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What is Bioenergetics?

What is Bioenergetics?. “…..the study of the flow and transformation of energy in and between living organisms and between living organisms and their environment”. Consumption = Metabolism + Waste + Growth. MUST!. What Goes In = What Comes Out. Why bioen is useful to fish ecology?

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What is Bioenergetics?

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  1. What is Bioenergetics? “…..the study of the flow and transformation of energy in and between living organisms and between living organisms and their environment”

  2. Consumption = Metabolism + Waste + Growth MUST! What Goes In = What Comes Out

  3. Why bioen is useful to fish ecology? • It is a mathematical representation of how a fish grows • Allows researchers, students and managers a way to understand …..

  4. Typical Energy Budgets Differ for Carnivores & Herbivores: Green Sunfish Muskellunge Largescale Stoneroller

  5. Bioenergetics ~ Economics Consumption = Metabolism + Waste + Growth Consumption = Income Metabolism = Rent (temp. and ritz) Wastes & Losses = Taxes Growth = Savings and Investments

  6. June 1 = June 1 Sept 1 Sept 1 ? Lake Salad Lake Cheeseburger Why would growth be different?

  7.  More food? Better quality food? Don’t work as hard for food? Lakes are different temperatures? Stress, contaminants, food webs Why would growth be different?

  8. Active Metabolism  Costs from activity Consumption ΔBiomass  Growth Respiration  Basal Metabolism Specific Dynamic Action  Costs from digestion Egestion-F& Excretion -U Gonads Reproduction (R + A + S) +(F + U) + (ΔB + G) C = Model Components:

  9. (R + A + S) +(F + U) + (ΔB + G) C = Independent C = consumption Losses R = respiration A = active metabolism S = specific dynamic action Losses F= egestion U = excretion Gains ΔB = Change in biomass (somatic growth) G = gonads /reproduction

  10. All processes are temp. and size dependent grams prey grams of body mass day 0.08 0.06 Specific Rate (g/g/d) 0.04 Grams body mass stays the same, so how much you eat is what is changing 0.02 0.00 30 15 25 20 10 5 Temperature (C)

  11. All processes are temp. and size dependent Too hot,body starts to fail SDA Max. Consumption = Cmax egestion excretion starvation Upper lethal growth respiration loss of growth “Golden Banana” 0.08 0.06 Specific Rate (g/g/d) 0.04 0.02 0.00 30 15 25 20 10 5 Temperature (C)

  12. Maximum consumption isn't realisticp-value = proportion of maximum consumption Maximum 0.08 80% of Maximum SDA 0.06 Specific Rate (g/g/d) egestion Pvalue = .80 excretion 0.04 growth 0.02 respiration 0.00 30 15 25 20 10 5 Temperature (C)

  13. Maximum consumption isn't realisticp-value = proportion of maximum consumption Maximum 0.08 Realistic Value 0.06 Specific Rate (g/g/d) P-value = .35 35% of Maximum SDA 0.04 egestion excretion growth 0.02 respiration 0.00 30 15 25 20 10 5 Temperature (C) starvation Upper lethal loss of growth

  14. How do we estimate these basic relationships? Yellow Perch Feed ad libitum – as much as possible Hot TooHot moderate Cold grams/grams/day Temperature Like the relationship between temperature and maximum consumption…..lab experiments and or field data

  15. Consumption and Respiration  Temperature Dependent Consumption = Cmax Respiration 0.08 0.06 Specific Rate (g/g/d) starvation Upper lethal 0.04 0.02 0.00 loss of growth 30 15 25 20 10 5 Temperature (C)

  16. Consumption and Respiration  Size Dependent 0.8 8 0.6 6 Specific Consumption g/g/d Maximum Consumptiong/d 0.4 4 0.2 2 5 10 15 25 20 Weight (g) FOR A GIVEN TEMPERATURE

  17. Solve for Consumption Can also organize equation in terms of growth.. ΔB = C - (R + A + S) -(F + U) - G Main equation can be organized in 2 ways (R + A + S) +(F + U) + (ΔB + G) C = Can solve for any of the different parameters of the equations….hence the power of bioenergetics

  18. What do we need to run the model?

  19. Length Frequency Growth: JUNE 1 40mm 16 12 8 # Caught 4 0 10 30 50 70 90 110 130 150 170 190 Length Category (10cm) SEPT 1 16 70mm 12 # Caught 8 4 0 10 30 50 70 90 110 130 150 170 190 Length Category (10cm) Mark and recap

  20. Measure growth: Calcified structuresfish parts: scales, otoliths, vertebrae, rays, opercula 15 mm – Age0 76 mm – Age1 123 mm – Age2

  21. What else do we need to run the model? Temperatures where fish live (optimal temperature, lethal limits) • alewife - 20° C • bluegill - 29° C • coho salmon - 15° C • largemouth bass – 27.5° C • muskellunge - 26° C • northern pike - 24° C • rainbow smelt - 13° C • rainbow trout - 20° C • striped bass – 21.6° C • walleye - 22° C • yellow perch - 26° C • smallmouth bass – 29 ° C • sea lamprey - 18° C • chinook salmon - 15° C

  22. What do we need to run the model? What a fish eats (Diets!)…

  23. What do we need to run the model? Prey and Predator Energy Densities … Snails – 1800 j/g dry mass Zooplankton – 2513 j/g wet mass Crayfish – 3766 j/g wet mass Yellow Perch – 5000 j/g dry mass Alewife – 7225 j/g wet mass Leech – 2400 j/g dry mass

  24. What do we need to run the model? Basic physiological parameters… • Egestion (size/temp dependent)  F • Excretion (size/temp dependent)  U • Specific Dynamic Action  SDA • Basal Metabolism  R • Active Metabolism  A Where do we get all these….? - We do painstakingly difficult lab experiments (imagine having to measure fish excrement or…)- We steal them, I mean “borrow” them!- Species borrowing is common, it can cause problems- Should evaluate and test if borrowing is appropriate

  25. How have bioenergetic models been used in fisheries?

  26. Sea Lamprey Control and Lake Trout Control of sea lamprey will result in a 5% increase in lake trout survival How much will rainbow smelt mortality increase? Calculate increase in trout biomass, then use bioenergetics to calculate increase in consumption of rainbow smelt Rainbow smelt mortality will increase by 3 percent (La Bar 1993)

  27. Alewives and P Cycling in Lake Michigan Alewives may promote phytoplankton growth by recycling P through egestion and excretion Question: How much do alewives in Lake Michigan contribute to phosphorus regeneration? Bioenergetics nutrient analysis was used to find that 12,000 metric tons of P were regenerated per year Strong year classes may contribute to algal blooms (Kraft 1993)

  28. Climate Change and Sockeye Salmon Production Climate change may result in increased temperatures and less upwelling How will this affect production of sockeye salmon? If temperatures increase 2-4 degrees C, the combination of lower food availability and higher temperatures would result in a 10-30% decrease in adult mass may also lead to lower juvenile survival (Hinch et al 1995)

