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1.
Insect societies are important models for evolutionary biology and sociobiology. The complexity of some eusocial insect societies
appears to arise from self-organized task allocation and group cohesion. One of the best-supported models explaining self-organized
task allocation in social insects is the response threshold model, which predicts specialization due to inter-individual variability
in sensitivity to task-associated stimuli. The model explains foraging task specialization among honeybee workers, but the
factors underlying the differences in individual sensitivity remain elusive. Here, we propose that in honeybees, sensory sensitivity
correlates with individual differences in the number of sensory structures, as it does in solitary species. Examining European
and Africanized honeybees, we introduce and test the hypothesis that body size and/or sensory allometry is associated with
foraging task preferences and resource exploitation. We focus on common morphological measures and on the size and number
of structures associated with olfactory sensitivity. We show that the number of olfactory sensilla is greater in pollen and
water foragers, which are known to exhibit higher sensory sensitivity, compared to nectar foragers. These differences are
independent of the distribution of size within a colony. Our data also suggest that body mass and number of olfactory sensilla
correlate with the concentration of nectar gathered by workers, and with the size of pollen loads they carry. We conclude
that sensory allometry, but not necessarily body size, is associated with resource exploitation in honeybees and that the
differences in number of sensilla may underlie the observed differences in sensitivity between bees specialized on water,
pollen and nectar collection. 相似文献
2.
In honeybees, as in other highly eusocial species, tasks are performed by individual workers, but selection for worker task
phenotypes occurs at the colony level. We investigated the effect of colony-level selection for pollen storage levels on the
foraging behavior of individual honeybee foragers to determine (1) the relationship between genotype and phenotypic expression
of foraging traits at the individual level and (2) how genetically based variation in worker task phenotype is integrated
into colony task organization. We placed workers from lines selected at the colony level for high or low pollen stores together
with hybrid workers into a common hive environment with controlled access to resources. Workers from the selected lines showed
reciprocal variation in pollen and nectar collection. High-pollen-line foragers collected pollen preferentially, and low-
pollen-line workers collected nectar, indicating that the two tasks covary genetically. Hybrid workers were not intermediate
in phenotype, but instead showed directional dominance for nectar collection. We monitored the responses of workers from the
selected strains to changes in internal (colony) and external (resource) stimulus levels for pollen foraging to measure the
interaction between genotypic variation in foraging behavior and stimulus environment. Under low-stimulus conditions, the
foraging group was over-represented by high-pollen-line workers. However, the evenness in distribution of the focal genetic
groups increased as foraging stimuli increased. These data are consistent with a model where task choice is a consequence
of genetically based response thresholds, and where genotypic diversity allows colony flexibility by providing a range of
stimulus thresholds.
Received: 3 May 1999 / Received in revised form: 22 December 1999 / Accepted: 23 January 2000 相似文献
3.
There has now been an abundance of research conducted to explore genetic bases that underlie learning performance in the honey bee (Apis mellifera). This work has progressed to the point where studies now seek to relate genetic traits that underlie learning ability to learning in field-based foraging problems faced by workers. Accordingly, the focus of our research is to explore the correlation between laboratory-based performance using an established learning paradigm and field-based foraging behavior. To evaluate learning ability, selected lines were established by evaluating queens and drones in a proboscis extension reflex (PER) conditioning procedure to measure learning in a laboratory paradigm—latent inhibition (LI). Hybrid queens were then produced from our lines selected for high and low levels of LI and inseminated with semen from many drones chosen at random. The genetically diverse worker progeny were then evaluated for expression of LI and for preference of pollen and/or nectar during foraging. Foragers from several different queens, and which had resulted from fertilization by any of several different drone fathers, were collected as they returned from foraging flights and analyzed for pollen and nectar contents. They were subsequently evaluated for expression of LI. Our research revealed that pollen foragers exhibited stronger learning, both in the presence (excitatory conditioning) and absence (LI) of reinforcement. The heightened overall learning ability demonstrated by pollen foragers suggests that pollen foragers are in general more sensitive to a large number of environmental stimuli. This mechanism could contribute toward explanations of colony-level regulation of foraging patterns among workers.Communicated by R. Page 相似文献
4.
