Enhancing Bioenergetics Models to Account for Dynamic Changes in Fish Body Composition and Energy Density

Fish proximate composition and energy density can influence growth, survival, and reproduction, so it is important to develop models to understand the patterns and predict dynamic changes. This paper presents three such models. Model 1 describes the general pattern of changes in lipid, protein, ash,...

Full description

Bibliographic Details
Published in:Transactions of the American Fisheries Society
Main Author: Breck, James E.
Format: Article in Journal/Newspaper
Language:unknown
Published: Wiley Periodicals, Inc. 2008
Subjects:
Online Access:http://hdl.handle.net/2027.42/141335
https://doi.org/10.1577/T05-240.1
id ftumdeepblue:oai:deepblue.lib.umich.edu:2027.42/141335
record_format openpolar
institution Open Polar
collection University of Michigan: Deep Blue
op_collection_id ftumdeepblue
language unknown
topic Natural Resources and Environment
Science
spellingShingle Natural Resources and Environment
Science
Breck, James E.
Enhancing Bioenergetics Models to Account for Dynamic Changes in Fish Body Composition and Energy Density
topic_facet Natural Resources and Environment
Science
description Fish proximate composition and energy density can influence growth, survival, and reproduction, so it is important to develop models to understand the patterns and predict dynamic changes. This paper presents three such models. Model 1 describes the general pattern of changes in lipid, protein, ash, and energy density that occur with changes in water content. The key assumption this model is that there is a fixed amount of water associated with each gram of protein and a much smaller fixed amount of water associated with each gram of lipid. In combination with a mass balance constraint, this explains the commonly observed linear relationship between the fraction lipid and the fraction water. Because energy density varies in direct proportion to the fractions lipid and protein, the linear relationship between body composition and fraction water makes energy density also a linear function of the fraction water. The model is fitted to data for lake trout Salvelinus namaycush and coho salmon Oncorhynchus kisutch for a limited range in wet weight. Model 2 describes the pattern of proximate composition and energy density that occurs with variation in body size. A strong pattern was found between the mass of water and the mass of protein, suggesting strict control of body water. The model is fitted to data for common carp Cyprinus carpio and bluegill Lepomis macrochirus. This analysis shows that the relationship between body composition, energy density, and fraction water is expected to vary with body size because both the water: Protein ratio and the fraction ash change with body size. Model 3 demonstrates how this approach can be used to predict changes in fish body composition and energy density during starvation, as might be done with a bioenergetics model. This model is fitted to data from a starvation experiment involving largemouth bass Micropterus salmoides. Peer Reviewed https://deepblue.lib.umich.edu/bitstream/2027.42/141335/1/tafs0340.pdf
format Article in Journal/Newspaper
author Breck, James E.
author_facet Breck, James E.
author_sort Breck, James E.
title Enhancing Bioenergetics Models to Account for Dynamic Changes in Fish Body Composition and Energy Density
title_short Enhancing Bioenergetics Models to Account for Dynamic Changes in Fish Body Composition and Energy Density
title_full Enhancing Bioenergetics Models to Account for Dynamic Changes in Fish Body Composition and Energy Density
title_fullStr Enhancing Bioenergetics Models to Account for Dynamic Changes in Fish Body Composition and Energy Density
title_full_unstemmed Enhancing Bioenergetics Models to Account for Dynamic Changes in Fish Body Composition and Energy Density
title_sort enhancing bioenergetics models to account for dynamic changes in fish body composition and energy density
publisher Wiley Periodicals, Inc.
publishDate 2008
url http://hdl.handle.net/2027.42/141335
https://doi.org/10.1577/T05-240.1
genre Arctic
genre_facet Arctic
op_relation Breck, James E. (2008). "Enhancing Bioenergetics Models to Account for Dynamic Changes in Fish Body Composition and Energy Density." Transactions of the American Fisheries Society 137(1): 340-356.
