Studying trophic ecology in marine ecosystems using fatty acids: A primer on analysis and interpretation

Marine Mammal Science

Volumn 22, Issue 4, 2006, Pages 759-801

  Budge, Suzanne M ^a   Iverson, Sara J ^a   Koopman, Heather N ^b  

^a DALHOUSIE UNIVERSITY   (Canada)

^b UNIVERSITY OF NORTH CAROLINA WILMINGTON   (United States)

Author keywords

[No Author keywords available]

Indexed keywords

FATTY ACID; MARINE ECOSYSTEM; QUALITATIVE ANALYSIS; QUANTITATIVE ANALYSIS; TROPHIC STRUCTURE;

EID: 33748930236     PISSN: 08240469     EISSN: 17487692     Source Type: Journal    
DOI: 10.1111/j.1748-7692.2006.00079.x     Document Type: Review

References (142)

1. AAREFJORD, H., A. J. BJØRGE, C. C. KINZE AND I. LINDSTEDT. 1995. Diet of harbour porpoises (Phocoena phocoena) in Scandinavian waters. Report of the International Whaling Commission (Special Issue 16):211-222.
2. ACKMAN, R. G. 1986. WCOT (capillary) gas-liquid chromatography. Pages 137-206 in R. J. Hamilton and J. B. Rossel, eds. Analysis of oils and fats. Elsevier, London, UK.
3. ACKMAN, R. G. 1989. Fatty acids. Pages 103-137 in R. G. Ackman, ed. Marine biogenic lipids, fats, and oils. Volume 1. CRC Press, Boca Raton, FL.
4. ACKMAN, R. G. 1994. Animal and marine lipids. Pages 292-328 in B. S. Kamel and Y. Kakuda, eds. Technological advances in improved and alternative sources of lipids. Chapman and Hall, London, UK.
5. ACKMAN, R. G. 1998. Comparison of lipids in marine and freshwater organisms. Pages 263-298 in M. T. Arts and B. C. Wainman, eds. Lipids in freshwater ecosystems. Springer-Verlag, New York, NY.
6. ACKMAN, R. G., AND C. A. EATON. 1966. Lipids of the fin whale (Balaenoptera physalus) from north Atlantic waters. III. Occurrence of eicosenoic and docosenoic fatty acids in the zooplankter Meganyctiphanes norvegica (M. Sars) and their effect on whale oil composition. Canadian Journal of Biochemistry 44:1561-1566.
7. ACKMAN, R. G., C. A. EATON AND P. M. JANGAARD. 1965. Lipids of the fin whale (Balaenoptera physalus) from North Atlantic waters I. Fatty acid composition of whole blubber and blubber sections. Canadian Journal of Biochemistry 43:1513-1520.
8. ACKMAN, R. G., S. N. HOOPER, S. EPSTEIN AND M. KELLEHER. 1972. Wax esters of barracudina lipid: A potential replacement for sperm whale oil. Journal of the American Oil Chemists' Society 49:378-382.
9. AITCHISON, J. 1986. The statistical analysis of compositional data. Chapman and Hall, New York, NY.
10. BACHORIK, P. S. 1982. Collection of blood samples for lipoprotein analysis. Clinical Chemistry 28:1375-1378.
11. BANDI, Z. L., AND H. K. MANGOLD. 1973. Substrate specificity of enzymes catalyzing interconversions of long-chain acids and alcohols in the rat. FEBS Letters 31:97-100.
12. BECK, C. A., W. D. BOWEN AND S. J. IVERSON. 2003. Sex differences in the seasonal patterns of energy storage and expenditure in a phocid seal. Journal of Animal Ecology 72:280-291.
13. BECK, C. A., S. J. IVERSON AN DW. D. BOWEN. 2005. Blubber fatty acids of gray seals reveal sex differences in the diet of a size-dimorphic marine carnivore. Canadian Journal of Zoology 83:377-388.
14. BEST, N. J., C. J. BRADSHAW, M. A. HINDELL AND P. D. NICHOLS. 2003. Vertical stratification if fatty acids in the blubber of southern elephant seals (Mirounga leonine): Implications for diet analysis. Comparative Biochemistry Physiology B 134:253-263.
