Abstract
Biologists in the last 50 years have increasingly emphasized the role of historical contingency in explaining the distribution and dynamics of biological systems. However, recent work in philosophy of biology has shown that historical contingency carries various interpretations and that we are still lacking a general understanding of “historicity,” i.e., a framework from which to interpret why and to what extent history matters in biological processes. Building from examples and analyses of the long-term experimental evolution (LTEE) project, this paper argues that historicity possess three essential conditions: (1) multiple possible pasts, (2) multiple possible outcomes at a given instant, and (3) a relationship of causal dependence between these two sets. These criteria can be further specified in two general forms of historicity: dependence on initial conditions and path dependence. More attention is devoted to developing a rigorous account of the latter, which captures the type of historicity displayed by stochastic processes. This paper also highlights that it is often more productive to adopt an instant-relative approach and think in terms of degree of historicity instead of trying to maintain a rigid and absolute dichotomy between historical and ahistorical (completely convergent) processes.
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Notes
Just to cite a few ones: Brown (1995), Strong (1984), Gould and Lewontin (1979), Gould (1970, 1980, 1989, 1991), Lenski et al. (1991), Lenski and Travisano (1994), Lewontin (1966, 1967), Pickett et al. (1994), Ricklefs and Schluter (1993), Szathmáry (2006), Travisano et al. (1995), Williams (1992) and Wilson (1992).
For reason of space, I will not be able to include all the relevant discussions of historicity in biology in this essay. I will focus on experimental evolution and thus leave aside many relevant examples from evolution and ecology (e.g., studies reporting that evolutionary and ecological processes are sometimes affected phylogenetic constraints (Price 2003); see also Sterelny and Griffiths (1999) for a more philosophical discussion on the role of history in ecology). I will also not discuss the position defended by Wimsatt (2001), according to which that history matters in (macro)evolution essentially because of the phenomenon of generative entrenchment. These topics are addressed more in detail in another, more extended manuscript (Desjardins 2009).
Note that there is a growing literature on “path dependence” in the social sciences (Bassanini and Dosi 1999; Castaldi and Dosi 2006; David 2001; Hodgson 1993; Mahoney 2006; Page 2006; Pierson 2004; Mahoney 2000), and that it has been recently applied in biology (Szathmáry 2006). Some differences between my and some of these accounts will be highlighted along the way.
The fitness of a derived (evolved) population is obtained by allowing the population to compete against the ancestral type. The relative fitness is then obtained by calculating the ratio of the competitors’ realized rates of increase.
And note that the populations have still not converged after 50,000 generations.
A very similar analysis was performed by Wahl and Krakauer (2000).
Note however that the variations in genotypes were not directly measured at this point. What they proved is that the fitness of all twelve populations were significantly different when introduced in a maltose limited environment. Although different genotypes could very well reach the same level of fitness, the reverse would be very surprising (Travisano et al. 1995, p. 88).
More precisely, two experiments were discussed in this paper, one with 24 (12 groups of 2) populations put in lower-temperature environment, the other with 36 (12 groups of 3) populations put in environment with different nutrient contents. But the idea was the same for both experiments: create different initial states (genotypes) and see whether these historical differences affect the evolutionary dynamics.
Recall that “chance” in these studies is usually interpreted as random mutations, drift or a combination of both.
This does not mean as we will see below that different initial states is necessary.
I base the following analysis of “tree” on Belnap et al. (2001), although the nodes are not “moments” in my account, but “states.”
Note also that, unlike many accounts of path dependence, this definition does not assume that outcomes are equilibrium states or stable attractors.
But see Ben-Menahem (1997) for a similar notion.
In this case, the order of environments was simply reversed for the second run of the model.
Dependence on initial conditions can also occur in stochastic processes. I chose the case of deterministic processes because they are simpler to represent.
Note that identifying alleles and genotypes is possible only for haploid organisms.
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Acknowledgments
I wish to thank John Beatty, Paul Bartha, Christopher Stephens, Robert Batterman, Gillian Barker and Christopher Smeenk, and the reviewers of Biology and Philosophy for their comments on earlier drafts of this paper.
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Desjardins, E. Historicity and experimental evolution. Biol Philos 26, 339–364 (2011). https://doi.org/10.1007/s10539-011-9256-4
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DOI: https://doi.org/10.1007/s10539-011-9256-4