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A Comparative Trial of the Pollen Subs: Half 6- Do We Must 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 Revealed in ABJ December 2021

Randy Oliver
ScientificBeekeeping.com

Bee nutritionists have lengthy cited the important amino acid ratios steered 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 food plan for honey bee colony development.

De Groot understood that what actually counted for bee diets meant for colony development 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 apart from Elton Herbert have adopted up on measuring the expansion of colonies (as opposed to some caged employees) fed synthetic diets, which might be the perfect and best strategy to decide whether or not de Groot’s important amino acid (EAA) ratios for grownup (non-nurse) bee improvement could be the identical as for total colony improvement, 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 type of utilized analysis that our business deserves.

Permit me to start by having a look how the amino acids in a food 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 varied sorts of physique and haemolymph proteins are all created from the metabolic pool of amino acids derived from their food plan, as illustrated by Determine 1 (revealed through 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 have been utilized by the physique. Picture from [[2]].

The protein in a food plan 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 food plan [[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 additionally 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 gasoline for power if sufficient carbohydrates (sugars) are usually not obtainable within the food plan.

One strategy to measure bee development is by drying and weighing particular person bees. However such weight achieve contains elements largely constructed from carbohydrates, comparable 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 food plan, which comes from the protein.

Sensible utility: 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. Examine the results of EAA deficiencies upon weight achieve (above) versus achieve in nitrogen (under), which signifies the precise incorporation of these EAAs into physique mass. Word extreme weight reduction in grownup bees if leucine, threonine, valine are poor. However the graph on the proper signifies that the load loss was because of the vital requirement of these EAAs for the method of constructing non-proteinaceous physique tissues, relatively than being included into these tissues.

Sensible utility: We are able to’t simply take a look at protein analyses of physique tissues estimated from nitrogen content material; we should additionally remember 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 improvement, a employee honey bee is anabolizing completely different physique elements, maybe consuming completely different meals, and should require completely different proportions of EAAs:

  1. Larval improvement — Main weight achieve consisting of a small quantity of integument, muscle, digestive tract, and tracheal system, however largely from giant quantities of storage protein in fats our bodies [[4]]. Protein supply — nurse-fed jelly.
  2. Pupal-to-adult improvement — Growth 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. Submit-emergence improvement — Main improvement of the hypopharyngeal glands and fats our bodies, Partial improvement of the wing muscular tissues and the sting. Protein supply — pollen and a few jelly.
  4. Nurse bee improvement — 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 improvement — Growth of the wax glands and wing muscular tissues. Protein supply —pollen, jelly, and autophagy of the fats our bodies and hypopharyngeal glands as they shed pounds in preparation for foraging [[5]].
  6. Forager improvement — Remaining development and upkeep of the wing muscular tissues and physique tissues. Protein supply — jelly begged from nurses.
  7. “Winter bee” improvement — The diutinus state taken by employees that emerge throughout pollen dearths may be thought-about as “arrested improvement” 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 utility: Much like the completely different diets offered for the optimum improvement of broiler chickens at every stage of development, we might discover that there is no such thing as a one-size-fits-all food plan for honey bees.

WINTER FEEDING

AUTUMN FEEDING FOR STORAGE PROTEINS

Earlier than I return to food plan for colony improvement, let’s first think about 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 turn out to be long-lived “winter bees,” but additionally to permit these emerged employees 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 food 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 get hold of the amino acids (and below situations of hunger, power) crucial for “physique upkeep.”

Sensible utility: 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 advisable EAA ratios might effectively 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 are usually not saved within the combs [[7], [8]], so the advantage of autumn feeding of pollen patties might largely be to advertise the synthesis of these storage proteins. That stated, these storage proteins permit for less than 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 must devour extra protein as a way to full its improvement).

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

So which EAAs are vital 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 often called methionine-rich storage proteins. Frustratingly, I used to be unable to search out data 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 improvement or “winter bees.”

Sensible utility: 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 could be concluded, that in distinction with findings of different authors the longevity of previous bees is elevated by supplementing the carbohydrate food plan with numerous protein containing proteins or pure protein in an appropriate focus. Clearly within the bee colony the necessity for protein is just not restricted to the brood rearing interval however proteins are required for upkeep of grownup bees as effectively. This discovering is in hanging distinction with the widespread opinion, that through the winter interval the bee requires solely carbohydrates. It is not uncommon use in beekeeping follow to substitute the honey for shops of pure sugar for overwintering bee colonies, which isn’t thought-about to be disadvantageous from the dietary standpoint. Nevertheless, honeys comprise 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 follow as we speak (Determine 3).

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

Sensible utility: Including some high-quality protein to sugar fed throughout winter would doubtless 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 discipline 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 robust colonies for early pollination contracts.

Sensible utility: We almond pollinators have discovered that midwinter feeding of pollen sub below the proper 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 improvement does a employee bee truly require probably the most protein in its food plan? It’s throughout their larval stage (Determine 4).

Fig. 4 Enhance in nitrogen content material of the our bodies of employee bees throughout development 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 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 vital so far as colony development. However the larvae profit from pollen sub not directly — it’s the nurse bees that truly devour the pollen sub and convert it to jelly (any pollen that the larvae may 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 the whole 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 “foreign money of protein” throughout the colony).

Sensible utility: So for colony development, what we actually want to find out is the optimum EAA ratio within the food 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 food plan consists primarily of harvested royal jelly [[19]]. Till a very synthetic food plan for larvae is developed, we have to keep on with determining diets that permit nurse bees to provide probably the most jelly.

