Both physiological and evolutionary criteria of biological individuality are underpinned by the idea that an individual is a functionally integrated whole. However, a precise account of functional integration has not been provided so far, and current notions are not developed in the details, especially in the case of composite systems. To address this issue, this paper focuses on the organisational dimension of two representative associations of prokaryotes: biofilms and the endosymbiosis between prokaryotes. Some critical voices have been raised against the (...) thesis that biofilms are biological individuals. Nevertheless, it has not been investigated which structural and functional obstacles may prevent them from being fully integrated physiological or evolutionary units. By contrast, the endosymbiotic association of different species of prokaryotes has the potential for achieving a different type of physiological integration based on a common boundary and interlocked functions. This type of association had made it possible, under specific conditions, to evolve endosymbionts into fully integrated organelles. This paper therefore has three aims: first, to analyse the organisational conditions and the physiological mechanisms that enable integration in prokaryotic associations; second, to discuss the organisational differences between biofilms and prokaryotic endosymbiosis and the types of integration they achieve; finally, to provide a more precise account of functional integration based on these case studies. (shrink)

The aim of this paper is to provide a theoretical framework to understand how multicellular systems realize functionally integrated physiological entities by organizing their intercellular space. From a perspective centered on physiology and integration, biological systems are often characterized as organized in such a way that they realize metabolic self-production and self-maintenance. The existence and activity of their components rely on the network they realize and on the continuous management of the exchange of matter and energy with their environment. One (...) of the virtues of the organismic approach focused on organization is that it can provide an understanding of how biological systems are functionally integrated into coherent wholes. Organismic frameworks have been primarily developed by focusing on unicellular life. Multicellularity, however, presents additional challenges to our understanding of biological systems, related to how cells are capable to live together in higher-order entities, in such a way that some of their features and behaviors are constrained and controlled by the system they realize. Whereas most accounts of multicellularity focus on cell differentiation and increase in size as the main elements to understand biological systems at this level of organization, we argue that these factors are insufficient to provide an understanding of how cells are physically and functionally integrated in a coherent system. In this paper, we provide a new theoretical framework to understand multicellularity, capable to overcome these issues. Our thesis is that one of the fundamental theoretical principles to understand multicellularity, which is missing or underdeveloped in current accounts, is the functional organization of the intercellular space. In our view, the capability to be organized in space plays a central role in this context, as it enables (and allows to exploit all the implications of) cell differentiation and increase in size, and even specialized functions such as immunity. We argue that the extracellular matrix plays a crucial active role in this respect, as an evolutionary ancient and specific (non-cellular) control subsystem that contributes as a key actor to the functional specification of the multicellular space and to modulate cell fate and behavior. We also analyze how multicellular systems exert control upon internal movement and communication. Finally, we show how the organization of space is involved in some of the failures of multicellular organization, such as aging and cancer. (shrink)

What are the prospects for a monistic view of biological individuality given the multiple epistemic roles the concept must satisfy? In this paper, I examine the epistemic adequacy of two recent accounts based on the capacity to undergo natural selection. One is from Ellen Clarke, and the other is by Peter Godfrey-Smith. Clarke’s position reflects a strong monism, in that she aims to characterize individuality in purely functional terms and refrains from privileging any specific material properties as important in their (...) own right. I argue that Clarke’s functionalism impairs the epistemic adequacy of her account compared to a middle-ground position taken by Godfrey-Smith. In comparing Clarke and Godfrey-Smith’s account, two pathways emerge to pluralism about biological individuality. The first develops from the contrast between functionalist and materialist approaches, and the second from an underlying temporal structure involved in using evolutionary processes to define individuality. (shrink)

This paper focuses on physiological integration in multicellular systems, a notion often associated with biological individuality, but which has not received enough attention and needs a thorough theoretical treatment. Broadly speaking, physiological integration consists in how different components come together into a cohesive unit in which they are dependent on one another for their existence and activity. This paper argues that physiological integration can be understood by considering how the components of a biological multicellular system are controlled and coordinated in (...) such a way that their activities can contribute to the maintenance of the system. The main implication of this perspective is that different ways of controlling their parts may give rise to multicellular organizations with different degrees of integration. After defining control, this paper analyses how control is realized in two examples of multicellular systems located at different ends of the spectrum of multicellularity: biofilms and animals. It focuses on differences in control ranges, and it argues that a high degree of integration implies control exerted at both medium and long ranges, and that insofar as biofilms lack long-range control (relative to their size) they can be considered as less integrated than other multicellular systems. It then discusses the implication of this account for the debate on physiological individuality and the idea that degrees of physiological integration imply degrees of individuality. (shrink)

The definition of biological individuality is one of the most discussed topics in philosophy of biology, but current debate has focused almost exclusively on evolution-based accounts. Moreover, several participants in this debate consider the notions of a biological individual and an organism as equivalent. In this paper, I show that the debates would be considerably enriched and clarified if philosophers took into account two elements. First, physiological fields are crucial for the understanding of biological individuality. Second, the category of biological (...) individuals should be divided into two subcategories: physiological individuals and evolutionary individuals, which suggests that the notions of organism and biological individual should not be used interchangeably. I suggest that the combination of an evolutionary and a physiological perspective will enable biologists and philosophers to supply an account of biological individuality that will be both more comprehensive and more in accordance with scientific practices. (shrink)

