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Entrenchment as a Theoretical Tool in Evolutionary Developmental Biology

  • Oct 9, 2014
  • William C Wimsatt
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Abstract

Inferences from dependencies in development, or from conserved traits, structures, or genes, have become a central and ubiquitous tool in evolutionary developmental biology. An element with many downstream consequences of its operation in an adaptive system should be conserved because its deletion or change would likely have massively deleterious effects. Thus, substantial observed dependencies should predict conservation, and vice versa. Dependencies and conservation are both matters of degree, generating an ordering of rates of evolutionary change. I trace four independent sources for this idea with subtle differences in their application. The origins of the use of genetic pleiotropy to predict conservation is complex and progressed in multiple stages, with partial awareness emerging in the 1960s, and grew erratically in use without becoming a major theme until the 1990s. Since 2000 it has grown to become an essential tool and applied in an expanding number of ways. I distinguish and illustrate “top-down” and “bottom up” approaches, though people now working from one direction often recruit collaborators from the other. Other approaches have had diverse inspirations and points of initial application. Rupert Riedl’s notion of “burden” was applied primarily to morphological traits, with an elaborated account of organizational factors modulating entrenchment. Wallace Arthur explored dependency in “morphogenetic trees,” utilizing as his inspiration cellular descent trees in metazoans. Bill Wimsatt looked for “generative entrenchment” in factors both internal and external to the developing system to analyze innate behavior. Entrenchment was later extended to analyze development in Drosophila and to construct multi-locus population genetic models of gene control networks, yielding new results that illuminate relations between micro and meso-evolution. Applications of generative entrenchment to cultural and technological evolution are also sketched, where development of individuals and other social entities must play a central role.

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