Beekeeping
Overview
Beekeeping is the maintenance of honey bee
Honey bee
Honey bees are a subset of bees in the genus Apis, primarily distinguished by the production and storage of honey and the construction of perennial, colonial nests out of wax. Honey bees are the only extant members of the tribe Apini, all in the genus Apis...

 colonies, commonly in hives
Beehive
A beehive is a structure in which bees live and raise their young.Beehive may also refer to:Buildings and locations:* Bee Hive, Alabama, a neighborhood in Alabama* Beehive , a wing of the New Zealand Parliament Buildings...

, by humans. A beekeeper
Beekeeper
A beekeeper is a person who keeps honey bees for the purposes of securing commodities such as honey, beeswax, pollen, royal jelly; pollinating fruits and vegetables; raising queens and bees for sale to other farmers; and/or for purposes satisfying natural scientific curiosity...

 (or apiarist) keeps bees in order to collect honey
Honey
Honey is a sweet food made by bees using nectar from flowers. The variety produced by honey bees is the one most commonly referred to and is the type of honey collected by beekeepers and consumed by humans...

 and other products of the hive (including beeswax
Beeswax
Beeswax is a natural wax produced in the bee hive of honey bees of the genus Apis. It is mainly esters of fatty acids and various long chain alcohols...

, propolis
Propolis
Propolis is a resinous mixture that honey bees collect from tree buds, sap flows, or other botanical sources. It is used as a sealant for unwanted open spaces in the hive. Propolis is used for small gaps , while larger spaces are usually filled with beeswax. Its color varies depending on its...

, pollen
Pollen
Pollen is a fine to coarse powder containing the microgametophytes of seed plants, which produce the male gametes . Pollen grains have a hard coat that protects the sperm cells during the process of their movement from the stamens to the pistil of flowering plants or from the male cone to the...

, and royal jelly
Royal jelly
Royal jelly is a honey bee secretion that is used in the nutrition of larvae, as well as adult queens. It is secreted from the glands in the hypopharynx of worker bees, and fed to all larvae in the colony....

), to pollinate
Pollination
Pollination is the process by which pollen is transferred in plants, thereby enabling fertilisation and sexual reproduction. Pollen grains transport the male gametes to where the female gamete are contained within the carpel; in gymnosperms the pollen is directly applied to the ovule itself...

 crops
Agriculture
Agriculture is the cultivation of animals, plants, fungi and other life forms for food, fiber, and other products used to sustain life. Agriculture was the key implement in the rise of sedentary human civilization, whereby farming of domesticated species created food surpluses that nurtured the...

, or to produce bees for sale to other beekeepers. A location where bees are kept is called an apiary
Apiary
An apiary is a place where beehives of honey bees are kept. Traditionally beekeepers paid land rent in honey for the use of small parcels. Some farmers will provide free apiary sites, because they need pollination, and farmers who need many hives often pay for them to be moved to the crops when...

 or "bee yard".
There are more than 20,000 species of wild bees. Many species are solitary (e.g.mason bee
Mason bee
Mason bee is a common name for species of bees in the genus Osmia, of the family Megachilidae. They are named from their habit of making compartments of mud in their nests, which are made in hollow reeds or holes in wood made by wood boring insects....

s), and many others rear their young in burrows and small colonies, (e.g.
Encyclopedia
Beekeeping is the maintenance of honey bee
Honey bee
Honey bees are a subset of bees in the genus Apis, primarily distinguished by the production and storage of honey and the construction of perennial, colonial nests out of wax. Honey bees are the only extant members of the tribe Apini, all in the genus Apis...

 colonies, commonly in hives
Beehive
A beehive is a structure in which bees live and raise their young.Beehive may also refer to:Buildings and locations:* Bee Hive, Alabama, a neighborhood in Alabama* Beehive , a wing of the New Zealand Parliament Buildings...

, by humans. A beekeeper
Beekeeper
A beekeeper is a person who keeps honey bees for the purposes of securing commodities such as honey, beeswax, pollen, royal jelly; pollinating fruits and vegetables; raising queens and bees for sale to other farmers; and/or for purposes satisfying natural scientific curiosity...

 (or apiarist) keeps bees in order to collect honey
Honey
Honey is a sweet food made by bees using nectar from flowers. The variety produced by honey bees is the one most commonly referred to and is the type of honey collected by beekeepers and consumed by humans...

 and other products of the hive (including beeswax
Beeswax
Beeswax is a natural wax produced in the bee hive of honey bees of the genus Apis. It is mainly esters of fatty acids and various long chain alcohols...

, propolis
Propolis
Propolis is a resinous mixture that honey bees collect from tree buds, sap flows, or other botanical sources. It is used as a sealant for unwanted open spaces in the hive. Propolis is used for small gaps , while larger spaces are usually filled with beeswax. Its color varies depending on its...

, pollen
Pollen
Pollen is a fine to coarse powder containing the microgametophytes of seed plants, which produce the male gametes . Pollen grains have a hard coat that protects the sperm cells during the process of their movement from the stamens to the pistil of flowering plants or from the male cone to the...

, and royal jelly
Royal jelly
Royal jelly is a honey bee secretion that is used in the nutrition of larvae, as well as adult queens. It is secreted from the glands in the hypopharynx of worker bees, and fed to all larvae in the colony....

), to pollinate
Pollination
Pollination is the process by which pollen is transferred in plants, thereby enabling fertilisation and sexual reproduction. Pollen grains transport the male gametes to where the female gamete are contained within the carpel; in gymnosperms the pollen is directly applied to the ovule itself...

 crops
Agriculture
Agriculture is the cultivation of animals, plants, fungi and other life forms for food, fiber, and other products used to sustain life. Agriculture was the key implement in the rise of sedentary human civilization, whereby farming of domesticated species created food surpluses that nurtured the...

, or to produce bees for sale to other beekeepers. A location where bees are kept is called an apiary
Apiary
An apiary is a place where beehives of honey bees are kept. Traditionally beekeepers paid land rent in honey for the use of small parcels. Some farmers will provide free apiary sites, because they need pollination, and farmers who need many hives often pay for them to be moved to the crops when...

 or "bee yard".

Origins

There are more than 20,000 species of wild bees. Many species are solitary (e.g.mason bee
Mason bee
Mason bee is a common name for species of bees in the genus Osmia, of the family Megachilidae. They are named from their habit of making compartments of mud in their nests, which are made in hollow reeds or holes in wood made by wood boring insects....

s), and many others rear their young in burrows and small colonies, (e.g. bumblebee
Bumblebee
A bumble bee is any member of the bee genus Bombus, in the family Apidae. There are over 250 known species, existing primarily in the Northern Hemisphere although they are common in New Zealand and in the Australian state of Tasmania.Bumble bees are social insects that are characterised by black...

s). Beekeeping, or apiculture, is concerned with the practical management of the social species of honey bees, which live in large colonies of up to 100,000 individuals. In Europe
Europe
Europe is, by convention, one of the world's seven continents. Comprising the westernmost peninsula of Eurasia, Europe is generally 'divided' from Asia to its east by the watershed divides of the Ural and Caucasus Mountains, the Ural River, the Caspian and Black Seas, and the waterways connecting...

 and America
United States
The United States of America is a federal constitutional republic comprising fifty states and a federal district...

 the species universally managed by beekeepers is the Western honey bee (Apis mellifera). This species has several sub-species or regional varieties, such as the Italian bee
Italian bee
Apis mellifera ligustica is the Italian bee which is a sub-species of the western honey bee .- Origin :The Italian honey bee is thought to originate from the continental part of Italy, South of the Alps, and North of Sicily...

 (Apis mellifera ligustica ), European dark bee
European dark bee
The European dark bee was domesticated in modern times, and taken to North America in colonial times. These small, dark-colored honey bees are sometimes called the German black bee, although they occurred originally from Britain to eastern Central Europe.There are three main races,...

 (Apis mellifera mellifera), and the Carniolan honey bee (Apis mellifera carnica). In the tropics, other species of social bee are managed for honey production, including Apis cerana
Apis cerana
Apis cerana, or the Asiatic honey bee , is a species of honey bee found in southern and southeastern Asia, such as China, India, Japan, Malaysia, Nepal, Bangladesh and Papua New Guinea. This species is the sister species of Apis koschevnikovi, and both are in the same subgenus as the Western honey...

.

All of the Apis mellifera sub-species are capable of inter-breeding and hybridizing. Many bee breeding companies strive to selectively breed and hybridize varieties to produce desirable qualities: disease and parasite resistance, good honey production, swarming behaviour reduction, prolific breeding, and mild disposition. Some of these hybrids are marketed under specific brand names, such as the Buckfast Bee
Buckfast bee
The Buckfast bee is a honey bee a cross of many species of bees into a stable race developed by "Brother Adam", , who was in charge of beekeeping at Buckfast Abbey, where the bees are still bred today.-Origin:...

 or Midnite Bee. The advantages of the initial F1 hybrids produced by these crosses include: hybrid vigor, increased honey productivity, and greater disease resistance. The disadvantage is that in subsequent generations these advantages may fade away and hybrids tend to be very defensive and aggressive.

