REVIEWS
"Lidgard and Nyhart have compiled a superb collection of essays concerning recent work on the concept and nature of biological individuals. . . . The overarching question ‘What is a biological individual?’ is demonstrated to have an intricate history, changing and evolving as biologists learned more about organismic structures and functions, life cycles, cellular systems, and ecological interactions. Notions of what counts as an individual are shown to have carried over into social and political identities and practices. In addition, philosophical, conceptual underpinnings and commitments are shown to interplay with these changing biological understandings. . . . The volume is marvelous! While not for the casual reader, it is exemplary of the best current work in biological philosophy. . . . Highly recommended."
— D. B. Boersema, Pacific University, Choice
“I found this collection a very enjoyable read and recommend it as a starting point to anyone interested in the notion of biological individuality broadly construed and interested in understanding what unifies and separates the different meanings of this term across time and disciplines.”
— Pierrick Bourrat, Macquarie University, Notre Dame Philosophical Reviews
“Individuality is one of the hottest topics in the philosophy of biology today. The recent, technologically driven explosion of knowledge about the microbial world has led both biologists and philosophers to reconsider the traditional idea that organisms are genetically uniform populations of cells with a clear physical boundary—bodies. Very up-to-date, containing a number of substantial original contributions, and with a good balance of historians, philosophers, and eminent scientists, Biological Individuality is well positioned to represent the most exciting strands of the current debate.”
— Paul Griffiths, University of Sydney, coauthor of "Genetics and Philosophy: An Introduction"
“A tour de force inquiry of individuality that proposes a fruitful perspective that potentially resolves many of its disparate definitions. Biological Individuality reflects the strengths of cross-disciplinary research on fundamental biological concepts.”
— Paul Lawrence Farber, Oregon State University, emeritus, author of "Finding Order in Nature: The Naturalist Tradition from Linnaeus to E. O. Wilson"
"...how has the discussion and the concept of biological individuality evolved over the past 25 years or so? Scott Lidgard and Lynn Nyhart’s volume aims to answer that question and succeeds admirably...This book pushes one hard to reexamine foundational ideas about what biology studies as its basic subject matter."
— James Strick, Journal of the History of Biology
TABLE OF CONTENTS
Introduction: Working Together on Individuality
DOI: 10.7208/chicago/9780226446592.003.0000
[biological individuality;interdisciplinarity;part-whole relations]
In this essay, we characterize the interdisciplinary work of this volume. We first sketch the different goals and assumptions of scientists, historians, and philosophers with respect to biological individuality, noting where their interests cross, where they diverge, and how different disciplinary perspectives might provide resources for each other. We then summarize the main themes of the different chapters, individually and collectively. Here, the discussion pays special attention to hierarchy and evolutionary transitions in individuality, to historical approaches to individuality that extended across multiple temporal scales and often took non-Darwinian evolutionary perspectives, and to the dynamic relations among parts and between parts and wholes that often call into question the very notion of individuality. (pages 1 - 16)
This chapter is available at:
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1. The Work of Biological Individuality: Concepts and Contexts
DOI: 10.7208/chicago/9780226446592.003.0001
[individuation;levels of individuality;biological hierarchy;biological systems;organism;symbiosis;holobiont;epistemology;problem space;pluralism]
Why are there so many concepts of biological individuality? This chapter shows that a wide range of definitional criteria of individuality has been integral to biological studies for at least 170 years. Using the notion of a dynamic problem space to explore biological, historical, and philosophical aspects of biological individuality, we argue that the work of biological individuality concepts must be understood contextually. These concepts are contextualized by specific research problems, biological systems, and epistemic goals; by different disciplinary norms; and by change over time. Concepts also address different categories of problems involving individuation, hierarchies or levels, temporal continuity or change, and what does or does not constitute an individual. To move our understanding of biological individuality forward productively, we recast it as a broad, stable, and growing domain of problems that each merit investigation in its own right, for which no single definition of individuality will suffice. (pages 17 - 62)
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2. Cells, Colonies, and Clones: Individuality in the Volvocine Algae
