Abstract
The adaptive landscape is a metaphorical device employed to depict the evolutionary change in a population or lineage undergoing natural selection. It is a powerful heuristic and didactic tool. This paper has two objectives. The first is to dig beneath the adaptive landscape in order to expose certain presuppositions about evolution concealed there. The second is to propose and motivate an alternative spatial metaphor, one that embodies a wholly different set of presuppositions. I develop the idea that adaptive evolution occurs on an ‘affordance landscape.’ The conception of adaptation—both the process and the product—that follows from adopting the affordance landscape metaphor is a significant departure from the conception of adaptation embodied in orthodox Modern Synthesis biology.
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Notes
- 1.
With characteristic modesty, Lewontin credits this dictum—an allusion to a similar saying about the condition or price of liberty—to Rosenblueth and Wiener. However, I was unable to find it in any of their co-authored papers. It appears unattributed, though enclosed in quotation marks, in Lewontin (1963, 230).
- 2.
- 3.
- 4.
There may be differences in landscape topology as we investigate different levels of detail (Wilkins and Godfrey-Smith 2009), but the processes are the same at every scale.
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- 6.
- 7.
Notice how naturally convergence falls out of the traditional conception of organism/environment relations.
- 8.
Nerlich (1991) calls the independence of spatial relations from spatial properties the ‘Detachment Thesis’: “thing-thing relations are logically independent of thing-space relations” (172).
- 9.
There is a further, related, Newtonian analogy to be considered. Most philosophers of biology seem to hold that adaptive space is inert. Consequently, extraneous causes or forces, like selection and drift, are required to propel form across the adaptive landscape.
- 10.
A Paramecium actually has three ‘gaits,’ only two which involve the asymmetric beating of cilia. See Hamel et al. (2011).
- 11.
Some of the implications of seeing evolution as a response to affordances are discussed in Walsh (2012).
- 12.
Which dispositional properties are represented as Reynold’s Number (Purcell 1977).
- 13.
A sympathetic reading of Gibson (1979), I believe, suggests the same.
- 14.
It is interesting that in those sciences in which the niche concept plays a genuine theoretical role, e.g. community and population ecology, the niche concept is often defined more in the way an affordance is. The niche concept seems to have originated with Elton (Hutchinson 1978) and was defined in terms of resource utilization. Odum (1959) likens a niche to an organism’s ‘profession.’ See Beatty (1995). I thank Sahotra Sarkar for pointing this out to me.
- 15.
One additional advantage of not seeing affordances as dispositional properties of an organism’s environment is that it relieves us of the temptation of thinking that all affordance-presenting features are external to organisms. Inner workings of organisms present affordances too.
- 16.
In the case of organisms, ‘value’ may be read, often enough, as ‘survival value.’
- 17.
I borrow the term ‘commingled’ from Haugeland (1998). The relation between organisms and their conditions of existence I envisage includes but extends beyond what Gillian Barker (2008) calls ‘selective interaction.’ The principal difference is that selective interaction emphasises the ways in which organisms causally influence their conditions of existence. The ‘commingling’ of organisms and their affordances underscores the way in which the capacities of organisms partly constitute those conditions. See Walsh (2012) for a discussion of the distinction.
- 18.
I would suggest an amendment to West-Eberhard’s definition. Plasticity should be seen as the capacity to react to an input from any source, not merely an environmental input.
- 19.
Confusion persists on this point. See, for example, Sterelny (2009, 101) who claims that novelties generated by phenotypic plasticity are “mere ecological events.” “Such novelties have no effects on the germline are not inherited [sic]” (2009, 94). He muses on how they can be transformed from “mere ecological events” into evolutionary events. My claim is that no transformation is needed; any ecological event that is intergenerationally stable is an evolutionary event.
- 20.
One salutary implication, for both relativity and its metaphorical extension to evolution, is that it is not necessary to posit an additional metaphysical category of force to propel bodies through space.
- 21.
Perhaps ironically, Conway Morris (2010) offers a number of these.
- 22.
I thank Chris Haufe for the quotation.
- 23.
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Acknowledgments
I would like to thank audiences in London (Ontario), Dubrovnik, and Stockholm for their lively responses. I am particularly grateful to Fermin Fulda and Susan Oyama for suggested improvements to an earlier draft. In addition, Jacob Stegenga, Cory Lewis, Michael Cournoyea, and Alex Djedovic provided a very helpful discussion. The editors of this volume also provided very helpful suggestions.
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Walsh, D.M. (2014). The Affordance Landscape: The Spatial Metaphors of Evolution. In: Barker, G., Desjardins, E., Pearce, T. (eds) Entangled Life. History, Philosophy and Theory of the Life Sciences, vol 4. Springer, Dordrecht. https://doi.org/10.1007/978-94-007-7067-6_11
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