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Natural Selection Rambling

2976 Views 21 Replies 7 Participants Last post by  GregB, 
Litsinger
Since it is relatively cold and quiet in the beeyard and gets dark early now, I have more time inside to read / study various random concepts in the treatment-free realm and roll them around in my mind.

One such concept I keep struggling to wrap my mind around is the division between so-called Natural versus Artificial Selection, at least in a managed apiary. An article recently making the rounds in TF circles penned by Mr. David Heaf entitled, 'Dealing with Varroa: natural selection or artificial selection?' touches on this idea:

https://www.naturalbeekeepingtrust.org/natural-selection

In it, Mr. Heaf makes the following statement: "Whereas artificial selective breeding is a justifiable approach in husbandry generally, it is questionable whether it is a sustainable approach with the honey bee, an essentially wild creature. Focusing on individual desirable traits that appeal to the beekeeper such as Varroa resistance (e.g. Varroa sensitive hygiene), docility, honey productivity etc. may displace traits for long term survival from the mix of traits that more holistic breeding by natural selection delivers. In short, in not knowing the direction in which natural selection is heading, killing colonies could be throwing away good genetics."

While I recognize that almost anything we do in relation to bees exerts some influence upon the colony and could thus technically be defined as artificial selection of a sort, what I am curious about is whether we can ever consistently and repeatably produce a bee with superior overall fitness with a grafting tool versus an innate colony-driven impulse to produce reproductive and/or supercedure cells?

One thing that has always troubled me is to read about folks who have been successfully TF for a few or several years only to experience increased rates of failure to the point it is no longer sustainable.

While I recognize there are many, many factors to consider as to what may be the root cause(s) in each specific situation I often wonder if increased beekeeper selection might be at least partially to blame in some situations?

I am way out of my depth asking this question, but part of what has me wondering this is a talk by Dr. Keith Delaplane (MSL previously posted it here) in which he spends quite a lot of time talking about the various genetic sub-families present in a diverse hive and how the colony as a whole will preferentially choose from these sub-families to develop a new queen when given the option:

https://www.youtube.com/watch?v=PglNInHcj1I

The whole talk is well worth the investment in my opinion, but at about the 50:00 mark he gets into the 'Royal Patriline' concept and it runs to about the 57:00 mark. The crux as I understand it is that reproductive swarm cells (in general) produce the best overall fitness, especially as compared to supercedure cells.

So while I make no judgment upon folks breeding for traits, is it safe to assume that (in general) colony-driven reproductive swarm cells will produce the overall fittest colonies all other factors (i.e. colony health, drone encounters, etc.) being equal?

Again, I am not advocating for any particular breeding approach or method but simply trying to better understand more about the overall impact of bee versus beekeeper-driven queens.
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J
Litsinger said:
https://www.youtube.com/watch?v=PglNInHcj1I

The whole talk is well worth the investment in my opinion, but at about the 50:00 mark he gets into the 'Royal Patriline' concept and it runs to about the 57:00 mark. The crux as I understand it is that reproductive swarm cells (in general) produce the best overall fitness, especially as compared to supercedure cells.
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Brother Adam wrote that supersedure cells make inferior queens compared to swarm cells.

Bees are able to select larvae for their like(favor one particular patriline), but the problem in natural selection concepts is mating. In dense beekeeping areas this leads to failure. Simple as that, but hard to accept for many.
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Litsinger
Juhani Lunden said:
Brother Adam wrote that supersedure cells make inferior queens compared to swarm cells.

Bees are able to select larvae for their like(favor one particular patriline), but the problem in natural selection concepts is mating. In dense beekeeping areas this leads to failure. Simple as that, but hard to accept for many.
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Juhani:

Thank you for your reply. I sincerely appreciate it. I respect and admire the breeding program that you have developed, and I put a lot of stock in what you have to say.

I suppose my question has less to do with open versus closed mating per se but more to do with egg selection- i.e. the process that produces a queen cell.

In his talk Dr. Delaplane makes reference to research suggesting that the resident queen is involved in the process behind selecting the patriline which is placed in individual swarm cells whereas a supercedure cell is worker-driven and thus runs the risk of evoking the expression of parasitic genetics.

I can do his talk no justice, but fundamentally it begs the question in my mind if we would be generally better-served to employ swarm cells as the primary basis for our propagation efforts?

