Task-specific expression of the foraging gene in harvester ants

In social insects, groups of workers perform various tasks such as brood care and foraging. Transitions in workers from one task to another are important in the organization and ecological success of colonies. Regulation of genetic pathways can lead to plasticity in social insect task behaviour. The colony organization of advanced eusocial insects evolved independently in ants, bees, and wasps and it is not known whether the genetic mechanisms that influence behavioural plasticity are conserved across species. Here we show that a gene associated with foraging behaviour is conserved across social insect species, but the expression patterns of this gene are not. We cloned the red harvester ant (Pogonomyrmex barbatus) ortholog (Pbfor) to foraging, one of few genes implicated in social organization, and found that foraging behaviour in harvester ants is associated with the expression of this gene; young (callow) worker brains have significantly higher levels of Pbfor mRNA than foragers. Levels of Pbfor mRNA in other worker task groups vary among harvester ant colonies. However, foragers always have the lowest expression levels compared to other task groups. The association between foraging behaviour and the foraging gene is conserved across social insects but ants and bees have an inverse relationship between foraging expression and behaviour.     

Thermoregulation of dancing bees: thoracic temperature of pollen and nectar foragers in relation to profitability of foraging and colony need

The thorax surface temperature of dancing honeybees (Apis mellifera carnica) recruiting nestmates to natural sources of nectar and pollen around Graz (Austria) was measured by real-time infrared thermography without touching them or disturbing social interactions. Thorax temperature during dancing was quite variable (31.4-43 degrees C). In the course of a foraging season it varied considerably and was always lower than in bees foraging from a highly profitable food source (2 molar sucrose 120 m from the hive). It averaged 38.0 degrees C (SD=2.24, n=224 dances) in the nectar foragers and 37.4 degrees C (SD=1.64, n=171) in the pollen foragers, resembling that of dancers foraging 0.5 molar sucrose from feeders with unlimited flow. Hive air temperature accounted only for about 3-8% of total variation. Foraging distance modulated dancing temperature in a way that, according to the decrease of the profitability of foraging with distance, maximum temperatures decreased and, in accordance with the increase of the dancing threshold with distance, minumum temperatures increased with distance, this way providing new support for the hypothesis that the dancing temperature is modulated by the profitability of foraging and the dancing and foraging motivation of the bees. Dancing temperature of both nectar and pollen dancers correlated with several parameters of the hive status, increasing with the amount of brood and decreasing with the amount of honey and pollen. These correlations are discussed with respect to literature reports on a colony's need for pollen and nectar, in particular the effect of brood and the amount of pollen on pollen foraging, and the effect of honey stores and demand for nectar on nectar foraging.     

基于RFID技术的西方蜜蜂采集行为研究

蜜蜂能够灵活调整出巢觅食行为,从而有效保证蜂群的正常发育与繁衍.为了探索采集蜂的行为特性,本文利用由江西农业大学蜜蜂研究所与广州市远望谷信息技术股份有限公司合作研发的蜜蜂无线射频识别(RFID)系统,对西方蜜蜂进行为期38 d的全天候监控记录.结果表明: 两蜂群中分别有63.4%和64.5%的采集蜂存在轮休现象,轮休时间比例为22.5%~26.4%;轮休与非轮休蜜蜂的采集工作总量差异不显著,但轮休蜜蜂寿命显著高于非轮休蜜蜂;提前发育的采集蜂的寿命显著低于正常采集蜂.本研究丰富了蜜蜂社会行为学内容,为进一步探索蜜蜂采集行为的形成机制奠定了基础.     

Percent lipid is associated with body size but not task in the bumble bee <Emphasis Type="Italic">Bombus impatiens</Emphasis>

In some group-living organisms, labor is divided among individuals. This allocation to particular tasks is frequently stable and predicted by individual physiology. Social insects are excellent model organisms in which to investigate the interplay between physiology and individual behavior, as division of labor is an important feature within colonies, and individual physiology varies among the highly related individuals of the colony. Previous studies have investigated what factors are important in determining how likely an individual is, compared to nestmates, to perform certain tasks. One such task is foraging. Corpulence (i.e., percent lipid) has been shown to determine foraging propensity in honey bees and ants, with leaner individuals being more likely to be foragers. Is this a general trend across all social insects? Here we report data analyzing the individual physiology, specifically the percent lipid, of worker bumble bees (Bombus impatiens) from whom we also analyze behavioral task data. Bumble bees are also unusual among the social bees in that workers may vary widely in size. Surprisingly we find that, unlike other social insects, percent lipid is not associated with task propensity. Rather, body size closely predicts individual relative lipid stores, with smaller worker bees being allometrically fatter than larger worker bees.     

