From where did the Western honeybee (Apis mellifera) originate?

The native range of the honeybee Apis mellifera encompasses Europe, Africa, and the Middle East, whereas the nine other species of Apis are found exclusively in Asia. It is therefore commonly assumed that A. mellifera arose in Asia and expanded into Europe and Africa. However, other hypotheses for the origin of A. mellifera have also been proposed based on phylogenetic trees constructed from genetic markers. In particular, an analysis based on >1000 single-nucleotide polymorphism markers placed the root of the tree of A. mellifera subspecies among samples from Africa, suggestive of an out-of-Africa expansion. Here, we re-evaluate the evidence for this and other hypotheses by testing the robustness of the tree topology to different tree-building methods and by removing specimens with a potentially hybrid background. These analyses do not unequivocally place the root of the tree of A. mellifera subspecies within Africa, and are potentially consistent with a variety of hypotheses for honeybee evolution, including an expansion out of Asia. Our analyses also support high divergence between western and eastern European populations of A. mellifera, suggesting they are likely derived from two distinct colonization routes, although the sources of these expansions are still unclear.     

Toxins induce ‘malaise’ behaviour in the honeybee (Apis mellifera)

To avoid poisoning and death when toxins are ingested, the body responds with a suite of physiological detoxification mechanisms accompanied by behaviours that in mammals often include vomiting, nausea, and lethargy. Few studies have characterised whether insects exhibit characteristic ‘malaise-like’ behaviours in response to intoxication. Here, we used the honeybee to investigate how intoxication produced by injection or ingestion with three toxins with different pharmacological modes of action quinine, amygdalin, and lithium chloride affected behaviour. We found that toxin-induced changes in behaviour were best characterised by more time spent grooming. Bees also had difficulty performing the righting reflex and exhibited specific toxin-induced behaviours such as abdomen dragging and curling up. The expression of these behaviours also depended on whether a toxin had been injected or ingested. When toxins were ingested, they were least 10 times less concentrated in the haemolymph than in the ingested food, suggesting that their absorption through the gut is strongly regulated. Our data show that bees exhibit changes in behaviour that are characteristic of ‘malaise’ and suggest that physiological signalling of toxicosis is accomplished by multiple post-ingestive pathways in animals.     

Pattern vision of the honeybee (Apis mellifera). What is an oriented edge?

Pairs of black patterns on a white background, one rewarded the other not, were presented vertically each in one arm of a Y-maze. During training the locations of the black areas were changed every 5 min to prevent the bees using them as cues, but cues from edges were kept consistent. Bees detect orientation even in a gradient that subtends 36° from black to white (normal to the edge). Orientation cues in short lengths of edge are detected and summed on each side of the fixation point, irrespective of the lay-out of the pattern. Edges at right angles reduce the total orientation cue. The polarity of edges in a sawtooth grating is weakly discriminated, but not the orientation of a fault line where two gratings meet. Edge quality can be discriminated, but is not recognised in unfamiliar orientations. When spot location is excluded as a cue, the orientation of a row of spots or squares which individually provide no net orientation cue is not discriminated. In conclusion, when locations of black areas are shuffled, the bees remember the sum of local orientation cues but not the global pattern, and there is no re-assembly of a pattern based on differently oriented edges. A neuronal model consistent with these results is presented. Accepted: 5 March 2000     

Pattern discrimination by the honeybee (Apis mellifera ) is colour blind for radial / tangential cues

Bees were trained to discriminate between two patterns, one of which was associated with a reward, in a Y-choice apparatus with the targets presented vertically at a distance at an angular subtense of 50°. Previous work with this apparatus has found discrimination between two patterns of coloured gratings or radial sectors that are fixed in different orientations during the training. When there was contrast to the blue receptors alone, gratings of period 6° were resolved, and 4° when there was contrast to the green receptors. In the present work, bees discriminate between a pattern containing tangentially arranged edges and one containing radially arranged edges, both with no average edge orientation. The targets were rotated every 5 min to make the locations of areas useless as cues. The edges remained consistently radial or tangential and were therefore the only cues. Tests with patterns of selected colours and various levels of grey show that for each colour there is a level of grey at which discrimination fails. Discrimination is therefore colour-blind. The same patterns were made with combinations of coloured papers that give no contrast to the green receptors or alternatively to the blue receptors. The bees discriminate only if the edges between colours present a contrast to the green receptors. The system that discriminates generalized radial and tangential cues is therefore colour blind because the inputs are restricted to the green receptors, not because receptor outputs are added together. The same result was obtained with a very coarse pattern of period 20°. Accepted: 10 January 1999     