DOI: 10.7208/chicago/9780226446592.003.0002
[progress;complexity;individuality;major transitions;Volvox;Chlamydomonas;Pleodorina]
The hierarchy of life results from a series of evolutionary transitions in individuality. In each case new, higher-level units emerged from coalitions of the lower-level units, resulting in entirely new kinds of individuals with new, emergent properties. A common feature of such transitions is the division of labor between reproductive and somatic functions, as in the castes of eusocial animals and differentiated cells in multicellular organisms. One of the best-studied transitions in individuality is the origin of differentiated multicellular organisms from unicellular ancestors in the volvocine green algae (Volvox and its close relatives). This chapter uses the volvocine algae as a case study to consider how evolutionary changes in development led to the emergence of a new kind of individual. Cells, colonies, and clones all have a legitimate claim to some degree of individuality, and those degrees vary across different concepts of individuality and different criteria relevant to specific concepts. (pages 63 - 83)
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3. Individuality and the Control of Life Cycles
DOI: 10.7208/chicago/9780226446592.003.0003
[mereology;cell differentiation;multicellularity;major transitions;scaffolding;material overlap;multilevel selection]
Evolutionary transitions in individuality occur when units of evolution at one level, such as single cells, associate together and evolve into higher-level units. I argue that multi-level selection theory, the leading method for explaining how and why evolutionary transitions happen, is not sufficient in itself. In particular, knowledge of the material and causal structure of the particular case is crucial for specifying the appropriate units and parameters for multi-level models and for explaining the evolvability of the higher-level unit after the transition. I develop a complementary approach based on the concept of a demarcator—a material entity or causal process responsible for determining what is and is not a part of the individual. I also show how demarcators provide a potentially level-neutral account of inheritance based on the concepts of material overlap and scaffolding and give examples from the inheritance of differentiated cell types (Griesemer 2000a; Wimsatt and Griesemer 2007). (pages 84 - 108)
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4. Discovering the Ties That Bind: Cell-Cell Communication and the Development of Cell Sociology
DOI: 10.7208/chicago/9780226446592.003.0004
[cell-cell communication;community effect;multicellularity;metaphors in science;cell sociology]
Social metaphors have long been a part of the cell theory. In the nineteenth century plants and animals were described as ‘cell-states’ (ZellenStaaten) composed of cell ‘citizens’ functionally arranged into specialized tissue ‘classes’ or ‘professions.’ This conception of the organism as a ‘society of cells’ was conceptually grounded by the principle of the division of (physiological) labor, an idea transferred from political economy to help explain the functional organization of a coherent system of semi-autonomous parts. More recently however cell biology has been reshaped by the idea of inter-cellular communication. This paper details how the science of cell communication provides a new explanatory account of how multicellular organisms and other ‘cell societies’ with their division of labor are created and maintained through the bonds of communication. The idea of ‘cell sociology’ is discussed as a particular reflection of this new communicative model of multicellular organization and function. (pages 109 - 128)
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5. Alternation of Generations and Individuality, 1851
DOI: 10.7208/chicago/9780226446592.003.0005
[alternation of generations;levels of individuality;biological hierarchy;life cycles;Johannes Japetus Steenstrup;Thomas Henry Huxley;Richard Owen;Wilhelm Hofmeister]
Alternation of generations historically referred to the phenomenon in which an organism asexually produced offspring that did not resemble itself; these offspring then sexually produced a new generation resembling the grandparental form. This presented problems for understanding individuality, with dissenting views passing through an intellectual bottleneck ca. 1851. Organisms as life cycles, transformation of bodies, different reproductive modes, and causal forces of development gripped biologists across Europe, engaging prominent theorists on individuality, including zoologists J. J. Steenstrup, Richard Owen, T. H. Huxley, and Johannes Müller, and botanists Matthias Schleiden, Alexander Braun, and Wilhelm Hofmeister. While these scientists reached no interpretive consensus, we show that they nonetheless generally shared a broad commitment to a temporal perspective, an overall development toward "perfection" (or completeness of variation through a life cycle) over time, and a nested hierarchy of individuality. Alternation of generations thus helped define the enduring problematic of biological individuality. (pages 129 - 157)
