# Drone genetics



## sqkcrk (Dec 10, 2005)

Favor? Dominant traits are still dominant traits. Consider that it seems as though an AHB queen that mates w/ nonAHB drones exhibits AHB characteristics and​ that a nonAHB queen that mates w/ AHB drones exhibits AHB characteristics too.


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## Phoebee (Jan 29, 2014)

You caught me while reading this:

http://www.glenn-apiaries.com/principles.html

My question comes because my teensie little apiary has only two hives, and I'm about 3 miles and one mountain gap from the nearest other hives that I know of. But they're recently-built nucs, and if I'm doing the bee math right, most of the drones presently flying are still the offspring of the colonies from which the nucs were built. In other words, it could well be I've got a lot of drones of mixed West Virginia ancestry, born on out on the almond groves, plus a few recent additions from the introduced queens. 

If, in fact, each drone is a random mix of the 16 chromosomes of his mom, and I've still got four moms in the mix, then there is pretty good diversity present from such a small apiary.

One of my queens evidently quit laying recently and the hive created a handful of supercedure cells. I'm planning to let them go ahead and raise a queen. Plus a friend is offering me a nuc that is supposed to have VSH ancestry. So maybe I'm not as bad off as I though considering the apiary size.

Eventually my drones should find the hives out past the mountain gap, and they will find mine, and maybe there will be drone yards that open up the opportunity for any queens I raise. So hopefully there are more genes around than I originally thought, possibly with some local selection pressure already applied.

None of them should be africanized. In fact, so far they're the gentlest little sweethearts you could ask to keep.


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## sqkcrk (Dec 10, 2005)

There are a lot more bees out there than anyone knows about. You'd have to be really secluded before you would end up w/ unmated queens or poorly mated queens.


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## JSL (Sep 22, 2007)

You are correct. There is a great deal of genetic diversity in your four colonies. Honey bees have one of the highest recombination rates, and as Mark pointed out, there are a lot of colonies out there, even if you can't see them.


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## Kamon A. Reynolds (Apr 15, 2012)

Shoot I know a guy who raise 2000 plus queens a year and doesn't have drone colonies at all. I feel this is negligence on his part. Yes there are alot of other bees around but I would hope that all producers/sellers would have drones of exceptional qualities to insure not only the best mated queens but also the toughest genetically. 

Unless I was trying to mate 50-100 at a time on a consistent basis I wouldn't bother.


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## Juhani Lunden (Oct 3, 2013)

Phoebee said:


> Do bee eggs favor the maternal contribution?


This would be an interesting point to study. I would like to know more of that myself. I once read (did not copy the text to my files!) about the "fight" of genes, the idea of that being that a gene wants its own replicas to the offpring. Sounds scifi, doesn´t it?

Human dna genealogy is my hobby. In these results it is interesting to notice that female persons have about 20-30% of their dna cousins having similarities in mitochondrial dna(this is coming from mother, in the egg cell, both to girl and boy babies), the tested males have around 5% of their dna cousins having similarities in mitochondrial dna. Females have more dna cousins coming from mothers side than males.


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## Phoebee (Jan 29, 2014)

Juhani Lunden said:


> Sounds scifi, doesn´t it?


Heh, I WRITE science fiction. No kiddin'. But with biology, I guarantee you that nature has more imagination than all the members of the Science Fiction Writers of America put together.


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## Juhani Lunden (Oct 3, 2013)

My e-mail is [email protected]


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## Phoebee (Jan 29, 2014)




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## Juhani Lunden (Oct 3, 2013)

All the experts in genetics on the forum, where are you? Here is a question to show off.

Do bee eggs favor the maternal contribution?


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## sqkcrk (Dec 10, 2005)

Juhani, let me show my ignorance. A queen comes from a fertilized egg. So she is the result of her Mother and her Father. Are you asking whether her Father's contribution is passed on in her eggs? Or if just her Mother's contribution/genes?

Dangerously thinking out loud here at 5AM. The unfertilized egg will produce a drone. The sperm came from a drone. When used to fertilize an egg (a drone in its most basic form in essence) queens or workers are produced. They're all drones. Just that some are twice the drones that drones are.

