Larger queens, higher queens - phase 0

That is the prequel to Larger queens, higher queens – phase 1 which was once all in regards to the maternal impact in honey bees.

The maternal impact, even though well known in different species, has most effective not too long ago been demonstrated in honey bees. Necessarily, it comes to the queen preferentially laying greater eggs in queen cells than she lays in employee cells. Those greater eggs transform greater and heavier queens.

In all probability unsurprisingly – despite the fact that additionally reassuringly – those greater queens have extra ovarioles and lay extra eggs than smaller queens. In the end, those greater queens seem to go on fascinating characteristics to their progeny; 2nd technology queens also are greater, as are the employees.

Phase 1 on this sequence of three (or in all probability now 4) posts described the demonstration of the maternal impact. On this put up I will discover – inevitably in brief because of the provision of one of the vital historic literature – why greater queens could be higher.

It’s a must to emphasise that none of the resources used for this put up have been acutely aware of the maternal impact in honey bees. Despite the fact that there could have been hints this type of factor befell in research relationship again to the 1970’s, the proof was once circumstantial or – at best possible – correlative, and it wasn’t demonstrated till 2019.

Table of Contents

Elements affecting the worth of a queen

This heading is lifted verbatim from Laidlaw and Eckert’s Rearing queens (1962). In spite of now being over 60 years outdated, nearly all of this ebook – a minimum of relating to the sensible beekeeping – stays related. Unsurprisingly, there is not any point out of Varroa which wasn’t recognized in the US till 1987 {{1}}.

The price of a queen relies, in large part, upon her skill to put a enough choice of eggs to care for a powerful colony suitable to the quite a lot of seasons of the 12 months (Laidlaw & Eckert, 1962).

The egg-laying skill of the queen is the number one measure of her worth. It must be evident why that is; a queen that lays too few eggs, or too many unfertilised eggs, or who does no longer get started laying early sufficient within the season, or who stops laying too quickly within the autumn, will lead to an undersized colony this is too small to thrive.

Sluggish spring enlargement signifies that the colony is not going to be large enough to swarm (reproduce).

As a beekeeper chances are you’ll suppose that is “a just right factor” … it’s not.

An undersized colony will acquire much less nectar, will likely be much less ready to protect itself in opposition to robbing, and would possibly not acquire enough retail outlets to live on the iciness. Iciness survival may be compromised by means of too few iciness bees … a lot of research have proven that enormous colonies (in overdue autumn, no longer essentially in midsummer) have higher overwinter survival.

High quality and amount

Laidlaw and Eckert certified their definition of the worth of a queen by means of additionally commenting on how she is answerable for the traits of the colony.

A queen that laid ‘enough choice of eggs to care for a powerful colony’ is of little worth if the colony is competitive, liable to illness, or that swarms once your again is became {{2}}.

Then again, those qualities are in large part inherited and/or made up our minds by means of environmental stipulations.

In case you permit an competitive colony to requeen itself, don’t be shocked if the colony stays competitive as soon as the employee inhabitants is changed with the brand new queen’s brood. On this case, the competitive trait is inherited.

In a similar way, if it is raining, or just about middle of the night (or, for impressive impact, each), and also you elevate the roof and crownboard, even a typically placid colony will react defensively. That is the affect of our environment. Precisely the similar issues applies in a nectar dearth, or if the colony is queenless.

Despite the fact that it would possibly be preferable to be confronted with a smaller choice of psychotic bees in the dark on a wet night time, the truth is that the qualitative characteristics – relating to colony survival and productiveness – are normally secondary to the manufacturing of enough eggs to care for a powerful colony.

Reproductive attainable

The reproductive attainable of the queen contains, however isn’t limited to, her egg laying. Reproductive attainable is outlined relating to:

fecundity – the quantity or price at which eggs are laid
fertility – (associated with above) however in all probability higher outlined because the quantity or share of fertilised eggs produced
longevity – the duration over which the queen is productive, outlined each relating to years, however (in my opinion) extra importantly, relating to the months of productiveness consistent with season {{3}}

No longer all queens are the similar dimension. My local darkish bees have reasonably small queens, while my bait hives every so often draw in swarms headed by means of monumental yellow queens. That is the herbal variation between other traces – and hybrids of the ones traces – of honey bees.

Extra related to reproductive attainable is the demonstration that queens of other sizes can also be constituted of a unmarried pressure of bees.

