Developing the genotype-to-phenotype relationship in evolutionary theory: A primer of developmental features

Emilie C. Snell-Rood, Sean M. Ehlman

Research output: Contribution to journalArticlepeer-review

1 Scopus citations


For decades, there have been repeated calls for more integration across evolutionary and developmental biology. However, critiques in the literature and recent funding initiatives suggest this integration remains incomplete. We suggest one way forward is to consider how we elaborate the most basic concept of development, the relationship between genotype and phenotype, in traditional models of evolutionary processes. For some questions, when more complex features of development are accounted for, predictions of evolutionary processes shift. We present a primer on concepts of development to clarify confusion in the literature and fuel new questions and approaches. The basic features of development involve expanding a base model of genotype-to-phenotype to include the genome, space, and time. A layer of complexity is added by incorporating developmental systems, including signal-response systems and networks of interactions. The developmental emergence of function, which captures developmental feedbacks and phenotypic performance, offers further model elaborations that explicitly link fitness with developmental systems. Finally, developmental features such as plasticity and developmental niche construction conceptualize the link between a developing phenotype and the external environment, allowing for a fuller inclusion of ecology in evolutionary models. Incorporating aspects of developmental complexity into evolutionary models also accommodates a more pluralistic focus on the causal importance of developmental systems, individual organisms, or agents in generating evolutionary patterns. Thus, by laying out existing concepts of development, and considering how they are used across different fields, we can gain clarity in existing debates around the extended evolutionary synthesis and pursue new directions in evolutionary developmental biology. Finally, we consider how nesting developmental features in traditional models of evolution can highlight areas of evolutionary biology that need more theoretical attention.

Original languageEnglish (US)
Pages (from-to)393-409
Number of pages17
JournalEvolution and Development
Issue number6
StatePublished - Nov 2023

Bibliographical note

Funding Information:
Many thanks to Olof Leimar, Thomas Flatt, Tobias Uller, Bernhard Voelkl, Susan Foster, John Baker, Sonia Sultan, and other participants in the 2015 “Evolving Plasticity” workshop at the Center for Interdisciplinary Research (ZIF) in Bielefeld, Germany for the discussions that led to the initial seeds of this review. Members of University of Minnesota's Biology Interest Group, the Snell‐Rood lab, and UMN's Developmental Biology Center provided valuable insights on manuscript drafts. Alan Love, Pim Edelaar, David Zarkower, and Ruth Shaw gave important feedback on earlier versions of the framework and manuscript. Insightful comments from the editor and two anonymous reviewers greatly improved the review. SME was supported in part by an NSF PRFB (grant number #1811708) and by the Deutsche Forschungsgemeinschaft under German's Excellence Strategy EXC 2002/1, “Science of Intelligence” (project number #390523135). We thank the journal for the opportunity to contribute to this special issue and the Konrad Lorenz Institute for Evolution and Cognition Research for hosting the Agency in Living Systems workshop June 2022 which helped to shape this manuscript.

Publisher Copyright:
© 2023 The Authors. Evolution & Development published by Wiley Periodicals LLC.


  • gene regulatory networks
  • macroevolution
  • plasticity

PubMed: MeSH publication types

  • Journal Article
  • Research Support, Non-U.S. Gov't
  • Research Support, U.S. Gov't, Non-P.H.S.


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