The Honeybee's Protrusible Glossa is a Compliant Mechanism

Many biological structures can perform highly-dexterous actions by using dynamic surfaces.To deal with the contradictive demands of high feeding efficiency and low energy expenditure during nectar feeding,the glossal surface of a honeybee undergoes shape changes,in which glossal hairs erect together with segment elongation in a drinking cycle.In this paper,we extracted a transmission link embedded in the glossa from postmortem examination and found that the compliance of the intersegmental membranes provides more possibilities for this highly kinematic synchronicity.According to the morphing phenomena of honeybee's glossa,we proposed a compliant mechanism model to predict the deformation behavior of honeybee considering elastic properties of the glossal intersegmental membranes.The increase of membrane stiffness may improve the capacity of elastic potential energy transfer,but will still result in the increase of mass.An index is introduced to evaluate the contradiction for optimizing structural parameters.This work may arouse new prospects for conceptual design of micro-mechanical systems equipped with bio-inspired compliant mechanisms.     

Preference of cabbage white butterflies and honey bees for nectar that contains amino acids

Summary Amino acids occur in most floral nectars but their role in pollinator attraction is relatively unstudied. Nectars of butterfly-pollinated flower tend to have higher concentrations of amino acids than do flowers pollinated by bees and many other animals, suggesting that amino acids are important attractants of butterflies to flowers. In order to determine whether amino acids are important in attracting butterflies and bees, we tested the preference of cabbage white butterflies (Pieris rapae) and honey bees (Apis mellifera) by allowing them to feed from artificial flowers containing sugar-only or sugar-amino acid mimics ofLantana camara nectar. Honey bees and female cabbage white butterflies consumed more sugar-amino acid nectar than sugar-only nectar. In addition, female cabbage white butterflies visited artificial flowers containing sugar-amino acid nectars more frequently than flowers containing sugar-only nectars; honey bees spent more time consuming the sugar-amino acid nectar. Male cabbage white butterflies did not discriminate between the two nectars. These results support the hypothesis that the amino acids of nectar contribute to pollinator attraction and/or feeding.     

Bees get a head start on honey production

Nectar concentration is assumed to remain constant during transport by honeybees between flowers and hive. We sampled crop contents of nectar foragers on Aloe greatheadii var. davyana, a major winter bee plant in South Africa. The nectar is dilute (approx. 20% w/w), but the crop contents of bees captured on flowers are significantly more concentrated. In returning foragers, the concentration increases further to 38–40%, accompanied by a volume decrease. The doubling of sugar concentration suggests that nectar is regurgitated onto the tongue and evaporated during foraging and on the return flight. Processing of the dilute nectar into honey thus begins early, aided by low ambient humidities. This has implications for honeybee thermoregulation, water balance and energetics during foraging, and for the communication of nectar quality to recruits.     

Impact of Food Availability,Pathogen Exposure,and Genetic Diversity on Thermoregulation in Honey Bees (Apis mellifera)

Accurate thermoregulation in honey bees is crucial for colony survival. Multiple factors influence how colonies manage in-hive temperature, including genetic diversity. We explored the influence of genetic diversity on thermoregulatory behavior under three conditions: natural foraging, supplemental feeding, and exposure to the fungal pathogen shown to induce a social fever in honey bees. Our data suggest that (1) the degree of genetic diversity expected under normal conditions is not predictive of thermoregulatory stability, (2) the social fever response of honey bees is not a simple stimulus–response mechanism but appears to be influenced by ambient temperature conditions, and (3) a temperature-based circadian rhythm emerges under high nectar flow conditions. Taken together, these data suggest that a richer, context-dependent thermoregulatory system exists in honey bees than previously understood.     

