Original Article printed in
American Bee Journal (1967) Vol. 107, No. 12: 461-462,464


The Life of the Honey Bee


Its Biology and Behavior with an Introduction to Managing the Honey-Bee Colony


by C. L. FARRAR

Entomology Research Division

Agric. Res.Serv., U.S.D.A.*


THE honey bee, Apis mellifera L., is a social organism unique in the insect world where so many kinds of insects destroy crops, bite, or are an irritating nuisance. In contrast to the insect pests, the honey bee is essential to the welfare of mankind. Not only does it provide man with honey-a delightful food with many delicate flavors-but, more important, the honey bee insures the fertility of many economic plant species man depends upon for his direct and indirect food supply.

Man could survive without honey for the table, although many would not like to do so. On the other hand, his diet would be pretty drab without the fruit and vegetables that require pollination for productivity and without the milk and meat from livestock that prosper so well on the legumes. These basic forage crops important in livestock production also must be pollinated by bees.


The honey bee reached its social climax many millions of years before man sought any level of society. Beginning with Aristotle about 300 B.C., the literature pertaining to honey bees is probably greater than that dealing with any other species except man himself. Philosophers, scientists , doctors, lawyers, ministers, and laymen have been interested in keeping and studying the honey bee. Only a small portion of those that keep bees do so with a business objective. The majority keep them for pleasure and study. How can one obtain pleasure in working with a creature that stings? Once one has gained some insight into the great depth of their biology and social behavior, their stings become of little consequence unless it is to study the intricate morphology of the stinger or some physiological effect resulting from a sting.


In a honey-bee society it is the colony that is representative of the species organism. Species perpetuation is accomplished by the colony reproducing through the process of swarming, that is by division. Over the centuries man has learned how to subordinate the reproductive instinct of the species by management practices to control swarming and to capitalize on their productive instincts. He has learned how to increase the number of colonies at will and to move them anywhere in the numbers required to meet his need.


Bees and Plant Pollination

The honey bee was not native to the Western Hemisphere-North and South America, Australia, and New Zealand. They were brought to these areas by the very earliest settlers.


How were the plants pollinated in these areas before the introduction of the honey bee? There are hundreds of kinds of bees that also pollinate plants and, in some cases' even more efficiently than the honey bee. However, when man increased his crop acreages, many nesting sites of the solitary and semicolonial bees were eliminated. The cultivation of specific crops in large acreages was as favorableto crop pests as it was detrimental to the native wild bees.The need for chemical control of these pests further upset the natural balance between the flowers that require pollination and the pollinators that were available to do the job. Honey bees in millions, or even billions, can be supplied more easily than a few thousand of the so-called wild bees.


Large numbers of honey bees are killed by the chemical pesticides, but the effect on survival of the species is less damaging. The reproductive members of the so-called wild bees are the flowers' visitors and where these are poisoned the species suffers. With the honey bee, usually only a part of the workers from the colony are killed, which permits the colony to survive even though its productivity may be destroyed for a period of time.


Incidentally, honey bees, like other bees, are wild rather than domesticated. The species would survive in the wild state without any interference from man and its behavior would continue to be unchanged. Any success man has attained in utilizing honey bees for their productivity is the result of his close study of their natural instincts and behavior. He has used this knowledge to develop management practices that permit him to operate them for his own benefit.


All bees, including the honey bee, have branched hairs covering their bodies. This is one of the important characteristics that distinguishes bees from other insects. These branched hairs become dusted with pollen grains as they visit the flowers. The pollen of a different flower of the same species usually competes well in the fertilization process with pollen from the flower being pollinated. Since a bee may visit 100 to 400 blossoms during each trip to the field, cross- pollination is effected by distribution of pollen grains from the anthers of one flower to the stigma of another. Many plants with perfect flowers, that is bearing both anthers and stigma, are self-sterile to their own pollen; in others the mate and female flowers are produced on different plants; and in others the male and female blossoms are separate on the same plant. All three situations make bees essential to the production of seed and fruit. Even self-fertile plants are usually more productive when crosspollinated.


