# Honeybee Genetics and Breeding Bees



## Stephenpbird

Thank you I enjoyed that video.


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## jbraun

This presentation helped me understand the feedback that I received about starting to breed for queens. It also explained why Rusty was having his problems with VSH in Florida.

Thanks Ray


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## RayMarler

I was just watching this video and I have a disagreement with a statement that Dean makes at the 5:35 time mark. He stated that "Queens and workers are genetically identical". I have a problem with that statement. The workers are made up of half of the queen's 32 genes and all of their particular father drone's 16 genes, giving each worker a total of 32 genes, and not every worker has the same drone father, and no workers have the same drone father as what their queen mother had (although they do all have the same drone grandfather). Therefore, I think that the Queen is not genetically identical to her worker daughters. Does anyone out there have any thoughts, insights, or comments on this?


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## RayMarler

I especially liked the chart and commentary at the 6:30 time mark through the 7:55 time mark. To me, this sums up honeybee genetics in a nutshell. As Dean said, this is the key to understanding honeybee genetics.


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## Rader Sidetrack

RayMarler said:


> I was just watching this video and I have a disagreement with a statement that Dean makes at the 5:35 time mark. He stated that "Queens and workers are genetically identical". I have a problem with that statement.


Lets look at that statement from a slightly different perspective. 

The virgin queen was just another run-of-the-mill worker larva before that larva was selected to be fed additional royal jelly (to enable that larva to develop into a virgin queen). Since she got all her genes as just a run-of-the-mill larva, it seems as though there can be a reasonable basis for Dean's assertion. The genes that 'that' virgin has are no different than those of a bunch of workers that were raised at the same time as that virgin was.


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## RayMarler

I agree Graham, in a generalized way, yes they are similar, but to say they are identical seems a bit of a stretch. That virgin you mention, does not have the same drone father genes that say 80 to 95 percent or so of her sisters that are raised to be worker bees. They are close, yes, but not identical, in my way of thinking about it.


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## RayMarler

RayMarler said:


> I was just watching this video and I have a disagreement with a statement that Dean makes at the 5:35 time mark. He stated that "Queens and workers are genetically identical". I have a problem with that statement. The workers are made up of half of the queen's 32 genes and all of their particular father drone's 16 genes, giving each worker a total of 32 genes, and not every worker has the same drone father, and no workers have the same drone father as what their queen mother had (although they do all have the same drone grandfather). Therefore, I think that the Queen is not genetically identical to her worker daughters. Does anyone out there have any thoughts, insights, or comments on this?


Someone just cleared me up on this in a private message. The focus of Dean's discussion at the time was diploid/haploid genes, and he was absolutely right of course, there is no difference between the queen and the workers, but there is a difference between the drones as compared to both the queen and the workers. Now I got it!


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## grozzie2

RayMarler said:


> I agree Graham, in a generalized way, yes they are similar, but to say they are identical seems a bit of a stretch. That virgin you mention, does not have the same drone father genes that say 80 to 95 percent or so of her sisters that are raised to be worker bees


No, but she does have the same father as the 5 to 20% of worker sisters that come from the same father. So, if you pull out the queen, and a worker sister that came from the same drone father, they will be identical in every way. The point being, there is nothing genetically different between a queen, and a worker, at the time the fertilized egg is placed in a cell. The difference happens later, due to how that particular cell is tended, compared to others of the exact same genetic makeup.


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## RayMarler

All good comments, thanks to you all for helping straighten out my way of thinking. I think I'm good now, at least for the moment!


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## JRG13

Not quite right, same drone father doesn't mean identical, have to remember recombination during meiosis when the queen is producing eggs


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## AstroBee

grozzie2 said:


> So, if you pull out the queen, and a worker sister that came from the same drone father, they will be identical in every way.


Please see the comment by JRG13. The statement above is technically not correct.


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## mike bispham

JRG13 said:


> Not quite right, same drone father doesn't mean identical, have to remember recombination during meiosis when the queen is producing eggs


Yes, every individual is unique. For bees a 50% combination of those genes held by the mother (of which the mother herself only used half) and 50% the fathers genes (of which he used all)[1]. No two individuals are the same. 

However, that doesn't mean traits can't be passed on... 

Mike (UK)

[1] I think that's right, but if its askew the point remains.... It probably means half-sisters are a bit genetically closer than i.e. human half sisters are.


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## deknow

I'm glad this has been helpful.

Remember that 2 workers that 'share a father' are fathered by sperm that is identical...all the sperm in a given drone is identical.

If you want to put this into a more familiar perspective, it is the queen that produces the drone that is the 'father'. The drones are the sperm racing to be the first (or 20th in the case of bees) to reach the egg.


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## RayMarler

Yes, thank you Dean for allowing this video to be posted publicly! 
It seems to me that the drone side of the mating is more important, or at least as important as the queen side. I think it a mistake that many queen breeders that I've run into over the years, only pay attention to the queen side of the equation. Not all breeders, but quite a few.


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## Lauri

Read this book and your perception may change on may things. Genomic Imprinting for one.

http://www.amazon.com/gp/product/0262525844/ref=oh_aui_detailpage_o09_s00?ie=UTF8&psc=1

I found it to be a most _excellent_ book, aside from the childish illustrations and slightly strange dialogue at the end of each chapter.

Quite easy to understand if you have any basic biology experience. It's made me decide to get II equipment sooner rather than later.

Drones from my Mountain line of VSH Carnie hybrids










Drone from a Pol-Line strain










Allele's passed to offspring from the drones go through _spermatogenesis_ and the results can be quite different, or have different expression than if the same allele was passed to offspring from the queen.


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## beepro

Lauri, does this book focus on the bee genetics or just genetics in general?
So do you think that II will help control the bee genetics better than the open mated queens?

I have notice that the pol-line bees have a bigger dark band at the ends compared to the regular Italian bees.
You notice that too?


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## Lauri

Genetics in general, but it touches on may aspects you will find fascinating.

II will help allow me to choose my crosses, help me get to a higher standard of my own choosing, develop more custom or more unique lines & I will then be more likely to be able to replicate with a higher rate of reliability. I'll see just what genes need to come from the drones and which are better inherited from the queens. It will take me a couple years of trials, but I am looking forward to doing the work.

I've spent many years breeding animals and some plants so I've come to trust my 'selection skills'. II will just allow me to actually implement those selections.


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## RayMarler

What unusual drones huh Lauri? 
I got drones just like that, two toned and striped, back 8 years ago or so when all season long I crossed back and forth between Carni's and ITA's. It was really something to me back then, so I imagine you have enjoyed seeing that as well. It was quite educational for me. The drones can help show the trueness of the strain of bee, at least to some extent.


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## kilocharlie

I.I. has many advantages over open mating. It allows the breeder to be highly selective of both drone and queen colonies when choosing trait combinations to begin a base bloodline of a stock, eliminating many undesired traits from the very get-go (only open-mating in an area devoid of feral bees, such as Brother Adam had at Dartmoor allows this same level of control) I.I. allows back-crossing, selfing, and many other techniques as well. 

Breeding in sufficient quantity, I.I. tends to improve a bloodline every generation, where open-mating has to wait for "luck of the draw", of which never (zero progress toward a genetic goal) is a subset of, and therefore an actual possibility for the open-mating schemes (tough not very likely).

Generally, faster progress toward a genetic goal can be made with fewer total colonies using I.I. than using more colonies using open-mating, but that does depend on the numbers used in each, and could be skewed greatly either way simply by starting one group with better stock than the other.

Also, on the topic of books - look for Dr. Ernesto Guzman-Novoa's book, Elemental Genetics and Breeding for the Honeybee, available at www.ontariobee.com/outreach/manuals-books-dvds or through Paul Kelly, University of Guelph, Honeybee Research Centre, School of Environmental Sciences, 50 Stone Road East, Guelph, Ontario, Canada, NIG 2W1.

Another good one for beginners to genetics is Introducing Genetrics by Steve Jones and Borin Van Loon, a fun cartoonish book that starts with Mendel, advances through the years and personalities involved, through Watson and Crick, and on to some more current foundations in genetics. Bees are, of course, a bit different with haplodiploidy and perhaps some thelytoky being expressed, but this little book gives a decent bit of background.


