Abstract
The concept of fitness has generated a lot of discussion in philosophy of biology. There is, however, relative agreement about the need to distinguish at least two uses of the term: ecological fitness on the one hand, and population genetics fitness on the other. The goal of this paper is to give an explication of the concept of ecological fitness by providing a reconstruction of the theory of natural selection in which this concept was framed, that is, based on the way the theory was put to use in Darwin’s main texts. I will contend that this reconstruction enables us to account for the current use of the theory of natural selection. The framework presupposed in the analysis will be that of metatheoretical structuralism. This framework will provide both a better understanding of the nature of ecological fitness and a more complete reconstruction of the theory. In particular, it will provide what I think is a better way of understanding how the concept of fitness is applied through heterogeneous cases. One of the major advantages of my way of thinking about natural selection theory is that it would not have the peculiar metatheoretical status that it has in other available views. I will argue that in order to achieve these goals it is necessary to make several concepts explicit, concepts that are frequently omitted in usual reconstructions.
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Notes
I will use this terminology. The use of “ecological” does not imply that this is a concept of ecology since, as I will show, the ecological fitness is the fitness from NST.
Of course, other authors have noted this ambiguity (e.g., West-Eberhard 1994; Burian 1994; Gould and Vrba 1982; Brandon 1990, pp. 39–44). I do not make use of any existing account because later on I will characterize adequations in a way that differs from those available. This characterization will be crucial, since it will influence our view of the very structure of NST.
The election of this name, ‘historical theory of natural selection’, presupposes that this historical explanation can be thought of as an application of a theory that can also be reconstructed by means of metatheoretical structuralism. Though I think this can be defended, this discussion will not be addressed here. There is a precedent analogous to the case at hand of two theories that are frequently confused, one of which has the standard form, and the other has a more historical character and iterates the former theory repeatedly and also adds some new concepts. This is the case of the distinction made by Lucía Federico between metabolic biochemistry and biochemistry of the metabolic pathways (Federico 2009, p. 98). Nevertheless, even if it was not the case and what I have been calling HNST is nothing more than the iteration of NNST, the distinction can still be thought of as two different ways of applying the same theory—instead of two different theories—and the main points of my paper remain unaltered. In any case, to get a better understanding of the historical explanations proposed by Darwin, a reconstruction of NNST—a theory that we find presupposed in his texts—is needed.
The informal application I make of structuralism leads to a presentation of the fundamental law by means of statements, which may cause the reader some confusion given that structuralism is a semantic metatheory. It must be remembered that I am not presenting NNST in a structuralist language, but rather using important concepts of this metatheory in an informal way.
In Sect. 4.5 I will focus on the notions of function and differential reproductive success and the reason why heritability is introduced in the fundamental law.
Although Darwin sometimes presents sexual and natural selection as if they were two distinct mechanisms, he at times also speaks of natural selection in a more comprehensive fashion. I have presented a defense of this point in a previous paper (Ginnobili 2011b). Endler (1986, pp. 11–12), Gayon (1998, pp. 51–54), and Ghiselin (1969, p. 215), amongst others who share this view.
One interesting issue with these sort of applications, which can also generate reasonable doubts about the attempt to reduce artificial selection to natural selection, is that in this case it may seem strange that the long tail of a dove produced by artificial selection has the function of improving the chances of crossings being attractive to the breeder. Nevertheless, it should be noted that in this sense the application does not seem different from the one in which a flower improves its chances of being crossed by being attractive to an insect. Nor is it different from the one that considers that a peacock improves its chances of crossing by looking attractive to females of their species. The issue deserves further discussion. Here I present these applications separately to adjust more to Darwin’s writing, which usually presents artificial and natural selection as separate mechanisms, and considers that the case of insect attraction through flowers is not a case of sexual selection. Some authors, however, have found closer relations between artificial and sexual selection—e.g., Ghiselin considers the possibility that sexual selection is a variant of artificial selection (Ghiselin 1969, p. 246), which goes in the same direction as some statements by Darwin himself (1871, v.I, 259).
Regarding the reconstruction of NST, group selection as it was considered by Darwin is especially interesting, since entities of different level are interrelated in the same application. Below, I include “performance of the community they belong to” as a specification of fitness, but the complete formal reconstruction requires a better way of dealing with group selection cases (see Ginnobili 2012b).
An interesting precedent for the kind of work that tries to propose terms for concepts that function implicitly in practice, and which the title of this work is based on, is the article on the exaptations of Gould and Vrba (1982).
