# Does colony have a sub-family preference in raising queens?



## MethowKraig (Aug 21, 2011)

I have been puzzling over whether colonies tend to favor larvae from one or more sub-families to raise queens from.

Assume ten drone mates, we have ten sub-families. If one has a distinct genetic resistance to an outside threat such as disease or parasite, it will have a higher survival rate. Thus, over time, its percentage of the worker population will increase, say to 20%.

But the queen still lays eggs from that family at a 10% rate. Is there any evidence or studies that shows the bees have any feedback method that increases the odds of the successful sub-family in raising queens?

Note that from my experience rearing queens, I would say no. They will accept larvae from any source. BUT most queen rearing is started under the emergency impulse, which is actually rare in nature. In a crisis, they may not be selective, and accept whatever larvae might save the hive.

Given the luxury of time, as in swarming or supercedure, perhaps they are more selective.

It is beyond the scope of a simple post, but this would help explain to me why drone laying is suppressed in a queenright hive. Darwin was extremely troubled by haplodiploidy and I am not convinced by Hamilton’s explanation.


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## indypartridge (Nov 18, 2004)

From Dr. John Connor's _Bee Sex Essentials_, "efforts to show that super-sisters favor queens from their sub-family over half-sister queens have failed to show there is a clear and significant bias." However, he also notes research which has shown that bees from the same sub-family can recognize each other, and that division of labor within the hive often corresponds to these sub-families (patrilines).


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## MethowKraig (Aug 21, 2011)

Any actual scientific studies?


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## kilocharlie (Dec 27, 2010)

MethowKraig - I'll shoot an email off to Doc (Dr. Lawrence John Connor) and ask about his bibliography for a starter reading list. My intuition says some patrilines will be more "popular" than others - and I'll bet the tendency starts early, favoring certain grubs when food is scarce, especially during supercedure response, etc., but I'd rather read a study and repeat an experiment, or design a new one and let the bees tell me the answer, any time!

I may send a few emails around the universities and Randy Oliver and others and see what studies are out there. Good question, good post! This is right at the core of rearing quality queens, especially for those of us who are running colony trait tests, mixing and concentrating semen from desirable drones, and getting into I.I. Thank you for starting this one!

Indypartridge - perhaps you could post a list from the bibliography of LJC's Bee Sex Essentials...?


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## MethowKraig (Aug 21, 2011)

Here is the basic conundrum:

Given a stable population of colonies (pre-human influence), a single queen will, on average, produce one successful daughter queen.

For ease of the math, assume an average of ten drones per typical colony, thus ten sub-families each colony.

First problem: 90% of genetic material contributed by drones is "dead-ended" at the colony because 9 out of ten sub-families do not produce a queen.

Also, note that each queen will, on average, have ten drone sons that contribute genetic material to other colonies. But only 10% of those drones will actually be lucky enough to have a daughter queen.

Second problem: Without going into the math of Hamilton's "relatedness index," if you multiply relatedness by the 10% success rate, the result is that drone laying workers would be a far more effective way of passing on genetic material. This method would also increase genetic diversity dramatically because the male side of the genome would be passed on. It would also be directly influenced by the sub-familiy's success rate. The more workers the more drones.

Third problem: Drone laying workers are the hive's last resort effort to pass on its genetics. They have tried supercedure. They have tried emergency rearing. The only thing left is drone laying workers. Given that the math says otherwise, there must be some extreme danger in this method of reproduction. What is the danger?

In summary, only when genetic material gets into the queen is it passed on in the lineage. Having a good drone mate or two and thus a successful colony is dead ended most of the time.

UNLESS, as I noted in my start of this thread, there is some sort of feed back mechanism to increase the success rate of queen rearing for the best sub-families. 

If the workers do not exhibit a preference, perhaps it is the queen herself. The increase in the sub-family pheromone, resulting from more viable workers surviving, could influence her directly.


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## cerezha (Oct 11, 2011)

MethowKraig said:


> .... But only 10% of those drones will actually be lucky enough to have a daughter queen.


 From evolution point of view, it is very high success rate!



MethowKraig said:


> ...In summary, only when genetic material gets into the queen is it passed on in the lineage.


