Form and Function in Developmental Evolution (Cambridge Studies in Philosophy and Biology)

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The only useful measure of adaptedness so construed is, as Cuvier thought, the very existence of the organisms, populations, species, and wider lineages whose presumptively integrated suites of traits sustain their development, life-styles, adaptability to environmental fluctuations, and reproductive prowess. True, Reiss thinks that even if one can conceptually distinguish selection from genetic drift—he doubts it —they are almost impossible to tell apart in practice. On this point, Reiss agrees without saying so with Matthen and Ariew [] and Walsh et al. His implication is that this is nice work if you can get it, but even if you succeed in getting it what you have done is find how the characters of organisms have enabled certain organisms to remain within the boundary conditions that define their conditions of existence.

The larger point is that both medium and narrow sense selection are in the service of broad sense selection. Seen in this way, the teleological fallacy should be far less tempting. Let me make two remarks about these claims. First, I offer a historical note. On this reading, Fisher counted average reproductive success after the fact as fitness and so avoided teleology. In this he differed from Wright, Dobzhansky, Simpson, and Bock, who discriminated between drift and selection in a vain effort to find causes at the population level and ended up instead falling into the teleological fallacy.

Unsurprisingly, Reiss rejects the so-called etiological or selected effects view of functions, according to which biological functions are adaptations that have been created by directional selection for the purpose of performing some current physiological or behavioral job that has a positive effect on reproduction.

Knowing the origins of traits that contribute to maintaining conditions for existence is, in any case, less important than the fact that they have arisen. Consequently, Reiss analyzes biological functions in roughly the system-maintaining way pioneered by the philosopher Robert Cummins and developed recently by Arno Wouters Wouters An implication, I think, is that without knowing that they are evolved adaptations the pre-evolutionary Harvey and Cuvier could still find and validate biological functions as well as Darwin or, say, Richard Dawkins.

In fact, they did just that.


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In addition, the Cummins-Wouters-Reiss analysis of biological functions has the further advantage for Reiss of evading thinking that organisms, like The Little Engine That Could , are constantly huffing and puffing their way up adaptive hills 25, Not looking obsessively for adaptations means that we do not have to engage in the nearly impossible task of finding their etiologies. We do not, Reiss writes, need to. Still, there are advantages for critics of design teleology to retaining consequence etiology of some sort, if that is possible.

Such accounts depend entirely on the amplification of past reproduction-enhancing events rather than on any future-oriented appeals to intentions of the sort that for Reiss lead to, or come with, design teleology. On the etiological or selected-effects view, traits spread through populations because their historical precursors had reproductive effects in the past that increased the proportion of organisms that carried them. I think an effort in this direction is warranted. I admit that there are many philosophical formulations of consequence etiology that do seem to project the offending picture Reiss sketches in the paragraph just quoted.

However, the main problem with these formulations lies primarily in their attempt to causally explain particular adaptations on the assumption that they are the evolutionary equivalent of discrete biological functions. To me these two concepts should not be assimilated to each other Depew When they are, two different claims, one misguided, the other not, get conflated. If you think evolving discrete adaptations is the most important process in evolution you will need detailed, but alas usually unavailable, knowledge of the history of a trait in order to determine whether it is an adaptation, an exaptation, or a concomitant result of selection for something else.

If, however, you decline to think of organisms as bundles of adaptations, as Reiss does, and if you analyze functions in a role-in-a-system-no-matter-where-they-come-from way, as he also does, it may still be possible to acknowledge that the overall adaptedness of organisms, populations, or lineages is maintained precisely because they have a history that sustains that adaptedness. The fact that one does not know that history, or at least much of it, makes no difference if one is not captured by the design paradigm and the search for adaptations.

All that matters is that it did happen and happened in such a way that it resulted in beings able to satisfy their conditions of existence in what usually turn out to be very flexible ways. To that extent, the adaptedness or fitness that is warranted by continued existence is etiologically grounded and caused. It is worth pointing out in this connection that some philosophers think that the consequence-etiological analysis of biological functions, precisely because it departs from the design paradigm, actually requires the kind of developmental holism about organisms that Reiss promotes McLaughlin To get rid of design and its adaptationist doubles in this way one may have to call on dynamical models of complex self-organization to provide a suitable framework for picturing and mathematizing organisms as inherently adaptive developmental systems Depew and Weber Empedoclean-Epicurean-Lucretian materialism certainly contrasts with and throws doubt on design talk.

But it also makes the etiology of organisms, species, and lineages a series of accidents.

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Adaptive traits do not arise in order to deal with environments, but just happen to have adaptive effects that are retained by environments ; see Depew If he did he would be begging the question about the inaccessibility of the past that his analysis strongly requires him to leave open. Intriguingly, Reiss ascribes Epicurean metaphysics, and so an implicitly accident-prone etiology, to Cuvier , However that may be, I doubt whether an evolutionary Cuvier would hold it.

I imagine that an evolutionary Cuvier, like an evolutionary Kant, would agree that the evolved ability of complex adaptive systems to adjust to changed circumstances rules out happy accidents just as much as it rules out thinking of organisms as assemblies of discrete adaptations by an analogue of conscious design. Such models typically do treat organisms as collections of independently optimized traits and do rely, as Reiss says, on a strong analogy between organisms and artifacts. In the process they probably do smuggle in under cover of seemingly innocent metaphors traces of the old theological doctrine of creatio ex nihilo.


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I am less sure, however, that practicing evolutionary biologists or the mathematical methods and models in which they are schooled are necessarily compromised by design teleology. It is not clear, in the first instance, how ontologically committed muddy-boots evolutionary biologists have to be in order to go about their work productively, as Reiss clearly wants to do, but with a clean philosophical conscience.