Where is the mitocondrial DNA in a honeybee? In the drones?


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## Juhani Lunden (Oct 3, 2013)

If there is some favoring it happens when egg is produced. Queen is diploid, and she uses all her genes to produce the eggs. Of course there is mothers and fathers contribution in the eggs, but are there more genes from mothers side? This would be an advantage, if some genes affect for instance egglaying capacity and it comes from her mother. 

Mitochondrial dna is in the egg cell and by cell divisions it goes to all cells of all creatures.


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## deknow (Jul 17, 2006)

I don't believe there is any bias towards contributuon...except for the ironic twist that all male traits come from the mother with no paternal contribution.


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## JSL (Sep 22, 2007)

Greg Hunt and others have been working on this concept. They may have a more recent publication of their findings, but below is a short abstract from a thesis out of Greg Hunt's lab. The work suggests a slight bias for certain genes from the male or female. 

http://gradworks.umi.com/35/56/3556196.html
*Characterizing the honey bee genome for parent-of-origin effects on gene expression*
by _Emore, Christine Michelle_, Ph.D., *PURDUE UNIVERSITY*, 2012, 126 pages; 3556196


*Abstract:*
Genomic imprinting has been studied extensively in higher plants and mammals where it plays a crucial role in development and has been linked to cancer and other diseases in mammals. The honey bee is one of the few invertebrate species in which genomic imprinting has been studied and could give new insights into parent-of-origin effects on behavior. The hypothesis tested was that parent-of-origin effects do exist in the honey bee and was done by using RNA-seq on two life stages of reciprocal crossed honey bees. Two different RNA-seq systems were used. The project was first done using the SOLiD 4 system, but validation of the results was poor due to a large number of false positives created from systematic and alignment biases that were not taken into account by the statistical methods used. The second system was the Illumina HiSeq 2 and the study was done using replicates and generalized linear mixed model statistics that could model and correct for systematic biases introduced during sequencing and alignment. The Illumina study showed that parent-of-origin effects do take place in the honey bee and the results were validated by sequencing PCR products of parent-of-origin effect transcripts using the Illumina MiSeq. The Illumina study showed that parent-of-origin effects are more evident in adult honey bees than in early larval stages, contrary to mammals and plants where the majority genomic imprinting takes place in early development. Regardless of life stage parent-of-origin effects in the honey bee are primarily biased towards maternal expression with the strongest maternal expression found in nuclear copies of mitochondrial genes or numts. The numts are not complete genes but gene fragments or pseudo-genes that are actively transcribed by the nuclear genome, which suggests a biological function and evolutionary selection for conserved maternal expression.


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## Juhani Lunden (Oct 3, 2013)

JSL said:


> Greg Hunt and others have been working on this concept.
> ...Genomic imprinting has been...


I´m not sure imprinting is what Phoebee meant, I sure didn´t. I meant the formation of an egg, and the genes which come in it.


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## Richard Cryberg (May 24, 2013)

Juhani Lunden said:


> All the experts in genetics on the forum, where are you? Here is a question to show off.
> 
> Do bee eggs favor the maternal contribution?


All critters have two kinds of DNA, nuclear and mitochondrial. Mitochondrial DNA is passed intact from mom to offspring with Dad seldom making any contribution at all. This DNA is not in the nucleus and does little besides run the energy chemistry in mitochondria.

Nuclear DNA is totally scrambled by crossovers before egg formation happens. So, by the time crossovers are done there are no maternal or paternal chromosomes left. There can be individual genes or DNA regions that carry epigenetic marks that define them as being maternal or paternal. And, it is always possible such marks can influence if the marked DNA region goes to the egg or to a polar body. In genetics it is a good bet if you can think up a scheme mom nature will already be using it someplace.

In drones there is no crossover process so chromosomes in haploid sperm are just like they are in the haploid drone.


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## Juhani Lunden (Oct 3, 2013)

Richard Cryberg said:


> And, it is always possible such marks can influence if the marked DNA region goes to the egg or to a polar body. In genetics it is a good bet if you can think up a scheme mom nature will already be using it someplace.