Wild honey harvesting

Collecting honey from wild bee colonies is one of the most ancient human activities and is still practiced by aboriginal societies in parts of Africa, Asia, Australia, and South America. Some of the earliest evidence of gathering honey from wild colonies is from rock paintings
Cave painting
Cave paintings are paintings on cave walls and ceilings, and the term is used especially for those dating to prehistoric times. The earliest European cave paintings date to the Aurignacian, some 32,000 years ago. The purpose of the paleolithic cave paintings is not known...

, dating to around 13,000 BCE
Upper Paleolithic
The Upper Paleolithic is the third and last subdivision of the Paleolithic or Old Stone Age as it is understood in Europe, Africa and Asia. Very broadly it dates to between 40,000 and 10,000 years ago, roughly coinciding with the appearance of behavioral modernity and before the advent of...

. Gathering honey from wild bee colonies is usually done by subduing the bees with smoke and breaking open the tree or rocks where the colony is located, often resulting in the physical destruction of the nest location.

Domestication of wild bees

At some point humans began to attempt to domesticate wild bees in artificial hives
Bee Hive
The term Bee Hive can refer to:*Bee hive, an alternate spelling of the word beehive*Bee Hive, Alabama, an unincorporated community*The Bee-Hive , a 19th century British newspaper*Bee Hive Records, a jazz record label...

 made from hollow logs, wooden boxes, pottery vessels, and woven straw baskets or "skeps". Honeybees were kept in Egypt
Egypt
Egypt , officially the Arab Republic of Egypt, Arabic: , is a country mainly in North Africa, with the Sinai Peninsula forming a land bridge in Southwest Asia. Egypt is thus a transcontinental country, and a major power in Africa, the Mediterranean Basin, the Middle East and the Muslim world...

 from antiquity. On the walls of the sun temple of Nyuserre Ini
Nyuserre Ini
Nyuserre Ini , was a Pharaoh of Egypt during the Fifth dynasty. He is frequently given a reign of 24 or 25 years and is dated from ca. 2445 BC to 2421 BC. His prenomen, Nyuserre, means "Possessed of Re's Power"...

 from the 5th Dynasty, before 2422 BCE, workers are depicted blowing smoke into hives as they are removing honeycombs. Inscriptions detailing the production of honey are found on the tomb of Pabasa
Pabasa
The Ancient Egyptian noble Pabasa was Chief Steward to Nitocris I, Divine Adoratrice of Amun, during the Saite Period . He is buried in tomb TT279, which is located in the El-Assasif, part of the Theban Necropolis, near Thebes....

 from the 26th Dynasty (circa 650
650s BC
-Events and trends:*Occupation begins at Maya site of Piedras Negras, Guatemala.*657 BC—Cypselus becomes the first tyrant of Corinth.*656 BC—Psamtik I extends his control over all of Egypt...

 BCE), depicting pouring honey in jars and cylindrical hives. Sealed pots of honey were found in the grave goods of Pharaohs such as Tutankhamun
Tutankhamun
Tutankhamun , Egyptian , ; approx. 1341 BC – 1323 BC) was an Egyptian pharaoh of the 18th dynasty , during the period of Egyptian history known as the New Kingdom...

.

In prehistoric Greece (Crete
Crete
Crete is the largest and most populous of the Greek islands, the fifth largest island in the Mediterranean Sea, and one of the thirteen administrative regions of Greece. It forms a significant part of the economy and cultural heritage of Greece while retaining its own local cultural traits...

 and Mycenae
Mycenae
Mycenae is an archaeological site in Greece, located about 90 km south-west of Athens, in the north-eastern Peloponnese. Argos is 11 km to the south; Corinth, 48 km to the north...

), there existed a system of high-status apiculture, as can be concluded from the finds of hives, smoking pots, honey extractors and other beekeeping paraphernalia in Knossos
Knossos
Knossos , also known as Labyrinth, or Knossos Palace, is the largest Bronze Age archaeological site on Crete and probably the ceremonial and political centre of the Minoan civilization and culture. The palace appears as a maze of workrooms, living spaces, and store rooms close to a central square...

. Beekeeping was considered a highly valued industry controlled by beekeeping overseers — owners of gold rings depicting apiculture scenes rather than religious ones as they have been reinterpreted recently, contra Sir Arthur Evans.

Archaeological finds relating to beekeeping have been discovered at Rehov
Rehov
Rehov , meaning "broad", "wide place", was an important Bronze and Iron Age city located at Tel Rehov , an archaeological site in the Jordan Valley, Israel, approximately 5 km south of Beit She'an and 3 km west of the Jordan River...

, a Bronze
Bronze Age
The Bronze Age is a period characterized by the use of copper and its alloy bronze as the chief hard materials in the manufacture of some implements and weapons. Chronologically, it stands between the Stone Age and Iron Age...

 and Iron Age
Iron Age
The Iron Age is the archaeological period generally occurring after the Bronze Age, marked by the prevalent use of iron. The early period of the age is characterized by the widespread use of iron or steel. The adoption of such material coincided with other changes in society, including differing...

 archaeological site in the Jordan Valley
Jordan Valley (Middle East)
The Jordan Valley forms part of the larger Jordan Rift Valley. It is 120 kilometers long and 15 kilometers wide, where it runs from Lake Tiberias in the north to northern Dead Sea in the south. It runs for an additional 155 kilometer south of the Dead Sea to Aqaba, an area also known as Wadi...

, Israel
Israel
The State of Israel is a parliamentary republic located in the Middle East, along the eastern shore of the Mediterranean Sea...

. Thirty intact hives, made of straw and unbaked clay, were discovered by archaeologist Amihai Mazar
Amihai Mazar
Amihai "Ami" Mazar is an Israeli archaeologist. Born in Haifa, Israel , he is currently Professor at the Institute of Archaeology of the Hebrew University of Jerusalem, holding the Eleazer Sukenik Chair in the Archaeology of Israel.Mazar has directed archaeological excavations at a number of...

 of the Hebrew University of Jerusalem
Hebrew University of Jerusalem
The Hebrew University of Jerusalem ; ; abbreviated HUJI) is Israel's second-oldest university, after the Technion – Israel Institute of Technology. The Hebrew University has three campuses in Jerusalem and one in Rehovot. The world's largest Jewish studies library is located on its Edmond J...

 in the ruins of the city, dating from about 900 BCE. The hives were found in orderly rows, three high, in a manner that could have accommodated around 100 hives, held more than 1 million bees and had a potential annual yield of 500 kilograms of honey and 70 kilograms of beeswax, according to Mazar, and are evidence that an advanced honey industry existed in ancient Israel 3,000 years ago. Ezra Marcus, an expert from the University of Haifa
University of Haifa
The University of Haifa is a university in Haifa, Israel.The University of Haifa was founded in 1963 by Haifa mayor Abba Hushi, to operate under the academic auspices of the Hebrew University of Jerusalem....

, said the finding was a glimpse of ancient beekeeping seen in texts and ancient art
Art
Art is the product or process of deliberately arranging items in a way that influences and affects one or more of the senses, emotions, and intellect....

 from the Near East
Near East
The Near East is a geographical term that covers different countries for geographers, archeologists, and historians, on the one hand, and for political scientists, economists, and journalists, on the other...

.

In ancient Greece
Ancient Greece
Ancient Greece is a civilization belonging to a period of Greek history that lasted from the Archaic period of the 8th to 6th centuries BC to the end of antiquity. Immediately following this period was the beginning of the Early Middle Ages and the Byzantine era. Included in Ancient Greece is the...

, aspects of the lives of bees and beekeeping are discussed at length by Aristotle
Aristotle
Aristotle was a Greek philosopher and polymath, a student of Plato and teacher of Alexander the Great. His writings cover many subjects, including physics, metaphysics, poetry, theater, music, logic, rhetoric, linguistics, politics, government, ethics, biology, and zoology...

. Beekeeping was also documented by the Roman
Ancient Rome
Ancient Rome was a thriving civilization that grew on the Italian Peninsula as early as the 8th century BC. Located along the Mediterranean Sea and centered on the city of Rome, it expanded to one of the largest empires in the ancient world....

 writers Virgil
Virgil
Publius Vergilius Maro, usually called Virgil or Vergil in English , was an ancient Roman poet of the Augustan period. He is known for three major works of Latin literature, the Eclogues , the Georgics, and the epic Aeneid...

, Gaius Julius Hyginus
Gaius Julius Hyginus
Gaius Julius Hyginus was a Latin author, a pupil of the famous Cornelius Alexander Polyhistor, and a freedman of Caesar Augustus. He was by Augustus elected superintendent of the Palatine library according to Suetonius' De Grammaticis, 20...

