Caste Differences in Polybia emaciata, a Mud-Nesting Eusocial Wasp (Hymenoptera: Vespidae), that Bear on Pre-Imaginal Determination but not Size Differences

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Caste differences in Polybia ematiata, a mud-nesting eusocial wasp (hymenoptera: Vespidae), that bear on pre-imaginal determination but not size differences

The Epiponini is a group of social wasps living in polygynic societies. The caste dimorphism varies from incipient to well distinct. Morphometric analyses on twelve body parts and ovarian development of Polybia emaciata Lucas 1879, were used to estimate the level of caste differentiation in seven colonies. Physiologic differences were found. Among queens the ovaries were inseminated and presented as highly developed; in intermediate females, ovaries were slightly developed and uninseminated, and for the workers the ovaries were not developed or inseminated. In addition, the coloration of the mesosoma and metasoma were darker in workers and intermediates. Morphometric analysis evidenced slight morphological differences. Even though body proportions were not enough to discriminate castes, the fifth sternite of the gaster had important characters that were able to differentiate queens from workers and intermediates. We propose that P. emaciata presents a case of pre-imaginal caste determination, which is not strictly based on size.

Caste Asymmetries in the Neotropical Swarm-Founding Wasp Polybia (Trychothorax) ignobilis (Hymenoptera, Vespidae, Epiponini)

Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)

Morphological caste differences in three species of the Neotropical genus Clypearia (Hymenoptera: Polistinae: Epiponini)

Clypearia is a rare genus of swarm-founding Neotropical wasp whose biology is very little known. Morphological castes differences, condition of ovaries, relative age, and color pattern differences were analyzed in three species of Clypearia. Physiological differences and low morphometric differentiation between queens and workers were present in all species studied, indicating that these species are characterized by physiological caste only. We suggest that caste determination in the three Clypearia species studied is postimaginal. © 2010 Giovanna Tocchini Felippotti et al.

Development and evolution of caste dimorphism in honeybees - a modeling approach

The difference in phenotypes of queens and workers is a hallmark of the highly eusocial insects. The caste dimorphism is often described as a switch-controlled polyphenism, in which environmental conditions decide an individual's caste. Using theoretical modeling and empirical data from honeybees, we show that there is no discrete larval developmental switch. Instead, a combination of larval developmental plasticity and nurse worker feeding behavior make up a colony-level social and physiological system that regulates development and produces the caste dimorphism. Discrete queen and worker phenotypes are the result of discrete feeding regimes imposed by nurses, whereas a range of experimental feeding regimes produces a continuous range of phenotypes. Worker ovariole numbers are reduced through feeding-regime-mediated reduction in juvenile hormone titers, involving reduced sugar in the larval food. Based on the mechanisms identified in our analysis, we propose a scenario of the evolutionary history of honeybee development and feeding regimes.

Causes and consequences of inbreeding in the ant Formica exsecta

Human actions cause destruction and fragmentation of natural habitats, predisposing populations to loss of genetic diversity and inbreeding, which may further decrease their fitness and survival. Understanding these processes is a main concern in conservation genetics. Yet data from natural populations is scarce, particularly on invertebrates, owing to difficulties in measuring both fitness and inbreeding in the wild. Ants are social insects, and a prime example of an ecologically important group for which the effects of inbreeding remain largely unstudied. Social insects serve key roles in all terrestrial ecosystems, and the division of labor between the females in the colonies queens reproduce, workers tend to the developing brood probably is central to their ecological success. Sociality also has important implications for the effects of inbreeding. Despite their relative abundance, the effective population sizes of social insects tend to be small, owing to the low numbers of reproductive individuals relative to the numbers of sterile workers. This may subject social insects to loss of genetic diversity and subsequent inbreeding depression. Moreover, both the workers and queens can be inbred, with different and possibly multiplicative consequences. The aim of this study was to investigate causes and consequences of inbreeding in a natural population of ants. I used a combination of long-term field and genetic data from colonies of the narrow-headed ant Formica exsecta to examine dispersal, mating behavior and the occurrence of inbreeding, and its consequences on individual and colony traits. Mating in this species takes place in nuptial flights that have been assumed to be population-wide and panmictic. My results, however, show that dispersal is local, with queens establishing new colonies as close as 60 meters from their natal colony. Even though actual sib-mating was rare, individuals from different but related colonies pair, which causes the population to be inbred. Furthermore, multiple mates of queens were related to each other, which also indicates localized mating flights. Hence, known mechanisms of inbreeding avoidance, dispersal and multiple mating, were not effective in this population, as neither reduced inbreeding level of the future colony. Inbreeding had negative consequences both at the individual and colony level. A queen that has mated with a related male produces inbred workers, which impairs the colony s reproductive success. The inbred colonies were less productive and, specifically, produced fewer new queens, possibly owing to effects of inbreeding on the caste determination of female larvae. A striking finding was that males raised in colonies with inbred workers were smaller, which reflects an effect of the social environment as males, being haploid, cannot be inbred themselves. The queens produced in the inbred colonies, in contrast, were not smaller, but their immune response was up-regulated. Inbreeding had no effect on queen dispersal, but inbred queens had a lower probability of successfully founding a new colony. Ultimately, queens that survived through the colony founding phase had a shorter lifespan. This supports the idea that inbreeding imposes a genetic stress, leading to inbreeding depression on both the queen and the colony level. My results show that inbreeding can have profound consequences on insects in the wild, and that in social species the effects of inbreeding may be multiplicative and mediated through the diversity of the social environment, as well as the genetic makeup of the individuals themselves. This emphasizes the need to take into account all levels of organization when assessing the effects of genetic diversity in social animals.

