Category Archives: Science

Dancing in the City

Beekeeping is an increasingly popular pastime. Since ~84% of the UK population live in urban areas (up from ~70% in 1950’s) it is not unsurprising that the number of hives kept in cities is increasing.

Of course, not everyone who lives in a town or city keeps their bees in the back garden. When I lived in the Midlands I lived on a small estate that was indisputably ‘urban’, although there was farmland within sight 1. My bees were on the nearby farmland and I just kept a few mating nucs and a bait hive in the back garden 2

Hive in a field margin

I kept my bees in the farmland because 3 I reasoned that there were both larger amounts and a greater range of forage available for them there.

But I was probably wrong.

It wouldn’t be the first time, and it certainly won’t be the last.

Too many bees?

Before discussing urban bees and forage in more detail I’ll digress a minute to mention the suggestions that the inexorable rise and rise of urban beekeeping is threatening our native pollinators.

Actually … more than suggestions.

There are a number of scientific reports and reviews that indicate that urban beekeeping harms – by outcompeting – native pollinators like solitary bees. A recent report by the Royal Botanic Gardens at Kew states:

‘Campaigns encouraging people to save bees have resulted in an unsustainable proliferation in urban beekeeping. This approach only saves one species of bee, the honeybee, with no regard for how honeybees interact with other, native species.’

‘In some places, such as London, so many people have established urban hives that the honeybee populations are threatening other bee species.’

Our bees (Apis mellifera) are generalists. They are not particularly well adapted to any flower or nectar/pollen source.

They are equal opportunists.

Individually, there are many solitary bee species or non-hymenopteran pollinators, that are ‘better’ adapted. They pollinate more efficiently, or collect more nectar, faster.

But honey bees arrive in the environment mob handed.

Thousands of them.

Actually, tens or hundreds of thousands of them if there are several hives in an apiary. They are a formidable force and can easily outcompete other pollinators that are either solitary or only live in small colonies.

Save the bees, save humanity

When you see the phrase “Save the bees” what it usually means is save the honey bees.

Save the bees ...

Save the bees …

What it should be encouraging is “Save anything but the honey bees, because they don’t need saving … actually, there are possibly too many of them altogether”.

But that’s a lot less catchy … and it won’t let the supermarket, or food manufacturer or whatever, illustrate their campaign with cute photos of pollen-laden honey bees returning to quaint white-painted WBC hives in some sort of idyllic rural scene 4.

I suppose a photo of a honey bee would be better than drawing of a wasp though … which is what the NRDC used for a (mis)information campaign on CCD (colony collapse disorder) a few years ago.

D’oh!

Anyway … saving the bees is usually “greenwash” or bee-washing as it was termed by MacIvor and Packer in a 2015 paper on bee hotels 5

I’ll return to this topic, and urban beekeeping, later in the winter 6.

The Town mouse bees and the Country mouse bees

Where was I?

Oh yes, my – as will shortly be clear – incorrect assumption that country bees have access to more diverse and richer sources of forage than their poor relatives living in the town.

There are many sorts of countryside and many sorts of urban environments.

A hive in an intensively farmed arable landscape could be located in hundreds of acres of wheat fields, where all of the hedgerows were grubbed up years ago and replaced – if they weren’t just removed altogether – with barbed wire 7

How different is that environment to the ‘concrete jungle’ of a modern city? 

Tokyo

Surely that must be a terrible environment for bees?

In contrast, the suburban sprawl that surrounds most cities is possibly a good place for bees to live. Lots of neat little gardens, each 8 with a profusion of flowering plants, each chosen to provide vibrant colour for a much longer period than native plants.

And I’m sure we can all think of forage-rich rural environments. Here the bees gorge on early crocus, then gorse, willow, oil seed rape, clover, lime, blackberry, fireweed and himalayan balsam, before finishing the season will full crops and corbiculae from the ivy.

Now, in a recent publication 9 scientists have compared the forage available to town and country bees, and the results are quite surprising.

Let the bees tell you

If you live in a country with enlightened right to roam laws (like Scotland) you could wander around the countryside recording all the forage available to bees.

But the laws in England are much less enlightened. However, your right to roam in either country does not extend to the land around a private house or building. 

So, although you might be able to determine the forage available in the Scottish countryside, you can’t be certain you would have good enough access to do the same in England. And in a city you could only map what forage was available by peering over fences from public roads. 

So Elli Leadbetter and colleagues let the bees do the work.

They established 20 observation hives. Ten were in the the centre of London and 10 in agricultural land to the North East and South West of London. The hives were 5 km from each other to avoid overlapping areas of forage. They used observation hives so that they could watch and record the waggle dances of foragers in the hives.

I’ve discussed the waggle dance before. It is used to communicate three important pieces of information about a forage source:

  • direction
  • distance
  • quality

The first two bits of information are encoded in the angle of the waggle run to the perpendicular and the duration of the waggle run. The quality is conveyed by the number of circuits a dancing working performs. It’s a case of ‘the more, the better’ … energetically higher quality resources 10 result in more circuits.

Having recorded thousands of these waggle dances, they used the direction and distance information to ‘map’ where the bees were foraging.

GIS data, land use and foraging bees

For many locations there exists a wealth of land use data (GIS data; Geographic Information Systems). Much of this is at high resolution. For each of the 20 observation hives they produced a map of land use within 2.5 km of the hive at a resolution of 25 m. 

Land use was defined in broad categories such as 11 buildings, woodland, arable, pasture, fruit, OSR (all in agricultural areas) and dense or sparse residential, parks, amenity grassland or railways (all in urban landscapes). 

They then used some clever mathematics to decode the waggle dances 12 to work out where the bees had been, converting the distance and direction components to geographic locations.

Urban (top) and rural foraging heat maps, overlaid on GIS land use maps (5 km diameter)

These locations were overlaid on the land use maps to produce ‘heat maps’ showing where the bees were foraging. The image above shows these heat maps. In the spring the urban bees (top left) were foraging intensively to the east and west of the hive and the rural bees (bottom left) were mainly foraging in two large areas to the south east and south west of the hive.

Foraging distance and nectar quality

Even a cursory look at the image above shows that the urban bees tended to forage closer to the hive than the rural bees. But remember, that’s just three snapshots during the season.

Waggle run duration – rural bees fly further

However, more detailed analysis confirmed that this was the case. Throughout the season, the bees in the agricultural landscape foraged further from the hive. I’m showing the log-transformed median waggle run duration (above) as this allows slightly easier comparison across the season. The further the wiggly ‘best fit’ line is above the horizontal axis, the longer the duration of the waggle dance run, and so the further the bees are flying to find the forage.

Interestingly, the median foraging distances were relatively short when compared with the maximum foraging distances from the decoded waggle dances. This applied wherever the bees lived. For urban bees the median was 492 m (max 9375 m) and for agricultural bees it was 743 m (max 8158 m 13 ).

Perhaps the agricultural bees were flying further because there was better quality nectar available at more distant sites?

Nectar concentration (% w/w) sampled from returning workers

They controlled for this by recovering returning foragers and robbing them of their nectar load before analysing the sugar content. There was no significant difference 14 between the nectar from agricultural or urban landscapes. The sugar content of the nectar was recorded as reducing through the season.

Foraging preferences

So where did the town bees and the country bees prefer to forage?

City rooftop bees

City rooftop bees …

In urban areas the bees exhibited a strong preference for residential gardens (the ‘sparse residential land’) … these are presumably the flower-rich urban gardens that the homeowners also tend to prefer.

In contrast, bees in an agricultural landscape showed an entirely unsurprising reliance upon mass-flowering crops like oil seed rape (in spring only). They also showed weaker preferences for arable land and fruit crops throughout the season.

Mid-April in the apiary ...

Mid-April in a Warwickshire apiary …

I’ve skipped over a host of additional observations from the study, and almost all of the controls. Two things that are worth mentioning though.

Firstly, hive strength had no influence on waggle dance duration (and hence foraging distance). It therefore wasn’t the case that stronger hives had a larger workforce that could survey and exploit more distant forage. 

Secondly, cities are warmer due to the urban heat island effect. However, temperature also did not affect waggle dance duration when it was factored in. So, the city bees aren’t foraging at shorter distances because the dance is truncated at higher temperatures.

Conclusions

So, although cities are predominantly filled with buildings and roads, city bees travel less far to forage when compared to bees in agricultural landscapes.

This strongly suggests that the urban landscape consistently provided more available forage 15

Conversely, the bees in agricultural landscapes had to fly further, not because there were better quality nectar sources available at long-range, but because there wasn’t enough nectar nearby.

There were a number of additional interesting points in the paper, some of which were known already from other studies. For example, the high sugar content in spring nectar was already known (and confirmed here). Similarly, foraging distances in midsummer were longer than those in spring or autumn. This could be predicted due to the reduced rainfall in summer, and consequently the reduced overall level of nectar available.

I need to think more about how this study contributes – if at all – to the ‘too many bees in cities’ argument. If anything, forage-rich towns should be able to support a greater number of bees 16 without the honey bees impacting on other species. In contrast, honey bees in agricultural landscapes might dominate the available nectar sources because they can forage at long distances and then communicate to their nestmates the location.

Perhaps it just shows that that heavily farmed land is actually very poor in terms of nectar availability? It’s either boom or bust … once the OSR has finished, or the clover has been ploughed in, or the fruit trees stop flowering then there’s nothing left 🙁

It would be interesting to conduct a similar study in a forage-rich, non-agricultural, rural landscape.

Access all areas

Finally, I think a particularly neat thing about this study is the use of bees to ‘map’ the forage availability using what the authors term “a large-scale search effort that has no access limitations”. The scientists interpreted the waggle dance to get information that would otherwise have been difficult or impossible to determine.

However, using the bee’s own perception of the distances flown might actually lead to inaccuracies in the calculations. As discussed previously, bees measure distance by optic flow. Optic flow increases in complex landscapes … and cities are likely (at least to us, and certainly when compared with arable farmland, to bees) to be complex landscapes. Increased optic flow leads to a perception of increased distance, and hence to longer waggle runs.

This means that bees in complex landscapes might overestimate the distance they have actually flown to forage. Conversely, those in uniform landscapes might underestimate.

Which means that the results of this study may be a conservative estimate of the differences in foraging distance.

And therefore a conservative estimate of the differences in forage availability in urban and agricultural landscapes.


Notes

Dancing in the City was a pretty cheesy song that reached #3 in the charts for the rock-pop duo Marshall Hain in 1978. Having remembered the track when thinking up a title for this post, I made the mistake of listening to it on Spotify.

I now can’t get the damned thing out of my head. 

But it gets worse. I rummaged through Wikipedia and discovered that Kit Hain, the vocalist, subsequently had a very successful songwriting career with people like Roger Daltrey, Chaka Khan and Fleetwood Mac. Impressive. 

In contrast, Julian Marshall, the keyboard player became a member of the Flying Lizards who had a minor hit with a cover version of Barrett Strong’s 17 Money (That’s What I Want)

… and now I can’t get that out of my head 🙁

Counting by numbers

Can bees count?

If they can count, what’s the highest number they can count to?

Do they understand the concept of ‘more than’ or ‘less than’? And what about zero? These are very much more complex concepts than simple numbers.

If bees have simple counting skills, can they do mathematics? Addition, subtraction, multiplication and division?

What about more complicated maths like differential equations or Fourier transforms? 1

And finally, if we can determine the answer to all these questions, do we know how bees count? Do they simply see things and think “Ah Ha! Two flowers each with four petals”, or do they count in a non-numerical way?

Numbers, arithmetic, more than, less than, zero etc. are complicated concepts that involve aspects of neuroscience, psychology and even philosophy. I’m a molecular biologist/virologist and so singularly ill-equipped to understand the more esoteric or advanced aspects of these topics.

Don’t say I didn’t warn you 😉

Can bees count?

Yes.

Bees, like many other animals have basic counting skills.

In studies going back 25 years, Lars Chittka 2 demonstrated that bees exhibited what he called protocounting. He positioned a feeder between distinctive landmarks (tents in a field) and showed – by altering the number of tents between the hive and the feeder – that the ability of the bees to find the feeder was at least partly due to their ability to count the landmarks.

Counting by bees

If he added more tents between the feeder 3 and the hive, the bees flew a shorter distance to where they thought the food supply was. If he removed some of the landmarks, the bees flew further.

Chittka used the term protocounting because he reasoned that the things that the bees were counting – in this case tents in a row in a field – may not be transferrable to different objects. For example, bees trained to a feeder between tents 3 and 4 in a series might not return to the same position in a series of trees in a row. He didn’t test this, but others now have.

Bees also measure distance flown and this had a greater influence on where the bees searched in the series of landmarks. Nevertheless, these early studies provided compelling evidence that bees had some basic counting skills … or numerosity as it is called.

Is this ‘just’ protocounting and how high can they count?

To understand numerosity in more detail scientists moved the experiments into a laboratory setting where they could control the environment much more tightly.

Landmark counting in a laboratory flight tunnel

These use uniform flight tunnels containing painted or taped on landmarks. For some of the detail read the legend in the figure above 4. By using flight tunnels they could change the landmarks – see (a) above – from stripes to dots to baffles. In doing so they demonstrated that the bees were counting landmarks per se, rather than a particular type of landmark.

For example, bees trained to search for food between the third and fourth set of stripes would look between the third and fourth set of dots. This is much more advanced than the protocounting proposed in the initial study by Chittka and colleagues. … it shows a degree of abstract numerosity.

The use of baffles prevented the bees ‘looking ahead’ (or behind) to visualise the entire set of landmarks together … they had to keep a running total in their heads.

Finally, the highly controlled environment of a flight tunnel enabled the researchers to determine that bees could count up to … wait for it … four.

Not a huge number but, in terms of cognition, a significant advance on none, one or more.

But let’s not get ahead of ourselves.

We’re going to tackle the concept of zero and ‘more than’ in a minute.

It has been argued that the upper limit of four – a limit shared with a number of other animals – might be a consequence of the small size of the bee’s brain 5 and the small percentage of the neurones in the brain being dedicated to counting 6.

However, it may represent a more fundamental limit of cognition and perception that reflects the capacity limit of working memory. This isn’t restricted to bees … it’s something shared with beekeepers. For more detail on this I recommend Nelson Cowan’s The magical number 4 in short-term memory: A reconsideration of mental storage capacity. 7

Go on … knock yourself out 😉

More or less … and zero is not a number

Actually, it is.

What’s more, it’s an even number … 0, 2, 4, 6, 8 are the first five even numbers from zero.

It’s the ’empty set’, smaller than one.

It’s also, conceptually, a difficult number to grasp. Children take much longer to understand that zero is a number (e.g. that they can do maths with) than they do to understand that zero means “nothing”.

There are four steps defined in the acquisition of the understanding of zero in human history, psychology, animal cognition and neurophysiology” 8, which are:

  1. defining zero as ‘nothing’
  2. the categorical definition of zero as ‘nothing’ versus ‘something’
  3. an appreciation that zero is a quantity at the low end of the positive integer numerical continuum (see above)
  4. the symbolic representation of zero i.e. 0, the Arabic number for use in maths

Can bees distinguish ‘more than’ or ‘less than’? And what about zero being ‘less than’ one?

