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Theoretical developments in the study and prediction of food intake

Published online by Cambridge University Press:  28 February 2007

Jonathan Yearsley*
Affiliation:
Macaulay Land Use Research Institute, Craigiebuckler, Aberdeen AB15 8QH, UK
Bert J. Tolkamp
Affiliation:
Animal Nutrition and Health Department, Animal Biology Division, Scottish Agricultural College, West Mains Road, Edinburgh EH9 3JG, UK
Andrew W. Illius
Affiliation:
Institute of Cell Animal and Population Biology, University of Edinburgh, West Mains Road, Edinburgh EH9 3JT, UK
*
*Corresponding author: Dr J. Yearsley, fax +44 1224 311556, email [email protected]
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Abstract

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The purpose of the present paper is to review recent theoretical developments in food intake modelling applied to animal science and ecology. The models are divided into those that have been developed for intensive agricultural systems, and those which consider more extensive systems and natural systems. For the most part the present paper discusses models that predict the food intake of herbivores. The mechanisms of each model are discussed, along with a brief mention of the experimental support for the most popular models. We include a discussion of models that approach the study of food intake behaviour from an evolutionary perspective, and suggest that lifetime models are especially useful when food intake carries an intrinsic cost. These long timescale evolutionary models contrast with the more common food intake models, whose timescale is usually much shorter. We conclude that the ‘eating to requirements’ model highlights an important food intake mechanism that provides an accurate predictive tool for intensive agricultural systems. The mechanisms of food intake regulation in extensive systems are less certain, and closer links between the ideas of animal science and ecology will be helpful for improving our understanding of food intake regulation.

Type
Nutrition and Behaviour Group Symposium on ‘Future Perspectives in Nutrition and Behaviour Research’
Copyright
Copyright © The Nutrition Society 2001

References

Agricultural Research Council (1980). The Nutrient Requirements of Livestock. Farnham Royal, Berks: Commonwealth Agricultural Bureaux.Google Scholar
Alexander, RM (1991) Optimization of gut structure and diet for higher vertebrate herbivores. Philosophical Transactions of the Royal Society 333B, 249255.Google Scholar
Allen, MS (1996) Physical constraints on voluntary intake of forages by ruminants. Journal of Animal Science 74, 30633075.CrossRefGoogle ScholarPubMed
Amer, P & Emmans, GC (1998) Predicting changes in food energy requirements due to genetic changes in growth and body composition of growing ruminants. Animal Science 66, 143153.CrossRefGoogle Scholar
Anderson, RA & Roitberg, BD (2000) Modelling trade-offs between mortality and fitness associated with persistent blood feeding by mosquitoes. Ecology Letters 2, 98105.CrossRefGoogle Scholar
Armstrong, HM, Gordon, IJ, Hutchings, NJ, Illius, AW, Milne, JA & Sibbald, AR (1997) A model of the grazing of hill vegetation by sheep in the UK. II. The prediction of offtake by sheep. Journal of Applied Ecology 34, 186207.CrossRefGoogle Scholar
Bajzer, N (1999) Gompertzian growth as a self-similar and allometric process. Growth Development and Aging 63, 311.Google ScholarPubMed
Baldwin, RL (1995) Modelling Ruminant Digestion and Metabolism. London: Chapman and Hall.Google Scholar