  29. What do bioenergetis have to do with the University of Wisconsin? Kitchell, J.F., and J.T. Windell. 1968. Rate of gastric digestion in pumpkinseed sunfish, Lepomis gibbosus. Trans. Am. Fish. Soc. 97(4):489-492. Kitchell, J.F. 1969. Thermophilic and thermophobic responses of snakes in a thermal gradient. Copeia 1969(1):189-191. Windell, J.T., D.O. Norris, J.F. Kitchell, and J.S. Norris. 1969. Digestive response of rainbow trout, Salmo gairdneri, to pellet diets. J. Fish. Res. Board Can. 26(7):1801-1812. Kitchell, J.F., and J.T. Windell. 1970. Nutritional value of algae to bluegill sunfish, Lepomis macrochirus. Copeia 1970(1):186-190. Windell, J.T., and J.F. Kitchell. 1970. A laboratory-discussion exercise on earthworm behavior. BSCS Newsletter. Soc. 41:2-6. Windell, J.T., D.O. Norris, J.H. Bock, and J.F. Kitchell. 1971. General Biology Laboratory II. Kendall/Hunt Publ. Co., Dubuque, Iowa. 129 p. Kitchell, J.F., and J.T. Windell. 1972. Energy budget for the lizard, Anolis carolinensis. Physiol. Zool. 45:178-188. Baumann, P.C., J.F. Kitchell, J.J. Magnuson, and T.B. Kayes. 1974. Lake Wingra, 1837-1973: a case history of human impact. Trans. Wis. Acad. Sci. Arts Lett. 62:57-94. Baumann, P.C., and J.F. Kitchell. 1974. Diel patterns of distribution and feeding of bluegill, Lepomis macrochirus, in Lake Wingra, Wisconsin. Trans. Am. Fish. Soc. 103:255-260. Kitchell, J.F., J.F. Koonce, R.V. O'Neill, H.H. Shugart, Jr., J.J. Magnuson, and R.S. Booth. 1974. Model of fish biomass dynamics. Trans. Am. Fish. Soc. 103:786-798. McCormick, A.J.A., O.L. Loucks, J.F. Koonce, J.F. Kitchell, and P.R. Weiler. 1974. An ecosystem model for the pelagic zone of a lake. In V.J. Yannacone, ed., Proceedings American Bar Association National Institute Environmental Litigation. American Bar Association, Chicago. Stein, R.A., J.F. Kitchell and B. Knezevic. 1975. Selective predation by carp (Cyprinus carpio L.) on benthic molluscs in Skadar Lake, Yugoslavia. J. Fish Biol. 1975:391-399. Kitchell, J.F., J.F. Koonce, and P.S. Tennis. 1975. Phosphorus flux through fishes. Verh. Internat. Verein. Limnol. 19:2478-2484. Windell, J.T., J.F. Kitchell, D.O. Norris, J.S. Norris, and J.W. Foltz. 1976. Temperature and rate of gastric evacuation by rainbow trout, Salmo gairdneri. Trans. Am. Fish. Soc. 105:712-717. Kitchell, J.F., M. Johnson, L. Greig, K. Loftus, K. Minns, and C. Olver. 1977. Percid habitat: the river analogy. J. Fish. Res. Board Can. 34:1936-1940. Kitchell, J.F., D.J. Stewart, and D. Weininger. 1977. Applications of a bioenergetics model to perch (Perca flavescens) and walleye (Stizostedion vitreum). J. Fish. Res. Board Can. 34:1922-1935. Kitchell, J.F., J.J. Magnuson, and W.H. Neill. 1977. Estimation of caloric equivalents for fish biomass. Environ. Biol. Fish. 2:185-189. Bartell, S.M., and J.F. Kitchell. 1978. Seasonal impact of planktivory on phosphorus release by Lake Wingra zooplankton. Verh. Internat. Verein. Limnol. 20:466-474. Gallepp, G.W., J.F. Kitchell, and S.M. Bartell. 1978. Phosphorus released from lake muds affected by chironomids. Verh. Internat. Verein. Limnol. 20:458-474. Kitchell, J.A., J.F. Kitchell, D.L. Clark, and L. Dangeard. 1978. Deep sea foraging behavior: its bathymetric potential in the fossil record. Science 200:1289-1291. Kitchell, J.F., R.A. Stein, and B. Knezevic. 1978. Utilization of filamentous algae by fishes in Skadar Lake, Yugoslavia. Verh. Internat. Verein. Limnol. 20:2159-2165. Kitchell, J.A., J.F. Kitchell, G.L. Johnson, and K. Hunkins. 1978. Abyssal traces and megafauna: comparison of productivity, diversity and density in the Arctic and Antarctic. Paleobiology 4:171-180. Kitchell, J.F., W.H. Neill, A.E. Dizon, and J.J. Magnuson. 1978. Bioenergetic spectra of skipjack and yellowfin tunas. Pages 357-368 in G.D. Sharp and A.E. Dizon (eds.), Physiological Ecology of Tunas. Academic Press, New York. Kitchell, J.F. 1979. In search of an optimal environment. Pages 31-33 in R.H. Stroud and H.E. Clepper (eds.), Predator-Prey Systems in Fisheries Management. Sport Fishing Institute, Washington, D.C. Kitchell, J.F., R.V. O'Neill, D. Webb, G.W. Gallepp, S.M. Bartell, J.F. Koonce, and B.S. Ausmus. 1979. Consumer regulation of nutrient cycling. BioScience 29:28-34. Lozano, S.L. ,and J.F. Kitchell. 1979. Response of a mobile invertebrate to heterothermal conditions. Pages 511-521 in J.H. Thorpe and J.W. Gibbons (eds.), Energy and Environmental Stress in Aquatic Systems. Selected papers from a symposium held at Augusta, Georgia. Technical Information Center, U.S. Department of Energy. Breck, J.E., and J.F. Kitchell. 1980. Effects of macrophyte harvesting on simulated predator-prey interactions. Pages 211-228 in J.E. Breck et al. (eds.), Aquatic Plants, Lake Management, and Ecosystem Consequence of Lake Harvesting. Proceedings of a conference at Madison, Wisconsin. Inst. for Env. Studies, UW-Madison. 435 pages. Loucks, O.L., M.S. Adams, J.E. Breck, J.F. Kitchell, R.D. Koegel, D. F. Livermore, R.T. Pretki, and J.E. Ross. 1980. Conference findings: an overview. Pages 421-434 in J.E. Breck et al. (eds.), Aquatic Plants, Lake Management, and Ecosystem Consequences of Lake Harvesting. Proceedings of a conference at Madison, Wisconsin. Inst. for Env. Studies, University of Wisconsin-Madison. 435 p. Kitchell, J.A., and J.F. Kitchell. 1980. Size selective predation, light transmission, and oxygen stratification: evidence from the recent sediments of manipulated lakes. Limnol. and Oceanogr. 25:389-402. Kitchell, J.F. 1980. Fish dynamics and phosphorus cycling in lakes. Pages 81-91 in D. Scavia and R. Moll (eds.), Nutrient Cycling in the Great Lakes: a Summarization of Factors Regulating the Cycling of Phosphorus. Special Report No. 83. NOAA, Great Lakes Env. Res. Lab., Ann Arbor, Michigan. 140 p. Kitchell, J.F., and J.E. Breck. 1980. A bioenergetics model and foraging hypothesis for sea lamprey. Can. J. Fish. Aquat. Sci. 37:2159-2168. Walters, C.J., G. Spangler, W.J. Christie, P. Marion, and J.F. Kitchell. 1980. A synthesis of knowns, unknowns and policy recommendations from the Sea Lamprey International Symposium. Can. J. Fish. Aquat. Sci. 37:2202-2208. Backiel, T., J. Thorpe, and J.F. Kitchell. 1981. Secondary production--fish. Pages 307-340 in E.D. LeCren and R.H. Lowe-McConnell (eds.), Functioning of Freshwater Ecosystems. Cambridge University Press, London. Kitchell, J.A., C.H. Boggs, J.F. Kitchell, and J.A. Rice. 1981. Prey selection by naticid gastropods: experimental tests and application to the fossil record. Paleobiology 7(4):533-552. Stewart, D.J., J.F. Kitchell, and L.B. Crowder. 1981. Forage fishes and their salmonid predators in Lake Michigan. Trans. Am. Fish. Soc. 110:751-763. Kitchell, J.F. 1983. Energetics. Pages 312-338 in P. Webb and D. Weihs (eds.), Fish Biomechanics. Praeger Publishers, New York. 398 p. Rice, J.A., J.E. Breck, S.M. Bartell, and J.F. Kitchell. 1983. Evaluating the constraints of temperature, activity and consumption on growth of largemouth bass. Env. Biol. Fish. 9:263-275. Boggs, C.B., J.A. Rice, J.A. Kitchell, and J.F. Kitchell. 1984. Predation at a snail's pace: what's time to a gastropod? Oecologia 62:13-17. Carpenter, S.R., and J.F. Kitchell. 1984. Plankton community structure and limnetic primary production. American Naturalist 124:159-172. Kitchell, J.F., H.F. Henderson, E. Grygierek, J. Hrbacek, S. Kerr, M.Pedini, T. Petr, J. Shapiro, R. Stein, J. Stenson, and T. Zaret. 1984. Management of Lakes by Food-Chain Manipulation. Food and Agricultural Organization of the United Nations, Rome, Italy. 83 p. Carpenter, S.R., J.F. Kitchell, and J.R. Hodgson. 1985. Cascading trophic interactions and lake productivity. BioScience 35:634-639. Kitchell, J.F. 1985. An ecological rationale for managing predator-prey systems in the Great Lakes. Pages 85-97 in R. L. Eshenroder (ed.), Presented Papers from the Council of Lake Committees Plenary Session on Great Lakes Predator-Prey Issues, March 20, 1985. Great Lakes Fishery Commission, Special Publication 85-3. Father of modern bioenergetics