Any studies, publications?


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## Michael Bush (Aug 2, 2002)

> But is it this random in bees? 

Yes.

>Do bee eggs favor the maternal contribution? 

That depends on what you mean by this sentence. The haploid egg produced by the queen is a random single set built from pairs (diploid). Those are just random but they are both maternal contributions to the egg. The other half of the worker or queen produced from an egg is the single set (haploid) that comes from the drone's sperm. Since the population of bees in the hive has half of their genetics from the queen (always the same queen except in rare circumstances) and the other half from a multitude of drones, the maternal contribution (as in "the current queen's contribution) is higher than any given drone.

So if by "maternal contribution" you mean what the queen who is laying the egg contributes, all the genes that go into the haploid egg come from here. How can there be any favor? If you mean the queen's mother's contribution to that egg getting any preference, no it does not.


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## JSL (Sep 22, 2007)

Juhani,

Imprinting as used in genetic research is not the same as imprinting in the case of young geese "imprinting" on their parents. It refers to the genetic influence imparted by the male or female, in this case bees. This is over my head, but appears to be the first described instance in honey bees where it is not the traditional 50% from mom and 50% from dad at least in terms of expression. I think Hunt did this work in the expression of behavior.


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## JSL (Sep 22, 2007)

As Richard pointed out, there is no crossover in drones, they are "flying sperm bags". The sperm cells from a single drone are identical.


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## Phoebee (Jan 29, 2014)

Michael,

That's what I suspected, but nature has exceptions and I wondered if bees had any way around this. Girls rule in the hive, and the queen's genes would have a survival advantage to being over-represented in the offspring.

According to the article I linked, there is a sex chromosome pair in bee genetics, but it works differently than in humans. The two components of the pair must differ for a female to develop, otherwise the larvae is destroyed. So one can imagine that the bees might cull drones that don't represent them well. In mammals, the example would be male lions killing offspring from a previous male.

Supposedly bees don't mate with drones from their own hive. I've never heard how they manage this ... I would think there must be frequent cases otherwise. Are the resulting offspring discarded?

The queen apparently does not chose her successor ... the workers pick diploid eggs to place in queen cups and feed a steady diet of royal jelly. So that selects particular eggs with particular drone daddies to become queens. Groups of workers share a father. Do they have a bias when selecting eggs?

As I understand the varroa life cycle, the female first produces a male offspring, and mates with it in a closed cell, before producing female offspring.

There are lots of ways to cheat, and I was wondering if bees do.


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## Michael Bush (Aug 2, 2002)

>That's what I suspected, but nature has exceptions

I don't know of any exceptions to the even distribution of genetics in mitosis...

>According to the article I linked, there is a sex chromosome pair in bee genetics, but it works differently than in humans. The two components of the pair must differ for a female to develop, otherwise the larvae is destroyed.

The larvae is destroyed because it is a male in a worker cell and probably also because it is a diploid male. Diploid males are not tolerated. Haploid males are.

>So one can imagine that the bees might cull drones that don't represent them well.

All drones in a hive represent no other single entity except the queen. All of their genes come from her.

> Supposedly bees don't mate with drones from their own hive. I've never heard how they manage this ... 

I'm sure they do on occasion. Supposedly the deck is stacked against it. How they manage it is that the queen tends to fly further than the drones do to mate. She also flies at a different altitude than the drones to get there. This tends to lead to a result where she doesn't mate with drones from her apiary as much as drones from other apiaries. There is no foolproof mechanism at work, just mechanisms that shift the odds.

>I would think there must be frequent cases otherwise. Are the resulting offspring discarded?

The resulting offspring often have that sex determination allele you alluded to earlier, match, resulting in a lot of diploid drone larvae in worker cells which are discarded.

>The queen apparently does not chose her successor ... the workers pick diploid eggs to place in queen cups and feed a steady diet of royal jelly.

Workers do not place eggs anywhere at anytime. The queen places eggs.

>So that selects particular eggs with particular drone daddies to become queens.

