Category Archives: Queen failure

It makes you go blind

Synopsis: There is a sexual arms race between the queen and the drones she mates with. The queen needs to mate with multiple drones to maximise colony fitness. Conversely, it’s in the interest of individual drones to reduce the number of additional partners who mate with the queen. Recent studies have demonstrated that drones reduce repeat mating flights by impairing the eyesight of the queen. Potential implications of this for practical beekeeping are discussed.

Introduction

Honey bee queens are described as polyandrous 1 because they copulate with multiple drones during one or more mating flights taken shortly after emergence.

These multiple matings are a risky business 2.

It takes longer to mate with multiple drones than it does to mate with one, but this time is minimised by reducing the number of mating flights. Rather than leaving the hive, mating once, returning and then repeating the process, the queen flies some distance to a drone congregation area and copulates with multiple drone before returning to the hive.

Shallow depth of field

One of many …

I’ve discussed the location and locating drone congregation areas previously and the distances the queens and drones respectively fly to reach these (which are different to avoid inbreeding).

Between the queen returning from the mating flight and the onset of egg laying there is a delay of a few days. During this period the queen is storing the sperm from the drones in her spermatheca. These are the sperm storage organs within which sperm stays active for years … a necessity as, after the onset of laying, the queen will not go on any more mating flights.

Perhaps surprisingly, only about 3-5% of the sperm transferred from each drone is stored by the queen.

I hope that makes you wonder why she bothers mating with so many drones … it should.

Polyandry and hyperpolyandry

Just before I explain why she only stores 3-5% of the sperm from each of several drones, rather than storing it all from one twentieth the number (and thereby reducing the risks of longer mating flights) of drones, I need to explain the poly bit of polyandry.

How many drones does the queen mate with?

The usual figures quoted are in the high teens, with a range extending from single digits into the low forties. These numbers are determined using a variety of different techniques, at least some of which are likely to underestimate the actual number of drones.

Marked queen surrounded by a retinue of workers.

Here’s one I made earlier …

Think of it like this, if you have a large population of something – like beekeepers – how many would you have to ‘sample’ to find one called ’David’.

Not many, it’s a common name.

But what about ’Atlas’ or ’Zebedee’?

You’d have to sample a lot more apiarists to find any with these rarer names, though I bet they’re out there somewhere. You might even have to use a different way to screen the population.

And it’s the same when determining the numbers of drones that the queen mates with.

Search and ye shall find – detecting rare patrilines

When you use a method that specifically looks for rare patrilines – essentially genetically distinct offspring fathered by different drones – you can find them. This suggests that the queen probably mates with more than the 15-19 drones usually quoted, and that hyperpolyandry is perhaps a better term to describe the mating behaviour of queen honey bees.

There’s evidence that these very rare patrilines (so-called ‘Royal patrilines’) are preferentially selected when rearing queens under the emergency response.

Colony fitness

So now we’ve defined what the poly in polyandry means … but we still don’t know why the queen risks all those aerial shenanigans to mate with so many different drones.

By mating with multiple drones she ensures that the workers in the colony are genetically diverse. This genetic diversity increases the rather-difficult-to-grasp concept of colony ‘fitness’. In this instance fitness is used to mean a combination of adaptability, resistance to stress or pathogens, increased foraging activity, better overwinter survival etc.

I’ve discussed this concept before and suggest you revisit that post for all the gory details.

The bottom line is that colonies that are headed by queens that are mated with very many drones (50+) produce more brood, have better disease resistance and have many other desirable traits (that benefit both the colony and the beekeeper).

The final piece of this introductory jigsaw I need to mention is that drone sperm is used randomly. It’s not a case of ‘first in, last out’. The 3-5% of sperm stored from each drone is mixed thoroughly in the spermatheca.

This makes sense in light of the comments above about colony fitness. If the sperm were used in batches from each drone you’d have cohorts of young bees being produced that had reduced genetic diversity, thereby potentially compromising colony fitness.

It takes two to tango

But let’s think about the poor drones for a moment.

Drones have two fates (excluding getting eaten by a bee eater); they either die while mating with a queen, or they get turfed out of the hive and starve to death towards the end of the season.

If the drone fails to mate with a queen he’s genetic dead end.

If he does mate with a queen there’s a good probability that the genes he carries will be passed on to the following generation.

There is therefore a lot of competition for the queen in the drone congregation areas (DCA).

The drones, once sexually mature, fly every (suitable) day to several DCAs, one after the other. In addition, they fly relatively short distances from the hive to maximise their time within the DCAs.

Heat map of the landscape used by drones – bright spots are DCA’s

This competition is intense, and it doesn’t stop once the drone has mated (and died).

If a queen mates with a relatively small number of drones – let’s say 10 for the sake of argument – the chance of the sperm from any one of those drones being used to fertilise an egg is much greater than if the queen had mated with 50 drones.

The fewer drones the queen mates with the better the chances that the genes from any one of her successful suitors will be passed on to the following generation.

Paradoxically, it therefore benefits the drone 3, if the queen mates with fewer other drones.

And, remarkably, drones have evolved a way to reduce the number of additional drones that a queen mates with.

A sexual arms race

Before I describe the mechanism, it’s worth emphasising here that best interests of the colony are served by the queen mating with many drones, but those of the drones are best achieved by limiting the polyandrous activity of the queen.

These two processes are therefore in direct competition.

There are some additional subtleties.

If the drone simply prevented the queen from mating again 4 it would be detrimental if that drone was the first with which the queen mated. The resulting colony would have little genetic resilience and would be unlikely to survive.

Any one drone must therefore allow the queen to mate with sufficient other drones to ensure colony fitness.

In addition, the more mating flights that a queen goes on, the greater the chances she will be predated by a passing bird, or get lost on the return flight.

From the drones point of view it would probably be beneficial for the queen to go on only one mating flight, but that she mates with sufficient (but no more than that) drones on that flight.