Waggle Dance Distances as Integrative Indicators of Seasonal Foraging Challenges

Even as demand for their services increases, honey bees (Apis mellifera) and other pollinating insects continue to decline in Europe and North America. Honey bees face many challenges, including an issue generally affecting wildlife: landscape changes have reduced flower-rich areas. One way to help is therefore to supplement with flowers, but when would this be most beneficial? We use the waggle dance, a unique behaviour in which a successful forager communicates to nestmates the location of visited flowers, to make a 2-year survey of food availability. We “eavesdropped” on 5097 dances to track seasonal changes in foraging, as indicated by the distance to which the bees as economic foragers will recruit, over a representative rural-urban landscape. In year 3, we determined nectar sugar concentration. We found that mean foraging distance/area significantly increase from springs (493 m, 0.8 km²) to summers (2156 m, 15.2 km²), even though nectar is not better quality, before decreasing in autumns (1275 m, 5.1 km²). As bees will not forage at long distances unnecessarily, this suggests summer is the most challenging season, with bees utilizing an area 22 and 6 times greater than spring or autumn. Our study demonstrates that dancing bees as indicators can provide information relevant to helping them, and, in particular, can show the months when additional forage would be most valuable.     

Senescence in the worker honey bee Apis Mellifera

Honey bees are social insects that exhibit striking caste-specific differences in longevity. Queen honey bees live on average 1-2 years, whereas workers live 2-6 weeks in the summer and about 20 weeks in the winter. It is not clear whether queen-worker differences in longevity are due to intrinsic physiological differences in the rate of senescence, to differential exposure to extrinsic factors such as predation and adverse environmental conditions, or both. To determine if the relatively short lifespan of worker bees involves senescence, we measured age-specific resistance to three different physiological stressors (starvation, thermal, and oxidative stress) while eliminating age-related differences in foraging activity and minimizing age-related differences in energy expenditure. Despite these manipulations, older worker bees were still significantly less resistant to all three stressors than were younger bees. These results indicate that the regulation of worker bee lifespan involves senescence, in addition to extrinsic factors.     

Generalist bees (Meliponina) and the reproductive success of the mass flowering tree Stryphnodendron pulcherrimum (Fabales: Mimosaceae) in the Atlantic Rainforest, Bahia

It is controversial the role played by Meliponina bees in the pollination of mass flowering trees with small generalized flowers (FMPG), very common group of trees in the tropical forest canopy. The species richness and relative abundance of flower visiting insects of the mass flowering tree Stryphnodendron pulcherrimum were measured to test the hypothesis of tight ecological association between these generalist bees and FMPG and to evaluate the effect of this relationship upon the reproductive success variation among tree crowns. The flower visiting insects were sampled on 10 flowering tree crowns at the Atlantic Rainforest in southern Bahia. Altogether, 553 visiting insects were collected during the flowering period of S. pulcherrimum: 293 (52%) Meliponina bees out of 438 bees (79.4%). All tree crowns were visited by Meliponina, with the proportion of these bees ranging from 27% to 87%. The tight ecological association between FMPG trees and Meliponina bees is supported by the observed pattern of spatial relationship. Both the relationship between variation of fruit set among tree crowns and species richness (r = 0.3579; P = 0.3098) or relative abundance (r = 0.3070; P = 0.3881) of Meliponina were not statistically significant. Likely a threshold of minimum relative abundance combined with the absolute abundance of these bees explain the fruit set variation among tree crowns of S. pulcherrimum, even by self-pollination. We tested this assumption with a preliminary analysis of Melipona bee genera distribution among the tree crowns.     

Bumble bee abundance and richness improves honey bee pollination behaviour in sweet cherry

Pollination service in agricultural crops increases significantly with pollinator diversity and wild pollinator abundance. Differences in the foraging behaviour of pollinating insects are one of the reasons why pollinator diversity and abundance enhances crop pollination. Here, we focused on the foraging behaviour of honey bees and bumble bees in sweet cherry orchards. In addition, we studied the influence of bee diversity and abundance on the foraging behaviour of honey bees and bumble bees. Honey bees were found to visit fewer flowers than bumble bees. Bumble bees also showed a higher probability of changing trees between rows than honey bees. Both visitation rate and probability of row changes of honey bees increased with bumble bee diversity and with bumble bee abundance. We also found that the probability of row changes of honey bees increased with increasing bumble bee abundance. These effects of bumble bee richness and abundance on the pollination behaviour of honey bees can improve the pollination performance of honey bees in crops that depend on cross pollination. Our results highlight the higher pollination performance of bumble bees and the facilitative effect of wild pollinators to crop pollination.     

