The Cambridge Companion to the Philosophy of Biology (Cambridge Companions to Philosophy)

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Readers who compare earlier collections of essays in the philosophy of biology with the present volume may notice the disappearance not only of evolutionary laws, but also of the received theory of explanation based on them. They may notice, that is, the disappearance of the notion that a scientific explanation, as distinct from a psychologically satisfying explication, is a deduction of a fact from a general law.

This once dominant view has been displaced by the idea that explanation is an inference to the best causal account of a given phenomenon, laws or no laws. This conception first emerged in connection with demands for greater realism than logical empiricism, with its phenomenalist roots, could ever muster.

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It allows for more realism in part because, as Lindley Darden puts it in her chapter on mechanisms and models, the view attributes to mechanisms "the roles attributed to general scientific theories. They provide explanations of puzzling phenomena. They enable biologists to make predictions" For Darden, mechanisms perform these roles because they reveal "productive continuity between stages [of a phenomenon] ….

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The entities and activities of each stage give rise to the next stage" Such mechanisms, she reports, are discovered through and subsequently represented by models. These include "model organisms" like the estimable fruit fly, which has done yeoman service in the development of evolutionary theory for over a century.

Inference to the best explanation is often invoked in order preserve the adequacy and autonomy of adaptationist explanations in the face of reductionist prophecies like those of Rosenberg. Does not natural selection remain the best explanation of adaptation even if it is not "covered by" laws? At the same time, neo-Paleyan intelligent design advocates, who are as quick as their creationist predecessors to turn the latest ideas of philosophers of science against evolution, have found their way to inference to the best explanation in order to push the notion that intelligent design, not natural selection, is the best explanation of putatively irreducibly complex biochemical mechanisms.

This argument depends in part on the notion that organisms contain "molecular machines. Nor are organisms as wholes mechanisms unless one arbitrarily imposes an engineering or artifact paradigm on entities that on closer inspection seem to be "built … by tinkering and satisficing" Nor, finally, is there anything machine-like about the mechanisms by which lineages of organisms are explained, "such as the isolating mechanisms leading to speciation" These considerations hark back to the issues about adaptation raised at the outset.

Inference to intentional design as the best explanation of organic functions depends on invoking efficient means-end reasoning. If that is what Darwin meant by teleology, intelligent design might have a prayer. But, Ariew argues, Darwin's teleology is more like Aristotle's than Paley's. No less than Aristotle, Darwin thought of organisms not as an assembly site where separate traits favored by a particular environment happened to collect, but as outcomes of an integrated, end-oriented process of development. To be sure, unlike Aristotle, Darwin thought that beneficial changes in this process, rather than its perpetual repetition or degeneration, needed explaining and he appealed to natural selection in conjunction with strong heredity to do the explaining.

Still, attending to the developmental context seems just the right antidote to the provincial, design-based conception of teleological explanation that for historical reasons dominates Anglophone cultures. Its ordered end-orientation undercuts the widespread assumption that the only alternative to design is pure chance, a misunderstanding that has dogged Darwinism from the outset. Nonetheless, I think Ariew fails fully to exploit the analogy to Aristotle when he claims that for Darwin natural selection is not the author of inherited biological form, but merely of traits that show tinkering with local constraints.

The Origin of Species does not draw the contrast this way. I turn now to chapters on the state of play in biological fields that the Modern Evolutionary Synthesis hoped to attract to its population genetical core. It made progress with adaptive radiation and speciation. But by the ls its expectation of eventually capturing developmental biology, ecology, and phylogeny -- the three core disciplines of what might be called the old, l9th century evolutionary synthesis -- began to stall.

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Partly as a result of the belated discovery that genetic architecture is highly conserved across lineages and the inference that differences must be due to the interaction between genes and other factors in the ontogenetic process, developmental biology has reemerged strongly in recent decades. Jane Maienschein shows that the epigenetic roots of developmentalism contrast with the weak preformationism of the "genetic program" trope that in the ls fused genetical Darwinism with post-Watson and Crick molecular genetics.