And finally, before I get to the mechanism by which all this is achieved – a compromise solution, like all the best solutions – I’ll remind you that studies have shown that queens go on about 5 mating flights spread over 3, usually successive, days.

Love is blind

At least, too much love is … 😉

Liberti and colleagues have recently published a snappily titled paper on how drones reduce the number of mating flights taken by a queen. The paper is Open Access so you can get all of the nitty-gritty details I don’t have time, energy or intelligence to include in the summary below.

The paper is:

Seminal fluid compromises visual perception in honeybee queens reducing their survival during additional mating flights by Joanito Liberti et al., (2019) eLife 2019;8:e45009

As with all science, the results published in this paper were a continuation of earlier studies of queen honey bees. In particular, these included studies by some of the same authors who had showed that seminal fluid contained proteins that had the ability to interact with neurons.

In addition, in Drosophila melanogaster (the fruit fly, and genetically best studied insect) there was evidence to suggest that seminal fluid promotes fast oviposition and reduces the willingness of females to seek additional copulations.

Drosophila mating in captivity

Now, Drosophila mating behaviour is very different to that of honey bees, but there was clearly a precedent here in which some of the components of seminal fluid – the ‘carrier’ that keeps sperm alive and motile and protects against pathogens – influenced subsequent mating in insects.

Or the lack of mating.

The study by Liberti et al., involves an elegant combination of hardcore molecular gene expression analysis coupled with electroretinography 5 and field work. I’ll skip briefly through the first two of these and provide a bit more detail on the last.

Analysis of gene expression

Virgin queen bees were instrumentally inseminated with seminal fluid (i.e. no sperm) or a control saline solution. Subsequent analysis of the brains of the bees – using a method called RNA-Seq which allows the qualitative and quantitative changes in gene expression to be accurately determined – demonstrated reproducible changes in the gene expression of dozens of genes.

Venn diagram of differential gene expression in instrumentally inseminated queen bees

Detailed analysis of which genes had changed in expression showed that several so-called signalling and metabolic cascades were modified in response to seminal fluid, and many of these mapped to the phototransduction pathways i.e. those involved in sight.

Several of the genes that were detected encoded proteins that were implicated in the conversion of light into the electrical signals in photosensitive electrical cells.

Inevitably, that one sentence has probably confused half the readers that have persevered to this point in the post …

Essentially what this means is that there are components within drone seminal fluid that change the ability of the queen to perceive light, or to see.

So, do they?

Visual perception of queens

The gene expression studies in this paper are complicated (for a molecular biologist). The electroretinography is an order of magnitude more complicated for this molecular biologist to understand … but here goes.

Electroretinography involves measuring the electrical signals generated by particular neurones that are connected to the compound eyes and ocelli 6. This allows the consequences of the changes in gene expression to be determined in terms of the vision of the queen bee.

These studies showed that queens instrumentally inseminated with seminal fluid had lower responses to low frequency flickering light, and that that this response (or lack of response) increased on the second day after insemination.

There were additional changes in the response of the ocelli in queens inseminated with seminal fluid.

Taken together, these results show that queens exposed to seminal fluid experience reduced visual performance.

They are not blinded, but their vision is impaired.

Does this visual impairment have any influence on their mating behaviour?

Mating flight behaviour

Finally, we come to something that’s a bit easier to comprehend, not least because I’ve previously discussed the technology used – the RFID tagging of individual bees to monitor their flight frequency and duration.

RFID-tagged queens (34 in total) were instrumentally inseminated (either mock, or seminal fluid or semen) and subsequently monitored when going on mating flights. Those receiving either seminal fluid or semen were more likely to get lost on these flights, and repeatedly triggered the hive entrance sensors, suggesting they were disorientated by sunlight after leaving the hive.

Of the 21 queens that returned, 81% went on mating flights of more than 7 minutes which was considered a conservative threshold for a completed mating flight i.e. flight to a DCA, mating(s) and return to the hive, and about 50% laid worker brood.

Notably, of the 17 queens that went on ‘successful’ (by duration, not necessarily by outcome) mating flights, those receiving the control saline solution left 1-2 days later than those that had received seminal fluid or semen.

Seminal fluid and semen induce alterations of mating flight behaviour in honeybee queens

These results show that exposure to seminal fluid induces significant changes in queen mating flight behaviour, presumably as a consequence of the alteration to the vision of the queen.

Therefore, the implication from these results is that proteins in the seminal fluid of drones impairs the visual perception of queens, thereby reducing the likelihood that the queen will embark on additional mating flights.

Queens that had already mated (or been instrumentally inseminated in this study) were more likely to get lost on subsequent mating flights, and embarked on these flights earlier.

But what about swarming?

The hive – or a natural nest site – is a low-luminance environment. Queens do not need fully functional eyesight once they have returned from their mating flights. In the hive communication is non-visual, mediated by pheromones, contact, vibrations and sound.

However, although a queen only goes on a few mating flights, she will also leave the colony if it swarms.

Swarm of bees

Swarm of bees

What are the implications for the this study on the eyesight of queens during swarming?

This isn’t really discussed in the paper, but I think there are two likely scenarios:

  • the changes in visual perception by the queen are transient and return to ‘normal’ after a few days, weeks or months
  • swarming is a fundamentally different activity in which thousand of bees leave the hive and for which accurate vision is not needed by the queen.

There’s a world of difference between embarking alone on a mating flight of several kilometres and having to return to the exactly the same location, and leaving on a one-way trip with a swirling mass of attendees with dozens of scout bees leading the way.

Further studies will be needed to determine whether the changes in vision are transient or permanent, as well as to identify the ‘active ingredient’ in seminal fluid that is responsible for the degradation of the mated queen’s vision.

I also think further studies will be required to determine the relationship between dose and timing of the response.

How long does it take for the reduction in visual perception? If the first and second mating flight are taken on successive days is the “return rate” greater than if they are taken a few days apart?

How many drone matings are needed to reduce the visual acuity of the queen? I would predict that this would be a number consistent with the lower estimates of polyandrous matings needed to generate fitness in the resulting colony.

