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A Comparative Trial of the Pollen Subs: Half 6- Do We Have to Revise De Groot?

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A COMPARATIVE TRIAL OF THE POLLEN SUBS

Half 6: DO WE NEED TO REVISE DE GROOT?

Beekeeper-Funded Analysis

First Printed in ABJ December 2021

Randy Oliver
ScientificBeekeeping.com

Bee nutritionists have lengthy cited the important amino acid ratios instructed by de Groot in 1953. However he himself identified the shortcomings of his strategies and certified his suggestions. I really feel that after almost seventy years, it might be time to meet up with the opposite livestock industries, and replace our data on what constitutes an optimum supplemental eating regimen for honey bee colony progress.

De Groot understood that what actually counted for bee diets meant for colony progress could be the necessities for the nurse bees producing the jelly essential to feed the larvae, queen, and older employees. However that’s not what he investigated. This has bugged me for years — that few researchers aside from Elton Herbert have adopted up on measuring the expansion of colonies (as opposed to a couple caged employees) fed synthetic diets, which might be the perfect and best technique to decide whether or not de Groot’s important amino acid (EAA) ratios for grownup (non-nurse) bee growth could be the identical as for total colony growth, which is what we beekeepers would need in our pollen subs. On this final article of this collection, I’m going to stay my neck out and make a stab at revising de Groot’s ratios and recommend the kind of utilized analysis that our trade deserves.

Enable me to start by looking how the amino acids in a eating regimen are literally utilized by an organism.

HOW PROTEIN AND AMINO ACIDS ARE USED

The physique tissues of grownup bees are largely composed of protein — proteins represent about 50% of a bee’s dry weight [[1]]. The varied kinds of physique and haemolymph proteins are all created from the metabolic pool of amino acids derived from their eating regimen, as illustrated by Determine 1 (printed in the course of the years through which de Groot was performing his analysis).

 

Fig. 1 Dr. Harold Mitchell was a pioneering animal nutritionist on the College of Illinois. In 1950 he printed this stream chart of how dietary proteins had been utilized by the physique. Picture from [[2]].

The protein in a eating regimen is first digested into its constituent amino acids, from which the physique then synthesizes new physique elements by the method of anabolism (the other is catabolism — the “harmful metabolism” of physique tissue). That course of is proscribed largely by the quantity of important amino acids (EAAs) within the eating regimen [[3]], in addition to their relative proportions to one another. The EAAs are used not just for constructing (or changing) muscle mass and inside organs, but in addition to create haemolymph and dispensable storage proteins. Some protein is interspersed in chitin to kind the bees’ cuticle and wings. And a few could also be burned as gas for vitality if sufficient carbohydrates (sugars) should not out there within the eating regimen.

One technique to measure bee progress is by drying and weighing particular person bees. However such weight achieve contains elements largely constructed from carbohydrates, similar to blood sugars, low-proteinaceous chitin, and lipids. With regard to synthetic diets, what we’re extra involved with is the utilization of the nitrogen within the eating regimen, which comes from the protein.

Sensible software: So maybe what we actually ought to focus upon is their improve in nitrogen content material.This consideration didn’t escape de Groot, so he measured each dry weight achieve in addition to nitrogen improve (Determine 2).

Fig 2. Evaluate the consequences of EAA deficiencies upon weight achieve (above) versus achieve in nitrogen (beneath), which signifies the precise incorporation of these EAAs into physique mass. Observe extreme weight reduction in grownup bees if leucine, threonine, valine are poor. However the graph on the proper signifies that the burden loss was because of the important requirement of these EAAs for the method of constructing non-proteinaceous physique tissues, reasonably than being included into these tissues.

Sensible software: We are able to’t simply take a look at protein analyses of physique tissues estimated from nitrogen content material; we should additionally needless to say some EAAs are important for the constructing of non-nitrogenous physique elements. I’ll return to this idea once I talk about “carcass evaluation.”