The experiments to determine this were simple but elegant (as all the best experiments are 9) and involved training and rewarding bees to select targets that displayed more items or fewer items.

Bees trained to select ‘more than’ targets would choose one displaying four items over one with two or three. Similarly, the ‘less than’ trainees 10 would select two items rather than three or four.

More than, less than and zero

These experiments confirmed that bees ‘understood’ the concepts of more than or less than. The experiments were controlled to exclude the possibility that the bees were responding to other stimuli that changed in the target e.g. signal density or colour intensity.

And if that wasn’t enough 11 they also showed that the ‘less than’ trainees would choose an empty target when offered the choice between ‘some’ and ‘none’.

But there’s more … or less

The ‘popularity’ of the empty set – zero items – was directly related to the difference in the number of items displayed on the non-empty targets. For example, bees more frequently chose the zero option when numbers were numerically more distant (0 vs. 5 or 0 vs. 6) than those closer together (0 vs. 1 or 0 vs. 2).

This relationship between accuracy and numerical distance is called Weber’s law.

This demonstrates a significant degree of understanding of the place of zero in a numerical continuum, an extraordinary feat in a honey bee.

This appreciation of the concept of zero as a number is similar in ability to that shown by African grey parrots, non-human primates … and pre-school children.

That’s impressive.

Counting by numbers

These numerical skills, amazing as they are 12 demonstrate an appreciation of numbers, but not the ability to manipulate numbers.

Can bees count?

For example, could bees solve this testing little equation?

Uh oh ... swarming ...

Maxwell’s equation – the foundation of classical magnetism, optics and electric circuits … but you knew that already

OK, not so fast … is that really a fair test?

These coupled partial differential equations are quite challenging. As bees have yet to be trained to hold a piece of chalk, they can’t write QED 13 at the bottom of the blackboard to indicate they’ve solved it.

So what about these instead?

1 + 1 = 2

3 – 1 = 2

To test this Adrian Dyer 14 and colleagues substituted colours for mathematical operands during the training e.g. blue for addition, yellow for subtraction, coupled with shapes for the numbers.

Y shaped maze to determine if bees can do maths

They trained the bees using a classical reward-punishment system. This sounds worse than it is. It means they received delicious syrup 15 when they got the right answer, but bitter-tasting quinine when they got the wrong answer.

The more training the bees received, the better they got at maths – both addition and subtraction. Once trained, the bees were tested.

Honey bees can do basic maths

In this subsequent testing phase the bees got the correct answer about 75% of the time. If they had been choosing the answer at random they would have achieved only ~50% of correct answers.

So, if more than, less than and an appreciation of zero aren’t enough, bees can also do mathematics.

Though probably not partial differential equations 😉

How do bees count?

It’s not yet clear how bees actually count objects – like the shapes scientists train them to recognise in the Y shaped maze shown further up this post.

Numerical cognition is considered a higher cognitive activity. In humans we’ve abstracted the process and – for at least the last 1100 years – used Arabic numbers to indicate quantities of ‘stuff’.

Of course, humans could count long before this, as could the apes from which we are descended. But they don’t use Arabic numerals and nor do bees.

3 + 1 = 4

Humans can look at a diagram like the one above and immediately recognise and count the similar shapes – this is a process called subitizing and is restricted to small numbers of shapes. You can train computers to do the same thing using neural networks, object detection and counting the instances of the objects detected.

However this is both computationally expensive and exploits symbolic mathematics, neither of which are achievable in the tiny brain of a honey bee. They cannot inspect a complete field of view and conclude “four apples”, or “three black circles”.

Bees cannot subitize … their brains don’t work fast enough and their visual acuity, and compound eyes, are probably incompatible with the process.

Instead, recent modelling studies have suggested that bees might count objects by serial sequential processing.

If you observe honey bees traversing a Y shaped maze they closely inspect each and every element in the patterns they are presented with.

Simple neural model for counting in honey bees

Using just four neurones, this method appears to allow the bee to achieve the known capabilities of their numerosity i.e. counting to four, concepts of more than or less than, understanding zero, and the fulfilment of Weber’s Law.

This area of the science is even more outside my comfort zone than Maxwell’s equation (see above) and I recommend you read the article cited in the legend to the figure above for further details.

Unanswered questions

There’s clearly a lot more to understand about the numerical and mathematical abilities of honey bees. This will take time, but good progress has been made over the last decade or so.

A full understanding of the neuronal processing involved in object discrimination and counting will probably require an ability to determine which neurones are firing, and the pathways of neuronal communication.

Scientists are years away from being able to do this with honey bees. It necessitates the ability to introduce specific reporter genes (for example that turn green when a neurone fires) into individual cell lineages. We don’t have the tools to do this (yet), but it can be done with fruit flies (Drosophila) and some of the same methods do – or might – work in bees.

In the meantime it’s worth thinking about what the ability to count contributes to the biology and behaviour of bees.

Do plants with more petals have more nectaries, and do bees follow these numerical clues rather than – or in addition to – other visual or scent stimuli?

And why on earth do bees have the ability to do basic maths?

I can’t think of an advantage that this would confer to bees.

Can you?


Notes

The title of this post was going to be 0 … 1 … 2 … 3 … 4 … more’ but that would have looked terrible as a URL and would have sunk without trace in Google searches.

Instead I’ve used a play on the title of the 1988 Peter Greenaway film Drowning by Numbers. This has a fairy tale-like plot that involves three generations of women – all called Cissie Colpitts – drowning their husbands. Bernard Hill plays Madgett, the coroner encouraged to cover up the crimes and involves a series of invented games, one of which is called ‘Bees in the Trees’ (which seemed appropriate).

Drowning by Numbers … going by the number in the background, a scene from about half way through the film

If you’ve not seen the film (and enjoy slightly surreal movies and black comedies) I can recommend it. See if you can count the numbers 1 to 100 that appear, almost all in sequence, either in the background or spoken by the characters. Michael Nyman wrote the music which is similarly highly structured, and based entirely upon a Mozart Sinfonia Concertante in E flat.

Of course Counting by numbers is also relevant as bees probably count in a different way altogether. Perhaps I should have ended the title of the post with a question mark?

Hydroxymethylfurfural

Excuse me?

Hydroxymethylfurfural which, for very obvious reasons is usually abbreviated to HMF, is an organic compound that forms in sugar-containing foods, often as a result of heating.

Hydroxymethylfurfural (HMF) Oxygen = red, Hydrogen = white, Carbon = black

HMF is relevant to bees because, at high levels, it is toxic for them. Since beekeepers often heat (or use ready-made feed that has been heated during production) sugar-containing syrups or fondants it’s worth being aware of it.

HMF is also relevant to beekeepers as high levels of it in honey are an indication of prolonged heating during storage and preparation or potential adulteration. For this reason there are legal limits on the levels of HMF in honey sold for human consumption.

I suspect beekeepers in the UK who know about HMF – and many may not – probably worry about it unduly. In tropical countries or regions where high fructose corn syrup is used as a bee food then HMF is likely to be of more immediate importance.

Natural occurrence of HMF

Hydroxymethylfurfural is essentially absent from fresh foods. However, in sugar-containing foods, particularly those that are acidic, HMF levels can build up. A chemical process called a Maillard reaction is responsible for HMF formation (there are other reactions that generate HMF as well, including caramelisation) and the reaction works about five times faster for every 10°C rise in temperature.

Therefore processes such as drying or cooking result in elevated HMF levels. The precise amount varies depending upon the foodstuff, the amount of heating and other factors; typical figures are bread 3 – 180 mg/kg, prunes 240 mg/kg, sugarcane syrup 100 – 300 mg/kg and roast coffee 900 mg/kg 1.

All of these foods can be consumed perfectly safely (at least in terms of their HMF content … prunes can have some adverse effects 😉  ).

It should therefore be obvious that the 40 mg/ml limit 2 of HMF in honey has nothing to do with its safety for human consumption.

Dietary HMF has been extensively studied as there were concerns it may be carcinogenic for humans. Several studies showed that non-physiological levels and/or prolonged exposure were cytotoxic or inhibited key enzymes in the cell such as DNA polymerase. However, no evidence for in vivo carcinogenic or genotoxic effects have been demonstrated 3.

HMF is currently considered safe and has been shown to have beneficial antioxidant activity, to protect against hypoxic (low oxygen) injury and to counteract the activities of some allergens.

HMF in honey

Readers familiar with the chemistry of honey will be aware that it is often rich in fructose (one of the sugars from which HMF is derived) and is acidic.

Add a little heat and you have near-perfect conditions for the production of HMF.

How much heat? 

It’s actually not just heat but a combination of heat and time.

The higher the temperature, the less time is required for the production of a certain amount of HMF. There are several studies of this, but one of the most frequently quoted is from White et al., in 1964 4 which has this slightly skewwhiff, but nevertheless useful, graph of the influence of storage temperature and time on HMF production in honey.

HMF production in honey – influence of storage temperature and time

That’s barely legible – check (and enlarge) the original if needed – but the approximate times/temperatures required to generate 30 mg/kg of HMF in honey 5 are as follows:

30°C ~250 days
50°C ~10 days
70°C ~10 hours

All of which is good news … heating a 15 kg bucket of rock-solid OSR honey overnight at 50°C to melt it before making soft-set honey is unlikely to significantly increase the HMF levels.

How to avoid the generation of HMF in honey

But, if you are worried about HMF levels, you could always produce creamed honey which just requires overnight warming at 33°C.

This is what I now do; not because of any concern over the HMF levels but because it’s:

  • faster
  • produces a honey with better batch-to-batch consistency of texture
  • generates a jarred honey much less susceptible to frosting

Long-term storage of honey results in the formation of HMF. The lower the temperature it is stored (and the shorter the time) the less HMF is produced. For an exhaustive list of HMF levels quantified in honey stored at different temperatures have a look at Table 1 in Shapla et al., (2018).

Store honey carefully in a cool place … or sell, eat or gift it quickly

It makes senses to store honey in a cool place with a relatively stable temperature.

Quantifying HMF

There are a variety of ways of detecting HMF. Unfortunately, all require laboratory equipment and none are really suitable for home use.

There are spectrophotometric methods – essentially detecting a colour change after adding an indicator that reacts to the presence of HMF – but these can lack both sensitivity and specificity. Some of the chemicals involved are carcinogenic.

More accurate and sensitive are methods use reversed-phase high-performance liquid chromatography. These have been in routine use for years.

Orbitrap ID-X Tribrid Mass Spectrometer

Probably the newest and most advanced methods involve the use of time-of-flight mass spectrometry (MS MALDI-TOF). These ionise the constituents of the sample and measure the time they take to reach a detector. Mass spectrometry is exquisitely sensitive and specific … and the equipment is eye-wateringly expensive. 

Since you’re unlikely to have one in your honey processing room 6 you’re better off doing your best to avoid conditions that lead to the build-up of HMF in the first place.

OK, enough about honey and humans, what about the bees?

HMF is toxic for both adult bees and developing larvae. The level of toxicity depends upon the concentration of the HMF, the duration of exposure and the developmental stage of the bee.

Krainer and colleagues 7 looked at toxicity of HMF to developing larvae and showed that concentrations up to 750 ppm (i.e. 750 mg/kg) did not reduce larval or pupal mortality.

Larval and pupal mortality when exposed to HMF at different concentrations.

They calculated that the LC50 (concentration that produced 50% mortality) at day 7 and day 22 was 4280 ppm and 2424 ppm respectively, with a calculated LD50 (dose per larva that resulted in 50% mortality) of 778 μg and 441 μg at day 7 and day 22 respectively.

Adult bees were less sensitive to HMF in the first week after emergence than during the first week of larval development.

Adult bee mortality when fed a diet containing HMF at the levels specified

What does all this mean?

It means that high levels of HMF are likely to have a significant impact on adult bees, but – at least until the levels are exceptionally high (grams, not milligrams, per kilogram) will probably not adversely impact brood levels.

Further validation of the adverse effects of HMF to adult bees

A similar study was recently conducted by Gregorc and colleagues 8 using lower concentrations of HMF.

Survival of adult bees fed with HMF-spike Apifonda

Again, there was a time/dose response, but note that only about 30% of the control bees survived 30 days and this was only double the number that had been fed the lowest level of HMF-spiked Apifonda. Note the clear evidence of a dose-response with increasing levels of HMF in the diet.

Dysentery

Several studies, dating back at least 50 years, report that high levels of HMF result in dysentery-like symptoms due to ulceration of the gastrointestinal tract of honey bees.

Gregorc and colleagues used immunohistochemistry to investigate the integrity of the gut tissues in the honey bees fed HMF. They stained cells red that were undergoing a process called ‘programmed cell death’ or apoptosis. This is a natural physiological response to damage. The more red staining, the worse the damage.

Midgut of formalin-fixed, paraffin-embedded tissue of worker bees exposed to HMF

At higher doses of HMF and/or longer exposure there was increased apoptosis in the gut tissues, presumably accounting for the dysentery-like symptoms often seen (though these were not recorded in this particular study).

Real world beekeeping

All of these bee corpses and fancy-dan immunohistothingamajiggery really just confirm that high levels of HMF are a bad thing™

In terms of honey processing and storage the allowed levels are nothing to do with human (or bee) health, but everything to do with evidencing overheated, poorly stored or doctored honey.

And since no readers of this blog do these things then there’s no need to be concerned 😉

Assuming your honey starts with low HMF levels (on extraction) then any reasonable levels of heating to liquify honey for filtering, blending or jarring should not result in HMF levels anywhere near to those that would prevent the honey being saleable 9.

Refer to the graph above from the 55 year old paper from White and Co. (shown above) for further validation.

If you’re making thick (2:1 by weight sugar to water) syrup to feed bees perhaps use warm rather than boiling water. However, considering the time involved and the absence of the acidity of honey, even with the latter HMF levels should not get close to high enough levels to endanger the bees.

If you’re making thin (1:1 by weight) syrup then use cold water. Just stir it a bit longer to dissolve it all.

However, take care – or avoid altogether – the use of high fructose corn syrups (HFCS) for feeding bees. I don’t know anyone who does this in the UK and have no experience of it myself. To learn more have a look at this article in Bee Culture. HFCS is high in fructose (the clue is in the name) and acidic, so HMF readily forms.

Studies of commercial HFCS show levels of HMF can start at 30 – 100 mg/kg before any long-term storage. 

Oxalic acid

The only time most beekeepers probably need to have concern about HMF levels is in the preparation and storage of oxalic acid solutions for trickle treating colonies in midwinter.

Oxalic acid is, er, acidic. For trickle treating it’s mixed with thin syrup to make a 3.2% solution. The combination of syrup and acidity means that HMF can be produced if stored – for a long time – in unsuitable conditions (under which there is an obvious colour change).

Stored OA solution and colour change

Stored OA solution and colour change …

So, if you’re preparing OA solutions for trickle treating either:

  • use it immediately and safely dispose of the excess
  • store it at 4°C and use then it as soon as possible (before safely disposing the excess)

Fondant

But what about fondant?

