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Apiculture, or beekeeping, is the semidomestication of honey bees (genus Apis, primarily A. mellifera) in human-made and managed structures (beehives) for the purposes of harvesting honey and other hive products, providing pollination services in natural and agroecosystems, raising and selling bees to others, and providing other benefits. The practice dates back millennia and has arisen multiple times throughout human history in many different cultures and with several different species. However, given the proclivity of temperate subspecies of A. mellifera to store honey, apiculture has largely adopted this species worldwide (including areas outside of its native range, most notably the New World, Australia, and East Asia) and in many cases on an industrial scale. Because they are so readily managed and highly amenable to manipulation through apicultural practices, honey bees have also become one of the main model systems for the scientific study of social insects and sociobiology.
Honey Hunting and Wild-Nest Tending
Other examples demonstrate that the activity of honey harvesting could be quite elaborate, with multiple individuals working in tandem, suggesting that the practice was commonplace and originated even earlier than is known from present archeological evidence. Because honey was the only natural sweetener and source of simple sugars (outside of ripened fruits) to early humans, some argue that honey harvesting was a communal activity (as seen in Fig. 1b), celebrated and even ritualized in religious rites. This connection between honey bees, the highly prized honey, and spirituality is a common theme woven throughout human culture and history. The practice of honey robbing occurs to this day in many parts of the globe, following many of the same practices as depicted in Paleolithic rock art (Fig. 1c).
Honey robbing of known “bee trees” became quite sophisticated in the European Middle Ages, resulting in laws to determine the ownership of particular wild nests, whether it was vested in the land owner or the one who harvested the honey. Trees containing wild colonies were often branded and revisited by honey robbers, some even cutting sections of the trunk so that the combs could be more readily removed repeatedly with minimal disruption to the colony . Locating the wild nests was accomplished using bee traps and beelining, where foraging bees could be captured in small handheld containers made of wood or animal horns. When released, the bees would fly in a “beeline” toward their nest, which could then be followed and triangulated to more easily locate their colony.
Locating wild nests can also be assisted by other animals, most notably the greater honey guide Indicator indicator of sub-Saharan Africa. Members of this Old World family (Indicatoridae) of nest-parasitic birds have coevolved with several mammalian species, including humans (but not honey badgers, despite widespread belief), to lead them to previously located honey bee nest through various songs. The birds then eat the remains of the destroyed nest once the human hunters have opened it. Honey bees were likely co-opted into agriculture by early Neolithic farmers, at least in Europe, as evidenced by the extensive and continuous use of beeswax .
Evolution of the Beehive
The prehistoric connection of humans and honey bees was an opportunistic one – locating wild nests in their native range and exploiting their resources. The practice of apiculture did not emerge until early peoples began to provide physical structures to living colonies and actively to manage them. Evidence from across the globe has shown this practice independently arising numerous times and in several different bee species, including the keeping of stingless bees, a practice known as meliponiculture.
Other hive designs were moveable-comb hives, since the beekeeper did not have to destroy the combs in order to manipulate them. Top-bar hives, for example, date back to ancient Greece, where beekeepers would place wooden slats or sticks across the top of a cavity from which the bees would build their combs without connecting them to the inner hive walls. The beekeepers could then move the sticks and hence the combs without destroying them. The combs were still relatively fragile, however, since they were attached only along the top. The first moveable-frame hives arose in the seventeenth century, with combs attached on all four sides, thereby providing additional structural support to avoid breaking combs.
The historical roots of modern apiculture date back to the latter half of the nineteenth century and specifically to four main innovations that arose independently but greatly complemented each other. In doing so, the standard beehive and apicultural practices that are used today derive almost directly from them. First, the incorporation of the bee space hive design was a critical breakthrough that was independently derived by Jan Dzierzon in Poland and Lorenzo L. Langstroth in the USA. The bee space is the distance between two adjacent combs (slightly less than 1 cm in A. mellifera) that optimizes the nest structure by maximizing the number of combs that can be built within a given cavity volume but also enables the two inner surfaces to be utilized by the bees for food storage and brood rearing. Adherence to bee space within a wood beehive greatly decreased the construction of connecting comb to the hive body and facilitated the beekeepers’ ability to manipulate frames without damage to the combs (Fig. 2d). The standardization of frame and hive dimensions also permitted interchangeable hive equipment within and among colonies, which facilitated the advance of larger-scale beekeeping and industrialization.