But even if their models and methods are inseparably connected to larger evolutionary theories, which are indeed ontologically committed, I have raised in the preceding section the possibility that the design-without-a-designer paradigm is merely one or at most a family of interpretations of genetic Darwinism.

There are also, among others, etiological interpretations. These evade the design paradigm even if Reiss is right that they fail to evade the deterministic notion that external environments mold adaptive traits. He has, however, made about as good a case for that proposition as anyone is likely to make.

Form and Function in Developmental Evolution (Cambridge Studies in Philosophy and Biology)

What, then, of the accusation that Gould himself covertly retained adaptationist assumptions? Gould thought of them as something that adaptive natural selection wants to find a way around. They are like old, deeply rooted, gnarly tree stumps around which farmers must plow.

The stumps themselves are largely the result of previously successful evolutionary novelties that selection has successfully locked in by entrenching them in the developmental programs of organisms. The teleology that Reiss ferrets out of Gould lurks in that counterfactual. Selection, Gould argues, is unable to get around developmental constraints. Hence, he concludes, adaptation ism is wrong. But, as Reiss rightly points out, the very optimizing assumptions against which Gould launches his glittering polemics are buried in a shallow grave in his way of conceiving of development Reiss argues that development enters into the evolutionary process not in the way it biases the distribution of available variation by the exertion of evolutionary forces such as selection, but in the way in which the developmental process itself is the source of the variation by which beings as presumptively adaptive as organisms maintain their conditions of existence.

I get the impression that for Reiss phylogenetic disparity and diversity are no more than the accumulated effects of forks in the road in which conditions of existence were or were not maintained at the organismic level. In one way or another, all these writers point out that if the developmental process and the complex way it as opposed to genes interacts with the environment is the presumptive and proximate source of potentially adaptive variation, then organisms must be seen as inherently adaptive developmental systems.

Where arguments arise among advocates of this new family of evolutionary theories is on how adaptive ontogenetic processes are in themselves and therefore on what role and importance remains to natural selection. Some have hardly any use for selection at all, proposing autopoietic self-formation or self-organization as a substitute.

I think Reiss, like Massimo Pigliucci, has a more positive view of natural selection than that. Pigliucci is as dubious about distinguishing drift from selection as Reiss Pigliucci and Kaplan , 29; Reiss, But Pigliucci sees the developmentalist turn from the perspective of quantitative genetics, developing themes about phenotypic plasticity and evolvability that go back to Dobzhansky and Lewontin Pigliucci To that extent he has a more positive view of the causal significance of population genetics than Reiss.

Perhaps for this reason he retains an etiological view of adaptedness Pigliucci and Kaplan , He and his philosophical co-author take consequence etiology to offer protection against the design paradigm in both its natural and theological senses. In doing so they do not seem to think that they have entangled themselves in the environmental determinism that Reiss thinks carries design talk with it. There are many other related views on offer at present and still more are likely to come.

I would even venture to say that the developmentalist turn in evolutionary theory is still at the stage of inventing a wide array of related theories rather than perhaps prematurely selecting against some of them. Reiss is also to be commended for realizing that evolutionary biologists cannot ignore, if they are to do their work productively, the conceptual history and philosophy of their discipline.

To be sure, anyone who goes back through history to find the roots of current problems runs a risk of dragging the past into an empirically advanced present or conversely of inscribing the present into the past. Still, finding the roots of present practice, if done as responsibly as Reiss does it, increases the likelihood that we may turn up something that throws an unexpected light on business as usual.

By reading and thinking hard about the historical roots of present practice, Reiss has written a challenging book that, especially when read in dialogue with other works on his themes, will contribute to facilitating the great transition that evolutionary theory is currently undergoing. I wish to thank an anonymous reviewer and Alan Love for penetrating and helpful comments on an earlier draft of this essay. This is an open-access article distributed under the terms of the Creative Commons Attribution-NonCommercial-NoDerivs license, which permits anyone to download, copy, distribute, or display the full text without asking for permission, provided that the creator s are given full credit, no derivative works are created, and the work is not used for commercial purposes.

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Form and Function in Developmental Evolution (Cambridge Studies in Philosophy and Biology)

Quick search:. Disciplinary disagreements over the interpretation and weighting of evidence need reconciliation, not adjudication, and recent research validates a synthetic approach to the problem Xu et al. One lesson is that productive evidential integration occurs more readily in the local context of a specific question avian digit homology , rather than at a global or theoretical level. The philosophical contributions cover a narrower territory than that tackled by the biologists, and this is an unfortunate drawback to the volume.

We are given just enough of a taste to want more. The thematic links are more tenuous for the chapter by Richards, which is focused on systematics and rehearses a dated methodological debate over functional analysis and its role in comprehending character transformations.

How this touches on the canvas of Evo-devo is opaque, and the unblinking treatment of a nonstandard approach using morphological hydrostatics adds to the confusion. There is no doubt that this volume stimulates reflection on questions of interdisciplinary integration. How much is supplied beyond stimulation is another matter. Many issues remain unaddressed, especially on the philosophical side: how do methodological and explanatory standards from the different disciplinary strands come together around form and function? How does one combine functional morphology and molecular developmental genetics?


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Tantalizing exemplars appear in the combinations of investigative techniques displayed by the scientific contributors, but mostly it is puzzle pieces without a box to know how they fit together. A more direct antidote would be a challenge to the very idea of an integrated theory of evolution. But these piecemeal models harbor the possibility that a pluralist stance on evolutionary theory is not a temporary state of affairs i.

Related Papers. Evolutionary novelty and the evo-devo synthesis: field notes. By Stavros Ioannidis. Conceptualizing evolutionary novelty: moving beyond definitional debates. Interdisciplinary lessons for the teaching of biology from the practice of evo-devo. By Alan C Love.

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