There are a number of attainable methods for researchers to take action. Permit 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 will be cheap to imagine that royal jelly produced by nurse bees would comprise the best ratio of EAAs for optimum larval development. I downloaded a number of analyses of the amino acids in royal jelly, and located that the EAA ratios have been remarkably constant, regardless of being from completely different international locations and the consumption of various pollens. A few of the older analyses lacked knowledge for tryptophan. Thankfully, in 2009 a brand new technique was developed to quantify tryptophan [ [20]], so I used to be in a position to insert a worth. I averaged the 5 most constant analyses for graphing functions.

Facet word: 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 Seventies verify beekeeper observations that colonies develop very effectively on almond pollen [[22]], so we are able to doubtless use the EAA ratio of almond pollen or different identified nutritious pollens (comparable to mustard or candy clover) as benchmarks.

COMPARSION TO OTHER ANIMALS AND WHAT THEY FEED THEIR OFFSPRING

Much like royal jelly being the “good” meals for queens and larvae, cows and chickens additionally produce proteins which might be good 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 may permit for full improvement of their offspring. Maybe not surprisingly, each work very effectively as protein sources for honey bees [[23],­­­­[24]].

CARCASS ANALYSIS

One other attainable indicator of EAA necessities for a food plan are the EAAs current within the animal’s physique. That is termed “carcass evaluation” [[25]], and will surely be a easy strategy to decide the best ratio. Sadly, EAA necessities change with an animal’s stage of development [[26], [27]], and whether or not the EAAs are literally included into the physique, or merely concerned within the means 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 food plan, whereas it ought to doubtless 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 have been crucial for tissue formation solely, relatively 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 purpose 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 have been in-hive “home bees” (wings eliminated) that had the chance to feed upon pollen within the combs, so their EAA modifications from pupae have been doubtless because of the improvement of grownup tissues after feeding upon pollen.
  • Tryptophan is required for tissue development, 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 stunning, 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 development, 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 in a position to detect tryptophan in larvae or pupae, and solely 0.03% in adults — additional purpose to not normalize EAA ratios to tryptophan.

Sensible utility: De Groot’s ratios for grownup bees seem like near these required for larval improvement, however the variations point out that the optimum EAA ratios for a really perfect food plan for colony developmentmay should 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, comparable to the truth that bees develop very effectively 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 levels, royal jelly, pollen, rooster egg and cow milk proteins (each of which have been proven to assist bee development). I’ve circled indicators of the place de Groot’s EAA ratios might should be adjusted for colony development versus emerged employee improve in nitrogen content material [[32]]. I’ve additionally (in yellow) inserted a steered revision of de Groot’s advisable ratios. Word that not all analyses mixed the fragrant amino acids.

Sensible utility: The above chart means that de Groot was actually not far off the mark. Nevertheless, for optimum egg-to-adult employee development, his steered 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 crimson) to de Groot’s suggestions for the proportional ratios of the EAAs for diets meant for colony development.
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 steered 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. Word how intently my steered 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 diet. … Our examine additionally highlights the significance of testing honey bee diet with discipline and laboratory research. Testing food plan remedies on caged bees as in contrast with complete colonies might yield completely different insights.

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

Ideas for researchers: That stated, I’m curious as as to if a modification of Haydak’s technique of extending nurse longevity by eradicating sealed brood [[35]], or the swarm field technique 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

Primarily based upon a overview of the literature, plus the findings from our discipline trial, one can arrive at steered proportions for an optimum synthetic food plan for exciting brood manufacturing.

Replace:  I don’t make suggestions.  My hope is that future analysis will advantageous tune the specs for pollen substitutes.  A wonderful overview is Black, J (2006) Honeybee Vitamin: Assessment 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 have been within the 15-20% vary (moist weight of the food 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 steadiness 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 effectively. Ghosh discovered that palmitic, stearic and oleic acids have been predominant in all the levels.

Nutritional vitamins, minerals, and sterols: Dietary necessities are usually not actually clear, and these could also be adequately offered by the uncooked feedstuffs (comparable to brewer’s yeast). However word that a minimum of some brewer’s yeasts could also be distasteful to bees [[40]]. Ghosh discovered that grownup bees comprise larger concentrations of the hint components iron, zinc, calcium, magnesium, and copper than these present in crimson meat, suggesting that any pollen sub ought to comprise ample ranges of those hint components. It might be sensible to guarantee that zinc is within the 75 ppm vary.

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

Updates:  Flours sifted by way of 200-mesh have a most particles measurement 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 food plan.  It was notably in a position to synthesize lysine, which seems to be extra vital for larvae than for grownup employees.  This discovering opens our eyes to new features 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 by way of lysine supplementation. bioRxiv.

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

A further picture.

Fig. 8 We’ve heard claims {that a} excessive proportion of pure pollen in a patty ends in the bees consuming it way more rapidly. So this fall my son Eric ran a take a look at, by putting one 4% pollen and one 15% pollen International patty side-by-side on 150 hives. He then checked again, taking pictures in any respect levels of consumption, and confirmed them to me as a blinded grader of consumption charge. 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 an excessive amount 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 realized, 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 troublesome to entry. These can obtain a duplicate 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 improvement. 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 Vitamin. Annual Assessment of Entomology 15:143-156. Haydak discovered that 45% of stomach 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 food 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, 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 strategies, 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 through 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 improvement 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 threat 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 crucial.

[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 development 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 development 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 advisable EAA ratios.

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

[34] Herbert E & H. Shimanuki (1978) Impact of the scale 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 in a position to 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 technique.

[36] Laidlaw, H & R Web page (1997) Queen Rearing and Bee Breeding. Wicwas Press. It may be attainable to measure what number of wholesome queens might be raised by a cohort of nurses fed a man-made food 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 results of dietary protein ranges on the inhabitants development, efficiency, and physiology of honey bee employees throughout early spring. Journal of Insect Science14(1).

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

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