
Observations on Pollen Subs:
Half 3
Placement of the Pollen Patty
First printed in ABJ October 2021
Randy Oliver
ScientificBeekeeping.com
There doesn’t look like a profit from feeding pollen sub when there’s already a considerable pure pollen move occurring. Alternatively, throughout a dearth, pollen subs might carry out higher when the foragers are nonetheless ready to herald a small quantity of pure pollen. The query then is, why would that small quantity of pollen make a distinction?
A variety of area trials of pollen subs have proven that if colonies are fed pollen sub when there’s a pure pollen move occurring, that there could also be little profit from offering a synthetic weight loss program [[1],[2]], and even patties of pure pollen [[3]]. However because the outcomes of this trial present, many people within the West acquire nice profit from feeding pollen sub throughout sure intervals of pollen dearth, corresponding to in autumn and early spring.
However “pollen dearth” is a relative time period. I used to be talking with a fellow beekeeper the opposite day about feeding pollen sub, and my ears perked up when he famous that his colonies reply to pollen sub a lot better when there may be additionally a light-weight pure pollen move occurring — one thing that I’ve additionally seen. So what precisely does that small quantity of pure pollen coming right into a hive present when there may be already a surplus of synthetic weight loss program being fed?
Entrepreneurs have achieved an ideal job at convincing us that “pure” is at all times higher. So it’s tempting to leap to the conclusion that the little little bit of incoming pure pollen provids some lacking vital dietary parts. However within the case of my pal, he already contains 5% pure pollen within the patties that he feeds, which would supply these lacking parts.
Sensible utility: On this area trial, regardless of feeding as much as 12 kilos of pollen sub formulations containing 15-20% pure pollen, these colonies nonetheless barely grew throughout the excessive pollen dearth over the course of the trial.
So I think that one thing else apart from some lacking “pure” nutrient is concerned. Maybe we must always look someplace apart from at vitamin alone, and as an alternative take into consideration bee habits in response to environmental cues. Merely offering a natural-pollen-rich patty to a colony doesn’t be sure that the colony will reply by increasing the broodnest. However the addition of somewhat little bit of incoming pure pollen might kick the colony into broodnest growth mode. Maybe we must always strive to determine what cue that pollen introduced in by the foragers offers to set off the colony to develop brood rearing in response.
THE NURSES RUN THE SHOW
The query to ask might then be, who makes the choice to develop the broodnest? Queens typically proceed to put loads of eggs throughout a pollen dearth, however are adopted by nurse bees who, when unable to provide sufficient jelly, devour these eggs to recycle the protein, asnicely defined by Heather Mattila [[4]]:
It appeared as if the queen was regularly laying eggs throughout the cluster in early spring, however whether or not the eggs have been reared to the sealed brood stage trusted the provision of pollen to the employees within the colony. This speculation is supported by the observations of Imdorf et al. (1998), who reported the presence of eggs in pollen-stressed colonies that lacked open brood, and by Schmickl et al. (2003), who noticed that the queen didn’t regulate her price of egg laying once they created synthetic pollen shortages in an commentary hive, regardless of giant adjustments in brood demography. It is sensible that eggs would proceed to be laid by the queen when colonies have restricted entry to pollen as a result of their presence within the colony represents comparatively little dietary funding, however offers a possibility to quickly recommence brood rearing as soon as favorable situations return.
This culling and recycling of eggs and larvae by the nurses has been termed “brood trimming” by Larry Connor[[5]], and demonstrates that it’s the nurses, relatively than the queen who really run the present, since they’re those that produce jelly — the forex of protein for your entire colony. It’s the nurses who resolve who will get fed, and who doesn’t. In late summer season, my sons and I monitor our colonies for the quantity of jelly that the nurses place round younger larvae. Again and again we observe an instantaneous improve within the quantity of jelly after we present pollen sub. However that improve remains to be not the identical as when the foragers are bringing in loads of pollen. Might the distinction in response be because of the place the pollen or patty is positioned?
THE PLACEMENT OF THE POLLEN
In my current dialog with my beekeeping buddy, he mentioned that he has best success with feeding delicate bulk patty, relatively than preformed patties in waxed paper. I observe the identical (Determine 1).
Fig. 1 A variety of us Western beekeepers want to feed chunks of “delicate” patty that can squish between the frames, particularly upward between the underside bars. Might it’s that offering the feed nearer to the cells of brood makes a distinction?
THE PROBLEM WITH “SURVIVORSHIP BIAS”
We beekeepers are accustomed to the look of the textbook brood body, having an ellipse of brood within the decrease center, a band of beebread across the brood, after which honey above and to the edges (Determine 2).
