Marine Biology

, Volume 151, Issue 1, pp 277–291 | Cite as

Phylogenetic, ecological, and ontogenetic factors influencing the biochemical structure of the blubber of odontocetes

  • Heather N. KoopmanEmail author
Research Article


To explore ecological, phylogenetic, and developmental factors affecting the structure of blubber in Odontocetes (toothed whales), lipid composition of this specialized adipose tissue was determined in 260 specimens (30 species representing all families except the river dolphins), most of which were collected between 1995 and 2005, from all over the world. In most odontocetes, blubber contained primarily triacylglycerols; the blubber of beaked and sperm whales was dominated by wax esters (WE), exhibiting ontogenetic patterns of deposition. WEs may represent an adaptation to deep diving for marine mammals that do not rely on blubber for stored energy. Fatty acid (FA) composition was stratified through blubber depth, with higher concentrations of dietary FA in the inner and endogenous FA in the outer layers of the blubber. Stratification can be considered a characteristic feature of odontocetes, and is likely the result of differential metabolism through the blubber. Small body size appears to constrain blubber lipid content to be high. Thermal habitat also represents an important selective pressure for blubber composition. Species inhabiting colder waters exhibited both higher lipid content and increased FA stratification in blubber, compared to species from warm/tropical habitats. The isolation of mobilization to inner blubber may permit metabolic enzymes to function without limitation by lower temperatures. The variation in composition and distribution of blubber lipids in odontocetes suggests that different species may have evolved slightly diverse arrays of secondary functions for this specialized tissue as adaptations for specific ecological niches.


Odontocete Blubber Fatty acid Wax ester Lipid Toothed whale Beaked whale Sperm whale Dolphin Porpoise Beluga Narwhal 



The author wishes to thank many individuals and organizations for their generosity in contributing samples and assisting with sample collection: Bill McLellan and Ann Pabst (University of North Carolina Wilmington); John Nicolas (NMFS, Northeast Fisheries Science Center); Charley Potter, Jim Mead, and Dee Allen (National Museum of Natural History, Smithsonian Institution); Thomas Jefferson and Susan Chivers (NMFS, Southwest Fisheries Science Center); Ellie Dickson and Elizabeth Slooten (University of Otago); Koen van Waerebeek and Julio Reyes; and many people from: state stranding networks from Massachusetts to Florida; Virginia Aquarium and Science Center; Hubbs-Sea World, Orlando, Florida; Mote Marine Laboratory, Sarasota, Florida; the Beaufort, North Carolina and Woods Hole, Massachusetts NMFS offices; the Marine Mammal Stranding Network at UNCW; the Duke University Marine Laboratory, Beaufort, North Carolina; the Canadian Department of Fisheries and Oceans; the Grand Manan Whale and Seabird Research Station, New Brunswick, Canada; the fishermen of the Bay of Fundy and Gulf of Maine; and the Alaska Department of Fish and Game. Much of the work carried out in this study was generously supported by Sara Iverson. This manuscript was improved by the comments of Ann Pabst, Sara Iverson, Andy Read, and Andrew Westgate, as well as by discussions with Sue Budge and Ted Cranford; Sue is also thanked for advice with laboratory procedures. All samples were imported into the US and Canada under appropriate and approved Marine Mammal Protection Act, US Fish & Wildlife Service and CITES permits, and all analytical methods complied with current US and Duke University regulations. This study was supported by postgraduate fellowships from the Canadian Natural Sciences and Engineering Research Council (NSERC) and the Duke University Marine Laboratory, as well as grants from the Duke University Graduate School and the Nicholas School of the Environment to the author; the University of North Carolina Wilmington; a Duke University Marine/Freshwater Biomedical Center Feasibility Study grant to Andy Read; and NSERC Research and Equipment Grants to Sara Iverson, Dalhousie University.

