Bioenergetics Model

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What is Bioenergetics?
..the study of the flow and transformation of
energy in and between living organisms and
between living organisms and their environment
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MUST!
What Goes In What Comes Out
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• 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 ..

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Typical Energy Budgets Differ for Carnivores
Herbivores
Green Sunfish
Muskellunge
Largescale Stoneroller
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Bioenergetics Economics
Consumption Metabolism Waste Growth
Consumption Income
Metabolism Rent
Wastes Losses Taxes
Growth Savings and Investments
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Lake Cheese
Lake Salad
Why would growth be different?
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Why would growth be different?
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Model Components
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(?B G)
(R A S)
(F U)
C
Independent C consumption
Losses R respiration A active
metabolism S specific dynamic action
Gains ?B Change in biomass G gonads
/reproductionn
Losses F egestion U excretion
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All processes are temp. and size dependent
0.08
0.06
Specific Rate (g/g/d)
0.04
0.02
Grams body mass stays the same, so how much you
eat is what is changing
0.00
15
25
30
20
10
5
Temperature (C)
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All processes are temp. and size dependent
0.08
0.06
Specific Rate (g/g/d)
0.04
0.02
0.00
15
25
30
20
10
5
Temperature (C)
13
Maximum consumption isn't realisticp-value
proportion of maximum consumption
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Maximum consumption isn't realisticp-value
proportion of maximum consumption
starvation
Upper lethal
loss of growth
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How do we estimate these basic relationships?
Like the relationship between temperature and
maximum consumption..lab experiments and or
field data
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Consumption and Respiration ? Temperature
Dependent
0.08
0.06
Specific Rate (g/g/d)
starvation
Upper lethal
0.04
0.02
0.00
loss of growth
30
15
20
25
10
5
Temperature (C)
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Consumption and Respiration ? Size Dependent
FOR A GIVEN TEMPERATURE
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Main equation can be organized in 2 ways
(?B G)
(R A S)
(F U)
C
Can solve for any of the different parameters of
the equations.hence the power of bioenergetics
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What do we need to run the model?
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Growth
Mark and recap
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Measure growth Calcified structures fish
parts scales, otoliths, vertebrae, rays, opercula
15 mm Age0
76 mm Age1
123 mm Age2
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What else do we need to run the model?
Temperatures where fish live

