# Bees criteria to choose queen larvae



## Eduardo Gomes (Nov 10, 2014)

I can explain the reason for my intuition. I read somewhere that the flowers convey the slopes bees to them " say " if it has been crafted by another bee. These tracks , subliminal to us, are "read" by the bees . So their work is much more efficient because it does not waste time on flowers that have nothing to give them. I've seen many times bees hanging around flowers without their reach it landed .

Does the larvae can also send any message to bees which to assess a more complex and full of its potential to be future queens? You know any research that has already specifically assessed this issue?


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## beepro (Dec 31, 2012)

What do you mean by age criterion of the larva? You
mean choosing a larva that is older than 4 days to make
a queen from it? If they have a choice they will select
younger larva so that it can feed on more RJ to develop into
better queen. But when all else fail they have no choice but
to pick a larva that hopefully will create a good queen for them.
I think it all depends on the situation at hand and what is available
to them. Don't you think so?


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## jbraun (Nov 13, 2013)

If I have somethings to eat and they are not all equal but similar, I always choose the best one first. That way the next one I choose is the best one and so on. Why would a hive do anything different? We all want the best outcome for us.


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## sqkcrk (Dec 10, 2005)

Workers will often pick larvae that are not the best right age for making into queens. Or so I understand.


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## drlonzo (Apr 15, 2014)

I've witnessed exactly what Mark states. If you put together a NUC and allow them to make their own queen, at times they will take older larva than a 4 cycle day old and turn it into queens. Although, at the same time, i've witnessed the fact that they will also use the correct age 4 cycle day old larvas to make queens.

I've myself put together several NUC's with only freshly lain eggs for them to make queens with, and they fail miserably... Not quite sure why, but after them not making a queen with the eggs, i've placed into those nuc's a frame with larvas and gotten great queens.. 

So I think, IMO that at times it depends upon the bees as to if they do or don't pick the correct age.. 

HENCE - NICOT system..


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## Eduardo Gomes (Nov 10, 2014)

beepro said:


> What do you mean by age criterion of the larva?


To explain better I 'll use an example to our human scale : when I choose a second hand car I use several criteria to select : Car age; kilometers ; price; consumption; model .... I do not use only one criterion . However someone who was watching my selection process this car could get the wrong idea that I bought that car only to be what had fewer years . But I took that decision after considering various criteria.

It may be that the bees because it has very different sensory thresholds of us, use beyond the age criteria , other clues that allow them to assess more fully the quality of small larvae who choose to be queens when they are in a natural situation .

Are only thoughts ... but all begins this way.


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## centrarchid (Jun 13, 2014)

I would look for workers to select larval sisters that are most similar to them to have a go at being the next queen once age criteria are met. It would not be surprising to me if they can estimate relatedness based on smell. They would have a choice between full sisters and half sisters to varying degrees depending on how closely related the drones were to the queen.


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## Marysia2 (May 23, 2014)

centrarchid said:


> I would look for workers to select larval sisters that are most similar to them to have a go at being the next queen once age criteria are met. It would not be surprising to me if they can estimate relatedness based on smell. They would have a choice between full sisters and half sisters to varying degrees depending on how closely related the drones were to the queen.


Seems reasonable, based on my own human prejudices (yes, if I can't be queen, then having one of my sisters become queen might be the next best thing in terms of being beneficial to ME). But how would it benefit a bee to have a full sibling as queen? Does the queen show a preference somehow to her siblings? 

Usually animal behavior is predicated by what will be best for the individual or the group or the species. Interesting question...


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## sqkcrk (Dec 10, 2005)

Or what is available? 

When I set things up for bees to raise their own queen from larvae they quite often will draw three or four cells right next to each other. I wonder if they do that to make sure that one of the "right" age gets a chance to make a viable queen. There must be something about the larvae that makes the bees pick certain ones and not others, makes them grow only a relatively small number and not 20 at a time. Maybe that's an energy and resources constraint. I bet there are real theories known to queen experts.


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## Marysia2 (May 23, 2014)

sqkcrk said:


> Or what is available?...There must be something about the larvae that makes the bees pick certain ones and not others...


Why does the position of the queen cup on the comb indicate its purpose: for a future swarm, or an overthrow of the monarchy (supercedure)? 

And for that matter...how do they all decide and agree on overthrowing the queen? Are there whispers in the hallway? Is the old queen not aware of the chatter going on about giving her the boot? And what if the old queen somehow manages to kill her young rival? Are the conspirators executed? Do the bees try again to get rid of the old battle-axe? So many questions, so few answers...


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## sqkcrk (Dec 10, 2005)

It is my understanding that the lack of queen scent/pheromone produced by the resident queen starts the superscedure process. Queens being supersceded don't defend themselves or attack virgins, as far as I know. Some times one will find a new mated queen and an old queen in the same hive. Mother and daughter queens. Only the new queen is the one laying eggs, I think.

If one sees a queen cell on the face of a comb usually that cell was made from a cell that was originally going to be a worker bee but the colony determined a need for a queen and made modifications. Those queen cells are due to an emergency need or a failing queen. Therefore we call them superscedure cells or emergency cells. I don't know what the bees call them, if anything.

As far as queen cells occurring along the bottom edge of combs, queen cups are often found along the bottom edges of combs. They are always present, but only put into use when swarm season occurs. The queen lays eggs in these cups because she has to, every other cell already being filled.


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## Michael Bush (Aug 2, 2002)

>I would look for workers to select larval sisters that are most similar to them

I've read research that goes both directions on that theory. Perhaps they do...


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## crofter (May 5, 2011)

In Snelgroves queen rearing he suggests that after you isolate open brood to initiate queen cell starting, to go back on day 4 and destroy any cells that are already capped since they must have been made from older larvae. This is to prevent creation of caste queens. I wonder if the bees, left to their own might cull such cells later on in development. Maybe they can judge a queens potential by smell. There obviously must be some mechanism generally in effect to keep the older age larvae from hatching first and killing the better potential queens.


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## Eduardo Gomes (Nov 10, 2014)

crofter said:


> In Snelgroves queen rearing he suggests that after you isolate open brood to initiate queen cell starting, to go back on day 4 and destroy any cells that are already capped since they must have been made from older larvae. This is to prevent creation of caste queens.


I do it whenever I can , but further tightening the mesh preferring to make the selection after 5 days. 



> I wonder if the bees, left to their own might cull such cells later on in development.


Very pertinent question undoubtedly. I hope so but I have no data to confirm that yes or no . Who has observation hives , has seen something interesting about this?


