Honey: Difference between revisions

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Honey use and production has a long and varied history, with its beginnings in [[Prehistory|prehistoric times]]. Several cave paintings in [[Cuevas de la Araña]] in [[Spain]] depict humans foraging for honey at least 8,000 years ago. While [[Western honey bee|''Apis mellifera'']] is an [[Old World]] [[insect]], large-scale meliponiculture of [[New World]] stingless bees has been practiced by [[Maya civilization|Mayans]] since [[Pre-Columbian era|pre-Columbian]] times.

== Formation ==

[[File:Bee on -calyx 935.jpg|thumb|A [[honey bee]] with its proboscis extended into a [[Calyx (botany)|calyx]] of [[goldenrod]]]]

[[File:Honey comb.jpg|thumb|[[Honeycomb]] displaying [[Hexagonal prismatic honeycomb|hexagonal prismatic]] [[Beeswax|wax]] cells in which [[honey bee]]s store honey]]

=== By honey bees ===

Honey is produced by bees who have collected [[nectar]] or [[honeydew (secretion)|honeydew]]. Bees value honey for its sugars, which they consume to support general [[metabolism|metabolic activity]], especially that of their flight muscles during [[forage (honey bee)|foraging]], and as a food for their [[larva]]e. To this end bees stockpile honey to provide for themselves during ordinary foraging as well as during lean periods, as in [[overwintering]]. During foraging bees use part of the nectar they collect to power their flight muscles. The majority of nectar collected is not used to directly nourish the insects but is instead destined for [[regurgitation (digestion)|regurgitation]], [[Enzyme catalysis|enzymatic digestion]], and finally long-term [[Food storage|storage]] as honey. During cold weather or when other food sources are scarce, adult and [[larva]]l bees consume stored honey, which is many times more energy-dense as the nectar from which it is made.

After leaving the hive, a foraging bee collects sugar-rich nectar or honeydew. Nectar from the flower generally has a water content of 70 to 80% and is much less viscous than finished honey, which usually has a water content around 18%. The water content of honeydew from aphids and other [[Insect|true bugs]] is generally very close to the sap on which those insects feed and is usually somewhat more dilute than nectar. One source describes the water content of honeydew as around 89%. Whether it is feeding on nectar or honeydew, the bee sucks these runny fluids through its [[proboscis]], which delivers the liquid to the bee's honey stomach or "honey crop". This cavity lies just above its food stomach, the latter of which digests pollen and sugars consumed by an individual honey bee for its own nourishment.

In ''[[Western honey bee|Apis mellifera]]'', the honey stomach holds about 40 mg of liquid. This is about half the weight of an unladen bee. Collecting this quantity in nectar can require visits to more than a thousand flowers. When nectar is plentiful, it can take a bee more than an hour of ceaseless work to collect enough nectar to fill its honey crop. [[Saliva]]ry enzymes and proteins from the bee's [[Insect mouthparts#Hypopharynx|hypopharyngeal gland]] are secreted into the nectar once it is in the bee's honey stomach. These [[Digestive enzyme|substances]] begin [[Hydrolysis|cleaving]] [[Sugar|complex sugars]] like [[sucrose]] and [[starch]]es into simpler sugars such as [[glucose]] and [[fructose]]. This process slightly raises the water content and the acidity of the partially digested nectar.

Once filled, the forager bees return to the hive. There they regurgitate and transfer nectar to hive bees. Once it is in their own honey stomachs, the hive bees regurgitate the nectar, repeatedly forming bubbles between their [[Mandible (insect mouthpart)|mandibles]], speeding its digestion and concentration. These bubbles create a large surface area per volume and by this means the bees evaporate a portion of the nectar's water into the warm air of the hive.