Why acquiesce? Worker reproductive parasitism in the Eastern honeybee (Apis cerana)

Most societies are vulnerable to rogue individuals that pursue their own interests at the expense of the collective entity. Societies often protect themselves from selfish behaviour by ‘policing’, thereby enforcing the interests of the collective over those of individuals. In insect societies, for example, selfish workers can activate their ovaries and lay eggs, exploiting the collective brood rearing system for individual benefit. Policing, usually in the form of oophagy of worker‐laid eggs, controls selfish behaviour. Importantly, once an effective system of policing has evolved, the incentive for personal reproduction is lost, and ‘reproductive acquiescence’ in which ovary activation is rare or absent is predicted to evolve. Studies of social Hymenoptera have largely supported the prediction of worker ‘acquiescence’; workers of most species where policing is well developed have inactive ovaries. However, the eastern honeybee Apis cerana appears to be an exception. A. cerana colonies are characterized by highly efficient policing, yet about 5% of workers have active ovaries, even when a queen is present. This suggests that the evolution of acquiescence is incomplete in A. cerana. We regularly sampled male eggs and pupae from four A. cerana colonies. Workers had high levels of ovary activation overall (11.7%), and 3.8% of assignable male eggs and 1.1% of assignable male pupae were worker‐laid. We conclude that workers with active ovaries lay their eggs, but these rarely survive to pupation because of intense policing. We then used our findings as well as previously published data on A. cerana and A. mellifera to redo the meta‐analysis on which reproductive acquiescence theory is based. Including data on both species did not affect the relationship between effectiveness of policing and levels of worker reproduction. Their inclusion did, however, seriously weaken the relationship between relatedness among workers and levels of worker reproduction. Our work thus suggests that relatedness among workers does not affect the probability that workers will attempt to reproduce, but that it is coercion by peers that limits worker reproduction.     

Individual versus social pathway to honeybee worker reproduction (Apis mellifera): pollen or jelly as protein source for oogenesis?

Honeybee workers, Apis mellifera, can reproduce in queenless colonies. The production of queen-like pheromones may be associated with their reproductive activity and induce nestmates to respond by feeding them. Such frequent trophallaxis could supply their protein needs for oogenesis, constituting a social pathway to worker reproduction. However, some individuals can develop ovaries without producing queen pheromones. The consumption of protein-rich pollen could be an alternative solitary pathway for them to satisfy this dietary requirement. In order to investigate the way in which workers obtain proteins for oogenesis, we created orphaned worker groups and determined ovarian and pheromonal development in relation to pollen consumption of selected workers. Individuals that did not consume pollen had significantly more developed ovaries and produced significantly more queen mandibular pheromone than workers that fed directly on pollen. Our results suggest that workers producing queen-like secretions are fed trophallactically. However, reproductive workers that lacked queen pheromones had consumed little or no pollen, suggesting that they also obtained trophallaxis. Although pollen consumption might contribute to sustaining oogenesis, it does not appear to be sufficient. Trophallaxis as a means of obtaining proteins seems to be necessary to attain reproductive status in queenless honeybee colonies.     

The control of the proventriculus in the honeybee (Apis mellifera carnica L.) I. A dynamic process influenced by food quality and quantity?

The control of crop emptying in foraging honeybees was investigated in individuals trained to collect defined amounts of sugar solutions. Following feeding, they were dissected after fixed periods of time in order to measure crop content and haemolymph sugar titers. Between feeding and dissection, the metabolic rate of every investigated forager was measured using open-flow respirometry, so as to assess the effects of both food quality (concentration, molarity and viscosity of the fed sugar solution) and food quantity on the transport rate through the proventriculus. The sugar transport rate through the proventriculus was observed to be mainly dependent on the metabolic expenditure of the individual. Bee foragers were able to precisely adjust the sugar transport rate to their metabolic rates, but under certain conditions, an excess of sugars was transported through the proventriculus, more than needed to cover the bee’s energetic demands. This excess depended on the nutritive value and quantity of the fed sugar solution, and on the time after feeding. It did not depend on the metabolic rate of the bee, the molarity, or the viscosity of the fed sugar solution. As long as the bees did not exhaust their crop contents, the haemolymph sugar titers were unaffected by this excess amount transported, by the time after feeding, the concentration and the viscosity of the fed sugar solution. For all feeding conditions assayed, the haemolymph trehalose titer remained constant, while the titers of other haemolymph sugars varied. It is suggested that the trehalose concentration in the haemolymph is regulated in honeybees, and that it represents the controlled variable in the feedback loop responsible for the transport rate through the proventriculus.     

