, Volume 48, Issue 6, pp 575–582 | Cite as

The biochemistry of natural fasting at its limits

  • M. A. Castellini
  • L. D. Rea
Multi-author Review Ecological Implications of Metabolic Biochemistry


There are several groups of animals that are adapted for extremely long duration fasting as part of their reproductive cycle. Penguins, bears and seals routinely fast without food or water for months at time. However, they do not ‘starve’, as the biochemical implications of starving are very different from those of successful fasting. There are distinct biochemical adaptations in lipid, carbohydrate and especially protein metabolism that allow these animals to survive. It appears, at least for penguins and seals, that the duration of the fast may be limited by changes that occur in biochemical regulation near the end of the fast. In all of these species, the biochemistry of fasting and the ecological and behavioral demands of their breeding cycles are closely interrelated.

Key words

Starvation penguins seals marine mammals bears 


Unable to display preview. Download preview PDF.

Unable to display preview. Download preview PDF.


  1. 1.
    Adams, S. H., Changes in protein metabolism and water conservation in northern elephant seals pups during the postweaning fast. MS Thesis, University of California, Santa Cruz, CA 1991.Google Scholar
  2. 2.
    Ahlquist, D. A., Nelson, R. A., Steiger, D. L., Jones, J. D. and Ellefson, R. D., Glycerol metabolism in the hibernating black bear. J. comp. Physiol.155 (1984) 75–79.CrossRefGoogle Scholar
  3. 3.
    Bailey, B. A., Downer, R. G. H., Lavigne, D. M., Drolet, G., and Worthy, G. A. J., Changes in fatty acid composition of plasma of the harp seal,Pagophilus groenlandicus during the post-weaning fast. J. comp. Physiol.70(B) (1981) 795–798.Google Scholar
  4. 4.
    Baumber, J., South, F. E., Ferren, L., and Zatman, M. L., A possible basis for periodic arousal during hibernation: accumulation of ketone bodies. Life Sci.10 (1971) 463–467.CrossRefGoogle Scholar
  5. 5.
    Bowen, W. D., Boness, D. J., and Oftedal, O. T., Mass transfer from mother to pup and subsequent mass loss by the weaned pup in the hooded seal,Cystophora cristata. Can. J. Zool.65 (1987) 1–8.Google Scholar
  6. 6.
    Bowen, W. D., Oftedal, O. T., and Boness, D. J., Birth to weaning in four days: remarkable growth in the hooded seal,Cytophora cristata. Can. J. Zool.63 (1985) 2841–2846.Google Scholar
  7. 7.
    Boyd, I. L., and Duck, C. D., Mass changes and metabolism in territorial male antarctic fur seals (Arctocephalus gazella). Physiol. Zool.64(1) (1991) 375–392.Google Scholar
  8. 8.
    Bryden, M. M., and Stokes, G. B., Metabolism of fatty acids in the southern elephant seal (Mirounga leonina). Can. J. Biochem.47(8) (1969) 757–760.PubMedGoogle Scholar
  9. 9.
    Burton, B. T., Human Nutrition. A Textbook of Nutrition in Health and Disease. McGraw-Hill, New York 1976.Google Scholar
  10. 10.
    Cahill, G. F., Herrera, M. G., Morgan, A. P., Soeldner, J. S., Steinke, J., Levy, P. L., Reichard, G. A., and Kipnis, D. M., Hormone-fuel interrelationships during fasting. J. clin. Invest.45(11) (1966) 1751–1769.PubMedGoogle Scholar
  11. 11.
    Castellini, M. A., and Costa, D. P., Relationships between plasma ketones and fasting duration in neonatal elephant seals. Am. J. Physiol.259 (1990) R1086-R1089.PubMedGoogle Scholar
  12. 12.
    Castellini, M. A., Costa, D. P., and Huntley, A. C., Fatty acid metabolism in fasting northern elephant seal pups. J. comp. Physiol.157(B) (1987) 445–449.Google Scholar
  13. 13.
    Cherel, Y., and Le Maho, Y., Five months of fasting in king penguin chicks: Body mass loss and fuel metabolism. Am. J. Physiol.249 (1985) R387-R392.PubMedGoogle Scholar
  14. 14.
    Cherel, Y., Robin, J.-P., and Le Maho, Y., Physiology and biochemistry of long-term fasting in birds. Can. J. Zool.66 (1988) 159–166.Google Scholar
