Honey

A jar of honey with a honey dipper and an American biscuit

Honey is a sweet and viscous substance made by several species of bees, the best-known of which are honey bees. Honey is made and stored to nourish bee colonies. Bees produce honey by gathering and then refining the sugary secretions of plants (primarily floral nectar) or the secretions of other insects, like the honeydew of aphids. This refinement takes place both within individual bees, through regurgitation and enzymatic activity, and during storage in the hive, through water evaporation that concentrates the honey's sugars until it is thick and viscous.

Honey bees stockpile honey in the hive. Within the hive is a structure made from wax called honeycomb. The honeycomb is made up of hundreds or thousands of hexagonal cells, into which the bees regurgitate honey for storage. Other honey-producing species of bee store the substance in different structures, such as the pots made of wax and resin used by the stingless bee.

Honey for human consumption is collected from wild bee colonies, or from the hives of domesticated bees. The honey produced by honey bees is the most familiar to humans, thanks to its worldwide commercial production and availability. The husbandry of bees is known as beekeeping or apiculture, with the cultivation of stingless bees usually referred to as meliponiculture.

Honey is sweet because of its high concentrations of the monosaccharides fructose and glucose. It has about the same relative sweetness as sucrose (table sugar). One standard tablespoon (14 mL) of honey provides around 180 kilojoules (43 kilocalories) of food energy. It has attractive chemical properties for baking and a distinctive flavor when used as a sweetener. Most microorganisms cannot grow in honey and sealed honey therefore does not spoil. Samples of honey discovered in archaeological contexts have proven edible even after millennia.

French honey from different floral sources, with visible differences in color and texture

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

Formation

A honey bee with its proboscis extended into a calyx of goldenrod

Honeycomb displaying hexagonal prismatic wax cells in which honey bees store honey

By honey bees

Honey is produced by bees who have collected nectar or honeydew. Bees value honey for its sugars, which they consume to support general metabolic activity, especially that of their flight muscles during foraging, and as a food for their larvae. 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, enzymatic digestion, and finally long-term storage as honey. During cold weather or when other food sources are scarce, adult and larval 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 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 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. Salivary enzymes and proteins from the bee's hypopharyngeal gland are secreted into the nectar once it is in the bee's honey stomach. These substances begin cleaving complex sugars like sucrose and starches 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 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 spores can reproduce in it, a process which, if left unchecked, would rapidly consume the new honey's sugars. To combat this, bees use an ability rare among insects: the 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 35 °C (95 °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 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 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 reasons such high concentrations of sugar are extremely unfavorable to microbiological reproduction and all 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 eusocial insects to produce honey. All non-parasitic bumblebees and stingless bees 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 denser source of food energy.

Human intervention

Human beings have 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, pollen, wax and 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 supers, 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.

Production

Collection

Sealed frame of honey

Extraction from a honeycomb

Filtering from a honeycomb

Honey is collected from wild bee colonies or from domesticated beehives. On average, a hive will produce about 29 kilograms (65 lb) of honey per year. Wild bee nests are sometimes located by following a honeyguide bird.

To safely collect honey from a hive, beekeepers typically pacify the bees using a bee smoker. The smoke triggers a feeding instinct (an attempt to save the resources of the hive from a possible fire), making them less aggressive, and obscures the pheromones the bees use to communicate. The honeycomb is removed from the hive and the honey may be extracted from it either by crushing or by using a honey extractor. The honey is then usually filtered to remove beeswax and other debris.

Before the invention of removable frames, bee colonies were often sacrificed to conduct the harvest. The harvester would take all the available honey and replace the entire colony the next spring. Since the invention of removable frames, the principles of husbandry led most beekeepers to ensure that their bees have enough stores to survive the winter, either by leaving some honey in the beehive or by providing the colony with a honey substitute such as sugar water or crystalline sugar (often in the form of a "candyboard"). The amount of food necessary to survive the winter depends on the variety of bees and on the length and severity of local winters.

Many animal species are attracted to wild or domestic sources of honey.

Preservation

Because of its composition and chemical properties, honey is suitable for long-term storage, and is easily assimilated even after long preservation. Honey, and objects immersed in honey, have been preserved for centuries. (However, no edible honey has been found in Egyptian tombs; all such cases have been proven to be other substances or only chemical traces.) The key to preservation is limiting access to humidity. In its cured state, honey has a sufficiently high sugar content to inhibit fermentation. If exposed to moist air, its hydrophilic properties pull moisture into the honey, eventually diluting it to the point that fermentation can begin.

The long shelf life of honey is attributed to an enzyme found in the stomach of bees. The bees mix glucose oxidase with expelled nectar they previously consumed, creating two byproducts – gluconic acid and hydrogen peroxide, which are partially responsible for honey acidity and suppression of bacterial growth.

Adulteration

Honey is sometimes adulterated by the addition of other sugars, syrups, or compounds to change its flavor or viscosity, reduce cost, or increase the fructose content to inhibit crystallization. Honey has been adulterated since ancient times, when honey was sometimes blended with plant syrups such as maple, birch, or sorghum and sold to customers as pure honey. Sometimes crystallized honey was mixed with flour or other fillers, hiding the adulteration from buyers until the honey was liquefied. In modern times, the most common adulterant became clear, almost-flavorless corn syrup; the adulterated mixture can be very difficult to distinguish from pure honey.

According to the Codex Alimentarius of the United Nations, any product labeled as "honey" or "pure honey" must not be adulterated, although labeling laws differ between countries. In the United States, according to the National Honey Board, "Ensuring honey authenticity is one of the great challenges facing the honey industry today. Over the past half century, a number of honey testing methods have been developed to detect food fraud. To date, there is no single universal analytical method available which is capable of detecting all types of adulteration with adequate sensitivity."

Isotope ratio mass spectrometry can be used to detect addition of corn syrup and cane sugar by the carbon isotopic signature. Addition of sugars originating from corn or sugar cane (C4 plants, unlike the plants used by bees, and also sugar beet, which are predominantly C3 plants) skews the isotopic ratio of sugars present in honey, but does not influence the isotopic ratio of proteins. In an unadulterated honey, the carbon isotopic ratios of sugars and proteins should match. Levels as low as 7% of addition can be detected.

Production

Honey production
2023, tonnes

China	463,500
Turkey	114,886
Ethiopia	84,591
Iran	80,389
Argentina	73,395
United States	62,855
World	1,893,805
Source: FAOSTAT of the United Nations

In 2023, world production of honey was 1.9 million tonnes, led by China with 24% of the total, and Turkey, Ethiopia, and Iran as secondary producers (table).

Modern uses

Food

Main article: Mellivory

Over its history as a food, the main uses of honey are in cooking, baking, desserts, as a spread on bread, as an addition to various beverages such as tea, and as a sweetener in some commercial beverages.

Due to its energy density, honey is an important food for virtually all hunter-gatherer cultures in warm climates, with the Hadza people ranking honey as their favorite food. Honey hunters in Africa have a mutualistic relationship with certain species of honeyguide birds.

Fermentation

Possibly the world's oldest fermented beverage, dating from 9,000 years ago, mead ("honey wine") is the alcoholic product made by adding yeast to honey-water must and fermenting it for weeks or months. The yeast Saccharomyces cerevisiae is commonly used in modern mead production.

Mead varieties include drinks called metheglin (with spices or herbs), melomel (with fruit juices, such as grape, specifically called pyment), hippocras (with cinnamon), and sack mead (high concentration of honey), many of which have been developed as commercial products numbering in the hundreds in the United States. Honey is also used to make mead beer, called "braggot".