The workers can decide which larvae in the queen cups they wish to raise or in an emergency which ones they want to choose. There has been much research on the workers preferring full sisters with research that proved that they did and research that proved that they did not.

>Groups of workers share a father. Do they have a bias when selecting eggs?

Pro:
http://link.springer.com/article/10.1007/BF00300521#page-1
http://onlinelibrary.wiley.com/doi/...sCustomisedMessage=&userIsAuthenticated=false
http://www.ingentaconnect.com/content/esa/aesa/1986/00000079/00000003/art00012


Sort of:
http://afrsweb.usda.gov/SP2UserFile...00/236-Oldroyd--Nepotism in the Honey Bee.pdf
"The degree of discrimination observed within honey-bee colonies is generally weak, never reaching a 2:1 preference for full sisters."

Con:
http://www.ingentaconnect.com/content/esa/aesa/1984/00000077/00000006/art00015
"The results of these experiments do not support the original hypothesis; no phenotypic preference in rearing emergency replacement queens or in choosing grafted larvae was noted. We conclude from these experiments that under normal conditions information concerning the phenotype of honey bee larvae does not allow discrimination between related (half- or full-sister) and completely unrelated individuals."


>As I understand the varroa life cycle, the female first produces a male offspring, and mates with it in a closed cell, before producing female offspring.

No. The foundress mite has already mated and will never mate again. She lays a male (haploid) offspring first and then a female (diploid) every 32-36 hours or so. Those females (her offspring), if they make it to maturity before the bee emerges, will mate with some male (which may be from some other foundress mite or from that same foundress mite). Only the female mites that manage to mature and mate will be viable when the bee emerges.


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## Richard Cryberg (May 24, 2013)

Juhani Lunden said:


> Any studies, publications?


There probably are well over 1,000 publications in the peer reviewed literature that show genes with epigenetic marks that id the parent the gene came from and govern the expression of that gene in the offspring. Maybe 10,000 by now? There are a bunch of human genetic diseases that differ tremendously depending on if the mutant came from Dad or Mom. Here is a real old summary on one sheep mutant written for the public:

Scientific American, December, 2003, pg 108.

You might also find the book "The Survival of the Sickest" interesting reading. It was written to be understandable by about an eight grader.

Off hand I do not know of any published findings that such markings influence if the marked gene goes to a polar body or reproductive cell. I know of examples where there is a strong suggestion this is happening. For example generation of wildly abnormal sex ratios in birds and fish as a result of both known and unknown environmental stimuli. I know of an autosomal recessive feather abnormality in birds that does not follow Mendel's 1/4 rule worth a hoot. It is more like 1/6 or 1/7 in very statistically meaningful quantities of data. This hints that this gene goes to the polar body more often than simple chance. The data clearly say it is not embryo death. I could likely think up other such oddities but in all cases I know of no one who has really chased the thing down to the point it is publishable today. Most professional geneticists denied such things were even possible until the last decade so blamed such observations on dumb students that could not do the experiment right or after fertilization environmental factors which curiously were nearly always unidentified even after extensive experiments to find what these environmental factors were.


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## Juhani Lunden (Oct 3, 2013)

JSL said:


> Juhani,
> 
> Imprinting as used in genetic research is not the same as imprinting in the case of young geese "imprinting" on their parents.


Wow, is that right  tell me something new, Joe.


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## Juhani Lunden (Oct 3, 2013)

Richard Cryberg said:


> Off hand I do not know of any published findings that such markings influence if the marked gene goes to a polar body or reproductive cell. I know of examples where there is a strong suggestion this is happening. For example generation of wildly abnormal sex ratios in birds and fish as a result of both known and unknown environmental stimuli. I know of an autosomal recessive feather abnormality in birds that does not follow Mendel's 1/4 rule worth a hoot. It is more like 1/6 or 1/7 in very statistically meaningful quantities of data. This hints that this gene goes to the polar body more often than simple chance. The data clearly say it is not embryo death. I could likely think up other such oddities but in all cases I know of no one who has really chased the thing down to the point it is publishable today. Most professional geneticists denied such things were even possible until the last decade so blamed such observations on dumb students that could not do the experiment right or after fertilization environmental factors which curiously were nearly always unidentified even after extensive experiments to find what these environmental factors were.