15. BISHOP, D. G., D. A. RITZ, G. W. HOSIE, J. R. KENRICK AND J. OLLEY. 1983. Fatty acid composition of the lipids of Puffinus Tenuirostris (Temminck) in relation to its diet. Journal of Experimental Marine Biology and Ecology 71:17-26.
16. BLIGH, E. G., AND W. J. DYER. 1959. Arapid method of total lipid extraction and purification. Canadian Journal of Biochemistry and Physiology 37:911-917.
17. BÖER, M., C. GANNEFORS, G. KATTNER, M. GRAEVE, H. HOP AND S. FALK-PETERSEN. 2005. The arctic pteropod Clione limacine: Seasonal lipid dynamics and life-strategy. Marine Biology 147:707-717.
18. BOWDEN, K., I. M. HEILBRON, E. R. H. JONES AND B. C. WEEDON. 1946. Researches on acetylenic compounds. Part I. The preparation of acetylenic ketones by oxidation of acetylenic carbinols and glycols. Journal of the Chemical Society 39-45.
19. BOWEN, W. D., S. J. IVERSON, D. J. BONESS AND O. T. OFTEDAL. 2001. Foraging effort, food intake, and lactation performance depend on maternal mass in a small phocid seal. Functional Ecology 15:325-334.
20. BOWEN, W. D., C. A. BECK, S. J. IVERSON, D. AUSTIN AND J. I. MCMILLAN. 2006. Linking predator foraging behaviour and diet with variability in continental shelf ecosystems: Grey seals of eastern Canada. Pages 63-81 in I. L. Boyd, S. Wanless and C. J. Camphuysen, eds. Top predators in marine Ecosystems: Their role in monitoring and Management. Cambridge University Press, Cambridge, U.K.
21. BRADSTREET, M. S. W. 1980. Thick-billed murres and black guillemots in the Barrow Strait area, N. W. T., during spring: Diets and food availability along ice edges. Canadian Journal of Zoology 58:2120-2140.
22. BRINDLEY, D. N. 1978. Regulation of fatty acid esterification in tissue. Pages 31-39 in R. Dils and J. Knudsen, eds. Regulation of fatty acid and glycerolipid metabolism. Pergamon, Oxford, UK.
23. BRINDLEY, D. N. 1991. Metabolism of triacylglycerols. Pages 171-203 in D. E. Vance and J. Vance, eds. Biochemistry of lipids, lipoproteins and membranes. Elsevier Science, New York, NY.
24. BROWN, D. J., I. L. BOYD, G. C. CRIPPS AND P. J. BUTLER. 1999. Fatty acid signature analysis from the milk of Antarctic fur seals and southern elephant seals from South Georgia: Implications for diet determination. Marine Ecology Progress Series 187:251-263.
25. BUDGE, S. M., AND S. J. IVERSON. 2003. Quantitative analysis of fatty acid precursors in marine samples: Direct conversion of wax ester alcohols and dimethylacetals to fatty acid methyl esters. Journal of Lipid Research 44:1802-1807.
26. BUDGE, S. M., C. C. PARRISH AND C. H. MCKENZIE. 2001. Fatty acid composition of phytoplankton, settling particulate matter and sediments at a sheltered bivalve aquaculture site. Marine Chemistry 76:285-303.
27. BUDGE, S. M., S. J. IVERSON, W. D. BOWEN AND R. G. ACKMAN. 2002. Among- and within-species variation in fatty acid signatures of marine fish and invertebrates on the Scotian Shelf, Georges Bank and southern Gulf of St. Lawrence. Canadian Journal of Fisheries and Aquatic Sciences 59:886-898.
28. BUDGE, S. M., M. H. COOPER AND S. J. IVERSON. 2004. Demonstration of the deposition and modification of dietary fatty acids in pinniped blubber using radiolabeled precursors. Physiological and Biochemical Zoology 77:682-687.
29. CABRINI, L., L. LANDI, C. STEFANELLI, V. BARZANTI AND A. M. SECHI. 1992. Extraction of lipids and lipophilic antioxidants from fish tissues: A comparison among different methods. Comparative Biochemistry and Physiology B 101:383-386.
30. CANTELLOPS, D., A. P. REID, R. R. EITENMILLER AND A. R. LONG. 1999. Determination of lipids in infant formula powder by direct extraction methylation of lipids and fatty acid methyl esters (FAME) analysis by gas chromatography. Journal of the AOAC International 82:1128-1139.
