Did you know that queen bees have evolved to be adaptable and flexible, and eggs laid in queen cells are larger than those laid in worker cells? Read on to discover the significance. My search for reasons why our queens fail is taking me on a convoluted journey which I share with you today.
English Market, Cork.
The planned adventures on my recent holiday to Cork were curtailed by a nasty respiratory virus which was thankfully not passed on to my host Tricia. We made it to the English market which I find fascinating and always enjoy visiting. What was interesting was the change in jar sizes years since my last visit three years ago. In the past, the stalls have been stacked high with lb jars, but there were only a handful this visit and the rest were ½ lb jars. The price range was variable with one jar retailing at €8.50 (£7.31) per ½ lb and another at €12 (£10.32) per ½ lb.
Biodynamic Farming.
Our next trip should have been to a local farmer’s market where I would have met a biodynamic beekeeper. However, the beekeeper’s wife kindly sent an article for me from the Star & Furrow which is the Biodynamic Association’s newsletter1. Little did she know how helpful this gesture would be in my quest for answers to queen problems. Welsh bee farmer Selwyn Runnett’s thought provoking article addressing the problems of modern beekeeping, including colony and queen failures, piqued my interest greatly. Runnett’s suggestion that Rudolf Steiner’s predictions, that artificial queen rearing and grafting might be harmful to beekeeping in the long-term, might be relevant spurred me on to search for answers myself. I want to share my findings and the latest research on how high-quality queens are produced. But first here is the background to Steiner’s philosophy and beliefs around managing honey bees.
Queen Rearing:Grafting Larvae.
During the first decade of last century, it became fashionable in beekeeping to rear queens by grafting newly hatched worker larvae under the emergency queen rearing impulse. In the US, Doolittle progressed commercial beekeeping through his method of making wax queen cups and grafting larvae from stock with chosen traits to raise queens. It is exactly one hundred years since Rudolf Steiner questioned the effect that this might have on the nature of bees during his eight lectures on honey bees to audiences in Dornach, Switzerland in 19232.
Rudolf Steiner.
To examine this in context we need to review Steiner’s philosophy. Rudolf Joseph Lorenz Steiner was an Austrian born in Croatia in 1861. He was educated in Austria and trained as an architect before going on to spend his working life in Dornach, Switzerland where he died in 1925. During his lifetime, Steiner developed himself through science, the arts and humanities and became a renowned social reformer and philosopher. He was a talented artist, public speaker, and lecturer, and he wrote widely sharing his knowledge in over 40 written works and 2,000 lectures.
Anthroposophy.
Steiner created the anthroposophical movement which is a means of closing the gap between science, the spiritual world, and the arts. Anthroposophy is derived from the Greek language where Anthropos refers to human beings, and Sophia means inherent wisdom. It is not a religion but rather a spiritual philosophy based a belief that that the human intellect has the ability to connect with spiritual worlds. Steiner defines anthroposophy as “a path of knowledge, which intends to lead what is spiritual in the human being to what is spiritual in the universe”. In other words, he believed that we humans are spiritual beings living in a physical existence with the ability to develop the self as individuals as well as transforming society. It explores the deep questions of humanity including our own basic needs for living and also our need to develop a relationship to the wider world in a way that gives us complete freedom to express our individuality.
Practical anthroposophy remains visible today in several fields including the Waldorf/Steiner education system, biodynamic gardening, farming, and beekeeping, and in medicine. It is a pathway to developing a conscious awareness of one’s humanity and self-awareness. Steiner embraced change and he didn’t try to keep things as they traditionally were. He was a scientist with a view that science has a soul and spirit and is not purely mechanistic. Rather than discounting the intellectual achievements of the scientific age he added another spiritual dimension to complement them.
Steiner’s Lectures.
Steiner was not a beekeeper but a researcher and he delivered eight public lectures on bees in 1923. They were attended by a wide audience that included some beekeepers. Among them was Mr Muller who was an experienced beekeeper and also gave lectures and contributed to Steiner’s presentations by asking relevant questions and proffering insights. Mr Muller was interested in rearing queens from grafting larvae into wax cups and a great proponent of this new way forward in beekeeping. Mr Muller didn’t understand why Steiner was against the raising of queens from worker larva which had been practised successfully for 15 years. Steiner explained that raising queens like this would have a detrimental effect on honey bees because the beekeepers were choosing the genetics and selecting for traits, rather than leaving the bees to do this for themselves.