Hive bees form honey-processing groups. These groups work in relay, with one bee subjecting the processed nectar to bubbling and then passing the refined liquid on to others. It can take as long as 20 minutes of continuous regurgitation, digestion and evaporation until the product reaches storage quality. The new honey is then placed in honeycomb cells, which are left uncapped. This honey still has a very high water content, up to 70%, depending on the concentration of nectar gathered. At this stage of its refinement the water content of the honey is high enough that ubiquitous [[yeast]] [[spore]]s can [[Asexual reproduction|reproduce]] in it, a process which, if left unchecked, would rapidly [[Fermentation|consume]] the new honey's sugars. To combat this, bees use an ability rare among insects: the [[Endogeny (biology)|endogenous]] generation of heat.

Bees are among the few insects that can create large amounts of body heat. They use this ability to produce a constant ambient temperature in their hives. Hive temperatures are usually around {{convert|35|C|F}} in the honey-storage areas. This temperature is regulated either by generating heat with their bodies or removing it through water evaporation. The evaporation removes water from the stored honey, drawing heat from the colony. The bees use their wings to govern hive cooling. Coordinated wing beating moves air across the wet honey, drawing out water and heat. Ventilation of the hive eventually expels both excess water and heat into the outside world.

The process of evaporating continues until the honey reaches its final water content of between 15.5% to 18%. This concentrates the sugars far beyond the [[Solubility#Factors affecting solubility|saturation point]] of water, which is to say there is far more sugar dissolved in what little water remains in honey than ever could be dissolved in an equivalent volume of water. Honey, even at hive temperatures, is therefore a [[Supercooling|supercooled]] solution of various sugars in water. These concentrations of sugar can only be achieved near room temperature by evaporation of a less concentrated solution, in this case nectar. For [[Osmotic pressure|osmotic reasons]] such high concentrations of sugar are extremely unfavorable to microbiological reproduction and all [[fermentation (food)|fermentation]] is consequently halted. The bees then cap the cells of finished honey with wax. This seals them from contamination and prevents further evaporation.

So long as its water concentration does not rise much above 18%, honey has an indefinite shelf life, both within the hive and after its removal by a [[beekeeper]].

=== By other insects ===

Honey bees are not the only [[Eusociality|eusocial insects]] to produce honey. All non-parasitic [[bumblebee]]s and [[stingless bee]]s produce honey. Some wasp species, such as ''[[Brachygastra lecheguana]]'' and ''[[Brachygastra mellifica]],'' found in South and Central America, are known to feed on nectar and produce honey. Other wasps, such as ''[[Polistes versicolor]]'', also consume honey. In the middle of their life cycles they alternate between feeding on protein-rich pollen and feeding on honey, which is a far [[Energy density|denser]] source of [[food energy]].

=== Human intervention ===

Human beings have [[Domestication|semi-domesticated]] several species of honey bee by taking advantage of their swarming stage. Swarming is the means by which new colonies are established when there is no longer space for expansion in the colony's present hive. The old queen lays eggs that will develop into new queens and then leads as many as half the colony to a site for a new hive. Bees generally swarm before a suitable location for another hive has been discovered by scouts sent out for this purpose. Until such a location is found the swarm will simply conglomerate near the former hive, often from tree branches. These swarms are unusually docile and amenable to transport by humans. When provided with a suitable nesting site, such as a commercial [[Langstroth hive]], the swarm will readily form a new colony in artificial surroundings. These semi-domesticated colonies are then looked after by humans practicing apiculture or meliponiculture. Captured bees are encouraged to forage, often in agricultural settings such as orchards, where pollinators are highly valued. The honey, [[Bee pollen|pollen]], [[Beeswax|wax]] and [[Plant defense against herbivory|resins]] the bees produce are all harvested by humans for a variety of uses.

The term "semi-domesticated" is preferred because all bee colonies, even those in very large agricultural apiculture operations, readily leave the protection of humans in swarms that can establish successful wild colonies. Much of the effort in commercial beekeeping is dedicated to persuading a hive that is ready to swarm to produce more honeycomb in its present location. This is usually done by adding more space to the colony with ''[[honey super]]s'', empty boxes placed on top of an existing colony. The bees can then usually be enticed to develop this empty space instead of dividing their colony through swarming.

