On a simplified model for pattern formation in honey bee colonies

We present a simplified version of a previously presented model (Camazine et al. (1990)) that generates the characteristic pattern of honey, pollen and brood which develops on combs in honey bee colonies. We demonstrate that the formation of a band of pollen surrounding the brood area is dependent on the assumed form of the honey and pollen removal terms, and that a significant pollen band arises as the parameter controlling the rate of pollen input passes through a bifurcation value. The persistence of the pollen band after a temporary increase in pollen input can be predicted from the model. We also determine conditions on the parameters which ensure the accumulation of honey in the periphery and demonstrate that, although there is an important qualitative difference between the simplified and complete models, an analysis of the simplified version helps us understand many biological aspects of the more complex complete model. Corresponding author     

Brood Pheromone Modulation of Pollen Forager Turnaround Time in the Honey Bee (<Emphasis Type="Italic">Apis mellifera</Emphasis> L.)

Foraging for pollen is an important behavior of the honey bee because pollen is their sole source of protein. Through nurse bees, larvae are the principal consumers of pollen. Fatty acid esters extractable from the surface of larvae, called brood pheromone, release multiple colony-level and individual foraging behaviors increasing pollen intake. In this study pollen forager turnaround time was measured in observation hives supplemented with brood pheromone versus a blank control treatment. Treatment with brood pheromone significantly decreased pollen forager turnaround time in the hive between foraging bouts by approximately 72%. Concurrently, brood pheromone increased the ratio of pollen to non-pollen foragers entering colonies. Brood pheromone has been shown to release most of the mechanisms known to increase pollen intake by colonies acting as an important regulator of colony foraging decisions and growth.     

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.     

Pollen Contamination of Honey by Bees inside the Hive

This paper aims to assess to what degree and in what way the brood chamber affects the pollen content of the honey. Twenty-nine pieces of comb containing only honey were cut from different frames of hives. The percentage of cells in each frame occupied by the brood chamber and the distance between these cells and the cut piece were recorded. A honey sample was extracted from each comb piece, avoiding any contamination with pollen, its sediment examined under the microscope and its botanical constituents identified and counted. The results show that the pollen content of honey was higher in samples from frames containing brood or pollen cells; in these samples the pollen content was positively correlated with the amount of these cells in the frame and the proximity of the honey to them. The proportion of pollen grains from principally nectariferous plants was lower in honeys with a high pollen content.     

The relationship between comb age and performance of honey bee (Apis mellifera) colonies

A study on the relationship between the age of comb and the activity of the hybrid Carniolan honey bee colonies in collecting pollen activity, worker brood production, colony strength, and honey yield was conducted. In comparison to colonies with combs aged 4-years, colonies with combs aged 1, 2 and 3-years significantly exceeded in the number returning workers, number returning workers with pollen loads, rate of storing pollen, rate of worker brood production, and size of colony population. Colonies with combs aged 1, 2 and 3-years produced significantly more honey than colonies with combs aged 4-years (5.25, 4.90 and 4.65 kg/colony vs. 4.45 kg/colony, respectively). It can be concluded that the foraging rate, gathering and storing pollen, brood production, colony population size, and honey yield significantly depended on the age of combs. Beekeepers can replace old combs with new ones to increase brood and honey production.     

Several workers lay eggs in the same brood cell in queenless honey bee (Apis mellifera) colonies

Queenless honey bee (Apis mellifera) colonies are often characterized by the presence of multiple eggs in brood cells. This is surprising because only one egg can be reared to maturity per cell. Moreover, worker honey bees cannot produce many eggs per day. There are several reasons that could explain the presence of multiple eggs in single cells: a) workers cannot control how many eggs they release; in this case we would expect all eggs to be from the same mother; b) excess eggs could be provided as food for the first larva to hatch in the absence of adequate brood care and this would again result in all eggs in one cell sharing the same mother; c) the number of cells available for oviposition may be limiting, obliging workers to lay eggs in cells that already contain eggs, resulting in eggs of mixed maternity. Here we show that the majority of brood cells in queenless colonies contain eggs from multiple mothers. Therefore our results suggest that the presence of multiple eggs in brood cells arises from a limitation on the number of suitable cells available for oviposition. Received 29 September 2008; revised 21 November 2008; accepted 24 November 2008.     

