Decentralized control of drone comb construction in honey bee colonies

Honey bee colonies furnish their nests with two types of comb distinguished by cell size: large cells for rearing males (drone comb) and small cells for rearing workers (worker comb). The bees actively regulate the relative quantity of each type, a behavior likely to be important in setting a colony's sex ratio. Experimental analysis of the information pathways and control mechanisms responsible for this regulation found the following results. The amount of drone comb in a nest is governed by negative feedback from drone comb already constructed. This feedback depends on the workers having direct contact with the drone comb in their nest, but does not depend on the queen's contact with the comb. The comb itself, rather than the brood within it, is sufficient to provide the negative feedback, although the brood may also contribute to the effect. These findings show that drone comb regulation does not depend on the queen acting as a centralized information gatherer and behavioral controller. Instead, the evidence points to a decision-making process distributed across the population of worker bees, a control architecture typical of colony organization in honey bees and other large-colony insect societies. Received: 24 May 1997 / Accepted after revision: 30 August 1997     

阿维菌素对意大利蜜蜂(Apis mellifera L.)的毒性影响

通过单次饲喂高浓度阿维菌素药液以及连续饲喂亚致死浓度阿维菌素药液进行意大利蜜蜂的经口染毒,从而探讨阿维菌素对意大利蜜蜂的急性及慢性毒性影响。结果表明,阿维菌素对意大利蜜蜂急性经口毒性48 h半数致死剂量(48 h-LD50)为0.00700μg a.i.·蜂-1,慢性经口毒性240 h每日半数致死剂量(240 h-LDD50)为0.000308μg a.i.·蜂-1·天-1。在亚致死效应方面,0.0233 mg a.i.·L-1和0.0467 mg a.i.·L-1处理组在168 h后摄食量出现明显的减少,说明阿维菌素中毒已经严重影响意大利蜜蜂的觅食和摄食能力。同时,由于摄食量的下降以及阿维菌素的毒性作用,造成了0.0467 mg a.i.·L-1处理组意大利蜜蜂体重的大幅度下降,试验前后的体重变化率达到-54.84%。意大利蜜蜂爬行能力的测定结果显示,各处理组的爬行通过率均低于对照组,特别是0.0117 mg a.i.·L-1处理组、0.0233 mg a.i.·L-1处理组和0.0467 mg a.i.·L-1处理组(P0.05)。综上所述,阿维菌素对意大利蜜蜂的急性经口毒性为高毒,较高剂量染毒会引起意大利蜜蜂的直接死亡;此外,长期接触较低浓度的阿维菌素,一方面会损害意大利蜜蜂的运动能力,如爬行、飞行能力的减弱;另一方面意大利蜜蜂生理方面也会遭到威胁,表现为摄食量下降、体重减轻,甚至死亡。因此,在施用该农药时应尽量避开蜜蜂栖息地,同时避免在蜜源植株花期时施用。     

Evolution of extreme polyandry in the honeybee Apis mellifera L.

A number of hypotheses have been proposed to explain the evolution of multiple mating in the honeybee queen. In particular, the consequences of reduced intracolonial relatedness provide plausible explanations for multiple mating with up to ten drones, but fail to account for the much higher mating frequencies observed in nature. In this paper, we propose an alternative mechanism which builds on non-linear relationships between intracolonial frequencies in genotypic worker specialization and colony fitness. If genes for any worker specialization confer an advantage on colony fitness only when they are rare, this would require a stable mix of sperm from a few drones which contribute that trait, and many which do not. To ensure both specific, low within-colony proportions of “rare specialist” genes, and to reduce random variation of these proportions would require mating with high numbers of drones. The quantitative implementation shows that moderate to very high numbers of matings are required to exploit colony advantages from genotypic allocation of workers to rare tasks. Extreme polyandry thus could result from colony selection dependent on the intracolonial frequency of rare genetic specialists. Received: 30 January 1998 / Accepted after revision: 7 October 1998     

