A Nearer Glance
Queen’s Court docket
By means of: Clarence Collison
When the queen is desk bound at the comb she is surrounded by means of a circle of staff referred to as the “courtroom” or “retinue” that face towards her and feed, palpate and lick her. Generally, the retinue consists of 8 or 10 staff. The purposes of the employees making up the queen’s courtroom (retinue) are to feed her, groom her, take away her waste and to lick pheromones from her frame. The queen hardly, if ever, feeds herself. Secretions from mandibular glands and hypopharyngeal glands shape the proteinaceous jelly this is fed to queens. The volume the queen is fed by means of grownup staff in her courtroom is expounded to the queen’s egg laying fee. “The queen produces QRP (Queen Retinue Pheromone) which accommodates QMP (Queen Mandibular Pheromone) and no less than 4 further pheromones. QRP elicits retinue conduct from more youthful staff, who contact, lick and groom the queen. Within the procedure they gain QMP and the opposite queen pheromones, which might be then disbursed all over the colony” (Schneider, 2015). Queen mandibular pheromone is a mix of 5 other pheromones produced by means of the mandibular glands (9–oxo-trans-2–decenoic acid [9-ODA], ± 9-hydroxydec-2-enoic acid [+/- 9-HAD], methyl-p-hydroxybenzoate [HOB] and 4-hydroxy-3-methyoxyphenylethanol [HVA]).
The queen mandibular pheromone (QMP), consisting of the 5 synergistic elements indexed in the past, is the one pheromone chemically recognized within the honey bee queen, however this pheromone does no longer totally replica the pheromonal task of a complete queen extract. To spot the rest unknown compounds for retinue appeal, honey bee colonies have been selectively bred to have low reaction to artificial QMP and top reaction to a queen extract in a laboratory retinue bioassay. Employees from those colonies have been then used within the bioassay to steer the isolation and identity of the rest lively elements. 4 new compounds have been recognized from a number of glandular assets that account for almost all of the adaptation in retinue appeal between artificial QMP and queen extract: methyl (Z)-octadec-9-enoate (methyl oleate), (E)-3-(4-hydroxy-3-methoxyphenyl)-prop-2-en-1-ol (coniferyl alcohol), hexadecan-1-ol and (Z9,Z12,Z15)-octadeca-9,12,15-trienoic acid (linolenic acid). Those compounds have been inactive on my own or together, and so they best elicited appeal within the presence of QMP. There was once nonetheless unidentified task final within the queen extract. The queen due to this fact produces a synergistic, multiglandular pheromone mix of no less than 9 compounds for retinue appeal, essentially the most complicated pheromone mix identified for inducing a unmarried conduct in any organism (Keeling et al., 2003).
Queen attendance conduct of staff from decided on honey bee colonies with low and high employee retinue reaction to artificial queen mandibular gland pheromone (QMP) was once investigated. Antennating, licking, grooming and feeding of the queen by means of staff from low and high responding colonies have been tested. Low and high QMP responding staff didn’t attend the queen otherwise. On the other hand, staff originating from other colonies antennated and licked the queen extra regularly than others, suggesting there could also be a genetic foundation for queen attendance conduct no longer essentially related to reaction to QMP. The median age of queen attendance was once unbiased of pressure (Pankiw et al., 1995).
The conduct of queen and employee honey bees was once noticed the usage of small colonies in commentary hives. Employees paid extra consideration to queens which were mated for 2 months or greater than to these which have been newly mated; virgin queens gained the least consideration. Queens gained maximum consideration after they have been desk bound and least after they have been strolling over the brush; virgin queens have been maximum lively. The queen pheromone element 9–oxo-trans-2–decenoic acid stimulated ‘courtroom’ conduct when offered on small polyethylene blocks, however staff replied aggressively to finish extracts of queens’ heads. Each the heads and abdomens of mated queens gained a lot consideration from courtroom staff, however the abdomens have been palpated by means of extra staff for longer and have been licked a lot more. The queens’ thoraces have been least attended. Stomach tergites posterior to tergite glands have been licked for longer than the ones anterior to the glands. Simplest employee bees very almost about the queen reacted to her and joined her ‘courtroom’. No proof was once discovered of a diel periodicity within the conduct of a queen or her ‘courtroom’. Throughout the Iciness the queen’s courtroom was once smaller than in Summer time and she or he walked much less and laid fewer eggs. When colonies have been fed with sucrose syrup in Iciness, their queens laid extra eggs and staff reared extra brood however there was once no trade within the consideration gained by means of the queens (Unfastened et al., 1992).