  30. Carpenter, S.R., and J.F. Kitchell. 1993. Simulation models of the trophic cascade: predictions and evaluations. Pages 310-331in S.R. Carpenter and J.F. Kitchell (eds.), The Trophic Cascade in Lakes. Cambridge Univ. Press, Cambridge, England.Kitchell, J.F., and S.R. Carpenter. 1993. Synthesis and new directions. Pages 332-350in S.R. Carpenter and J.F. Kitchell (eds.), The Trophic Cascade in Lakes. Cambridge Univ. Press, Cambridge, England.Kitchell, J. F., S. R. Carpenter, J. R. Hodgson, X. He and P. A. Soranno. 1993. Phosphorus in food webs: Compensatory responses in experimental lakes. Verh. Internat. Verein. Limnol. 25: 344-348.Carpenter, S. R., X. He, J. F. Kitchell and P. A. Soranno. 1993. Whole lake experiments: Is grazer functional response stable? Verh. Internat. Verein. Limnol. 25: 398-400.Hansen, M. J., D. Boisclair, S. B. Brandt, S. W. Hewett, J. F. Kitchell, M. C. Lucas, and J. J. Ney. 1993. Applications of bioenergetics models to fish ecology and management: Where do we go from here? Trans. Amer. Fish. Soc. 122: 1019-1030.Schindler, D. E., J. F. Kitchell, X. He, S. R. Carpenter, J. R. Hodgson and K. L. Cottingham. 1993. Food web structure and phosphorus cycling in lakes. Trans. Amer. Fish. Soc. 122:756-772.He, X., J. F. Kitchell, S. R. Carpenter, J. R. Hodgson and D. E. Schindler. 1993. Food web configuration and long-term phosphorus recycling: A simulation model evaluation. Trans. Amer. Fish. Soc. 122: 773-783.He, X., J. . Hodgson, J. R., J. F. Kitchell and R. A. Wright. 1994. Growth and diet composition of largemouth bass (Micropterus salmoides) from four experimental lakes. Verh. Internat. Verein. Limnol. 25: 2092-2097.Schindler, D. E., B. M. Johnson, N. A. MacKay, N. Bouwes and J. F. Kitchell. 1994. Crab:snail size-structured interactions and salt marsh predation gradients. Oecologia 97: 49-61. Carpenter, S. R., T. M. Frost, A. R. Ives, J. F. Kitchell and T. K. Kratz. 1994. Complexity, cascades and compensation in ecosystems. In Yasuna, M. and M. M. Watanabe (eds.) Biodiversity: Its Complexity and Role. Global Environmental Forum, Tokyo, Japan. Pp. 197-20Kitchell, J. F., E. A. Eby, X. He, D. E. Schindler and R. M. Wright. 1994. Predator-prey dynamics in an ecosystem context. J. Fish Biol. 45:209-226.Kitchell, J.F., and L.B. Crowder. 1986. Predator-prey interactions in Lake Michigan: model predictions and recent dynamics. Env. Biol. Fish. 16(1-3):205-211.Kraft, C.E., and J.F. Kitchell. 1986. Partitioning of food resources by sculpins in Lake Michigan. Env. Biol. Fish. 16(4):309-316.Kitchell, J.A., C.H. Boggs, J.A. Rice, J.F. Kitchell, A. Hoffman, and J. Martinell. 1986. Anomalies in naticid predatory behavior: a critique and experimental observations. Malacalogia 27(2):291-298.Kitchell, J.F., and S.R. Carpenter. 1987. Piscivores, planktivores, fossils and phorbins. Pages 132-146 in W.C. Kerfoot and A. Sih (eds.), Predation: Direct and Indirect Impacts on Aquatic Communities. University Press of New England, Hanover, N.H.Carpenter, S.R., and J.F. Kitchell. 1987. The temporal scale of variance in limnetic primary productivity. Am. Nat. 129(3):417-433.Hodgson, J.R., and J.F. Kitchell. 1987. Opportunistic foraging by largemouth bass (Micropterus salmonides). Am. Midl. Nat. 118(2):323-336.Carpenter, S.R., J.F. Kitchell, J.R. Hodgson, P.A. Cochran, J.J. Elser, M.M. Elser, D.M. Lodge, D. Kretchmer, X. He and C.N. von Ende. 1987. Regulation of lake ecosystem productivity by food web structure. Ecology 68(6):1863-1876.Kitchell, J.F., and S.W. Hewett. 1987. Forecasting forage demand and yield of sterile chinook salmon in Lake Michigan. Can. J. Fish. Aquat. Sci. 44(Suppl. 2):384-389.Carpenter, S.R., and J.F. Kitchell. 1988. Introduction. Pages 1-8. in S.R. Carpenter (ed.), Complex Interactions in Lake Communities. Springer-Verlag, New York Carpenter, S.R., and J.F. Kitchell. 1988. Consumer control of lake productivity. BioScience 38:764-769.Hartig, J.H., J.F. Kitchell, D. Scavia, and S.B. Brandt. 1988. Rehabilitation of Lake Ontario: the role of nutrient reduction and food web dynamics. Report to the Great Lakes Science Advisory Board, International Joint Commission. 75 p.Kitchell, J.F., and S.R. Carpenter. 1988. Food web manipulation in experimental lakes. Verh. Internat. Verein. Limnol. 23:351-358.Kitchell, J.F., M.S. Evans, D. Scavia, and L.B. Crowder. 1988. Food web regulation of water quality in Lake Michigan: report of a workshop. J. Great Lakes Res. 14:109-114.Scavia, D., G.A. Lang, and J.F. Kitchell. 1988. Dynamics of Lake Michigan plankton: a model evaluation of nutrient loading, competition and predation. Can. J. Fish. Aquat. Sci. 45:165-177.Kitchell, J.F., S.M. Bartell, S.R. Carpenter, D.J. Hall, D.J. McQueen, W.E. Neill, D. Scavia, and E.E. Werner. 1988. Epistemology, experimentation and pragmatism. Pages 263-280 in S.R. Carpenter (ed.), Complex Interactions in Lake Communities. Springer-Verlag, New York Leavitt, P.R., S.R. Carpenter, and J.F. Kitchell. l989. Whole lake experiments: the annual record of fossil pigments and zooplankton. Limnol. Oceanogr. 34:700-717.Cochran, P.A., and J.F. Kitchell. l989. A model of feeding by parasitic lampreys. Can. J. Fish. Aquat. Sci. 46:1845-52.Crowder, L., R. Francis, E. Fritz, D. Hoss, J. Kitchell, R. Lasker, P. Ortner, K. Sherman, and G. Stauffer. l989. Recruitment Fisheries Oceanography Program Development Plan. National Oceanic and Atmospheric Administration, Washington D.C. 102 p.Vanni, M.J., C. Luecke, J.F. Kitchell, and J.J. Magnuson. 