Within social insect colonies, sterile workers are responsible for brood care, while queens are the primary egg-layers. These reproductive caste differences are often accompanied by pronounced morphological distinctions. Queen and worker phenotypic differences are particularly remarkable considering caste is environmentally, rather than genetically, determined. Environmental caste determination can produce intralocus genetic conflict between castes, particularly when homologous traits are highly dimorphic. Therefore, when studying the evolution of social insect caste dimorphism, one must consider the genetic architecture underlying phenotypic expression as well as the selective pressures that have shaped caste morphology. This dissertation presents the results of four studies that investigated factors affecting caste morphology in the social wasp Vespula maculifrons. The first two studies focused on identifying queen morphological traits that were positively associated with queen fitness and would therefore be subject to selection. Queen length, specifically gaster length, was positively associated with overwintering survival and was consistently associated with mating success. Both of these findings suggest that queen gaster length is under selection during two life-history events, mating and overwintering, in which workers do not participate. These findings provide empirical support for the adaptive evolution of a caste dimorphic trait. The third and fourth studies used classical quantitative genetic and morphological analyses to examine the genetic architecture underlying caste dimorphism in V. maculifrons. I determined which traits were under caste-specific selection by analyzing trait allometries and the levels of genetic control, variation, and dimorphism of traits between castes. Little genetic variation for morphological trait size was detected for most worker and queen traits, suggesting a strong influence of environment on phenotypic variation. Additionally, analyses of trait allometries indicated that several queen traits (mass, thorax width and length) were under queen-specific selection. The relationship between thorax length, gaster length, and overall body size is further evidence of selection on length in queens. Overall, these studies provide evidence for the importance of queen-specific selection in the evolution of caste dimorphism. When placed in the broader context of caste evolution, they point to the importance of life-history in shaping the genetic architecture underlying caste dimorphism.

Ambedkar’s role as an Indian political leader during the late colonial period has attracted increased attention politically and historically. However, there is a startling disconnect between the modern, often mythological, construction of Ambedkar and the near forgotten historical figure. His broader programme for social uplift of the underprivileged is often lost in the record of his conflict with M. K. Gandhi and the Indian National Congress and their role as the dominant nationalist group in India at the time. The deification that has resulted from his use of Buddhism as an emancipatory identity has obscured his interpretation of it as a secular political tool in a political debate shaped and dominated by religious identity. This thesis will argue that the Buddhist conversion was a continuation of his political and social programme, not, as some have suggested, a retreat to religion after failing to secure reforms to Indian law and society.

Ambedkar’s role as an Indian political leader during the late colonial period has attracted increased attention politically and historically. However, there is a startling disconnect between the modern, often mythological, construction of Ambedkar and the near forgotten historical figure. His broader programme for social uplift of the underprivileged is often lost in the record of his conflict with M. K. Gandhi and the Indian National Congress and their role as the dominant nationalist group in India at the time. The deification that has resulted from his use of Buddhism as an emancipatory identity has obscured his interpretation of it as a secular political tool in a political debate shaped and dominated by religious identity. This thesis will argue that the Buddhist conversion was a continuation of his political and social programme, not, as some have suggested, a retreat to religion after failing to secure reforms to Indian law and society.