The concept of “autonomy”, once at the core of the original enactivist proposal in The Embodied Mind, is nowadays ignored or neglected by some of the most prominent contemporary enactivists approaches. Theories of autonomy, however, come to fill a theoretical gap that sensorimotor accounts of cognition cannot ignore: they provide a naturalized account of normativity and the resources to ground the identity of a cognitive subject in its specific mode of organization. There are, however, good reasons for the contemporary neglect (...) of autonomy as a relevant concept for enactivism. On the one hand, the concept of autonomy has too often been assimilated into autopoiesis and the implications are not always clear for a dynamical sensorimotor approach to cognitive science. On the other hand, the foundational enactivist proposal displays a metaphysical tension between the concept of operational closure, deployed as constitutive, and that of structural coupling ; making it hard to reconcile with the claim that experience is sensorimotorly constituted. This tension is particularly apparent when Varela et al. propose Bittorio as a model of the operational closure of the nervous system as it fails to satisfy the required conditions for a sensorimotor constitution of experience. It is, however, possible to solve these problems by re-considering autonomy at the level of sensorimotor neurodynamics. Two recent robotic simulation models are used for this task, illustrating the notion of strong sensorimotor dependency of neurodynamic patterns, and their networked intertwinement. The concept of habit is proposed as an enactivist building block for cognitive theorizing, re-conceptualizing mental life as a habit ecology, tied within an agent’s behaviour generating mechanism in coordination with its environment. Norms can be naturalized in terms of dynamic, interactively self-sustaining, coherentism. This conception of autonomous sensorimotor agency is put in contrast with those enactive approaches that reject autonomy or neglect the theoretical resources it has to offer for the project of naturalizing minds. (shrink)

Biological regulation is what allows an organism to handle the effects of a perturbation, modulating its own constitutive dynamics in response to particular changes in internal and external conditions. With the central focus of analysis on the case of minimal living systems, we argue that regulation consists in a specific form of second-order control, exerted over the core regime of production and maintenance of the components that actually put together the organism. The main argument is that regulation requires a distinctive (...) architecture of functional relationships, and specifically the action of a dedicated subsystem whose activity is dynamically decoupled from that of the constitutive regime. We distinguish between two major ways in which control mechanisms contribute to the maintenance of a biological organisation in response to internal and external perturbations: dynamic stability and regulation. Based on this distinction an explicit definition and a set of organisational requirements for regulation are provided, and thoroughly illustrated through the examples of bacterial chemotaxis and the lac-operon. The analysis enables us to mark out the differences between regulation and closely related concepts such as feedback, robustness and homeostasis. (shrink)

Individuality is an important concept in biology, yet there are many non-equivalent criteria of individuality expressed in different kinds of biological individuals. This article evaluates these different kinds in terms of their capacity to support explanatory generalizations over the systems they individuate. Viewing the problem of individuality from this perspective promotes a splitting strategy in which different kinds make different epistemic trade-offs that suit them for different explanatory roles. I argue that evolutionary individuals, interpreted as forming a functional kind, face (...) difficulties of individuation and explanatory power that are mitigated by relying on more structurally based properties and non-evolutionary kinds. 1Introduction2Kinds of Biological Individuals3Evolutionary Individuals and Functional Kinds4Evolutionary versus Non-evolutionary Kinds of Individuals 4.1Physiological individuality4.2Ecological individuality4.3Developmental individuality5Conclusion. (shrink)

Eusociality is broadly defined as: colonies consisting of overlapping generations, cooperative brood care, and a reproductive division of labour where sterile (or non-reproductive) workers help the reproductive members. Colonies of many complex eusocial insect species (e.g. ants, bees, termites) exhibit traits, at the collective level, that are more analogous to biological individuals rather than to groups. Indeed, due to this, colonies of the most complex species are typically a unit of selection, which has led many authors to once again apply (...) the concept of the superorganism to eusocial insects. However, unlike W. M. Wheeler, who originally employed the concept from a physiological and evolutionary perspective, today the superorganism is typically understood only from an evolutionary perspective, using group selection. This is because of the widely held view that eusocial colonies are self-organized systems. According to this view, even the most complex eusocial systems can be explained by appealing to a set of local interactions between parts of an initially disordered system (i.e. self-organization), without the need of any hierarchical control. In this paper we challenge the presupposition that hierarchical control and regulation is not possible in complex eusocial colonies. Using a case study of honey bees (Apis mellifera), we develop an alternative to the self-organization approach, that focuses on the hierarchical nature of the organization of complex eusocial systems—that we refer to as the hierarchical-organizational approach. In addition, we will analyse how colonies of eusocial insects show a complex set of interactions between the different organisms that bring forth a new cohesive collective organization, and how in turn the constitutive entities of this collective organization are transformed in this process. This paper argues that an inter-identity (namely the superorganism) emerges at the collective level in complex eusocial colonies, such as honey bees, due to the hierarchically-organized network of interactions within the colony. (shrink)