Supplementary material

227_2006_489_MOESM1_ESM.doc (71 kb)
Supplementary material


  1. Ackman RG (1991) Application of gas-liquid chromatography to lipid separation and analysis: qualitative and quantitative analysis. In: Perkins EG (ed) Analysis of fats, oils, and lipoproteins. American Oil Chemists’ Society, Champaign, IL, pp 270–300Google Scholar
  2. Ackman RG, Lamothe F (1989) Marine mammals. In: Ackman RG (ed) Marine biogenic lipids, fats, and oils, vol 2. CRC Press, Boca Raton, FL, pp 179–381Google Scholar
  3. Ackman RG, Eaton CA, Jangaard PM (1965) Lipids of the fin whale (Balaenoptera physalus) from north Atlantic waters. Can J Biochem 43:1513–1520CrossRefGoogle Scholar
  4. Ackman RG, Eaton CA, Litchfield CA (1971) Composition of wax esters, triglycerides and diacyl glyceryl ethers in the jaw and blubber fats of the Amazon River dolphin (Inia geoffrensis). Lipids 6:69–77CrossRefGoogle Scholar
  5. Ackman RG, Hingley JH, Eaton CA, Sipos JC, Mitchell ED (1975) Blubber fat deposition in mysticeti whales. Can J Zool 53:1332–1339CrossRefGoogle Scholar
  6. Aguilar A, Borrell A (1990) Patterns of lipid content and stratification in the blubber of fin whales (Balaenoptera physalus). J Mamm 71:544–554CrossRefGoogle Scholar
  7. Bauermeister A, Sargent JR (1979) Wax esters: major metabolites in the marine environment. Trends Biochem Sci 4:209–211CrossRefGoogle Scholar
  8. Beck CA, Iverson SJ, Bowen WD (2005) Blubber fatty acids of gray seals reveal sex differences in the diet of a size-dimorphic marine carnivore. Can J Zool 83:377–388CrossRefGoogle Scholar
  9. Bradshaw CJA, Hindell MA, Best NJ, Phillips KL, Wilson G, Nichols PD (2003) You are what you eat: describing the foraging ecology of southern elephant seals (Mirounga leonina) using blubber fatty acids. Proc Roy Soc Lond B 270:1283–1292CrossRefGoogle Scholar
  10. Budge SM, Iverson SJ (2003) Quantitative analysis of fatty acid precursors in marine samples: direct conversion of wax ester alcohols and dimethylacetals to FAMEs. J Lipid Res 44:1802–1807CrossRefGoogle Scholar
  11. Clarke R, Paliza O, Aguayo A (1988) Sperm whales of the Southeast Pacific, part IV: fatness, food and feeding. In: Pilleri G (ed) Investigations on Cetacea, vol 21. Brain Anatomy Institute, Berne, Switzerland, pp 53–195Google Scholar
  12. Cooper MH (2004) Fatty acid metabolism in marine carnivores: implications for quantitative estimation of predator diets. PhD thesis. Dalhousie University, Halifax, CanadaGoogle Scholar
  13. Costa DP, Ortiz CL (1982) Blood chemistry homeostasis during prolonged fasting in the northern elephant seal. Am J Physiol 242:R591–R595PubMedGoogle Scholar
  14. CRC Press (1975) In: Fasman GD (ed) Handbook of biochemistry and molecular biology, 3rd edn. CRC Press, Cleveland, OHGoogle Scholar
  15. Dahl TM, Lydersen C, Kovacs KM, Falk-Petersen S, Sargent J, Gjertz I, Gulliksen B (2000) Fatty acid composition of the blubber in white whales (Delphinapterus leucas). Polar Biol 23:401–409CrossRefGoogle Scholar
  16. Doidge DW (1990) Integumentary heat loss and blubber distribution in the beluga, Delphinapterus leucas, with comparisons to the narwhal, Monodon monoceros. In: Smith TG, St. Aubin DJ, Geraci JR (eds) Advances in research on the beluga whale, Delphinapterus leucas. Canadian Bulletin of Fisheries and Aquatic Sciences 224, pp 129–140Google Scholar