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## JRG13

Spermatogenesis is an interesting topic unto itself, but I wouldn't put much stock into it. That being said, you will find certain combinations of drone mothers and queen mothers probably out performing others but whose to say it's the effect of spermatogenesis or just simply better hybridity matching. You can look at mito types, haplotypes etc... but in the end you have to build your breeding/heterotic groups and map out the best combinations. Dean hit some great points on starting with stable lines. II would capitalize on that nicely as well, great tool for trait integration and creating know hybrids/crosses. I've spent the last two years putting together potential breeding pools, still have some work to do, but I'm excited about the ride it's about to take me.


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## deknow

Lauri said:


> Allele's passed to offspring from the drones go through _spermatogenesis_ and the results can be quite different, or have different expression than if the same allele was passed to offspring from the queen.


I'm not sure I'm understanding this statement. Spermatogenesis in the honeybee is quite a bit different than in most of nature. The process doesn't produce 4 sperm cells from a single germ cell (like it does in regular haploid systems)...it produces a single sperm cell and a single polar body. This appears well hardcoded into the honeybee, even diploid drone sperm is diploid.
The real differentiation from the original germ cell happens when the mother of the drone produces the egg...not when the drone produces the sperm (which is only during the pupal phase). Different drones from the same queen will be as different as different sperm from the same human.
There may well be something I don't understand here (or don't know about), but I'm not seeing how/why alleles passed to offspring frmo the drones going through spermatogenesis has any effect on expression..


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## deknow

I'm not a fan of II. I obviously understand its usefulness (or I understand it's obvious usefulness), but I tend to see this as a forest/trees problem, and I also see some real dangers.

Focusing on narrow lines and hard artificial selection is opposite of the bees reproductive nature. What has led to their long term success (most of which without humans), is their nature. Population level selection that holds traits in reserve for many generations due to multiple matings, massive inbreeding after die off events...massive outmating in times of plenty.

I can see II (as well as simply buying stock) as a way to begin breeding, but soon, I think that one must let things develop on a popluation level, and let some dirt into the system.

The other real danger I see (or at least imagine) is that clearly the drones are competing to mate with the queen. Eyesight, sense of smell, speed, agility, ..probably even manners, all go into deciding who the queen mates with (along with luck as well). Is this race to the queen really an unimportant step in the process? Is the inevitable result of II that the resulting drones are less competitive in the mating game when given the chance? Is that a result that leads us where we want to go?
deknow


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## Eduardo Gomes

deknow said:


> Eyesight, sense of smell, speed, agility, ..probably even manners, all go into deciding who the queen mates with (along with luck as well). Is this race to the queen really an unimportant step in the process? Is the inevitable result of II that the resulting drones are less competitive in the mating game when given the chance? Is that a result that leads us where we want to go?


Very good insights deknow, IMO.
Has anyone made ​​longitudinal experimental studies over several generations to evaluate these aspects? 
What is the impact of II in the following generations? Are we doing God's role of an uninformed and naive way?
Are we run exactly where?


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## Lauri

Once the II breeding lines are established, most of the resulting daughters will be open mated at a different location for production queens. Hybrid vigor is very important, I agree. My locations are quite remote and I should still have some decent control over the drone quality. I'm fortunate in Washington State not to be near any other apiaries and have quality gentle feral drones, all from survivor stock in higher elevation locations.

Like Jeff, I've been putting together potential breeding pools the last couple years and have some great overwintered stock to work with. Several queens now going into their third winter. I have a good stock of Glenn descendants from my original II queens purchased in 2011 and acquired feral genetics from wilderness areas near Mt. Rainier.

I also brought in some new VSH lines from VP Queens this spring (USDA based stock). Since I like my lines and want to enhance them, not change them, I bought virgins from Adam. They got mated here for an immediate cross with my current lines. They will provide me with pure drones for crossing in 2015. VP gets their USDA stock in somewhat of a raw form, then evaluates, tests the lines, selects from them and makes crosses using their experience. I will take that work and continue with it here at my place for a local, Northern flavor to the line.

But back to those coming 4 year old queens I've got cookin in the hives under the rain and snow. 
How much epigenetic influence will now be contributed from a 4 year old breeder queen? All these queens have at least one or two generations of daughters on the place, also overwintered & evaluated for long term performance under different conditions and their interesting traits. I suppose the first question would be, what conditions have they been repeadly exposed in the course of four years and how could that influence the development of their future offspring? 

A couple that come to mind right away are:

I have a LOT of colonies in one location and the bees have to work harder to bring in adequate stores. They are less likely to swarm, because they don't get a huge influx of feed all at once. I also use custom frames to influence them to store feed slightly differently. How will years of no swarming impulse effect future behavior?

With so much competition from other hives, are they flying in cooler weather? Flying longer distance? Flying earlier and later than most? How long before they develop a slightly longer tongue to take advantage of different & previously unavailable nectar sources? Is that even possible? As long as robbing is not being encouraged as a trait, a person could push that for years to see what develops. It's hard not to notice the darker bees are the ones I see in the dead of winter out foraging when temps are barely favorable & rain is not too hard.


Another thing daughters may be exposed to, is a short slight cooling of queen cells from the time I take them out of the cell builder and transfer them into the incubator. Sometimes slightly cooled again, as I place cells in mating nucs. Of course I am careful not to let them actually chill, but temps do fluctuate slightly for a short amount of time during their gestation period. I commonly have a disposable hand warmer in my pocket along with my cells to be placed. Or a warmer in an insulated lunch box along with the cells.

What effect may several generations of that slight temp variation have on future generations? I can control the hatch rate of my queens by a full day with one or two degrees of incubator variation ether way. WIth a gestation as short as 16 days, every minute, every degree - counts. 

How will my recipes for supplimental feed effect actual development? You all know I use a lot of cider vinegar, electrolytes & vitamins in my recipes.
Will the added acidity and added nutrients eventually stimulate changes? Most people know nutrition has a direct effect on the size of the creature in question. Generally, the better the nutrition, the larger the critter (Reaches full size potential). Sometimes the harsher the enviroment (Alaskan wildlife for example) the larger the critter.

So all the talk of cell size and smaller regressed bees VS larger cell size and larger bees actually misguided by the lack of consideration for nutritional supplimentation in a mono cultured environment? Are bees large in size _because _of good nutrition, able to keep mite populations in balance more naturally without problems on larger cell size such as 5.4 foundation? VS bees that are only large because they were forced onto large cell foundation, but are slightly nutritionally deprived and therefore not able to tolerate mites as well? Is cell size unjustly credited/blamed for mite tolerance, when in actuality it may have nothing to do with the size, but the nutrition factor?

One thing my bees are Not exposed to, are many of the pressures commercial bees have to endure. I have No commercial crop exposures, no migratory stresses, virtually no or low treatments. No antibiotics. Is that good or bad? Is the ability to handle those stresses inherited? My stock is healthier because of this lack of exposure, but if daughters are placed under that kind of stress, how will they respond?

Same goes for VSH traits. If my stock is secluded and not exposed to recurrent mite infestations from outside sources, will they loose the VSH expression if it not constantly being needed?
My stock is healthier because of low mite counts, but when daughters are exposed to colonies with high mite pressures, how will they respond?

My Carnie hybrids are well suited for a broodless period for several months during winter. How would they respond to a California climate?

This book talks about experiments with fruit flies when environmental exposures were involved and in some cases, the changes they were able to bring out, then establish quite quickly in the genetic memory and produce even without the continued outside influence. Not changing the DNA, but changing the way it's expressed. Exposing genes that were previously masked or turned off.

I'm no scientist, but do find it fairly understandable. You don't have to be a rocket scientist to rear queens, but there can be a LOT more to it if you choose to dig deeper, be observant, be creative. And although I consider my stock to be somewhat proven, they were only proven to the extent of my own conditions. Now, with some are scattered around the country, feedback will be helpful. IF the beekeepers that own them have enough experience to make use of their potential.
While most will be happy if they overwinter well and have good productivity & longevity, I'd like to know exactly what the pressures were in their location and how the bees ultimately responded.

I love the fact the bees can reproduce & come to sexual maturity so quicky. 
In a single season, I can get a couple generations to overwinter & evaluate long before I do any grafting in any sort of volume from the original breeder queen. With the longevity I am getting, its almost like looking into the future a couple years and still having grandma around long enough to actually use.





