Bannick, A, de Visser, H & France, J (1997) Impact of diet-specific input parameters on simulated rumen function. Journal of Theoretical Biology 184, 371384.CrossRefGoogle Scholar
Baumont, R, Prache, S, Meuret, M & Morand-Fehr, P (2000) How forage characteristics influence behaviour and intake in small ruminants: a review. Livestock Production Science 64, 1528.CrossRefGoogle Scholar
Belovsky, GE (1978) Diet optimization in a generalist herbivore: the moose. Theoretical Population Biology 14, 105134.CrossRefGoogle Scholar
Belovsky, GE (1984) Herbivore optimal foraging: a comparative test of three models. American Naturalist 124, 97115.CrossRefGoogle Scholar
Belovsky, GE, Fryxell, J & Schmidt, OJ (1999) Natural selection and herbivore nutrition: Optimal foraging theory and what it tells us about the structure of ecological communities. In Nutritional Ecology of Herbivores. Proceedings of the Vth International Symposium on the Nutrition of Herbivores, pp. 170 [Jung, HJG and Fahey, GC Jr, editors]. Savoy, IL: American Society of Animal Science.Google Scholar
Berthoud, HR (2000) An overview of neural pathways and networks involved in the control of food intake and selection. In Neural and Metabolic Control of Macronutrient Intake, pp. 361387 [Berthoud, HR and Seeley, RJ, editors]. Boca Raton, FL: CRC Press.Google Scholar
Blaxter, KL & Boyne, AW (1978) The estimation of the nutritive value of feeds as energy sources for ruminants and the derivation of feeding systems. Journal of Agricultural Science, Cambridge 90, 4768.CrossRefGoogle Scholar
Brown, JS (1999) Vigilance, patch use and habitat selection: Foraging under predation risk. Evolutionary Ecology Research 1, 4977.Google Scholar
Burrows, MT, Santini, G & Chelazzi, G (2000) A state-dependent model of activity patterns in homing limpets: balancing energy returns and mortality risks under constraints on digestion. Journal of Animal Ecology 69, 290300.CrossRefGoogle Scholar
Chadwick, L (1998) The Farm Management Handbook 1998/99. Edinburgh: Scottish Agricultural College.Google Scholar
Chan, MS & Godfray, HCJ (1993) Host-feeding strategies of parasitoid wasps. Evolutionary Ecology 7, 593604.CrossRefGoogle Scholar
Charnov, EL (1976) Optimal foraging, the marginal value theorem. Theoretical Population Biology 9, 129136.CrossRefGoogle ScholarPubMed
Danfaer, A (1990) A dynamic model of nutrient digestion and metabolism in lactating dairy cows. PhD Thesis, National Institute of Animal Science, Denmark.Google Scholar
Deag, JM (1996) Behavioural ecology and the welfare of extensively farmed animals. Applied Animal Behaviour 49, 922.CrossRefGoogle Scholar
Dijkstra, J, France, J & Tamminga, S (1998) Quantification of the recycling of microbial nitrogen in the rumen using a mechanistic model of rumen fermentation processes. Journal of Agricultural Science, Cambridge 130, 8194.CrossRefGoogle Scholar
Dijkstra, J, Neal, HDSC, Beever, DE & France, J (1992) Simulation of nutrient digestion, absorption and outflow in the rumen: Model description. Journal of Nutrition 122, 22392256.CrossRefGoogle ScholarPubMed
Downs, CT (1997) Sugar digestion efficiencies of Gurney's sugarbirds, malachite sunbirds, and black sunbirds. Physiological Zoology 70, 9399.CrossRefGoogle ScholarPubMed
Edwards, GP (1997) Predicting seasonal diet in the yellow-bellied marmot: success and failure for the linear programming model. Oecologia 112, 320330.CrossRefGoogle ScholarPubMed
Emmans, GC (1989) The growth of turkeys. In Recent Advances in Turkey Science. Poultry Science Symposium, pp. 135166 [Nixey, C and Grey, TC, editors]. London: Butterworth.Google Scholar
Emmans, GC (1997) A method to predict the food intake of domestic animals from birth to maturity as a function of time. Journal of Theoretical Biology 186, 189199.CrossRefGoogle Scholar
