About Honey
Nectar in the Scheme of Things
Nectar is the gold that entices insects to pollinate flowering plants. When it comes time for these plants to reproduce, the anthers of their flowers make pollen that must be conveyed to the stigma, where it germinates and triggers the development of seeds and fruits. Often, a plant will not accept it’s own pollen; it must devise means to acquire pollen from other individuals. Some plants rely on the wind; others employ pollinating insects such as bees.
To attract and reward these insects, plants produce nectar. Its principal ingredients are sugars – primarily sucrose, glucose and fructose. They are fabricated from carbon dioxide and water, using the energy of sunlight captured in the process of photosynthesis. The bonds between the atoms of these sugars store the energy of the sun for a rainy day or a cold winter.
In addition to these sugars, nectar contains minerals, amino acids, enzymes, and flavonoids. Each plant species produces nectar according to its own recipe, imparting a signature aroma and flavor designed to attract its preferred pollinators.
As honeybees move from blossom to blossom collecting nectar, the pollen that adheres to their bodies is conveyed to stigmas where it initiates the production of seeds and fruits. The resulting fruits, vegetables, seeds, and nuts compose about one third – a most nutrient-rich fraction - of the North American diet.
Honeybees also collect pollen for its own sake; they have tiny baskets (corbiculae) on their hind legs for this purpose. It is their major source of protein. Nectar and pollen provide complete nutrition for honeybees.
Honey from Nectar
Having supplied a crucial step in the life cycle of flowering plants and, by so doing, provided the animal kingdom with food, honeybees carry the nectar back to their hives. From it, they create honey: They add enzymes that convert most of the sucrose to glucose and fructose and they distill it by patiently fanning it with their wings to evaporate most of the water. When it is just right, they seal it in beeswax cells. By these means they convert nectar, which is perishable, to honey, which is not; it can be stored indefinitely without spoiling.
The bees consume honey throughout the winter, tapping its stored energy. Their specialized muscles metabolize its sugars to produce heat; honeybees can maintain portions of their hive above 90 degrees F, even when the outside air temperature falls below zero.
Happily for humans, honeybees do not have an off-switch; they collect nectar as long as it is available and there is a place to put it. A hive may store hundreds of pounds more honey than it can use. This unquenchable hoarding instinct permits humans to appropriate a portion of the bees’ treasure without jeopardizing their life-style.
Honey for Humans
A typical managed beehive comprises a stack of two deep wooden boxes, where the colony raises new bees, and a variable number of shallow boxes, called honey supers, where it stores extra honey. All of the boxes are filled with beeswax honeycomb set in frames that can be removed without damage to the hive.
Of several approaches to harvesting honey, Nectar of Ladue honey is typically harvested in the following way: Frames of sealed honeycomb are removed from the honey supers and gently brushed to remove adhering bees. The wax caps that seal the honey in the comb are sliced off with a serrated knife and the uncapped frames are whirled in a honey extractor – a stainless steel tank with a central rotor that holds the uncapped frames of honeycomb; the centrifugal force generated by the spinning rotor encourages the honey to separate from the comb and collect at the bottom of the tank. It is clarified by passage through a mesh strainer under the influence of gravity; this removes hive-debris without altering the composition of the honey. The honey rests for several days, allowing any remaining impurities to float to the surface to be removed. After the water content is measured to assure that it is within the optimal range, the honey is bottled and sealed in glass jars.
The Nature of Honey
Honey is the distilled essence of sunlight; like a solar storage battery, it is a reservoir for the energy of the summer sun that allows honeybees to survive the winter and humans to experience the benefits of liquid or crystalline sunlight.
Nectar Sources
Because each species of flowering plant produces nectar according to its own recipe, honeys from different floral sources differ in flavor and color. Honeybees tend to collect nectar exclusively from the most profitable source and store it in neighboring cells within their hive. Because of this, it is sometimes possible to harvest mono-floral honey that is predominantly the product of a single plant species, e.g., clover honey, orange blossom honey, buckwheat honey etc. For example, approximately once in five years, early spring weather conditions in Eastern Missouri permit bees to harvest nectar from the blossoms of the Black Locust tree (Robinia pseudoacacia) for an extended period. In those years, an early honey harvest can produce prodigious amounts of a remarkably light, almost clear honey with a delicate floral aroma and flavor.
