The influence of brood on the pollen consumption of worker bees (Apis mellifera L.)

(1) In midgut dry weight (tissue plus contents) of worker bees we found a representative parameter for pollen consumption. Midguts of bees of successive ages were analyzed and correlated with various parameters. The relative proportions of sugar, protein and water were either constant or negatively correlated with midgut weight. Only the relative pollen weight (percent of midgut dry weight) increased. (2) To investigate the influence of different levels of brood on pollen consumption of individual bees, midgut dry weights from 2 normally breeding control colonies and 2 brood-reduced experimental colonies were analyzed. In bees from control colonies the pollen consumption increased up to the nursing age (3-10d), remained on an elevated level in middle-aged-bees (10-18d) and decreased relatively sharply towards the foraging ages (>21d). When queens were caged in the experimental colonies, the following decline of brood cells affected the consumption of pollen differently. After 6 days of caging, with a reduction of open brood only, no effect was seen. After 15 days, and even more pronounced after 23 days when no brood was present, the pollen consumption in young and middle-aged (10, 14, 18d) worker bees was significantly reduced, while it was clearly elevated in older bees. We discuss pollen consumption as an adaptation to reduced necessity to nurse brood in young and middle-aged bees, and to enhance life span in older animals.     

Dwindling pollen resources trigger the transition to broodless populations of long-lived honeybees each autumn

Abstract.  1. Each autumn in northern regions, honeybee colonies shift from populations of short-lived workers that actively rear brood to broodless populations of long-lived winter bees. To determine if dwindling pollen resources trigger this transition, the natural disappearance of external pollen resources was artificially accelerated or delayed and colonies were monitored for effects on the decline in brood-rearing activity and the development of populations of long-lived winter bees.
2. Delaying the disappearance of pollen resources postponed the decline in brood rearing in colonies. Colonies with an extended supply of pollen reared workers longer into October before brood rearing ended than control colonies or colonies for which pollen supply was cut short artificially in autumn.
3. Colonies with extended pollen supply produced more workers throughout autumn than colonies with less pollen, but the development of the population of long-lived winter bees was delayed until relatively later in autumn. Colonies produced similar numbers of winter bees, regardless of the timing of the disappearance of pollen resources.
4. Mean longevity of autumn-reared workers was inversely related to the amount of brood remaining to be reared in colonies when workers eclosed. Consequently, long-lived workers did not appear in colonies until brood rearing declined, which in turn was controlled by the availability of pollen.
5. Dwindling pollen resources provide a powerful cue that initiates the transition to populations of broodless winter bees because it directly affects the brood-rearing capacity of colonies and indirectly indicates deteriorating environmental conditions associated with the approach of winter.     

Effects of clothianidin on Bombus impatiens (Hymenoptera: Apidae) colony health and foraging ability

We conducted laboratory experiments to investigate the lethal and sublethal effects of clothianidin on bumble bee, Bombus impatiens Cresson, colony health and foraging ability. Bumble bee colonies were exposed to 6 ppb clothianidin, representing the highest residue levels found in field studies on pollen, and a higher dose of 36 ppb clothianidin in pollen. Clothianidin did not effect pollen consumption, newly emerged worker weights, amount of brood or the number of workers, males, and queens at either dose. The foraging ability of worker bees tested on an artificial array of complex flowers also did not differ among treatments. These results suggest that clothianidin residues found in seed-treated canola and possibly other crops will not adversely affect the health of bumble bee colonies or the foraging ability of workers.     

Effect of primer pheromones and pollen diet on the food producing glands of worker honey bees (Apis mellifera L.)

