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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 advised by de Groot in 1953. However he himself identified the shortcomings of his strategies and certified his suggestions. I really feel that after practically seventy years, it could be time to meet up with the opposite livestock industries, and replace our data on what constitutes an optimum supplemental weight-reduction plan 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 staff. 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 staff) fed synthetic diets, which might be the very best 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 sequence, I’m going to stay my neck out and make a stab at revising de Groot’s ratios and recommend the form of utilized analysis that our trade deserves.
Enable me to start by looking how the amino acids in a weight-reduction plan 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 assorted kinds of physique and haemolymph proteins are all created from the metabolic pool of amino acids derived from their weight-reduction plan, as illustrated by Determine 1 (revealed in the course of the years during 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 revealed this move chart of how dietary proteins had been utilized by the physique. Picture from [[2]].
The protein in a weight-reduction plan is first digested into its constituent amino acids, from which the physique then synthesizes new physique elements by the method of anabolism (the alternative is catabolism — the “harmful metabolism” of physique tissue). That course of is restricted largely by the quantity of important amino acids (EAAs) within the weight-reduction plan [[3]], in addition to their relative proportions to one another. The EAAs are used not just for constructing (or changing) muscle mass and inner organs, but additionally to create haemolymph and dispensable storage proteins. Some protein is interspersed in chitin to type the bees’ cuticle and wings. And a few could also be burned as gas for power if sufficient carbohydrates (sugars) aren’t accessible within the weight-reduction plan.
One technique to measure bee progress is by drying and weighing particular person bees. However such weight acquire consists of elements largely constructed from carbohydrates, equivalent 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 weight-reduction plan, which comes from the protein.
Sensible software: So maybe what we actually ought to focus upon is their enhance in nitrogen content material.This consideration didn’t escape de Groot, so he measured each dry weight acquire in addition to nitrogen enhance (Determine 2).
Fig 2. Evaluate the consequences of EAA deficiencies upon weight acquire (above) versus acquire in nitrogen (under), which signifies the precise incorporation of these EAAs into physique mass. Be aware 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 essential requirement of these EAAs for the method of constructing non-proteinaceous physique tissues, somewhat than being integrated 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 take into account that some EAAs are important for the constructing of non-nitrogenous physique elements. I’ll return to this idea after I focus on “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:
- Larval growth — Main weight acquire consisting of a small quantity of integument, muscle, digestive tract, and tracheal system, however principally from massive quantities of storage protein in fats our bodies [[4]]. Protein supply — nurse-fed jelly.
- 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 the pupa is its personal protein supply, in order that evaluation of the amino acid ratio of its physique could also be informative.
- Put up-emergence growth — Main growth of the hypopharyngeal glands and fats our bodies, Partial growth of the wing muscle mass and the sting. Protein supply — pollen and a few jelly.
- Nurse bee growth — The nurses want to supply copious portions of protein-rich jelly for feeding the larvae and different members of the colony. Protein supply — pollen.
- Mid-age employee growth — Improvement of the wax glands and wing muscle mass. Protein supply —pollen, jelly, and autophagy of the fats our bodies and hypopharyngeal glands as they shed weight in preparation for foraging [[5]].
- Forager growth — Ultimate progress and upkeep of the wing muscle mass and physique tissues. Protein supply — jelly begged from nurses.
- “Winter bee” growth — The diutinus state taken by staff that emerge throughout pollen dearths is likely to be thought-about as “arrested growth” of nursing physiology. Protein supply — pollen.
- “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 could discover that there is no such thing as a one-size-fits-all weight-reduction plan for honey bees.
WINTER FEEDING
AUTUMN FEEDING FOR STORAGE PROTEINS
Earlier than I return to weight-reduction plan for colony growth, let’s first contemplate diets in preparation for, or throughout winter. Beekeepers feed pollen subs not solely to offer the required vitamins for the final rounds of brood that can turn into long-lived “winter bees,” but additionally to permit these emerged staff to totally develop their storage proteins as a way to survive the winter. Would the event of storage proteins require a distinct EAA ratio within the weight-reduction plan?
As identified by de Groot:
Many sorts of bugs can preserve life for appreciable intervals 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 receive the amino acids (and below situations of hunger, power) obligatory for “physique upkeep.”
