Skip to main content Accessibility help

Links between ruminants’ food preference and their welfare

  • J. J. Villalba (a1), F. D. Provenza (a1) and X. Manteca (a2)


Nutrition is an important aspect of welfare, and in most recommendations for the welfare of animals adequate nutrition is a primary requirement. However, in intensive livestock production systems the decision for adequate nutrition is made based on traditional paradigms of feeding monotonous rations or plant monocultures, frequently with excesses or imbalances of nutrients relative to the individuals’ physiology, which can compromise welfare. Individual ruminants can better meet their needs for nutrients and regulate their intake of secondary compounds when offered a variety of foods than when constrained to a single food, even if the food is nutritionally balanced. The concept of food variety is central because monotonous flavors and feeds and excess nutrients all cause animals to satiate, which in turn causes animals to eat a variety of foods. When offered a variety of foods, satiety for single foods stimulates the selection of a diverse diet and thus food intake, but when restricted to a monotonous diet satiety is aversive and limits food intake. Moreover, if a monotonous diet is aversive to animals then this could be stressful, even if monotony implies consuming a balanced diet. A diverse diet may also increase resistance to disease in ruminants, by allowing consumption of small amounts of compounds with antimicrobial/antiparasitic effects and immunity-enhancing properties. Herbivores also experience the benefits of ingesting compounds with medicinal (i.e. antiparasitic) benefits and they learn to prefer foods containing such compounds as their preferences are associatively conditioned by the food’s homeostatic utility to the body. Such learned patterns of behavior begin in utero and feeding experiences early in life cause changes – neurological, morphological and physiological – in animals, which influence on their subsequent behavior and welfare. Such experiences with the environment enable animals to adapt to local diets and stressors and reduce the levels of fear. Finally, feeding behavior in farm animals could be an aid in the early detection and mitigation of pain or sickness, and as such become an important tool in the identification of welfare and health of animals before the appearance of clinical signs. Management strategies in ruminant production systems could benefit by allowing animals to manifest their feeding preferences, thereby acknowledging the animals’ role as active players in feeding systems, instead of regarding them as passive entities that just respond to prescriptions and formulations.