Heartbeat rate modulation mediated by the ventral nerve cord in the honey bee,Apis mellifera

Summary Using impedance conversion as a recording technique, heartbeat patterns were established for intact adult honey bee workers during rest and during nectar feeding and locomotion, activities that both result in accelerating heartbeat. Heartbeat patterns were then described separately for bees in which assumed neural regulation was disrupted (the ventral nerve cord was transected), and for bees in which assumed neurohormonal regulation was disrupted (blood circulation to the abdomen was blocked). Heartbeat patterns from these latter two groups were compared with the baseline data from the former three groups. Results showed that normal modulatory patterns occurring during feeding and locomotion persisted when the neurohormonal pathway was disrupted, but the same modulation was absent when the neural pathway was disrupted. These results offer clear support for the conclusion that neural mechanisms provide a primary form of regulation of heartbeat in the honey bee.     

Regulation of nectar collection in relation to honey storage levels by honey bees, Apis mellifera

Honey bees collect distinct nutrient sources in the form ofnectar (energy) and pollen (nitrogen). We investigated the effectof varying energy stores on nectar and pollen foraging. We foundno significant changes in nectar foraging in response to changesin honey storage levels within colonies. Individual foragersdid not vary activity rates or nectar load sizes in responseto changes in honey stores, and colonies did not increase nectarintake rates when honey stores within the hive were decreased.This result contrasts with pollen foraging behavior, which isextremely sensitive to colony state. Our data show that individualforaging decisions during nectar collection and colony regulationof nectar intake are distincdy different from pollen foraging.The behavior of honey bees illustrates that foraging strategy(and therefore foraging models) can incorporate multiple currencies,including both energy and protein intake.[Behav Ecol 7: 286–291(1996)]     

Imidacloprid Alters Foraging and Decreases Bee Avoidance of Predators

Concern is growing over the effects of neonicotinoid pesticides, which can impair honey bee cognition. We provide the first demonstration that sublethal concentrations of imidacloprid can harm honey bee decision-making about danger by significantly increasing the probability of a bee visiting a dangerous food source. Apis cerana is a native bee that is an important pollinator of agricultural crops and native plants in Asia. When foraging on nectar containing 40 µg/L (34 ppb) imidacloprid, honey bees (Apis cerana) showed no aversion to a feeder with a hornet predator, and 1.8 fold more bees chose the dangerous feeder as compared to control bees. Control bees exhibited significant predator avoidance. We also give the first evidence that foraging by A. cerana workers can be inhibited by sublethal concentrations of the pesticide, imidacloprid, which is widely used in Asia. Compared to bees collecting uncontaminated nectar, 23% fewer foragers returned to collect the nectar with 40 µg/L imidacloprid. Bees that did return respectively collected 46% and 63% less nectar containing 20 µg/L and 40 µg/L imidacloprid. These results suggest that the effects of neonicotinoids on honey bee decision-making and other advanced cognitive functions should be explored. Moreover, research should extend beyond the classic model, the European honey bee (A. mellifera), to other important bee species.     

Bee visitation rates to cultivated sunflowers increase with the amount and accessibility of nectar sugars

Pollinators make foraging decisions based on numerous floral traits, including nectar and pollen rewards, and associated visual and olfactory cues. For insect‐pollinated crops, identifying and breeding for attractive floral traits may increase yields. In this study, we examined floral trait variation within cultivated sunflowers and its effects on bee foraging behaviours. Over 2 years, we planted different sunflower inbred lines, including male‐fertile and male‐sterile lines, and measured nectar volume, nectar sugar concentration and composition, and corolla length. During bloom, we recorded visits by both managed honey bees and wild bees. We then examined consistency in relative nectar production by comparing field results to those from a greenhouse experiment. Sunflower inbred lines varied significantly in all floral traits, including the amount and composition of nectar sugars, and in corolla length. Both wild bee and honey bee visits significantly increased with nectar sugar amount and decreased with corolla length, but appeared unaffected by nectar sugar composition. While wild bees made more visits to sunflowers providing pollen (male‐fertile), honey bees preferred plants without pollen (male‐sterile). Differences in nectar quantity among greenhouse‐grown sunflower lines were similar to those measured in the field, and bumble bees preferentially visited lines with more nectar in greenhouse observations. Our results show that sunflowers with greater quantities of nectar sugar and shorter corollas receive greater pollination services from both managed and wild bees. Selecting for these traits could thus increase sunflower crop yields and provide greater floral resources for bees.     

