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1.
The regulation of protein collection through pollen foraging plays an important role in pollination and in the life of bee
colonies that adjust their foraging to natural variation in pollen protein quality and temporal availability. Bumble bees
occupy a wide range of habitats from the Nearctic to the Tropics in which they play an important role as pollinators. However,
little is known about how a bumble bee colony regulates pollen collection. We manipulated protein quality and colony pollen
stores in lab-reared colonies of the native North American bumble bee, Bombus impatiens. We debut evidence that bumble bee colony foraging levels and pollen storage behavior are tuned to the protein quality (range
tested: 17–30% protein by dry mass) of pollen collected by foragers and to the amount of stored pollen inside the colony.
Pollen foraging levels (number of bees exiting the nest) significantly increased by 55%, and the frequency with which foragers
stored pollen in pots significantly increased by 233% for pollen with higher compared to lower protein quality. The number
of foragers exiting the nest significantly decreased (by 28%) when we added one pollen load equivalent each 5 min to already
high intranidal pollen stores. In addition, pollen odor pumped into the nest is sufficient to increase the number of exiting
foragers by 27%. Foragers directly inspected pollen pots at a constant rate over 24 h, presumably to assess pollen levels.
Thus, pollen stores can act as an information center regulating colony-level foraging according to pollen protein quality
and colony need.
An erratum to this article can be found at 相似文献
2.
Tanya Pankiw Robert E. Page Jr M. Kim Fondrk 《Behavioral ecology and sociobiology》1998,44(3):193-198
Foraging and the mechanisms that regulate the quantity of food collected are important evolutionary and ecological attributes
for all organisms. The decision to collect pollen by honey bee foragers depends on the number of larvae (brood), amount of
stored pollen in the colony, as well as forager genotype and available resources in the environment. Here we describe how
brood pheromone (whole hexane extracts of larvae) influenced honey bee pollen foraging and test the predictions of two foraging-regulation
hypotheses: the indirect or brood-food mechanism and the direct mechanism of pollen-foraging regulation. Hexane extracts of
larvae containing brood pheromone stimulated pollen foraging. Colonies were provided with extracts of 1000 larvae (brood pheromone),
1000 larvae (brood), or no brood or pheromone. Colonies with brood pheromone and brood had similar numbers of pollen foragers,
while those colonies without brood or pheromone had significantly fewer pollen foragers. The number of pollen foragers increased
more than 2.5-fold when colonies were provided with extracts of 2000 larvae as a supplement to the 1000 larvae they already
had. Within 1 h of presenting colonies with brood pheromone, pollen foragers responded to the stimulus. The results from this
study demonstrate some important aspects of pollen foraging in honey bee colonies: (1) pollen foragers appear to be directly
affected by brood pheromone, (2) pollen foraging can be stimulated with brood pheromone in colonies provided with pollen but
no larvae, and (3) pollen forager numbers increase with brood pheromone as a supplement to brood without increasing the number
of larvae in the colony. These results support the direct-stimulus hypothesis for pollen foraging and do not support the indirect-inhibitor,
brood-food hypothesis for pollen-foraging regulation.
Received: 5 March 1998 / Accepted after revision: 29 August 1998 相似文献
3.
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
4.
A fundamental requirement of task regulation in social groups is that it must allow colony flexibility. We tested assumptions
of three task regulation models for how honeybee colonies respond to graded changes in need for a specific task, pollen foraging.
We gradually changed colony pollen stores and measured behavioral and genotypic changes in the foraging population. Colonies
did not respond in a graded manner, but in six of seven cases showed a stepwise change in foraging activity as pollen storage
levels moved beyond a set point. Changes in colony performance resulted from changes in recruitment of new foragers to pollen
collection, rather than from changes in individual foraging effort. Where we were able to track genotypic variation, increases
in pollen foraging were accompanied by a corresponding increase in the genotypic diversity of pollen foragers. Our data support
previous findings that genotypic variation plays an important role in task regulation. However, the stepwise change in colony
behavior suggests that colony foraging flexibility is best explained by an integrated model incorporating genotypic variation
in task choice, but in which colony response is amplified by social interactions.