, Varro
Marcus Terentius Varro
Marcus Terentius Varro was an ancient Roman scholar and writer. He is sometimes called Varro Reatinus to distinguish him from his younger contemporary Varro Atacinus.-Biography:...

, and Columella
Columella
Lucius Junius Moderatus Columella is the most important writer on agriculture of the Roman empire. Little is known of his life. He was probably born in Gades , possibly of Roman parents. After a career in the army , he took up farming...

.

The art of beekeeping appeared in ancient China for a long time and hardly traceable to its origin. In the book "Golden Rules of Business Success" written by Fan Li
Fan Li
Fan Li was an ancient Chinese advisor in the state of Yue in the Spring and Autumn Period. He had been to the state of Wu as hostage together with King Goujian of Yue. Three years later they came back and he helped Goujian to carry on a reform. At last Yue was able to defeat the state of Wu. After...

 (or Tao Zhu Gong) during the Spring and Autumn Period there are some parts mentioning the art of beekeeping and the importance of the quality of the wooden box for bee keeping that can affect the quality of its honey.

Study of honey bees

It was not until the 18th century that European natural philosophers undertook the scientific study of bee colonies and began to understand the complex and hidden world of bee biology. Preeminent among these scientific pioneers were Swammerdam, René Antoine Ferchault de Réaumur
René Antoine Ferchault de Réaumur
René Antoine Ferchault de Réaumur was a French scientist who contributed to many different fields, especially the study of insects.-Life:Réaumur was born in a prominent La Rochelle family and educated in Paris...

, Charles Bonnet
Charles Bonnet
Charles Bonnet , Swiss naturalist and philosophical writer, was born at Geneva, of a French family driven into Switzerland by the religious persecution in the 16th century.-Life and work:Bonnet's life was uneventful...

, and the blind Swiss scientist Francois Huber
François Huber
François Huber was a Swiss naturalist.He was born at Geneva, of a family which had already made its mark in the literary and scientific world: his great-aunt, Marie Huber, was known as a voluminous writer on religious and theological subjects, and as the translator and epitomizer of The Spectator...

. Swammerdam and Réaumur were among the first to use a microscope and dissection to understand the internal biology of honey bees. Réaumur was among the first to construct a glass walled observation hive to better observe activities within hives. He observed queens laying eggs in open cells, but still had no idea of how a queen was fertilized; nobody had ever witnessed the mating of a queen and drone and many theories held that queens were "self-fertile," while others believed that a vapor or "miasma" emanating from the drones fertilized queens without direct physical contact. Huber was the first to prove by observation and experiment that queens are physically inseminated by drones outside the confines of hives, usually a great distance away.

Following Réaumur's design, Huber built improved glass-walled observation hives and sectional hives which could be opened, like the leaves of a book, to inspect individual wax combs; this greatly improved the direct observation of activity within a hive. Although he became blind before he was twenty, Huber employed a secretary, Francois Burnens, to make daily observations, conduct careful experiments, and to keep accurate notes over a period of more than twenty years. Huber confirmed that a hive consists of one queen who is the mother of all the female workers and male drones in the colony. He was also the first to confirm that mating with drones takes place outside of hives and that queens are inseminated by a number of successive matings with male drones, high in the air at a great distance from their hive. Together, he and Burnens dissected bees under the microscope
Microscope
A microscope is an instrument used to see objects that are too small for the naked eye. The science of investigating small objects using such an instrument is called microscopy...

 and were among the first to describe the ovaries
Ovary
The ovary is an ovum-producing reproductive organ, often found in pairs as part of the vertebrate female reproductive system. Ovaries in anatomically female individuals are analogous to testes in anatomically male individuals, in that they are both gonads and endocrine glands.-Human anatomy:Ovaries...

 and spermatheca
Spermatheca
The spermatheca , also called receptaculum seminis , is an organ of the female reproductive tract in insects, some molluscs, oligochaeta worms and certain other invertebrates and vertebrates...

, or sperm store, of queens as well as the penis
Penis
The penis is a biological feature of male animals including both vertebrates and invertebrates...

 of male drones. Huber is universally regarded as "the father of modern bee-science" and his "Nouvelles Observations sur Les Abeilles (or "New Observations on Bees") http://books.google.com/books?id=cQwAAAAAQAAJ&printsec=titlepage&dq=Nouvelles+Observations+sur+Les+Abeilles#PPP5,M1) revealed all the basic scientific truths for the biology and ecology of honeybees.

Invention of the movable comb hive

Early forms of honey collecting entailed the destruction of the entire colony when the honey was harvested. The wild hive was crudely broken into, using smoke to suppress the bees, the honeycomb
Honeycomb
A honeycomb is a mass of hexagonal waxcells built by honey bees in their nests to contain their larvae and stores of honey and pollen.Beekeepers may remove the entire honeycomb to harvest honey...

s were torn out and smashed up — along with the eggs, larvae and honey they contained. The liquid honey from the destroyed brood nest was crudely strained through a sieve or basket. This was destructive and unhygienic, but for hunter-gatherer
Hunter-gatherer
A hunter-gatherer or forage society is one in which most or all food is obtained from wild plants and animals, in contrast to agricultural societies which rely mainly on domesticated species. Hunting and gathering was the ancestral subsistence mode of Homo, and all modern humans were...

 societies this did not matter, since the honey was generally consumed immediately and there were always more wild colonies to exploit. But in settled societies the destruction of the bee colony meant the loss of a valuable resource; this drawback made beekeeping both inefficient and something of a "stop and start" activity. There could be no continuity of production and no possibility of selective breeding, since each bee colony was destroyed at harvest time, along with its precious queen. During the medieval period abbey
Abbey
An abbey is a Catholic monastery or convent, under the authority of an Abbot or an Abbess, who serves as the spiritual father or mother of the community.The term can also refer to an establishment which has long ceased to function as an abbey,...

s and monasteries
Monastery
Monastery denotes the building, or complex of buildings, that houses a room reserved for prayer as well as the domestic quarters and workplace of monastics, whether monks or nuns, and whether living in community or alone .Monasteries may vary greatly in size – a small dwelling accommodating only...

 were centers of beekeeping, since beeswax was highly prized for candles and fermented honey was used to make alcoholic mead
Mead
Mead , also called honey wine, is an alcoholic beverage that is produced by fermenting a solution of honey and water. It may also be produced by fermenting a solution of water and honey with grain mash, which is strained immediately after fermentation...

 in areas of Europe where vines would not grow.

The 18th and 19th centuries saw successive stages of a revolution in beekeeping, which allowed the bees themselves to be preserved when taking the harvest.

Intermediate stages in the transition from the old beekeeping to the new were recorded for example by Thomas Wildman in 1768/1770, who described advances over the destructive old skep-based beekeeping so that the bees no longer had to be killed to harvest the honey. Wildman for example fixed a parallel array of wooden bars across the top of a straw hive or skep (with a separate straw top to be fixed on later) "so that there are in all seven bars of deal" [in a 10 inches (254 mm) hive] "to which the bees fix their combs". He also described using such hives in a multi-storey configuration, foreshadowing the modern use of supers: he described adding (at a proper time) successive straw hives below, and eventually removing the ones above when free of brood and filled with honey, so that the bees could be separately preserved at the harvest for a following season. Wildman also described a further development, using hives with "sliding frames" for the bees to build their comb, foreshadowing more modern uses of movable-comb hives. Wildman's book acknowledged the advances in knowledge of bees previously made by Swammerdam, Maraldi, and de Réaumur — he included a lengthy translation of Réaumur's account of the natural history of bees — and he also described the initiatives of others in designing hives for the preservation of bee-life when taking the harvest, citing in particular reports from Brittany dating from the 1750s, due to Comte de la Bourdonnaye.

The 19th Century saw this revolution in beekeeping practice completed through the perfection of the movable comb hive by Lorenzo Lorraine Langstroth, a descendant of Yorkshire farmers who emigrated to the United States. Langstroth was the first person to make practical use of Huber's earlier discovery that there was a specific spatial measurement between the wax combs, later called "the bee space", which bees would not block with wax, but kept as a free passage. Having determined this "bee space" (between 5 and 8 mm, or 1/4 to 3/8"), Langstroth then designed a series of wooden frames within a rectangular hive box, carefully maintaining the correct space between successive frames, and found that the bees would build parallel honeycombs in the box without bonding them to each other or to the hive walls. This enables the beekeeper to slide any frame out of the hive for inspection, without harming the bees or the comb, protecting the eggs, larvae and pupae contained within the cells. It also meant that combs containing honey could be gently removed and the honey extracted without destroying the comb. The emptied honey combs could then be returned to the bees intact for refilling. Langstroth's classic book, The Hive and Honey-bee, published in 1853, described his rediscovery of the bee space and the development of his patent movable comb hive.