31. CARROLL, G. M., J. C. GEORGE, L. F. LOWRY AND K. O. COYLE. 1987. Bowhead whale (Balaena mysticetus) feeding near Point Barrow, Alaska, during the 1985 migration. Arctic 40:105-110.
32. CHAPELLE, S., AND A. A. BENSON. 1986. Studies on the plasmalogens of crustacean gills. Comparative Biochemistry and Physiology B 85:507-510.
33. CHAPMAN, M. J., AND J. P. KANE. 1975. Stability of the apoprotein of human serum low density lipoprotein: Absence of endogenous endopeptidase activity. Biochemical and Biophysical Research Communications 66:1030-1036.
34. CHRISTIE, W. W. 1982. Lipid Analysis. Pergamon Press, Oxford, UK.
35. CHRISTIE, W. W. 1989. Gas chromatography and lipids. A practical guide. Oily Press, Ayr, Scotland.
36. CHRISTIE, W. W. 1998. Gas chromatography-mass spectrometry methods for structural analysis of fatty acids. Lipids 33:343-353.
37. CHRISTIE, W. W., E. Y. BRECHANY, S. B. JOHNSON AND R. T. HOLMAN. 1986. A comparison of pyrrolidide and picolinyl ester derivatives for the identification of fatty acids in natural samples by gas chromatography-mass spectrometry. Lipids 21:657-661.
38. COOK, H. W. 1991. Fatty acid desaturation and chain elongation in eucaryotes. Pages 141-169 in D. E. Vance and J. Vance, eds. Biochemistry of lipids, lipoproteins and membranes. Elsevier Science, New York, NY.
39. COOPER, M. H. 2004. Fatty acid metabolism in marine carnivores: Implications for quantitative estimation of predator diets. Ph.D. thesis, Dalhousie University, Halifax, Canada. 228 pp.
40. COOPER, M. H., S. J. IVERSON AND H. HERAS. 2005. Dynamics of blood chylomicron fatty acids in a marine carnivore: Implications for lipid metabolism and quantitative estimation of predator diets. Journal of Comparative Physiology B 175:133-145.
41. COOPER, M. H., S. J. IVERSON AND K. ROUVINEN-WATT. 2006. Metabolism of dietary cetoleic acid (22:1n-11) in mink (Mustela vison) and grey seals (Halichoerus grypus) studied using radiolabelled fatty acids. Physiological and Biochemical Zoology 79:820-826.
42. DAHL, T. M., C. LYDERSEN, K. M. KOVACS, S. FALK-PETERSEN, J. SARGENT, I. GJERTZ AND B. GULLIKSEN. 2000. Fatty acid composition of the blubber in white whales (Delphinapterus leucas). Polar Biology 23:401-409.
43. DAHL, T. M., S. FALK-PETERSEN, J. SARGENT, G. GABRIELSEN, H. HOP AND R. M. MILLAR. 2003. Lipids and stable isotopes in common eider, black-legged kittiwake and northern fulmar - a trophic study from an Arctic fjord. Marine Ecology Progress Series 256:257-269.
44. DALSGAARD, J., M. ST JOHN, G. KATTNER, D. MÜLLER-NAVARRA AND W. HAGEN. 2003. Fatty acid trophic markers in the pelagic marine environment. Advances in Marine Biology 46:225-340.
45. DOIDGE, D. W. 1990. Integumentary heat loss and blubber distribution in the beluga, Delphinapterus leucas, with comparisons to the narwhal, Monodon monoceros. Pages 129-140 in T. G. Smith, D. J. St. Aubin and J. R. Geraci, eds. Advances in research on the beluga whale, Delphinapterus leucas. Canadian Bulletin of Fisheries and Aquatic Sciences, 224.
46. FALK-PETERSEN, S., T. HAUG, K. T. NILSSEN, A. WOLD AND T. M. DAHL. 2004. Lipids and trophic linkages in harp seal (Phoca groenlandica) from the eastern Barents Sea. Polar Research 23:43-50.
47. FOLCH, J., M. LEES AND G. H. SLOANE-STANLEY. 1957. A simple method for the isolation and purification of total lipids from animal tissues. Journal of Biological Chemistry 226:497-509.
48. GERACI, J. R., AND V. J. LOUNSBURY. 1993. Marine mammals ashore: A field guide for strandings. Texas A&M Sea Grant Publication, Galveston, TX.