 I am not sure I understood this. In my understanding, the drones determines the lineage because they are haploid and thus, "pure line". In queen, since she is diploid, Mendel's law would work efficiently "diluting" the lineage: only 25% will keep lineage in each generation (1/4 of 1/4 in 2nd and so on). Because many drones inseminate the queen, it is really difficult to keep lineage via queen. In this sense, drones are "lineage" keepers. 

Very interesting topic. I wish to know more on this subject. Selection may be at the queen and/or nurse-bees level. It is easy to imagine that nurse-bees could have some preferences who to choose to rise. From another hand, at normal circumstances, it is a queen who lays the egg into queen cell. It is known (?) that queen may choose the semen to be used - particularly from africanized drones. I do not know if queen could segregate semen from all drones participated in insemination or most of the semen is mixed? 

Sergey


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## MethowKraig (Aug 21, 2011)

MethowKraig said:


> Darwin was extremely troubled by haplodiploidy and I am not convinced by Hamilton’s explanation.


I used the wrong word. Should be, "Darwin was troubled by *eusocial* behavior." Eusocial behavior means some of the females give up reproducing to help the queen.

However, it is interesting to note that Bees belong to the Hymenoptora Order, along with ants and wasps. The entire order reproduces by haplodiploidy (haploid males). There seems to be a link between haploidiploidy and social behavior. Almost all social insects are in the Hymenoptera Order. The one big exception is termites.


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## MethowKraig (Aug 21, 2011)

Sergey,

The queen is inseminated by many drones and thus her *colony* is influenced by many drones. However, only sub-family lines that actually produce a queen extend their drone's influence. If only one or two sub-families produce a queen, the other drone lines are at a dead end in the colony.

"Met-How" Kraig


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## cerezha (Oct 11, 2011)

MethowKraig said:


> ... If only one or two sub-families produce a queen, the other drone lines are at a dead end in the colony.


 Agree. I think my wording was not good - I meant that two "pure" lineages (mother/father) presented only in the drones the queen produced- each drone in the hive inherited ether mother or father haploid set of genes. Each drones from worker bees (assuming random semen) would have one out of many genotype presented in the inseminated queen (plus mother's set). I agree that only one drone preserved the lineage via the queen, but it is happened in any diploid organisms. 
The subject of this thread is much more interesting if bees could recognize and possibly select the beneficial (?) genotype (or phenotype?).
Sergey


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## kilocharlie (Dec 27, 2010)

I'm in the beginning of trying to design an/some experiment(s). Using the centrifuge, homogenize the semen of two drones at a time, one having a visible trait such as Cordovan or Chartreuse Eyes, and the other not having the chosen visible trait. Instrumentally inseminate one group of queens with the visible trait and one group of queens without. Put the resulting queens in observer hives and keep quantitative records of resulting bees and daughter queens. Watch for and record as many behaviors related to patrilines / sub-families as you can think of.

Suggestions are welcome.


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## kilocharlie (Dec 27, 2010)

Sergey - a point of interest regarding if the semen is mixed. Instrumental insemination can utilize "homogenized" semen from multiple drones by putting the various semen samples into a centrifuge and spinning them all together, or semen directly collected from drones, unmixed. I doubt that nature mixes it as thoroughly as 20 G's in a centrifuge, but the question is a good one.

I do remember reading a study that said that several queens that had been instrumentally inseminated were observed going out on a mating flight anyways (after I.I.), and that many did not. Your question makes me wonder if laboratory mixing being different than "Mother Nature mixing" (and perhaps number of drone matings) has anything to do with an I.I. queen's behavior as contrasted to a naturally mated queen's behavior.

Either way, if Mama Queen has some kind of control over which husband's sperm she's laying, that would seem a surprise - but it is not out of the realm of possibility.


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## MethowKraig (Aug 21, 2011)

MethowKraig said:


> Third problem: Drone laying workers are the hive's last resort effort to pass on its genetics. They have tried supercedure. They have tried emergency rearing. The only thing left is drone laying workers. Given that the math says otherwise, there must be some extreme danger in this method of reproduction. What is the danger?


Let me propose a hypothesis for the danger in drone laying workers. I will call it the "arms race restriction."

If one sub family has 1% of its workers laying drones, the incentive will be for the second to have 2%, etc. Very quickly, within a dozen or so generations, the drone population will be dominated by one or two drone lines. This will feed back as a reduction in sex alleles and brood failure due to diploid drones.

The same would be true if a sub family is preferential to raising its own queens.