Experimental work on stem cells, for example, has been going on since the first decade of the 20th century; it only appears to have recently come out of nowhere because the preformationist bias of genetic Darwinism had screened this work from view. In a similar spirit, Manfred Laubichler gives a genealogy of evo-devo and devo-evo research programs, which hold that evolutionary changes cannot simply be functions of changes of gene frequencies in populations. I should mention, because the Cambridge Companion does not, that some philosophers who prefer Fisher's causally inert conception of fitness over the propensity interpretation do so in order to shift the explanatory focus from population dynamics back to variations that emerge in the course of individual development Walsh, For them population dynamics is really just population kinetics.

What about ecology? In the s, serious efforts were made to bring community ecology into the Modern Synthesis by mathematically modeling competition between species. In his chapter on the field's current state, however, Gregory Mikkleson argues against reductionist biases which, in treating ecologies as closed systems of competing species, miss the fact that even on islands these systems are usually open to migration.

In open systems, Mikkleson says, migration cannot simply be added as one more "force" to the see-saw between competing plants and animals. That is because when the reductionistic biases associated with closed systems are removed it is as likely as not that the appropriate level of causal interaction will also shift to higher levels and units, such as "metapopulations, metacommunities, landscapes, regions, biotic provinces, or at the largest scale to date, the entire biosphere or ecosphere" The relevance of development and the ecology of open systems, as well as the limits of reductionism, are also themes in Kim Sterelny's informative chapter on macroevolution.

By way of a fascinating review of literature on evolutionary trends and major evolutionary transitions, especially the Cambrian explosion of bilaterian metazoa, Sterelny shows that the Modern Synthesis's gambit of treating all macroevolution as extrapolation from population genetical microevolution is inadequate. To be sure, this "minimalist approach," as Sterelny calls it, is superior to Stephen Jay Gould's neo-saltationist notion that macroevolution is dominated by chance. In the Cambrian, however, developmental changes that were already afoot resulted in taxa that were able to affect, not just adjust to, environmental contingencies.

Their enhanced agency was made possible by preadaptive modular construction, increasingly powerful nervous systems and sensory organs, and complex life histories. Accordingly, Sterelny concludes that "the diversification pulse in the Cambrian represents a change in developmental program rather than change in selective regime" This is enough to undermine Dawkins's assumption that "these evolutionary changes can be understood as the replacement of one allele by another in the context of invariant systems of gene regulation and expression" Maureen Kearney's review of the triumph of phylogenetic systematics cladism over rival theories of classification contains echoes of the developmentalist themes to which I have been pointing.

Cladism says that branching points alone, not perceived similarities, should be used to construct classifications. Cladists rejected efforts by the rival pheneticist school to make classification more empirical as in logical empiricism by identifying and sorting through more characters than traditional practitioners of the art of taxonomy. Instead, cladists appealed to Popper's falsificationist norms by assuming that nature's branching points could always be counted on to take the most parsimonious path. Still, in constructing their parsimonious cladograms of ancestral and descendent species they typically used conventional diagnostic characters, if fewer.

Kearney reports, however, that this method is now being overtaken by efforts to "order things into systems on the basis of the natural process by which their parts [rather than sums of traits] are related" Jason Roberts's chapter on systems biology testifies to this anticipated shift. Having found their way through genome sequencing programs to a mother lode of information, Roberts reports that even molecular geneticists are now calling for a new "systems biology" that will integrate genetic with other sorts of data.

Since the systems in question are developmental, Roberts applauds "molecular biologists' apparent realization finally! I conclude this section by noting that, in contrast to the genocentrism that still dominates popular science writing about evolution, contemporary philosophers of biology sense that the link between phylogeny and ontogeny is returning to its previous centrality.

Nowhere in the Cambridge Companion, though, is one pressing issue squarely addressed: What precisely is the relation between population genetics and the new developmentalism? The role of developmental plasticity in negotiating the complex, dynamical interaction between genotypes and phenotypes, as in the work of Mary Jane West Eberhard, would be a good place to look.

I turn finally to essays on human evolution. Developmental themes are less prominent in these chapters. One reason might be that for the last several decades questions about human evolution have been dominated by attempts to capture this subject for gene-centered versions of the Modern Synthesis by appealing to kin selection and game theory to explain cooperative and altruistic behavior. The role of game theory in this work is well summarized in Zachary Ernst's chapter.

