Drone production

Drones usually are not present in the colony during the whole year. In moderate climate they can be found in most colonies during spring and summer [1]. Only in queenless colonies drones can be present during autumn and winter. Colonies with larger number of workers start drone production earlier [2]. Initiation of drone rearing occurs about three weeks before swarming but this time varies greatly between colonies [3]. Maximum number of drones occurs during swarming season [4][5][6][7][8]. At this time of year ratio of drone to worker brood is particularly high in swarming colonies [6]. In spring workers remove honey from drone cells in order to make space for drone brood [9]. Peak of drone production can occur at different time of year in different subspecies [10]. There is no correlation between time of production of first drones and time of swarming [11][12]. Drone brood area changes during a season in similar way to worker brood area [12]. Later during season production of drones is smaller and in queenright colonies they are not produced later than late summer [13]. At this time in temperate climate workers stop feeding drones, deny them access to honey stores and drag them from the nest (see: Drone eviction) [5]. Under starvation conditions production of drones is smaller [14] and they are removed from colony faster [15].

In natural conditions number of drones produced by one non-swarming colony during one year is about 20 thousand [4][6] (the exact estimate from one study was 22560 ± 8280 [6]). In managed colonies the number of drones produced during one season is smaller (between few hundred and few thousand) because of small number of drone cells which strongly affect production of drones [16][12]. If large amount of drone comb is provided up to 45 thousand of drones can be produced by one colony. Queenless colony can produce more than six thousand drones [6][17]. Queenless colonies with large number of patrilines produce fewer drones [18]. Colonies with bigger honey reserves produce more drones [19]. Small colonies in poor environmental conditions can produce no drones during whole season [12]. Drone comb is build only in colonies with more than 4000 workers [20]. In some studies larger colonies produce more drones [5] but see [4][6]. Number of drones produced in a colony can be determined genetically [21] and differs between subspecies [22]. Production of drones is smaller in presence of drone brood [5] or adult drones [23] but see [24].

There are suggestions that drones are costly to their colony [25], therefore, beekeepers often provide only worker foundation and remove combs with large number of drone cells. Colonies without drone combs produce more honey [26] but see [16][4][27][21]. It was not confirmed that colonies with smaller number of drone comb produce more workers [16][4][27]. Although swarming colonies tend to rear more drones [6] the presence of drones does not affect tendency to swarm [26].

Queen prefers to lay eggs in worker cells than in drone cells [28]. The proportion of eggs laid by the queen in drone cells was higher in spring than in summer [28]. Queens who were forced to lay only drone eggs subsequently laid fewer drone eggs [29]. This suggests that the queen can be involved in regulation of drone production. It was suggested that in colonies with older and unhealthy queens more drone cells are build [30], however, this needs to be verified. Drone cell construction is not regulated by queen mandibular pheromones because removal of queens mandibular gland does not affect drone cell production [31].

Theoretical models analysing possible proximate mechanisms of drone production control: [32][33].
For review of drone production see [30][34].
Other references: [35][36][37][38][39][40].

References

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