Now we are talking. So the whole phenomenon is argued, no studies in bees. Thanks!


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## JSL (Sep 22, 2007)

Juhani,

No offense meant, just clarifying.


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## Juhani Lunden (Oct 3, 2013)

Ok Joe, 
just came home, made some more mating hives, midsummer, temperature 10 C (=50F), rain and wind. And we got some snow couple days back. Old saying about the Finnish summer: Short but not very snowy.
Some queens have been waiting about 10 days for the weather to get warmer. Doesn´t matter how good genetics my drones have.


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## j.kuder (Dec 5, 2010)

what i want to know with out a whole bunch of sicentific speak is are the drones out of a particular queen an exact copy of that queens genetics. yes- no or maybe?


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## Juhani Lunden (Oct 3, 2013)

The drones of one particular queen are all diffrent, all have one haploid set of genes from the queen, and the queen only. But one drones sperm cells are all equal.


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## sqkcrk (Dec 10, 2005)

So all of the offspring drones from a queen are genetically different from each other? Their sperm is different from each other? But equal? What's the equal part? Same number of chromosomes? I probably don't know enough about genetics to be asking these questions or understanding the answers.


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## Juhani Lunden (Oct 3, 2013)

In meiosis (formation of egg cells and sperm) there is crossing over taking place, that makes each drone individual. But because the drone has only one haploid set of genes, he can make only one type of sperm cells. Sperm cell has to have a haploid set of genes. Drone cannot choose what to but in there, he has to put all his genes in every sperm.

(Crossing over is when the chromosomes set aside and change genes.)


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## j.kuder (Dec 5, 2010)

the reason i asked the question the way i did is i'm trying to develop a simple breeding plan for my small apiary by taking a couple of queens whose genetics i believe are worth saving . requeen all my hives with queens raised out of these 2 mother queens so next spring only their drones will be in my apiary then bring in new queen lines to raise next years queens from and mate with the drones from this years queens. my goal is to load up my bees with hygienic genes. will this work?


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## Juhani Lunden (Oct 3, 2013)

Depends on the isolation, or the saturation of drones in that area. If there are not much other bees, why wouldn´t it work. Raise a lof of drones.


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## BEES4U (Oct 10, 2007)

Here, I will share this with you.
http://www.beeculture.com/storycms/index.cfm?cat=Story&recordID=672


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## j.kuder (Dec 5, 2010)

sqkcrk said:


> So all of the offspring drones from a queen are genetically different from each other? Their sperm is different from each other? But equal? What's the equal part? Same number of chromosomes? I probably don't know enough about genetics to be asking these questions or understanding the answers.


 me either and i don't really feel like becoming a geneticist so i can figuire this out. i was hoping some one who is an expert would put out an article like Easy Genetics for the Sideliner and lay out a simple plan for small beekeepers to help improve the genetics not only in our own hives but also in our own areas.


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## Phoebee (Jan 29, 2014)

One thing to remember here is that, while the drones can have a mix of genes, those genes are from a limited source.

Assuming there is no tomfoolery to jigger the odds (my original question), a diploid bee (queen or worker) has 16 pairs of chromosomes with one in each pair from the mother, one from the father. Each drone has 16 haploid (single) chromosomes. In the absence of any mechanism for jiggering the odds, each of these 16 has an equal chance of being from the queen's paternal or maternal parent. In other words, each chromosome has only two possibilities. Overall there are 2^16 possible combinations (2^15 when you factor in that sex determination gene that must have two different copies), but for a given chromosome, only 2 possibilities. If there is a particular trait you are interested in and it is determined by a particular gene, that lies on one chromosome and you're still pretty darned limited in genetic diversity for that trait. If you have two hives and nature has provide some sort of "incest taboo" against the queen mating with her brothers, the genetic pickings are pretty slim.

So if you are hoping to breed tougher bees adapted for your local area but gentle, and honey-making machines, with a vicious dislike for varroa mites and SHB, you still need more than just a one other hive. If you have the perfect bees and don't want more genes, maybe you want to limit outside interaction. But breed some of those for us, please.