As a result, debates about human evolution still reflect old quarrels about whether cultural processes are products of natural selection, but now work autonomously -- the orthodox Dobzhanskyan view set forth in the Cambridge Companion by Francisco Ayala -- or whether human traits, including our sexual proclivities and cognitive quirks, are distinct adaptations. The latter hypothesis was first proposed in E. Wilson's Sociobiology As David Buller explains in his chapter, Wilson's sociobiology had two successor programs. One is Evolutionary Psychology, which postulates distinct mental modules for different psychological functions and regards each module as a domain-specific adaptation.

The other is human behavioral ecology, which, in Dobzhansky's spirit, treats the human mind as a highly plastic domain-general capacity for managing the trade-offs on which adaptedness in cultural niches depends. Buller expresses a decided preference for the latter.

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A similar preference can be found in Valerie Hardcastle's report on trends in neurobiology. Extreme modularism underestimates the noise-signal ratio in brain functions and the plasticity of the brain, leading Hardcastle to skepticism about too quickly adjusting public policy toward potential criminals for example under the influence of fads in cognitive neuroscience. This sort of cautionary skepticism can also be felt in chapters that deal with other issues where evolutionary science is supposed to ground public policies. Sahotra Sarkar, for example, points out the disconcertingly thin mathematical and empirical grounds on which policies aimed at protecting biodiversity actually rest.

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  7. Presumably, philosophers of biology can be persuasive witnesses on topics where biology meets ethics, law, and public policy because they are supposed to know their way around is-ought logics. Nonetheless, their interventions will fail to persuade the public unless philosophers bear in mind the evolution-religion matrix with which bioethical issues are hopelessly entangled. Robert Pennock's chapter on biology and religion is helpful because it rises above the debaters' points that dominate public discussion of creationism, sexuality, environmentalism, biotechnology, and other such issues.


    All these controversies, Pennock argues, have a common thread -- whether nature is so inherently value-laden that it dictates norms of behavior. In the modern West, religion except of the most liberal sort still sticks up for this natural law tradition. Oddly, however, in opposing its religious form Sociobiologists and Evolutionary Psychologists perpetuate the normative assumptions of this tradition. Consider homosexuality.

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    In his chapter on the subject Christopher Horvath summarizes evidence pointing to some heritability of male homosexuality, but puts more stress on the non-heritable, but still decidedly biological effects of birth order. A statistically higher level of homosexuality in younger sons correlates with immune responses in mothers who have already borne a number of children. Since ought implies can Horvath says these facts call for tolerance. They remind us "to treat all people with respect and dignity regardless of their sexual orientation" But this little sermon still leaves open whether homosexuality, compelled or not, is morally good, bad, or indifferent.

    This loaded question is usually, and badly, framed against an assumed background of masculinist heternormativity. The fact that sociobiologists and their successors often defend male centered heterosexuality as natural is not unconnected with the concessive tone of their claim that childless or gay uncles and aunts can still contribute to overall fitness through kin selection.

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    But those who take this line, like those who postulate a "gay gene," are assuming that moral goodness depends on adaptedness. Pennock points out that they are defending the normativity of nature no less than those who say that the heritability of religion means that there must be a "god gene. Indeed, it is the same game. It not only pulls ought out of is, but has a bad track record. In retrospect, it is clear that prejudices and pipedreams, not ineluctable facts, have generated and protected from criticism once cherished notions about the biological basis of free markets, eugenics, and, in the era of feminism, heterosexual male dominance.

    This Cambridge Companion, like others, is not a general survey. Rather, readers will encounter well-informed and analytically skilled minds grappling at close quarters with biology's most interesting issues. Since philosophy of biology is a generally naturalistic discipline -- it tends to treat conceptual and empirical claims as thoroughly intertwined -- readers will also discover that they are picking up a lot of up-to-date biology.

    In view of evolutionary biology's entanglement with questions of value, however, philosophers of biology should reflect as much on the cultural history of the life sciences as on biological science itself. As it is, Robert Richards's chapter on why it is unfair to blame Ernst Haeckel for Nazism is the only essay on this theme.

    The exoneration of a great scientist from the crude fallacy of reductio ad Hitlerum is fine. But it should not be allowed to displace the overall record of biologists who have called on nature to bless certain social arrangements. The volume could also use a bit more reflection on the nature and uses of philosophy of biology itself.

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    Steven Crowell (ed.), The Cambridge Companion to Existentialism - PhilPapers

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