NUTRIENT REQUIREMENTS CHANGE WITH STAGE OF DEVELOPMENT

At every stage of growth, a employee honey bee is anabolizing totally different physique elements, maybe consuming totally different meals, and will require totally different proportions of EAAs:

  1. Larval growth — Main weight achieve consisting of a small quantity of integument, muscle, digestive tract, and tracheal system, however principally from giant quantities of storage protein in fats our bodies [[4]]. Protein supply — nurse-fed jelly.
  2. Pupal-to-adult growth — Improvement of the chitinous exoskeleton (together with wings, legs, and eyes), grownup organs, and physique musculature. Protein supply — the fats our bodies of the pupa, and the repurposing of pupal tissue. Which means that the pupa is its personal protein supply, in order that evaluation of the amino acid ratio of its physique could also be informative.
  3. Publish-emergence growth — Main growth of the hypopharyngeal glands and fats our bodies, Partial growth of the wing muscle tissues and the sting. Protein supply — pollen and a few jelly.
  4. Nurse bee growth — The nurses want to provide copious portions of protein-rich jelly for feeding the larvae and different members of the colony. Protein supply — pollen.
  5. Mid-age employee growth — Improvement of the wax glands and wing muscle tissues. Protein supply —pollen, jelly, and autophagy of the fats our bodies and hypopharyngeal glands as they reduce weight in preparation for foraging [[5]].
  6. Forager growth — Remaining progress and upkeep of the wing muscle tissues and physique tissues. Protein supply — jelly begged from nurses.
  7. “Winter bee” growth — The diutinus state taken by employees that emerge throughout pollen dearths may be thought-about as “arrested growth” of nursing physiology. Protein supply — pollen.
  8. “Winter bee” upkeep — Little or no protein is required. Protein supply — a small quantity from honey, saved beebread, autophagy (“self-consumption” of storage proteins within the fats our bodies) [[6]].

Sensible software: Much like the totally different diets bought for the optimum growth of broiler chickens at every stage of progress, we might discover that there is no such thing as a one-size-fits-all eating regimen for honey bees.

WINTER FEEDING

AUTUMN FEEDING FOR STORAGE PROTEINS

Earlier than I return to eating regimen for colony growth, let’s first take into account diets in preparation for, or throughout winter. Beekeepers feed pollen subs not solely to supply the mandatory vitamins for the final rounds of brood that can develop into long-lived “winter bees,” but in addition to permit these emerged employees to completely develop their storage proteins as a way to survive the winter. Would the event of storage proteins require a special EAA ratio within the eating regimen?

As identified by de Groot:

Many sorts of bugs can keep life for appreciable durations within the absence of nitrogenous meals.

They do that by digesting their physique protein reserves (catabolism). “Winter bees” exhibiting diutinus physiology (just like nurses, however not producing jelly), as talked about earlier, break down their protein reserves as a way to get hold of the amino acids (and beneath situations of hunger, vitality) vital for “physique upkeep.”

Sensible software: De Groot studied the EAA necessities of younger employees not being uncovered to brood pheromone, suggesting that they may have gone into diutinus physiology. Thus, his beneficial EAA ratios might properly apply to diets fed to the final spherical of rising employees after broodrearing ceases in autumn.

Many beekeepers feed pollen subs in autumn as a way to put together the “winter bees” for longevity and late-winter brood rearing. In contrast to as with forager-collected pure pollen or dry-fed pollen sub, diets fed in patty kind contained in the hive should not saved within the combs [[7], [8]], so the good thing about autumn feeding of pollen patties might principally be to advertise the synthesis of these storage proteins. That stated, these storage proteins permit for under a really restricted quantity of brood rearing. Haydak’s [[9]] findings recommend that it requires the storage proteins of round 30 employee bees to rear a single substitute employee to maturity (at which level that employee would nonetheless have to eat further protein as a way to full its growth).

Most beekeepers have heard of the essential storage protein vitellogenin. Of curiosity is that Otis [[10]] discovered that winter bees saved extra of a hexamerin protein (presumably arylphorin) than vitellogenin. In addition they confirmed {that a} “winter bee” saved solely a fraction of the protein essential to feed even a single larva.

