Pheromones are intraspecific chemical signals. They can have profound effects on the behaviour and/or physiology of the receiver, and it is still common to hear pheromones described as controlling of the behaviour of the receiver. The discussion of pheromonal control arose initially from a close association between hormones and pheromones in the comparative physiological literature, but the concept of a controlling pheromone is at odds with contemporary signal evolution theory, which predicts that a manipulative pheromonal signal negatively affecting the receiver's fitness should not be stable over evolutionary time. Here we discuss the meaning of pheromonal control, and the ecological circumstances by which it might be supported. We argue that in discussing pheromonal control it is important to differentiate between control applied to the effects of a pheromone on a receiver's physiology (proximate control), and control applied to the effects of a pheromone on a receiver's fitness (ultimate control). Critically, a pheromone signal affecting change in the receiver's behaviour or physiology need not necessarily manipulate the fitness of a receiver. In cases where pheromonal signalling does lead to a reduction in the fitness of the receiver, the signalling system would be stable if the pheromone were an honest signal of a social environment that disadvantages the receiver, and the physiological and behavioural changes observed in the receiver were an adaptive response to the new social circumstances communicated by the pheromone.

No abstract

No abstract

The presence of the honey bee queen reduces worker ovary activation. When the queen is healthy and fecund, this is interpreted as an adaptive response as workers can gain fitness from helping the queen raise additional offspring, their sisters. However, when the queen is absent, workers activate their ovaries and lay unfertilized eggs that become males. Queen pheromones are recognised as a factor affecting worker ovary activation. Recent work has shown that queen mandibular pheromone composition changes with queen mating condition and workers show different behavioural responses to pheromone extracts from these queens. Here, we tested whether workers reared in colonies with queens of different mating condition varied in level of ovary activation. We also examined the changes in the chemical composition of the queen mandibular glands to determine if the pheromone blend varied among the queens. We found that the workers activated their ovaries when queens were unmated and had lower ovary activation when raised with mated queens, suggesting that workers detect and respond adaptively to queens of differing mating status. Moreover, variation in queen mandibular gland's chemical composition correlated with the levels of worker ovary activation. Although correlative, this evidence suggests that queen pheromone may act as a signal of queen mating condition for workers, in response to which they alter their level of ovary activation.

If a honeybee (Apis mellifera) colony loses its queen, worker bees develop their ovaries and produce male offspring [1]. Kin selection theory predicts that the degree of altruism in queenless colonies should be reduced because the relatedness of workers to a hivemate's offspring is less in queenless colonies than it is to the daughters of the queen in queenright colonies [2-4]. To explore this hypothesis, we examined the behavior and physiology of queenless egg-laying workers. Queenless bees engaged in both personal reproduction and the social foraging and defense tasks that benefited their colony. Laying workers also had larger brood-food-producing and wax glands, showing metabolic investments in both colony maintenance and personal reproduction. Whereas in queenright colonies there is a very clear age-based pattern of division of labor between workers, in queenless colonies the degree of individual specialization was much reduced. Queenless colonies functioned as a collective of reproductive and behaviorally generalist bees that cooperatively maintained and defended their nest. This social structure is similar to that observed in a number of primitively social bee species [5]. Laying workers therefore show a mix of selfish personal reproduction and altruistic cooperative behavior, and the queenless state reveals previously unrecognized plasticity in honeybee social organization.