Received: 17 October 1998 / Received in revised form: 11 March 1999 / Accepted: 12 March 1999 相似文献
5.
Benjamin P. Oldroyd Thomas E. Rinderer Steven M. Buco 《Behavioral ecology and sociobiology》1992,30(5):291-295
Summary Colonies of honey bees with two identifiable subfamilies were established. Returning foragers were captured and killed at two different sampling times. The mean volume and per cent soluble solids of crop contents were determined for each subfamily, as was the mean weight of the pollen pellets. No significant differences in nectar volume or concentration were detected between subfamilies within colonies. However, in a few colonies, significant subfamily by sampling-time interactions were present, suggesting that in these colonies subfamilies differed in their nectar and pollen collecting behavior at different times of day. The plant genera worked by pollen foragers were also determined. In four of six colonies, bees of different subfamilies were found to be majoring on different plant species (Fig. 1). Implications of this intra-colonial variance in foraging behavior for colony fitness are discussed.
Offprint requests to: B.P. Oldroyd 相似文献
6.
Two-way selection for quantities of stored pollen resulted in the production of high and low pollen hoarding strains of honey bees (Apis mellifera L.). Strains differed in areas of stored pollen after a single generation of selection and, by the third generation, the high strain colonies stored an average 6 times more pollen than low strain colonies. Colony-level organizational components that potentially affect pollen stores were identified that varied genetically within and between these strains. Changes occurred in several of these components, in addition to changes in the selected trait. High strain colonies had a significantly higher proportion of foragers returning with loads of pollen, however, high and low strain colonies had equal total numbers of foragers Colony rates of intake of pollen and nectar were not independent. Selection resulted in an increase in the number of pollen collectors and a decrease in the number of nectar collectors in high strain colonies, while the reciprocal relationship occurred in the low strain. High and low strain colonies also demonstrated different diurnal foraging patterns as measured by the changing proportions of returning pollen foragers. High strain colonies of generation 3 contained significantly less brood than did low strain colonies, a consequence of a constraint on colony growth resulting from a fixed nest volume and large quantities of stored pollen. These components represent selectable colony-level traits on which natural selection can act and shape the social organization of honey bee coloniesCommunicated by R.F.A. Moritz 相似文献
7.
Pollen is the sole source of protein for honey bees, most importantly used to rear young. Honey bees are adept at regulating
pollen stores in the colonies based on the needs of the colony. Mechanisms for regulation of pollen foraging in honey bee
are complex and remain controversial. In this study, we used a novel approach to test the two competing hypothesis of pollen
foraging regulation. We manipulated nurse bee biosynthesis of brood food using a protease inhibitor that interferes with midgut
protein digestion, significantly decreasing the amount of protein extractable from hypopharyngeal glands. Experimental colonies
were given equal amounts of protease inhibitor-treated and untreated pollen. Colonies receiving protease inhibitor treatment
had significantly lower hypopharyngeal gland protein content than controls. There was no significant difference in the ratio
of pollen to nonpollen foragers between the treatments. Pollen load weights were also not significantly different between
treatments. Our results supported the pollen foraging effort predictions generated from the direct independent effects of
pollen on the regulation of pollen foraging and did not support the prediction that nurse bees regulate pollen foraging through
amount of hypopharyngeal gland protein biosynthesis. 相似文献
8.
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
9.
Claudia Dreller Robert E. Page Jr. M. Kim Fondrk 《Behavioral ecology and sociobiology》1999,45(3-4):227-233
Pollen storage in a colony of Apis mellifera is actively regulated by increasing and decreasing pollen foraging according to the “colony's needs.” It has been shown that
nectar foragers indirectly gather information about the nectar supply of the colony from nestmates without estimating the
amount of honey actually stored in the combs. Very little is known about how the actual colony need is perceived with respect
to pollen foraging. Two factors influence the need for pollen: the quantity of pollen stored in cells and the amount of brood.