So which EAAs are essential for the formation of those hexamerin storage proteins? These wealthy within the fragrant amino acids (phenylalanine, tyrosine, and tryptophan) are known as arylphorins, whereas one other group of hexamerins are often known as methionine-rich storage proteins. Frustratingly, I used to be unable to search out info on the EAAs in vitellogenin. The extraordinarily excessive proportion of tyrosine in insect storage proteins [[11]] means that we’d wish to take note of this “conditionally important” amino acid in diets meant for colony growth or “winter bees.”

Sensible software: The findings of Haydak, Jeffree, and Otis level out that the flexibility of colonies to rear brood throughout winter is poorly understood. We clearly want extra analysis on this topic.

WINTER FEEDING FOR BODY MAINTENANCE

De Groot discovered that based mostly upon his preliminary analysis:

From the above outcomes it might be concluded, that in distinction with findings of different authors the longevity of outdated bees is elevated by supplementing the carbohydrate eating regimen with numerous protein containing proteins or pure protein in an appropriate focus. Clearly within the bee colony the necessity for protein just isn’t restricted to the brood rearing interval however proteins are required for upkeep of grownup bees as properly. This discovering is in putting distinction with the widespread opinion, that in the course of the winter interval the bee requires solely carbohydrates. It is not uncommon use in beekeeping apply to substitute the honey for shops of pure sugar for overwintering bee colonies, which isn’t thought-about to be disadvantageous from the dietary perspective. Nonetheless, honeys include small quantities of protein various within the area the place we discovered the protein containing materials to exert its longevity selling motion on outdated bees. … Due to this fact it’s apparent to recommend that sucrose shops enriched with an appropriate quantity of protein meals with a excessive organic worth could also be extra favorable for overwintering bee colonies than [the] shops of pure carbohydrate typically used at current in apiculture.

Winter feeding with straight sugar remains to be a standard apply as we speak (Determine 3).

Fig. 3 A wintering colony being fed dry granulated sugar. This methodology was described by Johansson in 1977 [[12]], and later popularized on Beesource because the “mountaincamp methodology,” after posts by a beekeeper from Mountain Camp Farm in New York. Observe that Johansson cautioned that colonies fed with dry sugar might have to forage for water to dilute the sugar. Photograph credit score: Phillip Cairns/ mudsongs.org

Sensible software: Including some high-quality protein to sugar fed throughout winter would possible assist the “winter bees” to take care of their physique reserves. A number of producers are providing protein-enriched “winter patties.” We may positive use some easily-performed subject trials to see whether or not enhancing that protein with the sure amino acids [[13]] could be of profit.

WHICH EAA RATIO IS BEST FOR COLONY DEVELOPMENT?

Winter feeding apart, most of us feed pollen sub to permit our colonies to develop. Jeffree [[14]] discovered that colonies in Scotland enter winter with out sufficient saved beebread to permit nurse bees to feed greater than a minimal quantity of larvae, but sporadically rear larvae all through the winter. Thus late-winter feeding of pollen sub may be of nice profit for these beekeepers who require robust colonies for early pollination contracts.

Sensible software: We almond pollinators have discovered that midwinter feeding of pollen sub beneath the correct situations can encourage severe brood rearing and colony buildup.

AT WHAT DEVELOPMENTAL STAGE DO BEES MOST REQUIRE PROTEIN?

So the query is, at what stage of growth does a employee bee truly require essentially the most protein in its eating regimen? It’s throughout their larval stage (Determine 4).

Fig. 4 Improve in nitrogen content material of the our bodies of employee bees throughout progress durations. Two-thirds of the incorporation of amino acids right into a bee’s physique happens when they’re consuming the jelly produced by nurse bees, versus the 1/3 added from the pollen that it consumes after emergence. Information from [[15], [16]].