Steiner did not wish to turn the clock back but he wanted beekeepers to have the insight and the freedom to keep bees in a way that was appropriate for them. He wanted bees to be able to regain life forces and to heal from the damage inflicted on them by humans. He told his audiences that it did not hurt to be aware and conscious of the fact that by introducing a mechanical, artificial, element such as grafting worker larvae, we are actually disturbing that which nature has produced in a such a wonderful manner. He believed that artificial bee breeding was disruptive to the strong generational bond in a “bee family” and that it would have a detrimental affect over the longer period. Queen rearing by grafting newly hatched worker larvae means that queens are always raised by workers under emergency situations. The beekeepers rather than the bees select for traits that they choose rather than it being a colony choice. Referring to the great changes in beekeeping during last century, Steiner acknowledges “that beekeepers can indeed be very happy with all the progress that beekeeping has recently experienced in such a short time, but this happiness will barely continue for one hundred years.”
Review in a Hundred Years!
Steiner tells Mr. Muller that they will have to wait to see how things are in 50-80 years but that it will not be possible to establish the intimate relationship between the queen that has just been purchased and the worker bees in the way it would have arisen by itself in nature. He explains that at the beginning, the effects on queen breeding would not be apparent, and he says to Mr. Muller, “Let’s talk to each other again in one hundred years, Mr. Muller, then we’ll see what kind of opinion you’ll have at that point.”
Have these predictions of Steiner’s come to pass? Steiner’s critics might be sceptical of his lack of beekeeping knowledge, and his independent citizen science research, but nobody will dispute that modern conventional beekeeping is not going so well one hundred years on from the Steiner lectures. One just needs to study colony loss figures and listen to complaints of queen failures across beekeeping globally to know that something, or many things, are fundamentally wrong at present. We are fortunate that scientists today are onto the case and investigating many aspects of this conundrum, including what good quality queens are, and how best they should be raised.
Raising High-Quality Queens.
Yu et al3 experimented with three different ways of raising queens. They acknowledge that artificial queen rearing is crucial to commercial beekeeping in order to regularly expand colonies, minimise swarming, enhance brood and honey production, increase colony numbers, and improve genetic characteristics. They also recognise that inappropriate use of artificial queen rearing techniques contributes to the decline of queens and colonies.
Qualities.
What are high-quality queens? They are usually raised in a longer queen cell and have more left-over royal jelly in the cell after emergence. They have a higher weight, larger thorax and ovaries, and more ovarioles. A high-quality mated queen has a strong egg-laying ability, more stored sperm, and higher sperm viability. She produces more pheromones which improves colony cohesion and growth in contrast to a lesser-quality queen that produces fewer pheromones resulting in poor colony labour and low production of products.
Reproductive Genes & Hormones.
Developmental and reproductive-related genes and hormones are important and are expressed at a higher level in high-quality queens. These include; hexamerin, vitellogenin, transferrin, juvenile hormone, abeacin, defensin, hymenteaicn, and insect specific cyt P450. Measurements of these traits can be used to assess the quality of queens.
The scientists explain that the development of queens in artificial queen rearing is influenced by the age of the grafted larvae and maternal effects. Maternal effects relate to the fact that queens can lay larger eggs in queen cells which results in high-quality offspring because they are much larger than eggs laid in worker cells. This might come as a surprise to learn that honey bee queens can vary egg size in response to genetic and environmental conditions. This is well documented4 however, and some queen ants and female mallard ducks do the same thing in laying selectively larger eggs. The mallard duck lays larger eggs if she has mated with a preferred male. If you have ever watched mallards during mating season you will know that often the female has more attention from drakes than she wants. Many drakes may mate with her and she can be held under the water for so long that she drowns.