1990. Effects of planktivorous fish mass mortality on the plankton community of Lake Mendota, Wisconsin: implications for biomanipulation. Hydrobiologia 200/201:329-336.Kitchell, J.F. l990. Fisheries in the Great Lakes: a brief history as related to research and management programs for Lake Champlain. Water Resources Res. Reports, Univ. Vermont. Pp. 97-108.Kitchell, J.F. l990. The scope for mortality caused by sea lamprey. Trans. Amer. Fish. Soc. 119:642-648.Vanni, M.J., C. Luecke, J.F. Kitchell, Y. Rentmeester, J. Temte and J.J. Magnuson. 1990. Cascading trophic interactions and phytoplankton abundance in a eutrophic lake: effects of massive fish mortality. Nature 344:333-335.66. He, X., and J.F. Kitchell. 1990. Direct and indirect effects of predation on a fish community: a whole lake experiment. Trans. Amer. Fish Soc. 119:825-835.67. Luecke, C., M.J. Vanni, J.J. Magnuson, J.F. Kitchell, and P.T. Jacobson. 1990. Seasonal regulation of Daphnia populations by planktivorous fish: implications for the spring clear water phase. Limnol. Oceanogr. 35(8):1718-1733.Hartig, J.H., J.F. Kitchell, D. Scavia, and S.B. Brandt. 1991. Rehabilitation of Lake Ontario: the role of nutrient reduction and food web dynamics. Can. J. Fish. Aquat. Sci. 48:1574-1580.Carpenter, S.R., T.M. Frost, J.F. Kitchell, T.K. Kratz, D.W. Schindler, J. Shearer, W.G. Sprules, M.J. Vanni, and A.P. Zimmerman. l991. Patterns of primary production and herbivory in 25 North American lake ecosystems. Pages 67-96.in J. Cole, S. Findlay, and G. Lovett (eds.), Comparative Analyses of Ecosystems: Patterns, Mechanisms and Theories. Springer-Verlag, New York. Kitchell, J.F. l991. Salmonid carrying capacity: estimates and experiences in the Great Lakes of North America. Pages 59-66 in. R.J. Svrjcek (ed.), Marine Ranching: Proceedings of the Seventeenth U.S.-Japan Meeting on Aquaculture, Ise, Japan, 1988. NOAA Technical Report NMFS 102. National Marine Fisheries Service, [Washington, D.C.].Kitchell, J.F., S.R. Carpenter, S.W. Bayley, K.C. Ewel, R.W. Howarth, S.M. Nixon, and D.W. Schindler. 1991. Aquatic ecosystem experiments in the context of global climate change: working group report. Pages 229-235.in H.A. Mooney et al. (eds.), Ecosystem Experiments. John Wiley and Sons, Sussex, England. Boggs, C.H., and J.F. Kitchell. l991. Tuna metabolic rate estimated from energy losses during starvation. Physiol. Zool. 64:502-524.Carpenter, S.R., S.G. Fisher, N.B. Grimm, and J.F. Kitchell. l992. Global change and freshwater ecosystems. Ann. Rev. Ecol. Syst. 23:119-139.Carpenter, S.R., and J.F. Kitchell. l992. Trophic cascade and biomanipulation: interface of research and management. Limnol. Oceanogr.37:208-213.Kitchell, J.F. (ed.). l992. Food Web Management: A Case Study of Lake Mendota. Springer-Verlag, New York. 553 p.Kitchell, J.F. l992. Introduction: the rationale and goals for food web management in Lake Mendota. Pages 1-5 in J.F.Kitchell (ed.). Food Web Management: A Case Study of Lake Mendota. Springer-Verlag, New York.Kitchell, J.F. and P.R. Sanford. l992. Paleolimnological evidence of food web structure. Pages 31-47 in J.F.Kitchell (ed.). Food Web Management: A Case Study of Lake Mendota. Springer-Verlag, New York.Kitchell, J.F., and S.R. Carpenter. l992. Summary: accomplishments and new directions of food web management in Lake Mendota. Pages 539-544 in J.F.Kitchell (ed.). Food Web Management: A Case Study of Lake Mendota. Springer-Verlag, New York.Johnson, B.M., R.S. Stewart, C. Luecke, M.D. Staggs, S.J. Gilbert and J.F. Kitchell. 1992. Forecasting effects of harvest regulations and stocking on prey fish communities in Lake Mendota, Wisconsin. N. Amer. J. Fish. Mgmt. 12:797-807.Johnson, B. L., S. R. Milliman, R. C. Bishop and J. F. Kitchell. 1992. Evaluating fishery rehabilitation under uncertainty: a bioeconomic analysis of quota management for the Green Bay yellow perch fishery. N. Amer. J. Fish. Mgmt. 12: 703-720.Carpenter, S.R., T.M. Frost, T.K. Kratz, and J.F. Kitchell. 1993. Species dynamics and global environmental change: a perspective from ecosystem experiments. Pages 267-269 in J. Kingsolver, P. Kareiva, and R. Huey (eds.), Biotic Interactions and Global Change. Sinauer Associates, Sunderland, Mass.Kitchell, J. F. and S. R. Carpenter. l993. Variability in lake ecosystems: complex responses by the apical predator. Pages 111-124 in M. McDonnell and S. Pickett (eds.) Humans as Components of Ecosystems. Springer-Verlag, New York.Carpenter, S.R., and J.F. Kitchell (eds.). 1993. The Trophic Cascade in Lakes. Cambridge Univ. Press, Cambridge, England.Kitchell, J.F., and S.R. Carpenter. Cascading trophic interactions. 1993. Pages 1-14 in S.R. Carpenter and J.F. Kitchell (eds.), The Trophic Cascade in Lakes. Cambridge Univ. Press, Cambridge, England.Carpenter, S.R., and J.F. Kitchell. 1993. Experimental lakes, manipulations and measurements. Pages 15-25in S.R. Carpenter and J.F. Kitchell (eds.), The Trophic Cascade in Lakes. Cambridge Univ. Press, Cambridge, England.Hodgson, J.R., X. He, and J.F. Kitchell. The fish populations. 1993. Pages 43-68 in S.R. Carpenter and J.F. Kitchell (eds.), The Trophic Cascade in Lakes. Cambridge Univ. Press, Cambridge, England.He, X., R.A. Wright, and J.F. Kitchell. Fish behavioral and community responses to manipulation. 1993. Pages 69-84in S.R. Carpenter and J.F. Kitchell (eds.), The Trophic Cascade in Lakes. Cambridge Univ. Press, Cambridge, England.He, X., J.F. Kitchell, J.R. Hodgson, R.A. Wright, P.A. Soranno, D.M. Lodge, P.A. Cochran, D.A. Benkowski, and N.A. Bouwes. 1993. Roles of fish predation: piscivory and planktivory. Pages 85-102in S.R. Carpenter and J.F. Kitchell (eds.), The Trophic Cascade in Lakes. Cambridge Univ. Press, Cambridge, England.Leavitt, P.R., P.A. Sanford, S.R. Carpenter, J.F. Kitchell, and D.A. Benkowski. Annual fossil pigment records of food web manipulation. 1993. Pages 278-309in S.R. Carpenter and J.F. Kitchell (eds.), The Trophic Cascade in Lakes. Cambridge Univ. Press, Cambridge, England.4