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## j.kuder (Dec 5, 2010)

ay yi yi yi yi. lets say 30 hives the way it sounds it is a waste of time for a small beekeeper to bother with selection because there are to many possible combinations that can effect outcome


Phoebee said:


> One thing to remember here is that, while the drones can have a mix of genes, those genes are from a limited source.
> 
> Assuming there is no tomfoolery to jigger the odds (my original question), a diploid bee (queen or worker) has 16 pairs of chromosomes with one in each pair from the mother, one from the father. Each drone has 16 haploid (single) chromosomes. In the absence of any mechanism for jiggering the odds, each of these 16 has an equal chance of being from the queen's paternal or maternal parent. In other words, each chromosome has only two possibilities. Overall there are 2^16 possible combinations (2^15 when you factor in that sex determination gene that must have two different copies), but for a given chromosome, only 2 possibilities. If there is a particular trait you are interested in and it is determined by a particular gene, that lies on one chromosome and you're still pretty darned limited in genetic diversity for that trait. If you have two hives and nature has provide some sort of "incest taboo" against the queen mating with her brothers, the genetic pickings are pretty slim.
> 
> So if you are hoping to breed tougher bees adapted for your local area but gentle, and honey-making machines, with a vicious dislike for varroa mites and SHB, you still need more than just a one other hive. If you have the perfect bees and don't want more genes, maybe you want to limit outside interaction. But breed some of those for us, please.


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## Juhani Lunden (Oct 3, 2013)

Phoebee said:


> Each drone has 16 haploid (single) chromosomes. In the absence of any mechanism for jiggering the odds, each of these 16 has an equal chance of being from the queen's paternal or maternal parent. In other words, each chromosome has only two possibilities. .


But but, crossing over is taking place when the egg (which is to become a drone) is made. Queen is shuffling the deck between her maternal and paternal kromosomes. Correct me if I´m wrong.


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## norton (Mar 19, 2005)

Yes I agree.


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## Juhani Lunden (Oct 3, 2013)

Norton, is that YOU, whom I have visited? (it is not allowed to call a person with a name unless he/she allows, that is the rule of the forum, I was told)


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## norton (Mar 19, 2005)

Yes.


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## Juhani Lunden (Oct 3, 2013)

What about the drone genetics (line) in Dartmore, is it in use this year?


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## Phoebee (Jan 29, 2014)

Juhani Lunden said:


> But but, crossing over is taking place when the egg (which is to become a drone) is made. Queen is shuffling the deck between her maternal and paternal kromosomes. Correct me if I´m wrong.


More like she's flipping 16 coins. But you need one side of each coin to make a drone, and there are only two sides available. So let's say you're trying to develop a line of TF bees and want varroa resistance. If you've got two hives, there are four versions of each chromosome in play. You can combine all you want, but if what you need is a particular variation gene that is not in the mix, you can shuffle all you want and never get lucky because the card (or coin) is not there.

This is why it is futile for a little guy like me to attempt breeding treatment free bees *from scratch *from 2-3 hives, especially from queens coming from one breeder in California. But Michael Bush keeps more like 200, and that may be enough. But that does not mean the little guy can't find queens from folks who HAVE found the lucky coin and gotten it in their mix. I've gotten suckered into a small group trying to breed queens from our best colonies. I'm being given a nuc built on a VSH queen, the daughter of a very successful queen from a Virginia breeder, bred with an uncontrolled population of mutt drones (including some Russian ancestry) in an apiary of about 13 hives. Anything could result from this mix, maybe even good. She stands a fair chance of having a couple of really special genes, and her daughters will have a mix of daddies. 

The group also is raising a few queens who trace their ancestry back to Michael Palmer's beeyard.

Whatever comes of this nuc, I'm satisfied it can't be worse than the California Girl who just failed and was superceded after less than two months laying. And she was my GOOD queen. I don't think I want to waste a lot of time on her genes.


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## BEES4U (Oct 10, 2007)

Re:
Do bee eggs favor the maternal contribution?
The following data explains why you can see different phenotypic drones and the queens daughters.