0002-8487
1548-8659
http://hdl.handle.net/2027.42/141335
doi:10.1577/T05-240.1
Transactions of the American Fisheries Society
M. S. Ridgway, B. J. Shuter and E. E. Post, 1991 The relative influence of body size and territorial behaviour on nesting asynchrony in male smallmouth bass, Micropterus dolomieui (Pisces: Centrarchidae), Journal of Animal Ecology, 60, Pages 665 – 681.
K. Schmidt‐Nielsen, 1975. In Animal physiology: Adaptation and environment, Cambridge University Press, London.
B. J. Shuter, J. A. MacLean, F. E. J. Fry and H. A. Regier, 1980 Stochastic simulation of temperature effects on first‐year survival of smallmouth bass, Transactions of the American Fisheries Society, 109, Pages 1 – 34.
D. G. Simpkins, W. A. Hubert, C. Martinez del Rio and D. C. Rule, 2003 Effect of swimming activity on relative weight and body composition of juvenile rainbow trout, North American Journal of Fisheries Management, 23, Pages 283 – 289.
S. M. Sogard and B. L. Olla, 2000 Endurance of simulated winter conditions by age‐0 walleye pollock: Effects of body size, water temperature, and energy stores, Journal of Fish Biology, 56, Pages 1 – 21.
D. J. Stewart and M. Ibarra, 1991 Predation and production by salmonine fishes in Lake Michigan, 1978‐1988, Canadian Journal of Fisheries and Aquatic Sciences, 48, Pages 909 – 922.
D. J. Stewart, D. Weininger, D. V. Rottiers and T. A. Edsall, 1983 An energetics model for lake trout, Salvelinus namaycush: Application to the Lake Michigan population, Canadian Journal of Fisheries and Aquatic Sciences, 40, Pages 681 – 698.
M. J. Tarby, 1977. In Energetics and growth of walleye (Stizostedion vitreum vitreum) in Oneida Lake, New York. Doctoral dissertation, Cornell University, Ithaca, New York.
J. M. Thompson, E. P. Bergersen, C. A. Carlson and L. R. Kaeding, 1991 Role of size, condition, and lipid content in the overwinter survival of age‐0 Colorado squawfish, Transactions of the American Fisheries Society, 120, Pages 346 – 353.
A. V. Tyler and R. S. Dunn, 1976 Ration, growth, and measures of somatic and organ condition in relation to meal frequency in winter flounder, Pseudopleuronectes americanus, with hypotheses regarding population homeostasis, Journal of the Fisheries Research Board of Canada, 33, Pages 63 – 75.
T. I. Van Pelt, J. F. Piatt, B. K. Lance and D. D. Roby, 1997 Proximate composition and energy density of some North Pacific forage fishes, Comparative Biochemistry and Physiology, 118A, Pages 1393 – 1398.
W. Van Winkle, B. J. Shuter, B. D. Holcomb, H. I. Jager, J. A. Tyler and S. Y. Whitaker, 1997, “ Regulation of energy acquisition and allocation to respiration, growth, and reproduction: Simulation model and example using rainbow trout ”, Pages 103 – 137. Edited by: R. C. Chambers, E. A. Trippel. In Early life history and recruitment in fish populations, Chapman and Hall, London.
A. H. Weatherley and H. S. Gill, 1983 Protein, lipid, water, and caloric contents of immature rainbow trout, Salmo gairdneri Richardson, growing at different rates, Journal of Fish Biology, 23, Pages 653 – 673.
G. J. Wege and R. O. Anderson, 1978, “ Relative weight ( W r ): A new index of condition for largemouth bass ”, Pages 79 – 91. Edited by: G. Novinger, J. Dillard. In New approaches to the management of small impoundments, American Fisheries Society, North Central Division, Bethesda, Maryland, Special Publication 5.
