Journal of Chemical Ecology

, Volume 20, Issue 3, pp 609–624 | Cite as

How goats learn to distinguish between novel foods that differ in postingestive consequences

  • Frederick D. Provenza
  • Justin J. Lynch
  • Elizabeth A. Burritt
  • Cody B. Scott


To better understand some of the mechanisms that control selection of novel foods differing in postingestive consequences, we offered goats current season's (CSG) and older (OG) growth twigs from the shrub blackbrush (Coleogyne ramosissima). CSG is higher than OG in nitrogen (1.04% v. 0.74%) and it is more digestible in vitro in goat rumen fluid (48% v. 38%). Nevertheless, goats acquire a preference for OG because CSG contains much higher levels of a condensed tannin that causes a learned food aversion. When CSG and OG were offered to goat naive to blackbrush, the goats did not choose either OG or CSG exclusively, but when they finally (1) ate more CSG than OG within a meal (averages of 44 g and 16 g, respectively) and (2) ate enough CSG within the meal to acquire an aversion (average of 44 g), they ingested less CSG than OG from then onward. Accordingly, the change in food selection resulting from postingestive feedback was influenced by the amount of each food ingested within a meal. This was further shown when we varied the amounts of CSG and OG that goats ingested within a meal, and then gave them by gavage the toxin lithium chloride (LiCl). They subsequently ate less of the food eaten in the greatest amount, regardless of whether it was CSG or OG. The salience of the flavor (i.e., taste and odor) of CSG and OG also played a role in the acquired aversion to CSG. Salience evidently was due to a flavor common to both OG and CSG that was more concentrated in CSG. We conclude that the relative amounts of different foods ingested within a meal, and the salience of the flavors of those foods, are both important variables that cause goats to distinguish between novel foods that differ in postingestive consequences.

Key Words

Toxin food selection food aversion secondary metabolite nutrition palatability lithium chloride ruminants goat Capra sp 