The HMF levels in commercially available fondant have recently been discussed on the Beekeeping Forum. I’m grateful to ‘loyal listener reader’ (to use Radio 4’s More or Less definition) Archie McLellan for bringing this to my attention.

The thread started with the challenging title The truth behind fondants.

Like all discussion groups, the contributions are many and varied.

Some wander off-topic.

Others use it as an opportunity to get a little dig in at the opposition.

Or a great big dig 😉

Novices and the naive ask simple questions and hope for straightforward answers 10.

Usernames often give no indication of who the poster actually is.

Is the poster a manufacturer or distributor of BeeCentric fondant™ “The best fondant for bees and a whole lot better than that cr*p they sell for ice buns”

Does the poster use 5 tonnes of fondant a year and buys anything s/he can get as long as it’s cheap enough?

Or does the contributor have a £576,000 Orbitrap MALDI-TOF mass spectrometer in their basement and a damned good idea of exactly how much HMF is present in every commercial source of fondant?

On the internet, nobody knows you’re a dog

Who knows?

I certainly don’t know all of the contributors to these threads.

But I know some of them 😉

Read the thread. It’s now 12 pages long and you’ll do well not to get lost or to disappear down a few cul-de-sacs

If you’ve ‘got a life’ and want to cut to the chase then have a look at this post in particular.

What do I do?

I use standard Baker’s fondant. It costs about £8-12 for 12.5 kg depending how much you need. I’ve used this type of fondant for a decade for 90% of my colony feeding (and 100% of my autumn feeding).

I’ve never seen any adverse effects from using this type of fondant for my bees.

I simply do not believe some of the negative marketing that is used to promote BeeCentric fondant™ costing £36 for 12.5 kg. It’s not that I can’t afford this 11 and it’s certainly not because I don’t care about my bees. I simply choose to trust experience over carefully-worded marketing ‘information’.

To convince me they’d need to publish the HMF levels in their products. They might be lower than bog-standard Baker’s fondant.

And I’d also want to know the HMF levels in standard Baker’s fondant 12.

If they were significantly higher 13, are they anywhere near high enough to damage my bees?


Note

A version of this article appeared in the November 2021 edition of An Beachaire – The Irish Beekeeper.

Cut more losses

This is a follow-on to the post last week, this time focusing on feeding and a few ‘odds and sods’ that failed to make it into the first 3000 words on reducing overwintering colony losses.

Both posts should be read in conjunction with one (or more 1 ) of my earlier posts on disease management for winter. Primarily this involves hammering down the mite levels before the winter bees are produced, so ensuring their longevity.

But also don’t forget to treat your colonies during a broodless period in midwinter to mop up mites that survived the autumn treatment, or have reproduced since then.

Why feed colonies?

All colonies need sufficient stores to get the colony through the winter until suitable nectar sources and good enough weather make foraging profitable the following spring.

How much the colony needs depends upon the bees themselves – some strains are more frugal than others – and the duration of the winter. If there is no forage available, or the weather is too poor for the bees to fly, then they will be dependent upon stores in the hive.

A reasonable estimate would probably be somewhere around 20 kg of stores, but this isn’t a precise science.

It’s better for the colony to have too much than too little. 

If the colony has stores left over at winter’s end you can always remove them and use them when you make up nucs later in the season. Just pull out the frames and store them safely until needed.

Unused winter stores

In contrast, if the colony starts the winter with too few stores there are only two possible outcomes:

  • the colony will starve to death, usually in late winter/early spring (see below)
  • you will spend your winter having to regularly check the colony weight and opening the hive to add “emergency rations” to get them through the winter

Neither of these is desirable, though you should expect to have to check the colony periodically in winter anyway.

Feeding honey for the winter … and meaningless anecdotes

By the end of the summer the queen has reduced her laying rate and the bees should be backfilling brood comb with honey stores. If you assume there’s about 5 kg of stores 2 in the brood box then they’ll need about another 15 kg. 

15 kg is about the amount of honey you can extract from a well-filled super. 

Convenient 😉

Some beekeepers leave a full super of honey on the hive, claiming the “it’s better for the bees than syrup”

Of course, it’s a free world, but there are two things wrong with doing this:

  • where is the evidence that demonstrates that honey is better than sugar-based stores?
  • it’s an eye-wateringly expensive way to feed your colonies

By evidence, I mean statistically-valid studies that show improved overwintering on honey rather than sugar.

Not ‘my hive with a honey super was strong in spring but I heard that Fred lost his colony that was fed syrup’ 3.

That’s not evidence, that’s anecdote.

If you want to get this sort of evidence you’d need to start with a lot of hives, all headed by queens of a similar age and provenance, all with balanced numbers of brood frames/strength, all with similar mite levels and other pathogens.

For starters I’d suggest 200 hives; feed 50% with honey, 50% with sugar … and then repeat the study for the two following winters.

Then do the stats 4.

The economics of feeding honey

If I were a rich man …

The 300 supers of honey used for that experiment would contain honey valued at about £80,000.

That’s profit, not sale price (though it doesn’t include labour costs as I – and many amateur beekeepers – work for free).

The honey in a single full super has a value of £250-275 … that’s an expensive way to feed your bees 5.

Particularly when it’s not demonstrably better than a tenner or so of granulated sugar 🙁

But there are more costs to consider

The economic arguments made above are simplistic in the extreme. However, there are other costs to consider when feeding colonies.

  • time taken to prepare and store whatever you will be feeding them with 6
  • feeders needed to dispense the food (and storage of these when not in use)
  • energetic costs for the colony in converting the food to stores

Years ago I stopped worrying (or even thinking much) about any of this and settled on feeding colonies fondant in the autumn.

Fondant mountain ...

Fondant mountain …

Fondant is ~78% sugar, so a 12.5 kg block contains about 9.75 kg of sugar.

This year I’m paying £11.75 for fondant which equates to ~£1.20 / kg for the sugar it contains.

In contrast, granulated sugar is currently about £0.63 / kg at Tesco.

The benefits of fondant

Although my sugar costs are about double this is a relatively small price I’m (more than) prepared to accept when you take into account the additional benefits.

  • zero preparation time and no container costs. Fondant comes ready-wrapped and stores for years in the box it is purchased in
  • no need for jerry cans, plastic buckets or anything to prepare or store it in before use
  • no need for expensive Ashforth-type feeders that sit around for 95% of the year unused When I last checked an Ashforth feeder cost £66 😯 
  • it takes less than 2 minutes to add fondant to a colony
  • no risk of spillages – in the kitchen, the car or the apiary 7.
  • fondant is taken down more slowly than syrup, so providing more space for the queen to continue laying. In addition, in the event of an early cold snap, fondant remains accessible whereas bees often stop taking syrup down

Regarding the energetic costs for the colony in storing fondant rather than syrup … I assume this is the case based upon the similarity of the water content of fondant to capped stores (22% vs. 18%), whereas syrup contains much more water and so needs to be ripened before capping to avoid fermentation.

Fondant block under inverted perspex crownboard – insulation to be added on top.

Whether this is correct or not 8, the colony has no problem taking down the fondant over a 2-4 week period and storing it.

What are the disadvantages of using fondant? 

The only one I’m really aware of is that the colony will not draw fresh comb when feeding on fondant (or at least, not enthusiastically). In contrast, bees fed syrup in the autumn and provided with fresh foundation will draw lovely worker brood comb. 

Do not underestimate this benefit.

They fancied that fondant

Brood frames of drawn comb are a very valuable resource. Every time you make up a nuc, or shift a nuc to a full-sized box, providing drawn comb significantly speeds up the expansion of the resulting colony.

Nevertheless, for me, the advantages of fondant far outweigh the disadvantages …

Finally, in closing, I’ve not purchased or used invert syrup for feeding colonies. Other than no prep time this has the same drawbacks as syrup made from granulated sugar. Having learnt to use fondant a decade or so ago from Peter Edwards (Stratford BKA) I’ve never felt the need to look at other options.

Let’s move on …

Ventilation and insulation

Bees can withstand very cold temperatures if healthy and provided with sufficient stores. In northern Canada bees may experience only 120 frost-free days a year, and cope with 3-4 week periods in winter when the temperature is -25°C (and colder if you consider the wind chill).

That makes anywhere in the UK look positively balmy.

Margate vs. the Maldives … a similar temperature difference to Margate vs. Manitoba in the winter

I’ve overwintered colonies in cedar or poly boxes for a decade and not noticed a difference in survival rates. Like the honey vs. sugar argument above, if there is a difference it is probably minor. 

However, colony expansion in poly boxes in the spring is usually better in my experience, and they often fill the outer frames with brood well before cedar boxes in the same apiary get there.

Whether cedar or poly I take care with three aspects of their insulation/ventilation:

  • the colonies have open mesh floors and the Varroa tray is only in place when I’m actively monitoring mite drop
  • all have insulation above the crownboard in the form of a 50 mm thick block of Kingspan (or Recticel, or Celotex), either integrated into the crownboard itself, placed above it or built into the roof
  • I ensure there is no upper ventilation – no matchsticks under the crownboard, no holes etc.
  • excess empty space in the brood box is reduced to minimise the dead air space the bees might lose heat to

In my experience bees actively dislike ventilation in the crownboard. They fill mesh with propolis …

Exhibit A … are you getting the message?

… and block up the holes in those over-engineered Abelo crownboards …

Exhibit B … ventilated hole in an Abelo crownboard

Take notice of what the bees are telling you … 😉

Insulation over the colony

I’ve described my insulated perspex crownboards before. They work well and – when inverted – can just about accomodate a flattened 9, halved block of fondant.

Perspex crownboard with integrated insulation

Finally, if it’s a small colony in a brood box 10 then I reduce the dead space in the brood box using a fat dummy

Fat dummy with integral feeder

Fat dummy …

I build these filled with polystyrene chips.

You don’t need this sort of high-tech solution … some polystyrene wrapped tightly in a thick plastic bag and sealed up with gaffer tape works just as well.

Insulation ...

Insulation …

I’ve even used bubblewrap or that air-filled plastic packaging to fill the space around a top up block of fondant in a super ‘eke’ before now.

However, remember that a small weak colony in autumn is unlikely to overwinter as well as a strong colony. Why is it weak? Would you be better uniting it before winter starts?

Nucleus colonies

Everything written above applies equally well to nucleus colonies.

A strong, healthy nuc should overwinter well and be ready in the spring for sale or promoting to a full colony.

Here's one I prepared earlier

Here’s one I prepared earlier … an overcrowded overwintered nuc in April

Although I have overwintered nucs in cedar boxes I now almost exclusively use polystyrene. This is another economic decision … a well made cedar nuc costs about double the price of the best poly nucs

I feed my nucs fondant in preparation for the winter, typically by adding 1-2 kg blocks to the integral feeder.

Everynuc fondant topup

Everynuc fondant topup

Because of the absence of storage space in the nuc brood box it’s not unusual to have to supplement this several times during the autumn and winter.

You can even overwinter queens in mini-mating nucs like Apidea’s and Kieler’s.

Kieler mini-nuc with overwintering queen

This deserves a post of its own. Briefly, the mini-nuc needs to be very strong and usually double- or triple- height. I build fondant frame feeders for Kieler’s that can be quickly swapped in/out to compensate for the limited amounts of stores present in the brood box.

Kieler mini-nuc frame feeders

My greatest success in overwintering these was in winters when I provided additional shelter by placing the nucs in an unheated greenhouse. A tunnel provided access to the outside. However, I know several beekeepers who overwinter them without this sort of additional protection (and have done so myself).

Just because this can be done doesn’t mean it’s the best thing to do.

I’d always prefer to overwinter a colony as a 5 frame nuc. The survival rates are much better, their resilience to long periods of adverse weather is significantly greater, and they are generally much more useful in the spring.

Miscellaneous musings

Hive weight

A colony starting the winter with ample stores can still starve if the bees are particularly extravagant, or if they rear lots of brood but cannot forage.

The rate at which stores are used is slow late in the year and speeds up once brood rearing starts again in earnest early the following spring (though actually in late winter).

Colony weight in early spring

As should be obvious, this is a Craptastic™ sketch simply to illustrate a point 😉

The inflection point might be mid-December or even early February.

The important message is that, once brood rearing starts, consumption of stores increases. Keep checking the colony weight overwinter and supplement with fondant as needed.

I’m going to return to overwinter colony weights sometime this winter as I’m dabbling with a weather station and set of hive scales … watch this space.

An empty super cuts down draughts

Periodically it’s suggested that an empty super under the (open mesh) floor of the hive ‘cuts down draughts’, and is therefore beneficial for the colony.

It might be.

But like the ‘overwintering on honey’ (and being a pedant scientist) I’d always want to see the evidence.

There are two claims being made here:

  • a super under the floor cuts down draughts
  • fewer draughts benefits the colony which consequently overwinters better

Really?

There are ways to measure draughts but has anyone ever done so? Remember, the key point is that the airflow around the winter cluster would be reduced if there are fewer draughts. 

Does a super reduce this airflow significantly over and above that already caused by the sidewalls of the floor?

And, even if it does, perhaps the colony ‘reshapes’ itself to accommodate the draught from an open mesh floor.

What shape is the winter cluster?

For example, in an uninsulated hive (including no insulation over the cluster) with a solid floor the cluster is likely to be roughly spherical. They minimise the surface area.

With an open mesh floor are they more ellipsoid, so avoiding draughts from below? If so, is this improved much by an empty super below the open mesh floor? Does the cluster change shape or position? I don’t know as I’ve not compared cluster shapes in solid vs. open mesh floors plus/minus a super underneath.

And anyway, an open mesh floor looks very like a baffle to me … how much better can it get? How draughty is it in the first place?

Is this example 8,639 for my ‘Beekeeping Myths’ book?

I do know that top insulation tends to flatten the cluster against the warm underside of the crownboard.

Midwinter cluster

A strong colony in midwinter

Having worked out that draughts are (or are not) reduced … you still need another couple of hundred hives to test whether overwintering success rates are improved!

More winter bees

Finally, always remember that the survival of the colony is dependent upon the winter bees. All other things being equal (stores, disease etc.), a colony with lots of winter bees will overwinter better than one with fewer.

This is one of the reasons I stopped using Apiguard for mite control in autumn. Apiguard contains thymol and quite regularly (30-50% of the time in my experience) stopped the queen from laying, particularly in warmer weather. 

Apiguard works well for mite control, but I became wary that I was potentially stopping the queen at a time critical for late-season colony development. I worried that, once treatment was finished, a cold snap would shut down brood rearing leaving it with suboptimal numbers of winter bees.

I never checked to see whether the queen ‘made good’ any shortfall after removal of the treatment … instead I moved to Scotland where it’s too cold to use Apiguard effectively 🙁


 

Absconding

One of the few principles I have ( 😉  ) is that the posts here should be based upon practical experience. When I write about swarm control I describe the methods that I use. When I write about Varroa management I discuss Apivar and oxalic acid in detail as I have a lot of experience using these compounds. I’ve not written about MAQS as I don’t use it.

For the same reasons, you won’t see a discussion about top bar hives or the Bee Guardian piezoelectric gadgets that causes the varroa mites stop to reproduce and go away from the hive” 1.

