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=== 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]].<ref name="Suarez Lighton Joos Roberts 1996 pp. 12616β12620">{{cite journal |last1=Suarez |first1=R. K. |last2=Lighton |first2=J. R. |last3=Joos |first3=B. |last4=Roberts |first4=S. P. |last5=Harrison |first5=J. F. |title=Energy metabolism, enzymatic flux capacities, and metabolic flux rates in flying honeybees. |journal=Proceedings of the National Academy of Sciences |volume=93 |issue=22 |date=29 October 1996 |issn=0027-8424 |doi=10.1073/pnas.93.22.12616 |pages=12616β12620 |pmid=8901631 | doi-access=free |pmc=38041 |bibcode=1996PNAS...9312616S}}</ref><ref name="NHB Bee facts" /> 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.<ref name="Suarez Lighton Joos Roberts 1996 pp. 12616β12620" /><ref name="cd">{{cite news |last=Binkley |first=D. |date=31 August 2014 |title=How bees make honey is complex process |no-pp=y |newspaper=[[The Columbus Dispatch]] |url=https://www.dispatch.com/story/news/technology/2014/08/31/how-bees-make-honey-is/23975471007/ |access-date=2 April 2022 |archive-date=14 October 2023 |archive-url=https://web.archive.org/web/20231014203304/http://www.dispatch.com/story/news/technology/2014/08/31/how-bees-make-honey-is/23975471007/ |url-status=live }}</ref> 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.<ref name="NHB Bee facts">{{cite web |url=http://www.honey.com/honey-at-home/learn-about-honey/how-honey-is-made |title=Honey and Bees |access-date=17 November 2015 |url-status=dead |archive-url=https://web.archive.org/web/20100305094736/http://www.honey.com/honey-at-home/learn-about-honey/how-honey-is-made |archive-date=5 March 2010}} National Honey Board</ref> 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%.<ref name="South Mountain Bees">{{Cite web |title=Did you know there's water in honey? |url=https://southmountainbees.com/blogs/on-bees-and-honey/how-much-water-is-there-in-honey |access-date=24 September 2022 |website=South Mountain Bees |archive-date=13 July 2024 |archive-url=https://web.archive.org/web/20240713173049/https://southmountainbees.com/blogs/on-bees-and-honey/how-much-water-is-there-in-honey |url-status=live }}</ref><ref>''Beekeeping: Everything You Need to Know to Start Your First Beehive'' by Joachim Petterson β Weldonowen 2015 Page 57</ref> 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%.<ref>{{Cite journal |last=Lamb |first=K. P. |date=1 February 1959 |title=Composition of the honeydew of the aphid Brevicoryne brassicae (L.) feeding on swedes (Brassica napobrassica DC.) |url=https://dx.doi.org/10.1016/0022-1910%2859%2990054-X |journal=Journal of Insect Physiology |language=en |volume=3 |issue=1 |pages=1β13 |doi=10.1016/0022-1910(59)90054-X |bibcode=1959JInsP...3....1L |issn=0022-1910}}</ref> 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".<ref name=cd /> 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.<ref name="Suarez Lighton Joos Roberts 1996 pp. 12616β12620" /><ref name="Rossano Larocca Polito Perna 2012 p. e49164">{{cite journal |last1=Rossano |first1=Rocco |last2=Larocca |first2=Marilena |last3=Polito |first3=Teresa |last4=Perna |first4=Anna Maria |last5=Padula |first5=Maria Carmela |last6=Martelli |first6=Giuseppe |last7=Riccio |first7=Paolo |title=What Are the Proteolytic Enzymes of Honey and What They Do Tell Us? A Fingerprint Analysis by 2-D Zymography of Unifloral Honeys |journal=PLOS ONE |volume=7 |issue=11 |date=7 November 2012 |issn=1932-6203 |doi=10.1371/journal.pone.0049164 |page=e49164 | doi-access=free |pmid=23145107 |pmc=3492327 |bibcode=2012PLoSO...749164R}}</ref> 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.<ref name="Suarez Lighton Joos Roberts 1996 pp. 12616β12620" /><ref name="cd" /><ref>{{cite web |last=Standifer |first=L. N. |title=Fact sheets |url=http://maarec.cas.psu.edu/bkCD/HBBiology/nutrition_supplements.htm |access-date=13 July 2023 |publisher=Mid-Atlantic Apiculture Research and Extension Consortium |date=2020 |archive-date=4 October 1999 |archive-url=https://web.archive.org/web/19991004222025/http://maarec.cas.psu.edu/bkCD/HBBiology/nutrition_supplements.htm |url-status=live }}</ref> 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.<ref name="cd" /> 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.<ref>{{Bulleted list|Beekeeping as a Business by Richard Jones β Commonwealth Secretariat 1999 Page 49 |The BBKA Guide to Beekeeping, Second Edition by Ivor Davis, Roger Cullum-Kenyon β Bloomsbury Publishing 2015 Page 173-174 |[https://web.archive.org/web/20121205012233/http://www.honey.com/honey-at-home/learn-about-honey/how-honey-is-made/ "Honey and Bees"]. Archived from the original on 5 March 2010. Retrieved 17 November 2015. National Honey Board}}</ref> 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%.<ref name="South Mountain Bees" /> 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.<ref name="NHB Bee facts" /><ref name="cd" /> The bees then cap the cells of finished honey with wax. This seals them from contamination and prevents further evaporation.<ref name="cd" /> 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]].<ref name="NHB Bee facts" />
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