De Groot’s estimates represented the EAA necessities for newly-emerged non-nursing grownup employees (which can have developed their ovaries because of the lack of suppressive queen pheromone [[17]]). The graph above signifies that the protein requirement of the larvae is extra essential so far as colony progress. However the larvae profit from pollen sub not directly — it’s the nurse bees that truly eat the pollen sub and convert it to jelly (any pollen that the larvae would possibly obtain from the nurses gives an insignificant quantity of protein total [[18]]).

Thus de Groot’s estimated ratios might not apply to the all-important nurse bees, that are chargeable for the digestion of proteins for your entire colony (and are thus the customers of pollen sub). The nurses then convert pollen sub into the jelly to feed the queen, the brood, and the remainder of the employees (jelly being the “forex of protein” throughout the colony).

Sensible software: So for colony progress, what we actually want to find out is the optimum EAA ratio within the eating regimen of nurse bees feeding larvae.

WAYS TO DETERMINE OPTIMAL EAA REQUIREMENTS

Though researchers have now developed strategies for efficiently rearing employee larvae in vitro, the eating regimen consists primarily of harvested royal jelly [[19]]. Till a very synthetic eating regimen for larvae is developed, we have to stick with determining diets that permit nurse bees to provide essentially the most jelly.

There are a number of doable methods for researchers to take action. Enable me to record another benchmark indicators to which we are able to evaluate de Groot’s suggestions. I’ll graph them out after explaining them.

COMPARE TO ROYAL JELLY

It could be cheap to imagine that royal jelly produced by nurse bees would include the perfect ratio of EAAs for optimum larval progress. I downloaded a number of analyses of the amino acids in royal jelly, and located that the EAA ratios had been remarkably constant, regardless of being from totally different nations and the consumption of various pollens. A number of the older analyses lacked knowledge for tryptophan. Happily, in 2009 a brand new methodology was developed to quantify tryptophan [ [20]], so I used to be capable of insert a price. I averaged the 5 most constant analyses for graphing functions.

Aspect observe: An fascinating examine by Webster [[21]] discovered that nurse bees that cannibalize brood quickly recycle the amino acids of these larvae again into royal jelly, and to a lesser extent into physique tissue. The take-home is that bees don’t waste amino acids, and that open brood can function a protein reserve for producing jelly.

COMPARE TO KNOWN NUTRITIOUS NATURAL POLLENS

Research from the Tucson Lab within the Nineteen Seventies verify beekeeper observations that colonies develop very properly on almond pollen [[22]], so we are able to possible use the EAA ratio of almond pollen or different identified nutritious pollens (similar to mustard or candy clover) as benchmarks.

COMPARSION TO OTHER ANIMALS AND WHAT THEY FEED THEIR OFFSPRING

Much like royal jelly being the “excellent” meals for queens and larvae, cows and chickens additionally produce proteins which can be excellent meals for his or her offspring — casein and albumin, respectively. Once more, each of these proteins are the results of evolutionary choice for an EAA composition that might permit for full growth of their offspring. Maybe not surprisingly, each work very properly as protein sources for honey bees [[23],­­­­[24]].

CARCASS ANALYSIS

One other doable indicator of EAA necessities for a eating regimen are the EAAs current within the animal’s physique. That is termed “carcass evaluation” [[25]], and will surely be a easy technique to decide the perfect ratio. Sadly, EAA necessities change with an animal’s stage of progress [[26], [27]], and whether or not the EAAs are literally included into the physique, or merely concerned within the strategy of tissue synthesis.

That stated, Ghosh [[28]] gives amino acid analyses for employee bee larvae, pupae, and adults (Determine 5).

 

Fig. 5 Comparability of the EAA ratios of the three developmental levels of honey bee employees, in comparison with these decided by de Groot. (I doubled de Groot’s ratio, since his EAA combination solely constituted 25.6% of the protein within the eating regimen, whereas it ought to possible have been ~50%.) Of curiosity is how the concentrations of leucine, phenylalanine, and threonine drop considerably upon pupation (after the larvae have defecated), indicating that these amino acids had been vital for tissue formation solely, reasonably than being included into the ensuing physique tissue.