Nadine C. Chapman Benjamin P. Oldroyd William O. H. Hughes 《Behavioral ecology and sociobiology》2007,61(8):1185-1194
Which task a social insect worker engages in is influenced by the worker’s age, genotype and the colony’s needs. In the honeybee,
Apis mellifera, genotype influences both the age a worker switches tasks and its propensity of engaging in specialist tasks, such as water
collecting, which only some workers will perform. In this study, we used colonies with natural levels of genetic diversity
and manipulated colony age demography to drastically increase the stimuli for the generalist tasks of foraging and nursing,
which all workers are thought to engage in at some point in their lives. We examined the representation of worker patrilines
engaged in nursing and foraging before and after the perturbation. The representation of patrilines among foragers and nurses
differed from that of their overall colony’s population. In the case of foraging, over- and underrepresentation of some patrilines
was not simply due to differences in rates of development among patrilines. We show that replacement foragers tend to be drawn
from patrilines that were overrepresented among foragers before the perturbation, suggesting that there is a genetic component
to the tendency to engage in foraging. In contrast, the representation of patrilines in replacement nurses differed from that
in the unperturbed nursing population. Our results show that there is a genetic influence on even the generalist tasks of
foraging and nursing, and that the way patrilines in genetically diverse colonies respond to increases in task stimuli depends
upon the task. The possible significance of this genetic influence on task allocation is discussed.
Electronic supplementary material Supplementary material is available in the online version of this article at doi: and is accessible to authorized users. 相似文献
5.
Solitary foragers can balance demands for food and safety by varying their relative use of foraging patches and their level of vigilance. Here, we investigate whether colonies of the ant, Formica perpilosa, can balance these demands by dividing labor among workers. We show that foragers collecting nectar in vegetation near their nest are smaller than are those collecting nectar at sites away from the nest. We then use performance tests to show that smaller workers are more likely to succumb to attack from conspecifics but feed on nectar more efficiently than larger workers, suggesting a size-related trade-off between risk susceptibility and harvesting ability. Because foragers that travel away from the nest are probably more likely to encounter ants from neighboring colonies, this trade-off could explain the benefits of dividing foraging labor among workers. In a laboratory experiment, we show that contact with aggressive workers results in an increase in the mean size of recruits to a foraging site: this increase was not the result of more large recruits, but rather because fewer smaller ants traveled to the site. These results suggest that workers particularly susceptible to risk avoid dangerous sites, and suggest that variation in worker size can allow colonies to exploit profitably both hazardous and resource-poor patches.Communicated by L. Sundström 相似文献
6.
Paul Schmid-Hempel 《Behavioral ecology and sociobiology》1984,14(4):263-271
Summary Observations and field experiments on the foraging behaviour of individual workers of Cataglyphis bicolor in a Southern Tunisian shrub desert are reported. The workers search singly for their food (mostly animal carcasses) and are singleprey loaders. The individuals differ to a great extent in their persistence to re-search the place of a find on a previous foraging excursion. The differences range continuously from thoroughly researching a place to just walking by. If, in an experiment, the same reward is offered farther from the nest, each ant persists more in re-searching the place than if food is offered close to the nest. In a further experiment, some individuals persisted less in searching near the former finding site if they had collected a fly than after collecting a piece of cheese. There is, however, evidence that individuals do not differ in their food preference. Persistent individuals, which re-search the place of a former find, are faster than non-persistent ones in retrieving food that is experimentally arranged in an aggregated manner. The experiment failed to demonstrate the (reverse) superiority of non-persistent individuals foraging on homogeneously distributed food. The observations of unmanipulated foraging excursions in the field suggest such an advantage for non-persistent foragers under natural conditions where food in general occurs widely dispersed. The colony as a whole retrieves more food within the same time from an experimental lay-out that is homogeneous than from an aggregated one. The behavioural differences between individuals could be caused by a training bias of the short-lived foragers, leading to a different assessment of the profitability of a searching method which implies returning to a formerly rewarding place. Thus, each worker uses the most promising behaviour according to its individual experience. Alternatively, the individually different searching methods could mainly contribute to the welfare of the colony as a whole rather than leading to a maximal short-term efficiency of each individual. In particular, the colony, disposing of only a few highly persistent foragers, could quickly exploit occasional short-lived, but unpredictible, clumps of food within its foraging range. 相似文献
7.