When there is not a dominant source, the bees store a mixture of nectar from a variety of plant species. The resulting honey is often designated “wildflower honey”, a euphemism that begs the question of the source. Wildflower honeys that are harvested at different times or from different locations may have distinctly different compositions and flavors.
Living Honey
Natural honey contains living microorganism such as yeasts. They are dormant because the properties of honey prevent their growth; it is too acidic (average pH of about 4) and too dry (the water content is insufficient to support growth). In addition, the bees add an enzyme, glucose oxidase, that can catalyze the production of hydrogen peroxide, a compound that is most unfriendly to microbes. Indeed, honey has been used medicinally as an antimicrobial agent since antiquity.
Water Content
Honey is a concentrated solution of sugars and other ingredients in water. Before they put the lid on the beeswax storage cells, honeybees usually remove sufficient water to prevent the growth of microbes. If the water content is too high (greater than about 18.6%) there is a danger that microorganisms will ferment the sugars and spoil the honey.
The water content also determines the consistency of honey – whether it is thick or thin. Honey that aspires to a rating of USDA grade A must contain no more than 18.6% water. In Canada, the requirement for the top grade is 17.5% or lower. The judges at the Missouri State Fair award maximum points for honey that contains between 16.0% and 17.0% water.
Crystallization
Sooner or later, most natural honey will crystallize; there is not enough water to hold the sugars – especially glucose – in solution indefinitely (i.e., it is a supersaturated solution). Honey that has crystallized can be re-liquefied by warming it in hot water; 120 degrees F is usually sufficient. Alternatively it can be enjoyed in its crystalline form. Indeed, much of the world prefers honey that has been induced to crystallize by the addition of tiny seed-crystals. The result – termed creamed (or spun or whipped) honey - differs from spontaneously crystallized honey in that the crystals are usually smaller and more uniform, creating a smoother, spreadable concoction.
Store-bought Honey
To prolong its liquid shelf life, honey that is manufactured for the mass retail market is usually pasteurized to insure that it will not ferment regardless of the water content; in addition, it is finely filtered under pressure to remove the tiniest particles that might serve as nuclei for crystal formation. These procedures alter the nature of the honey: they inactivate enzymes, kill living microorganisms, and may remove natural constituents such as pollen. Many local beekeepers, including Nectar of Ladue, strive obsessively to offer honey that is as close as practical to the way the bees made it.
Raw Honey
As applied to honey, the term "raw" implies "natural"; but raw has no standard definition. To some it means honey that has been neither heated nor strained. Such honey is generally cloudy and contains bits of wax, propolis, unidentifiable hive components, and, inevitably, fragments of bees. Others would include honey that has been clarified by course straining, but not finely filtered under pressure. To others, honey qualifies as "raw" if it has not been warmed to temperatures above those it may have experienced naturally within the hive: 115 or 120 degrees F are often considered limits.
Honey and Infants
Honey should not be fed to infants less than one year of age owing to the possibility that, as a natural product, it could contain spores of the bacterium Clostridium botulinum. Between approximately 6 weeks and six months of age, an infant's large intestine goes through a stage when it may be hospitable to these spores, allowing them to germinate and cause the rare but serious condition of infant botulism. These spores are remarkably resistant to heat, and would not necessarily be inactivated by Pasteurization. After one year of development, the human intestine is no longer susceptible to such spores.
About Honeybees
Those Who Make the Honey
A hive of honeybees is among the most complex of insect societies. It is a bit like a single organism in which the individual bees play the role of cells. It may contain as many as 50,000 bees belonging to three different classes: worker, queen and drone.
Workers
The vast majority of honeybees are workers. A worker begins life as a fertilized egg laid by the queen in an open wax cell of the honeycomb. After about three days, a larva emerges and begins to feed voraciously upon a fare produced from honey and nectar by the glands of young adult nurse bees. Over the course of about 5 days, the larva increases in weight by as much as 1,500 fold. Then adult bees seal the larva in its cell with a wax cap. In the privacy of its sealed cell, the larva spins a cocoon and becomes a pupa that undergoes metamorphoses, the larva changing, miraculously, into a bee. After about two weeks, a brand new worker bee emerges from the cell and begins a life of labor for the hive.
