# Organization in Biology

Matteo Mossio

► To cite this version:

Matteo Mossio (Dir.). Organization in Biology. Springer International Publishing, 33, 2023, History, Philosophy and Theory of the Life Sciences, Philippe Huneman; Charles Wolfe; Thomas Reydon, 978-3-031-38967-2. 10.1007/978-3-031-38968-9. hal-04335078

HAL Id: hal-04335078

https://hal.science/hal-04335078v1

Submitted on 11 Dec 2023

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 CC BY 4.0 - Attribution - International License
Matteo Mossio Editor

Organization
in Biology

OPEN ACCESS

<figure><img src="image_2.png" /></figure>

Springer
# History, Philosophy and Theory of the Life Sciences

## Volume 33

### Series Editors

Philippe Huneman, Institut d'Histoire et de Philosophie des Sciences et des Techniques (IHPST, CNRS/Paris1), Paris, France

Thomas A. C. Reydon, Institute of Philosophy & CELLS, Leibniz Universität Hannover, Hannover, Germany

Charles T. Wolfe, Département de Philosophie & ERRAPHIS, Université de Toulouse Jean-Jaurès, Toulouse, France

### Editorial Board Members

Marshall Abrams, University of Alabama, Birmingham, AL, USA

André Ariew, University of Missouri, Columbia, MO, USA

Domenico Bertoloni Meli, Indiana University, Bloomington, IN, USA

Richard Burian, Virginia Tech, Blacksburg, VA, USA

Minus van Baalen, Institut Biologie de l'Ecole Normale Supérieure, Paris, France

Pietro Corsi, University of Oxford, Oxford, UK

François Duchesneau, Université de Montréal, Montreal, QC, Canada

John Dupre, University of Exeter, Exeter, UK

Paul Farber, Oregon State University, Corvallis, OR, USA

Lisa Gannett, Saint Mary's University, Halifax, NS, Canada

Andy Gardner, University of St Andrews, St Andrews, UK

Jean Gayon, UFR de Philosophie, Université Paris 1 Panthéon-Sorbonne, Paris, France

Guido Giglioni, University of Macerata, Civitanova Marche, Italy

Paul Griffiths, University of Sydney, Sydney, NSW, Australia

Thomas Heams, AgroParisTech, Paris, France

James G. Lennox, University of Pittsburgh, Pittsburgh, PA, USA

Annick Lesne, Sorbonne Université, Paris, France

Tim Lewens, University of Cambridge, Cambridge, UK

Edouard Machery, University of Pittsburgh, Pittsburgh, PA, USA

Alexandre Métraux, Archives Poincaré, Nancy, France

Hans Metz, Leiden University, Leiden, The Netherlands

Roberta L. Millstein, University of California, Davis, CA, USA

Staffan Müller-Wille, University of Cambridge, Cambridge, UK

François Munoz, Université Montpellier 2, Montpellier, France

Dominic Murphy, University of Sydney, Camperdown, NSW, Australia
Stuart A. Newman, New York Medical College, Valhalla, NY, USA

Frederik Nijhout, Duke University, Durham, NC, USA

Samir Okasha, University of Bristol, Bristol, UK

Susan Oyama, The City University of New York, New York, USA

Kevin Padian, University of California, Berkeley, CA, USA

David Queller, Washington University in St. Louis, St. Louis, MO, USA

Stephane Schmitt, Archives Poincaré, Nancy, France

Phillip Sloan, University of Notre Dame, Notre Dame, IN, USA

Jacqueline Sullivan, Western University, London, ON, Canada

Giuseppe Testa, University of Milan, Milan, Italy

J. Scott Turner, SUNY College of Environmental Science and Forestry,
Syracuse, NY, USA

Denis Walsh, University of Toronto, Toronto, ON, Canada

Marcel Weber, University of Geneva, Geneva, Switzerland
**History, Philosophy and Theory of the Life Sciences** is a space for dialogue between life scientists, philosophers and historians – welcoming both essays about the principles and domains of cutting-edge research in the life sciences, novel ways of tackling philosophical issues raised by the life sciences, as well as original research about the history of methods, ideas and tools, which constitute the genealogy of our current ways of understanding living phenomena.

The series is interested in receiving book proposals that • are aimed at academic audience of graduate level and up • combine historical and/or philosophical and/or theoretical studies with work from disciplines within the life sciences broadly conceived, including (but not limited to) the following areas: • Anatomy & Physiology • Behavioral Biology • Biochemistry • Bioscience and Society • Cell Biology • Conservation Biology • Developmental Biology • Ecology • Evolution & Diversity of Life • Genetics, Genomics & Disease • Genetics & Molecular Biology • Immunology & Medicine • Microbiology • Neuroscience • Plant Science • Psychiatry & Psychology • Structural Biology • Systems Biology • Systematic Biology, Phylogeny Reconstruction & Classification • Virology
The series editors aim to make a first decision within 1 month of submission. In case of a positive first decision the work will be provisionally contracted: the final decision about publication will depend upon the result of the anonymous peer review of the complete manuscript. The series editors aim to have the work peer-reviewed within 3 months after submission of the complete manuscript. The series editors discourage the submission of manuscripts that contain reprints of previously published material and of manuscripts that are below 150 printed pages (75,000 words). For inquiries and submission of proposals prospective authors can contact one of the editors: Charles T. Wolfe: ctwolfe1@gmail.com Philippe Huneman: huneman@wanadoo.fr Thomas A.C. Reydon: reydon@ww.uni-hannover.de
Organization in Biology

<figure><img src="image_3.png" /></figure>

Springer
Editor

Matteo Mossio

Institut d'Histoire et de Philosophie
des Sciences et des Techniques

(IHPST, CNRS/Paris1)

Paris, France

<figure><img src="image_4.png" /></figure>

ISSN 2211-1948

ISSN 2211-1956 (electronic)

History, Philosophy and Theory of the Life Sciences

ISBN 978-3-031-38967-2

ISBN 978-3-031-38968-9 (eBook)

https://doi.org/10.1007/978-3-031-38968-9

© The Editor(s) (if applicable) and The Author(s) 2024. This book is an open access publication.

**Open Access** This book is licensed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license and indicate if changes were made.

The images or other third party material in this book are included in the book's Creative Commons license, unless indicated otherwise in a credit line to the material. If material is not included in the book's Creative Commons license and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder.

The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use.

The publisher, the authors, and the editors are safe to assume that the advice and information in this book are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the editors give a warranty, expressed or implied, with respect to the material contained herein or for any errors or omissions that may have been made. The publisher remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

This Springer imprint is published by the registered company Springer Nature Switzerland AG

The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland

Paper in this product is recyclable.
# Contents

1 **Introduction: Organization as a Scientific Blind Spot** 1
Matteo Mossio

2 **"Organization": Its Conceptual History and Its Relationship to Other Fundamental Biological Concepts** 23
Georg Toepfer

3 **Varieties of Organicism: A Critical Analysis** 41
Charles T. Wolfe

4 **Judging Organization: A Plea for Transcendental Logic in Philosophy of Biology** 59
Gertrudis Van de Vijver and Levi Haeck

5 **On the Organizational Roots of Bio-cognition** 85
Cliff Hooker

6 **Does Organicism Really Need Organization?** 103
Olivier Sartenaer

7 **Organisms: Between a Kantian Approach and a Liberal Approach** 127
Philippe Huneman

8 **The Fourth Perspective: Evolution and Organismal Agency** 159
Johannes Jaeger

9 **On the Evolutionary Development of Biological Organization from Complex Prebiotic Chemistry** 187
Kepa Ruiz-Mirazo and Alvaro Moreno

10 **Organization and Inheritance in Twenty-First-Century Evolutionary Biology** 219
Gaëlle Pontarotti
<table><tr><td>11</td><td>There Are No Intermediate Stages: An Organizational<br/>View on Development</td><td>241</td></tr><tr><td></td><td>Leonardo Bich and Derek Skillings</td><td></td></tr><tr><td>12</td><td>Modeling Organogenesis from Biological First Principles</td><td>263</td></tr><tr><td></td><td>Maël Montévil and Ana M. Soto</td><td></td></tr><tr><td>13</td><td>From the Organizational Theory of Ecological Functions<br/>to a New Notion of Sustainability</td><td>285</td></tr><tr><td></td><td>Charbel N. El-Hani, Felipe Rebelo Gomes de Lima,<br/>and Nei de Freitas Nunes-Neto</td><td></td></tr><tr><td></td><td>Index</td><td>329</td></tr></table>
Chapter 1

# Introduction: Organization as a Scientific
Blind Spot

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

Check for
updates

Matteo Mossio

**Abstract** For most of the twentieth century, biology forgot or largely neglected organization. By this term, I mean a certain mode of interaction among the parts of a system, which is by hypothesis distinctively realized by biological systems. While a systemic trend is progressively pervading various biological fields – notably Evolutionary Biology, Systems Biology and Origins of Life – I suggest that organization still remains a blind spot of biological thinking. Therefore, I submit, biology should be enriched by an explicit and specific notion of organization, drawing in particular on the theory of autonomy, of which I recall some central tenets. I conclude with a brief overview of the scientific and philosophical tradition which has explicitly elaborated on biological organization, and of the more recent literature to which this book aims to contribute.

## 1.1 The Neglect of Organization

For most of the twentieth century, biology forgot or largely neglected organization. Since the establishment of the Modern Synthesis in evolutionary biology in the 1930s and 1940s, and the flourishing of molecular biology in the 1950s and 1960s, biological research has focused almost exclusively on entities described at the population or molecular level, by adopting what is usually called “genocentrism,” the perspective that places strong emphasis on genes as the fundamental determinants of biological phenomena (Rosenberg, 2007; Fox Keller, 2000).

As Gilbert and Sarkar have pointed out, “for most of this century, the major project of biology has been to reinterpret living properties as being epiphenomena of genes” (Gilbert & Sarkar, 2000: 5).

M. Mossio (✉)

Institut d'Histoire et de Philosophie des Sciences et des Techniques (IHPST, CNRS/Paris1),
Paris, France

e-mail: matteo.mossio@univ-paris1.fr
The Modern Synthesis reconciled Darwin's theory of natural selection with Mendelian genetics through population genetics and put forward a reconceptualization of evolution as the change in allele frequencies in a population (Dobzhansky, 1937). Later, with the discovery of the structure of DNA in 1953, molecular biology underwent a comprehensive research program aimed at studying how the genetic program – a notion introduced simultaneously by Ernst Mayr on the one hand, and François Jacob and Jacques Monod on the other (Mayr, 1961; Monod & Jacob, 1961) – governs the synthesis of macromolecules and their interactions and, therefore, cell activity and functionality. The so-called “central dogma of molecular biology,” i.e., the idea that biological information flows unidirectionally from DNA to proteins, was thereby interpreted as the idea that genes are the primary (if not sole) determinants of form, function, and behavior of organisms (Crick, 1966). In both evolutionary and molecular biology, genocentrism has therefore consisted in a reductionist stance, resulting in the neglect of biological organization as such.

By the term “organization” I mean a certain mode of interaction among the parts of a system, distinctively realized by biological systems, when compared to other kinds of natural systems, or to artifacts. Broadly speaking (I will return to this below), organization refers to a regime in which a set of entities happen to be related to each other so as to constitute a system that displays both functional differentiation and integration. Moreover, the activity of the whole system plays a role in maintaining its constituents over time: organized systems self-maintain. Let me point out right away that organization is typically, but not exclusively realized by organisms. For instance, it might be argued that colonies, symbioses, or, at a higher level of description, ecosystems can be described as organized systems, although they would not necessarily count as organisms. Accordingly, the notion of “organization” and that of “organism” should not be straightforwardly conflated, although they are closely related: organisms are organized systems, but organized systems are not necessarily organisms.

## 1.2 Organization as an Explanandum and an Explanans of Biology

Before reductionist and genocentric approaches became mainstream, many eminent biologists and philosophers put emphasis on the centrality of organization in biology. For instance, in his *Modern Theories of Development* (1933), Ludwig von Bertalanffy argued that “all vital processes are so organized that they are directed to the maintenance, production, or restoration of the wholeness of the organism.” Therefore, he writes, “there is no ‘living substance’ because the characteristic of life is the organization of substances” (von Bertalanffy, 1933: 8).

At first sight, one might think that any biologist would easily agree with Bertalanffy. In a sense, it seems obvious that biological systems are organized and, hence, that biology should deal with their organization, its general principles and its
various concrete manifestations. After all, biology emerged at the turn of the nine-
teenth century precisely as the science of “organized beings” (Gambarotto, 2018).
Yet, several authors have noted that twentieth-century genocentric biology has lost
sight of organisms (Laublicher, 2000; Huneman & Wolfe, 2010; Nicholson, 2014;
Walsh, 2015) and thereby, I hold, of organization. What does this alleged neglect of
organization consist in? Philosophically, I submit – following Huneman (2010) –
that it has taken two different forms, related to the *explanandum* (what must be
explained) and the *explanans* (what explains) of biology, respectively.

Firstly, organization is not the *explanandum* of genocentric biology; at best, the
latter takes organization as a vague object of explanation. Indeed, the Modern
Synthesis has been classically criticized because of its *atomization* of organisms,
i.e., the fact of explaining biological traits separately as adaptations, without
accounting for their integration as organized systems (Gould & Lewontin, 1979). As
Huxley famously put it “every organism cannot be other than a bundle of adapta-
tions” (Huxley, 1942: 20). Even though some advocates of the Modern Synthesis
such as Mayr and Dobzhansky did disagree with such a radical view, it seems only
fair to claim that the organized complexity of biological systems is not what this
framework was designed to explain. Modern Synthesis’ main explanandum is the
evolution of adaptive traits as the result of differential selection acting on genes. In
the case of molecular biology, it could certainly be argued that organization is the
ultimate explanandum, insofar as the study of the parts taken separately is supposed
to lead, in the long run, to an explanation of the whole system. Nevertheless, such a
remote objective remained largely out of reach throughout the actual development
of the discipline, which has seemed de facto unable to provide a molecular under-
standing of biological organization in all its characteristic complexity.

Secondly, organization does not play any explanatory role in genocentric biol-
ogy. If it were the case, explanations would presuppose the fact that biological phe-
nomena occur *because* biological systems are organized, and would not aim at
explaining organization as such, by relying on more fundamental principles. Biology
would explain phenomena in the light of their organizational nature. In evolutionary
biology, the theoretical framework of the Modern Synthesis identifies natural selec-
tion as the main (and often sole) *explanans*, while the impact of organization in the
evolutionary dynamics of biological systems is seldom taken into account. As for
molecular biology, the rejection of organization as the explanans relies on more
explicit theoretical reasons. Indeed, molecular genocentrism has advocated the idea
that the specificity of biological phenomena is the fact that they are the result of the
expression of a genetic program. Genes code for protein synthesis which, in turn,
contributes to the realization of biological functions and, ultimately, of the whole
organization. Accordingly, organization itself is explained by the expression of the
genetic program (Mayr, 1961; Jacob, 1973).

It is worth noting that genocentrism has also influenced another research domain, that of the origins of life. Influenced by the flourishing of molecular biology, many researchers in this field have defended the idea that life on earth was initiated by the appearance of the first replicators, i.e., the first self-replicative molecules (the “first genes”) (Pereto, 2005). On the one hand, organization should constitute – as it does
for molecular biology – the ultimate explanandum, to the extent that self-replicative molecules are supposed to trigger an evolutionary process leading to the emergence of living systems endowed with organized complexity. Here again, however, the “replication first” approach has had a hard time in providing satisfactory explanations of the emergence of organized complexity. On the other hand, the “replication first” approach has explicitly excluded organization from the explanans, which seems at first sight an obvious move for a discipline that aims at explaining the *emergence* of life. Organized complexity cannot be adequately explained by appealing to an explanans that already implies organization. Rather, what could explain the emergence of organization would be the inherent capacity of (a population of) self-replicative molecules to generate more and more complex systems by evolving through natural selection – assuming that natural selection can operate before organized systems exist.

In recent times, genocentrism has been increasingly criticized. As it has been previously highlighted (Gilbert & Sarkar, 2000; Bateson, 2005), biology is paying more and more attention to the fact that biological phenomena should be studied by taking into account their distinctive complexity. Accordingly, a general trend toward a more organicist – or at least systemic – perspective can be easily perceived, not only in biology but also in related fields, like prebiotic chemistry. Importantly, recent organicist approaches challenge genocentrism not only on the side of the explanandum but also, and somehow more radically, on the side of the explanans: biology should be more inclusive and ambitious with the kind of phenomena to be explained and, at the same time, equip itself with more adequate explanatory tools.

## 1.3 The Anti-reductionist Trend in Evolutionary Biology

One of the fields in which an anti-reductionist reaction to genocentrism has been explicitly invoked is evolutionary biology, in the context of a lively debate about whether evolutionary theory needs a rethink (Laland et al., 2014). The debate puts into question the mainstream conceptual framework of the modern synthesis, leading to the call for an “extended evolutionary synthesis” (Pigliucci & Müller, 2010; Laland et al., 2015; Huneman & Walsh, 2017). The extended evolutionary synthesis puts emphasis on four key research themes whose implications, in the view of its advocates, converge on a different, “extended” understanding of the main factors that determine evolutionary change: constructive development, phenotypic plasticity, niche construction, and inclusive inheritance.

Constructive development refers to the idea that the development of organisms does not result from the execution of a genetic program but, rather, from multiple interactions between many factors within the developing system, as well as between the system and the environment (Oyama, 1985). Such a view has evolutionary implications, insofar as development is conceived as a process that facilitates the emergence of phenotypic variation (within and between individuals, populations, species, etc.) on which selection may operate (Minelli & Fusco, 2008). The classical
conception of “developmental bias” that reduces the range of possible variations is replaced by a vision according to which development produces phenotypic novelty and affects evolutionary trajectories (Gerhart & Kirschner, 2007).

Phenotypic (or developmental) plasticity is the capacity of a given organism to generate different phenotypes to adapt to different environmental conditions. Plasticity can be seen as a generalization of constructive development: as such, it is meant to lead to phenotypic innovations and, thereby, to play a role in modulating adaptive evolution. In particular, a lively debate exists on whether – and if so, how – phenotypic plasticity can be consolidated by means of genetic accommodation, beyond phenotypic accommodation. As a consequence, adaptive phenotypic novelties would induce genetic changes in populations, and not vice versa (West-Eberhard, 2003).

Niche construction refers to all those processes through which the organism systematically modifies the environment and, thereby, the selective pressures acting on itself. Typical examples are the construction of dams, nests or webs, or the active intervention in the composition or distribution of nutrients in the environment (Odling-Smee et al., 2003).

Inclusive (or extended) inheritance challenges the idea according to which genes are the sole factors involved in biological inheritance. Rather, increasing experimental evidence shows that trans-generational similarities also rely on a variety of transmission processes that do not seem to require a genetic ground. Examples that are often mentioned include epigenetic, symbiotic, ecological, and cultural inheritance. The main implication for evolutionary biology consists in the fact that nongenetic inheritance would allow the transmission of acquired characteristics, thus being responsible for adaptive variations (Jablonka & Lamb, 1995, 2005).

Taken together, these recent research themes are promoting a shift from a genetic-
to an organismic-centered view of evolution, insofar as each of them focus on phe-
nomena with respect to which explanations relying on genetic determinism do not
appear relevant or adequate. The general idea is that a better understanding of evo-
lutionary trajectories requires taking into account a number of capacities and pro-
cesses that cannot be described and explained by appealing only to genetic factors.
Rather, biological systems as wholes become the relevant object with respect to
which phenomena as development, plasticity, niche construction, and extended
inheritance can be described and explained. When compared to the conceptual
framework of the Modern Synthesis, it seems therefore clear that the organization
of biological systems is being brought back to the foreground by the extended evo-
lutionary synthesis, both as an explanandum and as an explanans (Nicholson, 2014).
On the one hand, the EES pays more attention to phenomena that reflect the inte-
grated activity of organized systems and their complexity; on the other hand, the
explanatory strategy does not consist in deriving organization from more fundamen-
tal principles but, rather, in relying on organized systems (including, of course,
organisms) and their characteristics in order to elaborate more adequate explana-
tions of these phenomena.