Allen (1960) investigated the conduct of the queen’s attendants. 100 ninety-six for my part marked bees have been observed to inspect the queen and the intervals in their visits have been recorded. All gradations of conduct various from an obvious need to steer clear of the queen to a powerful need to stay along with her have been discovered. The conduct of particular person bees numerous on other visits, however no common collection may well be discovered. Greater than part the overall choice of visits had a period of 30 seconds or much less whilst the remaining have been longer than 30 seconds, with a most of 41 mins. About 8% of the visits have been very quick and terminated rapidly and with indicators of alarm, however no reason why for this kind of conduct was once obvious. The ages of the attendants numerous from a couple of hours to 52 days however no glaring distinction in age-distribution between the attendants final for terribly quick classes and the ones final for longer was once discovered. Relating to the overall numbers of marked bees of various ages provide, the share of staff performing as attendants confirmed no systematic permutations, even supposing in a colony the place the common age was once low (as in Summer time), the true numbers of younger attendants would it appears very much exceed the ones of the older ones. The age-range of attendants licking the queen was once very similar to the total age-range of the attendants. The ages of the attendants feeding the queen in two colonies ranged from sooner or later to 23 days. No bee was once observed to feed the queen right through the day of its emergence, even supposing the best numbers recorded befell in the following couple of days of grownup existence. No systematic adjustments have been obvious within the period of feeds given by means of bees of various ages, the imply worth for 2 colonies being 44 seconds. Observations made right through the season of lively brood-rearing as much as swarming time (April-July) confirmed that the quantity and period of feeds provided to the queen tended to upward thrust till two to a few weeks prior to the swarm left, however on this latter duration each quantity and period reduced, roughly from the time that queen mobile formation began. Thus, the queen gained steadily diminishing amounts of meals within the duration of swarm preparation. Egg-production over the similar duration confirmed a an identical pattern.
In every other find out about, staff accumulated on cages containing a mated laying queen positioned above a typical queenright colony. Fewer staff accumulated on a cage when they may no longer contact the caged queen, or when she was once useless, or when her mandibular glands were got rid of. The choice of staff at the cage was once correlated with the quantity of 9-oxodecenoic acid (number one element of queen mandibular pheromone) within the queen’s head. Employees accumulated on cages containing best 9-oxodecenoic acid on filter-paper that the bees may no longer contact. ‘Court docket’ formation spherical a loose queen was once no longer decreased by means of eliminating her mandibular glands. The pinnacle of queens whose mandibular glands were got rid of contained lines of 9-oxodecenoic acid. Up until now, no unknown substance affecting employee bees’ conduct was once detected within the queen’s head. No sounds more likely to be in particular communicative have been heard from the queen or from staff that shaped a ‘courtroom’ around her. Employees didn’t transfer from afar to shape a ‘courtroom’ spherical a queen. It’s concluded that during an undisturbed colony with a mated laying queen there’s no appeal of staff in opposition to the queen except for over very quick distances (Butler et al., 1973).
Honey bee colonies deal with viable queens partially thru conversation with queen mandibular pheromone (QMP), a combination that alerts the queen’s presence and reproductive high quality to staff. In flip, staff are idea to supply retinue queen care or exchange queens in part in accordance with QMP profiles. The results of seasonal dearth have been tested (overwintering in a heat subtropical location) on queen-worker interactions. Retinue employee responses to ceaselessly ovipositing queens have been regarded as in view of QMP signaling and queen reproductive high quality. QMP signaling was once estimated from QMP residues recovered from nest employee our bodies, which is the principle mode of QMP switch from the queen to the colony at massive. QMP residues numerous seasonally however under no circumstances with queen reproductive high quality (spermatheca sperm garage, ovary protein and lipid contents). 9-HDA and 9-ODA have been decrease in January than different months. HOB reduced from July to January, whilst HVA, an element related to mated queens, larger sharply in January. Regardless of those seasonal signaling variations, retinue staff attended queens at an identical ranges throughout the months. In relation to reproductive high quality, queens didn’t fluctuate over the months in matedness (spermatheca sperm garage) or physiological age (protein carbonyl content material) however numerous in nutrient allocation to reproductive and non-reproductive tissues. Queen ovaries contained extra protein in September than in November, and extra lipid in July and September than in November and January. Queen fats our bodies had extra protein in July than September or November, however much less lipid in July and September than November or January. Retinue employee responses didn’t range with seasonal QMP adjustments, however mirrored total steady brood rearing efforts and queen matedness all over the yr. The absence of differences due to the season in employee responses to QMP must be regarded as within the broader context of continuing reproductive efforts in heat subtropical colonies (Carroll et al., 2023).
Honey bee queen attendants disperse queen pheromones to complement pheromone dispersal by means of direct queen-worker contacts. With time they lose their dispersal serve as exponentially due principally to volatilization of queen pheromones carried on their our bodies. The removing of the ones airborne pheromones along with the air whilst ventilating the hive is balanced by means of pheromone free up by means of the queen. This equilibrium ends up in a undeniable stage of queen pheromones within the broodnest. The trade of the pheromone stage (for instance, because of lack of the colony of its queen) can function a sign to vary the conduct of the employees and the state of the colony (Juška et al., 1981).