  31. Kitchell, J.F. l990. The scope for mortality caused by sea lamprey. Trans. Amer. Fish. Soc. 119:642-648. Vanni, M.J., C. Luecke, J.F. Kitchell, Y. Rentmeester, J. Temte and J.J. Magnuson. 1990. Cascading trophic interactions and phytoplankton abundance in a eutrophic lake: effects of massive fish mortality. Nature 344:333-335. 66. He, X., and J.F. Kitchell. 1990. Direct and indirect effects of predation on a fish community: a whole lake experiment. Trans. Amer. Fish Soc. 119:825-835. 67. Luecke, C., M.J. Vanni, J.J. Magnuson, J.F. Kitchell, and P.T. Jacobson. 1990. Seasonal regulation of Daphnia populations by planktivorous fish: implications for the spring clear water phase. Limnol. Oceanogr. 35(8):1718-1733. Hartig, J.H., J.F. Kitchell, D. Scavia, and S.B. Brandt. 1991. Rehabilitation of Lake Ontario: the role of nutrient reduction and food web dynamics. Can. J. Fish. Aquat. Sci. 48:1574-1580. Carpenter, S.R., T.M. Frost, J.F. Kitchell, T.K. Kratz, D.W. Schindler, J. Shearer, W.G. Sprules, M.J. Vanni, and A.P. Zimmerman. l991. Patterns of primary production and herbivory in 25 North American lake ecosystems. Pages 67-96.in J. Cole, S. Findlay, and G. Lovett (eds.), Comparative Analyses of Ecosystems: Patterns, Mechanisms and Theories. Springer-Verlag, New York. Kitchell, J.F. l991. Salmonid carrying capacity: estimates and experiences in the Great Lakes of North America. Pages 59-66 in. R.J. Svrjcek (ed.), Marine Ranching: Proceedings of the Seventeenth U.S.-Japan Meeting on Aquaculture, Ise, Japan, 1988. NOAA Technical Report NMFS 102. National Marine Fisheries Service, [Washington, D.C.]. Kitchell, J.F., S.R. Carpenter, S.W. Bayley, K.C. Ewel, R.W. Howarth, S.M. Nixon, and D.W. Schindler. 1991. Aquatic ecosystem experiments in the context of global climate change: working group report. Pages 229-235.in H.A. Mooney et al. (eds.), Ecosystem Experiments. John Wiley and Sons, Sussex, England. Boggs, C.H., and J.F. Kitchell. l991. Tuna metabolic rate estimated from energy losses during starvation. Physiol. Zool. 64:502-524. Carpenter, S.R., S.G. Fisher, N.B. Grimm, and J.F. Kitchell. l992. Global change and freshwater ecosystems. Ann. Rev. Ecol. Syst. 23:119-139. Carpenter, S.R., and J.F. Kitchell. l992. Trophic cascade and biomanipulation: interface of research and management. Limnol. Oceanogr.37:208-213. Kitchell, J.F. (ed.). l992. Food Web Management: A Case Study of Lake Mendota. Springer-Verlag, New York. 553 p. Kitchell, J.F. l992. Introduction: the rationale and goals for food web management in Lake Mendota. Pages 1-5 in J.F.Kitchell (ed.). Food Web Management: A Case Study of Lake Mendota. Springer-Verlag, New York. Kitchell, J.F. and P.R. Sanford. l992. Paleolimnological evidence of food web structure. Pages 31-47 in J.F.Kitchell (ed.). Food Web Management: A Case Study of Lake Mendota. Springer-Verlag, New York. Kitchell, J.F., and S.R. Carpenter. l992. Summary: accomplishments and new directions of food web management in Lake Mendota. Pages 539-544 in J.F.Kitchell (ed.). Food Web Management: A Case Study of Lake Mendota. Springer-Verlag, New York. Johnson, B.M., R.S. Stewart, C. Luecke, M.D. Staggs, S.J. Gilbert and J.F. Kitchell. 1992. Forecasting effects of harvest regulations and stocking on prey fish communities in Lake Mendota, Wisconsin. N. Amer. J. Fish. Mgmt. 12:797-807. Johnson, B. L., S. R. Milliman, R. C. Bishop and J. F. Kitchell. 1992. Evaluating fishery rehabilitation under uncertainty: a bioeconomic analysis of quota management for the Green Bay yellow perch fishery. N. Amer. J. Fish. Mgmt. 12: 703-720. Carpenter, S.R., T.M. Frost, T.K. Kratz, and J.F. Kitchell. 1993. Species dynamics and global environmental change: a perspective from ecosystem experiments. Pages 267-269 in J. Kingsolver, P. Kareiva, and R. Huey (eds.), Biotic Interactions and Global Change. Sinauer Associates, Sunderland, Mass. Kitchell, J. F. and S. R. Carpenter. l993. Variability in lake ecosystems: complex responses by the apical predator. Pages 111-124 in M. McDonnell and S. Pickett (eds.) Humans as Components of Ecosystems. Springer-Verlag, New York. Carpenter, S.R., and J.F. Kitchell (eds.). 1993. The Trophic Cascade in Lakes. Cambridge Univ. Press, Cambridge, England. Kitchell, J.F., and S.R. Carpenter. Cascading trophic interactions. 1993. Pages 1-14 in S.R. Carpenter and J.F. Kitchell (eds.), The Trophic Cascade in Lakes. Cambridge Univ. Press, Cambridge, England. Carpenter, S.R., and J.F. Kitchell. 1993. Experimental lakes, manipulations and measurements. Pages 15-25in S.R. Carpenter and J.F. Kitchell (eds.), The Trophic Cascade in Lakes. Cambridge Univ. Press, Cambridge, England. Hodgson, J.R., X. He, and J.F. Kitchell. The fish populations. 1993. Pages 43-68 in S.R. Carpenter and J.F. Kitchell (eds.), The Trophic Cascade in Lakes. Cambridge Univ. Press, Cambridge, England. He, X., R.A. Wright, and J.F. Kitchell. Fish behavioral and community responses to manipulation. 1993. Pages 69-84in S.R. Carpenter and J.F. Kitchell (eds.), The Trophic Cascade in Lakes. Cambridge Univ. Press, Cambridge, England. He, X., J.F. Kitchell, J.R. Hodgson, R.A. Wright, P.A. Soranno, D.M. Lodge, P.A. Cochran, D.A. Benkowski, and N.A. Bouwes. 1993. Roles of fish predation: piscivory and planktivory. Pages 85-102in S.R. Carpenter and J.F. Kitchell (eds.), The Trophic Cascade in Lakes. Cambridge Univ. Press, Cambridge, England. Leavitt, P.R., P.A. Sanford, S.R. Carpenter, J.F. Kitchell, and D.A. Benkowski. Annual fossil pigment records of food web manipulation. 1993. Pages 278-309in S.R. Carpenter and J.F. Kitchell (eds.), The Trophic Cascade in Lakes. Cambridge Univ. Press, Cambridge, England. Carpenter, S.R., and J.F. Kitchell. 1993. Simulation models of the trophic cascade: predictions and evaluations. Pages 310-331in S.R. Carpenter and J.F. Kitchell (eds.), The Trophic Cascade in Lakes. Cambridge Univ. Press, Cambridge, England. Kitchell, J.F., and S.R. Carpenter. 