Behavioral and life-history components of division of labor in honey bees (Apis mellifera L.)

Variability exists among worker honey bees for components of division of labor. These components are of two types, those that affect foraging behavior and those that affect life-history characteristics of workers. Variable foraging behavior components are: the probability that foraging workers collect (1) pollen only; (2) nectar only; and (3) pollen and nectar on the same trip. Life history components are: (1) the age the workers initiate foraging behavior; (2) the length of the foraging life of a worker; and (3) worker length of life. We show how these components may interact to change the social organization of honey bee colonies and the lifetime foraging productivity of individual workers. Selection acting on foraging behavior components may result in changes in the proportion of workers collecting pollen and nectar. Selection acting on life-history components may affect the size of the foraging population and the distribution of workers between within nest and foraging activities. We suggest that these components define possible sociogenic pathways through which colony-level natural selection can change social organization. These pathways may be analogous to developmental pathways in the morphogenesis of individual organisms because small changes in behavioral or life history components of individual workers may lead to major changes in the organizational structure of colonies. Correspondence to: R.E. Page, Jr.     

On being the right size: male contributions and multiple mating in social Hymenoptera

Summary A number of hypotheses for the occurrence of multiple mating by queens of social Hymenoptera are reviewed in the light of Cole's (1983) observation that polyandrous species tend to have larger colonies than single-mating ones. Most of these hypotheses cannot be definitively excluded, but only three of them appear sufficiently general, plausible and predictive to be useful guides to further research. These, and their predictions, are: (1) Caste-determination has a genetic basis and hence polyandry allows fuller expression of the potential caste system in each colony. Species with more complex caste differentiation should be more often polyandrous than species with simpler caste systems. (2) Polyandry maximises the production of divergent worker genotypes and hence the range of environmental conditions that a colony can tolerate. Broader-niched species should be more often polyandrous than species with narrower niches. (3) The reduction of the variance of diploid male production, under the heterozygosity sex-determination model, favors polyandry when sexuals are produced late during colony growth. Queens in species reproducing during the exponential phase of colony growth should tend to mate once, but queens should tend to be polyandrous in species with reproduction occurring further along the colony growth curve. Williams's (1975) observation that the level of genetic variation in a brood approaches a maximum very quickly with increasing polyandry is quantified for females; the initial increase is much greater for male-haploid than for male-diploid species.     

Visions for the future of transport: Approaching consensus?

Many cities and national authorities have made substantial progress in developing transport strategies to reconcile environmental pressures with the increasing demand for travel. However, there is still some doubt as to what are the key ingredients of a successful strategy and progress in implementing such changes has been slow. Research is needed to better understand what are the essential ingredients to a balanced transport policy, the constraints, which are preventing these from being implemented, and the acceptability of such changes. This paper reports on the outcomes of a project funded under the Rees Jeffreys Road Fund research programme 'From Realism to Reality', which aims to identify the elements necessary for a successful transport policy in the UK into the 21st century, The project seeks to develop a vision for the transport system of the future through an understanding of the extent of the current problems in the transport sector and an examination of the range and effectiveness of options and instruments available to resolve them. The paper examines a range of visions that are currently in the public domain. Results show that there is a degree of consistency in the general aims and objectives of such visions, but that there is rather less consensus in the means proposed to achieve those aims.     

Influences of a calcium gradient on soil inorganic nitrogen in the Adirondack Mountains, New York

Studies of the long-term impacts of acidic deposition in Europe and North America have prompted growing interest in understanding the dynamics linking the nitrogen (N) and calcium (Ca) cycles in forested watersheds. While it has been shown that increasing concentrations of nitrate (NO3-) through atmospheric deposition or through nitrification can increase Ca loss, the reciprocal effects of Ca on N transformation processes have received less attention. We studied the influence of soil Ca availability on extractable inorganic N (NO3- + NH4+) across a Ca gradient in the Adirondack Mountains, New York, USA. Our results did not show the direct Ca-N interaction that we had expected, but instead showed that exchangeable Ca coupled with soil moisture, soil organic matter, and ambient temperature accounted for 61% of the variability in extractable inorganic N across 11 sites over two growing seasons. Soil Ca concentrations were, however, positively related to sugar maple (Acer saccharum) and American basswood (Tilia americana) basal areas and negatively related to American beech (Fagus grandifolia) basal area. Based on litter chemistry differences among these tree species and reported potential N mineralization values, we suggest that the influence of Ca on soil inorganic N is through a multistep pathway: reciprocal interactions between soil Ca concentrations and species composition, which in turn affect the quality of litter available for N mineralization. If chronic soil Ca depletion continues, as reported in some forested ecosystems, potential shifts in biotic communities could result in considerable alterations of N cycling processes.     

