The beekeeping industry is essential to agribusiness, yet it relies on technology whose major breakthrough came in 1851 and whose every activity must accommodate the unalterable social universe of an insect.

By Roger A. Morse

As long as there has been technology it has been applied to agriculture. Every major advance throughout history, from metallurgy to steam power to computers, has been used to make the natural processes of growth more productive and efficient. Just as hunting and gathering have given way to neat rows of crops and pens filled with cattle, another area of food production-vital but easily overlooked-has been systematized in much the same fashion: apiculture, or the raising of bees. And as technology has revolutionized the way crops and animals are raised, so too has it made an enormous difference in the practice of beekeeping.

Consider, for example, the influence of the automobile. Early each May more than 100 truckloads of honeybees roll up Interstate 95 from several Southern states, but mostly Florida, to Maine for the purpose of pollinating blueberries. Each of these trucks, which are as large as any on the road, is covered with heavy, coarse netting to confine the bees while allowing air to flow over the hives and help the bees maintain a temperature of about 95 degrees. Beneath the netting a typical truck carries 400 to 500 hives, or colonies, of bees. Each colony contains a queen, a thousand or so males (known as drones), and perhaps 20,000 to 30,000 underdeveloped female worker bees. It also holds about 20,000 young, collectively known as brood, in the form of eggs, larvae, and pupae.

Many of these honeybee colonies have just finished harvesting a crop of orange-blossom honey in Florida. When they are done with their work in Maine, about half of them will be moved to Cape Cod to pollinate cranberries, which reach peak bloom in early July. Some of the rest will stay in Maine to collect honey from wild raspberries; others will go west, especially to Michigan, for cucumber pollination. In August many of these hardworking bees will be moved to western New York State to make a crop of goldenrod honey, after which they will go back south. If they start to get too hot while on the road, they can die if their hives are not showered with water or chipped ice (refrigerated trucks have proved expensive and hard to load and unload).

This pollination industry is a twentieth-century phenomenon; the earliest known example of bee rental occurred at a New Jersey apple orchard in 1909. Development of the automobile helped create both the problem and the solution: New suburbs cleared out woods and other natural habitats where bees had built their hives, but at the same time, the improvement of roads and spread of car ownership made it easy for domesticated bees to be taken wherever they were needed.

Maine's need for bees is small compared with that of California, where about 900,000 colonies are used for the pollination of almonds alone. More bees pollinate almonds than any other crop in the United States. If no bees were available to rent, the California almond industry (which produces more than half the world's total) would disappear. About 250,000 colonies are used for apple pollination in the Northern states; about the same number pollinate melons in Florida and the Southwest and alfalfa in the West.

All told, more than 1.1 million colonies of honeybees are rented each year to pollinate about 50 different crops in the United States. Most of them are used on two separate crops, for a total of more than 2.2 million rentals at fees varying from $35 to $60 per colony. American beekeepers maintain a total of almost 3 million colonies; more than a third of them are used in commercial pollination. They are generally lifted on and off trucks by specially designed forklifts, loaded in the dark or the rain, when they are mostly at home and quiet, and when they're let out at a new destination they can find their target flowers within minutes.

One difference between the free pollination provided by nature and the paid kind is that when farmers hire the service, they often get several hives per acre of crops to ensure maximum fertilization. This guarantees a good crop yield but makes a large number of bees compete for a limited amount of pollen and nectar. Thus, while commercial pollination might seem like free money for the beekeeper-getting paid for what he would be doing anyway-in fact it usually weakens a colony by making the bees work too hard for too little food. It also exposes them to insecticides, which can sometimes be harmful, though government restrictions have greatly reduced the problem over the last two or three decades. Therefore most commercial pollinators also feed their bees for free, at lower densities, on wild or cultivated plants. The typical commercial beekeeper still makes more money from honey sales than from colony rental, though the margin continues to shrink. A small but significant beeswax industry also exists, accounting for about 5 percent of beekeepers' income.

Bees have been studied for millennia, with ancient Egyptians, Greeks, and Romans making detailed observations of their behavior and social organization. Yet the role of bees in pollination was not elucidated until late in the nineteenth century and was incompletely understood for many years afterward.

Pollination is necessary because plants cannot move, which makes for a dull sex life. They need a go-between to transfer pollen, which is the equivalent of semen in animals, from male to female flower parts. Depending on the species, these parts may exist on the same flower (as with apples and clover), in separate male and female flowers on the same plant (as with pumpkins, squash, and cucumbers), or in separate male and female plants (as with kiwi fruit).