  17. Elsner R (1999) Living in water: solutions to physiological problems. In: Reynolds JE III, Rommel SA (eds) Biology of Marine Mammals. Smithsonian Institution Press, Washington, DC, pp 73–116Google Scholar
  18. Evans K, Kemper C, Hill M (2001) First records of the spectacled porpoise Phocoena dioptrica in continental Australian waters. Mar Mamm Sci 17:161–169CrossRefGoogle Scholar
  19. Fish FE (2000) Biomechanics and energetics in aquatic and semiaquatic mammals: platypus to whale. Physiol Biochem Zool 73:683–698CrossRefGoogle Scholar
  20. Folch J, Lees M, Sloan-Stanley GH (1957) A simple method for the isolation and purification of total lipids from animal tissues. J Biol Chem 226:497–509Google Scholar
  21. Gales NJ, Burton HR (1987) Ultrasonic measurement of blubber thickness of the southern elephant seal, Mirounga leonina (Linn). Aust J Zool 35:207–217CrossRefGoogle Scholar
  22. Geraci JR, Lounsbury VJ (1993) Marine mammals ashore: a field guide for strandings. Texas A&M, Galveston TX, Sea Grant Publication TAMU-SG-93-601Google Scholar
  23. Gurr MI, James AT (1975) Lipid biochemistry: an introduction, 2nd edn. Chapman and Hall, LondonGoogle Scholar
  24. Hadley NF (1985) The adaptive role of lipids in biological systems. Wiley, New YorkGoogle Scholar
  25. Haeckel E (1896) The evolution of man: a popular exposition of the principal points of human ontogeny and phylogeny. Appleton, New YorkCrossRefGoogle Scholar
  26. Henderson RJ, Kalogeropoulos N, Alexis MN (1994) The lipid composition of selected tissues from a Mediterranean monk seal, Monachus monachus. Lipids 29:577–582CrossRefGoogle Scholar
  27. Heyning JE, Lento GM (2002) The evolution of marine mammals. In: Hoelzel AR (ed) Marine Mammal Biology: an evolutionary approach. Blackwell Publishing, Durham UK, pp 38–72Google Scholar
  28. Iverson SJ (1988) Composition, intake, and gastric digestion of milk lipids in pinnipeds. PhD thesis, University of Maryland, College Park, MDGoogle Scholar
  29. Iverson SJ (1993) Milk secretion in marine mammals in relation to foraging: can milk fatty acids predict diet? Symp Zool Soc Lond 66:263–291Google Scholar
  30. Iverson SJ, Frost KJ, Lowry LF (1997) Fatty acid signatures reveal fine scale structure of foraging distribution of harbor seals and their prey in Prince William Sound, Alaska. Mar Ecol Prog Ser 151:255–271CrossRefGoogle Scholar
  31. Iverson SJ, Field C, Bowen WD, Blanchard W (2004) Quantitative fatty acids signature analysis: a new method of estimating predator diets. Ecol Mon 74:211–235CrossRefGoogle Scholar
  32. Johnson M, Madsen PT, Zimmer WMX, de Soto NA, Tyack PL (2004) Beaked whales echolocate on prey. Proc R Soc B 271:S383–S386CrossRefGoogle Scholar
  33. Katona S, Whitehead H (1988) Are Cetacea ecologically important? Oceanogr Mar Biol 26:553–568Google Scholar
  34. Kastelein RA, van Battum R (1990) The relationship between body weight and morphological measurements in harbour porpoises (Phocoena phocoena) from the North Sea. Aquat Mamm 16:48–52Google Scholar
  35. Kirsch PE, Iverson SJ, Bowen WD (2000) Effect of a low-fat diet on body composition and blubber fatty acids of captive juvenile harp seals (Phoca groenlandica). Physiol Biochem Zool 73:45–59CrossRefGoogle Scholar
  36. Koopman HN (2001) The structure and function of the blubber of odontocetes. PhD thesis, Nicholas School of the Environment, Duke University, Durham, NC, USAGoogle Scholar
