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

  • Paul Schmid-Hempel (a1), Mark L. Winston (a2) and Ron C. Ydenberg (a2)

Abstract

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. whether individuals should work harder or more workers should be allocated to a task that is in demand. We propose a simple scenario that suggests testable predictions. Finally, we discuss how colony state, individual work load, and the dependence of worker mortality rate on activity level may interact to generate different short-term foraging strategies that workers should adopt.

Chez les insectes sociaux, les ouvrières appartiennent à une colonie où la survie et la reproduction sont des phénomènes collectifs. Les activités comportementales de chacune des ouvrières font donc partie intégrante de l’état de la colonie. La relation entre l’état de la colonie et le comportement de quête de nourriture d’ouvrières particulières est définie et examinée dans le cadre de l’hypothèse selon laquelle la colonie fonctionne comme une unité. Nous croyons que l’état de la colonie peut être décrit par un certain nombre de variables qui n’ont de sens que si elles sont reliées aux composantes du "fitness." Parmi les variables possibles, la taille de la population dans la colonie semble avoir une importance capitale par plusieurs aspects, comme le démontre sa relation avec des composantes telles que la probabilité de survie et le succès de la reproduction. Parmi les autres variables importantes, il faut mentionner la démographie de la colonie, i.e. la répartition selon la caste ou la taille, le statut alimentaire ou le nombre de reines. Chacune de ces variables a un effet sur les composantes du "fitness," mais cependant les preuves en sont plutôt faibles. Nous examinons aussi l’hypothèse d’une relation entre les fluctuations des variables associées à l’état de la colonie et celles du comportement individuel des ouvrières. Le statut alimentaire (i.e. quantités faibles ou élevées de réserves de nourriture) et la taille de la colonie ont été reconnus à plusieurs reprises comme des facteurs qui affectent le comportement individuel. Cependant, les preuves à l’appui de cette affirmation viennent toutes d’études sur l’Abeille domestique. Certains travaux indiquent que les réactions comportementales dépendent de la hiérarchie sociale. Les mécanismes réels de la contribution des comportements individuels à l’état de la colonie doivent être soumis à des études plus poussées. Les processus qui régissent le recrutement aux sources de nourriture sont mieux connus maintenant. Nous concluons cette synthèse par l’examen de certaines théories et de problèmes qui devront être étudiés plus en détails, en particulier la théorie d’un état optimal que la colonie devrait retrouver après les perturbations entraînées par les activités comportementales des ouvrières. Des spéculations théoriques et des recherches empiriques pourront déterminer si cette théorie est nécessaire, voire utile. La question du nombre-versus-l’effort est toujours fortement négligée; chaque individu devrait-il travailler plus fort ou serait-il préférable que plus d’ouvrières soient affectées à là tâche requise. Nous proposons ici un scénario simple qui suppose des prédictions vérifiables par des tests. Finalement, nous examinons comment l’état de la colonie, la charge de travail de chaque individu et la relation entre le taux de mortalité et la somme de travail peuvent se combiner pour donner lieu à de nouvelles stratégies de recherche de nourriture à court terme que pourraient adopter les ouvrières.

[Traduit par la rédaction]

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