The Colony

The honey-bee colony consists of one queen, many thousand worker bees, and, at certain seasons, from a few to several thousand drone or male bees. It also includes a series of parallel honey combs made up of six-sided cells on both sides for the rearing of young bees (brood rearing) and for the storage of food-pollen and honey. The domicile to house these must be considered an element of the colony, since the colony would experience great difficulty in surviving without protection from the weather elements. Individual queens, workers, or drones cannot survive alone. Collectively, the bees of a colony cannot survive without combs for brood, honey, and pollen.


"The Life of the Honey Bee, Its Biology and Behavior with an Introduction to Managing the Honey-Bee Colony" was written to stimulate interest and use in secondary school Biology courses and to make available to beginners basic information about bees and their management. In this introduction to management, the whole procedure is portrayed with a series of pictures. In this first article in this issue, appears the story of the honey-bee colony and the three individuals that comprise the colony-the queen, the drones and the worker bees. This will be followed in the next issue by how bees communicate, bee stings, and the introduction to keeping bees. Finally, in the last issue, will appear a picture story of colony management.

The Queen

The queen is a large, slender individual whose sole function is to lay eggs. In this she is most remarkable, since she may produce up to 1,500 eggs per day, 250,000 per year, and, under some circumstances, more than a million during her lifetime. The eggs laid in a single day may weigh more than her own body weight. She lays two kinds of eggs-those that she fertilizes and places in small hexagonal cells that develop into worker bees and those that she does not fertilize, which she places only in the larger hexagonal cells that develop into drone bees. Occasionally she places fertilized eggs in round, peanut-shaped queen cells. These are temporary cells that hang down. The worker and drone cells lie in a horizontal position. Approximately 95 percent of the typical hexagonal cells of the honey comb are worker cells, 25 per square inch; the remainder are drone cells, 16 per square inch.


The queen develops from a fertilized egg the same as the worker bees. The egg hatches in 3 days, For 5 days the queen larva is fed a food secreted by the worker bees known as royal jelly. Then the larva is sealed in its cell where it spins a cocoon and goes through a number of transformations before emerging 7 days later as a fully developed queen bee. Five to 8 days after the queen emerges, she leaves the colony to mate with a number of drones or male bees. From these matings she is able to store 5 to 8 million sperm cells in a tiny organ called the spermatheca. She releases several sperm from the spermatheca each time she lays an egg destined to produce a worker or queen bee. When she places an egg in the drone cell, she does not release sperm to fertilize the egg. Once the mated queen commences egg laying-usually 8 to 10 days after she emerges-she never leaves the hive, except when the colony swarms. The queen is constantly attended and fed by the worker bees in the colony. She may live 1, 2, and sometimes as many as 5 to 7 years.


The Drones

The drones are the largest bees in the colony, heavy bodied, and more or less rectangular in body shape. They develop in 24 days from an unfertilized egg and thus have a grandfather but no father. They become sexually mature about 12 days after emergence and die instantly upon mating. The matings take place in flight, often several miles from the colony in which the queens and drones are produced. Like the pollen of flowers, there must be a great excess of drones to insure matings of queens that fly over a vast area.


Only a small percentage of the drones fulfill their basic function. At the close of the honey harvest, the drones remaining in the colony are driven off the combs until they become weakened from starvation. They are then carried out of the hive by the worker bees to perish. A few drones may develop late in the season and overwinter with the colony. The drones are mainly reared and tolerated by the colony during spring and summer.


The Worker Bees

The worker bees are sexually underdeveloped females smaller than the queen but capable of laying small numbers of eggs under some conditions. Worker bees that lay eggs are called laying workers, Their eggs, usually placed in worker cells, develop into undersized but functional drones.


Worker-bee larvae hatch from the eggs in 3 days, are fed royal jelly for 2 1/2 days, and then their diet is changed to include pollen and honey for 2 1/2 days. They are sealed in their cells for 12 days, during which period they spin a cocoon and transform from the larvae to the pupae, emerging as adult bees 20 days after the eggs were laid.


The difference in the cell and food environment causes the worker bees to require 5 days longer to develop the queen, yet their life expectancy is only 5 weeks during the summer and a few months during the winter. Any' worker larva under 24 to 48 hours old can be developed into a queen under the proper colony conditions that insures the nurse bees will construct a queen cell and feed royal jelly lavishly to the developing larva. The rearing of queens for market is a highly specialized field of beekeeping.