The topic today is absconding. My qualification to write about this is extremely limited, but just about sufficient. I think I’ve had only one colony abscond in the last decade. It’s not something I take any notice of (or precautions against) in my regular beekeeping. However, it’s an interesting subject as there’s some relevant science associated with absconding and honey bee migration, so it’s worth discussing.

And perhaps more science to do …

But first some definitions

Colony reproduction involves swarming. The colony rears one or more new queens. Once the queen cells are capped, the current queen and up to 75% of the adult bees leave the hive as a swarm. Prior to leaving, scout bees have scoured the environment for suitable new nest sites. These scouts lead the swarm to the chosen new location 2.

The swarm leaves behind all the brood and most of the stores. Together with the adult bees that remain, this colony has a good chance of survival (~80%) which is probably a reflection on queen mating success rates 3.

‘Most of the stores’ because the swarming bees gorge themselves on honey prior to leaving the hive (or nest site if it’s a feral colony). Something like 40% by weight of the swarm is honey stores. They need these stores to survive – to build new comb, to tide them over a period of bad weather and while they scout the environment for forage. Swarming is a risky business, only about 20% of natural swarms survive.

Absconding is very different. During this process the entire colony – the queen and all the flying bees – leave the nest site (hive). They usually leave behind almost nothing. There may be very limited amounts of capped brood/larvae or eggs remaining, but the stores are usually gone. Absconding therefore does not involve colony reproduction. There are no queen cells produced. You start with one colony and end with one.

However, although absconding is very different, it’s not completely different. It still involves scout bees and it still involves waggle dances to communicate distance and direction.

Like swarming, it’s also a completely natural process. In certain parts of the world there are annual cycles of absconding and colony migration.

In the discussion that follows I’m going to try and make a distinction between absconding by managed and unmanaged colonies. 

The consequences of either type of colony absconding are probably the same. 

The drivers that result in the colony absconding are sometimes different.

My experience

Let’s get this out of the way … 4

To my knowledge the only colony I have ever had abscond was from a Kieler mini-nuc. The mini-nuc had been primed with a mugful of bees and a queen cell a week or so earlier. The queen had emerged and may (or may not?) have gone on a mating flight 5.

An Apidea mini-nuc ‘catching a few rays’

One baking hot June afternoon I turned up at the apiary just as a small swirling mass of bees disappeared over the fence. 

Never to return 🙁

The mini-nuc was low on stores (but far from empty) and devoid of bees (or brood). There was drawn comb so the queen would have been able to lay (if she had been mated). I can’t remember whether there were eggs present … this was several years ago 6.

‘Natural’ absconding and colony migration

This mini-nuc wasn’t the one pictured, but it was similarly exposed. In full sun these can rapidly overheat and there is a real risk of the small colony absconding. I now always site my mini-nucs out of the heat of the full sun – even in Scotland – in dappled shade, at the bottom of a hedge or somewhere similar.

Of course, I don’t know that overheating caused this little colony to abscond, but it seems like a reasonable assumption.

Bees living in temperate and tropical regions exhibit gross behavioural differences that reflect the climate and availability of forage. Those in temperate climates swarm annually, coinciding with the predictable period of forage availability, and are quiescent over ‘winter’.

In contrast, bees in tropical climates have no ‘winter’ to survive as the temperature is high enough all year for brood rearing and comb building. What differs though is the availability of forage and water. If these are limiting the bees migrate to other areas.

This annual migration involves the colonies absconding … and it has been quite well studied by scientists.

Adverse environmental conditions are one of the recognised drivers of absconding. In addition to overheating, these include a dearth of resources during the wet season. 

The other major driver of absconding is disturbance, for example by predators such as ants (or beekeepers). Disturbance is a lot less predictable than environmental factors, and it is the latter that has been better studied.

Preparing to migrate

Absconding and migration appear to be a characteristic of strong, healthy colonies. Prior to absconding the colony reduces brood rearing drastically although the queen continues to lay a very limited number of eggs until the bulk of the worker brood has emerged 7.

Colonies tended to abscond within a day of this worker brood emerging, leaving almost nothing in the original nest site. 

So, this isn’t a spur of the moment decision, it’s a protracted process taking at least a fortnight from the near-cessation of brood rearing. This means the colony benefits from the resources they have invested in rearing brood, rather than leaving behind slabs of capped brood that would otherwise be doomed.

How does the colony know where to go when it absconds?

Actually, these preparations probably take more than a fortnight. Analysis of the waggle dances for several weeks prior to absconding show that the foraging area and distances were both increasing and becoming more variable. 

Schneider and McNally 8 showed that these waggle dances regularly communicated distances of up to 20 km from the nest site.

However, these weren’t typical dances … the distance component was variable, the dance occurred during periods of little flight activity and the dance was not associated with forage sources. They interpreted this as a generalised signal to fly for a long (but unspecified) distance in a particular direction, rather than to a specific location.

I’m not aware of follow-up studies to these. Do the bees go through the same sort of decision-making process to ‘agree’ on the final direction as the scout bees do when a colony swarms? I suspect not, the distance component was very variable and there was no direct evidence that the dancing bees ever made the entire journey anyway.

Perhaps these waggle dances simply indicate “Things are better a long way south of here. When we go, that’s the direction to take”.

Stopovers

If a colony absconds due to adverse environmental conditions – such as a lack of forage, or overheating – it seems unlikely that things would be much better only 20 km away. “environment” is local, but not necessarily that local.

In reality, colonies abscond and migrate much further than this when necessary, restoring in temporary stopover locations when necessary. In the case of Apis mellifera I’m not aware of any studies of these sites. However, in the Giant honey bee (Apis dorsata) some of these stopover locations appear to be re-used annually. 

Apis dorsata migrates up to 200 km and has even been reported crossing 50 km of open water between Sumatra and Malaysia. These long migrations take up to a month and the bees bivouac on trees, resting and replenishing their stores (by foraging locally) 9.

Giant honey bee (Apis dorsata) temporary stopover bivouac

Analysis of scout bee dancing activity on the surface of these bivouacked colonies show that this again determines the direction (and possibly distance) of the next stage of the journey. 

Absconding and managed colonies

I think it’s reasonable to assume that at least some of the factors that induce colonies to abscond in tropical regions also trigger absconding in our managed colonies in the UK. 

Very small colonies – like the mini-nuc described above – are poor at thermoregulation. There are simply too few bees present to cool the colony in very hot weather.

Although I’m aware that colonies may abscond due to disturbance – from wax moth, Varroa or small hive beetle infestation – I’ve no experience of this 10.

What about disturbance by beekeepers managing colonies? It’s a possibility I suppose. Clearly the regular weekly inspections are not sufficient disturbance to trigger absconding, but perhaps a daily rummage through the brood box might not be tolerated 11.

Absconding swarms

In temperate climates most beekeepers associate absconding with recently hives swarms.

Here’s a typical scenario …

The beekeeper is called out to a bivouacked swarm hanging – conveniently and precariously, just out of reach – in a tree.

By the time they’ve collected the ladder, the skep, the white sheet and the secateurs it’s late afternoon. Never mind … A swarm in May is worth a bale of hay etc.

A spring swarm in a skep

They drop the swarm into the skep, avoid toppling off the ladder, allow the flying bees to join the queen, wrap everything in the sheet and return triumphantly to their apiary 12.

In time honoured tradition they assemble a new hive, prop the entrance open and build a sheet-covered ramp onto which they unceremoniously dump the collected swarm.

'Walking' a swarm into a hive

‘Walking’ a swarm into a hive

And the bees calmly walk up the slope into the hive.

It’s one of the great sights in beekeeping … and one I now never bother to do.

I just dump the swarm into the hive and close it up. 

Boring, but quick 😉

Back to the absconding swarm scenario …

The beekeeper returns late the following morning to find the swarm has gone 🙁

Is this typical absconding?

Other than one or two typical circumstances such as a freshly painted (and still smelly) hive, I think that these swarms may abscond because they have already chosen an alternative nest site

The scout bees from the bivouacked colony (collected a day or two previously) had been busy surveying the environment for suitable nest sites. This process can take several days until a sufficient number of the scouts are convinced of the benefits of a particular site.

Once that decision is made the colony leaves the bivouac and flies to the new nest site. However, this flight tends to happen in the middle of the day, not late in the afternoon.

The beekeeper who hived our hypothetical swarm in the scenario above may have actually interrupted this process, which simply continued the following morning.

I don’t know if scout bees conduct waggle dances overnight to reinforce nest site choices (but the normal waggle dance for forage resources can occur during the night). If they do, this might account for the bees disappearing soon after being hived.

How do you stop hived swarms absconding?

There are three methods I’m aware of.

One is foolproof and I use every season. The other two are reported to work with variable levels of success, but which I have never used.

Adding a frame of open brood is reported to help stop the colony absconding. Alternatively, placing a queen excluder under the brood box (but above the floor) ‘traps’ the queen and prevents the colony leaving.

The first of these provides brood to care for, brood pheromones and the general ‘pong’ of a hive, all of which are likely to be beneficial. As I’ve not used this method I’m unsure how effective it is.

The queen excluder seems a heavy-handed and rather crude solution. The colony may well still try and abscond, but the queen will remain trapped. This seems like a great way to induce considerable stress in the colony.

And it’s unlikely to be successful long-term if the swarm is a cast with a virgin queen 😉

And the totally foolproof method?

Swarm arriving at bait hive ...

Swarm arriving at bait hive …

Bait hives.

I’ve never had a swarm that voluntarily arrived in a bait hive abscond. Even if I move the bait hive to another apiary, they still happily stay 🙂

Citizen science

I almost never hive bivouacked swarms these days as I am sufficiently successful in attracting swarms with bait hives 13.

I’m therefore unable to conduct the following experiment that I think would be quite interesting.

I’ve predicted above that a swarm collected from a bivouac that absconds may have already decided on the new nest site. By ‘hiving’ the swarm all the beekeeper is doing is moving the bivouac.

That being the case, I’d expect that collected swarms would be less likely to abscond if they’re moved to an area the scout bees have no knowledge of.

Scout bees survey the environment at least 3 km from the original nest site although swarms tend to occupy new nest sites well within this distance.

There are two things that would be interesting to monitor:

  1. Are swarms hived over 8 km from the location the swarm is collected less likely to abscond?
  2. Is the delay between hiving a swarm and it absconding related to the distance between the original bivouac and the initial location it is hived in?

I’ve chosen 8 km because you cannot always be certain where the bivouacked swarm came from (and because it’s a convenient 5 miles for these post-Brexit times). If you assume that the bivouac is always within a few dozen metres of the original nest site this ‘removal’ distance could be decreased to about 4 km.

The time delay addresses a slightly different question. I’m assuming here that the scout bees have yet to reach a quorum decision and are continuing to survey the environment. The further you move them, the more the environment changes, so potentially necessitating a longer decision making period.

As the 2021 season starts to wind down that’s something to think about for the year ahead.


Note

Please don’t email me with all the gruesome details of swarms you’ve had abscond in the past. It’s not that I’m not interested … I’m just completely swamped with correspondence.

If there’s sufficient interest in this post over the next few months (and as a bit of ‘Citizen Science’ experiment which are all the rage) – determined by page accesses and comments – I’ll create a simple web form to log everything to a database. No individual beekeeper is likely to collect sufficient swarms to generate a meaningful amount of data. I doubt even if an entire association could do so. However, the thousands of readers a week are surely able to have enough hived swarms abscond to test the hypothesis?

Growing old (dis)gracefully

… but in this world nothing can be said to be certain, except death and taxes.

So wrote Benjamin Franklin in 1789 1.

Taxes are a subject that I’ve not previously covered. Furthermore, I have no intention of writing about them in the future. My once a year late-night wrestle with the HMRC website is a distressing enough experience and one I’d prefer not to be reminded about.

So this week I’ll deal with that other certainty … death.

Eilean Fhianain burial ground, Loch Shiel

Sooner or later it will happen to us all.

No ifs or buts, and – unlike taxes – it really is a certainty.

What’s interesting about death is the ‘sooner or later’ element.

Some die young, due to bad luck, poor health or overindulgence.

Others live to a ripe old age, outliving their peers by many years or even decades.

Talking the talk

Beekeeping has been a relatively solitary pastime for the last 16 months. The restrictions imposed by lockdowns and social distancing have meant that beekeeping meetings have all been ‘virtual’. 

I’ve written about these recently and it seems likely that many associations are going to continue (at least some of the time) with Zoom talks.

A caffeine-fueled Q&A session

Whether in person, or online, one of the things that’s noticeable is that all beekeeping audiences are – how can I put this delicately? – not as young as they used to be.

By which I mean the individual beekeepers are not callow youths, but are instead older, wiser, and – of course – better looking. 

In my experience, giving talks over the last decade or so, beekeeping audiences have always had an older average age than a cross-section of society.

In addition, as I briefly mentioned recently, the average age of beefarmers in the UK is about 66 years old.

Why is this?

It seems there are two possibilities:

  1. Since beekeeping takes a reasonable amount of time, it’s largely people who have more spare time who start or stick with the hobby. At least at the start, beekeeping also costs quite a bit of money. Again, those who are a bit older probably have more disposable income (or fewer distractions like mortgages or kids to spend it on).
  2. Beekeepers live longer. The relatively high average age of a beekeeping audience – when compared with a similarly-sized cross-section of society – reflects their increased longevity.

Of course, both of those are rather simplistic explanations, but it’s a start.

Do beekeepers live longer?

When you start beekeeping you tend to be interested in honey and swarm control and pathogens.

Or just honey 😉

But after a few years of successful beekeeping you probably produce quite a bit of honey. Your success in honey production is partly due to your understanding and implementation of swarm control, and by your interventions that minimise disease.

And so your interests in beekeeping expand.

Some produce award-winning candles or wax flowers, some rear hundreds of queens a season, some explore esoteric hive designs, and some become interested in the history of beekeeping.

And one of the things that’s noticeable about the history of beekeeping is that several well known beekeepers lived to a remarkably old age.

With improvements in nutrition and healthcare, the average life expectancy of the population has been increasing for the last 150 years or so. 

UK life expectancy (from birth) 1765-2020

If you were born in the 18th Century you’d be expected to live (on average) about 40 years. However, in the last third of the 19th Century, life expectancy started to increase.

I was going to say ‘inexorably increase’ were it not for that little blip around 1918-1920. That wasn’t the First World War. It was the last significant global viral pandemic, the so-called Spanish ‘flu 2. Only recently has this increase in life expectancy started to plateau, and actually reverse.

Well known historical (old) beekeepers

My knowledge of the history of beekeeping is rather patchy so I did a quick search for ‘famous beekeepers‘.

Don’t bother with the first couple of hits 3, but the third is the ever-dependable and enjoyable Bad Beekeeping Blog by Ron Miksha.

Here’s a few picked at anything-but-random ( 😉 ) to support my hypothesis that beekeepers live longer. 

