{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,10]],"date-time":"2026-06-10T11:17:04Z","timestamp":1781090224952,"version":"3.54.1"},"reference-count":119,"publisher":"MDPI AG","issue":"7","license":[{"start":{"date-parts":[[2023,6,21]],"date-time":"2023-06-21T00:00:00Z","timestamp":1687305600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000001","name":"National Science Foundation","doi-asserted-by":"publisher","award":["2010290"],"award-info":[{"award-number":["2010290"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000001","name":"National Science Foundation","doi-asserted-by":"publisher","award":["205103\/Z\/16\/Z"],"award-info":[{"award-number":["205103\/Z\/16\/Z"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Wellcome Centre for Human Neuroimaging","award":["2010290"],"award-info":[{"award-number":["2010290"]}]},{"name":"Wellcome Centre for Human Neuroimaging","award":["205103\/Z\/16\/Z"],"award-info":[{"award-number":["205103\/Z\/16\/Z"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Entropy"],"abstract":"<jats:p>This paper introduces a variational formulation of natural selection, paying special attention to the nature of \u2018things\u2019 and the way that different \u2018kinds\u2019 of \u2018things\u2019 are individuated from\u2014and influence\u2014each other. We use the Bayesian mechanics of particular partitions to understand how slow phylogenetic processes constrain\u2014and are constrained by\u2014fast, phenotypic processes. The main result is a formulation of adaptive fitness as a path integral of phenotypic fitness. Paths of least action, at the phenotypic and phylogenetic scales, can then be read as inference and learning processes, respectively. In this view, a phenotype actively infers the state of its econiche under a generative model, whose parameters are learned via natural (Bayesian model) selection. The ensuing variational synthesis features some unexpected aspects. Perhaps the most notable is that it is not possible to describe or model a population of conspecifics per se. Rather, it is necessary to consider populations of distinct natural kinds that influence each other. This paper is limited to a description of the mathematical apparatus and accompanying ideas. Subsequent work will use these methods for simulations and numerical analyses\u2014and identify points of contact with related mathematical formulations of evolution.<\/jats:p>","DOI":"10.3390\/e25070964","type":"journal-article","created":{"date-parts":[[2023,6,22]],"date-time":"2023-06-22T02:29:18Z","timestamp":1687400958000},"page":"964","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":31,"title":["A Variational Synthesis of Evolutionary and Developmental Dynamics"],"prefix":"10.3390","volume":"25","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-7984-8909","authenticated-orcid":false,"given":"Karl","family":"Friston","sequence":"first","affiliation":[{"name":"Wellcome Centre for Human Neuroimaging, Institute of Neurology, University College London, London WC1E 6AP, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6232-9096","authenticated-orcid":false,"given":"Daniel A.","family":"Friedman","sequence":"additional","affiliation":[{"name":"Department of Entomology and Nematology, University of California, Davis, Davis, CA 95616, USA"},{"name":"Active Inference Institute, Davis, CA 95616, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1547-8803","authenticated-orcid":false,"given":"Axel","family":"Constant","sequence":"additional","affiliation":[{"name":"Theory and Method in Biosciences, The University of Sydney, Sydney, NSW 2006, Australia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"V. Bleu","family":"Knight","sequence":"additional","affiliation":[{"name":"Active Inference Institute, Davis, CA 95616, USA"},{"name":"Department of Biology, New Mexico State University, Las Cruces, NM 88003, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4812-0744","authenticated-orcid":false,"given":"Chris","family":"Fields","sequence":"additional","affiliation":[{"name":"Allen Discovery Center at Tufts University, Medford, MA 02155, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5108-5743","authenticated-orcid":false,"given":"Thomas","family":"Parr","sequence":"additional","affiliation":[{"name":"Wellcome Centre for Human Neuroimaging, Institute of Neurology, University College London, London WC1E 6AP, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"John O.","family":"Campbell","sequence":"additional","affiliation":[{"name":"Independent Researcher, Victoria, BC V8V 1X4, Canada"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2023,6,21]]},"reference":[{"key":"ref_1","unstructured":"Darwin, C. 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