Sucrose response thresholds of honey bee (Apis mellifera) foragers are not modulated by brood ester pheromone

Division of labor is a hallmark of eusocial insects and their ecological success can be attributed to it. Honey bee division of labor proceeds along a stereotypical ontogenetic path based on age, modulated by various internal and external stimuli. Brood pheromone is a major social pheromone of the honey bee that has been shown to affect honey bee division of labor. It elicits several physiological and behavioral responses; notably, regulating the timing of the switch from performing in-hive tasks to the initiation of foraging. Additionally, brood pheromone affects future foraging choice. In honey bees, sucrose response threshold is a physiological correlate of age of first foraging and foraging choice. Brood pheromone has been shown to modulate sucrose response threshold in young bees, but its effects on sucrose response thresholds of bees in advanced behavioral states (foragers) are not known. In this study we examined the sucrose response thresholds of two different task groups, foragers (pollen and non-pollen) and non-foraging bees, in response to honey bee brood pheromone. Sucrose response thresholds were not significantly different between brood pheromone treatment and controls among both non-pollen and pollen foragers. However, the sucrose response threshold of non-foraging bees was significantly higher in the brood pheromone treatment group than in the control group. The switch to foraging task is considered a terminal one, with honey bee lifespan being determined at least partially by risks and stress accompanying foraging. Our results indicate that foragers are physiologically resistant to brood pheromone priming of sucrose response thresholds.     

Static Electricity in the Spatial Orientation and Signaling of Honey Bees

A static electric charge that foraging bees gain by rubbing their body parts against the substrate surface plays a role in social communication of bees. Vibrations of the charged body of a forager with a frequency of approximately 14 Hz indicate the location of the forager bee in a rich variety of bees within a poorly lit nest. The perception of vibrations of the charged ventral body surface of a forager occurs due to the antennal sensory organs of hive mates. An electric charge produced as the foraging bee moves its wings enhances vibrations of the trichoid sensilla, which act as mechanoreceptors. This provides reliable communication between foraging bees and their hive mates.     

Can alloethism in workers of the bumblebee, Bombus terrestris, be explained in terms of foraging efficiency?

Bumblebee workers vary greatly in size, unlike workers of most other social bees. This variability has not been adequately explained. In many social insects, size variation is adaptive, with different-sized workers performing different tasks (alloethism). Here we established whether workers of the bumblebee, Bombus terrestris (L.) (Hymenoptera; Apidae), exhibit alloethism. We quantified the size of workers engaging in foraging compared to those that remain in the nest, and confirmed that it is the larger bees that tend to forage (X±SE thorax widths 4.34±0.01 mm for nest bees and 4.93±0.02 mm for foragers). We then investigated whether large bees are better suited to foraging because they are able to transport heavier loads of food back to the nest. Both pollen and nectar loads of returning foragers were measured, demonstrating that larger bees do return with a heavier mass of forage. Foraging trip times were inversely related to bee size when collecting nectar, but were unrelated to bee size for bees collecting pollen. Overall, large bees brought back more nectar per unit time than small bees, but the rate of pollen collection appeared to be unrelated to size. The smallest foragers had a nectar foraging rate close to zero, presumably explaining why foragers tend to be large. Why might larger bees be better at foraging? Various explanations are considered: larger bees are able to forage in cooler conditions, may be able to forage over larger distances, and are perhaps also less vulnerable to predation. Conversely, small workers are presumably cheaper to produce and may be more nimble at within-nest tasks. Further research is needed to assess these possibilities. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Racial Differences in Division of Labor in Colonies of the Honey Bee (Apis mellifera)

We measured the age at onset of foraging in colonies derived from three races of European honey bees, Apis mellifera mellifera, Apis mellifera caucasica and Apis mellifera ligustica , using a cross-fostering design that involved six unrelated colonies of each race. There was a significant effect of the race of the introduced bees on the age at onset of foraging: cohorts of A. m. ligustica bees showed the earliest onset, regardless of the race of the colony they were introduced to. There also was a significant effect of the race of the host colony: cohorts of bees introduced into mellifera colonies showed the earliest onset of foraging, regardless of the race of the bees introduced. Significant inter-trial differences also were detected, primarily because of a later onset of foraging in trials conducted during the autumn (September–October). These results demonstrate differences among European races of honey bees in one important component of colony division of labor. They also provide a starting point for analyses of the evolution of division of labor under different ecological conditions.     