The invention and development of the movable-comb-hive fostered the growth of commercial honey production on a large scale in both Europe and the USA (see also Beekeeping in the United States
Beekeeping in the United States
Beekeeping in the United States dates back to the 1860s.-Development of beekeeping in the United States:John Harbison, originally from Pennsylvania, successfully brought bee keeping to the US west coast in the 1860s, in an area now known as Harbison Canyon, California, and greatly expanded the...

).

Evolution of hive designs

Langstroth's design for movable comb hives was seized upon by apiarists and inventors on both sides of the Atlantic
Atlantic Ocean
The Atlantic Ocean is the second-largest of the world's oceanic divisions. With a total area of about , it covers approximately 20% of the Earth's surface and about 26% of its water surface area...

 and a wide range of moveable comb hives were designed and perfected in England
England
England is a country that is part of the United Kingdom. It shares land borders with Scotland to the north and Wales to the west; the Irish Sea is to the north west, the Celtic Sea to the south west, with the North Sea to the east and the English Channel to the south separating it from continental...

, France
France
The French Republic , The French Republic , The French Republic , (commonly known as France , is a unitary semi-presidential republic in Western Europe with several overseas territories and islands located on other continents and in the Indian, Pacific, and Atlantic oceans. Metropolitan France...

, Germany
Germany
Germany , officially the Federal Republic of Germany , is a federal parliamentary republic in Europe. The country consists of 16 states while the capital and largest city is Berlin. Germany covers an area of 357,021 km2 and has a largely temperate seasonal climate...

 and the United States
United States
The United States of America is a federal constitutional republic comprising fifty states and a federal district...

. Classic designs evolved in each country: Dadant hives and Langstroth hive
Langstroth hive
The Langstroth bee hive, patented in October 1852, is the standard beehive used in many parts of the world for beekeeping. The advantage of this hive is that the bees build honeycomb into frames, which can be moved with ease. The frames are designed to prevent bees from attaching honeycombs where...

s are still dominant in the USA; in France the De-Layens
Georges de Layens
Georges de Layens was a French botanist and apiculturalist. He was the creator of a popular mobile beehive called the "Layens hive".He published numerous works on beekeeping and botany...

 trough-hive became popular and in the UK a British National Hive became standard as late as the 1930s although in Scotland the smaller Smith hive is still popular. In some Scandinavian countries and in Russia the traditional trough hive persisted until late in the 20th Century and is still kept in some areas. However, the Langstroth and Dadant designs remain ubiquitous in the USA and also in many parts of Europe, though Sweden
Sweden
Sweden , officially the Kingdom of Sweden , is a Nordic country on the Scandinavian Peninsula in Northern Europe. Sweden borders with Norway and Finland and is connected to Denmark by a bridge-tunnel across the Öresund....

, Denmark
Denmark
Denmark is a Scandinavian country in Northern Europe. The countries of Denmark and Greenland, as well as the Faroe Islands, constitute the Kingdom of Denmark . It is the southernmost of the Nordic countries, southwest of Sweden and south of Norway, and bordered to the south by Germany. Denmark...

, Germany, France and Italy
Italy
Italy , officially the Italian Republic languages]] under the European Charter for Regional or Minority Languages. In each of these, Italy's official name is as follows:;;;;;;;;), is a unitary parliamentary republic in South-Central Europe. To the north it borders France, Switzerland, Austria and...

 all have their own national hive designs. Regional variations of hive evolved to reflect the climate, floral productivity and the reproductive characteristics of the various subspecies of native honey bee in each bio-region.

The differences in hive dimensions are insignificant in comparison to the common factors in all these hives: they are all square or rectangular; they all use movable wooden frames; they all consist of a floor, brood-box, honey-super, crown-board and roof. Hives have traditionally been constructed of cedar, pine, or cypress
Cypress
Cypress is the name applied to many plants in the cypress family Cupressaceae, which is a conifer of northern temperate regions. Most cypress species are trees, while a few are shrubs...

 wood, but in recent years hives made from injection molded dense polystyrene
Polystyrene
Polystyrene ) also known as Thermocole, abbreviated following ISO Standard PS, is an aromatic polymer made from the monomer styrene, a liquid hydrocarbon that is manufactured from petroleum by the chemical industry...

 have become increasingly important.

Hives also use queen excluders between the brood-box and honey supers to keep the queen from laying eggs in cells next to those containing honey intended for consumption. Also, with the advent in the 20th century of mite pests, hive floors are often replaced for part of (or the whole) year with a wire mesh and removable tray.

Pioneers of practical and commercial beekeeping

The 19th Century produced an explosion of innovators and inventors who perfected the design and production of beehives, systems of management and husbandry, stock improvement by selective breeding
Selective breeding
Selective breeding is the process of breeding plants and animals for particular genetic traits. Typically, strains that are selectively bred are domesticated, and the breeding is sometimes done by a professional breeder. Bred animals are known as breeds, while bred plants are known as varieties,...

, honey extraction
Honey extraction
Honey extraction is the central process in beekeeping of removing honey from honeycomb so that it is isolated in a pure liquid form.Normally, the honey is stored by the bees on a very regular honeycomb they build on a frame. The frames are typically harvested late summer, when they will be most...

 and marketing. Preeminent among these innovators were:

Jan Dzierżon
Jan Dzierzon
Johann Dzierzon, in Polish Jan Dzierżon or Dzierżoń , also John Dzierzon , was a pioneering apiarist who discovered the phenomenon of parthenogenesis in bees and designed the first successful movable-frame beehive.Dzierzon came from a Polish family in Silesia...

, was the father of modern apiology and apiculture. All modern beehives are descendants of his design.

L. L. Langstroth
L. L. Langstroth
Rev. Lorenzo Lorraine Langstroth , apiarist, clergyman and teacher, is considered the "Father of American Beekeeping."L. L. Langstroth was born in Philadelphia, Pennsylvania...

, revered as the "father of American apiculture", no other individual has influenced modern beekeeping practice more than Lorenzo Lorraine Langstroth. His classic book The Hive and Honey-bee was published in 1853.

Moses Quinby
Moses Quinby
Moses Quinby was one of the first commercial beekeepers in the United States, a native of St. Johnsville, New York.Quinby established his business in his early 20s and expanded it to own about 1,200 hives in the Mohawk Valley, New York....

, often termed 'the father of commercial beekeeping in the United States', author of Mysteries of Bee-Keeping Explained.

Amos Root
Amos Root
Amos Ives Root developed innovative beekeeping techniques in the United States during the mid-19th century, a period when the industry played an important role in the economy of many communities. He founded his own company, which continues in business to the present day...

, author of the A B C of Bee Culture which has been continuously revised and remains in print to this day. Root pioneered the manufacture of hives and the distribution of bee-packages in the United States.

A.J. Cook, author of The Bee-Keepers' Guide; or Manual of the Apiary, 1876.

Dr. C.C. Miller was one of the first entrepreneurs to actually make a living from apiculture. By 1878 he made beekeeping his sole business activity. His book, Fifty Years Among the Bees, remains a classic and his influence on bee management persists to this day.

Major Francesco De Hruschka was an Italian military officer who made one crucial invention that catalyzed the commercial honey industry. In 1865 he invented a simple machine for extracting honey from the comb by means of centrifugal force. His original idea was simply to support combs in a metal framework and then spin them around within a container to collect honey as it was thrown out by centrifugal force. This meant that honeycombs could be returned to a hive undamaged but empty, saving the bees a vast amount of work, time, and materials. This single invention greatly improved the efficiency of honey harvesting and catalysed the modern honey industry.

Walter T. Kelley
Walter T. Kelley
Walter T. Kelley was beekeeper who created a large bee equipment supply and queen breeding company based in Clarkson, Kentucky. Kelley also wrote extensively about apiculture and published the bee journal Modern Beekeeping.-Early life:...

was an American pioneer of modern beekeeping in the early and mid 20th century. He greatly improved upon beekeeping equipment and clothing and went on to manufacture these items as well as other equipment. His company sold via catalog worldwide and his book, How to Keep Bees & Sell Honey, an introductory book of apiculture and marketing, allowed for a boom in beekeeping following World War II
World War II
World War II, or the Second World War , was a global conflict lasting from 1939 to 1945, involving most of the world's nations—including all of the great powers—eventually forming two opposing military alliances: the Allies and the Axis...

.

In the U.K. practical beekeeping was led in the early 20th century by a few men, pre-eminently Brother Adam and his Buckfast bee
Buckfast bee
The Buckfast bee is a honey bee a cross of many species of bees into a stable race developed by "Brother Adam", , who was in charge of beekeeping at Buckfast Abbey, where the bees are still bred today.-Origin:...

 and R.O.B. Manley
R.O.B. Manley
Robert "Bert" Orlando Beater Manley was a British Beekeeper, an authority on commercial honey farming and developer of the popular Manley moveable frame hives and frame systems.-Achievements:...

, author of many titles, including 'Honey Production In The British Isles' and inventor of the Manley frame, still universally popular in the U.K. Other notable British pioneers include William Herrod-Hempsall and Gale.