1993. Synthesis and new directions. Pages 332-350in S.R. Carpenter and J.F. Kitchell (eds.), The Trophic Cascade in Lakes. Cambridge Univ. Press, Cambridge, England. Kitchell, J. F., S. R. Carpenter, J. R. Hodgson, X. He and P. A. Soranno. 1993. Phosphorus in food webs: Compensatory responses in experimental lakes. Verh. Internat. Verein. Limnol. 25: 344-348. Carpenter, S. R., X. He, J. F. Kitchell and P. A. Soranno. 1993. Whole lake experiments: Is grazer functional response stable? Verh. Internat. Verein. Limnol. 25: 398-400. Hansen, M. J., D. Boisclair, S. B. Brandt, S. W. Hewett, J. F. Kitchell, M. C. Lucas, and J. J. Ney. 1993. Applications of bioenergetics models to fish ecology and management: Where do we go from here? Trans. Amer. Fish. Soc. 122: 1019-1030. Schindler, D. E., J. F. Kitchell, X. He, S. R. Carpenter, J. R. Hodgson and K. L. Cottingham. 1993. Food web structure and phosphorus cycling in lakes. Trans. Amer. Fish. Soc. 122:756-772. He, X., J. F. Kitchell, S. R. Carpenter, J. R. Hodgson and D. E. Schindler. 1993. Food web configuration and long-term phosphorus recycling: A simulation model evaluation. Trans. Amer. Fish. Soc. 122: 773-783. He, X., J. . Hodgson, J. R., J. F. Kitchell and R. A. Wright. 1994. Growth and diet composition of largemouth bass (Micropterus salmoides) from four experimental lakes. Verh. Internat. Verein. Limnol. 25: 2092-2097. Schindler, D. E., B. M. Johnson, N. A. MacKay, N. Bouwes and J. F. Kitchell. 1994. Crab:snail size-structured interactions and salt marsh predation gradients. Oecologia 97: 49-61. Carpenter, S. R., T. M. Frost, A. R. Ives, J. F. Kitchell and T. K. Kratz. 1994. Complexity, cascades and compensation in ecosystems. In Yasuna, M. and M. M. Watanabe (eds.) Biodiversity: Its Complexity and Role. Global Environmental Forum, Tokyo, Japan. Pp. 197-207. Kitchell, J. F., E. A. Eby, X. He, D. E. Schindler and R. M. Wright. 1994. Predator-prey dynamics in an ecosystem context. J. Fish Biol. 45:209-226. Leavitt, P.R., P.R. Sanford, S.R. Carpenter, and J.F. Kitchell. l994. An annual fossil record of production, planktivory and piscivory during whole lake experiments. J. Paleolimnology 11:133-149. Schindler, D. E., S. T. Carpenter; K. L. Cottingham, X. He, J. R. Hodgson, J. F. Kitchell and P. A. Soranno. 1995. Food web structure and littoral zone coupling to pelagic trophic cascades. In G. Polis and K. O. Winemiller (eds). Food Webs: Integration of Patterns and Dynamics. Chapman Hall, NY. Pp. 96-105. Carpenter, S. R., D. L. Christensen, J. J. Cole, K. L Cottingham, X. He, J. R. Hodgson, J. F. Kitchell, S. E. Knight, M. L. Pace, D. M. Post, D. E. Schindler and N. Voichick. 1995. Biological control of eutrophication in lakes. Env. Sci. and Tech. 29: 784-786. Kitchell, J. F. 1995. Toxic contaminants in the Great Lakes: A polythetic view of the issues and evidence. Ecol. Appl. 5:304-307. Rudstam, L.G., P.E. Peppard, T.W. Fratt, R.E.Bruesewitz, D.W.Coble, F.A. Copes and J. F. Kitchell. 1995. Prey consumption by the burbot (Lota lota) population in Green Bay, Lake Michigan, based on a bioenergetics model. Can. J. Fish. Aquat. Sci. 52:1074-1082. Eby, L. A., L. G. Rudstam and J. F. Kitchell. 1995. Predator responses to prey population dynamics: an empirical analysis based on lake trout growth rates. Can. J. Fish. Aquat. Sci. 52.1564-1571. Post, D. M., T. M. Frost and J. F. Kitchell. 1995. Morphological responses by Bosmina longirostris and Eubosmina tubicen to changes in copepod predator populations during a whole-lake acidification experiment. J. Plankton Res. 17:1621-1632. Hansson, S., L. G. Rudstam, J. F. Kitchell, M. Hilden, B. L. Johnson and P. E. Peppard. 1996. Predation rates by North Sea cod (Gadus morhua) - predictions from models on gastric evacuation and bioenergetics. ICES J. Marine Sci. 51:107-114. Cochran, P.S., and J.F. Kitchell. 1986. Use of modeling to investigate feeding strategies of parasitic lampreys. Env. Biol. Fish. 16(1-3):219-223. Leavitt, P.R., P.R. Sanford, S.R. Carpenter, and J.F. Kitchell. l994. An annual fossil record of production, planktivory and piscivory during whole lake experiments. J. Paleolimnology 11:133-149. Schindler, D. E., S. T. Carpenter; K. L. Cottingham, X. He, J. R. Hodgson, J. F. Kitchell and P. A. Soranno. 1995. Food web structure and littoral zone coupling to pelagic trophic cascades. In G. Polis and K. O. Winemiller (eds). Food Webs: Integration of Patterns and Dynamics. Chapman Hall, NY. Pp. 96-105. Carpenter, S. R., D. L. Christensen, J. J. Cole, K. L Cottingham, X. He, J. R. Hodgson, J. F. Kitchell, S. E. Knight, M. L. Pace, D. M. Post, D. E. Schindler and N. Voichick. 1995. Biological control of eutrophication in lakes. Env. Sci. and Tech. 29: 784-786. Kitchell, J. F. 1995. Toxic contaminants in the Great Lakes: A polythetic view of the issues and evidence. Ecol. Appl. 5:304-307. Rudstam, L.G., P.E. Peppard, T.W. Fratt, R.E.Bruesewitz, D.W.Coble, F.A. Copes and J. F. Kitchell. 1995. Prey consumption by the burbot (Lota lota) population in Green Bay, Lake Michigan, based on a bioenergetics model. Can. J. Fish. Aquat. Sci. 52:1074-1082. Eby, L. A., L. G. Rudstam and J. F. Kitchell. 1995. Predator responses to prey population dynamics: an empirical analysis based on lake trout growth rates. Can. J. Fish. Aquat. Sci. 52.1564-1571. Post, D. M., T. M. Frost and J. F. Kitchell. 1995. Morphological responses by Bosmina longirostris and Eubosmina tubicen to changes in copepod predator populations during a whole-lake acidification experiment. J. Plankton Res. 17:1621-1632. Hansson, S., L. G. Rudstam, J. F. Kitchell, M. Hilden, B. L. Johnson and P. E. Peppard. 1996. Predation rates by North Sea cod (Gadus morhua) - predictions from models on gastric evacuation and bioenergetics. ICES J. Marine Sci. 51:107-114.