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  1. Berenbaum, M. 1986. Postingestive effects of phytochemicals on insects: OnParacelsus and plant products, pp. 121–153,in J.R. Miller andT.A. Miller (eds.). Insect-Plant Interactions. Springer-Verlag, New York.Google Scholar
  2. Bernays, E.A., andChapman, R.F. 1987. Evolution of plant deterrence to insects, pp. 159–173,in R.F. Chapman, E.A. Bernays, and J.G. Stoffolano (eds.). Perspectives in Chemoreception and Behavior. Springer-Verlag, New York.Google Scholar
  3. Bernays, E.A., andCornelius, M. 1992. Relationship between deterrence and toxicity of plant secondary compounds for the alfalfa weevilHypera brunneipennis.Entomol. Exp. Appl. 64:289–292.Google Scholar
  4. Bond, N., andDiGuisto, E. 1975. Amount of solution drunk is a factor in the establishment of taste aversion.Anim. Learn. Behav. 3:81–84.Google Scholar
  5. Bryant, J.P., Provenza, F.D., Pastor, J. Reichardt, P.B., Clausen, T.P., andDuToit, J.T. 1991. Interactions between woody plants and browsing mammals mediated by secondary metabolites.Annu. Rev. Ecol. Syst. 22:431–446.Google Scholar
  6. Burritt, E.A. andProvenza, F.D. 1989. Food aversion learning: Ability of lambs to distinguish safe from harmful foods.J. Anim. Sci. 67:1732–1739.PubMedGoogle Scholar
  7. Burritt, E.A. andProvenza, F.D. 1991. Ability of lambs to learn with a delay between food ingestion and consequences given meals containing novel and familiar foods.Appl. Anim. Behav. Sci. 32:179–189.Google Scholar
  8. Burritt, E.A., andProvenza, F.D. 1992. Lambs form preferences for non-nutritive flavors paired with glucose,J. Anim. Sci. 70:1133–1136.PubMedGoogle Scholar
  9. Cannon, D.S., Berman, R.F., Baker, T.B., andAtkinson, C.A. 1975. Effect of preconditioning unconditioned stimulus experience on learned taste aversions.J. Exp. Psycho. Anim. Behav. Processes 104:270–284.Google Scholar
  10. Cannon, D.S., Best, M.R., Batson, J.D., Brown, E.R., Rubenstein, J.A., andCarrell, L.E. 1985. Interfering with taste aversion learning in rats: The role of associative interference.Appetite 6:1–19.PubMedGoogle Scholar
  11. DuToit, J.T., Provenza, F.D, andNastis, A. 1991. Conditioned food aversions: How sick must a ruminant get before it learns about toxicity in foods?Appl. Anim. Behav. Sci. 30:35–46.Google Scholar
  12. Egan, A.R., andRogers, Q.R. 1978. Amino acid imbalance in ruminant lambs.Aust. J. Agric. Res. 29:1263–1279.Google Scholar
  13. Feeny, P. 1992. The evolution of chemical ecology: contributions from the study of herbivorous insects, pp. 1–44, in G.A. Rosenthal and M.R. Berenbaum (eds.). Herbivores: Their Interactions with Secondary Plant Metabolites, 2nd ed. Academic Press, New York.Google Scholar
  14. Freeland, W.J., andJanzen, D.H. 1974. Strategies in herbivory by mammals: The role of plant secondary compounds.Am. Nat. 108:269–289.Google Scholar
  15. Garcia, J. 1989. Food for Tolman: Cognition and cathexis in concert, pp. 45–85,in T. Archer and L. Nilsson (eds.). Aversion, Avoidance and Anxiety. Erlbaum Hillsdale, New Jersey.Google Scholar
  16. Garcia, J., Lasiter, P.A., Bermudez-Rattoni, F., andDeems, D.A. 1985. A general theory of aversion learning, pp. 8–21,in N.S. Braveman and P. Bronstein (eds.). Experimental Assessments and Clinical Applications of Conditioned Food Aversions. New York Academy of Science, New York.Google Scholar
  17. Hay, M.E., andW. Fenical. 1988. Marine plant-herbivore interactions: The ecology of chemical defense.Annu. Rev. Ecol. Syst. 19:111–145.Google Scholar
  18. Kalat, J.W., andRozin, P. 1970. “Salience:” A factor which can override temporal contiguity in taste-aversion learning.J. Comp. Physiol. Psychol. 71:192–197.Google Scholar
  19. Kalat, J.W., andRozin, P. 1971. Role of interference in taste-aversion learning.J. Comp. Physiol. Psychol. 77:53–58.PubMedGoogle Scholar
  20. Kalat, J.W., andRozin, P. 1973. “Learned safety” as a mechanism in long-delay taste-aversion learning in rats.J. Comp. Physiol. Psychol. 83:198–207.PubMedGoogle Scholar
  21. Lane, M.A., Ralphs, M.A., Olsen, J.D., Provenza, F.D., andPfister, J.A. 1990. Conditioned taste aversion: Potential for reducing cattle loss to larkspur.J. Range. Manage. 43:127–131,Google Scholar