And implications for practical beekeeping?

Perhaps none directly, though I’m interested in the answers to the questions I posed in the paragraphs above.

In an area with low drone densities and those with shall we say ‘variable’ weather – such as my apiaries on the west coast of Scotland (or for that matter, any beekeepers living in remote northerly areas with just a few hives) – is colony fitness compromised by reduced matings?

An isolated apiary

Conversely, is mating success lower because more queens fail to return from subsequent mating flights that they have to take to try and mate with enough drones?

Can mating success and colony fitness be increased by boosting drone numbers?
And is this achievable at a scale meaningful to a small-scale beekeeper?

If a measurable increase in mating success took a 1000-fold increase in drone numbers it’s probably not achievable.

However, if all it took was an extra frame of drone comb in every hive in the apiary, then that’s quick win.


 

Tragedies and triumphs

Synopsis: Beekeeping shouldn’t be “a series of calamities then winter”, though it sometimes feels like that. In the first of a two-part post I look at the real and imagined disasters that can befall you during the season. The reality is that the observant and well-prepared beekeeper can avoid most of the ‘tragedies’, and recover from almost all of them.

Introduction

A few weeks ago I did a live-streamed Q&A with Laurence Edwards from Black Mountain Honey. Some of the questions were both good and interesting, some of the answers were perhaps less so. Before any readers think I’m being rude here I should point out that Laurence was asking the questions – often on behalf of others – and I was answering them.

There were quite a few questions on non-chemical treatment which I was singularly ill-equipped to deal with. Not because I don’t know anything about it, but because I don’t practice it 1 and because I suspect I’m not a good enough beekeeper to be successful if I did try it. There’s clearly a lot of interest in the topic, though I fear much of this is also from beekeepers who are not sufficiently experienced to succeed with it either.

However, there were two questions – or perhaps it was one merged question – that went something like this:

What is your greatest beekeeping success and your biggest beekeeping disaster?

I’m paraphrasing here. I can’t remember the precise wording and daren’t review it on YouTube as I’d then have to listen to my erudite insights inchoate waffle … which would be excruciating.

My answer probably involved asking whether I was restricted to just one disaster … 😉

Let’s get some perspective first

New beekeepers in particular are likely to worry about the “disasters” and overlook some of the “successes” in their first season or two. I therefore thought I’d discuss what I consider are the highs and lows – abbreviated to tragedies and triumphs’ to give the post a snappy title – of the first few years of beekeeping.

Obviously this is biased and based upon my own experience, and from mentoring others. Your experience may be very different … or you may have yet to experience the highs and lows of a beekeeping season.

But before I start using superlatives to describe the chaos of my early efforts at swarm control it’s worth remembering – particularly as the war in Ukraine enters its third week – that I’m only talking about beekeeping here.

In the overall scheme of things it’s simply not very important.

What might feel like a disaster of biblical proportions in the apiary … isn’t.

Yes, it might threaten the productivity, or even the survival, of the colony, but it is only beekeeping 2.

So, having got that out of the way, which do you want first?

The good news or the bad news?

The bad news … how mature 😉

The loss of a hive tool

Clearly I’m being flippant here.

The loss of a hive tool is a minor inconvenience rather than a tragedy.

There you are!

Unless you don’t have a spare and/or you’re about to inspect a dozen heavily-supered hives in the apiary … in which case it’s a major inconvenience.

It’s remarkably easy for a hive tool to fall out of those tall, thin pockets in the sleeve or thigh of your beesuit. Inevitably it falls, not onto closely cropped sward, but into tangled tussocks of rarely-mown grass.

You will probably find it again.

You could spend 15 minutes on your hands and knees retracing your steps since you left the car or you could become a detectorist and conduct a grid-based search, sweeping the area for metal objects.

Neither method is guaranteed to work.

To be certain, you must cut the grass.

But be careful. A glancing contact with the lawnmower or brush cutter and a half-buried hive tool will be damaging at best, and potentially a lot worse 🙁

Hive tools soaking

Hive tools soaking in a solution of soda crystal

Or you can avoid all this grief by keeping a covered bucket in the apiary – a honey bucket is ideal – containing a strong solution of soda crystals. You know exactly where the hive tools are and you soon get into the habit of dropping it back in after an inspection.

Better still, keep two hive tools in the bucket and alternate them as you look at your colonies. The soaking in soda will clean the hive tool, reduce any potential cross-contamination and improve your apiary hygiene.

The loss of a queen

This can be anything from a minor inconvenience to a bit of a calamity.

It very much depends upon the:

  • time of the season
  • whether you notice she’s missing
  • availability of a spare colony

How do you lose a queen? Other than by losing a swarm (see below) the two most likely reasons are cackhanded beekeeping or a queen that fails due to being poorly mated.

Returning a marked and clipped queen to a nuc

Losing a queen mid-season, for whatever reason, should be little more than a minor inconvenience. Assuming you notice she’s missing in action you can remove unwanted queen cells, leaving a single charged (i.e. known to contain a fat larva lounging around on a comfortable bed of Royal Jelly) cell, and wait while she pupates, emerges, mates and starts laying.

Nerve racking? Perhaps slightly, but it’s usually a pretty safe bet that things will work out OK.

If, through clumsiness or stupidity 3, you kill the queen during an inspection there should be ample eggs and young larvae for the colony to use when rearing one or more replacements.

Keep your eyes peeled …

But what if you don’t notice she’s missing? You assume she’s there and blithely knock back all the queen cells you can find 4.

Sealed queen cells

You return the next week … all looks good, no more queen cells.

But wait a minute … there are no eggs either 🙁

Under these circumstances you realise the importance of having at least two colonies. You can rescue the queenless colony by donating a frame of eggs from a queenright colony.