Second, wax foundation helped to facilitate comb construction by the bees by providing them with a template and starting wax secured within the wooden frames. John Mehring first generated such foundation in 1857 by pressing soft or molten beeswax between two engraved wooden planks, but later metal dowels embossed with a hexagonal pattern were used to roll sheets of wax. Eventually, J. F. Hetherington added vertical wires within the foundation that, together with horizontal wires strung through the wooden frames, provided significant structural support for the wax combs.
Third, the bellowed smoker365体育网站 was invented by Moses Quinby in 1870. The use of smoke to pacify honey bees was known from the dawn of beekeeping and honey robbing, but it was difficult to maintain a reliable source of smoke. By fashioning bellows to the back of a cylindrical metal canister, beekeepers can stoke the fire of combustible materials from below and direct the smoke from a nozzle on the top.
Finally, the radial honey extractor was invented in 1865 by Franz Endler von Hruschka. It consists of a large metal cylinder with a pivoting rank that vertically holds several honey frames. Prior to loading the frames, beekeepers use a hot metal knife to cut off the wax cappings from the honeycombs, enabling the honey to drip out. Because it is so viscous, the frame rack is then spun at relatively high speed (either hand-cranked or motorized for larger units) to allow centrifugal force to cause the honey to sling out of the combs onto the inner walls of the extractor, drain down, and collect on the bottom from where the beekeeper can more easily strain and bottle it. Moreover, because of the moveable frames and wire-embedded foundation, the combs remain intact and can be reused by the beekeeper for the bees to store more honey.
Beekeeping Population and Demographics
365体育网站Apiculture has tended to follow the greater trends in agriculture. For much of the modern agrarian era, small family farms would typically include a few beehives to provide honey, with crop pollination benefits as an incidental additional benefit. As agriculture became more consolidated, monocultural, and large-scale, apiculture followed suit and is now a major service industry to the farming systems.
Apiculture consolidation has resulted in inverse demographics of beekeepers and the population of managed beehives. One way to categorize beekeeping operations is by their number of hives, where those with fewer than 50 hives are considered backyard hobbyists, those with 50–500 are sideliners, and those with more than 500 are considered commercial . This metric is fairly specific to the USA, however, and thus may be more applicable to characterize beekeepers into part-time beekeepers (those who do not make their living on the practice) and full-time beekeepers (those whose sole or main living is from apiculture) [1365体育网站]. Irrespective of how to categorize beekeepers, the vast majority (~90%) of the beekeeping community tends to be part-time (hobbyist) beekeepers that collectively own a small fraction (~10%) of the managed beehives, while about 10%) of beekeepers are full-time (professional) beekeepers that own most (~90%) of the managed hives.
Honey is certainly the best known product of apiculture and the original reason for its development. The most common form of honey is liquid, or extracted honey (Fig. 3c), which is fully removed from the honeycomb, strained, and bottled. However, other variants can be found as cut-comb honey (sections of honeycomb complete with wax cappings), whipped or creamed honey (liquid honey that is heat-treated and purposefully seeded with crystals to cause the sugars to come out of solution), and chunk honey (a combination of a cut-comb section within extracted honey).
Beeswax is another widely known product from managed beehives. It may well have been the most valuable product in Europe during the Middle Ages, when candles were in very high demand as the main or sole source of household lighting. Brace or burr comb – wax connecting combs that bees built but are disfavored by beekeepers – is often scraped during hive inspections and stored by beekeepers for later use. Wax cappings following honey extraction is also another major source of beeswax. Because of its relatively low melting temperature (65 °C), one popular means of melting wax is solar heaters. These boxes are built with an inclined, metal floor and glass ceiling, which uses the greenhouse effect to passively melt the wax inside so that it can drain down and collect in a pan placed on the bottom. While candles have long been a popular application of beeswax, the largest contemporary use of beeswax is the cosmetics industry, where it is used as a foundation for balms and creams because of its favorable texture and stability.
