The multiple realizability of biological individuals

Journal of Philosophy

Volumn 110, Issue 8, 2013, Pages 413-435

  Clarke, Ellen ^a  

^a ALL SOULS COLLEGE   (United Kingdom)

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EID: 84890071592     PISSN: 0022362X     EISSN: 19398549     Source Type: Journal    
DOI: 10.5840/jphil2013110817     Document Type: Review

References (105)

1. Leo W. Buss, The Evolution of Individuality (Princeton: University Press, 1987);
2. John Maynard Smith and Eörs Szathmáry, The Major Transitions in Evolution (New York: Freeman, 1995), pp. 1-12;
3. Richard E. Michod, Darwinian Dynamics: Evolutionary Transitions in Fitness and Individuality (Princeton: University Press, 1999), pp. 81-107;
4. Samir Okasha, Evolution and the Levels of Selection (New York: Oxford, 2006), pp. 218-41.
5. Ellen Clarke, "Plant Individuality: A Solution to the Demographer's Dilemma," Biology and Philosophy, xxvii, 3 (May 2012): 321-61, at p. 332.
6. Elisabeth Lloyd, "Units and Levels of Selection," The Stanford Encyclopedia of Philosophy (Winter 2012 Edition), Edward N. Zalta, ed., URL: 〈http://plato.stanford.edu/archives/win2012/entries/selection-units/〉;
7. H. Kern Reeve and Laurent Keller, "Levels of Selection: Burying the Units-of-Selection Debate and Unearthing the Crucial New Issues," in Keller, ed., Levels of Selection in Evolution (Princeton: University Press, 1999), pp. 3-14.
8. David Sloan Wilson, "Altruism and Organism: Disentangling the Themes of Multilevel Selection Theory," The American Naturalist, cl, S1 ( July 1997): S122-34.
9. The problem I treat here is one of finding necessary and sufficient conditions for membership of the class, rather than of finding identity conditions for particular organisms. In other words, my concern is not whether my definition will serve to re-identify one biological individual over space and/or time. For a treatment of this sort, see Thomas Pradeu, "What Is an Organism? An Immunological Answer," History and Philosophy of the Life Sciences, xxxii, 2-3 (2010): 247-68;
10. Charles T. Wolfe, "Do Organisms Have an Ontological Status?," History and Philosophy of the Life Sciences, xxxii, 2-3 (2010): 195-232.
11. A detailed investigation of the different concepts that biologists use goes beyond the scope of this paper, so I will do just enough here to show that there is a genuine multiplicity of candidate solutions on the table, and that there is no simple way of conjoining or otherwise assimilating them. See Clarke, "The Problem of Biological Individuality," Biological Theory, v, 4 (2010): 312-25, for a full list and references.
12. Mitosis occurs when a cell divides to produce two identical copies.
13. Thomas H. Huxley, "Upon Animal Individuality," Proceedings of the Royal Institution of Great Britain, xi (1852): 184-89;
14. Alexander Braun, "The Vegetable Individual, in Its Relation to Species," trans. C. F. Stone, Annals and Magazine of Natural History, published in three parts: xvi (1855): 233-56, xvi (1855) 333-54, and xviii (1857): 363-86;
15. Daniel H. Janzen, "What Are Dandelions and Aphids?," The American Naturalist, cxi, 979 (May-June 1977): 586-89;
16. Robert E. Cook, "Asexual Reproduction: A Further Consideration," The American Naturalist, cxiii, 5 (May 1979): 769-72.
17. John Bonner, On Development (Cambridge: Harvard, 1974); Richard Dawkins, The Extended Phenotype: The Long Reach of the Gene (New York: Oxford, 1982), pp. 254-65;
18. Maynard Smith and Szathmáry, op. cit., p. 244;
19. Peter Godfrey-Smith, Darwinian Populations and Natural Selection (New York: Oxford, 2009), pp. 91-108.
20. Janzen, op. cit., p. 588.
21. The two views also diverge with respect to monozygotic twins. A sexual (and genetic) view treats them as two halves of a whole, while a bottleneck view can separate them, on the grounds that the zygote splits into two sufficiently small pieces. See Julian S. Huxley, The Individual in the Animal Kingdom (Cambridge, UK: University Press, 1912), p. 70.