The new worker engages in a succession of age-appropriate tasks. For the first three weeks, these involve domestic housekeeping duties: cleaning the beeswax cells, making them ready to receive new eggs; playing nursemaid to larva, providing them with food; removing dead bees from the hive; tending and feeding the queen bee; receiving, processing, and storing nectar and pollen brought into the hive by the foraging bees; producing beeswax flakes by means of an abdominal gland that converts honey into beeswax – it takes about eight pounds of honey to make one pound of beeswax; using the new beeswax to construct the intricate honeycomb and cap its cells; guarding the hive entrance, allowing only bees with the signature odor of the hive to enter – though foreign bees may slip by if they carry a sufficiently attractive bribe of honey or pollen. The new bee knows how and when to perform each of these complex housekeeping tasks without being taught.
After about three weeks, the new bee graduates from domestic chores and enters the ranks of the field bees. It begins foraging for nectar and pollen, using the position of the sun and other cues to navigate over an area of about 30 square miles. The amount of nectar that a single bee collects in its lifetime of foraging may produce less than 0.1 teaspoon of honey; a pint of honey may require visitation of 5 million flowers. After about three weeks of foraging, the worker bee wears out and dies.
The Queen
A hive usually has just one queen. She is responsible for producing all of the eggs. She is about 1/3 longer than a worker bee, and, unlike a worker, has a fully functional reproductive system. And yet, a queen bee originates from a larva that - like Cinderella - had it not been selected for special treatment, would have become just another worker bee; queens and workers are both female, and have the same genes. What makes them different is what they eat. Soon after the larva-that-would-be-queen emerges from her egg, she is plied with copious quantities of royal jelly - a concoction produced in the hypopharyngeal and mandibular glands in the heads of young worker bees. It contains substances – a protein named royalactin has been proposed as a major culprit – that trigger the development of queen characteristics such as functional ovaries and a receptacle for sperm, the spermatheca; the development of worker characteristics, such as pollen baskets, is suppressed. (Worker larva are fed a similar concoction, but in smaller quantities and for a shorter period of time.) The beeswax cell in which a queen develops is elongated to accommodate her extra length; it hangs vertically in the hive and is easily recognized.
When a new queen emerges from her cell, her first order of business is to eliminate the competition; she attempts to kill, by stinging, any other queen in the hive, including those that have not yet emerged from their cells.
When her primacy has been assured, she attends to the business of mating, which must be accomplished within the first several weeks of her emergence. She leaves the hive and flies to a “drone congregation area” high in the air, where male bees, drones, are drawn to await a receptive queen. Over the course of several flights, a queen may mate with more than a dozen drones. She stores their sperm - as many as six million - in her spermatheca. She draws upon this supply of sperm to fertilize each worker/queen egg that she lays for the rest of her life– as long as five years. At the peak of her egg-laying season, she may lay nearly her own weight of eggs in a day - as many as 2000. She relies upon a retinue of attendant workers to produce and feed her a sufficient quantity of royal jelly to maintain her egg-laying capacity.
Because the queen mates with multiple drones, workers and queens that develop from her fertilized eggs are related to each other as groups of half sisters, all sharing the same queen mother, but each group sired by a different drone father.
Drones
A hive in late spring and summer includes several hundred male bees, called drones. Larger than workers, drones lack a stinger, and have large upturned eyes. They are specialized for finding and mating with queens in flight in order to propagate the hive's genes. They do not participate in most maintenance chores of the hive. On warm afternoons, they are drawn by unknown forces to "drone congregation areas", high in the air, where they hang out with drones from other hives. When a receptive queen enters the area, the drones give chase and compete for the opportunity to mate with her on the fly. The act requires only seconds. Because the transfer of sperm to the queen involves the rupture and detachment of essential internal organs, the successful drone dies shortly after its consummation. At the end of the mating season, as winter approaches, the remaining unsuccessful drones are usually excluded from the hive, conserving resources.