abstract: Gene-centric theories of evolution by natural selection have been popularized and remain generally accepted in both scientific and public paradigms. While gene-centrism is certainly parsimonious, its explanations fall short of describing two patterns of evolutionary and social phenomena: the evolution of sex and the evolution of social altruism. I review and analyze current theories on the evolution of sex. I then introduce the conflict presented to gene-centric evolution by social phenomena such as altruism and caste sterility in eusocial insects. I review gene-centric models of inclusive fitness and kin selection proposed by Hamilton and Maynard Smith. Based their assumptions, that relatedness should be equal between sterile workers and reproductives, I present several empirical examples that conflict with their models. Following that, I introduce a unique system of genetic caste determination (GCD) observed in hybrid populations of two sister-species of seed harvester ants, Pogonomyrmex rugosus and Pogonomyrmex barbatus. I review the evidence for GCD in those species, followed by a critique of the current gene-centric models used to explain it. In chapter two I present my own theoretical model that is both simple and extricable in nature to explain the origin, evolution, and maintenance of GCD in Pogonomyrmex. Furthermore, I use that model to fill in the gaps left behind by the contributing authors of the other GCD models. As both populations in my study system formed from inter-specific hybridization, I review modern discussions of heterosis (also called hybrid vigor) and use those to help explain the ecological competitiveness of GCD. I empirically address the inbreeding depression the lineages of GCD must overcome in order to remain ecologically stable, demonstrating that as a result of their unique system of caste determination, GCD lineages have elevated recombination frequencies. I summarize and conclude with an argument for why GCD evolved under selective mechanisms which cannot be considered gene-centric, providing evidence that natural selection can effectively operate on non-heritable genotypes appearing in groups and other social contexts. / Dissertation/Thesis / M.S. Biology 2012

A relação entre nutrição e fenótipo é uma questão especialmente desafiadora em casos de polifenismo facultativo, como no caso das castas de insetos sociais, por exemplo, a abelha melífera Apis mellifera. Após estudos de vias de sensoriamento de nutrientes, modificações inesperadas nestas vias de sinalização revelaram a resposta à hipóxia como um possível mecanismo subjacente à regulação do tamanho corporal e crescimento de órgãos. Uma vez que essa resposta está intimamente ligada a condições metabólicas das células, o presente estudo foi concebido para investigar possíveis alterações no metabolismo mitocondrial e oxidativo no processo de diferenciação de castas em A. melífera na fase larval, com foco no corpo gorduroso, que é o centro metabólico dos insetos. Partindo da hipótese de que rainhas e operárias são criadas em células de cria abertas sob condições iguais de disponibilidade de oxigênio, nós investigamos o número e a distribuição mitocondrial, bem como as taxas de consumo de oxigênio em mitocondrias de células de corpo gorduroso durante estágios larvais críticos. Por meio de análises de imunofluorescência e microscopia eletrônica encontramos maior densidade de mitocôndrias no corpo gorduroso larval da rainha, dado corroborado pela quantificação de unidades funcionais mitocondriais por ensaio de citrato sintase. Medições de consumo de oxigênio por respirometria de alta resolução revelaram que as larvas de rainha têm capacidades máximas mais altas de produção de ATP, com menor demanda fisiológica, mas com mesma eficiência mitocondrial que operárias. A análise da expressão de fatores relacionados à mitogênese mostrou que os homólogos dos genes codificadores dos fatores de transcrição TFB1 e TFB2 e de um regulador nutricional, ERR, estão mais expressos em larvas de rainha. Apesar das diferenças encontradas na respiração mitocondrial entre as duas castas, as mesmas apresentaram níveis similares de produção de lactato e peróxido de hidrogênio, sem grandes alterações referentes à condições de estresse oxidativo e ao estado redox das célula. Encontramos altos níveis de expressão de genes codificadores das enzimas do sistema antioxidante MnSOD e catalase em células do corpo gorduroso larval de rainha, que garantem a redução de eventuais danos oxidativos. Estes resultados são fortes evidências de que a nutrição diferencial das larvas pelas operárias adultas, como estímulo externo da indução do desenvolvimento das castas, diferencialmente afeta a dinâmica e funcionalidade mitocondrial como elemento intrínseca da plasticidade fenotípica neste inseto social. / The connection between nutrition and phenotype is a particularly challenging issue in cases of facultative polyphenism, as for instance in the honeybee Apis mellifera. After studies on nutrient sensing pathways found unexpected modifications in these signaling pathways, a response to hypoxia was revealed as a possible mechanism underlying regulation of body size and organ growth. Since this response is closely linked to the metabolic conditions of the cells, the present study was designed to investigate the role of the mitochondrial and oxidative metabolism in the fat body of honeybee larvae, which is the metabolic center in insects, in the context of the caste differentiation process in A. mellifera. Based on the fact that honey bee larvae are reared in open brood cells, the queen and worker larvae should be exposed to equal oxygen diffusion conditions, and hence we investigated mitochondrial number and intracellular distribution, as well as rates of mitochondrial oxygen consumption in fat body cells during the larval stages critical for caste differentiation. By means of immunofluorescence and electron microscopy we found a higher density of mitochondria in the fat body of queen larvae. This result was corroborated by the quantification of mitochondrial functional units using a citrate synthase assay. Measurements of oxygen consumption obtained by high resolution respirometry revealed that queen larvae have higher maximum capacities of ATP production, with less physiological demand and higher mitochondrial efficiency than workers. Analysis of the expression of genes related to mitogenesis showed that Apis homologs of the transcription factors TFB1 and TFB2 and of the nutritional regulator, ERR are higher expressed in queen larvae. Despite differences in mitochondrial respiration, the two castes presented similar levels of lactate and hydrogen peroxide production, and without major chang in oxidative stress and cellular redox status. The high transcriptional levels of genes encoding enzymes of the antioxidant system, MnSOD and catalase observed in fat body cells of queen larvae guarantee that oxidative damage is reduced during larval development. These results are strong evidence that the differential nutrition of honey bee larvae by the adult worker, as the external stimulus for caste induction, differentially affects mitochondrial dynamics and functionality as an intrinsic element of phenotypic plasticity in this social insect.