Fig 2. A textbook instance of how honey, beebread, and brood are organized on a comb. However no bee has ever seen this view (it’s darkish contained in the hive). And simply because we generally see beebread in an arc round the brood, that doesn’t imply that that’s the pollen the nurses take note of.
The situation of the band of beebread is an ideal instance of “survivorship bias,” resulting in the inaccurate conclusion that pollen foragers preferentially retailer incoming pollen there. In actuality, that band of beebread represents the leftovers — surplus pollen that was not instantly consumed by the nurses. In a well-nourished, rapidly-growing colony on a average pollen move, there might not be any beebread displaying across the brood in any respect.
Sensible utility: On this case it’s not what we people see that’s vital to the nurses, it’s what we don’t see. In Determine 2 discover the pollen scattered between the cells containing eggs or sealed brood. That’s the pollen that might be first consumed by the nurses. Any pollen within the “pollen band” is surplus past the wants of the nurses, and is fermented into beebread for storage.
The basic instance of survivorship bias occurred throughout World Battle II, when the Air Pressure recorded the places of bullet holes within the bottoms of returned plane (Determine 3).
Fig. 3 Mechanic’s notations of the place bullet holes have been present in planes after bombing missions over Germany. The brass interpreted that to imply that these have been the areas calling for reinforcement. However mathematician Abraham Wald identified the error of survivor bias, since they have been solely recording the presence of bullet holes in planes that made it dwelling (the survivors). The vital locations to armor have been the areas missing bullet holes. Picture from Wikipedia.
It’s the identical factor with the pollen band on a brood body. A rapidly-growing colony might not include any pollen band in any respect if the nurses are consuming each little bit of incoming pollen. The one factor {that a} pollen band signifies is that there’s a non permanent surplus of pollen within the hive, left over after the nurses have already consumed their fill.
THE CUE TO THE NURSES THAT A POLLEN FLOW IS OCCURRING
There are a number of very fascinating research on the interplay between the place incoming pollen foragers place their hundreds, and the place nurse bees search for pollen to devour.
In 1969, Doull discovered that the efficient olfactory vary of the nurse bees was lower than 2 inches [[6]]. After which in 1974 [[7]], utilizing an elegantly easy experimental design, he demonstrated the profound impact of distance from brood for nurse bee response to provided patty of pure pollen (Determine 4).
Fig. 4 Word how the response of nurse bees to positioned pollen quickly drops off at even very brief distance. (Graph plotted from knowledge from Doull).
Wow! Even two inches of distance makes an enormous distinction to the nurses, who want to seek out pollen within the darkness of the hive by their sense of scent.
Doull instructed:
These findings have some implications in using pollen for feeding to bees within the hive. Pollen saved across the brood would usually be throughout the vital distance, or can be positioned by bees after solely a brief interval of random motion. Pollen saved in outer combs of the hive, nonetheless, can be accessible solely to these bees that moved away from the brood space. It might be that, in the primary, pollen saved in outer combs might be used solely when the brood nest expands to combs close to it.
Throughout a considerable pollen move, one can typically observe how the foragers place pollen proper in the midst of the brood space (Determine 5).
Fig. 5 This pollen positioned adjoining to hungry larvae might properly disappear in a single day, as it’s preferentially consumed by the nurse bees. The convenience with which the nurses discover pollen seems to be the cue that there’s a pollen move occurring, indicating that the colony can go into buildup mode.
Word that the pollen in Determine 5 is positioned adjoining to older larvae about to be capped. In 1973 Steve Taber [[8]] had issues to say about this:
The information point out that there’s a pollen consumption space. Because the eggs hatch, the rising larvae are fed increasingly more. Queens generally tend to put eggs in adjoining cells if these can be found, relatively than laying in a random style all through the brood space. Because the teams of larvae mature, the consumption of pollen positioned close to larvae aged 5 and 6 days would most likely be so fast that no incoming pollen would or might accumulate in that a part of the hive.
My bees are maintained beneath intensive administration for queen and drone rearing and are given greater than 40 kg of pollen per colony every year. Subsequently, after I hear that bees won’t eat the pollen they’re provided, I conclude solely that the pollen was positioned unsuitably within the hive. Except positioned inside 4-6 cm of the brood space, the pollen might be taken very slowly or by no means …
The normally accepted speculation regarding pollen in combs is that the pollen is saved and consumed in a band across the brood space (Farrar, 1963). My speculation is that the belt or band of pollen which surrounds the brood nest in periods of pollen surplus is an space of non-consumption. Since pollen is consumed within the areas of unsealed brood, most new incoming pollen is consumed there. Thus, there may be little or no probability of any pollen accumulating contained in the brood space.