Corresponding author


Hide All
Atwood, SB, Provenza, FD, Wiedmeier, RD, Banner, RE 2001a. Changes in preferences of gestating heifers fed untreated or ammoniated straw in different flavors. Journal of Animal Science 79, 30273033.
Atwood, SB, Provenza, FD, Wiedmeier, RD, Banner, RE 2001b. Influence of free-choice versus mixed-ration diets on food intake and performance of fattening calves. Journal of Animal Science 79, 30343040.
Balleine, BW, Dickinson, A 1998. The role of incentive learning in instrumental outcome revaluation by sensory-specific satiety. Learning & Behavior 26, 4659.
Barry, TN, McNeill, DM, McNabb, WC 2001. Plant secondary compounds; their impact on nutritive value and upon animal production. Proc. XIX Int. Grass. Conf., SaoPaulo, Brazil. 19, 445–452.
Bechara, A, Tranel, D, Damasio, H, Adolph, R, Rockland, C, Damasio, AR 1995. Double dissociation of conditioning and declarative knowledge relative to the amygdale and hippocampus in humans. Science 269, 11151118.
Berman, DG, Johnson, DE, Phillips, RW, Barry, BP 1980. Physiological and urinary metabolite responses to cold shock and confinement of sheep. Journal of Animal Science 50, 713722.
Berridge, KC, Robinson, TE 1998. What is the role of dopamine in reward: hedonic impact, reward learning, or incentive salience? Brain Research Reviews 28, 309369.
Bevans, DW, Beauchemin, KA, Schwartzkopf-Genswein, KS, McKinnon, JJ, McAllister, TA 2005. Effect of rapid or gradual grain adaptation on subacute acidosis and feed intake by feedlot cattle. Journal of Animal Science 83, 11161132.
Boissy, A 1998. Fear and fearfulness in determining behavior. In Genetics and the behavior of domestic animals (ed. T Grandin), pp. 67111. Academic Press, San Diego, USA.
Boissy, A, Le Neindre, P 1997. Behavioral, cardiac and cortisol responses to brief peer separation and reunion in cattle. Physiology & Behaviour 61, 693699.
Broom, DM 1986. Indicators of poor welfare. The British Veterinary Journal 142, 524526.
Burritt, EA, Provenza, FD 1997. Effect of an unfamiliar location on the consumption of novel and familiar foods by sheep. Applied Animal Behaviour Science 54, 317325.
Carlstead, K, Shepherdson, D 2000. Alleviating stress in zoo animals with environmental enrichment. In The biology of animal stress. Basic principles and implications for animal welfare (ed. GP Moberg and JA Mench), pp. 337354. CAB International, Wallingford.
Chapple, RS, Lynch, JJ 1986. Behavioural factors modifying acceptance of supplementary foods by sheep. Research and Development in Agriculture 3, 113123.
Coetzee, JF, Gehring, R, Bettenhausen, AC, Lubbers, BV, Toerber, SE, Thomson, DU, Kukanich, B, Apley, MD 2007. Attenuation of acute plasma cortisol response in calves following intravenous sodium salicylate administration prior to castration. Journal of Veterinary Pharmacology and Therapeutics 30, 305313.
Colpaert, FC, De Witte, P, Mondi, AN, Awouters, F, Niemegeers, CJE, Jansen, PAJ 1980. Self-administration of the analgesic suprofen in arthritic rats: evidence of mycobacterium butyricum-induced arthritis as an experimental model of chronic pain. Life Sciences 27, 921928.
Cooper, SD, Kyriazakis, I, Oldham, JD 1996. The effects of physical form of feed, carbohydrate source, and inclusion of sodium bicarbonate on the diet selections of sheep. Journal of Animal Science 74, 12401251.
Cozier, A, Clifford, MN, Ashihara, H 2006. Plant secondary metabolites: occurrence, structure and role in the human diet. Blackwell Publishing, Oxford, UK.
Critchley, HD, Rolls, ET 1996. Hunger and satiety modify the responses of olfactory and visual neurons in the primate orbitofrontal cortex. Journal of Neurophysiology 75, 16731686.
Danbury, TC, Weeks, CA, Chambers, JP, Waterman-Pearson, AE, Kestin, SC 2000. Self-selection of the analgesic drug carprofen by lame broiler chickens. The Veterinary Record 146, 307311.