49. GOEDKOOP, W., L. SONESTEN, G. AHLGREN AND M. BOBERG. 2000. Fatty acids in profundal benthis invertebrates and their major food resources in Lake Erken, Sweden: Seasonal variation and trophic indications. Canadian Journal of Fisheries and Aquatic Sciences 57:2267-2279.
50. GRAHL-NIELSEN, O., AND O. MJAAVATTEN. 1991. Dietary influence on fatty acid composition of blubber fat of seals as determined by biopsy: A multivariate approach. Marine Biology 110:59-64.
51. GROB, R. L. 1995. Modern practices of gas chromatography. Wiley and Sons, New York, NY.
52. GROSCOLAS, R. 1990. Metabolic adaptations to fasting in emperor and king penguins. Pages 269-295 in L. S. Davis and J. T. Darby, eds. Penguin Biology. Acedemic Press, New York, NY.
53. GUILLOU, A., P. SOUCY AND M. KHALIL. 1996. Preparation of fatty acid methyl esters from brook charr tissues: Comparison of a classical and a direct method. Fett Lipid 98:18-21.
54. GUNSTONE, F. D., J. L. HARWOOD AND F. B. PADLEY. 1986. The lipid handbook. Chapman and Hall, London, UK.
55. GURR, M. I., AND J. L. HARWOOD. 1991. Lipid biochemistry: An introduction. Chapman and Hall, New York, NY.
56. HARWOOD, J. L., AND N. J. RUSSELL. 1984. Lipids in plants and microbes. Allen and Unwin, London, UK.
57. HERMAN, D. P., D. G. BURROWS, P. R. WADE, J. W. DURBAN, C. O. MATKIN, R. G. LE DUC, L. G. BARRETT-LENNARD AND M. M. KRAHN. 2005. Feeding ecology of eastern North Pacific killer whales Orcinus orca from fatty acid, stable isotope and organochlorine analyses of blubber biopsies. Marine Ecology Progress Series 302:275-291.
58. HOBSON, K. A. 1993. Trophic relationships among high Arctic seabirds: Insights from tissue-dependent stable-isotope models. Marine Ecology Progress Series 95:7-18.
59. HOOKER, S. K., S. J. IVERSON, P. OSTROM AND S. C. SMITH. 2001. Diet of northern bottlenose whales inferred from fatty-acid and stable-isotope analyses of biopsy samples. Canadian Journal of Zoology 79:1442-1454.
60. HORGAN, I. E., and J. A. BARRETT. 1985. The use of lipid profiles in comparing the diet of seabirds. Pages 493-497 in W. R. Siegfried, P. R. Condy, and R. M. Laws, eds. Antarctic nutrient cycles and food webs. Springer-Verlag, Berlin, Germany.
61. HORROCKS, L. A. 1972. Content, composition, and metabolism of mammalian and avian lipids that contain ether groups. Pages 177-272 in F. Snyder, ed. Ether lipids: Chemistry and biology. Academic Press, New York, NY.
62. IVERSON, J. L., and A. J. SHEPPARD. 1977. Butyl ester preparation for gas-liquid chromatographic determination of fatty acids in butter. Journal of the AOAC 60:284-288.
63. IVERSON, S. J. 1988. Composition, intake and gastric digestion of milk lipids in pinnipeds. Ph.D. thesis, University of Maryland, College Park, MD. 198 pp.
64. IVERSON, S. J. 1993. Milk secretion in marine mammals in relation to foraging: Can milk fatty acids predict diet? Symposium of the Zoological Society of London 66:263-291.
65. IVERSON, S. J. 2002. Blubber. Pages 107-112 in W. F. Perrin, B. Würsig and H. G. M. Thewissen, eds. Encyclopedia of marine mammals. Academic Press, San Diego, CA.
66. IVERSON, S. J., AND O. T. OFTEDAL. 1992. Fatty acid composition of black bear (Ursus americanus) milk during and after the period of winter dormancy. Lipids 27:940-943.
67. IVERSON, S. J., AND A. M. SPRINGER. 2002. Estimating seabird diets using fatty acids: Protocol development and testing of ReFER hypotheses. Report to the National Pacific Marine Research Program. University of Alaska, Fairbanks, AK.
