Almost 30% of Dutch beekeepers work with what are disparagingly called bastard bees. Jokers also refer to them as F-16s because of the supposed aggressiveness of these bees. The impression is created that only those who work with purebred bees have reached a higher beekeeping level, while the ‘bastard beekeepers’ just muddle along. This image does not always correspond with reality. In this article I want to discuss what it means for a beekeeper to work with local bees and local mating. Is it possible to keep bees successfully and pleasantly in this way?
Introduction
In my beekeeping library I have an old book by Wolfgang Golz (whom I mentioned elsewhere in these pages – (see ‘2 x 9 = Golz’) about beekeeping with ‘the’ landrace. In this booklet he advocates beekeeping with bees that do not belong to a known race and also open mating.
According to Wikipedia, a landrace is a dynamic population of plants or animals that is maintained on site without significant selection and has adapted to its environment over the years. The central question in this article is whether such a landrace could form and/or maintain itself in the current Dutch beekeeping landscape.
Open mating is the mating of young queens in an open population, hence with the drones they choose themselves. Is open mating a good idea? This is actually a strange question. The honeybee has survived for about 30 million years via mating in an open population. Apparently it doesn’t work that badly.
In 2015, Kees van Heemert published a fascinating series of review articles in which he discussed a European study that showed that bee colonies of local origin were best adapted to the local situation and therefore had the greatest chance of survival (European bee project: the genetic diversity of bee colonies, see ‘Bijenhouden’ 2015 no. 3, 4, 5 and 6 (in Dutch)). Colonies with a locally bred queen lived on average 83 days longer than colonies with a queen that came from another location. It turned out that the interaction between environment and genotype, rather than the genetic origin of the queens, was the main cause of this. This study was artificial in the sense that the queens studied were in experimental situations. In the reality of beekeeping, in which queens are purchased or in which beekeepers in the area participate in joint queen rearing projects, nothing remains of this local characteristic.
The misconception that pure bred bees are superior lies in the fact that breeding success is often defined in terms of improving a number of characteristics such as honey yield, gentleness and steadiness on the comb, but not vitality. These characteristics are not necessarily linked (Bienefeld et al., 2016).
Selection
We can only call our beekeeping breeding if we sort the material produced in such a way that we only keep the best colonies. In fact, we all perform some sort of selection when we requeen the worst of two colonies and make a nuc from the best. But because we have little or no influence on the paternal side during the local mating process, in almost all cases we cannot achieve real breeding progress for the characteristics that many beekeepers are interested in, such as honey yield and gentleness.
We can influence the paternal side a tiny bit by ensuring that we cut away the drone brood from inferior colonies, in anticipation of their imminent requeening, and allow as many drones as possible to be produced by the good colonies, but even then the effect is very small, since mating takes place at a distance from the apiary, which mainly involves drones from other apiaries. It would help enormously if all beekeepers in the area adopted a similar approach and applied strict criteria with regard to the quality of their colonies.
Strictly speaking, selection also means that we should not unite colonies and not combat Varroa. Colonies should be able to survive on their own, just as is done in the testing of queens in selection programmes.
Furthermore, nature helps out with selection: colonies that are too weak do not survive the winter. They are attacked and robbed by wasps before the winter (this can be seen on the bottom board by the rough wax mullet and dissected workers). Winter losses are indeed annoying, but they are an essential part of natural selection.
Bienefeld modelling study
A 2016 modelling study (Bienefeld et al., 2016) concluded that there is no ‘progress’ to be made by mating in an open population. At the same time, it was found that the absence of controlled mating results in a shift towards maternal genetic influence.
In the long term, breeding schemes with uncontrolled mating will lead to a lower degree of inbreeding and therefore to less loss of genetic variation than schemes in which mating is controlled. On the other hand, there are breeding schemes available that can minimize such loss.
Selection programs for honeybees are often defined by quantitative traits, such as honey production or gentleness, but, as mentioned above, it seems that these traits have little or no influence on the fitness of a colony. Incidentally, it is difficult to define what fitness actually means.
Female dominance?
At the time, Golz believed that inheritance was due to maternal genetic dominance, which would naturally be very convenient for a stationary beekeeper! However, this reasoning doesn’t hold true for nuclear DNA: a worker bee receives half of its genes from its mother and half from its father. However, the matter is a bit more complex. Two things need to be considered: mitochondrial DNA and the microbiome.
Mitochondrial DNA, or mtDNA, is a small, ring-shaped DNA found not in the nucleus, but in the mitochondria, which are found in every cell and function as energy production centres for the cell’s function. Mitochondrial DNA is normally inherited exclusively through the female line in both animals and plants, and therefore not passed on with sperm or pollen. In animals, the mitochondria of the male sperm cell are located in the tail, which usually does not enter the egg during fertilisation. The influence of mitochondrial inheritance is not yet clear, but it is known, for example, that certain human conditions are passed on through mitochondrial inheritance, and therefore through the mother. So perhaps Golz was right after all…
By microbiome we mean symbiotic bacteria, the micro-organisms that live together with and within the bees and the colony, and which play important roles in the metabolism of the entire colony.