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## Michael Bush (Aug 2, 2002)

>I wonder if the bees, left to their own might cull such cells later on in development.

That was the conclusion of Jay Smith and C.C. Miller and Snelgrove...


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## crofter (May 5, 2011)

Would the cell length tend to be greater if the bees had longer to play with the construction rather than being rushed with a rapidly developing larvae? It certainly seems to be popular that length is an indicator for better potential when the beekeeper is doing the choosing rather than the bees. Would the bees use size to judge or something more sophisticated like pheremones?


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## sqkcrk (Dec 10, 2005)

Isn't cell length influenced by how much the larvae was fed, how much royal jelly was put in the cell by the bees?


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## dsegrest (May 15, 2014)

centrarchid said:


> I would look for workers to select larval sisters that are most similar to them to have a go at being the next queen once age criteria are met. It would not be surprising to me if they can estimate relatedness based on smell. They would have a choice between full sisters and half sisters to varying degrees depending on how closely related the drones were to the queen.


Last month's ABJ had an article that proposed bees would select a queen from the same father as the bees making the choice.


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## sqkcrk (Dec 10, 2005)

I have heard of Nurse Bees and House Cleaner Bees, but never a Geneology Bee.  Just funnin' ya.


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## dsegrest (May 15, 2014)

They only know who's who when their mama calls them by their last name.


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## beepro (Dec 31, 2012)

Confusing! If mama call then everyone with the same last name would responded. 
YESss....mommy! 


No, the cell size or length will not indicate a good queen or not. The only way to get exceptional queens is to
have lots of royal jelly producing workers to feed the queen cell. But having a cup big enough to fit all those RJ will
help too. When you see some RJ left at the bottom of the cup after the queen had hatched then you know she
is well fed. So it might be a potential good queen after evaluation. After 3 seasons of Spring and late summer queen 
grafting I have some interesting observation of these cups.


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## JRG13 (May 11, 2012)

I tend to do what Crofter mentions as well, I'll come back and check for cells 4-5 days after a split and cull any capped ones sinc there are usually newer cells being made from eggs that have hatched after making the split. I know I've read posts on Beesource where people refer to research that concluded supersisters tend to select their own supersisters, but I don't know how bees put together a genetic sequencer to figure it all out. I have a strain of bees from a swarm that will not requeen with donated cells either.... they have to make their own.


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## GusK (Jan 24, 2013)

Royal allele hypothesis. Could often play a role in emergency queens. Some of my past cut down splits have produced incredible queens. However, if the first queen out is a dud, then all bets are off. 

I think it's better to graft for consistent results. Just let the bees have at the frame for a half day, then graft from the best fed larva, as they are most likely the chosen ones. Just check on their size, it must make sense. Work with the bees, get their input, and correct their mistakes.

BTW, I'm still new at this and still experimenting, FWIW. 


http://www.ncbi.nlm.nih.gov/pubmed/16151795


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## Eduardo Gomes (Nov 10, 2014)

GusK said:


> Royal allele hypothesis. http://www.ncbi.nlm.nih.gov/pubmed/16151795


"_*Here we show that honeybee queens are not reared at random but are preferentially reared from rare "royal" subfamilies, which have extremely low frequencies in the colony's worker force but a high frequency in the queens reared.*_" Rare royal families in honeybees, Apis mellifera. Moritz RF1, Lattorff HM, Neumann P, Kraus FB, Radloff SE, Hepburn HR.
Thank you Gusk. More material to study.:thumbsup:


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## JRG13 (May 11, 2012)

I'll read it, but how does that explain grafting success then?


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## Eduardo Gomes (Nov 10, 2014)

JRG13 said:


> I'll read it, but how does that explain grafting success then?


I have no doubt that it is a technique successfully ... for the professional queens breeders at least.
For many beekeepers is also a success because so must conveniently and timely have all the queens they need.

But some of us have some doubts. And there is no need to be afraid of doubts because they force us to think, to leave the usual frame, to create and improving solutions.

A french queens breeder raises some questions on this problem.
"Dans la technique du picking, c'est l'apiculteur qui sélectionne des larves. Mais que sait-il de l'état de la larve qu'il prélève ? Les abeilles elles, choisissent celles qui deviendront leurs reines. 
D'autre part, il faut savoir que dès son éclosion, les larves choisies par les abeilles pour devenir reines, bénéficient d'un régime alimentaire différent de celui des larves abeilles. Ces dernières sont nourries à la bouillie de gelée royale alors que les larves sélectionnées pour devenir reines sont nourries à la gelée royale pure. Par conséquent, en prélevant des jeunes larves (aussi jeunes soient-elles), l'apiculteur qui pratique la technique de picking sélectionne des larves qui n'ont pas été nourries correctement pour devenir reines. Certes, cela donnera de toutes manières des reines.. mais "inférieures" quand même. Faut-il dès lors s'étonner de l'essaimage précoce de certaines colonies ? Faut-il s'étonner si les reines vivent moins longtemps ? Faut-il s'étonner si après une saison et demi, elles tombent en panne de ponte plus rapidement, n'assurant plus la rentabilité attendue ? etc.." in http://www.abeille-et-nature.com/index.php?cat=essaims&page=elevage_de_reines

I understand very well the successful result for both parts ( sellers and buyers ) for the reasons I mentioned above.
As far as I am concerned I have always had doubts about picking and still have them after everything I've ever read . I have produced many hundreds of queens for myself , I have good queens, good productions for the portuguese standard , low mortality ... and never made a picking. I think this year I will need making some, but if I can I will avoid.

Gilles Fert a great European queens breeder recommends picking for beekeepers to do over 200 queens per year. It costs me a little in Portugal a beekeeper with 4 hives is led by the most experienced to learn to do picking for half a dozen queens. As if there were no other alternative and more adjusted to the size of the task !


