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The diving reflex and asphyxia: working across species in physiological ecology

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Abstract

Beginning in the mid-1930s the comparative physiologists Laurence Irving and Per Fredrik (Pete) Scholander pioneered the study of diving mammals, particularly harbor seals. Although resting on earlier work dating back to the late nineteenth century, their research was distinctive in several ways. In contrast to medically oriented physiology, the approaches of Irving and Scholander were strongly influenced by natural history, zoology, ecology, and evolutionary biology. Diving mammals, they argued, shared the cardiopulmonary physiology of terrestrial mammals, but evolution had modified these basic adaptive processes in extreme ways. In particular, seals’ remarkable ability to hold breath, lower metabolism, produce energy anaerobically, and resist asphyxiation, provided a sharp contrast with terrestrial mammals, including humans. This diving physiology was an extreme elaboration of a general regulatory mechanism that allowed seals and other diving mammals to remain active underwater for extended periods. The decrease in heart rate referred to as bradycardia or the “diving reflex” was highly developed in diving mammals, but also found in less developed form in many other organisms faced by asphyxia. It therefore served as a kind of “master switch” for lowering metabolism in diving, hibernation, parturition, drowning, and other physiological responses involving lack of oxygen. Studying bradycardia unified a wide diversity of physiological phenomena, while also providing a context for contrasting the physiological responses of various species, including humans. Conducted in the laboratory and the field, this research served as a bridge between a comparative physiological ecology focused on non-human species and a human-centered general physiology.

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Notes

  1. Scholander directed his own institute of zoophysiology at the University of Oslo from 1955 to 1958; see his CV in box 1, folder 1 of the Per Scholander Collection, Archives, University of California at San Diego. Irving planned and directed a laboratory of zoophysiology modeled on Krogh’s institute when he moved to the University of Alaska in 1962 (Dawson 2007). Plans for the zoophysiology lab are described in a proposal submitted by Irving to the U.S. National Science Foundation, July 31, 1963, series 2, box 18, folder 202, Laurence Irving Collection, University of Alaska Archives.

  2. See letters from Krogh to Irving concerning Scholander’s fellowship dated May 31, 1939 and November 25, 1939 in series 2, box 3, folder 11, Irving Collection.

  3. See correspondence between Irving and Swarthmore president, Frank Aydellotte, leading to the official offer of a contract to Irving on June 4, 1937, series 2, box 3, folder 9, Irving Collection.

  4. Letter from Irving to Krogh, February 3, 1938, series 2, box 3, folder 11, Irving Collection. The lecture series aimed at both professional scholars and the general public, brought international figures from business, politics, arts, the humanities, and sciences to the Swarthmore campus.

  5. The cordial professional and personal relationship between Krogh and Irving had important and far-reaching consequences for the later development of physiological ecology. After World War II Irving also hired Krogh’s daughter Bodil and her husband Knut Schmidt-Nielsen, who was a student of Krogh, as post-doctoral researchers (Hagen 2015). Scholander and the Schmidt-Nielsens became influential leaders in the development of post-war physiological ecology that flourished in the United States (Elsner 2000; Dawson 2007).

  6. During this same period, Bert was also conducting experiments on various animals (including humans) in barometric chambers to simulate the effects of high altitude. Vanessa Heggie (2013) provides a detailed account of how these early studies stimulated later biomedical experiments on Mount Everest, but also the criticism that physiologists and mountaineers directed at the artificiality of Bert’s pioneering research in the laboratory.

  7. Irving seems to have taken it for granted that breathing oxygen gave the Japanese swimmers an advantage, but he was primarily interested in the similarities with the equally artificial practice of forced breathing that all competitive swimmers used before a race. This behavior was in marked contrast to the natural behavior of seals and other diving mammals, which exhaled before going underwater.

  8. Irving traced the history of this observation to Robert Boyle who suggested that the ability of newborn mammals to tolerate anoxia was a requirement for live birth.

  9. Although Scholander and Iriving described these experiments as “dives” not all of the sloths were fully submerged. In some cases, the scientists prevented breathing by clamping the hose connected to an air mask sealed to the face of the experimental animal. This latter procedure was necessary because some of the sloths continued to try to breath underwater. In contrast to the docile three toed-sloths (Bradypus tridactylus) which made good research subjects, two-toed sloths (Choloepus hoffmanni) struggled when submerged and could stay underwater for only a short time.

  10. Letter from Revelle to Scholander, June 5, 1957, box 2, folder 24, Scholander Collection.

  11. See Scholander’s report to Revelle and the advisory board for the Scripps Institution of Oceanography Division of Physiology, May 13, 1963, Box 4, Scholander Collection.

  12. Despite the reference to pearl divers in the title of the article, the subjects in the study actually dived for trochus shells which were used to make buttons.

  13. Despite the late date of this article, the previously unpublished data had been collected by Irving and Scholander in 1940. At the time of publication, Irving was not actively involved in diving research.

  14. See memo from Elsner to Scholander, Dec. 27, 1965, box 6, folder 11, Scholander Collection.

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  1. Department of Biology, Radford University, Radford, VA, 24142, USA

    Joel B. Hagen

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Hagen, J.B. The diving reflex and asphyxia: working across species in physiological ecology. HPLS 40, 18 (2018). https://doi.org/10.1007/s40656-018-0188-z

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