D. W. Willis, C. S. Guy and B. R. Murphy, 1991 Development and evaluation of a standard weight ( W s ) equation for yellow perch, North American Journal of Fisheries Management, 11, Pages 374 – 380.
M. J. Wuenschel, A. R. Jugovich and J. A. Hare, 2006 Estimating the energy density of fish: The importance of ontogeny, Transactions of the American Fisheries Society, 135, Pages 379 – 385.
S. D. Gerking, 1955 Influence of rate of feeding on body composition and protein metabolism of bluegill sunfish, Physiological Zoology, 28, Pages 267 – 282.
S. M. Adams, J. E. Breck and R. B. McLean, 1985 Cumulative stress‐induced mortality of gizzard shad in a southeastern U.S. reservoir, Environmental Biology of Fishes, 13, Pages 103 – 112.
K. P. Anderson and E. Ursin, 1977 A multispecies extension to the Beverton and Holt theory, with accounts of phosphorus circulation and primary production, Meddelelser fra Danmarks Fiskeri‐ og Havundersogelser, N.S, 7, Pages 319 – 435.
R. O. Anderson and S. J. Gutreuter, 1983, “ Length, weight, and associated structural indices ”, Pages 283 – 300. Edited by: L. A. Nielsen, D. L. Johnson. In Fisheries techniques, American Fisheries Society, Bethesda, Maryland.
J. A. Anthony, D. D. Roby and K. R. Turco, 2000 Lipid content and energy density of forage fishes from the northern Gulf of Alaska, Journal of Experimental Marine Biology and Ecology, 248, Pages 53 – 78.
S. M. Bartell, J. E. Breck, R. H. Gardner and A. L. Brenkert, 1986 Individual parameter perturbation and error analysis of fish bioenergetics models, Canadian Journal of Fisheries and Aquatic Sciences, 43, Pages 160 – 168.
D. E. Barziza and D. M. Gatlin III., 2000 An evaluation of total body electrical conductivity to estimate body composition of largemouth bass, Micropterus salmoides, Aquatic Living Resources, 13, Pages 439 – 447.
D. Black and R. M. Love, 1986 The sequential mobilization and restoration of energy reserves in tissues of Atlantic cod during starvation and refeeding, Journal of Comparative Physiology B, 156, Pages 469 – 479.
J. E. Breck, 1993 Hurry up and wait: Growth of young bluegills in ponds and simulations with an individual‐based model, Transactions of the American Fisheries Society, 122, Pages 467 – 480.
J. E. Breck, 1996. In Mechanisms of recruitment failure in bluegill ponds, Michigan Department of Natural Resources, Ann Arbor, Fisheries Research Report 2024.
J. R. Brett, 1995, “ Energetics ”, Pages 1 – 68. Edited by: C. Groot, L. Margolis, W. C. Clarke. In Physiological ecology of Pacific salmon, UBC Press, Vancouver.
J. R. Brett and T. D. D. Groves, 1979, “ Physiological energetics ”, Pages 279 – 352. Edited by: W. S. Hoar, D. J. Randall, J. R. Brett. In Fish physiology, Academic Press, New York, Vol. 8.
J. R. Brett, J. E. Shelbourn and C. T. Shoop, 1969 Growth rate and body composition of fingerling sockeye salmon, Oncorhynchus nerka, in relation to temperature and ration size, Journal of the Fisheries Research Board of Canada, 26, Pages 2363 – 2394.
N. Broekhuizen, W. S. C. Gurney, A. Jones and A. D. Bryant, 1994 Modelling compensatory growth, Functional Ecology, 8, Pages 770 – 782.
M. L. Brown and B. R. Murphy, 1991 Relationship of relative weight ( W r ) to proximate composition of juvenile striped bass and hybrid striped bass, Transactions of the American Fisheries Society, 120, Pages 509 – 518.
L. M. Cargnelli and M. R. Gross, 1997 Fish energetics: Larger individuals emerge from winter in better condition, Transactions of the American Fisheries Society, 126, Pages 153 – 156.