68. IVERSON, S. J., J. SAMPUGNA AND O. T. OFTEDAL. 1992. Positional specificity of gastric hydrolysis of n-3 polyunsaturated fatty acids of seal milk triglycerides. Lipids 27:870-878.
69. IVERSON, S. J., O. T. OFTEDAL, W. D. BOWEN, D. J. BONESS AND J. SAMPUGNA. 1995. Prenatal and postnatal transfer of fatty aicds from mother to pup in the hooded seal. Journal of Comparative Physiology B 165:1-12.
70. IVERSON, S. J., J. P. Y. ARNOULD AND I. L. BOYD. 1997a. Milk fatty acid signatures indicate both major and minor shifts in the diet of lactating Antarctic fur seals. Canadian Journal of Zoology 75:188-197.
71. IVERSON, S. J., K. J. FROST AND L. F. LOWRY. 1997b. Fatty acid signatures reveal fine scale structure of foraging distribution of harbor seals and their prey in Prince William Sound, Alaska. Marine Ecology Progress Series 151:255-271.
72. IVERSON, S. J., S. L. C. LANG AND M. H. COOPER. 2001a. Comparison of the Bligh and Dyer and Folch methods for total lipid determination in a broad range of marine tissue. Lipids 36:1283-1287.
73. IVERSON, S. J., J. E. MCDONALD AND L. K. SMITH. 2001b. Changes in the diet of free ranging black bears in years of contrasting food availability revealed through milk fatty acids. Canadian Journal of Zoology 79:2268-2279.
74. IVERSON, S. J., K. J. FROST AND S. L. C. LANG. 2002. Fat content and fatty acid composition of forage fish and invertebrates in Prince William Sound, Alaska: Factors contributing to among and within species variability. Marine Ecology Progress Series 241:161-181.
75. IVERSON, S. J., B. S. STEWART AND P. K. YOCHEM. 2003. Captive validation and calibration of fatty acid signatures in blubber as indicators of prey in Hawaiian monk seal diet. NOAA Technical Report, San Diego, CA, 22 pp.
76. IVERSON, S. J., C. FIELD, W. D. BOWEN AND W. BLANCHARD. 2004. Quantitative fatty acid signature analysis: A new method of estimating predator diet. Ecological Monographs 74:11-235.
77. IVERSON, S. J., I. STIRLING AND S. L. C. LANG. 2006. Spatial and temporal variation in the diets of polar bears across the Canadian Arctic: Link with and indicators of changes in prey populations. Pages 98-117 in I. L. Boyd, S. Wanless and C. J. Camphuysen, eds. Top predators in marine Ecosystems: Their role in monitoring and Management. Cambridge University Press, Cambridge, U.K.
78. JOSEPH, J. D. 1982. Lipid composition of marine and estuarine invertebrates. Part II: Mollusca. Progress in Lipid Research 21:109-153.
79. KÄKELÄ, R., AND H. HYVÄRINEN. 1996. Site-specific fatty acid composition in adipose tissues of several northern aquatic and terrestrial mammals. Comparative Biochemistry and Physiology B 115:501-514.
80. KATTNER, G., C. ALBERS, M. GRAEVE AND S. B. SCHNACK-SCHIEL. 2003. Fatty acid and alcohol composition of the small polar copepods Oithona and Oncaea: Indication of feeding modes. Polar Biology 26:666-671.
81. KIRSCH, P. E., S. J. IVERSON AND W. D. BOWEN. 2000. Effect of a low-fat diet on body composition and blubber fatty acids of captive juvenile harp seals (Phoca groenlandica). Physiological and Biochemical Zoology 73:45-59.
82. KOOPMAN, H. N. 1998. Topographical distribution of the blubber of harbour porpoises (Phocoena phocoena). Journal of Mammalogy 79:260-270.
83. KOOPMAN, H. N. 2001. The structure and function of the blubber of odontocetes. Ph.D. thesis, Duke University, Durham, NC. 408 pp.
84. KOOPMAN, H. N., S. J. IVERSON AND D. E. GASKIN. 1996. Stratification and age-related differences in blubber fatty acids of the male harbour porpoise (Phocoena phocoena). Journal of Comparative Physiology, B 165:628-639.
85. KOOPMAN, H. N., S. J. IVERSON AND A. J. READ. 2003. High concentrations of isovaleric acid in the fats of odontocetes: Stability in the melon vs. variation and patterns of accumulation in blubber. Journal of Comparative Physiology B 173:247-261.
