Risk-sensitive foraging: the effect of relative variability

Animals are sometimes sensitive and sometimes indifferent to variability in amount of reward. I analyzed a large set of published studies to test whether these differences in risk-sensitivity may be influenced by the coefficient of variation (s.d./mean) of the variable reward and by the presence of empty rewards. The coefficient of variation had a strong effect on the strength of risk-sensitivity among studies with nectarivores, both invertebrates and vertebrates. The effect was not significant for studies with non-nectarivores, but these studies included only a limited range of coefficients of variation, and risk-sensitivity in these studies was based on fewer subjects per experimental condition than in studies of nectarivores. The presence of empty rewards had no effect on the strength of risk-sensitivity. The apparent effect of empty rewards seems to be an artifact of the tendency of distributions that include zero rewards to have larger coefficients of variation than distributions that do not include zero rewards. I introduce the CV model, a simple descriptive model that accounts for a large proportion of the variability in levels of risk-sensitivity among experiments. My analysis resolves the apparent discrepancies among studies that report risk-sensitivity and risk-indifference. It is necessary to control for the magnitude of the coefficient of variation when testing risk-sensitivity under different conditions (e.g., energy budgets). A successful descriptive model of risk-sensitive choice behavior is useful for guiding future research both in ecology and in the underlying mechanism of cognitive processes.

[1]  L. Real,et al.  Risk-sensitive foraging: choice behaviour of honeybees in response to variability in volume of reward , 1999, Animal Behaviour.

[2]  C. Herrera,et al.  Population-level estimates of interannual variability in seed production : what do they actually tell us ? , 1998 .

[3]  N. Waser Pollination, angiosperm speciation, and the nature of species boundaries , 1998 .

[4]  Peter Dayan,et al.  A Neural Substrate of Prediction and Reward , 1997, Science.

[5]  E. Weber,et al.  Perceived risk attitudes: relating risk perception to risky choice , 1997 .

[6]  R. Hilborn,et al.  The Ecological Detective: Confronting Models with Data , 1997 .

[7]  P. D. Smallwood,et al.  RISK-SENSITIVE BEHAVIOR : WHERE DO WE GO FROM HERE ? , 1996 .

[8]  A. Kacelnik,et al.  Risky Theories—The Effects of Variance on Foraging Decisions , 1996 .

[9]  K. Waddington,et al.  Carpenter bee (Xylocopa micans) risk indifference and a review of nectarivore risk-sensitivity studies , 1996 .

[10]  Peter Dayan,et al.  Bee foraging in uncertain environments using predictive hebbian learning , 1995, Nature.

[11]  M. Hammer,et al.  Learning and memory in the honeybee , 1995 .

[12]  K. Waddington,et al.  Risk-sensitive foraging in honey bees: no consensus among individuals and no effect of colony honey stores , 1994, Animal Behaviour.

[13]  Juan C. Reboreda,et al.  Risk sensitivity in starlings: variability in food amount and food delay , 1991 .

[14]  R. Cartar A TEST OF RISK-SENSITIVE FORAGING IN WILD BUMBLE BEES' , 1991 .

[15]  J. Ha Risk-sensitive foraging: the role of ambient temperature and foraging time , 1991, Animal Behaviour.

[16]  T. Caraco,et al.  Risk-sensitivity: ambient temperature affects foraging choice , 1990, Animal Behaviour.

[17]  J. Wunderle,et al.  Constant vs. Variable Risk‐Aversion in Foraging Bananaquits , 1988 .

[18]  L. Real,et al.  Why are Bumble Bees Risk Averse , 1987 .

[19]  Anne Lohrli Chapman and Hall , 1985 .

[20]  T. Caraco Aspects of risk-aversion in foraging white-crowned sparrows , 1982, Animal Behaviour.

[21]  R. Hogarth Judgement and choice: The psychology of decision , 1982 .

[22]  B. Heinrich,et al.  Floral preferences of bumblebees (Bombus edwardsii) in relation to intermittent versus continuous rewards , 1981, Animal Behaviour.

[23]  L. Real UNCERTAINTY AND POLLINATOR-PLANT INTERACTIONS: THE FORAGING BEHAVIOR OF BEES AND WASPS ON ARTIFICIAL FLOWERS' , 1981 .

[24]  T. Caraco,et al.  An empirical demonstration of risk-sensitive foraging preferences , 1980, Animal Behaviour.

[25]  Thomas Caraco,et al.  ON FORAGING TIME ALLOCATION IN A STOCHASTIC ENVIRONMENT , 1980 .

[26]  C. C. Perkins,et al.  The relation between mean reward and mean reinforcement. , 1959, Journal of experimental psychology.

[27]  K. Waddington Foraging behavior of nectarivores and pollen collectors , 1997 .

[28]  J. Bacon,et al.  THE EVOLUTION OF ARTHROPOD NERVOUS SYSTEMS , 1997 .

[29]  S. M. Perez The risk-sensitive foraging behavior of carpenter bees (Xylocopa micans) , 1996 .

[30]  D. DeAngelis,et al.  Individual-Based Models and Approaches in Ecology , 1992 .

[31]  C. Gallistel The organization of learning , 1990 .

[32]  Robin M. Hogarth,et al.  Judgement and choice: The psychology of decision, 2nd ed. , 1987 .

[33]  J. Kagel,et al.  Animals' Choices over Uncertain Outcomes: Some Initial Experimental Results , 1985 .

[34]  D. Norman Learning and Memory , 1982 .

[35]  L. Real CHAPTER 2 – On Uncertainty and the Law of Diminishing Returns in Evolution and Behavior , 1980 .