M. K. Cox and K. J. Hartman, 2005 Nonlethal estimation of proximate composition in fish, Canadian Journal of Fisheries and Aquatic Sciences, 62, Pages 269 – 275.
J. F. Craig, 1977 The body composition of adult perch, Perca fluviatilis, in Windermere, with reference to seasonal changes and reproduction, Journal of Animal Ecology, 46, Pages 617 – 632.
J. F. Craig, M. J. Kenley and J. F. Talling, 1978 Comparative estimations of the energy content of fish tissue from bomb calorimetry, wet oxidation, and proximate analysis, Freshwater Biology, 8, Pages 585 – 590.
G. T. Crossin and S. G. Hinch, 2005 A nonlethal, rapid method for assessing the somatic energy content of migrating adult Pacific salmon, Transactions of the American Fisheries Society, 134, Pages 184 – 191.
A. J. Danylchuk and M. G. Fox, 1994 Age‐ and size‐dependent variation in the seasonal timing and probability of reproduction among mature female pumpkinseed (Lepomis gibbosus), Environmental Biology of Fishes, 39, Pages 119 – 127.
S. I. Doroshov, 1985, “ Biology and culture of sturgeon Acipenseriformes ”, Pages 251 – 274. Edited by: J. F. Muir, R. J. Roberts. In Recent advances in aquaculture, Croom Helm, London.
J. D. Dutil, 1986 Energetic constraints and spawning interval in the anadromous Arctic charr (Salvelinus alpinus), Copeia, 1986, Pages 945 – 955.
J. M. Elliott, 1976 Body composition of brown trout (Salmo trutta L.) in relation to temperature and ration size, Journal of Animal Ecology, 45, Pages 273 – 289.
R. U. Fischer, J. D. Congdon and M. Brock, 1996 Total body electrical conductivity (TOBEC): A tool to estimate lean mass and nonpolar lipids of an aquatic organism?, Copeia, 1996, Pages 459 – 462.
U. Focken and K. Becker, 1993, “ Body composition of carp (Cyprinus carpio L.) ”, Pages 269 – 288. Edited by: T. Braunbeck, W. Hanke, H. Segner. In Fish ecotoxicology and ecophysiology, VCH Publishers, New York.
J. From and G. Rasmussen, 1984 A growth model, gastric evacuation, and body composition in rainbow trout, Salmo gairdneri Richardson, 1836, Dana, 3, Pages 61 – 139.
K. J. Hartman and S. B. Brandt, 1995 Estimating energy density of fish, Transactions of the American Fisheries Society, 124, Pages 347 – 355.
op_rights IndexNoFollow
op_doi https://doi.org/10.1577/T05-240.1
container_title Transactions of the American Fisheries Society
container_volume 137
container_issue 1
container_start_page 340
op_container_end_page 356
_version_ 1774713605465309184
spelling ftumdeepblue:oai:deepblue.lib.umich.edu:2027.42/141335 2023-08-20T04:03:12+02:00 Enhancing Bioenergetics Models to Account for Dynamic Changes in Fish Body Composition and Energy Density Breck, James E. 2008-01 application/pdf http://hdl.handle.net/2027.42/141335 https://doi.org/10.1577/T05-240.1 unknown Wiley Periodicals, Inc. Taylor & Francis Group Breck, James E. (2008). "Enhancing Bioenergetics Models to Account for Dynamic Changes in Fish Body Composition and Energy Density." Transactions of the American Fisheries Society 137(1): 340-356. 0002-8487 1548-8659 http://hdl.handle.net/2027.42/141335 doi:10.1577/T05-240.1 Transactions of the American Fisheries Society M. S. Ridgway, B. J. Shuter and E. E. Post, 1991 The relative influence of body size and territorial behaviour on nesting asynchrony in male smallmouth bass, Micropterus dolomieui (Pisces: Centrarchidae), Journal of Animal Ecology, 60, Pages 665 – 681. K. Schmidt‐Nielsen, 1975. In Animal physiology: Adaptation and environment, Cambridge University Press, London. B. J. Shuter, J. A. MacLean, F. E. J. Fry and H. A. Regier, 1980 Stochastic simulation of