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.

A morphologically specialized soldier caste improves colony defense in a neotropical eusocial bee

Division of labor among workers is common in insect societies and is thought to be important in their ecological success. In most species, division of labor is based on age (temporal castes), but workers in some ants and termites show morphological specialization for particular tasks (physical castes). Large-headed soldier ants and termites are well-known examples of this specialization. However, until now there has been no equivalent example of physical worker subcastes in social bees or wasps. Here we provide evidence for a physical soldier subcaste in a bee. In the neotropical stingless bee Tetragonisca angustula, nest defense is performed by two groups of guards, one hovering near the nest entrance and the other standing on the wax entrance tube. We show that both types of guards are 30% heavier than foragers and of different shape; foragers have relatively larger heads, whereas guards have larger legs. Low variation within each subcaste results in negligible size overlap between guards and foragers, further indicating that they are distinct physical castes. In addition, workers that remove garbage from the nest are of intermediate size, suggesting that they might represent another unrecognized caste. Guards or soldiers are reared in low but sufficient numbers (1-2% of emerging workers), considering that <1% usually perform this task. When challenged by the obligate robber bee Lestrimelitta limao, an important natural enemy, larger workers were able to fight for longer before being defeated by the much larger robber. This discovery opens up opportunities for the comparative study of physical castes in social insects, including the question of why soldiers appear to be so much rarer in bees than in ants or termites.

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.

Impacts of ocean acidification on multiplication and caste organisation of parasitic trematodes in their gastropod host

Ocean acidification is predicted to impact the structure and function of all marine ecosystems in this century. As focus turns towards possible impacts on interactions among marine organisms, its effects on the biology and transmission potential of marine parasites must be evaluated. In the present study, we investigate two marine trematode species (Philophthalmus sp. and Parorchis sp., both in the family Philophthalmidae) infecting two marine gastropods. These trematodes are unusual in that their asexually multiplying stages within snails display a division of labour, with two distinct castes, a large-bodied morph producing infective stages and a smaller morph playing a defensive role against other competing parasites. Using a potentiometric ocean acidification simulation system, we test the impacts of acidified seawater (7.8 and 7.6 pH) on the production of free-living infective stages (cercariae), the size and survival of encysted resting stages (metacercariae), and the within-host division of labour measured as the ratio between numbers of the two morphs. In general, low pH conditions caused an increase in cercarial production and a reduction in metacercarial survival. The ratio of the two castes within snail hosts tended to shift towards more of the smaller defensive morphs under low pH. However, the observed effects of reduced pH were species specific and not always unimodal. These results suggest that ocean acidification can affect the biology of marine parasites and may also impact transmission success and parasite abundance of some trematodes, with possible consequences for marine communities and ecosystems.

Caste, land, and migration : a preliminary analysis of a village survey in an underdeveloped state in India

This paper explores migration from Bihar, one of the most underdeveloped states in India, by paying particular attention to social class (caste) and landholdings. After describing details of individual migrants, we present our preliminary findings on the determinants of migration, based on our field survey of 200 households in four villages in 2011. In terms of social class, Muslims are more likely to migrate, but Scheduled Castes do not show a high propensity to migrate as is stated in some of the existing literature where the underclass is said to be more mobile. In terms of landholdings, the probability that someone will migrate is high among the landless and smaller landholders but it decreases as the size of the landholding increases. However, as the size of the landholding increases still further, a reverse effect of landholding on decisions regarding migration moves in, with the decline in probability becoming less and less. This result confirms a non-linear relationship between landholdings and the decision to migrate. Some further research questions are raised in the paper.