Pollen can also be harvested from managed colonies. Unlike honey, pollen stored in the wax combs (= beebread) cannot be easily extracted. Instead, beekeepers affix special traps at the hive entrance to dislodge and capture pollen loads from the returning foragers. Most involve a narrow mesh or screen through which foragers must pass in entering the hive; in doing so they lose the collected pollen from the corbiculae on their hind legs. The dislodged pellets fall down into a collecting tray (Fig. 3c365体育网站). Raw pollen is often used as a health supplement, although its nutritional value for human consumption is not well demonstrated. In fact, it is not even certain that humans can digest pollen.
Propolis, or bee glue, has been used in human medicine for millennia in several different cultures, as it has been shown to exhibit antimicrobial properties. Beekeepers can harvest it from their hives by placing a propolis trap on the hive, typically underneath the inner cover. The narrow slits in the trap are too small for bees to pass and thus provide a super-stimulus for them to fill with propolis. A small subset of foragers usually performs this task by flying from the hive to collect tree and other plant resins, return to the hive with their loads on their corbiculae, mix it with wax, then place it into the cracks and spaces provided to create a water-tight seal. Beekeepers then remove the trap, freeze it to make the propolis brittle, and twist the trap to break the hardened propolis from the plastic.
Royal jelly365体育网站 is a glandular secretion from nurse bees that is fed to developing queen larvae as a key factor in caste differentiation. It too has been used for centuries in folk medicine, particularly in East Asia. Beekeepers harvest it by grafting young worker larvae into queen cups that are placed into populous queenless colonies, where the nurse bees provision mass quantities of royal jelly into each cell. Several days later, the larvae are removed and the royal jelly harvested. However, royal jelly consists of little more than water, amino acids, and a few minerals, so that it is quite harmless when consumed by humans but probably of no nutritional value.
Other hive products include venom, which is mass-extracted by having workers sting a rubber bladder so that the venom is deposited on a glass plate, dried, and collected after crystallization. Venom of honey bees and other stinging insects is used in immunotherapy and other treatments of sting allergies. Queens are also produced by beekeepers to sell to other beekeepers for replacements or making new colonies (Fig. 3c), and there are entire sub-industries in many countries for the breeding and large-scale production of queens of different genetic stocks. Live adult workers can also be raised or sold for their use in apitherapy365体育网站, the use of stings to relieve pain and other ailments.
While honey bees are by far the most extensively used managed pollinators in commercial production agriculture, they are but one member of the entire pollinator community upon which these industries rely (Fig. 3d) . Most of the approximately 20,000 species of bees worldwide are solitary, and most are more efficient at pollinating than honey bees on an individual bee basis. However, honey bees are well suited for commercial pollination because they are generalist pollinators that visit a wide variety of flowering crops, live in large colonies (30,000–50,000 individuals) that forage up to about 3 km from the hive, are active year-round in warm climates, and can be transported and relocated in and out of areas as pollination requirements demand.
365体育网站Growers of various bee-dependent crops often enter into contractual agreements with beekeepers to rent beehives during the blooming period. In doing so, beekeepers usually move their hives at night when the foragers are at home inside the hive. When placed in their new location, scout bees locate new floral resources the following morning when foraging resumes and in doing so often lock into the more proximate of those available (usually the target crop). Beekeepers and growers tend to follow the “10% rule” in deciding when to move the hives; fewer blooms run the risk of the newly translocated bees of overlooking the target crop, but waiting for more blooms would waste them because the bees would not be there to pollinate. Because honey bees exhibit flower constancy, they lock into a favorable resource once they find it and keep collecting that one source until it is no longer in bloom.
The optimal density and location of beehives varies widely depending on crop, populations of local wild pollinators, floral density, and quality of the nectar/pollen . Honey bees favor resources that are higher in quality (sugar concentration for nectar), lower in handling time (more easily collected from each blossom), higher in quantity, and closer to the hive. Crops that have high densities of blossoms usually require hives to be placed in and around the crop, especially fields with large acreages, so that all blossoms across the field have a chance to be pollinated rather than just the ones near the apiary. The default density for most cropping systems is 2.5 hives per hectare, but this can range from one-fifth to five times that value, depending on pollination demands.
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