The pollination efficiency of a pollinator depends on its foraging strategy,flowering phenology,and the flower characteristics of a plant species

Honey bees and stingless bees are generalist visitors of several wild and cultivated plants. They forage with a high degree of floral fidelity and thereby help in the pollination services of those plants. We hypothesized that pollination efficiency might be influenced by flowering phenology, floral characteristics, and resource collection modes of the worker bees. In this paper, we surveyed the foraging strategies of honey bees (Apis cerana, Apis dorsata, and Apis florea) and stingless bees (Tetragonula iridipennis) concerning their pollination efficiencies. Bees showed different resource gathering strategies, including legitimate (helping in pollination as mixed foragers and specialized foragers) and illegitimate (serving as nectar robbers and pollen thieves) types of flower visitation patterns. Foraging strategies are influenced by the shape of flowers, the timing of the visitation, floral richness, and bee species. Honey bees and stingless bees mainly acted as legitimate visitors in most plants studied. Sometimes honey bees served as nectar robbers in tubular flowers and stingless bees as pollen thieves in large-sized flowers. Among the legitimate categories, mixed foragers have a comparatively lower flower visitation rate than the specialized nectar and pollen foragers. However, mixed foragers have greater abundance and higher values of the single-visit pollination efficiency index (PEi) than nectar and pollen foragers. The value of the combined parameter ‘importance in pollination (PI)’ was thus higher in mixed foragers than in nectar and pollen foragers.     

Flowers morphology and nectar concentration determine the preferred food source of stingless bee,Heterotrigona itama

Heterotrigona itama is a stingless bee species from Meliponini tribe. The bee collects nectar, pollen and resin to produce honey, bee bread, and propolis. The bee is also known to visit and collect nectar from various types of flowers but there are limited studies on why this species of bee prefers to visit certain types of flowers. This study was conducted to identify the nectar concentration in selected flowers favoured by H. itama and the relationship between the bee and the morphology of the flowers. Nectar was obtained from different species of flowers and the concentrations were measured using a digital refractometer. The tube length of each flower species and the tongue length of the bees were also measured. The results revealed that flowers preferred by H. itama have high nectar concentrations. The tube lengths of the preferred flowers were between 2.0 and 4.0 mm, which is compatible with the tongue length of the bee. This study revealed that both nectar concentration and flower morphology are important factors for the bees in choosing their food sources. The results from this study will benefit the beekeepers in the identification of flowers that should be planted in their farms to improve stingless bee beekeeping activities. Understanding the relationship between the bees and their flower preferences could also help us to understand the importance of conserving both the bee colonies and the various species of flowering plants to ensure the sustainability of flora and fauna in the ecosystem.     

Honey bees and wild pollinators differ in their preference for and use of introduced floral resources

Introduced plants may be important foraging resources for honey bees and wild pollinators, but how often and why pollinators visit introduced plants across an entire plant community is not well understood. Understanding the importance of introduced plants for pollinators could help guide management of these plants and conservation of pollinator habitat. We assessed how floral abundance and pollinator preference influence pollinator visitation rate and diversity on 30 introduced versus 24 native plants in central New York. Honey bees visited introduced and native plants at similar rates regardless of floral abundance. In contrast, as floral abundance increased, wild pollinator visitation rate decreased more strongly for introduced plants than native plants. Introduced plants as a group and native plants as a group did not differ in bee diversity or preference, but honey bees and wild pollinators preferred different plant species. As a case study, we then focused on knapweed (Centaurea spp.), an introduced plant that was the most preferred plant by honey bees, and that beekeepers value as a late‐summer foraging resource. We compared the extent to which honey bees versus wild pollinators visited knapweed relative to coflowering plants, and we quantified knapweed pollen and nectar collection by honey bees across 22 New York apiaries. Honey bees visited knapweed more frequently than coflowering plants and at a similar rate as all wild pollinators combined. All apiaries contained knapweed pollen in nectar, 86% of apiaries contained knapweed pollen in bee bread, and knapweed was sometimes a main pollen or nectar source for honey bees in late summer. Our results suggest that because of diverging responses to floral abundance and preferences for different plants, honey bees and wild pollinators differ in their use of introduced plants. Depending on the plant and its abundance, removing an introduced plant may impact honey bees more than wild pollinators.     