The nearer the better? Drones (Apis mellifera) prefer nearer drone congregation areas

Summary. At 2 drone congregation areas (DCA) the relation between drone presence and distance to the apiary of origin was studied. Two methods were applied. First, drones were caught and marked on the DCA and later recovered in the colonies. Second, drones which were marked before at the apiary (in the colonies) were subsequently recaptured on both DCAs. The 2 methods led to identical conclusions. Consistently in each of 3 years the majority of the drones from each of the 3 apiaries was found at the nearer DCA. There was, however, no direct correlation between the flight distances and the ratio of drones visiting from each apiary. Thus some other factors (attractiveness of the DCA) may also have influenced the choice of the drones. Our findings support the idea that there is an orientation phase during which drones explore several DCAs before each drone stays at 1 DCA, and energetic choices made by drones in relation to flight distances seem to be important. The choice of the nearer DCA would permit the drone to prolong his presence at the DCA and increase his chances to mate: the nearer the better!The drones strategy to chose the nearer DCA would boost the genetic representation of local colonies and this drone clumping would increase genetic differences among the DCAs within an area. In this context the choice of virgin queens among DCAs is of great significance.Received 1 December 2003; revised 9 April 2004; accepted 7 June 2004.     

Do feral honey bees (Apis mellifera) and regent parrots (Polytelis anthopeplus) compete for nest sites?

Abstract Surveys of nesting sites of feral honey bees (Apis mellifera) and regent parrots (Polytelis anthopeplus) were made in the red gum/black box woodlands of Wyperfeld National Park, Victoria, Australia. Data on tree species and size, and number of hollows were collected from all trees within seven 500 × 100m plots. Nest site characteristics were quantified for both bees and parrots. We found 27 feral honey bee colonies, suggesting a density of 77.1 colonies per km². The average occupation rate for bees was 1.3% of trees and 0.7% of available hollows. The height, aspect and entrance characteristics of honey bee nests at Wyperfeld were not qualitatively different to those reported elsewhere. We found 15 pairs of nesting regent parrots. Nest sites chosen by these birds overlapped those chosen by honey bees, but 52% of bee nests were in cavities unsuitable for regent parrots. We suggest that honey bee population growth may be limited in the park by a lack of water.     

Ultrastructural characterization of the ovarian stages of the worker bee, Apis mellifera mellifera Linné (Hymenoptera, Apidae)

The classification of the ovarian stages remains still vague. The ultrastructure of the basal follicle made it possible to solve this problem. From the sixth to the eighth day of the worker bee's adult life, the length of the ovariole hardly exceeds 2 mm. The basal oocytes of stages 1 and 2 are characterized by the presence of many ranges of granules encircled by the dilated cylinders of the granular endoplasmic reticulum. These cylinders decondensate the proteinic granules and the lipidic globules generating multivesicular bodies. The basal oocyte, at the third stage of development, measures 0.35+/-0.05 mm in length. This stage of the previtellogenesis' end is characterized by the appearance, in the ooplasm, of many tubules, whose dilated end constitutes a lysosome. At the beginning of the second week, vitellogenesis starts. Stage 4 is defined by the appearance, in the ooplasm, of the yellow granules in relationship to the tubular formations. At the 14th to 16th days of the adult life, 25+/-7.1% of the worker bees reach the end vitellogenesis, or stage 5. The ovariole, about 7 mm in length, contains 11 follicles. This stage 5 is distinguished by the synthesis from the vitelline membrane, the regression of the size of yellow granules and the accessory cores. The sixth stage or oocyte maturation is reached by 12.5+/-3.5% of the workers aged 18 to 20 days. It is defined by the presence of the chorion, the dissolution of the yellow granules, the disappearance of the accessory nuclei and the total degeneration of the follicular cells.     