  15. 15.
    Cherel, Y., Stahl, J.-C., and Le Maho, Y., Ecology and physiology of fasting in king penguin chicks. Auk104 (1987) 254–262.Google Scholar
  16. 16.
    Condit, R. S., and Oritz, C. L., The physiological transition from fasting to feeding in weaned elephant seal pups. Mar. Mamm. Sci.3(3) (1987) 207–219.Google Scholar
  17. 17.
    Costa, D. P., Le Boeuf, B. J., Ortiz, C. L., and Huntley, A. C., The energetics of lactation in the northern elephant seal. J. Zool. Lond.209 (1986) 21–33.Google Scholar
  18. 18.
    Costa, D. P., and Ortiz, C. L., Blood chemistry homeostasis during prolonged fasting in northern elephant seals. Am. J. Physiol.242 (1982) R591-R595.PubMedGoogle Scholar
  19. 19.
    Depocas, F., Hart, J. S., and Fisher, H. D., Sea water drinking and water flux in starved and fed harbor seals,Phoca vitulina. Can. J. Physiol. Pharmac.49 (1971) 53–62.Google Scholar
  20. 20.
    Derocher, A. E., Nelson, R. A., Stirling, I., and Ramsay, M. A., Effects of fasting and feeding on serum urea and serum creatinine levels in polar bears. Mar. Mamm. Sci.6(3) (1990) 196–203.Google Scholar
  21. 21.
    Deutsch, C. J., Haley, M. P., and Le Boeuf, B. J., Reproductive effort of male northern elephant seals: estimates from mass loss. Can. J. Zool.68 (1990) 2580–2593.Google Scholar
  22. 22.
    Florant, G. L., Tokuyama, K., and Rintoul, D. A., Carbohydrate and lipid utilization in hibernators, in: Living in the Cold II, pp. 137–145. Eds A. Malan and B. Cangiulhem. John Libbey Eurotext Ltd, London 1989.Google Scholar
  23. 23.
    Friedman, M. I., Tordoff, M. G., and Kare, M. R., (Eds), Chemical Senses, vol. 4. Appetite and Nutrition. Marcel Dekker, New York 1991.Google Scholar
  24. 24.
    Goodman, M. N., Larsen, P. R., Kaplan, M. M., Aoki, T. T., Young, V. R., and Ruderman, N. B., Starvation in the rat. II. Effect of age and obesity on protein sparing and fuel metabolism. Am. J. Physiol.239 (1980) E277-E286.PubMedGoogle Scholar
  25. 25.
    Grande, F., Man under caloric deficiency, in: Handbook of Physiology. Adaptation to the Environment, pp. 911–937. Eds D. B. Dill, E. F. Adolph and C. G. Wilber. American Physiological Society, Washington, D.C. 1964.Google Scholar
  26. 26.
    Groscolas, R., Changes in body mass, body temperature and plasma fuel levels during the natural breeding fast in male and female emperor penguinsAptenodytes forsteri. J. comp. Physiol.156(B) (1986) 521–527.Google Scholar
  27. 27.
    Groscolas, R., Metabolic adaptations to fasting in emperor and king penguins, in: Penguin Biology, pp. 269–296. Eds L. S. Davis and J. T. Darby. Academic Press, New York 1990.Google Scholar
  28. 28.
    Harlow, H. J., Beck, T. D. I., Walters, L. M., and Greenhouse, S. S., Seasonal serum glucose, progesterone, and cortisol levels of black bears (Ursus americanus). Can. J. Zool.68 (1990) 183–187.Google Scholar
  29. 29.
    Henry, C. J. K., Rivers, J. P. W., and Payne, P. R., Protein and energy metabolism in starvation reconsidered. Eur. J. clin. Nutr.42 (1988) 543–549.PubMedGoogle Scholar
  30. 30.
    Hochachka, P. W., and Guppy, M., Metabolic Arrest and the Control of Biological Time. Harvard University Press, Cambridge 1987.Google Scholar
  31. 31.
    Hoffer, L. J., Starvation, in: Modern Nutrition in Health and Disease, pp. 774–794. Eds M. E. Shils and V. R. Young. Lea & Febiger, Philadelphia 1988.Google Scholar
  32. 32.
    Kopple, J. D., Nutrition and the kidney, in: Nutrition, Metabolic and Clinical Applications, pp. 409–457. Ed R. E. Hodges. Plenum Press, New York 1979.Google Scholar
  33. 33.
    Le Boeuf, B. J., Elephant seals, in: The Natural History of Ano Nuevo, pp. 326–374. Eds B. J. Le Boeuf and S. Kaza. Boxwood Press, Pacific Grove, CA 1981.Google Scholar
  34. 34.