To elucidate the mechanisms of perception, we changed the environment within normal-sized colonies by adding pollen or young
brood and measured the pollen-foraging activity, while foragers had either direct access to them or not. Our results show
that the amount of stored pollen, young brood, and empty space directly provide important stimuli that affect foraging behavior.
Different mechanisms for forager perception of the change in the environment are discussed.
Received: 13 June 1998 / Accepted after revision: 25 October 1998 相似文献
10.
Summary In a controlled laboratory experiment, we re-examined the question of bumble bee risk-sensitivity. Harder and Real's (1987) analysis of previous work on bumble bee risk aversion suggests that risk-sensitivity in these organisms is a result of their maximizing the net rate of energy return (calculated as the average of expected per flower rates). Whether bees are risk-sensitive foragers with respect to minimizing the probability of energetic shortfall is therefore still an open question. We examined how the foraging preferences of bumble bees for nectar reward variation were affected by colony energy reserves, which we manipulated by draining or adding sucrose solution to colony honey pots. Nine workers from four confined colonies of Bombus occidentalis foraged for sucrose solution in two patches of artificial flowers. These patches yielded the same expected rate of net energy intake, but floral volumes were variable in one patch and constant in the other. Our results show that bumble bees can be both risk-averse (preferring constant flowers) and risk-prone (preferring variable flowers), depending on the status of their colony energy reserves. Diet choice in bumble bees appears to be sensitive to the target value a colony-level energetic requirement.
Offprint requests to: R.V. Cartar 相似文献
11.
Brian R. Johnson 《Behavioral ecology and sociobiology》2005,58(3):219-226
Caste theory predicts that social insect colonies are organized into stable groups of workers specialized on particular task sets. Alternative concepts of organization of work suggest that colonies are composed of extremely flexible workers able to perform any task as demand necessitates. I explored the flexibility of workers in temporal castes of the honey bee Apis mellifera by determining the ability of colonies to reorganize labor after a major demographic disturbance. I evaluated the flexibility of temporal castes by comparing the foraging rates of colonies having just lost their foragers with colonies having also lost their foragers but having been given a week to reorganize. The population sizes and contents of the colonies in each group were equalized and foraging rates were recorded for one week. Colonies given a weeks initial recovery time after the loss of their foragers were found to forage at significantly higher rates than those colonies given no initial recovery time. This result was consistent for nectar and pollen foraging. These results suggest that honeybee workers lack sufficient flexibility to reorganize labor without compromising foraging. This finding is consistent with the caste concept model of organization of work in insect societies. 相似文献
12.
The regulation of pollen foraging by honey bees: how foragers assess the colony's need for pollen 总被引:2,自引:0,他引:2
Scott Camazine 《Behavioral ecology and sociobiology》1993,32(4):265-272
Summary The honey bee colony presents a challenging paradox. Like an organism, it functions as a coherent unit, carefully regulating its internal milieu. But the colony consists of thousands of loosely assembled individuals each functioning rather autonomously. How, then, does the colony acquire the necessary information to organize its work force? And how do individuals acquire information about specific colony needs, and thus know what tasks need be performed? I address these questions through experiments that analyze how honey bees acquire information about the colony's need for pollen and how they regulate its collection. The results demonstrate features of the colony's system for regulating pollen foraging: (1) Pollen foragers quickly acquire new information about the colony's need for pollen. (2) When colony pollen stores are supplemented, many pollen foragers respond by switching to nectar foraging or by remaining in the hive and ceasing to forage at all. (3) Pollen foragers do not need direct contact with pollen to sense the colony's change of state, nor do they use the odor of pollen as a cue to assess the colony's need for pollen. (4) Pollen foragers appear to obtain their information about colony pollen need indirectly from other bees in the hive. (5) The information takes the form of an inhibitory cue. The proposed mechanism for the regulation of pollen foraging involves a hierarchical system of information acquisition and a negative feedback loop. By taking advantage of the vast processing capacity of large numbers of individuals working in parallel, such a system of information acquisition and dissemination may be ideally suited to promote efficient regulation of labor within the colony. Although each individual relies on only limited, local information, the colony as a whole achieves a finely-tuned response to the changing conditions it experiences. 相似文献
13.
