Common Name: Honey Bee – While there are numerous insects that produce honey from plant nectar, the bee is the most prolific and accomplished and therefore assigned the mellifluous honorific. Both honey (honag) and bee (bīa) are of Old High German origination.
Scientific Name: Apis mellifera – Latin for bee, honey-bearing, the species and genus were assigned by Carolus Linnaeus in 1756.
Potpourri: Honey bees are not native, they were introduced to North America by the English colonists of Jamestown in 1622. The importance of honey to humanity as a source of sweetness to offset the blandness of tubers and bushmeat is profound; there is evidence of its consumption 3 million years ago (MYA). Domestication of bees was a harbinger of civilization; tomb paintings of ancient Egypt depict apiaries and Minoan accounts in Linear B include reference to a honey market in the eastern Mediterranean. The Jewish Talmud of the Babylonian exile from 597 to 539 BCE proscribed against gathering honey on the sabbath, which presumes someone would want to do so. The Danish warriors of the English epic poem Beowulf drank the honey brew called mead in halls of valor, a legend that permeates Germanic and Celtic culture. It is no wonder that the early colonists could not long live without the hallowed honey; they carried it westward in migration. Thomas Jefferson in his Notes on the State of Virginia reported that Native Americans called the honey bee “the white man’s fly” and considered its appearance to be a warning to the onslaught of settler encroachment into their territories.
Bees evolved from wasps in consonance with the flowering angiosperms that first appeared in the Cretaceous Period at the end of the Mesozoic Era some 100 MYA. Just as the human farmer followed the hunter-gatherer in the biblical allegory of Cain and Abel, the nectar and pollen harvesting herbivore bees emerged from the mostly arthropod eating wasps; about 20,000 bee species have since evolved to fill diverse niche habitats. The predominant adaptations were a long lower lip called a labium with a long tongue-like glossa to extract nectar and feathery hairs on enlarged hind legs for the collection of pollen. The nest-building bees of the genus Apis first appeared at the dawn of the Cenozoic Era 65 MYA in coincidence with mammalian diversification. Primitive single-comb nests became multi-combed hives managed by complex social organizations epitomized by the honey bee. It was long thought that Apis mellifera, the Western honey bee that now predominates globally due in whole or in part to domestication, originated in Eurasia along with the other seven odd honey bee species. Modern genomic analysis postulates an African provenance, a matter of some irony as “Africanized honey bees” are the evil doppelgangers of their docile cousins.
The bee hive as trope for industry is trite but true; it literally hums with activity. With as many as 60,000 individuals, it is an arthropod city where division of labor is a matter of necessity. The singular queen is attended by the female workers who attend to foraging, feeding, and housekeeping; the louche male drones comprise about two percent the population and do nothing but stay warm and mate. Each bee goes through the entire metamorphosis of egg, larva, pupa, and adult that characterizes the Class Insecta. The female workers prepare the brood cells into which the queen deposits an egg, feed the resultant larva for about a week and then seal it up to pupate for a second week before it emerges as a member of the sisterhood. The nascent worker bees tend to the hive for about ten days before graduating to building new waxwork comb cells for the deposition of honey, pollen or eggs. After the apprentice and journeywoman experiences are mastered, the now mature and competent worker spends the balance of her fleeting time as a forager. The average worker only lives for about 6 weeks; the entire hive population is therefore replaced by turnover every 6 months. The hive is a worker bee factory made of wax provisioned by continuous airborne logistics that operates autonomously with no manager. How this is done has been a matter of considerable interest to scientists, notably Charles Darwin.
Eusociality is the name given to cohabiting animals that employ cooperative division of labor with overlapping generations; it applies mostly to ants, termites and bees that live in nests though there are a few mammal and crustacean examples. The renowned biologist Edward O. Wilson postulates that eusocial organization is the evolutionary response to the establishment of a defensible nest. The diversity of body forms, workers, soldiers and drone males evolved due to “extreme plasticity of certain genes, programmed so that the altruistic workers have the same genes as the mother queen, even though they differ drastically from the queen and each other in these traits.” Wilson’s “group theory” approach was favored by Darwin, who rationalized that altruism could result when “selection may be applied to a family as well as to the individual and thus gain the desired end.” The nest centric theory supplants its altruistic predecessor which posited relative genetic benefit as the forcing function of individuals; one would be more likely to risk diving into a raging torrent to rescue a brother than a cousin as a matter of enhancing one’s own gene propagation. The genetic argument with eusocial insects was based on female workers sharing three quarters of their genes with their sisters; the haploid drone fathers provided one half and the diploid queen provided a quarter in what is euphonically called haplodiploidy. It turns out that some eusocial insects have this trait, and some don’t; it is not a sine qua non for eusocial organization. The “selfish gene” theory promoted by Richard Dawkins is also confounded by the Cirque du Soleil of bee sex, which involves incest, rape and suicide.