temperature effects on first‐year survival of smallmouth bass, Transactions of the American Fisheries Society, 109, Pages 1 – 34. D. G. Simpkins, W. A. Hubert, C. Martinez del Rio and D. C. Rule, 2003 Effect of swimming activity on relative weight and body composition of juvenile rainbow trout, North American Journal of Fisheries Management, 23, Pages 283 – 289. S. M. Sogard and B. L. Olla, 2000 Endurance of simulated winter conditions by age‐0 walleye pollock: Effects of body size, water temperature, and energy stores, Journal of Fish Biology, 56, Pages 1 – 21. D. J. Stewart and M. Ibarra, 1991 Predation and production by salmonine fishes in Lake Michigan, 1978‐1988, Canadian Journal of Fisheries and Aquatic Sciences, 48, Pages 909 – 922. D. J. Stewart, D. Weininger, D. V. Rottiers and T. A. Edsall, 1983 An energetics model for lake trout, Salvelinus namaycush: Application to the Lake Michigan population, Canadian Journal of Fisheries and Aquatic Sciences, 40, Pages 681 – 698. M. J. Tarby, 1977. In Energetics and growth of walleye (Stizostedion vitreum vitreum) in Oneida Lake, New York. Doctoral dissertation, Cornell University, Ithaca, New York. J. M. Thompson, E. P. Bergersen, C. A. Carlson and L. R. Kaeding, 1991 Role of size, condition, and lipid content in the overwinter survival of age‐0 Colorado squawfish, Transactions of the American Fisheries Society, 120, Pages 346 – 353. A. V. Tyler and R. S. Dunn, 1976 Ration, growth, and measures of somatic and organ condition in relation to meal frequency in winter flounder, Pseudopleuronectes americanus, with hypotheses regarding population homeostasis, Journal of the Fisheries Research Board of Canada, 33, Pages 63 – 75. T. I. Van Pelt, J. F. Piatt, B. K. Lance and D. D. Roby, 1997 Proximate composition and energy density of some North Pacific forage fishes, Comparative Biochemistry and Physiology, 118A, Pages 1393 – 1398. W. Van Winkle, B. J. Shuter, B. D. Holcomb, H. I. Jager, J. A. Tyler and S. Y. Whitaker, 1997, “ Regulation of energy acquisition and allocation to respiration, growth, and reproduction: Simulation model and example using rainbow trout ”, Pages 103 – 137. Edited by: R. C. Chambers, E. A. Trippel. In Early life history and recruitment in fish populations, Chapman and Hall, London. A. H. Weatherley and H. S. Gill, 1983 Protein, lipid, water, and caloric contents of immature rainbow trout, Salmo gairdneri Richardson, growing at different rates, Journal of Fish Biology, 23, Pages 653 – 673. G. J. Wege and R. O. Anderson, 1978, “ Relative weight ( W r ): A new index of condition for largemouth bass ”, Pages 79 – 91. Edited by: G. Novinger, J. Dillard. In New approaches to the management of small impoundments, American Fisheries Society, North Central Division, Bethesda, Maryland, Special Publication 5. D. W. Willis, C. S. Guy and B. R. Murphy, 1991 Development and evaluation of a standard weight ( W s ) equation for yellow perch, North American Journal of Fisheries Management, 11, Pages 374 – 380. M. J. Wuenschel, A. R. Jugovich and J. A. Hare, 2006 Estimating the energy density of fish: The importance of ontogeny, Transactions of the American Fisheries Society, 135, Pages 379 – 385. S. D. Gerking, 1955 Influence of rate of feeding on body composition and protein metabolism of bluegill sunfish, Physiological Zoology, 28, Pages 267 – 282. S. M. Adams, J. E. Breck and R. B. McLean, 1985 Cumulative stress‐induced mortality of gizzard shad in a southeastern U.S. reservoir, Environmental Biology of Fishes, 13, Pages 103 – 112. K. P. Anderson and E. Ursin, 1977 A multispecies extension