We know that the microbiome plays a significant role in the identity of a bee colony, including the mutual recognition of its members. Bees consider bees with a different bacterial strain to be “foreign” and refuse them access to the hive. When the primary swarm leaves, this nest odor persists in the main colony.
According to Blacquière (Blacquière, 2015), the DNA of the queen and drones together represents only 10% of the hereditary information in a bee colony. Perhaps 90% is provided by the parasites, yeasts, bacteria, archaea, fungi, and viruses in that colony. All of these together influence what happens in a bee colony.
To determine the strongest colony, we must consider not only the genes of the queen, drones, and workers, but all the genes present within the colony. This also applies to selection and breeding: a chosen colony is not just a chosen queen, but a chosen whole.
Morphological or behavioral characteristics?
Open mating is at the heart of the search for a new landrace. The morphological and genetic characterization of varieties isn’t decisive in the eyes of the stationary beekeeper. Behavioral traits, on the other hand, are certainly given considerable weight. With all these open matings, all sorts of new gene combinations arise—good ones, but also, of course, very bad ones. Therefore, selection must continue, generation after generation, without much hope of “improvement,” but with the maintenance of vitality. Incidentally, even in selection programs, the work is never finished. As soon as selection stops, improvement quickly declines. Breeding programs assume that growth of a trait is always possible. But should the honey yield per colony or the size of a colony always increase or expand, or can a balance be defined somewhere between the environmental carrying capacity and the number of interventions the beekeeper must perform to maintain the colony?
Breeding through open mating is actually an expression of the conviction of stationary beekeepers that not everything can be managed and controlled. They let nature take its course and select on-site what is best adapted to the local situation. Whether this will endure in subsequent generations is, unfortunately, very questionable. If everyone follows this approach with minimal intervention, allowing swarms (or only creating artificial swarms when closed queen cells are present in the colony), feeding, and pest control, an equilibrium will be created. This is rewilding, or beekeeping in an organic way.
Open mating
We all know how much effort and energy goes into the logistics of mating unfertilized queens in controlled mating situations, at land or island mating stations. This effort can certainly be saved with open mating. Here, mating takes place in an open population. Stationary beekeepers often don’t distinguish between breeding and propagation, since a large proportion of the colonies participate in the selection process.
In normal situations, we don’t need to worry about inbreeding at all. Generally, there are sufficient colonies within a radius of about 10 km to produce enough drones. If you’re the only beekeeper within a radius of, say, 10 kilometers year-round, inbreeding could eventually occur if you consistently bred from the same queen, both on the mother’s and father’s side. However, this is purely hypothetical in the Dutch situation.
By mating with many different drones in an open mating setting, the virgin queen fulfills her natural urge for genetic diversity. We know that honeybees are very sensitive to inbreeding. This, and the fact that only the most vital drones will succeed in fertilizing the queen, creates the foundation for the vitality of the colonies.
F1
The term is often used—but not necessarily!—to refer to breeding purebred queens in their own apiary. These queens produce purebred drones, but the workers are crossbreds. This is sometimes called the F1 method.
During the 2021 queen rearing webinar, Marie José Duchateau advocated for beekeeping with F1 colonies, which are genetically homogeneous if the parents are purebred. The F1 generation is gentle and productive, ideally. However, if you continue breeding them in an open mating setting —the F2 generation and beyond—increased defensiveness occurs. She argued that this stems from the mismatch between the various, genetically distinct, half-sister groups in a colony. Half-sisters, because a young queen mates with multiple, unrelated drones during open mating. The resulting workers are therefore related only through their mother’s side. The heterosis effect we find in the F1 generation fades away with continued open mating in subsequent generations. Therefore, you must continually create the F1 generation, otherwise you will face the negative consequences. With open mating, of course, there is no F1 generation.
But there’s another side to the story. When multiple, genetically less related, half-sister groups occur within a colony, that colony has a broader behavioral and reaction spectrum. This allows it to respond more flexibly to environmental stimuli. Greater genetic variability within a colony means: more stable temperature regulation, better nest defense, better division of labour, and lower susceptibility to disease. In short: the colony is more vital.
In the previously mentioned study by Kees van Heemert, vitality increased as the queen mated with more drones from other colonies. The relationship between half-sisters follows a sliding scale: from one drone to 20 drones, the relationship decreases from 75% to 25%.
Beekeeping with the landrace
The basic principles of breeding with non-purebred bees and open mating are therefore: no mating stations and no selection based on morphological traits. Only selection based on behavioral traits. No distinction is made between breeder and production colonies.
A major advantage, in my view, lies in the fact that a locally adapted bee still performs reasonably well even under suboptimal conditions and doesn’t need to be pampered. What good is a bee that only performs well when it is constantly placed in the midst of abundant forages, or in the absence of such forages, requires constant supplementary feeding, as it is unable to adjust its brood size to temporarily more challenging circumstances?