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 Honey use and production has a long and varied history, with its beginnings in [[Prehistory|prehistoric times]]. Several cave paintings in [[Cuevas de la Araña]] in [[Spain]] depict humans foraging for honey at least 8,000 years ago. While [[Western honey bee|''Apis mellifera'']] is an [[Old World]] [[insect]], large-scale meliponiculture of [[New World]] stingless bees has been practiced by [[Maya civilization|Mayans]] since [[Pre-Columbian era|pre-Columbian]] times.
 {{TOC limit|3}}
+== Formation ==
+[[File:Bee on -calyx 935.jpg|thumb|A [[honey bee]] with its proboscis extended into a [[Calyx (botany)|calyx]] of [[goldenrod]]]]
+[[File:Honey comb.jpg|thumb|[[Honeycomb]] displaying [[Hexagonal prismatic honeycomb|hexagonal prismatic]] [[Beeswax|wax]] cells in which [[honey bee]]s store honey]]
+=== By honey bees ===
+Honey is produced by bees who have collected [[nectar]] or [[honeydew (secretion)|honeydew]]. Bees value honey for its sugars, which they consume to support general [[metabolism|metabolic activity]], especially that of their flight muscles during [[forage (honey bee)|foraging]], and as a food for their [[larva]]e. To this end bees stockpile honey to provide for themselves during ordinary foraging as well as during lean periods, as in [[overwintering]]. During foraging bees use part of the nectar they collect to power their flight muscles. The majority of nectar collected is not used to directly nourish the insects but is instead destined for [[regurgitation (digestion)|regurgitation]], [[Enzyme catalysis|enzymatic digestion]], and finally long-term [[Food storage|storage]] as honey. During cold weather or when other food sources are scarce, adult and [[larva]]l bees consume stored honey, which is many times more energy-dense as the nectar from which it is made.
+After leaving the hive, a foraging bee collects sugar-rich nectar or honeydew. Nectar from the flower generally has a water content of 70 to 80% and is much less viscous than finished honey, which usually has a water content around 18%. The water content of honeydew from aphids and other [[Insect|true bugs]] is generally very close to the sap on which those insects feed and is usually somewhat more dilute than nectar. One source describes the water content of honeydew as around 89%. Whether it is feeding on nectar or honeydew, the bee sucks these runny fluids through its [[proboscis]], which delivers the liquid to the bee's honey stomach or "honey crop". This cavity lies just above its food stomach, the latter of which digests pollen and sugars consumed by an individual honey bee for its own nourishment.
+In ''[[Western honey bee|Apis mellifera]]'', the honey stomach holds about 40&nbsp;mg of liquid. This is about half the weight of an unladen bee. Collecting this quantity in nectar can require visits to more than a thousand flowers. When nectar is plentiful, it can take a bee more than an hour of ceaseless work to collect enough nectar to fill its honey crop. [[Saliva]]ry enzymes and proteins from the bee's [[Insect mouthparts#Hypopharynx|hypopharyngeal gland]] are secreted into the nectar once it is in the bee's honey stomach. These [[Digestive enzyme|substances]] begin [[Hydrolysis|cleaving]] [[Sugar|complex sugars]] like [[sucrose]] and [[starch]]es into simpler sugars such as [[glucose]] and [[fructose]]. This process slightly raises the water content and the acidity of the partially digested nectar.
+Once filled, the forager bees return to the hive. There they regurgitate and transfer nectar to hive bees. Once it is in their own honey stomachs, the hive bees regurgitate the nectar, repeatedly forming bubbles between their [[Mandible (insect mouthpart)|mandibles]], speeding its digestion and concentration. These bubbles create a large surface area per volume and by this means the bees evaporate a portion of the nectar's water into the warm air of the hive.
+Hive bees form honey-processing groups. These groups work in relay, with one bee subjecting the processed nectar to bubbling and then passing the refined liquid on to others. It can take as long as 20 minutes of continuous regurgitation, digestion and evaporation until the product reaches storage quality. The new honey is then placed in honeycomb cells, which are left uncapped. This honey still has a very high water content, up to 70%, depending on the concentration of nectar gathered. At this stage of its refinement the water content of the honey is high enough that ubiquitous [[yeast]] [[spore]]s can [[Asexual reproduction|reproduce]] in it, a process which, if left unchecked, would rapidly [[Fermentation|consume]] the new honey's sugars. To combat this, bees use an ability rare among insects: the [[Endogeny (biology)|endogenous]] generation of heat.