*The success of the colony would seem to depend on the complete suppression of any competition between families.* After all, by its very nature, competition would tear apart the very cooperation that makes a beehive successful.

*However, this still leaves open the concept of the queen having an influence.* Sergey noted there is evidence of queens being preferential to africanized drone semen. Perhaps queens have been influencing survival all along but it was just never noticed.

Maybe she really is a "Queen" and not just an egg laying machine!


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## kilocharlie (Dec 27, 2010)

MethowKraig said:


> If one sub family has 1% of its workers laying drones, the incentive will be for the second to have 2%, etc. Very quickly, within a dozen or so generations, the drone population will be dominated by one or two drone lines. This will feed back as a reduction in sex alleles and brood failure due to diploid drones.
> 
> The same would be true if a sub family is preferential to raising its own queens.


Usually only 1 to 2 workers become laying workers. I wonder if 2% of a 60,000 bee colony (=1,200 Laying Workers) would tolerate each other laying eggs in the same colony? Would they fight? Wouldn't their pheromones suppress other workers from becoming LW's? I think even 1% is too high a number.

I think that the greater likelihood would be workers from a patriline preferring it's own brood, if they can indeed tell the difference. In the supercedure and swarming responses, this would make sense, because the workers choose to supercede a queen, even to the point of killing Mama once the VQ has mated and begun laying. Swarming response is likely a group decision - workers and possibly the queen, too, involving several factors. Swarming is probably a goal of a colony. 

In the emergency response, they are more likely to choose *any* healthy grub the right age, or closest to it, and tend to try to make many queens at first, only continuing feeding the one or two that are best suited to becoming a queen after the second day. Emergency queens are often superceded quickly anyways, likely because they were fed worker jelly for a day or two before being switched back to royal jelly in the queenless emergency event, and thus became an intercaste queen/worker. This would tend to reduce the number of ovarioles in a "queen's" ovaries, causing her to lay fewer eggs over her lifetime, and possibly her rate per day.


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## MethowKraig (Aug 21, 2011)

Charlie, 

I see your point. 

I guess I am puzzling why drone laying workers are suppressed at all in terms of evolution since it would seem to be a better way to pass on genetics, at least by the math. 

But if one family started producing lw's, couldn't it set off an arms race, where the result after a few generations would be one or two families dominating and reducing the number of sex alleles.

In other words, competition between families, in and of itself, would defeat the integrity of the colony. This would be true even if the competition was in the form of favoring their own larvae for starting a queen cell.

Competition between families thus might be very dangerous and the honeybee has evolved to suppress any urge that would threaten the cooperation between families. Sort of a bargain with the devil, "let's agree not to fight so we are stronger as a group." I realize that is an anthropomorphism.

The queen on the other hand, is sort of impartial. She might be responding to the pheromones in the colony. If one family is more successful, their numbers would be higher and she could then favor their sperm in producing queens. 

Researchers have apparently looked to see if workers favor queen larvae from their own family and not found any evidence. 

Maybe they just have never looked at the queen because it seems so impossible to imagine she can select sperm.

I'm not sure how you could design an experiment. If queens are selecting sperm, it might be hard to duplicate with I.I. because perhaps the sperm inside of her is actually in globules of mucus that keep the different lines separate. There is no way an instrument can duplicate what happens in flight.

I'm thinking this is a topic for an advanced lab to investigate. With genetic testing, I'm sure they would figure out a way.

-Met-How Kraig

ps where is Ojai, CA


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## kilocharlie (Dec 27, 2010)

MHK - I believe a healthy queen can lay more drones to pass male-passed traits along than LW's can, as the LW's time window before the hive dies out is about 44 days at most, minus 24 days to raise a drone, minus how many days it takes him to mature to mating ability. Also, a laying queen will lay plenty of worker grubs who will become nurse bees a few days later to help raise them and later feed them. These workers seem to know that drones are a luxury, and without a queen and no hope of making one, they are a last resort. I would probably wager that is why LW's laying drones is suppressed until the very end.

I doubt the math favors LW's over a healthy laying queen in evolution - the LW drones are often underfed and probably have a lower % chance of sucessfully mating and producing a colony that survives if conditions are less than ideal when the queenless / no viable grubs event occurred. Too late in the season - they'll lose a robbing fight and die off, etc. A queen that lives 4 years and makes one swarm plus a batch of healthy drones a year has done her tribe proud in terms of evolution.


