by Larry Connor
Before obtaining the first bee colony, the future sustainable apiculturist must master key aspects of bee biology. Here we look at the activities of queen bees.
Queen Activity, Behavior and Lifespan
When everything is working in a colony there is usually just one queen bee. This queen is a female bee that has been selected by her sister bees and is the only female bee that is fully reproductive. The queen is sexually active during the early part of her life, mating with multiple drones before spending the rest of her life laying eggs.
Worker bees feed and groom the queen, as well as take care of her waste products. She produces odors, chemical signals called pheromones and which we also call the ‘queen substance’ or ‘queen signal’. There may be a link between the number of eggs a queen lays and the amount of these chemicals she produces.
Who Decides Queen Activities?
New beekeepers often assume that the queen bee is in charge inside the hive, but she has no such power. In fact, the queen is chemically reactive to the needs of the entire colony. Queen feeding, waste removal, and her eventual supersedure replacement are all the results of the collaborative decision-making nature of worker bees. These decisions are based on chemical information (feedback) the bees receive from the body of the queen. She also produces eggs that hatch into larvae and pupae. This is the brood. Both open and sealed brood influence worker bee behavior. The queen decides very little.
Developmental Time and What it Means
Queens are one of two female castes of bees found inside the hive, the other caste being the worker bees. Queens and worker bees develop from apparently identical eggs that are deposited into cells by their mother queen following successful mating with multiple drones. These eggs have two sets of chromosomes, making them diploid individuals. Worker bees are unable to mate and, in queenless and broodless situations, produce eggs with a single set of chromosomes. These become drones. Both queens and worker bees produce haploid bees.
Queen bees have the shortest developmental time, running 15.5 to 16 days from the time the egg is placed into the cell to the new queen’s emergence from her queen cell. Some strains have shorter developmental times; African queens are known to develop in just 14 days.
Once a queen emerges from the cell, she will feed herself and is fed by nurse bees inside the hive. After a week or so, the queen will make orientation flights, then mating flights, coupling with 12 to 20 different drones. After several days of grooming and feeding by nurse bees inside the hive, the queen will start to lay eggs into worker cells which have been emptied and polished by the bees in the brood nest. Once she begins laying eggs, the queen does not mate again. Any shortage of sperm will not be corrected, and the fate of the queen, and her hive, is set.
In Nature, old and inferior queens are replaced through a process called supersedure. This happens when the queen’s pheromone and brood production drops to about half of its normal level. Then several larvae are selected, their cells are enlarged, and peanut shaped queen cells are built on the surface of the comb. There are three to nine supersedure cells produced in the average colony, and these cells may be located anywhere on the surface of the brood frame. The production of queen cells requires the contributions of many worker bees. Nurse bees are required for the production of royal jelly, the substance key to the development of new queens. Other bees are concerned with temperature stability to ensure proper queen development, wax secretion and cell building.
Mating, Egg-laying and Sperm Storage
Queens and drones fly on warm and calm afternoons to Drone Congregation Areas (DCAs) where the queen is receptive to the many drones that follow the queen’s pheromone plume and dark form against the sky. DCAs may be located anywhere around an apiary, and can be found by careful tracking with helium balloons or kites or radar and lures containing queen pheromone. Mating occurs 50 to 150 feet off the ground, and are thus rarely seen by humans. These are often associated with geographical ‘edges.’ Tree lines near a field, bottoms of hills, openings in heavily wooded areas and the like.
Once laying, queen bees in the wild produce about 150,000 eggs per year and depend upon two large ovaries that nearly fill her abdomen. The ovaries are made up of about 370 thin tubes called ovarioles that produce eggs on a continuous basis. In the peak of the season, a queen will produce about 1,500 or more eggs per day. Favorable weather, food supply and genetic programming stimulate her productivity. Reports of queens with egg-laying rates of 3,000 eggs per day may be a reflection of a second queen in the colony (a mother queen and her supersedure daughter, an event that occurs in over ten percent of vigorous spring colonies.)
Sperm are stored at the end of the queen’s abdomen in a clear, fluid-filled sac or sphere located called the spermatheca. This structure is covered with a fine network of breathing tubes, called trachea, that bring oxygen to the sperm stored there. The spermatheca floats in the blood (called hemolymph) of the queen and receives constant nutrition. The spermatheca holds five to eight million sperm, but a failing queen may only have a few thousand sperm and are identified by drone cells within the worker brood pattern in the hive.
Research has shown that when the queen finishes her reproductive flights, her median and two lateral oviducts are filled with sperm. The nurse bees massage her body and remove the surplus sexual fluids, while about 10 percent of the sperm successfully migrate through a spermathecal duct into the spermatheca. In one to four, days the queen will begin to deposit eggs into worker-prepared cells.
Longevity of Queens
Some queens only live a few weeks before the worker bees decide – for reasons we do not completely understand – to replace her with another. Sometimes queens stop laying eggs after several days, and no queen cells are produced from the eggs and larvae in the hive. Other queens produce a good brood pattern for several weeks before the colony replaces her with a daughter.
