{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T05:29:12Z","timestamp":1792042152914,"version":"4.4.0"},"reference-count":26,"publisher":"National Academy of Sciences","issue":"50","license":[{"start":{"date-parts":[[2019,11,25]],"date-time":"2019-11-25T00:00:00Z","timestamp":1574640000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000006","name":"DOD | United States Navy | Office of Naval Research","doi-asserted-by":"publisher","award":["N000141912455"],"award-info":[{"award-number":["N000141912455"]}],"id":[{"id":"10.13039\/100000006","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100010868","name":"SU | School of Humanities and Sciences, Stanford University","doi-asserted-by":"publisher","award":["Terman Fellowship"],"award-info":[{"award-number":["Terman Fellowship"]}],"id":[{"id":"10.13039\/100010868","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["www.pnas.org"],"crossmark-restriction":true},"short-container-title":["Proc. Natl. Acad. Sci. U.S.A."],"published-print":{"date-parts":[[2019,12,10]]},"abstract":"<jats:p>\n                    The biology of the blue whale has long fascinated physiologists because of the animal\u2019s extreme size. Despite high energetic demands from a large body, low mass-specific metabolic rates are likely powered by low heart rates. Diving bradycardia should slow blood oxygen depletion and enhance dive time available for foraging at depth. However, blue whales exhibit a high-cost feeding mechanism, lunge feeding, whereby large volumes of prey-laden water are intermittently engulfed and filtered during dives. This paradox of such a large, slowly beating heart and the high cost of lunge feeding represents a unique test of our understanding of cardiac function, hemodynamics, and physiological limits to body size. Here, we used an electrocardiogram (ECG)-depth recorder tag to measure blue whale heart rates during foraging dives as deep as 184 m and as long as 16.5 min. Heart rates during dives were typically 4 to 8 beats min\n                    <jats:sup>\u22121<\/jats:sup>\n                    (bpm) and as low as 2 bpm, while after-dive surface heart rates were 25 to 37 bpm, near the estimated maximum heart rate possible. Despite extreme bradycardia, we recorded a 2.5-fold increase above diving heart rate minima during the powered ascent phase of feeding lunges followed by a gradual decrease of heart rate during the prolonged glide as engulfed water is filtered. These heart rate dynamics explain the unique hemodynamic design in rorqual whales consisting of a large-diameter, highly compliant, elastic aortic arch that allows the aorta to accommodate blood ejected by the heart and maintain blood flow during the long and variable pauses between heartbeats.\n                  <\/jats:p>","DOI":"10.1073\/pnas.1914273116","type":"journal-article","created":{"date-parts":[[2019,11,25]],"date-time":"2019-11-25T20:27:33Z","timestamp":1574713653000},"page":"25329-25332","update-policy":"https:\/\/doi.org\/10.1073\/pnas.cm10313","source":"Crossref","is-referenced-by-count":83,"title":["Extreme bradycardia and tachycardia in the world\u2019s largest animal"],"prefix":"10.1073","volume":"116","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-4170-7294","authenticated-orcid":false,"given":"J. A.","family":"Goldbogen","sequence":"first","affiliation":[{"name":"Hopkins Marine Station, Department of Biology, Stanford University, Pacific Grove, CA 93950;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3641-1242","authenticated-orcid":false,"given":"D. E.","family":"Cade","sequence":"additional","affiliation":[{"name":"Hopkins Marine Station, Department of Biology, Stanford University, Pacific Grove, CA 93950;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"J.","family":"Calambokidis","sequence":"additional","affiliation":[{"name":"Cascadia Research Collective, Olympia, WA 98501;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"M. F.","family":"Czapanskiy","sequence":"additional","affiliation":[{"name":"Hopkins Marine Station, Department of Biology, Stanford University, Pacific Grove, CA 93950;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"J.","family":"Fahlbusch","sequence":"additional","affiliation":[{"name":"Hopkins Marine Station, Department of Biology, Stanford University, Pacific Grove, CA 93950;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"A. S.","family":"Friedlaender","sequence":"additional","affiliation":[{"name":"Institute of Marine Sciences, University of California, Santa Cruz, CA 95064;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"W. T.","family":"Gough","sequence":"additional","affiliation":[{"name":"Hopkins Marine Station, Department of Biology, Stanford University, Pacific Grove, CA 93950;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"S. R.","family":"Kahane-Rapport","sequence":"additional","affiliation":[{"name":"Hopkins Marine Station, Department of Biology, Stanford University, Pacific Grove, CA 93950;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"M. S.","family":"Savoca","sequence":"additional","affiliation":[{"name":"Hopkins Marine Station, Department of Biology, Stanford University, Pacific Grove, CA 93950;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"K. V.","family":"Ponganis","sequence":"additional","affiliation":[{"name":"Scripps Institution of Oceanography, University of California San Diego, La Jolla, CA 92093"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"P. J.","family":"Ponganis","sequence":"additional","affiliation":[{"name":"Scripps Institution of Oceanography, University of California San Diego, La Jolla, CA 92093"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"341","published-online":{"date-parts":[[2019,11,25]]},"reference":[{"key":"e_1_3_4_1_2","doi-asserted-by":"crossref","DOI":"10.1017\/CBO9781139045490","volume-title":"Diving Physiology of Marine Mammals and Seabirds","author":"Ponganis P. J.","year":"2015","unstructured":"P. J. Ponganis, Diving Physiology of Marine Mammals and Seabirds (Cambridge University Press, 2015)."