With two hives a crisis is rarely a disaster

Queens also fail because they are poorly mated. They either stop laying, or they stop laying fertilised eggs (i.e.they continue to lay unfertilised ones, leaving you with ever-increasing numbers of drones in the colony). The colony might realise and supersede her, or you might be able to rescue the situation with a donated frame of eggs.

I’ll deal with the consequences of a failed or slaughtered queen at the extremities of the season – early or late – below.

The loss of a swarm

It happens to the best of us, and it sometimes seems to happen even if you do your swarm prevention and control by the book 5.

I’ve turned up in the apiary on a warm May afternoon to discover a whirling mass of bees swarming from one of my hives 6.

It’s not a disaster … in fact it’s one of the greatest sights in beekeeping.

With luck the swarm will bivouac nearby and you’ll be able to collect them in a skep and re-hive them late in the afternoon.

A small swarm

A small swarm …

At least it shouldn’t be a disaster, but Sod’s Law usually dictates that …

  1. if you’re there when the swarm emerges, and
  2. you have a skep and sheet with you

… the swarm will alight 45 feet up a Leylandii 🙁

Even then it might end well if you’ve got a suitable bait hive set out nearby.

The time when losing a swarm is a disaster 7 is when you don’t realise you’ve lost a swarm. You find some queen cells, hurriedly knock them all back 8 and then wonder why there are no eggs the following week.

Déjà vu

At which point you’re in a similar situation to the ‘loss of the queen’ I described above … except you’ve also lost up to 75% of the workers from the colony. The situation is still rescuable with a frame of eggs from your other hive 9 but you’re likely to miss out on the major nectar flow.

Could the situation be any worse?

Oh no it can’t … Oh yes it can!

You miss the lost the swarm, you knock back all those queen cells and you then fail to realise there are no eggs or young larvae in the colony until only sealed brood remains (i.e at least 9 days).

Or worse still, until no brood remains (i.e at most 21 days).

With no brood pheromone being produced there’s now a real danger that the colony will develop laying workers. Things now get an order of magnitude more difficult as a colony with laying workers is very difficult to requeen (and generally will not even attempt to rear their own if presented with a frame of eggs).

Drone laying workers ...

Multiple eggs per cell = laying workers (usually)

You’re fast approaching the next of the beekeeping ‘disasters’ …

The loss of a colony

How do you lose a colony?

What was it Elizabeth Barrett Browning said? ’Let me count the ways’ 10.

Natural disasters such as falling trees, winter gales, raging floods, woodpeckers, honey badgers and stampeding elephants 11 can destroy a colony.

Hive toppled by a summer storm

However much care you take – avoiding floodplains, strapping the hives down, seeking shelter (but not near shallow-rooted trees) – sometimes sh1t just happens 12.

You did your best and nature did her worst.

See what you can rescue and try again next year.

Queen loss at the start or end of the season

Losing a queen very early or very late in the season – for whatever reason – is a problem. There’s no chance of the colony rearing another – it’s too cold and/or there are no drones available. I suppose there’s an outside chance you could requeen the colony – if you had a queen available 13 – but doing so involves quite a bit of risk.

If the queen fails overwinter, all the bees in the box will be very old by the time your colony inspection confirms she’s firing blanks, or not firing at all. The chances of successfully requeening the hive are slim at best.

Although that colony is effectively lost – at least if it happens late in the season – it’s not an unmitigated disaster if you have another hive 14. You can unite the queenless colony over a queenright colony very late into the autumn, strengthening the latter and (at least) using the bees from the former, rather than condemning them to a lingering death.

An Abelo/Swienty hybrid hive ...

An Abelo/Swienty hybrid hive … uniting colonies in midsummer

I wouldn’t bother trying to unite a queenless colony (or one with a failed queen) at the very beginning of the season. The remaining bees will be pretty decrepit and there won’t be many of them. It’s unlikely they would contribute in a meaningful way to the successful build-up of another colony.

Winter losses through starvation

These are unfortunately common and often entirely avoidable.

Small-scale surveys from the BBKA and SBA often report winter colony losses of 20-30%, and up to 50% in some years. Large scale surveys, like the Bee Informed Partnership (BIP) one in the USA, have reported annual colony losses – the majority of which occur in the winter – exceeding 40% in all but two years since 2013.

Bee Informed Partnership loss and management survey

I’ve lost colonies through both starvation and disease.

In both cases it was entirely my fault 🙁

It was a disaster for the bees and it was a sobering and educational experience for me.

I discussed starvation, and how to avoid it, in winter weight a couple of weeks ago. I won’t rehash it here, but I will repeat again that the bees are still in the ‘danger zone’.

Time for another?

Time for another? Definitely.

There’s little nectar available and they are busy rearing brood. Their need for stores is probably higher now than at any time over the last 4-5 months.

At best, a shortage will hold the colony back. At worst they’ll die of starvation.

All of which is completely avoidable by ensuring they have ample stores at the beginning of the winter, and then by keeping an eye on the weight of the colony as they enter the spring. If you’re adding fondant in late December it’s likely the colony had insufficient stores to start with … but at least you’re keeping a check on the weight of the colony.

Winter losses due to disease

I suspect that the majority of winter losses are not due to starvation but are instead due to inadequate or incorrect Varroa management.

This is a topic that has been covered numerous times in posts here. The most recent overarching review of the topic is probably Rational Varroa Control. Versions of this appeared in the August 2020 BBKA Newsletter and in The Scottish Beekeeper in the same month.

Successful Varroa control requires an understanding of the treatments available and the pros and cons of using them on your bees and in your location/climate. Too many beekeepers simply want to know whether they should add Apiguard in the third week of August or middle of September.

Apivar strip on wire hangar

Unfortunately, it’s not quite that simple.

But that doesn’t mean it’s particularly difficult either.

Unlike many of the other diseases of honey bees – e.g. chronic bee paralysis virus (CBPV), Nosema and the foulbroods – there are effective treatments to control Varroa and the damaging viruses that it transmits.

Losing a colony in June to CBPV is possibly unavoidable (it’s just bad luck) but losing one to Varroa/DWV in January – which is largely avoidable – might well be bad beekeeping.