  32. Carpenter, S. R., J. F. Kitchell, K. L. Cottingham, D. E. Schindler, D. L. Christensen, D. M. Post and N. Voichick. 1996. Chlorophyll variability, nutrient input and grazing: evidence from whole-lake experiments. Ecology:725-735. Kitchell, J. F. 1996. Where do we go from here? An outsider's view of the Symposium on Pacific salmon and Their Ecosystem. Pages 657-663. In D. J. Stouder, P. A. Bisson and R. J. Naiman (eds.). Pacific Salmon and Their Ecosystem: Status and Future Options. 685 pp. Carpenter, S. R., L. M. Jackson, J. F. Kitchell, and C. R. Stow. 1996. That was then, but this is now--A reply to Gilbertson and Ludwig. Ecol. Appl. 6: In press. Johnson, T. B., and J. F. Kitchell. 1996. Long-term changes in zooplantivorous fish community composition: Implications for food webs. Can. J. Fish. Aquat. Sci. 53:2792-2803. Kitchell, J. F., D. E. Schindler, R. Ogutu-Ohwayo and P. M. Reinthal. 1997. The Nile perch in Lake Victoria: interactions between predation and fisheries. Ecol. Appl. 7: 653-664. Post, D. M., and J. F. Kitchell. 1997. Trophic ontogeny and life history effects on interactions between age-0 fish and zooplankton. Arch. Hydrobiol. Spec. Issues Advanc. Limnol. 49:1-12. Schindler, D.E., J.R. Hodgson and J.F. Kitchell. 1997. Density-dependent changes in individual foraging specialization of largemouth bass. Oecologia 110:592-600. Post, D. M., S. R. Carpenter, D. L. Christensen, K. L. Cottingham, J. R. Hodgson, J. F. Kitchell and D. E. Schindler. 1997. Seasonal effects of variable recruitment of a dominant piscivore on pelagic food web structure. Limnol. Oceanogr. 42:722-729. Hansson, S., D. M. Post, J. F. Kitchell and T. S. McComish. 1997. Predation by yellow perch (Perca flavescens) in southern Lake Michigan--a model analysis. International Symposium on Forage Fish Dynamics, Univ. of Alaska Sea Grant, Anchorage, Alaska. Eby, L. A., C. A. Stow, R. J. Hesselberg and J. F. Kitchell. 1997. Modeling changes in growth and diet on polychlorinated biphenyl (PCB) bioaccumulation in bloater (Coregonus hoyi). Ecol. Appl. 7:981-990. Hanson, P.C., T. B. Johnson, D. E. Schindler, and J. F. Kitchell. 1997. Fish Bioenergetics 3.0. Sea Grant Technical Report, University of Wisconsin Sea Grant Institute, Madison, WI. 119 pages. Schindler, D. E., S. R. Carpenter, J. J. Cole, J. F. Kitchell and M. L. Pace. 1997. Food web structure alters carbon exchange between lakes and the atmosphere. Science 277:248-251. Hodgson, J. R., X. He, D. E. Schindler, and J. F. Kitchell. 1997. Diet overlap in a piscivore community. Ecol of Freshwater Fish 6: 34-42. Kitchell, J.F., and M.L. Pace. 1998. Understanding human modifications of the ocean: Opportunities for large-scale experiments. www.joss.ucar.edu/joss_psg/project/oce_workshop/oeurve/kitchell_pace.html Carpenter, S. R., J. J. Cole, J. F. Kitchell and M. L. Pace. 1998. Impact of dissolved organic carbon, phosphorus and grazing on phytoplankton biomass and production in experimental lakes. Limnol. and Oceanogr. 43:73-80. Schindler, D.E., J.F. Kitchell and R. Ogutu-Ohwayo. 1998. Ecological consequences of alternative gill net fisheries for Nile perch in Lake Victoria. Conservation Biology 12:56-64. Post, D. M., J. P. Taylor, J. F. Kitchell, M. H. Olson, D. E. Schindler and B. E. Herwig. 1998. The role of migratory waterfowl as nutrient vectors in a managed wetland. Conservation Biology 12:910-920. Kitchell, J. F. 1998. Physiological ecology. Pages 164-198 In S.I. Dodson (ed.), Ecology. Oxford Univ. Press. 434 p. Mason, D. M., T. B. Johnson and J. F. Kitchell. 1998. Consequences of prey fish community dynamics on lake trout (Salvelinus namaycush) foraging efficiency in Lake Superior. Can. J. Fish. Aquat. Sci.55:1273-1284. Johnson, T.B., D.M. Mason, C.R. Bronte, and J.F. Kitchell. 1998. Estimation of invertebrate production from patterns of forage fish consumption in western Lake Superior. Trans. Amer. Fish. Soc. 127:496-506. Carpenter, S. R., J. Cole, T. E. Essington, J. R. Hodgson, J. Houser, J. F. Kitchell and M. L. Pace. 1998. Evaluating alternative explanations in ecosystem experiments. Ecosystems 1:335-344. Post, D. M., J. F. Kitchell and J. R. Hodgson. 1998. Interactions of spawning date, growth rates, predation and overwinter mortality in largemouth bass recruitment. Can. J. Fish. Aquat. Sci. 55: 2588-2600. Essington, T. E., and J. F. Kitchell. 1999. New perspectives in the analysis of fish distributions: A case study on the spatial distribution of largemouth bass. Can. J. Fish. Aquat. Sci. 56:52-60. Johnson, T. B., D. M. Mason, S. T. Schram and J. F. Kitchell. 1999. Ontogenetic and seasonal patterns in the energy content of piscivorous fishes in Lake Superior. J. Great Lakes Res. 25:275-281. Kitchell, J. F., D. E. Schindler, B. R. Herwig, D. M. Post, M. H. Olson and M. Oldham. 1999. Nutrient cycling at the landscape scale: The role of diel foraging migrations by geese at the Bosque del Apache Wildlife Refuge, New Mexico. Limnology and Oceanography 44:828-836. Kitchell, J. F., C. Boggs, X. He and C. J. Walters. 1999. Keystone predators in the Central Pacific. Pages 665-683 In Proc. 12th Wakefield Symposium on Ecological Considerations in Fisheries Management. Univ. of Alaska Sea Grant, Anchorage, Alaska. 