caption><thead><tr><th>Performance of the agentive action</th><th>Autonomous organisation</th><th>Goal-oriented choice</th><th>Inventiveness</th></tr></thead><tbody><tr><th>Self-initiative</th><td>+</td><td>+</td><td>+</td></tr><tr><th>Locus of the direct impact</th><td>Agent</td><td>Relationship with the environment</td><td>Agent & its relationship with the environment</td></tr><tr><th>Control</th><td>+</td><td>++</td><td>+++</td></tr></tbody></table>

<table><caption>Chart 2. The role of IPM in different types of agency</caption><thead><tr><th>Agent</th><th>Autonomous organisation</th><th>Goal-oriented choice</th><th>Inventiveness</th></tr></thead><tbody><tr><th>Individuality</th><td>++</td><td>+</td><td>+</td></tr><tr><th>Playful flexibility</th><td>+</td><td>+</td><td>+++</td></tr><tr><th>Memory</th><td>+</td><td>+</td><td>+</td></tr></tbody></table>
**Figure 1.** A visual diagram titled “Figure 1. Properties of the agentive actions.” The diagram is structured as a network of four distinct approaches—Intuitive definition, Organisational approach, Ethological approach, and Evolutionary approach. Each approach lists several properties that define agentive actions, with arrows connecting similar or related properties across approaches. In the centre of the diagram, text emphasises that agentive actions are initiated by the agent itself, have consequences, and are within the agent’s control.

**Figure 2.** A visual diagram titled “Figure 2. Core characteristics of biological agency (IPM).” The diagram is structured similarly to the previous one, with four approaches—Intuitive definition, Organisational approach, Ethological approach, and Evolutionary approach. Individuality, Playful Flexibility, and Memory are highlighted in a green oval at the centre, connected by arrows to the different approaches. The overall design illustrates how biological agency integrates flexibility, memory, and individuality across various theoretical frameworks.

**Figure 3.** A visual diagram titled “Figure 3. Autonomous organisation.” The diagram is set within an orange oval representing Autonomous organisation, showing the developmental progression of a biological agent to a self-organising autonomous adult. On the left, a smaller circle labelled "Biological agent" contains the components I (Individuality), P (Playful flexibility), and M (Memory). Arrows indicate interactions between the agent and its environment, constraints, and internal goals. The middle section represents Development with key terms such as Playful flexibility, Adaptive improvisation, and Successive organisations in the course of development. To the right, the Self-organising autonomous adult section includes concepts like Closure, Organisation, Internal norms, and Internal goals. Black arrows mark external influences such as genetics and environmental constraints.

**Figure 3-b.** A visual diagram titled “Figure 3-b. Autonomous organisation without any other forms of agency”. The diagram displays the relationships between Autonomous organisation and the constitution of the Behavioural Repertoire and the Natural Goals. On the left, an orange section labelled Autonomous organisation contains smaller circles representing the development of a biological agent into a self-organising autonomous adult, influenced by IPM, genetics and the environment. To the right, a larger yellow oval labelled Behavioural repertoire highlights its uniqueness to the individual, its flexibility, and its historical dimension. Below it, the oval labelled Goals is connected to the Behavioural Repertoire through the Ability of choice. The Ability of choice depicted with dashed yellow arrows indicating that these choices may not be performed, leading to a separate black oval labelled Non-agentive actions, meaning that these actions are fully determined by genetics and/or the environment.
**Figure 4.** A visual diagram titled "Figure 4. Goal-directed choice". The diagram illustrates the relationships between Autonomous organisation, Behavioural repertoire, and Goals, framed within the concept of agency as Goal-directed choice. On the left, an orange oval represents Autonomous organisation, showing the development of a biological agent through environmental and genetic constraints. Arrows lead to the right, connecting the Autonomous organisation to both the yellow circle labelled Goals, and the one labelled Behavioural Repertoire. Goals and Behavioural Repertoire connects through the Ability of choice and then to Choice, all influenced by constraints. An arrow departs from Choice to connect to Autonomous organisation to indicate the effects of Choice on organisation.