In India, N. K. Garg is known as the pioneer of starting apiculture. He is the one who helped people in taking apiculture commercially. He started it in early 1990's.

Traditional beekeeping

Fixed comb hives

A fixed comb hive is a hive in which the combs cannot be removed or manipulated for management or harvesting without permanently damaging the comb. Almost any hollow structure can be used for this purpose, such as a log gum, skep or a clay pot. Fixed comb hives are no longer in common use in most places, and are illegal in some places that require inspection for problems such as varroa
Varroa
Varroa is a genus of parasitic mites associated with honey bees, placed in its own family, Varroidae. The genus was named for Marcus Terentius Varro, a Roman scholar who was also a beekeeper.-History and behavior:...

 and American foulbrood
American foulbrood
American foulbrood , caused by the spore- forming Paenibacillus larvae ssp. larvae , is the most widespread and destructive of the bee brood diseases. Paenibacillus larvae is a rod-shaped bacterium, which is visible only under a high power microscope...

.

Movable frame hives

In the United States
United States
The United States of America is a federal constitutional republic comprising fifty states and a federal district...

, the Langstroth hive
Langstroth hive
The Langstroth bee hive, patented in October 1852, is the standard beehive used in many parts of the world for beekeeping. The advantage of this hive is that the bees build honeycomb into frames, which can be moved with ease. The frames are designed to prevent bees from attaching honeycombs where...

 is commonly used. The Langstroth was the first successful top-opened hive with movable frames, and other designs of hive have been based on it. Langstroth hive was however a descendant of Jan Dzierzon’s Polish hive designs. In the United Kingdom, the most common type of hive is the British National Hive, which can hold Hoffman, British Standard or popular Manley frames, but it is not unusual to see some other sorts of hive (Smith, Commercial and WBC, rarely Langstroth). Straw skeps, bee gums, and unframed box hives are now unlawful in most US states, as the comb and brood cannot be inspected for diseases. However, straw skeps are still used for collecting swarms by hobbyists in the UK, before moving them into standard hives.

Top-bar hives

A growing number of amateur beekeepers are adopting various top-bar hive
Top-bar hive
Top-bar hives are a style of beehive used for beekeeping. They are especially useful in areas where resources are limited, but are also increasingly popular among hobby beekeepers in industrialized nations...

s similar to the type commonly found in Africa. Top bar hives were originally used as traditional beekeeping a method in both Greece and Vietnam. These have no frames and the honey-filled comb is not returned to the hive after extraction, as it is in the Langstroth hive. Because of this, the production of honey is likely to be somewhat less than that of a Langstroth hive. Top bar hives are mostly kept by people who are more interested in having bees in their garden than in honey production per se.

Some of the most well known top-bar hives are the Kenyan Top Bar Hive (KTBH) with sloping sides, the Tanzanian Top Bar Hive, which has straight sides and the Vertical Top Bar Hives such as the Warre or "People's Hive" designed by Abbe Warre in the mid 1900's.

The initial costs and equipment requirements are far lower. Often scrap wood or #2 or #3 pine is able to be used with a nice hive as the outcome. Top-bar hives also offer some advantages in interacting with the bees and the amount of weight that must be lifted is greatly reduced. Top-bar Hives are being widely used in developing countries in Africa and Asia as a result of the 'Bees For Development' program. There is a growing number of beekeepers in the U.S. using various top-bar hives within the past 2 years or so.

Protective clothing

While knowledge of the bees is the first line of defense, most beekeepers also wear some protective clothing. Novice beekeepers usually wear gloves and a hooded suit or hat and veil. Experienced beekeepers sometimes elect not to use gloves because they inhibit delicate manipulations. The face and neck are the most important areas to protect, so most beekeepers will at least wear a veil.

Defensive bees are attracted to the breath, and a sting on the face can lead to much more pain and swelling than a sting elsewhere, while a sting on a bare hand can usually be quickly removed by fingernail scrape to reduce the amount of venom injected.

The protective clothing is generally light colored (but not colorful) and of a smooth material. This provides the maximum differentiation from the colony's natural predators (bears, skunks, etc.) which tend to be dark-colored and furry.

The 'stings' retained in the fabric of the clothing will continue to pump out an alarm pheromone that actually attracts aggressive action and further stinging attacks. Washing suits regularly, and rinsing gloved hands in vinegar will minimize attraction.

Smoker

Smoke is the beekeeper's third line of defense. Most beekeepers use a "smoker" — a device designed to generate smoke from the incomplete combustion of various fuels. Smoke calms bees; it initiates a feeding response in anticipation of possible hive abandonment due to fire. Smoke also masks alarm pheromones released by guard bees or when bees are squashed in an inspection. The ensuing confusion creates an opportunity for the beekeeper to open the hive and work without triggering a defensive reaction. In addition, when a bee consumes honey the bee's abdomen distends, supposedly making it difficult to make the necessary flexes to sting, though this has not been tested scientifically.

Smoke is of questionable use with a swarm, because swarms do not have honey stores to feed on in response. Usually smoke is not needed, since swarms tend to be less defensive, as they have no stores to defend, and a fresh swarm will have fed well from the hive.

Many types of fuel can be used in a smoker as long as it is natural and not contaminated with harmful substances. These fuels include hessian, twine
Twine
Twine is a light string or strong thread composed of two or more smaller strands or yarns twisted together. More generally, the term can be applied to any thin cord....

, burlap
Burlap
Hessian , or burlap in the US, is a woven fabric usually made from skin of the jute plant or sisal fibres, or may be combined with other vegetable fibres to make rope, nets, and similar products...

, pine needles, corrugated cardboard, and mostly rotten or punky wood. Indian beekeepers, especially in Kerala, often use coconut fibers as they are readily available, safe, and of negligible expense. Some beekeeping supply sources also sell commercial fuels like pulped paper and compressed cotton, or even aerosol cans of smoke. Other beekeepers use sumac
Sumac
Sumac is any one of approximately 250 species of flowering plants in the genus Rhus and related genera, in the family Anacardiaceae. Sumacs grow in subtropical and temperate regions throughout the world, especially in Africa and North America....

 as fuel because it ejects lots of smoke and doesn't have an odor.

Some beekeepers are using "liquid smoke" as a safer, more convenient, alternative. It is a water-based solution that is sprayed onto the bees from a plastic spray bottle.

Torpor may also be induced by the introduction of chilled air into the hive - while chilled carbon dioxide may have harmful long-term effects.

Effects of stings and of protective measures

Some beekeepers believe that the more stings a beekeeper receives, the less irritation each causes, and they consider it important for safety of the beekeeper to be stung a few times a season. Beekeepers have high levels of antibodies (mainly IgG
Immunoglobulin G
Immunoglobulin G are antibody molecules. Each IgG is composed of four peptide chains — two heavy chains γ and two light chains. Each IgG has two antigen binding sites. Other immunoglobulins may be described in terms of polymers with the IgG structure considered the monomer.IgG constitutes 75%...

) reacting to the major antigen of bee venom, phospholipase A2 (PLA)
Phospholipase
A phospholipase is an enzyme that hydrolyzes phospholipids into fatty acids and other lipophilic substances. There are four major classes, termed A, B, C and D, distinguished by the type of reaction which they catalyze:*Phospholipase A...

. Antibodies correlate with the frequency of bee stings.

The entry of venom into the body from bee-stings may also be hindered and reduced by protective clothing which allows the wearer to remove stings and venom sacs simply with a tug on the clothing.

Natural beekeeping

There is a current movement that eschews chemicals in beekeeping and believes that health issues in bees can most effectively be addressed by reversing trends that disrespect the needs of the bees themselves. Crop spraying, unnatural conditions in which bees are moved thousands of miles to pollinate commercial crops, frequent opening of the hive for inspection, artificial insemination of queens, routine medication and sugar water feeding are all thought to contribute to a general weakening of the constitution of the honey bee.

Practitioners of 'natural beekeeping' tend to use variations of the top-bar hive, which is a simple design that retains the concept of movable comb without the use of frames or foundation. The horizontal top-bar hive, as championed by Marty Hardison, Michael Bush, Philip Chandler, Dennis Murrell and others, can be seen as a modernization of hollow log hives, with the addition of wooden bars of specific width from which bees hang their combs. Its widespread adoption in recent years can be attributed to the publication in 2007 of The Barefoot Beekeeper by Philip Chandler, which challenged many aspects of modern beekeeping and offered the horizontal top-bar hive as a viable alternative to the ubiquitous Langstroth-syle movable-frame hive.

The most popular vertical top-bar hive is probably the Warré hive, based on a design by the French priest Abbé Émile Warré (1867-1951) and popularized by Dr. David Heaf in his English translation of Warré's book L'Apiculture por Tous as Beekeeping For All.

