Biology & Philosophy

, Volume 28, Issue 5, pp 719-745

First online:

More on how and why: cause and effect in biology revisited

  • Kevin N. LalandAffiliated withSchool of Biology, University of St Andrews Email author 
  • , John Odling-SmeeAffiliated withMansfield College, University of Oxford
  • , William HoppittAffiliated withSchool of Biology, University of St Andrews
  • , Tobias UllerAffiliated withDepartment of Zoology, Edward Grey Institute, University of Oxford

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In 1961, Ernst Mayr published a highly influential article on the nature of causation in biology, in which he distinguished between proximate and ultimate causes. Mayr argued that proximate causes (e.g. physiological factors) and ultimate causes (e.g. natural selection) addressed distinct ‘how’ and ‘why’ questions and were not competing alternatives. That distinction retains explanatory value today. However, the adoption of Mayr’s heuristic led to the widespread belief that ontogenetic processes are irrelevant to evolutionary questions, a belief that has (1) hindered progress within evolutionary biology, (2) forged divisions between evolutionary biology and adjacent disciplines and (3) obstructed several contemporary debates in biology. Here we expand on our earlier (Laland et al. in Science 334:1512–1516, 2011) argument that Mayr’s dichotomous formulation has now run its useful course, and that evolutionary biology would be better served by a concept of reciprocal causation, in which causation is perceived to cycle through biological systems recursively. We further suggest that a newer evolutionary synthesis is unlikely to emerge without this change in thinking about causation.


Niche construction Nongenetic inheritance Evo-devo Cultural evolution