    Le Boeuf, B. J., and Peterson, R. S., Social status and mating activity in elephant seals. Science163 (1969) 91–93.Google Scholar
  35. 35.
    Le Maho, Y. The emperor penguin: A strategy to live and breed in the cold. Am. Sci.65 (1977) 680–693.Google Scholar
  36. 36.
    Le Maho, Y., Robin, J.-P., and Cherel, Y., The metabolic features of starvation, in: Comparative Physiology of Environmental Adaptations 2. Adaptations to Extreme Environments, pp. 177–187. Ed P. Dejours. Karger, Basel 1987.Google Scholar
  37. 37.
    Le Maho, Y., Robin, J.-P., and Cherel, Y., Starvation as a treatment for obesity: The need to conserve body protein. NIPS3 (1988) 21–24.Google Scholar
  38. 38.
    Le Maho, Y., Vu Van Kha, H., Koubi, H., Dewasmes, G., Girard, J., Ferre, O. and Cagnard, M., Body composition, energy expenditure, and plasma metabolites in long-term fasting geese. Am. J. Physiol.241 (1981) E342-E354.PubMedGoogle Scholar
  39. 39.
    Linder, M. C., Nutrition and metabolism of proteins, in: Nutritional Biochemistry and Metabolism with Clinical Applications, pp. 51–68. Ed. M. C. Linder. Elsevier, New York 1985.Google Scholar
  40. 40.
    Lundberg, D. A., Nelson, R. A., Wahner, H. W., and Jones, J. D., Protein metabolism in the black bear before and during hibernation. Mayo clin. Proc.51 (1976) 716–722.PubMedGoogle Scholar
  41. 41.
    Lyman, C. P., Willis, J., Malan, A., and Wang, L., Hibernation and Torpor in Mammals and Birds. Academic Press, New York 1982.Google Scholar
  42. 42.
    Nelson, R. A., Urea metabolism in the hibernating black bear. Kid. Intl.13, Suppl. 8 (1978) S177-S179.Google Scholar
  43. 43.
    Nelson, R. A., Protein and fat metabolism in hibernating bears. Fedn Proc.39 (1980) 2955–2958.Google Scholar
  44. 44.
    Nelson, R. A., Black bears and polar bears — Still metabolic marvels. Mayo clin. Proc.62 (1987) 850–853.PubMedGoogle Scholar
  45. 45.
    Nelson, R. A., Beck, T. D. I., and Steiger, D. L., Ratio of serum urea to serum creatinine in wild black bears. Science226 (1984) 841–842.PubMedGoogle Scholar
  46. 46.
    Nelson, R. A., Folk, G. E., Pfeiffer, E. W., Craighead, J. J., Jonkel, C. J., and Steiger, D. L., Behavior, biochemistry, and hibernation in black, grizzly, and polar bears. Int. Conf. Bear Res. Manage.5 (1980) 284–290.Google Scholar
  47. 47.
    Nelson, R. A., Jones, W. D., Wahner, H. W., McGill, D. B., and Code, C. F., Nitrogen metabolism in bears: Urea metabolism in summer starvation and in winter sleep and role of urinary bladder in water and nitrogen conservation. Mayo clin. Proc.50 (1975) 141–146.PubMedGoogle Scholar
  48. 48.
    Nelson, R. A., Wahner, H. W., Jones, J. D., Ellefson, R. D., and Zollman, P. E., Metabolism of bears before, during, and after winter sleep. Am. J. Physiol.224(2) (1973) 491–496.PubMedGoogle Scholar
  49. 49.
    Nicolaidis, S., What determines food intake? The ischymetric theory. NIPS2 (1987) 104–107.Google Scholar
  50. 50.
    Nordoy, E. S., and Blix, A. S., Energy sources in fasting grey seal pups evaluated with computed tomography. Am. J. Physiol.249 (1985) R471-R476.PubMedGoogle Scholar
  51. 51.
    Nordoy, F. S., and Blix, A. S., Glucose and ketone body turnover in fasting grey seal pups. Acta physiol. scand.4 (1991) 565–571.Google Scholar
  52. 52.
    Nordoy, F. S., Ingebretsen, O. C., and Blix, A. S., Depressed metabolism and low protein catabolism in fasting grey seal pups. Acta physiol. scand.139 (1990) 361–369.PubMedGoogle Scholar
  53. 53.
    Oftedal, O. T., Boness, D. J., and Tedman, R. A., The behavior, physiology and anatomy of lactation in the Pinnepedia, in: Current Mammology, pp. 175–245. Ed. H. H. Genoways. Plenum Press, New York 1987.Google Scholar
  54. 54.
    Oomura, Y., Regulation of feeding by neural responses to endogenous factors. NIPS2 (1987) 199–203.Google Scholar
  55. 55.
