43
Chapter two: Oxygen stores and diving
Forced submersion: An experimental technique in which an animal is constrained and
held underwater for a duration unknown to it. This approach was utilized in early
studies on diving physiology.
Hypoxemia: A low blood O 2 level; in humans, arterial hemoglobin saturations less than
90% are considered hypoxemic; hypoxemic tolerance refers to the ability of an animal or tissue to tolerate and survive such low O 2 levels.
Lung collapse: The process of alveolar collapse and lack of gas exchange at depth secondary to the increase in ambient pressure during a dive; see Chapter 4.
O 2 –Hb dissociation curve: The curve that describes the relationship between hemoglobin saturation and the partial pressure of O 2 (P O2 ).
Peripheral vasoconstriction: The constriction of arterial blood vessels. Such constriction
maintains blood pressure when heart rate and cardiac output decrease.
References
Andrews, R.D., D.R. Jones, J.D. Williams et al. 1997. Heart rates of northern elephant seals diving at
sea and resting on the beach. Journal of Experimental Biology 200:2083–2095.
Baker, D.A. and D.A. Gough. 1995. A continuous, implantable lactate sensor. Analytical Chemistry
67:1536–1540.
Blix, A.S., R.W. Elsner, and J.K. Kjekhus. 1983. Cardiac output and its distribution through capillaries
and A-V shunts in diving seals. Acta Physiologica Scandanavica 118:109–116.
Bryden, M.M. 1972. Body size and composition of elephant seals (Mirounga leonina): Absolute measurements and estimates from bone dimensions. Journal of Zoology (London) 167:265–276.
Burns, J.M., K.C. Lestyk, M.O. Hammill, L.P. Folkow, and A.S. Blix. 2007. Size and distribution of oxygen stores in harp and hooded seals from birth to maturity. Journal of Comparative Physiology
B 177:687–700.
Butler, P.J. and D.R. Jones. 1997. The physiology of diving of birds and mammals. Physiological
Reviews 77:837–899.
Castellini, M.A., G.L. Kooyman, and P.J. Ponganis. 1992. Metabolic rates of freely diving Weddell
seals: Correlations with oxygen stores, swim velocity, and diving duration. Journal of
Experimental Biology 165:181–194.
Cross, J.P., C.G. Mackintosh, and J.F.T. Griffin. 1988. Effect of physical restraint and xylazine sedation
of haemotological values in red deer (Cervus elaphus). Research in Veterinary Science 45:281–286.
Davis, R.W. 2014. A review of the multi-level adaptations for maximizing aerobic dive duration in
marine mammals: From biochemistry to behavior. Journal of Comparative Physiology B 184:23–53.
Davis, R.W., M.A. Castellini, G.L. Kooyman, and R. Maue. 1983. Renal GFR and hepatic blood flow
during voluntary diving in Weddell seals. American Journal of Physiology 245:R743–R748.
Davis, R.W. and T.M. Williams. 2012. The marine mammal dive response is exercise modulated to
maximize aerobic dive duration. Journal of Comparative Physiology A 198:583–591.
El-Sayed, H., S.R. Goodall, and R. Hainsworth. 1995. Re-evaluation of Evans Blue dye dilution
method of plasma volume measurements. Clinical and Laboratory Haematology 17:189–194.
Elsner, R. 1965. Heart rate response in forced versus trained experimental dives of pinnipeds.
Hvalradets Skrifter 48:24–29.
Elsner, R. 2015. Diving Seals and Meditating Yogis: Strategic Metabolic Retreats. Chicago, IL: Universtiy
of Chicago Press.
Elsner, R., D.L. Franklin, R.L. Van Citters, and D.W. Kenney. 1966. Cardiovascular defense against
asphyxia. Science 153:941–949.
Elsner, R., J.T. Shurley, D.D. Hammond, and R.E. Brooks. 1970. Cerebral tolerance to hypoxemia in
asphyxiated Weddell seals. Respiration Physiology 9:287–297.
Fahlman, A., S.H. Loring, M. Ferrigno et al. 2011. Static inflation and deflation pressure–volume
curves from excised lungs of marine mammals. Journal of Experimental Biology 214:3822–3828.
Gallivan, G.J., J.W. Kanwisher, and R.C. Best. 1986. Heart rates and gas exchange in the Amazonian
manatee (Trichecus manutus) in relation to diving. Journal of Comparative Physiology B 156:415–423.