Natural beekeeping is characterized by a willingness to hand most of the control to the bees themselves, and to minimize interference in their lives. Practitioners expect to take honey only when the bees' needs have first been taken care of, and the feeding of sugar is discouraged except as an emergency measure.

Urban or backyard beekeeping

Related to natural beekeeping, urban beekeeping is an attempt to revert to a less industrialized way of obtaining honey by utilizing small-scale colonies that pollinate urban gardens. Urban apiculture has undergone a renaissance in the 2000s. Paris, Berlin, London, Tokyo and Washington, D.C., are among beekeeping cities. Until 2010, beekeeping was banned in New York City and punishable with a $2,000 fine. Urban beekeeping is commonly practiced in areas that have a pesticide
Pesticide
Pesticides are substances or mixture of substances intended for preventing, destroying, repelling or mitigating any pest.A pesticide may be a chemical unicycle, biological agent , antimicrobial, disinfectant or device used against any pest...

 ban. This includes Paris, as well as 156 municipalities in Canada and 3 of 10 Canadian provinces
Pesticides in Canada
The use of pesticides in Canada is regulated by Health Canada via the Pest Control Products Act to provide effective management of pest infestations when other methods of pest control are not succeeding for the health of the community...

. Beekeeping was illegal in Vancouver until 2003, for example, but by 2010 there were bees on the roof of Vancouver City Hall.

Castes

A colony of bees consists of three castes of bee:
  • a queen bee
    Queen bee
    The term queen bee is typically used to refer to an adult, mated female that lives in a honey bee colony or hive; she is usually the mother of most, if not all, the bees in the hive. The queens are developed from larvae selected by worker bees and specially fed in order to become sexually mature...

    , which is normally the only breeding female in the colony;
  • a large number of female worker bee
    Worker bee
    A Worker bee is any female eusocial bee that lacks the full reproductive capacity of the colony's queen bee; under most circumstances, this is correlated to an increase in certain non-reproductive activities relative to a queen, as well...

    s, typically 30,000–50,000 in number;
  • a number of male drones
    Drone (bee)
    Drones are male honey bees. They develop from eggs that have not been fertilized, and they cannot sting, since the worker bee's stinger is a modified ovipositor .-Etymology:...

    , ranging from thousands in a strong hive in spring to very few during dearth or cold season.


The queen is the only sexually mature female in the hive and all of the female worker bees and male drones are her offspring. The queen may live for up to three years or more and may be capable of laying half a million eggs or more in her lifetime. At the peak of the breeding season, late spring to summer, a good queen may be capable of laying 3,000 eggs in one day, more than her own body weight. This would be exceptional however; a prolific queen might peak at 2,000 eggs a day, but a more average queen might lay just 1,500 eggs per day. The queen is raised from a normal worker egg, but is fed a larger amount of royal jelly
Royal jelly
Royal jelly is a honey bee secretion that is used in the nutrition of larvae, as well as adult queens. It is secreted from the glands in the hypopharynx of worker bees, and fed to all larvae in the colony....

 than a normal worker bee, resulting in a radically different growth and metamorphosis. The queen influences the colony by the production and dissemination of a variety of pheromone
Pheromone
A pheromone is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting outside the body of the secreting individual to impact the behavior of the receiving individual...

s or "queen substances". One of these chemicals suppresses the development of ovaries in all the female worker bees in the hive and prevents them from laying eggs.

Mating of queens

The queen emerges from her cell after 15 days of development and she remains in the hive for 3–7 days before venturing out on a mating flight. Mating flight is otherwise known as 'nuptial flight'. Her first orientation flight may only last a few seconds, just enough to mark the position of the hive. Subsequent mating flights may last from 5 minutes to 30 minutes, and she may mate with a number of male drones on each flight. Over several matings, possibly a dozen or more, the queen will receive and store enough sperm
Sperm
The term sperm is derived from the Greek word sperma and refers to the male reproductive cells. In the types of sexual reproduction known as anisogamy and oogamy, there is a marked difference in the size of the gametes with the smaller one being termed the "male" or sperm cell...

 from a succession of drones to fertilize hundreds of thousands of eggs. If she does not manage to leave the hive to mate — possibly due to bad weather or being trapped within part of the hive — she will remain infertile and become a 'drone layer', incapable of producing female worker bees. Worker bees will sometimes kill a non-performing queen, and produce another. Without a properly performing queen, the hive is doomed.

Mating takes place at some distance from the hive and often several hundred feet up in the air; it is thought that this separates the strongest drones from the weaker ones - ensuring that only the fastest and strongest drones get to pass on their genes.

Female worker bees

Almost all the bees in a hive are female worker bees. At the height of summer when activity in the hive is frantic and work goes on non-stop, the life of a worker bee may be as short as 6 weeks; in late autumn, when no brood is being raised and no nectar is being harvested, a young bee may live for 16 weeks, right through the winter. During its life a worker bee performs different work functions in the hive which are largely dictated by the age of the bee.
Period Work activity
Days 1-3 Cleaning cells and incubation
Day 3-6 Feeding older larvae
Day 6-10 Feeding younger larvae
Day 8-16 Receiving honey and pollen from field bees
Day 12-18 Wax making and cell building
Day 14 onwards Entrance guards; nectar and pollen foraging

Male bees (drones)

Drones are the largest bees in the hive (except for the queen), at almost twice the size of a worker bee. They do no work, do not forage for pollen or nectar and have no other known function than to mate with new queens and fertilize them on their mating flights. A bee colony will generally start to raise drones a few weeks before building queen cells in order to supersede a failing queen or in preparation for swarming. When queen raising for the season is over, the bees in colder climates will drive the drones out of the hive to die, biting and tearing their legs and wings.

Differing stages of development

Stage of development Queen Worker Drone
Egg 3 days 3 days 3 days
Larva 8 days 10 days 13 days
Pupa 4 days 8 days 8 days
Total 15 days 21 days 24 days

Structure of a bee colony

A domesticated bee colony is normally housed in a rectangular hive body, within which eight to ten parallel frames house the vertical plates of honeycomb which contain the eggs, larvae, pupae and food for the colony. If one were to cut a vertical cross-section through the hive from side to side, the brood nest would appear as a roughly ovoid ball spanning 5-8 frames of comb. The two outside combs at each side of the hive tend to be exclusively used for long-term storage of honey and pollen.

Within the central brood nest, a single frame of comb will typically have a central disk of eggs, larvae and sealed brood cells which may extend almost to the edges of the frame. Immediately above the brood patch an arch of pollen
Pollen
Pollen is a fine to coarse powder containing the microgametophytes of seed plants, which produce the male gametes . Pollen grains have a hard coat that protects the sperm cells during the process of their movement from the stamens to the pistil of flowering plants or from the male cone to the...

-filled cells extends from side to side, and above that again a broader arch of honey-filled cells extends to the frame tops. The pollen is protein-rich food for developing larvae, while honey is also food but largely energy rich rather than protein rich. The nurse bees which care for the developing brood secrete a special food called 'royal jelly' after feeding themselves on honey and pollen. The amount of royal jelly which is fed to a larva determines whether it will develop into a worker bee or a queen.

Apart from the honey stored within the central brood frames, the bees store surplus honey in combs above the brood nest. In modern hives the beekeeper places separate boxes, called 'supers', above the brood box, in which a series of shallower combs is provided for storage of honey. This enables the beekeeper to remove some of the supers in the late summer, and to extract the surplus honey harvest, without damaging the colony of bees and its brood nest below. If all the honey is 'stolen', including the amount of honey needed to survive winter, the beekeeper must replace these stores by feeding the bees sugar or corn syrup in autumn.

Annual cycle of a bee colony

The development of a bee colony follows an annual cycle of growth which begins in spring with a rapid expansion of the brood nest, as soon as pollen is available for feeding larvae. Some production of brood may begin as early as January, even in a cold winter, but breeding accelerates towards a peak in May (in the northern hemisphere), producing an abundance of harvesting bees synchronized to the main 'nectar flow' in that region. Each race of bees times this build-up slightly differently, depending on how the flora of its original region blooms. Some regions of Europe have two nectar flows: one in late spring and another in late August. Other regions have only a single nectar flow. The skill of the beekeeper lies in predicting when the nectar flow will occur in his area and in trying to ensure that his colonies achieve a maximum population of harvesters at exactly the right time.
The key factor in this is the prevention or skillful management of the swarming impulse. If a colony swarms unexpectedly and the beekeeper does not manage to capture the resulting swarm, he is likely to harvest significantly less honey from that hive, since he will have lost half his worker bees at a single stroke. If, however, he can use the swarming impulse to breed a new queen but keep all the bees in the colony together, he will maximize his chances of a good harvest. It takes many years of learning and experience to be able to manage all these aspects successfully, though owing to variable circumstances many beginners will often achieve a good honey harvest.