Effects of brood pheromone (SuperBoost) on consumption of protein supplement and growth of honey bee (Hymenoptera: Apidae) colonies during fall in a northern temperate climate

Honey bee, Apis mellifera L. (Hymenoptera: Apidae), nutrition is vital for colony growth and maintenance of a robust immune system. Brood rearing in honey bee colonies is highly dependent on protein availability. Beekeepers in general provide protein supplement to colonies during periods of pollen dearth. Honey bee brood pheromone is a blend of methyl and ethyl fatty acid esters extractable from cuticle of honey bee larvae that communicates the presence of larvae in a colony. Honey bee brood pheromone has been shown to increase protein supplement consumption and growth of honey bee colonies in a subtropical winter climate. Here, we tested the hypothesis that synthetic brood pheromone (SuperBoost) has the potential to increase protein supplement consumption during fall in a temperate climate and thus increase colony growth. The experiments were conducted in two locations in Oregon during September and October 2009. In both the experiments, colonies receiving brood pheromone treatment consumed significantly higher protein supplement and had greater brood area and adult bees than controls. Results from this study suggest that synthetic brood pheromone may be used to stimulate honey bee colony growth by stimulating protein supplement consumption during fall in a northern temperate climate, when majority of the beekeepers feed protein supplement to their colonies.     

Division of Labor Associated with Brood Rearing in the Honey Bee: How Does It Translate to Colony Fitness?

Division of labor is a striking feature observed in honey bees and many other social insects. Division of labor has been claimed to benefit fitness. In honey bees, the adult work force may be viewed as divided between non-foraging hive bees that rear brood and maintain the nest, and foragers that collect food outside the nest. Honey bee brood pheromone is a larval pheromone that serves as an excellent empirical tool to manipulate foraging behaviors and thus division of labor in the honey bee. Here we use two different doses of brood pheromone to alter the foraging stimulus environment, thus changing demographics of colony division of labor, to demonstrate how division of labor associated with brood rearing affects colony growth rate. We examine the effects of these different doses of brood pheromone on individual foraging ontogeny and specialization, colony level foraging behavior, and individual glandular protein synthesis. Low brood pheromone treatment colonies exhibited significantly higher foraging population, decreased age of first foraging and greater foraging effort, resulting in greater colony growth compared to other treatments. This study demonstrates how division of labor associated with brood rearing affects honey bee colony growth rate, a token of fitness.     

The provision of supplementary food to hive bees

Sucrose syrup is less satisfactory than honey as a winter and spring food for bees: a mixture of approximately equal parts of honey and syrup is of almost the same value as honey alone.
Feeding confined to early autumn induced the strongest spring development of colonies; syrup feeding in the spring may retard colony development, and food supplied at this time is apparently wasted. Feeding with syrup and pollen is advantageous only when a colony is lacking in stores of carbohydrate and protein.
A total of four British Standard brood combs full of pollen provides sufficient protein for a colony on B.S. equipment from autumn until April: the best results were obtained by providing colonies on eleven B.S. combs, in early autumn, with 35-40 lb. of honey, or honey and concentrated sucrose syrup, and four brood combs full of pollen.     

Longevity and reproductive success of Aethina tumida (Coleoptera: Nitidulidae) fed different natural diets

The longevity and reproductive success of newly emerged, unfed adult Aethina tumida Murray assigned different diets (control = unfed; honey-pollen; honey; pollen; empty brood comb; bee brood; fresh Kei apples; and rotten Kei apples) were determined. Longevity in honey-fed small hive beetle adults (average maximum: 167 d) was significantly higher than on other diets. Small hive beetles fed empty brood comb lived significantly longer (average maximum: 49.8 d) than unfed beetles (average maximum: 9.6 d). Small hive beetle offspring were produced on honey-pollen, pollen, bee brood, fresh Kei apples, and rotten Kei apples but not on honey alone, empty brood comb, or in control treatments. The highest reproductive success occurred in pollen fed adults (1773.8 +/- 294.4 larvae per three mating pairs of adults). The data also show that A. tumida can reproduce on fruits alone, indicating that they are facultative parasites. The pupation success and sex ratio of small hive beetle offspring were also analyzed. Larvae fed pollen, honey-pollen, or brood had significantly higher pupation success rates of 0.64, 0.73, and 0.65 respectively than on the other diets. Sex ratios of emerging adults fed diets of pollen or brood as larvae were significantly skewed toward females. Because small hive beetle longevity and overall reproductive success was highest on foodstuffs located in honey bee colonies, A. tumida are efficient at causing large-scale damage to colonies of honey bees resulting in economic injury for the beekeeper. Practical considerations for the control of A. tumida are briefly discussed.     