756 pp. Carpenter, S. R., J. F. Kitchell, J. J. Cole and M. L. Pace. 1999. Predicting responses of chlorophyll and primary production to changes in phosphorus, grazing and dissolved organic carbon. Limnology and Oceanography 44:1179-1182. Pace, M. L., J. J. Cole, S. R. Carpenter and J. F. Kitchell. 1999. Trophic cascades revealed in diverse ecosystems. Trends in Ecology and Evolution 14:483-488. Walters, C. J., D. M. Pauly, V. Christensen and J. F. Kitchell. 2000. Representing density dependent consequences of life history strategies in aquatic ecosystems: EcoSim II. Ecosystems 3:70-83. Harvey, C.J., and J.F. Kitchell. 2000. A stable isotope evaluation of the structure and spatial heterogeneity of a Lake Superior food web. Can. J. of Fish. Aquat. Sci. 57:1395-1403. Essington, T.E., J.R. Hodgson and J.F. Kitchell. 2000. The role of satiation in the functional response of a piscivore, largemouth bass. Can. J. Fish. Aquat. Sci. 57:548-556. Kitchell, J. F., S. P. Cox, C. J. Harvey, T. B. Johnson, D. M. Mason, K. K. Schoen, K. Aydin, C. Bronte, M. Ebener, M. Hansen, M. Hoff, S. Schram, D. Schreiner and C. J. Walters. 2000. Sustainability of the Lake Superior fish community: Interactions in a food web context. Ecosystems 3:545-560. Walters, C. J., and J. F. Kitchell. 2001. Cultivation/depensation effects on juvenile survival and recruitment. Can. J. Fish. Aquat. Sci. 58:39-50. Carpenter, S.R., J.J. Cole, J.R. Hodgson, J.F. Kitchell, M.L. Pace, D. Bade, K.L. Cottingham, T.E. Essington, J.N. Houser and D.E. Schindler. 2001. Trophic cascades, nutrients and lake productivity: experimental enrichment of lakes with contrasting food webs. Ecological Monographs 71(2):163-86. Carpenter, S.R., and J.F. Kitchell. 2001. Resolution of respect: Arthur Davis Hasler, 1908-2001. Bulletin of the Ecological Society of America 82:172-3. Cole, J. J., M. L. Pace, S. R. Carpenter, and J. F. Kitchell. 2000. Persistence of net heterotrophy in lakes during nutrient addition and food web manipulations. Limnology and Oceanography 45:1718-1730. Petersen, J., and J. F. Kitchell. 2001. Climate regimes and water temperature changes in the Columbia River: Effects on predators of juvenile salmon. Can. J. Fish. Aquat. Sci.: 58:1831-1841. Harvey, C.J., P.C. Hanson, T.E. Essington, P.B. Brown, and J.F. Kitchell. 2002. Using bioenergetics models to predict stable isotope ratios in fishes. Canadian Journal of Fisheries and Aquatic Sciences 59:115-124. Essington, T. E., J. F. Kitchell and C. J. Walters. 2001. The Von Bertalanffy growth function, bioenergetics and the consumption rates of fish. Canadian Journal of Fisheries and Aquatic Sciences 58: 2129-2138. Essington, T.E., D.E. Schindler., R. J. Olson, J. F. Kitchell, C. J. Boggs and R. Hilborn 2002. Alternative fisheries and the predation rate of yellowfin tuna (Thunnus albacares) in the eastern Pacific Ocean. Ecological Applications 12: 724-734. Lathrop, R.C., B.M. Johnson, T.B. Johnson, M.T. Vogelsang, S.R. Carpenter, T.R. Hrabik, J.F. Kitchell, J.J. Magnuson, L.G. Rudstam, and R.S. Stewart. 2002. Stocking piscivores to improve fishing and water clarity: a synthesis of the Lake Mendota biomanipulation project. Freshwat. Biol. 47:2410-2424. Kitchell, J. F., T. E. Essington, C. H. Boggs, D. E. Schindler and C. J. Walters. 2002. The role of sharks and longline fisheries in a pelagic ecosystem of the central Pacific. Ecosystems 5: 202-216. Schindler, D. E., T. E. Essington, J. F. Kitchell, C. J. Boggs and R. Hilborn. 2002. Sharks and tunas: Fisheries impacts on predators with contrasting life histories. Ecological Applications 12: 735-748. Harris, G. P., S. W. Bigelow, J. J. Cole, H. Cyr, L. L. Janus, A. P. Kinzig, J. F. Kitchell, G. E. Likens, K. H. Reckhow, D. Scavia, D. Soto, L. M. Talbot, and P. H. Templer. 2002. The role of models in ecosystem management In C. D. Canham, J. J. Cole and W. K. Laurenroth (eds.) The Role of Models in Ecosystem Science. Princeton Univ. Press. Princeton NJ. Olson, R.J., G.M. Watters, K.Y. Audin, C.H. Boggs, T.E. Essington, R.C. Francis, J.F. Kitchell, J.J. Polovina, and C.J. Walters. 2002. Interactive effects of climate variability and fishing: a modelling analysis for the eastern tropical Pacific pelagic ecosystem. In Christensen, V., G. Reck, and J.L. Maclean (eds.), Proceedings of the INCO-DC Conference Placing Fisheries in their Ecosystem Context. Galápagos Islands, Ecuador, 4-8 December 2000. ACP-EU Fish. Res. Rep. 12: 31-32. Harvey, C.J., S.T. Schram, and J.F. Kitchell. 2003. Trophic relationships among lean and siscowet lake trout in Lake Superior. Trans. Amer. Fish. Soc. 132:219-228. Cox, S. P., S. J. D. Martell, C. J. Walters, T. E. Essington, J. F. Kitchell, C. H. Boggs, and I. Kaplan. 2002. Reconstructing ecosystem dynamics in the central Pacific Ocean, 1952-1998: I. Estimating population biomass and recruitment of tunas and billfishes Can. J. Fish. Aquat. Sci. 59:1724-1735. Cox, S. P., T. E. Essington, J. F. Kitchell, S. J. D. Martell, C. J. Walters, C. H. Boggs and I. Kaplan. 2002. Reconstructing ecosystem dynamics in the central Pacific Ocean, 1952-1998: II. a preliminary assessment of the trophic impacts of fishing and effects on tuna dynamics. Can. J. Fish. Aquat. Sci. 59:1736-1747. Cole, J. J., S. R. Carpenter, J. F. Kitchell and M. L. Pace. 2002. Pathways of organic C utilization in small lakes: Results from a whole-lake 13C addition and coupled model. Limnology and Oceanography 47:1664-1674. 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