Читать книгу Honey Bee Medicine for the Veterinary Practitioner - Группа авторов - Страница 84

The honey bee queen largely regulates colony activities, producing a variety of pheromones, known collectively as the queen signal, to support many hive functions such as cleaning, brood rearing, comb‐building, guarding, and foraging (Free 1987; Slessor et al. 2005; Bortolotti and Costa 2014). Such pheromones also influence drone mating, swarm clustering and the queen retinue behavior, while also suppressing worker bee ovary function and egg laying, queen supersedure, and limiting the potential for queen rearing (Figure 4.4). When the queen is removed from a colony or dies, her absence is discovered with little delay and the colony swiftly initiates queen rearing because of the loss of queen pheromones (Free 1987; Slessor et al. 2005; Bortolotti and Costa 2014). After an extended time without a queen, the colony's worker bees stop completing their hive tasks, begin laying unfertilized eggs that mature into drone bees, and the colony declines toward collapse.

Queen mandibular pheromone, previously believed only secreted from the mandibular gland, has been most studied and is comprised of many different chemical compounds affecting many of the aforementioned hive behaviors (Free 1987; Slessor et al. 2005; Bortolotti and Costa 2014). Queen mandibular pheromone is now recognized as queen retinue pheromone because of its multiglandular origin (Slessor et al. 2005). This multicomponent queen pheromone attracts a continually shifting cohort of 6–10 young worker bees that feed and groom the queen (Free 1987). These young bees form the queen's retinue who, in addition to caring for the queen, collect and distribute her pheromones about the colony via antennal contacts and trophallaxis (Slessor et al. 2005; Jarriault and Mercer 2012). Queen retinue pheromone has many stimulatory properties beyond shaping the queen's court or retinue, including formation and steadiness of the swarm cluster, attracting drone bees and mating, influencing worker tasks and development, comb building, brood rearing, as well as foraging behavior. In addition, this queen pheromone is vital for the regulation of colony reproduction and worker physiology, inducing worker sterility when the structure of the hive – having a productive, fertile queen – favors the long‐term genetic interests of the worker bees (Slessor et al. 2005; Princen et al. 2019).

Figure 4.4 A colony of honey bees survives in large part because of the presence of a healthy queen. The queen honey bee produces several pheromones, known collectively as the queen signal, helping to regulate colony activities. Such pheromones support many important worker bee hive functions including cleaning, comb building, brood rearing, foraging behavior, and hive defense. Further, the queen signal is necessary for drone bee attraction and mating, maintaining the swarm cluster and queen retinue behaviors, as well as inhibiting worker bee reproduction and egg laying, queen supersedure, and queen rearing.

Source: © Lauren D. Sawchyn, DVM, CMI. Chapter: Physiology of the honey bee, authored by Rolfe M. Radcliffe and illustrated by Lauren D. Sawchyn.

Significant queen signal redundancy has been identified throughout the pheromone communication repertoire of the honey bee colony (Slessor et al. 2005; Princen et al. 2019). Besides the queen mandibular gland, other sources of the queen signal have been isolated from the tergal, labial, tarsal, Dufour's, and Koschevnikov glands (Slessor et al. 2005; Bortolotti and Costa 2014). Located beneath the abdominal tergites and more developed in the queen, the tergal glands secrete pheromones that support the function of the queen retinue pheromone. The tarsal glands, present in all three bee castes, release a footprint pheromone that in the queen inhibits queen cell construction by the worker bees (Bortolotti and Costa 2014). The Dufour's gland, closely associated with the sting apparatus in the female honey bee, is another part of the queen signal and also provides worker bees with information about queen fertility and reproductive potential (Dor et al. 2005; Bortolotti and Costa 2014). The Koschevnikov gland, located near the sting shaft of females, is yet another source of queen signal, and with gland degeneration beginning at one year of age contributes to the failing of aging queens (Bortolotti and Costa 2014).