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## Eduardo Gomes (Nov 10, 2014)

"Finally, *there is the important question of who decides whether an egg is a queen-to-be or a common worker, and how this decision is made.* Workers construct especially large “queen cups” out of wax throughout the season, but they often remain empty. New queens are reared when the old queen dies, or in preparation for colony fission [3]. Eggs are often laid by the queen directly into the cups, in which case she presumably has made a decision that this particular egg should be raised into a queen. *However, workers also move eggs from regular worker cells into these queen cups, and queen larvae and pupae are frequently destroyed by workers *[4]. Thus, *workers can substantially interfere with royal destiny*, and this raises the question of whether there is some nepotism involved. While caste differentiation into queens and workers is largely mediated by nutrition in honeybees, *there appears to be a genetic influence as well* [1]. Because of the promiscuous habits of the queen, a honeybee hive typically contains multiple subfamilies of workers each fathered by a different drone, and some subfamilies can be substantially overrepresented in queen production, presumably mediated by preferential treatment of certain larvae and selective abortion of others. *This adds a level of complexity to the interplay between genomic and environmental factors: the provisioning of epigenetic factors via larval nutrition might in turn be controlled by genetic factors that control provisioning behaviour.*" in http://www.plosbiology.org/article/info:doi/10.1371/journal.pbio.1000532


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## Eduardo Gomes (Nov 10, 2014)

And , in a way, to close the circle , we can see in this short video ( What Does Epigenetics Have to Do with Honeybees ?) As an existing component in royal jelly can make all the difference (http://www.nature.com/scitable/topicpage/epigenetics-fundamentals-26099600)

The world of bees is a world to another scale , is a world of subliminal signals to us, is a world full of clues to which we are blind . In this world pretend that we do exactly what the bees would at its best, can only say something about us.

The huge work of Mr. Doolittle was and continues to be a great help to the current beekeeping. Begins however to be the time to change the paradigm , IMO.


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## Michael Palmer (Dec 29, 2006)

Eduardo Gomes said:


> *However, workers also move eggs from regular worker cells into these queen cups,.... *


They do? That's debatable.


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## GusK (Jan 24, 2013)

I just want to make a couple things clear about the royal allele hypothesis. I first heard about it in a lecture given by Dave Tarpy - on the ABF website a while back. I then did more research. Overall, research shows that in an emergency situation, royal cells are less likely to be torn down, while older larva cells are more likely to be torn down. Nonetheless, some non-royal age inappropriate larva do make the final cut and become the first queen out thereby making the bees' royal selection moot.

Another question: Does that mean that bees raised randomly by grafting (larva age only criteria) are not as good as royal larva? I have no idea. It should take me a year or two to figure that one out. Luckily, I don't graft hundreds at a time, so I can take my time and focus by setting aside larva that appear to be chosen and random, yet age appropriate larva... then compare the two groups over the course of a year or so.

Randy Oliver recently alluded to royal larva in his recent queens for pennies slideshow. It's in a footnote on the pdf version if I recall correctly. He chooses the best fed larva after leaving the frame in the starter hive a few hours.


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## Eduardo Gomes (Nov 10, 2014)

Michael Palmer said:


> They do? That's debatable.


Hi Michael
First of all I want to tell you that I already follow some of the lecture you on Youtube and I'm a great admirer of your knowledge and ability to communicate.

Everything is questionable , no doubt. And I'm sure that you will have excellent reasons to question the statement in question .

Quite honestly if you are available to present yours arguments I will certainly learn with you. One thing you can also be right of me: you have all my attention .
Thank you!


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## Michael Bush (Aug 2, 2002)

>However, workers also move eggs from regular worker cells into these queen cups, and queen larvae and pupae are frequently destroyed by workers 

I have yet to see any evidence to support that theory. I seriously doubt they have the right glue to glue an egg into a cell. When the queen lays the egg it's wet and it sticks. When workers remove eggs I have never seen any evidence they do anything but eat them. Huber and Burnens spent thousands of hours and many experiments trying to set up scenarios where the bees desperately needed to move eggs and never got them to do it.


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## dsegrest (May 15, 2014)

Eduardo Gomes said:


> "_*Here we show that honeybee queens are not reared at random but are preferentially reared from rare "royal" subfamilies, which have extremely low frequencies in the colony's worker force but a high frequency in the queens reared.*_" Rare royal families in honeybees, Apis mellifera. Moritz RF1, Lattorff HM, Neumann P, Kraus FB, Radloff SE, Hepburn HR.
> Thank you Gusk. More material to study.:thumbsup:


That sounds reasonable for swarm cells and supercedure cells, but what if there is no egg/larvae from the royal family when an emergency queen is needed?


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## Eduardo Gomes (Nov 10, 2014)

dsegrest said:


> That sounds reasonable for swarm cells and supercedure cells, but what if there is no egg/larvae from the royal family when an emergency queen is needed?


From what I have observed also make a queen.

In this situation the balance between having a queen, albeit in a family " suspicion" and not have any queen , prefer first option. From an evolutionary point of view it makes sense so be it.

The Doolittle method , to the best reading, what it does is emergency queens : 1 ) the bees have a limited choice by choosing the beekeeper ; 2 ) create their new queen , at least in first moments of the process, in the absence of queen.


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## Eduardo Gomes (Nov 10, 2014)

Michael Bush said:


> Huber and Burnens spent thousands of hours and many experiments trying to set up scenarios where the bees desperately needed to move eggs and never got them to do it.


Michael Bush can you send me the reference of Huber and Burnens? ThanK you!

For the statement: "workers also move eggs from regular worker cells into these queen cups" the reference is Winston M. L (1987) The biology of the honeybee. Cambridge, Massachusetts: Harvard University Press.

You can read here at the bottom of the page what he states: http://books.google.pt/books?id=-5i...ar worker cells into these queen cups&f=false


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## Michael Bush (Aug 2, 2002)

>Michael Bush can you send me the reference of Huber and Burnens?

Keep in mind this is in August of 1791 which is 54 years before Dzierzon proposed parthenogenesis (in 1845) and 114 years before parthenogenesis was proven (in 1906). The prevailing theory at the time was that drone eggs and worker eggs were separated in some way in the body of the queen either in sequence in the ducts or some other mechanism.

http://www.bushfarms.com/huber.htm#commonbeesdonot

He mentions evidence of bees not moving eggs several times between volume I and volume II but here is the first and longer reference (in volume I):

"The worms proceeding from them pass their vermicular state in the same place where the eggs were deposited, which proves that bees are not charged with the care of transporting the eggs as has been supposed...Bees, I say, are not charged with the care of transporting into cells, the eggs misplaced by the queen: and, judging by the single instance I have related, you will think me well entitled to deny this feature of their industry. However, as several authors have maintained the reverse, and even demanded our admiration of them in conveying the eggs, I should explain clearly that they are deceived.

I had a glass hive constructed of two stages; the higher was filled with combs of large cells, and the lower with those of common ones. A kind of division, or diaphragm, separated these two stages from each other, having at each side an opening for the passage of the workers from one stage to the other, but too narrow for the queen.