Emmans, GC & Kyriazakis, I (1995) The idea of optimisation in animals: uses and dangers. Livestock Production Science 44, 189197.CrossRefGoogle Scholar
Emmans, GC & Kyriazakis, I (2001) Consequences of genetic change in farm animals on food intake and feeding behaviour. Proceedings of the Nutrition Society 60, 115125.CrossRefGoogle ScholarPubMed
Engen, S & Stenseth, NC (1989) Age-specific optimal diets and optimal foraging tactics. Theoretical Population Biology 36, 281295.CrossRefGoogle ScholarPubMed
Farnsworth, KD & Illius, AW (1996) Large grazers back in the fold: generalizing the prey model to incorporate mammalian herbivores. Functional Ecology 10, 678680.Google Scholar
Farnsworth, KD & Illius, AW (1998) Optimal diet choice for large herbivores: an extended contingency model. Functional Ecology 12, 7481.CrossRefGoogle Scholar
Forbes, J & France, J (editors) (1993). Quantitative Aspects of Ruminant Digestion and Metabolism. Wallingford, Oxon: CAB International.Google Scholar
Forbes, JM (1996) Integration of regulatory signals controlling forage intake in ruminants. Journal of Animal Science 74, 30293035.CrossRefGoogle ScholarPubMed
Forbes, JM (1999) Minimal total discomfort as a concept for the control of total food intake and selection. Appetite 33, 371.CrossRefGoogle ScholarPubMed
Friggens, NC, Emmans, GC & Veerkamp, RF (1999) On the use of simple ratios between lactation curve coefficients to describe parity effects on milk production. Livestock Production Science 39, 1318.Google Scholar
Gibb, MJ & Treacher, TT (1983) The performance of lactating ewes offered diets containing different proportions of fresh perennial ryegrass and white clover. Animal Production 37, 433440.Google Scholar
Gill, M, Thornley, JHM, Black, JL, Oldham, JD & Beever, DE (1984) Simulation of the metabolism of absorbed energy yielding nutrients in young sheep. British Journal of Nutrition 52, 621649.CrossRefGoogle Scholar
Ginnett, TF & Demment, MW (1995) The functional response of herbivores: analysis and test of a simple mechanistic model. Functional Ecology 9, 376384.CrossRefGoogle Scholar
Gross, JE, Hobbs, NT & Wunder, BA (1993 a) Independent variables for predicting intake rate of mammalian herbivores: biomass density, plant density and bite size. OIKOS 68, 7581.CrossRefGoogle Scholar
Gross, JE, Shipley, LA, Hobbs, NT, Spalinger, DE & Wunder, BA (1993 b) Function-response of herbivores in food-concentrated patches – Tests of a mechanistic model. Ecology 74, 778791.CrossRefGoogle Scholar
Hilton, GM, Furness, RW & Houston, DC (2000) A comparative study of digestion in North Atlantic seabirds. Journal of Avian Biology 31, 3646.CrossRefGoogle Scholar
Hilton, GM, Houston, DC & Furness, RW (1998) Which components of diet quality affect retention time of digesta in seabirds? Functional Ecology 12, 929939.CrossRefGoogle Scholar
Hirakawa, H (1997) Digestion-constrained optimal foraging in generalist mammalian herbivores. Oikos 78, 3747.CrossRefGoogle Scholar
Hobbs, NT (1990) Diet selection by generalist herbivores: A test of the linear programming model. In Behavioural Mechanisms of Food Selection. NATO ASI Series G: Ecological S, ciences, pp. 395413 [Hughes, RN, editor]. Berlin: Springer–Verlag.CrossRefGoogle Scholar
Holling, CS (1959) Some characteristics of simple types of predation and parasitism. Canadian Entomologist 91, 385398.CrossRefGoogle Scholar
Horn, MH & Messer, KS (1992) Fish guts as chemical reactors: a model of the alimentary canals of marine herbivorous fishes. Marine Biology 113, 527535.CrossRefGoogle Scholar