This obviously is but a small slice of the overall concept of 'natural selection', and I concede your point about controlled mating.
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crofter
You propose that the workers have potential selection opportunity in supercedure or emergency conditions, and that the queen could have input only in swarm cell requeening.

Do we know that workers do make active choices or is this only considered a possibility? I find it difficult to accept that the queen can keep track of separate egg and sperm lineage and also implement the combination with intent. This would require knowledge on her part about the significance of a deliberate supercedure or swarm cell vs a regular brood cell.

I can see that the answer to these questions would have a big potential impact on the wisdom of the grafting process compared to leaving all such choices to the bees.

Do we have the wisdom and foresight to know that we would be making the best decisions? Left up to the bees discretion would they move in directions we would not find to our liking?

You pose a difficult question!
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GregB
Everything in this life seems to run in a spiral trajectory.
Most all new inventions/discoveries - had something related already been found/thought about (and often ignored/not-understood) in the past.

For the purposes of the science, it'd be really useful to resort to keeping the bees in the most natural setting possible (a log or a skep) AND let them run the normal life cycle (let them swarm if they do so) AND apply the latest technologies/tools to study the normal processes then.

Why don't the "researchers" get this, rather obvious, experiment setting?
Obvious to me, anyhow.
Even the standard foundation/standard frame usage in the experiment setups - is polluting and invalidating the experiments to a large degree (if not totally).

What about hit a pause button.
Setup a good baseline (use the latest tech against the natural, unaltered bee environment).
Then review everything.

Am I an idiot thinking this?
Maybe I am.
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crofter
Greg, I think the hollow tree scenario would be a difficult condition to replicate in any meaningful quantity compared to the total existent population. The sheer numbers and population density of the real world of bees is of such overwhelming influence that it makes the log or langstroth decision trivial. Forced pre-industrial conditions could easily point out that a more self sufficient strain could be selected for, but the associated disposition and productivity characteristics would not make it fly.
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GregB
crofter said:
Greg, I think the hollow tree scenario would be a difficult condition to replicate in any meaningful quantity compared to the total existent population. The sheer numbers and population density of the real world of bees is of such overwhelming influence that it makes the log or langstroth decision trivial. Forced pre-industrial conditions could easily point out that a more self sufficient strain could be selected for, but the associated disposition and productivity characteristics would not make it fly.
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Given the tools, tech, cheap wood available - production of several hundred simulated log hives for experimental purposes is rather a trivial project.
No one cared to look into it, to my knowledge.
Yes - need some funds and someone who cares to do it (nothing new).

Doing few independent and objective baselines (trying to quantify and collect data) using such setups should produce lots of curious findings, I imagine.
(no need to connect to existing populations or look for immediate practical applications - not even a point)
If for academic purposes only - totally fine.
But who knows - the usual case, again.

Sam Ramsie findings - who cared to look if it was easier to theorize and assume, until he actually DID look.
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crofter
New Zealand had a large number of feral colonies in trees; still do, but according to what I read on their forums the ferals do not seem to be any bastion of disease resistance. They do manage to survive while continually reinforced by swarms from commercial colonies but are considered a great part of the problem rather than a potential solution.

Pastoral nostalgia!
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GregB
The strict tree dwelling adherence is not the thing.

Unmanaged bee in a sufficiently closely simulated tree dwelling should be sufficient for data collection and baseline settings.
Similar to this - John Moerschbacher from Alberta(?), Canada has it:
Bee Beehive Honeybee Honeycomb Apiary


Standard lumber; standard tooling; some high tech gizmos for in-place data collection; some crude nest destructions/manipulations will be needed for more data collection (genetic material sampling, etc).
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J
Wasn't there a study or paper saying that the worker bees tend to select the least represented sister group to make queencells from? But I also recall a study or paper saying that super-sister groups tend to promote themselves when making queencells.
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Litsinger
crofter said:
You propose that the workers have potential selection opportunity in supercedure or emergency conditions, and that the queen could have input only in swarm cell requeening.
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Frank:

Thank you for your thoughtful reply. To be fair, I don't consider myself either intelligent enough or experienced enough to propose anything, but am rather trying to think through the implications of assertions made by those much more qualified than me.

That said, I would suggest that this is exactly what I understand that Dr. Delplane is asserting.

His point (as I understand it) is that the process of natural requeening is more nuanced than we generally think and that this process reflects an evolutionary struggle between the castes.