Cooperative brood care is highly developed in the honey bee such that workers called nurses use their hypopharyngeal and mandibular glands to biosynthesize proteinaceous secretions that are progressively provisioned to larvae. The role that honey bee primer pheromones play in the functional physiology of food producing glands was examined. The combined and separate effects of queen mandibular pheromone (QMP) and brood pheromone (BP) on amount of protein extractable from hypopharyngeal and mandibular glands of workers reared for 12 days with and without pollen diets was measured. In rearing environments with a pollen diet, BP, and QMP + BP pheromone treatments significantly increased extractable protein from both glands. Bees reared with QMP + pollen had amounts of protein extractable from both glands that were not significantly different from control bees (no pheromones, no pollen). Pollen in the diet alone significantly increased amounts of protein extractable from glands versus control. In rearing environments without pollen, QMP + BP had a synergizing effect on amount of protein in both glands. The QMP + BP treatment was the only rearing environment without a pollen diet where protein amounts were significantly greater than the control. The synergizing effect of QMP + BP on extractable mandibular and hypopharyngeal gland protein suggests a highly derived role for the combined effect of these two primer pheromones on honey bee cooperative brood care. Mandibular gland area was significantly and positively correlated with extractable protein. Amounts of extractable protein from both glands declined significantly with age of workers in all treatments. However, treatment significantly affected rate of decline. The adaptive significance of gland protein amounts in response to pheromones and pollen diet are discussed.     

Regulation of hypopharyngeal gland activity and oogenesis in honey bee (Apis mellifera) workers

In the honey bee, vitellogenin has several functions in addition to egg provisioning. Among others, it serves as a precursor to brood food proteins secreted by the hypopharyngeal glands of worker bees. In queenless workers with developing gonads, oogenesis and development of the hypopharyngeal glands are correlated. Here we describe two experiments that explored whether this relationship also exists in non-reproductive workers, and investigated a possible role of ecdysteroid hormones in the regulation of vitellogenin uptake. In the first experiment, the correlation between oocyte length and hypopharyngeal gland development was measured in workers before and after de-queening. In the second experiment, we induced middle-aged bees with resting glands to suddenly initiate brood care behaviour, and measured haemolymph ecdysteroid and vitellogenin titres. A strong positive relationship existed between morphometrical parameters of hypopharyngeal glands and ovaries in both queenless and queenright (functionally sterile) workers. No response of ecdysteroid titres to the addition of brood was detected in experiment 2, but high concentrations were measured in a small group of bees characterised by the possession of oocytes on the brink of yolk incorporation. We conclude that hypopharyngeal glands may belong to a previously described group of reproduction-related traits that are pleiotropically regulated in workers. A possible role for ecdysteroids in honey bee reproduction is discussed.     

THE INFLUENCE OF STORED POLLEN AND OF COLONY SIZE ON THE BROOD REARING OF HONEYBEES

Four hundred and thirty records of the numbers of bees in honeybee colonies and of the amounts of brood and pollen present have been kept during various months of the years 1945-53, and the data have been used to calculate total and partial regression coefficients showing the influence of stored pollen and of colony size on brood rearing throughout the year.
It was found that pollen storage and colony size were correlated but that, even allowing for this, colony size and pollen both independently influenced brood rearing.
The annual distribution of the total regression coefficients of brood on pollen was somewhat similar to the brood curve itself, rising from a minimum in October and November to a maximum in midsummer, while the partial regression coefficients showed less clearly marked but similar features.
Both total and partial regression coefficients showing the influence of colony size on the amount of brood reared were also at a minimum in October and November, but reached their peaks in May.
The quantities of brood present in these colonies at Aberdeen, Scotland, followed a pattern similar to that given by Nolan for colonies near Washington, D.C.     

