Bees collecting pollen
Honey bee swarm
Honey bee nectaring
A honey bee (or honeybee) is any bee member of the genus Apis, primarily distinguished by the production and storage of honey and the construction of perennial, colonial nests from wax.
Currently, only seven species of honey bee are recognized, with a total of 44 subspecies, though historically, from six to eleven species have been recognized. The best-known honey bee is the Western honey bee which has been domesticated for honey production and crop pollination. Honey bees represent only a small fraction of the roughly 20,000 known species of bees. Some other types of related bees produce and store honey, including the stingless honey bees, but only members of the genus Apis are true honey bees.
Two species of the honey bee, A. Mellifera and A. Cerana Indica, are often maintained, fed, and transported by beekeepers. Modern hives also enable beekeepers to transport bees, moving from field to field as the crop needs pollinating and allowing the beekeeper to charge for the pollination services they provide, revising the historical role of the self-employed beekeeper, and favouring large-scale commercial operations.
Males, or drones, are typically haploid, having only one set of chromosomes. They are produced by the queen if she chooses not to fertilize an egg; or by an unfertilized laying worker. Diploid drones may be produced if an egg is fertilized but is homozygous for the sex-determination allele. Drones take 24 days to develop and may be produced from summer through autumn. Drones have large eyes used to locate queens during mating flights. They do not have a stinger.
Workers have two sets of chromosomes. They are produced from an egg that the queen has selectively fertilized from stored sperm. Workers typically develop in 21 days. A typical colony may contain as many as 60,000 worker bees. Workers exhibit a wider range of behaviours than either queens or drones. Their duties change upon the age of the bee in the following order (beginning with cleaning out their own cell after eating through their capped brood cell): feed brood, receive nectar, clean hive, guard duty, and foraging. Some workers engage in other specialized behaviours, such as “undertaking” (removing corpses of their nestmates from inside the hive).
Workers have morphological specializations, including the pollen basket (corbicula), abdominal glands that produce beeswax, brood-feeding glands, and barbs on the sting. Under certain conditions (for example, if the colony becomes queenless), a worker may develop ovaries.
Queen honey bees are created at the decision of the worker bees by feeding a larva only royal jelly throughout its development, rather than switching from royal jelly to a mixture of honey and pollen known as bee bread once the larva passes three days of age. Queens are produced in oversized cells and develop in only 16 days; they differ in morphology and behaviour from worker bees. In addition to the greater size of the queen, she has a functional set of ovaries, and a spermatheca, which stores and maintains sperm after she has mated. Apis queens practise polyandry, with one female mating with multiple males. The highest documented mating frequency for an Apis queen is in Apis nigrocincta, where queens mate with an extremely high number of males with observed numbers of different matings ranging from 42 to 69 drones per queen. The sting of queens is not barbed like a worker’s sting, and queens lack the glands that produce beeswax. Once mated, queens may lay up to 2,000 eggs per day. They produce a variety of pheromones that regulate the behaviour of workers and helps swarms track the queen’s location during the migratory phase.
As in a few other types of eusocial bees, a colony generally contains one queen bee, a fertile female; seasonally up to a few thousand drone bees, or fertile males; and tens of thousands of sterile female worker bees. Details vary among the different species of honey bees, but common features include:
Eggs are laid singly in a cell in a wax honeycomb, produced and shaped by the worker bees. Using her spermatheca, the queen actually can choose to fertilize the egg she is laying, usually depending on into which cell she is laying. Drones develop from unfertilised eggs and are haploid, while females (queens and worker bees) develop from fertilised eggs and are diploid. Larvae are initially fed with royal jelly produced by worker bees, later switching to honey and pollen. The exception is a larva fed solely on royal jelly, which will develop into a queen bee. The larva undergoes several moultings before spinning a cocoon within the cell and pupating.