In both cases of course it’s a disaster for the colony 🙁

Disaster

The meaning of disaster is ‘An event or occurrence of a ruinous or very distressing nature; a calamity; esp. a sudden accident or natural catastrophe that causes great damage or loss of life’. Its origins date back to the mid-16th Century.

Some of the ‘disasters’ I’ve described above involve the loss of just one life – that of the queen. For the reasons I describe, they’re not really disasters at all, or shouldn’t be for the observant and well prepared beekeeper.

Locally bred queen ...

Locally bred queen …

They become disasters i.e. causing great damage or loss of life, if you miss the tell-tale signs and so contribute to the eventual demise of the colony.

The avoidable loss of a queen or a colony is a distressing experience, or at least it should be 15.

If it is distressing then it will probably also be a learning experience.

Analyse what went wrong and work out how you might prevent it happening again in the future.

We have a duty of care for the bees we manage. I don’t like losing colonies, but it still happens infrequently. When it does I try and determine whether it was just fate … or my incompetence (or – let’s be generous – my actions or inactions) that caused the loss.

And the times you manage to work out where you went wrong are the foundations for your beekeeping triumphs in the future … which is what we’ll return to next week.


 

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’ 🙂

Little dramas

This post was originally titled Drama queens.

Apposite … it’s mostly about queens.

However, the term drama queen refers to someone who overreacts to a minor setback 1 … which is almost the complete opposite of what I’m intending to discuss.

Instead, this post is about the – sometimes unseen – little dramas in the apiary. Things that go wrong, or could go wrong but eventually go OK because you gently intervene … or often because you don’t intervene at all 😉

It’s also about observing rather than doing. It’s sometimes surprising what you see, and – with a little application – you can learn something about your bees 2.

Of course, in the end some things do not end well … but there’s no point in being a drama queen about it 😉

Swarmtastic

There’s a certain predictability to the beekeeping year. It’s dictated by the climate and latitude, by the forage available, by the need for bees to reproduce (swarm) and by our efforts as beekeepers to corral them and keep them producing honey 3.

All of which means that June has been pretty manic. 

After a record-breakingly cold spring things finally warmed up. Here in Scotland this was 2-3 weeks into May.

Since then it’s been a near-constant round of queen rearing, swarm control, making up nucs and adding supers. Most of the OSR supers are now off, meaning that I’ll be hunched over the extractor for hours when I’m not with the bees 🙁

All the OSR near my bees is well and truly over – this lot is sadly just out of range

The rapid warming in late spring triggered a lot of swarming activity. I found my first charged queen cell on the 18th of May and, in at least one or two colonies, at every subsequent inspection since then.

Visits to the apiaries have been hard work. Inspecting a double brood colony with four full supers involves a lot of lifting 4.

And the lifting is necessary because I need to check whether there are any queen cells in the brood chamber.

I know some beekeepers simply prise the two brood boxes apart and expect to see queen cells at the junction.

That certainly works … sometimes.

However, I’ve found several colonies with queen cells in the middle of frames, or otherwise in positions I would not see them if I just looked at the interface between the boxes. 

Queen cell … and what else?

And I would still have to remove the supers to prise the brood boxes apart.

Although I’ve invested in some better quality hive tools, I’d need a crowbar to separate the boxes if there was 80 kg of supers on top 5.

So, if I have to take the supers off, I might as well look through the box carefully.

More haste, less speed

But before I fire up the smoker and start rushing around prising off crownboards I always try and simply observe what’s happening in the apiary.

Are all the colonies equally busy? If it’s the time of day when the new foragers are going on orientation flights are any colonies much less active? Have they had a brood break?

Which direction are the bees flying off or returning from? Has the main forage changed?

Are there any drones on orientation flights yet?

What’s happening at the hive entrances?

Is there pollen going in?

Any sign of fighting?

Or robbing?

It’s surprising what a few minutes observation can tell you about the local forage, the state of the colonies and their relative strength.

If you’ve not already read it (and even if you have) it’s worth finding a copy of At the Hive Entrance by Prof. H. Storch 6. The book’s strap-line is “How to know what happens inside the hive by observation on the outside”. Recommended.

And, now and again, you notice something unusual …

Queen under the open mesh floor

Like – in my peripheral vision – a single bee flying out from underneath an open mesh floor.

My queens are generally clipped. If the colony swarms the queen often finds her way back to the hive stand after crashing – very unregally 7 – to the ground. She crawls up the leg of the stand and ends up underneath the open mesh floor (OMF).

The bees then join her. It’s not unusual to find a large cluster of bees under the hive floor, with lots of activity, and lots of bees flying to and fro from underneath the OMF 8.

But last Friday, by chance I noticed a single bee and this prompted me to investigate.

A quick peek confirmed that there wasn’t a swarm under the OMF.

But there was a queen.

I spy with my little eye … you can just see the marked and clipped queen under this Abelo floor.

Almost completely alone.

I presume the colony had swarmed, the queen had got as far as she could and the swarm had eventually abandoned her and returned to the hive. 

When I inspected the colony I found a single sealed queen cell and confirmed that the queen I found was the one that was missing.

This colony was one of my ‘middle third’ ones 9i.e. destined for requeening with better stock if I had any spares.

There’s a near-to-eclosion queen cell under there …

I did.

I had half a dozen ‘spare’ queen cells almost ready to emerge from grafting at the start of June. I removed the queen cell in the hive and carefully checked I’d not missed any others. I then added the grafted cell, seating it in a thumb-sized depression over some brood. She will have emerged the following day and might even be mated when I check early next week.

Had I not seen the bee emerge from under the floor I’d have never otherwise checked. There are always a few bees under an OMF, but it’s rare to find a queen all alone there.

Queen in the grass

In another apiary the previous week I’d found a satsuma-sized cluster of bees in long grass about 10 metres from the hives. The application of a little gentle smoke and some prodding around with my index finger resulted in a clipped and marked queen calmly walking up onto my hand.