Notes for Ghosh’s findings:

  • Non-essential tyrosine and cysteine might substitute for important phenylalanine and methionine (respectively) if they’re briefly provide. For some cause Ghosh didn’t measure methionine, whereas de Groot didn’t measure cysteine or tyrosine, so the comparative columns for these AAs are questionable. In lots of present EAA analyses, authors mix the 2 pairs of complimentary amino acids, as do I.
  • The grownup bees analyzed had been in-hive “home bees” (wings eliminated) that had the chance to feed upon pollen within the combs, so their EAA adjustments from pupae had been possible because of the growth of grownup tissues after feeding upon pollen.
  • Tryptophan is required for tissue progress, however not a lot for upkeep. Ghosh’s evaluation was unable to detect tryptophan in larvae or pupae, and it barely registered within the adults — that is shocking, since de Groot decided that tryptophan ought to account for a full 4% of the EAAs. And bear in mind Determine 2, which signifies that the necessities for leucine, threonine, and valine could also be underestimated by carcass nitrogen analyses alone.

From the above graph, it seems that for larval progress, in comparison with grownup weight achieve, larvae require a bit extra isoleucine, lysine, phenylalanine, and threonine, and rather less valine than do grownup bees. Of curiosity is that Ghosh was not capable of detect tryptophan in larvae or pupae, and solely 0.03% in adults — additional cause to not normalize EAA ratios to tryptophan.

Sensible software: De Groot’s ratios for grownup bees look like near these required for larval growth, however the variations point out that the optimum EAA ratios for a super eating regimen for colony progresswould possibly have to be barely tweaked.

I’ve since engaged in reviewing a considerable amount of analysis on the amino acid composition of pure pollens famous as being of excessive dietary worth to bees, analyses of royal jelly, and the amino acid necessities of different bugs and animals. I’ve come to the conclusion that de Groot’s experimentally-determined ratios ought to be checked out in comparison with different findings, similar to the truth that bees develop very properly on the proteins casein (from milk) [[29]], albumen (from hen eggs) [[30], [31]], royal jelly, and a few pollen sorts. So I in contrast the EAA ratios of the above protein sources, in addition to the EAA ratio in bee brood itself within the chart beneath (Determine 6).

Fig. 6 De Groot’s suggestions, in comparison with carcass analyses of employee developmental levels, royal jelly, pollen, hen egg and cow milk proteins (each of which have been proven to help bee progress). I’ve circled indicators of the place de Groot’s EAA ratios might have to be adjusted for colony progress versus emerged employee improve in nitrogen content material [[32]]. I’ve additionally (in yellow) inserted a instructed revision of de Groot’s beneficial ratios. Observe that not all analyses mixed the fragrant amino acids.

Sensible software: The above chart means that de Groot was definitely not far off the mark. Nonetheless, for optimum egg-to-adult employee progress, his instructed ratios could also be a bit excessive for isoleucine and valine, and low for lysine and phenylalanine. We clearly want additional experimental outcomes to verify. However permit me to be daring sufficient to recommend a revision (Desk 1):

Desk 1. Urged revisions (in purple) to de Groot’s suggestions for the proportional ratios of the EAAs for diets meant for colony progress.
de Groot Revised? Revised normalized to leucine
Arginine 3 3.25 0.72
Histidine 1.5 1.5 0.33
Isoleucine 4 3 0.67
Leucine 4.5 4.5 1.00
Lysine 3 4.5 1.00
Methionine & cysteine 1.5 1.5 0.33
Phenylalanine + tyrosine 2.5 4 0.89
Threonine 3 3 0.67
Tryptophan 1 1 0.22
Valine 4 3.5 0.78

 

OK, that’s a bunch of numbers, however I think that what you’d actually wish to know is how do my instructed ratios evaluate to these of the top-performing pollen subs in my trial? See Determine 7.

Fig. 7 A comparability of the EAA ratios, normalized to leucine, from de Groot’s suggestion, my suggestion, and the 2 top-performing examined pollen subs. Observe how carefully my instructed ratios (yellow) match these of the highest performers.