In 1977 Jerzy Woyke studied brood cannibalism [[9]], noting the upper price of cannibalism of younger larvae in autumn over that in summer season.
The upper survival price of brood when the pollen provide was good, and in colonies to which pollen combs have been added … means that scarcity of pollen releases cannibalism within the honeybees … Pollen could also be out there in a single comb, and never in one other, and even in one other a part of the identical comb. So the bees with no pollen close by, eat brood as an alternative, and are thus capable of produce meals for the queen and for larvae.
One in every of my favourite researchers, Scott Camazine, in 1991 rejected the “blueprint speculation” and as an alternative laid out an exquisite clarification (self-organizing sample formation) of how particular person bees working at nighttime create the textbook association of brood combs, with none bee being conscious of a “blueprint” [[10]]:
Every day honey and pollen from tens of 1000’s of foraging journeys are deposited into the cells, shops of honey and pollen cells are regularly consumed, tons of of eggs are laid, mature grownup bees emerge from their cells, and diseased brood is eliminated. Regardless of the common turnover by which cells are sometimes refilled with one thing apart from what they just lately held, the soundness of the sample is maintained … there isn’t a blueprint to specify places for brood, pollen and honey; the dynamic relationships among the many element processes of deposition and elimination are ample to arrange a sample on the combs.
Camazine introduced up the idea of “interface zones,” which have been an aha idea for me for hive inspections and studying to “learn the combs” [[11]].
Sensible purposes:
- Contemporary pollen, randomly positioned by returning foragers within the cells all through the broodnest, permits the nurses to provide ample jelly.
- The convenience with which the nurses discover recent pollen subsequent to the brood that they’re feeding cues them to feed, relatively than devour younger larvae, thus placing the colony into “buildup mode.”
- The presence of a pollen band signifies that there’s a surplus of incoming pollen. The inside fringe of that band signifies the gap from the brood that the nurse bees wanted (or have been keen) to journey till they bought their fill.
- All of the above findings point out the significance of putting supplemental pollen or synthetic weight loss program as near cells of brood as doable.
- Contemporary nectar can be positioned all through the broodnest by mid-age nectar receivers, once more offering the nurses with vital sugar, and the massive quantity of water vital to provide jelly (I’ll focus on this in my subsequent article).
BACK TO MY TRIAL
The colonies in my trial, regardless of being supplied pollen subs advert libitum, typically didn’t develop throughout the late summer season and fall. However this isn’t to say that they weren’t regularly rearing brood – simply to not sufficient extent to develop their populations (Determine 6).
Fig. 6 This photograph of a brood comb was taken on 15 October, from a weak colony (3-frame cluster measurement) that had been utterly consuming its pollen sub. As you possibly can see, the larvae have been in a superb sample, and well-fed with jelly. However be aware that though there are just a few cells containing pure pollen on the periphery; there may be not a single cell with pollen in the midst of the brood space.
Sensible utility: So might the good thing about even a minimal pollen move whereas feeding pollen sub be because of it being positioned by the foragers into the center of the brood space, even when the beekeeper won’t discover it as a result of it’s being so quickly consumed? Might it’s that that little little bit of pollen proper subsequent to the brood offers the cue for the nurse bees to permit for growth of the broodnest, after which additional devour patty provided on the periphery of the brood space? As researchers in search of funding are fond of claiming, this clearly requires extra analysis.
This brings to thoughts an experiment that I carried out in 2010. I fed fluorescent-labeled pollen sub to double-deep hives, putting the patties on the highest bars of both the decrease or higher brood chamber. After the bees had fed for a few days, I froze samples of fifty bees from every comb, crushed their guts out, positioned them beneath UV gentle, and recorded the proportion that contained fluorescent pigment (Determine 7).
Fig. 7 To my nice dismay, a bear ripped the lid off the freezer off earlier than I might course of all of the samples, however listed below are two consultant outcomes, with the colony fed between the brood chambers to the left, and that fed under the hive cowl to the fitting. H, P, or B point out honey, pollen, or brood within the comb, respectively. Word that nurse bees that had consumed the labeled pollen sub have been current all through the hive, and that extra of them had achieved so when fed in the midst of the cluster.
Let’s return to my 2020 area trial. It occurred to me that since we had recorded the variety of frames of bees (FOB) individually for the decrease and higher brood chambers, that I might see whether or not there was a correlation between the placement of the cluster relative to the position of the pollen sub (because it was at all times positioned on the highest bars of the decrease brood chamber), and the expansion of the colony (indicated by cluster measurement) (Determine 8).