Associations between reproduction and work in workers of the Asian hive bee Apis cerana

If a honey bee (Apis spp.) colony becomes queenless, about 1/3 of young workers activate their ovaries and produce haploid male-producing eggs. In doing so queenless workers maximize their inclusive fitness because the normal option of vicarious production of relatives via their queen’s eggs is no longer available. But if many workers are engaged in reproduction, how does a queenless colony continue to feed its brood and forage? Here we show that in the Asian hive bee Apis cerana hypopharyngeal gland (HPG) size is larger in queenless workers than in queenright workers and that bees undertaking brood-rearing tasks have larger HPG than same-aged bees that are foraging. In queenless colonies, workers with a smaller number of ovarioles are more likely to have activated ovaries. This reinforces the puzzling observation that a large number of ovarioles reduces reproductive success in queenless A. cerana. It further suggests that reproductive workers either avoid foraging or transition to foraging later in life than non-reproductive workers. Finally, our study also showed that ovary activation and larger-than-average numbers of ovarioles had no statistically detectable influence on foraging specialization for pollen or nectar.     

Interadult feeding of jelly in honeybee (Apis mellifera L.) colonies

Summary Honeybee nurses (8 days old) were injected with ¹⁴C-phenylalanine. These bees then dispensed the ¹⁴C-labelled protein-rich products of their hypopharyngeal glands to the queen and the brood, and also to young drones and workers of all age classes. In small colonies containing 400–800 bees, nearly one-quarter of the radioactivity which could not be recovered in the nurses was fed by them in a protein-bound form to other members of the worker caste. During one night, one nurse fed an average of 4–5 foragers with proteinaceous food. The role of nurses in the work allotment system of honeybee colonies therefore needs a new, extended definition. Nurses are largely responsible for preparing nutrients from pollen, which is difficult to digest. They then distribute the nutritionally valuable protein produced by their hypopharyngeal glands to practically all hive mates.Dedicated to Professor Dr. O. Kepka on the occasion of his 65th birthday     

The queen in relation to wax secretion and comb building in honeybees

Summary The role of the queen in relation to wax secretion and comb building in honeybees was analyzed with respect to queen status (mated, virgin and dead queens and queenlessness), and pheromones of the head and abdominal tergite of queens. Worker variables considered were colony size, percentage of bees bearing wax scales, wax scale weight, and weight of constructed combs.The amount of wax recovered from festoon bees and the percentage of festoon bees bearing wax were independent of queen status, the pheromones of queens and access to the queen. Colonies with full access to freely moving mated queens always constructed significantly more comb than those headed by virgin or dead queens as well as all permutations of caged and division board queens whose mandibular glands and/or abdominal tergite glands were operative or not.Despite pheromonal similarity of virgin queens to mated ones, colonies headed by virgin queens constructed as little comb as did queenless colonies. The bouquets of the mandibular glands did not differ significantly among queens nor was the amount of comb constructed correlated with pheromonal bouquet. Comb building is greatest among colonies having full access to freely moving queens but the stimulus for such building is not attributable to the 90DA, 9HDA and 10HDA components of the queen's mandibular gland secretions.     

The relationship between population size,amount of brood,and individual foraging behaviour in the honey bee,Apis mellifera L.

This study experimentally examines the relationship between colony state and the behaviour of individual pollen and nectar foragers in the honey bee, Apis mellifera L. In the first experiment we test the prediction that individual pollen foragers from colonies with higher brood quantities should exhibit a greater work effort for pollen resources than individual pollen foragers from colonies with low brood quantities. Eight colonies were assigned into two treatment groups; HIGH brood colonies were manipulated to contain 9600±480 cm² brood area; LOW brood colonies were manipulated to contain 1600±80 cm² brood area. We measured colony brood levels over the course of the experiment and collected individual pollen loads from returning pollen foragers. We found that, while colonies remained significantly different in brood levels, individual pollen foragers from HIGH brood colonies collected larger loads than individuals from LOW brood colonies. In the second experiment we investigated the influence of colony size on the behaviour of individual nectar foragers. We assigned eight colonies to two treatment groups; LARGE colonies were manipulated to contain 35000±1700 adult workers with 3500±175 cm² brood area, and SMALL colonies were manipulated to contain 10000±500 adult workers with 1000±50 cm² brood area. We observed foraging trips of individually marked workers and found that individuals from LARGE colonies made longer foraging trips than those from SMALL colonies (LARGE: 1666.7±126.4 seconds, SMALL: 1210.8±157.6 seconds), and collected larter nectar loads (LARGE: 19.2±1.0 l, SMALL: 14.6±0.8 l). These results indicate that individual nectar foragers from LARGE colonies tend to work harder than individuals from SMALL colonies. Both experiments indicate that the values of nectar and pollen resources to a colony change depend on colony state, and that individual foragers modify their behaviour accordingly.     