Microswarm? … or more likely the remains of a much larger one …

Again, I wouldn’t have seen this had I not been taking my time checking the hive entrances and the activity in the apiary. I was being even more leisurely than normal as there was rain threatening and I was trying to decide whether to start the inspections or not

Because of the known state of other colonies in the apiary – most were nucs with virgin or recently-mated queens – it was obvious which colony the queen had come from. 

The ‘threatening rain’ looked like it would soon become a certainty. I ran the queen in through the front entrance of the hive and the remaining bees eventually returned to the hive, fanning madly at the entrance.

Bees fanning at the entrance

When I next checked the hive the queen had gone 🙁

There was no sign the colony had swarmed, but there was a recently opened queen cell in there. I assumed there’s a newly emerged virgin queen running about in there with ‘blood on her hands’ having done away with the original queen.

We’ll find out next week.

Again, a few minutes just watching things in the apiary meant I found the queen. Had I not done so I’d have only seen the end result – a queenless colony – not the events that led to it.

Preventative and reactive swarm control

I should emphasise that the majority of my colonies are a little more under control than the two described above, both of which clearly attempted to swarm.

In both cases the clipped queen saved the day, even though she may not have lived to fight another day.

My swarm control (and success thereof) this season has been in stark contrast to last year’s ‘lockdown beekeeping’.

Then the priority was minimising travel and guaranteeing I wasn’t haemorrhaging swarms that might cause problems for the the public or other beekeepers.

I therefore used the nucleus method of swarm control on all my colonies. I implemented it well in advance of the peak swarming period. By doing so, I undoubtedly weakened my colonies. I produced less honey and did no queen rearing.

But I didn’t lose a single swarm 🙂

This year the priority has been to maintain strong colonies. Some are being used for honey production 10 and others are being split to make up nucs.

Inevitably a few have got a little ‘overcooked’ … but the clipped queen has usually ensured the bees remain in the hive.

I don’t think I’ve lost a swarm, but I have lost a few queens.

Queen in the cage

One of my colonies went queenless in mid May. This was well before I’d got any spare queens – mated or otherwise. I’d hoped that they would rear another, but it was too cold for the new queen to mate and the colony started to look a little pathetic.

I considered uniting them but, for a variety of reasons, never got round to it.

When I finally had a spare mated queen (in early June) I popped her into a JzBz introduction cage. I’d already plugged the tube with candy and placed a plastic cap over the end. 

The bees could feed the queen through the cage, but could not release her.

This is my usual method for queen introduction. I check the cage a day or so after hanging it between the frames. If the bees are showing aggression to the queen I leave it and check again 24 hours later.

Once they’re no longer showing any aggression I remove the plastic cap. The bees chew through the candy and release the queen.

Job done 🙂

I then leave the colony at least a week before inspecting, by which time I expect to see eggs and larvae.

JzBz queen introduction & shipping cage with removable plastic cap

On returning a week after removing the plastic cap I was dismayed to find the queen still in the cage. Most of the candy had gone, but there was a plug at one end that was rock hard. Clearly the bees had been unable to release her.

The colony had now been broodless almost a month. Brood pheromone is really important in suppressing laying worker activity in the hive. Queen pheromone is no substitute for brood pheromone 11 and I was very concerned about the additional lost week due to my stupidity 12.

But there was no point in being a drama queen … I opened the cage and gently released the queen onto a seam of bees. Five days later there are eggs and larvae (and the queen) in the hive, though I also think there are a few laying workers as there’s a smattering of drone pupae in worker cells (a classic sign).

Fingers crossed 🙂

Queen failure

The final ‘little drama’ was played out in full view over almost two months. Its eventual unsatisfactory conclusion was largely due to my procrastination … though I suspect a swallow or house martin may have hastened events at the end.

In late April, during one of the rare warm days it was possible to actually open a colony, I noticed some strange egg laying behaviour in one hive. 

The colony was queenright. The queen was marked and clipped and laying. However, although she was laying single eggs in worker comb, she was laying multiple eggs in about 10% of cells, almost all of which were in drone comb.

A fortnight or so later she was still doing the same thing. Even if it wasn’t obvious to me, it was clearly obvious to the bees that the queen was failing as they started a couple of queen cells. Here’s an enlargement of an earlier photo in this post – blue arrows mark single eggs, red arrows indicate multiples.

SIgns of a failing queen

I removed the queen and added a near-mature queen cell from my first round of grafting. She had emerged when I next checked, but was not yet laying (and I didn’t bother looking for her).

But, unlike the queen stuck in the cage, this didn’t have a happy ending.

By early June there was no sign of the queen and I fear she failed to return from a mating flight. There’s a big pond bear the apiary and it’s a magnet for swallows and house martins 13.

I added a frame of open brood (including both young larvae and eggs) in the hive, but they ignored it 14.

Frames showing the characteristic dispersed bullet brood of laying workers

When I next checked it was clear there were laying workers and I cut my losses and shook the colony out. 

In retrospect what should I have done? 

I should have united the colony in mid-May.

It was obvious then – at least to the bees – that the queen was failing. I’d never seen a queen laying singles in worker comb 15 but multiples in drone cells. 

Uniting would have immediately provided both brood pheromone and a laying queen. This would have suppressed the development of laying workers.

My notes go something like:

  • 18/5 – Still laying singles in worker and multiples in drone. Weird. QC looks like supersedure. Give them a week.
  • 26/5 – Q out. Didn’t check further. Decision time next week.
  • 3/6 – Strange. Increasing drone brood. Behaving queenright. Decision time next week.
  • 12/6 – Laying workers. Shook them out. Will I ever learn? EEJIT 16

The second rule of beekeeping

Anytime I write Decision time next week (or variants thereof, like Give them another weekin two successive weeks then it’s almost always going to end in tears 🙁

If it happens three times in succession it’s a nailed on certainty.