Houston, AI & McNamara, JM (1999). Models of Adaptive Behaviour: An Approach Based on State. Cambridge: Cambridge University Press.Google Scholar
Houston, AI, McNamara, JM & Hutchinson, JMC (1993) General results concerning the trade-off between gaining energy and avoiding predation. Philosophical Transactions of the Royal Society 341B, 375397.Google Scholar
Huggard, DJ (1994) A linear programming model of herbivore foraging: imprecise, yet successful? Oecologia 100, 470474.CrossRefGoogle ScholarPubMed
Hughes, RN (editor) (1993). Diet Selection: An Interdisciplinary Approach to Foraging Behaviour. Oxford: Blackwell.Google Scholar
Hume, ID (1989) Optimal digestive strategies in mammalian herbivores. Physiological Zoology 62, 11451163.CrossRefGoogle Scholar
Hutchings, MR, Kyriazakis, I, Gordon, IJ & Jackson, FJ (1999) Trade-offs between nutrient and faecal avoidance in herbivore foraging decisions: the effect of animal parasitic status, level of feeding motivation and sward nitrogen content. Journal of Animal Ecology 68, 310323.CrossRefGoogle Scholar
Hutchinson, JMC & McNamara, JM (2000) Ways to test stochastic dynamic programming models empirically. Animal Behaviour 59, 665676.CrossRefGoogle ScholarPubMed
Hutchinson, JMC, McNamara, JM & Cuthill, IC (1993) Song, sexual selection, starvation and strategic handicaps. Animal Behaviour 45, 11531177.CrossRefGoogle Scholar
Hyer, JC, Oltjen, JW & Galyean, ML (1991 a) Development of a model to predict forage intake by grazing cattle. Journal of Animal Science 69, 827835.CrossRefGoogle Scholar
Hyer, JC, Oltjen, JW & Galyean, ML (1991 b) Evaluation of a feed intake model for the grazing beef steer. Journal of Animal Science 69, 836842.CrossRefGoogle ScholarPubMed
Illius, AW & Allen, MS (1994) Assessing forage quality using integrated models of intake and digestion by ruminants. In Forage Quality, Evaluation and Utilization, pp. 869890 [Fahey, GC Jr, editor]. Lincoln, NE: University of Nebraska.Google Scholar
Illius, AW, Clark, DA & Hodgson, J (1992) Discrimination and patch choice by sheep grazing grass-clover swards. Journal of Animal Ecology 61, 183194.CrossRefGoogle Scholar
Illius, AW & Gordon, IJ (1991) Prediction of intake and digestion in ruminants by a model of rumen kinetics integrating animal size and plant characteristics. Journal of Agricultural Science, Cambridge 116, 145157.CrossRefGoogle Scholar
Illius, AW & Jessop, NS (1995) Modelling metabolic costs of allelochemical ingestion by foraging herbivores. Journal of Chemical Ecology 21, 693719.CrossRefGoogle ScholarPubMed
Illius, AW, Jessop, NS & Gill, M (2000) Mathematical models of food intake and metabolism in ruminants. In Proceedings of the IXth International Symposium on Ruminant Physiology. Wallingford, Oxon: CABI–Publishing.Google Scholar
Ingvartsen, KL (1994) Models of voluntary food intake in cattle. Livestock Production Science 39, 1938.CrossRefGoogle Scholar
Jumars, PA (2000 a) Animal guts as ideal chemical reactors: Maximising absorption rates. American Naturalist 155, 527543.CrossRefGoogle Scholar
Jumars, PA (2000 b) Animal guts as nonideal chemical reactors: partial mixing and axial variation in absorption kinetics. American Naturalist 155, 544555.CrossRefGoogle ScholarPubMed
Kennedy, M & Gray, RD (1993) Can ecological theory predict the distribution of foraging animals? A critical review of experiments on the Ideal Free Distribution. OIKOS 68, 158166.CrossRefGoogle Scholar
Ketelaars, JJMH & Tolkamp, BJ (1992 a) Toward a new theory of feed-intake regulation in ruminants. 1. Causes of difference in voluntary feed-intake – critique of current views. Livestock Production Science 30, 269296.CrossRefGoogle Scholar