},{"key":"e_1_3_4_2_2","first-page":"1","article-title":"Experimental investigations on the respiratory function in diving mammals and birds","volume":"22","author":"Scholander P. F.","year":"1940","unstructured":"P. F. Scholander, Experimental investigations on the respiratory function in diving mammals and birds. Hvalradets Skrifter 22, 1\u2013131 (1940).","journal-title":"Hvalradets Skrifter"},{"key":"e_1_3_4_3_2","doi-asserted-by":"crossref","first-page":"283","DOI":"10.1002\/jcp.1030180302","article-title":"Significance of the heart rate to the diving ability of seals","volume":"18","author":"Irving L.","year":"1941","unstructured":"L. Irving, P. Scholander, S. Grinnell, Significance of the heart rate to the diving ability of seals. J. Cell. Comp. Physiol. 18, 283\u2013297 (1941).","journal-title":"J. Cell. Comp. Physiol."},{"key":"e_1_3_4_4_2","doi-asserted-by":"crossref","first-page":"6055","DOI":"10.1038\/ncomms7055","article-title":"Exercise at depth alters bradycardia and incidence of cardiac anomalies in deep-diving marine mammals","volume":"6","author":"Williams T. M.","year":"2015","unstructured":"T. M. Williams ., Exercise at depth alters bradycardia and incidence of cardiac anomalies in deep-diving marine mammals. Nat. Commun. 6, 6055 (2015).","journal-title":"Nat. Commun."},{"key":"e_1_3_4_5_2","doi-asserted-by":"crossref","first-page":"583","DOI":"10.1007\/s00359-012-0731-4","article-title":"The marine mammal dive response is exercise modulated to maximize aerobic dive duration","volume":"198","author":"Davis R. W.","year":"2012","unstructured":"R. W. Davis, T. M. Williams, The marine mammal dive response is exercise modulated to maximize aerobic dive duration. J. Comp. Physiol. A Neuroethol. Sens. Neural Behav. Physiol. 198, 583\u2013591 (2012).","journal-title":"J. Comp. Physiol. A Neuroethol. Sens. Neural Behav. Physiol."},{"key":"e_1_3_4_6_2","doi-asserted-by":"crossref","first-page":"e44854","DOI":"10.1371\/journal.pone.0044854","article-title":"Metabolic expenditures of lunge feeding rorquals across scale: Implications for the evolution of filter feeding and the limits to maximum body size","volume":"7","author":"Potvin J.","year":"2012","unstructured":"J. Potvin, J. A. Goldbogen, R. E. 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Goldbogen ., Mechanics, hydrodynamics and energetics of blue whale lunge feeding: Efficiency dependence on krill density. J. Exp. Biol. 214, 131\u2013146 (2011).","journal-title":"J. Exp. Biol."},{"key":"e_1_3_4_18_2","doi-asserted-by":"crossref","first-page":"227","DOI":"10.1111\/j.1748-7692.1986.tb00043.x","article-title":"Estimating feeding rates of marine mammals from heart mass to body mass ratios","volume":"2","author":"Innes S.","year":"1986","unstructured":"S. Innes, D. Lavigne, W. Earle, K. Kovacs, Estimating feeding rates of marine mammals from heart mass to body mass ratios. Mar. Mamm. Sci. 2, 227\u2013229 (1986).","journal-title":"Mar. Mamm. Sci."},{"key":"e_1_3_4_19_2","doi-asserted-by":"crossref","first-page":"453","DOI":"10.1152\/jappl.1967.22.3.453","article-title":"Scaling of respiratory variables in mammals","volume":"22","author":"Stahl W. R.","year":"1967","unstructured":"W. R. Stahl, Scaling of respiratory variables in mammals. J. Appl. Physiol. 22, 453\u2013460 (1967).","journal-title":"J. Appl. Physiol."},{"key":"e_1_3_4_20_2","doi-asserted-by":"crossref","first-page":"704","DOI":"10.1152\/ajplegacy.1968.215.3.704","article-title":"Ventricular volumes and body weight in mammals","volume":"215","author":"Holt J. P.","year":"1968","unstructured":"J. P. Holt, E. A. Rhode, H. Kines, Ventricular volumes and body weight in mammals. Am. J. Physiol. 215, 704\u2013715 (1968).","journal-title":"Am. J. Physiol."},{"key":"e_1_3_4_21_2","doi-asserted-by":"crossref","first-page":"259","DOI":"10.1093\/icesjms\/36.3.259","article-title":"Body weights of some species of large whales","volume":"36","author":"Lockyer C.","year":"1976","unstructured":"C. Lockyer, Body weights of some species of large whales. ICES J. Mar. Sci. 36, 259\u2013273 (1976).","journal-title":"ICES J. Mar. 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Circulation 110, 2802\u20132808 (2004).","journal-title":"Circulation"},{"key":"e_1_3_4_26_2","doi-asserted-by":"crossref","first-page":"263","DOI":"10.1016\/j.cbpb.2004.05.003","article-title":"The diving paradox: New insights into the role of the dive response in air-breathing vertebrates","volume":"138","author":"Davis R. W.","year":"2004","unstructured":"R. W. Davis ., The diving paradox: New insights into the role of the dive response in air-breathing vertebrates. Comp. Biochem. Physiol. A Mol. Integr. Physiol. 138, 263\u2013268 (2004).","journal-title":"Comp. Biochem. Physiol. A Mol. Integr. 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