22. We now know this assumption to be false in the aphid case. See Hugh Loxdale, "Was Dan Janzen (1977) Right about Aphid Clones Being a 'Super-organism', i.e. a Single 'Evolutionary Individual'?," Mitteilungen der Deutschen Gesellschaft für Allgemeine und Angewandte Entomologie, xvi (2008): 437-49.
23. In the terms of Price's equation, between-insect selection will increase the amount of transmission bias at the clone level-assuming that the two levels of selection are antagonistic, that is, that the two components of the covariance term in the multilevel Price equation are opposite in sign. Without taking this into account, our clone-counting scientist will end up overestimating the amount of evolutionary change that is going to take place. See George R. Price, "Selection and Covariance," Nature, ccxxvii, 5257 (Aug. 1, 1970): 520-21;
24. Okasha, op. cit., p. 70.
25. See Clarke, "Plant Individuality," p. 354
26. for additional ways in which organism counts can go wrong. For a recent explanation of the way in which selection at one level can bring about transmission bias or lowered heritability at a higher level see S. A. Frank, "Natural Selection. III. Selection versus Transmission and the Levels of Selection," Journal of Evolutionary Biology, xxv, 2 (February 2012): 227-43.
27. Dawkins, op. cit., p. 260
28. Garrett Hardin, "The Tragedy of the Commons," Science, clxii, 859 (Dec. 13, 1968): 1243-48.
29. When I say "lower-level selection" I always refer to selection amongst the lowerlevel entities within partitions. This is a terminological point made clear by Elliott Sober, "Realism, Conventionalism, and Causal Decomposition in Units of Selection: Reflections on Samir Okasha's Evolution and the Levels of Selection," Philosophy and Phenomenological Research, lxxxii, 1 ( January 2011): 221-31.
30. Dawkins, op. cit., p. 260.
31. Maynard Smith and Szathmáry, op. cit., p. 244.
32. Godfrey-Smith, op. cit., p. 83.
33. These views originate in the work of August Weismann, "The Continuity of the Germ-Plasm as the Foundation of a Theory of Heredity," in Essays upon Heredity and Kindred Biological Problems (Oxford: Clarendon, 1891), pp. 161-254
34. A germ-soma view is implicit in Ratcliff et al.'s recent claim to have produced a new multicellular organism in the laboratory. See William C. Ratcliff et al., "Experimental Evolution of Multicellularity," Proceedings of the National Academy of Sciences, cix, 5 ( Jan. 31, 2012): 1595-600.
35. David L. Kirk, "Evolution of Multicellularity in the Volvocine Algae," Current Opinion in Plant Biology, ii, 6 (December 1999): 496-501.
36. Buss, op. cit., especially pp. 6-9.
37. This is because a cell's fate (that is, whether it will differentiate into soma or not) depends on relational properties such as its position within the developing embryo, rather than on any of its own characteristics. Godfrey-Smith, op. cit., pp. 102-03
38. In Michod's view it is more natural to say that germ-soma separation eliminates withinorganism selection by preventing genetic variation from giving rise to fitness differences, because somatic cells do not really have fitness at all. See Michod and Aurora Nedelcu, "On the Reorganization of Fitness During Evolutionary Transitions in Individuality," Integrative and Comparative Biology, xliii, 1 (February 2003): 64-73.
39. Maynard Smith and Szathmáry, op. cit., p. 244.
40. Francis L. W. Ratnieks, "Reproductive Harmony via Mutual Policing by Workers in Eusocial Hymenoptera," The American Naturalist, cxxxii, 2 (August 1988): 217-36;
41. Robert Boyd and Peter J. Richerson, "Punishment Allows the Evolution of Cooperation (or Anything Else) in Sizable Groups," Ethology and Sociobiology, xiii, 3 (May 1992): 171-95;
42. Frank, "Mutual Policing and Repression of Competition in the Evolution of Cooperative Groups," Nature, ccclxxvii, 6549 (Oct. 12, 1995): 520-22;
43. Michod, Darwinian Dynamics; Stephen Jay Gould, The Structure of Evolutionary Theory (Cambridge: Harvard, 2002), p. 1291;
44. Okasha, op. cit.;
45. Godfrey-Smith, op. cit.;
46. Claire El Mouden, Stuart West, and Andy Gardner, "The Enforcement of Cooperation by Policing," Evolution, lxiv, 7 ( July 2010): 2139-52.
