The Colony and Its Organization
Honey bees are social insects, which means that they live together in large, well-organized family groups. Social insects are highly evolved insects that engage in a variety of complex tasks not practiced by the multitude of solitary insects. Communication, complex nest construction, environmental control, defense, and division of the labor are just some of the behaviors that honey bees have developed to exist successfully in social colonies. These fascinating behaviors make social insects in general, and honey bees in particular, among the most fascinating creatures on earth.
A honey bee colony typically consists of three kinds of adult bees: workers, drones, and a queen. Several thousand worker bees cooperate in nest building, food collection, and brood rearing. Each member has a definite task to perform, related to its adult age. But surviving and reproducing take the combined efforts of the entire colony. Individual bees (workers, drones, and queens) cannot survive without the support of the colony.
In addition to thousands of worker adults, a colony normally has a single queen and several hundred drones during late spring and summer (Figure 1). The social structure of the colony is maintained by the presence of the queen and workers and depends on an effective system of communication. The distribution of chemical pheromones among members and communicative “dances” are responsible for controlling the activities necessary for colony survival. Labor activities among worker bees depend primarily on the age of the bee but vary with the needs of the colony. Reproduction and colony strength depend on the queen, the quantity of food stores, and the size of the worker force. As the size of the colony increases up to a maximum of about 60,000 workers, so does the efficiency of the colony.
Each colony has only one queen, except during and a varying period following swarming preparations or supersedure. Because she is the only sexually developed female, her primary function is reproduction. She produces both fertilized and unfertilized eggs. Queens lay the greatest number of eggs in the spring and early summer. During peak production, queens may lay up to 1,500 eggs per day. They gradually cease laying eggs in early October and produce few or no eggs until early next spring (January). One queen may produce up to 250,000 eggs per year and possibly more than a million in her lifetime.
A queen is easily distinguished from other members of the colony. Her body is normally much longer than either the drone’s or worker’s, especially during the egg-laying period when her abdomen is greatly elongated. Her wings cover only about two-thirds of the abdomen, whereas the wings of both workers and drones nearly reach the tip of the abdomen when folded. A queen’s thorax is slightly larger than that of a worker, and she has neither pollen baskets nor functional wax glands. Her stinger is curved and longer than that of the worker, but it has fewer and shorter barbs. The queen can live for several years—sometimes for as long as 5, but average productive life span is 2 to 3 years.
The second major function of a queen is producing pheromones that serve as a social “glue” unifying and helping to give individual identity to a bee colony. One major pheromone—termed queen substance—is produced by her mandibular glands, but others are also important. The qualities of the colony depend largely on the egg-laying and chemical production capabilities of the queen. Her genetic makeup—along with that of the drones she has mated with—contributes significantly to the quality, size, and temperament of the colony.
About one week after emerging from a queen cell, the queen leaves the hive to mate with several drones in flight. Because she must fly some distance from her colony to mate (nature’s way of avoiding inbreeding), she first circles the hive to orient herself to its location. She leaves the hive by herself and is gone approximately 13 minutes. The queen mates, usually in the afternoon, with seven to fifteen drones at an altitude above 20 feet. Drones are able to find and recognize the queen by her chemical odor (pheromone). If bad weather delays the queen’s mating flight for more than 20 days, she loses the ability to mate and will only be able to lay unfertilized eggs, which result in drones.
After mating the queen returns to the hive and begins laying eggs in about 48 hours. She releases several sperm from the spermatheca each time she lays an egg destined to become either a worker or queen. If her egg is laid in a larger drone-sized cell, she does not release sperm. The queen is constantly attended and fed royal jelly by the colony’s worker bees. The number of eggs the queen lays depends on the amount of food she receives and the size of the worker force capable of preparing beeswax cells for her eggs and caring for the larva that will hatch from the eggs in 3 days. When the queen substance secreted by the queen is no longer adequate, the workers prepare to replace (supersede) her. The old queen and her new daughter may both be present in the hive for some time following supersedure.
New (virgin) queens develop from fertilized eggs or from young worker larvae not more than 3 days old. New queens are raised under three different circumstances: emergency, supersedure, or swarming. When an old queen is accidentally killed, lost, or removed, the bees select younger worker larvae to produce emergency queens. These queens are raised in worker cells modified to hang vertically on the comb surface (Figure 2). When an older queen begins to fail (decreased production of queen substance), the colony prepares to raise a new queen. Queens produced as a result of supersedure are usually better than emergency queens since they receive larger quantities of food (royal jelly) during development. Like emergency queen cells, supersedure queen cells typically are raised on the comb surface. In comparison, queen cells produced in preparation for swarming are found along the bottom margins of the frames or in gaps in the beeswax combs within the brood area.