The substance produced by means of the mandibular gland complicated (HQMC) of queen honey bees is thought of as to be liable for retinue formation in honey bees. The retinue reaction comprises the licking and antennating conduct which alerts the presence of a dominant reproductive queen and thereby establishes and stabilizes the social cloth of the colony. Regardless of 25 years of study, not one of the identified constituents of the mandibular gland complicated extract has been efficient in eliciting complete retinue conduct. We record that the mandibular-gland-based retinue reaction is mediated by means of 5 semio-chemicals. Every element is weakly lively on my own, however your entire mix imparts task similar to HQMC extract at a degree as little as 10–7 of that found in a queen (Slessor et al., 1988).
Ferguson and Unfastened (1980) observations equipped robust circumstantial proof that staff in a queen’s courtroom download queen pheromone on their antennae, and that queen pheromone is shipped throughout the colony right through antennal touch between staff. Employees that had simply left the courtroom of a mated or virgin queen had an larger tendency to make reciprocated antennal contacts with different staff. This tendency was once bolstered when the employees involved licked the queen along with palpating her with their antennae, most probably as a result of they spent longer within the courtroom. The primary staff contacted by means of the ones leaving the courtroom additionally had an larger tendency to make reciprocated antennal contacts. Antennal contacts have been much more likely to be initiated by means of bees rather then the ones from the queen’s courtroom. Employees from the courtroom and the ones they first contacted participated in meals switch greater than did staff decided on at random. Employees that licked a virgin queen therefore participated in meals switch greater than those who palpated her with their antennae best. Inside of about 5 mins of leaving the courtroom, a employee’s participation in reciprocated antennal contacts and in meals switch decreased to the extent of non-court staff. Employees selected at random made extra brief and non-reciprocated antennal contacts when in a colony with a mated queen than when in a colony with a virgin queen.
The ontogeny of the 5 queen mandibular gland semiochemicals that begin and deal with the retinue conduct of employee honey bees was once investigated by means of quantitative splitless capillary gasoline chromatography. No detectable pheromone is provide on the time of eclosion, however decenoic acid ranges building up hastily right through the primary week of the queen’s existence. Two fragrant elements reach detectable ranges later, with the extra considerable methylp-hydroxybenzoate previous the 4-hydroxy-3-methoxyphenylethanol. Pheromone ranges are maximal in mature, mated, laying queens. The ratio of (R,E)-(−)-9-hydroxy-2-decenoic acid to the (S,E)-(+) enantiomer will increase with the age of the queen. Pheromone ranges in queen mandibular glands are in large part unaffected by means of queen banking, restraint with staff in mailing cages, and restricted garage on dry ice. All primary frame portions of standard queens, particularly the top and legs, have enough mandibular exudate to be extremely sexy to employee bees (Slessor et al., 1990).
References
Allen, M.D. 1960. The honey bee queen and her attendants. Anim. Behav. 8: 201-208.
Butler, C.G., R.Okay. Callow, C.G. Koster and J. Simpson 1973. Belief of the queen by means of staff within the honey bee colony. J. Apic. Res. 12: 159-166.
Carroll, M.J., N.J. Brown, Z. Ruetz, V.A. Ricigliano, and Okay.E. Anderson 2023. Honey bee retinue staff reply in a similar fashion to queens in spite of differences due to the season in Queen Mandibular Pheromone (QMP) signaling. PLoS ONE 18(9): e0291710
Ferguson, A.W. and J. B. Unfastened 1980. Queen pheromone switch inside of honey bee colonies. Physiol. Entomol. 5: 358-366.
Unfastened, J.B., A.W. Ferguson and J.R. Simpkins 1992. The behaviour of queen honey bees and their attendants. Physiol. Entomol. 17: 43-55.
Juška, A., T.D. Seeley and H.H.W. Velthuis 1981. How honey bee queen attendants turn out to be peculiar staff. J. Insect. Physiol. 27: 515-519.
Keeling, C.I., Okay.N. Slessor, H.A. Higo and M.L.Winston 2003. New elements of the honey bee (Apis mellifera L.) queen retinue pheromone. Proc. Natl. Acad. Sci. USA 100 (8): 4486-4491.
Pankiw, T., M.L. Winston and Okay.N. Slessor 1995. Queen attendance conduct of employee honey bees (Apis mellifera L.) which are low and high responding to queen mandibular pheromone. Insect. Soc. 42: 371-378.
Schneider, S.S. 2015. The honey bee colony: existence historical past. In: The Hive And The Honey Bee (Ed. J.M. Graham), Dadant and Sons, Hamilton, IL, pp. 73-109.
Slessor, Okay.N., L.-A. Kaminski, G.G.S. King, J.H. Borden and M.L. Winston 1988. Semiochemical foundation of the retinue reaction to queen honey bees. Nature 332: 354-356.
Slessor, Okay.N., L.A. Kaminski, G.G.S. King and M.L. Winston 1990. Semiochemicals of the honey bee queen mandibular glands. J. Chem. Ecol. 16: 851-860.
Clarence Collison is an Emeritus Professor of Entomology and Division Head Emeritus of Entomology and Plant Pathology at Mississippi State College, Mississippi State, MS.