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6. Spencer’s Evolutionary Entanglement: From Liminal Individuals to Implicit Collectivities
DOI: 10.7208/chicago/9780226446592.003.0006
[Herbert Spencer;organism;environment;hierarchy;complexity;individual;collectivity]
Herbert Spencer’s notions of biological individuality as expounded from the 1850s until the late 1860s are presented. I argue that: 1) Spencer developed a basic epistemological and methodological tripartite frame of analysis of living nature, whereby the elements of that frame—"organism", “environment", and their evolving interactions and their evolved effects—are not separable. An organism can therefore never be viewed as a passive receiver of changes; 2) living nature is hierarchically ordered, and the main feature of Spencerian evolution is the growing complexity of interrelations 3) the Spencerian biological individual is the product of evolutionary work; 4) the same applies to Spencerian collectivities. Thus, Spencer’s biological individuals harbor within them plurality. Furthermore, hybrid categories of collective individuals and of collectivities composed of differing individuals are conceptually necessary for Spencer’s arguments. In a concluding section the possible contemporary interest in and relevance of the Spencerian problématique is addressed briefly. (pages 158 - 183)
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7. Biological Individuality and Enkapsis: From Martin Heidenhain’s Synthesiology to the Völkisch National Community
DOI: 10.7208/chicago/9780226446592.003.0007
[Martin Heidenhain;enkapsis;hierarchy;histology;ecology;phylogeny;biological individuality]
Enkapsis shows biological individuality to be hierarchically structured, and to be grounded not in intrinsic essential properties, but in relational properties, most notably the property of descent, either ontogenetically, or phylogenetically, from divisible parts. During the first half of the 20th century the enkaptic hierarchy developed by the Tübigen anatomist Martin Heidenhain became the central concept capturing biological individuality from a holistic-organicist perspective in German biology. The paradigmatic example of an enkaptic hierarchy was the organism with the cells as its most fundamental divisible subunits. The cells combine to form organs, these to form organ systems, ultimately the organism itself which thus represents a nested hierarchy of complex wholes with emergent properties manifest at different levels of complexity. The concept of the enkaptic hierarchy was soon extrapolated to ecosystems, to the phylogenetic system, ultimately to the socio-political conception of the National Community of National Socialist Germany. (pages 184 - 205)
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8. Parasitology, Zoology, and Society in France, ca. 1880–1920
DOI: 10.7208/chicago/9780226446592.003.0008
[Casimir Davaine;Charles Gide;Edmond Perrier;organicism;parasitism;Raphaël Blanchard;Solidarism;symbiosis;France]
The phenomena of parasitism, mutualism, commensalism, and symbiosis complicate our ideas of parts and wholes. Focusing primarily on the French Third Republic (1879-1940), this chapter examines the terms parasite and parasitism, their definitions, and broader connections to part-whole relations and dependence across three overlapping realms: biological theorizing about parts and wholes, medical parasitology, and social and political theory. The term parasite, which in antiquity connoted common dining activity free of harm to the participating organisms, acquired new valences in this period. Ultimately in biology, the term has come to imply two things: that parasites cause harm to their hosts and that parasitism is largely, if not exclusively, an interspecific relationship. But the appropriation of biological concepts by promoters of Solidarism, a theory of social cohesion infused with organicism, was selective as regards parasitism where it was applied to human-to-human relationships and thus violated the interspecific consensus established by biologists. (pages 206 - 224)
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9. Metabolism, Autonomy, and Individuality
DOI: 10.7208/chicago/9780226446592.003.0009
[metabolism;autonomy;individuality;nutrition;Claude Bernard;Justus von Liebig;homeostasis;microbiome;epigenetics]
This chapter discusses the interdependence of metabolism, autonomy and individuality. The work of physiologist Claude Bernard illustrates the logic of total conversion that emerged in nineteenth century physiology. This logic posited that consumed organisms were completely broken down and transformed into nutritive reserves of the consumer. This conversion was the basis of autonomy in a fluctuating environment. Similarly, in twentieth century intermediary metabolism individuation—the separation of an individual from the surrounding world—was theorized as due to metabolic conversion of the world into the self. Throughout, this framework depended on a near-total loss of agency on the part of food vis-à-vis the eating organism. The chapter ends with examples from contemporary science in which food maintains various degrees of agency in relation to the eater. These examples break with and simultaneously highlight often-implicit assumptions about conversion and individuality inherited from the science and theory of metabolism. (pages 225 - 248)