The effect of diet on protein concentration, hypopharyngeal gland development and virus load in worker honey bees (Apis mellifera L.)

Elucidating the mechanisms by which honey bees process pollen vs. protein supplements are important in the generation of artificial diets needed to sustain managed honeybees. We measured the effects of diet on protein concentration, hypopharyngeal gland development and virus titers in worker honey bees fed either pollen, a protein supplement (MegaBee), or a protein-free diet of sugar syrup. Workers consumed more pollen than protein supplement, but protein amounts and size of hypopharyngeal gland acini did not differ between the two feeding treatments. Bees fed sugar syrup alone had lower protein concentrations and smaller hypopharyngeal glands compared with the other feeding treatments especially as the bees aged. Deformed wing virus was detected in workers at the start of a trial. The virus concentrations increased as bees aged and were highest in those fed sugar syrup and lowest in bees fed pollen. Overall results suggest a connection between diet, protein levels and immune response and indicate that colony losses might be reduced by alleviating protein stress through supplemental feeding.     

Division of labor inPonera pennsylvannica (Formicidae: Ponerinae)

Summary We examined division of labor and colony demography in the antPonera pennsylvannica. Observation of three colonies with individually marked workers revealed a high degree of interindividual behavioral variation and a rough but consistent division of labor between brood tenders and foragers. This division was present both in colonies consisting entirely of workers produced in the previous summer and in colonies containing freshly eclosed ants. Two colonies showed typical age-based polyethism, with young ants focusing on brood care and overwintered ants on foraging. No such age basis was detected in the third colony. This difference may relate to variability in brood production schedules. Colonies showing temporal polyethism had two peaks of brood production and thus had relatively large brood populations when the first young workers eclosed, while the third colony had only one peak and little brood for the young workers to tend. Even if young ants have a lower threshold for brood care, it may have been concealed in the latter situation. Demographic data indicate that natural colonies produce one brood per year and that workers typically eclose into colonies with relatively low brood care demands. This suggests that overwintered workers do most of a colony's work and that the division of labor among overwintered ants is the more important one under natural conditions. The basis of this division is as yet unknown. These results also suggest that small colony size, univoltine brood schedule and a close association between foraging and brood care do not preclude division of labor among specialized castes, as has been suggested for another ponerine species (Traniello 1978).     

Eine schnelle biologische Methode zum Nachweis der Phytotoxizität von Wirkstoffen in Pestiziden mit Hilfe eines Blattaufstiegtestes bei Sinapis alba L. Kotyledonen: (Kurze Mitteilung)

The present work was carried out to study the relationship between brood rearing activity, bee density and stored pollen grains. Fifteen colonies, 1st. Carniolan hybrid nearly in equal strength were put under investigation from June 1994 to May 1995.

The results revealed that the major peak of brood activity and higher rate of stored pollen were in May, consequently the maximum number of house bees was recorded during June and July. However no correlation existed, the lowest production of brood, bees and gathered pollen occurred during February. It was found that high significant differences were in quantity of brood and house bees, while low significant differences were in quantity of stored pollen. Correlations between brood activity, house bees density and stored pollen were observed in winter.     