Drones (male bees) are the largest bees in the colony. They are generally present only during late spring and summer. The drone’s head is much larger than that of either the queen or worker, and its compound eyes meet at the top of its head. Drones have no stinger, pollen baskets, or wax glands. Their main function is to fertilize the virgin queen during her mating flight. Drones become sexually mature about a week after emerging and die instantly upon mating. Although drones perform no useful work for the hive, their presence is believed to be important for normal colony functioning.
While drones normally rely on workers for food, they can feed themselves within the hive after they are 4 days old. Since drones eat three times as much food as workers, an excessive number of drones may place an added stress on the colony’s food supply. Drones stay in the hive until they are about 8 days old, after which they begin to take orientation flights. Flight from the hive normally occurs between noon and 4:00 p.m. Drones have never been observed taking food from flowers.
When cold weather begins in the fall and pollen/nectar resources become scarce, drones usually are forced out into the cold and left to starve. Queenless colonies, however, allow them to stay in the hive indefinitely.
Workers are the smallest and constitute the majority of bees occupying the colony. They are sexually undeveloped females and under normal hive conditions do not lay eggs. Workers have specialized structures, such as brood food glands, scent glands, wax glands, and pollen baskets, which allow them to perform all the labors of the hive. They clean and polish the cells, feed the brood, care for the queen, remove debris, handle incoming nectar, build beeswax combs, guard the entrance, and air-condition and ventilate the hive during their initial few weeks as adults. Later as field bees they forage for nectar, pollen, water, and propolis (plant sap).
The life span of the worker during summer is about 6 weeks. Workers reared in the fall may live as long as 6 months, allowing the colony to survive the winter and assisting in the rearing of new generations in the spring before they die.
When a colony becomes queenless, the ovaries of several workers develop and workers begin to lay unfertilized eggs. Development of the workers’ ovaries is believed to be inhibited by the presence of brood and the queen and her chemicals. The presence of laying workers in a colony usually means the colony has been queenless for one or more weeks. However, laying workers also may be found in normal “queenright” colonies during the swarming season and when the colony is headed by a poor queen. Colonies with laying workers are recognized easily: there may be anywhere from five to fifteen eggs per cell (Figure 3) and small-bodied drones are reared in worker-sized cells. In addition, laying workers scatter their eggs more randomly over the brood combs, and eggs can be found on the sides of the cell instead of at the base, where they are placed by a queen. Some of these eggs do not hatch, and many of the drone larvae that do hatch do not survive to maturity in the smaller cells.
All three types of adult honey bees pass through three developmental stages before emerging as adults: egg, larva, and pupa. The three stages are collectively labeled brood. While the developmental stages are similar, they do differ in duration (see Table 1). Unfertilized eggs become drones, while fertilized eggs become either workers or queens Nutrition plays an important part in caste development of female bees; larvae destined to become workers receive less royal jelly and more a mixture of honey and pollen compared to the copious amounts of royal jelly that the queen larva receives.
Honey bee eggs are normally laid one per cell by the queen. Each egg is attached to the cell bottom and looks like a tiny grain of rice. When first laid, the egg stands straight up on end (Figure 4). However, during the 3-day development period the egg begins to bend over. On the third day, the egg hatches into a tiny grub and the larval stage begins.
Healthy larvae are pearly white in color with a glistening appearance. They are curled in a “C” shape on the bottom of the cell (Figure 5). Worker, queen, and drone cells are capped after larvae are approximately 5 ½, 6, and 6 ½ days old, respectively. During the larval stage, they are fed by adult worker (nurse) bees while still inside their beeswax cells. The period just after the cell is capped is called the prepupal stage. During this stage the larva is still grub-like in appearance but stretches itself out lengthwise in the cell and spins a thin silken cocoon. Larvae remain pearly white, plump, and glistening during the prepupal stage.
Within the individual cells capped with a beeswax cover provided by adult worker bees, the prepupae begin to change from their larval form to adult bees (Figure 6). Healthy pupae remain white and glistening during the initial stages of development, even though their bodies begin to take on adult forms. Compound eyes are the first feature begin to take on color; changing from white to brownish-purple. Soon after this, the rest of the body begins to take on the color of an adult bee. New workers, queens, and drones emerge approximately 12, 7 ½, and 14 ½ days, respectively, after their cells are capped.
Healthy brood patterns are easily recognized when looking at capped brood. Frames of healthy capped worker brood normally have a solid pattern with few cells missed by the queen in her egg laying. Cappings are medium brown in color, convex, and without punctures (Figure 7). Because of developmental time, the ratio should be four times as many pupae as eggs and twice as many as larvae; drone brood is usually in patches around the margins of comb.