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10. Bodily Parts in the Structure-Function Dialectic
DOI: 10.7208/chicago/9780226446592.003.0010
[structure;function;activities;part-whole relations;hierarchy;individuation;biological systems;homology;evolutionary novelty]
Understanding the organization of an organism by individuating meaningful parts and accounting for organismal properties by studying the interaction of bodily parts is a central practice in many areas of biology. While structures are obvious bodily parts and structure and function have often been seen as antagonistic principles in the study of organismal organization, my tenet is that structures and functions are on a par. I articulate a notion of function (functions as activities), according to which functions are bodily parts just as structures are. Recognizing part-whole relations among an organism’s various structures and functions permits fruitful investigation and multilevel explanation of organismal properties and functioning, across both developmental and evolutionary time. I show how my perspective clarifies debates surrounding homology and evolutionary novelty that stem from an alleged structure-function dichotomy. (pages 249 - 276)
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11. Distrust That Particular Intuition: Resilient Essentialisms and Empirical Challenges in the History of Biological Individuality
DOI: 10.7208/chicago/9780226446592.003.0011
[biological individuality;essentialism;Darwin;intuitions;parasitism;chimerism;levels of individuality;holobiont]
There seem to be three intuitions about biological individuality that are held in western culture. This paper calls these intuitions about essences anatomical essentialism, physiological essentialism, and developmental essentialism. These intuitions reinforce one another. They work to a certain extent, and any explanation using one of the essentialisms tends to be accepted by an audience more easily. Yet when held unreflectively such intuitions lead to an impoverished understanding of individuality. These intuitions are highly resilient—histories of biology show how they have been challenged both analytically and empirically, yet they still persist to this day. I suggest this is because they both depend on and reinforce notions of purity, stability, indivisibility and autonomy. (pages 277 - 296)
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12. Biological Individuality: A Relational Reading
DOI: 10.7208/chicago/9780226446592.003.0012
[symbiosis;holobiont;part-whole;co-development;individual]
Biology is a study of relationships between parts and wholes. An individual at one level is a part at another. In the late nineteenth century, embryologist E. B. Wilson wrote, “There is at present no biological question of greater moment than the means by which the individual cell-activities are co-ordinated, and the organic unity of the body maintained; for upon this question hangs… our conception of life itself.” This question of individuality gains prominence in the twenty-first century, as processes that unite parts into new wholes are becoming accessible to study. This essay reviews a newly discerned level of individuality, the holobiont, which may be a critical unit of anatomy, physiology, development, and evolution. This supraorganismal individual constitutes the eukaryotic organism plus its persistent symbionts. The essay then reflects on some of the “parts” of this whole volume. Each reader gets the privilege of reconstituting them into a new whole. (pages 297 - 317)
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13. Philosophical Dimensions of Individuality
DOI: 10.7208/chicago/9780226446592.003.0013
[individuality;individuation;monism;parts;pluralism;wholes]
While individuality has been a perennial concern because of biological examples that are difficult to categorize, contemporary discussions among philosophers of biology are dominated by individuality’s crucial role for understanding aspects of evolution. These debates include whether species are individuals or kinds, what counts as a unit of selection, and how transitions in individuality occur evolutionarily. Philosophical analyses are often conducted in terms of metaphysics (“what is an individual?”) rather than epistemology (“how can and do researchers conceptualize individuals so as to address scientific goals?”). We review conceptual distinctions in light of contributions to the volume, highlighting the way metaphysical assumptions have animated controversies, and explicitly shift attention from metaphysics to epistemology. We argue that many controversies involve epistemological differences (e.g., a diversity of legitimate scientific goals) rather than metaphysical disagreement, which implies that a pluralist stance about individuality in biology is warranted. (pages 318 - 348)
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Acknowledgments
List of Contributors
Index