86. KOOPMAN, H. N., S. M. BUDGE, D. R. KETTEN AND S. J. IVERSON. 2006. The topographical distribution of lipids inside the mandibular fat bodies of odontocetes: Remarkable complexity and consistency. Journal of Oceanographic Engineering Special Issue on Effects of Sound in the Marine Environment 31:95-106.
87. LAYTON, H., K. ROUVINEN AND S. J. IVERSON. 2000. Body composition in mink (Mustela vison) kits during 21-42 days postpartum using estimates of hydrogen isotope dilution and direct carcass analysis. Comparative Biochemistry and Physiology A 126:295-303.
88. LIN, J.-H., L.-Y. LIU, M.-H. YANG AND M.-H. LEE. 2004. Ethyl acetate/ethyl alcohol mixtures as an alternative to Folch reagent for extracting animal lipids. Journal of Agricultural and Food Chemistry 52:4984-4986.
89. LITCHFIELD, C., A. J. GREENBERG AND J. G. MEAD. 1976. The distinctive character of Ziphiidae head and blubber fats. Cetology 23:1-10.
90. LITCHFIELD, C., A. J. GREENBERG, R. G. ACKMAN AND C. A. EATON. 1978. Distinctive medium chain wax esters, triglycerides, and diacyl glyceryl ethers in the head fats of the Pacific Beaked Whale, Berardius bairdi. Lipids 13:860-866.
91. LIU, K.-S. 1994. Preparation of fatty acid methyl esters for gas-chromatographic analysis of lipids in biological materials. Journal of the American Oil Chemists' Society 71:1179-1187.
92. LOWRY, L. F. 1993. Foods and feeding ecology. Pages 201-238 in J. J. Burns, J. J. Montague, and C. J. Cowles, eds. The bowhead whale, Special Publication No. 2, Society for Marine Mammalogy. Allen Press, Lawrence, KS.
93. MAYZAUD, P., E. ALBESSARD, P. VIRTUE AND M. BOUTOUTE. 2000. Environmental constraints on the lipid composition and metabolism of euphausiids: The case of Euphausia superba and Meganyctiphanes norvegica. Canadian Journal of Fisheries and Aquatic Sciences 57:91-103.
94. MORII, H., AND T. KANEDA. 1982. Biosynthesis of branched-chain fatty acids from branched-chain amino acids in subcutaneous tissue of the marine little toothed whale, Stenella caeruleo-alba. Comparative Biochemistry and Physiology B 71:357-365.
95. NAJDEK, M., S. PUKARIC AND A. B. BOCHDANSKY. 1994. Contribution of zooplankton lipids to the flux of organic matter in the northern Adriatic Sea. Marine Ecology Progress Series 111:241-249.
96. NERINI, M. 1984. A review of gray whale feeding ecology. Pages 423-450 in M. L. Jones, S. L. Swartz and S. Leatherwood, eds. The grey whale, Eschrichtius robustus. Academic Press, Orlando, FL.
97. NORSETH, J. AND B. O. CHRISTOPHERSEN. 1978. Chain shortening of erucic acid in isolated liver cells. FEBS Letters 88:353-357.
98. OFTEDAL, O. T. 1997. Lactation in whales and dolphins: Evidence of divergence between baleen- and toothed-species. Journal of Mammary Gland Biology and Neoplasia 2:205-230.
99. OSMUNDSEN, H., C. E. NEAT AND K. R. NORUM. 1979. Peroxisomal oxidation of long chain fatty acids. FEBS Letters 99:292-296
100. PABST, D. A. 1996. Springs in swimming animals. American Zoologist 36:723-735.
101. PALTAUF, F. 1994. Ether lipids in biomembranes. Chemistry and Physics of Lipids 74:101-139.
102. PAPAMANDJARIS, A. A., D. E. MACDOUGALL AND P. J. H. JONES. 1998. Medium chain fatty acid metabolism and energy expenditure: Obesity treatment implications. Life Sciences 62:1203-1215.
103. PASCAL, J. C., AND R. G. ACKMAN. 1975. Occurrence of 7-methyl-7- hexadecenoic acid, the corresponding alcohol, 7-methyl-6-hexadecenoic acid, and 5-methyl-4-tetradecenoic acid in sperm whale oils. Lipids 10:478-482.