CAGE AND SEMI-FIELD STUDIES

In-lab cage research are restricted by the issue of sustaining the mandatory jelly-producing nurse bees and rearing the produced brood to maturity for a number of generations on synthetic diets. I’m in full settlement with Hendricksma [[33]] that:

We advise that colony supplementation ought to goal nurse bee vitamin. … Our examine additionally highlights the significance of testing honey bee vitamin with subject and laboratory research. Testing eating regimen therapies on caged bees as in contrast with entire colonies might yield totally different insights.

Sensible software: The gold customary semi-field bioassay for testing the dietary effectivity of honey bee diets is on the colony stage through the use of nucs with free-flight entry in screened cages (4x4x8 ft), developed on the Beltsville Lab [[34]]. I strongly encourage extra analysis on synthetic diets utilizing this methodology.

Options for researchers: That stated, I’m curious as as to whether a modification of Haydak’s methodology of extending nurse longevity by eradicating sealed brood [[35]], or the swarm field methodology for producing a number of rounds of queen cells from a cohort of nurse bees may very well be used [[36]].

BASIC FORMULA FOR HONEY BEE ARTIFICIAL DIETS

Based mostly upon a assessment of the literature, plus the findings from our subject trial, one can arrive at instructed proportions for an optimum synthetic eating regimen for exciting brood manufacturing.

Replace:  I don’t make suggestions.  My hope is that future analysis will fantastic tune the specs for pollen substitutes.  A superb assessment is Black, J (2006) Honeybee Diet: Evaluate of analysis and practices.  RIRDC Publication No 06/052  https://www.agrifutures.com.au/wp-content/uploads/publications/06-052.pdf

Crude protein: Different analysis means that 25-30% (dry weight) could also be optimum [[37], [38]].  The very best-performing subs on this trial had been within the 15-20% vary (moist weight of the eating regimen, moisture content material was not measured).  EAAs ought to make up about 50% of the crude protein content material. One can use my calculator [[39]] to stability the EAAs by including concentrates.

Sugar content material: round 40-50%.

Lipid content material: within the 1-5% vary, with canola oil seeming to work properly. Ghosh discovered that palmitic, stearic and oleic acids had been predominant in the entire levels.

Nutritional vitamins, minerals, and sterols: Dietary necessities should not actually clear, and these could also be adequately offered by the uncooked feedstuffs (similar to brewer’s yeast). However observe that no less than some brewer’s yeasts could also be distasteful to bees [[40]]. Ghosh discovered that grownup bees include greater concentrations of the hint parts iron, zinc, calcium, magnesium, and copper than these present in purple meat, suggesting that any pollen sub ought to include ample ranges of those hint parts. It could be clever to ensure that zinc is within the 75 ppm vary.

Particle dimension: most bee-collected pollens are lower than 50 micrometers in diameter [[41]], suggesting that every one feedstuffs ought to possible be floor to not bigger than that particle dimension. (I’ve noticed colonies kicking out, reasonably than consuming, bigger particles.)

Updates:  Flours sifted by means of 200-mesh have a most particles dimension of 74 micrometers (microns).

Affect of the intestine microbiota:  An eye fixed-opening examine from Nancy Moran’s lab signifies that the symbiotic bacterium Bombella apis, which is present in affiliation with the jelly fed to queens and larvae, is ready to synthesize all important amino acids from an insufficient eating regimen.  It was notably capable of synthesize lysine, which seems to be extra essential for larvae than for grownup employees.  This discovering opens our eyes to new features of bee vitamin.

Parish, A.J., Rice, D.W., Tanquary, V.M., Tennessen, J.M., & Newton, I.L. (2022). A honey bee symbiont buffers larvae towards dietary stress by means of lysine supplementation. bioRxiv.

Pure pollen: Once more, any profit just isn’t clear. De Groot discovered that the enzymes in honey or beebread appeared to not be important in a eating regimen [[42]]. For phagostimulation, pollen must make up no more than 5% of the patty (Determine 8).

