Entrenchment as a Theoretical Tool in Evolutionary Developmental Biology

  • William C. WimsattEmail author
Part of the Boston Studies in the Philosophy and History of Science book series (BSPS, volume 307)


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.


Fitness Class Evolutionary Developmental Biology Population Genetic Model Niche Construction Theory Darwinian Fitness 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.



In addition to the formative influence of many of the people in this, or in the original 1981 Dahlem volume, I would like to acknowledge the help and influence of Alan Love for extended and penetrating editorial commentary, and the pleasures of co-teaching Evo-devo and working together. Günter Wagner pointed me to the work of Frietson Galis when I was in a quandary as to how to trace the growing influence of pleiotropy in assessments of evolutionary conservatism in the last 15 years. Thanks to Frietson herself for many illuminating articles and conversations. Scott Gilbert confirmed and amplified my intuitions on C.H. Waddington. The remarkably constructive and open atmosphere of the “Dahlem revisited” conference in Berlin in July 2010 owes a great deal to the participants, to the charge of its organizers to us, and to the hospitality of the Max Planck Institute for History of Science, and the support of the Minnesota Center for Philosophy of Science and the Konrad Lorenz Institute for the Study of Evolution, Development and Cognition. Finally, the Minnesota Center for Philosophy of Science and the support of my Winton Professorship in the College of Liberal Arts at the University of Minnesota during this period have been invaluable, and the environment as challenging and stimulating as anyone could wish.


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Authors and Affiliations

  1. 1.Department of PhilosophyUniversity of MinnesotaMinneapolisUSA
  2. 2.Department of Philosophy and Committee on Evolutionary BiologyUniversity of ChicagoChicagoUSA

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