Dantzer, R, Arnone, M, Mormède, P 1980. Effects of frustration on behaviour and plasma corticosteroid levels in pigs. Physiology & Behavior 24, 14.
Distel, RA, Villalba, JJ, Laborde, HE 1994. Effects of early experience on voluntary intake of low-quality roughage by sheep. Journal of Animal Science 72, 11911195.
Distel, RA, Villalba, JJ, Laborde, HE, Burgos, MA 1996. Persistence of the effects of early experience on consumption of low-quality roughage by sheep. Journal of Animal Science 74, 965968.
Dumont, B, D’hour, P, Petit, M 1995. The usefulness of grazing tests for studying the ability of sheep and cattle to exploit reproductive patches of pastures. Applied Animal Behaviour Science 45, 7988.
Dumont, B, Dutronc, A, Petit, M 1998. How readily will sheep walk for a preferred forage? Journal of Animal Science 76, 965971.
Duncan, AJ, Elwert, C, Villalba, JJ, Yearsley, J, Pouloupoulou, I, Gordon, IJ 2007. How does pattern of feeding and rate of nutrient delivery influence conditioned food preferences? Oecologia 153, 617624.
Early, D, Provenza, FD 1998. Food flavor and nutritional characteristics alter dynamics of food preference in lambs. Journal of Animal Science 76, 728734.
Engel, C 2002. Wild Health. Houghton Mifflin Co, Boston.
Ewing, SA, Lay, DC Jr, von Borell, E 1999. Farm animal well-being. Stress physiology, animal behavior and environmental design. Prentice Hall, New Jersey, NY, USA.
FAWC 1993. Second report on priorities for research and development in farm animal welfare. Farm animal welfare council. MAFF Tolworth, UK.
Fell, LR, Shutt, DA 1986. Use of salivary cortisol as an indicator of stress due to management practices in sheep and calves. Proceedings of the Australian Society of Animal Production 16, 203206.
Fish, EW, Shahrokh, D, Bagot, R, Caldji, C, Bredy, T, Szyf, M, Meaney, MJ 2004. Epigenetic programming of stress responses through variations in maternal care. Annals of the NewYork Academy of Sciences 1036, 167180.
Forbes, JM 2003. The multifactorial nature of food intake control. Journal of Animal Science 81, E139E144.
Fraser, AF, Broom, DM 1997. Farm animal behaviour and welfare. CAB International, New York, NY, USA.
Freeland, W, Janzen, DH 1974. Strategies in herbivory by mammals: the role of plant secondary compounds. American Naturalist 108, 269286.
Gluckman, PD, Hanson, MA, Spencer, HG 2005. Predictive adaptive responses and human evolution. Trends in Ecology & Evolution 20, 527533.
Gordon, IJ, Illius, AW, Milne, JD 1996. Sources of variation in the foraging efficiency of grazing ruminants. Functional Ecology 10, 219226.
Grade, JT, Tabuti, JRS, Van Damme, P, Arble, BL 2007. Deworming efficacy of Albizia anthelmintica in uganda: preliminary findings. African Journal of Ecology 45, 1820.
Grade, JT, Tabuti, JRS, Van Damme, P 2009. Four footed pharmacists: indications of self-medicating livestock in karamoja, uganda. Economic Botany 63, 2942.
Gregory, NG 2004. Physiology and behaviour of animal suffering. UFAW Animal Welfare Series. Blackwell Publishing, Oxford, UK.
Hill, J, Chapman, DF, Cosgrove, GP, Parsons, AJ 2009. Do ruminants alter 1 their preference for pasture species in response to the synchronisation of delivery and release of nutrients? Rangeland Ecology & Management In Press.
Hocquemiller, R, Cortes, D, Arango, GJ, Myint, SH, Cave, A, Angelo, A, Munoz, VFournet, A 1991. Isolation and synthesis of espintanol, a new antiparasitic monoterpenes. Journal of Natural Products 54, 445452.
Huber, TL 1976. Physiological effects of acidosis on feedlot cattle. Journal of Animal Science 43, 902909.
Huffman, MA 2003. Animal self-medication and ethno-medicine: exploration and exploitation of the medicinal properties of plants. Proceedings of the Nutrition Society 62, 371381.