There is therefore no doubt that the EES is more attentive to the organized com-
plexity of the entities which are involved in evolutionary processes, like other
authors remarked (Callebaut et al., 2007;<fnref n="1" /> Wagner & Laublicher, 2000). Nevertheless, I submit, a conceptual characterization of biological organization as such has not yet been elaborated. In other words, the EES does not rely on a characterization of what is an organized system, which rather consists in a particular *level of description* at which phenomena are described and explanations are provided. The fact that the systemic shift is not accompanied by a characterization of organization has important implications for the explanatory scope and power of evolutionary biology. Some of the contributions to this book explore these implications. Here, I want to emphasize that, in spite of the increasing focus on it, organization remains a blind spot from the current EES views.

## 1.4 The Anti-reductionist Trend in Systems Biology

The systemic trend is also explicit in molecular biology with the emergence, in the early 2000s, of systems biology. Broadly speaking, systems biology was promoted in reaction to the genocentrism of classic molecular biology, as an approach that aims at shifting the view from genes to the larger systems in which genes are embedded. Rather than studying how gene expression generates biological complexity, systems biology focuses on how the biological system (typically, a cell) works and, in particular, how it regulates gene expression itself. It is now quite common to make a distinction, initially proposed by O'Malley and Dupré (2005), between two attitudes within systems biology: a “pragmatic” one and a “theoretic” one.

The first attitude includes the great majority of research in the field. Pragmatic systems biology can be described as an extension of molecular biology, which studies the interactions and dynamics of large molecular networks. Pragmatic (or “molecular”) systems biology challenges genocentrism by considering large systems within which genes are a (crucial) component among many others. Different sets of molecular components (typically biopolymers) belonging to specific cellular organisms, under specific environmental conditions, are studied by different sub-fields: e.g., full cell genomes, as studied by genomics, the whole pool of RNA transcripts (awaiting translation, at a given time) by transcriptomics, the diverse proteins operating in the system by proteomics, metabolites by metabolomics, membrane lipids by lipidomics, and so on. Collectively, these sub-fields generate huge datasets that systems biologists are nowadays trying to interpret and integrate by using mathematical models and computer simulations (Nicholson, 2014: 352).

In spite of these innovations, systems biology can be said to place itself in continuity with the reductionist bottom-up strategies of classical molecular biology, insofar as the general objective consists in obtaining knowledge about the structure, function, organization, and dynamics of whole biological systems by elaborating

<footnote n="1">¹Callebaut, Müller, and Newman, for instance, propose an “Organismic Systems Approach” to Evo-Devo, which explicitly elaborates on biological organization as a core notion.</footnote>
models that integrate the parts and their relations (Saetzler et al., 2011: 3). Accordingly, the explanatory strategy operative in the field assumes organization as something that must be explained, but that does not play any explanatory role. More generally, as it has been pointed out by O'Malley and Dupré, molecular systems biology relies on a generic and theoretically ungrounded notion of “system,” which designates the network of interacting molecules of different kinds. The central question is whether molecular systems biology can succeed in providing an adequate understanding of organized complexity through such a bottom-up explanatory strategy. Recently, several researchers have expressed skepticism in this respect (Mesarović & Sreenath, 2006; Bertolaso, 2011; Saetzler et al., 2011; Noble, 2017), arguing that the quantity and complexity of available data make more and more difficult their interpretation and integration. Data – the criticism points out – do not speak by themselves, and biologists are in trouble in asking them the relevant questions so as to get an adequate understanding of the whole organization.

This is where theoretical systems biology steps in. Instead of producing models that include more and more experimental data, theoretical systems biology looks for what Green and Wolkenhauer call “organizing principles” (Green & Wolkenhauer, 2013), which are used to select relevant data. One may say that while pragmatic systems biology aims at getting knowledge by adding more details, theoretical systems biology pursues the same objective by abstracting from details. The principles on which theoretical systems biology focuses are mathematical descriptions of recurrent constraints, relations, and patterns that are similar (“isomorphic”) in different systems, not necessarily or exclusively biological. A classic example is homeostasis, the capacity to maintain an internal steady state in spite of external perturbation (Cannon, 1929). Other organizing principles have been applied to explain phenomena as flows and oscillations (see for instance the classical work of Goodwin, 1963, and recent developments). A milestone in this respect is the first mathematical model of the heart rhythm (Noble, 1962).

When compared to the pragmatic approach, theoretical systems biology makes a further step in challenging the reductionist perspective. Systemic principles are understood as general hypotheses, which means that they are supposed to explain the data (top-down) and not be explained by them (bottom-up). For instance, if one elaborates a model for accounting for a homeostatic behavior of a biological system, the question would not be “why is the system homeostatic?”, but rather “how does the system manage to maintain that specific variable steady, given its (hypothetical) homeostatic capacities”? Accordingly, the notion of “system” is theoretically enriched, and its explanatory role enhanced by the expression of the principles.

Yet those principles are not meant to be distinctive to the biological realm. They are usually elaborated and formalized in other domains, as engineering and graph theory. For instance, negative feedbacks were formalized by cybernetics to account for homeostatic behavior in both animal and machines (Wiener, 1948). The transdisciplinary application of the principles is taken to be a fruitful explanatory strategy, which allows getting insights into the properties of biological systems by looking for analogies with physical systems or machines.
The strategy has proven successful and could certainly foster the acquisition of further knowledge. Nevertheless, it raises the question whether a full-fledged theoretical systems biology can rely on principles that are not designed to capture specific biological features, but more general systemic ones. While it is certainly true that biological systems are a class of physical systems, and therefore share with them (including artifacts and machines) common principles, a satisfactory theoretical framework of biological complexity should also look for *distinctive* biological principles.

One of the objectives of this book consists precisely in promoting the idea that theoretical biology should be endowed with a distinctive principle of organization, which would characterize biological complexity as such, above and beyond the different systemic principles that account for some of its features.

## 1.5 The Anti-reductionist Trend in the Origins of Life

In the origins of life field, the anti-reductionist alternative to the “replication first” view, which assumes that the individuals of an evolving population can be bare molecules, has taken the form of what is usually called the “metabolism first” view (Pereto, 2005). According to this perspective, the relevant starting point of the emergence of life is the spontaneous appearance of primitive self-catalytic metabolic networks, which would be a condition for the subsequent synthesis of replicators and genes.

The central objection of the “metabolism first” perspective to the “replication first” one is that a process of evolution by natural selection faces “bottlenecks” when it starts from a population of “naked” molecular species. In sharp contrast to the underlying assumptions of the competing view, the anti-reductionist approach argues that molecular replicators alone cannot generate the relevant kind of complexity leading to the appearance of life in the form of organized chemical systems.

Accordingly, the anti-reductionist view challenges the reductionist one with respect to the *explanans* that is adopted. Although it might seem obvious, the idea that we should not presuppose organization to explain the emergence of life (given that organization is taken as an inherent characteristic of life) leads to an impasse: if the *explanans* is too simple (in terms of its complexity), it cannot generate entities that are complex enough for our explanatory purposes. That is why the “metabolism first” approach does presuppose organization as an *explanans*, under the general hypothesis that some degree of organized complexity is actually required to boot-strap an evolutionary process leading to the appearance of living systems as we know them (Hordijk et al., 2011).

The general strategy consists in characterizing chemical systems (usually referred to as “protocells” or “proto-organisms,” Rasmussen et al., 2008) that can appear spontaneously in plausible prebiotic environments and are endowed with the capacity of dynamically self-maintain and increase their functional complexity. Of course, organization cannot be the *explanans* of this very initial phase, in which
organized systems emerge from the integration of different kinds of preexisting
processes and components. Once these systems have appeared, however, the pro-
cess toward primitive living cells implies a set of intermediate forms of organiza-
tion, each playing a role in the emergence of the next, more complex one. During
that long process, thus, each form of organization plays both the role of *explanans*
of the next one and of *explanandum* of the previous one.

Although the systemic perspective on the origins of life is taking momentum,
after the flourishing of “systems chemistry” as a research field important criticisms
have also been addressed to it (Vasas et al., 2010), which explains that the debate is
still lively in the field. One important issue is that – here again – the very notion of
organization is not expressed in explicit conceptual and theoretical terms and, there-
fore, does not provide a sufficiently precise guidance in the elaboration of relevant
protocells models (see Ruiz-Mirazo et al., 2017, for a discussion of this issue and a
proposal).

1.6 What Is Organization?

The central objective of this book is to make a contribution to the current anti-
reductionist trend in biology, by putting organization to the foreground. I submit
that the systemic, or even organicist-thinking that is progressively pervading various
biological fields should be enriched by an explicit and specific notion of organiza-
tion, understood as both a fundamental explanandum and explanans in biology.

Adopting organization as an explanandum means that the object of biological
explanation should be the phenomenon of organization itself in its various realiza-
tions, as well as its evolution. A satisfactory biological explanation should aim at
making explicit how any specific phenomenon under scrutiny is to be understood as
a manifestation or an aspect of organized complexity and, for this precise reason,
biological. Beyond the fragmentation of reductionist approaches, anti-reductionist
ones should hence aim at explaining biological phenomena by locating them into
integrated organized wholes.

Adopting organization as an explanans, in turn, means conceiving it as a *theoreti-
cal principle* (Mossio et al., 2016). A theoretical principle is an overarching hypoth-
esis that frames the intelligibility of the objects within a scientific domain. All
biological systems, in all their diversity and richness of forms and kinds, comply
with the principle, and are therefore organized. The crucial implication here is that
theoretical principles enable explanations, but are not themselves the object of an
explanation. Accordingly, the adoption of organization as an explanans means that
biology presupposes the principle without trying to deduce it from something else,
which would be precisely the reductionist stance.

A recent example of the adoption of organization as both an explanandum and an
explanans is given by the model of a self-maintaining metabolic system, inspired by
Robert Rosen's idea of (M,R)-systems (Piedrafita et al., 2010). The model is a com-
putational simulation of a chemical network, made of three interlocking catalytic
cycles, in which catalysts are produced by the very network that they contribute to maintain. Here, organization is the explicit explanandum in the sense that the model explores the conditions under which the system as a whole exhibits properties of organized – and therefore biologically relevant – systems, such as steady autoca-talysis, robustness, and bi-stability. At the same time, organization is an explanans to the extent that the mutual dependence between the catalysts is presupposed as a background hypothesis and not obtained as a result. The model does not try to explain how and why catalysts get organized in the first place (i.e., mutually depen-dent); rather, it takes the very fact of being organized as an unexplained premise (an explanatory principle) and aims at explaining the properties of a system that realizes a specific instantiation of biological organization.

It is worth reflecting on this apparent contradiction, stemming from the double role of organization as both the explanandum and the explanans of biology. In the two cases, the term designates in fact different conceptual entities: as an explanan-dum, organization refers to its various concrete realizations in nature; as an explan-ans, it designates the general “regime” or “set of relations” that are common to all realizations. To make a (somehow very perilous) analogy, the dual role of organiza-tion as explanans and explanandum can be grasped by thinking to the role played by Newton’s laws of motion in Classical Mechanics. On the one hand, the laws of motion are principles that are presupposed (i.e., not explained within the field), so as to provide explanation of physical phenomena; on the other hand, Classical Mechanics provide explanations of phenomena which realize (are in conformity with) the laws. Classical Mechanics presuppose the laws of motion so as to explain specific instantiations of these laws. In this very general sense, I submit that organi-zation could play an analogous role in the biological domain.

Needless to say, for organization to play such a role, it should be characterized in a way that is more precise than the general and intuitive notions of “system” or even “organism” as they are currently employed in evolutionary and systems biology. So, the main question is: what does organization mean? To answer this question, let me spell out some guidelines that will be explored in the book. These guidelines are mostly inspired by the theory of biological autonomy, one of the contemporary heirs of the organicist tradition, to which I have myself contributed (Moreno and Mossio, 2015).

As already mentioned, organization designates a specific kind of complexity, a specific set of relations among elements. To a first approximation, organization refers to the differentiation of functional roles (i.e., division of labor) among the parts of a system and, at the same time, to their integration and coordination as a whole. Furthermore, and crucially, organization involves a generative dimension in the form of a mutual dependence, such that the very activity and existence of each organized part depends on its mutual relationship with the others. Organized parts have functions (Mossio et al., 2009; Saborido et al., 2011), which means that organization most basically designates *functional* complexity. Overall, biological organization is capable of self-determination, insofar as functional constraints collectively contribute to determine their conditions of existence. As I have argued
elsewhere (Mossio & Bich, 2017), the capacity of self-determination provides a
naturalized ground for purposiveness: biological organization can be legitimately
said to be cause and effect of itself and thereby an intrinsically purposive regime.

Biological organization goes along with thermodynamic openness, which is the
fact that organized systems continuously exchange energy and matter with the sur-
roundings. The connection is theoretically deep, insofar as only thermodynami-
cally open systems can possibly comply with the organization principle, although
not any thermodynamically open system does. As all open systems, indeed, be they
physical or chemical, biological systems are traversed by a flow of energy and mat-
ter, which takes the form of processes and reactions occurring in open thermody-
namic conditions. In this respect, biological systems do not differ from other
natural open systems. Yet, unlike “self-organizing” dissipative structures, they con-
strain and canalize the thermodynamic flow through the collective activity of their
functional parts, which realize a specific form of mutual relationship, i.e.,
organization.

Because of their distinctive functional complexity, furthermore, organized systems (usually) do not appear spontaneously when some specific boundary conditions are met, as self-organizing structures do. Rather, organized systems are the result of a long historical evolutionary process of increase and preservation of complexity, which means in particular that any individual biological system is generated through the reproduction of other biological systems. In spite of their common thermodynamic grounding, hence, (biological) organization cannot be conflated with (physical) self-organization.

As the result of an historical process, biological complexity raises the central
question of understanding how biological systems manage to maintain their stabil-
ity while continuously undergoing variation (Montévil et al., 2016a; Longo et al.,
2012, argue that biological variation is theoretically unprestatable). In this respect,
I submit that the notion of organization plays a twofold explanatory role: on the one
hand, organization allows explaining the stability of biological systems (both at the
individual and evolutionary scale), and the maintenance of their constitutive dynam-
ics over time; on the other hand, it provides a ground to understand how quantitative
and qualitative innovations can be produced, and then preserved though its func-
tional integration.

Lastly, let me emphasize again that biological organization should be conceived
as a broader notion than that of “organism.” Although organisms are – by hypothe-
sis – organized systems, not all organized systems are necessarily organisms. For
instance, some contributions to this book explore the idea that ecosystems might
also be described as organized systems, without implying that they constitute a kind
of organisms. Reciprocally, additional specifications should be added to characterize
organisms among the broader set of organized systems. In this respect, a possible
route is traced by the theory of autonomy, according to which organisms are autono-
mous systems, i.e., organized systems endowed with agential and adaptive capaci-
ties (see for instance Moreno and Mossio, 2015, section 4.4). As autonomous
systems, in a word, organisms are organized adaptive agents.
## 1.7 Historical Overview

The historical roots of the notion of biological organization I refer to can be traced back to Immanuel Kant. In his *Critique of the Power of Judgment* (1790/1987, Kant argues that, unlike any other kind of system, the parts of biological systems do not and cannot exist by themselves, but only insofar as they constitute an organized whole which, in turn, is itself a condition of their own existence and functioning. In this sense, biological systems display self-organizing features that are absent in machines. In a watch, for example, every part is organically arranged in relation to the others, but the watch does not *produce* them. It “is certainly present for the sake of the other but not because of it.” Hence the producing cause of the watch is the watchmaker, not the watch itself: “one wheel in the watch does not produce the other, and even less does one watch produce another, using for that purpose other matter (organizing it); hence it also cannot by itself replace parts that have been taken from it, or make good defects in its original construction by the addition of other parts, or somehow repair itself when it has fallen into disorder: all of which, by contrast, we can expect from organized nature.” Based on these considerations, Kant claims that “an organized being is thus not a mere machine, for that has only a *motive* force, while the organized being possesses in itself a *formative force* (Bildungskraft), and indeed one that it communicates to the matter, which does not have it (it organizes the latter): thus it has a self-propagating formative power, which cannot be explained through the capacity for movement alone (that is, mechanism)” (Kant, 1790/1987: §65).

The Kantian focus on biological organization had continuity in the (mostly Continental) Biology of the nineteenth century, notably in the work of Goethe (1995) and Cuvier (1817). Cuvier’s principle of the “condition of existence,” for instance, claims that “the different parts of each being must be coordinated in such a way as to render possible the existence of the being as a whole” (1817 i., 6, quoted and translated by Reiss, 2009). By implying that the different parts are linked and coordinated, Cuvier’s principle grounds and guides his empirical investigations in comparative anatomy and paleontology (Cuvier, 1805; see also Huneman, 2006, for an analysis).

Kant’s and Cuvier’s perspectives further influenced German organicist tradition leading to Johannes Müller’s physiology (1837–1840) and Karl von Baer’s embryology (1828). They both consider that, as Huneman writes “the proper object of life sciences should be a set of parts organizing itself as a whole, the development and the functioning of this specific kind of entity being the proper field of, respectively, embryology and physiology” (Huneman, 2010: 342).

Claude Bernard explicitly invokes Cuvier’s view and claims that biological systems are to be conceived as organized entities, whose parts are interdependent and mutually generative. In his words, “The physiologist and the physician must never forget that the living being comprises an organism and an individuality… If we decompose the living organism into its various parts, it is only for the sake of experimental analysis, not for them to be understood separately. Indeed, when we wish to
ascribe to a physiological quality its value and true significance, we must always
refer to this whole and draw our final conclusions only in relation to its effects in the
whole” (Bernard, 1865/1984, II, ii, x 1, 137, quoted and translated by Wolfe, 2010).
Bernard’s main focus is on the contribution of the organized parts – that must be
investigated through the experimental method to the conservation of the internal
milieu, in spite of the continuous variations taking place in the external milieu.

An important moment in the history of the scientific treatment of biological orga-
nization is represented by the so-called Theoretical Biology Club, that refers to a
group of researchers including Woodger, Needham, and Waddington (Etxeberria &
Umerez, 2006; Peterson, 2010). The Theoretical Biology Club promoted a scientific
organicist perspective for biology and underwent a rigorous conceptual and theo-
retical treatment of various dimensions of the very idea of organization, including
the analysis of internal relations (Woodger, 1929) and hierarchies (Needham, 1937).
Another particularly relevant contribution is due to Ludwig von Bertalanffy (1952),
who was one of the first authors that made explicit the fact that biological systems
as thermodynamically open systems. Initially used by Bertalanffy as an argument
against both vitalism and mechanism, the thermodynamic openness of biological
systems remains central aspect role in the subsequent elaborations on
organization.

Later on, the notion of organization played a central role in the organicist per-
spective that permeated embryology in the first half of the twentieth century. In
particular, Paul Weiss refers to organization as the “coordinating principle” (Weiss,
1963: 190) that characterizes biological systems beyond local components and pro-
cesses and that grounds their stability in the face of internal or external perturba-
tions (Rosslenbroich, 2011; Bich & Arnellos, 2013; Nicholson & Gawne, 2015).

In the second half of the twentieth century, the conceptualization and scientific
treatment of biological organization entered into a new phase, characterized by an
increasing coherence and theoretical refinement. A milestone in this tradition is the
account put forward by Jean Piaget (Piaget, 1967), whose core idea was to integrate
into a single coherent picture thermodynamic openness and organizational closure.
On the one hand, as emphasized by Bertalanffy, organisms are thermodynamically
open systems, traversed by a continuous flow of matter and energy. On the other
hand, they realize “closure,” i.e., the mutual dependence between a set of constitu-
ents which maintain each other through their interactions and which could not exist
in isolation.

In Piaget’s view, closure captures a fundamental aspect of the very idea of “orga-
nization,” through the association between division of labor and mutual dependence
that it implies. In other words, biological organisms are organized precisely because
they realize closure. The centrality of closure and its connection to organization, as
well as its distinction from (and, yet, complementarity to) thermodynamic openness
have become givens in most subsequent accounts of biological organization (Letelier
et al., 2011).

