INVITATION PAPER (C.P. ALEXANDER FUND): FORAGING OF INDIVIDUAL WORKERS IN RELATION TO COLONY STATE IN THE SOCIAL HYMENOPTERA

Workers of social insects are members of colonies that survive and reproduce together. Therefore, the behavioral activities of individual workers should be integrated with colony state. We here summarize and discuss the relationship between colony state and foraging behavior of individual workers under the provisional assumption that the colony is a unit. We argue that colony state can be described by a number of variables that should relate to fitness components in order to be meaningful. Among the possible candidates, colony population size seems to have an overriding importance in many respects, as shown by its relation to fitness components such as survival probability and reproductive performance. Other important variables include colony demography, i.e. caste or size distributions, nutritional status, or queen number. Each of these variables has been shown to affect fitness components; however, the evidence is rather scanty. We also discuss the evidence that variation in colony state variables relates to variation in individual worker behavior. Nutritional status (i.e. low or high levels of food stores) and colony size have been shown repeatedly to affect individual behavior. However, most of the evidence comes from the honey bee. Some studies suggest that behavioral responses are hierarchically structured. More work needs to be done to investigate the actual mechanisms of integration of individual behavior with colony state. Some knowledge has accumulated about the processes that govern recruitment to food sources. We conclude this review by discussing some concepts and problems for further research. These include the concept of a preferred colony state to which the colony should return after disturbance through the behavioral activities of the workers. Further theoretical elaboration and empirical investigations may help to elucidate whether this concept is useful and necessary. A largely neglected issue concerns the number versus effort problem, i.e. whethe