This has been exploited to review the reproductive attainable of honey bees. Those medical research in large part post-date the overall acceptance that ‘larger queens are higher queens’, however supply tangible proof supporting this view.

All that attainable

The destiny of a fertilised egg relies fully upon its surroundings. That is termed polyphenism – a mixture of poly- (a couple of) and phenotype (the observable traits of a person) – which means that:

The capability of a genotype to provide two or extra distinct phenotypes according to an environmental issue comparable to temperature, photoperiod, or diet.

In honey bees, it’s diet that determines the destiny of the egg.

If the egg is laid in a horizontal cellular – a employee cellular – it’s fed employee jelly and, after pupation, emerges as a feminine employee (the blue line within the cool animated film beneath).

Against this, if the cellular orientation is vertical, the nurse bees feed the growing larvae on a wealthy nutrition of royal jelly (the purple traces). This induces an in depth vary of DNA adjustments (methylation) which lead to adjustments within the expression of quite a lot of genes. After pupation, a queen emerges.

It’s (predominantly, however no longer solely) the further sugar within the royal jelly that results in queen building

Thankfully for clumsy beginner beekeepers and the ones folks who rear queens {{4}} (and for the honey bee colony) there’s a bit of suppleness all through the primary 0 to a few days of larval building that permits older larvae to even be reared into queens, beneath sure cases.

Queen (purple) and employee (blue) building – numbers are days after the egg is laid

If a tender larva is transferred from a horizontal employee cellular to a vertically-oriented queen cellular, it is going to be fed royal jelly and reared as a queen. In a similar way, if the queen is misplaced, the bees will re-engineer a employee cellular to be vertically orientated, feed the larva it incorporates royal jelly, and thereby produce the queen.

It is this ‘magic’ that permits us to rear queens and that permits the colony to in most cases recuperate from the – in a different way catastrophic – lack of a queen because of cackhanded beekeeping or illness.

After 3 days of larval building, this skill to reprogramme a employee right into a queen is misplaced.

Serve as no longer shape

Then again, there may be an instantaneous dating between the quantity of royal jelly fed to a growing larva, and the way queen-like the ensuing grownup bee is.

The longer a larva is fed royal jelly, and due to this fact the extra she consumes, the extra queen-like the grownup will likely be (indicated by means of the thickness of the purple traces).

Growing queen larvae are in a perpetual ‘all you’ll consume’ eating place, while growing staff ‘experience’ much less of a much less wealthy nutrition.

In fact, it is a bit extra difficult than that, however at a primary approximation that’ll do.

Seminal research carried out by means of Jerzy Woyke (1971) demonstrated that queens raised from employee larvae of accelerating age had decreased reproductive attainable; outlined relating to their weight, ovariole quantity, spermatheca diameter, and saved sperm counts.

Because of this it is very important use very younger larvae when queen rearing. It is usually why colonies preferentially make a choice 3 day outdated eggs when rearing queens beneath the emergency reaction.

Sadly, Woyke’s research are hidden in the back of the Magazine of Apiculture Analysis‘s paywall or even my ‘get right of entry to all spaces’ college account cannot get to a duplicate of the paper.

By no means thoughts … Woyke was once finding out the phenotype – the bodily traits – of the queen. So far as sensible beekeeping is anxious, it is what the queen does with that additional weight, the ones additional ovarioles and the higher saved sperm that issues.

And, it seems, there is not essentially a direct dating between the bodily attributes of a queen and her reproductive attainable. Some next research have produced contradictory effects; greater or extra ovarioles do not all the time equate to extra eggs, despite the fact that Nelson & Gary (1983) demonstrated an instantaneous dating between heavy queens, larger brood space and higher honey manufacturing.

Colony health and queen dimension

In most cases, those types of research are executed by means of rearing queens from younger (day 0) or outdated (day 2) larvae and evaluating the efficiency of the ensuing queens, or the colonies that they head.

What different traits of a colony might be measured as a method of figuring out whether or not greater queens are fascinating?

I have up to now mentioned polyandry and colony health, and the fascinating thought of hyperpolyandry and larval variety:

Those posts are value (re)studying … my writing could be as deficient as ever, however the underlying science is superb, and the consequences are attention-grabbing.

Queens move on a number of mating flights and mate with more than one drones. It is necessary that they do because it will increase the genetic range throughout the colony. This genetic range is significant to verify colony health; this can be a catch-all time period that describes how effectively the colony survives and reproduces.