One of the best-known accounts of biological organization is the one centered on
the concept of autopoiesis (Varela et al., 1974; Varela, 1979) which, among other
aspects, places heavy emphasis on the generative dimension of closure: biological
systems determine themselves in the sense that they “make themselves” (auto-
poiein). Precisely because of their dissipative nature, the components of biological
organisms undergo degradation over time; the whole system preserves its coherence
and identity only insofar as it maintains and stabilizes not just some internal states
or processes but the autopoietic system itself as an organized unity. In spite of its
qualities, however, a central weakness of the concept of autopoiesis is that it does
not provide a sufficiently explicit characterization of closure (Montévil & Mossio,
2015). Biological systems are at the same time thermodynamically open and orga-
nizationally closed, but no details are given regarding how the two dimensions are
interrelated, what constituents are involved in closure, and at what level of descrip-
tion. In the absence of such specifications, it remains unclear in what precise sense
closure would constitute a causal regime that distinctively characterizes biological
organization.

A concerted attempt to answer this question was made by Robert Rosen. In *Life Itself* (Rosen, 1991), Rosen reinterprets the Aristotelian categories of causality and claims that the distinction between closure and openness should be grounded on a distinction between efficient cause and material cause. By relying on this distinction, Rosen's central thesis is that: "a material system is an organism [a living system] if, and only if, it is closed to efficient causation" (Rosen, 1991: 244). In turn, a natural system is closed to efficient causation if, and only if, all components having the status of efficient causes within the system are materially produced by the system itself. What matters here is that closure is located at the level of efficient causes: what constitutes the organization is the set of efficient causes subject to closure, and its maintenance (and stability) is the maintenance of the closed network of efficient causes.

Although Rosen's account represents a crucial step forward in the theoretical
understanding of organization, I think that it still remains too abstract, and therefore
hardly applicable as a guiding principle for biological theorizing, modeling, and
experimentation. Rosen defines closure as involving efficient causes, but, without
additional specifications, it might be difficult to identify efficient causes in the sys-
tem: what entities actually play the role of efficient causes in a biological system?
To deal with this issue, decisive insights have emerged from more recent literature
that elaborates more explicitly on the "thermodynamic grounding" of biological
systems (Bickhard, 2000; Christensen & Hooker, 2000; Moreno & Ruiz Mirazo,
1999) and the relations between closure and openness. In particular, Stuart Kauffman
(2000) argues that biological organization implies a circular relationship between
work and constraints, in the form of what he labels a "work-constraint (W-C) cycle."
When a (W-C) cycle is realized, constraints that apply to the system are produced
and maintained by the system itself. Hence, the system needs to use the work gener-
ated by the constraints in order to generate those very constraints, by establishing a
mutual relationship – a cycle – between constraints and work.

More recently, a characterization of biological organization as “closure of con-
straints,” which puts together many of the central ideas evoked above, has been
proposed (Montévil & Mossio, 2015). Most of the contributions to this book actu-
ally rely on this characterization to further develop its implications.
1.8 The Current Context, and The Place of the Book

Nowadays, what is generically called “organicism” is undergoing resurgence, as an
increasing number of philosophical, theoretical, and even formal accounts have
advocated it as an integrative and fecund framework for biology (Gilbert & Sarkar,
2000). Among these accounts, the aforementioned theory of biological autonomy –
originally elaborated by Varela (Varela, 1979) – is gaining momentum (Moreno &
Mossio, 2015).

Recently, several studies have relied on the pioneering work mentioned above, and tried to further elaborate on the central notion of organization. Some have investigated its philosophical (Mossio & Moreno, 2010) and theoretical (Letelier et al., 2011; Wolkenhauer & Hofmeyr, 2007) implications, while others have developed applications to various domains, such as the already mentioned metabolic networks (see also Cornish-Bowden et al., 2013), physiological regulation (Bich et al., 2020), the transition from unicellular to multicellular organisms (Arnellos et al., 2014), organogenesis (Montévil et al., 2016b), ecology (Nunes et al., 2014), agency (Barandiaran et al., 2009), cognition (Barandiaran & Moreno, 2006), and the origins of life (Ruiz-Mirazo & Moreno, 2004). These are just a few examples showing the existence of a scientific perspective that aims at establishing an organizational framework for biology – whether or not they stem from the theory of autonomy – and elaborates on some recurrent theoretical themes such as openness, closure, constraints, agency, and circularities, as well as their connections with philosophical issues as teleology, functionality, normativity, historicity, and individuation.

The main objective of the book is to assess the prospects and the fecundity of the concept of organization in biological research, both as a philosophical foundation and as a theoretical principle able to generate models and experimental protocols. The various chapters deal with a variety of issues with respect to which an organizational perspective can be adopted and discussed. Collectively, they show that the notion of organization can nourish the current anti-reductionist trend, by guiding the elaboration of models and the connection with experimental biology.

In the second chapter (Chap. 2), Georg Toepfer recounts the history of the con-
cept of organization, as used in relation to organic bodies. Toepfer underscores that
organization becomes a defining feature of life from the seventeenth century and
plays a central role in the establishment of biology as an autonomous scientific dis-
cipline. During the nineteenth and twentieth centuries, then, it has been supple-
mented by the concepts of evolution and regulation, which refer to the transformation
and stabilization of organized systems, respectively. In its more recent formulations
(notably in terms of constraints closure) – Toepfer argues – the specificity of orga-
nization is more explicitly tied to the specificity of forms that enable its realization.
As he writes: “The only life-forces that exist are life-forms.”

Charles Wolfe (Chap. 3) discusses the challenges that a naturalistic and non-
foundationalist – and thereby scientifically workable – organicist project should
take up. In his view, some versions of organicism suffer from at least three main
hesitations or “instabilities,” which relate to the interpretation of organismal
properties (epistemological vs. ontological, irreducible vs. empirical), as well as the opposition with mechanism. Wolfe argues that “one more effort” should be made to overcome these instabilities, without giving in to the symmetrical temptations of objectification and subjectification of organisms. The very concept of organization is likely to play a crucial role in making this effort successful.

In Chap. 4, Gertrudis Van de Vijver and Levi Haeck focus on one of the instabilities noted by Wolfe, that between the epistemological and ontological interpretations of organisms as organized systems. They put forward an original transcendental stance, inspired by Kant's treatment of biological organization, according to which both the subject and the object involved in our understanding of organisms should be treated as organized living systems. Thereby, the enquiry about the properties of living organization is simultaneously an enquiry about the subject and the object of biological knowledge. Insofar as our rational capacities are a manifestation of life, studying the latter “folds back onto” the former and reveals that our cognition shares fundamental organizational properties with biological phenomena, starting with their purposiveness.

Cliff Hooker (Chap. 5) shares with Van de Vijver and Haeck the idea that cognition and life display common properties. As life, cognition is organized in a narrow, theoretically precise sense, which implies in particular the realization of agency, purposiveness, and anticipation. In a word, both life and cognition realize autonomy. Yet, instead of focusing on the epistemic loop between cognition and life, Hooker emphasizes that autonomy comes in degrees: in particular, cognition (and notably human cognition) relies on much more sophisticated anticipatory and adaptive capacities, when compared to noncognitive biological autonomy. The take home message is “unification without reduction”: the concept of organization can be usefully put to work to provide a general understanding of cognition and life, while preserving their specific features.

Chapter 6 provides a counterpoint to the general message of the book. Olivier Sartenaer argues that organicism does not need organization to remain “chauvinist about organisms and autonomist about biology.” Organicism can vindicate the irreducibility of organisms – and justify its epistemological autonomy – by showing that they comply with the requirements of transformational emergence. Sartenaer’s argument is not that organization is an illegitimate concept, but that organicism could stand while discarding it. To the objection that, without organization, transformational emergence does not capture what distinguishes organisms from other emergents, Sartenaer replies that their specificity could be grounded in their being the outcome of specific transformational transitions during biological evolution. Yet, this solution begs the question whether biological evolution can be the evolution of anything else than organized systems.

Philippe Huneman (Chap. 7) offers a comparison between organizational and evolutionary approaches of organisms, that he labels “Kantian” and “liberal,” respectively. While the former aim at characterizing organisms by appealing to a distinctive set of organizational properties, the latter situate organisms in a larger, continuous spectrum of biological individuals understood as units of selection. In liberal approaches, organisms are not the only biological individuals and, in
addition, “being an organism comes by degrees.” The comparison raises the ques-
tion of the connections between the two conceptions of individuality, as well as
between the two underlying theoretical frameworks. Huneman addresses this ques-
tion and explores the prospects of a fruitful reconciliation between Kantian and
liberal approaches.

In Chap. 8, Johannes Jaeger provides an answer to the question raised by Huneman. He advocates the radical idea according to which biological evolution cannot be but the evolution of organized systems. Evolutionary theory should move toward a fourth perspective, which would complement and succeed existing structural, functional, and processual perspectives. In particular, the fourth perspective is an agential perspective, centered on the hypothesis that evolving organisms are organized purposive agents. As such, not only organisms are what evolutionary processes operate on, but they also modulate such processes. As Jaeger, following Walsh, puts it: “Some things in evolution happen because organisms make them happen.” The agential perspective relies on a naturalized understanding of purposiveness, provided by recent characterizations of organization as constraints closure. Importantly, Jaeger underscores that such an organizational purposiveness applies to individual organisms exclusively, and not to evolutionary processes as such. The existence of macroevolutionary trends is a different issue that should be explored separately.

Sharing Jaeger’s perspective on the role of organization in shaping evolution – one might argue – still leaves room to the assumption that, while looking at the origins of life, organization initially emerged from evolutionary processes. In Chap. 9, however, Kepa Ruiz-Mirazo and Alvaro Moreno argue that things are more complicated. They put forward an account of biogenesis that also ascribes an explanatory role to the concept of organization. Again, the emphasis is placed on the interplay between individual organization and evolutionary processes, which take place at a different spatial and temporal scale. Their main thesis is that, to result in the emergence of complex biological organisms (as we observe them), biogenetic trends require complex enough, organized self-maintaining systems as a starting point. Complexity begets complexity, in the sense of generating functional variety and more sophisticated forms of control. In particular, Ruiz-Mirazo and Moreno emphasize the evolutionary significance of forms of regulation and heredity relying on dynamical decoupling, whose emergence has drastically enhanced individual adaptivity and cross-generation stability.

Gaëlle Pontarotti (Chap. 10) specifically deals with the connection between
organization and heredity, a key ingredient of evolution. Pontarotti argues that the
general trend beyond genocentrism implies a shift from a heuristic of replication,
which sees evolution in terms of a competition among self-replicating objects, to a
heuristic of collaboration, which emphasizes the mutual dependence of objects
belonging to integrated wholes. The heuristic of collaboration can be applied to
elaborate an organizational account of heredity, which characterizes the latter as the
“trans-generational conservation of functional networks.” Pontarotti submits that
the organizational account allows expanding heredity beyond genes, while keeping
the concept conceptually bounded. The extension avoids then the dilution of
heredity into a too general concept of biological cross-generation stability. The chapter also discusses how the organizational account of heredity impacts some of the central tenets of evolutionary theory.

Chapter 11 shifts to the individual scale and explores the explanatory role that the concept of organization can have with regard to development. Leonardo Bich and Derek Skillings put forward an organizational view on development that, here again, makes determination reciprocal: not only development determines the establishment of biological organization but, reciprocally, organization enables developmental processes. As Bich and Skillings emphasize, the organizational view “favors a switch in perspective,” whereby each stage of development is understood as an organized system aiming at its own maintenance, rather than being an intermediate step of a process tending to a final state (typically identified with the achievement of reproductive capabilities). Bich and Skillings argue that the organizational view accounts not only for maintenance but also for change, which is genuinely developmental only if it is controlled by regulatory functions exerted by the organized system. Regulation also draws the boundaries of development, which starts when regulatory functions appear and ends when the organized system ceases to undergo regulated change.

Maël Montévil and Ana Soto (Chap. 12) further explore developmental processes by discussing their recent efforts to model morphogenesis, and more specifically mammary ductal morphogenesis. In their model, Montévil and Soto have applied two principles: cells default state and organization. According to the default state, cells move and proliferate when unconstrained, in the presence of sufficient nutrients and space, while the organization of the multicellular system that they constitute exerts the constraints that canalize or inhibit the default state. In particular, Montévil and Soto show that the formation of mammal ducts is determined by the interplay between the constraints exerted by proliferating cells on the extracellular matrix (notably on collagen fibers), which in turn constrains cells proliferation and motility. The chapter also addresses important issues raised by the modeling practice relying on organization. These include the choice of those organized parts and constraints expected to play a role in determining the target phenomenon, and their insertion into a description of the whole organismal organization.

The last chapter (Chap. 13) shifts again to a different scale and discusses some theoretical and ethical implications stemming from the application of the organizational framework to the ecological domain. Charbel El-Hani, Felipe Lima, and Nei Nunes-Neto argue that the concept of organization provides a relevant tool to individuate ecosystems and to ascribe functions to their parts (both to items of biodiversity and abiotic items). In particular, they provide a detailed reply to some objections recently raised against the organizational account of ecosystemic functions. El-Hani, Lima and Nunes-Neto’s contribution is particularly important because it shows that the concept of organization may be pertinently used to characterize biological systems in general, and not only organisms. Also, it opens the way to future research directions, which would explore the relations and interplay between nested levels of organization. Lastly, El-Hani, Lima, and Nunes-Neto argue that the concept of
organization, by naturalizing intrinsic purposiveness, provides a ground to ascribe
intrinsic value to ecosystems. In turn, this supports an original conception of sus-
tainability, which is alternative to the usual anthropocentric interpretation.

**Acknowledgments** I am indebted to many colleagues and friends who contributed to this project in different ways. First of all, I thank all authors for their contributions, as well as for taking the time to read and to comment on other chapters. The cross-reading effort certainly helped to improve the global coherence of the book. I also warmly thank the external reviewers for their detailed and constructive remarks: Argyris Arnellos, Mike Beaton, James DiFrisco, Antoine Dussault, James Griesemer, Francesca Merlin, Guglielmo Militello, Auguste Nahas, Dan Nicholson, Juli Peretó, Cyril Rauch, and Denis Walsh.

I am thankful to my research unit, the Institut d'Histoire et de Philosophie des Sciences et des Techniques (IHPST), and to the Université Paris 1 Panthéon – Sorbonne for funding and hosting, a few years ago, the workshop from which this book has been subsequently elaborated. A special mention goes to Andrea Gambarotto, who co-organized the workshop, and encouraged me to undertake the project of this book.

I would also like to thank the Labex “Who Am I? Exploring identity: from molecules to indi-
viduals” for its support, and to express my gratitude to the John Templeton Foundation for gener-
ously funding the open access publication of the book through the Agency, Directionality and
Function program (#62220), led by Alan Love. The opinions expressed in this book are those of
the authors and not those of the John Templeton Foundation.

References

Arnellos, A., Moreno, A., & Ruiz-Mirazo, K. (2014). Organizational requirements for multicel-
lular autonomy: Insights from a comparative case study. *Biology and Philosophy, 29*, 851–884.

Barandiaran, X., & Moreno, A. (2006). On what makes certain dynamical systems cognitive. *Adaptive Behavior, 14*, 171–185.

Barandiaran, X., Di Paolo, E., & Rohde, M. (2009). Defining agency. Individuality, normativity, asymmetry and spatio-temporality in action. *Adaptive Behavior, 17*(5), 367–386.

Bateson, P. (2005). The return of the whole organism. *Journal of Biosciences, 30*(1), 31–39.

Bernard, C. (1865/1984). *Introduction à l'étude de la médecine expérimentale*. Baillière.

Bertolaso, M. (2011). Hierarchies and causal relationships in interpretative models of the neoplas-
tic process. *History and Philosophy of the Life Sciences, 33*, 515–536.

Bich, L., & Arnellos, A. (2013). Autopoiesis, autonomy and organizational biology: Critical remarks on “Life after Ashby”. *Cybernetics and Human Knowing, 19*(4), 75–103.

Bich, L., Mossio, M., & Soto, A. (2020). Glycemia regulation: From feedback loops to organiza-
tional closure. *Frontiers in Physiology, 11*, 69.

Bickhard, M. H. (2000). Autonomy, function, and representation. *Communication and Cognition Artificial Intelligence, 17*(3–4), 111–131.

Callebaut, W., Müller, G. B., & Newman, S. A. (2007). The organismic systems approach: Evo-
devo and the streamlining of the naturalistic agenda. In R. Sansom & R. N. Brandon (Eds.),
*Integrating evolution and development: From theory to practice* (pp. 25–92). MIT Press.

Cannon, W. B. (1929). Organization for physiological homeostasis. *Physiological Reviews, 9*(3), 399–431.

Christensen, W. D., & Hooker, C. A. (2000). An interactivist-constructivist approach to intelli-
gence: Self-directed anticipative learning. *Philosophical Psychology, 13*, 5–45.

Cornish-Bowden, A., Piedrafita, G., Morán, F., Cárdenas, M.-L., & Montero, F. (2013). Simulating a model of metabolic closure. *Biological Theory, 8*(4), 383–390.

Crick, F. (1966). *Of molecules and man*. University of Washington Press.
Cuvier, G. (1805). *Leçons d'anatomie comparée*. Baudoin.

Cuvier, G. (1817). *Le règne animal distribué d'après son organization, pour servir de base à l'histoire naturelle des animaux et d'introduction à l'anatomie compare*. Déterville.

Dobzhansky, T. (1937). *Genetics and the origin of species*. Columbia University Press.

Etxeberria, A., & Umerez, J. (2006). *Organización y organismo en la Biología Teórica ¿Vuelta al organicismo? Ludus Vitalis, 26, 3–38*.

Fox Keller, E. (2000). *The century of the gene*. Harvard University Press.

Gambarotto, A. (2018). *Vital forces, teleology and organization: Philosophy of nature and the rise of biology in Germany*. Springer.

Gerhart, J. C., & Kirschner, M. W. (2007). The theory of facilitated variation. *Proceedings of the National Academy of Sciences of the United States of America, 104*, 8582–8589.

Gilbert, S. F., & Sarkar, S. (2000). Embracing complexity: Organicism for the 21st century. *Developmental Dynamics, 219*(1), 1–9.

Goethe, J. W. (1995). *Collected works, XII. Scientific studies*. Princeton University Press.

Goodwin, B. C. (1963). *Temporal Organization in Cells*. Academic.

Gould, S. J., & Lewontin, R. C. (1979). The spandrels of San Marco and the Panglossian paradigm: A critique of the adaptationist programme. *Proceedings of the Royal Society of London - Series B: Biological Sciences, 205*(1161), 581–598.

Green, S., & Wolkenhauer, O. (2013). Tracing organizing principles – Learning from the history of systems biology. *History and Philosophy of the Life Sciences, 35*, 553–576.

Hordijk, W., Kauffman, S. A., & Steel, M. (2011). Required levels of catalysis for emergence of autocatalytic sets in models of chemical reaction systems. *International Journal of Molecular Sciences, 12*(5), 3085–3101.

Huneman, P. (2006). Naturalizing purpose: From comparative anatomy to the “adventures of reason”. *Studies in History and Philosophy of Life Sciences, 37*(4), 621–656.

Huneman, P. (2010). Assessing the prospects for a return of organisms in evolutionary biology. *History and Philosophy of the Life Sciences, 32*, 341–372.

Huneman, P., & Walsh, D. M. (2017). *Challenging the modern synthesis: Adaptation development, and inheritance*. Oxford University Press.

Huneman, P., & Wolfe, C. T. (Eds.). (2010). The concept of organism: Historical, philosophical, scientific perspectives. *History and Philosophy of the Life Sciences, 32*(2–3), 145–424.

Huxley, J. (1942). *Evolution: The modern synthesis*. Allen & Unwin.

Jablonka, E., & Lamb, M. J. (1995). *Epigenetic inheritance and evolution: The Lamarckian dimension*. Oxford University Press.

Jablonka, E., & Lamb, M. J. (2005). *Evolution in four dimensions*. MIT Press.

Jacob, F. (1973). *The logic of life: A history of heredity*. Princeton University Press.

Kant, I. (1790/1987). *Critique of judgment*. Hackett Publishing.

Kauffman, S. (2000). *Investigations*. Oxford University Press.

Laland, K., Uller, T., Feldman, M., Sterelny, K., Muller, G. B., Moczek, A., & Odling-Smee, J. (2014). Does evolutionary theory need a rethink? Yes, urgently. *Nature, 514*, 161–164.

Laland, K. N., Uller, T., Feldman, M. W., Sterelny, K., Muller, G. B., Moczek, A., & Odling-Smee, J. (2015). The extended evolutionary synthesis: Its structure, assumptions, and predictions. *Proceedings of the Royal Society B, 282*(1813).