In no particular order:

  • François Huber (1750-1831, 81 years), Swiss, ~40 years 4. Huber was an extraordinary individual 5. Despite being blind his ‘observations’ worked out many details of the life cycle of the queen and he developed one of the first observation hives.
  • Lorenzo Langstroth (1810-1895, 85 years), US, ~40 years. Langstroth combined an understanding of ‘bee space’ with the movable frame ‘leaf hive’ (developed by Huber) to develop and patent the first removable frame hive.
  • Brother Adam (1898-1996, 98 years), German (lived in UK), ~47 years. Brother Adam (Karl Kehrle) was an authority on bee breeding and the developer of the Buckfast strain of honey bees. He resigned his post of beekeeper at Buckfast Abbey at the age of 93.
  • Charles Dadant (1817-1902, 85 years), French (lived in US), ~40 years. Dadant invented the Dadant hive, ran thousands of hives (after failing as a vintner) and founded the – still flourishing – Dadant & Sons beekeeping business.
  • Eva Crane (1912-2007, 95 years), UK, ~52 years. Eva Crane was a mathematician/physicist who spent much of her (long) life doing research on bees. She founded the Bee Research Association (still flourishing as the International Bee Research Association) and wrote hundreds of papers, and notable books, on bees and beekeeping.
  • Harry Laidlaw (1907-2003, 96 years), US, ~51 years. Harry Laidlaw was one of the pioneers of studies on bee genetics and optimised methods for instrumental insemination of queens. 
  • Karl von Frisch (1886-1982, 96 years), German, ~39 years. Karl von Frisch deciphered the waggle dance of honey bees and received the Nobel Prize in 1973.

And there are many others … including many much less famous but equally old 6

Correlation not causation

Just because (some) beekeepers live a long time doesn’t mean that beekeeping is responsible for their longevity. 

Perhaps they’ve just got ‘good genes’ and they’d have lived into their 80’s or 90’s whether they’d been beekeepers or BASE jumpers.

BASE jumping, Half Dome, Yosemite

Maybe it takes that long to become an acknowledged expert at beekeeping? 

How many famous beekeepers can you name who died young?

If Harry Laidlaw had only lived to his mid-60’s (still older than the average for his year of birth) perhaps he’d have been unknown?

Unlikely … he published his first book at the age of 25, was elected a fellow of the American Association for the Advancement of Science at 48 and was the first Associate Dean for Research in UC Davis in his late 50’s.

All of the individuals listed enjoyed a near-lifelong association with bees and were clearly exceptional beekeepers well before they also achieved an exceptional age (considering the year that they were born).

So perhaps there is something about bees or beekeeping that makes beekeepers live longer?

One possibility is that honey is good for you. 

Although some beekeepers don’t like honey 7, most undoubtedly do. Honey has a host of antimicrobial, antiviral, antiparasitic, anti-inflammatory and antioxidant effects, as well as being a guaranteed ( 😉 ) way to prevent hay fever.

Or perhaps it’s bee stings? 8

Caveat

Treat most of what I’ve written above with some caution.

My selection of famous (old) beekeepers is extremely selective. 

The reason audiences in beekeeping association meetings have a high average age is almost certainly due to spare time and disposable incomes … and because all the young ones are out partying.

It’s not my sort of science, but a proper study of the association between longevity and beekeeping would be quite interesting. 

Do beekeepers actually live longer than non beekeepers? 

Is there a causative association? Is it associated with beekeeping per se, or do non-beekeepers who eat honey also live longer? How many hive/years do you need to keep bees to increase your longevity? If it’s bee stings that are beneficial, do beekeepers who keep stingless bees also live long and healthy lives?

There is literally a certain finality about studying the age at death. Are there perhaps other markers of longevity that could be investigated a little earlier in a beekeeper’s life?

There certainly are …

Chromosomes and DNA replication

We (beekeepers) have 23 pairs of chromosomes 9 that consist of DNA and proteins and contain the majority of our genetic material 10

The chromosomes are in nucleus of the cell.

Cells associate to form tissue (like muscle or nerves) which associate to form organs (like the heart or brain).

As humans grow – from egg, to embryo, to foetus, to adult – these cells have to divide. And the chromosomes have to be duplicated to ensure that all cells end up with the required 23 pairs. 

Chromosomes are not circular but are essentially linear strands of DNA. This introduces a problem. 

The enzymes that copy and make new DNA (the DNA polymerase) only ‘work’ in one direction. Since DNA consists of two antiparallel strands this means that the polymerase copies one strand directly to make one continuous product, but it copies the other strand in small pieces, and then joins them together. 

The details really don’t matter … but the consequences do. 

The small piece synthesised at the very end of the discontinuously copied strand isn’t quite at the very end of the strand 11

Frankly, this is a bit of a design flaw 🙁

Telomeres

As a consequence of this discontinuous copying, one of the strands of the DNA molecule gets a little bit shorter every time it is copied.

The DNA of chromosomes contains all the genes that make all the proteins that makes all the cells that get together to form all the tissues that create the organs that make beekeepers.

Phew!

So, if the little bit of the chromosome that’s lost during replication happened to contain an essential gene, things would go very seriously wrong™.

But chromosomes have a sort of ‘get out of jail card’.

The ends of the chromosome contain a region of highly repetitive and non-coding 12 DNA called telomeres. You can imagine the telomere as a sort of ‘cap’ at the end of the chromosome.

During replication, little bits of this cap are lost – the cap gets shorter – but this truncation does not result in the loss of any essential genes.

Telomere shortening and cell division

And all this telomere shortening is rather predictable.

As cells divide – during growth or tissue repair – the telomeres shorten. Therefore, if you measure telomere length you can get an idea of how many division it has undergone and therefore how old it is.

Telomere length is therefore a measure of biological age.

Except it’s not quite that simple

There are a bunch of things that also influence telomere length.

For example, the age of the father influences the length of the child’s telomeres. 

Telomeres also accumulate damage – and so shorten – through oxidative stress. This is a process that results from the excess production of oxidants such as peroxides, free radicals and reactive oxygen species. These are chemical intermediates in normal biochemical processes. The cell can cope with small amounts of oxidants. However, the protection mechanisms become swamped if they are in excess, resulting in cellular damage.

Some diets are rich in antioxidants which can (or at least are hypothesised to) reduce oxidative stress. 

Honey can contain high levels of polyphenols, these are well known antioxidants that are also found in some fruits, vegetables and olive oil.

Finally … we’re getting somewhere 😉

There are studies that demonstrate that eating honey every day increases the levels of antioxidants 13. However, I’m not aware that these or other studies were extended to investigate whether the test cohort also exhibited a reduced rate of telomere shortening.

This isn’t surprising … the inherent variation between individuals and the relatively slow rate at which telomeres shorten means thousands of individuals would need to be analysed. Potentially over many years.

But there is also a study of telomere length in beekeepers … which is the reason for the 2,200 word introduction above.

Beekeepers and telomeres

A Malaysian research team 14 have measured telomere length in 30 beekeepers and the same number of age-matched non-beekeepers.

The beekeepers chosen had all been keeping bees for at least five years. The non-beekeepers didn’t just not keep bees 15, they also did not consume any bee products (honey, propolis 16 or royal jelly 17 ). In addition to being age-matched (average age ~42 years) both groups excluded individuals with known disease.

And I wouldn’t have been telling you all this unless the telomere length did differ significantly. The non-beekeeper’s had telomeres that were ~30% shorter.

Chronologically they were the same age, but biologically they were older.

This small study also investigated whether there was a correlation between the period of beekeeping, the number of bee products consumed, or the period or frequency of bee product consumption, and telomere length.

Telomere length only correlated with the frequency and duration of consumption of bee products, not with the types of products or the number of years of beekeeping.

The significance of these sorts of population-based studies is related to the scale of the study. This is a small study, and the only one I’m aware of that specifically investigates telomere length in beekeepers.

It has only been cited 7 times and has not been repeated.

It remains firmly in the ‘interesting observation’ rather than ‘acknowledged fact’ category. 

So all those afternoons hunched over the hive appear to make no difference 18, but having honey every day on your porridge might actually make you younger.

Biologically younger … you’ll still look old 😉


 

Rational Varroa control

It’s the end of July … in the next two to three weeks the first eggs will be laid that will develop into the winter bees that get your colonies through to next spring. Protecting these winter bees is necessary to prevent overwintering colony losses.

I’ve written and lectured extensively on Varroa control and related topics for at least 5 years. The following article is published in August’s BBKA Newsletter and The Scottish Beekeeper. It provides an overview of what I term rational Varroa control.

I define this as effective mite management based upon our current understanding of the biology of bees and Varroa. The goal of this control is to minimise winter losses due to Varroa and viruses.

It is not a recipe with easy to follow if this, then that instructions. Neither does it provide a calendar-based guide of what to do and when to do it.

It does not even tell you what you should use for mite control.

Instead it focuses on the principles … understanding these will enable you to implement control strategies that help your bees, in your environment, survive.

This version is hyperlinked to additional, more expansive, posts on particular topics, is slightly better illustrated than those that appeared in print and contains some additional footnotes with caveats and exceptions.


Introduction

Despite almost 30 years experience of Varroa in the UK, this ectoparasitic mite of honey bees remains the greatest threat to bees and beekeeping. With the exception of those fortunate to live in mite-free regions, all beekeepers must manage the mite population in their hives or risk losing the colony to the viruses transmitted when Varroa feeds on developing pupae. 

Fortunately, Varroa control is relatively straightforward; there are a range of approved and effective miticides that – used appropriately – reduce mite infestation levels significantly. The key words in that last sentence are ‘approved’, ‘effective’ and ‘used appropriately’. In reality annual colony losses, primarily occurring in the winter, often exceed 20% (Figure 1) and may be significantly higher in long or harsh winters 1. Many of these losses are attributable to Varroa and viruses. It is therefore clear that many beekeepers are not successful in managing Varroa; either they are not treating at all, or they are treating inappropriately.

Figure 1. BBKA winter survival survey – larger studies (COLOSS and BIP) often show much higher losses

This article is primarily aimed at relatively inexperienced beekeepers, but may also help the more experienced who still suffer with high levels of winter losses. It emphasises the importance of two, correctly timed, appropriate miticide treatments per season that should ensure colony survival. It is not going to deal with treatments of questionable or minor efficacy. These include the use of small cell foundation, drone brood culling or sugar dusting. These may reduce mite levels, but insufficiently to benefit colony health. Nor will it discuss the use of any miticides (or application methods) that are not approved by the Veterinary Medicines Directorate. I will also not discuss treatment-free beekeeping, selection of mite-resistant bees or advanced colony manipulations like queen trapping. In my view any or all of these could or should be tried … but only once a beekeeper can routinely successfully overwinter colonies using strategies similar to those described here.

The problem

Varroa is an ectoparasitic mite that feeds on developing honey bee pupae. During feeding it transmits a range of honey bee viruses, the most important of which is deformed wing virus (DWV). DWV is present in honey bees in the absence of Varroa. In our studies, using sensitive PCR-based detection methods, we never detect bees – even those from mite-free regions of Scotland – without DWV. The virus is transmitted horizontally between bees during trophallaxis, and vertically from drones or the queen through sperm or eggs. These routes of transmission are rarely if ever associated with any significant levels of disease and virus only replicates to modest levels (perhaps 1-10 thousand viruses per bee). However, when Varroa transmits DWV the virus bypasses the bee’s natural defence mechanisms and replicates to very high levels in recipient pupae (billions per pupa, 1 – 10 million times higher than in unparasitised pupae). Studies from our laboratory have shown that ~75% of pupae with these high virus loads either do not emerge, or emerge exhibiting the characteristic “deformed wings” that give the virus its name (Figure 2; Gusachenko et al., Viruses 2020, 12, 532; doi:10.3390/v12050532). The ~25% of bees that do emerge and appear ‘normal’ exhibit a range of symptoms including reduced fitness, impaired learning and reduced foraging. However, most importantly they also exhibit reduced longevity. During the summer this is probably not critical; the lifespan of a worker is only ~6 weeks and, assuming the queen is laying well, there are thousands of half-sisters around with more being produced every day.

DWV symptoms

Figure 2. DWV symptoms

But during the winter, brood rearing either stops completely or drops to a very low level. The bees reared from late summer onwards are physiologically very different. These are the ‘winter bees’, also termed the diutinus bees (from the Latin meaning long-lived). Physiologically these bees resemble juvenile workers and they can survive for many months. And they need to … it is these bees that get the colony through the autumn, winter and into the following spring. They protect the queen, they thermoregulate the hive and, usually around the winter solstice, they start to rear small amounts of new brood for the season ahead.

The longevity of the bees in the hive in winter is critical to colony survival. If the winter bees have high DWV levels their longevity is reduced (in addition to the reduced numbers due to overt disease or non-viability). This means that the winter cluster shrinks in size faster than it would do otherwise. With reduced numbers of bees it cannot keep brood warm enough and so the colony fails to expand early the following season. In cold spells it may be unable to reach the food stores resulting in the colony perishing from ‘isolation starvation’. It may not be able to maintain sufficient warmth to protect the queen, or may simply freeze to death.

The goal of rational Varroa control

Successful overwintering requires lots of winter bees. The size of the winter cluster is directly related to its survival chances. Therefore the goal of rational Varroa control is to prevent the winter bees from being exposed to mites and mite-transmitted viruses during their development. Winter bee production is induced by a range of factors including photoperiod, nectar and pollen availability, brood and forager pheromones. Together these induce slowed behavioural maturation of the winter bees. This is not like flicking a switch, instead it is a seamless transition occurring as late summer segues into early autumn (Figure 3). Winter bee production is also influenced by the queen. Young queens lay later into the autumn, so increasing the numbers of winter bees. 

Figure 3. Colony age structure from August to December.

It is important to note that these events are environment-driven, not calendar-driven. It will not happen at precisely the same time each year, or at the same time in different locations (or latitudes) each year.

To protect these winter bees the colony needs to be treated with an effective miticide before the majority of the winter bees are produced. This ensures that the developing winter bee pupae are not parasitised by virus-laden mites and so do not suffer from reduced longevity. 

When are winter bees produced in the UK? 

Unfortunately, I’m not aware of any direct studies of this. Scientists in Bern (49.9°N) in 2007/08, where the average temperatures in November and December were ~3°C, showed that the Varroa- and virus-reduced longevity of bees was first measurable in mid-November, 50 days after emergence. By extrapolation, the eggs must have been laid in the first week of September. 

Doing large scale experiments of Varroa control is time-consuming and subject to the vagaries of the climate (and, as a molecular virologist, beyond me in terms of the resources needed). I have therefore used the well-established BEEHAVE program of colony development (from scientists in the University of Exeter; https://beehave-model.net/) to model the numbers of developing and adult bees, and the mite numbers in a colony. BEEHAVE by default uses environmental parameters (climate and forage) based upon data from Rothamsted (51.8°N). Using results from this model system, the bees present in the hive at the end of December – by definition the diutinus winter bees – were produced from eggs laid from early/mid August (Figure 4).

Whatever the precise date – and it will vary from season to season as indicated above – at some point in September the adult bee population starts to be entirely replaced with winter bees. Large numbers of these need to live until the following February or March to ensure the colony survives and is able to build up again once the queen starts laying.