A number of go-betweens can transport the pollen, including wind, water, birds, and small mammals. Most pollen transfer, however, is done by insects, especially bees. Bees differ from all other insects in two ways. First, their bodies are usually covered with plumose (branched) hairs, in which pollen grains are easily caught and carried from one flower to another; second, and equally important, they feed exclusively on flower nectar (which is their source of carbohydrates) and pollen (which provides protein and the small amount of fat they eat). As they feed, their bodies become coated with dustlike pollen, which they unwittingly carry to the next flower they visit. That's one way in which bees are useful to humans. The other way, of course, is by converting the nectar into honey to feed themselves and their young.

Wild honeybees have always had their hives raided for honey and wax, and attempts to domesticate the process go back to ancient times. Man-made bee houses have ranged from wood or earthenware cylinders,which were used in ancient Egypt, to the British skep, a dome made of straw, and the "gum," a hollow log with a rudimentary cover that was popular on the American frontier.

With all these variations, however, one problem remained: To harvest the honey, keepers had to tear apart a hive, and usually kill the bees with sulfur fumes. There was no effective way to make hives (or the bees they held) reusable. Slow and inefficient harvesting by repeated destruction continued only because wax and honey were always in demand.

The solution to this problem, and the greatest advance in several millennia of beekeeping, was discovered by Lorenzo L. Langstroth in 1851. Langstroth, a minister, was the principal of a girls' school in Philadelphia who kept bees as a hobby. He experimented with combs that hung from wooden bars, and after trying different sizes and arrangements, he discovered one extremely important fact: If he left a space no less than a quarter inch but no more than three-eighths inch around the edges of his bars, the bees would not join them to the outer case of the box. This allowed him to remove the cover and peer down into his hives.

A few weeks later, on October 31, on the way home from his apiary, he realized that if he left the same amount of space between and around the combs, then the frames would not stick to one another either. Each frame could be removed, stripped of honey, and replaced. When he came upon this realization, he later wrote, "I could scarcely refrain from shouting 'Eureka' in the open streets." With this momentous idea, modern beekeeping was born.

The habits of bees living in a hive explain why the spacing was so critical. If Langstroth left less than a quarter inch of space, the bees would fill it with propolis-naturally occurring gum or resin collected from plants. Bees use propolis, a natural antiseptic also known as "bee glue," to varnish the insides of their hive and the beeswax cells in combs and to fill cracks and crevices that might harbor microorganisms, fungi, and insects.

On the other hand, if Langstroth left too wide a space-more than three-eighths inch-the bees would fill it with comb. An intermediate gap of about five-sixteenths inch, however, was too wide for propolis and too narrow for comb, so the bees left it open for walking and working. Langstroth's five-sixteenths inch has come to be known, logically enough, as "bee space."
The Egyptians had come close to discovering bee space 5,000 or more years ago. They built round, horizontal beehives approximately eight inches in diameter out of mud. The combs were perpendicular to the side of the hive and were spaced the correct distance apart. The combs still had to be cut from the hive to harvest the honey, but they could be put back in place if the spacing between them was respected. This advance was never adopted by beekeepers in the Greek and Roman civilizations that followed. A small number of these old-fashioned pipe hives are still used in Egypt today.

Langstroth's advance changed beekeeping from a cottage industry into a commercial business. He patented his discovery in 1852, but he earned no royalties because the patent was widely infringed. He did manage to earn a small amount of money from Langstroth on the Hive and the Honeybee, the first practical book on beekeeping. It was published in 1853 and is still available today, more than 40 editions later.

In Langstroth's original design the frames hung freely from bars and had to be spaced by hand. Some time later Julius Hoffman of Canajoharie, New York, invented the self-spacing frame by adding shoulders to the end bars. Thus modified, Langstroth's basic arrangement remains in use. The standard modern beehive is 9.625 inches deep because that was the width of the board Langstroth had on hand when he made his first movable-frame hive. The outside dimensions of the box are 16.25 by 20 inches because Langstroth found that a convenient size to carry. A standard Langstroth hive accommodates 10 parallel combs or frames. A great number of other sizes and dimensions have been tried, but American beekeepers long ago decided that Langstroth's notions were practical and correct.

Three other inventions stimulated by Langstroth's discovery did much to encourage the development of the American honey industry after the Civil War. The need to extract honey from honeycomb gave rise to the first of these. Some honey is sold in the comb, giving the buyer a guarantee against adulteration, but comb honey is awkward to store and transport and cannot be pasteurized, leaving it vulnerable to crystallization. Bulk liquid honey is far more convenient for both producer and consumer, yet for centuries there was no practical way to collect it in large quantities.

In 1865 a Major Hruschka of the Italian army invented a centrifuge that would quickly extract the honey from an uncapped comb by spinning it at high speed. (Since bees keep their brood and their food separate, consumers need not worry about finding larvae in their honey.) Except for the use of electrical power, today's extractors follow Hruschka's basic design.