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## kilocharlie (Dec 27, 2010)

MethowKraig said:


> Assume ten drone mates, we have ten sub-families. If one has a distinct genetic resistance to an outside threat such as disease or parasite, it will have a higher survival rate. Thus, over time, its percentage of the worker population will increase, say to 20%.


I think this intra-hive evolutionary force of sub-family attrition through natural selection due to stresses will have a lot to do with who gets to become queen over an acceptible evolutionary time period.


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## MethowKraig (Aug 21, 2011)

Charlie,

I think this thread got sidetracked on the LW issue. I agree with how you describe LW's as they are now. I was trying to understand why honeybees evolved that way.

But going back to the original topic. Given a hive with an exceptionally healthy sub family. In my example, growing from 10% to to 20% of the population through attrition of the other bees. It seems like there must be some mechanism for the hive to take advantage of this.

I can think of only two possibilities. Either could be controlled by pheromones.

1. Nurse bees influence the outcome of queen larvae by favoring the thriving family when raising queens.
2. The queen influences the outcome by favoring the family's sperm when laying in a queen cell.

Sounds like the first may have been studied but not the second.

I wish some experts would weigh in on this thread. Do you have any contacts with better minds than ours?

Thanks for your thoughts. Always enjoy your posts.

Kraig


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## kilocharlie (Dec 27, 2010)

Dr. Larry Connor emailed me back, saying that there were conflicting results. He gave a brief list of study leads from the entomologists at UC Davis, Berkley, North Carolina State, and some names to look for - Harry Laidlaw, Robert Page, Radniks, others, David Tarpy, Otto Mackenson - and suggested that I ask Dr. Tom Rinderer at Baton Rouge Bee Lab. I have not yet heard back from Dr. Rinderer. It could be a while searching these studies without titles, but the topic is interesting.

My guess would be that hypothesis #1 is indeed worthy of investigating if it has not been done already, and that possibly in hypothesis #2 that Daddy Drone may have flavored his own sperm with whatever nectar / pollen beer his working ladies fed him, along with his mama's gene mix.

And Ojai, CA is in Ventura County (Just "North" err, West, really of LA) about 12 miles inland from the city of Ventura. I'm a short ways out in the sticks from Ojai.


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## kilocharlie (Dec 27, 2010)

MethowKraig said:


> ....Darwin was extremely troubled by haplodiploidy and I am not convinced by Hamilton’s explanation.


Kraig - I understand you backed off from the haplodiploidy wording for eusocial behavior, but could you expound a bit further regarding Darwin and Hamilton? Again, this is interesting stuff.


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## MethowKraig (Aug 21, 2011)

Go to:

http://en.wikipedia.org/wiki/Evolution_of_eusociality

The basis of Hamilton's thesis for the evolution of eusocial behavior is that a worker gives up her ability to reproduce because her own offspring would be related by 1/2 but helping her mother queen produces super-sisters that are related by 3/4.

Where Hamilton breaks down for me is she is also cooperating in raising half-sisters and drone brothers who are related only by 1/4. 

This is noted in the above reference under: *Current Therories: Haplodiploidy/Kin selection* in the statement, "Other problems with this theory are when there are multiple males breeding with the queen then siblings are less related."

So, that is why I am wondering why workers wouldn't have evolved to raise their own drones, which would be related by 1/2. Obviously, over evolutionary time, they would have evolved to be well fed, etc. You could imagine a system where the workers laid haploid drones and the queen laid diploid workers.

There is some huge impediment to this happening. Again, I'm thinking there is some feedback mechanism, perhaps the sex allele, that stops this evolution.

See my next post;


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## MethowKraig (Aug 21, 2011)

Charlie,

The favoring of queen larvae from the successful sub-family is a separate, but related, question to why the workers don't produce drones of their own.

It seems likely to me that two forces are in play.

1. There is an extreme danger of some sort in drone laying workers. Probably some sort of in-breeding danger.
2. The advantage of cooperating with half sisters is in some mechanism that reinforces the genetic influence of the best sub-families.

Exactly what the danger is in #1, and the feedback mechanism in #2 is, remains to be explored by scientists.