**Figure 5.** A visual diagram titled "Figure 5. Inventiveness". The diagram explores agency as inventiveness in relation to the Autonomous organisation, and agency as Goal-directed choice. On the left, the orange section represents Autonomous organisation, with arrows connecting it to Goal-directed choice, shown in a smaller circle to the right. However the Ability of choice is labelled by dashed lines: the Ability of choice is not performed when agents invent. From Goals, dashed red lines lead to Invention, indicating that sometimes Goals play a role in Invention. Arrows depart from Invention to connect to Behavioural repertoire and Autonomous organisation to show the effects of Invention on those.

**Figure 6.** A visual diagram titled "Figure 6. Autonomous organisation and evolution.". The diagram depicts the evolution of organisations across three generations, when organisational processes affect evolution. On the left, the first generation is represented by an orange oval labelled Autonomous organisation, showing the development of a biological agent into a self-organising autonomous adult, influenced by IPM environmental and genetic constraints. An arrow labelled Heredity leads from this first generation to the second generation, represented by a green oval. The green oval depicts similar developmental processes with constraints and selective pressures impacting the second generation. A further arrow, again labelled Heredity, points to the third generation, represented by a yellow oval. Each generation builds upon the previous one, with selective pressures from the environment continuously influencing and influenced by the Autonomous organisation.

**Figure 7.** A visual diagram titled "Figure 7. Goal-directed choice and evolution.". The diagram depicts the evolution of goal-directed agents across three generations, when choices affect evolution. At the bottom left, the first generation is represented by a pink oval, labelled Goal-directed choice, showing the development of choice processes influenced by IPM, constraints, goals, and behavioural repertoire. Arrows representing heredity and selective constraints lead to the second generation, depicted in a blue-green oval in the centre. A yellow arrow links Choice and Heredity and another one links Choice with selective pressures, showing the effect of choice on the next generations. The second generation is linked in the same way to the third, shown in a red-orange oval at the top right. Each generation builds upon the previous
one, with heredity and pressures from the environment continuously influencing and influenced by the Choices.

**Figure 8.** A visual diagram titled "Figure 8. Inventiveness and evolution". The diagram illustrates the progression of inventive agents and goal-directed agents across multiple generations. Beginning on the left, the first generation is represented by a blue oval labelled Inventiveness, showing the process of invention emerging from IPM, influenced by environmental and genetic constraints. Purple arrows link Invention to Autonomous organisation, Heredity and selective pressures, showing the effects of Inventiveness on these different processes. Arrows lead to the second generation, depicted in a green oval labelled Goal-directed choice, and depicting an agent capable of choice. The same elements of choice, behavioural repertoire, and constraints continue, influenced by ecology and selective pressures. Arrows link choice with Autonomous organisation, Heredity and Selective pressures. A further arrow representing heredity points to the third generation, shown in a red oval representing again an agent with goal-directed choices, and similar processes. gothic third generation is linked to a the fourth generation depicted in a blue oval labelled Inventiveness. Purple arrows link the first oval to the selective pressures across all the generations, displaying the continuous effects of inventions on selective pressures.

**Chart 1.** A table titled "Chart 1. Performance of the agentive action". The table compares three types of agentive action: Autonomous organisation, Goal-oriented choice, and Inventiveness. These are assessed across three rows: Self-initiative, Locus of the direct impact, and Control. For Self-initiative, each action type is marked with a plus sign (+), indicating presence. For Locus of the direct impact, Autonomous organisation affects the Agent, Goal-oriented choice affects the Relationship with the environment, and Inventiveness affects the Agent and its relationship with the environment. For Control, Autonomous organisation is marked with one plus (+), Goal-oriented choice with two pluses (++), and Inventiveness with three pluses (+++), indicating increasing levels of control.

**Chart 2.** A table titled "Chart 2. The role of IPM in different types of agency". The table compares three types of agency: Autonomous organisation, Goal-oriented choice, and Inventiveness. These are assessed across three categories: Individuality, Playful flexibility, and Memory. For Individuality, Autonomous organisation is marked with two pluses (++), while Goal-oriented choice and Inventiveness are marked with one plus (+). For Playful flexibility, Autonomous organisation and Goal-oriented choice are marked with one plus (+), and Inventiveness is marked with three pluses (+++), indicating a higher emphasis on playfulness. For Memory, all three types of agency—Autonomous organisation, Goal-oriented choice, and Inventiveness—are marked with one plus (+), indicating their equal reliance on memory.