4种烟嘧磺隆多元复配除草剂对意大利蜜蜂和玉米螟赤眼蜂的急性毒性及初级风险评估

为明确4种烟嘧磺隆复配除草剂对意大利蜜蜂和玉米螟赤眼蜂的急性毒性影响并评估其初级风险。采用饲喂法、点滴法和玻璃管药膜法,分别测定了8%烟嘧·氯吡嘧磺隆、16%烟嘧·硝磺·氯吡嘧磺隆、36%烟嘧·莠去津和22%烟嘧·氯吡·氯氟吡氧乙酸异辛酯对意大利蜜蜂成年工蜂和玉米螟赤眼蜂的急性毒性,并分别采用危害商值和安全系数进行初级风险评估。结果表明,上述4种制剂对意大利蜜蜂成年工蜂的经口毒性48 h-LD_(50)(半致死剂量,median lethal dose)分别为68.03、1.60×10~2、1.40×10~2和100μg a.i.·蜂-1,接触毒性48 h-LD_(50)分别为18.53、53.01、89.98和100μg a.i.·蜂-1,危害商值均小于50,对意大利蜜蜂均为低毒且低风险。4种制剂对玉米螟赤眼蜂的急性毒性24 h-LR50(半致死用量,median lethal rate)分别为1.88×10~(-4)、5.46×10~(-4)、2.12×10~(-3)和1.68×10~(-3)mg a.i.·cm~(-2),安全系数分别为0.16、0.23、0.59和0.51,其中8%烟嘧·氯吡嘧磺隆和16%烟嘧·硝磺·氯吡嘧磺隆对玉米螟赤眼蜂为高风险性,36%烟嘧·莠去津和22%烟嘧·氯吡·氯氟吡氧乙酸异辛酯为中等风险性。因此,这4种复配制剂对蜜蜂成年工蜂的风险性较低,但对赤眼蜂的风险性较高,所以在喷洒防除期间应与赤眼蜂释放期错开。     

Floral odor learning within the hive affects honeybees’ foraging decisions

Honeybees learn odor cues quickly and efficiently when visiting rewarding flowers. Memorization of these cues facilitates the localization and recognition of food sources during foraging flights. Bees can also use information gained inside the hive during social interactions with successful foragers. An important information cue that can be learned during these interactions is food odor. However, little is known about how floral odors learned in the hive affect later decisions of foragers in the field. We studied the effect of food scent on foraging preferences when this learning is acquired directly inside the hive. By using in-hive feeders that were removed 24 h before the test, we showed that foragers use the odor information acquired during a 3-day stimulation period with a scented solution during a food-choice situation outside the nest. This bias in food preference is maintained even 24 h after the replacement of all the hive combs. Thus, without being previously collected outside by foragers, food odors learned within the hive can be used during short-range foraging flights. Moreover, correct landings at a dual-choice device after replacing the storing combs suggests that long-term memories formed within the colony can be retrieved while bees search for food in the field.     

Attraction,deterrence or intoxication of bees (Apis mellifera) by plant allelochemicals

The influence of 63 dietary allelochemicals (alkaloids, terpenes, glycosides,etc.) on the feeding behaviour of bees (Apis mellifera) was tested in terms of deterrency and attraction. For 39 compounds a deterrent (mostly alkaloids, coumarins and saponins) and for 3 compounds an attractive response (mostly terpenes) was obtained in choice tests, which allowed the calculation of respective ED₅₀-values. Under no-choice conditions, 17 out of 29 allelochemicals caused mortality at concentrations between 0.003 and 0.6%. Especially toxic were alkaloids, saponins, cardiac glycosides and cyanogenic glycosides. These data show that bees which are confronted with plant allelochemicals in nectar and pollen, are not especially adapted (i.e. insensitive) to the plants' defence chemistry. GLC and GLS-MS data are given on the alkaloid composition of nectar and pollen ofBrugmansia aurea, Atropa belladonna andLupinus polyphyllus.     