Colony reproduction: swarming and supersedure

All colonies are totally dependent on their queen, who is the only egg-layer. However, even the best queens live only a few years and one or two years longevity is the norm. She can choose whether or not to fertilize an egg as she lays it; if she does so, it develops into a female worker bee; if she lays an unfertilized egg it becomes a male drone. She decides which type of egg to lay depending on the size of the open brood cell which she encounters on the comb; in a small worker cell she lays a fertilized egg; if she finds a much larger drone cell she lays an unfertilized drone egg.

All the time that the queen is fertile and laying eggs she produces a variety of pheromones which control the behavior of the bees in the hive; these are commonly called 'queen substance' but in reality there are various pheromones with different functions. As the queen ages she begins to run out of stored sperm and her pheromones begin to fail. At some point, inevitably, the queen begins to falter and the bees will decide to replace her by creating a new queen from one of her worker eggs. They may do this because she has been damaged (lost a leg or an antenna), because she has run out of sperm and cannot lay fertilized eggs (has become a 'drone laying queen') or because her pheromones have dwindled to a point where they cannot control all the bees in the hive anymore.

At this juncture the bees will produce one or more queen cells by modifying existing worker cells which contain a normal female egg. However, there are two distinct behaviors which the bees pursue:
  1. Supersedure: queen replacement within one hive without swarming
  2. Swarm cell production: the division of the hive into two colonies by swarming


Different sub-species of Apis mellifera exhibit differing swarming characteristics which reflect their evolution
Evolution
Evolution is any change across successive generations in the heritable characteristics of biological populations. Evolutionary processes give rise to diversity at every level of biological organisation, including species, individual organisms and molecules such as DNA and proteins.Life on Earth...

 in different ecotope
Ecotope
Ecotopes are the smallest ecologically-distinct landscape features in a landscape mapping and classification system. As such, they represent relatively homogeneous, spatially-explicit landscape functional units that are useful for stratifying landscapes into ecologically distinct features for the...

s of the European continent. In general the more northerly black races are said to swarm less and supersede more, whereas the more southerly yellow and grey varieties are said to swarm more frequently. The truth is complicated because of the prevalence of cross-breeding and hybridization of the sub species and opinions differ.

Supersedure is highly valued as a behavioral trait by beekeepers because a hive that supersedes its old queen does not swarm and so no stock is lost; it merely creates a new queen and allows the old one to fade away, or alternatively she is killed when the new queen emerges. When superseding a queen the bees will produce just one or two queen cells, characteristically in the center of the face of a broodcomb.

In swarming, by contrast, a great many queen cells are created — typically a dozen or more — and these are located around the edges of a broodcomb, most often at the sides and the bottom.

Once either process has begun, the old queen will normally leave the hive with the hatching of the first queen cells. When she leaves the hive the old queen is accompanied by a large number of bees, predominantly young bees (wax-secreters), who will form the basis of the new hive. Scouts are sent out from the swarm to find suitable hollow trees or rock crevices and as soon as one is found the entire swarm moves in, building new wax brood combs within a matter of hours using the honey stores which the young bees have filled themselves with before leaving the old hive. Only young bees can secrete wax from special abdominal segments and this is why there tend to be more young bees than old in swarms. Often a number of virgin queens accompany the first swarm (the 'prime swarm'), and the old queen is replaced as soon as a daughter queen is mated and laying. Otherwise, she will be quickly superseded in their new home.

Factors that trigger swarming

It is generally accepted that a colony of bees will not swarm until they have completed all of their brood combs, i.e. filled all available space with eggs, larvae and brood. This generally occurs in late Spring at a time when the other areas of the hive are rapidly filling with honey stores. So one key trigger of the swarming instinct is when the queen has no more room to lay eggs and the hive population is becoming very congested. Under these conditions a prime swarm may issue with the queen, resulting in a halving of the population within the hive, leaving the old colony with a large number of hatching bees. The queen who leaves finds herself in a new hive with no eggs and no larvae but lots of energetic young bees who create a new set of brood combs from scratch in a very short time.

Another important factor in swarming is the age of the queen. Those under a year in age are unlikely to swarm unless they are extremely crowded, while older queens have swarming predisposition.

Beekeepers monitor their colonies carefully in spring and watch for the appearance of queen cells, which are a dramatic signal that the colony is determined to swarm.

When a colony has decided to swarm, queen cells are produced in numbers varying to a dozen or more. When the first of these queen cells is sealed, after eight days of larval feeding, a virgin queen will pupate and be due to emerge seven days after sealing. Before leaving, the worker bees fill their stomachs with honey in preparation for the creation of new honeycombs in a new home. This cargo of honey also makes swarming bees less inclined to sting, and a newly issued swarm is noticeably gentle for up to 24 hours — often capable of being handled without gloves or veil by a beekeeper.

This swarm is looking for shelter. A beekeeper may capture it and introduce it into a new hive, helping to meet this need. Otherwise, it will return to a feral
Feral
A feral organism is one that has changed from being domesticated to being wild or untamed. In the case of plants it is a movement from cultivated to uncultivated or controlled to volunteer. The introduction of feral animals or plants to their non-native regions, like any introduced species, may...

 state, in which case it will find shelter in a hollow tree, an excavation, an abandoned chimney, or even behind shutters.

Back at the original hive, the first virgin queen to emerge from her cell will immediately seek to kill all her rival queens who are still waiting to emerge from their cells. However, usually the bees deliberately prevent her from doing this, in which case, she too will lead a second swarm from the hive. Successive swarms are called 'after-swarms' or 'casts' and can be very small, often with just a thousand or so bees, as opposed to a prime swarm which may contain as many as ten to twenty thousand bees.

Small after-swarms have less chance of survival and, depleting the original hive, may threaten its survival as well. When a hive has swarmed despite the beekeeper's preventative efforts, a good management practice is to give the depleted hive a couple frames of open brood with eggs. This helps replenish the hive more quickly, and gives a second opportunity to raise a queen, if there is a mating failure.

Each race or sub-species of honey bee has its own swarming characteristics. Italian bees are very prolific and inclined to swarm; Northern European black bees have a strong tendency to supersede their old queen, without swarming. These differences are the result of differing evolutionary pressures in the regions where each sub-species evolved.

Artificial swarming

When a colony accidentally loses its queen, it is said to be 'queenless'. The workers realize that the queen is absent after as little as an hour, as her pheromones fade in the hive. The colony cannot survive without a fertile queen laying eggs to renew the population. So the workers select cells containing eggs aged less than three days and enlarge these cells dramatically to form 'emergency queen cells'. These appear similar to large peanut-like structure about an inch long, which hangs from the center or side of the brood combs. The developing larva in a queen cell is fed differently from an ordinary worker-bee, receiving in addition to the normal honey and pollen a great deal of royal jelly, a special food secreted by young 'nurse bees' from the hypopharyngeal gland. This special food dramatically alters the growth and development of the larva so that, after metamorphosis and pupation, it emerges from the cell as a queen bee. The queen is the only bee in a colony which has fully developed ovaries and she secretes a pheromone which suppresses the normal development of ovaries in all her workers.

Beekeepers use the ability of the bees to produce new queens in order to increase their colonies, a procedure called splitting a colony. In order to do this, they remove several brood combs from a healthy hive, taking care that the old queen is left behind. These combs must contain eggs or larvae less than three days old which will be covered by young 'nurse bees' which care for the brood and keep it warm. These brood combs and attendant nurse bees are then placed into a small 'nucleus hive' along with other combs containing honey and pollen. As soon as the nurse bees find themselves in this new hive and realize that they have no queen they set about constructing emergency queen cells using the eggs or larvae which they have in the combs with them.

World apiculture

World honey production and consumption in 2005
Country Production
(1000 metric tons)
Consumption
(1000 metric tons)
Number
of beekeepers
Number
of bee hives
Europe and Russia
Ukraine
Ukraine
Ukraine is a country in Eastern Europe. It has an area of 603,628 km², making it the second largest contiguous country on the European continent, after Russia...

71.46 52
Russia
Russia
Russia or , officially known as both Russia and the Russian Federation , is a country in northern Eurasia. It is a federal semi-presidential republic, comprising 83 federal subjects...

52.13 54
Spain
Spain
Spain , officially the Kingdom of Spain languages]] under the European Charter for Regional or Minority Languages. In each of these, Spain's official name is as follows:;;;;;;), is a country and member state of the European Union located in southwestern Europe on the Iberian Peninsula...

37.00 40
Germany
Germany
Germany , officially the Federal Republic of Germany , is a federal parliamentary republic in Europe. The country consists of 16 states while the capital and largest city is Berlin. Germany covers an area of 357,021 km2 and has a largely temperate seasonal climate...

 (*2008)
21.23 89 90,000* 1,000,000*
Hungary
Hungary
Hungary , officially the Republic of Hungary , is a landlocked country in Central Europe. It is situated in the Carpathian Basin and is bordered by Slovakia to the north, Ukraine and Romania to the east, Serbia and Croatia to the south, Slovenia to the southwest and Austria to the west. The...