Drone genetics

See also: Haplodiploid sex-determination system


Question book-new.svg 


Drones carry only one type of allele because they are haploid (containing only one set of chromosomes from the mother); thus, they are also called hemizygous. During the queen's egg developing process, a diploid cell with 32 chromosomes divides to generate haploid cells called gametes with 16 chromosomes. The result is a haploid egg, with chromosomes having a new combination of alleles at the various loci. This process is also called arrhenotokous parthenogenesis or simply arrhenotoky. There is much debate in the scientific literature about the dynamics and apparent benefit of the combined forms of reproduction in honey bees and other social insects. The drones have two reproductive functions. They convert and extend the queen's single unfertilized egg into about 10 million genetically identical male sperm cells. Secondly, they serve as a vehicle to mate with a new queen to fertilize her eggs. Female worker bees develop from fertilized eggs and are diploid in origin, which means that the sperm from a father provides a second set of 16 chromosomes for a total of 32 - one set from each parent. Since all the sperm cells produced by a particular drone are genetically identical, sisters are more closely related than full sisters of other animals where the sperm is not genetically identical. A laying worker bee will exclusively produce unfertilized eggs, which develop into drones. As an exception to this rule, laying worker bees in some sub-species of honey bees may also produce diploid (and therefore female) fertile offspring in a process called thelytoky. In thelytoky the second set of chromosomes comes not from sperm, but from one of the three polar bodies during anaphase II of meiosis. In honey bees, the genetics of offspring can best be controlled by artificially inseminating a queen with drones collected from a single hive, where the drones' mother is known. In the natural mating process, a queen mates with multiple drones, which may not come from the same hive. Therefore, in the natural mating process, batches of female offspring will have fathers of different genetic origin. Source: 
Harbo, John R.; Rinderer, Thomas E. Breeding and Genetics of Honey Bees USDA Mar 1993 
Page, Robert E. Jr.; Gadaub, Jürgen; Beye, Martin The Emergence of Hymenopteran Genetics Genetics (journal), Vol. 160, 375-379, Feb 2002


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## Richard Cryberg (May 24, 2013)

Phoebee said:


> More like she's flipping 16 coins.
> 
> 
> She is certainly NOT flipping 16 coins. You are not accounting for crossovers. She is flipping more like 10s or 100s of thousands of coins. She has 16 pairs of chromosomes or 32 chromosomes total. But, before the egg is formed each of those pairs of chromosomes swap parts in what is called crossovers. Each chromosome swaps several parts with its mate. Crossovers happen at semi random places along the length of the chromosome. The net result is a single queen can potentially make millions of drones, no two of which are the same genetically. After crossovers happen talking about a chromosome being from the queens mother or father is simply nonsense. The chromosomes are each a mix of the genetics the queen got from her mother and father.
> ...


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## peterloringborst (Jan 19, 2010)

I think folks are overlooking two important factors. 1) a colony is a population, not an individual though it behaves as an individual. 2) nobody has a handle on the genetic basis of bee behavior. 



> As we all know, Darwin stated that evolution results from survival of the fittest individual, the fittest having been determined by competition. There is much truth in this concept, but at the same time, emphasis on the individual is an unwarranted limitation. There is also competition between products of an individual, between populations, and between species, and all these competitions may lead to the selection of survivors.
> 
> Populations are not merely groups of individuals. The fittest population is characterized by diversity of its members and by collaboration between them. The strong population will have leaders and followers. Human populations may have engineers, inventors, soldiers, and farmers, who all are not necessarily the fittest individuals.


Werner Kalow, 2001. Pharmacogenetics in Perspective



> We already know nearly the entire DNA sequence of the Apis genome and most of
> its neural proteome, but the predicted behaviors of the organism do not emerge
> from this knowledge. The trajectories from genotype to complex phenotypes are
> indirect, multi-level and virtually unknown. Most types of behaviors depend on
> ...


Ryszard Maleszka, 2012 Elucidating the Path from Genotype to Behavior in Honey Bees: Insights from Epigenomics


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