Bruce J. Eckholm Kirk E. Anderson Milagra Weiss Gloria DeGrandi-Hoffman 《Behavioral ecology and sociobiology》2011,65(5):1037-1044
Multiple mating by honeybee queens results in colonies of genotypically diverse workers. Recent studies have demonstrated
that increased genetic diversity within a honeybee colony increases the variation in the frequency of tasks performed by workers.
We show that genotypically diverse colonies, each composed of 20 subfamilies, collect more pollen than do genotypically similar
colonies, each composed of a single subfamily. However, genotypically similar colonies collect greater varieties of pollen
than do genotypically diverse colonies. Further, the composition of collected pollen types is less similar among genotypically
similar colonies than among genotypically diverse colonies. The response threshold model predicts that genotypic subsets of
workers vary in their response to task stimuli. Consistent with this model, our findings suggest that genotypically diverse
colonies likely send out fewer numbers of foragers that independently search for pollen sources (scouts) in response to protein
demand by the colony, resulting in a lower variety of collected pollen types. The cooperative foraging strategy of honeybees
involves a limited number of scouts monitoring the environment that then guide the majority of foragers to high quality food
sources. The genetic composition of the colony appears to play an important role in the efficiency of this behavior. 相似文献
8.
James F. A. Traniello Marty S. Fujita Rhys V. Bowen 《Behavioral ecology and sociobiology》1984,15(1):65-68
Summary When prey of two sizes (6 and 32 mg) were offered in a choice situation to foragers of the ant Formica schaufussi at different ambient temperatures, significantly more workers rejected the smaller prey at low temperatures, whereas at high temperatures workers accepted the less profitable smaller item. Foragers scavenge for arthropod prey over a temperature range of 15–40°C, and increasing temperature significantly increases a forager's oxygen consumption, an index of energy expenditure. 相似文献
9.
Thomas D. Seeley 《Behavioral ecology and sociobiology》1986,19(5):343-354
Summary To understand how a colony of honeybees keeps its forager force focussed on rich sources of food, and analysis was made of how the individual foragers within a colony decide to abandon or continue working (and perhaps even recruit to) patches of flowers. A nectar forager grades her behavior toward a patch in response to both the nectar intake rate of her colony and the quality of her patch. This results in the threshold in patch quality for acceptance of a patch being higher when the colonial intake rate of nectar is high than when it is low. Thus colonies can adjust their patch selectivity so that they focus on rich sources when forage is abundant, but spread their workers among a wider range of sources when forage is scarce. Foragers assess their colony's rate of nectar intake while in the nest, unloading nectar to receiver bees. The ease of unloading varies inversely with the colonial intake rate of nectar. Foragers assess patch quality while in the field, collecting nectar. By grading their behavior steeply in relation to such patch variables as distance from the nest and nectar sweetness, foragers give their colony high sensitivity to differences in profitability among patches. When a patch's quality declines, its foragers reduce their rate of visits to the patch. This diminishes the flow of nectar from the poor patch which in turn stimulates recruitment to rich patches. Thus a colony can swiftly redistribute its forager force following changes in the spatial distribution of rich food sources. The fundamental currency of nectar patch quality is not net rate of energy intake, (Gain-Cost)/Time, but may be net energy efficiency, (Gain-Cost)/Cost. 相似文献
10.
Michael Simone-Finstrom Joel Gardner Marla Spivak 《Behavioral ecology and sociobiology》2010,64(10):1609-1617
Honeybees harvest and use plant resins in a mixture called propolis to seal cracks and smooth surfaces in the nest architecture.