The worker bees differ markedly from the queen in many respects other than function, length of life, and behavior. Structurally they have a longer tongue for gathering nectar, modified mandibles (jaws) especially designed for comb building, special glands for secreting royal jelly, enzymes for the conversion of nectar into honey, and glands that function in communication; highly specialized leg struc- I tures for gathering and carrying pollen, four pairs of wax glands on the underside of their abdomen for the secretion of wax, and a straight barbed sting for the defense of the colony. The queen's sting is curved and smooth and is used only to destroy rival queens.


The worker bees exhibit a well-defined division of labor based primarily upon their physiological age but modified to some degree by the needs of the colony. The physio- age of bees is similar to their actual age during the active season when the colony is raising brood and storing food. During dearth periods, especially in winter, a 60-day old bee may be younger physiologically than a 20-day old bee in summer.


In a general way, bees under 3 days old clean and polish the cells for the queen to lay in and for food storage; those 3 to 7 days old feed the older larvae; those 7 to 14 days old secrete royal jelly for feeding the queen, younger worker larvae, and queen larvae of any age, and they secrete wax for comb building; those 14 to 21 days old forage primarily for pollen; and those over 21 days old forage for nectar. All the bees in the colony probably contribute to the process of changing nectar into honey and in the air conditioning of the colony to maintain a suitable temperature and hu midity. Other labor activities include gathering water and propolis, and defense of the colony.

There is considerable overlapping of the age groups engaged in the various duties. When the age groups are not in normal balance, bees of any age can do the work necessary but not so efficiently. Bees under 3 days old and the field bees can feed the queen and raise brood or they can secrete wax and build comb even though their glands are not fully developed or they have degenerated from lack of use. Similarly, very young bees can forage for pollen and perhaps nectar when there are no field bees of normal age to do this work.


Worker bees inherit many skills man employs that they manifest purely on a behavioral basis whereas man has had to develop these through intellectual inquiry, learning, and experience. They are skilled architects and craftsmen, qualified dieticians and nurses, proficient housekeepers, experts in heating and air conditioning, and fully qualified to police and defend their colony.


Their architectural skill and craftsmanship is exemplified by the beauty of the honey comb, its structural strength, economy of material, and the rapidity with which they construct the uniform hexagonal cells. The building of comb is accomplished by first "plastering" the wax into approxi- mate position in the form of round cells, and then thinning down the wax walls to a uniform thickness to produce the hexagonal cells for strength and economy of wax. As dietitians they prepare one kind of food for the queen larvaeand another for the worker and drone larvae. Each larvae receives approximately 10,000 visits from the nurse bees during development. The hive is maintained immaculately clean at all times, and the guard bees with their stingers for armor protect the hives against all intruders.


Honey bees, like other insects, are cold-blooded and have a body temperature close to that of their environment. However, the honey-bee colony functioning as a single organism can maintain uniform hive temperatures under northern winter conditions identical with those in summer or in the tropics. Only recently has man accomplished this by developing elaborate heating and air-conditioning equipment. By clustering together, they generate and conserve heat, or they lower the temperature by evaporating moisture and establishing air currents through the colony to maintain a uniform temperature of 93' F. within the cluster, even though the outside temperature is at -50' F. or 120' F. Under low temperatures, the cluster temperature ranges from 45' F. on the surface to as high as 93' F. within when brood is being reared.


The most conspicuous characteristic dominant in honey bees is their great industry. Honey bees do not procrastinate by doing tomorrow what they can do now. They may fly 50,000 miles and visit 5,000,000 blossoms to gather enough nectar to produce one pound of honey, which is stored not for themselves but for the survival of the colony. The bees that gather this food do not live long enough to enjoy it. One bee, of course, cannot fly such a distance, yet the bees of a colony may store 5, 10, or even 20 pounds of honey in a day. They must gather 200 to 300 pounds of honey and 50 pounds of pollen (10 gallons) to meet the colony's needs each year. The beekeeper also expects to harvest a surplus of 100 or more pounds of honey for his efforts. The bees have to be industrious to gather so much food, rear so many young, build comb, air condition the hive, and perform all the other duties peculiar to the colony.



(Continued in the January 1968 issue.)