"I put a considerable number of bees into this hive; and, in the upper part, confined a very fertile queen that had just finished her great laying of male eggs; therefore she had only those of workers to lay, and she was obliged to deposit them in the surrounding large cells from the want of others. My object in this arrangement will already be anticipated. My reasoning was simple. If the queen laid workers eggs in the large cells, and the bees were charged with transporting them if misplaced, they would infallibly take advantage of the liberty allowed to pass from either stage: they would seek the eggs deposited in the large cells, and carry them down to the lower stage containing the cells adapted for that species. If, on the contrary, they left the common eggs in the large cells, I should obtain certain proof that they had not the charge of transporting them.

"The result of this experiment excited my curiosity extremely. We observed the queen several days without intermission. During the first twenty-four hours, she persisted in not laying a single egg in the surrounding cells; she examined them one after another, but passed on without insinuating her belly into one. She was restless, and traversed the combs in all directions: her eggs appeared an oppressive burden, but she persisted in retaining them rather than they should be deposited in cells of unsuitable diameter. The bees, however, did not cease to pay her homage, and treat her as a mother. I was amused to observe, when she approached the edges of the division separating the two stages, that she gnawed at them to enlarge the passage: the workers approached her and also laboured with their teeth, and made every exertion to enlarge the entrance to her prison, but ineffectually. On the second day, the queen could no logger retain her eggs: they escaped in spite of her, and fell at random. Then we conceived that the bees would convey them into the small cells of the lower stage, and we sought them there with the utmost assiduity; but I can safely affirm there was not one. The eggs that the queen still laid the third day disappeared as the first. We again sought them in the small cells, but none were there. The fact is, they are ate by the workers; and this is what has deceived the Naturalists, who supposed them carried away. They have observed the misplaced eggs disappear, and, without farther investigation, have asserted that the bees convey them elsewhere: they take them, indeed, not to convey them anywhere, but to devour them.

"Thus nature has not charged bees with the care of placing the eggs in the cells appropriated for them, but she has inspired females themselves with sufficient instinct to know the species of eggs they are about to lay, and to deposit them in suitable cells. This has already been observed by M. de Reaumur, and here my observations correspond with his. Thus it is certain that in the natural state, when fecundation takes place at the proper time, and the queen has suffered from nothing, she is never deceived in the choice of the cells where her eggs are to be deposited; she never fails to lay those of workers in small cells, and those of males in large ones. "--Francis Huber, New Observations on the Natural History of Bees Letter III, August 1791

This is also available in a more modern translation by C.P. Dadant in "Hubers new Observations On Bees", Volume I, Letter III, page 77 of the 2012 edition. There are also letters from Huber detailing more on some of these experiments that were published in French and serialized in ABJ from 1922 to 1924.


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## Daniel Y (Sep 12, 2011)

beepro said:


> What do you mean by age criterion of the larva? You
> mean choosing a larva that is older than 4 days to make
> a queen from it? If they have a choice they will select
> younger larva so that it can feed on more RJ to develop into
> ...


Using only what is on hand is not a selection.

Are you certain they would select a younger larva? How do you know this?

How can you be certain that here are not other factors the bees select by? Such as location of the cell, Health or other potential attributes of the larva. Maybe they select the largest of a given age.

I have not been able to do any tracking of acceptance of various larva. Might be interesting what it might reveal though.


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## Eduardo Gomes (Nov 10, 2014)

Michael Bush said:


> "I put a considerable number of bees into this hive; and, in the upper part, confined a very fertile queen that had just finished her great laying of male eggs; therefore she had only those of workers to lay, and she was obliged to deposit them in the surrounding large cells from the want of others.


Thank you Michael . You were very kind to put your information available.

My English is not famous and therefore may have escaped me something, but to me this experience, very well designed, does not seem sufficient to clarify this debate, since the conditions are different: In the experiment we have a fertile queen that not is to be replaced ; the experience nothing refers to the presence of queens cups.
In my opinion this experiment was not designed to test the hypothesis : *the bees change / carrying eggs for/out queens cups* .


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## Eduardo Gomes (Nov 10, 2014)

Eduardo Gomes said:


> In my opinion this experiment was not designed to test the hypothesis : *the bees change / carrying eggs for/out queens cups* .


For a deeper analysis of the case see: http://link.springer.com/article/10.1007/BF02223969#page-1

"*Apart from what is known about queen rearing, relatively little is understood concerning its potential regulation during phases of queen replacement.*" in Worker regulation of emergency queen rearing in honey bee colonies and the resultant variation in queen quality, S. Hatch, D. R. Tarpy 1, * and D. J. C. Fletcher, 1999
This phrase summarizes a very careful view , as I understand it must be for now , of our knowledge about this process.


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## Michael Bush (Aug 2, 2002)

In Volume two he has a few more situations where the bees have good reasons to move eggs but never do. Also he talks more about some of these experiments in his letters. Basically, though, I have yet to hear of a situation that couldn't be explained by other behavior that HAS been observed nor have I ever heard of someone actually seeing the bees put an egg in a cell and anchor it there, nor move a larvae and put it in a cell. It's easy enough to see them remove an egg or remove a larvae, but I have never heard of a eye witness to them putting one in a cell from somewhere else. I would be more inclined to believe they could move a larvae than an egg but I doubt that as well.


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## Eduardo Gomes (Nov 10, 2014)

The nature and its processes do not mind anything that we hear about them or read about them. Nature is what it is and we must study it carefully and with humility.

I will not make my point saying that after the Huber studies already much water has passed under the bridge .

Michael I'm not saying anything you do not already know . I value your work and your convictions . But I do not read and hear only what you say . I know you are able to accept this my position.


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## antonioh (Oct 15, 2014)

> "Here we show that honeybee queens are not reared at random but are preferentially reared from rare "royal" subfamilies, which have extremely low frequencies in the colony's worker force but a high frequency in the queens reared." Rare royal families in honeybees, Apis mellifera. Moritz RF1, Lattorff HM, Neumann P, Kraus FB, Radloff SE, Hepburn HR.


.

No question about that, but a doubt came into my mind...

" ... preferentially reared ..." that means usually but not always . It means that if there is nothing more available, any larva will do. On the other hand, if there are many larva available, only the "chosen ones"will be used. In an orfan colony, the bees rear two, three , well, less than ten queens. 

So, how to explain when rearing queens and doing picking, in fourty larva, they raise 80% ? Surely the beekeeper didn´t pick only the royal subfamilies. Why the bees don´t take only these ones and reject all the others ? Why this high level of acceptance ?


