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

[MethowKraig]

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

[kilocharlie]

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.

[MethowKraig]

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.

[MethowKraig]

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

[cerezha]

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?

[kilocharlie]

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.

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

[kilocharlie]

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.

[MethowKraig]

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

[kilocharlie]

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.

[kilocharlie]

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!

[MethowKraig]

Charlie,
Thanks for researching this.
-Kraig

[kilocharlie]

*** 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.

[kilocharlie]

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.

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?...

Kraig

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.

[MethowKraig]

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

[kilocharlie]

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