Honeybee larvae and pupae are extremely stenothermic, i.e. they strongly depend on accurate regulation of brood nest temperature for proper development (33–36°C). Here we study the mechanisms of social thermoregulation of honeybee colonies under changing environmental temperatures concerning the contribution of individuals to colony temperature homeostasis. Beside migration activity within the nest, the main active process is “endothermy on demand” of adults. An increase of cold stress (cooling of the colony) increases the intensity of heat production with thoracic flight muscles and the number of endothermic individuals, especially in the brood nest. As endothermy means hard work for bees, this eases much burden of nestmates which can stay ectothermic. Concerning the active reaction to cold stress by endothermy, age polyethism is reduced to only two physiologically predetermined task divisions, 0 to ∼2 days and older. Endothermic heat production is the job of bees older than about two days. They are all similarly engaged in active heat production both in intensity and frequency. Their active heat production has an important reinforcement effect on passive heat production of the many ectothermic bees and of the brood. Ectothermy is most frequent in young bees (<∼2 days) both outside and inside of brood nest cells. We suggest young bees visit warm brood nest cells not only to clean them but also to speed up flight muscle development for proper endothermy and foraging later in their life. Young bees inside brood nest cells mostly receive heat from the surrounding cell wall during cold stress, whereas older bees predominantly transfer heat from the thorax to the cell wall. Endothermic bees regulate brood comb temperature more accurately than local air temperature. They apply the heat as close to the brood as possible: workers heating cells from within have a higher probability of endothermy than those on the comb surface. The findings show that thermal homeostasis of honeybee colonies is achieved by a combination of active and passive processes. The differential individual endothermic and behavioral reactions sum up to an integrated action of the honeybee colony as a superorganism.

SUMMARY: In order to survive cold northern winters, honeybees crowd tightly together in a winter cluster. Present models of winter cluster thermoregulation consider the insulation by the tightly packed mantle bees as the decisive factor for survival at low temperatures, mostly ignoring the possibility of endothermic heat production. We provide here direct evidence of endothermic heat production by `shivering' thermogenesis. The abundance of endothermic bees is highest in the core and decreases towards the surface. This shows that core bees play an active role in thermal control of winter clusters. We conclude that regulation of both the insulation by the mantle bees and endothermic heat production by the inner bees is necessary to achieve thermal stability in a winter cluster.

The Western honeybees Apis mellifera carnica and A. m. ligustica are closely related subspecies living in neighbouring regions. Metabolism and the upper lethal thermal limits are crucial physiological traits, adapted in the evolutionary process to environment and climate conditions. We investigated whether samples from these two ecotypes differ in these traits. The standard metabolic rate was higher in the A. m. ligustica population only at a high temperature (T _a ~ 40 °C; dVCO₂ = 12 nl s⁻¹; P < 0.05), probably due to a higher body temperature (dT_thorax = 1.5 °C; P < 0.01). The critical thermal maximum of activity and respiration was similar (difference activity CT_max = 0.8 °C, respiratory CT_max = 1.1 °C). The lethal temperature (LT_{50, 8h}) revealed higher tolerance and survival rates of the Ligustica bees (Carnica 50.3 °C; Ligustica 51.7 °C; P < 0.02). Results reveal the adaptation of the two subspecies to their historic climate conditions, possibly favouring Ligustica in a warming environment.

The worldwide distribution of honeybees and their fast propagation to new areas rests on their ability to keep up optimal ‘tropical conditions’ in their brood nest both in the cold and in the heat. Honeybee colonies behave like ‘superorganisms’ where individuals work together to promote reproduction of the colony. Social cooperation has developed strongly in thermal homeostasis, which guarantees a fast and constant development of the brood. We here report on the cooperation of individuals in reaction to environmental variation to achieve thermal constancy of 34–36 °C. The measurement of body temperature together with bee density and in-hive microclimate showed that behaviours for hive heating or cooling are strongly interlaced and differ in their start values. When environmental temperature changes, heat production is adjusted both by regulation of bee density due to migration activity and by the degree of endothermy. Overheating of the brood is prevented by cooling with water droplets and increased fanning, which start already at moderate temperatures where heat production and bee density are still at an increased level. This interlaced change and onset of different thermoregulatory behaviours guarantees a graded adaptation of individual behaviour to stabilise the temperature of the brood.