Optimal colony fissioning in social insects: testing an inclusive fitness model with honey bees

In most species of social insects, when a queen departs from her parental nest to found a new colony, she leaves on her own. In some species, however, the departing queen leaves accompanied by a portion of the parental colony’s workers and there is a permanent fissioning of the worker force. Little is known about how the adult workers in colonies of fissioning species distribute themselves between the old and the new colonies. We examined this problem, building on Bulmer’s (J Theor Biol 100: 329–339, 1983) model for the optimal splitting of a colony’s adult workforce during colony reproduction. We first created an inclusive fitness model of optimal colony fissioning that applies to species in which fissioning gives rise to two autonomous colonies. The model predicts the optimal “swarm fraction”, which we define as the proportion of the adult workers in a fissioning colony that join the departing queen. We then tested the model by comparing the predicted and observed swarm fractions in honey bees. We found a close match between predicted (0.76–0.77) and observed (0.72 ± 0.04) swarm fractions. Evidently, worker honey bees distribute themselves between the old and new colonies in a way that jointly maximizes the inclusive fitness of each worker. We conclude by discussing additional ways to test the model.     

Colony size evolution and the origin of eusociality in corbiculate bees (Hymenoptera: Apinae)

Recently, it has been proposed that the one of the main determinants of complex societies in Hymenoptera is colony size, since the existence of large colonies reduces the direct reproductive success of an average individual, given a decreased chance of being part of the reproductive caste. In this study, we evaluate colony size evolution in corbiculate bees and their relationship with the sociality level shown by these bees. Specifically i) the correlation between colony size and level of sociality considering the phylogenetic relationship to evaluate a general evolutionary tendency, and ii) the hypothetical ancestral forms of several clades within a phylogeny of corbiculate bees, to address idiosyncratic process occurring at important nodes. We found that the level of social complexity in corbiculate bees is phylogenetically correlated with colony size. Additionally, another process is invoked to propose why colony size evolved concurrently with the level of social complexity. The study of this trait improves the understanding of the evolutionary transition from simple to complex societies, and highlights the importance of explicit probabilistic models to test the evolution of other important characters involved in the origin of eusociality.     

Colony founding and initial nest design of honey bees, Apis mellifera L

Thirty-two colonies of bees with varying populations were established in 1·2-m³ boxes for 24 days. At the end of this time, the bees were killed, and the comb structure, design, and placement were examined to obtain information about the initiation of bee nests. The analysis of variance of three dimensions of the comb (height, width, and length) showed no statistical differences when all thirty-two colonies were considered as a group.Comb built by queenless bees differed from any described in the literature. We think it is comb characteristically built by queenless bees that is neither worker nor drone in size but intermediate.The occurrence of multiple clusters building comb with and without queen in the same box is recorded but unexplained at present.An explanation is offered for the curl appearing in the lower edges of straight combs.     

European foulbrood in honey bees

European foulbrood (EFB) is a severe bacterial brood disease caused by the Gram-positive bacterium Melissocccus plutonius. The disease has a worldwide distribution and is an increasing problem in some areas. Although the causative agent of EFB was described almost a century ago, many basic aspects of its pathogenesis are still unknown. This review presents both historical results and recent molecular data to synthesize present knowledge of this enigmatic honey bee disease.     

Benefits of recruitment in honey bees: effects of ecology and colony size in an individual-based model

Why do some social insects have sophisticated recruitment systems,while other species do not communicate about food source locationsat all? To answer this question, it is necessary to identifythe social or ecological factors that make recruitment adaptiveand thus likely to evolve. We developed an individual-basedmodel of honey bee foraging to quantify the benefits of recruitmentunder different spatial distributions of nondepleting resourcepatches and with different colony sizes. Benefits of recruitmentwere strongly dependent on resource patch quality, density,and variability. Communication was especially beneficial ifpatches were poor, few, and variable. A sensitivity analysisof the model showed that under conditions of high resource densityrecruitment could even become detrimental, especially if foragingduration was short, tendency to scout was high, or recruitsneeded a long time to find communicated locations. Colony size,a factor often suspected to influence recruitment evolution,had no significant effect. These results may explain the recentexperimental findings that in honey bees, benefits of waggledance recruitment seem to vary seasonally and with habitat.They may also explain why some, but not other, species of socialbees have evolved a strategy to communicate food locations tonest mates.     