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## Eduardo Gomes (Nov 10, 2014)

antonioh said:


> .Why this high level of acceptance ?


There are two hypotheses according Wisnton (1987) :
1 ) picking is unnatural situation , wich bees were confronted with totally/or most unrelated individuals to rear ( when there is no dog hunt with a cat! Evolutionary wisdom??!! ) ;
2 ) larvae were transferred to cells containing royal jelly , wich may mask kin-related odors .

Putting the question around : why in more natural situations bees prefer to rear the larvae closely related to her patrilines? (Visscher, 1985; Page and Erikson, 1984).

To clarify I am not following the line of inquiry about the royal subfamilies. The line of research that I know best and that seems more worked up is the " kin recognition functions in colonies " .

To conclude I liked to say that in the absence of the best, orphaned bees will try to make a queen from a nail. I have seen with my eyes , queenless hives where bees make royal cells with larvae not fertilized, created from laying eggs bees.


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## centrarchid (Jun 13, 2014)

What is the meaning of rare in the context of honey bee families? If a queen mates with a dozen or more drones, then all families based on sire / drone will appear to be in the minority relative to the whole of the queens fertilized egg output.


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## Eduardo Gomes (Nov 10, 2014)

The honeybee queen, Apis mellifera, mates with extremely large numbers of males resulting in up to 45 subfamilies that coexist in the colony (Palmer and Oldroyd 2000).

You can read more yourself. The link is available and is written in English.opcorn:


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## centrarchid (Jun 13, 2014)

If the number of patrilines (drone based subfamilies) is high and roughly equally represented in the queens fertilized broods, then any one will appear rare relative to the whole. What is it that makes one of the rare subfamilies more likely to be used to generate the new queen? Numerical dominance of her subfamilies worker force that has say in selection process by picking out like smelling larva? Or do the workers, regardless of their subfamily pick the "rare royal subfamily" based on some combination of alleles they can measure by smell? Can they pick subfamilies that are more or less homozygous for the special alleles.


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## antonioh (Oct 15, 2014)

> larvae were transferred to cells containing royal jelly , wich may mask kin-related odors .


 No. I do dry picking.



> wich bees were confronted with totally/or most unrelated individuals to rear


 No , when you use the larvae from the same hive.

The question remains , why 80% acceptance instead of 10 or 20% ?


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## GusK (Jan 24, 2013)

antonioh said:


> No. I do dry picking.
> 
> The question remains , why 80% acceptance instead of 10 or 20% ?



Simple enough... because when you graft, you place those grafts in cups which are upside down. They mimic swarm cells or supercedure cells laid by the queen, they are not worker larva in worker cells anymore. In other words, they are not selected from worker cells which are not upside down. Hence the 80% acceptance rate, instead of 10 or 20%.

In the experiment, they did not graft. They just removed the queen of an existing hive. In such a situation, the bees have 100% control over who gets to become a queen larva. And in those circumstances, not all, but a large amount of queen larvae are not well represented in the worker population of the colony, if at all.

Thus the royal allele hypothesis. The authors' conclusions from the paper:



> If larvae from rare subfamilies are preferentially reared to queens there
> are two possible explanations. The first is that workers prefer
> to rear uncommon genotypes to queens in a frequency
> dependent selection process. Although we cannot exclude
> ...


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## Eduardo Gomes (Nov 10, 2014)

GusK said:


> In the experiment, they did not graft. They just removed the queen of an existing hive. In such a situation, the bees have 100% control over who gets to become a queen larva. And in those circumstances, not all, but a large amount of queen larvae are not well represented in the worker population of the colony, if at all.


In addition to saying Gusk, and that explains much, I would add , in a orphaned beehive, the bees will do everything in their power to make queens, virtually all that looks like an egg ( see more closely what I said about the actual cells made from not fertilized eggs in hives with bees laying).

But the question that responds to the core of this thread I opened must be another : *why in a more natural situation bees will not randomly pick the larvae of their future queens *? That is the question!!

António, I ask myself if this issue that has accompanied me almost since I started in beekeeping, it is a folly and a delusion on my part, why is starting to appear much research around it? Are all these researchers deluded try to answer how many angels fit on the head of a pin as was done in the middle ages? Surely not !. Place the keywords in google and please confirm what I am about to say.


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## antonioh (Oct 15, 2014)

> Simple enough... because when you graft, you place those grafts in cups which are upside down. They mimic swarm cells or supersedure cells laid by the queen, they are not worker larva in worker cells anymore.


 Indeed . In fact, if you rotate the frames around an axis perpendicular to their surface at regular time laps, they will not raise queens.

But the question remains :Why the bees do not raise only the larva of the lineage but instead raise all the larva?

The link to pubmed site only displays the abstract. Here is the link to the full text:

Rare royal families in honeybees, Apis mellifera

The experiment was made by orphanising a colony, thus leading the bees to raise an emergency queen…

Traditionally, beekeepers establish a ranking of queens , according to their origin. Maybe a myth, as beekeeping is full of myths among old (and young) beekeepers.

So, the best ones would be the supersedure queens, raised in the presence of the old decaying queen. Then the swarming queens and, in the end of the row…the emergency queens, raised by anxious orfans in a hurry from any egg they find…



> why in a more natural situation bees will not randomly pick the larvae of their future queens ? That is the question!!


 Well Eduardo, it seems that in swarming is absolutely random. In supersedure we don´t know, which leaves us with the selection being done only in the emergency queens… those of the end of the row…

Eduardo, you know I am not an investigator, but a practical beekeeper. Even my job is based on practice. So on the practical side , what should we conclude ?

Does this means that the chosen lineage in emergency queens is not the best one ? Or , on the other hand, if is the best one , why the poorer performance of emergency queens, when compared with artificial raised queens ? 

And I can include in this “artificial raising” also the more “natural” methods indicated by Mr. Gilles Fert, for those raising queens is small scale : Miller method and Alley method, to which I add Perret-Maisonneuve method, this last one giving the beekeeper a more accurate selection of larvae. But all these restrain the larvae available to the bees.

Should we consider swarming as the offspring of the hive ? We know that in animal kingdom, the best is picked for the offspring, so, in swarming, the bees do their best to their offspring ? Or if they don´t do their best, shouldn´t they?


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## antonioh (Oct 15, 2014)

Sorry , the link didn´t work.~

Here it is :


http://www.researchgate.net/publica...Apis_mellifera/links/0fcfd50e7f9f516f5e000000


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## Daniel Y (Sep 12, 2011)

antonioh said:


> But the question remains :Why the bees do not raise only the larva of the lineage but instead raise all the larva?