黄猄蚁对大蜜蜂的捕食行为及其潜在影响

捕食作用会对访花昆虫的种群、行为以及植物适合度产生影响,是植物与传粉者相互关系研究中常被忽视的因素.本文报道了黄猄蚁对大蜜蜂的捕食行为,并模拟捕食的关键环节研究了捕食过程对重要访花昆虫行为的影响.结果表明,黄猄蚁能够利用局部环境主动攻击猎物,利用群体合作捕获采集过程中的体型较大的大蜜蜂,捕食威胁是其影响植物-访花者关系的重要机制.大蜜蜂具有感知花上危险的能力,模拟处理的个体会逃离危险的花或植株并在花上留下标记,将危险信息传递给其它个体.其它拜访者对具有危险信号花的采集频次明显减少,采集时间缩短;模拟处理的影响会随时间推移而较快地消失.此外,该实验没有发现大蜜蜂在花上停留采集过程中具有明显的防御行为.     

Modeling Beehive Microclimate at the End of Wintering

Biophysics - Abstract—Beekeepers use electric heating of hives to overcome the difficult wintering period of bees, which is observed in late winter and early spring. The data previously given...     

A Strong Immune Response in Young Adult Honeybees Masks Their Increased Susceptibility to Infection Compared to Older Bees

Honeybees, Apis mellifera, show age-related division of labor in which young adults perform maintenance (“housekeeping”) tasks inside the colony before switching to outside foraging at approximately 23 days old. Disease resistance is an important feature of honeybee biology, but little is known about the interaction of pathogens and age-related division of labor. We tested a hypothesis that older forager bees and younger “house” bees differ in susceptibility to infection. We coupled an infection bioassay with a functional analysis of gene expression in individual bees using a whole genome microarray. Forager bees treated with the entomopathogenic fungus Metarhizium anisopliae s.l. survived for significantly longer than house bees. This was concomitant with substantial differences in gene expression including genes associated with immune function. In house bees, infection was associated with differential expression of 35 candidate immune genes contrasted with differential expression of only two candidate immune genes in forager bees. For control bees (i.e. not treated with M. anisopliae) the development from the house to the forager stage was associated with differential expression of 49 candidate immune genes, including up-regulation of the antimicrobial peptide gene abaecin, plus major components of the Toll pathway, serine proteases, and serpins. We infer that reduced pathogen susceptibility in forager bees was associated with age-related activation of specific immune system pathways. Our findings contrast with the view that the immunocompetence in social insects declines with the onset of foraging as a result of a trade-off in the allocation of resources for foraging. The up-regulation of immune-related genes in young adult bees in response to M. anisopliae infection was an indicator of disease susceptibility; this also challenges previous research in social insects, in which an elevated immune status has been used as a marker of increased disease resistance and fitness without considering the effects of age-related development.     

Do foraging bumblebees scent-mark food sources and does it matter?

Summary The foraging of worker bees of Bombus terrestris visiting artificial feeders in a climatic test chamber was investigated. The behaviour of worker bees visiting rewarding and unrewarding feeders is completely different. Of all flower visits to rewarding feeders 94% are probing-visits, i.e. the bees land on the flower and probe for nectar. In contrast, only 0.3% of all visits to unrewarding feeders are probing-visits, whereas 47% are approach-visits, i.e., the bees approach the feeders without landing. Exchanging feeder discs proves that the signal used for discrimination must be associated with the plastic disc used as landing platform. Most probably it involves scent-marking of the rewarding feeders with components of high and low volatility. The mean foraging efficiency of bees in a scent-marked foraging arena is 5.7 mg sugar/min and drops to 2.8 mg sugar/min after the scent marked discs are replaced by clean ones. Three components generate this drop in foraging efficiency: (1) the between-flower flight time increases, i.e. the bees search for a longer time before landing on flowers, (2) the bees no longer discriminate between rewarding and unrewarding feeders, and (3) the bees probe empty feeders longer than necessary; obviously they expect to find nectar.     