Ant genomics sheds light on the molecular regulation of social organization.

Ants are powerful model systems for the study of cooperation and sociality. In this review, we discuss how recent advances in ant genomics have contributed to our understanding of the evolution and organization of insect societies at the molecular level.

Effect of larval food amount on ovariole development in queens of Trigona spinipes (Hymenoptera, Apinae)

Caste determination in Trigona spinipes Fabricius (Hymenoptera, Apidae, Meliponini) is trophogenic. Larvae that eat about 360 mu l of food become queens, while those who consume 36 mu l develop into workers. We studied the effect of larval nutrition on the number and length of ovarioles and on ovarian development in fifth instar larvae, white eyed, pink eyed and black-eyed pupae as well as newly emerged adults. All larvae have four ovarioles per ovary, while in queen pupae this number ranged from 8 to 15. Cyst formation, the cell death and other characteristics of ovary morphogenesis were the same regardless of the quantity of food consumed. These results are discussed in relation to caste differentiation in other bees.

Nutritional and social hierarchy establishment of the primitively eusocial wasp Mischocyttarus cassununga (Hymenoptera, Vespidae, Mischocyttarini) and related aspects

The results on the nutritional influence in the social hierarchy establishment among females from six colonies of Mischocyttarus cassununga show that: - there was no caste determination in the pre-imaginal period concerning ingested food quantities, however, in relation to qualification (proteins and carbohydrate) the Mann-Whitney test applied showed variation among colonies: in two colonics (colony 3, p=0.0209; colony 4, p=0.0090) the larvae which ingested a higher quantity of solid food became adults of hierarchically higher position, however in two other colonies, colony 1 (p=0.6744) and colony 2 (p=0.3367) this difference was not statistically significant; in two experimentally manipulated colonies the caste determination was observed in the imaginal phase (it was verified that the larger the food quantity consumed, the higher the assumed hierarchical position by the individuals); - adult individuals of hierarchically higher position in two colonies ingested a greater amount of solid than liquid food (Mann-Whitney; p ≤ 0.05), however in the other three colonies there was no difference. It can be seen that for all colonies (2, 3, 4, 5 and 6) the high tendency to solid food ingestion occurs. In colony 1, this data was not collected because the observation purpose was different); - inside the colony hierarchy, the older individuals were the ones that occupied higher hierarchical positions; - more than one inseminated female per colony was found in the two experimentally manipulated colonies ; - ovarian development is greater in inseminated females; - the average height of the 3rd tergite was greater for inseminated females, indicating an adaptation to accommodate larger ovary development (t-Student test; p ≤ 0.05).

The behavior and natural history of Chartergellus, a little-known genus of neotropical social wasps (Vespidae Polistinae Epiponini)

This study reviews published data on the behavior and natural history of Chartergellus and presents the first observations on social interactions in this genus of tropical swarm-founding wasps. Observations of Chartergellus golfitensis in Costa Rica and C. punctatior in Colombia showed that queens perform a post-oviposition egg-guarding vigil, and a bending display like that characteristic of epiponine social wasps that lack consistent morphological differences between workers and queens and have caste determination in the adult stage. Young, old, and queen (egg-laying) females of C. golfitensis showed small differences that indicate color changes with age, and structural differences that could be due to seasonal or colony-cycle changes in developmental conditions, but do not rule out the possibility of pre-adult caste determination, a phenomenon that needs to be carefully distinguished from pre-adult caste bias. Sexual dimorphism and the behavior of males at the nest in C. golfitensis is described, as well as the aggressive and avoidance behavior of females toward males. Nest structure in both species is as described previously for Chartergellus species, but some anomalies and their possible evolutionary significance are discussed. Cell initiation by an egg-laying queen, a behavior never seen by workers, and by a young female with slightly developed ovaries, may be vestiges of ancestral solitary reproductive traits where developed ovaries are associated with cell construction. © 2010 Dipartimento di Biologia Evoluzionistica dell'Università, Firenze, Italia.