Ketelaars, JJMH & Tolkamp, BJ (1992 b) Toward a new theory of feed-intake regulation in ruminants. 3. Optimum feed-intake in search of a physiological background. Livestock Production Science 31, 235258.CrossRefGoogle Scholar
Ketelaars, JJMH & Tolkamp, BJ (1996) Oxygen efficiency and the control of energy flow in animals and humans. Journal of Animal Science 74, 30363051.CrossRefGoogle Scholar
Knap, PW (1999) Simulation of growth in pigs: evaluation of a model to relate thermoregulation to body protein and lipid content and deposition. Animal Science 68, 655679.CrossRefGoogle Scholar
Knap, PW & Schrama, JW (1996) Simulation of growth in pigs: approximation of protein turn-over parameters. Animal Science 63, 533547.CrossRefGoogle Scholar
Laca, EA & Demment, MW (1992) Modelling intake of a grazing ruminant in a heterogeneous environment. In Proceedings of the International Symposium on Vegetation-Herbivore Relationships, pp. 5776 [Okubo, T, Hubest, B and Arnold, G, editors]. New York: Academic Press.Google Scholar
Laca, EA, Ungar, ED & Demment, MW (1994) Mechanisms of handling time and intake rate of a large mammalian grazer. Applied Animal Behaviour Science 39, 319.CrossRefGoogle Scholar
Laca, EA, Ungar, ED, Seligman, NG, Ramey, MR & Demment, MW (1992) An integrated methodology for studying short-term grazing behaviour of cattle. Grass and Forage Science 47, 8192.CrossRefGoogle Scholar
Laredo, MA & Minson, DJ (1973) The voluntary intake, digestibility, and retention time by sheep of leaf and stem fractions of five grasses. Australian Journal of Agricultural Research 24, 875888.CrossRefGoogle Scholar
Lima, SL (1998) Stress and decision making under the risk of predation: Recent developments from behavioral, reproductive, and ecological perspectives. Stress and Behaviour 27, 215290.CrossRefGoogle Scholar
Lima, SL & Bednekoff, PA (1999) Temporal variation in danger drives antipredator behaviour: the predation risk allocation hypothesis. American Naturalist 153, 649659.CrossRefGoogle ScholarPubMed
Lima, SL & Dill, L (1990) Behavioural decisions made under the risk of predation: a review and prospectus. Canadian Journal of Zoology 68, 619640.CrossRefGoogle Scholar
López-Calleja, MV, Bozinovic, F & Martínez del Rio, C (1997) Effects of sugar concentrations on hummingbird feeding and energy use. Comparative Biochemistry and Physiology 118A, 12911299.CrossRefGoogle Scholar
McNamara, JM & Houston, AI (1996) State-dependent life histories. Nature 380, 215221.CrossRefGoogle ScholarPubMed
McNamara, JM, Mace, RH & Houston, AI (1987) Optimal routines of singing and foraging in a bird singing to attract a mate. Behavioural Ecology and Sociobiology 20, 399405.CrossRefGoogle Scholar
McNamara, JM, Webb, JN & Collins, EJ (1995) Dynamic optimisation in fluctuating environments. Proceedings of the Royal Society 258B, 2328.Google Scholar
Mangel, M & Clark, CW (1988) Dynamic Modelling in Behavioural Ecology. Princeton, NJ: Princeton University Press.Google Scholar
Mertens, DR (1994) Regulation of forage intake. In Forage Quality, Evaluation and Utilization, pp. 450493 [Fahey, GC Jr, editor]. Lincoln, NE: University of Nebraska.Google Scholar
Metz, JAJ, Nisbet, RM & Geritz, SAH (1992) How should we define 'fitness' for general ecological scenarios? Trends in Ecology and Evolution 7, 198202.CrossRefGoogle ScholarPubMed
Newman, JA, Parsons, AJ & Harvey, A (1993) Not all sheep prefer clover: diet selection revisited. Journal of Agricultural Science Cambridge 119, 275283.CrossRefGoogle Scholar
Newman, JA, Parsons, AJ & Penning, PD (1994) A note on the behavioural strategies used by grazing animals to alter their intake rates. Grass and Forage Science 49, 502505.CrossRefGoogle Scholar