Comparison of the activity and productivity of Carniolan (Apis mellifera carnica Pollmann) and Yemeni (Apis mellifera jemenitica Ruttner) subspecies under environmental conditions of the Al-Ahsa oasis of eastern Saudi Arabia

This study was conducted at the apiary of the Agricultural and Veterinary Training and Research Station of King Faisal University in the Al-Ahsa oasis of eastern Saudi Arabia. We performed a comparison between Carniolan (Apis mellifera carnica Pollmann) and Yemeni (Apis mellifera jemenitica Ruttner) honeybee races to determine the monthly fluctuations in foraging activity, pollen collection, colony growth and honey yield production under the environmental conditions of the Al-Ahsa oasis of eastern Saudi Arabia. We found three peaks in the flight activity of the two races, and the largest peaks occurred during September and October. Compared to Carniolan bee colonies, the performance of Yemeni bee colonies was superior in terms of stored pollen, worker and drone brood rearing, and the adult population size. The Carniolan bee colonies produced 27.77% and 27.50% more honey than the Yemeni bee colonies during the flow seasons of alfalfa and sidir, respectively, with an average increase of 27.64%. It could be concluded that the race of bees is an important factor affecting the activity and productivity of honeybee colonies. The Yemeni bee race produced more pollen, a larger brood and more bees, which exhibited a longer survival. The imported Carniolan bees can be reared in eastern Saudi Arabia, but the Yemeni bee race is still better.     

The relationship between comb age and performance of honey bee (Apis mellifera) colonies

A study on the relationship between the age of comb and the activity of the hybrid Carniolan honey bee colonies in collecting pollen activity, worker brood production, colony strength, and honey yield was conducted. In comparison to colonies with combs aged 4-years, colonies with combs aged 1, 2 and 3-years significantly exceeded in the number returning workers, number returning workers with pollen loads, rate of storing pollen, rate of worker brood production, and size of colony population. Colonies with combs aged 1, 2 and 3-years produced significantly more honey than colonies with combs aged 4-years (5.25, 4.90 and 4.65 kg/colony vs. 4.45 kg/colony, respectively). It can be concluded that the foraging rate, gathering and storing pollen, brood production, colony population size, and honey yield significantly depended on the age of combs. Beekeepers can replace old combs with new ones to increase brood and honey production.     

Body size differs in workers produced across time and is associated with variation in the quantity and composition of larval food in <Emphasis Type="Italic">Nannotrigona perilampoides</Emphasis> (Hymenoptera,Meliponini)

Although variation in body size has been recently reported in stingless bees (Meliponini), empirical evidence evaluating possible factors related to such variation is lacking, and thus it is not clear if it may have an adaptive significance. We evaluated if variation in the body size and weight of workers of stingless bees fluctuates across a seasonal pattern and if this could be related to characteristics of the food consumed during the larval stage. The weight of larval provisions, their protein, and sugar content were evaluated in four colonies of Nannotrigona perilampoides every 2 months across 1 year. Worker-destined larvae from the same combs were allowed to develop and were sampled as callow workers to determine their weight and size using morphometric data. The weight and size of workers were highly correlated and varied across the seasons in established colonies, suggesting that size variation cycles across the year in stingless bees. An increase in the protein content and, to a lesser degree, the quantity of larval food were positively linked to variation in body weight and size; food with richer protein content resulted in larger and heavier workers. This study provides the first evidence of an effect of the quantity and composition of larval food on the size of workers in stingless bees. Although body weight and size of workers differed across seasons, they were not readily noticeable as changes seem to occur as a continuum across the year. Since size polymorphism was of a larger magnitude across time but not within age cohorts and as it was highly determined by food resources, it may not be an adaptive feature in stingless bees. However, more studies are needed to determine the role of the cyclical change in worker body size on colony performance and thus its adaptive significance in stingless bees.     