Melipona scutellaris (Apidae: Meliponini) é uma espécie de abelha sem ferrão popularmente conhecida como abelha Uruçu. Como outros meliponíneos, forma colônias perenes e apresenta diferenciação de castas. É encontrada na zona litorânea do Sul da Bahia ao Ceará, e em regiões do interior da Bahia e Pernambuco. Os ninhos são construídos somente em troncos ocos de árvores e apresentam arquitetura elaborada. As células de cria são verticais, arranjadas em favos horizontais formando placas que se sobrepõe. Seus principais recursos alimentares são pólen e néctar. A quantidade de alimento estocada nos potes está relacionada com a manutenção e produção de operárias, rainhas virgens e machos. Por não apresentar células de cria diferenciadas para o desenvolvimento de rainhas e operárias, não é possível estabelecer seguramente os fatores responsáveis pela determinação das castas neste grupo de abelhas. Estudos indicam que mecanismos genéticos e tróficos, incluindo a qualidade do alimento larval, somados a fatores ambientais interferem na produção de rainhas. Este estudo avaliou a variação do teor proteico e composição polínica do alimento larval ao longo de um ano. Os resultados mostraram que o valor proteico do alimento larval variou de forma equivalente para todas as colônias em todos os meses, com elevação significativa no mês de julho. O valor proteico do alimento larval não apresentou correlação com o valor proteico dos tipos polínicos. O valor proteico dos tipos polínicos não apresentou relação significativa com sua ocorrência no alimento larval, o que indica hábitos generalistas para a coleta de recursos alimentares. Através de bioensaios, foi testada a interferência do volume e da suplementação proteica do alimento larval na determinação de rainhas, cujos resultados demonstraram uma ocorrência de rainhas significativamente maior nos tratamentos com suplementação proteica. Conclui-se que embora os tipos polínicos não influenciem diretamente o teor proteico do alimento larval, os resultados encontrados sugerem que a alteração do valor proteico do alimento larval depositado nas células seja um fator importante na determinação de castas nesta espécie / Melipona scutellaris (Apidae: Meliponini) is an indigenous stingless bee species. Like other Meliponine, colonies are perennial and have female caste differentiation. This species is found from the cost area of Bahia to Ceará states. The nests are built only in hollow trees and have an elaborate architecture. The brood cells are vertical, arranged in horizontal overlapping combs. Pollen and honey are the main food source. The quantity of food stored in the pots is associated with the maintenance and production of workers, queens and males. Since there are no differences between the cells that queens and workers are reared, the factors responsible for caste determination are still unknown. Studies suggest that both genetic and trophic mechanisms, including the larval food quality, combined with environmental factors, interfere in the production of queens. This study evaluated the variation protein content and the pollinic composition of larval food among colonies of M. scutellaris across a year. The results showed that the protein content of the larval food varied equivalently for all the colonies at all months, but presenting an elevation on the protein content in July. The protein content in the larval food had no correlation with the pollen types. The protein content of the pollen types showed no significant relation with the pollen types occurring in the larval food, indicating that the species M. scutellaris presents generalist habits for food gathering. The interference of volume and protein supplementation of larval food on queen rearing determination was tested via bioassays, which results showed a significantly higher occurrence of queens in the treatments with protein supplementation. We concluded that although the pollen types did not influence directly the protein content of the larval food, the results obtained suggest that the variation of protein content of the larval food deposited in the cells may be an important factor in caste determination in this species