Some of you guys on Beesource are highly educated & I appreciate your input. I'm not to your caliber, but still can pursue progress at my own level. I probably know just enough to make me dangerous . I'd like to look into some online genetic/biology courses in my 'spare time'. The more I learn about it, the more specific I become when it comes to selection and reliable replication. At least tip the scales from pure luck to some sort of control over results.

If anyone is running experiments, I might like to be involved with your project in my neck of the woods and give you some feed back. I've got about 200 hives (Equivelent to double deeps),run about 180 mating nucs and have enjoyed nearly 100% overwintering success the last couple years. Enough resources to devote a few to some good research, other than my own.


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## zhiv9

I haven't had a chance to watch Dean's presentation, but thought that I would pass along that I found Ernesto Guzman's Elemental Genetics and Breeding for the Honey Bee to be worthwhile introduction to honeybee genetics and breeding.

You can purchase it through the OBA here:

http://www.ontariobee.com/outreach/manuals-books-dvds


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## Lauri

zhiv9 said:


> You can purchase it through the OBA here:
> 
> http://www.ontariobee.com/outreach/manuals-books-dvds


It's sold out.


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## zhiv9

Lauri said:


> It's sold out.


Sorry about that. I picked it up at queen rearing course this summer and quite enjoyed it. Unfortunately, i don't think it is available anywhere else.


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## beepro

Yes, the bee genetics is an interesting subject indeed. I'm watching you tube vids on this subject on my
spare time. At the same time I'm doing my little selection experiments while expanding my hive numbers.
While we are thinking about this subject there are many experiments already being done 
of what we are trying to accomplish. Here is an interesting read http://dencor.ca/BCBBA/index8db2.html?page_id=17
when I did a search. I'm sure the II way will help to speed up on your queen selection process at least to eliminate the
virgin mating flight schedule so she will not get lost. I wonder if once a 2nd year queen became a drone layer will II help her to become a
worker layer again? This way I will not have to kill her because she is not flying again to mate. As in any tools use it wisely on a positive outcome
will benefit a lot.


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## SRatcliff

No such thing as a biological free lunch. Best conform to nature rather than try to force it a certain direction.


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## kilocharlie

Eduardo Gomes said:


> Very good insights deknow, IMO.
> Has anyone made ​​longitudinal experimental studies over several generations to evaluate these aspects?
> What is the impact of II in the following generations? Are we doing God's role of an uninformed and naive way?
> Are we run exactly where?


Dr. Susan Cobey did a paper summarizing many studies on I.I. vs. N.M. (Naturally Mated) queens. It is available on her website, www.honeybeeinsemination.com

I do not recall if the paper covers later generations of bees after the first. I do know that the paper has many clues to the factors making I.I. viable, enumerating many common errors in the processes. It is highly recommended reading for those considering it.

I do know that many have had bad experiences with improperly inseminated queens or queens shipped with far too much banking / transit time affecting their performance quite negatively.

Even still, the superceding queen a single generation after the I.I. queen is positively affected by her mother's genetics, especially if she (and daddy) were well-chosen for their environment, in many cases probably worth the investment of the poor-performing I.I. mother.

I agree with deknow that the competition of suitor drones is a helpful factor, as anyone familiar with evolutionary forces probably would. I do think that the single-generation "boost" that one selective mating using I.I. gives to a bloodline can be far more beneficial than the "drone skills" lost on a single drone father's contribution had that mating been natural. Another thing to remember is that queens Inseminated in the lab often go out and mate as well. Breeders have no problem with that! Any severely undesireable traits are eliminated the following generation. 

Usually, an entire colony's drones have their semen harvested - around 200 drones - the semen is allowed to mix by diffusion for 24 hours, then is concentrated in a centrifuge, where I assume it mixes further. This reduces deknow's complaint somewhat - the competition between brother drones is accounted for.

Another method is often utilized, a good number of drones from similar and/or dissimilar colonies with but a few common traits are harvested for male germplasm, so drones with some common trait, but many unlike each other, are represented in the dose administered to the queen. More drones are represented in germplasm dose than have ever been observed in nature mating a single queen! So the only part of deknow's complaint is that some of the would-be "loser" drones that wouldn't have gotten to mate naturally *do* show up in a very genetically diverse hive of the resulting I.I. queens.

This is why I design my mating program to include both I.I. and N.M queens, and why I spend a lot of time testing & observing breeder colonies, both male and female.

Perhaps the biggest advantage is the ability of the geneticist/breeder to almost ensure the expression of double-recessive traits that might be lost on naturally-mated bees. This is a highly unlikely occurrence in nature, and quite valuable to the beekeeper. Many might-fighting traits are recessive, some double-recessive.

Beepro - I hadn't thought about trying to re-inseminate an old queen, perhaps worth a try?


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## beepro

KC maybe the next time I have one I can send her to you
for this experiment, eh? But she has to be a valuable queen 1st.
If she is a breeder queen it is even more interesting to see
what is the outcome. If it is a prolific production queen then
she might be able to head the colony for another year or 2. Or 
just let her lay more good drones to saturate the entire DCA. The value
has to justify the work put in otherwise just forget it.


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## kilocharlie

i may give that a try while I'm up there at U. C. Davis under her direction. If that goes well, that would be very cool. If donig I.I. at home, that saves the price of a queen at least once, perhaps more.


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## AstroBee

kilocharlie said:


> i may give that a try while I'm up there at U. C. Davis under her direction.


I thought that she left UC Davis in 2012? Last I knew she's at Washington State. Are you saying that she's offering an II class in 2015 at UC Davis?


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## AstroBee

deknow said:


> *Focusing on narrow lines and hard artificial selection* is opposite of the bees reproductive nature.


Don't you think there's more to bee breeding using II that what is quoted above? That to me seems to do a huge disservice to breeders who are constantly evaluating stock, making tweaks, and managing diversity to achieve a broad spectrum of traits.


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## Eduardo Gomes

kilocharlie said:


> Dr. Susan Cobey did a paper summarizing many studies on I.I. vs. N.M. (Naturally Mated) queens. It is available on her website, www.honeybeeinsemination.com
> I do not recall if the paper covers later generations of bees after the first.


Thank you for referring to this article. I've been reading with some attention the article by Dr . Susan Cobey , one of the leading experts in II, and as much as I understand these studies are not longitudinal , ie, do not evaluate the effect of II over several generations. This fundamental issue continues , it seems to me , without being evaluated.

Take this opportunity to mention an interesting excerpt of Dr. Susan Cobey's article , which raises questions , not being fundamentals, deserve our best attention : "_However, beekeeping pratices designed to reduce labor, increase efficiency and provide convenience in scheduling, often provide suboptimal conditions for queen development and sperm storage in the spermatheca._" (p.405).


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## beepro

I once got a very good laying queen that turned drone layer. Could of save her
with II if I have the right equipments and know how. If Sue is doing a class at UCD o II then
I would like to join too. Any infos you can provide on her schedule, KC?
On queen selection you don't have to narrow down on a specific trait but try to broaden your
selection on multiple good traits to keep and enhance. II will shorten the breeding process within
a season or 2.


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## mike bispham

Eduardo Gomes said:


> Take this opportunity to mention an interesting excerpt of Dr. Susan Cobey's article , which raises questions , not being fundamentals, deserve our best attention : "_However, beekeeping pratices designed to reduce labor, increase efficiency and provide convenience in scheduling, often provide suboptimal conditions for queen development and sperm storage in the spermatheca._" (p.405).


Can you tell us where we can read this Eduardo?

Thanks,

Mike (UK)


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## Eduardo Gomes

Hi Mike!
Here is the link: http://www.honeybeeinsemination.com/uploads/RedIIvsNM.pdf


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## apis maximus

Reading the paragraph posted by Eduardo, out of the context, one can interpret that Cobey is somehow tilting towards the idea that II has a lot of possible shortcomings...as one can see though, in her own words, in the same paragraph, she says: "*The unfounded reputation for poor performance of IIQs has been difficult to dispel"*...."*although the scientific community has repeatedly demonstrated their equal and sometimes higher performance compared to NMQs*."

Reading the whole thing though is critical...But yes, "cutting corners"...especially when it comes to precise protocols, techniques, scheduling, etc., has its own consequences.