47. Some authors merely demand a reduction of lower-level selection (for example, Sober and Wilson, Unto Others: The Evolution and Psychology of Unselfish Behavior (Cambridge: Harvard, 1998)) while others insist on complete elimination. In section iv of this paper I explain why the former view of policing is superior.
48. Michod and Denis Roze, "Cooperation and Conflict in the Evolution of Multicellularity," Heredity, lxxxvi, 1 ( January 2001): 1-7, at p. 5.
49. Andy Gardner and Alan Grafen, "Capturing the Superorganism: A Formal Theory of Group Adaptation," Journal of Evolutionary Biology, xxii, 4 (April 2009): 659-71;
50. Gardner, "Adaptation as Organism Design," Biology Letters, v, 6 (2009): 861-64.
51. David C. Queller and Joan E. Strassmann, "Beyond Society: The Evolution of Organismality," Philosophical Transactions of the Royal Society B: Biological Sciences ccclxiv, 1533 (Nov. 12, 2009): 3143-55;
52. Strassmann and Queller, "The Social Organism: Congresses, Parties, and Committees," Evolution, lxiv, 3 (March 2010): 605-16.
53. Godfrey-Smith, op. cit.;
54. Henri J. Folse III and Joan Roughgarden, "What Is an Individual Organism? A Multilevel Selection Perspective," The Quarterly Review of Biology, lxxxv, 4 (December 2010): 447-72.
55. Egbert Giles Leigh, Jr., Adaptation and Diversity: Natural History and the Mathematics of Evolution (San Francisco: Freeman, Cooper, 1971).
56. Frank, "Repression of Competition and the Evolution of Cooperation," Evolution, lvii, 4 (April 2003): 693-705.
57. Buss, op. cit.
58. Wei-Jen Chung et al., "Endogenous RNA Interference Provides a Somatic Defense against Drosophila Transposons," Current Biology, xviii, 11 ( Jun. 3, 2008): 795-802.
59. Frank, "Models of Symbiosis," The American Naturalist, cl, S1 ( July 1997): S80-99.
60. Leigh, "The Group Selection Controversy," Journal of Evolutionary Biology, xxiii, 1 ( January 2010): 6-19.
61. Michod, "Cooperation and Conflict in the Evolution of Individuality. II. Conflict Mediation," Proceedings of the Royal Society B: Biological Sciences, cclxiii, 1372 ( Jul. 22, 1996): 813-22.
62. Buss, op. cit.
63. Ratnieks and P. Kirk Visscher, "Worker Policing in the Honeybee," Nature, cccxlii, 6251 (Dec. 14, 1989): 796-97.
64. Janzen, op. cit., Nature, pp. 586-89.
65. 46Weismann, The Germ-Plasm: A Theory of Heredity, trans. W. N. Parker (London: Walter Scott, 1893).
66. David L. Hull, "A Matter of Individuality," Philosophy of Science, xlv, 3 (September 1978): 335-60;
67. Hull, "Individuality and Selection," Annual Review of Ecology and Systematics, xi, 1 (1980): 311-32;
68. Clive Brasier, "A Champion Thallus," Nature, ccclvi, 6368 (Apr. 2, 1992): 382-83;
69. Huxley, The Individual in the Animal Kingdom; Gould, The Flamingo's Smile: Reflections in Natural History (New York: Norton, (1991);
70. Kim Sterelny and Paul Griffiths, Sex and Death: An Introduction to Philosophy of Biology (Chicago: University Press, 1999);
71. Leigh, "The Group Selection Controversy"; Buss, op. cit.
72. Leo Loeb, "Transplantation and Individuality," The Biological Bulletin, xl, 3 (Mar. 1, 1921): 143-80;
73. Loeb, "The Biological Basis of Individuality," Science, lxxxvi, 2218 (Jul. 2, 1937): 1-5;
74. Peter Medawar, The Uniqueness of the Individual (New York: Basic, 1957); Pradeu, op. cit.;
75. Alfred Tauber, "The Biological Notion of Self and Non-self," The Stanford Encyclopedia of Philosophy (Summer 2012 Edition), Zalta, ed., URL: 〈http://plato.stanford.edu/archives/sum2012/entries/biology-self/〉;
76. Frank M. Burnet, "Self and Not-self," in Cellular Immunology (London: Cambridge, 1969);
77. Élie Metchnikoff, Immunity in Infective Diseases, trans. Francis G. Binnie (Cambridge, UK: University Press, 1905).
78. V. L. Scofield et al., "Protochordate Allorecognition Is Controlled by a MHClike Gene System," Nature, ccxcv, 5849 (Feb. 11, 1982): 499-502.
79. Buss, "Evolution, Development, and the Units of Selection," Proceedings of the National Academy of Sciences, lxxx, 5 (Mar. 1, 1983): 1387-91;
80. Douglas Stoner and Irving Weissman, "Somatic and Germ Cell Parasitism in a Colonial Ascidian," Proceedings of the National Academy of Sciences, xciii, 26 (Dec. 24, 1996): 15254-59.