Colony pollen reserves affect body size,sperm production and sexual development in males of the stingless bee <Emphasis Type="Italic">Melipona beecheii</Emphasis>

The production of male sexual offspring by social insect colonies is often strongly seasonal or resource-dependent. In stingless bees, males are produced in smaller numbers under conditions of low colony food reserves; whether such males are negatively affected in traits related to reproductive success is not known. We compared body size, sperm production and sexual maturity in Melipona beecheii males reared with experimentally supplemented or reduced pollen reserves, but with otherwise equal numbers of workers and equal quantities of honey reserves. We also studied the same traits in males collected from non-manipulated colonies with pollen reserves intermediate between the supplemented or reduced groups but with more workers and honey reserves. Males reared under experimentally reduced pollen reserves had significantly smaller bodies and lower sperm counts compared to those reared in colonies with experimentally supplemented pollen reserves. There was also a significantly positive relationship between the number of sperm and body size in males across all colony treatments. The maximum number of sperm in seminal vesicles was recorded 2 days later in males from colonies with reduced pollen compared to males from colonies with supplementary pollen. Males from non-manipulated colonies were intermediate in size, sperm count and speed of maturation. Our study documents for the first time the existence of large size variation in males of stingless bees that is related with the amount of pollen reserves in their natal colony. We conclude that a colony’s pollen reserves have a major impact on male body size, sperm production and speed of sexual maturity in this stingless bee, which may be the case in other social insects. Stingless bees are a good model system to study the balance between colony-level selection and individual-level selection on male sexually selected traits such as body size.     

Managing honey bees (Hymenoptera: Apidae) for greenhouse tomato pollination

Although commercially reared colonies of bumble bees (Bombus sp.) are the primary pollinator world-wide for greenhouse tomatoes (Lycopersicon esculentum Mill.) previous research indicates that honey bees (Apis mellifera L.) might be a feasible alternative or supplement to bumble bee pollination. However, management methods for honey bee greenhouse tomato pollination scarcely have been explored. We 1) tested the effect of initial amounts of brood on colony population size and flight activity in screened greenhouses during the winter, and 2) compared foraging from colonies with brood used within screened and unscreened greenhouses during the summer. Brood rearing was maintained at low levels in both brood and no-brood colonies after 21 d during the winter, and emerging honey bees from both treatments had significantly lower weights than bees from outdoor colonies. Honey bee flight activity throughout the day and over the 21 d in the greenhouse was not influenced by initial brood level. In our summer experiment, brood production in screened greenhouses neared zero after 21 d but higher levels of brood were reared in unscreened greenhouses with access to outside forage. Flower visitation measured throughout the day and over the 21 d the colonies were in the greenhouse was not influenced by screening treatment. An economic analysis indicated that managing honey bees for greenhouse tomato pollination would be financially viable for both beekeepers and growers. We conclude that honey bees can be successfully managed for greenhouse tomato pollination in both screened and unscreened greenhouses if the foraging force is maintained by replacing colonies every 3 wk.     

Functional flexibility of the honey bee hypopharyngeal gland in a dequeened colony

The role of the worker honey bee Apis mellifera L. changes depending on age after eclosion (age polyethism): young workers (nurse bees) take care of their brood by synthesizing and secreting brood food (royal jelly), while older workers (foragers) forage for nectar and process it into honey. Previously, we showed that the major proteins synthesized in the hypopharyngeal gland of the worker change from brood food proteins to alpha-glucosidase at the single secretory cell level in parallel with this age polyethism [Kubo et al., J. Biochem. 119, 291-295 (1996); Ohashi et al., Eur. J. Biochem. 249, 797-802 (1997)]. Here, we examined whether the function of the hypopharyngeal gland has flexibility depending on the colony conditions, by creating a dequeened colony in which the older workers were compelled to feed the drone larvae. It was found that most of the older workers in the dequeened colony synthesized brood food proteins as did nurse bees. Furthermore, the percentage of workers that synthesized brood food proteins was maintained at 80-90% of the total workers for at least two months, as in a normal colony. These results indicate that the function of the hypopharyngeal gland cells of the worker has flexibility and can, if necessary, be maintained as that of the nurse bee, depending on the condition of the colony.