Soldier caste-specific gene expression in the mandibular glands of Hodotermopsis japonica (Isoptera: Termopsidae)

Although “polymorphic castes” in social insects are well known as one of the most important phenomena of polyphenism, few studies of caste-specific gene expressions have been performed in social insects. To identify genes specifically expressed in the soldier caste of the Japanese damp-wood termite Hodotermopsis japonica, we employed the differential-display method using oligo(dT) and arbitrary primers, compared mRNA from the heads of mature soldiers and pseudergates (worker caste), and identified a clone (PCR product) 329 bp in length termed SOL1. Northern blot analysis showed that the SOL1 mRNA is about 1.0 kb in length and is expressed specifically in mature soldiers, but not in pseudergates, even in the presoldier induction by juvenile hormone analogue, suggesting that the product is specific for terminally differentiated soldiers. By using the method of 5′- and 3′-rapid amplification of cDNA ends, we isolated the full length of SOL1 cDNA, which contained an ORF with a putative signal peptide at the N terminus. The sequence showed no significant homology with any other known protein sequences. In situ hybridization analysis showed that SOL1 is expressed specifically in the mandibular glands. These results strongly suggest that the SOL1 gene encodes a secretory protein specifically synthesized in the mandibular glands of the soldiers. Histological observations revealed that the gland actually develops during the differentiation into the soldier caste.

Caste in a peasant society ; a case study in the dynamics of caste.

Mode of access: Internet.

Sull'esistenza delle antiche caste egiziane : negata da J.-J. Ampère /

Cavagna 6363: Some leaves unopened.

Income mobility and the U.S. economy : open society or caste system?

"January 1992."

Caste and slavery in the American church.

Mode of access: Internet.

Census of India, 1901. Volume I. India. Ethnographic appendices, being the data upon which the caste chapter of the Report is based,

Mode of access: Internet.

Developmental Pathways in Microcerotermes turneri (Termitidae: Termitinae): Biometric Descriptors of Worker Caste and Instar

The foraging process of location and exploitation of food in complex termite societies is in part reliant upon unequal division of specific tasks amongst its members (polyethism). To conduct studies assessing the role of individuals in foraging activities it is necessary to have descriptors of worker caste and instar. Here we provide biometric descriptors of specific caste and instar for worker caste and instars of Microcerotermes turneri (Froggatt) (Termitidae: Termitinae) for the worker castes (male and female) for the identification of individuals in laboratory assays applicable across multiple nests. The use of head width for determining sex of workers was successful across multiple nests. The length of the first three flagellum segments of the antenna and tibia three could be used to determine worker instar.

Caste, class and Trade Unionism in India

The caste system in India and its exploitative nature has been well researched (Siddique 2011 Gupta 2000). However, the role of caste in Indian employment relations and in particular its role in the labor movement in India is yet to be fully explored. The primary aim of this paper is to examine the rise of caste- based trade unions in India over the past decade. Specifically, we aim to examine why the lower-caste workers (historically treated as untouchables, referred to as ‘Dalits’and officially designated as Scheduled Caste and Tribes) are leaving established trade unions to organize their own unions along caste lines? While fragmentation of trade unions is a well-known phenomenon both in India and in the Western World (Shyam Sundar 2015; Connolly et al. 2014), the rise of caste based trade unions is a relatively new phenomenon which is yet to be fully explored. Caste based trade unionism appears to be counter-intuitive when the conventional logic suggests that unions are class based collective institutions which represent the interest of the working classes (Ramaswamy 1976). The Indian trade union movement has historically been fragmented along political ideological lines ranging from moderate unions affiliated to the Congress Party to the militant unions affiliated to the Communist and Socialist parties. However, the rise of caste-based trade unions of the lower caste workers is a relatively new phenomenon. Our findings from surveys and interviews with mainstream unions and caste-based trade unions suggest that the caste-based trade unions are unique in at least three ways. First, these unions are breaking away from well-established radical and militant union federations such as those affiliated to the Communist and Socialist parties. Second, these unions are predominantly organized on caste identities and not occupational identities or political ideologies. Third in unionized workplaces, lower caste workers are forming their own separate unions along caste lines with membership restricted only to workers of their own caste instead of joining the mainstream unions where present. We examine these issues using the analytical framework of Paulo Freire (1970) – dialogics, praxis and cultural oppression and relate it with the insights from comparative politics which examine the role of actors & their interests within institutions (Peters 2011).