One of the most famous battles of ideas in comparative biology was that between Etienne Geoffroy St. Hilaire and George Cuvier over the existence, or not, of a common plan of construction (or Bauplan) for animals (Appel, 1987). Geoffroy was of the opinion, previously developed by Buffon (1753), that all animals are built according to a single plan or archetype, but Cuvier, France's most illustrious morphologist, recognized at least four different types. Their disagreement erupted into the public sphere when Geoffroy in 1830 endorsed the view that the ventral nerve cord of invertebrates is directly comparable (today we say 'homologous') to the spinal cord of vertebrates. Cuvier responded that Geoffroy was speculating far beyond the available data, and he reasserted publicly that the major types of animals could not be linked by intermediate forms or topological transformations. This Cuvier-Geoffroy debate was followed closely by comparative biologists all across Europe, who were already flirting with the idea of biological evolution or, as they called it, the transmutation of species. If Cuvier was right, then evolution was impossible. On the other hand, some of Geoffroy's hypotheses (e.g., his proposal that insect legs correspond to vertebrate ribs) did seem a trifle fanciful. Thus, the Cuvier-Geoffroy debate embodied much of the ambivalence surrounding evolution in the first half of the nineteenth century.
After Darwin offered a plausible mechanism for the transmutation of species, namely, natural selection (Darwin, 1859), the idea of biological evolution took hold and, by extension, Geoffroy's ideas gained currency. Innumerable homologies were sought and, frequently, revealed (Russel, 1916). Most impressive was the discovery of extensive molecular homologies between species that span the metazoan family tree (Schmidt-Rhaesa, 2003). It was striking, for example, to discover that many of the genes critical for early brain development are homologous between insects and vertebrates (Sprecher and Reichert, 2003). Indeed, the invertebrate and vertebrate genes are sometimes functionally interchangeable (Halder et al., 1995; deRobertis and Sasai, 1996). Those discoveries supported Geoffroy's view that all animals were built according to a common plan, which could now be understood to be a common genetic blueprint or 'program' (Gehring, 1996). Indeed, many biologists proceeded to search for molecular genetic homologies that could reveal previously unimagined morphological homologies (Janies and DeSalle, 1999). Geoffroy would have been thrilled. There are, however, problems with the view that animals are all alike.
The most serious problem, in my view, is that homologous genes may sometimes be involved in the development of adult structures that are clearly not homologous (Striedter and Northcutt, 1991). For example, insect wings and vertebrate nervous systems both depend on hedgehog function for normal development, but this does not make neural tubes and insect wings homologous (Bagufia and Garcia-Fernandez, 2003). Instead, findings such as this suggest that evolution tends to work with highly conserved 'master genes' (Gehring, 1996) or, more accurately, tightly knit assemblies of crucial genes (Nilsson, 2004), which it occasionally reshuffles by altering their upstream regulatory elements and/or downstream targets. Evolution is a terrific tinkerer that manages to create novelty from conserved elements. This conclusion echoes Geoffroy's arguments insofar as it acknowledges that ''Nature works constantly with the same materials'' (Geoffroy, 1807), but it does not mesh with the view that evolution built all animals according to a single plan. What we have, then, is at least a partial rapprochement of the positions held by Cuvier and Geoffroy: adult organisms do conform to several different body plans, but they are built by shuffling repeatedly a highly conserved set of genes (Raff, 1996). Therefore, a crucial question for research is how evolutionary changes in networks of develop-mentally important genes influence adult structure and function.
Implicit in the preceding discussion has been the idea that adult species differences arise because of evolutionary changes in development (Garstang, 1922). This idea is commonly accepted now, but, back in the nineteenth century, Haeckel (1889) used to promote its polar opposite, namely, the notion that phylogeny creates ontogeny (see Gould, 1977). Haeckel also promoted the idea that all vertebrates pass through a highly conserved phylotypic stage of embryonic development (Slack et al., 1993). Studies have, however, challenged the phylotypic stage idea by showing that the major groups of vertebrates can be distinguished at all stages of embryogenesis (Richardson et al., 1997). An intriguing aspect of that early embryonic variability is that it consists mainly of differences in the timing of developmental processes (Richardson, 1999). Little is known about the genes that generate those changes in developmental timing (also known as heterochrony), but some of them, at least, are likely to be fairly well conserved across species (Pasquinelli and Ruvkun, 2002). More importantly, the notion that adult diversity is based on evolution changing the temporal relationships of conserved processes represents another reconciliation of Cuvier's insistence on adult diversity with Geoffroy's belief in a common plan. Thus, the field of evolutionary developmental biology (evo-devo for short) has overcome the once so prominent dichotomy between conservation and diversity. Its major challenge now is to discover the mechanistic details of how conserved genes and processes are able to produce such diverse adult animals.
Evo-devo thinking has also invaded neuroscience, but evo-devo neurobiology still emphasizes conservation over diversity. For example, we now have extensive evidence that all vertebrate brains are amazingly similar at very early stages of development (Puelles et al., 2000; Puelles and Rubenstein, 2003). However, we still know very little about how and why brain development diverges in the various vertebrate groups after that early, highly conserved stage or period. Looking beyond vertebrates, we find that insect brain development involves at least some genes that are homologous to genes with similar functions in vertebrates (Sprecher and Reichert, 2003). This is remarkable but does not prove that insects and vertebrates are built according to a common plan - if by that we mean that the various parts of adult insect brains all have vertebrate homologues. For example, the finding that several conserved genes, notably Pax6, are critical to eye development in both invertebrates and vertebrates, does not indicate that all those eyes are built according to a common plan. The crucial question, which we are just beginning to explore, is how the conserved genes are tinkered with (reshuffled, co-opted, or redeployed) to produce very different adult eyes (Zuber et al., 2003; Nilsson, 2004). This, then, seems to be the future of evo-devo neurobiology: to discover how highly conserved developmental genes and processes are used to different ends in different species. As I have discussed, this research program has ancient roots, but it is just now becoming clear.
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