An extra picture.

Fig. 8 We’ve heard claims {that a} excessive proportion of pure pollen in a patty leads to the bees consuming it far more rapidly. So this fall my son Eric ran a take a look at, by inserting one 4% pollen and one 15% pollen International patty side-by-side on 150 hives. He then checked again, taking images in any respect levels of consumption, and confirmed them to me as a blinded grader of consumption fee. Neither of us may see any distinction.

A POSSIBLE INEXPENSIVE DIET

THAT’S IT!

This has been a for much longer write up on pollen subs than I imagined, however since so many people skilled beekeepers spend quite a lot of cash on them, I hope that ABJ’s readers really feel that this collection has been informative. I particularly hope that what we’ve discovered, in addition to my remaining questions, encourages our producers to enhance their formulations, and our researchers to take up the place de Groot and Elton Herbert, Jr. left off.

DE GROOT’S PAPER

I’ve talked about that de Groot’s monumental examine is tough to entry. These can obtain a replica at https://scientificbeekeeping.com/de-groot-1953/

REFERENCES AND NOTES

[1] Ghosh, S, et al. (2016) Dietary worth and chemical composition of larvae, pupae, and adults of employee honey bee, Apis mellifera ligustica as a sustainable meals supply. Journal of Asia-Pacific Entomology 19: 487–495.

[2] Mitchell, H, in Albanese, A (1950) Protein and amino acid necessities of mammals. Tutorial Press.

[3] Church, D, et al. (2020) Important amino acids and protein synthesis: insights into maximizing the muscle and whole-body response to feeding. Vitamins 12, 3717.

[4] Chan, Q & L Foster (2008) Modifications in protein expression throughout honey bee larval growth. Genome Biol. 9(10): R156.

[5] Corby-Harris V, et al. (2019) Fats physique lipolysis connects poor vitamin to hypopharyngeal gland degradation in Apis mellifera. J Insect Physiol. 116:1-9.

[6] Haydak, M (1970) Honey Bee Diet. Annual Evaluate of Entomology 15:143-156. Haydak discovered that 45% of belly nitrogen could also be misplaced by long-lived employees starved for protein.

[7] https://scientificbeekeeping.com/where-does-fed-pollen-sub-go/

[8] Noordyke, E, et al. (2021) Tracing the destiny of pollen substitute patties in western honey bee (Hymenoptera: Apidae) Colonies. Journal of Financial Entomology 114(4): 1421–1430.

[9] Haydak, M. H. (1935). Brood rearing by honey bees confined to a pure carbohydrate eating regimen. Journal of Financial Entomology, 28, 657-660.

[10] Otis, G, et al. (2004) Storage proteins in winter honey bees. Apiacata 38: 352-357.

[11] Scheller Okay, et al. (1990) Molecular properties, features and developmentally regulated biosynthesis of arylphorin in Calliphora vicina. In: Hagedorn H, et al. (eds) Molecular Insect Science.

[12] Johansson, T & M Johansson (1977) Beekeeping methods, Feeding Sugar to Bees. 3. Dry Sugar and Sweet. Bee World, 58:2, 49-52.

[13] The “fragrant” amino acids listed later on this artlcle.

[14] Jeffree, E (1956) Winter brood and pollen in honeybee colonies. Ins. Soc 3: 417–422.

[15] Haydak, M (1934) Modifications in whole nitrogen content material in the course of the lifetime of the imago of the employee honeybee. J. Agricultural Analysis 49(1): 21-28.

[16] Straus, J (1911) Die chemische zusammensetzung der arbeitsbienen und drohnen während verschiedenen entwicklungsstadien. Ztschr. Biol. 56: 347-397.

[17] De Groot, A (1954) On the ovary growth in queenless employee bees (Apis mellifica L.) Experientia Vol. X/9: 384-385.

[18] Babendreier, D, et al. (2004) Pollen consumption in honey bee larvae: a step ahead within the danger evaluation of transgenic crops. Apidologie 35: 293–300.