Resins in the nest may be important in maintaining a healthy colony due to their antimicrobial properties. This study had
two main objectives: (1) Provide initial insight on the learning capabilities of resin foraging honeybees; (2) analyze the
sensitivity of resin foraging honeybees to tactile stimuli to elucidate its possible role as a mechanism behind resin foraging.
The first objective provides insight into the phenotype of these bees as compared to other forager types, while the second
creates a starting point for further work on behavioral mechanisms of resin foraging. Using tactile proboscis extension response
conditioning, we found that resin foragers learned to associate two different tactile stimuli, the presence of a gap between
two plates and a rough sandpaper surface, with a sucrose reward significantly better than pollen foragers. The results of
differential tactile conditioning exhibited no significant difference in the ability of resin foragers to discriminate between
smooth and rough surfaces as compared to pollen foragers. We also determined that the sucrose response thresholds (SRTs) of
returning resin foragers were lower compared to returning pollen foragers, but both resin foragers and pollen foragers learned
a floral odor equally well. This is the first study to examine SRTs and conditioning to tactile and olfactory stimuli with
resin foraging honeybees. The results provide new information and identify areas for future research on resin collectors,
an understudied foraging phenotype. 相似文献
11.
Promiscuous honeybee queens generate colonies with a critical minority of waggle-dancing foragers 总被引:1,自引:1,他引:0
Honeybees present a paradox that is unusual among the social Hymenoptera: extremely promiscuous queens generate colonies of
nonreproducing workers who cooperate to rear reproductives with whom they share limited kinship. Extreme polyandry, which
lowers relatedness but creates within-colony genetic diversity, produces substantial fitness benefits for honeybee queens
and their colonies because of increased disease resistance and workforce productivity. However, the way that these increases
are generated by individuals in genetically diverse colonies remains a mystery. We assayed the foraging and dancing performances
of workers in multiple-patriline and single-patriline colonies to discover how within-colony genetic diversity, conferred
to colonies by polyandrous queens, gives rise to a more productive foraging effort. We also determined whether the initiation
by foragers of waggle-dance signaling in response to an increasing sucrose stimulus (their dance response thresholds) was
linked to patriline membership. Per capita, foragers in multiple-patriline colonies visited a food source more often and advertised
it with more waggle-dance signals than foragers from single-patriline colonies, although there was variability among multiple-patriline
colonies in the strength of this difference. High-participation patrilines emerged within multiple-patriline colonies, but
their more numerous foragers and dancers were neither more active per capita nor lower-threshold dancers than their counterparts
from low-participation patrilines. Our results demonstrate that extreme polyandry does not enhance recruitment effort through
the introduction of low-dance-threshold, high-activity workers into a colony’s population. Rather, genetic diversity is critical
for injecting into a colony’s workforce social facilitators who are more likely to become engaged in foraging-related activities,
so boosting the production of dance signals and a colony’s responsiveness to profitable food sources. 相似文献
12.
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. 相似文献
13.
Sean O’Donnell 《Behavioral ecology and sociobiology》1996,38(2):83-88
Genetic variability within insect societies may provide a mechanism for increasing behavioral diversity among workers, thereby
augmenting colony efficiency or flexibility. In order to assess the possibility that division of labor has a genetic component
in the eusocial wasp Polybia aequatorialis, I asked whether the genotypes of workers within colonies correlated with behavioral specialization. Workers specialized
by foraging for one of the four materials (wood pulp, insect prey, nectar, or water) gathered by their colonies. I collected
foragers on 2 days from each of three colonies and identified the material the foragers were carrying when collected. I produced
random amplified polymorphic DNA (RAPD) markers from the genomic DNA of these foragers and estimated genotypic similarity
of foragers based on sharing of variable RAPD marker bands. Contingency tests on 20 variable loci per colony showed statistically
significant (P <0.05) biases in RAPD marker frequencies among forager types in the three colonies. Patterns of association of RAPD marker
bands with specializations were constant in two colonies, but changed between collection days in one colony. RAPD marker biases
suggest that division of labor among workers includes a genetic component in P. aequatorialis. Colony-level selection on variation in division of labor is a possible factor favoring the evolutionary maintenance of high
genotypic variability (low relatedness) in epiponine wasp colonies and in other eusocial insects.