Influence of pollen diet in spring on development of honey bee (Hymenoptera: Apidae) colonies

The effects of changes in spring pollen diet on the development of honey bee, Apis mellifera L. (Hymenoptera: Apidae), colonies were examined in a 3-yr study (2002-2004). Pollen-supplemented and pollen-limited conditions were created in colonies every spring, and brood rearing and honey yields were subsequently monitored throughout the summer. In all 3 yr, colonies that were supplemented with pollen or a pollen substitute in the spring started rearing brood earlier than colonies in other treatment groups and produced the most workers by late April or early May. In 2002, these initial differences were reflected by a two-fold increase in annual honey yields by September for colonies that were pollen-supplemented during the spring compared with pollen-limited colonies. In 2003 and 2004, differences between treatment groups in the cumulative number of workers produced by colonies disappeared by midsummer, and all colonies had similar annual honey yields (exception: in one year, productivity was low for colonies supplemented with pollen before wintering). Discrepancies between years coincided with differences in spring weather conditions. Colonies supplemented with pollen or a substitute during the spring performed similarly in all respects. These results indicate that an investment in supplementing the pollen diet of colonies would be returned for situations in which large spring populations are important, but long-term improvement in honey yields may only result when spring foraging is severely reduced by inclement weather. Beekeepers should weigh this information against the nutritional deficiencies that are frequently generated in colonies by the stresses of commercial management.     

Obtaining Specimens with Slowed,Accelerated and Reversed Aging in the Honey Bee Model

Societies of highly social animals feature vast lifespan differences between closely related individuals. Among social insects, the honey bee is the best established model to study how plasticity in lifespan and aging is explained by social factors.The worker caste of honey bees includes nurse bees, which tend the brood, and forager bees, which collect nectar and pollen. Previous work has shown that brain functions and flight performance senesce more rapidly in foragers than in nurses. However, brain functions can recover, when foragers revert back to nursing tasks. Such patterns of accelerated and reversed functional senescence are linked to changed metabolic resource levels, to alterations in protein abundance and to immune function. Vitellogenin, a yolk protein with adapted functions in hormonal control and cellular defense, may serve as a major regulatory element in a network that controls the different aging dynamics in workers.Here we describe how the emergence of nurses and foragers can be monitored, and manipulated, including the reversal from typically short-lived foragers into longer-lived nurses. Our representative results show how individuals with similar chronological age differentiate into foragers and nurse bees under experimental conditions. We exemplify how behavioral reversal from foragers back to nurses can be validated. Last, we show how different cellular senescence can be assessed by measuring the accumulation of lipofuscin, a universal biomarker of senescence.For studying mechanisms that may link social influences and aging plasticity, this protocol provides a standardized tool set to acquire relevant sample material, and to improve data comparability among future studies.     

Honey bee (Apis mellifera) preference towards micronutrients and their impact on bee colonies

Honey bees are important pollinators and take micronutrients from different natural floral resources and turbid water to adequately meet their nutritional requirements. But the role of micronutrients for honey bee health is not well understood. Here, the present study was conducted to determine honey bees' micronutrients preference in summer and winter seasons. Also, the impact of micronutrients on foraging behaviour and brood increase was studied in different honey bee colonies. The results elucidated that honey bees exhibited a strong preference for a salt solution compared to deionized water during the summer and winter seasons. However, there was a notable switch in salt preference between seasons. Overall, honey bees showed significantly more foraging activity, more pollen collection, and increased brood area after sodium consumption compared to other minerals in the summer season. Further, pollen collection and brood area were significantly higher after the use of potassium in the winter season. Thus, the food preference of honey bees is strongly linked with the seasons and the availability of the floral resources. Our data suggested that honey bees may seek specific nutrients during variation of the seasonal conditions.     

Ratios of template responses as the basis of semivision.

