# Natural Selection Rambling



## Juhani Lunden (Oct 3, 2013)

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.


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.


----------



## Litsinger (Jun 14, 2018)

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.


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.


----------



## crofter (May 5, 2011)

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!


----------



## GregB (Dec 26, 2017)

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.


----------



## crofter (May 5, 2011)

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.


----------



## GregB (Dec 26, 2017)

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.


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.


----------



## crofter (May 5, 2011)

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!


----------



## GregB (Dec 26, 2017)

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:








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).


----------



## JRG13 (May 11, 2012)

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.


----------



## Litsinger (Jun 14, 2018)

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.


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.


----------



## JRG13 (May 11, 2012)

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....


----------



## msl (Sep 6, 2016)

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.


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?


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).


----------



## crofter (May 5, 2011)

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.


----------



## GregB (Dec 26, 2017)

msl said:


> Going with swarm cells would be good, *if it didn't lend its self to swarmy stocks*..


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.


----------



## Litsinger (Jun 14, 2018)

msl said:


> here is the study people are talking about
> https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0199124


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._


----------



## Litsinger (Jun 14, 2018)

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.


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...


----------



## Litsinger (Jun 14, 2018)

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....


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


----------



## AR1 (Feb 5, 2017)

Litsinger said:


> Frank:
> 
> 
> 
> ...


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.


----------



## GregB (Dec 26, 2017)

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....


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.


----------



## AR1 (Feb 5, 2017)

GregV said:


> 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.
> 
> ...


5 or 10, I don't know enough to know. My thought is that 5 frames is very tight and would encourage swarming a lot more. Also, 5 frames is small and tight and easy for the bees to control warmth. On the other hand, it might be too small to have the resources? It really depends on how many hives survive the winter and if I have enough hives to play with. If several hives survive, I'll try both.


----------



## GregB (Dec 26, 2017)

AR1 said:


> 5 or 10, I don't know enough to know. My thought is that 5 frames is very tight and would encourage swarming a lot more. Also, 5 frames is small and tight and easy for the bees to control warmth. On the other hand, it might be too small to have the resources? It really depends on how many hives survive the winter and if I have enough hives to play with. If several hives survive, I'll try both.


40 litre volume has more flexibility built-in, so that you can run it as compressed 5-6 frames if desired OR expand to the max if needed.
Yet you can pack many more bees onto a small number of frames and still keep them inside - a very good thing.
Yet you have some spill over space so to reduce a chance to ACTUALLY loosing a swarm. 
As they explode in spring, the 40 litre volume is not much more to slow you down (in fact, for you and me and our terrible springs - we don't want them to jump out of the box too early).


----------