When to treat – late summer

The numbers of pupae and adult bees present in the colony are plotted in Figure 4 using dashed lines. Adult bee number decrease in early spring until new brood is reared. The influence of the ‘June gap’ on pupal numbers is obvious. Brood rearing gradually tails off from early July and stops altogether sometime in late October or early November. The shaded area represents the period of winter bee production – from early/mid August until brood rearing stops. 

Figure 4. Winter bee production and mite levels – see key and text for further details

Mite levels are indicated using solid lines. The impact on the mite population of treating in the middle of each month from July to November is shown (arrowed and labelled J, A, S, O and N) using the colours green, blue, red, cyan and black respectively. The absolute numbers of bees or mites is irrelevant, but bees (pupae and adults) are plotted on the left, and mites on the right hand axis, so they cannot be directly compared. The miticide treatment modelled was ‘applied’ for one month and was 95% effective, reproducing many licensed and approved products.

Mite levels peak in the colony in late September to October. If treatment does not occur until this time of the season then the majority of winter bees will have been reared in the presence of large amounts of mites. Unsurprisingly, the earlier the treatment is applied, the lower the mite levels during the period of winter bee production. 

Rational Varroa control therefore involves treatment soon after the summer blossom honey is removed from the hive, so maximising the winter bees produced in the presence of low mite numbers. If you leave treatment until mid-September, you risk exposing the majority of winter bees to high levels of Varroa in the hive. If your primary crop is heather honey, which is not harvested until September, you may need to consider treating earlier in the summer – for example during the brood break when requeening or during swarm control.

Why treat in midwinter?

A key point to notice from Figure 4 is that, paradoxically, the earlier the miticide is applied, the higher the mite levels are at the end of the year. Compare the August (blue) and October (cyan) lines at year end for example. This is because mites that survive treatment – and some always do – subsequently reproduce in the small amount of brood reared late in the season. This is what necessitates a ‘midwinter’ treatment. Without it, mite levels increase inexorably year upon year, and cannot be controlled by a single late-summer treatment. Beekeepers bragging on social media that their mite drop after the winter treatment was zero probably applied the summer treatment too late to effectively protect their winter bees.

And when is midwinter?

Historically beekeepers apply the ‘midwinter’ treatment between Christmas and New Year. This is probably too late. The usual miticide used at this time is oxalic acid, a ‘one shot’ treatment that is ineffective against mites in capped cells. For maximum efficacy this must be applied when the colony is broodless. Brood rearing usually starts (if it ends at all, again this is climate-dependent) around the winter solstice. By delaying treatment until a lull in the Christmas festivities or even early January some mites will already be inaccessible in capped cells. 

Figure 5. Biscuit coloured (or a bit darker) cappings indicating brood rearing in this colony

I check my colonies for brood – either by looking for biscuit-coloured cappings on the Varroa tray (Figure 5) or by quickly inspecting frames in the centre of the cluster – and usually treat in November or very early December. If I cannot check visually I apply the treatment during the first extended cold spell of the winter. By treating when the colony is broodless I can be certain my intervention will have maximal effect.

What to treat with?

I have deliberately avoided – other than mentioning oxalic acid – specific miticides. Rational Varroa control involves the choice of an appropriate miticide and its correct application. Examples of incorrect or inappropriate miticide choice include; use of Apistan when resistance is known to be very widespread, use of Apiguard when the average ambient temperature is below 15°C (which makes Apiguard of little use for effective control in much of Scotland) or the use of Api-Bioxal when there is capped brood present. In addition, use of a half-dose or a reduced period of application will both reduce efficacy and potentially lead to the selection of resistance in the mite population. Used correctly – the right dose at the right time and for the right duration – the majority of the currently licensed miticides are be capable of reducing mite levels by over 90%. If they do not, use one that does. Miticide choice should be dictated by your environment and the state of the colony.

All together now

Most beekeepers grossly underestimate the movement of bees (and their phoretic mites) between colonies. Numerous studies have shown that drifting and (to an even greater extent) robbing can result in the transfer of large numbers of mites from adjacent and, in the case of robbing, more distant colonies. 

Gaffer tape apiary

Figure 6. Gaffer tape apiary …

Rational Varroa control therefore involves treating all colonies within an apiary, and ideally the wider landscape, in a coordinated manner. In communal association apiaries (Figure 6), where beekeeping experience and therefore colony management and health can vary significantly, this is particularly important. Coordinated treatment is only relevant in late summer when bees are freely flying.

Swarms

Swarms originating from unmanaged or poorly managed colonies will have high mite levels. The bee population in a swarm is biased towards younger bees; these are the bees that phoretic mites preferentially associate with. Studies have shown that ~35% of the mite population of a colony leaves with the swarm.

Figure 7. Varroa treatment of a new swarm in a bait hive…

Since swarms contain no sealed brood until ~9 days after they are hived oxalic acid is the most appropriate treatment. I usually treat them using vaporised oxalic acid late in the evening soon after they are hived (Figure 7). Even casts get this treatment and I have not experienced any issues with the queen not subsequently mating successfully. I’d prefer to have a queenless low-mite colony than a queenright one potentially riddled with Varroa.

Midseason mite treatment

The text above describes the mite management strategies I have used for several years. I apply Apivar immediately the summer honey is removed and treat with oxalic acid when broodless before the end of the year. Doing this has almost never required any additional midseason treatments; if mite levels are sufficiently low at the beginning of the season they cannot rise to dangerous levels before the late summer treatment. I still get winter colony losses, but they are almost always due to poor queen mating and rarely due to Varroa and viruses.

Figure 8. Queenright splits and the window(s) of opportunity

However, if midseason treatments are required – either because there are signs of overt infestation, because regular mite counts have shown there is a problem, or to have low mite colonies after the heather honey is collected – then there are two choices. Treat with MAQS which is approved for use when there are supers on the hive and, more importantly, is effective against mites in capped cells 2. Alternatively, treat during swarm control. With care, the majority of splits (e.g. the Pagden artificial swarm or the nucleus method) can be performed to give a broodless period for both the queenright (Figure 8) and queenless partitions. That being the case, a single application of an oxalic acid-containing miticide can be very effective in controlling the mite population.

Costs

Many beekeepers complain about the cost of licensed and approved miticides. However, some perspective is needed. A colony with low levels of mites will be more likely to survive overwinter, so reducing the costs of replacement bees. In addition, a healthy colony will be a stronger colony, and therefore much more likely to produce a good crop of honey (and potentially an additional nuc). Over the last 5-6 years my miticide costs are equivalent to one jar of honey per colony per year. This is an insignificant amount to pay for healthy colonies.

Summary

Rational Varroa control requires an understanding of the goals of treatment – protecting the winter bees and minimising mite levels for the beginning of the following season – and an appreciation of how this can best be achieved using miticides appropriate for the environment and the state of the colony. Like so much of beekeeping, it involves judgement of the colony and will vary from season to season and your location. I’ve applied my midwinter treatment as early as the end of October or as late as mid-December, reflecting variation in timing of the broodless period. Rational Varroa control also involves an understanding of the biology of bees and an awareness of the influence of beekeeping (e.g. crowding colonies in apiaries which increases mite and disease transmission) on our bees. However, none of this is difficult, expensive or time consuming … and the benefits in terms of strong, healthy, productive colonies are considerable.


 

Superinfection exclusion

Alpha

Beta

Gamma

Delta 

The majority of readers will identify these as the current circulating variant strains of Covid 1. The World Health Organisation decided upon this naming system as being easier and more practical to discussed (sic) by non-scientific audiences 2.

All viruses vary, and viruses with genomes made from ribonucleic acid (RNA) vary more than those which have deoxyribonucleic acid (DNA) genomes. This is because the enzymes that replicate RNA virus genomes do not have an error correction facility. 

The virus that causes Covid-19 is an RNA virus and so is Deformed wing virus (DWV), probably the most significant virus to bees and beekeepers (other than those who have Covid that is).

Virus variation

Why does virus variation matter? 

Or, asking the same question in a more roundabout way, why don’t RNA viruses evolve error correcting enzymes (after all, the DNA viruses have these … can it be that difficult?).

If the enzyme makes errors then that’s surely a bad thing?

Actually … for the particular ‘lifestyle’ that these viruses practice, errors and variation are a good thing.

They benefit the virus.

But you know all this already, even if you don’t think you do.

Covid cases caused by the delta variant

The SARS-Cov2 delta variant now accounts for at least 99% of cases of Covid-19 in the UK. It accounted for just 0.1% of cases in late February.

The delta variant is much more transmissible. It carries errors, or mutations as they’re more correctly known, that – for whatever reason – means it can be passed from person to person much more efficiently 3.

These mutations benefit the virus and allow it to spread further and faster 4.

If the mutations (‘errors’) that the virus acquires are beneficial – by increasing transmission, by expanding the cell, tissue or host range, by helping evade the immune response in a partially vaccinated population (!) for example – then the virus will successfully replicate and produce more viruses carrying the same mutation.

And a bunch of additional ones as well … acquired during the last round of replication.

Strains and types

At some point a virus acquires sufficient mutations from an earlier incarnation that it’s identified as a distinct strain.

For example, SARS-Cov2 is ~82% identical at the genome level to SARS-Cov1 which caused the SARS pandemic in 2003 5. SARS-Cov2 did not evolve from SARS-Cov1, but they share a common ancestor. They are different strains or types of coronavirus.

There are no hard and fast rules that define when a virus is considered a different strain or type. Often it’s historical, reflecting the geographic origin, or the source from which the virus was isolated. Different strains may exhibit different phenotypes – host range, transmission, disease etc. – but don’t have to.

For example (before we get back to honey bee viruses) there are three ‘types’ of poliovirus that are about 80% identical at the genetic level. They all cause exactly the same disease (poliomyelitis) and they replicate in an identical manner – same cell, tissue and host range for example. However, to the human immune system they ‘look’ different. The immune response to poliovirus type 1 will not protect you from infection with poliovirus type 3. That’s why the poliovirus vaccine contains a mixture of all three types, to protect you from all polioviruses.

You can even get infected with two types of poliovirus simultaneously, and the virus can replicate in the same cells in your gut … or, if you’re unfortunate, your brain.

I’ll return to dual infections shortly as it’s an important topic … and related to the study I’m going to discuss.

Deformed wing virus

There are two types of DWV, designated type A and type B 6.

Originally these had names that reflected their original isolation.

DWV type A was termed Deformed wing virus and was isolated from honey bees displaying the characteristic symptoms of developmental deformities shown in the image below.

DWV type B was termed Varroa destructor virus type 1 and was isolated from the ectoparasitic mite Varroa destructor.

It “does what is says on the tin” … DWV symptoms in a recently emerged worker

These viruses are very similar to each other. They are something like 85% identical at the level of the RNA genome. More importantly than this genetic identity (or perhaps similarity would be a better term to use here) is the fact that they appear to cause very similar diseases in honey bees.

Although early studies suggested there were some differences in their virulence, more recent work from Prof. Rob Paxton in Germany, from my lab, and from Dr. Eugene Ryabov and colleagues in the USA suggests these two types of DWV are actually very similar, at least in pathogenesis.

There do appear to be some differences, with the suggestion that type A does not replicate in Varroa whereas (full disclosure, the following study is from my lab) type B does. Undoubtedly, the other genetic differences between the types will confer some subtle variation in phenotype (effectively what they ‘do’), but – as far as beekeeping is concerned – they should probably be considered the same.

A protective, non-lethal type A DWV? 

All of which made a 2015 colony-level study 7 of DWV infection rather intriguing.

This reported the survival of colonies that were infected with a “non-lethal” type B strain which were protected from infection with the “lethal” type A strain 8. The authors summarised the significance of this study like this:

We propose that this novel stable host-pathogen relationship prevents the accumulation of lethal variants, suggesting that this interaction could be exploited for the development of an effective treatment that minimises colony losses in the future.

At the time there was a flurry of excitement and discussion about this 9

Superinfection exclusion

They proposed that the mechanism that prevented the infection with the type A strain was superinfection exclusion.

Virologists love mechanisms … 🙂 

Which brings me back to virus variation. 

Imagine a population of variant viruses trying to infect a new host … like a bee.

Survival of the fittest – selection for better replicating viruses from a mixed population

In mixed infections, a virus that has an advantage over the others in the population ends up ‘winning’ the competition for the resources of the host. They therefore make more progeny viruses.

One of the advantages could be that the virus simply replicates faster

Another – more subtle, but the same outcome – is that the virus prevents other viruses from infecting the same cell (and, by extension, host).

By excluding competing viruses it effectively ‘wins’ the competition.

Superinfection exclusion – one virus (type) can prevent infection by related but different viruses

And some viruses do exactly this using a variety of cellular mechanisms.

For example, many viruses turn off the expression of the cellular receptor (think of this as the door) they use to enter the cell. If there’s no receptor (no door) then another virus cannot enter.

With no other viruses to compete with in the same cell there can only be one type of virus produced 10.

There are other mechanisms as well, but we’ll stick with the receptor one as it’s easy to comprehend.

Superinfection of a cell containing a virus that has the ability to turn off the cellular receptor it uses will effectively exclude the second virus from replicating … hence superinfection exclusion.

What don’t we know about DWV?

A lot 🙁

We don’t know how it gets into cells. We don’t know a huge amount about its replication and we know precious little about the way it interacts with the cellular machinery (the ‘stuff’ in the cell that the virus hijacks to make more viruses) of the host. 

However, since 2015 we do know that all the types of DWV that have been carefully studied appear to be more or less equally virulent. None appear to be ‘non-lethal’ as claimed for the type A virus in the superinfection exclusion paper.

This prompted us to look in a bit more detail at the consequences of dual or sequential infections with DWV in the laboratory. 

Is there a precedence at work?

In mixed infections, does one virus always ‘win’ and predominate in the new virus population?

In sequential infections, does it matter the order in which the viruses are acquired?

And mixed infections are pretty much the norm for DWV infection. All bees, whether previously exposed to Varroa or in Varroa-free regions, appear to have low levels of DWV already present. If parasitised by the mite, these bees must experience a mixed infection.

In addition, studies we published several years ago 11 showed that recombinant viruses – essentially hybrids between type A and type B DWV – often predominated in heavily Varroa-infested colonies 12.

If mixed infections cannot occur, how do such hybrids form?

Some of the answers to these questions are in our recently paper published in the ISME Journal. Gusachenko, O. et al., (2021) First come, first served: superinfection exclusion in Deformed wing virus is dependent upon sequence identity and not the order of virus acquisition. ISME J (2021). https://doi.org/10.1038/s41396-021-01043-4

Mixed DWV infections

I don’t propose to give a pupa-by-pupa account of the studies we conducted. You can read the paper – it’s open access and (because Olesya ‘Alex’ Gusachenko, the lead author, did most of the writing) relatively easy to comprehend 😉

But here are a few highlights.

Over the last few years we have produced reagents that allow us to produce almost ‘pure’ stocks of type A, type B or hybrid type A/B 13 strains of DWV.

At least 99.99% of these stocks are of one DWV type. Note that there will still be variation within this population as the replication errors probably generate one mutation per virus in the population. We therefore refer to these virus stocks as near clonal.

