{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,21]],"date-time":"2026-08-21T11:57:11Z","timestamp":1787313431776,"version":"build-2736575974"},"reference-count":223,"publisher":"Oxford University Press (OUP)","issue":"5","license":[{"start":{"date-parts":[[2022,9,16]],"date-time":"2022-09-16T00:00:00Z","timestamp":1663286400000},"content-version":"vor","delay-in-days":41,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000001","name":"National Science Foundation","doi-asserted-by":"publisher","award":["NSF IOS 2141592"],"award-info":[{"award-number":["NSF IOS 2141592"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2022,12,6]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Larger animals studied during ontogeny, across populations, or across species, usually have lower mass-specific metabolic rates than smaller animals (hypometric scaling). This pattern is usually observed regardless of physiological state (e.g., basal, resting, field, and maximally active). The scaling of metabolism is usually highly correlated with the scaling of many life-history traits, behaviors, physiological variables, and cellular\/molecular properties, making determination of the causation of this pattern challenging. For across-species comparisons of resting and locomoting animals (but less so for across populations or during ontogeny), the mechanisms at the physiological and cellular level are becoming clear. Lower mass-specific metabolic rates of larger species at rest are due to (a) lower contents of expensive tissues (brains, liver, and kidneys), and (b) slower ion leak across membranes at least partially due to membrane composition, with lower ion pump ATPase activities. Lower mass-specific costs of larger species during locomotion are due to lower costs for lower-frequency muscle activity, with slower myosin and Ca++ ATPase activities, and likely more elastic energy storage. The evolutionary explanation(s) for hypometric scaling remain(s) highly controversial. One subset of evolutionary hypotheses relies on constraints on larger animals due to changes in geometry with size; for example, lower surface-to-volume ratios of exchange surfaces may constrain nutrient or heat exchange, or lower cross-sectional areas of muscles and tendons relative to body mass ratios would make larger animals more fragile without compensation. Another subset of hypotheses suggests that hypometric scaling arises from biotic interactions and correlated selection, with larger animals experiencing less selection for mass-specific growth or neurolocomotor performance. An additional third type of explanation comes from population genetics. Larger animals with their lower effective population sizes and subsequent less effective selection relative to drift may have more deleterious mutations, reducing maximal performance and metabolic rates. Resolving the evolutionary explanation for the hypometric scaling of metabolism and associated variables is a major challenge for organismal and evolutionary biology. To aid progress, we identify some variation in terminology use that has impeded cross-field conversations on scaling. We also suggest that promising directions for the field to move forward include (1) studies examining the linkages between ontogenetic, population-level, and cross-species allometries; (2) studies linking scaling to ecological or phylogenetic context; (3) studies that consider multiple, possibly interacting hypotheses; and (4) obtaining better field data for metabolic rates and the life history correlates of metabolic rate such as lifespan, growth rate, and reproduction.<\/jats:p>","DOI":"10.1093\/icb\/icac136","type":"journal-article","created":{"date-parts":[[2022,8,6]],"date-time":"2022-08-06T20:59:00Z","timestamp":1659819540000},"page":"1395-1418","source":"Crossref","is-referenced-by-count":38,"title":["White Paper: An Integrated Perspective on the Causes of Hypometric Metabolic Scaling in Animals"],"prefix":"10.1093","volume":"62","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-5223-216X","authenticated-orcid":false,"given":"Jon F","family":"Harrison","sequence":"first","affiliation":[{"name":"School of Life Sciences, Arizona State University , Tempe, AZ 85287-4501 , USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Andrew","family":"Biewener","sequence":"additional","affiliation":[{"name":"Department of Organismic and Evolutionary Biology, Harvard University , Cambridge, MA 02138 , USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Joanna R","family":"Bernhardt","sequence":"additional","affiliation":[{"name":"Department of Zoology, University of British Columbia , Vancouver, BC V6T 1Z4 , Canada"},{"name":"Yale Institute for Biospheric Studies , New Haven, CT 06520 , USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Joseph R","family":"Burger","sequence":"additional","affiliation":[{"name":"Department of Biology, University of Kentucky , Lexington, KY 40506 , USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"James H","family":"Brown","sequence":"additional","affiliation":[{"name":"Center for Evolutionary and Theoretical Immunology, The University of New Mexico , Albuquerque, NM 87131 , USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zach N","family":"Coto","sequence":"additional","affiliation":[{"name":"Department of Biology, Boston University , Boston, MA 02215 , USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Meghan E","family":"Duell","sequence":"additional","affiliation":[{"name":"Department of Biology, The University of Western Ontario , London, ON N6A 3K7 , Canada"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Michael","family":"Lynch","sequence":"additional","affiliation":[{"name":"Biodesign Center for Mechanisms of Evolution, Arizona State University , Tempe, AZ 85281 , USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9891-6217","authenticated-orcid":false,"given":"Emma R","family":"Moffett","sequence":"additional","affiliation":[{"name":"Department of Ecology and Evolution, University of California , Irvine, CA 92697 , USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4323-7254","authenticated-orcid":false,"given":"Tommy","family":"Norin","sequence":"additional","affiliation":[{"name":"DTU Aqua | National Institute of Aquatic Resources, Technical University of Denmark , Anker Engelunds Vej 1 Bygning 101A, 2800 Kgs. 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