Sensible software: De Groot studied the EAA necessities of younger staff not being uncovered to brood pheromone, suggesting that they could have gone into diutinus physiology. Thus, his beneficial EAA ratios could nicely apply to diets fed to the final spherical of rising staff 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 type contained in the hive aren’t saved within the combs [[7], [8]], so the advantage of autumn feeding of pollen patties could principally be to advertise the synthesis of these storage proteins. That stated, these storage proteins enable 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 extra protein as a way to full its growth).
Most beekeepers have heard of the necessary 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 necessary for the formation of those hexamerin storage proteins? These wealthy within the fragrant amino acids (phenylalanine, tyrosine, and tryptophan) are referred to as arylphorins, whereas one other group of hexamerins are generally known as methionine-rich storage proteins. Frustratingly, I used to be unable to seek out info on the EAAs in vitellogenin. The extraordinarily excessive proportion of tyrosine in insect storage proteins [[11]] means that we’d need 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 power 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 primarily based upon his preliminary analysis:
From the above outcomes it could be concluded, that in distinction with findings of different authors the longevity of previous bees is elevated by supplementing the carbohydrate weight-reduction plan with numerous protein containing proteins or pure protein in an appropriate focus. Clearly within the bee colony the necessity for protein will not be restricted to the brood rearing interval however proteins are required for upkeep of grownup bees as nicely. This discovering is in hanging distinction with the widespread opinion, that in the course of the winter interval the bee requires solely carbohydrates. It’s common 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 viewpoint. 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 previous 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 usually used at current in apiculture.
Winter feeding with straight sugar continues to be a typical 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. Be aware that Johansson cautioned that colonies fed with dry sugar could have to forage for water to dilute the sugar. Picture 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 might positive use some easily-performed area 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 might be of nice profit for these beekeepers who require sturdy colonies for early pollination contracts.
Sensible software: We almond pollinators have discovered that midwinter feeding of pollen sub below the appropriate 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 probably the most protein in its weight-reduction plan? It’s throughout their larval stage (Determine 4).
Fig. 4 Improve in nitrogen content material of the our bodies of employee bees throughout progress intervals. 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 staff (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 necessary 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 supplies an insignificant quantity of protein total [[18]]).
Thus de Groot’s estimated ratios could not apply to the all-important nurse bees, that are chargeable for the digestion of proteins for your entire colony (and are thus the shoppers 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 “foreign money of protein” inside the colony).
Sensible software: So for colony progress, what we actually want to find out is the optimum EAA ratio within the weight-reduction plan 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 weight-reduction plan consists primarily of harvested royal jelly [[19]]. Till a totally synthetic weight-reduction plan for larvae is developed, we have to follow determining diets that enable nurse bees to supply probably 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 will evaluate de Groot’s suggestions. I’ll graph them out after explaining them.
COMPARE TO ROYAL JELLY
It might be affordable to imagine that royal jelly produced by nurse bees would include the best 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 international locations and the consumption of various pollens. Among the older analyses lacked knowledge for tryptophan. Fortuitously, in 2009 a brand new methodology was developed to quantify tryptophan [ [20]], so I used to be in a position to insert a price. I averaged the 5 most constant analyses for graphing functions.
Aspect word: An fascinating research 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 affirm beekeeper observations that colonies develop very nicely on almond pollen [[22]], so we will possible use the EAA ratio of almond pollen or different recognized nutritious pollens (equivalent 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 are 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 will enable for full growth of their offspring. Maybe not surprisingly, each work very nicely as protein sources for honey bees [[23],[24]].
CARCASS ANALYSIS
One other doable indicator of EAA necessities for a weight-reduction plan are the EAAs current within the animal’s physique. That is termed “carcass evaluation” [[25]], and would definitely be a easy technique to decide the best ratio. Sadly, EAA necessities change with an animal’s stage of progress [[26], [27]], and whether or not the EAAs are literally integrated into the physique, or merely concerned within the strategy of tissue synthesis.
That stated, Ghosh [[28]] supplies amino acid analyses for employee bee larvae, pupae, and adults (Determine 5).
Fig. 5 Comparability of the EAA ratios of the three developmental phases of honey bee staff, 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 weight-reduction plan, 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 obligatory for tissue formation solely, somewhat than being integrated into the ensuing physique tissue.
Notes for Ghosh’s findings:
- Non-essential tyrosine and cysteine could 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 modifications 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 keep 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 acquire, 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 in a position to 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 seem like near these required for larval growth, however the variations point out that the optimum EAA ratios for a really perfect weight-reduction plan 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 must be checked out in comparison with different findings, equivalent to the truth that bees develop very nicely on the proteins casein (from milk) [[29]], albumen (from rooster eggs) [[30], [31]], royal jelly, and a few pollen varieties. 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 under (Determine 6).