Mating for convenience or genetic diversity? Mating patterns in the polygynous ant Plagiolepis pygmaea

Several genetic and nongenetic benefits have been proposed toexplain multiple mating (polyandry) in animals, to compensatefor costs associated with obtaining additional mates. The mostprominent hypotheses stress the benefits of increased geneticdiversity. In social insects, queens of most species mate onlyonce or have effective mating frequencies close to one. Yet,in a few species of ants, bees, and wasps, polyandry is therule. In these species, colonies are usually headed by a singlequeen, whereas multiple queening adds diversity in several ofthe remaining species, especially in ants. Here we investigatedmating frequency, inbreeding and relatedness between the queensand their mates in the polygynous ant Plagiolepis pygmaea, andthe effect of polyandry on the genetic diversity as a functionof the effective population size of individual colonies. Ourresults show that polyandry occurs frequently in the species.However, queens are frequently inseminated by close relatives,and additional sires add little genetic diversity among offspringof individual queens. In addition, the increase in diversityat the colony level is only marginal. Hence, contrary to establishednotions, polyandry in P. pygmaea seems not to be driven by substantialbenefits of genetic diversity. Nonetheless, very small or asyet unidentified genetic benefits to one party (males, workers,queens) in conjunction with low costs of mating may favor polyandry.Alternatively, nongenetic factors, such as convenience polyandry,may be more important than genetic factors in promoting polyandryin P. pygmaea.     

意大利蜜蜂(Apis mellifera ligustica)与中华蜜蜂(Apis cerana ceraca)的生态位比较

为了明确中华蜜蜂和意大利蜜蜂在皖南山区生态适应性,研究两种蜜蜂的食物生态位、时间生态位和空间生态位及其差异,结果是中蜂与意蜂食物(蜜源植物)资源生态位宽度分别是 0.923、0.765,中蜂对蜜源植物采集喜好性差异小,而意蜂差异大,中蜂对意蜂生态位重迭为0.160,中蜂对意蜂生态位相似性为0.755;油菜花期,中蜂与意蜂时间资源生态位宽度分别是0.879、0.801,枇杷花期,分别是0.760、0.677,中蜂与意蜂的空间资源生态位宽度分别是0.797、0.670.中蜂3种生态位宽度均大于意蜂,中蜂三维生态位值是意蜂的1.61倍和1.57倍.表明中蜂在皖南山区生态适应性比意蜂强.     

E-β-ocimene, a volatile brood pheromone involved in social regulation in the honey bee colony (Apis mellifera)

BACKGROUND: In honey bee colony, the brood is able to manipulate and chemically control the workers in order to sustain their own development. A brood ester pheromone produced primarily by old larvae (4 and 5 days old larvae) was first identified as acting as a contact pheromone with specific effects on nurses in the colony. More recently a new volatile brood pheromone has been identified: E-β-ocimene, which partially inhibits ovary development in workers. METHODOLOGY AND PRINCIPAL FINDING: Our analysis of E-β-ocimene production revealed that young brood (newly hatched to 3 days old) produce the highest quantity of E-β-ocimene relative to their body weight. By testing the potential action of this molecule as a non-specific larval signal, due to its high volatility in the colony, we demonstrated that in the presence of E-β-ocimene nest workers start to forage earlier in life, as seen in the presence of real brood. CONCLUSIONS/SIGNIFICANCE: In this way, young larvae are able to assign precedence to the task of foraging by workers in order to increase food stores for their own development. Thus, in the complexity of honey bee chemical communication, E-β-ocimene, a pheromone of young larvae, provides the brood with the means to express their nutritional needs to the workers.     

Mating system and nesting biology of the Red-necked Phalarope Phalaropus lobatus: what constrains polyandry?

In this five-year study of Red-necked Phalaropes on the coast of Hudson Bay, Canada, only 8% (5/59) of females were polyandrous. Seventy-five percent were monogamous, and 17% did not obtain mates. All males obtained mates readily. To understand the low incidence of polyandry in this sex-role reversed, non-territorial mating system, aspects of the pair bond, nesting biology, and size and plumage dimorphism were examined.
Pair bonds lasted an average of 11 days and ended as soon as the clutch was completed. Polyandrous females initiated their second nests only seven days later. Since in most years nest initiation for the population as a whole spanned three weeks, there was ample time for multiple clutches. Energetic constraints may have played a limited role in constraining polyandry, but many females that had previously laid clutches courted incubating males or harassed pairs.
The major factor limiting polyandry appeared to be availability of mates. Because phalaropes generally breed in habitats that are less defensible than those of other role-reversed shorebirds, they are less able to monopolize males in populations with equal sex ratios. Sexual size dimorphism is correspondingly smaller than in territorial role-reversed shorebirds, although plumage variation between and within sexes in this study was strong (and invariant with age). Thus, despite the phalaropes' role-reversal, the evidence indicates that monogamy may predominate, with sexual selection driven by the ability of females to produce second clutches opportunistically when receptive males are available.     