Back to the whole paragraph:

"The data presented in Table I overwhelmingly demonstrates that instrumental insemination is a reliable and practical technique.
_*However, beekeeping practices designed to reduce labor, increase efficiency and provide convenience in scheduling, often provide suboptimal conditions for queen development and sperm storage in the spermatheca*_. The reported lower performance levels of IIQs can probably be attributed to such factors. With the ability to control mating, IIQs generally have been selected for superior genotypes, which may mask the possible disadvantages of instrumental insemination concerning their performance. The unfounded reputation for poor performance of IIQs has been difficult to dispel, although the scientific community has repeatedly demonstrated their equal and sometimes higher performance compared to NMQs."


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## JSL

Deknow brings up an interesting point. When I teach instrumental insemination to beekeepers, I caution them that II is a tool. It is a tool that can cause harm to a population or enhance it. Too much control over mating in a small population quickly leads to inbreeding and reduced performance. II allows beekeepers to perform single drone inseminations, but I do not see a useful application of this in the broader scheme of things for long term production and survival.


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## JRG13

I'm not sure if the spermatheca remains receptive after initial mating or the first few weeks of the queens life.


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## JSL

Queens can be re-inseminated. Harbo and perhaps others looked at this.


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## Eduardo Gomes

apis maximus said:


> _*However, beekeeping practices designed to reduce labor, increase efficiency and provide convenience in scheduling, often provide suboptimal conditions for queen development and sperm storage in the spermatheca*_. The reported lower performance levels of IIQs can probably be attributed to such factors. With the ability to control mating, IIQs generally have been selected for superior genotypes, which may mask the possible disadvantages of instrumental insemination concerning their performance. The unfounded reputation for poor performance of IIQs has been difficult to dispel, although the scientific community has repeatedly demonstrated their equal and sometimes higher performance compared to NMQs."


With the quote I presented I just meant to emphasized if the agenda of queens II breeder overlaps a strict protocol, the results are not so good, this according to Dr . Susan own, and maybee this is one of the reasons for low reputation of II queens.

Full quote that you present underlines further this point. Researchers the strictly comply with the protocol get in rule queens of good quality. As far I had understand in USA, Canada, like here in Portugal beekeepers usually do not buy their queens to researchers, or am I wrong?

In the same year in which the article of Dr . Susan Cobey was published (2007) was published this article: Effects of Insemination Quantity on Honey Bee Queen Physiology, Freddie-Jeanne Richard, David R. Tarpy, Christina M. Grozinger. In the abstract we can read: "_Here, we demonstrate for the first time that insemination quantity significantly affects mandibular gland chemical profiles, queen-worker interactions, and brain gene expression. *Further research will be necessary to elucidate the mechanistic bases for these effects*: insemination volume, sperm and seminal protein quantity, and genetic diversity of the sperm may all be important factors contributing to this profound change in honey bee queen physiology, queen behavior, and social interactions in the colony._"

What this abstract is saying, as well as others, is that II is not an area of closed investigation by the very beautiful work of Dr. Susan Cobey . IMO , much of the research points in the direction that there is still much to analyze and much to learn. It is an open field , and it is too early to draw definitive conclusions and in one sense that artificial II is heaven on earth.


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## kilocharlie

Kind of awesome...I've mentioned that 2007 Apimondia article by Dr. Cobey probably 8 times before on Beesource, and this is the first time that there is any evidence that anyone even read read it, let alone understood it. Mostly everyone gets the ideas, too. Dr. Latshaw is even in on the thread! I'm impressed, and pleased.

Fantastic to hear that Dr. John Harbo found that queens can, indeed, be re-inseminated. Is why they don't go out and mate again in nature is a puzzle yet to be completed? (Now I plan to try it for certain, on one of my better "retiring" queens, and with distinctly different sperm, to try further determine male-passed traits from female-passed traits.)

Last I heard, Dr. Cobey had a dual assignment, early spring at U.C. Davis, later at WSU. She also offers private classes for 1 or 2 people at a time at her home on Whidbey Island, WA, in the summer.

Dr. Cobey's New World Carniolan bees could serve as a study group for long-term effects of I.I. on a population of bees. I'll will try to ask her how many generations of the original stocks were back-crossed (or what ever other techniques used???) how many times, leading to this highly-successful bloodline of bees. If I get an answer, I'll do my best to report it back here. Feel free to remind me in a PM.

Another idea surfaces from Eduardo's point: there may need to be a standard protocol set for the entire I.I. process, measurements of it's various aspects, as well as a system of enumerating which revision of the protocol, as it will evolve, corresponds to a study being published. Good show, Eduardo!

Could you please post a link to the Richard/Tarpy /Grozinger article regarding effects of insemination volume on queen physiology? I'd really appreciate that. Thank you.


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## JRG13

Now I'm wondering if swarm queens remate during swarming at any detectable rate...


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## kilocharlie

RayMarler said:


> ... I have a disagreement with a statement that Dean makes at the 5:35 time mark. He stated that "Queens and workers are genetically identical". I have a problem with that statement. The workers are made up of half of the queen's 32 genes and all of their particular father drone's 16 genes, giving each worker a total of 32 genes, and not every worker has the same drone father, and no workers have the same drone father as what their queen mother had (although they do all have the same drone grandfather). Therefore, I think that the Queen is not genetically identical to her worker daughters...


I think he meant a queen is genetically identical to her worker (and queen) SUPER-SISTERS, not her worker daughters, nor her worker "half-sisters". Still, the concept of the definition of "identical" or "congruent" seems almost absurd in nature. I'm sure it can happen, but upon closer investigation, it almost never does.

Like religious kooks who describe themselves as Creationists, whom do not yet know that the ONLY thing we really know is that we KNOW nothing, do not know that all people are agnostics whether thy like it or not (forgetting that we must first convene on what assumptions are being made), we are mostly all genetic mutants, even "identical twins", who are most likely individuals that share a rather high degree of similarities rather than being truly identical.


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## Lauri

If I had to choose, I'd rather have frozed drone semen from my best older breeder queens than keep the original queens producing with new semen. 

I know the frozen semen is tricky to work with, but if you were quick and deliberate before your II queen was superceded, you'd get your new live, fresh source of specific line of virgins to continue your work.
You could keep your original frozen semen around for _years_ for occasional influence back to your lines.

Those are your Heirloom seeds


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## Eduardo Gomes

kilocharlie said:


> I'll will try to ask her how many generations of the original stocks were back-crossed (or what ever other techniques used???) how many times, leading to this highly-successful bloodline of bees. If I get an answer, I'll do my best to report it back here. Feel free to remind me in a PM.


Thank you kilo. Have no doubt that I will remember you if you forget



> Could you please post a link to the Richard/Tarpy /Grozinger article regarding effects of insemination volume on queen physiology? I'd really appreciate that. Thank you.


Just because you are a kind person : http://www.plosone.org/article/info:doi/10.1371/journal.pone.0000980

If you notice any rudeness here or there if I did not take it personally . My English is rude but I am more than my English.


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## kilocharlie

I'm afraid my Portugese is a bit worse than rude.. but it is a good thing that beekeepers with scientific minds get along well and discuss the topic rather than burning cars and attacking each other with hammers! I'd far rather read and learn here that read the latest news from Ferguson, Missouri on Yahoo!  I don't know what is worse, the misbehavior or the "reporting" (pronounced "race baiting")?

Good to have a buddy in Portugal! And thank you for the link.

Good idea, Lauri! Heirloom is a great concept that probably also needs a protocol. I suppose that Brother Adam's Buckfast bees of yesteryear would be more like the "Buckfast" bees of today, at least the ones here in the USA, had there been an heirloom stock kept continuously (fortunately, his Dartmoor mating yard is back in business, and the Karl Kehrle Society keeps his methods alive). Better for baseline control groups for multi-year studies, too. I hear they have made recent leaps in cryogenic storage of bee germplasm, too. Things are indeed looking up. Big Thank You.


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## Brandy

Excuse me coming into this late, but when your talking about re-inseminations are you talking a day or two, or a year or two, and where was the link that this is referencing... Thanks for the help,

Also, are Sue's NWC's treated or able to remain untreated for varroa?


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## AstroBee

kilocharlie said:


> I hear they have made recent leaps in cryogenic storage of bee germplasm, too.


Got a reference? Last I heard (2013) was that they were successful in using stored semen to inseminate queens, but the resulting supercedure rates were very high.