Laublichler, M. D. (Ed.). (2000). The organism in philosophical focus. *Philosophy of Science, 67*(3), S256–S321.

Letelier, J. C., Cárdenas, M., & Cornish-Bowden, A. (2011). From L'Homme machine to metabolic closure: Steps towards understanding life. *Journal of Theoretical Biology, 286*(1), 100–113.

Longo, G., Montévil, M., & Kauffman, S. (2012). No entailing laws, but enablement in the evolution of the biosphere. In *Proceedings of the 14th international conference on genetic and evolutionary computation conference*, pp. 1379–1392.

Mayr, E. (1961). Cause and effect in biology. *Science, 134*(3489), 1501–1506.

Mesarović, M., & Sreenath, S. N. (2006). Beyond the flat earth perspective in systems biology. *Biological Theory, 1*, 33–34.
Minelli, A., & Fusco, G. (Eds.). (2008). *Evolving pathways. Key themes in evolutionary developmental biology*. Cambridge University Press.

Monod, J., & Jacob, F. (1961). Genetic regulatory mechanisms in the synthesis of proteins. *Journal of Molecular Biology*, **3**(3), 318–356.

Montévil, M., & Mossio, M. (2015). Biological organization as closure of constraints. *Journal of Theoretical Biology*, **372**, 179–191.

Montévil, M., Mossio, M., Pocheville, A., & Longo, G. (2016a). Theoretical principles for biology: Variation. *Progress in Biophysics and Molecular Biology*, **122**, 36–50.

Montévil, M., Speroni, L., Sonnenschein, C., & Soto, A. M. (2016b). Modelling mammary organogenesis from biological first principles: Cells and their physical constraints. *Progress in Biophysics and Molecular Biology*, **122**(1), 58–69.

Moreno, M., & Mossio, M. (2015). *Biological autonomy. A philosophical and theoretical enquiry*. Springer.

Moreno, A., & Ruiz Mirazo, K. (1999). Metabolism and the problem of its universalization. *Biosystems*, **49**(1), 45–61.

Mossio, M., & Bich, L. (2017). What makes biological organization teleological? *Synthese*, **194**, 1089–1114.

Mossio, M., & Moreno, A. (2010). Organizational closure in biological organisms. *History and Philosophy of Life Sciences*, **32**, 269–288.

Mossio, M., Saborido, C., & Moreno, A. (2009). An organizational account of biological functions. *British Journal for the Philosophy of Science*, **60**, 813–841.

Mossio, M., Montévil, M., & Longo, G. (2016). Theoretical principles for biology: Organization. *Progress in Biophysics and Molecular Biology*, **122**, 24–35.

Müller, J. (1837–1840). *Handbuch der Physiologie des Menschenfür Vorlesungen* (Vol. 2 Vols). Verlag von J. Hölscher.

Needham, J. (1937). *Integrative levels: A revaluation of the idea of progress*. Clarendon Press.

Nicholson, D. J. (2014). The return of the organism as a fundamental explanatory concept in biology. *Philosophy Compass*, **9**, 347–359.

Nicholson, D. J., & Gawne, R. (2015). Neither logical empiricism nor vitalism, but organicism: What the philosophy of biology was. *History and Philosophy of the Life Sciences*, **37**, 345–381.

Noble, D. (1962). A modification of the Hodgkin-Huxley equations applicable to Purkinje fibre action and pace-maker potentials. *Journal of Physiology*, **160**, 317–352.

Noble, D. (2017). Systems biology beyond the genome. In S. Green (Ed.), *Philosophy of systems biology*. Springer.

Nunes, N., Moreno, A., & El Hani, C. (2014). Function in ecology: An organizational approach. *Biology and Philosophy*, **29**(1), 123–141.

O'Malley, M. A., & Dupré, J. (2005). Fundamental issues in systems biology. *BioEssays*, **27**, 1270–1276.

Odling-Smee, J., Laland, K., & Feldman, M. (2003). *Niche construction: The neglected process in evolution*. Princeton University Press.

Oyama, S. (1985). *The ontogeny of information*. Cambridge University Press.

Pereto, J. (2005). Controversies on the origin of life. *International Microbiology*, **8**, 23–31.

Peterson, E. (2010). *Finding mind, form, organism, and person in a reductionist age*. PhD Dissertation, 2 vols., Program in History and Philosophy of Science, University of Notre Dame.

Piaget, J. (1967). *Biologie et connaissance. Éditions de la Pléiade*.

Piedrafita, G., Montero, F., Morán, F., Cárdenas, M.-L., & Cornish-Bowden, A. (2010). A simple self-maintaining metabolic system: Robustness, autocatalysis, Bistability. *PLOS Computational Biology*, **6**(8): e1000872.

Pigliucci, M., & Müller, G. B. (Eds.). (2010). *Evolution: The extended synthesis*. MIT Press.

Rasmussen, S., Bedau, M. A., Liaohai, C., Deamer, D., Krakauer, D. C., Packhard, N. H., & Stadler, P. F. (Eds.). (2008). *Protocells: Bridging nonliving and living matter*. MIT Press.

Reiss, J. O. (2009). *Not by design: Retiring Darwin's watchmaker*. University of California Press.

Rosen, R. (1991). *Life itself. A comprehensive enquiry into the nature, origin and fabrication of lif*. Columbia University Press.
Rosenberg, A. (2007). Reductionism (and anti-reductionism) in biology. In D. L. Hull & M. Ruse (Eds.), *The Cambridge companion to the philosophy of biology* (pp. 120–138). Cambridge University Press.

Rosslenbroich, B. (2011). Outline of a concept for organismic systems biology. *Seminars in Cancer Biology, 21*, 156–164.

Ruiz-Mirazo, K., & Moreno, A. (2004). Basic autonomy as a fundamental step in the synthesis of life. *Artificial Life, 10*(3), 235–259.

Ruiz-Mirazo, K., Briones, C., & Escoura, A. (2017). Chemical roots of biologicalevolution: The origins of life as a process of development of autonomous functional systems. *Open Biology*, 7, 170050.

Saborido, C., Mossio, M., & Moreno, A. (2011). Biological organization and cross-generation functions. *The British Journal for the Philosophy of Science, 62*, 583–606.

Saetzler, K., Sonnenschein, C., & Soto, A. M. (2011). Systems biology beyond networks: Generating order from disorder through self-organization. *Seminars in Cancer Biology, 21*, 165–174.

Varela, F. J. (1979). *Principles of biological autonomy*. North Holland.

Varela, F. J., Maturana, H., & Uribe, R. (1974). Autopoiesis: The organization of living systems, its characterization and a model. *Biosystems, 5*, 187–196.

Vasas, V., Szathmary, E., & Santos, M. (2010). Lack of evolvability in self-sustaining autocatalytic networks constraints metabolism-first scenarios for the origin of life. *Proceedings of the National Academy of Sciences of the United States of America, 107*, 1470–1475.

von Baer, K. E. (1828). *Über Entwicklungsgeschichte der Thiere: Beobachtung und Reflexion*. Bornträger.

von Bertalanffy, L. (1933). *Modern theories of development*. Oxford University Press.

von Bertalanffy, L. (1952). *Problems of life: An evaluation of modern biological thought*. Watts & Co.

Wagner, G. P., & Laublicher, M. D. (2000). Character identification: The role of the organism. *Theory in Biosciences, 119*, 20–40.

Walsh, D. M. (2015). *Organisms, agency, and evolution*. Cambridge University Press.

Weiss, P. (1963). The cell as unit. *ICSU Review, 5*, 185–193.

West-Eberhard, M. J. (2003). *Developmental plasticity and evolution*. Oxford University Press.

Wiener, N. (1948). *Cybernetics: Or control and communication in the animal and the machine*. The MIT Press.

Wolfe, C. (2010). Do organisms have an ontological status? *History and Philosophy of the Life Sciences, 32*(2–3), 195–232.

Wolkenhauer, O., & Hofmeyr, J.-H. (2007). An abstract cell model that describes the self-organization of cell function in living systems. *Journal of Theoretical Biology, 246*, 461–476.

Woodger, J. H. (1929). *Biological principles. A critical study*. Routledge and Kegan.

**Open Access** This chapter is licensed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license and indicate if changes were made.

The images or other third party material in this chapter are included in the chapter's Creative Commons license, unless indicated otherwise in a credit line to the material. If material is not included in the chapter's Creative Commons license and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder.

<figure><img src="image_6.png" /></figure>
<figure><img src="image_7.png" /></figure>

Check for
updates

Chapter 2

**“Organization”**: Its Conceptual History
and Its Relationship to Other Fundamental
Biological Concepts

Georg Toepfer

**Abstract** The conceptual history of the term “organization” begins in Medieval times with the reception and transformation of Aristotle’s philosophy of life. It designates the corporeal structure and conditions of identity of natural “organic bodies,” a term that had been used to refer to living beings since antiquity. The term played an important role in specifying the ontological status of living beings. At the same time, it offered a basis for their mechanistic understanding. Starting with mechanistic models of life in the second half of the seventeenth century, “organization” and “life” were increasingly used interchangeably. This conjunction of meaning transformed “living beings” into “organisms.” Within physiological accounts of the eighteenth century, the living organization was compared to a causal cycle of interdependency. Philosophically, this conjunction was adapted at the end of the century in Kant’s philosophy of “organized beings of nature” in which he located the idea of causal cyclicity within a teleological framework and specified an “organized being” in causal terms as a system of interacting and interdependent parts characterized by functional closure. Thus, “organization” refers to the constitution of living beings as a particular kind of causal system. In the nineteenth century, the term achieves the status of a signal word for the life sciences and starts being applied in a wide variety of contexts, from comparative anatomy to physiology and ecology. It was supplemented by two other fundamental notions, namely, “regulation” and “evolution,” the first referring to the stabilization and the second to the long-term transformation of natural organizations. The twentieth century saw a further intensification of the complementarity of the perspectives associated with these three terms. Finally, in recent years, a substantial improvement in understanding the causal structure of “organization” was achieved by analyzing it in terms of the “closure of constraints.”

<footnote>G. Toepfer (✉)
Berlin, Germany
e-mail: toepfer@zfl-berlin.org</footnote>
## 2.1 Introduction

For a very long time, “organization” has been a central concept in biology. Since antiquity, the material basis of a living being has been called “organic body” (“corpus organicum”). For Aristotle, this meant that the body is an instrument (“organon”) of the soul (Bos, 2003). However, since late antiquity, especially since the writings of Galen, an “organic body” was understood as an integrated system in which the parts mutually depend on one another. Thus, they were seen as instruments not only for the soul but also for their own interdependent activity. Galen compared the working order of the organic body to a “symphony” and explained it as “sympathy” or “synergy” in the sense of a “functional organization” (Siegel, 1973, p. 129; Toepfer, 2011a, vol. 2, p. 779). However, the term “organization” was not applied to this functional organization prior to the Middle Ages. Surprisingly, there was apparently no original semantic connection between the Greek expressions for “organization” and “organ”: whereas the former was used in the sense of “forming,” the latter had a functional meaning from the beginning. Both expressions have therefore been described as “semantically autonomous” (Wolf, 1971, p. 31). Only in the course of their later development were the two notions semantically unified. Starting with mechanistic models of vital processes in the second half of the seventeenth century, “organization” and “life” were increasingly used interchangeably. Since the end of the eighteenth century, “organization” has thus predominantly been understood as a characteristic of living beings, becoming a signal word for the animate world and its scientific analysis. The assumption that it was the organization of their body that constituted the defining characteristic of living beings led to the transformation of “living beings” into “organisms.” However, on the level of individual organisms, this did not happen before the end of the eighteenth century; before that, “organism” was used (in parallel to “mechanism”) in the sense of “organization,” referring to the abstract structure of organisms, not specific individuals (Cheung, 2006). In the nineteenth century, it became common practice to equate life with organization: in the words of the early Neo-Kantian philosopher Kuno Fischer, “Life is organization, self-organization.” (Fischer, 1865, p. 534) A few decades later, at the beginning of the twentieth century, “biology,” the label that had been given to the science of life, was defined as the “study of the organization of the living” with “organization” being the name for “the association of different elements according to a uniform plan for a common effect” (Uexküll, 1903, p. 269). However, for the longer part of its history, “organization” has functioned as a dummy concept or placeholder for a theory of the living still under negotiation. It was embedded in an explanatory approach in which living beings were seen as functional systems composed of interacting parts. It was only during the last decades that a full-fledged theory of organization was proposed, enabling the term to fulfill the theoretical role it was meant to fulfill since the 1800s.
## 2.2 The Conceptual History of “Organization”

The term “organization” first appeared in medieval Latin. The word is related to the Greek expression “δργάνωσις” meaning “formation, arrangement.” The Greek term was used, for example, by Sextus Empiricus and Porphyrios in late antiquity, in the second and third century AD, respectively. Whereas Sextus Empiricus used the Greek expression in his treatise *Against the Logicians* in the more general sense of “arrangement” or “conjunction” (Adversus dogmaticos, 7, 126; Engl. transl. Bett, 2005, p. 114), Porphyrios applied it to the more specific context of bodily structures that living beings of distinct categories have in common (in order to argue against the practice of eating meat): “Almost everyone agrees that animals are like us in perception and in organisation generally with regard both to sense-organs and to the flesh” (De abstinentia 3, 7; Engl. transl. Clark, 2000, p. 84).

Later, in the High Middle Ages, the Latin term “organization” appears in several texts by Thomas Aquinas, especially in reference to Aristotle’s second book of *On the Soul*. Here, Thomas closely associated the term with the formation of organic bodies (“formatio et organizatio corporis”; In III Sententiarum distinctio (1254–56): dist. 3, qu. 2, art. 1; dist. 4, qu.2, art. 1) and claimed that, for Aristotle, “organization” was the basic principle of living bodies (“de ratione corporis vivi est organization”; In IV Sententiarum distinctio (1254–56) dist. 10, qu. 1, art. 2, quaestiuncula 3, sed contra 2). Furthermore, he stated that the term referred to a multitude of parts and was relevant to the form of the body. In these and other passages, Thomas attributed the concept of an “organic body” to Aristotle, in the sense of a body consisting of a diversity of organs; only those bodies which feature this inner diversity are “organic bodies”; this diversity was said to be “necessary” for living bodies: “dicitur corpus organicum, quod habet diversitatem organorum” (Commentarius in libros de anima II et III: 2, 1, 20 (No 230)). Following these lines of thought, future authors even referred to Aristotle as the “father of animal organization” (Schiller, 1978, p. 84).

However, there is some debate as to whether Aristotle actually saw living beings as organisms (Bolton, 1978; Bos, 2003). In *On the Soul*, the book referred to by Thomas Aquinas, Aristotle claimed that the soul is the form of a natural body that potentially bears life. However, according to Aristotle, the soul as the principle of life was not directly related to the disposition of the organs in the body. For Aristotle, the body was an organ or an instrument of the soul but not necessarily an organized being consisting of a diversity of interacting organs. He did not elucidate the relationship of the soul to the diversity of its organs. “Organic,” a term Aristotle apparently introduced into the Greek language, was used by him in reference to something “instrumental.” Here, the term does not refer to a diversity of organs and does not even imply an “endowment with organs.” Consequently, in the modern and contemporary sense, the concept of organization was not an inherent part of Aristotle’s philosophy of living beings.

Aristotle had no specific term for the arrangement or disposition of the organs that provided essence and unity to living beings. In Aristotle’s terminology, this
function is fulfilled by the soul. However, there are good reasons to assume that the Aristotelean “soul” corresponds well with the medieval and modern idea of organization since both refer to the essence and unity of a living system. There are four particular parallels between the concepts of “soul” and “organization” (Quarantotto, 2010): (1) like the soul, the organization is not itself a body, but a property belonging to the body; (2) soul (or organization) and body do not exist independently of what possesses them, they are not self-sufficient autonomous entities; (3) both are considered the principle of unity and identity of the body that they organize/endow with soul; and (4) soul and organization are both explanatory principles for fundamental organic activities such as movement—the reason why animals have the capacity to move is found in their organization or endowment with a soul.

Thus, prior to the scientific revolution in the seventeenth century, “soul” was a concept perfectly equipped to fulfill the explanatory role that was later taken on by “organization.” In fact, the latter term only came into frequent use during the seventeenth century. Until then, it was employed nearly exclusively in the context of scholastic discussions on the changing arrangement of parts in embryonic development.

That situation changed in the early seventeenth century, when, in a commentary on Aristotle’s *On the Soul* (1600), “organized” was equated with the state of “potentially having life” (Collegium Conimbricense, 1600, p. 55; for the context, see Des Chene, 2000). Henceforth, the term “organization” entered the academic language, especially as a result of debates on the value of mechanistic models for living beings.

At the beginning of this debate, “organization” was not yet in the position to become the fundamental principle of life. For the Cambridge Platonist Henry More, for example, “organization” was not equivalent to the living state of a being. He postulated a “Plastical Power” that “organized” “duly-prepared Matter,” as he called it, “into life” (More, 1659, p. 46). Thus, the “mere organization of the Body” (ibid., p. 107) was not enough to constitute life; this was a merely *mechanical* organization—or, in More’s terms, “matter mechanically organized” (ibid., p. 109). Thus, More still differentiated between life as “being ensouled” and life as “organization.”

Three years later, in 1662, Joachim Jungius, a mathematician and philosopher of science from Hamburg, announced that “true organization alone” was at least sufficient for plants to perform their life functions of nutrition, growth, and reproduction. Jungius followed Descartes as he denied plants a soul, arguing that their life functions could be explained by the mere disposition and arrangement of their parts: “vero organisatio sola sufficiat” (Jungius, 1662, part. 2, sect. 3). To my knowledge, this is the first instance in which “organization” and “life” were used interchangeably. The exact wording was later resumed by La Mettrie in his description of the relationship between the mental and the material aspect of the brain: “The organization, is it really sufficient for everything? Once again, yes” (La Mettrie, 1747, p. 180).

In the period of dominant mechanistic thinking from the mid-1660s onward, several authors accepted the equation of life and organization. Robert Hooke, for example, used the term “organization” in English. Presumably inspired by his
microscopic observations of plant tissues, he claimed that there was an organization common to all vegetables ("the same Schematism or Organization that is common to all Vegetables"; Hooke, 1665, p. 116). In a similar vein, Francis Glisson argued in 1672 that the difference between plants and animals and other bodies could be deduced from their "organization." Thus, life had no cause other than the organization of bodies (Glisson, 1672, p. 226; for the context, see Hartbecke, 2006, p. 165). Additionally, in an important and well-known ontological argument, John Locke reflected on the conditions of identity of living beings whose parts are constantly being exchanged whereas the entirety of the system remains the same. Locke used the term "organization" to refer to this bodily property that always persists even as its parts are exchanged (Locke, 1689, p. 331).

In the 150 years between 1650 and 1800, which could be viewed as the formative period of biology, the ancient principle of life, the "soul," was gradually replaced by "organization." Organization became the central explanatory concept for biology. This was the conceptual revolution at the beginning of biology, which at the same time maintained the ontological specificity of life phenomena, and their mechanistic explainability starting from uniform principles transformed the study of life into an explanatory endeavor that maintained rests on a unifying principle that provided: life was equated with being organized (see also Jacob, 1973, Chap. 2). An important element of this revolution was a reversal of the relationship between the concepts of "life" and "organization": in the seventeenth century, "life" was the more fundamental notion and the living state was thought to somehow determine the organization of the body. However, during the eighteenth century, it was established that it actually works the other way around, a notion that persists until today: "organization" now forms the basic concept from which the determination and consequently the analysis starts; "life" becomes a phenomenon derived from "organization" (see Schiller, 1978, p. 24).

In the first half of the eighteenth century, however, this equation was not accepted by all authors. In order to integrate the living world into the nonliving (and thus advocate for the possibility of a spontaneous generation of living beings), some considered all parts of nature to be organized. This was the position held by Leibniz, for example. He reasoned that since everything is arranged by God, every piece of matter is organized ("la matière arrangée par une sagesse divine doit estre essentiellement organisée partout"; Leibniz, 1705, p. 342). In the 1720s, this view is supported by microscopic investigations and descriptions of the regular crystalline structure in minerals (see, e.g., Bourguet, 1729, p. 58: "tout est organisé dans la matière"). Thus, until the middle of the eighteenth century, there were influential authors for whom the concept of organization served to unify rather than to separate realms of nature.