81. Dawkins, op. cit.;
82. Godfrey-Smith, op. cit., p. 100;
83. Strassmann and Queller, "The Social Organism: Congresses, Parties, and Committees," p. 605
84. Richard Lewontin, "The Units of Selection," Annual Review of Ecology and Systematics, 1 (1970): 1-18.
85. Eva Jablonka and Marion Lamb, Evolution in Four Dimensions: Genetic, Epigenetic, Behavioral, and Symbolic Variation in the History of Life (Cambridge: MIT, 2006).
86. Scott O'Neill, Ary Hoffmann, and John H. Werren, eds., Influential Passengers: Inherited Microorganisms and Arthropod Reproduction (New York: Oxford, 1997); Pradeu, op. cit
87. This view is also suggested by Sober, "Organisms, Individuals, and Units of Selection," in Tauber, ed., Organism and the Origins of Self (Boston: Kluwer, 1991), pp. 275-96;
88. Bernabé Santelices, "How Many Kinds of Individual Are There?," Trends in Ecology and Evolution, xiv, 4 (Apr. 1, 1999): 152-55;
89. Godfrey-Smith, op. cit.;
90. Queller and Strassmann, Beyond Society: The Evolution of Organismality"
91. In other work I suggest a way to use Price's equation to measure a collection's degree of individuality, by quantifying the relative strength of selection at different hierarchical levels. See Clarke, "Biological Individuality and the Levels of Selection" (PhD dissertation, University of Bristol, October 2010), p. 86.
92. This could be useful, for example, in compiling comparative measures of different lineages' progress through a transition; see Matthew D. Herron et al., "Cellular Differentiation and Individuality in the 'Minor' Multicellular Taxa," Biological Reviews (2013), DOI: 10.1111/brv.12031.
93. I am very grateful to an anonymous referee for making this point clear to me. 60 Sober and Wilson, Unto Others, p. 97;
94. Gardner and Grafen, op. cit., p. 9.
95. Such feedback processes have now been modeled, for example in Thomas Garcia and Silvia De Monte, "Group Formation and the Evolution of Sociality," Evolution, lxvii, 1 ( January 2013): 131-41;
96. Simon Powers, Alexandra Penn, and Richard Watson, "The Concurrent Evolution of Cooperation and the Population Structures that Support It," Evolution, lxv, 6 (June 2011): 1527-43.
97. Queller, "Cooperators Since Life Began," review of Maynard Smith and Szathmáry's The Major Transitions in Evolution, The Quarterly Review of Biology, lxxii, 2 ( June 1997): 184-88.
98. Godfrey-Smith, op. cit., pp. 86-89.
99. Godfrey-Smith's definition of a reproducer as possessing a bottleneck, germ separation, and functional integration is explicitly meant to apply only to collectives. Gardner and Grafen's treatment of individuality is also restricted to "superorganisms"- that is, collective organisms. Rob Wilson separates out the issue of life; see Robert A. Wilson, "The Biological Notion of Individual," The Stanford Encyclopedia of Philosophy (Fall 2007 Edition), Zalta, ed., URL: 〈http://plato.stanford.edu/archives/fall2007/ entries/biology-individual/〉.
100. I am deliberately avoiding using the word "reproduction" here, because we need not think of this as full-blown reproduction, of the sort that occurs in paradigm, derived organisms. Some kind of growth, with a tendency to fragment on reaching a threshold size, ought to be sufficient. For a discussion of how to define fitness in the absence of reproduction, see Frédéric Bouchard, "Causal Processes, Fitness, and the Differential Persistence of Lineages," Philosophy of Science, lxxv, 5 (December 2008): 560-70.
101. Francisco Varela and Humberto Maturana, Autopoiesis: La Organización de la Vida (Santiago, Chile: University Press, 1972);
102. Stuart Kauffman, At Home in the Universe: The Search for Laws of Self-Organization and Complexity (New York: Oxford, 1995);
103. Matteo Mossio and Alvaro Moreno, "Organisational Closure in Biological Organisms," History and Philosophy of the Life Sciences, xxxii, 2-3 (2010): 269-88.
104. Erwin Schrödinger, What Is Life? The Physical Aspect of the Living Cell (Cambridge, UK: University Press, 1944);
105. Carol E. Cleland and Christopher F. Chyba, "Defining 'Life'," Origins of Life and Evolution of Biospheres, xxxii, 4 (August 2002): 387-93