Hydrodynamic characterization of Corpus Christi Bay through modeling and observation

Christi Bay is a relatively flat, shallow, wind-driven system with an average depth of 3–4 m and a mean tidal range of 0.3 m. It is completely mixed most of the time, and as a result, depth-averaged models have, historically, been applied for hydrodynamic characterization supporting regulatory decisions on Texas coastal management. The bay is highly stratified during transitory periods of the summer with low wind conditions. This has important implications on sediment transport, nutrient cycling, and water quality-related issues, including hypoxia which is a key water quality concern for the bay. Detailed hydrodynamic characterization of the bay during the summer months included analysis of simulation results of 2-D hydrodynamic model and high-frequency (HF) in situ observations. The HF radar system resolved surface currents, whereas an acoustic Doppler current profiler (ADCP) measured current at different depths of the water column. The developed model successfully captured water surface elevation variation at the mouth of the bay (i.e., onshore boundary of the Gulf of Mexico) and at times within the bay. However, large discrepancies exist between model-computed depth-averaged water currents and observed surface currents. These discrepancies suggested the presence of a vertical gradient in the current structure which was further substantiated by the observed bi-directional current movement within the water column. In addition, observed vertical density gradients proved that the water column was stratified. Under this condition, the bottom layer became hypoxic due to inadequate mixing with the aerated surface water. Understanding the disparities between observations and model predictions provides critical insights about hydrodynamics and physical processes controlling water quality.     

Division of labor during honey bee colony defense

Summary Some worker honey bees respond to major disturbances of the colony by flying around the assailant and possibly stinging; they are a subset of the bees involved in colony defense. These defenders have an open-ended age distribution similar to that of foragers, but defensive behavior is initiated at a younger age than foraging is. Behavioral and genetic evidence shows that defenders and foragers are distinct groups of older workers. Behaviorally, defenders have less worn wings than foragers, suggesting less flight activity. Genetically, defenders differ in allozyme frequencies, demonstrating different subfamily composition from foragers in the same colony. They also differ in allozyme frequencies from guards in the same colony, providing further evidence for division of labor associated with colony defense. We use this information to develop a model for honey bee colony defense involving at least two distinct groups of workers and we propose that the non-guard defenders be called soldiers, due to their important role in colony defense.Offprint requests to: M.D. Breed     

Genotypic differences in brood rearing in honey bee colonies: context-specific?

Summary Three experiments were performed to determine whether brood care in honey bee colonies is influenced by colony genetic structure and by social context. In experiment 1, there were significant genotypic biases in the relative likelihood of rearing queens or workers, based on observations of individually labeled workers of known age belonging to two visually distinguishable subfamilies. In experiment 2, no genotypic biases in the relative likelihood of rearing drones or workers was detected, in the same colonies that were used in experiment 1. In experiment 3, there again were significant genotypic differences in the likelihood of rearing queens or workers, based on electrophoretic analyses of workers from a set of colonies with allozyme subfamily markers. There also was an overall significant trend for colonies to show greater subfamily differences in queen rearing when the queens were sisters (half- and super-sisters) rather than unrelated, but these differences were not consistent from trial to trial for some colonies. Results of experiments 1 and 3 demonstrate genotypic differences in queen rearing, which has been reported previously based on more limited behavioral observations. Results from all three experiments suggest that genotypic differences in brood care are influenced by social context and may be more pronounced when workers have a theoretical opportunity to practice nepotism. Finally, we failed to detect persistent interindividual differences in bees from either subfamily in the tendency to rear queen brood, using two different statistical tests. This indicates that the probability of queen rearing was influenced by genotypic differences but not by the effect of prior queen-rearing experience. These results suggest that subfamilies within a colony can specialize on a particular task, such as queen rearing, without individual workers performing that task for extended periods of time.