Honey bee foragers specialize on collecting pollen and nectar. Pollen foraging behavior is modulated by at least two stimuli
within the nest: the presence of brood pheromone and young larvae and the quantity of stored pollen. Genetic variation in
pollen foraging behavior has been demonstrated repeatedly. We used selected high and low pollen-hoarding strains of bees that
differ dramatically in the quantity of pollen collected to determine if the observed differences in foraging could be explained
by differential responses to brood stimuli. Workers from the high and low pollen-hoarding strains and wild-type bees were
co-fostered in colonies with either brood or no brood. As expected based on previous studies, returning high pollen-hoarding
foragers collected heavier pollen loads and lighter nectar loads than low pollen-hoarding bees. Effects of brood treatment
were also observed; bees exposed to brood collected heavier pollen loads and initiated foraging earlier than those from broodless
colonies. More specifically, brood treatment resulted in increased pollen foraging in high pollen-hoarding bees but did not
affect pollen foraging in low pollen-hoarding bees, suggesting that high pollen-hoarding bees are more sensitive to the presence
of brood. However, response to brood stimuli does not sufficiently explain the differences in foraging behavior between the
strains since these differences persisted even in the absence of brood. 相似文献
14.
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
15.
We examined whether the quality (concentration) of incoming sucrose solutions returned by foraging honey bees affected the response thresholds of pre-foraging members of the colony. Six pairs of colonies were given ad libitum access to sucrose solution feeders. A colony from each pair was switched from 20–50% sugar concentration feeders while the other continued to have access to 20% sucrose feeders. Proboscis extension response (PER) scores to an increasing series of sucrose concentrations were significantly higher in pre-foragers of colonies foraging on 20% sucrose throughout compared to pre-foragers in colonies where foraging was switched to 50% sucrose. Although all colonies had honey stores, the concentration of sugar solution in non-foraging bees crops were significantly lower in bees from colonies foraging on 20% sucrose compared to those from colonies foraging on 50% sucrose. Because response thresholds to sugar of young bees were modulated by the concentration of sucrose solution returned to colonies, we repeated the 2000 study of Pankiw and Page that potentially confounded baseline response thresholds with modulated scores due to experience in the colony. Here, we examined PER scores to sucrose in bees within 6 h of emergence, prior to feeding experience, and their forage choice 2 to 3 weeks later. Pollen foragers had higher PER scores as newly emerged bees compared to bees that eventually became nectar foragers. These results confirm those of the 2000 study by Pankiw and Page. Combined, these experiments demonstrate that variation in pre-forager sucrose response thresholds are established prior to emerging as adults but may be modulated by incoming resources later on. Whether this modulation has long-term effects on foraging behavior is unknown but modulation has short-term effects and the potential to act as a means of communication among all bees in the colony.Communicated by M. Giurfa 相似文献
16.
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
17.
Deborah M. Gordon 《Behavioral ecology and sociobiology》1992,31(6):417-427
Summary Colonies of the harvester ant, Pogonomyrmex barbatus, adjust the direction and length of foraging trails in response to the foraging behavior of their conspecific neighbors. In the absence of any interaction with its neighbor, a mature colony expands its foraging range at a rate of 0.85 ± 0.15 m per day. Exclusion experiments show that if a colony is prevented from using its foraging trails, the neighbors of that colony will enter its foraging range within 10 days. Exclusion experiments were performed with three age classes of colonies: young (1 year old), intermediate (3–4 years old), and old (5 years old or more). Colonies 3–4 years old are most likely to expand foraging ranges, and to retain newly-gained areas. To examine the relation of colony age (in years) and colony size (in numbers of workers), colonies of known age were excavated. Colonies increase greatly in size in years 3 and 4. Foraging area may be of greater current or prospective value for younger, smaller, quickly growing colonies than for older, larger ones of stable size.