Polliniferous plants and foraging strategies of Apis mellifera (Hymenoptera: Apidae) in the Yucatán Peninsula,Mexico

A study of the most important polliniferous plants for European and Africanized honeybees (Apis mellifera L.) was made in Quintana Roo state. Comparisons were made between the plants visited by both bee types in order to determine whether there were qualitative or quantitative differences in their choice of plant species. Also some foraging strategies of the honeybees were analysed. Pollen from pollen load samples was acetolysed and mounted on slides. Subsequently the pollen grains were identified, counted and photographed. A total of 206 pollen load samples were collected at Palmas and St. Teresa during two years. The most frequent species in the pollen load samples from European and Africanized honeybees were Cecropia peltata, Metopium brownei, Lonchocarpus sp. 2, Viguiera dentata, Eragrostis sp. 1, Bursera simaruba and Eupatorium albicaule. Both types of honey bees show a high reliance on pollen from only a few species, the first five named above comprised around 50% of all the mean percentage frequencies. Families that contributed with the largest number of pollen species were Fabaceae, Asteraceae, Boraginaceae, Convolvulaceae, Euphorbiaceae, Sapindaceae, Poaceae, Myrtaceae, Sapotaceae and Tiliaceae. C. peltata, Trema micrantha, B. simaruba, Eugenia sp. 1, Thouinia canesceras, Pouteria sp. 1, Mimosa bahamensis and V. dentata, were the pollen species with the largest percentages of occurrence in both European and Africanized bee pollen load samples, and also represent a "long-term" food resources during the year.     

Does fumagillin control the recently detected invasive parasite Nosema ceranae in western honey bees (Apis mellifera)?

Western honey bee (Apis mellifera) colonies in Nova Scotia, Canada were sampled in spring and late summer 2007 to evaluate efficacy of fumagillin dicyclohexylammonium (hereafter, fumagillin) against Nosema ceranae. Colonies treated with fumagillin in September 2006 (n = 94) had significantly lower Nosema intensity in spring 2007 than did colonies that received no treatment (n = 51), but by late summer 2007 no difference existed between groups. Molecular sequencing of 15 infected colonies identified N. ceranae in 93.3% of cases, suggesting that fumagillin is successful at temporarily reducing this recent invasive parasite in western honey bees.     

分离自蜜蜂(Apis mellifera)的三株螺原体的基本特性

【目的】调查我国蜜蜂螺原体的种类,研究它们的基本生物学特性,初步确定其分类地位,为研究螺原体在自然界中的传播途径提供依据。【方法】螺原体的分离、培养方法,应用暗视野显微镜和透射电子显微镜观察螺原体形态,运用分子生物学方法(选16S rDNA、ITS、rpoB基因进行系统发育分析)和血清学方法(生长抑制试验、代谢抑制试验、菌体变形试验)研究螺原体分离菌株可能的分类地位。【结果】从健康的意蜂(Apis mellifera)体内分离到3株螺原体MF0903、MF0904、MF0905。3株螺原体都呈典型的螺旋状,但菌株MF0905的菌体短小,螺旋数较少;MF0903和MF0904菌落呈规则的圆形,MF0905菌落近圆形、较大;它们都能利用葡萄糖、D-果糖作为碳源,不能利用尿素;菌株MF0903、MF0904能强烈代谢精氨酸、不能利用蔗糖作为碳源,而MF0905不能代谢精氨酸、能利用蔗糖作为碳源;根据16S rDNA、ITS、rpoB基因序列构建系统发育树显示,分离菌株MF0903、MF0904与Spiroplasma melliferum聚类较近,而MF0905与Spiroplasma clarkii聚类较近。生长抑制试验、代谢抑制试验、菌体变形试验结果均表明标准菌株Spiroplasma melliferum CH-1的抗血清对菌株MF0905没有抑制作用,而能抑制菌株MF0903和MF0904生长。【结论】分离菌株MF0903、MF0904属于Spiroplasma melliferum,而MF0905可能是Spiroplasma clarkii,这表明我国蜜蜂中存在的螺原体不仅仅是Spiroplasma melliferum。