The most obvious answer to that specific questions would be that no such Royal selection in fact exists.

It is like doing an experiment designed to test a theory. and then when it does not calling the experiment flawed.

We all know that this extra selection must take place if only it where not for all these instances that no such thing happens.

In comparison to the claim that bees will rear queens from cells that are upside down. Now that I see happen. and I see no influence that it was some infrequent larvae.

Now I do have an additional question. How would they possibly know that any larva reared as a queen has come from any particular genetic line. much less that it is a genetic line more or less suitable as a queen?


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## GusK (Jan 24, 2013)

Daniel Y said:


> Now I do have an additional question. How would they possibly know that any larva reared as a queen has come from any particular genetic line. much less that it is a genetic line more or less suitable as a queen?


Sorry, I can't find a link for the entire paper. I downloaded the pdf a while ago. Anyway, to answer your question, they determined worker subfamilies' frequencies and distribution by genotyping worker brood. After the queen was removed, they did the same to the queen cells that were built after de-queening. 

You would think that the queens would be somewhat of a statistical sample of the workers, that is, more or less the same patrilines would be represented in the queens. That was not the case. Read above, where I posted the authors' conclusions. 

On the ABF website I listened to a Dave Tarpy lecture from I believe 2013. He did a similar study and got the same results. He said that he did not publish it because he didn't know what to make of it (yet?).

Now, if these larvae are more suitable as queens, is anyone's guess. Mating aside, I would think nutrition is the number one factor. But I do think this warrants further study.


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## Acebird (Mar 17, 2011)

I don't think there is any thinking going on when reproduction plays out. Honeybees are clearly a cast system so queens from queen lineage would make sense to me if that was an option. I don't see any advantage to this for the future queen though. The offspring of slaves have as much a chance for healthy offspring as do a king and a queen, maybe more.


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## JRG13 (May 11, 2012)

It's not a true caste system Ace, all of the bees are related to the queen maternally, and some will be related more on the paternal side depending on how related the queen's drone father was to the ones she mated with. Also, the role in the hive is more usually related to age, not genetics. Here's my thoughts on it, I think the grafting issue, Gus hit it on the head, it's a pure environmental response to finding larvae in a vertical cup to raise the queen. Royal selection may or may not happen depending on a lot of factors. From what I've seen, there's some factors that need to be addressed in any further studies. First, when did the workers get genotyped, before, after, or during the queen rearing. For me, you would have to genotype the workers laid surrounding the queencell, and then randomly select others laid at the same time elsewhere in the hive that were laid around the same time the selected queens were. I don't know if it's known how a queen stores sperm (i.e. is it all mixed, or stored in batches etc..) or chooses what sperm to fertilize eggs with or if it's completely random, but if she cycles through sperm in batches it's possible at the time they had to make cells, genetic selection was limited. Second, queencell placement in a hive under emergency conditions isn't always random, they tend to make queencells in certain parts of the comb. Ever notice if you use horizontal wires, they will almost always start cells where the wires pass through so the comb so the wire is always at the base of the queencells, and there's typically another batch of queencells just opposite each other on the same comb. So comb structure needs to be homogenized because they tend to make queencells at spots where there's imperfections in the comb. Lastly, I still need to read the paper, but it seems that the royal alleles are more linked to the drones passing them on so you could set up a study there using a mixture of drones related to the mother and some totally different line and you would see a preference in daughter queens.


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## Rader Sidetrack (Nov 30, 2011)

Acebird said:


> The offspring of slaves have as much a chance for healthy offspring as do a king and a queen, maybe more.


You appear to think that worker bees are slaves to the queen! :no: 

Try this on for size ....


Michael Bush said:


> It doesn't take long watching bees in an observation hive to conclude that the workers are in charge and the queen is their slave who they will replace at the drop of a hat if she doesn't do her job.


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## centrarchid (Jun 13, 2014)

Daniel Y said:


> Now I do have an additional question. How would they possibly know that any larva reared as a queen has come from any particular genetic line. much less that it is a genetic line more or less suitable as a queen?


If like similar done in mammals, then it would be based on scent where patriines will differ based on the bowkay of volatile compounds produced.


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## beepro (Dec 31, 2012)

I find that the subject of bee genetics rather facinating. And want to know more when I do queen grafting.
When I was raising fancy guppy the selection process is pretty much
the same trying to find the best traits to keep. As for the sperm storage, my observation on the guppy and the bees are much alike. Maybe in all insect species they store it the same way? For example, when the new male guppy mated with the female a new generation is produced that reflects the new male's traits. Once the new male's sperm is used up judging by the color of the young guppy, the older sperm from another male is use to produce the next generation. Female guppy like the queen bees is able to store the male sperm for many generations to come. In my bee yard I have noticed that every new batch of young bees have the same color trait coming from the production queen that is open mated. Some worker bees are Italians, Cordovan and pol-line in certain months. So my conclusion for now is that the queen will store the sperm of the drones in a LIFO (last in first out) basis in an organized way. Once one batch of sperms are used up then the next will follow coming from a different drone. It is not a random mixed of different sperms when she lay for the mixed batch of young bees. If it was then I would of seen many mixed of bee colors in any one batch that hatched. So far the young bees are all uniform in color.


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## Seven Hills (Apr 7, 2011)

I don't know if someone has already said this but could they find the larve they like and build the queen cell around it.


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## jonathan (Nov 3, 2009)

beepro said:


> So my conclusion for now is that the queen will store the sperm of the drones in a LIFO (last in first out) basis in an organized way. Once one batch of sperms are used up then the next will follow coming from a different drone. It is not a random mixed of different sperms when she lay for the mixed batch of young bees. If it was then I would of seen many mixed of bee colors in any one batch that hatched. So far the young bees are all uniform in color.


That has been disproved experimentally many times. Sperm in the queen spermatheca is completely mixed.

*Does Patriline Composition Change over a Honey Bee Queen’s Lifetime?
Robert Brodschneider Gérard Arnold Norbert Hrassnigg and Karl Crailsheim*

Insects 2012, 3(3), 857-869



> All these studies clearly demonstrate that many patrilines are present simultaneously in a honey bee colony, and therefore the scenario of “total” sperm clumping, i.e., the usage of most of one drone’s spermatozoa before using the spermatozoa of other drones, is rightly considered to be wrong.


http://www.mdpi.com/2075-4450/3/3/857


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## Marysia2 (May 23, 2014)

jonathan said:


> That has been disproved experimentally many times. Sperm in the queen spermatheca is completely mixed.