Under natural conditions, a colony would produce several swarms per year. Through grafting, it is theoretically possible to breed thousands of queens from a single colony. This would lead to a genetic bottleneck in the population. Stationary beekeepers ensure that between a quarter and half of the colonies are used for reproduction. This guarantees diversity.
When Golz described his approach, the beekeeping landscape looked quite different than it does today. He, too, looked back with some nostalgia to the Lüneburg heathland beekeepers. There, basic breeding was the norm, because everyone kept bees in a similar way and with the same species.
In our current fragmented beekeeping landscape, it is virtually impossible to establish and maintain even a relatively homogeneous population. The influence of foreign queens is greater today than in the 1980s. We can undoubtedly demonstrate a significant influence of Buckfast today, in addition to the ubiquitous Carnica genes. All these influences can be clearly visualized by mapping wing geometry and through DNA analysis. Beekeepers who think they’re working with a chosen breed would be amazed at how far their colonies deviate from the breed’s characteristics when they examine the morphology and genome of their purebred bees. Note for Carnica beekeepers: it could very well be that your Carnicas are ‘contaminated’ with Buckfast genes.
We might wish it were otherwise, but this is the reality today. Beekeepers have been dragging queens around for decades, and that’s here to stay.
So, breeding progress is practically zero, if not zero, but how bad is this really? “Nature is patient, now it’s our turn,” as a seed company in the north of the country rightly puts it. And what Dutch beekeeper these days still cares about maximizing honey yield per colony?
It can happen that a particular colony is so inundated with negative external influences (defensive bees, Buckfast) that it’s impossible to establish even one decent colony. At such times, beekeepers with multiple apiaries have a clear advantage. But then again, how many beekeepers would be in this position? It makes you envious, to be sure: multiple apiaries make beekeeping so much easier…
Many beekeepers are very pleased with their expensive purebred queen and try to breed from her for years, sometimes at a mating station. But the reality is somewhat disconcerting: after four generations of breeding such a top queen, approximately 6.25% of her genes are still represented in her descendants. The remaining 93.75% comes from the various drone colonies at a station, or from the drone-producing colonies in the vicinity of the virgin queen. After all, in each generation, the mother contributes 50%, and so does the mother of the drones. If a breeder believes they discover good traits in the descendants of the top queen, this is not simply the result of inheritance from this queen, but of conscious or unconscious selection applied to her offspring. In short, if you want to improve your breed, you’d be better off focusing more on the presence of top drones than buying top queens for a fortune.
Where do we actually stand now?
The problem lies in the patchwork of genetic material buzzing around the Dutch landscape. Within this patchwork, the genetic composition of the population is not stabilized, as the constant infusion of foreign genes repeatedly creates a kind of zero balance. The balance in the gene pool is continually disrupted.
It might work in a situation like the Lüneburg Heath, where everyone kept the same species of bees. But the Dutch beekeeping landscape is very different: everyone does their own thing, and there’s total anarchy in the air and in the apiary. Incidentally, when the Lüneburg beekeeper went out with his colonies, he often brought back a queen from elsewhere. Even a practically closed population can thrive with the measured introduction of new genes.
All in all: if honey production isn’t your primary concern, working with purebred colonies is unnecessary. Stationary beekeeping is the beekeeping method for beekeepers who simply want to keep bees with colonies in their garden, adapted to their environment, where they originated/formed as a swarm. So: open mating and no traveling. The colonies should preferably be spaced apart and not in a row: “Darwinian” beekeeping.
Within your own colonies, you select those colonies that can survive the winter without much intervention, produce sufficient honey for themselves and also a container of honey for the beekeeper, and are manageable. Working with F1s doesn’t fit this picture, because you’re always starting from scratch: no selection for anything. But again, you shouldn’t be under the illusion that you can achieve any kind of breeding progress, because that’s not possible with such a setup. Purchasing an F0 queen and then not taking the daughters to a mating station only increases the hybridization rate of the bee population and that doesn’t benefit anyone.
There are conceivable cases where beekeeping with pure breeds, and therefore controlled mating, is indeed appropriate: commercial beekeepers, who aim for maximum yields, and beekeepers concerned with breed preservation.
All in all, acquiring a new, relatively homogeneous landrace is no longer feasible in our current era. Due to the constant influx of foreign genes, through the purchase of foreign queens by nearby beekeepers and mating with the foreign drones of those purchased queens, the population cannot stabilize.
But there really should be more research into the potential evolutionary advantages and disadvantages of hybridized populations. The new allele combinations could be essential for responding to new selective pressures, such as the introduction of all sorts of new parasites and diseases. In that case, it might be advantageous to have so many “foreign” genes floating around.
Further reading
- Bienefeld, K. et al., 2016. Breeding success or genetic diversity in honey bees? Bee World 93(2):40-44.
- Golz, W., 1982. On the way to a new breed. Brochure 5. Zeidel Publishing.
- Blacquière, T., 2015. Resilient bee: Exploration of initiatives and scientific literature on the natural defense of bee colonies against diseases and pests. Report 2015, Project BO-20-003-023 Ministry of Economic Affairs.