Philosophically, I like the premise that is the queen who is pushing the sub-family advantage through the frequency of sub-family pheromones. If the workers are doing it, seems like it could set off an "arms race" within the hive that might threaten the very cooperation that is the basis of the honeybee's success.


Kraig


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## kilocharlie (Dec 27, 2010)

Kraig - The drones are genetically clones (or perhaps "1/2 clones") of their mother, not their father, having been made from *UN*-fertilized eggs, so there are no 1/4-related nor 3/4-related bees. Daddy Drone's papa does not count. The evolutionary differences must be studied comparing and contrasting to their closest relatives in the hymnoptera group that reproduce differently. My guess is that the setup has a lot to do with queens not liking each other and killing by pre-emptive strike sting before their sister queens hatch or fighting to the death if they have both hatched. The system is dependent on the lone egg-layer, and is therefore susceptible to disaster, so groups that re-queen well (fast enough, often enough, and well-timed in the season, etc.) and those that developed Laying Workers had a survival advantage.

Workers do not voluntarily "give up" their reproductivity, they are starved out of it before they are born by being fed "worker jelly", not royal jelly. Worker food is lower in sugars and proteins, requiring them to develop for 19 to 22 days, as opposed to 14 to 17 days for a queen.The ovarioles on the worker bees' ovaries are not developed - the ovaries are diminutive, and their pheromone production is minor, if even existent at all.


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## MethowKraig (Aug 21, 2011)

The relationship ratio system is used by biologists. It goes like this:

Daughter to Mother = 1/2 
Daughter to Father = 1/2
Sister to Sister (rare in bees, common in humans) =1/2
Half Sister to Half Sister = 1/4
Worker to Drone = 1/4
Super Sister to Super Sister = 3/4
Laying Worker to its own Drone = 1/2

In humans, there are no super-sisters, but you would add:
Identical Twins = 1
Son to Mother (or Father) =1/2
Son to Father = 1/2

I'm not sure about Brother to Sister in humans. Seems like it would be one chromosome less than 1/2 (The X/Y)
And, Brother to Brother and Sister to Sister; seems like it would be one chromosome more than 1/2.

The relationships are identical either way. Daughter to Mother = 1/2 = Mother to Daughter.

As I understand Darwin, it is a tenant that creatures will evolve to most efficiently transfer their own DNA genetics to the next generation. Eusocial behavior seems to challenge that tenant because workers give up their ability to reproduce.

Hamilton noted that by helping their mother raise super-sisters, sterile workers are helping the queen pass on 3/4 of their genome, instead of 1/2 by raising their own daughters.


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## MethowKraig (Aug 21, 2011)

Charlie,
I hadn't thought about the process of queen's killing other queen cells. I suppose that is another situation where the sub family influence could affect survival rates of a better sub-family in some way. Not sure how.
Kraig


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## cerezha (Oct 11, 2011)

MethowKraig said:


> The relationship ratio system is used by biologists. It goes like this:
> 
> Daughter to Mother = 1/2
> Daughter to Father = 1/2...


 I do not understand how daughter could pass 1/2 genome to the mother or father? Again,are you talking about diploid organisms or bees, who are diploid and haploid?


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## kilocharlie (Dec 27, 2010)

Kraig- It is a temptation to fall for the fallacy to think of them all as "half-sisters" in that while it is true that half of their genetics are from the same source, it is a matter of "Which half?" 

While any two _may_ be indeed identical, usually no two _are_ identical, it is highly unlikely because each sperm is usually different from the other, as is each egg. But a trait that is not present in one parent is extremely unlikely (~nearly impossible, except for a mutation) to show up in an offspring. 

Some traits are of a "yes/no" or qualitative nature, where other traits are of a "how much?" or quantitave nature, still others are complex groups of traits, and some are recessive and some are dominant, and perhaps some are conditional. The fraction system you mention is a "good enough" gloss-over for discussion sake, but one must keep in mind these "caveats" of the truth about what is actually happening. 

The original question is valid, and if there is an answer that leans on the "yes" side, there likely is a mechanism - such as the probable link of octopamine to the modulation of bees' assessment of resources such as pollen and nectar (BTW, octopamine would be a good thread - lots of different people are starting to look at it to try lots of things for lots of different reasons, and it could have hellish repercussions on bees!)