Newman, JA, Parsons, AJ, Thornley, JHM, Penning, PD & Krebs, JR (1995) Optimal diet selection by a generalist grazing herbivore. Functional Ecology 9, 255268.CrossRefGoogle Scholar
Orzack, SH (1995) Tests of optimality models. Trends in Ecology and Evolution 10, 121122.Google Scholar
Orzack, SH & Sober, H (1996) How to formulate and test adaptionism. American Naturalist 148, 202210.CrossRefGoogle Scholar
Owen-Smith, N (1993) Assessing the constraints for optimal diet models. Evolutionary Ecology 7, 530531.CrossRefGoogle Scholar
Owen-Smith, N (1994) Foraging responses of kudus to seasonal changes in food resources: Elasticity in constraints. Ecology 75, 10501062.CrossRefGoogle Scholar
Owen-Smith, N (1996) Circularity in linear programming of optimal diet. Oecologia 108, 259261.CrossRefGoogle ScholarPubMed
Owen-Smith, N (1997) How successful was Edwards' linear programming model for marmots? Oecologia 112, 331332.CrossRefGoogle ScholarPubMed
Owen-Smith, N & Novellie, P (1982) What should a clever ungulate eat? American Naturalist 119, 151178.CrossRefGoogle Scholar
Pandian, TJ & Marian, MP (1985) Estimation of absorption efficiency in polychaetes using nitrogen content of food. Journal of Experimental Marine Biology 90, 289295.CrossRefGoogle Scholar
Parsons, AJ, Thornley, JHM, Newman, J & Penning, PD (1994) A mechanistic model of some physical determinants of intake rate and diet selection in a two-species temperate grassland sward. Functional Ecology 8, 187204.CrossRefGoogle Scholar
Pastor, J, Standke, K, Farnsworth, K, Moen, R & Cohen, Y (1999) Further development of the Spalinger-Hobbs mechanistic foraging model for free-ranging moose. Canadian Journal of Zoology 77, 15051512.CrossRefGoogle Scholar
Penning, PD, Parsons, AJ, Ord, RJ & Treacher, TT (1991) Intake and behaviour responses by sheep to changes in sward characteristics under continuous stocking. Grass and Forage Science 46, 1528.CrossRefGoogle Scholar
Penry, DL (1993) Constraints on diet selection. In Diet Selection: An Interdisciplinary Approach to Foraging Behaviour, pp. 3255 [Hughes, RN, editor]. Oxford: Blackwell.Google Scholar
Penry, DL & Jumars, PA (1986) Chemical reactor analysis and optimal digestion theory. Bioscience 36, 310315.CrossRefGoogle Scholar
Penry, DL & Jumars, PA (1987) Modeling animal guts as chemical reactors. American Naturalist 129, 6996.CrossRefGoogle Scholar
Perry, G & Pianka, ER (1997) Animal foraging: Past, present and future. Trends in Ecology and Evolution 12, 360364.CrossRefGoogle ScholarPubMed
Pierotti, R & Annett, CA (1991) Diet choice in the herring gull: Constraints imposed by reproductive and ecological factors. Ecology 72, 319328.CrossRefGoogle Scholar
Pitroff, W & Kothmann, MM (1999) Regulation of intake and diet selection by herbivores. In Nutritional Ecology of Herbivores. Proceedings of the Vth International Symposium on the Nutrition of Herbivores, pp. 366422 [Jung, HJG and Fahey, GC Jr, editors]. Savoy, IL: American Society of Animal Science.Google Scholar
Poppi, DP (1996) Predictions of food intake in ruminants from analyses of food composition. Australian Journal of Agricultural Research 47, 489504.CrossRefGoogle Scholar
Poppi, DP, Gill, M & France, J (1994) Integration of theories of intake regulation in growing ruminants. Journal of Theoretical Biology 167, 129145.CrossRefGoogle Scholar
Pulliam, HR (1975) Diet optimization with nutrient constraints. American Naturalist 109, 765768.CrossRefGoogle Scholar
Rapport, DJ (1980) Optimal foraging for complimentary resources. American Naturalist 116, 324346.CrossRefGoogle Scholar