Fig. 8 Let me stroll you thru this. Colony energy is on the y-axis — the upper the information level, the stronger the colony. Cluster location is on the x-axis — these colonies with the cluster being under the patty being to the left, these with the cluster above the patty to the fitting, and people with the cluster (and presumably the broodnest) centered on the patty lifeless middle.
Word that over time, the strongest colonies (thus that grew probably the most) tended to be these by which the cluster was evenly centered above and under the pollen sub patties. This in fact is barely a correlation, nevertheless it positive makes me marvel in regards to the significance of precisely the place supplemental protein weight loss program needs to be provided to the colony. And likewise whether or not the rationale that delicate patty squished between the frames seems to be extra stimulatory.
Sensible utility: Might this be telling us one thing? Within the near-complete absence of a pure pollen move, maybe it’s vital to put pollen sub nearer to the brood. Or possibly present an extra synthetic pollen move?
BACK TO DRY FEEDING
Maybe we’d have gotten higher outcomes if we had concurrently fed somewhat dry pollen sub for the foragers to put within the brood space. In earlier analysis, I’ve discovered that bees enthusiastically forage upon provided dry powdered pollen sub provided within the apiary (Determine 9).
Fig. 9 There are a variety of feeders out there to maintain the powdered sub dry. Once I’ve calculated consumption charges, the typical per-colony gathering price of dry sub provided in an apiary feeding station could be equal to feeding a 1-lb. patty per week to every hive.
And after I provided foragers fluorescent-labeled dry sub for a single hour, after which inspected combs of brood, it was clear that the returning foragers packed their corbicular a great deal of dry pollen sub straight adjoining to cells of larvae (Determine 10).
Fig. 10 Underneath black gentle, it’s straightforward to see the bluish-white larvae, and the inexperienced fluorescent pollen sub saved after a single hour of providing it within the apiary. This photograph confirms Camazine’s observations that pollen foragers randomly place their hundreds all through the broodnest.
I hope to experiment additional with dry-fed sub concurrent with patty-fed. Dry fed can definitely stimulate jelly manufacturing, however I discovered that it was much less constant colony-to-colony than patty fed. Not solely that, however some industrial beekeepers report unfavorable outcomes with dry-fed sub within the fall, supported by an experiment that I ran with some USDA researchers [[12]].
Sensible utility: I think that we nonetheless have a lot to study in regards to the optimum method to feed synthetic diets to colonies. There may be in fact the tradeoff between practicality and labor prices vs. profit — it’s tough to put sub into particular person cells within the broodnest with a purpose to stimulate the nurses. Maybe patties together with dry-fed may do the trick. Or another resolution — please let me know in the event you give you one!
COMING NEXT
In any case, there may be but yet another issue presumably concerned in why our colonies grew so poorly throughout the trial. Subsequent month I’ll look into the significance of nectar, moisture, and humidity as limiting components for increasing the broodnest.
REFERENCES
[1] https://scientificbeekeeping.com/when-to-feed-pollen-sub/
[2] Lamontagne-Drolet, M, et al. (2019) The impacts of two protein dietary supplements on industrial honey bee (Apis mellifera L.) colonies. Journal of Apicultural Analysis 58(5): 800-813.
[3] Mattila, H & G Otis (2007) Dwindling pollen assets set off the transition to broodless populations of long-lived honeybees every autumn. Ecological Entomology 32: 496–505.
[4] Mattila, HR & GW Otis (2006) Affect of pollen weight loss program in spring on growth of honey bee (Hymenoptera: apidae) colonies. J. Econ. Entomol. 99: 604-613.
[5] Connor, L (2008) Bee Intercourse Necessities. Wicwas Press (p. 40).
[6] Keith M. Doull & L. N. Standifer (1969) A way for measuring feeding responses of honeybees of their hive. Journal of Apicultural Analysis 8(3): 153-157.
[7] DOULL, Ok M (1974) Impact of distance on the attraction of pollen to honeybees within the hive. Journal of Apicultural Analysis 13(1): 27–23.
[8] Taber, S (1973) Affect of pollen location within the hive on its utilization by the honeybee colony. J. Apic. Res. 12(1): 17-20.
[9] J. Woyke (1977) Cannibalism and brood-rearing effectivity within the honeybee. Journal of Apicultural Analysis 16(2): 84-94.
[10] Camazine, S (1991) Self-organizing sample formation on the combs of honey bee colonies. Behav Ecol Sociobiol 28:61-76.
[11] https://scientificbeekeeping.com/first-year-care-for-your-nuc/ (seek for the phrase “interface”).
[12] Maes, P, et al. (2016) Weight-reduction plan-related intestine bacterial dysbiosis correlates with impaired growth, elevated mortality and Nosema illness within the honeybee (Apis mellifera). Molecular Ecology 25(21): 5439-5450.
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