to the Beverton and Holt theory, with accounts of phosphorus circulation and primary production, Meddelelser fra Danmarks Fiskeri‐ og Havundersogelser, N.S, 7, Pages 319 – 435. R. O. Anderson and S. J. Gutreuter, 1983, “ Length, weight, and associated structural indices ”, Pages 283 – 300. Edited by: L. A. Nielsen, D. L. Johnson. In Fisheries techniques, American Fisheries Society, Bethesda, Maryland. J. A. Anthony, D. D. Roby and K. R. Turco, 2000 Lipid content and energy density of forage fishes from the northern Gulf of Alaska, Journal of Experimental Marine Biology and Ecology, 248, Pages 53 – 78. S. M. Bartell, J. E. Breck, R. H. Gardner and A. L. Brenkert, 1986 Individual parameter perturbation and error analysis of fish bioenergetics models, Canadian Journal of Fisheries and Aquatic Sciences, 43, Pages 160 – 168. D. E. Barziza and D. M. Gatlin III., 2000 An evaluation of total body electrical conductivity to estimate body composition of largemouth bass, Micropterus salmoides, Aquatic Living Resources, 13, Pages 439 – 447. D. Black and R. M. Love, 1986 The sequential mobilization and restoration of energy reserves in tissues of Atlantic cod during starvation and refeeding, Journal of Comparative Physiology B, 156, Pages 469 – 479. J. E. Breck, 1993 Hurry up and wait: Growth of young bluegills in ponds and simulations with an individual‐based model, Transactions of the American Fisheries Society, 122, Pages 467 – 480. J. E. Breck, 1996. In Mechanisms of recruitment failure in bluegill ponds, Michigan Department of Natural Resources, Ann Arbor, Fisheries Research Report 2024. J. R. Brett, 1995, “ Energetics ”, Pages 1 – 68. Edited by: C. Groot, L. Margolis, W. C. Clarke. In Physiological ecology of Pacific salmon, UBC Press, Vancouver. J. R. Brett and T. D. D. Groves, 1979, “ Physiological energetics ”, Pages 279 – 352. Edited by: W. S. Hoar, D. J. Randall, J. R. Brett. In Fish physiology, Academic Press, New York, Vol. 8. J. R. Brett, J. E. Shelbourn and C. T. Shoop, 1969 Growth rate and body composition of fingerling sockeye salmon, Oncorhynchus nerka, in relation to temperature and ration size, Journal of the Fisheries Research Board of Canada, 26, Pages 2363 – 2394. N. Broekhuizen, W. S. C. Gurney, A. Jones and A. D. Bryant, 1994 Modelling compensatory growth, Functional Ecology, 8, Pages 770 – 782. M. L. Brown and B. R. Murphy, 1991 Relationship of relative weight ( W r ) to proximate composition of juvenile striped bass and hybrid striped bass, Transactions of the American Fisheries Society, 120, Pages 509 – 518. L. M. Cargnelli and M. R. Gross, 1997 Fish energetics: Larger individuals emerge from winter in better condition, Transactions of the American Fisheries Society, 126, Pages 153 – 156. M. K. Cox and K. J. Hartman, 2005 Nonlethal estimation of proximate composition in fish, Canadian Journal of Fisheries and Aquatic Sciences, 62, Pages 269 – 275. J. F. Craig, 1977 The body composition of adult perch, Perca fluviatilis, in Windermere, with reference to seasonal changes and reproduction, Journal of Animal Ecology, 46, Pages 617 – 632. J. F. Craig, M. J. Kenley and J. F. Talling, 1978 Comparative estimations of the energy content of fish tissue from bomb calorimetry, wet oxidation, and proximate analysis, Freshwater Biology, 8, Pages 585 – 590. G. T. Crossin and S. G. Hinch, 2005 A nonlethal, rapid method for assessing the somatic energy content of migrating adult Pacific salmon, Transactions of the American Fisheries Society, 134, Pages 184 – 191. A. J. Danylchuk and M. G. Fox, 1994 Age‐ and size‐dependent variation in the