    Oritsland, N. A., Pasche, A., Markussen, N. H., and Ronald, K., Weight loss and catabolic adaptations to starvation in grey seal pups. Comp. Biochem. Physiol.82(A) (1985) 931–933.CrossRefGoogle Scholar
  56. 56.
    Ortiz, C. L., Costa, D. P., and Le Boeuf, B. J., Water and energy flux in elephant seal pups fasting and under natural conditions. Physiol. Zool.51 (1978) 166–178.Google Scholar
  57. 57.
    Ortiz, C. L., Le Boeuf, B. J., and Costa, D. P., Milk intake of elephant seal pups: an index of parental investment. Am. Nat.124 (1984) 416–422.CrossRefGoogle Scholar
  58. 58.
    Pernia, S. D., Hill, A., and Ortiz, C. L., Urea turnover during prolonged fasting in the northern elephant seal. Comp. Biochem. Physiol.65(B) (1980) 731–734.CrossRefGoogle Scholar
  59. 59.
    Ramsay, M. A., Nelson, R. A., and Stirling, I., Seasonal changes in the ratio of serum urea to creatinine in feeding and fasting polar bears. Can. J. Zool.69 (1991) 298–302.Google Scholar
  60. 60.
    Rea, L. D., and Costa, D. P., Changes in standard metabolism during long-term fasting in northern elephant seal pups (Mirounga angustirostris). Physiol. Zool. (1992) in press.Google Scholar
  61. 61.
    Rea, L. D., and Castellini, M. A., Plasma fuel levels during prolonged fasting in elephant seal pups (Mirounga angustirostris). 3rd IUBS (1991) 163 (Abstract).Google Scholar
  62. 62.
    Reilly, J. J., Adaptations to prolonged fasting in free-living weaned gray seal pups. Am. J. Physiol.260(2) (1991) R267-R272.PubMedGoogle Scholar
  63. 63.
    Reiter, J., Stinson, N. L., and Le Boeuf, B. J., Northern elephant seal development: the transition from weaning to nutritional independence. Behav. Ecol. Sociobiol.3 (1978) 337–367.CrossRefGoogle Scholar
  64. 64.
    Riedman, M., The Pinnipeds. Seals, Sea Lions, and Walruses. University of California Press, Berkeley 1990.Google Scholar
  65. 65.
    Ritter, R. C., Ritter, S., and Barmes, C. D., (Eds), Feeding Behavior. Neural and Humoral Controls, Academic Press, Orlando 1986.Google Scholar
  66. 66.
    Robin, J.-P., Frain, M., Sardet, C., Groscolas, R., and Le Maho, Y., Protein and lipid utilization during long-term fasting in emperor penguins. Am. J. Physiol.254 (1988) R61-R68.PubMedGoogle Scholar
  67. 67.
    Shoden, R. J., and Griffin, W. S., Fundamentals of Clinical Nutrition. McGraw-Hill, New York 1980.Google Scholar
  68. 68.
    Stewart, W. K., and Fleming, L. W., Features of a successful therapeutic fast of 382 days' duration. Post-grad med. J.49 (1973) 203–209.Google Scholar
  69. 69.
    Van Itallie, T. B., and Yang, M. U., Cardiac dysfunction in obese dieters: a potentially lethal complication of rapid, massive weight loss. Am. J. clin. Nutr.39 (1984) 695–702.PubMedGoogle Scholar
  70. 70.
    Williams, J. D., Crocker, D. E., and Costa, D. P., Free fatty acid and ketone levels in post-partum female elephant seals (Mirounga angustirostris). Marine Mammal Society (1991) Abstract.Google Scholar
  71. 71.
    Worthy, G. A. J., Metabolism and growth of young harp and grey seals. Can. J. Zool.65 (1987) 1377–1382.Google Scholar
  72. 72.
    Worthy, G. A. J., and Lavigne, D. M., Energetics of fasting and subsequent growth in weaned harp seal pups,Phoca groenlandica. Can. J. Zool.61(2) (1983) 447–456.Google Scholar
  73. 73.
    Worthy, G. A. J., and Lavigne, D. M., Mass loss, metabolic rate, and energy utilization by harp and gray seal pups during the postweaning fast. Physiol. Zool.60(3) (1987) 352–364.Google Scholar
  74. 74.
    Young, V. R., and Scrimshaw, N. S., The physiology of starvation. Sci. Amer.225(4) (1971) 14–21.PubMedGoogle Scholar

Copyright information

© Birkhäuser Verlag 1992

Authors and Affiliations

  • M. A. Castellini
    • 1
  • L. D. Rea
    • 1
  1. 1.Institute of Marine ScienceUniversity of AlaskaFairbanksUSA

Personalised recommendations