Sex Determination
Female honeybees - workers and queens - arise from fertilized eggs. They have 16 pairs of chromosomes - 32 chromosomes in all. One chromosome of each pair is provided by the queen-mother via the egg, the other by the drone-father via the sperm. This condition - two copies of each chromosome - is called the diploid condition.
When a queen lays an egg, she has the option not to fertilize it. Unfertilized eggs have only one copy of each chromosome - all from the queen mother - just 16 total. This is the haploid condition. Unfertilized haploid eggs develop into drones. Because all of a drone’s genes come from its queen-mother, it has no father and can have no sons; its closest male progenitor is the queen’s drone father - its grandfather.
The mechanism behind this idiosyncratic method of sex determination (called haplodiploidy and shared by bees, ants, wasps, and others) involves a gene designated csd (for complementary sex determiner) located on chromosome #3. To become a female(worker or queen), an egg must have two copies of this gene and the two must be different variants (alleles) of the gene.
An unfertilized (haploid) egg has only one copy of chromosome #3 (hence only one allele of the csd gene) and, as a consequence, will become a male. When both csd genes in a fertilized diploid egg are identical (this is bound to happen because there are a finite number of alleles of csd) the egg also produces a drone; however, workers recognize and destroy diploid drones when they are young larva.
Considering the peculiarity - from a human perspective - of this mechanism of sex determination, it is curious that bees (along with birds - as in "the birds and the bees") have been singled out as the gold standard model for discussions of human reproductive facts of life).
Communication
The survival of a hive depends upon the coordinated activity of thousands of bees, each doing the right job at the right time. On some occasions, the whole population must change the direction of their activities in concert. It requires robust communication between the individuals. Bees talk to each other in at least two different languages: a chemical language and a physical language.
Chemical language
Honeybees are immersed in a sea of intensely persuasive chemical language. It is mediated by pheromones produced in their glands. When these are released and combine with specific receptors - often located in the antennae - of recipient bees, they activate molecular machinery that regulates the recipients' behavior in a manner that accommodates the needs of the hive. The chemical language of bees is analogous to that of multicellular organisms where exchange of molecules (transmitters, hormones, cytokines and growth factors) between the cells serves to coordinate their activities for the good of the organism.
The queen is a major emitter of pheromones. She exudes them from numerous glands in her head, abdomen and feet. Within the hive, these act to regulate a variety of behaviors of the workers: they serve to attract a retinue of attendants to feed and groom her; they suppress the inclination of worker bees to begin the production of new queens or to lay eggs (workers possess the anatomical machinery to lay unfertilized eggs, and in the absence of a queen and brood, they do so); they stimulate the enthusiasm of workers to carry out the tasks of the hive such as foraging and building comb. Outside of the hive, the queen’s pheromones serve to attract drones on her mating flights; they also serve to organize a swarm of bees around the queen when it sets out from the hive to set up shop in a new location.
Pheromones are also used by worker bees to talk amongst themselves: When a bee stings, it releases an alarm pheromone that suggests to recipients that the target is worthy of additional punishment; Pheromones released by the older forager bees regulate the rate at which the younger house-bees develop, thus maintaining the proper proportion of bees to perform each of the tasks of the hive; A pheromone from the Nasonov gland at the tip of the abdomen has a "come-hither" effect upon its recipients - for example, bees at the entrance to a new hive hold their abdomens high in the air and fan their wings to waft this pheromone as an aid to their hive mates in identifying the new residence.
In addition, larva release pheromones that elicit age-appropriate care and feeding from the nurse bees and inhibit the laying of eggs by worker bees. Pheromones produced by drones facilitates their assembling in drone congregation areas where they await receptive queens.