Huffman, MA, Seifu, M 1989. Observations on the illness and consumption of a possibly medicinal plant Vernonia amygdalina by a wild chimpanzee in the Mahale Mountains National Park, Tanzania. Primates 30, 51–63.
Huffman, MA, Ohigashi, H, Kawanaka, M, Page, JE, Kirby, GC, Gasquet, M, Murakami, A, Koshimizu, K 1998. African great ape self-medication: a new paradigm for treating parasite disease with natural medicines? In Towards natural medicine research in the 21st century (ed. H Ageta, N Aimi, Y Ebizuka, T Fujita and G Honda), pp. 113123. Elsevier Science, Amsterdam, The Netherlands.
Jones, RB 1997. Fear and distress. In Animal welfare (ed. MC Appleby, BO Hughes), pp. 7587. CAB International, Wallingford, UK.
Kayser, O, Kiderlen, AF, Croft, SL 2003. Natural products as antiparasitic drugs. Parasitology Research 90, S55S62.
Kilgour, RJ, Szantar-Coddington, MR 1997. The arena test and cortisol response of sheep as indirect selection criteria for the improvement of lamb survival. Animal Reproduction Science 46, 97108.
Konarzewski, M, Diamond, J 1994. Peak sustained metabolic rate and its individual variation in cold-stressed mice. Physiological Zoology 67, 11861212.
Krachun, C, Rushen, J, de Passille, AM 2010. Play behaviour in dairy calves is reduced by weaning and by a low energy intake. Applied Animal Behaviour Science 122, 7176.
Kyriazakis, I, Oldham, JD 1993. Diet selection in sheep: the ability of growing lambs to select a diet that meets their crude protein (nitrogen × 6.25) requirements. The British Journal of Nutrition 69, 617629.
Kyriazakis, I, Savory, J 1997. Hunger and thirst. In Animal welfare (ed. MC Appleby and BO Hughes), pp. 4962. CAB International, Wallingford, UK.
Landgraf, R, Wigger, A, Holsboer, F, Neumann, D 1999. Hyper-reactive hypothalamo-pituitary-adrenocortical axis in rats bred for high-anxiety-related behaviour. Journal of Neuroendocrinology 11, 405407.
Launchbaugh, KL, Provenza, FD, Werkmeister, MJ 1997. Overcoming food neophobia in domestic ruminants through addition of a familiar flavor and repeated exposure to novel foods. Applied Animal Behaviour Science 54, 327334.
LeDoux, J 2002. Synaptic self: how our brains become who we are. Viking Penguin, New York, NY, USA.
Lim, SS, Norman, RJ, Clifton, PM, Noakes, M 2009. Hyperandrogenemia, psychological distress, and food cravings in young women. Physiology & Behavior 98, 276280.
Lisonbee, LD, Villalba, JJ, Provenza, FD, Hall, JO 2009. Tannins and self-medication: implications for sustainable parasite control in herbivores. Behavioural Processes In Press.
Lobley, GE, Milano, GD 1997. Regulation of hepatic nitrogen metabolism in ruminants. Proceedings of the Nutrition Society 57, 547563.
Lozano, GA 1998. Parasitic stress and self-medication in wild animals. In Advances in the study of behavior (ed. AP Moler, M Milinski and PJB Slater), pp. 291317. Elsevier Science, London, UK.
Manteca, X, Villalba, JJ, Atwood, SB, Dziba, L, Provenza, FD 2008. Is dietary choice important to animal welfare? Journal of Veterinary Behavior: Clinical Applications and Research 3, 229239.
Matteri, RL, Carroll, JA, Dyer, CJ 2000. Neuroendocrine responses to stress. In The biology of animal stress: basic principles and implications for animal welfare (ed. GP Moberg and JA Mench), pp. 4376. CAB International, Wallingford, CT, USA.
Meaney, M 2009. Epigenetic programming of behavior and physiology. Journal of Animal Science 87 (E-suppl. 2/J), 365.
Mellor, DJ, Stafford, KJ 2004. Animal welfare implications of neonatal mortality and morbidity in farm animals. Veterinary Journal 168, 118133.
Middleton, E Jr, Kandaswami, C, Theoharides, T 2000. The effects of plant flavonoids on mammalian cells: implications for inflammation, heart disease, and cancer. Pharmacological Reviews 52, 673751.
Min, BR, Hart, SP 2003. Tannins for suppression of internal parasites. Journal of Animal Science 81 (E-suppl. 2), E102E109.