The template model starts with a layer of receptors that in the case of vision are leaky detectors or counters of photons. In many animals, the ratio of the responses of a few spectral types is the basis of colour vision irrespective of intensity. Ratios of template responses are now introduced as the basis of form discrimination. In insects, the second-order neurons on the visual pathway appear to detect temporal contrast at the spatial resolution of the retina. At the next level, in the optic medulla, we find a large number of small local neurons in a column on each visual axis. The template theory is a hypothesis about how the above system functions. All possible combinations of positive, indeterminate or negative temporal contrast are considered, at two adjacent visual axes at two successive instants, giving 81 possible local templates. These templates are therefore phasic detectors of all the possible spatiotemporal contrast combinations. Some of the template responses indicate polarity of edge, flicker, or direction of motion and other abstracted features of the stimulus pattern with the maximum spatial and temporal resolution. The ratios of numbers of template responses, in higher fields at a higher level, yield quantitative measures of the qualities of edges independently of the number of edges, but taking ratios causes a corresponding loss of the spatiotemporal resolution and the pattern within each field. Templates respond to transients without computation, are readily modified or selected in evolution and can be simulated in artificial vision.     

Exploring the structure of opioid receptors with homology modeling based on single and multiple templates and subsequent docking: A comparative study

Opioid receptors are the principal targets for opioids, which have been used as analgesics for centuries. Opioid receptors belong to the rhodopsin family of G-protein coupled receptors (GPCRs). In the absence of crystal structures of opioid receptors, 3D homology models have been reported with bovine rhodopsin as a template, though the sequence homology is low. Recently, it has been reported that use of multiple templates results in a better model for a target having low sequence identity with a single template. With the objective of carrying out a comparative study on the structural quality of the 3D models based on single and multiple templates, the homology models for opioid receptors (mu, delta and kappa) were generated using bovine rhodopsin as single template and the recently deposited crystal structures of squid rhodopsin, turkey β-1 and human β-2 adrenoreceptors along with bovine rhodopsin as multiple templates. In this paper we report the results of comparison between the refined 3D models based on multiple sequence alignment (MSA) and models built with bovine rhodopsin as template, using validation programs PROCHECK, PROSA, Verify 3D, Molprobity and docking studies. The results indicate that homology models of mu and kappa with multiple templates are better than those built with only bovine rhodopsin as template, whereas, in many aspects, the homology model of delta opioid receptor with single template is better with respect to the model based on multiple templates. Three nonselective ligands were docked to both the models of mu, delta and kappa opioid receptors using GOLD 3.1. The results of docking complied well with the pharamacophore, reported for nonspecific opioid ligands. The comparison of docking results for models with multiple templates and those with single template have been discussed in detail. Three selective ligands for each receptor were also docked. As the crystallographic structures are not yet known, this comparison will help in choosing better homology models of opioid receptors for studying ligand receptor interactions to design new potent opioid antagonists.     

Effect of primer pheromones and pollen diet on the food producing glands of worker honey bees (Apis mellifera L.)

Cooperative brood care is highly developed in the honey bee such that workers called nurses use their hypopharyngeal and mandibular glands to biosynthesize proteinaceous secretions that are progressively provisioned to larvae. The role that honey bee primer pheromones play in the functional physiology of food producing glands was examined. The combined and separate effects of queen mandibular pheromone (QMP) and brood pheromone (BP) on amount of protein extractable from hypopharyngeal and mandibular glands of workers reared for 12 days with and without pollen diets was measured. In rearing environments with a pollen diet, BP, and QMP + BP pheromone treatments significantly increased extractable protein from both glands. Bees reared with QMP + pollen had amounts of protein extractable from both glands that were not significantly different from control bees (no pheromones, no pollen). Pollen in the diet alone significantly increased amounts of protein extractable from glands versus control. In rearing environments without pollen, QMP + BP had a synergizing effect on amount of protein in both glands. The QMP + BP treatment was the only rearing environment without a pollen diet where protein amounts were significantly greater than the control. The synergizing effect of QMP + BP on extractable mandibular and hypopharyngeal gland protein suggests a highly derived role for the combined effect of these two primer pheromones on honey bee cooperative brood care. Mandibular gland area was significantly and positively correlated with extractable protein. Amounts of extractable protein from both glands declined significantly with age of workers in all treatments. However, treatment significantly affected rate of decline. The adaptive significance of gland protein amounts in response to pheromones and pollen diet are discussed.     