Taste perception in honey bees

Taste is crucial for honeybees for choosing profitable food sources, resins, water sources, and for nestmate recognition. Peripheral taste detection occurs within cuticular hairs, the chaetic and basiconic sensilla, which host gustatory receptor cells and, usually a mechanoreceptor cell. Gustatory sensilla are mostly located on the distal segment of the antennae, on the mouthparts, and on the tarsi of the forelegs. These sensilla respond with varying sensitivity to sugars, salts, and possibly amino acids, proteins, and water. So far, no responses of receptor cells to bitter substances were found although inhibitory effects of these substances on sucrose receptor cells could be recorded. When bees are free to express avoidance behaviors, they reject highly concentrated bitter and saline solutions. However, such avoidance disappears when bees are immobilized in the laboratory. In this case, they ingest these solutions, even if they suffer afterward a malaise-like state or even die from such ingestion. Central processing of taste occurs mainly in the subesophageal ganglion, but the nature of this processing remains unknown. We suggest that coding tastants in terms of their hedonic value, thus classifying them in terms of their palatability, is a basic strategy that a central processing of taste should achieve for survival.     

Chalkbrood disease in honey bees

Chalkbrood is a fungal disease of honey bee brood caused by Ascosphaera apis. This disease is now found throughout the world, and there are indications that chalkbrood incidence may be on the rise. In this review we consolidate both historic knowledge and recent scientific findings. We document the worldwide spread of the fungus, which is aided by increased global travel and the migratory nature of many beekeeping operations. We discuss the current taxonomic classification in light of the recent complete reworking of fungal systematics brought on by application of molecular methods. In addition, we discuss epidemiology and pathogenesis of the disease, as well as pathogen biology, morphology and reproduction. New attempts at disease control methods and management tactics are reviewed. We report on research tools developed for identification and monitoring, and also include recent findings on genomic and molecular studies not covered by previous reviews, including sequencing of the A. apis genome and identification of the mating type locus.     

Juvenile hormone and aggression in honey bees

We determined whether defense by individual bees against non-nestmates in honey bees (Apis mellifera) is correlated with their juvenile hormone (JH) titers, which are known to vary developmentally and seasonally. We bioassayed winter and summer bees for aggressive and non-aggressive individuals. Bees in winter could not be distinguished by task group, but bees in summer were segregated into nurses and guards. JH titers were correlated with aggressive behavior at two levels. First, winter bees and summer nurses, known to have lower JH titers, both showed less aggression toward foreign bees than did summer guards. Second, aggressive individuals had significantly higher JH titers than did non-aggressive bees within each colony. Inter-colonial variation in aggressiveness was maintained during summer and winter, suggesting a genetic basis for these differences. An alarm pheromone test further substantiated the existence of inter-colonial differences. We found significant variation in JH titers among different colonies, but this variation was not significantly associated with colony-level aggressiveness. The correlation between JH and levels of aggressiveness within a colony suggests a regulatory role for JH, but variation among colonies involves factors other than JH.     

Mortality rates of honey bees in the wild

Senescence, defined as an age-specific decrease in physiological performance accompanied by an increase in mortality rate, has been studied in a wide range of animals including social insects. It is not clear, however, whether honey bees in the wild live long enough to exhibit senescent decline. I tested for the effects of senescence on honey bees foraging in natural settings and documented the predicted pattern of exponential increase in mortality rate with forager age. These data indicate that, in spite of high rates of external mortality, senescence is an important factor determining the performance of insects such as honey bees in the wild. Received 16 November 2007; revised 1 February 2008 and 10 March 2008; accepted 18 March 2008.     