Injection of honey bee pupae with any of these viruses resulted in very similar levels and kinetics of replication – all the viruses replicate as far and as fast as each other.

In mixed injections, when two viruses were administered simultaneously, both replicated to equivalent levels. 

We therefore found no evidence for the dominance of one strain over another.

Sequential DWV infections

But it got more interesting when we did sequential injections. We did these by injecting with one virus, waiting 24 hours and then injecting with a second virus.

Using type A and type B DWV both viruses had replicated to similar high levels (billions of virus per pupa) within 48 hours, irrespective of the order of addition. 

If superinfection exclusion was operating we would have perhaps expected type A to have prevented or reduced the replication of type B. However, that didn’t appear to be the case.

Competition between sequential infecting DWV isolates. VVV is type B, VDD is type A and VVD is a hybrid between them.

But, when we looked at sequential infections between type A and a type A/B hybrid we did see that replication of the second virus was delayed.

Delayed, but not stopped altogether.

It would take a complete post to describe the figure above 🙁 . We’ve quantified the virus present 5-7 days after sequential injections with type A (VDD), type B (VVV) or a hybrid virus (VVD) 14.

The columns labelled VDD→VVV or VVV→VDD show the viruses and order of addition. The dots represent the amount of virus present at 5 or 7 days post injection. When the viruses were more similar to each other – for example, the VVV→VVD or VVD→VVV pairs on the right – there was a greater impact on the replication of the virus added second.

The same but different

We extended these studies to look at sequential infections with two viruses that only differed by 4 nucleotides (the building blocks) of the 10,140 nucleotides in the RNA genome of DWV i.e. 99.6% identical.

Cunningly, these four differences allowed us to unambiguously identify which virus was replicating.

In this part of the study the virus added second did not replicate to detectable levels. 

So … our data clearly demonstrates that viruses that were more similar to each other were more likely to inhibit replication during sequential infections.

In addition, no individual virus type was dominant over any others. 

This didn’t look much like classic superinfection exclusion to us.

Red or green viruses

Not content with generating graphs and tables we went on to take photographs of virus infected pupae. 

You can’t beat a nice colour image when trying to impress the peer reviewers 😉 .

Remember that DWV is too small to see with even the most powerful light microscope. You could fit several billion on the head of a pin.

We therefore engineered the virus genome to ‘show’ us where it was replicating.

Green bees

We did this by introducing an additional gene that fluoresced green or red when under UV light. I’ve discussed green viruses before … the red version uses similar technology, but using a different fluorescent reporter gene.

DWV replication (showed by green fluorescent signal) in the head, wing and abdomen of honey bee pupae

Using the red or green viruses we showed very similar results to those described above. When we superinfected with a genetically similar virus, its replication was inhibited. When it was genetically more divergent it could replicate (and we could visualise it as red or green foci of infection in a variety of tissues of the developing pupa).

Red and green viruses

We also infected bee with the red and green viruses simultaneously. Most of the fluorescent foci of infection were red or green, but a small number were both red and green. 

Green (EGFP) and red (mCherry) expressing DWV coinfecting a honey bee pupa. Arrow indicates dually infected cells.

The most likely explanation for having both colours overlapping in the photograph was because the virus were replicating in the same cells in the honey bee pupa.

Since this was exactly the sort of situation that was needed to generate recombinants (hybrids or chimeras) between the two different DWV viruses we specifically looked for them 15.

And there were lots and lots of recombinants …

Recombination between DWV viruses. The size and position of ‘bubbles’ indicate the location and number of junctions.

The bubble plot above shows the location and frequency of junctions. One virus is plotted on the horizontal and one on the vertical axis. It’s a sort of two-dimensional map of the virus. Think of a junction as where one virus ends and the other starts. They are located throughout the DWV genome – hundreds of them.

This suggests that pupae infected with both type A and type B DWV will act as ‘factories’ for the production of thousands more different hybrid variants between the two viruses.

Most of these hybrids will grow poorly.

Many will be uncompetitive.

But some – like the delta variant of SARS-Cov2 – might be more transmissible.

And some could be more pathogenic.

Or – the nightmare scenario – both 😯 .

What’s this got to do with practical beekeeping?

Every time a beekeeper moves bees about s/he is also moving viruses about.

This happens when you move bees to out apiary, when selling a nuc or when importing a queen.

Double brood ...

Moving viruses (and bees) to a new apiary …

This will contribute to the constant mixing of DWV variants that occurs when bees drift between hives, when drones mate with queens, when phoretic Varroa jumps onto a bee that is robbing a collapsing colony.

There’s a difference of course.

All those bee-driven mixing events are local and small scale … a few bees and a few miles.

But if you import a nuc from Greece via Northern Ireland both the distance and number of bees (and hence number of viruses) is much greater.

Of course, most of this mixing will just generate more mixtures of viruses.

It will also generate more recombinants.

But there’s always the possibility it might throw up a highly virulent, highly transmissible variant.

Which would not be a ‘good thing’.

And if it does, a ‘non-lethal type A strain’ (should such a thing actually exist) is not going to help prevent infection by the mechanism of superinfection exclusion I’m afraid 🙁 .

Without doubt the best way to prevent infection is to minimise the mite numbers in your colonies. This is a subject I’ll be tackling again in a couple of weeks.

But, before I go, do we understand how the more closely related strains of DWV prevent superinfection? 

Yes … probably, and it’s all to do with the immune response of the bee

I’ll discuss this in the future as it’s a mechanism that could be exploited to produce bees immune to the ravages of DWV.


 

Queen introduction

I’m probably less qualified to write about queen introduction than almost any other aspect of beekeeping. This is not because I’ve not introduced any queens. Quite the opposite, it’s something I do more or less routinely many times a season. 

The reason(s) I’m really not qualified to discuss the topic are:

  • I almost exclusively use the method I first used and I’ve not done any side-by-side comparisons with other methods to determine which work ‘best’. I have a method that works well enough i.e. somewhere between most of the time and almost always. That’s good enough for me.
  • I’m not aware of any recent scientific studies on the subject so cannot use those to make informed decisions – or interpretations – of why some methods work and others don’t 1.

Nevertheless, not being qualified has never stopped me before 2 and it’s a topic that some beekeepers struggle with and many beekeepers worry about.

Successful introduction ...

Successful introduction …

So here goes …

Art or science?

David Cushman/Roger Patterson make the point that: 

” … you can have two colonies in the same condition, in the same apiary, on the same day and if you introduce a queen in the same condition into each, one will succeed and the other will fail.”

This doesn’t mean that 50% of introductions fail (although it reads that way). What he/they mean is that there appears to be no rhyme or reason why one succeeds and the other does not.

On another day, both might succeed … or both might fail 🙁

Is it therefore an art or a science?

I don’t know. All you can do is get the basics correct and cross your fingers …

For understandable reasons, beekeepers feel rather precious about their queens. In particular, beekeepers who do not rear their own queens (and so have no spares waiting in the wings) can get a bit paranoid about queen introduction. 

What if it goes wrong?

The colony will potentially be left irretrievably queenless and – if you purchased the queen – you’ll be £40 out-of-pocket 3.

If you do rear your own queens you can perhaps be a bit more blasé about queen introductions. Potentially you can also do the sort of side-by-side comparisons I mentioned above … though there aren’t many studies where this has been done in a rigorous way. 

Most seem to find a method that works for them and then stick with it … which is what I’ve done and what I’m going to describe.

This is what I mean by ‘get the basics correct’.

I’ll also mention an alternate method I irregularly use for what I consider to be really difficult situations and/or really valuable queens.

But before we get into the methodology, it’s worth making some general comments about the state of the recipient colony and the queen being introduced.

Is the colony really queenless?

Trying to introduce a new queen into a colony that is not actually queenless will not end well.

One or both of the queens will probably not survive the experience. Either the workers will reject (and slaughter) the incoming queen, or the queens will fight and may both be damaged and lost.

It is therefore important that the recipient colony is queenless.

By queenless I mean that there is no queen present.

I do not mean no laying queen present. If you try and introduce a new queen into a colony with a failed (non laying) queen or a virgin (unmated) queen you will have problems.

Sod’s Law is explicit in these instances … the valuable new mated laying queen will be lost 🙁

Queen above the QE

A virgin queen (in this instance on the wrong side of the queen excluder)

The very best way to be sure the colony is queenless is to remove the current queen before introducing the new one. That necessitates finding the queen in the first place. 

What if you can’t find the queen but you’re sure that the colony is queenless?

Well, there are only two possibilities if you can’t find the queen, these are:

  1. The colony is queenless … you’re good to go.
  2. The colony is not queenless … but you’ve looked so hard for so long they’re now disturbed and running manically around the frames, getting more and more agitated and angry. Neither the bees or you are any sort of state to allow the queen to be discovered. Close the hive up. Have a cup of tea. Try again tomorrow.

I discussed methods of determining whether the colony is queenright (though not by extrapolation, the opposite i.e. queenless – see below) last season. Towards the end of that post I described the addition of a ‘frame of eggs’ to determine if the colony is queenright or not. I won’t repeat all the details here.

If the colony draw queen cells on the introduced frame then you can be sure that the colony is queenless. See (1) above … you’re good to go 🙂

Not queenless, but not queenright

That same post describes the concepts of queenright and queenless.

A colony that is queenright has a mated queen capable of laying fertilised eggs (though she may temporarily not be laying, for example due to a dearth of nectar).

A queenless colony contains no queen.

But there’s an intermediate stage … or potentially two intermediate stages if you allow me a little leeway.

A colony containing a failed queen that’s either not laying at all (and not going to restart), or only laying drone (unfertilised) eggs is neither queenright not queenless. This colony will not draw queen cells on the introduced frame. You cannot safely introduce a new queen into such a colony before first finding and removing the failed queen.

A colony containing laying workers will also not 4 produce queen cells from the introduced frame of eggs. 

Laying workers ...

Laying workers …

A colony with laying workers behaves as though it’s queenright but is actually queenless. It’s not really an intermediate stage, but the consequences are the same. Again, they are highly unlikely to accept an introduced queen.

Deal with the laying workers first and then requeen … and good luck, laying workers can be a nightmare 🙁

OK … let’s assume the colony really is queenless … what’s the easiest way to introduce a new queen?

Add a sealed queen cell

Almost without exception, a queenless colony can be requeened by adding a sealed queen cell. The virgin queen will emerge, go on one or two mating flights and return and head the colony. This method of queen introduction is almost foolproof in my experience. 

Where do you get the queen cell from? Another colony, your mentor, a friend in your beekeeping association, a local queen rearer … necessity is the mother of invention 5.

Assuming the cell is a natural queen cell … cut the queen cell out of the comb with a generous amount of surrounding comb. Don’t risk damaging the queen cell. Keep it vertical … there are stages during development when the pupa is susceptible to damage. Ideally choose and use a cell 24-48 hours from emergence as they’re a lot more robust late in the development cycle.

Use your thumb to make an indentation towards the top of a frame near the centre of the broodnest, above some capped and emerging brood. Using the generous ‘edge’ of comb surrounding your chosen queen cell push this into the indentation so the cell is secure. Close up the colony and a) check for emergence in 48 hours or so 6 and b) a fortnight later for successful mating.

Adding a grafted queen to a colony

If the cell is from a grafted larvae it is even easier … press the plastic cell cup holder into the comb and push the frames together. I describe this in a recent discussion of grafting.

How successful is this method of ‘queen’ introduction?

I’d estimate at least 85%.

A very small percentage of queen cells fail to emerge (or rather, the queen fails to emerge from the cell … but you knew what I meant 😉 ).

A slightly larger percentage of queens fail to mate (or fail to return from a mating flight). But, even in a bad season, it’s rarely more than 10-15%.

The new queen is accepted by the colony because she emerged there and they all live happily ever after 😉 .

What?

I know, I know … that’s not really queen introduction.

You’re right. But it works. Very well.

These are the two methods I use for queen introduction.

Candy-plugged queen cage

I have a large supply 7 of JzBz queen introduction and shipping cages. 

JzBz queen cages

JzBz queen cages

I really like them because they were free they are reusable, they have a tube-like entrance that can be plugged with candy/fondant and they have a central region to protect the queen from aggressive workers outside the cage. 

Some cages offer no areas of refuge for the queen and workers can damage the queen through the perforations. Avoid cages that are all perforations.

The JzBz cages can be purchased with a removable plastic cap (shown below the cage in the image). These fit over the end of the tube and can seal the cage until you judge the colony is likely to gracefully receive the new queen … as described below 8.

JzBz queen introduction and shipping cage

Using a JzBz cage for queen introduction:

  • Plug the tube of the JzBz cage with queen candy or fondant. Queen candy can be purchased commercially and kept frozen for long periods. I almost always use fondant these days as I have spare boxes of the stuff from autumn feeding.
  • Add a short piece of wire or a cocktail stick through the perforations at one end of the cage to hang the cage – entrance tube pointing downwards – between two frames. Do this before adding the queen to avoid risking skewering the queen at a later stage 9
  • Place the queen in the cage without any attendants (see below for comments on removing them). Close and seal the cage. Seal the candy tube with the plastic cap.
  • Hang the cage in the centre of the broodnest, above some emerging brood. Leave the colony for 24 hours.

The idea here is that the colony gets the chance to accept the new queen without getting the opportunity to slaughter her.

Look for signs of aggression

Colonies that have been queenless for a few hours (say 2-24) before adding the new queen are usually very willing to accept a replacement. Adding a queen immediately after removing the old queen is likely to result in some aggression to the caged queen.

Check the colony after 24 hours. I usually lift the cage out and place it gently on the top bars to observe the interaction of the workers and the queen.

Checking for aggression

If the colony show no aggression to the caged queen – look for bees trying to sting through the cage or biting at the cage – then remove the plastic cap and re-hang the cage between the frames.

If they show aggression leave them another 24 hours and check again 10

Once you remove the cap the queen will be released by the workers after they eat through the candy/fondant. This takes just a few hours. 

Check again a week later to ensure the colony has accepted the queen.

Nicot introduction cage

I use the method described above for almost every queen I introduce. 

The only exception is if I have to requeen a colony that has previously not accepted a queen using the method described above. Usually such a colony will also be broodless (just based on the timings of determining they are queenless and failing once to successfully introduce a queen). 

Under these circumstances I use a Nicot queen introduction cage.

Nicot queen introduction cages

I find a frame from another colony with a hand-sized patch of emerging brood. The comb needs to be level so that the cage can sit on top without gaps for the queen to escape.

Then do the following:

  1. Remove all the bees from the frame and place the Nicot cage over the brood using the short plastic ‘legs’ to hold it into the comb 11.
  2. Secure the cage in place using one or two elastic bands.
  3. Introduce the queen through the removable – and eminently losable 12 – door.

In practice it’s easier to do this in the order 3-1-2 … place the queen on the frame, cover with the cage and then secure it with the elastic band.

Add the frame and cage to the hive, locating it centrally. Push the frames together. 

The emerging workers will immediately accept the queen and feed her. Other workers will feed the queen through the edges of the cage.