Adult pollen diet essential for egg maturation by a solitary Osmia bee

Reproduction is a nutritionally costly activity for many insects, as their eggs are rich in lipids and proteins. That cost seems especially acute for non-social bees, which for their size, lay enormous eggs. All adult female bees visit flowers, most of them to collect pollen and nectar, or sometimes oils, to feed their progeny. For adult bees, the need for pollen feeding has only been detailed for the honey bee, Apis mellifera. To experimentally test for the reproductive value of adult pollen feeding by a non-social bee, Osmia californica (Hymenoptera: Apiformes: Megachilidae), young female bees plus males were released into large glasshouse cages provided with either a male-fertile sunflower cultivar or a pollen-less one. Females regularly visited and drank nectar from flowers of both cultivars. Abundant orange pollen was seen regularly in guts of females confined with the male-fertile sunflowers, indicative of active pollen ingestion. All females’ terminal oocytes (next egg to be laid) were small at emergence. Oocytes of females confined with the pollen-less sunflowers remained small, despite frequent nectaring and exposure to other floral stimuli. In contrast, the basal oocytes of female O. californica with access to pollen had swelled to full size within ten days following emergence, enabling them to lay eggs in provided nest tubes. Adult females of this solitary bee required dietary pollen to reproduce; nitrogen stores acquired as larvae were inadequate. Early and regular pollen feeding in part paces the onset and maximum tempo of solitary bees’ lifetime reproductive output.     

Comparative Sucrose Responsiveness in Apis mellifera and A. cerana Foragers

In the European honey bee, Apis mellifera, pollen foragers have a higher sucrose responsiveness than nectar foragers when tested using a proboscis extension response (PER) assay. In addition, Africanized honey bees have a higher sucrose responsiveness than European honey bees. Based on the biology of the Eastern honey bee, A. cerana, we hypothesized that A. cerana should also have a higher responsiveness to sucrose than A. mellifera. To test this hypothesis, we compared the sucrose thresholds of pollen foragers and nectar foragers in both A. cerana and A. mellifera in Fujian Province, China. Pollen foragers were more responsive to sucrose than nectar foragers in both species, consistent with previous studies. However, contrary to our hypothesis, A. mellifera was more responsive than A. cerana. We also demonstrated that this higher sucrose responsiveness in A. mellifera was not due to differences in the colony environment by co-fostering two species of bees in the same mixed-species colonies. Because A. mellifera foragers were more responsive to sucrose, we predicted that their nectar foragers should bring in less concentrated nectar compared to that of A. cerana. However, we found no differences between the two species. We conclude that A. cerana shows a different pattern in sucrose responsiveness from that of Africanized bees. There may be other mechanisms that enable A. cerana to perform well in areas with sparse nectar resources.     