Nolte, DL, Provenza, FD 1992a. Food preferences in lambs after exposure to flavors in milk. Applied Animal Behaviour Science 32, 381389.
Nolte, DL, Provenza, FD 1992b. Food preferences in lambs after exposure to flavors in solid foods. Applied Animal Behaviour Science 32, 337347.
Oliver, G, Wardle, J 1999. Perceived effects of stress on food choice. Physiology & Behavior 66, 511515.
Palme, R, Robia, C, Messmann, S, Hofer, J, Mostl, E 1999. Measurement of faecal cortisol metabolites in ruminants: a non-invasive parameter of adrenocortical function. Wiener Tierarztliche Monatsschrift 86, 237241.
Picman, AK 1986. Biological activities of sesquiterpene lactones. Biochemical Systematics and Ecology 14, 255281.
Polivy, JC, Herman, P, Coelho, JS 2008. Caloric restriction in the presence of attractive food cues: external cues, eating, and weight. Physiology & Behavior 94, 729733.
Provenza, FD 1991. Behavior and nutrition are complementary endeavors. Proceedings 2nd Grazing Livestock Nutrition Conference. Agricultural Experiment Station MP-133. Oklahoma State University, Stillwater, OK, USA, pp. 157–169.
Provenza, FD 1996. Acquired aversions as the basis for varied diets of ruminants foraging on rangelands. Journal of Animal Science 74, 20102020.
Provenza, FD, Balph, DF 1990. Applicability of five diet-selection models to various foraging challenges ruminants encounters. In Behavioural mechanisms of food selection. NATO ASI Series G: ecological sciences (ed. Hughes RN). vol. 20, pp. 423459. Springer-Verlag, Berlin, Heildelberg, Germany.
Provenza, FD, Villalba, JJ 2006. Foraging in domestic vertebrates: linking the internal and external milieu. In Feeding in domestic vertebrates: from structure to function (ed. VL Bels), pp. 210240. CABI Publisher, Oxfordshire, UK.
Provenza, FD, Burritt, EA, Clausen, TP, Bryant, JP, Reichardt, PB, Distel, RA 1990. Conditioned flavor aversion: a mechanism for goats to avoid condensed tannins in blackbrush. American Naturalist 136, 810828.
Provenza, FD, Scott, CB, Phy, TS, Lynch, JJ 1996. Preferences of sheep for foods varying in flavors and nutrients. Journal of Animal Science 74, 23552361.
Provenza, FD, Villalba, JJ, Dziba, LE, Atwood, SB, Banner, RE 2003. Linking herbivore experience, varied diets, and plant biochemical diversity. Small Ruminant Research 49, 257274.
Redbo, I, Nordblad, A 1997. Stereotypies in heifers are affected by feeding regime. Applied Animal Behaviour Science 53, 193202.
Robles, M, Arguellin, M, West, J, Rodriguez, E 1995. Recent studies on zoopharmacognosy, pharmacology and neurotoxicology of sesquiterpene lactones. Planta Medica 61, 199203.
Rolls, BJ 1986. Sensory-specific satiety. Nutrition Reviews 44, 93101.
Rolls, BJ, Rolls, ET, Rowe, EA, Sweeney, K 1981. Sensory specific satiety in man. Physiology & Behavior 27, 137142.
Rolls, BJ, van Duijvenvoorde, PM, Rowe, EA 1983. Variety in the diet contributes to the development of obesity in the rat. Physiology & Behavior 30, 2127.
Rosenthal, GA, Janzen, DH 1979. Herbivores: their interaction with secondary plant metabolites. Academic Press, New York, NY, USA.
Rutter, SM 2006. Diet preference for grass and legumes in free-ranging domestic sheep and cattle: current theory and future application. Applied Animal Behaviour Science 97, 1735.
Schepetkin, IA, Quinn, MT 2006. Botanical polysaccharides: macrophage immunomodulation and therapeutic potential. International Immunopharmacology 6, 317333.
Schwartzkopf-Genswein, K, González, L, Gibb, D, McAllister, T 2009. Application of feeding behavior as an indicator of pain and morbidity in feedlot cattle. Journal of Animal Science 87 (E-suppl. 2/J), 319.
Scott, LL, Provenza, FD 1998. Variety of foods and flavors affects selection of foraging locations by sheep. Applied Animal Behaviour Science 61, 113122.