A healthier colony will show off higher illness resistance, will overwinter effectively and can increase strongly to swarm (reproduce) the next season.

So, how do ‘top’ and ‘low’ high quality queens – reared from younger or outdated larvae respectively – evaluate when their mating good fortune is analysed?

It takes two to tango

One find out about, by means of Tarpy et al., (2011), aimed to research the mating behaviour and good fortune of low and high high quality queens, a very powerful useful high quality associated with colony health.

Tarpy measured the bodily traits of the queens (reared by means of grafting 0 day or 2 day outdated larvae, however in a different way handled in an an identical way).

It is value citing right here that the ‘low high quality’ queens have been so small that many may get throughout the queen excluder at the hive entrances, so combating correct recording of the whole choice of mating flights.

The low high quality queens have been considerably lighter (~87% that of top quality queens), had a narrower thorax (~94%) and a discounted spermatheca diameter (~92%) and therefore quantity. As a outcome, the ‘low high quality’ queens saved considerably much less sperm (~65%), despite the fact that the entire degree of sperm viability was once the similar.

Why was once there much less sperm? Most certainly since the queens had mated with fewer drones. Each the noticed mating quantity and the efficient paternity frequency – made up our minds by means of genetic research of ensuing brood to quantify the distinct patrilines within the colony – have been considerably decrease within the ‘low high quality’ queens (at ~82% and ~78% respectively).

Do those variations subject?

Probably.

Despite the fact that no longer measured on this find out about, colony health – as outlined above – is expounded to the genetic range of the colony. If the queen pals with fewer drones the variety is inevitably diminished, which could be unfavorable. Moreover, if the quantity of sperm saved is inadequate it is going to prohibit the reproductive longevity of the queen.

Penalties for the colony

In an extension of those research, Tarpy and co-workers (Rangel et al., 2013) measured sure traits of colonies headed by means of ‘low high quality’ or ‘top quality’ queens.

The queens have been reared the usage of usual ways after which used to requeen a bundle of bees (US readers will likely be accustomed to those … it is a field containing bees however no comb or brood). The ensuing applications have been hived on frames without or with basis, in order that the bees may draw employee or drone comb primarily based upon the wishes of the colony. Colonies have been monitored from Would possibly to September, and their next overwintering survival recorded. An extra 40 colonies (50/50 low or top quality queens) have been arrange the next season to copy the research of overwintering survival.

Colony efficiency (from Rangel *et al.,* 2013)

Colonies headed by means of ‘top quality’ queens (forged traces within the graphs) constructed ~34% extra employee comb (a) and ~144% extra drone comb (b) and saved extra pollen and nectar (c), all of that have been statistically important. Despite the fact that there have been numerical variations in each employee and drone brood manufacturing (e and f, respectively) those weren’t statistically important. In the end, colonies headed by means of ‘top quality’ queens have been estimated to include ~41% extra staff (f) than the ones headed by means of ‘low high quality’ queens.

Larger queens, larger colonies 😄.

Survival of the fittest?

Then again, the overwinter colony survival is, frankly, just a little of a multitude and is confounded by means of the way in which it’s introduced within the paper. Necessarily, within the first 12 months maximum of the colonies perished, and there was once no important distinction between colonies headed by means of ‘top’ or ‘low’ high quality queens.

No longer just right 😔.

Within the repeat of this a part of the find out about, 75% of the colonies headed by means of ‘top quality’ queens survived overwinter, against this to just 30% of the ‘low high quality’ queen colonies.

Complicated 😕. Such diametrically opposing effects can’t be meaningfully interpreted.

I feel this a part of the find out about must be repeated.

One fascinating statement was once the choice of occasions that queens have been outmoded.

My expectation would were that the colonies headed by means of ‘low high quality’ queens would were outmoded a lot more regularly. This could are compatible with earlier observations at the supersedure of poorly mated queens, or the ones failing because of illness. Then again, even though supersedure was once noticed, there was once no important distinction within the numbers of ‘top’ or ‘low’ high quality queens that have been changed.

Relevance to sensible beekeeping

All beekeepers rear queens. No longer all know they rear queens, however the truth that queens have a finite lifespan and the beekeeper proceed to stay bees signifies that queens will have to be being reared {{5}}.

In case you rear queens passively, through which I imply merely letting colonies requeen all through swarm regulate, or by means of splitting colonies, then it is value remembering that the bees will preferentially make a choice 3 day outdated eggs and that the ensuing queens wish to be well-fed all through their early building.