If I can be permitted to 'dumb down' his point, I think he is saying that all of the luminaries in our field have understood that reproductive swarm queens are better than supercedure queens but that we have not fully-understood or appreciated the underlying dynamics, to whit, the presence or absence of the resident queen in the process.

Like I said, I am way over my head in all this, but it does raise an interesting question about the implications of various methods utilized to propagate.
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J
On supercedures though, many are planned where the queen lays in a queen cup, but I don't know if a queen can select certain sperm from different matings and how would she know the best one to use anyways....
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msl
here is the study people are talking about
https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0199124

In his talk Dr. Delaplane makes reference to research suggesting that the resident queen is involved in the process behind selecting the patriline which is placed in individual swarm cells whereas a supercedure cell is worker-driven and thus runs the risk of evoking the expression of parasitic genetics.
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What was meant is when the queen lays an egg in a swarm cell, or a human picks up a larva with a grafting tool the chance of it being one of the rare parasitic genetics is very, very low and you have a much better chance of transferring traits the colony is expressing. But when the bees chose by making E cells the chance is 40%.

Ie from the study
" Because swarm queens are raised directly from queen-laid eggs, there is little opportunity for workers to bias the outcome. Such activities would be limited to preferentially transferring eggs/larvae from worker cells into queen cups [52], or selectively rejecting certain queen-deposited eggs/larvae cannibalism or destruction of completed swarm cells prior to queen emergence [21]. Lattorff and Moritz [22] found that swarm queens do not significantly differ from worker patriline distributions,"

"Similarly, supersedure queens (produced when a queen is failing but still present and laying) are presumably also laid by queens in queen cups [53] and so should not undergo significant selection by workers. Because of this, the “royal” patriline bias is a primary factor in only a minority of requeening events."

This is key to many TF types
"Because of this, the “royal” patriline bias is a primary factor in only a minority of requeening events"
The split culture(make them queen less, put cells in nucs) causes this to become a major player in the game, as 40% of the queens you make from a "good" hive won't have the genetics that made the hive good, and because of them being E Cells another 60% or so of those queens will be sub par. While trying to be "natural" They are propagating unnatural genetic lines and pushing those drones out.


So while I make no judgment upon folks breeding for traits, is it safe to assume that (in general) colony-driven reproductive swarm cells will produce the overall fittest colonies all other factors (i.e. colony health, drone encounters, etc.) being equal?
Click to expand...
Going with swarm cells would be good, if it didn't lend its self to swarmy stocks..
Tarpy's work shows grafted cells on par with swarm cells for queen quality.

If we wish to make good queens, from genetics of our choosing, when we need them. We need to select larva of the correct age(grafting, cell punch, cut strips, nicot, etc).
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crofter
From many past discussions here on Beesource, I dont think there is a great consensus that the issue from a swarm motivated queen cell is measurably different than that from grafting optimum aged larvae raised in a well populated and well supplied colony. Is it more than conjecture, that there might be some benefit from allowing the workers and queens possible input? I am asking this as a question rather than stating. If it is true, it would be a real forehead slapping event if we find for sure that we have been discarding such potentially consequential factors.
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GregB
msl said:
Going with swarm cells would be good, if it didn't lend its self to swarmy stocks..
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Not necessarily.
You simply force the queen into a swarm situation and they/she will create the proper swarm QCs.
Good queen in a small hive.
Check weekly and steal the QCs.
Should do it.

Not always possible, but something I try for in practice - timing the split to when they set a batch of swarm cells is about perfect.
It depends, but most any bee (however NOT swarmy) will try to swarm if kept in 40 liters or less.
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Litsinger
msl said:
here is the study people are talking about
https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0199124
Click to expand...
MSL:

Thank you for your helpful and detailed write-up. I also appreciate you posting the referenced research paper for our collective review.

After reading the paper, my take-aways were three:

1. The Background:

A total of 6 colonies produced sufficient data to analyze, yielding a total of 649 emergency queens sampled over a 10–14 week period.

Genotype analysis of the 6 experimental colonies identified 327 total subfamilies (34–77 per colony) from a total of 552 workers and 512 queens. This included 108 subfamilies (4–40 per colony) exclusively detected in workers and 130 subfamilies (5–55) exclusively detected in queens. An average of 40.21% of the queens produced per colony were from queen exclusive “cryptic” subfamilies.

2. Hyperpolyandry:

The identification of these cryptic “royal” subfamilies reveals that honey bee queens, already considered “hyperpolyandrous,” are mating with even more males than has been previously recognized. These results alter our understanding of reproductive behavior in honey bees, raising questions about the evolutionary implications of this phenomenon.