Scott, LL, Provenza, FD 1999. Variation in food selection among lambs: effects of basal diet and foods offered in a meal. Journal of Animal Science 77, 23912397.
Smith, J, Boyd, K 1991. Lives in the Balance. Oxford University Press, Oxford, UK.
Sowell, BF, Bowman, JGP, Branine, ME, Hubbert, ME 1998. Radio frequency technology to measure feeding behavior and health of feedlot steers. Applied Animal Behaviour Science 59, 277284.
Swithers, SE, Hall, WG 1994. Does oral experience terminate ingestion ? Appetite 23, 113138.
Tizard, IR, Carpenter, RH, McAnalley, BH, Kemp, MC 1989. The biological activities of mannans and related complex carbohydrates. Molecular Biotherapy 1, 290296.
Van Roijen, J 1991. Predictability and boredom. Applied Animal Behaviour Science 31, 283287.
Van Vuuren, AM, Meijas, JAC 1987. Effects of herbage composition and supplement feeding on the excretion on nitrogen dung and urine by grazing cows. In Animal manure on grassland and forage crops (ed. Van der Meer), pp. 1725. Martinus Nijhoff, Dordrecht, Netherlands.
Villalba, JJ, Provenza, FD 1996. Preference for flavored wheat straw by lambs conditioned with intraruminal administrations of sodium propionate. Journal of Animal Science 74, 23622368.
Villalba, JJ, Provenza, FD 1997a. Preference for wheat straw by lambs conditioned with intraruminal infusions of starch. The British Journal of Nutrition 77, 287297.
Villalba, JJ, Provenza, FD 1997b. Preference for flavored foods by lambs conditioned with intraruminal administrations of nitrogen. The British Journal of Nutrition 78, 545561.
Villalba, JJ, Provenza, FD 1999. Nutrient-specific preferences by lambs conditioned with intraruminal infusions of starch, casein, and water. Journal of Animal Science 77, 378387.
Villalba, JJ, Provenza, FD 2007. Self-medication and homeostatic behaviour in herbivores: learning about the benefits of nature’s pharmacy. Animal 1, 13601370.
Villalba, JJ, Provenza, FD 2009. Learning and dietary choice in herbivores. Rangeland Ecology & Management In Press.
Villalba, JJ, Provenza, FD, Hall, JO, Lisonbee, LD 2009a. Selection of tannins by sheep in response to gastro-intestinal nematode infections. Journal of Animal Science 87 (E-suppl. 2/J), 494.
Villalba, JJ, Manteca, X, Provenza, FD 2009b. Relationship between reluctance to eat novel foods and open-field behavior in sheep. Physiology & Behavior 96, 276281.
Waghorn, GC 1990. Beneficial effects of low concentrations of condensed tannins in forages fed to ruminants. In Microbial and plant opportunities to improve lignocellulose utilization by ruminants (ed. DE Akin, LG Ljungdahl, JR Wilson and PJ Harris), pp. 137247. Elsevier Science Publishing Co., New York, NY, USA.
Westoby, M 1974. An analysis of diet selection by large generalist herbivores. American Naturalist 108, 290304.
Westoby, M 1978. What are the biological bases of varied diets? American Naturalist 112, 627631.
Wilmshurst, JF, Fryxell, JM, Hudson, RJ 1995. Forage quality and patch choice by wapiti (Cervus elaphus). Behavioral Ecology 6, 209217.
Zobel, G, Schwartzkopf-Genswein, KS, Veira, DM, Genswein, BMA, Zobel, AM, Gibb, DJ, von Keyserlingk, MAG 2006. Beef feedlot cattle prefer more forage in their ration when given a choice. Canadian Journal of Animal Science 86, 596597 (abstract).
Recommend this journal

Email your librarian or administrator to recommend adding this journal to your organisation's collection.

  • ISSN: 1751-7311
  • EISSN: 1751-732X
  • URL: /core/journals/animal
Please enter your name
Please enter a valid email address
Who would you like to send this to? *



Full text views

Total number of HTML views: 0
Total number of PDF views: 0 *
Loading metrics...

Abstract views

Total abstract views: 0 *
Loading metrics...

* Views captured on Cambridge Core between <date>. This data will be updated every 24 hours.

Usage data cannot currently be displayed