This would seem logical to the survival of a species. If the sperm of one drone were tainted, mutated or non-viable for some reason, the hive would die out if only his sperm were used for several generations. That would be like putting all your eggs in one basket, so to speak, no pun intended.


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## Daniel Y (Sep 12, 2011)

GusK said:


> Sorry, I can't find a link for the entire paper. I downloaded the pdf a while ago. Anyway, to answer your question, they determined worker subfamilies' frequencies and distribution by genotyping worker brood. After the queen was removed, they did the same to the queen cells that were built after de-queening.
> 
> You would think that the queens would be somewhat of a statistical sample of the workers, that is, more or less the same patrilines would be represented in the queens. That was not the case. Read above, where I posted the authors' conclusions.
> 
> ...


I have one other thought in regard to why bees woudl rear a high percentage of grafted cells and abandon any such preferences for a Royal line. Grafting and altering the cell that a larva is in has then leap frogged or over ridden any selection process. A larva's presence in a queen cup indicates to the bees a selection has been made. just who or what made that selection is not apparent to the bee. The bees are no longer paying attention to lineage. Selections was supposedly made they now apply themselves to rearing a queen.


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## Michael Bush (Aug 2, 2002)

>That has been disproved experimentally many times. Sperm in the queen spermatheca is completely mixed.

My observation is that it is somewhat mixed, not completely mixed. Neither is it exactly layered. Too often you see a hive change proportions of color over time. If it was completely mixed this would not happen. If it was completely layered you would have all of one color for a while and then a shift to a different color. This does not happen either.


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## jonathan (Nov 3, 2009)

Didn't Bernhard post something recently about brood nest temperature having some influence on bee colouration/darkness. Maybe there are other factors in play as well.
Another possible explanation for a change in colour of the workers could be an undetected supersedure of a queen.


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## Lauri (Feb 1, 2012)

If you have ever tried to replicate certain behaviors (Such as a 2 queen colony going into winter) Basically, Cells raised under a specific stimulus, you'll find it is most difficult.
There are different situations or conditions queen cells are raised. 
-Supercedure cells -to replace an older, poorly mated or failing queen.
-Swarm cells
-Grafted (human selection)
-walk away nucs

If you compare supercedure cells and swarm cells, the first and most obvious difference is the shape and placement of the cells. 

Your colony induced supercedure cells are usually near the center of a frame. A reliably warm location within the hive.










Now I can manipulate my queen cell hatch in the incubator a full day one way or the other by slightly adjusting the temperature. Even one degree warmer means a shorter gestation. One degree cooler and I can delay the hatch a day. Why would I do that? I've done it just to see if there is any impact on the virgin. Sometimes outside weather conditions can make me want to change a hatch time slightly..Also in order to find the perfect temp for my incubator, I had to play with my temps a bit. My gestation rate is perfect, almost to the hour at 92 degrees. 
After hatching, I will hold and harden off my virgins at a slightly lower temp.

I've found no obvious difference in the queens from a slightly varied hatch. But looks aren't everything. Performance and longevity are also importaint and woudl need to be observed over a period of time to be sure.
When your average gestation is only 16 days, a full day of variant may be significant. 

With my strain of bees, my supercedure virgins don't kill the queen upon hatching. I often find newly hatched virgins and the queen together in the colonies and only when they new young queen is successfully mated and laying does the older queen finally disappear. I see this in early spring. I also see it late summer/fall when I have a marked queen and laying daughter going into winter together. By next spring the daughter is the only queen residing in the hive.

Is this tolerance for multiple Queen in the hive due to selection of like or favored genetics or something else- like temperature or length of gestation perhaps? It's likely many things we don't understand or realize are significant. It should all be taken under consideration if you really want to rear the best queens possible. I believe good observation & adaptation can be very importaint when trying to improve your efforts.


Now lets compare the swarm cells. Bottom of the frame, usually in the lower brood chamber. Cooler. Slightly longer gestation? Built under a swarm impulse, which likely has the entire colony excited to a degree. The queen and swarm are usually gone before the virgins hatch, but depending on the situation, multiple virgins may also hatch & throw cast swarms without being destroyed by the first virgin. Also quite tolerant of multiple queens in some instances. 










Then you have our human selection, grafts by age of larva only. Randomly selected by humans with none of the other critical fine detail considered, exposed to temp fluctuations during handling. Started for the first 24 hours under emergency conditions, at least in the bees mind.










Handling of the cells, whether it is the transfer to incubator or transfer to nucs will expose them to a few slight temp variations during their gestation period. 
How are these resulting cells or virgins ultimately received & perceived by the new colony? Unlike the supercedure and swarm cells, these queen cells were not there idea.

I notice I seem have better success with direct release introduction of my virgins compared to many other people. Is that due to careful temperature control and resulting gestation length? 
Anytime I place capped cells, I always have a small disposable hand warmer with the cells until they are placed.











Top shelf in my incubator is set at 92 degrees. Bottom shelf is 90-91 degrees. That variance makes a difference in my hatch rate. I generally incubate cells on the top shelf, Place virgins on the bottom shelf as they hatch. If I have no other cells in the incubator, I'll slowly drop my temp slightly until I place the virgins. 




























Anyone that has banked queens, both mated and virgins knows there are those that are highly favored by nurse bees right long side of those queens that are less favored. Right down to some that are totally ignored. Photo below isn't the best to show this, but the queen on the far right is obviously favored over the others. Although I generally don't bank my queens, I have seen a benefit to doing so for a short period for selection purposes on my end, especially if I am short on available mating nucs and long on virgins. I'll always place the favored queens first. 










And then there are the cells from the walk away nucs:






















As far as is drone semen being totally mixed or not, I have to say my experience with grafting & the grid system leads me to believe, while semen is somewhat mixed, it is still slighly clumped and the queen will lay patches of eggs that are genetically identical (semen from only one drone) How big those patches are exactly I have not yet tried to determine. 










For example: 
I have a favorite queen that is quite dark. I was surprised and a little disappointed when her first batch if daughters were nearly all blond-tiger tail.
Her second batch was all jet black. 
Third batch was dark striped. 
That queen is going into her third winter and I have probably grafted 50 batches from her. Results are predictably different. I get four colors out of her, but each individual batch is remarkably alike.