If these results accurately reflect the subfamilial makeup in colonies of Apis mellifera, they cast into doubt the established concepts of honey bee reproductive behavior such as the degree of polyandry in honey bee queens.

Given that the validity of these findings appears to be robust, there are two immediate implications that revise our understanding of honey bee reproduction: (1) there is a cryptic population of queen-biased patrilines within colonies that are consistently underrepresented in molecular-based estimates of honey bee polyandry, and (2) honey bee queens are mating more than previously recognized, as traditional mating estimates are calculated from worker populations using biased samples that often exclude the rare “royal” patrilines.

3. The Tension Between Castes:

A complex behavior such as queen selection—with its huge implications on future colony survival, relative subfamily fitness, and individual fitness of the candidate larvae—is certainly evolving under the pressures of multiple forces of selection. While many of the specific details and mechanisms are still to be determined, at this point we may safely conclude that, while inclusive fitness for nepotism may favor the individual level during emergency queen rearing, that advantage is profoundly overridden by opposing selective forces acting at multiple levels favoring cooperation and altruism.

… the conflict between individual- and colony-level selection is clearly visible when the colony engages in the process of selecting a new queen.

One factor countering this affect is the lack (or at least significant reduction) of worker-induced patriline bias during swarming and supersedure queen rearing.
Emergency queen selection may be viewed as larval competition as much as worker selection, as any advantage in attracting “royal treatment” from nurse bees during the critical hours of larval selection would confer a huge potential fitness boost on those larvae.
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Litsinger
crofter said:
Is it more than conjecture, that there might be some benefit from allowing the workers and queens possible input? I am asking this as a question rather than stating.
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Frank:

I could not have said it better myself- while I do not presume to have any answers, my mind wanders and I am left wondering whether there is more to the process of a colony selecting a queen than we might have previously anticipated.

If one assumes so, it leads to the inevitable question of whether I can do better than a colony in selecting larvae for queen rearing for the purposes of overall colony fitness?

Again, I concede the point that it is appropriate and maybe even wise to direct queen rearing when selecting for traits- that said (and getting more to the TF concept at hand), it seems that TF viability might be based at least in part on this nebulous concept of overall colony 'fitness' which appears to be the complex interaction of various traits being expressed in myriad sub-families and expressed in varying degrees based on the selection pressure being brought to bear on the local population.

Ultimately, makes me come back to the practical implication of such an assumption- one should primarily propagate via colony-driven swarm cells?

I have no answers, just questions...
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Litsinger
JRG13 said:
... but I don't know if a queen can select certain sperm from different matings and how would she know the best one to use anyways....
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JRG13:

Thank you for your responses- I appreciate your input. I certainly do not know either. It is kind of interesting to think through the implications if we determine that she does in-fact know the best one(s) to use...

Thanks again for your feedback.

Russ
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A
Litsinger said:
Frank:



Ultimately, makes me come back to the practical implication of such an assumption- one should primarily propagate via colony-driven swarm cells?

I have no answers, just questions...
Click to expand...
Well, my plan for next spring is to keep a few queens in 5-frame nucs and feed the heck out of them, pushing them to make as many swarm cells as they can. I then pull those swarm cells into as many new hives as I can manage to screw together this winter. If all goes well it should be easy to get enough swarm cells to make up the 20 or so hives I want next year. All chosen by the hive/queen via whatever system they use. If I get to July and still have empty boxes I'll do some fly-back splits and try to fill them.

People argue which type of queen is 'better'. I don't know enough to have any real idea. But I SUSPECT that letting the bees do the choosing MIGHT work better for what I want. I am not producing bees for maximum honey production, pollination or as a business. The two traits I want are survival, and not too nasty disposition. I'll cull any too-nasty queens and let the bees handle selection for survival.

Right now I have one queen line, a swarm I got in May 2019 and her 4 daughters. Not a lot to work with genetically unless there are some good drones out there somewhere.
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GregB
AR1 said:
Well, my plan for next spring is to keep a few queens in 5-frame nucs and feed the heck out of them, pushing them to make as many swarm cells as they can....
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Good plan.
If my #1 makes it into the next year (fingers crossed) - will try to do the same.
All she will be doing - making swarm QCs and pumping drones.

One BUT - using a 5-frame nuc for QC generator is a little too small I feel.
40 litre unit is good.
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