One corner of the cage has an entrance tunnel that can be filled with candy/fondant. I don’t think I’ve ever used this. In my experience the colony releases the queen by burrowing under one edge of the cage after a few days. If they don’t, check and remove the cage a week later.

I don’t think I’ve ever failed to successfully introduce a queen using one of these cages, but it’s a relatively small sample size.

Thorne’s sell a metal mesh version of this cage that has integral ‘legs’. I’ve not used it, but the principle is the same. Keep it in a box or the sharp cut metal edges will butcher your fingers – it’s difficult picking up queens with heavily bandaged digits.

You could also ‘fold’ your own from mesh floor material. One with deeper ‘sides’ could be pushed down to the midrib of the comb, so reducing the chances of the bees burrowing under the edge of the cage.

Mated or virgin? 

I use the JzBz cage for introducing either mated or virgin queens. I’m not aware of any significant difference in the acceptance rate between them. 

However, it’s worth noting that acceptance is dependent upon essentially ‘matching’ the expectations of the colony with the state of the queen. 

A virgin queen will be less likely to be accepted by a colony from which a mated laying queen has recently been removed. Leave them 24-48 hours. 

Likewise, I remove nearly mature queen cells from a colony I’m requeening with a mated queen. I don’t want to risk an early-emerged virgin queen from ‘raining on the parade’ of the introduced queen.

I’ve only used the Nicot cage for mated queens. Since the latter is usually used for a broodless colony I want the minimum possible delay before there is new brood in the colony.

Alone or with attendants?

If you purchase a queen and receive her by post there will be a few workers caged with her.

I always remove these although some suggest that they do not adversely influence acceptance rates 13. I remove them because I’m a bit paranoid about viruses … these workers come from an ‘unknown’ hive (quite possibly not the same one that the queen came from) and will carry a potentially novel range of Deformed wing virus variants (and possibly others as well).

I don’t want these in my hive so I remove the workers

It’s also worth noting that Wyatt Mangum has an interesting report in American Bee Journal indicating that the presence of attendants significantly increases the acceptance time 14 for an introduced queen 15. In some cases the presence of attendants resulted in the colony showing aggression for longer than it took for the bees to eat through the candy plug … that’s not going to end well for the queen.

The safest way to remove attendants is to open the caged queen in a dim room with a single closed window. The bees will fly to the window (perhaps with a little encouragement).

A mated queen probably will not fly at all and can be re-caged. A virgin queen can fly well and will also end up at the window. Gently grab her by her wings and re-cage her.

You can do all this in the apiary … it requires confidence and dexterity. I know this because I recently tried it with a virgin queen in my apiary, using lashings of overconfidence and hamfistedness.

She flew away 🙁

Inevitably you can buy a gadget to help you with this – the queen muff

Conclusions

There is always a slight risk that queen introductions will not be successful. The queen pheromones have such a fundamental role in colony maintenance that disrupting them – or suddenly changing them – may lead to rejection. 

However, the methods described above are sufficiently successful that I’ve not found the need to look for better alternatives. They’re also sufficiently fast that I’m not tempted to try some of the ‘quick and dirty’ approaches 16 to save time.

Finally, it’s worth noting that it is usually easier to requeen a nucleus colony than a full hive. If I ever bought one of those €500 breeder queens I’d introduce her to a nuc first and then unite the nuc back with the original colony.

But that’s not going to happen 😉


 

Fainting goats … and queens

Myotonia congenita is a genetic disorder that affects the muscles used for movement. Myotonia refers to the delayed relaxation of these skeletal muscles, resulting in a variety of obvious symptoms including temporary paralysis, stiffness or transient weakness.

In humans these symptoms are often manifest as difficulty in swallowing, gagging and frequent falls. Children are affected more than adults. One of the most dramatic manifestations are the falls (‘fainting’) that can occur as a result of a hasty movement. 

Although physiologically distinct, ‘fainting’ is a reasonably accurate description of the sudden loss of movement and the transient nature of the disorder. Like fainting, loss of movement is usually quickly resolved. However, unlike fainting, myotonia congenita involves muscular rigidity or stiffness, so more closely resembles catalepsy.

Genes

There are two types of myotonia congenita, termed Thomsen disease and Becker disease, both of which are usually associated with mutations in the gene CLCN1 1. This encodes a chloride channel (a ‘hole’ through the cell membrane that allows the transfer of chloride ions) critical for muscle fibre activity. 

Cartoon of a transmembrane chloride channel.

With loss-of-function mutations in CLNC1 the muscle fibre continues to to be activated. When stimulated, for example if the fibre is triggered to suddenly contract for jumping or running (or  to stop a fall), the muscle fibre is hyper-excitable and continues to contract, and shows delayed relaxation

Around 1 in 100,000 people exhibit myotonia congenita, though it is about ten times more common in northern Scandinavia. Treatment involves use of a number of anticonvulsant drugs.

The same loss-of-function CLCN1 mutation in humans is seen in symptomatically similar horses, dogs … and goats.

Goats

In the late 19th century four goats were imported to Marshall County, Tennessee. Their strange behaviour when startled was first described in 1904 and defined as a congenital myotonia by Brown and Harvey in 1939. 

The eponymous Tennessee fainting goat

These pre-war studies formed the basis of of our understanding of both the physiology and genetics of myotonia congenita, though the specific mutation in the CLCN1 gene was only confirmed several years after it had been identified in humans.

Since then myotonic goats have become an internet staple, with any number of slightly distressing (for me at least, if not for the goats) YouTube videos showing their characteristic fainting when surprised or frightened 2.

Don’t bother watching them.

If you want to see a fainting goat in action watch little ‘Ricky’ jump up onto a swinging seat on the National Geographic website.

It’s a perfect example.

He jumps up, gets a mild fright as the swing moves, goes stiff legged and simply rolls over and falls to the ground. A few moments later he’s back on his feet again, looking slightly shaken perhaps, but none the worse for wear.

Queens

All of that preamble was to introduce the topic of fainting queens. 

A fainting queen

This was a subject I’d heard about, but had no experience of until last week.

Periodically it gets discussed on Beesource or the Beekeepingforum – usually the topic is raised by a relatively small-time amateur beekeeper (like me) and it gets a little airtime before someone like Michael Palmer, Michael Bush, Hivemaker or Into the Lion’s Den 3 shuts down the conversation with a polite “Yes, I see it a few times a year. They recover”, or words to that effect.

Since these commercial guys handle hundreds or perhaps thousands of queens a year I think we can safely assume it’s a relatively rare phenomenon. 

Since I don’t handle hundreds or thousands of queens a year – and you probably don’t either – I thought the incident was worth recounting, so you know what to expect should it ever happen.

And to do that I have to first explain the fun I had with the first of the two queens in the hive I was inspecting.

A two queen colony

It was late afternoon and I was inspecting the last of our research colonies in the bee shed.

The hive had two brood boxes and a couple of supers. Nothing particularly surprising in that setup at this time of the season; the colony was quite strong, the spring honey had been extracted and a couple of supers had been returned to the hive for cleaning.

However, it wasn’t quite that straightforward. 

The lower brood box had been requeened ~3 weeks earlier with a mature queen cell from one of my queen rearing attempts. I’d seen that the virgin had emerged and restricted her to the lower box at my last visit. 

I’d added a queen excluder (QE) over the lower box with the intention of removing all the old frames above the QE once the brood had emerged.

However, at that last visit I’d ended up with a good looking 4 ‘spare’ virgin queen. Although I had no need for her at the time, and no time to make up a nuc 5, I decided to put her in a fondant-plugged introduction cage in this upper box.

This ‘upper’ queen couldn’t fly and mate in the week I was away, but I reasoned that I could merge the colony with the bottom box if the ‘lower’ queen failed to mate 6.

So, after adding the virgin queen to the top box I added a second QE and the two supers.

She can fly …

Having removed the supers and the upper QE I carefully inspected the upper box looking for the virgin queen who had been released from the cage

No sign of her 🙁

I went through the box again.

Time to try some of the ‘queen finding tricks’.

I moved three frames out of the way having examined them very carefully. The remaining 8 frames were then spaced out as four, well separated, pairs. I let the colony settle for a few minutes and then looked at the inner face of each pair of frames.

No sign of her 🙁

I looked again … nada, rien, niets, nunda, dim byd and sod it 7.

The obvious conclusion was that the colony had killed the queen after releasing her from cage. 

How uncharitable.

I reassembled the upper brood box and lifted it off the lower QE, in preparation to leave it outside the shed door while I went through the lower box. 

As I carried the brood box to the door I briefly looked up and saw a 8 virgin queen climbing up the inner pane of one of the shed windows, flapping frantically and fast approaching the opening that would allow her escape.

For obvious reasons I have no photographs of the next few minutes.

Bee shed window ...

Bee shed window …

For those unfamiliar with the bee shed windows, these have overlapping outer and inner panes, so are always open. They provide a very effective ‘no moving parts’ solution to clearing the shed of bees very quickly.

Which was the very last thing I wanted at that moment 😉

… rather well

I had a brood box and hive tool in my hands, the shed door was wide open, there was all sorts of stuff littering the floor and the virgin queen was inches away from making a clean getaway.

It’s worth noting that when virgin queens are disturbed and fly they almost always return to the hive. However, the hives in the shed have a single entrance and all the hives were already occupied with queens. I couldn’t let her fly and hope for the best … it probably wouldn’t end well.

By balancing half the brood box on an unoccupied corner of an adjacent hive roof I made a largely ineffective swipe for the queen, but disturbed her enough she flew away from the window in spirals around my head.

I    s  t  r  e  t  c  h  e  d    to reach the shed door and pulled it close, so reducing the possible exits from eight to seven. A small victory.

I put the brood box safely on the floor, leaning at an angle against the hive stand 9, and abandoned the hive tool.

The next 5 minutes were spent ineptly trying to catch the queen. When she wasn’t flying around the shed (where the lighting isn’t the best) she usually made for the same window.

The one behind the hive with four supers stacked on top 🙁

After a few more laps of the shed, dancing around the precariously balanced brood box and reaching around the hive tower for the window, I finally caught her.

And caged her 10.

I’m looking for publisher for my latest book, ‘Slapstick beekeeping’. If any readers know of a publisher please ask them to contact me.

After all that I should have had a little rest. I’d had enough excitement for the afternoon 11.

But there was still the queen in the bottom box to find and mark.

Feeling faint

The queen in the bottom box was mated and laying well. 

I made a near-textbook example of finding her 12.

After moving aside a few frames I should have announced (to the non-existent audience), She’s on the other side of the next frame … ” (the big reveal) ” … ah ha! There you are my beauty!”.

Holding the frame in one hand I checked my pockets for my marking cage 13.

All present and correct.

I then calmly picked her up by her wings. She was walking towards me, bending slightly as she crossed over another bee, so her wings were pushed up and away from her abdomen.

A perfect ‘handle’.

I didn’t touch her abdomen, thorax or head.

A swooning queen

And, as soon as I lifted her from the frame, she fell into a swoon and ‘dropped dead’.

This is an ex-parrot

Her wings were extended to the sides, her abdomen was curled round in a foetal position and she appeared completely motionless.

It is pining for the fjords

I dropped her into the marking cage and took the photo further up the page.

It was 6:49 pm.

For several minutes there was no obvious movement at all. Her legs and antennae were immobile. She showed no sign of breathing.

I gently shook her out onto a small piece of Correx on a nuc roof to watch and photograph her. I picked her up by the wing and held her in my palm … perhaps she needed some warmth to ‘come round’.

Was that a twitch?

Or was that me shaking slightly because I’d inadvertently killed her? 

Several more minutes of complete catatonia 14 passed … and then a gentle abdominal pulsing started.

This was now 10-11 minutes after I’d first picked her up.

Which got a bit stronger and was accompanied by a feeble waggle of the antennae.

And was followed a minute or so later by a bit of uncoordinated leg flexing.

And after 15 minutes she took her first steps.

It looked like she’d been on an ‘all nighter’ and was still rather the worse for wear.

I slipped her into a JzBz queen cage, sealed it with a plastic cap, and left it hanging between a couple of brood frames.

From picking her up to placing the caged queen into the brood box had taken 24 minutes.

Caged queen after fainting (and recovering … more or less)

I reasoned that if …

  • she fully recovered they’d feed her through the cage and I could release her the following morning
  • I’d released her immediately and she’d acted abnormally the colony might have killed her off
  • she did not recover I would at least be able to find the corpse easily ( 🙁  ) and so could confidently requeen the colony (with the virgin I’d tucked away safely in my pocket)

The following morning the cage was covered in bees and she looked just fine, so I released her. 

Somewhere under that lot is the recovered queen – still caged

She walked straight down between the frames as though nothing untoward had happened.

I didn’t have the heart to mark and clip her … I didn’t want to risk her ‘fainting’ again and, if she had, didn’t have the time to hang around while she recovered 15.

So was this ‘fainting’ myotonia congenita?

I suspect not.

Another name for the Tennessee fainting goat is the ‘stiff-legged’ goat. This reflects the characteristic rigidity in the limbs when the muscles fail to relax. The queen’s legs were curled under her, rather than being splayed out rigidly.

However, this interpretation may simply reflect my near complete ignorance of the musculature of honey bees 😉

However, I do know that the basics of muscle contraction and relaxation are essentially the same in invertebrate and vertebrate skeletal muscle. There are differences in the innervation of muscle fibres, but the fundamental role of chloride channels in allowing muscle relaxation is similar.

Therefore, for this fainting queen to be affected by myotonia congenita she should have a mutation in the CLCN1 gene encoding the chloride channel.

Although the honey bee genome has been sequenced a direct homolog for CLCN1 appears not to have been identified, though there are plenty of other chloride channels present 16

The majority of the 60 or so mapped mutations associated with myotonia congenita (in humans) are recessive. Two copies of the mutated gene (in diploids, like humans or female honey bees) are needed for the phenotype to occur.

Of course, drones are haploid so it should be easier to detect the phenotype.

I’ve never heard of drones ‘fainting’ when beekeepers practise their queen marking skills on them. Have you?

Repeated fainting

I’ll try to mark and clip this queen again.

It will be interesting to see if she behaves in the same way 17.

A quick scour of the literature (or what passes for the ‘literature’ on weird beekeeping phenomena i.e. the discussion fora) failed to turn up examples of the same queen repeatedly fainting.

Or any mention of daughter queens showing the same behaviour.

All of which circumstantially argues against this being myotonia congenita.

However, there are many other causes of sudden fainting (from the NHS website):

  • standing up too quickly – (low blood pressure)
  • not eating or drinking enough
  • being too hot
  • being very upset, angry, or in severe pain
  • heart problems
  • taking drugs or drinking too much alcohol

… though I can exclude the last one as my bees are teetotal 😉

So, there you have it, a brief account of a cataleptic queen … and her recovery.


Notes

A fortnight after the events described above I clipped and marked the queen. I did everything the same – picked her up by the wings in the shed (so again not exposed to bright sunlight – which may be relevant, see the comment by Ann Chilcott).

She (the queen) didn’t faint. She behaved just like the remaining 4 queens I marked on the same afternoon.

So no repeat of the ‘amateur dramatics’ 🙂