Neste estudo, foi investigado o efeito da dieta larval no desenvolvimento de características morfológicas casta-específicas e na expressão de alguns genes durante o desenvolvimento da abelha sem ferrão Scaptotrigona aff. depilis . Nesta espécie, as castas femininas são determinadas pela quantidade de alimento consumida durante o desenvolvimento larval. Grupos experimentais larvas criadas in vitro foram feitos com duas quantidades diferentes de alimento larval, obtido de favos recém provisionados nesta espécie. As larvas do primeiro grupo receberam 32l de alimento larval, correspondente a quantidade média de alimento recebida por larvas de operárias naturalmente, já as larvas do segundo grupo receberam 130l, correspondente a quantidade média encontrada naturalmente em células reais. Todas as larvas criadas com 130l de alimento larval se desenvolveram em rainhas, como esperado; da mesma forma, a maioria das larvas criadas com 32l se desenvolveram em operárias. Interessantemente, porém, algumas larvas deste grupo se desenvolveram em rainhas miniaturas, sugerindo que outros fatores, além do trófico, estejam envolvidos na determinação de castas em S. aff. depilis. Subsequentemente, analisamos os títulos de hormônio juvenil (HJ) na hemolinfa por radioimunoensaio durante quatro estágios do último instar larval e encontramos que as larvas do grupo de 130l de dieta apresentaram maiores títulos no estágio defecante (LD) (p=0,034, t-test), se comparado com as larvas do grupo de 32l. Os níveis de expressão de determinados genes que foram previamente descritos em outras espécies como expressos preferencialmente em rainhas. Um destes genes, dnmt-3, codifica a DNA metiltransferase envolvida na metilação do DNA. Para este gene, encontramos maiores níveis de transcritos nas larvas alimentadas como rainhas (130l) nos estágios LPD e LD (p=0,029, Mann-Whitney), do que as larvas que receberam 32l. Analisamos também dois genes envolvidos no metabolismo do HJ, e para um deles, jheh, codificador de uma epóxido-hidrolase do hormônio juvenil, não encontramos diferença nos níveis de mRNA entre larvas de rainhas e operárias. Para jhe, o qual codifica a esterase do hormônio juvenil, encontramos maiores níveis de transcritos no estágio LD nas larvas criadas com 130l de alimento. Isso indica que a expressão de jhe pode ser induzida pelo aumento dos títulos de HJ neste estágio. Os dois genes, hmgr e mfe, envolvidos no passo inicial e final da síntese do HJ, respectivamente, o primeiro mostrou uma pequena variação nos níveis de expressão, sendo que a expressão de mfe foi menor nas larvas criadas com 32l de alimento larval e foi encontrado um pico de expressão no estágio LPD nas larvas alimentadas com 130l. O produto desse gene, metilfarnesoato epoxidase, está envolvido no passo limitante da síntese de HJ em Apis mellifera. Dois outros genes analisados foram o homólogo ao EcR, codificador do receptor de ecdisona e o usp, codificador do potencial receptor do HJ. Seus níveis de expressões foram maiores nos estágios LPD e LD, respectivamente, nas larvas alimentadas com 130l de alimento e os níveis de EcR se correlacionaram com o perfil dos títulos de ecdisona e os de usp com os títulos de HJ, publicados para esta espécie. Não houve diferença nos níveis de expressão dos genes selecionados que pudessem relacionar ou permitir distinguir as rainhas miniaturas das operárias durante os estágios do desenvolvimento estudados. Uma análise morfométrica de adultos faratos indica claramente que as rainhas miniaturas são rainhas autênticas. / In this work we investigated the role played by the larval diet in the development of caste-specific morphological traits and in the expression of candidate genes during the development of the stingless bees, Scaptotrigona aff. depilis. In this species, the female castes are determinate by the amount of food consume during larval development. Experimental groups of larvae were reared in vitro on two different quantities of larval food obtained from newly provisioned brood cells of this species. Larvae of the first group received 32l of larval food, corresponding to the quantity usually received by the workers larva, whereas larvae of the second group received 130l, corresponding to the quantity normally deposited in queen cells. All larvae reared on 130l of larval food developed into queens, as expected, and similarly, most of the larvae reared on 32l developed into workers. Interestingly, however, some larvae of this group developed into miniature queens, suggesting that factors additional to the trophic ones may be involved in caste determination in S. aff. depilis. We subsequently analyzed the hemolymph juvenile hormone (JH) titers by radioimmunoassay during four stages of the last larval instar and we found that larvae of the 130l diet group had higher JH titer in the defecating stange (LD) (p=0,034, t-test) than larvae of the 32l diet