Sewall and Rodgveller: Changes in body composition and fatty acid profile during embryogenesis of Sebastes maliger 
219 
and it was present at relatively low levels in quillback 
rockfish embryos. 
Although both lipid and protein are consumed during 
quillback rockfish embryogenesis, lipid is used more 
rapidly and contributes a greater portion of the total 
energy expended. Lipid is typically the most variable 
dry mass component of fish eggs, showing significant 
differences between populations and within a popula- 
tion over time; additionally, lipid concentration has been 
used as an indicator of larval viability for several spe- 
cies (reviewed in Kamler, 1992). Given the importance 
of lipid as an energy source, the strong relationship be- 
tween OGV and lipid levels confirms the utility of OGV 
as an indicator of differences in condition of quillback 
rockfish embryos and preparturition larvae. Differences 
in FA profiles of early embryos and preparturition lar- 
vae indicate FAs are depleted at different rates during 
embryogenesis. More rapidly used FAs may contribute 
more to lipid energy use or serve as precursors in the 
synthesis of other FAs, while conserved FAs likely are 
incorporated into tissues or hormone-like compounds. 
The conservation of 20:4n-6, the most abundant n-6 
PUFA, indicates that this essential fatty acid may well 
reflect the quality of maternal provisioning. The high 
degree of interspecific variability in body composition 
and energy use patterns among rockfish illustrates the 
need for data gathered from the species of interest, in 
order to make the most accurate models of energy use 
and most appropriate indicators of condition. 
Acknowledgments 
We thank R. Heintz for advice on study methodology and 
data analysis, M. Larsen for generating the fatty acid 
data, and R. Bradshaw for generating the protein, mois- 
ture, and ash content data. We also thank J. Maselko for 
developing the computer code for creating the Aitchison 
distance matrix and ANOSIM statistical analysis, C. 
Lunsford for assisting with field sampling, and J. Heifetz 
and L. Schaufler for providing helpful commentary and 
edits on earlier drafts. 
Literature cited 
Aitchison, J. 
1992. On criteria for measures of compositional 
difference. Math. Geol. 24:365-379. 
Bell, J. G., and J. R. Sargent. 
2003. Arachidonic acid in aquaculture feeds: current 
status and future opportunities. Aquacult. 218:491- 
499. 
Berkeley, S. A., C. Chapman, and S. M. Sogard. 
2004. Maternal age as a determinant of larval growth and 
survival in a marine fish, Sebastes melanops. Ecology 
85(5):1258-1264. 
Brett, J. R. 
1995. Energetics. In Physiological ecology of Pacific 
salmon (Groot, C., Margolis, L., and W.C. Clarke, eds.), 
p. 1-68. UBC Press, Vancouver, Canada. 
Christie, W. W. 
2003. Lipid analysis: Isolation, separation, identification 
and structural analysis of lipids, 416 p. Oily Press, 
Bridgwater, U.K. 
Clarke, K. R., and R. M. Warwick. 
1994. Change in marine communities: An approach to 
statistical analysis and interpretation, 144p. Plymouth 
Marine Laboratory, Plymouth, U.K. 
Craig, J. F., M. J. Kenley, and J. F. Tailing. 
1978. Comparative estimations of the energy content of 
fish tissue from bomb calorimetry, wet oxidation and 
proximate analysis. Freshw. Biol. 8:585-590. 
Eldridge, M. B., E. C. Norton, B. M. Jarvis, and R. B. 
MacFarlane. 
2002. Energetics of early development in the viviparous 
yellowtail rockfish. J. Fish Biol. 61:1122-1134. 
Gunasekera, R. M., S. S. De Silva, and B. A. Ingram. 
1999. Early ontogeny-related changes of the fatty acid 
composition in the percichthyid fishes trout cod, Maccullo- 
chella macquariensis and Murray cod, Maccullochella 
pelii peelii. Aquat. Living Resour. 12:219-227. 
Horwitz, W., ed. 
2002. Official methods of analysis, 17 th ed. AOAC 
(Association of Analytical Communities) International, 
Gaithersburg, MD. 
Kamler, E. 
1992. Early life history of fish, an energetics approach. 
267 p. Chapman and Hall, London. 
Lee, S. M. 
2001. Review of the lipid and essential fatty acid require- 
ments of rockfish ( Sebastes schlegeli). Aquacult. Res. 
32 ( s 1 ) : 8— 17. 
MacFarlane, R. B., and M. J. Bowers. 
1995. Matrotrophic viviparity in the yellowtail rockfish 
Sebastes flavidus. J. Exp. Biol. 198:1197-206. 
MacFarlane, R. B., and E. C. Norton. 
1999. Nutritional dynamics during embryonic develop- 
ment in the viviparous genus Sebastes and their applica- 
tion to the assessment of reproductive success. Fish. 
Bull. 97:273-281. 
Matala, A. P., A. K. Gray, J. Heifetz, and A. J. Gharrett. 
2004. Population structure of Alaskan shortraker rock- 
fish, Sebastes borealis, inferred from microsatellite 
variation. Environ. Biol. Fishes 69:201-210. 
Mourent.e, G., and D. R. Tocher. 
1993. Incorporation and metabolism of 14 C-labeled poly- 
unsaturated fatty acids in wild-caught juveniles of golden 
grey mullet, Liza aurata, in vivo. Fish Physiol Bio- 
chem. 12(2):119— 130. 
Mourente, G., and R. Vazquez. 
1996. Changes in the content of total lipid, lipid classes 
and fatty acids of developing eggs and unfed larvae of 
the Senegal sole, Solea senegalensis Kaup. Fish Physiol. 
Biochem. 15(3):221-235. 
Norton, E. C., R. B. MacFarlane, and M. S. Mohr. 
2001. Lipid class dynamics during development in early 
life stages of shortbelly rockfish and their application 
to condition assessment. J. Fish. Biol. 58:1010—1024. 
Parker, S. J., S. A. Berkeley, J. T. Golden, D. R. Gunderson, J. 
Heifetz, M. A. Hixon, R. Larson, B. M. Leaman, M. S. Love, 
J. A. Musick, V. M. O’Connell, S. Ralston, H. J. Weeks, and 
M. M. Yoklavich. 
2000. Management of Pacific rockfish. Fisheries 25(3): 
22-30. 
Ralston, S., and D. F. Howard 
1995. On the development of year-class strength and cohort 