Correspondence to the second address 相似文献
18.
Summary To place social insect foraging behavior within an evolutionary context, it is necessary to establish relationships between individual foraging decisions and parameters influencing colony fitness. To address this problem, we examined interactions between individual foraging behavior and pollen storage levels in the honey bee, Apis mellifera L. Colonies responded to low pollen storage conditions by increasing pollen intake rates 54% relative to high pollen storage conditions, demonstrating a direct relationship between pollen storage levels and foraging effort. Approximately 80% of the difference in pollen intake rates was accounted for by variation in individual foraging effort, via changes in foraging activity and individual pollen load size. An additional 20% resulted from changes in the proportion of the foraging population collecting pollen. Under both high and low pollen storage treatments, colonies returned pollen storage levels to pre-experimental levels within 16 days, suggesting that honey bees regulate pollen storage levels around a homeostatic set point. We also found a direct relationship between pollen storage levels and colony brood production, demonstrating the potential for cumulative changes in individual foraging decisions to affect colony fitness.
Offprint requests to: J.H. Fewell at the current address 相似文献
19.
The age at which worker honey bees begin foraging varies under different colony conditions. Previous studies have shown that
juvenile hormone (JH) mediates this behavioral plasticity, and that worker-worker interactions influence both JH titers and
age at first foraging. These results also indicated that the age at first foraging is delayed in the presence of foragers,
suggesting that colony age demography directly influences temporal division of labor. We tested this hypothesis by determining
whether behavioral or physiological development can be accelerated, delayed, or reversed by altering colony age structure.
In three out of three trials, earlier onset of foraging was induced in colonies depleted of foragers compared to colonies
depleted of an equal number of bees across all age classes. In two out of three trials, delayed onset of foraging was induced
in colonies in which foragers were confined compared to colonies with free-flying foragers. Finally, in three out of three
trials, both endocrine and exocrine changes associated with reversion from foraging to brood care were induced in colonies
composed of all old bees and devoid of brood; JH titers decreased and hypopharyngeal glands regenerated. These results demonstrate
that plasticity in age-related division of labor in honey bee colonies is at least partially controlled by social factors.
The implications of these results are discussed for the recently developed ‘‘activator-inhibitor” model for honey bee behavioral
development.
Received: 8 November 1995/Accepted after revision: 10 May 1996 相似文献
20.
The impact of a parasitic infestation may be influenced by nutritional state, in both individuals and colonies. This study
examined the interaction between pollen storage and the effects of an infestation by the mite, Varroa jacobsoni Oudemans, in colonies of the honey bee, Apis mellifera L. We manipulated the pollen storage and mite infestation levels of colonies, and measured pollen foraging and brood rearing.
Increased pollen stores decreased both the number of pollen foragers and pollen load size, while initially at least foragers
from colonies with moderate infestations carried smaller pollen loads than those from lightly infested colonies. Over the
course of the experiment, all colonies significantly increased pollen-foraging rates and pollen consumption, which was presumably
a seasonal effect. Lightly infested colonies exhibited a larger increase in pollen forager number than moderately infested
colonies, suggesting that more intense mite infestations compromised forager recruitment. Brood production was not affected
by the addition of pollen, but moderately infested colonies were rearing significantly less brood by the end of the experiment
than lightly infested colonies. Furthermore, the efficiency with which colonies converted pollen to brood decreased as the
pollen storage level decreased and the infestation level increased. The results of this study may indicate that honey bee
colonies adaptively alter brood-production efficiency in response to parasitic infestations and seasonal changes.
Received: 3 May 1999 / Received in revised form: 14 September 1999 / Accepted: 25 September 1999 相似文献