group. Next we analyzed the expression levels of set of candidate genes that had previously been described as preferentially expressed in queens of others bee species. One of these genes dnmt-3, encodes DNA methyltransferase involved in DNA methylation. For this gene we found higher transcripts levels in prospective queens during the final larval stage (both in pre-defecating larvae LPD and defecating larvae- LD) (p=0,029, Mann-Whitney). We also analyzed two genes involved in JH metabolism, and for one of them jheh, encoding a juvenile hormone epoxide hydrolase, we did not find any differences in mRNA levels for queens and workers larvae. For jhe, which codes juvenile hormone esterase we found higher transcript levels in the LD stage in larvae reared on 130l of larval food. This indicates that jhe expression may be induced by an elevated JH titer during this stage. For two genes, hmgr and mfe, involved in an initial and a final step of JH synthesis, respectively, the first one showed little variation in expression levels, whereas mfe expression was lower in larvae reared on 32l of larval food and we found an expression peak in the LPD stage (p=0,029, Mann-Whitney) of larvae reared on 130l larval food. The product of this gene, a methylfarnesoate epoxidase, has been shown to be involved in a rate-limiting step of JH sunthesis in the honey bee, Apis mellifera. The other two genes analyzed were the EcR homolog encoding an ecdysone receptor and usp coding for potencial JH receptor. Their expressions were higher during LPD (p=0,006, t-test) stages and LD (p=0,026, t-test), respectively, in larvae fed with 130l of food and the levels of transcripts of EcR correlated with changes in the ecdysone titer and usp with the JH titer published for this species. We did not find differences in expressions levels for any of these candidate genes that could related to and allow to distinguish between prospective miniature queens and workers in these stages. A morphometric analysis of pharate adults, however, clearly demonstrated that the miniature queens are authentic queens.

Neste estudo, foi investigado o efeito da dieta larval no desenvolvimento de características morfológicas casta-específicas e na expressão de alguns genes durante o desenvolvimento da abelha sem ferrão Scaptotrigona aff. depilis . Nesta espécie, as castas femininas são determinadas pela quantidade de alimento consumida durante o desenvolvimento larval. Grupos experimentais larvas criadas in vitro foram feitos com duas quantidades diferentes de alimento larval, obtido de favos recém provisionados nesta espécie. As larvas do primeiro grupo receberam 32l de alimento larval, correspondente a quantidade média de alimento recebida por larvas de operárias naturalmente, já as larvas do segundo grupo receberam 130l, correspondente a quantidade média encontrada naturalmente em células reais. Todas as larvas criadas com 130l de alimento larval se desenvolveram em rainhas, como esperado; da mesma forma, a maioria das larvas criadas com 32l se desenvolveram em operárias. Interessantemente, porém, algumas larvas deste grupo se desenvolveram em rainhas miniaturas, sugerindo que outros fatores, além do trófico, estejam envolvidos na determinação de castas em S. aff. depilis. Subsequentemente, analisamos os títulos de hormônio juvenil (HJ) na hemolinfa por radioimunoensaio durante quatro estágios do último instar larval e encontramos que as larvas do grupo de 130l de dieta apresentaram maiores títulos no estágio defecante (LD) (p=0,034, t-test), se comparado com as larvas do grupo de 32l. Os níveis de expressão de determinados genes que foram previamente descritos em outras espécies como expressos preferencialmente em rainhas. Um destes genes, dnmt-3, codifica a DNA metiltransferase envolvida na metilação do DNA. Para este gene, encontramos maiores níveis de transcritos nas larvas alimentadas como rainhas (130l) nos estágios LPD e LD (p=0,029, Mann-Whitney), do que as larvas que receberam 32l. Analisamos também dois genes envolvidos no metabolismo do HJ, e para um deles, jheh, codificador de uma epóxido-hidrolase do hormônio juvenil, não encontramos diferença nos níveis de mRNA entre larvas de rainhas e operárias. Para jhe, o qual codifica a esterase do hormônio juvenil, encontramos maiores níveis de transcritos no estágio LD nas larvas criadas com 130l de alimento. Isso indica que a expressão de jhe pode ser induzida pelo aumento dos títulos de HJ neste estágio. Os dois genes, hmgr e mfe, envolvidos no passo inicial e final da síntese