For the life sciences, "organization" increasingly served as an important explanatory concept. This is especially true for mechanical approaches toward the generation and transformation of living beings. In fact, in the mid-eighteenth century, it was preformationism, i.e., the idea that the forms of living beings are already existent in their germs, that mainly contributed to the diffusion of the concept (Schiller, 1978, p. 40). In reference to the preexistent structures in the germ, its
"organization," it was possible to explain the emergence of complex adult forms as "development."

In the eighteenth century, the idea of organization was mainly associated with mechanical understandings of life and preformationist accounts of individual development. In this context, the term mainly referred to the specific body plan of organisms and could therefore be used not only in developmental studies but also in natural history for the classification of organisms into taxonomic groups. Linnaeus, the master of this approach, defended the view that "organization" was a specific concept within the life sciences, presumably because matter in living beings, in contrast to nonliving ones, was specifically arranged into recurrent, taxonomically significant structures. In the tenth edition of his *Systema Naturae*, the association of "organized" and "living" was made explicit and formalized typographically by characterizing plants and animals as "organized and living" (*organisata* & *viva*), whereas stones were seen as merely "composite" (*congesta*) (Linnaeus 1758, p. 6). In the mid-eighteenth century, several authors stressed the explanatory value of "organization" in different fields within the life sciences: in 1750, John Turberville Needham argued that vitality, sensation, and thinking appear to be an immediate consequence of "organization" (Needham, 1750, p. 375). In 1772, Voltaire famously defined life as organization: "La vie est organisation avec capacité de sentir" (Voltaire, 1772, p. 55). Some years later, Diderot claimed that the soul is nothing but organization and life: "L'organisation et la vie, voilà l'âme" (Diderot, 1778, p. 358). At the end of the century, Christoph Girtanner also directly connected the living state with being organized: "Les mots organisé & vivant sont, selon moi, des synonimes" (Girtanner, 1790, p. 150). Even in Kant, in his *Opus postumum*, one can find this equation of "being alive" and "having an organization" (Kant, OP., AA XXI, p. 66). In his major work on the epistemology of biology, the *Critique of Judgement*, Kant (AA V) called living beings "organized beings in nature," although he avoided the term "living" because it had a terminological use in his ethical writings.<fnref n="1" />

Since the beginning of the nineteenth century, “organization” has been regarded as one of life’s most essential aspects. It proved to be valuable for the self-understanding of biology as an independent natural science. This was particularly evident in situations where life was either reduced to the merely mechanical or explained with additional supernatural “life forces.” In the mid-nineteenth century, Claude Bernard essentially relied on the concept of organization when he rejected vitalistic approaches. According to Bernard, all manifestations of life are not to be derived from a mysterious life force, but from “the phenomena of organization” (Bernard, 1867, p. 138). At the beginning of the twentieth century, in light of the growing struggle between mechanism and vitalism, the study of life was simultaneously confronted with the postulation of mysterious vital principles and reductionist views that denied any peculiarity of vital phenomena and their scientific explanations. In this situation, “organization” was propagated as a concept that offered a

<footnote n="1">¹However, things are complicated in Kant as he also has the concept of nonliving natural purposes which are the plants. Hence, for Kant, not all organized beings are living. I thank Gertrudis Van de Vijver for pointing to this; see also Piché, 2001.</footnote>
third way between these two metaphysical positions and, thus, was seen as a way
out of the fundamental dispute in theoretical biology. On the one hand, the assump-
tion of a central guiding vital force was considered unnecessary, because, if seen as
functional organizations, living beings and the orderly processes within them could
be described as the outcome of a decentralized structure of interacting parts. On the
other hand, "organization" was understood (in the Kantian sense) as an additional
principle that is not part of a purely mechanical approach, because it added the
aspect of integrating isolated causal relations into a coherent functional whole. In
1900, Oscar Hertwig argued that the explanation of life should neither introduce
mysterious forces nor follow the "mechanistic dogma," according to which "life
with all its complicated phenomena is nothing than a physico-chemical problem"
(Hertwig, 1900, p. 24). Instead, Hertwig argued for recognizing "that life is based
on a peculiar organization of the substance" (ibid, p. 4). Via the concept of organiza-
tion, biology could thus take a third path and navigate between the approaches of
vitalism and mechanism, thus securing biology's status as a natural science and its
methodological autonomy from physics (see also Wolfe's contribution to this vol-
ume). In the twentieth century, biological research programs that aimed to find life
on other planets and create life in the lab found the organizational approach to be
more stable than any purely material or molecular characterization of life: "The
peculiarity of life is not due to some chemical mystery but to organization"
(Bertalanffy, 1928, p. 68-9).

Hence, at least for 250 years, “organization” has been one of biology’s basic if
not one of its most fundamental concepts that which explains what life is. This
poses a question: What is “organization”? What does the term actually mean?

## 2.3 The Meaning of “Organization”

A fairly good, but still very open, definition can be found in the *Encylopédie*:
“Organization” is defined as “the arrangement of parts that constitute a living body”
(Anonymous, 1765, p. 629). Thirty years later, Kant, in a letter to Sömmering, con-
tributed an important amendment: he expanded this definition by including purpo-
siveness. For him, “organization” was “the purposeful and in its form persistent
arrangement of parts” (Kant, 1795, p. 33). In his works on natural philosophy, Kant
had a very specific understanding of purposefulness. As is well known, Kant stated
that in a thing as a natural purpose, the parts are “reciprocally the cause and effect
of their form” (Kant, 1790, p. 373). This means in a thing as a natural purpose the
parts’ *very existence* depends on the system’s other parts. Kant stresses this onto-
logical dependency in another passage: “For a body [...] which is to be judged as a
natural end in itself and in accordance with its internal possibility, it is required that
its parts reciprocally produce each other, as far as both their form and their combina-
tion is concerned, and thus produce a whole out of their own causality” (ibid.).

Kant did not indicate the origin of his idea of reciprocity as a condition for a
thing to be a natural purpose. However, similarities in the wording and his personal
contacts suggest that the idea was inspired by the Leiden physiologist Herman Boerhaave (see Toepfer, 2011b). In 1727, Boerhaave provided a definition for the concept of an “organic body” in which the decisive moment is the interdependence of the parts (*harum partium actiones ab invicem dependent*; Boerhaave, 1727, p. 3). To be sure, the emphasis on reciprocity as a hallmark of organic systems has its roots in Antiquity (see Toepfer, 2011a, vol. 3, pp. 738–763). However, only with the physiological theories since the end of the seventeenth century did it acquire a fundamental role in the identification and definition of living beings. This process took place in parallel with the introduction of the concept of “organism.” Georg Ernst Stahl, who proposed this notion in 1684, already described the relationship of the parts in an organism as an “adaptation of forms” (*aptatio configurationis*) and a dynamic interaction of a single part with the others (*cum aliis partibus cohaerens, conspirans, atque communicans*) (Stahl, 1707, p. 17). The parts in an organism would act “reciprocally and together” (*mutua & socia*) and thus be interrelated (ibid.). According to Stahl, the whole complex of the diverse organs in an organism forms a functional unity since the ultimate purpose of all movements is to preserve the body. The concept of “organism” thus establishes a causal model for a functionally closed and self-referential system of heterogeneous components. During the first half of the eighteenth century, Boerhaave and other mechanistically minded physiologists invoked the image of a “circle” (*circulo quasi*) for the causal pattern of organic systems, consequently firmly anchoring any discussions of causal reciprocity in physiological language (*mutuas causæ vices & effectuum gerant*) (Boerhaave, 1708, p. 11).

This physiological understanding of the interactions within an organic body formed the background for Kant's understanding of “organization” in terms of causal reciprocity. Kant's philosophical contribution was the integration of teleology into this causal understanding of natural systems of interdependent parts as well as the clarification of the peculiar metaphysical and ontological status of organisms or, in his terms, “things as natural purposes,” with respect to the explanatory level of causal mechanisms. In doing so, in combining teleology and cyclicity, Kant gave a justification of teleology within biology as the science of cyclical organized systems: the teleological way of thinking by focusing on outcomes of processes is justified in biology because biology is the study of systems consisting of interdependent parts in which the final state of one process is important for the existence of the other parts of the system and ultimately for its own maintenance (as a type of process or part).

Kant's philosophy of the organic was widely received in the years around 1800. Disciples of Kant gave explicit definitions of “organization” with Kant's philosophy in mind, for example: “Organization is the disposition of a body in which every part is at the same time means and ends to all the others” (Schmid, 1799, p. 274). Not only philosophers but also practicing biologists accepted this foundational role of reciprocity and teleology for the specification of their objects of study. One example by a famous author: “A living body is a natural organized body composed of different kinds of parts which act and react upon each other” (Lamarck, 1797, pp. 249–50). Lamarck repeatedly formulated concise sentences which express the close
connection between the state of being alive and organization or order. For him, life constitutes a “physical phenomenon resulting from the order of things and from the state of the parts,” their “organization” itself being a “physical phenomenon” (Lamarck, 1815, p. 60; 122; see Schiller, 1978, p. 70). “Life” was explained as an “ensemble of functions” with the functions being nothing but “acts of the organization and its pars” (ibid., p. 59). As these short quotations make clear, Lamarck had a dynamical understanding of “organization”; in his view, it is linked to movements caused by the arrangement of the parts within a body.

In the first half of the nineteenth century, however, the static interpretation of “organization” proved to be at least as important as the dynamic view.<fnref n="2" /> In comparative anatomy, one of the dominant research areas at that time, “organization” was understood as the “disposition” of the parts in an organic body; it referred to the configuration of the parts, the “body plan.” Anatomy with its focus on the spatial arrangement of parts within a body has even been called “the science of organization” (Schiller, 1978, p. 88). In this context, the analysis of the “organization” of body plans formed the foundation for the classification of animals into larger groups. For Georges Cuvier, one of its main representatives, comparative anatomy is the study of “the laws of organization of animals and of the modifications this organization shows in different species” (Cuvier, 1817, vol. 1, p. v). In Cuvier’s taxonomic system, the arrangement of the nervous system was of particular importance; it provided the basis for the classification of all animals into four main “branches” (see Figlio, 1976; Guillo, 2003). Here, “organization” was an important notion because it stressed the interdependence of the parts. In comparative anatomy, this interdependence was not primarily a causal notion but referred to the observation that traits of the body plan covary and do not exist independently from one another. Besides that, “organization” was also used as a measure for the complexity or “degree of perfection” of body plans. Even Charles Darwin, who was generally skeptical of the idea of progress in the history of life on earth, held the view that natural selection would result in an “improvement” that inevitably led to “the gradual advancement of the organization” (Darwin, 1860, p. 117).

For the philosophy of biology and its reflection on the ontological peculiarity of living beings, the important aspect was not this morphological concept of organization but the physiological meaning of the term. It was in the years around 1800 that the essential aspects of the concept were established—those that have persisted ever since. “Organization” now referred to the disposition of the parts in a certain type of causal system; the pattern of causal interactions has the form of a cycle because the parts mutually depend on one another’s influence, resulting in a functional “closure.”<fnref n="3" />

<footnote n="2">²Therefore, most of the historical accounts of the concept of “organization” in the history of biology focus on this aspect (see Figlio, 1976, Schiller 1978, and Guillo, 2003).</footnote>

<footnote n="3">³This currently prominent term that is most often derived from Piaget (1967, p. 182) also has its roots in early nineteenth-century reflections on the ontological status of organisms. Georges Cuvier, for example, put it this way: “Tout être organisé forme un ensemble, un système unique et clos, dont toutes les parties se correspondent mutuellement, et concourent à la même action définitive par une réaction réciproque” (Cuvier 1812, vol. 1, p. 58).</footnote>
Later definitions elaborated on these points, especially by highlighting the self-referential character of organizations. In 1928, Helmuth Plessner explained: “Organization is the mode of existence of the living body, which must differentiate itself and through which it generates the inner teleology according to which it is formed and functions” (Plessner, 1928, p. 170). Thus, an organization consists of differentiated functional parts, which, through their activity, generate and permanently regenerate the entire system. The same point was expressed in the theory of autopoiesis since the early 1970s, in which the “living organization” was characterized as a perpetual self-regeneration (Maturana et al., 1974). In this tradition, organization was defined as “the complex of interaction and properties of structure that make the perpetuation of structure possible” (Kolasa & Pickett, 1989, p. 8837).

Many authors have stressed the close connection between the concept of organization and teleology (except Maturana and his co-workers). Following Kant, one could say that “organization” and “function” or “purpose” go hand in hand: wherever there is organization in nature there is function and vice versa. As John von Neumann once said in conversation with Colin Pittendrigh, “Organization has purpose; order does not” (Pittendrigh, 1993, p. 20). Since functional reasoning and exploring purposes are essential to the domain of biology, it makes sense that “organization” has become a fundamental concept for that science—in contrast to physics, as “the physical sciences don’t deal in function” (Wicken, 1987, p. 40). Consequently, “organization” and “function” are frequently regarded as crucial to any attempt to justify the autonomy of biology as a science. Since Kant, this position has been defended by many authors, and teleology was even defined as the “philosophy of biology” because “the organism requires teleological consideration” (Kühnemann, 1924, p. 494).

It is a striking feature of biology that functions have been ascribed to living systems long before the causal pattern of their working order had been understood. Surprisingly, the basic inventory and supposition of functions have changed very little throughout the long history of biology. In fact, Aristotle had already named them: *nutrition, growth, movement, sensation, and reproduction*. However, it took more than 2000 years before biology began to understand the way in which they are realized in living beings. Functional knowledge is therefore a one-way kind of knowledge; it reduces the complexity of a system without necessarily having a complete understanding of it. Or, in other words: “organization [and hence function] emerges as a problem when there is too much knowledge in one direction and too little in another” (Beckner, 1959, p. 10).

The integrative power of the concept of “organization” in biology can also be demonstrated by explicating the fundamental properties of living beings as based on this concept. Metabolism, reproduction, development, metamorphosis, and evolution are fundamental aspects of life that can be described and analyzed, respectively, as maintenance, transmission, expansion, individual transformation, modification, or as the supraindividual transformation of organization. Based on this universal applicability, from cell theory to evolution, “organization” has been called “the key concept at all levels” for the life sciences (Figlio, 1976, p. 34).
## 2.4 "Organization" as One of the Three Basic Principles of Biology

"Organization" refers to the constitution of a system of interdependent parts. Two other important aspects of such a system that are related but not solely reducible to its constitution refer to the permanence and the transformation of the system. In well-known biological terms, they are called regulation and evolution. The three concepts refer to related but different aspects of organized systems: the causal pattern of their constitution, the capacity to control their relationship to the environment, and the potential for long-term transformation. The general meaning of all three principles operates on the same level of abstraction.<fnref n="4" />

*Organization*, in the systems-theoretical, Kantian tradition, essentially refers to the mutual dependence of parts in a system. *Regulation* refers to the stabilization of an organization by controlling environmental influences. Basically, regulation covers three processes: (1) supplying the system with necessary materials and other factors from its surroundings, (2) protecting it from detrimental influences, and (3) coordinating and integrating all the processes within the organized system. Taken together, they ensure the maintenance of the system, its preservation, and its perpetuation through time by managing the system's relationship with the environment. However, controlling the relation to the environment is not a conceptually necessary feature of organized systems. We can think of organized systems that are not regulated. Ecosystems might be an example for systems that are most certainly organized as their parts depend on each other. However, at least conceptually, we may think of them as not being controlled but more vulnerable to disturbances than organisms. In simple terms, *evolution* can be defined as the transgenerational transformation of organizations due to differential reproduction that is due to selection or genetic drift. The distinctness of "evolution" as a fundamental concept might be less controversial. Of course, we can imagine organisms that do not evolve, and most biologists did so until 1859.

The introduction of these three fundamental concepts in biology, organization, evolution, and regulation can be related to three conceptual turns or even “revolutions” in biology. They took place in the eighteenth, nineteenth, and twentieth century, respectively. The first is the revolution that established biology as a distinct scientific discipline at the end of the eighteenth century. It resulted in the conception

<footnote n="4">⁴There have been several attempts to separate the aspects of organization, regulation, and evolution of systems. Especially the relation between organization and regulation has been investigated from different angles, i.e., from the angle of economics with the distinction between internal order (organization) and external interventions (regulation) (Sombart, 1925) or the attempt to distinguish general systems theory from cybernetics (Bertalanffy 1951) or the efforts of autopoiesis theory to differentiate between (internal) self-organization and (external) control (Varela 1979). In all these cases, organization concerns the system-constituting internal structure of a dynamic entity, the regulation of its relation to the environment, especially the mechanisms of maintenance in the face of disturbances. For recent attempts to connect organization, closure, and regulation, see Di Paolo (2005) and Bich et al. (2016).</footnote>
of living beings as organisms. The second revolution was connected to evolution, to the insight that all life on earth is united in one all-encompassing process of transformation. The third revolution, the regulation revolution, took place mainly in the middle of the twentieth century and resulted in the description of organisms as cybernetic systems of control and information flow comparable to man-made machines.

One may think of organization, regulation, and evolution as a rather symmetrical trio: “organization” as the central category concerns the constitution of a system, “regulation” its stabilization, and “evolution” its transformation. However, it is also possible to derive the concepts from one linear argument. This argument begins with “organization” as the fundamental descriptive term for the constitution of living systems. It basically identifies a cycle of interdependent processes. Apart from this internal cycle, which grounds Kant’s internal purposiveness, there is an external cycle, a cybernetic feedback cycle that relates the system to its environment and stabilizes the system—the fundamental point of regulation. Regulation is directed toward the perpetuation of the system in time. This can be realized through two mechanisms: by stabilizing the individual system or by its multiplication, by the production of similar systems. Thus, we have two forms of self-preservation in organized systems: One is the regulation that consists in the preservation of individual systems, the maintenance of the dynamic state of an individual organism by devices for nutrition and protection. The other is preservation by multiplication of organizational types, which biologists call *reproduction*, the maintenance of the organizational structure of an individual by its multiplication in new individuals with a similar organization. In this view, reproduction is a preservation strategy by means of perpetuating an organizational type. Ironically, this most effective way of preservation has resulted in the vast process of transformation we call evolution.<fnref n="5" /> Therefore, reproduction leads to two contradictory consequences: On the one hand, it emerged as the most efficient means of self-preservation (“preservation by multiplication”). On the other hand, however, since it allows for variation (as mutations are inevitable and often even functional), this eventually results in the transformation of these systems. Ironically, preserving organization therefore means to transform it. As Paul Valéry surmised in an elegant and paradoxical formula: “Bios. Se transformer et transformer pour conserver” (Valéry, 1933, p. 755).

Thus, according to this argument, evolution is a derived feature of organized systems and regulated with respect to their maintenance. By seeking to maintain their systems through the most effective means at their disposal—reproduction—they initiate the process of transformation, which, in the long run, will erode their organization, at least in its original form.

<footnote n="5">⁵For the analysis of the relationship between organization and evolution, see also Ruiz-Mirazo et al. (2004) and Walsh (2006).</footnote>
## 2.5 Organization, Constraints, and Morphology

However, for evolution to begin in the first place, there must be an organized system of interdependent parts directed toward its own (or its type's) maintenance (for a critique of this view, see Ruiz-Mirazo et al., 2017). To understand the embodiment of such a system, the central concept of “constraints” has proven useful. Its conceptual history goes back to Gauss' principle of least constraint in classical mechanics. In the context of organization, it refers to the material structure or configuration of parts in a system that has a harnessing or channeling influence on the flow of energy within the system, for example, the structure of an organism's body, which serves as a boundary condition for physical laws. This general idea was described by Franz Reuleaux in his *Theoretical Kinematics* (1875).

Reuleaux is considered the founder of what has later been called “machine morphology.” According to Reuleaux, a machine is a “compound of resistant bodies, which is disposed in such a way that mechanical laws of nature are constrained to be effective under certain conditions” (Reuleaux, 1875, p. 38). This means that the effectiveness of a machine depends on the disposition of its parts, the structure of the whole, or its morphology. Morphology works by constraining the laws of nature. The machine does not introduce additional laws of nature; it simply channels or harnesses general laws through its morphology.