  37. Koopman HN, Iverson SJ, Gaskin DE (1996) Stratification and age-related differences in blubber fatty acids of the male harbour porpoise (Phocoena phocoena). J Comp Physiol B 165:628–639CrossRefGoogle Scholar
  38. Koopman HN, Pabst DA, McLellan WA, Dillaman RM, Read AJ (2002) Changes in blubber distribution and morphology associated with starvation in the harbour porpoise (Phocoena phocoena): evidence for regional differences in blubber structure and function. Physiol Biochem Zool 75:498–512CrossRefGoogle Scholar
  39. Koopman HN, Iverson SJ, Read AJ (2003) High concentrations of isovaleric acid in the fats of odontocetes: variation and patterns of accumulation in blubber vs. stability in the melon. J Comp Physiol B 173:247–261PubMedGoogle Scholar
  40. Koopman HN, Budge SM, Ketten DR, Iverson SJ (2006) The topographical distribution of lipids inside the mandibular fat bodies of odontocetes: remarkable complexity and consistency. IEEE J Ocean Eng 31:95–106CrossRefGoogle Scholar
  41. Krahn MM, Herman DP, Ylitalo GM, Sloan CA, Burrows DG, Hobbs RC, Mahoney BA, Yanagida GK, Calambokidis J, Moore SA (2004) Stratification of lipids, fatty acids and organochlorine contaminants in blubber of white whales and killer whales. J Cetacean Res Manag 6:175–189Google Scholar
  42. Käkelä R, Hyvärinen H (1996) Site-specific fatty acid composition in adipose tissues of several northern aquatic and terrestrial mammals. Comp Biochem Physiol B 115:501–514CrossRefGoogle Scholar
  43. Käkelä R, Hyvärinen H, Vainiotalo P (1993) Fatty acid composition in liver and blubber of the Saimaa ringed seal (Phoca hispida saimensis) compared with that of the ringed seal (Phoca hispida botnica) and grey seal (Halichoerus grypus) from the Baltic. Comp Biochem Physiol B 105:553–565CrossRefGoogle Scholar
  44. Leatherwood S, Reeves RR (1983) The Sierra Club handbook of whales and dolphins. Sierra Club Books, San Francisco, CAGoogle Scholar
  45. Lee RF, Patton JS (1989) Alcohol and waxes. In: Ackman RG (ed) Marine biogenic lipids, fats and oils, vol 1. CRC Press, Boca Raton, FL, pp 73–102Google Scholar
  46. Litchfield C, Greenberg AJ, Caldwell DK, Caldwell MC, Sipos JC, Ackman RG(1975) Comparative lipid patterns in acoustical and nonacoustical fatty tissues of dolphins, porpoises and toothed whales. Comp Biochem Physiol B 50:591–597CrossRefGoogle Scholar
  47. Litchfield CA, Greenberg AJ, Mead JG (1976) The distinctive character of ziphiidae head and blubber fats. Cetology 23:1–10Google Scholar
  48. Lockyer C (1991) Body composition of the sperm whale, Physeter catodon, with special reference to the possible functions of fat depots. Rit Fisk 12:1–24Google Scholar
  49. Lockyer CH, McConnell LC, Waters TD (1984) The biochemical composition of fin whale blubber. Can J Zool 62:2553–2562CrossRefGoogle Scholar
  50. McLellan WA, Koopman HN, Rommel SA, Read AJ, Potter CW, Nicolas JR, Westgate AJ, Pabst DA (2002) Ontogenetic allometry and body composition of harbour porpoises (Phocoena phocoena L.) from the western north Atlantic. J Zool Lond 257:457–472CrossRefGoogle Scholar
  51. Mead JF, Alfin-Slater RB, Howton DR, Popják G (1986) Lipids: chemistry, biochemistry and nutrition. Plenum Press, New YorkCrossRefGoogle Scholar
  52. Nevenzel JC (1970) Occurrence, function and biosynthesis of wax esters in marine organisms. Lipids 5:308–319CrossRefGoogle Scholar
  53. Nordøy ES (1995) Do minke whales (Balaenoptera acutorostrata) digest wax esters? Brit J Nutr 74:717–722CrossRefGoogle Scholar