Chapter two: Oxygen stores and diving
Forced submersion: An experimental technique in which an animal is constrained and
held underwater for a duration unknown to it. This approach was utilized in early
studies on diving physiology.
Hypoxemia: A low blood O 2 level; in humans, arterial hemoglobin saturations less than
90% are considered hypoxemic; hypoxemic tolerance refers to the ability of an animal or tissue to tolerate and survive such low O 2 levels.
Lung collapse: The process of alveolar collapse and lack of gas exchange at depth secondary to the increase in ambient pressure during a dive; see Chapter 4.
O 2 –Hb dissociation curve: The curve that describes the relationship between hemoglobin saturation and the partial pressure of O 2 (P O2 ).
Peripheral vasoconstriction: The constriction of arterial blood vessels. Such constriction
maintains blood pressure when heart rate and cardiac output decrease.
References
Andrews, R.D., D.R. Jones, J.D. Williams et al. 1997. Heart rates of northern elephant seals diving at
sea and resting on the beach. Journal of Experimental Biology 200:2083–2095.
Baker, D.A. and D.A. Gough. 1995. A continuous, implantable lactate sensor. Analytical Chemistry
67:1536–1540.
Blix, A.S., R.W. Elsner, and J.K. Kjekhus. 1983. Cardiac output and its distribution through capillaries
and A-V shunts in diving seals. Acta Physiologica Scandanavica 118:109–116.
Bryden, M.M. 1972. Body size and composition of elephant seals (Mirounga leonina): Absolute measurements and estimates from bone dimensions. Journal of Zoology (London) 167:265–276.
Burns, J.M., K.C. Lestyk, M.O. Hammill, L.P. Folkow, and A.S. Blix. 2007. Size and distribution of oxygen stores in harp and hooded seals from birth to maturity. Journal of Comparative Physiology
B 177:687–700.
Butler, P.J. and D.R. Jones. 1997. The physiology of diving of birds and mammals. Physiological
Reviews 77:837–899.
Castellini, M.A., G.L. Kooyman, and P.J. Ponganis. 1992. Metabolic rates of freely diving Weddell
seals: Correlations with oxygen stores, swim velocity, and diving duration. Journal of
Experimental Biology 165:181–194.
Cross, J.P., C.G. Mackintosh, and J.F.T. Griffin. 1988. Effect of physical restraint and xylazine sedation
of haemotological values in red deer (Cervus elaphus). Research in Veterinary Science 45:281–286.
Davis, R.W. 2014. A review of the multi-level adaptations for maximizing aerobic dive duration in
marine mammals: From biochemistry to behavior. Journal of Comparative Physiology B 184:23–53.
Davis, R.W., M.A. Castellini, G.L. Kooyman, and R. Maue. 1983. Renal GFR and hepatic blood flow
during voluntary diving in Weddell seals. American Journal of Physiology 245:R743–R748.
Davis, R.W. and T.M. Williams. 2012. The marine mammal dive response is exercise modulated to
maximize aerobic dive duration. Journal of Comparative Physiology A 198:583–591.
El-Sayed, H., S.R. Goodall, and R. Hainsworth. 1995. Re-evaluation of Evans Blue dye dilution
method of plasma volume measurements. Clinical and Laboratory Haematology 17:189–194.
Elsner, R. 1965. Heart rate response in forced versus trained experimental dives of pinnipeds.
Hvalradets Skrifter 48:24–29.
Elsner, R. 2015. Diving Seals and Meditating Yogis: Strategic Metabolic Retreats. Chicago, IL: Universtiy
of Chicago Press.
Elsner, R., D.L. Franklin, R.L. Van Citters, and D.W. Kenney. 1966. Cardiovascular defense against
asphyxia. Science 153:941–949.
Elsner, R., J.T. Shurley, D.D. Hammond, and R.E. Brooks. 1970. Cerebral tolerance to hypoxemia in
asphyxiated Weddell seals. Respiration Physiology 9:287–297.
Fahlman, A., S.H. Loring, M. Ferrigno et al. 2011. Static inflation and deflation pressure–volume
curves from excised lungs of marine mammals. Journal of Experimental Biology 214:3822–3828.
Gallivan, G.J., J.W. Kanwisher, and R.C. Best. 1986. Heart rates and gas exchange in the Amazonian
manatee (Trichecus manutus) in relation to diving. Journal of Comparative Physiology B 156:415–423.