+Bees are among the few insects that can create large amounts of body heat. They use this ability to produce a constant ambient temperature in their hives. Hive temperatures are usually around {{convert|35|C|F}} in the honey-storage areas. This temperature is regulated either by generating heat with their bodies or removing it through water evaporation. The evaporation removes water from the stored honey, drawing heat from the colony. The bees use their wings to govern hive cooling. Coordinated wing beating moves air across the wet honey, drawing out water and heat. Ventilation of the hive eventually expels both excess water and heat into the outside world.
+The process of evaporating continues until the honey reaches its final water content of between 15.5% to 18%. This concentrates the sugars far beyond the [[Solubility#Factors affecting solubility|saturation point]] of water, which is to say there is far more sugar dissolved in what little water remains in honey than ever could be dissolved in an equivalent volume of water. Honey, even at hive temperatures, is therefore a [[Supercooling|supercooled]] solution of various sugars in water. These concentrations of sugar can only be achieved near room temperature by evaporation of a less concentrated solution, in this case nectar. For [[Osmotic pressure|osmotic reasons]] such high concentrations of sugar are extremely unfavorable to microbiological reproduction and all [[fermentation (food)|fermentation]] is consequently halted. The bees then cap the cells of finished honey with wax. This seals them from contamination and prevents further evaporation.
+So long as its water concentration does not rise much above 18%, honey has an indefinite shelf life, both within the hive and after its removal by a [[beekeeper]].
+=== By other insects ===
+Honey bees are not the only [[Eusociality|eusocial insects]] to produce honey. All non-parasitic [[bumblebee]]s and [[stingless bee]]s produce honey. Some wasp species, such as ''[[Brachygastra lecheguana]]'' and ''[[Brachygastra mellifica]],'' found in South and Central America, are known to feed on nectar and produce honey. Other wasps, such as ''[[Polistes versicolor]]'', also consume honey. In the middle of their life cycles they alternate between feeding on protein-rich pollen and feeding on honey, which is a far [[Energy density|denser]] source of [[food energy]].
+=== Human intervention ===
+Human beings have [[Domestication|semi-domesticated]] several species of honey bee by taking advantage of their swarming stage. Swarming is the means by which new colonies are established when there is no longer space for expansion in the colony's present hive. The old queen lays eggs that will develop into new queens and then leads as many as half the colony to a site for a new hive. Bees generally swarm before a suitable location for another hive has been discovered by scouts sent out for this purpose. Until such a location is found the swarm will simply conglomerate near the former hive, often from tree branches. These swarms are unusually docile and amenable to transport by humans. When provided with a suitable nesting site, such as a commercial [[Langstroth hive]], the swarm will readily form a new colony in artificial surroundings. These semi-domesticated colonies are then looked after by humans practicing apiculture or meliponiculture. Captured bees are encouraged to forage, often in agricultural settings such as orchards, where pollinators are highly valued. The honey, [[Bee pollen|pollen]], [[Beeswax|wax]] and [[Plant defense against herbivory|resins]] the bees produce are all harvested by humans for a variety of uses.
+The term "semi-domesticated" is preferred because all bee colonies, even those in very large agricultural apiculture operations, readily leave the protection of humans in swarms that can establish successful wild colonies. Much of the effort in commercial beekeeping is dedicated to persuading a hive that is ready to swarm to produce more honeycomb in its present location. This is usually done by adding more space to the colony with ''[[honey super]]s'', empty boxes placed on top of an existing colony. The bees can then usually be enticed to develop this empty space instead of dividing their colony through swarming.