We argue that the transition from unicellular to multicellular systems raises important conceptual challenges for understanding agency. We compare several MC systems displaying different forms of collective behavior, and we analyze whether these actions can be considered organismically integrated and attributable to the whole. We distinguish between a ‘constitutive’ and an ‘interactive’ dimension of organizational complexity, and we argue that MC agency requires a radical entanglement between the related processes which we call “the constitutive-interactive closure principle”. We explain in detail (...) that this is not possible without a regulatory center functionally integrating the two dimensions, and we also argue that, in turn, this type of regulation would not be possible without a special type of organization between the cells required for the development and maintenance of systems capable of integrated behavior. (shrink)

Living systems employ several mechanisms and behaviors to achieve robustness and maintain themselves under changing internal and external conditions. Regulation stands out from them as a specific form of higher-order control, exerted over the basic regime responsible for the production and maintenance of the organism, and provides the system with the capacity to act on its own constitutive dynamics. It consists in the capability to selectively shift between different available regimes of self-production and self-maintenance in response to specific signals and (...) perturbations, due to the action of a dedicated subsystem which is operationally distinct from the regulated ones. The role of regulation, however, is not exhausted by its contribution to maintain a living system’s viability. While enhancing robustness, regulatory mechanisms play a fundamental role in the realization of an autonomous biological organization. Specifically, they are at the basis of the remarkable integration of biological systems, insofar as they coordinate and modulate the activity of distinct functional subsystems. Moreover, by implementing complex and hierarchically organized control architectures, they allow for an increase in structural and organizational complexity while minimizing fragility. Finally, they endow living systems, from their most basic unicellular instances, with the capability to control their own internal dynamics to adaptively respond to specific features of their interaction with the environment, thus providing the basis for the emergence of minimal forms of cognition. (shrink)

A crucial question for a process view of life is how to identify a process and how to follow it through time. The genidentity view can contribute decisively to this project. It says that the identity through time of an entity X is given by a well-identified series of continuous states of affairs. Genidentity helps address the problem of diachronic identity in the living world. This chapter describes the centrality of the concept of genidentity for David Hull and proposes an (...) extension of Hull’s view to the ubiquitous phenomenon of symbiosis. Finally, using immunology as a key example, it shows that the genidentity view suggests that the main interest of a process approach is epistemological rather than ontological and that its principal claim is one of priority, namely that processes precede and define things, and not vice versa. (shrink)

This paper criticizes the role of intuition-based ascriptions of cognition that are closely related to the ascription of mind. This practice hinders the explication of a clear and stable target domain for the cognitive sciences. To move forward, the proposal is to cut the notion of cognition free from such ascriptions and the intuition-based judgments that drive them. Instead, cognition is reinterpreted and developed as a scientific concept that is tied to a material domain of research. In this reading, cognition (...) becomes a changeable theoretical concept that can and must be adapted to the findings within this target domain. Taking humans as the best-established existing example of the relevant material target domain, this central case is extended to include all living systems. To clarify what it is about living systems that warrants their role as cognitive target domain, the new concept of cobolism is introduced as a complement to metabolism. Cobolism refers to the systematic ways in which each living system encompasses structures, processes and external events that maintain the fundamental metabolic processes that constitute the core of each living system. Cobolism is perfectly general, applies to bacterial and human cases alike, and provides a general format to describe wildly different cognitive organizations. It provides a clear target for the cognitive sciences to work on, turning them into what we can call the cognitive life sciences. (shrink)

The essay review summarizes the intention as well as some of the major topics from the book of A. Moreno and M. Mossio and discusses them against the background of recent considerations on the general understanding of organisms. The authors see themselves in the organicist tradition in biology and propose that a new understanding of living beings can be developed around the notion of organismic autonomy, which enables biological systems to maintain themselves in an environment through directed behavior.

Usually, the organization of living systems is explained by appealing to an intrinsic purpose that is based on the biological survival. However, paradigmatically, it is inevitable to observe that the final destiny of all living organisms is death. In this work, we defend that, from an organizational approach, there is a form of death—Regulatory Cell Death—that, far from being a mere "absence of life", is a process of biological regulation and a feature of self-maintenance in multicellular organisms.

In On the Origin of Autonomy; A New look at the Major Transitions in Evolution, Bernd Rosslenbroich argues that an increase of the relative autonomy of individual organisms is one of the central large-scale patterns in evolution. I begin by presenting how Rosslenbroich understands the notion of autonomy in biology and how he correlates its increase to different sets of morphological, physiological, and behavioral characteristics of various biological systems. I briefly discuss his view of directionality in evolution with respect to (...) its ontological and epistemological status. Then, I discuss the advantages of his thesis, and especially the emphasis on the organism as the subject rather than the object of evolution. I argue in detail that his account could benefit from a more exact conception of autonomy, and I discuss six problems related to the operationalization of his concept as applied to various theoretical issues. (shrink)