  22. Launchbaugh, K.L., andProvenza, F.D. 1993. Can plants practice mimicry to avoid grazing by mammalian herbivores?Oikos 66:501–504.Google Scholar
  23. Launchbaugh, K.L., Provenza, F.D., andBurritt, E.A. 1993. How herbivores track variable environments: Response to variability of phytotoxins.J. Chem. Ecol. 19:1047–1056.Google Scholar
  24. Olsen, J.D., andRalphs, M.H. 1986. Feed aversion induced by intraruminal infusion with larkspur extract in cattle.Am. J. Vet. Res. 47:1829–1833.PubMedGoogle Scholar
  25. Pfister, J.A., Provenza, F.D., andManners, G.D. 1990. Ingestion of tall larkspur by cattle: Separating the effects of flavor from post-ingestive consequences.J. Chem. Ecol. 16:1697–1705.Google Scholar
  26. Provenza, F.D. 1994a, A functional explanation for food selection and the nutritional wisdom of ruminants.J. Range Manage. Accepted.Google Scholar
  27. Provenza, F.D. 1994b. Ontogeny and social transmission of food selection in domesticated ruminants.In P. Valsecchi andB.G. Galef, Jr. (eds.). Ontogeny and Social Transmission of Food Preferences in Mammals: Basic and Applied Research. In press.Google Scholar
  28. Provenza, F.D., andBalph, D.F. 1990. Applicability of five diet-selection models to various foraging challenges ruminants encounter, pp. 423–459,in R.N. Hughes (ed.). Behavioral Mechanisms of Food Selection. NATO ASI Series G: Ecological Sciences, Vol. 20. Springer-Verlag, New York.Google Scholar
  29. Provenza, F.D., andCincotta, R.P. 1993. Foraging as a self-organizational learning process: Accepting adaptability at the expense of predictability, pp. 78–101in R.N. Hughes (ed.). Diet Selection. Blackwell, London.Google Scholar
  30. Provenza, F.D., andMalechek, J.C. 1984. Diet selection by domestic goats in relation to blackbrush twig chemistry.J. Appl. Ecol. 21:831–841.Google Scholar
  31. Provenza, F.D., Pfister, J.A., andCheney, C.D. 1992. Mechanisms of learning in diet selection with reference to phytotoxicosis in herbivores.J. Range Manage. 45:36–45.Google Scholar
  32. Provenza, F.D., Malechek, J.C., Urness, P.J., andBowns, J.E. 1983a. Some factors affecting twig growth in blackbrush.J. Range Manage. 36:518–520.Google Scholar
  33. Provenza, F.D., Bowns, J.E., Urness, P.J., Malechek, J.C., andButcher, J.E. 1983b. Biological manipulation of blackbrush by goat browsing.J. Range Manage. 36:513–518.Google Scholar
  34. Provenza, F.D., Burritt, E.A., Clausen, T.P., Bryant, J.P., Reichardt, P.B., andDistel, R.A. 1990. Conditioned flavor aversion: A mechanism for goats to avoid condensed tannins in blackbrush.Am. Nat. 136:810–828.Google Scholar
  35. Provenza, F.D., Lynch, J.J. andNolan, J.V., 1993a. The relative importance of mother and toxicosis in the selection of foods by lambs.J. Chem. Ecol. 19:313–323.Google Scholar
  36. Provenza, F.D., Lynch, J.J. andCheney, C.D., 1994a. An experimental analysis of the effects of a salient flavor and food deprivation on the intake of novel foods by sheep.J. Anim. Sci. Submitted.Google Scholar
  37. Provenza, F.D., Lynch, J.J., andNolan J.V., 1994b. Food aversion conditioned in anesthetized sheep (Ovis aries).Physiol. Behav. In press.Google Scholar
  38. Provenza, F.D., Nolan, J.V., andLynch, J.J. 1993b. Temporal contiguity between food ingestion and toxicosis affects the acquisition of food aversions in sheep.Appl. Anim. Behav. Sci. 38:269–281.Google Scholar
  39. Provenza, F.D., Ortega-Reyes, L., Scott, C.B., Lynch, J.J., andBurritt, E.A. 1994c. Antiemetic drugs attenuate food aversions in sheep.J. Anim. Sci. accepted.Google Scholar
  40. Revusky, S.H., andBedarf, E.W. 1967. Association of illness with prior ingestion of novel foods.Science 155:219–220.PubMedGoogle Scholar
  41. Rogers, Q.R., andEgan, A.R. 1975. Amino acid imbalance in the liquid-fed lamb.Aust. J. Biol. Sci. 28:169–181.PubMedGoogle Scholar
  42. Winer, B.V. 1971. Statistical Principles in Experimental Design. McGraw-Hill, New York, 907 pp.Google Scholar

Copyright information

© Plenum Publishing Corporation 1994

Authors and Affiliations

  • Frederick D. Provenza
    • 1
  • Justin J. Lynch
    • 2
  • Elizabeth A. Burritt
    • 1
  • Cody B. Scott
    • 1
  1. 1.Range Science DepartmentUtah State UniversityLogan
  2. 2.CSIRO Division of Animal ProductionArmidaleAustralia

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