The hive is in some ways like a single organism rather than the sum of many; it multiplies by division. At some point around mid-summer based on environmental factors like food availability and colony size, the female hive workers begin to create a new queen. About a dozen larger royal chambers are erected in the wax comb into which the reigning queen deposits one fertilized egg as part of her daily 1,000 + oviposition routine. Rather than feed the regent larvae the secreted high protein royal jelly for only three days, as is the standard fare for workers, proto-queens feast on it through the entire larval stage to create the egg-filled abdomens that are the morphological royal scepter. It is notable that the only difference between a worker and a queen is diet, the ultimate in you are what you eat. Near the end of these pre-coronation preparations, the reigning queen departs with about half of her cohorts to start a new hive. As the first of the replacement queens emerge as adults, their first instinctual action is to either destroy the competition if caught pupating or do battle with any others that succeed to full queenly form and function. When all is said and done, one new queen reigns with all the eggs she will ever lay. The second instinctual queen activity is sex – drone D-day.
Drone male bees result from parthenogenesis, when an egg is laid but not fertilized; it is an instinctual choice made by the queen during oviposition. Absent a father’s genetic input, drones have only 16 of the queen bee’s 32 chromosome compliment. Their only function is to pass along her genes to the next generation, a mission of penultimate importance to species adaptation and perpetuity. It is a task that they carry out with singular focus and dedication matched only by the industry of their many sisters. Their significance as an investment for hive continuity is manifest in their numbers, size and faculties. Drones can comprise several percent of a hive’s population; they must be fed adequately to maintain their larger girth and to sustain their excursions. Starting in late spring and extending into summer, drones transit to and occupy various concentration areas which are geographically fixed air volumes about 10 meters tall and 100 meters wide. One concentration area was analyzed in 1963 with an estimated 25,000 drones from 200 different nests, a Cirque du Soleil indeed. Each individual drone can only last about 30 minutes before returning to the nest for food. Day after day, they travel to and from different zones in their region hoping against hope for the appearance of a virgin queen; only about 1 in every 1,000 will succeed. Their mission is very like that of the pilotless military aircraft named by analogy; the assault on the queen is executed with the precision and force of a hellfire missile. Using their oversize and sensitive eyes, the observant drone kamikaze swoops down to execute the 2 second sexual act. An audible explosive ejaculation of the endophallus into the queen is so dynamic that castration results with the eviscerated but consummated drone crashing and dying, his life fulfilled. The no longer virgin queen will repeat this existential act up to 16 additional times with drones from as many as 16 different nests from as far away as 6 miles. When the full measure of sperm necessary to fertilize the eggs is collected, the queen returns to the nest triumphant to produce workers with as many as 17 different fathers, diversity redux.
The bees’ hive is their castle, and it would be as cold and as drafty as Elsinore without colonial cooperation. The apian version of heating, ventilating and air-conditioning (HVAC) maintains core nest temperature at about 93 degrees Fahrenheit; a gradient extends to the ambient temperature periphery. While this may seem a trice warm, it approximates the warm-blooded temperature (98.6ºF) that human metabolism maintains; the cold-blooded but busy bees are most efficient at about the same point (95ºF), maintaining that temperature largely by shivering motions to supplement kinetic activity. Heating is a matter of group thermodynamics as the workers congregate in the center of the nest to protect the queen and the very young; even the drones participate by putting their hulking bodies to good use. A laboratory experiment with 1,000 bees placed in an enclosure and left overnight at 60ºF revealed that the clustered bees had an average elevated body temperature (thermogenesis) of 90ºF with the youngest bees at the center. Cooling to prevent mid-summer overheating is a combination of ventilation and AC; thousands of micro-wafts created by bee wing flaps of up to 200 times a second are supplemented by the evaporation of water collected for this purpose. An ill-constructed nest is more difficult to maintain as gaps allow ambient temperatures to prevail. Honey bees gather a resinous material from tree buds and wood sap that they process to create a sticky sealant called propolis (colloquially bee glue) that is meticulously applied to stop leaks.
The bee factory workers that maintain and inhabit the hive also make beeswax for construction and honey for food. Bees are very busy; they must be. Beeswax is so culturally pervasive that it has become a single word, euphonious in the playground retort to mind your own. It is an excretion from eight glands on the ventral side of the abdomen of worker bees, a complex amalgam of hydrocarbons, esters and fatty acids. The individual exudate flakes are collected, chewed and coalesced with pollen and propolis to form comb building material at a rate of about 1,000 flakes per gram. The temperature controls maintained by the hive are in part a matter of its building materials; bees require a temperature in the 93ºF range to make beeswax flakes; the hexagonal comb structure becomes brittle and unmalleable when too cold and loses shear strength causing creep flow when too hot. Beeswax was one of the first plastics used by humans, taking advantage of its temperature dependent rigidity for sealants and combustibility for candles. While many insects and birds make nests from gathered and processed materials, the wax house is uniquely apian. One can only marvel at the intricacy of nature and be humbled by its inexorable evolution; the bees’ knees.
