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!
The queen does not mate with drones from her own colony due to problems caused by inbreeding. She flies to a drone congregation area, although how she finds it is a mystery – her mother does not show her the way!
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 queen produces a scent (queen substance) that causes the workers to turn to face and attend her as she walks through the hive.
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.
The queen spends her days window-shopping for cells that meet her satisfaction to lay eggs.
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.
A queen cell nurturing a new queen who will eventually replace the old queen of the hive.
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.
Related links
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
Olympic Queens!
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
Recommended reading
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
Mrs Apis Mellifera 
This is just fascinating. I went to a bee class yesterday and my teacher affirmed what you wrote about who decides the gender…according to him, the worker bees decide what gender they want by building the cell a certain size. When the queen backs into the cell to lay an egg, if it’s a narrow cell, the sperm sack is squeezed to release some sperm so that bee will become a worker. (Diploid) If the cell is wide, the sperm sack isn’t affected so that bee will be a drone. (Haploid)
I borrowed that beautiful video to send to my bee club president.
Thanks for posting this.
Bee classes are fun, aren’t they? I hadn’t heard that about the worker-sized cell squeezing the queen’s abdomen to release sperm and fertilise the egg, but it makes sense. The more I learn and observe about the queen and workers, the more the queen seems like a cosseted, captive egg layer for the hive. It is an interesting relationship. Still getting my head around haploid and diploid – genes are next on my revision list! I hope your bee club enjoys the video, it’s on YouTube so you could share it on your blog too. I look forward to reading about your bee classes 🙂
After I sent it to the bee club president who gave me a call to thank me, I sent it to a university research lab and one of the scientists thought it was ‘awesome.’ Thanks again for posting it.
That’s fantastic. I really hope scientists can do more research into honeybees, they are fascinating creatures. I know many beekeepers would be keen to help with research by reporting their weekly observations and goings on inside the hive.
What a fab post, especially considering the weekend you’ve had! The video is amazing and must have taken a huge amount of work to get the mating shots. Hope you are feeling a little stronger today.
I had nothing to do all weekend but read and rest! But spending yesterday in hospital was the worst. I hate being sick. The doctors were great though and I feel fully recovered today, just rescheduling my flight out for tomorrow morning. I can look forward to Thanksgiving again 🙂
I am so enjoying learning this. I never saw a video of bees mating like that, plus never knew the drone died losing his manly part. This is so interesting. The workers making different size cells too. So fascinating.
Thank you and apologies for the late reply, I have been really ill with a bout of gastroenteritis and am just recovering and catching up. The video is amazing isn’t it? Wouldn’t you love to work on something like that? I would love to know how they got those shots in the air? Camera-man bees?
Yes, those workers are crafty!
I imagined that the workers would be sisters to the new virgin queen since they have the same mother. Or am I getting things all mixed up?
The video is breath-taking ! I have never seen a DCA let alone mating on the wing ! Truly amazing!! Thanks for sharing that with us!
I think you’re right and I’ve got myself confused – there is an overlap generation in the hive when a virgin queen supersedes the old queen and the daughters of the old queen become aunts to her daughters… bee family trees are so confusing! The video is amazing, I think it is part of a series that I’m going to see if I can buy. Just breathtaking!
Hi Emma, enjoyed reading your blog as always, there’s always a little detail or two to learn from you.
Thanks, John! As I always say, there are no secrets in beekeeping except the secrets the bees keep! Hope you enjoyed the video – Markus Imhoof deserves a Bee Oscar!
You and Emily will be able to open your own beekeeping school soon! Great post!
He he! Well, Emily’s much further ahead on the exam path but I don’t feel ready to open a bee school yet 😉
I so much enjoy reading your posts that really make the life of the bees come alive for me – and increase my fascination. The video was incredible, I cannot imagine the technology that went into making that video, let alone the patience. I loved the ladies tending the bees in the mountains and sorting them out bare-handed with such confidence!
Thank you, and again apologies for the late reply, I have been really sick.
The video blew me away. I really want to know how they filmed the flight sequences, incredible. I loved the lady lighting up a cigar to smoke the bees, you can tell that she is just at one with them. I think perhaps those honeybees must be pure bred (which is why their keepers can sell newly mated queens) because they are so gentle to handle. In remote locations like that it is much easier to ensure purer bred bees, whereas in a city like London all honeybees are mongrels within a couple of generations.
I am not sure it was a cigar, I thought it was another type of smoker. I must have another look. Also I want to see if I can read the boxes. Could they be another sub-species like carnica?
You are right it is a cigar!
The boxes do have Carnica written on them. Do you think this means they are Carniolan bees?
It must be lovely to keep pure bred gentle bees – a bit like bumble bees.:)
Ah, Carniolan bees might explain it. Our bees are gentle but we couldn’t handle them like that.
The cigar is my favourite bit.
You wrote, “Her aunts, daughters of the old queen, initially ignore her”
Surely these are also her sisters, since they have the same mother? It’s just that they’re different ‘types’ of daughter, worker/queen. Eventually as she lays her eggs her own offspring, daughters will run the hive.
I have already shared the link on Youtube with my association as it is totally amazing!
Thanks so much for sharing that with us!!
sorry I didn’t see that it had posted my comment! disregard this post.
No worries, a good comment is worth posting twice!
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Thanks for another educational post on bees.
The mystery of how knowledge is being passed on location of where drones and Queen will meet to mate intrigues me.
I would like to solve that mystery too. It seems the drones choose the same areas each year to congregate and the queen somehow know they are there. Perhaps the area has a scent or something that attracts them, and perhaps the queens are attracted by a scent of the drones? No one really knows.
I think it’s an instinct for drones and queens to fly to certain areas. The DCAs tend to be over open ground and sheltered from the wind, although some are not. It’s even possible that the earth’s magnetic field is important, as drones develop large quantities of magnetite in their abdomens.
Yes, it could be suitability of location instinctively attracts them and I’ve read something similar about magnetic fields. Although I prefer to believe that bees have secret chronicles (like beekeepers records) and pass the knowledge down through mysterious ceremonies like a DCA dance! 😉
Navigation via magnetic fields is something I have considered of bees, for I find it amazing an individual bee can find their hive again amongst dozens or hundreds from miles away.
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