The first rule is – of course – Knocking off queen cells is not swarm control 😉


 

Acting on Impulse

Men just can’t help acting on Impulse … 

This was the advertising strapline that accompanied the 1982 introduction of a new ‘body mist’ perfume by Fabergé. It was accompanied by a rather cheesy 1 set of TV commercials with surprised looking (presumably fragrant) women being accosted by strange men proffering bouquets of flowers 2.

Men just can’t help acting on Impulse …

And, it turns out that women – or, more specifically, female worker honey bees – also act on impulse

In this case, these are the ‘impulses’ that result in the production of queen cells in the colony.

Understanding these impulses, and how they can be exploited for queen rearing or colony expansion (or, conversely, colony control), is a very important component of beekeeping.

The definition of the word impulse is an ‘incitement or stimulus to action’.

The action, as far as our bees are concerned, is the development of queen cells in the colony.

If we understand what factors stimulate the production of queen cells we can either mitigate those factors – so reducing the impulse and delaying queen cell production (and if you’re thinking ‘swarm prevention‘ here you’re on the right lines) – or exploit them to induce the production of queen cells for requeening or making increase.

But first, what are the impulses?

There are three impulses that result in the production of queen cells – supersedure, swarm and emergency.

Under natural conditions i.e. without pesky meddling by beekeepers, colonies usually produce queen cells under the supersedure or swarm impulse.

The three impulses are:

  1. supersedure – in which the colony rears a new queen to eventually replace the current queen in situ
  2. swarm – during colony reproduction (swarming) a number of queen cells are produced. In due course the current queen leaves heading a prime swarm. Eventually a newly emerged virgin queen remains to get mated and head the original colony. In between these events a number of swarms may also leave headed by virgin queens (so-called afterswarms or casts).
  3. emergency – if the queen is lost or damaged and the colony rendered queenless, the colony rears new queens under the emergency impulse.

Many beekeepers, and several books, state that you can determine the type of impulse that induced queen cell production by the number, appearance and location of the queen cells.

And, if you can do this, you’ll know what to do with the colony simply by judging the queen cells.

If only it were that simple

Wouldn’t it be easy?

One or two queen cells in the middle of frame in the centre of the brood nest? Definitely supersedure. Leave the colony alone and the old queen will be gently replaced over the next few weeks. Brood production will continue uninterrupted and the colony will stay together and remain productive.

A dozen or more sealed queen cells along the bottom edge of a frame? The colony is definitely  in swarm mode and – since the cells are already capped – has actually already swarmed. Time to thin out the cells and leave just one to ensure no casts are also lost.

But it isn’t that simple 🙁

Bees haven’t read the textbooks so don’t necessarily behave as expected.

I’ve found single open queen cells in the middle of a central frame, assumed it was supersedure, left the colony alone and lost a swarm from the hive a few days later 🙁

D’oh!

Or I’ve found loads of capped queen cells on the edges of multiple frames in a hive, assumed that I’d missed a swarm … only to subsequently find the original marked queen calmly laying eggs as I split the brood box up to make several nucleus colonies  🙂

Not all queen cells are ‘born’ equal

It’s worth considering what queen cells are … and what they are not. And how queen cells are started.

There are essentially two ways in which queen cells are started.

They are either built from the outset as vertically oriented cells into which the queen lays an egg, or they start their life as horizontally oriented 3 worker cells which, should the need arise, are re-engineered to face vertically.

Play cup or queen cell?

Play cup or are they planning their escape …?

Queen cells started under the supersedure or swarming impulse are initially created as ‘play cups‘. A play cup looks like a small wax version of an acorn cup – the woody cup-like structure that holds the acorn nut. In the picture above the play cup is located on the lower edge of a brood frame, but they are also often found ‘centre stage‘ in the middle of the frame.

Play cups

A colony will often produce many play cups and their presence is nothing to be concerned about. In fact, I think it’s often a rather encouraging sign that the colony is sufficiently strong and healthy that it might be thinking of raising a new queen. 

Before we leave play cups and consider how emergency queen cells start life it’s worth emphasising the differences between play cups and queen cells.

Play cups are not the same as queen cells

Until a play cup is occupied by an egg it is not a queen cell.

At least it’s not as far as I’m concerned 😉

And, even if it contains an egg there’s no guarantee it will be supported by the workers to develop into a new queen 4.

However, once the cell contains a larva and it is being fed by the nurse bees – evidenced by the larva sitting in an increasingly thick bed of royal jelly – then it is indisputably a queen cell.

Charged queen cell ...

Charged queen cell …

And to emphasise the fundamental importance in terms of colony management I usually refer to this type of queen cell as a ‘charged queen cell’.

Once charged queen cells appear in the colony, all other things being equal, they will be maintained by the workers, capped and – on the 16th day after the egg was laid – will emerge as a new queen.

And it is once charged queen cells are found in the colony that swarm control should be considered 5.

But let’s complete our description of the queen cells by considering those that are produced in response to the emergency impulse.

Emergency queen cells

Queen cells produced under the emergency impulse differ from those made under the swarm or supersedure impulse. These are the cells that are produced when the colony is – for whatever reason – suddenly made queenless. 

Without hamfisted beekeeping it’s difficult to imagine or contrive a scenario under which this would occur naturally 6, but let’s not worry about that for the moment 7

The point is that, should a colony become queenless, the workers in the colony can select one or more young larvae already present in worker cells and rear them as new queens.

So, although the eggs are (obviously!) laid by the queen 8, they have been laid in a normal worker cell. To ensure that they get lavished with attention by the nurse bees, feeding them a diet enriched in Royal Jelly, the cell must be re-engineered to project vertically downwards.

Location, location

Queen cells can occur anywhere in the hive to which the queen has access.

Queen cell on excluder

Queen cell on underside of the excluder …

But they are most usually found on the periphery of the frame, either along the lower edge …

Queen cells ...

Queen cells …

… or a vertical side edge of the frame …

Sealed queen cells

… but they can also be found slap, bang in the middle of a brood frame.