The repellent effect of two pyrethroid insecticides on the honey bee*

ABSTRACT. . An improved model for the repellent effect of pyrethroid insecticides on insects was developed using small colonies of honey bees, Apis mellifera L., in flight cages. Conditioning to scented feeders allowed the separation of foraging bees from a single colony into paired treatment and control groups. The repellent response was characterized as a sublethal toxic effect resulting in transitory inhibition of activity. Permethrin and cypermethrin were shown to be contact repellents to honey bees; exposure was primarily to the tarsi and abdominal venter. Repellency was fully reversible within 24 h. No permanent effects on either memory function or foraging efficiency were observed following acute exposure.     

Perception and the Characteristics of the Response of Honey Bees,Paper Wasps,and Red Wood Ants to a Low-Frequency Electric Field

Bees, wasps, and ants have no specialized receptors for the perception of an electric field. An appropriate response to naturally occurring electric fields in bees and ants is associated with atmospheric exposure, amplified by the approach of the front of a thunderstorm. The primary transducers of mechanoreceptors that respond to displacement are related to the perception of low-frequency electric fields of high intensity by insects. The non-specific mechanism of perception of electric fields is based on irritation by induced currents that flow in the locations of their contact with each other and/or conductive surfaces. The frequency dependence of the electric field sensitivity is determined mainly by the magnitude of the current induced by it and the intensity of its contact action. The magnitude of the current induced in the outer part of the insect body is non-linearly related to the frequency of the electric field. The region with the highest sensitivity to electric fields is close to 500 Hz, which is consistent with the maximum magnitude of the induced current. At the same time, the threshold of the sensitivity to an electric field in wasps is approximately 0.04 kV/m, while in bees it is 0.45 kV/m. Ants react to the action of an electric field of 7–10 kV/m by slowing their movement. Magnetic fields and ionization, which accompany the generation of an electric field whose intensity reaches 15–20 kV/m, do not stimulate changes in the behavior of insects.

     

Foraging behavior of honeybees on white clover

Summary Measurements of several aspects of foraging behavior of honeybees on white clover and of factors that might influence it were made at one-to-two-week intervals during the season. Other measurements were made on intervening days. Individual blossoms yielded a mean of from 0.02 to 0.08 l of nectar containing 42 to 65% sugar during the season. On one day when the nectar averaged 65% sugar, bees with pollen on their corbiculae carried a mean of 5.2 mg of pollen and 37.3 mg of nectar; bees without pollen carried a mean of 37.9 mg of nectar. On 10 occasions each of 50 bees were timed while foraging a single blossom; the mean foraging speed differed significantly on the 10 occasions, and varied from 1.5 to 3.0 seconds per blossom. Each of 18 bees was timed during visits to 25 blossoms, and the total time required for the same bees to forage from the blossoms on 25 racemes was also measured. The mean foraging speed of these bees varied from 1.1 to 4.5 seconds per blossom; the differences were highly significant. These bees foraged 7.0 to 26.0 blossoms per minute, and there was a strong negative correlation between the mean seconds required for bees to forage blossoms and the mean number of blossoms foraged per minute. Fifty one bees observed for a total of 314.5 minutes foraged an average of 18.4 blossoms per minute and 3.4 blossoms per inflorescence. Sixty one per cent of the flights of foraging bees were made to inflorescences estimated to be less than 5 inches from the starting point.Published as Technical Contribution N^o 4244 from theTexas Agricultural Experiment Station.     

Effects of selection for honey bee worker reproduction on foraging traits

The “reproductive ground plan” hypothesis (RGPH) proposes that reproductive division of labour in social insects had its antecedents in the ancient gene regulatory networks that evolved to regulate the foraging and reproductive phases of their solitary ancestors. Thus, queens express traits that are characteristic of the reproductive phase of solitary insects, whereas workers express traits characteristic of the foraging phase. The RGPH has also been extended to help understand the regulation of age polyethism within the worker caste and more recently to explain differences in the foraging specialisations of individual honey bee workers. Foragers that specialise in collecting proteinaceous pollen are hypothesised to have higher reproductive potential than individuals that preferentially forage for nectar because genes that were ancestrally associated with the reproductive phase are active. We investigated the links between honey bee worker foraging behaviour and reproductive traits by comparing the foraging preferences of a line of workers that has been selected for high rates of worker reproduction with the preferences of wild-type bees. We show that while selection for reproductive behaviour in workers has not altered foraging preferences, the age at onset of foraging of our selected line has been increased. Our findings therefore support the hypothesis that age polyethism is related to the reproductive ground plan, but they cast doubt on recent suggestions that foraging preferences and reproductive traits are pleiotropically linked.