*I'd strongly encourage all who take interest in these discussions to email off to the universities asking for research and (gasp) READ THE STUDIES. I'd further encourage people to coordinate with science teachers in public schools to replicate the experiments, and encourage students to question if the results are linked to the conclusions, and if so, how? I'd even encourage sponsoring related experiment design competitions.*

Darwin's mistake is that workers do not "give up" their reproductive ability, they were starved out of it before they grew up, as "betas, gammas, deltas, epsilons" in (I think it was) Huxley's Brave New World were cloned to various working classes, and their development of their ovaries are suppressed by Mama Queen's substances such as pheromones.

It's funny to see Hamilton use that system, thinking that super-sisters are 3/4 related...they are full sisters in human terms! Half sisters are 1/2 sisters (only same mama). The difference is two "___sisters" with same mamma but two unrelated paternal grandmothers vs. two "___sisters" with two different drone fathers that were brothers born to the same drone mother vs. two "___sisters" with the same daddy and mommy. How you fill in the blanks is a matter of nomenclature. There is also the issue of was either mom or dad inbred, or sisters of an out-crossed vs. an inbred maternal vs. paternal grandmother?, or even multiply inbred and how many generations back and on which sides of the family? It can get complicated...but I'd bet they have a clue if they don't know outright, and I'd bet it doesn't take them very long to know.

One thing is likely - they evloved that way because an ancestor mutated into a form that gave rise to the present generation, and the succeeding generations survived in the environments of their day. Creationists, please don't burn me at the stake, stone me, crucify me, nor apply any other religious purification rite, as you could go to the theological place of eternal punishment for your actions.:no:

Sergey - I believe the order may be reversed. I doubt we can pass anything but headaches and bills along to our parents!


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## kilocharlie (Dec 27, 2010)

Regarding your post #5, third problem, I'd agree that emergency queen (an intergrade worker-queen due to food change, rendering a less-than-ideal queen) then supercede her soon afterward with a premium daughter queen is a more favorable result than laying workers.


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## MethowKraig (Aug 21, 2011)

Charlie,

The sperm that a drone produces is all the same, a clone of his haploid genome. Millions of them, all identical. Absolutely no variation.

The queen produces a different egg, genetically, every time.

The result is two super-sisters, coming from the same drone, share an identical genome on the male side. The side that comes from the mother is not the same. But because each egg came from the same mom and inherited half of her genome, they have genes in common half the time. The 1/2 relationship on the mother's side of the worker's genome is an average. Some will be more related, some less. But the average is 50%. The father's side is a clone of the drone father and is the same 100% every time. 

So, in comparing super-sisters, the drone half is always identical, the mother side averages 50%. The result is called 3/4.

A true sister relationship in the hive would occur if the queen had two drone mates that originated from the same queen. In that case, the male side is different but from the same source(the drone mother). The female side is different but from the same source (the hive's queen). So in comparing sisters, each half has genes in common 50% of the time. True sisters are rare in a hive, but do occur. They are related by 1/2.

Two workers are half sisters when they have drone fathers from different sources. In that case, the fathers side of the genome is unrelated and the mother's side averages 50%. So, half sisters are related by 1/4.

A worker to any drone in the hive is related by 1/4. The worker's male side is totally foreign to the hive's drone. The queen side of the worker's genome will have genes in common with the drone 50% of the time on average.

A queen and any son or daughter are related by 1/2 because the offspring only has half of her genome.

Finally, the relationship fraction is about the *source* of the genes. We share 25% of our genes with a banana. We are not related by 1/4.

"Met-How" Kraig


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## kilocharlie (Dec 27, 2010)

I understand and agree, except that even "identical twins" are different. They grow from a single sperm/egg combination that splits, and are supposedly "identical", having the same source of DNA, but nature is not perfect, not every time. There is almost always a difference, almost to the point that there is "no such thing" as identical. I'm not saying that they cannot be identical, as that single possibility is a subset of the total. 

Think about it...that DNA double helix is one big old molecule...hard to copy it _exactly_ right every time, but admittedly, the smaller the dna the higher the probability of it happening. We have genetic mutations in us all (probably), somewhere on that giant double helix. It would be better to speak of "similarity" than "identicality" (would "congruence" would be a better word?), where relatives share a "high degree of similarity".