do HJ, respectivamente, o primeiro mostrou uma pequena variação nos níveis de expressão, sendo que a expressão de mfe foi menor nas larvas criadas com 32l de alimento larval e foi encontrado um pico de expressão no estágio LPD nas larvas alimentadas com 130l. O produto desse gene, metilfarnesoato epoxidase, está envolvido no passo limitante da síntese de HJ em Apis mellifera. Dois outros genes analisados foram o homólogo ao EcR, codificador do receptor de ecdisona e o usp, codificador do potencial receptor do HJ. Seus níveis de expressões foram maiores nos estágios LPD e LD, respectivamente, nas larvas alimentadas com 130l de alimento e os níveis de EcR se correlacionaram com o perfil dos títulos de ecdisona e os de usp com os títulos de HJ, publicados para esta espécie. Não houve diferença nos níveis de expressão dos genes selecionados que pudessem relacionar ou permitir distinguir as rainhas miniaturas das operárias durante os estágios do desenvolvimento estudados. Uma análise morfométrica de adultos faratos indica claramente que as rainhas miniaturas são rainhas autênticas. / In this work we investigated the role played by the larval diet in the development of caste-specific morphological traits and in the expression of candidate genes during the development of the stingless bees, Scaptotrigona aff. depilis. In this species, the female castes are determinate by the amount of food consume during larval development. Experimental groups of larvae were reared in vitro on two different quantities of larval food obtained from newly provisioned brood cells of this species. Larvae of the first group received 32l of larval food, corresponding to the quantity usually received by the workers larva, whereas larvae of the second group received 130l, corresponding to the quantity normally deposited in queen cells. All larvae reared on 130l of larval food developed into queens, as expected, and similarly, most of the larvae reared on 32l developed into workers. Interestingly, however, some larvae of this group developed into miniature queens, suggesting that factors additional to the trophic ones may be involved in caste determination in S. aff. depilis. We subsequently analyzed the hemolymph juvenile hormone (JH) titers by radioimmunoassay during four stages of the last larval instar and we found that larvae of the 130l diet group had higher JH titer in the defecating stange (LD) (p=0,034, t-test) than larvae of the 32l diet group. Next we analyzed the expression levels of set of candidate genes that had previously been described as preferentially expressed in queens of others bee species. One of these genes dnmt-3, encodes DNA methyltransferase involved in DNA methylation. For this gene we found higher transcripts levels in prospective queens during the final larval stage (both in pre-defecating larvae LPD and defecating larvae- LD) (p=0,029, Mann-Whitney). We also analyzed two genes involved in JH metabolism, and for one of them jheh, encoding a juvenile hormone epoxide hydrolase, we did not find any differences in mRNA levels for queens and workers larvae. For jhe, which codes juvenile hormone esterase we found higher transcript levels in the LD stage in larvae reared on 130l of larval food. This indicates that jhe expression may be induced by an elevated JH titer during this stage. For two genes, hmgr and mfe, involved in an initial and a final step of JH synthesis, respectively, the first one showed little variation in expression levels, whereas mfe expression was lower in larvae reared on 32l of larval food and we found an expression peak in the LPD stage (p=0,029, Mann-Whitney) of larvae reared on 130l larval food. The product of this gene, a methylfarnesoate epoxidase, has been shown to be involved in a rate-limiting step of JH sunthesis in the honey bee, Apis mellifera. The other two genes analyzed were the EcR homolog encoding an ecdysone receptor and usp coding for potencial JH receptor. Their expressions were higher during LPD (p=0,006, t-test) stages and LD (p=0,026, t-test), respectively, in larvae fed with 130l of food and the levels of transcripts of EcR correlated with changes in the ecdysone titer and usp with the JH titer published for this species. We did not find differences in expressions levels for any of these candidate genes that could related to and allow to distinguish between prospective miniature queens and workers in these stages. A morphometric analysis of pharate adults, however, clearly demonstrated that the miniature queens are authentic queens.

