The regulatory anatomy of honeybee lifespan.

Honeybee workers (Apis mellifera) may be classified as either short-lived summer bees or long-lived winter bees in temperate zones. The protein status appears to be a major determinant of honeybee lifespan, and the lipoprotein vitellogenin seems to play a crucial role. Here, we give a review of the role of the vitellogenin in honeybee workers, and present a data-driven mathematical model describing the dynamics of this representative protein in the individual bee as a function of its task profile under various regimes. The results support the hypothesis that vitellogenin is a true storage protein that is utilized for various metabolic purposes including the synthesis of brood food. Except for workers having been foragers for many days, they also suggest that the previous life histories of workers do not constrain them from becoming winter bees as long as they get ample food and time to build up their protein reserves before wintering. The results also indicate that it may not be necessary to introduce the ovary as a storage organ for vitellogenin in order to generate normal winter bees. The insights gained from these results are then discussed in a broader gerontological and life history context. Remarkably similar features concerning regulation of ageing in Caenorhabditis elegans, Drosophila melanogaster and honeybees are pointed out and discussed. Furthermore, we show that in contrast to the "mutation accumulation" and the "antagonistic pleiotropy" evolutionary theories of ageing, the "disposable soma" theory is capable of explaining the bimodal longevity distribution of honeybees when interpreted in a group selection context. Finally, by showing that depletion of nutrient stores can be actively controlled by pathways connected to regulation of ageing, we strengthen the claim that age-based division of labour, with performance of risky tasks delayed until late in life by workers with depleted nutrient stores, may have evolved as an energy-saving mechanism in insect colonies.