I'm curious how the fraction system represents an offspring of inbred bees...queens / workers, and drones? Using eighths for an inbred parent? Sixteenths, thirty-seconds, etc., as the generations go back? What about multiple-inbred lines? And how does it account for the mutations? I know mutations do not make you NOT a brother or sister of your sibling, I'm just reminding everyone that "1/2 genetically related" is approximate, not absolute (a machinist will make the joke +/- .005" Hahahaha). Instrumental insemination allows breeders to use techniques such as back-crossing and inbreeding to create lines that strongly promote a single (or a small group of) trait(s). This often produces bees that are somehow weakend, disease-prone, or suffer some other malady, but successive generations of out-crossing may create vigorous bees that still display that one, strong, very favorable trait.

Again, the fraction system is for theoretical discussions, usually an (admittedly VERY close) approximation. It would be very useful to continue the system down to smaller fractions for the purposes of tracking inbred lines and their progeny.


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## kilocharlie (Dec 27, 2010)

Dr. Tom Rinderer replied and said that bees DO NOT SHOW ANY PREFERENCE in patrilines when making queens. One study showed a bias, but was re-evaluated and found to show no bias. 

Thank you, Dr. Rinderer!


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## MethowKraig (Aug 21, 2011)

Charlie,
Thanks for researching this.
-Kraig


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## kilocharlie (Dec 27, 2010)

*** HOT NEWS FLASH ***
Dr. Tom Rinderer just sent an update. The inquiry got him curious, he re-visited the data last week, and noticed that *some* colonies *do* tend to show a patriline preference sometimes. He called it a "qualified yes", reversing his position. He was thankful it was brought up, he will likely study it again, soon.

I replied to him how much I love an interesting answer! I hope to follow his study, and will probably go ahead and look up the other papers published elsewhere, mentioned by Dr. Larry Connor.

The thanks are not deserved, Kraig - I did not research anything, merely shot off some emails. I will, however, begin to research soon. I need help from a seasoned researcher, though, regarding the question, "How does one find *ALL* the previously-published papers?". To study a subject incompletely is to skew any conclusion. Anyhow, the thanks go to Dr. Connor for a list of suggested studies and for recommending my asking Dr. Rinderer because it is "better to get a view from inside the research than from outside looking in", and very especially to Dr. Rinderer for taking a second look at what he called "muddled data" and Larry called "conflicting data". This could get very interesting.


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## kilocharlie (Dec 27, 2010)

MethowKraig said:


> Charlie,
> 
> ...But going back to the original topic. Given a hive with an exceptionally healthy sub family. In my example, growing from 10% to to 20% of the population through attrition of the other bees. It seems like there must be some mechanism for the hive to take advantage of this.
> 
> ...


regarding this in post #17...let us exaggerate your percentages to more clearly illustrate what probably happens most of the time.

Suppose one patriline (sub-family with the same daddy) has a far superior genetic advantage, say 66.7% survival rate against some very real stressor pathogen, while nine other patrilines bite the dust in great numbers, say 99% death rate. Now the question of "Who gets all the royal jelly?" is a mathematical no-brainer. We expect the patriline with the superior tolerance will be around 66.6 times more likely to count a successful new queen from their numbers than any other patriline, especially in an emergency response. Statistics strongly and consistently going against this tide would suggest a tendency toward favoritism of some kind (probably while dooming the hive) and would most certainly perk up the numeric sense of any good scientist.

That nurse bees can recognize super-sisters is very likely. That they spend even slightly more time attending super-sister larvae is somewhat probable, and therefore the more likely to be researched.

The queen somehow "knowing" whose sperm is passing while fertilizing is probably less likely, some researchers might consider the idea to be preposterous, so it is less likely to be researched. It remains a possibility until someone disproves it.


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## MethowKraig (Aug 21, 2011)

Charlie,

Not sure how to send a secure email on this site. If you know, email me. Otherwise, I will post my email.

I'd like to contact Dr. Rinderer with some of my puzzling thoughts.

This could be ground breaking. If there is some way the hive influences the success rate of a patrilineal line, then in theory, the breeder could use that to his(her) advantage. 

-Kraig


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## kilocharlie (Dec 27, 2010)

Along the top of the page, look for *Notifications*. Click on it and send a private message to me. Doctor Rinderer took a month to get back to me. He is busy. If he and I continue to communicate, I'll suggest he read this thread, which he likely will. I'd bet you'd get better responses from him if we use this "protocol" (for lack of a better term). Feel free to private message ("PM") me. - Casey


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