A hundred years later, Michael Polanyi applied this line of reasoning to biological systems. According to Polanyi, an organism has the same general makeup as a machine: its bodily structure serves as a boundary condition harnessing physical-chemical processes. In the case of the organism, this harnessing serves its organic functions (1968, p. 1308). Thus, quite surprisingly, life's irreducible structure rests on the very thing living beings have in common with machines: their specific structure (or morphology) that functions as a boundary condition in constraining laws of nature.

Starting in the late 1960s, Howard Pattee elaborated on this by stressing that it is not just the possession of functional constraints that is unique to living beings but the production and coordination of these constraints *by* the system itself. As he put it in 1971: “Life is distinguished from inanimate matter by the co-ordination of its constraints” (Pattee, 1971, p. 273). Organisms are embodied structures that produce their own structure, which feeds back on itself by maintaining and regenerating it. Thus, the boundary conditions or “constraints” of the system are, in the case of organisms, self-imposed. The structure of organismic bodies channels the energy in such a way that the body is preserved or rebuilt. The basic pattern is that of a cycle.

In recent years, a more detailed and precise rendering of this pattern has been provided by Alvaro Moreno and his collaborators. Since the early 1990s, his group has been describing the circular organization of systems on the basis of their components functioning as constraints. In their view, local constraints within the system are generated by the activity of components of the system (Moreno et al., 1994, p. 17; see also Ruiz-Mirazo & Moreno in this volume). They call the resulting
system an “autonomous organization” since the system itself generates and regenerates its constraints.

Stuart Kauffman called this circular organization “a virtuous cycle,” a cycle of works and constraints: The work of the system generates constraints, namely, precisely those constraints required for the work to be done. According to Kaufman, this cycle is “the heart of a new concept of ‘organization’” (Kauffman, 2000, p. 4).

In the last few years, Matteo Mossio has further elaborated on these matters. According to Mossio and his colleagues, biological organization is characterized by the fact that it realizes a specific kind of causal regime. This regime is based on nothing but the material structure of an organism acting as constraints for the physical laws. Because the constraints within the organization are mutually determined, the “organizational closure” of organisms consists in a “closure of constraints” and therefore in a biological self-determination (Montévil & Mossio, 2015, p. 180).

An important consequence of this account is that biological autonomy, in the double sense of biology as a distinct discipline and of organisms as self-determining systems, is entirely based on structures. Nothing but the structure of an organism embodies the constraints that effectively control the boundary conditions for the laws of nature. Biological autonomy is grounded in the material form of living bodies. Biology’s distinct causal regimes, once referred to as “life-forces,” are embodied in the forms of organisms.

With this emphasis on form, the biological subdiscipline of *morphology*, which has been marginalized for over a century now, is again taking center stage. Ultimately, it is morphology that provides the basis for the organism as an integrated autonomous system because it provides the only factor beyond the laws of nature that is specific to organisms. Insofar as organisms are considered to be autonomous, they are determined by their form. Form is the only additional factor that distinguishes organisms from inorganic bodies. This is true, at least at the explanatory level, because forms provide the only specific biological causal factor. To put it bluntly: The only life forces that exist are life forms.

Thus, morphology, the study of forms, is the fundamental explanatory principle of biology. It has always been fairly easy for biologists to identify functions in living beings. Aristotle was famously prolific at it, and his well-known functional categories, such as nutrition, growth, and reproduction, are still being used today. However, although they may still define what it means to be alive, functions do not necessarily provide causal understanding of processes. In biology, this is done by identifying mechanisms; and mechanisms are based on morphology, because it is morphology that identifies the structures that function as (self-)constraints within natural organizations, establishing them as a distinct type of material bodies.

Organic forms, then, are the mediators for the realization of biological organizations. They instantiate these organizations in specific living bodies; in their function as particular “constraints,” they enable the causal interdependence of the components and the self-referentiality of the whole system. Thus, “organization” and “form” are two complementary aspects of living bodies: the first refers to the causal pattern that constitutes the unity of the system and the second to the individuality of the system and the specific material “constraints” by which this causal pattern is
realized and instantiated in concrete living beings. Or, in other words, “organization” provides the law-like universal feature of all living beings (existent and potential), “forming” the physical realization of its causal pattern in distinct instances. Biological explanations demonstrate how forms are effective as functions, i.e., how they are integrated in functional closure. The fixed form of the heart chambers (in a particular individual or in the type of individuals of a certain class) explains how this form constrains the general laws in order to achieve closure.

"Organization" and "form" both fulfill *descriptive* and *explanatory* functions in biology, albeit at different levels: The first visible feature of living beings is, of course, their form; forms are *described* and are the basis for biological classifications (as they indicate genealogical relationships). However, forms also *explain* (and only they can do that) how the forces and energy flows in an organism are channeled to realize the functional closure that is characteristic of every living being. "Organization," on the other hand, is *descriptive* with respect to the causal pattern that all living beings have in common; this pattern, causal "cyclicity" or functional "closure," is identified when an entity is described as being organized.<fnref n="6" /> However, on a more abstract level, "organization" gives the *explanation* to the fact that all living beings share a certain functional order.<fnref n="7" /> This explanation consists in the specification of the causal pattern or the working order of every organism, namely, the self-referentiality of all their activities, their orientation toward self-maintenance (what the developmental biologist Wilhelm Roux once called the "autophely" or "self-utility" of organisms; Roux, 1895, p. 58). Since "organization" specifies this causal and functional pattern common to all living beings, it grounds the biological approach toward nature. At the same time, as there are living beings on earth that do not always behave autophelically, "organization" also marks the end of the biological perspective focused on autophely: "Man has reached a level of existence that stands above purpose. It is his distinctive value that he can act without purpose" (Simmel, 1918, p. 28).

## References

* Anonymous. (1765). Organisation. In *Encyclopédie ou Dictionnaire raisonné des sciences, des arts et des métiers* (Vol. 11, p. 629). Briasson.

* Beckner, M. (1959). *The biological way of thought*. Columbia University Press.

<footnote n="6">⁶For Kant, this identification procedure takes place in reflective judgments, and he is very clear about his view that they are not explanatory, but descriptive: "positing ends of nature in its products, insofar as it constitutes a system in accordance with teleological concepts, belongs only to the description of nature" (Kant, 1790, p. 417).</footnote>

<footnote n="7">⁷"Form," however, is also a concept that involves abstractions. In most biological contexts, it does not refer to individual bodies but to structural aspects that numerous individuals (of one species or of another taxonomic unit) have in common. Nevertheless, for every specific causal interaction, it is the realization of one form in an individual organism that serves as constraint.</footnote>
Bernard, C. (1867). *Rapport sur les progrès et la marche de la physiologie générale en France. L'Imprimerie Impériale.*

Bich, L., Mossio, M., Ruiz-Mirazo, K., & Moreno, A. (2016). Biological regulation: Controlling the system from within. *Biology and Philosophy*, **31**, 237–265.

Boerhaave, H. (1708). *Institutiones medicae*. Linden.

Boerhaave, H. (1727). *Historia plantarum*. Gonzaga.

Bolton, R. (1978). Aristotle’s definitions of the soul: *De anima* II, 1–3. *Phronesis*, **23**, 258–278.

Bos, A. P. (2003). *The soul and its instrumental body. A reinterpretation of Aristotle's philosophy of living nature*. Brill.

Bourguet, L. (1729). *Lettres philosophiques sur la formation des sels et des crystaux et sur la génération & le mécanisme organique des plantes et des animaux*. Honore.

Cheung, T. (2006). From the organism of a body to the body of an organism: Occurrence and meaning of the word ‘organism’ from the seventeenth to the nineteenth centuries. *The British Journal for the History of Science*, **39**, 319–339.

Collegium Conimbricense. (1600). *Commentarii Collegii Conimbricensis Societatis Iesv, in tres libros De anima Aristotelis Stagiritae Coimbra*. Zetzner.

Cuvier, G. (1812). *Recherches sur les ossemens fossiles de quadrupèdes* (4 vols.). Deterville.

Cuvier, G. (1817). *Le règne animal, distribué après son organisation* (4 vols.). Deterville.

Darwin, C. (1860). *On the origin of species* (rev ed.). Appleton.

de La Mettrie, J. O. (1747). L'Homme machine (A. Vartanian, Ed.). Princeton University Press 1960.

de Lamarck, J. B. (1815). *Histoire naturelle des animaux sans vertèbres* (Vol. 1). Verdière.

Des Chene, D. (2000). *Life's form. Late Aristotelian conceptions of the soul*. Cornell University Press.

Di Paolo, E. A. (2005). Autopoiesis, adaptivity, teleology, agency. *Phenomenology and the Cognitive Sciences*, **4**(4), 429–452.

Diderot, D. (1778). *Éléments de physiologie* (P. Quintili, Ed.). Champion 2004.

Figlio, K. M. (1976). The metaphor of organization: an historiographical perspective on the bio-medical sciences of the early nineteenth century. *History of Science*, **14**, 17–53.

Fischer, K. (1865). *System der Logik und Metaphysik oder Wissenschaftslehre* (2nd ed.). Bassermann.

Girtanner, C. (1790). Mémoires sur l'irritabilité, considérée comme principe de vie dans la nature organisée. *Observ Phys History Natural Arts*, **37**, 139–154.

Glisson, F. (1672). *Tractatus de natura substantiae energetica, seu de vita naturae*. Flesher.

Guillo, D. (2003). *Les figures de l'organisation. Sciences de la vie et sciences sociales au XIXe siècle*. Presses universitaires de France.

Hartbecke, K. (2006). Metaphysik und Naturphilosophie im 17. Jahrhundert. Francis Glissons Substanztheorie in ihrem ideengeschichtlichen Kontext. Niemeyer.

Hertwig, O. (1900). *Die Entwicklung der Biologie im 19. Jahrhundert*. Fischer.

Hooke, R. (1665). *Micrographia*. Martyn.

Jacob, F. (1973). *The logic of life* (1970), transl. by Betty E. Spillmann. Pantheon.

Jungius, J. (1662). *Doxoscopiae physicae minores*. Naumann.

Kant, I. (1790). *Critique of the power of judgment* (P. Guyer, Ed.). Cambridge University Press 2004.

Kant, I. (1795). [Letter to Samuel Thomas Soemmerring, 10 Aug. 1795]. In *Akademie-Ausgabe*, vol. XII. de Gruyter 1922, pp. 30–35.

Kant, I. (OP). Opus postumum. In: A. Buchenau (Ed.), Kant's Opus postumum, vol. 1 (= Akademie Ausgabe, vol. XXI). de Gruyter 1936.

Kauffman, S. (2000). *Investigations*. Oxford University Press.

Kolasa, J., & Pickett, S. T. A. (1989). Ecological systems and the concept of biological organization. *Proceedings of the National Academy of Sciences of the United States of America*, **86**, 8837–8841.

Kühnemann, E. (1924). Kant (Vol. 2). Das Werk Kants und der europäische Gedanke. Beck.
Lamarck, J. B. (1797). *Mémoires de physique et d'histoire naturelle*. Auteur.

Leibniz, G. W. (1705). Considerations sur les principes de vie, et sur les natures plastiques. Philosophische Schriften, Vol. 4. Suhrkamp 1996, pp. 327-347.

Locke, J. (1689). *An essay concerning human understanding*. Clarendon Press 1979.

Maturana, H. R., Varela, F. J., & Uribe, R. (1974). Autopoiesis: The organisation of living systems, its characterization and a model. *Biosystems*, *5*, 187-196.

Montévil, M., & Mossio, M. (2015). Biological organization as closure of constraints. *Journal of Theoretical Biology*, *372*, 179-191.

More, H. (1659). *The immortality of the soul*. Nijhoff 1987.

Moreno, A., Umerez, J., & Fernandez, J. (1994). Definition of life and the research program in artificial life. *Ludus Vitalis*, *2*, 15-33.

Needham, J. T. (1750). *Nouvelles observations microscopiques, avec des découvertes intéressantes sur la composition et la décomposition des corps organisés*. Ganeau.

Pattee, H. H. (1971). Physcial theories of biological co-ordination. *Quarterly Review of Biophysics*, *4*, 255-276.

Piaget, J. (1967). *Biologie et connaissance*. Gallimard.

Piché, C. (2001). Kant et les organismes non vivants. In L. Cournarie & P. Dupond (Eds.), *Préparer l'agréation de philosophie. La nature* (pp. 83-93). Ellipses.

Pittendrigh, C. S. (1993). Temporal organization: Reflections of a Darwinian clock-watcher. *Annual Review of Physiology*, *55*, 16-54.

Plessner, H. (1928). *Die Stufen des Organischen und der Mensch*. de Gruyter 1975.

Polanyi, M. (1968). Life's irreducible structure. *Science*, *160*, 1308-1312.

Porphyrios, De abstinentia. In *Opuscula selecta* (A. Nauck, Ed.). Leipzig 1886, Engl.: On absti-nence from killing animals, transl. by G. Clark. Ithaca. Cornell University Press 2000.

Quarantotto, D. (2010). Aristotle on the soul as a principle of biological unity. In S. Föllinger (Ed.), *Was ist Leben? Aristoteles' Anschauungen zur Entstehung und Funktionsweise von Leben* (pp. 35-53). Steiner.

Reuleaux, F. (1875). *Theoretische Kinematik. Grundzüge einer Theorie des Maschinenwesens*. Vieweg.

Roux, W. (1895). Ziele und Wege der Entwickelungsmechanik. In *Gesammelte Abhandlungen über Entwickelungsmechanik der Organismen* (Vol. 2, pp. 55-94). Engelmann.

Ruiz-Mirazo, K., & Moreno, A. (this volume). On the evolutionary development of biological organization from complex prebiotic chemistry. In M. Mossio (Ed.), *Organization in biology*. Springer.

Ruiz-Mirazo, K., Peretó, J., & Moreno, A. (2004). A universal definition of life: Autonomy and open-ended evolution. *Origins of Life and Evolution of Biospheres*, *34*, 323-346.

Ruiz-Mirazo, K., Briones, C., & Escoura, A. (2017). Chemical roots of biological evolution: The origins of life as a process of development of autonomous functional systems. *Open Biology*, *7*, 170050.

Schiller, J. (1978). *La notion d'organisation dans l'histoire de la biologie*. Maloine.

Schmid, C. C. E. (1799). *Physiologie philosophisch bearbeitet* (Vol. 2). Akademische Buchhandlung.

Sextus Empiricus, Adversus dogmaticos libros quinque (= Adversus mathematicus VII-XI). Against the Logicians, transl. by R. Bett. Cambridge University Press 2005.

Siegel, R. E. (1973). *Galen on psychology, psychopathology, and function and diseases of the nervous system*. Karger.

Simmel, G. (1918). *The view of life. Four metaphysical essays with journal aphorisms* (J. A. Y. Andrews, & D. N. Levine, Trans.). The University of Chicago Press 2010.

Sombart, W. (1925). *Die Ordnung des Wirtschaftslebens*. Springer.

Stahl, G. E. (1707). *De vera diversitate corporis mixti et vivi*. Orphanotropheum.

Toepfer, G. (2011a). *Historisches Wörterbuch der Biologie* (3 vols.). Metzler.

Toepfer, G. (2011b). Kant's teleology, the concept of the organism, and the context of contemporary biology. *Logical Analysis and History of Philosophy*, *14*, 107-124.
Valéry, P. (1933). *Bios. In Cahiers, Vol. 2. Gallimard, 1974*.

Varela, F. J. (1979). *Principles of biological autonomy*. North-Holland.

Voltaire (1772). *Vie. In Questions sur l'encyclopédie, vol. 9. pp. 55–58*.

von Bertalanffy, L. (1928). *Kritische Theorie der Formbildung*. Borntraeger.

von Bertalanffy, L. (1951). Towards a physical theory of organic teleology. *Human Biology*, *23*, 346–361.

von Uexküll, J. (1903). Studien über den Tonus, I. Der biologische Bauplan von *Sipunculus nudus*. *Zeitschrift für Biologie*, *44*, 269–344.

Walsh, D. (2006). Organisms as natural purposes: The contemporary evolutionary perspective. *Studies in History and Philosophy of Biological and Biomedical Sciences*, *37*, 771–791.

Wicken, J. S. (1987). *Evolution, thermodynamics, and information: Extending the Darwinian program*. Oxford University Press.

Wolf, J. H. (1971). Der Begriff »Organ« in der Medizin. Grundzüge der Geschichte seiner Entwicklung. Fritsch.

Wolfe, C. T. (this volume). Varieties of organicism: A critical analysis. In M. Mossio (Ed.), *Organization in biology*. Springer.

**Open Access** This chapter is licensed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license and indicate if changes were made.

The images or other third party material in this chapter are included in the chapter's Creative Commons license, unless indicated otherwise in a credit line to the material. If material is not included in the chapter's Creative Commons license and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder.

<figure><img src="image_8.png" /></figure>
Chapter 3

Varieties of Organicism: A Critical
Analysis

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

Check for
updates

Charles T. Wolfe

**Abstract** In earlier work I wrestled with the question of the “ontological status” of organisms. It proved difficult to come to a clear decision, because there are many candidates for what such a status is or would be and of course many definitions of what organisms are. But what happens when we turn to theoretical projects “about” organisms that fall under the heading “organicist”? I first suggest that organicist projects have a problem: a combination of invoking Kant, or at least a Kantian “regulative ideal,” usually presented as the *epistemological* component (or alternately, the complete overall vision) of a vision of organism – as instantiating natural purposes, as a type of “whole” distinct from a merely mechanistically specifiable set of parts, etc. – and a more ontological statement about the inherent or essential features of organisms, typically presented according to a combination of a “list of heroes” or “laundry list” of properties of organisms. This amounts to a category mistake. Other problems concern the too-strict oppositions between mechanism and organi ci sm, and symmetrical tendencies to “ontologize” (thus objectifying) properties of organisms and to “subjectify” them (turning them into philosophies of subjectivity). I don’t mean to suggest that no one should be an organicist or that Kant is a name that should be banished from civilized society. Rather, to borrow awkwardly from Sade, “organicists, one more effort!” if one wants a naturalistic, non-foundationalist concept of organicism, which is indeed quite active in recent theoretical biology, and which arguably was already alive in the organismic and even vitalist theories of thinkers like Goldstein and Canguilhem.

In an earlier paper (Wolfe, 2010), I wrestled with the question of the “ontological status” of organisms. It proved difficult to come to a decision on this matter, because there are many candidates for what such a status is or would be and of course many definitions of what organisms are (see also Pepper and Herron (2008), Pradeu (2010a), and Pradeu (2016), Bouchard & Huneman, 2013) – the “fungus problem,” as it were. But I did not focus in any detail on actual contemporary projects that rely on, or even assert, a certain concept of organism – namely, antireductionist,

<footnote>C. T. Wolfe (✉)
Université de Toulouse 2 Jean-Jaurès, Toulouse, France</footnote>
organicist projects (I’m fully aware that I haven’t defined organicism yet). And I wish to show that these projects have an “issue,” a sort of conceptual confusion, at least in part. My concern is thus not to argue for or against organicism, or to reconstruct a variant of organicism that is immune to some recent objections (a common enough practice), but rather, echoing two very different authors, to say “One more effort to achieve an organicism worth wanting.”<fnref n="1" />

## 3.1 Organicism as Ontology, as Epistemology or as a Blurry Mix

A curious consensus without examination of presuppositions reigns today in the various subfields concerned with the properties (organizational, systemic, etc.) of living beings, at least those which define themselves as more or less “organicist.” There is no strictly defined group of organicists, but attempts are regularly made to group various antireductionist trends in biology under this heading (e.g., Gilbert & Sarkar, 2000; Nicholson, 2014). Descriptions of how “the whole is greater than the sum of the parts” or, slightly less generically, how specific “system-level” or “higher-level” properties need a set of explanations (and perhaps an ontology) appropriate to them abound and have been repeated almost identically since the early twentieth century, with predecessors such as Aristotle and Kant invoked almost as often. At times, the concept of organism at work in these discussions is mostly a placeholder, such as when Lewontin, and developmental systems theory more generally, stresses that the focus is on organism-environment relations (“Just as there is no organism without an environment, so there is no environment without an organism”<fnref n="2" />). But more current organicism included various recent proclaimed “returns” of the organism concept<fnref n="3" /> and does not just treat the concept as a placeholder, a functional or a heuristic term. They move toward what I’ll call an *ontologization* of the notion: at that point, organicism has an ontology, whether it is clearly stated as such or not – it is then an *objectified* account of the nature of organisms.