Brood pheromone regulates foraging activity of honey bees (Hymenoptera: Apidae)

Brood pheromone modulated the foraging behavior of commercial honey bee, Apis mellifera L., colonies pollinating a 10-ha market garden of cucumber, Cucurbita pepo L., and zucchini, Cucumis saticus L., in Texas in late autumn. Six colonies were randomly selected to receive 2000 larval equivalents of brood pheromone and six received a blank control. The ratio of pollen to nonpollen foragers entering colonies was significantly greater in pheromone-treated colonies 1 h after treatment. Pheromone-treated foragers returned with pollen load weights that were significantly heavier than controls. Pollen returned by pheromone-treated foragers was 43% more likely to originate from the target crop. Number of pollen grains washed from the bodies of nonpollen foragers from pheromone-treated colonies was significantly greater than controls and the pollen was 54% more likely to originate from the target crop. Increasing the foraging stimulus environment with brood pheromone increased colony-level foraging and individual forager efforts. Brood pheromone is a promising technology for increasing the pollination activity and efficiency of commercial honey bee colonies.     

European bee-eaters (<Emphasis Type="Italic">Merops apiaster</Emphasis>) and apiculture: understanding their interactions and the usefulness of nonlethal techniques to prevent damage at apiaries

Finding a balance between wildlife conservation and land use protection when conflicts arise requires applying measures based on scientific knowledge and applicability. This is because of the complexity of interactions and the social dimensions of human–wildlife relationships. One of the conflicts in the Mediterranean basin is between European bee-eaters Merops apiaster and beekeeping. The objective of this study was to evaluate the activity of bee-eaters in relation to honey bees Apis mellifera in Extremadura (W Spain), and the possible correlation with traits and viability of beehives. We also aimed to assess the effectiveness of damage prevention measures, to be able to predict whether they could assist in the future sustainability of apicultural management. A monitoring program was conducted in 58 apiaries to observe bee and bee-eater activity and abundance. Additionally, we measured the effect of environmental variables and the amount of resources in the hives, by means of honey, pollen, and brood. We found that bee-eater negatively correlated to honey bee activity and the more abundant it was the fewer resources (honey, pollen, and brood) and lower abundance of bees in the hives, mainly during their migratory season (August). However, there were no negative effects on the survival and viability of the hives. The preventive and deterrent measures offered promising results, especially the installation of shading meshes around the apiary that prevented the predation of bees in the immediate surroundings of the beehives. Further studies on the effectiveness of preventive techniques and their practical implementation through different mechanisms of economic compensation must be enhanced to reduce this social conflict.     

Regular pattern formation in real ecosystems

Localized ecological interactions can generate striking large-scale spatial patterns in ecosystems through spatial self-organization. Possible mechanisms include oscillating consumer-resource interactions, localized disturbance-recovery processes and scale-dependent feedback. Despite abundant theoretical literature, studies revealing spatial self-organization in real ecosystems are limited. Recently, however, many examples of regular pattern formation have been discovered, supporting the importance of scale-dependent feedback. Here, we review these studies, showing regular pattern formation to be a general phenomenon rather than a peculiarity. We provide a conceptual framework explaining how scale-dependent feedback determines regular pattern formation in ecosystems. More empirical studies are needed to better understand regular pattern formation in ecosystems, and how this affects the response of ecosystems to global environmental change.     

Honey bee (Apis mellifera jemenitica) colony performance and queen fecundity in response to different nutritional practices

Honey bee colony nutritional dynamics depend on the availability of floral resources throughout a countryside with varying forage circumstances. Few studies quantify the queen fecundity and colony performance about certain management approaches on a broad scale. The present study was conducted to investigate the queen bee fecundity and various colony performance parameters in response to different nutritional practices, i.e., Group-I, supplied with sucrose solution (1:1; w/v), Group-II, provided with locally available commercial pollen substitute, Group-III, supplied with both sucrose solution + locally available commercial pollen substitute, and Group-IV without any sugar solution and pollen substitute. Our results demonstrated that eggs laid by queen bees were significantly higher (1241.83 ± 46.24) in Group-III than in other groups over the time of observations. Similarly, a significant difference was noticed in the mean sealed worker brood area and honey store area between the different groups of management practices. Both, the max mean sealed worker brood area (2153.53 ± 29.18 cm²) and max mean honey store area (1713.33 ± 12.06 cm²) were observed in Group-III while, the mini mean sealed worker brood area (1066.53 ± 20.18 cm²) and mini mean honey store area (1058.86 ± 4.07 cm²) were observed in Group-IV. In contrast, a non-significant difference was observed in pollen stores between Group-II and Group-III (p > 0.005). Current findings add to our understanding of the mechanisms that underpin large-scale controlled colony performance when the natural pollens resources are insufficient.