By
Dr. Clarence Collison
Honey bees are social insects that exhibit striking caste specific differences in longevity. All honey bee castes exhibit different developmental times and life spans. Drones have a developmental time of 24 days, and they live on average 21-32 days in the spring and summer, the only times when drones are produced (Winston 1987). Drones under some conditions can live at least 59 days (Page and Peng 2001). Workers develop in 21 days and their life spans vary depending on season. Workers live on average 15-38 days in the summer (Free and Spencer- Booth 1959), 30-60 days in the spring and fall, and 150-200 days in the winter (Remolina and Hughes 2008). Queens take the shortest time to develop (16 days) and have the longest life span. Queens in a commercial hive are generally replaced after 1-2 years (for productivity reasons) and have adult life spans of 2-5 years (Sammataro and Avitabile 1998) with the longest recorded lifespan for a honey bee queen being 8 years (Bozina 1961 cited by Page and Peng 2001).
Caste differentiation for larvae that become queens or workers has been associated with the quantity and quality of food that larvae receive before the third larval instar. Larvae fed a rich diet composed of royal jelly develop into queens; whereas larvae fed a diet composed of glandular secretions, honey, and pollen develop into workers (Winston 1987). The dimorphism observed in the honey bee female caste is particularly interesting because workers and queens have the same genotype yet exhibit a 10- fold difference in life span.
With the exception of the mating flights queens take early in life, they do not leave the protected hive environment except during colony swarming. Adult queens are protected from predators and environmental extremes by the physical structure of their nests and defensive behaviors of non- reproductive workers, which may increase their chances of having a longer life span. The physiological and molecular basis of queen longevity and the dramatic lifespan differences between queens and workers is just beginning to be explored.
Worker honey bees undergo age development after they become adults. Workers usually initiate foraging behavior when they are 2-3 weeks old. The age at which a worker initiates foraging is a strong determinant of her length of life. This is presumed to be a result of the hazards of foraging, but natural senescence also occurs (Page and Peng 2001). Senescence can be defined demographically as an age-dependent increase in mortality risk, or functionally as a decline in performance. The relationship between the two phenomena is central for understanding the biological aging process (Rueppell et al. 2007b).
Studies of senescence in the honey bee have focused on establishing the importance of extrinsic mortality factors (predation, weather) and behavior (nursing and foraging) in worker honey bee longevity. Senescence can be delayed or speeded up by preventing workers from leaving the nest or forcing them to do so earlier (Rueppell et al. 2007a). Hive bees, which are prevented from foraging, can live over eight times longer than foragers (Neukirch 1982, Rueppell et al. 2007a, Amdam et al. 2004), and non-flying winter bees in the ‘diutinus stage’ (stress resistant form) can live up to eight months (Amdam et al. 2004, Omholt and Amdam 2004). Senescence obviously starts with foraging: workers show a decline in physiological resistance to stress, such as heat, desiccation and starvation (Remolina et al. 2007) and their hemocyte count drops (Amdam et al. 2005), while their behavioral performance appears to remain unchanged (Rueppell et al. 2007b).
In an explicit effort to compare intrinsic vs extrinsic mortality factors in the determination of worker lifespan, Rueppell et al. (2007a) compared the lifespan of workers that foraged in a protected environment (flight cage) and workers that foraged in a natural setting. They also varied the amount of time bees were allowed to forage in a flight cage. Free-foraging mortality was higher than flight- cage mortality, but both groups showed increasing mortality rates with age consistent with senescence. Limited foraging opportunities in the cage had no overall effect on lifespan. There was also a negative correlation between age at first foraging and foraging lifespan, suggestive of pre-foraging senescence. In a separate study, Rueppell et al. (2007b) assessed age-dependent mortality and behavioral performance of foragers. They found that workers experienced an increase in mortality with chronological age, but their performance in behavioral assays related to foraging activity did not decline with age.
To eliminate the confounding effects of increased extrinsic hazard and energy expenditure faced by foraging bees, Remolina et al. (2007) used a social manipulation to prevent nurse bees from transitioning to foraging. They tested whether older nurses were more susceptible to different kinds of stress: starvation, heat and oxidative damage. In this study, all forms of stress resistance decreased in older bees, and this manifestation of senescence was evident by 30 days of age. They therefore concluded that intrinsic senescence affects nurse lifespan independently of extrinsic mortality force. Although there are a few contradictory reports, overall the evidence supports the proposition that worker bees (both nurses and foragers) exhibit senescence.
References:
Amdam, G.V., A. Aase, S.C. Seehuus, M.K. Fondrk and K. Hartfelder 2005. Social reversal of immunosenescence in honey bee workers. Exp. Gerontol. 40: 939-947.
Amdam, G.V., Z.L.P. Simões, A. Hagen, K. Norberg, K. Schrøder, Ø. Mikkelsen, T.B.L. Kirkwood and S.W. Omholt 2004. Hormonal control of the yolk precursor vitellogenin regulates immune function and longevity in honeybees. Exp. Gerontol. 39: 767- 773.
Bozina, K.D. 1961. How long does the queen live? Pchelovodstvo 38: 13. Free, J.B. and Y. Spencer-Booth 1959. The longevity of worker honey bees (Apis mellifera). Proc. R. Entomol. Soc. 34: 141- 150.
Neukirch, A. 1982. Dependence of the lifespan of the honey bee (Apis mellifica) upon flight performance and energy consumption. J. Comp. Physiol. 146: 35- 40.
Omholt, S.W. and G.V. Amdam 2004. Epigenetic regulation of aging in honeybee workers. Sci. Aging Knowl. Environ. 26: pe28. Bee. Harvard University Press, Cambridge, MA.
Beekeepers Welcome! Page, R.E. and C.Y.-S. Peng 2001. Aging and development in social insects with emphasis on the honey bee, Apis mellifera L. Exp. Gerontol. 36: 695-711.
Remolina, S.C. and K.A. Hughes 2008. Evolution and mechanisms of long life and high fertility in queen honey bees. Age 30: 177-185.
Remolina, S.C., D.M. Hafez, G.E. Robinson and K.A. Hughes 2007. Senescence in the worker honey bee Apis mellifera. J. Insect Physiol. 53: 1027-1033. Rueppell, O., C. Bachelier, M.K. Fondrk and R.E. Page, Jr. 2007a. Regulation of life history determines lifespan of workers honey bees (Apis mellifera L.) Exp. Gerontol. 42: 1020-1032.
Rueppell, O., S. Christine, C. Mulcrone and L. Groves 2007b. Aging without functional senescence in honey bee workers. Curr. Biol. 17: R274-R275.
Sammataro, D. and A. Avitabile 1998. The Beekeeper’s Handbook, Third Edition, Cornell Univ. Press., Ithaca, NY.