[19] Schmehl, D, et al. (2016). Protocol for the in vitro rearing of honey bee (Apis mellifera L.) employees. Journal of Apicultural Analysis 55(2): 113-129. I’ve corresponded with Dr. Schmehl about his inclusion of yeast extract — different researchers have discovered that it’s not vital.

[20] Zhang, J, et al. (2009) Dedication of tryptophan in bee pollen and royal jelly by high-performance liquid chromatography with fluorescence detection. Biomed. Chromatogr. 23: 994–998.

[21] Webster, T, et al. (1987). Conservation of vitamins in larval tissue by cannibalizing honey bees. Physiological entomology 12(2): 225-231.

[22] Loper, G & R Berdel (1980) The impact of 9 pollen diets on brood rearing of honey bees. Apidologie 11: 351-359.

[23] de Groot, A. (1953) Protein and amino acid necessities of the honey bee. Physiol. Comp. Oecol. 3, 197–285.

[24] Standifer, L, et al. (1960) Relative availability of varied proteins to the honey bee. Annals of the Entomological Society of America 53(5): 618–625.

[25] Williams, H, et al. (1954) Estimation of progress necessities for amino acids by assay of the carcass. J. Biol. Chem 208: 277-256.

[26] Ericson, L (1961) A criticism of the carcass evaluation process for the dedication of amino acid necessities. Acta physiol. Scand. 52: 90-98.

[27] Sklan D & Y Noy (2005) Direct dedication of optimum amino acid consumption for upkeep and progress in broilers. Poultry Science 84(3): 412-418.

[28] Ghosh, S, et al. (2016) Dietary worth and chemical composition of larvae, pupae, and adults of employee honey bee, Apis mellifera ligustica as a sustainable protein supply. Journal of Asia-Pacific Entomology 19(2): 487-495.

[29] de Groot, op. cit.

[30] Standifer, L, et al. (1960) Relative availability of varied proteins to the honey bee. Annals of the Entomological Society of America 53(5): 618–625.

[31] McNally, J, et al. (1965). Partition of excreted nitrogen from honey bees fed numerous proteins. The Journal of vitamin 85(1): 113-116.

[32] The premise for his beneficial EAA ratios.

[33] Hendriksma, H, et al. (2019) Results of important amino acid supplementation to advertise honey bee gland and muscle growth in cages and colonies. Journal of Insect Physiology 117, 103906.

[34] Herbert E & H. Shimanuki (1978) Impact of the dimensions of outside flight cages on brood rearing and meals consumption by honeybees. Journal of Apicultural Res 17(3): 114-117.

[35] Haydak, MH (1963) Age of nurse bees and brood rearing. Journal of Apicultural Analysis 2(2):101-103. Haydak was capable of get a cohort of nurses to repeatedly rear larvae for as much as 138 days. One may take a look at the dietary efficiency of synthetic diets by this methodology.

[36] Laidlaw, H & R Web page (1997) Queen Rearing and Bee Breeding. Wicwas Press. It may be doable to measure what number of wholesome queens may very well be raised by a cohort of nurses fed a synthetic eating regimen.

[37] Herbert, E, et al. (1977). Optimum protein ranges required by honey bees (Hymenoptera, Apidae) to provoke and keep brood rearing. Apidologie 8(2): 141-146.

[38] Zheng, B, et al. (2014). The consequences of dietary protein ranges on the inhabitants progress, efficiency, and physiology of honey bee employees throughout early spring. Journal of Insect Science14(1).

[39] https://scientificbeekeeping.com/7333-2/

[40] https://scientificbeekeeping.com/a-comparative-test-of-the-pollen-sub/

[41] Hao, Okay, et al. (2020) Pollen grain dimension related to pollinator feeding technique. Proc. R. Soc. B 287: 20201191.

[42] De Groot, A (1950) The affect of temperature and sort of meals on the rise within the nitrogen content material of the younger employee honeybee (Apis mellifica L.). forty fifth communication of the “Werkgemeenschap voor Endocrinologie”

 

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