  54. Pabst DA, Rommel SA, McLellan WA (1999) The functional morphology of marine mammals. In: Reynolds JE III, Rommel SA (eds) Biology of Marine Mammals. Smithsonian Institution Press, Washington, DC, pp 15–72Google Scholar
  55. Patton JS, Benson AA (1975) A comparative study of wax ester digestion in fish. Comp Biochem Physiol B 52:111–116CrossRefGoogle Scholar
  56. Place AR (1992) Comparative aspects of lipid digestion and absorption: physiological correlates of wax ester digestion. Am J Physiol 263:R464–R471PubMedGoogle Scholar
  57. Pond CM (1998) The fats of life. Cambridge University Press, Cambridge, UKCrossRefGoogle Scholar
  58. Read AJ, Hohn AA (1995) Life in the fast lane: the life history of harbor porpoises from the Gulf of Maine. Mar Mamm Sci 11:423–440CrossRefGoogle Scholar
  59. Rice DW (1998) Marine mammals of the world: systematics and distribution. Special Publication Number 4, Society for Marine Mammalogy, Lawrence, KSGoogle Scholar
  60. Samuel AM, Worthy GAJ (2004) Variability in fatty acid composition of bottlenose dolphin (Tursiops truncatus) blubber as a function of body site, season and reproductive state. Can J Zool 82:1933–1942CrossRefGoogle Scholar
  61. Sargent JR (1976) The structure, metabolism and function of lipids in marine organisms. In: Malins DC, Sargent JR (eds) Biochemical and biophysical perspectives in marine biology, vol 3. Academic Press, London, pp 149–212Google Scholar
  62. Sargent JR, Lee RF, Nevenzel JC (1976) Marine waxes. In: Kolattukudy PE (eds) Chemistry and biochemistry of natural waxes. Elsevier, Amsterdam, pp 49–91Google Scholar
  63. Savory P (1971) The action of pure pig pancreatic lipase upon esters of long-chain fatty acids and short-chain primary alcohols. Biochim Biophys Acta 248:149–155CrossRefGoogle Scholar
  64. Schreer JF, Kovacs KM (1997) Allometry of diving capacity in air-breathing vertebrates. Can J Zool 75:339–358CrossRefGoogle Scholar
  65. SPSS (1997) SPSS Base 7.5 for Windows user’s guide. SPSS Inc., Chicago, ILGoogle Scholar
  66. Steel RGD, Torrie JH (1980) Principles and procedures of statistics: a biometrical approach, 2nd edn. McGraw-Hill Inc., New YorkGoogle Scholar
  67. Struntz DJ, McLellan WA, Dillaman RM, Blum JE, Kucklick JR, Pabst DA (2004) Blubber development in bottlenose dolphins (Tursiops truncatus). J Morph 259:7–20CrossRefGoogle Scholar
  68. Tabachnick BG, Fidell LS (1996) Using multivariate statistics, 3rd edn. Harper Collins College Publishers, New YorkGoogle Scholar
  69. West GC, Burns JJ, Modafferi M (1979a) Fatty acid composition of blubber from the four species of Bering Sea phocid seals. Can J Zool 57:189–195CrossRefGoogle Scholar
  70. West GC, Burns JJ, Modafferi M (1979b) Fatty acid composition of Pacific walrus skin and blubber fats. Can J Zool 57:1249–1255CrossRefGoogle Scholar
  71. Worthy GAJ, Edwards EF (1990) Morphometric and biochemical factors affecting heat loss in a small temperate cetacean (Phocoena phocoena) and a small tropical cetacean (Stenella attenuata). Physiol Zool 63:432–442CrossRefGoogle Scholar
  72. Worthy GAJ, Morris PA, Costa DP, LeBoeuf BJ (1992) Moult energetics of the northern elephant seal (Mirounga angustirostris). J Zool Lond 227:257–265CrossRefGoogle Scholar

Copyright information

© Springer-Verlag 2006

Authors and Affiliations

  1. 1.Department of Biology & Marine BiologyUniversity of North Carolina WilmingtonWilmingtonUSA

Personalised recommendations