Honey is the lifeblood of the bee brood; pollen is its daily bread. In human discourse, honey is synonymous with goodness and widely used as a term of endearment; Baltimoreans are noted for the terse “hey hon.” Flowers make nectar to attract pollinators and bees take full advantage. Workers make about 15 sorties a day visiting some 100 blossoms per flight to extract the proffered nectar and obligingly transport the male pollen from one flower’s stamen to the female pistil of the next. Pollination is the bee part of the “birds and the bees;” the egg is for the birds. Making honey from nectar is a matter of bee physiology. The extracted floral essence of about 80 percent water with sugar mostly in the form of sucrose is enzymatically converted by the worker bees to fructose and glucose and then spread to thicken by evaporation into honey with only 20 percent water. Each bee’s lifetime contribution of ½ gram of honey provides a miniscule portion of the hive’s 25-kilogram annual honey requirement. Given the magnitude of the honey acquisition requirement and the dire consequences of failure, the quest for floral nectar sources has high selective potential. Pollen is also collected and brought back to the hive and mixed with honey to make bee bread, an anthropocentric name that implies a food staple. As the pollen collected varies considerably as to sourcing, so does its derivative food; bee bread is generally high in proteins and sugars with some vitamins and minerals. Suffice it to say that maintaining a bee hive is a taxing proposition, flower powered by nature’s interactive ingenuity.
Knowing where to find nectar and how to get there first in the competitive airspace of arthropods is a matter of survival; each flower has just a taste of nectar and some have none. Honey bees have evolved what is known as the honey dance, first worked out by Karl von Frisch for which the Nobel Prize in physiology was awarded in 1973. A food source within 80 yards is indicated by a circular movement and one more distant with a modified tail-wagging movement; the number of cycles proportional to range. Direction is conveyed by the angle of motion relative to the vertical on the hive wall to be the same as the angle of departure relative to the sun; tail up meaning toward the sun and tail down meaning away. When overcast, the sun’s position is estimated by shadows; its diurnal movement is accounted for as the afternoon passes with rotational regularity. How this singular behavior arose is a matter of the speculations of psychology; it would seem so much easier to simply follow a scout bee to the beckoning blooms. The bee dance may be evolving, as some recent observations suggest that successful foragers return to the hive to shake one or more of their cohorts; presumably they then all go back together.
The honey bee’s sting is a vestige of its wasp provenance. The mostly carnivorous wasps and hornets sting offensively to kill prey and defensively to deter intruders. The bees, pacifist by comparison, fight only when flight is impractical as a matter of nest defense. Honey bee stingers differ from those of other hymenopterans in that they are barbed at the end, a trait thought to have evolved as a means of repelling mammalian invaders whose elastic skin can be punctured. When this occurs, the barb is implanted, and the bee is essentially eviscerated and dies as the venom sac is torn out with its associated musculature to allow for continued venom insertion, a matter of enhanced deterrence. Bee venom or apitoxin is a complex mix of chemicals; the pain-inducing peptide melittin is the primary constituent augmented by histamines that promote an allergic response. It was designed by wasps to kill which is what it will do depending on the dose; it is estimated that 500 stings constitute a lethal dose for adults. About 50 people die annually in the United States due to stings from bees and wasps, mostly due to the allergic reaction of anaphylaxis. Bee venom has a long history of medicinal usage that dates to the Greek Hippocrates who used it to treat arthritis. Apitherapy is one of the burgeoning natural cures that social media has exploited and hard science largely discounts; bee venom is promoted for the treatment of nerve-related conditions of neuralgia and multiple sclerosis. Whatever else it may do, bee venom did exactly what the bees had intended, it kept people and other marauding mammals away from their nests so that they achieved global success as a species. So much so, that beehives are now created, nurtured, and protected for the pollination and honey that their resident bees so assiduously provide.
Beekeeping is as old as recorded history, a natural progression of domestication of animals by humans that enabled agricultural settlements that succeeded nomadic hunter-gatherer cultures. Collecting honey from wild bee colonies is primordial and was the predominant practice in North America until the advent of large commercial crop production in the early 20th century. The first documented instance of domestic beekeeping was in the Massachusetts Bay Colony in 1641, the object of a lawsuit that likely resulted from a dispute between neighbors; the litigious society has deep roots. The first instance of crop pollination using artificial and movable bee colonies was in New Jersey apple orchards in 1909. One hundred years later, about 3 million bee colonies are moved from field to field and state to state to connect pollinator to pollinated in the brief window of florescence. Honey production has become a $300 million collateral business to the $13 billion annual value of pollination services; bees are big business. Almond pollination alone requires 2 hives per acre; the annual California ritual comprises the single largest pollination event in the world. In 2016, 1.9 million honeybee colonies were needed to pollinate 940,000 acres of almonds, the crop worth $21 billion to the state’s economy. Things are not going well with the bee business; apian pandemics such as colony collapse disorder dominate agricultural news – annual colony losses have averaged 38 percent over the past 10 years. According to one official at USDA, “we are one poor weather event away from pollination disaster.” One promising yet ironic alternative to domestic honey bees delivered by the truckload is the promotion of indigenous wild bees. Attempts to determine the cause of colony collapse disorder have proved inconclusive to date. One not improbable reason is that making honey bees domestic and genteel made them less robust, subject to the onslaught of other arthropods and bacterial infestations. The success stories of nature are its survivors, adapting over time with the genetic diversity of sexual selection. For Homo sapiens, one can only say so far so good and hope for the best.