Fig. 6 De Groot’s suggestions, in comparison with carcass analyses of employee developmental phases, royal jelly, pollen, rooster egg and cow milk proteins (each of which have been proven to assist bee progress). I’ve circled indicators of the place de Groot’s EAA ratios could have to be adjusted for colony progress versus emerged employee enhance in nitrogen content material [[32]]. I’ve additionally (in yellow) inserted a advised revision of de Groot’s beneficial ratios. Be aware 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 advised 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 enable me to be daring sufficient to recommend a revision (Desk 1):
| Desk 1. Instructed 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 believe that what you’d actually need to know is how do my advised 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. Be aware how carefully my advised 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 required 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 diet. … Our research additionally highlights the significance of testing honey bee diet with area and laboratory research. Testing weight-reduction plan remedies on caged bees as in contrast with complete colonies could 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 degree by utilizing 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.
Ideas 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 might be used [[36]].
BASIC FORMULA FOR HONEY BEE ARTIFICIAL DIETS
Based mostly upon a overview of the literature, plus the findings from our area trial, one can arrive at advised proportions for an optimum synthetic weight-reduction plan for exciting brood manufacturing.
Replace: I don’t make suggestions. My hope is that future analysis will high quality tune the specs for pollen substitutes. A superb overview 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 most effective-performing subs on this trial had been within the 15-20% vary (moist weight of the weight-reduction plan, 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 nicely. Ghosh discovered that palmitic, stearic and oleic acids had been predominant in the entire phases.
Nutritional vitamins, minerals, and sterols: Dietary necessities aren’t actually clear, and these could also be adequately supplied by the uncooked feedstuffs (equivalent to brewer’s yeast). However word that not less than some brewer’s yeasts could also be distasteful to bees [[40]]. Ghosh discovered that grownup bees include increased concentrations of the hint components 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 components. It might 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 each one feedstuffs ought to possible be floor to not bigger than that particle dimension. (I’ve noticed colonies kicking out, somewhat than consuming, bigger particles.)
Updates: Flours sifted via 200-mesh have a most particles dimension of 74 micrometers (microns).
Affect of the intestine microbiota: An eye fixed-opening research 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 weight-reduction plan. It was notably in a position to synthesize lysine, which seems to be extra necessary for larvae than for grownup staff. This discovering opens our eyes to new facets of bee diet.
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 via lysine supplementation. bioRxiv.
Pure pollen: Once more, any profit will not be clear. De Groot discovered that the enzymes in honey or beebread appeared to not be important in a weight-reduction plan [[42]]. For phagostimulation, pollen must make up no more than 5% of the patty (Determine 8).
A further photograph.
Fig. 8 We’ve heard claims {that a} excessive proportion of pure pollen in a patty ends in the bees consuming it rather more shortly. So this fall my son Eric ran a check, by putting one 4% pollen and one 15% pollen International patty side-by-side on 150 hives. He then checked again, taking photographs in any respect phases of consumption, and confirmed them to me as a blinded grader of consumption price. Neither of us might 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 sequence 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 research 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 diet 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 staff 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 weight-reduction plan. 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, capabilities 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 complete 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 vegetation. Apidologie 35: 293–300.
[19] Schmehl, D, et al. (2016). Protocol for the in vitro rearing of honey bee (Apis mellifera L.) staff. 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 obligatory.
[20] Zhang, J, et al. (2009) Willpower 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 willpower of amino acid necessities. Acta physiol. Scand. 52: 90-98.
[27] Sklan D & Y Noy (2005) Direct willpower 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 diet 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 scale of out of doors 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 in a position to get a cohort of nurses to repeatedly rear larvae for as much as 138 days. One might check the dietary efficiency of synthetic diets by this methodology.
[36] Laidlaw, H & R Web page (1997) Queen Rearing and Bee Breeding. Wicwas Press. It is likely to be doable to measure what number of wholesome queens might be raised by a cohort of nurses fed a man-made weight-reduction plan.
[37] Herbert, E, et al. (1977). Optimum protein ranges required by honey bees (Hymenoptera, Apidae) to provoke and preserve 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 staff throughout early spring. Journal of Insect Science, 14(1).
[39] https://scientificbeekeeping.com/7333-2/
[40] https://scientificbeekeeping.com/a-comparative-test-of-the-pollen-sub/
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