This winter study post is based on BBKA module 6 honeybee behaviour syllabus items 6.2 and 6.3, which follow the life of the queen. I’m not taking the exams though they make for fascinating reading. First, a video with dazzling nature photography, ‘More than honey’, by Academy Award-nominated director Markus Imhoof takes a global examination of endangered honeybees. Here, he follows the mating behaviour of the queen and the drones.
6.2 The mating behaviour of the honeybee queen and drone including an account of the pheromones involved and the concept of drone congregational areas.
On a warm, sunny day in spring a swarm of honeybees fly away from the hive with the queen and leave behind a virgin queen to emerge from her cell. The virgin is fully sexually mature within a few days and she must mate within the next three to four weeks if she is to fulfil her duties to the hive. Her sisters, daughters of the old queen, initially ignore her, but as time passes the workers hassle her to take her mating flight. So on another warm, sunny day the virgin queen leaves the hive.
In a neighbouring hive, a big bug-eyed, fat-bottomed drone hatches from his cell. He spends his early days being fed and taking short flights. He doesn’t mature until around 12–14 days old – when the urge takes him to join his brothers and drones from the surrounding apiaries in a flight to a drone congregation area.
The drones travel to the same spot each year – an area that is about 100m away from their apiaries where they swarm around 10–40m above the ground. They take routes known as flyways that can reach 21m above the ground. How the newly hatched drones, and the virgin queens, find the same location to mate every year is a mystery. The older drones do not survive the season to tell the younger drones their secret!
Here the drones congregate noisily in the middle of the day waiting for the virgin queens. Our young queen arrives and flies higher and higher until she reaches the ‘drone layer’. Her mandibular glands release an intoxicating scent (the pheromone 9 oxydecenoic acid) which is irresistible to the drones. They form a comet tail behind the queen and chase her through the air.
Our drone is the strongest and fastest. He catches the queen with his six legs hovering above her abdomen and inserts his endophallus inside her open sting chamber, which causes him to become paralysed and flip backwards as he ejaculates. The endophallus breaks off and he falls to the ground mortally wounded. The queen continues on her flight with the drone’s endophallus, now called the ‘mating sign’, plugging the semen inside her abdomen. She will mate with 15–20 drones from different hives to her own. Each drone who mates with her will first remove the ‘mating sign’ and then insert his own endophallus before meeting the same fate as his predecessor.
That done, the newly mated queen returns to the hive. She may take several more mating flights and mate with up to 40 drones, but after her task is accomplished she will start laying eggs within a few days.
The workers are now very attentive to their new queen. She produces a scent that causes them to turn and face her as she passes. It is a wave effect that ripples through the colony creating a constant ring of workers to surround her wherever she passes – her ‘retinue’.
6.3 The queen honeybee’s egg-laying behaviour and its relationship to changing circumstances in the hive and external factors relating to climate and season.
The queen spends her days inside the hive laying around 1,500 eggs a day and taking short breaks of around 5 or 10 minutes. She will not leave the hive again unless it is to swarm like her mother.
She is very picky about the cells in which she chooses to lay an egg. First she will inspect the cell and peer closely inside to ensure it has been cleaned to her satisfaction. Then she will walk past the cell to look at other cells, before making up her mind and walking back to lay an egg in the first cell.
That decided, she sticks her bottom in the cell, produces a sticky substance to keep the egg in place and lays an egg. The egg is deposited standing upright at the bottom centre of the cell. She is very precise and lays each egg in exactly the same way, so it is easy to identify eggs that are laid by the queen and eggs that are produced by a rebellious laying worker who will lay one or more eggs along the cell wall.
The queen stores eggs and sperm separately in her abdomen and controls the release of sperm by opening and closing a small valve. The sperm from her mating flights is kept alive throughout her lifetime by the protein that she eats.
The egg that she has just laid is destined to become a female worker, so she deposits a single sperm on the egg to fertilise it. If the egg was destined to become a male drone she would leave it unfertilised.
Who decides whether the queen lays worker or drone? The workers build differently sized cells for worker and drone brood: small cells for workers and large cells for drones; larger still are the peanut-shaped cells made for new queens. The queen uses her front legs to measure the size of the cell before laying an egg – if it is a worker-size cell then she will lay an egg and fertilise it to become a worker. So it seems that the workers decide the demographics, in terms of gender and size, of the population. This is part of an ongoing debate in beekeeping: who rules the hive – the queen or the workers?
While the colony cannot survive without a queen to lay eggs, her egg-laying behaviour is often determined by her workers. For example, if the colony decides that it is time to swarm then the workers will starve the queen, which reduces her egg laying, to make her small and light enough to fly with them to a new home.
The queen’s egg laying is also seasonal and dependent on the weather. When the days grow warm in spring, her egg laying increases as the colony grows in size and she is well fed. In poor weather or when food is scarce, her egg laying decreases as she is fed less food. As the year turns into autumn, her egg laying decreases to ensure a smaller-sized colony for overwintering.
Our queen lives for around three years though she could live up to five years. In her second year she swarms, like her mother, to establish a new home and leaves behind virgin daughters to claim her old throne. However, her egg laying naturally decreases with each year of her life, as does her queen substance, the pheromone that she produces to control the colony. One day the workers choose a worker egg to become a new queen by building a queen cell around it and feeding the young larva highly nutritious royal jelly. Thus the old queen is succeeded by a young rival in a process called supersedure.
Emily and I have had plenty of dramas with our queens!
The Great Escape
In space no one can hear you scream
Myrrh, queen of the monsoon
The Bad Beekeepers Club
The red-headed queen of the Diamond Jubilee
Visit my blog index for more winter study posts.
Related revision posts from Emily Heath of Adventures in Beeland:
2nd Honeybee behaviour revision post: honeybee mating & Chelsea Physic Gardens visit
3rd Honeybee behaviour revision posts: the queen’s egg laying behaviour & seasonal variations in the size of a colony
Mid Buck Beekeepers Association Blog’s excellent revision notes for BBKA module 6
Celia F Davis. The Honey Bee Inside Out. Bee Craft Ltd, ISBN-10: 0900147075
Ted Hooper. Guide to Bees and Honey. Northern Bee Books, ISBN-10: 1904846513