<footnote n="1">¹ I am thinking both of Sade’s “Français, encore un effort si vous voulez être républicains” (a pamphlet read out in the fifth dialogue of *La philosophie dans le boudoir*: Sade 1795/1998) and Dennett’s elegant notion (the subtitle of his *Elbow Room*) of a notion of free will “worth wanting” (e.g., a pure unconditioned free will might be a lovely notion, but not accessible in our physical, causal universe, so which notions are *worth wanting*? Dennett, 1984).</footnote>

<footnote n="2">² Lewontin, 1983/1985, 99. Pradeu has argued that in certain formulations of DST, the notion of “organism” is lost when it is posited that what evolves is actually the “organism-environment system”; he speaks of the “loss of the uniqueness of the organism” (Pradeu, 2010b, 219). Thanks to Alejandro Fábregas Tejeda for this reference.</footnote>

<footnote n="3">³ See Laubichler, 2000a, b and Baedke, 2019 (as well as Baedke’s project at Bochum titled “The Return of the Organism in the Biosciences: Theoretical, Historical and Social Dimensions” (https://rotorub.wordpress.com/ although this aspect is primarily an effect of the ‘project description’ language), and the less commemorative or exhortatory special issue I coedited (Huneman and Wolfe, eds., 2010).</footnote>
Treatments of organisms as more than just placeholders in a theoretical biologi-
cal system, indeed, as entities with objective properties that should be the object of
an ontology do not pose a problem as such – except perhaps to the instrumentalist,
for whom it is a mistake to attribute any kind of innate meaning or definition to
biological entities, as these are merely instrumental (Wolfe, 2010). What seems
more curious to me is the way these treatments variously blend epistemological and
ontological versions of organicism, sometimes in awkward ways. Generally, these
accounts involve a combination of:

(a) an invocation of Kant, or at least a Kantian “regulative ideal,” usually presented as the *epistemological* component (e.g., organicism as a theory about what constitutes “knowledge of life”) or alternately, as the complete account of a vision of organism – as instantiating natural purposes, as a type of “whole” distinct from a merely mechanistically specifiable set of parts, interdependency, self-organization, and so on (Weber & Varela, 2002; Longo et al., 2012).

(b) a more *ontological* statement about the inherent or essential features of organ-
isms (organicism as a theory about the actual properties of living systems) –
essentially a “laundry list” of core features, e.g., “reproduction, life-cycles,
genetics, sex, developmental bottlenecks, germ-soma separation, policing
mechanisms, spatial boundaries or contiguity, immune response, fitness maxi-
mization, cooperation and/or conflict, codispersal, adaptations, metabolic
autonomy, and functional integration” (Clarke, 2013, 415), itself usually tied to
a kind of special tradition of thinkers who had a “feeling for the organism”:
typically, Claude Bernard,<fnref n="4" /> Sherrington and Waddington, and more recently
Maturana and Varela with the notion of autopoiesis, Rosen’s (M,R) systems,
Gánti’s chemoton, or Luisi’s minimal cell, all of which at one time or another
are presented as the defining “characteristics of life,” or “life criteria” (Griesemer
& Szathmáry, 2008). These authors, or perhaps their theoretical constructs, are
“heroes of homeostasis,” as it were, or “masters of metabolism,” given the latter
concept’s close association with definitions of life, e.g., when it is defined as
“the collection of chemical reactions that define a living organism and allow it
to make its components and obtain the energy required for staying alive”
(Cardenas & Cornish-Bowden, 2011). To be precise, Bernard and others named
above do not all perfectly match Clarke’s “laundry list” of core features of life,
e.g., fitness maximization. I used her list because it is one of the more expansive
yet precise such lists I’ve seen.

The combination of (a) and (b) poses several problems. Note that I am not sug-
gesting that a theory of organism has to be either epistemological or ontological. It
can be both, or neither (what one might term constructivism, i.e., the view that
organisms are constructs, and furthermore inasmuch as they are “sense-making”

<footnote n="4">⁴See Bechtel and Turner’s contributions to Normandin and Wolfe eds. (2013) (which strongly present Bernard as a founder of the tradition they work in), as well as Denis Noble’s influential lecture (Noble, 2008) which literally speaks in its title of Bernard as the “father of systems biology.”</footnote>
entities, they need to be understood via acts of cognitive construction).<fnref n="5" /> What seems problematic to me is to combine epistemological and ontological claims in an “ungrounded” way, not always clearly presented as such (but, conversely, presented as straightforwardly empirical).

Perhaps the most obvious difficulty created by the unreflexive combination of (a) and (b) is something like a category mistake, in this case treating an epistemological property as an ontological property;<fnref n="6" /> it is not quite right to invoke the authority of the Kantian “projective” approach to organisms (Huneman, 2007) in order to assert a set of ontological specificities about organisms. Because this is precisely what the Kantian regulative ideal concept was designed to avoid, in explicit contrast to what he would have called “rational metaphysics.” That is to say, to provide an empirical set of criteria for why living beings are special and to claim that this supports or is supported by a Kantian framework is not a good idea given that the latter framework explicitly rejects the idea of giving empirical definitions of organism, inasmuch as Kant’s organism concept is explicitly built around his notion of regulative ideal.<fnref n="7" /> For Kant, organism is a “reflective” construct rather than a “constitutive” feature of reality, and reflective judgments are “incapable of justifying any objective assertions.”<fnref n="8" /> The purposive character of organisms, on this view, imposes itself upon us by empirical experience, forcing us to (cognitively) recognize such entities as “organisms,” rather than as machines or mere bundles of matter. Note that this more “constructivist” approach is by no means ruled out in biology, where such regulative concepts can serve as heuristics, e.g., regulative teleology;<fnref n="9" /> the problem lies more in claiming to have an “empirical Kantian” foundation for organicism, because of the inherent tension therein – as when, e.g., Weber and Varela state that Kantian teleology is somehow “true” of organisms; “Our immodest conclusion is that Kant, although foreseeing the impossibility of a purely mechanical, Newtonian account of life, nonetheless was wrong in denying the possibility of a coherent explanation of the organism” (Weber & Varela, 2002, 120). As noted earlier, I am not suggesting that all organicists are Kantians or (a narrower claim) that all organicists make a category mistake with respect to Kantian notions; indeed, some contemporary organicists call for a return to Aristotle, while earlier organicists could also place themselves under the patronage of Hegel (thereby, by the way, eliminating

<footnote n="5">⁵I have developed this idea of a constructivist standpoint on organisms further with reference to Canguilhem and Goldstein in Wolfe (2015b, 2017) and more broadly in Wolfe (2014a).</footnote>

<footnote n="6">⁶I am using this term to simply mean “The error of assigning to something a quality or action which can only properly be assigned to things of another category, for example treating abstract concepts as though they had a physical location” (Stevenson, 2010), rather than the more specific sense of a semantic error in a syntactically valid statement, like Chomsky’s famous “colorless green ideas sleep furiously.”</footnote>

<footnote n="7">⁷Kant, 1987, § 73, p. 276.</footnote>

<footnote n="8">⁸Kant, 1987, § 67, p. 259; § 73, p. 277.</footnote>

<footnote n="9">⁹On regulative concepts as heuristics, see Sloan (2007) (discussing Grene) and Grene (1968, 235 f.); for a more critical view of the utility of Kantian regulative concepts in biology, see Zammito (2006); and for a more “objectivist” view, see Mossio and Bich (2017).</footnote>
the category mistake in one stroke, as that is a full-blown ontological realism about organisms) (for a rare recent Hegelian move in organicism, see Gambarotto and Illetterati (2020)).

The second difficulty in certain organicist arguments lies in their empirical stance: it is not clear in any case why it counts as an argument against “mechanism” or “reductionism” to present a list of key features (such as the “generative” circular causality in organisms: Mossio, 2020, 55, 57), especially from a standpoint that curiously combines Kantianism, or at least Kantian elements, and the ontologization of organisms. I called this the “laundry list” problem above;<fnref n="10" /> notice that the problem with this form of organicism – a kind of literal, empirical claim listing key definitory features of organisms – is very close to that found in defenses of “definitions of life,” particularly “criterial” definitions of life (Malaterre, 2010; see Bich and Green (2017) for a defense of definitions of life).

A different reason why the laundry list of the organic is not the argument-stopper some treat it as is the flexibility of mechanism: mechanism is not reducible to the strictures of say early modern mechanistic ontology (in which what is real is what is specifiable in terms of size, shape, and motion) or to a “toy model” ontology in which what is real in a system is its decomposition into parts (indeed, decomposition is typically presented as an *explanatory* strategy in contemporary mechanism, e.g., in Bechtel & Richardson, 1993, 30; here I primarily mean decomposition into parts). Organicism in this sense neglects, or almost forces itself to neglect, the reality of “expanded mechanism,” that is, the existence of nondogmatic, or non-foundationalist, or ontologically open mechanistic programs. In this sense organicism can be overly polarized (this is my disagreement with Nicholson, specifically Nicholson (2013); for a more open-ended view, reflecting its historical focus, see Toepfer, this volume). By extension, organicism can at times run the risk of verging on antinaturalism, although antimechanism is obviously not synonymous with antinaturalism (I’ll return to this below). Either organicism neglects the possibility of the reducibility of the laundry list (i.e., that it is potentially reducible to more basic features), or conversely, of the existence of expanded mechanism: the “remit,” the “span” of mechanism is broader than we are accustomed to think when we repeat the mechanism-vitalism or mechanism-teleology opposition. One could adduce the sheer historical complexity of mechanism and its various “expansions,” or the concept of teleomechanism – not in its historically “restricted” sense as in the Lenoir Thesis (Lenoir, 1982), but in a broader sense (Wolfe, 2014c), which stresses that even paradigm cases of “mechanism” or “the mechanical philosophy” (from Descartes and iatromechanism onward) are filled with functional language, including “function,” “use,” and the “office” of an organ. And they are concerned with properties of the organism such as health and survival, that is, not just with

<footnote n="10">¹⁰Within the autopoiesis discussions, I note that E. Di Paolo diagnosed this problem (see Di Paolo, 2018). But he also suggests that Varela himself became critical of the autopoietic theory for this “lazily empirical” character and refers to the 1996 preface Varela added to the second Spanish edition of the canonical text *De Máquinas y Seres Vivos*.</footnote>
microstructure or size, shape, and motion, if indeed, “medicine is the most useful of the sciences” for Descartes (*Discourse on method*<fnref n="11" />).

## 3.2 A Current Organicist Consensus?

Of course, the theoretical biologist interested in articulating a concept of organizational closure<fnref n="12" /> might say: none of this matters, because what these “positions” are, are simply theoretical constructs, *bricolages* intended to facilitate the articulation of an ongoing research project. This can be seen in an example from the earlier part of the twentieth century, Kurt Goldstein’s *The Structure of the Organism* [Goldstein, 1939/1995], a work which combines a “bottom-up” study of brain-damaged soldiers from World War I with a “top-down” set of ontological, not just tacit commitments but explicit *claims* about the “whole”: if we treat Goldstein’s references to Goethe and *Naturphilosophie* seriously, we can charge him with “romanticism,” with being “antimodern science,” and so on. But if we treat these references as simply a typical case of an educated German scientist of the early twentieth century displaying his “humanistic” breadth, then we can study the overall argument in much more naturalistic, or naturalism-friendly terms.

But the goal of the present remarks is to clarify the possible or actual commitments of these organicist projects in current research, to ask if they hang together successfully or not, and to distinguish between them in a way that has not previously been attempted. This is justified also by the fact that many of the actors involved go out of their way to claim a Kantian (or in some cases Aristotelian<fnref n="13" />) pedigree for their experimental, modeling, or otherwise “empirical” projects.

Now, it is not easy to produce a typology of current organicist views, partly because the actors involved waver between positions that we might consider clearly demarcated (notably, epistemological vs ontological positions on organisms and their “reality,” as in the (a) and (b) positions above). Varela, for one, sometimes stresses an “epistemological” standpoint but sometimes also speaks the language of

<footnote n="11">¹¹Descartes writes that the preservation of health is the most important value and the foundation of all other values; that states of mind are dependent on the organization of bodily organs; and that (hence) medicine is the most useful and necessary science (*Discourse*, VI, AT VI, 62–63).</footnote>

<footnote n="12">¹²Mossio et al., 2009; Mossio and Moreno 2010; Ruiz-Mirazo et al., 2000; and discussion in Bechtel, 2007; Moreno & Mossio, 2015.</footnote>

<footnote n="13">¹³The work of Marjorie Grene (who called for an Aristotelian perspective in biology and ‘philosophical biology” in the 1960s), James Lennox and Denis Walsh is influential here: see notably Walsh, 2021. In the context of the present paper, Aristotelian perspectives are squarely ontological. As Walsh puts it, Aristotelian biology (indeed, any “comparative biology”) “seeks to account for the fit and diversity of organismal form” (Walsh, 2021, 281). While Walsh also speaks of explanations, I would understand his insistence on the need for an ‘agent theory” with which to understand organisms (because it is able to accommodate teleological explanations), rather than an ‘object theory”, as ontologically revisionary (in Strawson’s sense).</footnote>
the *key features of organisms*. Rosen (1991) says he is speaking about “models” but then calls his book *Life Itself* and indeed complains that mechanistic science has not grasped “life itself.” He also holds that mechanistic laws are the special cases, the outliers, whereas biology gives us a truer picture of nature; but that is not our concern here.

## 3.3 Organicism Strong and Weak, and the Ghost of Vitalism

Interestingly, as regards the “autonomy and organizational closure,” researchers never seem to make this particular move (different from what I called a category mistake above) of reifying “life” or “organization” in a foundationalist way. The concept that emerges here which seems to overcome the epistemological/ontological divide is *organization*. Because, as can be seen with the case of Bechtel, both “organicists” and “mechanists” can help themselves to the concept of organization; differently put, organization can be a key concept for both of these approaches, and it can end up being more a matter of perspective than of some purported deep ontology that one specifies this concept in mechanistic terms (especially, expanded-mechanist terms) or in organicist terms (especially, non- or weakly ontologized organicist terms: Wolfe, 2010) – the same can be said of the notion of “structure.” I use the term “perspective” because the choice of studying a system as mechanistic and/or as organizational is also a choice of perspective, or “standpoint” as Needham put it: “all things are organisms and all things are atomic systems also. You choose your standpoint” (Needham, 1930, 85, longer citation in Wolfe, 2014a). However, it is nevertheless possible that any robust notion of organization, unlike older notions of systems à la von Bertalanffy (1933), does rely on a concept of life (and thus is not tantamount to mechanism).<fnref n="14" />

Further, the Kantian/epistemological view can also be transfigured into something “ontological,” not as a claim about the world (i.e., entities such as livers and hearts or microbes or finches or coral reefs) but as a claim about the unique nature of the subject (interiority, subjectivity, first-person knowledge, etc.,<fnref n="15" /> in which these are distinguished from the physical world as a whole). This is not so surprising if we consider Kant’s famous proclamation that “there will never be a Newton of a blade of grass”: a point which is both about our cognitive capacities *and* – at least the ambiguity is unresolved – about the uniqueness of organic entities in comparison with the entities studied by physics and the physicomechanical sciences more broadly. In the view of some notable interpreters, “the third *Critique* essentially proposed the reduction of life science to a kind of pre-scientific descriptivism, doomed never to attain authentic scientificity, never to have its ‘Newton of the blade

<footnote n="14">¹⁴ I am grateful to Bohang Chen for this suggestion.</footnote>

<footnote n="15">¹⁵ Varela, 1996; Varela & Shear, 1999; Thompson, 2007.</footnote>
of grass”” (Zammito, 2006, 755). Actually, Newtonianism was very fruitful as a methodology and an analogy in eighteenth-century life science, both in obvious cases like Albrecht von Haller’s physiology—the first sentence of his textbook on physiology famously proclaims that “the fibre is to the physiologist what the line is to the geometrician”— and in less obvious cases like Montpellier vitalism, which is also filled with Newtonian methodological invocations (Wolfe, 2014b).

This slippage from the epistemological to the ontological is exemplified in earlier twentieth-century theoretical biology in work like Uexküll’s, in which the “epistemic” (the states of knowledge of the world of given biological entities such as the tick: Uexküll, 1934/2010) becomes a feature of the organic as such. Similarly, Goldstein’s theory of organism (Goldstein, 1939/1995), which is a really a theory of the organismic (actually, holistic) capacities of meaning-making and world-making by agents such as humans; it is really more about holism and personhood than about biological organisms per se, but in any case Goldstein’s theory actively moves between these two positions, but that should not prevent us from trying to articulate the difference for the sake of clarification. When Grene discusses phenomenological and philosophico-anthropological approaches to life, including Goldstein’s (esp. Grene, 1966), like Goldstein himself, she elides the difference between states and processes of “the knower” (the subject) and objective properties of “the known” (the object). While most phenomenologists would claim to only be describing the former, Grene shifts the ground to describing the latter: matters of “ontological import” (Grene, 1968, 239).

When organicism is a strong ontology (e.g., Grene, 1966, 1968), which I have also discussed under the heading of “ontologization,” is it so different from its yet-stronger cousin, vitalism? That is to say, organicism recurrently tries to present itself as a “safe word,” the reasonable version of antireductionism (Gilbert & Sarkar, 2000: vitalism is the weird version of our view, so let’s call it organicism), while some biologists will playfully invoke “molecular vitalism” in mainly provocative ways (Kirschner et al., 2000). Indeed, this attempt to present organicism as, typically, the less metaphysical version of vitalism – today one would tend to say the “naturalized” version is an old, perhaps defining feature of this theory (or family of theories, since early twentieth-century organicism is not exactly cohesive in those terms). For instance, Needham wrote that organicism combined “the insistence of vitalism on the real complexity of life with the heuristic virtues of the mechanistic practical attack.”<fnref n="16" />

<footnote n="16">¹⁶Needham (1936), 9; cf. also Needham (1928), discussed in Nicholson and Gawne (2015); see also Peterson (2017). Autopoietic theory also will state that its novelty lies in its being neither reductionism nor vitalism (Varela, 1991). Interestingly, Beckner (1974) thought almost the perfect opposite of Needham, in the sense that for him, organicism only really was defensible (or could survive) if supplemented by vitalism. Beckner also uses the language of “weak” and “strong” organicism, but in his case these terms are indexed to the concept of emergence, that is, they mean the same as “weak emergence” and “strong emergence.” O. Sartenaer (discussion) notes nicely that anything in between reductionism and (strong) vitalism is emergentism.</footnote>
Now, it would be unfortunate if organicism operated with as caricatural a presentation of vitalism as the Cricks and Monods do, for vitalism itself lives in serious tension between different meanings, which indifferent history of science or philosophy often neglects. An eighteenth-century Montpellier vitalist, Bichat, Bernard (both of whom combine denunciations of earlier forms of vitalism with insistence on the existence of uniquely vital properties, Wolfe (2022)), or Hans Driesch – not to mention Bergson or Canguilhem, moving into the twentieth century – are defending very different and at times incompatible programs (Wolfe, 2015a). Indeed, vitalism itself exhibits a tension or plurality between more metaphysical and less metaphysical views and, contrary to a view still found, does not always maintain the existence in an ontologically specific sense of a vital principle, force, entelechy, élan vital, etc. In that sense, when organicism states “there is legitimate non-reductionist investigation of the core properties of life/organization/organism, and there is illegitimate, metaphysical speculation about or hypostatization of these properties,” it is just replaying the standard exclusionary game like mechanism in earlier times; the organicist who treats vitalism as the absolute cardinal error, in a self-congratulatory act of exclusion, should remember the warning “Don’t throw stones in glass houses” (see also Oyama (2010), and for a different version of this point on how organicism may have difficulty demarcating itself from vitalism, especially the less metaphysical variants of vitalism, see Chen (2019)).

In a standard case of organicism defining itself over and against a “bad version of itself” called vitalism, Cardenas and Cornish-Bowden insist that the recognition of metabolism is tantamount to a decline in vitalism, which they see in the nineteenth century: “Although a living organism is identified and recognized by its physical appearance, and hence by its structure, its status as living is defined by its chemistry, and thus by its metabolism” (Cardenas & Cornish-Bowden, 2011, 1016). This approach to metabolism emphasizes that the processes occurring in a living organism are fundamentally chemical reactions and implicitly echo nineteenth-century affirmations (Bernard, Wöhler) according to which chemistry rules out vitalism.