For honey bees, this adaptive behaviour could result in significant differences between queens raised from eggs laid in queen cells and those raised from eggs laid in worker cells. We know that the quality of artificially reared queens decreases as larval age increases.
Queen-Raising Experiment.
In the experiments3, three different methods of queen raising were studied and some of the queens were later dissected and measured. One group comprised queens raised from eggs laid in queen cells (QE), the second was eggs laid in worker cells (WE), and the third group comprised queens raised from 2-day old larvae (2-L). They also compared the first and second generations offspring from each group.
The results show that the highest quality queens were raised from the first group, QE. Interestingly the quality of queens raised by WE were higher than those raised from worker larvae, 2-L. So, a colony prefers to raise queens from larger eggs laid in queen cells. If left to their own devices, under emergency situations, workers will raise queens from worker eggs rather than larvae. So, does this mean that we should be transferring eggs in their cells to queen cups?
The Findings.
The authors reviewed the size of cell space for development. They concluded that a larger queen cell allowed for a larger queen to develop, as demonstrated in the offspring from the QE group, that had the longest cells and the heaviest queens with wider thoraces. Residual royal jelly was higher for this group and both generations of offspring had the highest newly emerged weight and expression of developmental, immune system, and detoxifying genes. Interestingly, the second generation of the lowest quality group 2-L were even lower in quality than in their first-generation offspring.
Conclusion.
These findings provide new information which can be of practical help to beekeepers involved with queen rearing. We already know that the age of grafted larvae is important along with good nutrition at colony level for the production of quality queens. However, knowing what we do now about how honey bees produce the best queens themselves; we might decide to allow our colonies to requeen naturally, or we might look at better ways to artificially raise queens by transferring eggs into queen cells.
I hope you enjoyed reading this as much as I have done finding and examining the evidence. Thank you to my friends; Tricia in Cork, Maria in Northampton, Marie-Helene in Quebec, Canada, and Jenny in Forres for supplying references and inspiring the production of this blog.
Happy festive season to you all and thank you for supporting the blog.
References:
1Runnett, S. (2023) The Crisis for Honey Bees: Was Rudolf Steiner Right? The Star & Furrow, Journal of the Biodynamic Association, ISSN No: 1472-4634, p 12-13.
2Steiner, R. (1923), Bees: Lectures by Rudolf Steiner, 1998, Anthroposophic Press, Hudson, NY.
3Yu, L., Shi, X., Zeng, Z., Yan, W., & Wu, X. (2022) High-Quality Queens Produce High-Quality Offspring Queens.Insects 2022, 13, 486. https://doi.org/10.3390/insects 13050486.
4Amiri, E., Le, K., Melendez, C., Strand, M., Tarpy, D., & Rueppell, O. (2020) Egg-Size Plasticity in Apis Mellifera: Honey Bee Queen Alters Egg Size in Response to Both Genetic & Environmental Factors. Journal of Evolutionary Biology, 2020, DOI: 10.1111/jeb.13589.
Now that (amongst many, many other things) I did not know. For years I have been saying that the only difference between producing a queen from a fertilized egg, instead of a worker, was how the egg was treated after laying. I can see how it would be beneficial to start a queen from a larger egg, and I can see how the queen could distinguish a queen cell from a worker cell at the time of laying, but what is puzzling is how she can have a larger egg instantly available in her oviduct when she arrives at a queen cell. I know – read the paper (which I will). It seems to imply that either she scouts out the queen cell first and comes back to it with an appropriate egg, or she pauses at the queen cell long enough to produce an appropriate egg, or she produces an appropriate egg and then goes looking for a queen cell to put it in, or that she already has a proportion of larger eggs in her oviduct and can select one of them as required, or that she already has a proportion of larger eggs in he oviduct and some of them happen to get laid in queen cells – which would mean that some worker cells would have larger eggs in them (and become larger workers?) I suppose the last is the most likely. Or am I missing something here? They never cease to amaze. I don’t raise queens ‘artificially’ so it’s all just happening in my colonies – except, as you say, when there’s an emergency. It’s an explanation for smaller queens in addition to the cell size. Or is it the other way round? Does the cell size reflect that the queen inside it was already smaller? And does the queen make ‘mistakes’ by not always laying a larger egg in a queen cell? Which brings us back to the possibility that she doesn’t ‘deliberately’ match the larger egg size to laying it in a queen cell but some proportion of the eggs in her oviduct are larger and some of those happen to be laid in a queen cell? … etc… got to read that paper. Thank you Ann (again).