• 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

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What do we need to run the model?
What a fish eats
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What do we need to run the model?
Prey and Predator Energy Densities
Snails 18000 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 24000 j/g dry mass
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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
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How have bioenergetic models been used in
fisheries?
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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)
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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 result in algal blooms
(Kraft 1993)
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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)
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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.
412-6. 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. 45178-188. 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. 6257-94. 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.
103255-260. 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. 103786-798. 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.
1975391-399. Kitchell, J.F., J.F. Koonce, and
P.S. Tennis. 1975. Phosphorus flux through
fishes. Verh. Internat. Verein. Limnol.
192478-2484. 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. 105712-717. 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. 341936-1940. 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. 341922-1935. Kitchell, J.F., J.J.
Magnuson, and W.H. Neill. 1977. Estimation of
caloric equivalents for fish biomass. Environ.
Biol. Fish. 2185-189. Bartell, S.M., and J.F.
Kitchell. 1978. Seasonal impact of planktivory on
phosphorus release by Lake Wingra zooplankton.
Verh. Internat. Verein. Limnol.
20466-474. Gallepp, G.W., J.F. Kitchell, and
S.M. Bartell. 1978. Phosphorus released from lake
muds affected by chironomids. Verh. Internat.
Verein. Limnol. 20458-474. 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 2001289-1291. Kitc
hell, J.F., R.A. Stein, and B. Knezevic. 1978.
Utilization of filamentous algae by fishes in
Skadar Lake, Yugoslavia. Verh. Internat. Verein.
Limnol. 202159-2165. 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 4171-180. 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 2928-34. 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. 25389-402. 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. 372159-2168. 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. 372202-2208. 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. 110751-763. 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. 9263-275. 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 6213-17. Carpenter, S.R., and J.F.
Kitchell. 1984. Plankton community structure and
limnetic primary production. American Naturalist
124159-172. 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 35634-639. Kitchel
l, 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
32
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. 122756-772.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.Schindle
r, D. E., B. M. Johnson, N. A. MacKay, N. Bouwes
and J. F. Kitchell. 1994. Crabsnail
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.
45209-226. 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.Carpen
ter, 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 38764-769.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. 23351-358.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. 14109-114.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. 45165-177.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.
34700-717.Cochran, P.A., and J.F. Kitchell.
l989. A model of feeding by parasitic lampreys.
Can. J. Fish. Aquat. Sci. 461845-52.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
200/201329-336.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. 119642-648.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 344333-335.66. 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.
119825-835.67. 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.
481574-1580.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. 64502-524.Carpenter,
S.R., S.G. Fisher, N.B. Grimm, and J.F.
Kitchell. l992. Global change and freshwater
ecosystems. Ann. Rev. Ecol. Syst.
23119-139.Carpenter, S.R., and J.F. Kitchell.
l992. Trophic cascade and biomanipulation
interface of research and management. Limnol.
Oceanogr.37208-213.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. 12797-807.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
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Rehabilitation of Lake Ontario the role of
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and A.P. Zimmerman. l991. Patterns of primary
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Stewart, C. Luecke, M.D. Staggs, S.J. Gilbert and
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398-400. Hansen, M. J., D. Boisclair, S. B.
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Fratt, R.E.Bruesewitz, D.W.Coble, F.A. Copes and
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52.1564-1571. Post, D. M., T. M. Frost and J. F.
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whole-lake acidification experiment. J. Plankton
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Fratt, R.E.Bruesewitz, D.W.Coble, F.A. Copes and
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G. Rudstam and J. F. Kitchell. 1995. Predator
responses to prey population dynamics an
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52.1564-1571. Post, D. M., T. M. Frost and J. F.
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changes in copepod predator populations during a
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E. Peppard. 1996. Predation rates by North Sea
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34
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Long-term changes in zooplantivorous fish
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532792-2803. Kitchell, J. F., D. E. Schindler,
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Nile perch in Lake Victoria interactions between
predation and fisheries. Ecol. Appl. 7
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Trophic ontogeny and life history effects on
interactions between age-0 fish and zooplankton.
Arch. Hydrobiol. Spec. Issues Advanc. Limnol.
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Kitchell. 1997. Density-dependent changes in
individual foraging specialization of largemouth
bass. Oecologia 110592-600. 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.
42722-729. 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
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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. 7981-990.
Hanson, P.C., T. B. Johnson, D. E. Schindler,
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Sea Grant Technical Report, University of
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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
277248-251. Hodgson, J. R., X. He, D. E.
Schindler, and J. F. Kitchell. 1997. Diet overlap
in a piscivore community. Ecol of Freshwater Fish
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Understanding human modifications of the ocean
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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.
4373-80. 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 1256-64.
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 12910-920. Kitchell, J. F. 1998.
Physiological ecology. Pages 164-198 In S.I.
Dodson (ed.), Ecology. Oxford Univ. Press. 434
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community dynamics on lake trout (Salvelinus
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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.
127496-506. 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 1335-344. 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.
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of fish distributions A case study on the
spatial distribution of largemouth bass. Can. J.
Fish. Aquat. Sci. 5652-60. 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. 25275-281. 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 44828-836. Kitchell, J. F., C.
Boggs, X. He and C. J. Walters. 1999. Keystone
predators in the Central Pacific. Pages 665-683
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Considerations in Fisheries Management. Univ. of
Alaska Sea Grant, Anchorage, Alaska. 756
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and M. L. Pace. 1999. Predicting responses of
chlorophyll and primary production to changes in
phosphorus, grazing and dissolved organic carbon.
Limnology and Oceanography 441179-1182. 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 14483-488. 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 370-83. 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. 571395-1403. 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. 57548-556. 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 3545-560. Walters, C.
J., and J. F. Kitchell. 2001. Cultivation/depensat
ion effects on juvenile survival and recruitment.
Can. J. Fish. Aquat. Sci. 5839-50. 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 82172-3. 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
451718-1730. 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. 581831-1841. 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
59115-124. 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. 472410-2424. 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
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Bigelow, J. J. Cole, H. Cyr, L. L. Janus, A. P.
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assessment of the trophic impacts of fishing and
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Sci. 591736-1747. Cole, J. J., S. R. Carpenter,
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pelagic ecosystem in the eastern tropical
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Sci. In press. Johnson, T. B., M. H. Hoff3, A.
S. Trebitz, C. R. Bronte, W. P. Brown, T. D.
Corry , J. F. Kitchell, S. T. Lozano, D. M.
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structures of pelagic forage fish and zooplankton
in western Lake Superior. J. Great Lakes Res. In
press Pace, M. L. J. J. Cole, S. R. Carpenter,
J. F. Kitchell, J. R. Hodgson, M. Van de Bogert,
D. L. Bade, E. S. Kritzberg and D. Bastviken.
2004. Whole lake carbon-13 additions reveal
terrestrial support of aquatic food webs. Nature
427240-243. Kitchell, J. F., I. Kaplan, S. P.
Cox, S. J. D. Martell, T. E. Essington, C. H.
Boggs and C. J. Walters. 2004. Ecological and
economic components of alternative fishing
methods to reduce by-catch of marlin in a
tropical pelagic ecosystem. Bull. Mar. Sci. In
press Hinke, J. T., I. C. Kaplan, K. Aydin, G.
M. Watters, R. J. Olson, and J. F. Kitchell.
2004. Visualizing the food-web effects of fishing
for tunas in the Pacific Ocean. Ecology and
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online http//www.ecologyandsociety.org/vol9/iss
1/art10 Sass, G. G., S. W. Hewett, T. D. Beard,
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of density-dependence in growth patterns of ceded
territory walleye populations of Northern
Wisconsin effects of changing management
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