19.71 4
Romania
Romania
Romania is a country located at the crossroads of Central and Southeastern Europe, on the Lower Danube, within and outside the Carpathian arch, bordering on the Black Sea...

19.20 10
Greece
Greece
Greece , officially the Hellenic Republic , and historically Hellas or the Republic of Greece in English, is a country in southeastern Europe....

16.27 16
France
France
The French Republic , The French Republic , The French Republic , (commonly known as France , is a unitary semi-presidential republic in Western Europe with several overseas territories and islands located on other continents and in the Indian, Pacific, and Atlantic oceans. Metropolitan France...

15.45 30
Bulgaria
Bulgaria
Bulgaria , officially the Republic of Bulgaria , is a parliamentary democracy within a unitary constitutional republic in Southeast Europe. The country borders Romania to the north, Serbia and Macedonia to the west, Greece and Turkey to the south, as well as the Black Sea to the east...

11.22 2
Serbia
Serbia
Serbia , officially the Republic of Serbia , is a landlocked country located at the crossroads of Central and Southeast Europe, covering the southern part of the Carpathian basin and the central part of the Balkans...

3 to 5 6.3 30,000 430,000
Denmark
Denmark
Denmark is a Scandinavian country in Northern Europe. The countries of Denmark and Greenland, as well as the Faroe Islands, constitute the Kingdom of Denmark . It is the southernmost of the Nordic countries, southwest of Sweden and south of Norway, and bordered to the south by Germany. Denmark...

 (*1996)
2.5 5 *4,000 *150,000
North America
United States of America
United States
The United States of America is a federal constitutional republic comprising fifty states and a federal district...

 (*2006, **2002)
70.306* 158.75* 12,029** (210,000 bee keepers) 2,400,000*
Canada
Canada
Canada is a North American country consisting of ten provinces and three territories. Located in the northern part of the continent, it extends from the Atlantic Ocean in the east to the Pacific Ocean in the west, and northward into the Arctic Ocean...

45 (2006); 28 (2007) 29 13,000 500,000
Latin America
Argentina
Argentina
Argentina , officially the Argentine Republic , is the second largest country in South America by land area, after Brazil. It is constituted as a federation of 23 provinces and an autonomous city, Buenos Aires...

93.42 (Average 84) 3
Mexico
Mexico
The United Mexican States , commonly known as Mexico , is a federal constitutional republic in North America. It is bordered on the north by the United States; on the south and west by the Pacific Ocean; on the southeast by Guatemala, Belize, and the Caribbean Sea; and on the east by the Gulf of...

50.63 31
Brazil
Brazil
Brazil , officially the Federative Republic of Brazil , is the largest country in South America. It is the world's fifth largest country, both by geographical area and by population with over 192 million people...

33.75 2
Uruguay
Uruguay
Uruguay ,officially the Oriental Republic of Uruguay,sometimes the Eastern Republic of Uruguay; ) is a country in the southeastern part of South America. It is home to some 3.5 million people, of whom 1.8 million live in the capital Montevideo and its metropolitan area...

11.87 1
Oceania
Australia
Australia
Australia , officially the Commonwealth of Australia, is a country in the Southern Hemisphere comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands in the Indian and Pacific Oceans. It is the world's sixth-largest country by total area...

18.46 16
New Zealand
New Zealand
New Zealand is an island country in the south-western Pacific Ocean comprising two main landmasses and numerous smaller islands. The country is situated some east of Australia across the Tasman Sea, and roughly south of the Pacific island nations of New Caledonia, Fiji, and Tonga...

9.69 8 2602 313,399
Asia
China
China
Chinese civilization may refer to:* China for more general discussion of the country.* Chinese culture* Greater China, the transnational community of ethnic Chinese.* History of China* Sinosphere, the area historically affected by Chinese culture...

299.33 (average 245) 238 7,200,000
Turkey
Turkey
Turkey , known officially as the Republic of Turkey , is a Eurasian country located in Western Asia and in East Thrace in Southeastern Europe...

82.34 (average 70) 66 4,500,000
Iran
Iran
Iran , officially the Islamic Republic of Iran , is a country in Southern and Western Asia. The name "Iran" has been in use natively since the Sassanian era and came into use internationally in 1935, before which the country was known to the Western world as Persia...

3,500,000
India
India
India , officially the Republic of India , is a country in South Asia. It is the seventh-largest country by geographical area, the second-most populous country with over 1.2 billion people, and the most populous democracy in the world...

52.23 45 9,800,000
South Korea
South Korea
The Republic of Korea , , is a sovereign state in East Asia, located on the southern portion of the Korean Peninsula. It is neighbored by the People's Republic of China to the west, Japan to the east, North Korea to the north, and the East China Sea and Republic of China to the south...

23.82 27
Vietnam
Vietnam
Vietnam – sometimes spelled Viet Nam , officially the Socialist Republic of Vietnam – is the easternmost country on the Indochina Peninsula in Southeast Asia. It is bordered by China to the north, Laos to the northwest, Cambodia to the southwest, and the South China Sea –...

13.59 0
Turkmenistan
Turkmenistan
Turkmenistan , formerly also known as Turkmenia is one of the Turkic states in Central Asia. Until 1991, it was a constituent republic of the Soviet Union, the Turkmen Soviet Socialist Republic . Turkmenistan is one of the six independent Turkic states...

10.46 10
Africa
Ethiopia
Ethiopia
Ethiopia , officially known as the Federal Democratic Republic of Ethiopia, is a country located in the Horn of Africa. It is the second-most populous nation in Africa, with over 82 million inhabitants, and the tenth-largest by area, occupying 1,100,000 km2...

41.23 40 4,400,000
Tanzania
Tanzania
The United Republic of Tanzania is a country in East Africa bordered by Kenya and Uganda to the north, Rwanda, Burundi, and the Democratic Republic of the Congo to the west, and Zambia, Malawi, and Mozambique to the south. The country's eastern borders lie on the Indian Ocean.Tanzania is a state...

28.68 28
Angola
Angola
Angola, officially the Republic of Angola , is a country in south-central Africa bordered by Namibia on the south, the Democratic Republic of the Congo on the north, and Zambia on the east; its west coast is on the Atlantic Ocean with Luanda as its capital city...

23.77 23
Kenya
Kenya
Kenya , officially known as the Republic of Kenya, is a country in East Africa that lies on the equator, with the Indian Ocean to its south-east...

22.00 21
Egypt
Egypt
Egypt , officially the Arab Republic of Egypt, Arabic: , is a country mainly in North Africa, with the Sinai Peninsula forming a land bridge in Southwest Asia. Egypt is thus a transcontinental country, and a major power in Africa, the Mediterranean Basin, the Middle East and the Muslim world...

 (*1997)
16* 200,000* 2,000,000*
Central African Republic
Central African Republic
The Central African Republic , is a landlocked country in Central Africa. It borders Chad in the north, Sudan in the north east, South Sudan in the east, the Democratic Republic of the Congo and the Republic of the Congo in the south, and Cameroon in the west. The CAR covers a land area of about ,...

14.23 14
Morocco
Morocco
Morocco , officially the Kingdom of Morocco , is a country located in North Africa. It has a population of more than 32 million and an area of 710,850 km², and also primarily administers the disputed region of the Western Sahara...

 (*1997)
4.5* 27,000* 400,000*
Source: Food and Agriculture Organization of the United Nations (FAO), August 2007.


Sources:
  • Denmark: beekeeping.com (1996)
  • Arab countries: beekeeping.com (1997)
  • USA: University of Arkansas National Agricultural Law Center (2002), Agricultural Marketing Resource Center (2006)
  • Serbia

See also

  • Agriculture
    Agriculture
    Agriculture is the cultivation of animals, plants, fungi and other life forms for food, fiber, and other products used to sustain life. Agriculture was the key implement in the rise of sedentary human civilization, whereby farming of domesticated species created food surpluses that nurtured the...

  • Biosecurity
    Biosecurity
    Biosecurity is a set of preventive measures designed to reduce the risk of transmission of infectious diseases, quarantined pests, invasive alien species, living modified organisms...

  • Western honey bee life cycle
  • Beekeeping in New Zealand
    Beekeeping in New Zealand
    Beekeeping in New Zealand started as a home craft in the 1850s, not long after initial European settlement and is now an established industry as well as being a hobby activity.-Industry:...

  • Beekeeping in the United Kingdom
    Beekeeping in the United Kingdom
    Beekeeping in the United KingdomIn the winter of 2008, about 20% of the UK's bees died. The losses were highest in the north of England and lowest in the east...

  • Beekeeping in the United States
    Beekeeping in the United States
    Beekeeping in the United States dates back to the 1860s.-Development of beekeeping in the United States:John Harbison, originally from Pennsylvania, successfully brought bee keeping to the US west coast in the 1860s, in an area now known as Harbison Canyon, California, and greatly expanded the...


External links

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