Diploid drones

Normally drones develop from unfertilized eggs and are haploid. Diploid drones (called also "biparental males") develop from fertilized eggs [1][2] which are homozygous at sex locus. In nature diploid drones do not survive until the end of larval development. The larvae of diploid drones are eaten by workers [3] within few hours after hatching from egg [4] despite the fact that they are viable [5][6].

Adult (imago) diploid drones can be reared in laboratory by hatching eggs in incubator and feeding larvae with royal jelly without workers [7][8]. The larva can be transferred to colony after 2-3 days. At this age workers feed them normally. Diploid drones can be reared also in autumn in mating nuclei with about 1000 workers [9].

Externally adult diploid drones are similar to haploid drones. In comparison to haploid drones diploid once are larger, heavier [10][11][12] but see [13][14], have smaller testes [15][14], fewer testicular tubules [16], fewer wing hooks [14] and lower vitellogenin concentration [14]. Diploid drone larvae produce more cuticular hydrocarbons than workers but less than haploid drones [17] but see [18].
Diploid drones produce diploid spermatozoa [19] containing twice as much DNA as haploid spermatozoa [20][14]. Diploid spermatozoa are longer than haploid spermatozoa; their head is particularly long [21]. Ultrastructure of haploid and diploid drones is similar [22]. In theory triploid honey bees can be obtained by inseminating queen with diploid spermatozoa [21], however, this was not achieved so far because of small number of sperm produced by diploid drones.

Workers recognize the diploid drones larvae using substances present at their bodies [23]. It was suggested that diploid drones produce pheromone called "cannibalism substance" which is a signal to workers that they should be destroyed [23] see also [24]. Such self-destructive behaviour of diploid drones can evolve because they are neither able to reproduce nor help their relatives. Eating of the diploid drones at early stage of larval development allows to save valuable resources and produce bigger number of their relatives. However, no cuticular compound specific for diploid drone larvae was found [17]. First instar larvae of haploid and diploid drones differ in relative amount of cuticular compounds [17] and the difference can be used by workers for detection of diploid drones. In older larvae the differences in cuticular compounds are smaller [14].

In natural conditions frequency of diploid drones (before destruction by workers) in a colony is 0.05±0.03 (mean±SD) [25]. The frequency can be much higher in case of inbreeding. In colonies with large proportion of diploid drones there is "shot brood" - brood of different ages scattered irregularly on a comb [26][27][28][29]. Multiple mating by the queen leads to reduced variance of proportion of diploid drones present in the colony [30]. When a queen is artificially inseminated with semen of one drone which is her brother, half of her female offspring develop into diploid drones [26].

Reviews: [31][32]
Other references: [33][34][35][36][37][38][39][40][41][42]

References

  1. Woyke J., Knytel A. (1966) The chromosome number as proof that drones can arise from fertilized eggs of the honeybee. Journal of Apicultural Research 5:149–154.
  2. Woyke J., Knytel A., Bergandy K. (1966) The presence of spermatozoa in eggs as proof that drones can develop from inseminated eggs of the honeybee. Journal of Apicultural Research 5:71–78.
  3. Woyke J. (1963) What happens to diploid drone larvae in a honeybee colony. J. Apic. Res. 2:73-75.
  4. Woyke J. (1962) The hatchability of "lethal" eggs in a two sex allele fraternity of honeybees. J. Apic. Res. 1:6-13.
  5. Woyke J. (1963) Rearing and viability of diploid drone larvae. J. Apic. Res. 2:77–84.
  6. Woyke J. (1965) Study on the comparative viability of diploid and haploid larval drone honeybees. Journal of Apicultural Research 4:12–16.
  7. Woyke J. (1969) A method of rearing diploid drones in a honeybee colony. J. Apic. Res. 8:65-74.
  8. Woyke J. (1969) Rearing diploid drones on royal jelly or bee milk. Journal of Apicultural Research 8:169-173.
  9. Polaczek B., Neumann P., Schricker B., Moritz R.F.A. (2000) A new, simple method for rearing diploid drones in the honeybee (Apis mellifera L.). Apidologie 31:525–530.
  10. Woyke J. (1977) Comparative biometrical investigation on diploid drones of the honeybee. I. The head. J. Apic. Res 16:131–142.
  11. Woyke J. (1978) Comparative biometrical investigation on diploid drones of the honeybee. II. The thorax. Journal of Apicultural Research 17:195–205.
  12. Woyke J. (1978) Comparative biometrical investigation on diploid drones of the honeybee. III. The abdomen and weight. J. Apic. Res 17:206–217.
  13. Chaud-Netto J. (1975) Sex determination in Bees. II. Additivity of maleness genes in Apis mellifera. Genetics 79:213-217.
  14. Herrmann M., Trenzcek T., Fahrenhorst H., Engels W. (2005) Characters that differ between diploid and haploid honey bee (Apis mellifera) drones. Genet. Mol. Res. 4:624-641.
  15. Woyke J. (1974) Genic balance, heterozygosity and inheritance of testis size in diploid drone honeybees. Journal of Apicultural Research 13:77–85.
  16. Woyke J. (1973) Reproductive organs of haploid and diploid drone honeybees. J. Apic. Res. 12:35-51.
  17. Santomauro G., Oldham N.J., Boland W., Engels W. (2004) Cannibalism of diploid drone larvae in the honey bee (Apis mellifera) is released by odd pattern of cuticular substances. Journal of Apicultural Research 43:69–74.
  18. Bienefeld K., Mattausch A., Möller U., Pritsch G. (1994) Ursache von Kannibalismus bei Arbeiterinnen der Honigbiene (Apis mellifera L.) an diploider Drohnenbrut. Verhandlungen der Deutschen Zoologischen Gesellschaft 87:30.
  19. Woyke J., Skowronek W. (1974) Spermatogenesis in diploid drones of the honeybee. Journal of Apicultural Research 13:183–190.
  20. Woyke J. (1975) DNA content of spermatids and spermatozoa of haploid and diploid drone honeybees. Journal of Apicultural Research 14:3–8.
  21. Woyke J. (1983) Lengths of haploid and diploid spermatozoa of the honeybee and the question of the production of triploid workers. J Apic Res 22:146-149.
  22. Woyke J. (1984) Ultrastructure of single and multiple diploid honeybee spermatozoa. Journal of Apicultural Research 23:123–135.
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  24. Dietz A., Lovins R.W. (1975) Studies on the ‘cannibalism substance’ of diploid drone honey bee larvae. Journal of the Georgia Entomological Society 10:314-315.
  25. Adams J., Rothman E.D., Kerr W.E., Paulino Z.L. (1977) Estimation of the number of sex alleles and queen matings from diploid male frequencies in a population of Apis mellifera. Genetics 86:583-596.
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