104. PHLEGER, C. F., P. D. NICHOLS AND P. VIRTUE. 1997. The lipid, fatty acid and fatty alcohol composition of the myctophid fish Electrona antarctica: High levels of wax esters and food chain implications. Antarctic Science 9:258-265.
105. PHLEGER, C. F., M. M. NELSON, B. D. MOONEY AND P. D. NICHOLS. 1999. Wax esters versus triacylglycerols in myctophid fishes from the Southern Ocean. Antarctic Science 11:436-444.
106. POND, C. M. 1998. The fats of life. Cambridge University Press, Cambridge, UK.
107. POND, C. M., C. A. MATTACKS, R. H. COLBY AND M. A. RAMSAY. 1992. The anatomy, chemical composition, and metabolism of adipose tissue in wild polar bears (Ursus maritimus). Canadian Journal of Zoology 70:326-341.
108. POND, C. M., C. A. MATTACKS AND P. PRESTRUD. 1995. Variability in the distribution and composition of adipose tissue in arctic foxes (Alopex lagopus) on Svalbard. Journal of Zoology, London 236:593-610.
109. RACLOT, T. 2003. Selective mobilization of fatty acids from adipose tissue triacylglycerols. Progress in Lipid Reseach 42:257-288.
110. RACLOT, T., R. GROSCOLAS AND Y. CHEREL. 1998. Fatty acid evidence for the importance of myctophid fishes in the diet of king penguins, Aptenodytes patagonicus. Marine Biology 132:523-533.
111. RATNAYAKE, W. N., AND R. G. ACKMAN. 1979. Fatty alcohols in capelin, herring and mackerel oils and muscle lipids: 1. Fatty alcohol details linking dietary copepod fat with certain fish depot fats. Lipids 14:795-803.
112. RECCHIA, C. A., AND A. J. READ. 1989. Stomach contents of harbour porpoises, Phocoena phocoena (L.), from the Bay of Fundy. Canadian Journal of Zoology 67:2140-2146.
113. REEVES, R. R., B. S. STEWART, P. J. CLAPHAM AND J. A. POWELL. 2002. National Audobon Society Guide to Marine Mammals of the World. Alfred A. Knopf, New York, NY.
114. ROBY, D. D., A. R. PLACE AND R. E. RICKLEFS. 1986. Assimilation and deposition of wax esters in planktivorous seabirds. Journal of Experimental Zoology 238:29-41.
115. ROOD, D. 1995. A practical guide to the care, maintenance, and troubleshooting of capillary gas chromatographic systems. Hüthig, Heidelberg, Germany.
116. ROUVINEN, K., AND T. KIISKINEN. 1989. Influence of dietary fat source on the body fat composition of mink (Mustela vison) and blue fox (Alopex lagopus). Acta Agricultura Scandinavia 39:279-288.
117. RYG, M., T. G. SMITH AND N. A. ØRITSLAND. 1988. Thermal significance of the topographical distribution of blubber in ringed seals (Phoca hispida). Canadian Journal of Fisheries and Aquatic Sciences 45:985-992.
118. SANTOS, M. B., AND G. J. PIERCE. 2003. The diet of harbour porpoises (Phocoena phocoena) in the Northeast Atlantic. Oceanographic and Marine Biology - an Annual Review 41:355-390.
119. SARGENT, J. R. 1976. The structure, metabolism and function of lipids in marine organisms. Pages 149-212 in D. C. Malins and J. R. Sargent, eds. Biochemical and biophysical perspectives in marine biology. Academic Press, London, UK.
120. SARGENT, J. R., R. F. LEE AND J. C. NEVENZEL. 1976. Marine waxes. Pages 49-91 in P. E. Kolattukudy, ed. Chemistry and biochemistry of natural waxes. Elsevier, Oxford, UK.
121. SARGENT, J. R., R. J. PARKES, I. MUELLER-HARVEY AND R. J. HENDERSON. 1987. Lipid biomarkers in marine ecology. Pages 119-138 in M. A. Sleigh, ed. Microbes in the sea. Wiley and Sons, New York, NY.
122. SHAIKH, N. A. 1986. Extraction, purification, and analysis of lipids from animal tissues. Pages 289-302 in H. A. Fozzard, E. Haber, R. B. Jennings, A. M. Katz and H. E. Morgan, eds. The heart and cardiovascular system. Raven Press, New York, NY.