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## kilocharlie

Brandy - We were talking about re-inseminating older queens, and I need to study up on Dr. John Harbo's work before I say too much more. I KNOW that I DON'T KNOW about this topic yet, and Dr. Latshaw says that Dr. Harbo worked on that, so we'll see what he says 

Astro, - I think I saw it mentioned in a scientific paper quite recent, like in the last 3 or 4 months. If I spot it again, I'll make an effort to drag this thread out of the past and post it here. My apologies for not having it at the tip of my brain. Up until now, germplasm has been viable at room temperature for 2 weeks, refrigerated for almost a year. I think it was a recent award mentioning a female researcher for her work advancing cryogenic methods with drone germplasm, perhaps one of the speakers at the California State Beekeepers' Association meeting. I'll check.


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## kilocharlie

Astrobee - my mistake. It was a brief about a speaker for the recent CSBA convention, Megan Taylor, a PhD candidate at Washington State University. The mention was that her masters dissertation at Universtiy of Guelph involved improved methods of cryostorage for bee germplasm, no big breakthroughs, but I will seek out her info and report back.


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## Michael Bush

> Is why they don't go out and mate again in nature is a puzzle yet to be completed?

Some people believe they do. I have always assumed they don't (as has everyone else) but I don't have any definitive proof of that.


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## SRatcliff

Michael Bush said:


> > Is why they don't go out and mate again in nature is a puzzle yet to be completed?
> 
> Some people believe they do. I have always assumed they don't (as has everyone else) but I don't have any definitive proof of that.


Those queens that live 4-5 years; do the bees supersede her because of her health or because she runs out of sperm? I thought the latter.


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## Michael Bush

When a queen is in a booming hive and laying a lot, her life is much shorter than if she is in a nuc and not laying much. Jay Smith claims to have had a queen he was breeding from that was eight years old. His methods were to confine a queen mother to a small area to lay and I would say that's the reason for the long life. I think they run out of sperm.

http://www.bushfarms.com/beesbetterqueens.htm#Queen Alice


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## Lauri

Don't forget the number if eggs are not infinite.

Trying to keep a prolific queen in a smallish colony is difficult to do. That hive ends up being a brood frame and feed donor to a lot of other nucs, unless there is another way of doing it. I have done simlulated swarms with many of my older queens that worked _great_. Still, their colony was difficult to keep small enough to access her on a regular basis easily by late summer. 
But those breeder queens are always of fresh, soft, clean comb. Plus no mite treatments needed if you do it right & you have good groomers. (One or two frames of_ only_ open brood-the rest of the frames new-undrawn) The only way I know how to give a colony a short brood break with an established laying queen. 
I've got many queens going into their third winter I've handled that way. Still they end up in double or triple deeps.

They'll draw out a & fill a single deep in about a week if you have a good number of foragers. Then add a second deep (with a feeder if there is no flow.)
The colony below has a 2012 queen and 3 or 4 year old colony. Off the old frames and onto new. A good chance to refinish the old hive box's too.



















That four deep hive is the one I changed out last March. Almost lost it here.










Heres a photo of that queen










Here she is when first mated, May of 2012










One of the few Buckskin colored Carnie hybrids I use. Or at least plan to use. I tend to favor the darks so I have several queens like this I've just kept around & never quite gotten to.


Heres how that freshened hive looked this fall. I've actually never grafted from this queen because when I've gone to do it, the hives gotten too big again.


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## AstroBee

Michael Bush said:


> > Is why they don't go out and mate again in nature is a puzzle yet to be completed?
> 
> Some people believe they do. I have always assumed they don't (as has everyone else) but I don't have any definitive proof of that.


Queen excluders prior to insemination, plus clipped wings during the insemination pretty much rules out any chance of natural mating.


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## Eduardo Gomes

SRatcliff said:


> Those queens that live 4-5 years; do the bees supersede her because of her health or because she runs out of sperm? I thought the latter.


"The surviving virgin queen will fly out on a sunny, warm day to a "drone congregation area" where she will mate with 12-15 drones. If the weather holds, she may return to the drone congregation area for several days until she is fully mated. Mating occurs in flight. *The young queen stores up to 6 million sperm from multiple drones in her spermatheca*. She will selectively release sperm for the remaining 2–7 years of her life.[3]" in http://en.wikipedia.org/wiki/Queen_bee . I've read something similar in a scientific study that passed through my hands a few days ago that also confirms these figures. If the find will reference it here.

Several experts refer to a queen by year lays about 250,000 eggs. At the end of four years lay about 1 million eggs. Having about 6 million sperm should live about 24 years if their life expectancy was mainly determined by this factor. I do not think is to disregard the possibility of their mandibular pheromone decrease.


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## Lauri

Eduardo Gomes said:


> .Several experts refer to a queen by year lays about 250,000 eggs.


I wonder just how many calories the queen has to consume over a season's worth of production. And there are many that think nutrition doesn't matter. Then there are accumulative toxin exposures to consider.
If you want to good longevity, need to keep them clean and on a superstar's diet.

Along with good genetics and mated well of course.


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## Eduardo Gomes

My mutt bees have a five star diet: nectar, honey, pollen and water. I rarely food with artificial feeding. In Portugal and with my bees things have not gone so bad so far.


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## Lauri

Eduardo Gomes said:


> My mutt bees have a five star diet


I have mutts too, LOL I've let my customers name some of them. One of the more complicated lines is called:

'Miller Ultimutt hybrid' (II Pol-Line based/Russian/VSH/ mated with Carniolans (2014 cross)
This is from Velberts Pol-Line II breeder (originating from VP Queens I am told) Daughters mated at his place with his Russian and pure VSH stck. One of those daughters was grafted at my place. THOSE daughters were late summer mated with my own line of VSH Carniolan hybrids. Overwintering right now. New crop 2015 test daughters from the best overwintered stock will also be mated here. _That_ is a mouthfull. And a touch of diversity, I'd say.
I like the looks of them so far, this is a virgin queen here:










They've already been impressive in the mating nucs, some of the nucs with serious issues to ovecome (Laying workers, no stores, low populations, very late summer. I had such a good crop of these I placed them all as virgins and was able to be very selective.

Pic below is a mid September photo. Virgin shown above was placed 30 days prior.









I enjoy the natural wide variation of nectar and pollen producing plants in my area, with no commercial crops around. But I still feed them a brew or two of my own.
I give my feeding plan a lot of credit for accomplishing so much is such a short amout of time. I also give the feed credit for producing very good sized virgins.


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## Eduardo Gomes

Lauri said:


> 'Miller Ultimutt hybrids'II Pol-Line based/Russian/VSH/ mated with Carniolans (2014 cross)



Only then lack the iberiensis . Lauri If you want i send you a queen of the nicest I have. 

In these diet thing and feeding bees as in many other things in beekeeping there are no doctrines to follow a strict form.
The beekeeping Bible is yet to be written. If ever written I do not know it but I do not feel a need to know. I'm fine and my bees too.


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## Snowhitsky

Lauri said:


> I enjoy the natural wide variation of nectar and pollen producing plants in my area, with no commercial crops around. But I still feed them a brew or two of my own.


We have a very mild winters (16ºC/61ºF at 21.20) with a reasonable amount of flowers and apis iberiensis is very well adapted to the climate which is why we usually don't have to do anything special to get the hives through winter.


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## kilocharlie

I've read testimonials to queens mating with from 1 to 55 drones. Many have quoted an average of 17 stated by one study, but averages aren't individuals, are they? NMQ performance may be poor due to not enough drones flying, by not finding a DCA, and especially by getting eaten by a bird, dragonfly, or other predator! I always hope to breed too many of every combination of queen colonies and drone colonies I can get my hands on in order to get at least some good variety of bees.

Some queens max out on a poor year laying about 1,000 eggs a day in the peak of the season, 2,000 eggs/day is considered good by most standards, 2,500 a day very good. A hot AHB queen in a large hive in premium conditions may lay as many as 5,000 eggs/day, and for a fairly long season, but usually interrupted by an abscond/usurp event or one or more swarming events, or a combination thereof. Russian queens sometimes lay 3,500 eggs/day.

I'm not so sure about 250,000 eggs per year, that sounds pretty low. That's only 100 days at 2,500 eggs/day - a good springtime with zero left for rearing brood the rest of the year. Also, a queen may release 5 to 7 sperm at the time the egg goes out the oviducts. Both of these factors reduce the "sperm longevity" of the queen.