But I find that with other queens too. Leading me to believe it is the small mixed up semen 'clumps' that are the reason for the similar phenotypes. 
Now I usually graft from a small portion of a frame and of course the grid is limited on space. If I really wanted to get a more mixed variety, I would graft from other areas on a full frame or use 2 frames for my donor larva.










So when creating queen cells, you have different motives, different temps, slightly altered gestation length, even different shapes & placements of the cell itself. The size of the cell may not make a difference in the size of the resulting queen, but it the size, shape & placement of the cell something the bees recognise and respond to more than we realize? Not to mention the original question in the OP. How is selection determined in the first place? 

Personally I believe they know when vigor or inbreeding is an issue and if left to naturally select on their own, they will facilitate beneficial changes on their own. 

When human selection is the only method, the necessary changes may not occur and result in eventual lack of vigor.

That could well be why II queens have a reputation for being superceded quickly. If not enough drones were used from different, unrelated sources, there could be not enough genetic variation to satisfy the colony.

I see that in horses too. My stallion loves most mares, but his full sister and he quite obviously dislike one another. Natures way of avoiding inbreeding.

I can tell when a mare is very close to actually ovulating by his behavior. I thought he was drooling at first, but when a mare is ready to be bred, liquid just poors out of his nose. In a 7 day heat cycle, he'll only do this for one day.If I see his nose dripping, I know I better get that mare bred ASAP or I'll miss her. 

My point is there are selection methods humans will never be tuned into. But if we take our clues from the critters, we can still get the job done.


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## Eduardo Gomes (Nov 10, 2014)

Lauri said:


> It's likely many things we don't understand or realize are significant. It should all be taken under consideration if you really want to rear the best queens possible. I believe good observation & adaptation can be very importaint when trying to improve your efforts.


I found your post very complete and clear. The photos are great .
I quote this phrase because it is precisely this attention of us all, in particular for researchers and queens breeders, that will make us go a step further.


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## centrarchid (Jun 13, 2014)

I very much like the effort some invested in this thread as it was quite educational. 

The problem I see that has been apparent in other threads as well involves the discussion drifting aware from the core of thread question and usually gravitating towards culture methods beyond or not considering context of the original question.


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## beepro (Dec 31, 2012)

Michael and Lauri's worker bees observation is pretty much inline with mine so far.
If thousands of semen from the same drone is stored in batches then grafting from the same full
size frame should give the same color of the daughter queens. If the daughter queens are of different
color then the sperm of the drone is not store in batches but rather it is randomly fertilized. Either that or
the queen has a way to select the sperm from different drones herself to fertilize them. So far I have 3 hives,
one Cordovan, 1 Italians and 1 pol-line bees to monitor the brood changes over months of observation. Once my full size
non-grafting frame is done I will do this experiment on 4 days larvae. It is exciting to be able to do your own
little experiments to learn from them. Keep up the good work and thanks for sharing your results right on topic.


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## Acebird (Mar 17, 2011)

I have been reading this thread and don't understand much of it but I don't see why a queen couldn't have genes in her makeup that would be for multiple color expression so the idea that different colored offspring have to come from the drones sperm I don't get.


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## centrarchid (Jun 13, 2014)

Acebird said:


> I have been reading this thread and don't understand much of it but I don't see why a queen couldn't have genes in her makeup that would be for multiple color expression so the idea that different colored offspring have to come from the drones sperm I don't get.


That can explain about half of the variation unless she is homozygous for a recessive light colored set of alleles and bred to a range of drone genotypes.


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## Daniel Y (Sep 12, 2011)

Ace, The queen would carry two genes. She will only pas one or the other to her offspring. while each sperm only carries one that mix of sperm could have come from many drones. S0 say she mated with 20 drones. there are a potential 20 various colors in that mix.

Now at this point it looks as if the queen is outnumbers 10 to one but that is not the entire story. The queen will in fact contribute 50% of the genetic makeup of every bee in the colony. While 1 out of 20 drones in the mix would contribute on average 2.5%. So the entire colony as a whole is 50% the queens genetics and 50% a combined total of all the drones she mated with.

So any layering in the sperm should be expressed enough to be observable. That is due to the idea that an influence that should on averages be only 2.5% has for a period of time nearly that entire 50% share of the drone influence.

The queen still has her contribution. so what you would be seeing is a difference in the remainder of the contribution.

That additional contribution could and would be more or less influenced by the queens particular genes. Say for example both genes of the queen are dominant. the influence of the drones variety would possibly be less pronounced.


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## beepro (Dec 31, 2012)

Ahh, ha. That makes more sense now. We're back to the
dominant and recessive genes again. Here's my observation for this year. The DCA is 
contributed by a majority of Italians, Cordovan and Pol-line bees. No Carnis drones are observed in
this late Fall.
So a Cordovan (recessive) queen vs an Italian (dominate) queen locally open mated would be like:
Cordovan x Cordovan drone = a Cordovan worker bee.
Cordovan x Italian drone = an Italian worker bee.
Cordovan x Pol-line drone = a Pol-line worker bee.
Italian x Cordovan drone = an Italian worker bee.
Italian x Italian drone = a regular Italian worker bee.
Italian x Pol-line drone = a strong Pol-line worker bee.
This year there are no Carnis drones because of the late Fall mating of the queens. So what do you think is
the color of the worker bees if the Cordovan queen got mated with the local Carnis drones in the early Spring time?


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## jonathan (Nov 3, 2009)

Honeybee abdomen colour is not governed by a single gene dominant-recessive scenario.
Jerzy Woyke published a paper on this years ago.
In general terms yellow is dominant over black but only as a rule of thumb as there are several genes and modifiers in play.
This means that if you have a pure race black queen such as Apis mellifera mellifera and she mates with yellow drones you should see some yellow banding in the workers.

International Symposium on bee biology,
MOSCOW, USSR, August 1976
THE HEREDITY OF COLOUR PATTERNS IN THE HONEYBEE
Jerzy Woyke
Results show clearly that yellow coloration is dominant over black
Inheritance of colour patterns in the honeybee is governed by 3 major allelic genes
having their expression modified by 6-7 polygenes with alternative alleles for light
and dark.

This paper used to be online but I cannot find it any more.


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## Rader Sidetrack (Nov 30, 2011)

The paper Jonathon referenced is available here:

http://www.researchgate.net/publica...e_Heredity_of_Color_Patterns_in_the_Honey_Bee

but requires registration to view (and possibly _may _require a fee).


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## antonioh (Oct 15, 2014)

The image of the full text, appears on the right side, below publicity.

Click on the full text image and it will open the full text pdf.


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