A second important advance came in the early 1870s, when a machine that looks like a laundry wringer was invented to help bees make straight, flat honeycombs. The rollers of this device, which is called a foundation machine, are imprinted with six-sided cells like those made naturally in the honeycomb by bees. When a thin sheet of bees-wax-often comb that is being reused after draining-is passed through the rollers, the cell bases are embossed onto it, and bees use the sheet as a foundation for their honeycomb.

Artificial foundation saves time for the bees and lets them produce more honey, since they no longer have to create and secrete as much wax to build their cells. It also ensures that the combs will be straight and parallel, which is convenient for the beekeeper. Several Europeans and Americans contributed to the invention of foundation machinery, but the first working, practical design was marketed by the A. I. Root Company of Medina, Ohio.

The third major invention that followed Langstroth's discovery was the smoker, which is used to keep bees from attacking the beekeeper. Under normal circumstances, when a beehive is attacked, guard bees release a volatile chemical substance known as alarm odor. Within seconds middle-aged bees, which carry the largest amount of venom, are ready to descend on the intruder. But if smoke is blown into the hive beforehand, the guard bees' sensory receptors will be deadened, and they will not emit the alarm odor. At the same time, the disturbance caused by the smoke will make the other bees fill their bodies' honey sacs as a precautionary measure. Gorging themselves in this way has a pacifying effect on the bees.

Ancient Egyptian beekeepers would hold a mass of smoldering, dried cow dung in one hand and blow its smoke into their hives. While the technique is effective, it also creates a lot of unwanted smoke. There was little advance on the Egyptian method until 1875, when Moses Quinby of St. Johnsville, New York, invented the modern bee smoker. It is a simple firepot with bellows and a top nozzle attached. The bellows force air through the fuel in the firepot, and the nozzle directs the smoke where it is needed with little leakage. Similar apparatus is used to simulate a smoky fire in the movies.

Much as agricultural machinery like the reaper and the cotton gin helped America to move from subsistence to commercial farming, so too did the inventions of Langstroth and his successors allow beekeeping to be pursued as a full-time occupation instead of a sideline. By 1880, for example, William L. Coggshall of Ithaca, New York, owned 4,000 hives of bees, more than anyone else in the world. He typically left home on Monday mornings with a wagon and a team of horses. His hired men followed behind. (By the turn of the century they would be traveling on bicycles.) They took a circuitous route, visiting apiary after apiary, which were usually two to three miles apart and contained 30 to 50 hives apiece. They slept in barns and took prearranged meals with farmers along the road, returning home on Saturday night. The honey was harvested in August and September, placed into wooden barrels, and brought into a shipping center at Ithaca on bobsleds in the fall and early winter.

Today's honeybee colonies are still invariably owner-operated, with no large, remotely owned corporations involved. The reason is that honeybees need close attention. They are not respecters of eight-hour days or five-day weeks. As a result, beekeepers often are forced into long workdays that do not fit a corporate schedule. Many full-time family operations manage between 500 and 1,000 colonies, though the two largest ones have maintained as many as 60,000 colonies-more than a billion bees-in a single year.

Such close, attentive care is necessary because of the demands created by honeybees' life cycle and organization. Honeybees are social insects that cannot live alone, yet no individual bee or group of bees controls life in a honeybee colony. Despite her title, the queen is not in charge; she is merely an egg-laying machine. So how do the thousands of bees in a hive know what to do? The present understanding is that bees are encoded to graduate from one activity to another as they age.

During their first day of adult life, all worker bees appear to spend their time cleaning cells like those from which they recently emerged. Following this, most workers feed the older larvae. While they do so, glands develop in their heads that after a few days produce a special secretion, called royal jelly, that is fed to young larvae and the queen. The next activities for house bees may be wax secretion and comb repair or building. Some bees become morticians, removing any dead bees they may find. Others warm the hive by clustering, or cool it by fanning their wings, as the outside temperature changes. Wing fanning is also used to vent excess carbon dioxide. Foraging for food and water, which is the most dangerous task, is done only by older bees.

Honeybees, like other insects, do not replace or repair the cells in their bodies as they age. This causes two problems: Their cells accumulate waste products that cannot be discharged, and their bodies get worn out and increasingly useless. A flying bee's wings become frayed. Workers may live for only four to six weeks in the active summer season, though they can survive four or even five months in the winter, when they endure less wear on their bodies and fewer predators.

We often think of honeybees as workaholics who never rest. In fact many bees loaf and wander around the hive looking for work. This reserve of workers who are capable of doing a variety of tasks with little or no notice makes colony life a success. Still, bees are proverbially busy because they are constantly rearing brood and developing replacement bees for those that pollinate and gather honey, who live only about a week after they become foragers. Foraging is hard and dangerous work, with predators-spiders, dragonflies, assassin flies, and birds, to name but a few-a constant problem. The bee's sting offers some protection, but many species have learned how to avoid the weapon.