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## Eduardo Gomes

kilocharlie said:


> I'm not so sure about 250,000 eggs per year, that sounds pretty low. That's only 100 days at 2,500 eggs/day - a good springtime with zero left for rearing brood the rest of the year. Also, a queen may release 5 to 7 sperm at the time the egg goes out the oviducts. Both of these factors reduce the "sperm longevity" of the queen.


Hello Kilocharlie
The numbers are huge and I will not give you my opinion, because I did not dare to have an opinion about this. I will introduce you to some other recent data that I have no reason to doubt them.

_A newly established queen bee flies out of the hive and will mate with one or several "drones." Most likely, this one mating event will allow her to lay eggs for the rest of her life. *A queen bee may lay up to 2,000 eggs a day and up to 1,000,000 in her lifetime*._ in http://www.colostate.edu/Dept/CoopExt/4dmg/Pests/bees.htm

"_Virgin queens mate early in their lives and only attend one mating flight. After several matings during this flight, *a queen stores up to 100 million sperm within her oviducts*. However, *only five to six million are stored within the queen’s spermatheca. The queen uses only a few of these sperm at a time in order to fertilize eggs throughout her life*._" In http://www.orkin.com/stinging-pests/...ey-bee-mating/


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## Eduardo Gomes

"*Those queens that live 4-5 years; do the bees supersede her because of her health or because she runs out of sperm? I thought the latter.*"
This is the original question that I answered.



kilocharlie said:


> but averages aren't individuals, are they?


At this point I completely agree with you. The average of a population or sample does not necessarily reflect the reality of each individual.




> NMQ performance may be poor due to not enough drones flying, by not finding a DCA, and especially by getting eaten by a bird, dragonfly, or other predator!


In this respect I can tell you according to my experience that a poorly mated queen is quickly replaced by bees , not getting 2 or 3 months , and therefore does not reach the 4 or 5 years as asked .
Those that are eaten by birds etc ... these are out of these accounts it seems to me. Thus nor live 4 or 5 years.



> A hot AHB queen in a large hive in premium conditions may lay as many as 5,000 eggs/day, and for a fairly long season, but usually interrupted by an abscond/usurp event or one or more swarming events, or a combination thereof. Russian queens sometimes lay 3,500 eggs/day.


If the scenario was this I could not read as well as you between the lines.



> I'm not so sure about 250,000 eggs per year, that sounds pretty low


"Queens lay the greatest number of eggs in the spring and early summer. During peak production, queens may lay up to 1,500 eggs per day. They gradually cease laying eggs in early October and produce few or no eggs until early next spring (January). *One queen may produce up to 250,000 eggs per year* and possibly more than a million in her lifetime." in https://agdev.anr.udel.edu/maarec/honey-bee-biology/the-colony-and-its-organization/

I'm not the family of flying brothers. I have this tendency to believe in the research that speaks to me of things that I do not know or I can not count. And I have given to me well so far and so I think that will not change.


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## kilocharlie

Eduardo Gomes said:


> "*Those queens that live 4-5 years; do the bees supersede her because of her health or because she runs out of sperm? I thought the latter.*"
> This is the original question that I answered. ...
> 
> "Queens lay the greatest number of eggs in the spring and early summer. During peak production, queens may lay up to 1,500 eggs per day. They gradually cease laying eggs in early October and produce few or no eggs until early next spring (January). *One queen may produce up to 250,000 eggs per year* and possibly more than a million in her lifetime." in https://agdev.anr.udel.edu/maarec/honey-bee-biology/the-colony-and-its-organization/...


The 4- and 5-year old queens are rarely discussed anymore. Since the advent of large-scale die-offs, we are very happy with queens that lay sufficiently past one year, and re-queen whenever necessary. Those living to 6 years are lucky girls indeed, and usually live in an environment that does not have the stresses that occur in the continental U.S.A. near monoculture agribusiness, pesticides, and varroa. Places like Pitcairn Island come to mind, I'm sure the gal there gets some queens that live a long time. I had one from 2008 to 20012 (she went 5 years + 7 months) that was my best producer, and some others went 4 1/2 years, but nothing over 3 years since, and recently only some 1-year- and a single 2-year-old queens. Conditions are precluding the expression of longevity in the genetics.

The 1,500 eggs a day figure sounds like A.M. ligurica, the north Italian strain of honeybee. I've seen well-bred queens of this race laying 2,000 to 2,200 eggs a day for several months in good conditions, filling up half a side of a deep frame in 2 days. 

My AHB queens used to go well over twice that egg-laying rate, filling up both sides of a comb in less than 2 days, but they'd go stir crazy when they ran out of brood comb. I had them under 3 layers of queen excluder cage so they could not swarm, nor send out drones. If I could have kept them a little less remote, I could have done more intense management and gotten better data off these amazing bees. I'm sad to have lost them when their cave collapsed.

Here in Southern California, in early December, I inspected a friend's hive yesterday that had 5 1/2 frames of brood. The weather got colder suddenly with a storm October 31, and one rain last Tuesday & Wednesday (Dec. 2nd & 3rd), and the drones are gone (first time in 2 years with no drones over the winter), but that queen is laying a nice, solid brood pattern, and dealing with a varroa population (he has not treated for them yet). From this I conclude that laying season is a local thing, having much more to do with local temperature and nectar / pollen flow than with scientific studies.


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## beepro

kilocharlie said:


> If I could have kept them a little less remote, I could have done more intense management and gotten better data off these amazing bees. I'm sad to have lost them when their cave collapsed.
> 
> Here in Southern California, in early December, I inspected a friend's hive yesterday that had 5 1/2 frames of brood. The weather got colder suddenly with a storm October 31, and one rain last Tuesday & Wednesday (Dec. 2nd & 3rd), and the drones are gone (first time in 2 years with no drones over the winter), but that queen is laying a nice, solid brood pattern, and dealing with a varroa population (he has not treated for them yet). From this I conclude that laying season is a local thing, having much more to do with local temperature and nectar / pollen flow than with scientific studies.


KC, I don't know the laws on keeping AHB in CA (California.) In Florida any swam that they caught need to be requeened with the EHB queen. Keep in mind that the queen will slim down so she can fly in a swarm so 3-5 layers of qe may not hold her in. Maybe to raise big head queens with smaller qE openings and preventing the brood nest from back filling which is hard to do on a flow. If you would like to have a source of AHB genetics I know who has them. Maybe to open up the cave again with railroad ties to reinforce the cave opening. My first thought was to pinch this aggressive queen in which we did. After dealing with them my first season and vid on this process I finally had enough. Now I only keep the gentle type bees. 
To have a long live queen with the quality genetics that you would like to graft from she must be confined in a single frame or 2. Look at the example on oldtimer's queen nuc thread. This will prolong her lifespan and all the good eggs that she carried. Of course, naturally being fed all those good RJ there is a detox mechanism built into every bees including the queen, I think. Then good nutrition and the nice local environment will help tremendously too.
With the changing weather pattern this year, we are warmer than usual with lots of rains so far. Like KC's friend, I did not let my Italians goto sleep or on a brood break this year. Already 4 frames of young winter bees are hatching with the mites they have to dealt with also. I did not treat hoping to test these late Fall queens to see their mite resistant capability as advertised. Because this year we have a drought just like the last 3 years, there isn't much pollen to be stored. Now they are picking on the eucalyptus pollen and nectar with some overwintering Borage too. So far they are brooding up nicely feeding on Lauri's sugar bricks and patty. Yes, the local environment has many to do than with any scientific studies since we don't know the local conditions at which they have done these studies on. But still a good read to get an idea though.


Today's hive check:


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## Eduardo Gomes

kilocharlie said:


> The 4- and 5-year old queens are rarely discussed anymore


I answered a rare question then. I did not know that . Rare matters not deserve the most accurate answer possible in your opinion?



> Since the advent of large-scale die-offs, we are very happy with queens that lay sufficiently past one year, and re-queen whenever necessary.


Sorry to hear that. In your opinion what are the causes?



> The 1,500 eggs a day figure sounds like A.M. ligurica, the north Italian strain of honeybee. I've seen well-bred queens of this race laying 2,000 to 2,200 eggs a day for several months in good conditions, filling up half a side of a deep frame in 2 days.