The application of this principle is always ceteris paribus given the existence of other evolutionary mechanisms. A problem of this formulation arises in the vacuous satisfaction of the conditional. If the adequation of the trait does not produce an increment in the fitness, then the law will be true. An option is to replace the first conditional with a conjunction. But then the idea that the adequation that is relevant to evolution produces an increment in the fitness is missed. Another option is to replace the material conditional with another kind of counterfactual conditional.
Elsewhere I have reconstructed NNST formally in structuralist terms making explicit all the concepts necessary to account for this structure in a more meticulous way (Ginnobili 2012a). For reasons of space I only provide the informal version, which is sufficient for the purposes of this paper. There is another structuralist formal reconstruction of NNST in Mario Casanueva (2011).
The relation between functional biology and evolutionary biology in Darwin’s texts is extremely interesting; nevertheless, treating it properly is not possible here. I have discussed this issue somewhere else (Ginnobili 2014).
The claim that the NNTS’ fundamental law includes at least these three concepts does not imply that every application includes only three elements. It is possible to have applications in which more traits interact, with more functions, with diverse specification of fitness. Special laws do not have to preserve the form of the fundamental law, even though they do have to maintain the same concepts and preserve certain essential relations among them.
This theory-net arises only from the examined cases, that do not exhaust all the applications Darwin makes of NNST in the Origin, nor all the specializations used by Darwin in other texts or discovered by other biologists. These might change the structure in question. I think, for instance, that it is possible to include as specialization the principle of divergence, but this discussion would take up too much space. I do not expand the branch of group selection, because all the cases treated in this work imply an improvement in the survival of the group, but maybe it is possible to find the same subclasses that we find in the branch of individual selection. The theory-net presented, therefore, is neither complete nor the only way to present the available information, but rather a possible way to organize the cases we have been studying, which allows us to show the complex and unifying structure of NNST.
Tim Lewens (2007, pp. 58–62) thinks that contemporary NST differs from Darwinian NST, since in contemporary NST the struggle for existence is not essential, as it is in Darwinian NST. In my reconstruction of Darwinian NST the struggle for existence—in Darwinian terms, the fact that more individuals are born than those that can survive—does not appear. This may seem a problem, but I think I have given enough bibliographic support to the idea that the struggle for existence understood as a struggle for survival is not always involved. If we think of the struggle for existence in a broad and metaphorical sense—as Darwin asks us to do (1859, p. 62)—as a struggle for leaving progeny, then there would be no difference between contemporary NST and Darwinian Theory. This metaphorical sense is covered by my reconstruction. Maybe the point raised by Lewens has to do with the role of limitations of resources that appear in the abstract way in which Darwin usually presents natural selection—as an inference from the geometric growth of populations, the arithmetic growth of food, the variation and the tendency to inherit variations, among other things (Darwin 1859, pp. 80–81). There are two points to be made about this. First, there is no reason to think that natural selection can only work in cases of limited resources (Barbadilla 1990, p. 172; Sober 1993, p. 194). Second, Darwin himself did not think that the cited argument results the theory of natural selection, but is an argument that makes natural selection probable (Darwin 1883, v. I p. 9; Gayon 1998, p. 23). This argument, moreover, should not be confused with the structure of the theory (Kitcher 1993, pp. 34–37).
There are two ways in which we can say that Darwinian theories have unifying power, both present in Darwin’s texts. The first one has to do with the fact that Darwinian evolutionary biology can unify data from different disciplines (geology, embryology, systematic, biogeography, etc.; see e.g., Ruse 1998, p. 3). The second one has to do with the fact that NNST has a wide and heterogeneous set of intended applications, in the same sense that Classical Particle Mechanics has (for example, Kitcher 1981, 1993). This latter sense is the one alluded to in this work.
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Acknowledgments
I would like to thank Martín Ahualli, Daniel Blanco, José Díez, Pablo Lorenzano, Andrea Melamed, Luciano Piazza and Ariel Roffé for their helpful comments on previous versions of this paper. Also, I am especially grateful to Staffan Mueller-Wille for his careful work as an editor with this paper. This research was supported by the research projects PICT-2014-1741, PICT-2012-2662 (ANPCyT, Argentina), PIP No. 112-201101-01135 (CONICET, Argentina) and 32/15 255 (UNTREF, Argentina).
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Ginnobili, S. Missing concepts in natural selection theory reconstructions. HPLS 38, 8 (2016). https://doi.org/10.1007/s40656-016-0109-y
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DOI: https://doi.org/10.1007/s40656-016-0109-y