The reproductive dilemmas of queenless red dwarf honeybee (Apis florea) workers

Honeybee (Apis) workers cannot mate, but retain functional ovaries. When colonies have lost their queen, many young workers begin to activate their ovaries and lay eggs. Some of these eggs are reared, but most are not and are presumably eaten by other workers (worker policing). Here we explore some of the factors affecting the reproductive success of queenless workers of the red dwarf honeybee Apis florea. Over a 2-year period we collected 40 wild colonies and removed their queens. Only two colonies remained at their translocated site long enough to rear males to pupation while all the others absconded. Absconding usually occurred after worker policing had ceased, as evidenced by the appearance of larvae. Dissections of workers from eight colonies showed that in A. florea, 6% of workers have activated ovaries after 4 days of queenlessness, and that 33% of workers have activated ovaries after 3 weeks. Worker-laid eggs may appear in nests within 4 days and larvae soon after, but this is highly variable. As with Apis mellifera, we found evidence of unequal reproductive success among queenless workers of A. florea. In the two colonies that reared males to pupation and which we studied with microsatellites, some subfamilies had much higher proportions of workers with activated ovaries than others. The significance of absconding and internest reproductive parasitism to the alternative reproductive strategies of queenless A. florea workers is discussed.     

Why are sand lizard males (Lacerta agilis) not equally green?

A honey bee (Apis mellifera) queen mates with about ten haploid drones, thus producing colonies composed of about ten subfamilies of super-sisters. An increasing but controversial body of literature supports the views that: (1) Members of each subfamily within a colony can recognise each other, and distinguish supersisters from half-sisters. (2) Members of each subfamily use this recognition information and increase the reproductive fitness of their own subfamily at the expense of half-sisters through behaviour termed nepotism. A mathematical model is developed that shows that task specialisation by subfamilies, and bees that repeatedly undertake the behaviour within subfamilies, can influence the numbers of interactions among super-sisters, relative to the numbers of interactions between half-sisters. The model is then evaluated using a data set pertaining to trophallaxis behaviour in a two-subfamily colony. It is concluded that with this data set, task specialisation and subfamily recognition were indeed confounded, suggesting that the apparent subfamily recognition could easily have been an artefact of task specialisation. Correspondence to: B.P. Oldroyd     

Effects of intracolony variability in behavioral development on plasticity of division of labor in honey bee colonies

There is a genetic component to plasticity in age polyethism in honey bee colonies, such that workers of some genotypes become precocious foragers more readily than do workers of other genotypes, in colonies lacking older bees. Using colonies composed of workers from two identifiable genotype groups, we determined that intracolony differences in the likelihood of becoming a precocious forager are a consequence of differences in rates of behavioral development that are also evident under conditions leading to normal development. An alternative hypothesis, that differences in the likelihood of becoming a precocious forager are due to differences in general sensitivity to altered colony conditions, was not supported. In three out of three trials, workers from the genotype group that was more likely to exhibit precocious foraging in single cohort colonies also foraged at relatively younger ages in colonies in which workers exhibited normal behavioral development. In contrast, in three out of three trials, workers from the genotype group that was more likely to exhibit precocious foraging in single-cohort colonies did not show disproportionately more overaged nursing in colonies in which workers exhibited delayed development. These results indicate that genotypic differences in plasticity in age-related division of labor are based on genotypic differences in rates of behavioral development.     

Paying for information: partial loads in central place foragers

Information about food sources can be crucial to the success of a foraging animal. We predict that this will influence foraging decisions by group-living foragers, which may sacrifice short-term foraging efficiency to collect information more frequently. This result emerges from a model of a central-place forager that can potentially receive information on newly available superior food sources at the central place. Such foragers are expected to return early from food sources, even with just partial loads, if information about the presence of sufficiently valuable food sources is likely to become available. Returning with an incomplete load implies that the forager is at that point not achieving the maximum possible food delivery rate. However, such partial loading can be more than compensated for by an earlier exploitation of a superior food source. Our model does not assume cooperative foraging and could thus be used to investigate this effect for any social central-place forager. We illustrate the approach using numerical calculations for honeybees and leafcutter ants, which do forage cooperatively. For these examples, however, our results indicate that reducing load confers minimal benefits in terms of receiving information. Moreover, the hypothesis that foragers reduce load to give information more quickly (rather than to receive it) fits empirical data from social insects better. Thus, we can conclude that in these two cases of social-insect foraging, efficient distribution of information by successful foragers may be more important than efficient collection of information by unsuccessful ones.