123. SMEDES, F. 1999. Determination of total lipid using non-chlorinated solvents. Analyst 124:1711-1718.
124. SMEDES, F., AND T. K. ASKLAND. 1999. Revisiting the development of the Bligh and Dyer total lipid determination method. Marine Pollution Bulletin 38:193-201.
125. SMITH, R. J., K. A. HOBSON, H. N. KOOPMAN, AND D. M. LAVIGNE. 1996. Distinguishing between populations of fresh- and salt-water harbour seals (Phoca vitulina) using stable isotope ratios and fatty acid profiles. Canadian Journal of Fisheries and Aquatic Sciences 53:272-279.
126. SMITH, S. J., S. J. IVERSON AND W. D. BOWEN. 1997. Fatty acid signatures and classification trees: New tools for investigating the foraging ecology of seals. Canadian Journal of Fisheries and Aquatic Sciences 54:1377-1386.
127. STENSTROM, M. K., S. FAM AND G. S. SILVERMAN. 1986. Analytical methods for quantitative and qualitative determination of hydrocarbons and oil and greases in water and wastewater. Environmental Technology Letters 7:625-636.
128. STEVENS, J. 1986. Applied multivariate statistics for the social sciences. Lawrence Erlbaum Associates, NJ.
129. STEWART, C. 2005. Inference on the diet of predators using fatty acid signatures. Ph.D. thesis, Dalhousie University, Halifax, NS; 191 pp.
130. TABACHNICK, B. G., AND L. S. FIDELL. 1989. Using Multivariate Statistics. Harper Collins Publishers, New York, NY.
131. THIEMANN, G. W., S. M. BUDGE AND S. J. IVERSON. 2004. Fatty acid composition of blubber: A comparison of in situ direct and traditional extraction methods. Marine Mammal Science 20:284-295.
132. THIEMANN, G. W., S. J. IVERSON AND I. STIRLING. 2006. Seasonal, sexual, and anatomical variability in the adipose tissue composition of polar bears (Ursus maritimus). Journal of Zoology London 269:65-76.
133. UNDELAND, I., M. HÄRRÖD AND H. LINGNERT. 1998. Comparison between methods using low-toxicity solvents for the extraction of lipids from herring (Clupea harengus). Food Chemistry 61:355-365.
134. VARANASI, U., AND D. C. MALINS. 1971. Unique lipids of the porpoise (Tursiops gilli): Differences in triacylglycerols and wax esters of acoustic (mandibular canal and melon) and blubber tissues. Biochimica et Biophysica Acta 231:415-418.
135. VOLPE, J. J., AND P. R. VAGELOS. 1973. Saturated fatty acid biosynthesis and its regulation. Annual Review of Biochemistry 42:21-60.
136. VOSS, A., M. REINHART, S. SANKARAPPA AND H. SPRECHER. 1991. The metabolism of 7,10,13,16,19-docosapentaenoic acid to 4,7,10,13,16,19- docosahexaenoic acid in rat liver is independent of a 4-desaturase. Journal of Biological Chemistry 266:19995-20000.
137. WALTON, M. J., R. J. HENDERSON AND P. P. POMEROY. 2000. Use of blubber fatty acid profiles to distinguish dietary differences between grey seals Halichoerus grypus from two UK breeding colonies. Marine Ecology Progress Series 193:210-208.
138. WAMBERG, S., C. R. OLESEN AND H. O. HANSEN. 1992. Influenece of dietary sources of fat on lipid synthesis in mink (Mustela vison) mammary tissue. Comparative Biochemistry and Physiology A 103:199-204.
139. WETZEL, D. L., AND J. E. REYNOLDS. 2004. Definitive identification of fatty acid constituents in marine mammal tissues. Canadian Journal of Fisheries and Aquatic Science 61:554-560.
140. WEST, G. C., J. J. BURNS AND M. MODAFFERI. 1979. Fatty acid composition of Pacific walrus skin and blubber fats. Canadian Journal of Zoology 57:1249-1255.
141. ZAR, J. H. 1999. Biostatistical analysis. Prentice-Hall, Upper Saddle River, NJ.
142. ZWEIG, G., AND J. SHERMA. 1972. CRC handbook of chromatography. CRC Press, Cleveland, OH.