Nectar robbery by bees Xylocopa virginica and Apis mellifera contributes to the pollination of rabbiteye blueberry

Honey bees, Apis mellifera L., probe for nectar from robbery slits previously made by male carpenter bees, Xylocopa virginica (L.), at the flowers of rabbiteye blueberry, Vaccinium ashei Reade. This relationship between primary nectar robbers (carpenter bees) and secondary nectar thieves (honey bees) is poorly understood but seemingly unfavorable for V. ashei pollination. We designed two studies to measure the impact of nectar robbers on V. ashei pollination. First, counting the amount of pollen on stigmas (stigmatic pollen loading) showed that nectar robbers delivered fewer blueberry tetrads per stigma after single floral visits than did our benchmark pollinator, the southeastern blueberry bee, Habropoda laboriosa (F.), a recognized effective pollinator of blueberries. Increasing numbers of floral visits by carpenter bee and honey bee robbers yielded larger stigmatic loads. As few as three robbery visits were equivalent to one legitimate visit by a pollen-collecting H. laboriosa female. More than three robbery visits per flower slightly depressed stigmatic pollen loads. In our second study, a survey of 10 commercial blueberry farms demonstrated that corolla slitting by carpenter bees (i.e., robbery) has no appreciable affect on overall V. ashei fruit set. Our observations demonstrate male carpenter bees are benign or even potentially beneficial floral visitors of V ashei. Their robbery of blueberry flowers in the southeast may attract more honey bee pollinators to the crop.     

Foraging behavior affects pollen removal and deposition in Impatiens capensis (Balsaminaceae)

Flowers of most plant species are visited by a variety of animals. Some of these visitors are effective pollinators while others remove resources without transferring pollen. Studies comparing the effectiveness of different visitors as pollinators often compare taxa without considering variation in behavior within a taxon. Wilson and Thomson (Ecology 72: 1503-1507, 1991) documented the effects of honey bees and bumble bees on the pollination dynamics of Impatiens capensis. They found that pollen-collecting honey bees removed large numbers of pollen grains from anthers but deposited little of it on stigmas; bumble bees, which sought nectar, removed less pollen but deposited more of it on stigmas. It is unclear whether the low pollen transfer efficiencies of honey bees are explained by their morphology or by their pollen-collecting behavior. We repeated the work of Wilson and Thomson at a site where honey bees were foraging for nectar, not pollen. We measured the quantity of pollen remaining in anthers, the number of pollen grains deposited on stigmas, and seed production after single visits by honey bees and bumble bees. The differences between the taxa disappeared when they were foraging in a similar manner. Our results clearly demonstrate the importance of foraging behavior on the pollination effectiveness of floral visitors.     

Accumulation of human EGF in nectar of transformed plants of Nicotiana langsdorffii x N. sanderae and transfer to honey by bees

Honey has been used successfully in wound healing for thousands of years. The peptide hormone human epidermal growth factor (hEGF) is also known to have a beneficial effect in various wound healing processes via mechanisms that differ from those for honey. In this study, we show that hEGF can be incorporated into honey via nectar. Plants of Nicotiana langsdorffii × N. sanderae were transformed with the gene for hEGF, equipped with a nectary‐targeted promoter and a signal sequence for secretion to nectar. These plants accumulated hEGF in the nectar. The maximum hEGF concentration recorded with ELISA in these plants is 2.5 ng·ml^?1. There is a significant linear relationship (P < 0.001) between hEGF concentration and induction of hEGF‐receptor phosphorylation. Since the flower morphology of these plants did not allow production of honey from their nectar, we used feeding solutions, spiked with synthetic hEGF, to study transfer of this peptide into honey through bee activity. Transfer of hEGF from a feeding solution to honey by bees occurred with retention of the hEGF concentration and the capacity to induce hEGF‐receptor phosphorylation. These observations indicate that plants can function as a production platform for honey containing biologically active peptides, which may enhance wound healing and other biological processes.     

Nectar utilization and pollination by Australian honeyeaters and insects visiting Calothamnus quadrifidus (Myrtaceae)

Nectar availability in Calothamnus quadrifidus flowers was studied at Wongamine Nature Reserve in late spring (November). Despite some overnight depletion by moths and other invertebrates, more nectar was present in flowers at dawn than at the preceding dusk. Significant nectar depletion occurred within a few hours after dawn, mainly due to foraging by two honeyeater species. Lichmera indistincta and Phylidonyris nigra. Thereafter, nectar availability was maintained at relatively low levels, principally because of foraging by honeyeaters and honey bees. Apis mellifera, that became active during the warmer part of the day. Although individual honeyeaters consumed more nectar than A. mellifera, honey bees were so abundant that their total impact was greater than that of either honeyeater species for much of the day. Transfer of C. quadrifidus pollen between flowers is necessary in order to achieve a high level of seed set, as the flowers appear to be protandrous. Honeyeaters appeared to be considerably more significant pollen vectors than A. mellifera.     