Techniques for the In Vitro Production of Queens in Stingless Bees (Apidae, Meliponini)

Considering the ecological importance of stingless bees as caretakers and pollinators of a variety of native plants makes it necessary to improve techniques which increase of colonies' number in order to preserve these species and the biodiversity associated with them. Thus, our aim was to develop a methodology of in vitro production of stingless bee queens by offering a large quantity of food to the larvae. Our methodology consisted of determining the amount of larval food needed for the development of the queens, collecting and storing the larval food, and feeding the food to the larvae in acrylic plates. We found that the total average amount of larval food in a worker bee cell of E varia is approximately 26.70 +/- 3.55 mu L. We observed that after the consumption of extra amounts of food (25, 30, 35 and 40 mu L) the larvae differentiate into queens (n = 98). Therefore, the average total volume of food needed for the differentiation of a young larva of F. varia queen is approximately 61.70 +/- 5.00 mu L. In other words; the larvae destined to become queens eat 2.31 times more food than the ones destined to become workers. We used the species Frieseomelitta varia as a model, however the methodology can be reproduced for all species of stingless bees whose mechanism of caste differentiation depends on the amount of food ingested by the larvae. Our results demonstrate the effectiveness of the in vitro technique developed herein, pointing to the possibility of its use as a tool to assist the production of queens on a large scale. This would allow for the artificial splitting of colonies and contribute to conservation efforts in native bees.

Morphological caste differences and non-sterility of workers in Brachygastra augusti (Hymenoptera, Vespidae, Epiponini), a neotropical swarm-founding wasp

Morphological caste differences and ovary conditions were analyzed in four colonies of Brachygastra augusti collected in different stages of the colony cycle. Differences between castes are distinct suggesting pre-imaginal determination. Many intermediates (non-inseminated ovary-developed females) were found. Because these intermediate females were morphologically similar to workers, it is suggested that they are young ovary-developed workers. Because there is a more evident overlap between queens and workers in the colony in pre-emergence of workers stage, and in three colonies in latter stages, especially in worker-production colonies, it is suggested that smaller queens are probably less viable than larger queens as observed in previously studied epiponines.

Molecular determinants of caste differentiation in the highly eusocial honeybee Apis mellifera

Abstract Background In honeybees, differential feeding of female larvae promotes the occurrence of two different phenotypes, a queen and a worker, from identical genotypes, through incremental alterations, which affect general growth, and character state alterations that result in the presence or absence of specific structures. Although previous studies revealed a link between incremental alterations and differential expression of physiometabolic genes, the molecular changes accompanying character state alterations remain unknown. Results By using cDNA microarray analyses of >6,000 Apis mellifera ESTs, we found 240 differentially expressed genes (DEGs) between developing queens and workers. Many genes recorded as up-regulated in prospective workers appear to be unique to A. mellifera, suggesting that the workers' developmental pathway involves the participation of novel genes. Workers up-regulate more developmental genes than queens, whereas queens up-regulate a greater proportion of physiometabolic genes, including genes coding for metabolic enzymes and genes whose products are known to regulate the rate of mass-transforming processes and the general growth of the organism (e.g., tor). Many DEGs are likely to be involved in processes favoring the development of caste-biased structures, like brain, legs and ovaries, as well as genes that code for cytoskeleton constituents. Treatment of developing worker larvae with juvenile hormone (JH) revealed 52 JH responsive genes, specifically during the critical period of caste development. Using Gibbs sampling and Expectation Maximization algorithms, we discovered eight overrepresented cis-elements from four gene groups. Graph theory and complex networks concepts were adopted to attain powerful graphical representations of the interrelation between cis-elements and genes and objectively quantify the degree of relationship between these entities. Conclusion We suggest that clusters of functionally related DEGs are co-regulated during caste development in honeybees. This network of interactions is activated by nutrition-driven stimuli in early larval stages. Our data are consistent with the hypothesis that JH is a key component of the developmental determination of queen-like characters. Finally, we propose a conceptual model of caste differentiation in A. mellifera based on gene-regulatory networks.

Determination of preventive maintenance lead time using hybrid analysis

The time for conducting Preventive Maintenance (PM) on an asset is often determined using a predefined alarm limit based on trends of a hazard function. In this paper, the authors propose using both hazard and reliability functions to improve the accuracy of the prediction particularly when the failure characteristic of the asset whole life is modelled using different failure distributions for the different stages of the life of the asset. The proposed method is validated using simulations and case studies.