Physical language
Bees engage in variety of physical interactions - jostling, shaking, buzzing, vibrating, bumping, a "spasmodic dance" - each conveying information from the perpetrator to the recipient. But they are most famous among humankind for their "waggle dance", the basis of a symbolic language that conveys the location of a point of interest outside of the hive, as well as a qualitative assessment of what is to be found there. For example, when a scout bee identifies a superior source of nectar - say a grove of linden trees just coming into bloom - she returns to the hive and recruits her sister foragers by means of this dance: she runs for a short distance in a straight line on the vertical surface of the comb while vibrating (waggling) her abdomen. The angle between the line-of-dance and the "up" direction defined by gravity indicates the angle at which a bee must fly, relative to the sun, to arrive at the nectar source. The duration of the run reflects the distance of the source from the hive, each second corresponding to about 0.6 miles. At the end of each run, the dancer circles back to the starting point and repeats the choreography. If the foragers attending to the dance are impressed, they will use the information to find the site, leading to the efficient harvest of the nectar source.
A dancing bee may remain in the dark hive for several hours. Over this time period, her dance adjusts to reflect the changing position of the sun. Bees rely on an internal clock to keep track of the position of the sun even when they cannot see it.
The Swarm
Hives of bees reproduce by a process called swarming. When conditions are favorable, the workers initiate the production of new queens by feeding young female larva with the correct dosage of royal jelly. Some time after the new queens reach the pupal stage of development, when their survival seems assured, the old queen and about half of the workers exit the hive in a dramatic swirling of bees. The queen alights on a nearby tree branch, bush, telephone wire, or any handy object; the workers cluster around her creating an aggregate that may be the size of a basketball. Only then do they address the problem of finding a new home.
The decision is made by a committee of the oldest foragers, several hundred in number. From these, scouts set out to identify potential hive-sites. When a scout finds a promising cavity, she measures it and evaluates it; a 40 liter cavity high in a living tree with a small opening would receive high marks. Having formed her opinion, she returns to the cluster and reports the location to the committee by means of the waggle dance, performed on the surface of the cluster. Her enthusiasm for the site is expressed in the persistence of her dance. Those witnesses to the dance who are impressed visit the site and then return to express their own degree of enthusiasm for the site by means of the dance. If these advertisements for the site are more popular than those for the other sites under consideration, committee members accumulate at the site. When a quorum is achieved, the swarm is alerted; it takes to the air and, directed by the committee, flies to occupy the new site.
Like the northern lights, swarming bees is a natural event that is rare to witness and well worth the experience. Without a hive and brood to defend, swarming bees are on their most docile behavior, and, if undisturbed, are most unlikely to sting. Beekeepers are often eager to collect swarms to populate new hives.
The Sting
Hoarders of gold do well to find an inconspicuous hiding place or to carry a big stick. Though honeybees do not advertise their riches, they have embraced the latter approach to good effect. In the course of the past million years or so, they have concocted a venom that inflicts a pain proportional to the value of the honey and the hive that it serves to protect.
The pain of a honeybee sting is exquisite in its purity. It begins with the sensation of a pin prick followed by a crescendo of pain the color of blue ice. When it can’t get any more intense, it does. It is unbearable.
Thankfully, for most people it lasts for only a matter of minutes. Some experience swelling that can last for several days. A small number, perhaps about 1%, respond with a serious allergic reaction, anaphylaxis, that can be life-threatening if not promptly treated - e.g., by an injection of epinephrine.
Fortunately, the European races of the honeybee Apis mellifera rarely sting except when subjected to physical injury, or to a threat to their hives. Even then, it is quite normal for a bare-handed beekeeper to spend a day disassembling and reassembling hives full of bees without experiencing a sting. It is more common to be stung by wasps, such as yellow jackets (that closely resemble honeybees), whose nests, often in holes in the ground, can be less obvious, and hence easy to disturb inadvertently.
The stinger of a worker bee is barbed; when it is set in skin it often detaches from the bee along with essential internal organs; for lack of these, the bee dies. When stung by a honeybee, it is prudent to remove the stinger - for example, with a scrape of a thumbnail - in a timely fashion; it has a mind of its own and may continue to pump venom if allowed to linger.
Honeybee venom comprises a mixture of proteins, peptides, and small molecules that collaborate to wreak havoc within cells. The most abundant component is melittin, a cleverly designed polypeptide 26 amino acids in length. Its structure – lipophilic at one end, polar at the other – allows it to insert into the membranes of cells and organelles, disrupting their function as a barrier, and causing them to disintegrate. Melittin’s antipathy for cells has been used in laboratory experiments to kill cancer cells, viruses and parasites.