You still insist on the idea that 250,000 is a low number. Let's put the numbers in another way: in the spermatheca are usually 5 million to 6 million sperm. For a queen exhausts this reserve in 5 years would by 1 to 1.2 million eggs per year . Laying 4000 eggs per day was to keep this standard approach for about 300 days year. Do you think this is credible? Do you think there is any place in the world one that can happen?



> From this I conclude that laying season is a local thing, having much more to do with local temperature and nectar / pollen flow than with scientific studies.


I have seven winter apiaries. 2 of them are in one side of my country and the other 5 are in the other side of the country . In recent queens stop laying. In the other two have my strongest Lang with 7 to 8 frames with compact brood. And there are still a peak. 4 years ago that I see it every year. No wonder nothing what you say. But I think that was not the point of our analysis.

"Those queens that live 4-5 years; do the bees supersede her because of her health or because she runs out of sperm? I thought the latter."
This is the original question that I answered. ... Your answer to this question you still not given .

Can you give me the reference on the assumption "a queen may release 5 to 7 sperm at the time the egg goes out the oviducts". Thank you!


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## mike bispham

Eduardo Gomes said:


> You still insist on the idea that 250,000 is a low number. Let's put the numbers in another way: in the spermatheca are usually 5 million to 6 million sperm. For a queen exhausts this reserve in 5 years would by 1 to 1.2 million eggs per year .


Eduardo, this assumes that the queen uses a single sperm to fertilise each egg. It may be that does happen, but do we know it does?

Mike (UK)


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## Eduardo Gomes

mike bispham said:


> Eduardo, this assumes that the queen uses a single sperm to fertilise each egg. It may be that does happen, but do we know it does?


"Virgin queens mate early in their lives and only attend one mating flight. After several matings during this flight, a queen stores up to *100 million sperm within her oviducts. However, only five to six million are stored within the queen’s spermatheca. The queen uses only a few of these sperm at a time in order to fertilize eggs throughout her life.*" In http://www.orkin.com/stinging-pests/...ey-bee-mating/

Mike, so far this is the best answer I have for your question. But I will research the research of others. I am a beekeeper , I am no researcher. When I can not find the solution for my questions or problems I look for the answers or the most experienced beekeepers or the researchers. You already know my method of approach. This matter does not seem to be answered by the former. So will be the latter.


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## mike bispham

Eduardo Gomes said:


> "Virgin queens mate early in their lives and only attend one mating flight. After several matings during this flight, a queen stores up to *100 million sperm within her oviducts. However, only five to six million are stored within the queen’s spermatheca. The queen uses only a few of these sperm at a time in order to fertilize eggs throughout her life.*" In http://www.orkin.com/stinging-pests/...ey-bee-mating/


Thanks Eduardo. It was kind of an idle question on my part! I'm not really burning to know the answers to this stuff, so don't do lots of work on my behalf!

Mike (UK)


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## mbc

kilocharlie said:


> Also, a queen may release 5 to 7 sperm at the time the egg goes out the oviducts. Both of these factors reduce the "sperm longevity" of the queen.





Eduardo Gomes said:


> The queen uses only a few of these sperm at a time in order to fertilize eggs throughout .


Hi Eduardo and kilo, my googling has failed me, do you have any links to work on this = how many sperm a queen releases to fertilize an egg, (your above link wouldn't work for me Eduardo)
It seems more likely to me she would release a squirt each time, containing many sperm.


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## Eduardo Gomes

mbc said:


> your above link wouldn't work for me Eduardo).


http://www.orkin.com/stinging-pests/bees/mechanic-sof-honey-bee-mating/
Sorry but after a while it seems that the link no longer works . I'll try to find another access.


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## mbc

Many thanks 

Having looked over the piece in the link, it's not really provided the enlightenment I was hoping for, anything a bit more authoritative?


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## kilocharlie

Eduardo Gomes said:


> I answered a rare question then. I did not know that . Rare matters not deserve the most accurate answer possible in your opinion?


Good to hear somebody still has 4 & 5 year old queens! I note the good 2-year-olds for candidates go to the drone yards, if they score high enough. Strong colonies of 2nd-year queens seem to always have more drones.




Eduardo Gomes said:


> Sorry to hear that. In your opinion what are the causes?


My opinion is that Americans are screwing up so much, so bad, and so fast we can't even identify, nor measure all the causes. This attitude of, "What's in it for me?", "If it doesn't agree with my opinion, you must suck!", "Not my problem", "That kind of thinking is so obsolete, I already saw it on YouTube last week!", "I'm better than you", "Don't inconvenience me, I need to get high" - it seems to be at, or very near the cause of many of the problems. The national personality is and has been going away from sustainability at high speed for a long time. Some are starting to talk about it, but to begin practicing it won't happen until the last tree is a national park. Pesticides are a problem, perhaps carbaryls and pyrethrins are a couple that are worse than neonicotinoids, but don't tell the liberal non-beekeeper I said that. They only want to have a march against neonic's down Main Street, then go have an organic beer or 12 and tell everyone how *****in' they are. Agribusinessmen have tripled the amount of fungicides they used a few years ago, coincided with huge die-offs in 2012. Related? Maybe, bees do need fungi in their digestive tracts to digest the proteins from the pollens. I have no studies to prove nor disprove either way. I'd like to see it *NOT HAPPEN,* but's too late now - 2012 is gone with wind and the dead bees. They made a small increase in yield, and many beekeepers got 60% losses across the country.

Chief Seattle, of the Dwamish and Squamish tribes concluded a speech, "...One thing is certain, continue to defecate in your own bed, and one day you will not awaken, having suffocated in your own waste." 

Many Americans seem to do a lot of things because they can, because it's cheaper, or because the saw it on TV (OK, the internet nowadays), or because they think it's "cool", not because they "should", nor do they listen to the Buddhist when he says, "Do nothing, time is precious." The don't give a thought about, "Is it _wise_?" If they do, wisdom to them is in a magazine about Feng Shui, or a cool line in a movie, or "how many Karma points do I get for it?". Few of them first master the self, few become caretakers of the earth and it's creatures, few are the seekers of truth, as passionate about wisdom as a drowning man is for a breath of air. Beekeepers are a shade to the brighter side. They are actually doing something that, done properly, leads toward "wisdom"! Especially those who study and ask lots of questions while practicing and revising this trade / craft / art, that is, they seek the wisdom in it and practice it. A yogi, a bodhisatva, or a guru would smile.




Eduardo Gomes said:


> You still insist on the idea that 250,000 is a low number. Let's put the numbers in another way: in the spermatheca are usually 5 million to 6 million sperm. For a queen exhausts this reserve in 5 years would by 1 to 1.2 million eggs per year . Laying 4000 eggs per day was to keep this standard approach for about 300 days year. Do you think this is credible? Do you think there is any place in the world one that can happen?


The part about 4 to 7 million sperm in the spermatheca I have read many studies with similar numbers, so agreed there, but the math applied to the days per year and number of years for EGGS seems skewed a bit. I think you assume one sperm per egg. The queen probably releases several sperm (5 to 7 released at a time was mentioned in a study - though I don't have it handy to back up the statement) with each egg to be fertilized when laying in a worker or a queen cell (my guess is perhaps more variability than this within a queen, and from queen to queen) If this is true, that immediately brings either the number of days per year or the number of years down to numbers consistent with what what we recognize as observed queen longevity - as much as 3 to up to 8 years. 2,000 eggs a day for 8 months (~240 days) is about 480,000 eggs, close to half a million eggs. Multiply that by 5, and you have about 2.4 million sperm per year. But she slows down that 3rd year, so she runs out in her 4th year if she is very prolific, and in good conditions for a lot of months, like I get to see in good years here, and what my friends report farther south in Mexico. In today's scene, with so many stressors (seen or unseen by us beekeepers or scientists) acting against the success of the hive, she gets superceded in year 2 or 3 before she runs out of sperm. Different locales, different numbers. Poorly mated queen, lower numbers. I'll have to get back into gambler mode and run a spreadsheet on all the sperm-to-longevity scenarios. I think queens get 86'ed (superceded) for running out of sperm, running out of eggs, or just plain slowing down too much to keep the colony viable (call it poor health or just old age). 

Tropical queens seem to be prolific, laying year-round, and tend to not live as long as temperate zone queens, who lay fewer eggs per year but last a few more years.