Honeybees forage on at least 2,000 plant species in North America. About 35 of these plants account for 95 percent of the nectar collected. Chief among them is white Dutch clover and its several varieties, which grow so well in lawns and pastures, especially in the North. Clover and alfalfa are responsible for about 55 percent of the honey produced in North America. Their honeys are light in color and mild in flavor. Goldenrod, sage, tupelo, sourwood, orange blossom, raspberry, blackberry, and another two dozen plants are also important for honey production.

With proper management a colony should produce 80 to 120 pounds of honey a year above and beyond its own needs. The total United States harvest is more than 250 million pounds a year. The honey sold in supermarkets is a blend of mostly clover honey from many states and sometimes Canada. It is packaged by fewer than a dozen firms that annually process 10 million or more pounds apiece. The chief source is 1,500 full-time beekeepers scattered across the country.

A few large packers also market varietal honeys, such as orange blossom, tupelo, fall flower (goldenrod and aster), buckwheat, sage, etc. Hobby or semicommercial beekeepers are more likely than large packers to prepare varietal honeys for the market. Some rare honeys have remarkable flavors and a variety of colors but are unknown outside the areas in which they are produced. These include false bamboo in the Northeast, wild thyme in parts of New York and eastern Massachusetts, and tulip poplar in the central Southern states.

Almost all honey packers, large and small, pasteurize their honey to protect it from fermenting and prevent premature granulation (honey is a supersaturated sugar solution). About half of it is sold for use in the home; most of the rest goes into commercial baked goods, principally graham crackers and breakfast cereals, with perhaps 5 percent used in beverages, including honey-flavored beers and honey wines.

When we think of bees, we think of hives, but not all bees live in groups. In fact most of the bees on earth (and there are probably 20,000 or more species worldwide) are solitary, living alone and nesting in the ground or in hollow stems and twigs. Males and females meet to mate and never see each other again. The females build nests, gather food, and lay eggs; the males spend most of their time harassing the females and attempting to mate. In most instances parents never see their young.

Solitary bees play an occasional role in modern agriculture. In the Western states one type of twig-nesting, gregarious solitary bee is sometimes used to pollinate alfalfa. In New York State a solitary bee called Peponapis pruinosa, the squash bee, which is about the size of a honeybee and nests in the ground, is almost exclusively responsible for pollinating the state's more than ten million dollars' worth of Halloween pumpkins grown each year. (The Halloween pumpkin crop is probably worth more than a hundred million dollars nationwide.)

Honeybees, bumblebees, and other species also visit pumpkin flowers, but the squash bees are there earlier in the morning and are more persistent in their foraging. The honeybees used in this country are native to Europe and were brought across the Atlantic sometime after Columbus, while both pumpkins and squash bees are indigenous Americans. Since they have had so much longer to get acquainted with each other, it's no surprise that squash bees work with pumpkins more efficiently than imported European bees do.

Some more recent and unwanted imports have severely damaged the continent's bee population. In the past 30 years three new bee diseases and one pest have been brought into North America. Two of the diseases, one caused by a fungus and the other by a mite, originated in Europe. The pest is a beetle from Africa. Thousands of colonies of bees have died as a result, but most of those that remain have developed resistance.

The third disease, also caused by a mite, is native to Asia and was originally found only on an Asian honeybee species. It turned up in Wisconsin in the fall of 1987. The mites, called Varroa mites, are pinhead size and attack both larval and adult honeybees. Beekeepers, who generally shy away from putting chemicals in their hives, have been distraught at needing a pesticide to control Varroa. The chemical of choice in the United States has been a synthetic pyrethroid that is fairly innocuous, but in the fall of 1997 it was found that the mites had become resistant. Mites are notorious for the speed with which they gain resistance to pesticides.

Commercial beekeepers, who have always had to know bee diseases and understand how to treat them, have managed to cope with these modern problems, but hobby beekeepers, who once numbered 200,000 in the United States, have suffered great losses and probably own and manage fewer than a quarter as many colonies as they had 10 years ago. In the wild the effects of introduced diseases have been even more dramatic. At one time hundreds of thousands of honeybee colonies lived in hollow trees, caves, stone walls, and the sides of buildings throughout the United States. These bees have been almost completely wiped out. They will rebound someday, but gaining resistance through evolution is a tedious process that may take a decade or more.

For beekeepers, these diseases are a serious problem but not a crippling one. There are a number of safe ways to control mites. The future for the American beekeeping industry looks good. As farms grow larger, the need for pollination will increase, and no other bee species is so well known or adaptable. Honeybees have not changed significantly in at least 10,000 years. They are as untamed as ever, but the more knowledge we gain about their biology, the better we can manipulate them for our benefit.

Roger A. Morse was a professor of apiculture at Cornell University in Ithaca, New York.