Received: 18 July 1995/Accepted after revision: 1 October 1995 相似文献
14.
Tamar Drezner-Levy Brian H. Smith Sharoni Shafir 《Behavioral ecology and sociobiology》2009,64(1):135-148
Honey bee foragers may collect nectar, pollen, water, or propolis, and their foraging specialization has been associated with
several behavioral traits. By conditioning of the proboscis extension response (PER), we compared the performance of foragers
that collected nectar, pollen, both nectar and pollen, or water in several learning and choice assays. Foragers were first
tested in a three-trial olfactory associative learning assay. For further tests, we selected only good learners that responded
in two out of three conditioning trials. One group was tested in an additional olfactory associative learning assay involving
different reward volumes and concentrations. Another group was tested for risk sensitivity in a two-alternative forced-choice
PER procedure and then in a latent inhibition (LI) assay. Levels of acquisition in olfactory associative learning were highest
in pollen and water foragers, and better acquisition was associated with collection of heavier pollen loads and smaller and
lighter nectar loads of lower sugar concentration. Among the good learners, pollen foragers still showed better acquisition
than nectar foragers when rewarded with several volumes and concentrations of sucrose solution. Pollen and nectar foragers
were equally risk averse, preferring a constant reward to a variable one, and choice was not affected by pollen load weight.
Contrary to a previous study, pollen and nectar foragers were similarly affected by LI. We discuss possible explanations for
the discrepancy between the two studies. Overall, our results suggest that differences between foraging groups in sensitivity
to various stimuli may not correspond to differences in choice behavior. 相似文献
15.
We have found that foraging bumblebees (Bombus hortorum, B. pascuorum, B. pratorum and B.␣terrestris) not only avoid flowers of Symphytum officinale that have recently been visited by conspecifics but also those that have been recently visited by heterospecifics. We propose
that the decision whether to reject or accept a flower is influenced by a chemical odour that is left on the corolla by a
forager, which temporarily repels subsequent foragers. Honeybees and carpenter bees have previously been shown to use similar
repellent forage-marking scents. We found that flowers were repellent to other bumblebee foragers for approximately 20 min
and also that after this time nectar levels in S. officinale flowers had largely replenished. Thus bumblebees could forage more efficiently by avoiding flowers with low rewards. Flowers
to which extracts of tarsal components were applied were more often rejected by wild B. terrestris workers than flowers that had head extracts applied, which in turn were more often rejected than flowers that had body extracts
applied. Extracts from four Bombus species were equally repellent to foragers. The sites of production of the repellent scent and its evolutionary origins are
discussed.
Received: 24 November 1997 / Accepted after revision: 8 March 1998 相似文献
16.
Summary The foragers in honeybee colonies cooperate by sharing information about rich sources of food. This study examines three hypotheses about the benefits of this cooperation: (H1) it decreases foragers' costs in finding new food sources, (H2) it increases the quality of the food sources located by foragers, and (H3) it increases the ability of a colony's foragers to compete for high-quality food sources. To test each hypothesis, we identified a critical pattern in the foraging process which, if observed, would cast doubt on that hypothesis, and then gathered data to check for these patterns. Our observations do not support the first hypothesis, but do support the second and third. These results, in addition to helping us understand the functional significance of the honeybee's dance language, provide insights into the colonial organization of foraging by honeybees. 相似文献
17.
Summary Food-sharing experiments were performed with laboratory colonies of Solenopsis invicta containing 1000, 10,000, or 20,000 workers and starved for 0, 3, 7, or 14 days. The effect of these variables was measured on the uptake of radioactive sugar water (1 M) by 1% of the colony's workers and on the trophallactic flow of food from these foragers to the remainder of the colony.Patterns of food distribution in small colonies differed significantly from those in larger nests. In 1000-ant nests, small workers more frequently received food than large workers, but in bigger colonies the opposite occurred.Fire ants were adept at distributing sugar water, with food from a few workers rapidly reaching the majority of the colony as foragers donate their crop contents to groups of recipients and these recipients may themselves act as donors.Foragers respond to colony starvation by individually taking up more food and sharing this fluid with a greater proportion of nestmates. Even foragers from satiated colonies can retrieve at least small amounts of liquid.The forager's state of hunger plays an important role in regulating food distribution. In sugar-satiated nests, previously starved foragers are highly successful at passing on labelled sugar whereas prviously fed foragers are not. 相似文献
18.