Invasive ant (<Emphasis Type="Italic">Anoplolepis gracilipes</Emphasis>) disrupts pollination in pumpkin

Yellow crazy ant (Anoplolepis gracilipes (F. Smith); “YCA”) is known for its aggressive predatory ability and ability to exert exploitation competition on both native and other invasive ants via floral nectar. We argue that YCA invasion can exert both interference and exploitation competition on legitimate pollinators. In pumpkin fields (Cucurbita maxima L.) of south India, YCA infested the flowers, particularly the pistillate flowers, for nectar foraging. Pumpkin is a honey bee-mediated cross-pollinated monoecious plant that produces disproportionately very few pistillate flowers. We hypothesize that YCA presence in the flowers can affect the visitation rate and foraging time of honey bees in the flowers, the fruit set in pumpkins, and can exert predatory pressure on the honey bees if the bees linger in ant-colonized flowers. Both YCA and honey bees preferred to forage on the limited pistillate flowers in the plants. After colonizing the flowers, YCA did not retreat for hours, even upon disturbance by competitors, such as honey bees. Both the visitation frequency and the foraging time of honey bees were drastically reduced in ant-colonized flowers, and none of the ant-colonized flowers developed into fruits, suggesting that the YCA exert both an ecological and evolutionary pressure on pumpkin. The ants preyed upon about 17% of the honey bees that lingered in ant-colonized flowers, and the time the bees spent foraging predicted the fate of the bees. Exploitation competition exerted by the YCA on pumpkin may have far-reaching consequences for the pollination and productivity of this cash crop.     

Post‐ingestive effect of plant phenolics on the feeding behaviour of the honeybee Apis cerana

As the staple food of honey bees, honey is rich in plant phenolics derived from pollen, nectar and resin. Most studies concentrate on the temporary response of bees' peripheral chemoreceptors to these chemicals, and the post‐ingestive effects of plant phenolics are largely ignored. In the present study, a series of feeding experiments are conducted to test whether plant phenolics modulate the response thresholds and rhythmic behaviour of the honeybee Apis cerana (Ruttner). The results of the study demonstrate that bees fed with syrup containing high concentrations of phenlics reduce their response thresholds greatly, and shift their feeding rhythms significantly. Because the forager response thresholds determine their foraging choice, and their rhythmic behaviour is required for timing visits to flowers, enhanced plant phenolics as a result of global environment change may change the bees' pollination service in our changing world.     

Nutritional limitation and resistance to opportunistic Aspergillus parasites in honey bee larvae

Honey bees are threatened by land use changes which reduce the availability and diversity of pollen and nectar resources. There is concern that poor nutrition may be involved in recent population declines, either directly or due to indirect effects on immunocompetence. The larval stage is likely to be the most vulnerable to a poor diet, but the effects of larval nutrition on the disease susceptibility of bees are not well known. In this study we used laboratory-reared honey bee larvae to investigate the effects of diet quality on disease susceptibility to the opportunistic fungal parasites Aspergillus flavus, Aspergillus phoenicis and A. fumigatus. Larvae fed on a nutritionally poor diet were found to be significantly more susceptible to A. fumigatus. Larval resistance to A. fumigatus was enhanced by feeding with a diet supplemented with either dandelion or polyfloral pollens. This indicates that dandelion and polyfloral pollens contain elements that enhance resistance to this fungal disease, illustrating an interaction between nutrition and parasitism and emphasising the benefit of diverse floral resources in the environment to maintain honey bee health.