One need only contrast Jonas (see Jonas, 1966/2011) and Cornish-Bowden and Cardenas on metabolism to see the way organicism can be more or less “ontologized,” more or less subjectivist. That is to say, metabolism and homeostasis are good examples of important physiological properties or processes which allow of a kind of philosophical “overdetermination” or “overinvestment” (as happens with Jonas, when a notion like metabolism cannot just mean what it means to a biologist but has to mean *more*). Notably, metabolism is taken to raise the problem of “subjectification,” i.e., when an organic property becomes construed as requiring a “self” or “subject.” Thus, Jonas presents his work as “a new reading of the biological record [to] recover the inner dimension” and describes metabolism as constituting the organization of organisms for “inwardness, for internal identity, for individuality” while also turning the organism outward “toward the world in a peculiar relatedness of dependence and possibility”; life is “self-centered individuality, being for itself and in contraposition to all the rest of the world, with an essential boundary
dividing ‘inside’ and ‘outside’ –notwithstanding, nay, on the very basis of the actual exchange.”<fnref n="17" />

One imagines that most physiologists from Bernard and Sherrington onward would find it quite strange to have metabolism equated with “inwardness”; doubt- lessly, Jonas is (i) stressing that metabolic entities have an inner/outer relation and (ii) hypostasizing this relation as a philosophical justification of interiority. Organicism is not, then, automatically vaccinated against the excesses of metaphys- ical vitalism (whether it is Jonas on metabolism, Merleau-Ponty on the flesh, writ- ing “Just as the sacrament not only symbolizes … an operation of Grace, but is also the real presence of God … in the same way the sensible has not only a motor and vital significance but is nothing other than a way of being in the world that our body takes over [...] sensation is literally a communion” (Merleau-Ponty, 1945, 245),<fnref n="18" /> or Evan Thompson’s reworking of Merleau-Ponty in an even more explicitly dualist direction: “Life is not physical in the standard materialist sense of purely external structure and function. Life realizes a kind of interiority, the interiority of selfhood and sense-making. We accordingly need an expanded notion of the physical to account for the organism or living being” (Thompson, 2007, 238)<fnref n="19" />), and because its reasonable variants themselves vary on issues like holism (itself just a word, to be sure).

Of course, organicism (or at least some forms of organicism) seems to steer clear of considerations concerning “subjectivity,” “the transcendental,” “embodiment,” and so on, due to its resolute focus on organization (for a different perspective, see Van de Vijver & Haeck, this volume). Granted, the latter notion is not always strictly defined, but even if it is related to “closure,” it does not seem to require a “control- ler,” a “hegemon,” as the notion of constraint indicates. However, it allows of more “abstract” and more “literally real” renditions.<fnref n="20" /> But organicism comes in different strengths, different degrees of ontologization. Some contributors to the history of organicism in the twentieth century, e.g., Jonas, seem to present organisms (which in

<footnote n="17">¹⁷ Jonas 1966/2011, xxiii, 84, 79. For discussion of these passages and more general reflections on metabolism as a concept that go in a direction quite different from that of this paper, see Landecker (2013).</footnote>

<footnote n="18">¹⁸ It falls outside the scope of the present paper to present a critique of the phenomenology of embodiment in terms of its revival of subjectivity, but I note that Canguilhem diagnosed this prob- lem in what was actually a “friendly critique,” in a late lecture on health, commenting on the way this sense of privacy and ineffability in Merleau-Ponty presented a kind of “wall,” an end to discus- sion (Canguilhem (2008b); see Wolfe (2015b) for brief discussion).</footnote>

<footnote n="19">¹⁹ I realize that early twentieth-century organicism (like that of E.S. Russell or J.H. Woodger), mid- twentieth century philosophy of biology with an antireductionist focus (like Jonas and some Merleau-Ponty), and contemporary organicism (like Moreno and Mossio, with Varela lying some- where in between) are not carrying out one and the same monolithic “program” (indeed, this monolithic, somehow uncontextualized view is something I’ve criticized organicism for in the present paper. But I suggest that my criticism of (a) an uncritically asserted empirical-realist appro- priation of the Kantian perspective and (b) an unfounded demarcation between organicism and vitalism in which the former is the legitimate, naturalism-friendly project can apply to some differ- ent moments of the organicist history.</footnote>

<footnote n="20">²⁰ Bich and Damiano (2008) Moreno, and Mossio et al. describe organization as a “specific causal régime” (Mossio & Bich, 2017; Moreno & Mossio, 2014; DiFrisco & Mossio, 2020).</footnote>
this case are never nematodes or slime molds, but typically humans or creatures resembling them) as almost outside the realm studied by natural science. But even the “weak organismic view” will sometimes rest its claims on the existence of certain irreducible, specific, *key features* of organisms (à la Claude Bernard) – what I called the “laundry list” approach above.

Now, this approach remains “biochauvinist,” to use Ezequiel Di Paolo’s suggestive term:<fnref n="21" /> it is essentially a naturalized form of (substance) vitalism, that is, claims about the “uniqueness” of biological entities. Indeed, besides seeking to define itself in opposition to the purportedly metaphysical version, vitalism, organicism can also subdivide into warring *groupuscules*: to cite one example among many, Needham distinguishes between “dogmatic” and “legitimate” organicism (regarding the concept of organization).<fnref n="22" /> One could also ask whether all forms of organicism are naturalist; how naturalist can one be here?<fnref n="23" /> This is a delicate matter, because most or all contemporary organicists either accept naturalism in a broad sense or at least would be wary of endorsing some antinaturalist position. But at the same time, when they invoke, inter alia, Merleau-Ponty or Jonas, they are explicitly using conceptual accounts of, e.g., life and embodiment, which insist that “life,” “mind,” “selfhood,” “subjectivity,” and “interiority” are somehow not in physical space, outside of physical space, and require almost a separate ontology: on most accounts, that is antinaturalism. If naturalism is a desideratum – if it is desirable for the position argued for in the context of discourses on biology and organisms to be naturalistic or at least broadly compatible with naturalism – then organicism has two different ways of achieving it: it can opt for the epistemological view, as Goldstein does, stressing that organisms are “knowers” who construct their worlds, or it can opt for naturalizing, e.g., declaring teleology to be integrated in a naturalistic framework (e.g., Mossio & Bich, 2017).

## 3.4 Ontology and Ontologization

One organicist response to my raising the issue of ontologization could be that in fact, they have moved on, and the issue is really *explanations*: that is, what is unique about biological entities is the type of *explanations* they require (an idea that might remind us of the epistemological position). So, for instance, Arno Wouters writes

<footnote n="21">²¹ Di Paolo (2009). Di Paolo does not actually define or expand on the term, but it is close, e.g., to Andy Clark’s discussion of “pressing the flesh” (Clark, 2008) in terms of distinguishing different “strengths” of embodiment as a perspective on cognition (embodied cognition), what I would term different degrees of ontologization of the biological, the living body, the flesh, etc.</footnote>

<footnote n="22">²² Needham (1936), 18, cit. in Nicholson and Gawne (2015). In the earlier *The Great Amphibium* (1932), Needham had spoken of the organicists who accept the notion of organization without seeking to investigate it scientifically as “passive organicists” (thanks to Alejandro Fábregas Tejeda for this reference).</footnote>

<footnote n="23">²³ On the idea of “naturalism friendly,” see notably the discussions concerning “naturalized teleology” (e.g., Walsh (2007, 2013). Keijzer (2016) comments briefly, with regard to Moreno and Mossio’s concept of autonomy, on the pros and cons of “naturalizing.”</footnote>
that “Functional explanations are, nevertheless, crucial to understand life, because they show us how the characteristics of an organism fit into the requirements for being alive.”<fnref n="24" /> Of course, such specific explanations may in turn be “generated” or “induced” by features that are claimed to be ontologically real (explanatory autonomy is not an argument-stopper). Indeed Wouters does not seem content to dwell at the level of explanations, as the dimension of organization he emphasizes is ontological: “Organization can be defined in terms of dependence on the composition, arrangement and timing of the relevant system, its parts and their activities.”<fnref n="25" /> When Dan Nicholson writes that “Organicists, in contrast, are committed to the belief that the integrated and inherently purposive nature of organisms reflects the very essence of what they actually are in reality,”<fnref n="26" /> I think this is right about a number of forms of organicism (and this leads to some of the problems surrounding “ontologization” I have discussed above), but it does not apply, for instance, to the organicism of Kurt Goldstein (together with another important figure who I have not discussed here, Georges Canguilhem<fnref n="27" />), because the latter form of organicism seeks to integrate – or better, it *builds on* – the irreducible existence of a feature of organisms one might call existential or perspectival: the fact that an organism always has an existential attitude, a perspective on other organisms, and that, thereby, part of its vital activity involves acts of (cognitive) construction.

It is as if the Kantian constructive concept of organisms had been ontologized without being subsumed under a potentially refutable or reducible list of vital features. Because, as Canguilhem argues powerfully in his essay “Aspects of vitalism,” vitalism (and here we could replace this term with “organicism,” *if this term is not construed in the strictly empirical, ontologized, criterial, “laundry list” sense*) is not refutable like geocentrism or the phlogiston theory are. It is a different kind of theory. If organicism were like geocentrism or the phlogiston theory, it would be because it was a strictly empirical (and thus refutable) set of claims. But, on the constructivist view, according to which organisms both *require* acts of cognitive construction and *perform* such acts, organicism is not reducible to an empirical set of claims (a.k.a. definitions of life). Goldstein and Canguilhem do not give a set of empirical features of organisms (purposiveness/teleology, integration, reflexivity, etc.) and then state “Here are organisms and here is why organicism is the theory that does justice to them.” They argue less in empirical, “realist” terms and more in existential, constructivist terms.

Now, reflecting on this existential or perspectival dimension (which will be familiar to students of Uexküll, of course) also helps us with our rough typology of organicisms, since it leads us to nuance, not so much the question of ontologization and weak or strong organicism, but at the other end of the spectrum, the question of

<footnote n="24">²⁴Wouters (2013a). Compare Walsh (2015).</footnote>

<footnote n="25">²⁵Wouters (2013b), 463, 460; see also Wouters 2013c.</footnote>

<footnote n="26">²⁶Nicholson (2010), Chap. 2, 63n.</footnote>

<footnote n="27">²⁷For an excellent discussion of the connection between the two thinkers in this context, see Gayon (1998); I have developed this point a bit further in Wolfe (2015b, 2017).</footnote>
the “subject.” That is to say, if earlier I tried to diagnose a problem of “ontologiza-
tion,” in that sense, *objectification* (of properties of organisms), there is a kind of
symmetrical problem, the other horn of the dilemma, which is *subjectification*. By
this I mean the kind of “romantic” vision of organism as deeply opposed to machines,
or inert matter, because it is a version of subjectivity, because it has a “self” (this is
explicit in Hegel’s philosophy of nature, Schelling’s and I think Jonas’s – as
defended again in Michelini et al. (2018); it is – debatably and controversially – also
present in the later Varela (Varela (1991), Varela (1996), Varela & Shear, (1999)),
and definitely in Evan Thompson). Arguably, the organizational perspective on
organisms allows the organicist to escape the danger of subjectification, but so does
the Goldstein-Canguilhem approach, i.e., the “constructivist” vision of organism.

3.5 Conclusion

To sum up, my point is not to assert or deny the reality or pertinence of organisms
or organismic theories in biology. I notice that no one seems to bother to do this
anymore: Bickle (2003) argues aggressively for reductionism (of cognitive neuro-
science to cellular-level neuroscience), but otherwise it has become more than
rare – irrelevant – to argue “reductionism” against “organicism” (contrast Nagel in
the 1960s). Present-day neomechanists need not worry, if we consider Bechtel’s
own “organizational” commitments. For Bechtel, mechanism and organization
stand in a complex and fruitful interaction, and mechanism can provide an adequate
account of organization by “placing as much emphasis on understanding the par-
ticular ways in which biological mechanisms are organized as it has on discovering
the component parts of the mechanisms and their operations”; “It is the fact that the
system is organized (and the type of organization it has) that makes it amenable to
mechanistic description and analysis” (Bechtel 2007, 270; Levy & Bechtel 2013,
244). My point is not to defend “neomechanism” in extenso (it has received a num-
ber of quite different, and significant criticisms) but to observe that one of its most
useful features in the present context is to not just oppose mechanistic and organi-
zational approaches. As Moreno puts it, “holism and mechanistic decomposition
can be combined for the purposes of biological explanation,” increased complexity
creates “selective functional constraints,” giving rise to “levels of organization in
which a mechanistic decompositional strategy might be locally applicable.”<fnref n="28" />

Similarly, my point is not to simply put forth a “safe,” reductionist critique of all
or any such programs. On the contrary, I view it as extremely useful to track such
articulations, whether or not we fully commit to them (in this I feel kinship with the
“epistemology of Life” in Canguilhem; see Canguilhem (2008a) and Wolfe (2017)).
To clarify one potential ambiguity, my point here is deflationary (about organicism,

<footnote n="28">²⁸ Moreno (2013, 901); for critical discussion of Bechtel’s way of articulating mechanism and organization, see Eronen, 2015 and Mossio (2020, 73).</footnote>
i.e., not reductionist in the sense of being committed to a more minimal ontology, but deflationary inasmuch as I challenge some higher-level concepts in favor of stripped-down versions thereof) but does not take a stand on organisms (their ontological status); I've taken several different stances on organism in earlier papers, e.g., in my 2010 "Do organisms have an ontological status?" arguing for a weakly ontological status – indeed rather consonant with what I'm saying here about organicism – and in my 2014 "The organism as ontological go-between" emphasizing rather the "nomadic concept" aspect, the hybridity in its fertility (Wolfe, 2010, 2014a). Organisms as conceptual hybrids are not so directly opposed to mechanism(s) as in the usual organicist party line. They are also so not ontologically solid or essentialist (that's what the word hybrid indicates).

I have tried to suggest (a) that organicism is not necessarily so far removed from the more naturalistic forms of vitalism (and this, to be clear, is not a bad thing); but (b) that when organicism relaunches itself as a fully respectable scientific theory – no one’s mistress, it might say to Haldane – it creates for itself several potential conceptual problems. One is the unstable combinations of the Kantian epistemological perspective with deliberately empirical, ontologized claims – as if organicists wanted to play both sides in an unprincipled way (sometimes claiming features must really be part of organisms, other times claiming they are only regulative ideals).<fnref n="29" /> The other is, in a sense, another kind of instability resulting from the reduction of organicism to a “merely empirical,” criterial set of definitions (the “laundry list”). This instability is, ironically, the more Kantian component of the present remarks: faced with the confidence of thinkers asserting the “the soul and the world exist,” Kant suggested an approach that would not be founded on directly refutable empirical claims. Founding claims for the irreducible nature of organisms on directly refutable empirical definitions create this instability.<fnref n="30" /> Not all contemporary organicists make this Kantian move, or simply the (ontologized) empirical criteria move; I noted in addition (c) that symmetrically to this danger of the excess, the addiction to ontologization, there is the potential to reintroduce into organicism a kind of Romantic love affair with subjectivity, ineffability (as we saw with Merleau-Ponty and Canguilhem’s sensitive commentary) and the first-person perspective. It would be a good idea to try and achieve some conceptual clarity so that the observer can have an idea of what is involved in the current smorgasbord of mildly or strongly organismic projects.

**Acknowledgments** Earlier versions of this paper were presented in Vienna and Paris. Thanks to Sebastjan Vörös and Matteo Mossio for their invitations and comments and to Arantza Etxeberria, Alejandro Fábregas Tejeda, Phillip Honenberger, and Andrea Gambarotto for further suggestions.

<footnote n="29">²⁹ I thank Phillip Honenberger for this formulation.</footnote>

<footnote n="30">³⁰ While this is not a reflection on the “definitions of life” debates (e.g., Machinery, 2012 versus Bich & Green, 2017), one can extrapolate from my position here: the idea would then be not necessarily to side with Machinery’s deflationary position, but to acknowledge the Sisyphean (or Pyrrhic?) character of attempts to define life.</footnote>
References

Baedke, J. (2019). O organism, where art thou? Old and new challenges for organism-centered biology. *Journal of History of Biology, 52*, 293–324.

Bechtel, W. (2007). Biological mechanisms: Organized to maintain autonomy. In F. Boogerd, F. J. Bruggeman, J.-H. S. Hofmeyr, & H. V. Westerhoff (Eds.), *Systems biology: Philosophical foundations* (pp. 269–302). Elsevier.

Bechtel, W., & Richardson, R. C. (1993). *Discovering complexity: Decomposition and localization as strategies in scientific research*. Princeton University Press.

Beckner, M. (1974). Reduction, hierarchies and organicism. In F. Ayala & T. Dobzhansky (Eds.), *Studies in the philosophy of biology: Reduction and related problems* (pp. 163–177). University of California Press.

Bich, L., & Damiano, L. (2008). Order in the nothing: Autopoiesis and the organizational characterization of the living. In I. Licata & A. J. Sakaji (Eds.), *Physics of emergence and organization* (pp. 343–373). World Scientific.

Bich, L., & Green, S. (2017). Is defining life pointless? Operational definitions at the frontiers of biology. *Synthese, 195*, 3919. https://link.springer.com/article/10.1007/s11229-017-1397-9

Bickle, J. (2003). *Philosophy and neuroscience: A ruthlessly reductive account*. Kluwer.

Bouchard, F., & Huneman, P. (Eds.). (2013). *From groups to individuals: Evolution and emerging individuality*. MIT Press, Vienna Series in Theoretical Biology.

Canguilhem, G. (2008a). *Knowledge of life* (S. Geroulanos, & D. Ginsburg, Trans.). Fordham University Press (A translation of *La connaissance de la vie*, 1965).

Canguilhem, G. (2008b). “Health: Crude concept and philosophical question” (translation of “La santé, concept vulgaire et question philosophique” (1988), by T. Meyers and S. Geroulanos). *Public Culture*, 20(3), 467–477.

Cardenas, M., & Cornish-Bowden, A. (2011). Metabolism (Biological). In M. Gargaud et al. (Eds.), *Encyclopedia of astrobiology* (pp. 1016–1017). Springer.

Chen, B. (2019). *A historico-logical study of vitalism: Life and matter*. PhD thesis, Department of Philosophy and Moral Sciences, Ghent University.

Clark, A. (2008). Pressing the flesh: A tension in the study of the embodied embedded mind? *Philosophy and Phenomenological Research*, 76(1), 37–59.

Clarke, E. (2013). The multiple realizability of biological individuals. *The Journal of Philosophy*, 110(8), 413–435.

Dennett, D. (1984). *Elbow room. The varieties of free will worth wanting*. MIT Press.

Di Paolo, E. (2009). Extended life. *Topoi*, 28, 9–21.

Di Paolo, E. A. (2018). The enactive conception of life. In A. Newen, S. Gallagher, & L. de Bruin (Eds.), *The Oxford handbook of 4E cognition: Embodied, embedded, enactive and extended* (pp. 71–94). Oxford University Press.

DiFrisco, J., & Mossio, M. (2020). Diachronic identity in complex life cycles: An organizational perspective. In A. S. Meincke & J. Dupré (Eds.), *Biological identity. Perspectives from meta-physics and the philosophy of biology* (pp. 177–199). Routledge.

Eronen, M. I. (2015). Levels of organization: A deflationary account. *Biology and Philosophy*, 30, 39–58.

Gambarotto, A., & Illetterati, L. (2020). Hegel’s philosophy of biology? A programmatic overview. *Hegel Bulletin*, 41(3), 349–370.

Gayon, J. (1998). The concept of individuality in Canguilhem’s philosophy of biology. *Journal of the History of Biology*, 31(3), 305–325.

Gilbert, S. F., & Sarkar, S. (2000). Embracing complexity: Organicism for the twenty-first century. *Developmental Dynamics*, 219, 1–9.

Goldstein, K. (1939/1995). *The Organism: A holistic approach to biology derived from pathological data in man*. American Book Company/Zone Books. (A translation of *Der Aufbau des Organismus*, 1934).

Grene, M. (1966). *The Knower and the Known*. London: Faber & Faber.