Single queen cell in the centre of a frame

And remember that bees have a remarkable ability to hide queen cells in inaccessible nooks and crannies on the frame … and that finding any queen cells is much more difficult when the frame is covered with a wriggling mass of worker bees.

Location and impulses

Does the location tell us anything about the impulse under which the bees generated the queen cell?

Probably not, or at least not reliably enough that additional checks aren’t also needed 🙁

Many descriptions will state that a small number (typically 1-3) of queen cells occupying the centre of a frame are probably supersedure cells. 

Whilst this is undoubtedly sometimes or even often true, it is not invariably the case.

The workers choose which larvae to rear as queens under the emergency impulse. If the only larvae of a suitable age are situated mid-frame then those are the ones they will choose.

In addition, since generating emergency cells requires re-engineering worker cells, newer comb is likely more easily manipulated by the workers.

Some beekeepers ‘notch’ comb under suitably aged larvae to induce queen cell production at particular sites on the frame 9. The photograph shows a frame of eggs with a notch created with the hive tool. It’s better to place the notch underneath suitably aged larvae, not eggs. Clearly, the age of the larvae is more critical than the ease with which the comb can be reworked. Those who use this method [PDF] properly/extensively claim up to a 70% ‘success’ rate in inducing queen cell placement on the frame. This can be very useful if the plan is to cut the – well separated – queen cells out and use them in mating nucs or for requeening other colonies.

Eggs in new comb ...

Eggs in new comb …

Comb at the bottom or side edges of the frame often has space adjacent and underneath it. Therefore the bees might favour these over sites mid-frame (assuming ample suitable aged larvae) simply because the comb is easier to re-work in these locations.

And don’t forget … under the emergency impulse the colony preferentially chooses the rarest patrilines to rear as new queens 10.

Not all larvae are equal, at least when rearing queens under an emergency impulse.

Active queen rearing and the three impulses

By ‘active’ queen rearing I mean one of the hundreds of methods in which the beekeeper is actively involved in selecting the larvae from which a batch of new queens are reared.

This doesn’t necessarily mean grafting , towering cell builders and serried rows of Apidea mini nucs.

It could be as simple as taking a queen out of a good colony to create a small nuc and then letting the original colony generate a number of queen cells.

Almost all queen rearing methods use either the emergency or supersedure impulses to induce new queen cell production 11.

For example, let’s consider the situation described above.

Active queen rearing and the emergency impulse

A strong colony with desirable traits (calm, productive, prolific … choose any three 😉 ) is made queenless by removing the queen on a frame of emerging brood into a 5 frame nucleus hive. With a frame of stores and a little TLC 12 the queen will continue to lay and the nuc colony will expand.

Everynuc

Everynuc …

But the, now queenless, hive will – under the emergency impulse – generate a number of new queen cells. These will probably be distributed on several frames if the queen was laying well before she was removed.

The colony will select larvae less than ~36 hours old (i.e. less than 5 days since the egg was laid) for feeding up as new queens.

If the beekeeper returns to the hive 8-9 days later it can be split into several 5 frame nucs, each containing a suitable queen cell and sufficient emerging and adherent bees to maintain the newly created nucleus colony 13.

Active queen rearing and the supersedure impulse

In contrast, queenright queen rearing methods such as the Ben Harden system exploit the supersedure impulse.

Queen rearing using the Ben Harden system

In this method suitably aged larvae are offered to the colony above the queen excluder. With reduced levels of queen pheromones present – due to the physical distance and the fact that queen cannot leave a trail of her footprint pheromone across the combs above the QE – the larvae are consequently raised under the supersedure impulse.

Capped queen cells

Capped queen cells produced using the Ben Harden queenright queen rearing system

I’m always (pleasantly) surprised this works so well. Queen cells can be produced just a few inches away from a brood box containing a laying queen, with the workers able to move freely through the queen excluder. 

Combining impulses …

Finally, methods that use Cloake or Morris boards 14 use a combination of the emergency and supersedure impulses.

Cloake board ...

Cloake board …

In these methods the colony is rendered transiently queenless to start new queen cells. About 24 hours later the queenright status is restored so that cells are ‘finished’ under the supersedure response.

The odd one out, as it’s not really practical to use it for active queen rearing, is the swarming impulse. Presumably this is because the conditions used to induce swarming are inevitably rather difficult to control. Active queen rearing is all about control. You generally want to determine the source of the larvae used and the timing with which the queen cells become available.

Environmental conditions can also influence colonies on the brink of swarming … literally a case of rain stopping play.

Acting on impulse

If there are play cups in the colony then you don’t need to take any action 15, but if there are charged queen cells present then your bees are trying to tell you something.

Precisely what they’re trying to tell you depends upon the number and position of the queen cells, the state or appearance of those cells, and the state of the colony – whether queenright or not.

What you cannot do 16 is decide what action to take based solely on the number, appearance or position of the queen cells you find in the colony. 

Is the colony queenright?

Are there eggs present in the comb?

Does the colony appear depleted of bees?

If there are lots of sealed queen cells, no eggs, no sign of the queen and a depleted number of foragers then the colony has probably swarmed. 

Frankly, this is pretty obvious, though it’s surprising the number of beekeepers who cannot determine whether their colony has swarmed or not.

But other situations are less clear … 

If there are a small number of charged queen cells, eggs, a queen and a good number of bees in the hive then it might be supersedure.

Or the colony might swarm on the day the first cell is sealed 🙁

How do you distinguish between these two situations? 

Is it mid-May or mid-September? Swarming is more likely earlier in the season, whilst supersedure generally occurs later in the season.

But not always 😉

Is the queen ‘slimmed down’ and laying at a reduced rate?

Much trickier to determine … but if she is then they are likely to swarm.

Decisions, decisions 😉 … and going by the number of visits to my previous post entitled Queen cells … don’t panic! there are lots of beekeepers trying to make these decisions right now 🙂