I think that bees exposed to some chemicals have been shown to possibly lose some of their sensitivity to the quality of nectar and/or pollen, causing the bees to take interest only in very sweet, high-protein sources of nectar/pollen, thereby reducing both the total amount of nectar/pollen brought home and most certainly the total range of varieties down to a much narrower band of "acceptable" foods. This may be interpreted by the bees to render a "not enough bees in the house, therefore supercede the queen" reaction. Beekeepers interpret it as poor queen performance, when the reality was possibly imidicloprid or other chemical. If this is true, you see why I think more like 1 million to 2 million sperm per year are used by queens in strong colonies situated in mild climates, yet the hives aren't performing like the amount of brood would make me think they should. More bees bringing in less food. Plus, there's mites and other cooties subtracting from the score.




Eduardo Gomes said:


> I have seven winter apiaries. 2 of them are in one side of my country and the other 5 are in the other side of the country . In recent queens stop laying. In the other two have my strongest Lang with 7 to 8 frames with compact brood. And there are still a peak. 4 years ago that I see it every year. No wonder nothing what you say. But I think that was not the point of our analysis.
> 
> "Those queens that live 4-5 years; do the bees supersede her because of her health or because she runs out of sperm? I thought the latter."
> This is the original question that I answered. ... Your answer to this question you still not given .
> 
> Can you give me the reference on the assumption "a queen may release 5 to 7 sperm at the time the egg goes out the oviducts". Thank you!


I don't clip all my queen's wings, so I rarely keep one that many years. I do lose some to swarming, mostly drone mothers. I call that "giving back to mother nature". But my oldest queens were both tired AND running out of sperm (or perhaps eggs). Which ever it is, or perhaps both reasons, she gets superceded. If I catch this in time, I'm harvesting supercedure cells and making nuc's or re-queening hives that need it for as long as I can. I will feed profusely and newspaper combine several weak hives to keep this going. Supercedure cells usually make good queens (probably average better than grafted E-cells from queenless starters), usually in low numbers like I want late in the season, when I have some time to play with this.

I do not recall the study that mentioned in passing that "several" sperm are released with the eggs to be fertilized, nor the study referenced in the footnotes that explained it. I will ask the next PhD's I talk to, someone will likely say, "Oh, that's Dr. X over at the University of Gilligan's Island. Here, I'll text him." That, or they forward my email to a scientist who was in on that experiment. Then I get a study in a pdf download the next day with an answer. Gotta love people like that! So far, a lot of them seem to have a good idea about who knows what, and it's been fairly easy to get incredible help. Most of them seem delighted to help us beekeepers. Wish me luck, though, that can't hurt. I've certainly been very lucky so far


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## Eduardo Gomes

kilocharlie said:


> My opinion is that Americans are


Kilo thanks for your good slow reply. Also in Portugal and in Europe there are many people with this "style" . Alessandro Baricco , an Italian writer, calls these "characters" new barbarians.

Bees teach us to be more " slow ", more patients, more observant, more respectful of the rhythms of the natural cycles .

On the question in debate I'm from an assumption that is capable of being wrong . I will also do my research. Good luck to you and good luck to us all!


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## Juhani Lunden

JSL said:


> Queens can be re-inseminated. Harbo and perhaps others looked at this.


Really, after first weeks and initial mating, as JRG13 doubted?

Do you have any studies to link?

Insemination becomes trickier.


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## Michael Bush

Americans have been in a hurry since the pilgrims arrived. The Europeans farmed with oxen. They were too slow for us. We used horses (2 1/2 times faster). But that wasn't fast enough so we invented tractors and bigger tractors and thrashing machines and combines until we were plowing, not just an acre a day (the amount a team of oxen can plow in a day) but now we can plow 100 acres a day with a 10 bottom plow... some of the modern tractors are 900 horse power...

Yes the attitude spills over into beekeeping...


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## JRG13

Don't forget about natural attrition of stored sperm, I'm betting there's a % loss of viability every few months etc...


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## kilocharlie

I spotted a different source of the "several" sperm released with each egg, and the reference was to Dr. John Harbo, 1977 

...typically 5 to 7 sperm are released as the egg passes through the oviducts... 

but the exact study is not cited. Anyone down in Dr. Harbo's area know him? Probably published in an entomological journal, apidologie, or proceedings of a society? I'd love to read more of his work, and Otto Mackinson's, and several others.

JRG13 - Yeah, and the queens we replace twice a year probably didn't get enough royal jelly as a larva, got bumped as a pupa, didn't mate with enough boys, their daughters weren't bringing in much pollen, and then the sperm they did get goes bad in the tank, too!:waiting:


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## Eduardo Gomes

In my research on the net I found this study dedicated to the issue at hand.

"_Queen age did not affect sperm use (r= 0.334, n= 7, p= 0.464) (see Figure 4); both the young (≤ 6 months) and old (≥ 18 months) queen groups had a median of three sperm per egg.[…] Overall, *queens seemed to be very economic in using their sperm supply to fertilize eggs; an overall median of only three sperm per egg was found*. None of the queens had a median over ten sperm per egg, however individual queens varied significantly in their sperm use (Kruskal-Wallis test, H= 93.873, df= 6, p<0.001) (see Figure 3). For example, queen 9, 10, and 17 used a median of two sperm per egg, while queen 18 used a median of nine sperm per egg. […] It would also be very interesting to run a separate experiment looking at the effect of this factor in which queens would be introduced and followed throughout their settling to see *if sperm use really does decrease as the queen becomes more comfortable in the colony and as the colony has more related individuals and is less likely to reject her*.[…] This method could also be used to see *if sperm use patterns have any hereditary components* and if related queens use a similar number of sperm per egg. If this is the case, *it would be extremely useful to beekeepers to be able to breed queens who use the fewest number of sperm possible* without laying patchy brood (as workers remove any eggs that have been mistakenly unfertilized). *This would lengthen queens' lifetimes and reduce the need for queen replacement, which costs keepers precious time and money*._"

in Sperm Use During Egg Fertilization in the Honeybee (Apis Mellifera), Maria Rubinsky, 2010.


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## Juhani Lunden

JRG13 said:


> I'm not sure if the spermatheca remains receptive after initial mating or the first few weeks of the queens life.


Answer:



JSL said:


> Queens can be re-inseminated. Harbo and perhaps others looked at this.


I´m still eager to know more about this. Is it really possible after initial mating and first weeks of queens life? 
What I do know, is that
- freemated queens are very tricky to inseminate, if it is possible at all to get the tube in
- queens can be inseminated several times, if done with one days interval


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## mbc

mbc said:


> I was hoping for anything a bit more authoritative?


Super! I think that's comprehensively answered the "how many sperm per egg" question, thanks guys


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## mike bispham

Eduardo Gomes said:


> _"....This method could also be used to see *if sperm use patterns have any hereditary components* and if related queens use a similar number of sperm per egg. If this is the case, *it would be extremely useful to beekeepers to be able to breed queens who use the fewest number of sperm possible* without laying patchy brood (as workers remove any eggs that have been mistakenly unfertilized). *This would lengthen queens' lifetimes and reduce the need for queen replacement, which costs keepers precious time and money*._"


Possibly. I think the assumption that more economical queens are an improvement over less is dubious without further work. In many natural features there is a spread between two extremes (or something multifactorial) going on for a sound reason - because it aids the population. Can we be sure that, for example, there is not a competition locating fittest sperm going on? 

Mike (UK)


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## apis maximus

mike bispham said:


> Can we be sure that, for example, there is not a competition locating fittest sperm going on?


Good point Mike.
I think that the more we look at the issue, the more it becomes evident that a competition does exist. 
Baer talks about such potential mechanisms, namely *sperm competition* and *cryptic female choice*.
http://www.apidologie.org/articles/apido/pdf/2005/02/M4072.pdf


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## Eduardo Gomes

Good thinking you both bring to the table.
Apis maximus do you know if this line of research (sperm competition and cryptic female choice) had latest developments?


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## apis maximus

Eduardo Gomes said:


> ... do you know if this line of research (sperm competition and cryptic female choice) had latest developments?


No, I don't. I am sure though that they are ongoing.
But I would love to be a fly on the wall at a nice, amicable dinner table with the Koenigers, Harbo, Tarpy...and maybe a few others for good measure...discussing and debating topics like this and listening to their findings and observations, that never get to be published.


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