seasonal timing and probability of reproduction among mature female pumpkinseed (Lepomis gibbosus), Environmental Biology of Fishes, 39, Pages 119 – 127. S. I. Doroshov, 1985, “ Biology and culture of sturgeon Acipenseriformes ”, Pages 251 – 274. Edited by: J. F. Muir, R. J. Roberts. In Recent advances in aquaculture, Croom Helm, London. J. D. Dutil, 1986 Energetic constraints and spawning interval in the anadromous Arctic charr (Salvelinus alpinus), Copeia, 1986, Pages 945 – 955. J. M. Elliott, 1976 Body composition of brown trout (Salmo trutta L.) in relation to temperature and ration size, Journal of Animal Ecology, 45, Pages 273 – 289. R. U. Fischer, J. D. Congdon and M. Brock, 1996 Total body electrical conductivity (TOBEC): A tool to estimate lean mass and nonpolar lipids of an aquatic organism?, Copeia, 1996, Pages 459 – 462. U. Focken and K. Becker, 1993, “ Body composition of carp (Cyprinus carpio L.) ”, Pages 269 – 288. Edited by: T. Braunbeck, W. Hanke, H. Segner. In Fish ecotoxicology and ecophysiology, VCH Publishers, New York. J. From and G. Rasmussen, 1984 A growth model, gastric evacuation, and body composition in rainbow trout, Salmo gairdneri Richardson, 1836, Dana, 3, Pages 61 – 139. K. J. Hartman and S. B. Brandt, 1995 Estimating energy density of fish, Transactions of the American Fisheries Society, 124, Pages 347 – 355. IndexNoFollow Natural Resources and Environment Science Article 2008 ftumdeepblue https://doi.org/10.1577/T05-240.1 2023-07-31T20:32:40Z Fish proximate composition and energy density can influence growth, survival, and reproduction, so it is important to develop models to understand the patterns and predict dynamic changes. This paper presents three such models. Model 1 describes the general pattern of changes in lipid, protein, ash, and energy density that occur with changes in water content. The key assumption this model is that there is a fixed amount of water associated with each gram of protein and a much smaller fixed amount of water associated with each gram of lipid. In combination with a mass balance constraint, this explains the commonly observed linear relationship between the fraction lipid and the fraction water. Because energy density varies in direct proportion to the fractions lipid and protein, the linear relationship between body composition and fraction water makes energy density also a linear function of the fraction water. The model is fitted to data for lake trout Salvelinus namaycush and coho salmon Oncorhynchus kisutch for a limited range in wet weight. Model 2 describes the pattern of proximate composition and energy density that occurs with variation in body size. A strong pattern was found between the mass of water and the mass of protein, suggesting strict control of body water. The model is fitted to data for common carp Cyprinus carpio and bluegill Lepomis macrochirus. This analysis shows that the relationship between body composition, energy density, and fraction water is expected to vary with body size because both the water: Protein ratio and the fraction ash change with body size. Model 3 demonstrates how this approach can be used to predict changes in fish body composition and energy density during starvation, as might be done with a bioenergetics model. This model is fitted to data from a starvation experiment involving largemouth bass Micropterus salmoides. Peer Reviewed https://deepblue.lib.umich.edu/bitstream/2027.42/141335/1/tafs0340.pdf Article in Journal/Newspaper Arctic University of Michigan: Deep Blue Transactions of the American Fisheries Society 137 1 340 356