Young worker bees, sometimes called “nurse bees”, clean the hive and feed the larvae. When their royal jelly-producing glands begin to atrophy, they begin building comb cells. They progress to other within-colony tasks as they become older, such as receiving nectar and pollen from foragers and guarding the hive. Later still, a worker takes her first orientation flights and finally leaves the hive and typically spends the remainder of her life as a forager.
Worker bees cooperate to find food and use a pattern of “dancing” (known as the bee dance or waggle dance) to communicate information regarding resources with each other; this dance varies from species to species, but all living species of Apis exhibit some form of the behaviour. If the resources are very close to the hive, they may also exhibit a less specific dance commonly known as the “round dance”.
Honey bees also perform tremble dances, which recruit receiver bees to collect nectar from returning foragers.
Virgin queens go on mating flights away from their home colony to a drone congregation area and mate with multiple drones before returning. The drones die in the act of mating. Queen honey bees do not mate with drones from their home colony.
Colonies are established not by solitary queens, as in most bees, but by groups known as “swarms”, which consist of a mated queen and a large contingent of worker bees. This group moves en masse to a nest site which was scouted by worker bees beforehand and whose location is communicated with a special type of dance. Once the swarm arrives, they immediately construct a new wax comb and begin to raise new worker brood. This type of nest founding is not seen in any other living bee genus, though several groups of vespid wasps also found new nests by swarming (sometimes including multiple queens). Also, stingless bees will start new nests with large numbers of worker bees, but the nest is constructed before a queen is escorted to the site, and this worker force is not a true “swarm”.
Species of Apis are generalist floral visitors and pollinate a large variety of plants. Honey bees obtain all of their nutritional requirements from a diverse combination of pollen and nectar. Pollen is the only natural protein source for honey bees. Adult worker honey bees consume 3.4-4.3 mg of pollen per day to meet a dry matter requirement of 66-74% protein. The rearing of one larva requires 125-187.5 mg pollen or 25-37.5 mg protein for proper development.
Pollen is also a lipid source for honey bees ranging from 0.8% to 18.9%. Lipids are metabolized during the brood stage for precursors required for future biosynthesis. Fat-soluble vitamins A, D, E, and K are not considered essential but have shown to significantly improve the number of brood reared. Honey bees ingest phytosterols from pollen to produce 24-methylenecholesterol and other sterols as they cannot directly synthesize cholesterol from phytosterols. Nurse bees can selectively transfer sterols to larvae through brood food.
Nectar is collected by foraging worker bees as a source of water and carbohydrates in the form of sucrose. The dominant monosaccharides in honey bee diets are fructose and glucose but the most common circulating sugar in hemolymph is trehalose which is a disaccharide consisting of two glucose molecules. Adult worker honey bees require 4 mg of utilizable sugars per day and larvae require about 59.4 mg of carbohydrates for proper development.
Honey bees require water to maintain osmotic homeostasis, prepare liquid brood food, and to cool the hive through evaporation. A colony’s water needs can generally be met by nectar foraging as it has high water content. Occasionally on hot days or when nectar is limited, foragers will collect water from streams or ponds to meet the needs of the hive.
Nectar, a liquid high in sucrose, is produced in plant glands known as nectaries. It is an important energy resource for honey bees and plays a significant role in foraging economics and evolutionary differentiation between different subspecies.
Worker bees of a certain age secrete beeswax from a series of glands on their abdomens. They use the wax to form the walls and caps of the comb. As with honey, beeswax is gathered by humans for various purposes.
Bees collect pollen in their pollen baskets and carry it back to the hive. In the hive, pollen is used as a protein source necessary during brood-rearing. In certain environments, excess pollen can be collected from the hives of A. Mellifera and A. Cerana. It is often eaten as a health supplement. It also has been used with moderate success as a source of pollen for hand pollination. However, pollen collected by bees and harvested for pollination must be used within a few hours because it loses its potency rapidly, possibly because of the effects of enzymes or other chemicals from the bees.
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