Abstract
Tinbergen’s classic “On Aims and Methods of Ethology” (Zeitschrift für Tierpsychologie, 20, 1963) proposed four levels of explanation of behavior, which he thought would soon apply to humans. This paper discusses the need for multilevel explanation; Huxley and Mayr’s prior models, and others that followed; Tinbergen’s differences with Lorenz on “the innate”; and Mayr’s ultimate/proximate distinction. It synthesizes these approaches with nine levels of explanation in three categories: phylogeny, natural selection, and genomics (ultimate causes); maturation, sensitive period effects, and routine environmental effects (intermediate causes); and hormonal/metabolic processes, neural circuitry, and eliciting stimuli (proximate causes), as a respectful extension of Tinbergen’s levels. The proposed classification supports and builds on Tinbergen’s multilevel model and Mayr’s ultimate/proximate continuum, adding intermediate causes in accord with Tinbergen’s emphasis on ontogeny. It requires no modification of Standard Evolutionary Theory or The Modern Synthesis, but shows that much that critics claim was missing was in fact part of Neo-Darwinian theory (so named by J. Mark Baldwin in The American Naturalist in 1896) all along, notably reciprocal causation in ontogeny, niche construction, cultural evolution, and multilevel selection. Updates of classical examples in ethology are offered at each of the nine levels, including the neuroethological and genomic findings Tinbergen foresaw. Finally, human examples are supplied at each level, fulfilling his hope of human applications as part of the biology of behavior. This broad ethological framework empowers us to explain human behavior—eventually completely—and vindicates the idea of human nature, and of humans as a part of nature.
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Acknowledgments
I am grateful to Marjorie Shostak, Dora Venit, Gerald Henderson, Herbert Perluck, Irven DeVore, Richard Lee, Jerome Kagan, Ernst Mayr, Jane Lancaster, Walter Abelmann, S. Boyd Eaton, and Irenäus Eibl-Eibesfeldt for early encouragement. Emory University has been my welcoming academic home for four decades, and its students have honed my thinking about levels of explanation. Ann Cale Kruger taught me much of what I know about human ontogeny. Sarah Blaffer Hrdy, James Chisholm, Randolph Nesse, James Rilling, and Louis Alvarado made helpful comments on earlier versions of this paper, as did three very thoughtful anonymous reviewers. It is dedicated to Nicholas G. Blurton Jones, mentor and friend, who introduced me to Tinbergen in more ways than one.
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Konner, M. Nine Levels of Explanation. Hum Nat 32, 748–793 (2021). https://doi.org/10.1007/s12110-021-09414-8
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DOI: https://doi.org/10.1007/s12110-021-09414-8