Читать книгу Honey Bee Medicine for the Veterinary Practitioner - Группа авторов - Страница 85

Worker honey bees also make use of a variety of pheromones, and most glands in both the queen and worker honey bees are fully developed yet secrete different pheromones (Free 1987; Ritter 2014; Bortolotti and Costa 2014). Further, gland maturity in worker bees follows a temporal pattern that parallels the changing tasks and activities of the worker bee within the colony. Worker bees employ several pheromones during their lifetime including alarm, Nasonov, footprint, and forager signals to help in colony defense, for orientation and marking sites, and for identifying the colony (Bortolotti and Costa 2014; Ritter 2014). Worker mandibular pheromone is also produced when needed in queenless colonies to suppress ovarian development in other workers when egg‐laying workers or psuedoqueens emerge (Bortolotti and Costa 2014).

The defense behavior of a honey bee colony is well known and alarm pheromones released from worker bees help guide this response (Bortolotti and Costa 2014; Free 1987; Breed et al. 2004). Defensive behavior tasks occur prior to foraging in worker bees and two types of behaviors are recognized: guarding and defending (Breed et al. 2004). Guard bees watch and monitor the hive entrance, inspect all who wish to enter, and reject non‐nestmates; soldier bees fly out in the face of danger, and chase, bite, or sting intruders. Honey bees are able to recognize nestmates of the same colony or non‐nestmates of other colonies through chemical signals present in the lipid covering of the insect cuticle. Both the sting apparatus, including the Koschevnikov gland, as well as the mandibular gland produce pheromones that evoke defensive behavior through the recruitment and amplification of other worker bees (Bortolotti and Costa 2014).

Worker honey bees produce a pheromone from the Nasonov gland that is restricted only to this caste (Bortolotti and Costa 2014; Free 1987). The gland secretion is composed of several volatile compounds that function to attract other bees, and is released during three main contexts: during swarm clustering, marking their hive entrance, and marking of water and foraging sources (Bortolotti and Costa 2014; Free 1987). The worker footprint pheromone, secreted from the tarsal glands of worker honey bees, may share the role of marking the hive entrance and food sources. The Nasonov pheromone may also be used for recruiting other workers to help develop queen cups during the process of queen rearing.

A honey bee colony has a social organization characterized by a division of labor among the worker honey bees – worker polyethism – that changes over time (Winston 1987; Seeley 1995). During their early life (0–20 days) they work inside the colony cleaning cells and caring for the brood, receiving nectar and handling pollen, comb building, and tending the queen among other tasks; as they age (20–45 days) they begin working outside the nest ventilating the hive and guarding its entrance, as well as commencing foraging flights (Winston 1987). The control of worker polyethism has been a mystery, yet recent evidence suggests that a pheromone, produced by older foraging sister worker bees, delays the age of onset of foraging and other age‐related tasks (Leoncini et al. 2004; Slessor et al. 2005).