Once a queen is well-established in a hive, we expect her to remain there for a year or more. Reports of older queens are common, some as old as five years. Commercial beekeepers usually replace queens once a year or once every two years in non-migratory, northern operations. Small-scale beekeepers often keep queens in a hive for a longer time period if the queen continues to perform well for the colony. Bee breeders attempt to select queens that maintain egg laying for as long as possible in an attempt to select for genetic longevity within the bloodline. With selection, breeders keep productive queens for five years.
Behavior of Queens and Workers
As queen cells develop, the fully formed adult queen confined inside the queen cell produces some of the chemicals that make up part of her queen substance (pheromones). Worker bees surrounding the queen cell to keep it warm and remove the wax tip of the queen cell to expose the silk cocoon tip. It is widely thought that the workers will keep these cells under close surveillance, monitoring the development of the queen inside the cells. When the queen is ready to emerge, she will use her sharp mandibles to cut her way out of the cells. Almost immediately, she will move to other queen cells, her sisters, and chew a hole into the side with her mandibles and sting the queen inside the cell. Worker bees do not interfere with this behavior, but will remove the dead queen and her cell.
Sometimes supernumerary queens are produced in a colony and held hostage inside their cells until the bees determine the proper time for their emergence. The worker bees add beeswax to the incision the queen makes to cut herself free from the cell. While preventing her emergence, the workers carefully feed such queens to keep her healthy.
Newly Emerged Queens
After a newly emerged queen has finished killing her sisters, she moves rapidly over the combs. She does not produce as much pheromone as she will when she is a laying queen and, for the first twelve hours or so after emergence, her odor level is quite reduced. After 12 hours her queen substance production is enough for the workers to respect her as an unmated queen and to attract drones to her in the DCA for multiple mating.
Some beekeepers try smoke, strong odors and other techniques to introduce virgin queens. These may work under certain conditions but, as a general rule, virgin queens should be introduced in a queen cage with a candy release plug. This candy can be a mixture of honey and powdered sugar or common baking fondant. Virgin queens are able to fly and may escape while being handled, unlikely to return to the hive. Though a virgin queen is unmated she is a queen and is producing the pheromones and she should be treated as a queen by the hive. I place the virgin queen in a cage for three to five days before I allow the bees to remove the candy for liberation!
Virgin Queens at the Time of Mating
Worker bees may fly with the queen when she leaves for the mating flight. I have not learned of a reason for this mating swarm, but it is common in other social insects – perhaps it is a method of increasing security against predators. Back at the colony, there is a change in the behavior of the house bees while mating is underway: where bees had been storing pollen and nectar, they remove these products and polish them as a place for the queen to lay. Even the sharpest-eyed beekeeper may not be able to find the virgin queen before her abdomen starts to swell with egg laying (This is a hormonal response to the mating process.) Once mated, there should be a large area of polished brood cells for the queen to use. From the time of the last mating flight to the first eggs, queens may require one to three days for the hormonal changes and heavy feeding by workers to stimulate egg production.
Newly Mated Queens
From the time she emerges from her queen cell, it takes at least four weeks for a queen to fully mature, mate and start to lay. During this month-long period, it is possible to disrupt the delicate balance between the queen and her colony (remember, these bees are not her daughters but usually sisters). If the queen was introduced to the colony from another hive, she may not be genetically related to the queen and the balance is even more fragile. There are reports of poor introduction and early rejection of queens introduced into unrelated stocks, like putting a Russian queen into a yellow Italian hive. There are undoubtedly genetically determined variations in pheromone production, as well as key queen behaviors that worker bees monitor which we know very little about.
Once established, a queen only needs to be checked every three or four weeks to make sure she is doing her job. I like to have a queen that is clipped and/or marked so I am able to confirm her bloodline. If you find eggs and young larvae and a nice brood pattern, you have seen evidence that the queen is doing her job. This means you do not need to see the queen on every inspection! For many small-scale beekeepers some colonies may only require a queen check once or twice a year; commercial beekeepers rarely check their queens.
Grand Old Ladies!
Many beekeepers develop favorite queens and want to keep them forever. Other beekeepers want to have a set schedule of queen replacement. I view older queens, those two years or older, as Grand Old Ladies. In breeding programs older queens get special respect when they continue to produce a quality brood pattern and a gentle, productive, winter-hearty hive in their third and fourth season. She can be converted over to drone production if she is not used for grafting to introduce longevity traits in your apiary – stock development is a never-ending challenge in beekeeping.
Sometimes beekeepers move older queens into smaller hives and keep an eye on them and use them for grafting. A two-deep five-frame nucleus is great for this. The older queen can be used to establish a five-frame nuc and then a super added as the colony expands. If the colony gets too strong, remove a frame of graftable larvae and give it to someone who is producing queen bees. This reduces the population of bees, spreads good genes to other colonies and keeps the older queen in balance with her reduced egg laying. Pull out frames with supersedure cells and make increase hives with them to keep her genetics in your apiary. This is part of the Sustainable Art of Beekeeping that provides me with so much satisfaction. Letting these Grand Old Ladies die a natural death seems like a fair trade for a number of highly productive seasons. It has nothing to do with being a business person, but says a great deal about your appreciation of genetic diversity, longevity and productivity.
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