This research paper uses Critical Discourse Analysis (CDA) to analyse texts produced by the political party Bharatiya Janata Party (BJP) in India. The analysis use Machin and Mayr’s (2012) concepts of Language and Identity and Nominalisation and Presupposition with the aim to understand how the BJP can influence the democratic society in India through discourse. The texts analysed was taken from BJPs website and from parts of their 2014 manifesto. The theoretical framework and literature review are built on the role of Hinduism in the democratization of India. In this research, Hinduism act as an important factor in defining identity in India and Hindutva as an important factor in defining identity for the BJP. The analysis concludes that when looking at identity, the BJP demonstrate that their texts can have both a positive and a negative effect on the democracy in India. The BJP strongly use ‘India First’ to state that they want to unify the country under one identity and similarities can be drawn to their previous use of ‘Hindutva’. By promoting ‘India First’ the BJP includes a large audience and a somewhat tolerant outlook by stating to include all castes and ethnicities. However, the analysis demonstrates that their strong promotion of ‘India First’ conceal who is responsible to uphold this identity and that in turn could affect the tolerance in society. The analysis also shows that their definition of ‘India First’ is left vague and this can conceal certain interest. Their use of ‘India First’ as an identity can lead to a fear that everything that does not belong under this category is a threat. This combined with the diffuse definition of what ‘India First’ mean can have a negative effect on the pluralistic and tolerant society that was needed for India to transform to a democracy. The research also explores whether the strong promotion of ‘India First’ can be compared to a religious or spiritual movement and touch upon the implications that could follow from that.

Les Hyménoptères sociaux (abeilles, fourmis et guêpes) doivent leur succès écologique à une division dutravail marquée par l’existence de castes. Les reines sont spécialisées dans la reproduction, tandis que les ouvrières sont stériles et prennent en charge la construction du nid et sa défense, la récolte de nourriture et l’élevage des jeunes. Le développement d’un œuf femelle en une reine ou une ouvrière est généralement régi par des facteurs environnementaux, tels que la qualité ou la quantité de la nourriture.En réalisant une étude de génétique des populations chez la fourmi Cataglyphis hispanica, nous avons mis en évidence un mode de reproduction inhabituel appelé hybridogenèse sociale. Cette espèce est caractérisée par la co-existence de deux lignées génétiques au sein des populations. Les reines de chaque lignée s’accouplent systématiquement avec un mâle de l’autre lignée génétique. Ainsi, les reines de la lignée 1 s’accouplent toujours avec un mâle de la lignée 2, et les reines de la lignée 2 s’accouplent avec un mâle de la lignée 1. Les ouvrières sont issues du croisement entre les deux lignées :il s’agit donc d’individus hybrides. A l’inverse, les individus reproducteurs (nouvelles reines et mâles) sont produits exclusivement par reproduction asexuée. La caste femelle (reine ou ouvrière) est donc déterminée génétiquement; les ouvrières portent un génome hybride, alors que les reproducteurs possèdent un génome non-hybride hérité exclusivement de la mère.Les travaux réalisés dans le cadre de cette thèse de doctorat visent à comprendre l’origine et l’évolution de l’hybridogenèse sociale chez les fourmis Cataglyphis par le biais d’approches variées (génétique des populations, phylogéographie et manipulation de colonies en laboratoire). Ils sont articulés selon plusieurs axes complémentaires :la description du phénomène, l’étude de sa distribution géographique et phylogénétique au sein du genre Cataglyphis, et l'étude de ces mécanismes génétiques sous-jacents. / In eusocial Hymenoptera, such as bees, wasp and ants, it is commonly accepted that the diploid female eggs are bipotent and develop either into a queen or a worker depending on environmental factors. While conducting a population genetic study of the ant Cataglyphis hispanica, we discovered an unusual reproductive system called social hybridogenesis. Under this system, queens and workers develop from eggs with different genetic make-up. Two divergent genetic lineages coexist within a population. Queens of each lineage mate with males originating from the other lineage. Workers are produced by sexual reproduction; hence, they are first generation hybrids of the two lineages. By contrast, new queens and males are produced by thelytokous and arrhenotokous parthenogenesis, respectively. By using alternative modes of reproduction, queens appear to maximize their reproductive success by increasing the transmission rate of their genes to the reproductive offspring while benefiting from genetic diversity and heterosis effects in their worker force. Males, however, do not contribute genes to the next generation as they only sire sterile workers. This system is expected to be evolutionary short-lived as lineages may be selected to stop the production of males with null direct fitness. This PhD project aims at understanding the origin and evolution of social hybridogenesis in Cataglyphis. Using several different approaches including population genetics methods based on microsatellite makers, phylogeography and experimental manipulations on lab colonies, I explored tthe characteristics of social hybridogenesis, the genetic mechanisms underlying the system and its distribution within the genus Cataglyphis. My results, together with other recent findings, question the paradigm of environmental caste determination in eusocial Hymenoptera and suggest that genetic influences on caste determination may be more common than previously thought. / Doctorat en Sciences / info:eu-repo/semantics/nonPublished