Hello Avery. Thank you for responding to this and for sharing your thoughts.It’s great to see such interest raised through this topic. The size of the cell in terms of length is key and is the adaptable factor. I imagine that this is controlled by the size of the larvae and the workers will keep building on to the end of the cell as she is grows. I discovered recently, from one of Professor Jeff Pettis’ presentations,that a very long queen cell often results from the larvae detaching from the bed of royal jelly and dropping down, so I presume the workers just keep adding wax to the end of the cell as she slips further down. I have had a couple of these very long cells and early on got excited thinking that I would see a massive queen emerge. However, the queen did not emerge around day 16 and when I opened the cell a week later it was a gooey mass of decaying larva.
I attended a Rudolf Steiner School. Did a Class Project on Honey Bees. Ended up having x4 Hives. And sending a ‘Note’ to Brother Adam. He Wrote back !
I was only a x14yrs old School Kid. . . Learning about ‘Bees’.
Four decades later, I know the importance of Brother Adam . And now Rudolf Steiner (!)
Bees are still very important in my Life.
(And those earlier experiences have an effect on today too.)
I will continue letting the Hives produce ‘Queens’ and it won’t be via the ‘Grafting’ process. Bees know best. 😎
Happy Beekeeping 2024 🐝
Interesting to hear about your link to Rudolph Steiner and Brother Adam.
Fascinating – larger eggs laid in a queen cell. We moved away from grafting as too many of the queens were superseded; our interest was in fitness and survivorship. Now, we only use naturally produced queens which has worked really well. Ambitions to mass produce queens reduces the quality – no surprises there. Too much of a disconnect with nature.
Thank you Ann for the research you surface for the bee community. Merry Christmas to you and your family.
Steve
Hello Steve. I thought of you and your colleagues when I was writing this and figured that you were ahead of the curve with this. Thank you for responding and sharing your experiences. Also thank you for your good wishes. Same to you, and all the best to Westerham Beekeepers for the new season. Ann.
This is a sort-of answer to the issues I was puzzled by. For my fellow geeks out there, as regards egg size, Reference no 4 above refers back to:
Wei et al., 2019, Current Biology 29, 2208–2213: July 8, 2019
https://doi.org/10.1016/j.cub.2019.05.059
This is what it says about egg size:
“We do not here propose that there is a special class of queen-destined eggs. The distribution of egg masses sampled from queen and worker cells was continuous, normal, and unimodal. Rather, we propose that fecund queens at any one time have more than one egg ready for laying [10] and that queens may lay the largest available egg in queen cells. Alternatively, queens may pause oviposition prior to laying in queen cells, since delaying oviposition causes bigger eggs with more yolk protein [37], but this possibility needs to be investigated.
An important inference of our data, however, is that queens can actively select larger fertilized eggs for oviposition in queen cells. It has been demonstrated previously that queens can control and withhold fertilization of eggs prior to laying in male (haploid) drone cells and that queens measure the larger drone cell with their foreleg prior to laying [38]. This is the first evidence that queens can select among fertilized eggs and that they differentiate between queen and worker cells.”
Is there any scientific paper that doesn’t qualify itself with some version of ‘further investigation is necessary’? The researchers then are suggesting either the ‘choose the biggest egg’ option, or the ‘hang around until I produce a bigger egg’ option. Still – however the trick is done – it certainly seems to be that queens lay larger eggs in queen cells. As Steve says: fascinating.
Thank you, Avery, for taking this further and clarifying for readers what is known now about how the egg gets bigger. I know that I thought they all came down from the ovarioles the same size before I read these papers. I doubt that all will be known about honey bees at the end of another hundred years!
Hi Ann. Sorry to take over the thread a bit. Yes, we (by which I mean they) can measure so many things, but the behavior that goes with them is much harder to know. ‘Further research is required.’