Virus infections in Brazilian honey bees

This work describes the first molecular-genetic evidence for viruses in Brazilian honey bee samples. Three different bee viruses, Acute bee paralysis virus (ABPV), Black queen cell virus (BQCV), and Deformed wing virus (DWV) were identified during a screening of RNAs from 1920 individual adult bees collected in a region of southeastern Brazil that has recently shown unusual bee declines. ABPV was detected in 27.1% of colony samples, while BQCV and DWV were found in 37% and 20.3%, respectively. These levels are substantially lower than the frequencies found for these viruses in surveys from other parts of the world. We also developed and validated a multiplex RT-PCR assay for the simultaneous detection of ABPV, BQCV, and DWV in Brazil.     

Beneficial microorganisms for honey bees: problems and progresses

Nowadays, honey bees are stressed by a number of biotic and abiotic factors which may compromise to some extent the pollination service and the hive productivity. The EU ban of antibiotics as therapeutic agents against bee pathogens has stimulated the search for natural alternatives. The increasing knowledge on the composition and functions of the bee gut microbiota and the link between a balanced gut microbiota and health status have encouraged the research on the use of gut microorganisms to improve bee health. Somehow, we are assisting to the transfer of the “probiotic concept” into the bee science. In this review, we examine the role of the honey bee gut microbiota in bee health and critically describe the available applications of beneficial microorganisms as pest control agents and health support. Most of the strains, mainly belonging to the genera Lactobacillus, Bifidobacterium and Bacillus, are isolated from honey bee crop or gut, but some applications involve environmental strains or formulation for animal and human consumption. Overall, the obtained results show the favourable effect of applied microbial strains on bee health and productivity, in particular if strains of bee origin are used. However, it is actually not yet possible to conclude whether this strategy will ever work. In particular, many aspects regarding the overall setup of the experiments, the dose, the timing and the duration of the treatment need to be optimized, also considering the microbiological safety of the hive products (i.e. pollen and honey). In addition, a deep investigation about the effect on host immunity and physiology is envisaged. Lastly, the final users of the formulations, i.e. beekeepers, should be taken into account for the achievement of high-quality, cost-effective and easy-to-use products.     

New allozyme variability in Italian honey bees

Adult workers of the honey bee, Apis mellifera ligustica, from Italy were assayed for enzyme polymorphism using a variety of electrophoretic conditions. Three polymorphic enzyme systems are described, two of which, malic enzyme and an esterase, were previously unknown in indigenous A. m. ligustica. In addition, a new allozyme for the Mdh locus is reported.     

Memory and sun compensation by honey bees

Summary Experiments with two species of honey bees (Apis mellifera andA. cerana) have revealed that bees form a detailed memory of the spatial and temporal pattern of the sun's azimuthal movement, using local landmarks as a reference for the learning. These experiments were performed on overcast days, and consisted of removing a hive from one site in which bees had been trained to find food by flying along a prominent landmark, and displacing it to a similar site in which the landmark was aligned in a different compass direction. On overcast days, bees which flew along the landmark in the new site oriented their waggle dances in the hive as if they had actually flown in the training site. Thus, they confused the two sets of landmarks and set their dance angles according to a memory of the sun's position relative to the original landmarks. Furthermore, the dances changed in correspondence with the sun's azimuthal shift over several hours, even reflecting (approximately) the regular temporal variations in the rate of shift; such features of the sun's course must therefore be stored in memory. The primary mechanism underlying the learning of this pattern is probably similar to that proposed by New and New (1962): bees store in memory several time-linked solar azimuthal positions relative to features of the landscape, and refer to this stored array when they need to determine an unknown azimuth intermediate between two known positions.During the cloudy-day displacement experiments, celestial cues often appeared to bees in the new site, contradicting the stored information on which they had been basing their dances. Although most bees quickly adopted the dance angle reflecting their actual direction of flight relative to the sun, some later reverted to the original dance angle, indicating that the information on which it was based had remained in memory when the new information was being expressed; other bees performed bimodal dances which expressed both sets of information in alternate waggle runs. The separation in memory implied by these behaviors may reflect a neural strategy for updating a previously stored relationship between celestial and terrestrial references with new information presented by seasonal changes in the sun's course or by newly learned landmarks.