Division of labor during honey bee colony defense 总被引:5,自引:0,他引:5
Michael D. Breed Gene E. Robinson Robert E. Page Jr. 《Behavioral ecology and sociobiology》1990,27(6):395-401
Summary Some worker honey bees respond to major disturbances of the colony by flying around the assailant and possibly stinging; they are a subset of the bees involved in colony defense. These defenders have an open-ended age distribution similar to that of foragers, but defensive behavior is initiated at a younger age than foraging is. Behavioral and genetic evidence shows that defenders and foragers are distinct groups of older workers. Behaviorally, defenders have less worn wings than foragers, suggesting less flight activity. Genetically, defenders differ in allozyme frequencies, demonstrating different subfamily composition from foragers in the same colony. They also differ in allozyme frequencies from guards in the same colony, providing further evidence for division of labor associated with colony defense. We use this information to develop a model for honey bee colony defense involving at least two distinct groups of workers and we propose that the non-guard defenders be called soldiers, due to their important role in colony defense.Offprint requests to: M.D. Breed 相似文献
19.
Madeleine Beekman Amy L. Gilchrist Michael Duncan David J. T. Sumpter 《Behavioral ecology and sociobiology》2007,61(7):985-995
A honeybee colony needs to divide its workforce so that each of the many tasks it performs has an appropriate number of workers
assigned to it. This task allocation system needs to be flexible enough to allow the colony to quickly adapt to an ever-changing
environment. In this study, we examined possible mechanisms by which a honeybee colony regulates the division of labor between
scouts (foragers that search for new food sources without having been guided to them) and recruits (foragers that were guided
via recruitment dances toward food sources). Specifically, we examined the roles that the availability of recruitment dances
and worker genotype has in the colony-level regulation of the number of workers engaged in scouting. Our approach was threefold.
We first developed a mathematical model to demonstrate that the decision to become a scout or a recruit could be regulated
by whether a potential forager can find a recruitment dance within a certain time period. We then tested this model by investigating
the effect of dance availability on the regulation of scouts in the field. Lastly, we investigated if the probability of being
a scout has a genetic basis. Our field data supported the hypothesis that scouts are those foragers that have failed to locate
a recruitment dance as predicted by our model, but we found no effect of genotype on the propensity of foragers to become
scouts. 相似文献
20.
In an experimental set-up, a colony of the stingless bee Melipona fasciata demonstrated its ability to choose the better of two nectar sources. This colony pattern was a result of the following individual
behavioural decisions: continue foraging, abandon the feeder, restart foraging and initiate foraging. Only very rarely did
individuals switch from one feeder to the other. With the first combination of a rich (2.7 M) and a poor (0.8 M) feeder M. fasciata behaved differently from Apis mellifera. Recruitment occurred to both feeders and the poor feeder was not abandoned completely. When the poor feeder was set to 0.4 M,
M. fasciata abandoned the poor feeder rapidly and allocated more foragers to the rich feeder. These patterns were similar to those reported
for A. mellifera with the first combination of feeders. Over a sequence of 4 days, experienced bees increasingly determined the colony patterns,
and the major function of communication between workers became the reactivation of experienced foragers. The foragers modulated
their behaviour not only according to the profitability of the feeder, but also according to previous experience with profitability
switches. Thus, experience and communication together regulated colony foraging behaviour. These findings and the results of studies with honeybees suggest
that M. fasciata and honeybees use similar decision-making mechanisms and only partly different tools.
Received: 21 December 1998 / Accepted: 5 January 1999 相似文献