Sæther, B-E & Gordon, IJ (1994) The adaptive significance of reproductive strategies in ungulates. Proceedings of the Royal Society 256B, 263268.Google Scholar
Sauvant, D, Baumont, R & Faverdin, P (1996) Development of a mechanistic model of intake and chewing activities of sheep. Journal of Animal Science 74, 27852802.CrossRefGoogle ScholarPubMed
Shipley, LA, Gross, JE, Spalinger, DE, Hobbs, NT & Wunder, BA (1994) The scaling of intake rate in mammalian herbivores. American Naturalist 143, 10551082.CrossRefGoogle Scholar
Shipley, LA & Spalinger, DE (1992) Mechanics of browsing in dense patches – effects of plant and animal morphology on intake rate. Canadian Journal of Zoology 70, 17431752.CrossRefGoogle Scholar
Spalinger, DE & Hobbs, NT (1992) Mechanisms of foraging in mammalian herbivores: New models of functional response. American Naturalist 140, 325348.CrossRefGoogle ScholarPubMed
Speakman, JR (2000) The cost of living: Field metabolic rates of small mammals. Advances in Ecological Research 30, 177297.CrossRefGoogle Scholar
Stephens, DW & Krebs, JR (1986) Foraging Theory. Princeton, NJ: Princeton University Press.Google Scholar
Taghon, GL & Greene, RR (1990) Effects of sediment-protein concentration on feeding and growth rates of Abarenicola pacifica Healy et Wells (Polchaeta: Arenicolidae). Journal of Experimental Marine Biology and Ecology 136, 197216.CrossRefGoogle Scholar
Thomas, RJ (1999 a) The effect of variability in the food supply on the daily singing routines of European robins: a test of a stochastic dynamic programming model. Animal Behaviour 57, 365369.CrossRefGoogle Scholar
Thomas, RJ (1999 b) Two tests of a stochastic dynamic programming model of daily singing routines in birds. Animal Behaviour 57, 277284.CrossRefGoogle ScholarPubMed
Thomas, RJ (2000) Strategic diel regulation of body mass in European robins. Animal Behaviour 59, 787791.CrossRefGoogle ScholarPubMed
Thornley, JHM, Parsons, AJ, Newman, J & Penning, PD (1994) A cost-benefit model of grazing intake and diet selection in a two-species temperate grassland sward. Ecology 8, 516.Google Scholar
Tolkamp, BJ & Ketelaars, JJMH (1992) Toward a new theory of feed-intake in ruminants. 2. Costs and benefits of feed consumption – an optimization. Livestock Production Science 30, 297317.CrossRefGoogle Scholar
Van de Meer, J & Ens, BJ (1997) Model of interference and their consequences for the spatial distribution of ideal free predators. Journal of Animal Ecology 66, 846858.CrossRefGoogle Scholar
Ward, D (1992) The role of satisfying in foraging theory. OIKOS 63, 312317.CrossRefGoogle Scholar
Ward, D (1993) Foraging theory, like all other fields of science, needs multiple hypotheses. OIKOS 67, 376378.CrossRefGoogle Scholar
Ward, JF, Austin, RM & MacDonald, DW (2000) A simulation model of foraging behaviour and the effect of predation risk. Journal of Animal Ecology 69, 1630.CrossRefGoogle Scholar
Webster, AJF (1993) Energy partitioning, tissue growth and appetite control. Proceedings of the Nutrition Society 52, 6976.CrossRefGoogle ScholarPubMed
Woodward, SJR (1997) Formulae for predicting animals' daily intake of pasture and grazing time from bite weight and composition. Livestock Production Science 52, 110.CrossRefGoogle Scholar
Xue, RD, Edman, JD & Scott, TW (1995) Age and body size effects on blood meal size and multiple blood feeding by Aedes aegypti (Diptera: Culicidae). Journal of Medical Entomology 32, 471474.CrossRefGoogle ScholarPubMed
Yang, Y & Joern, A (1994) Influence of diet quality, developmental stage, and on food residence time in the grasshopper Melanoplus differentialis . Physiological Zoology 67, 598616.CrossRefGoogle Scholar