If there’s a dearth of nectar or a scarcity of pollen, feed the colony. Agree with the decisions the bees make relating to the larvae they rear as queens. They know what they’re doing.

Then again, if the colony has very little eggs or very younger larvae then supply some … they may be able to most effective paintings with what they’ve within the hive and if, for no matter explanation why, the one larvae are 2-3 (or extra) days outdated they’ll most effective have the ability to rear small, low high quality, queens.

Lively queen rearing

In case you rear queens actively i.e. by means of making ready a cellular elevating colony, including larvae after which transferring the ensuing queens (or cells) to mating nucs, then it is not value going to all that palaver except you’ll be sure that; a) the larvae are as younger as imaginable, and b) the cellular elevating colony is full of younger bees and has plentiful nectar and pollen.

Grafted larvae flanked by means of open brood, a whole lot of pollen and a body feeder

I take advantage of the Ben Harden queenright queen rearing means. This comes to setting up a four-frame ‘chimney’ above a hectic brood field. One of the most frames incorporates the larvae – in most cases grafted, despite the fact that they do not have to be – one is filled with open brood, to attract up nurse bees from the brood field, and the remainder two are pollen-filled. And subsequent to that could be a frame-feeder to which I upload syrup every day till the cells are sealed.

All types of different methods were used to take a look at and ensure just right larval diet. Laidlaw and Eckert (1962) and David Woodward (in his ebook Queen Bee: Biology, Rearing and Breeding (2010), Northern Bee Books) each describe ‘double grafting’. This technique is designed to verify the larvae are effectively fed from the instant they’re transferred to the cellular elevating colony, in all probability even earlier than they get the eye of the nurse bees.

Larvae are grafted into cellular cups and added to the cellular elevating colony for 2 days. Right through this era they’re fed copious quantities of royal jelly. After two days, those preliminary larvae are got rid of (however the royal jelly isn’t) and changed with some other very younger larva.

While I perceive the common sense, this is not one way I take advantage of. Laidlaw and Eckert suggest that larval feeding earlier than grafting is most probably extra necessary than within the speedy duration after grafting earlier than the nurse bees get busy.

Contemporary research on larval selection by means of staff means that the bees additionally believe well-fed larvae are the most efficient beginning subject matter for brand spanking new queens … the colony from which larvae are sourced must even be effectively fed if wanted.

Notes

Phase 2 on this poorly looked after sequence will practice subsequent month as soon as a couple of ultimate experiments were carried out. I feel it is virtually sure that I will be able to want a ultimate put up (phase 3?, I have misplaced depend) to speak about one of the vital gaps in our working out of the maternal impact.

Informative? Helpful? Entertaining? … make a choice any 3.
Please improve additional articles by means of turning into a supporter or investment the caffeine that fuels my overdue night time writing …

Thanks

References

Nelson, D.L., and Gary, N.E. (1983) Honey Productiveness of Honeybee Colonies in Relation to Frame Weight, Good looks and Fecundity of the Queen. Magazine of Apicultural Analysis 22: 209–213 https://doi.org/10.1080/00218839.1983.11100589.

Rangel, J., Keller, J.J., and Tarpy, D.R. (2013) The results of honey bee (Apis mellifera L.) queen reproductive attainable on colony enlargement. Insect Soc 60: 65–73 https://doi.org/10.1007/s00040-012-0267-1.

Tarpy, D.R., Keller, J.J., Caren, J.R., and Delaney, D.A. (2011) Experimentally triggered variation within the bodily reproductive attainable and mating good fortune in honey bee queens. Insect Soc 58: 569–574 https://doi.org/10.1007/s00040-011-0180-z.

Woyke, J. (1971) Correlations Between the Age at Which Honeybee Brood was once Grafted, Traits of the Resultant Queens, and Result of Insemination. Magazine of Apicultural Analysis https://www.tandfonline.com/doi/abs/10.1080/00218839.1971.11099669.

{{1}}: Despite the fact that it was once definitely there earlier than then because it was once present in a minimum of 12 states that autumn.

{{2}}: That isn’t a systematic definition, however person who applies all too regularly 😉 .

{{3}}: A 7 12 months outdated queen that most effective laid eggs in Would possibly and June is of little use to any person.

{{4}}: I belong to each teams.

{{5}}: OK, sure they may be purchasing them in from Greece each and every season, however they might be nuts to try this, would not they?

Larger queens, higher queens – phase 0