Made available in DSpace on 2015-03-26T13:30:23Z (GMT). No. of bitstreams: 1 texto completo.pdf: 464995 bytes, checksum: da8a31699fc6432c9841c8e8d888ad17 (MD5) Previous issue date: 2008-09-29 / Coordenação de Aperfeiçoamento de Pessoal de Nível Superior / Leafcutter ants live in a world of complex interactions, the most notable being their cultivation of a mutualistic fungus on vegetable subtrates. These ants, and their mutualistic fungus gardens, are at risk from attack by parasitic microorganisms. To defend themselves and their mutualist, they have antimicrobial defences, such as substances secreted from their metapleural glands. As this defence is subject to selective pressures according to the risk of parasite attack, variations in the amount of defence may reflect variations in the risk. Meanwhile, there may be variation in the value of individual workers to the colony, dependent in part upon the area in which they work, and investment in their defences may also reflect this. In this study, the volume of the externally visible part (bulla) of the reservoir of the metapleural gland of workers was compared betwen species, colonies and tasks, to determine variations in defence strategies against pathogens. Additionally, colony sizes of a bacterial mutualist on the workers thoraxes, were estimated, looking for a possible relationship with metapleural gland bulla size. A total of eleven laboratory colonies were used, of four species of leafcutter ants: Atta laevigata (F.Smith, 1858), Atta sexdens rubropilosa Forel, 1908, Acromyrmex subterraneus brunneus (Forel, 1911) and Acromyrmex disciger (Mayr, 1887). Workers were collected from three areas of the colonies: foraging areas, symbiotic fungus gardens and the refuse deposits. Parameters measured were: the volume of the bulla of the metapleural gland, the weight of ants and an estimate of the olony sizes of the symbiotic bacteria. Analyses of covariance were conducted using individual mass as a covariate, to determine the contributions of a range of parameters in explaining variation in the size of this organ. The two species of Atta were found to have larger bulla when compared to the two species of Acromyrmex., probably reflecting not a greater degree of exposure to parasites but a strategy of greater investment in defences. Gardeners and foragers of the species A. laevigata and Ac. subterraneus brunneus had larger bulla than did waste workers, revealing a lower quality (in terms of parasite defence) of the waste workers. In this case, we expect that these individuals, being of lower quality, are allocated to more dangerous tasks, to die more rapidly. Acromyrmex subterraneus brunneus was the only species with visible colonies of the bacterial mutualist on the thorax. There was a positive relation between the size of this colony and the volume of the bulla of the metapleural gland. These variations in degrees of defence imply that the ants have different strategies to defend their colonies against pathogens and parasites. / Formigas cortadeiras apresentam interações complexas. São cultivadoras de fungo o qual é alimentado por substratos vegetais. Tanto as formigas como o fungo do jardim correm o risco de serem parasitados por microrganismos. Para defenderem tanto o fungo quanto elas mesmas, as formigas utilizam defesas antimicrobianas, como substâncias secretadas através da glândula metapleural. Sendo que esta defesa está sujeita a pressão de seleção de acordo com o risco de ataque por parasitas, variações no investimento nela podem ser consequência deste risco. Ao mesmo tempo, pode haver indivíduos com diferentes valores para a colônia, dependendo em parte na área onde são encontradas, e podem ter maior ou menor investimento em defesas contra parasitas. Diante disso o trabalho propôs verificar a variação do volume do reservatório da glândula metapleural (bula) entre espécies, colônias e tarefas de formigas cortadeiras para verificar possíveis estratégias de defesas das operárias contra patógenos. Avaliou-se também a presença de uma bactéria mutualista, que é utilizada também como defesa contra parasitismo, e a possível relação entre esta e o volume do reservatório da glândula metapleural. Foram utilizadas um total de 11 colônias de quatro espécies de formigas cortadeiras, Atta laevigata (F.Smith, 1858), Atta sexdens rubropilosa Forel, 1908, Acromyrmex subterraneus brunneus (Forel, 1911) e Acromyrmex disciger (Mayr, 1887). Formigas foram coletadas de três áreas das colônias: as áreas de forrageamento, os jardins do fungo simbionte e o lixo. As informações tomadas foram: volume da bula da glândula metapleural, massa das formigas e uma estimativa do tamanho das colônias de bactéria mutualista. Análises de covariância foram utilizadas, com massa dos indivíduos como covariável, para determinar as contribuições de diversos fatores para explicar o tamanho desse órgão. Pelas análises obtidas, as duas espécies de Atta possuem bulas maiores em relação às duas espécies de Acromyrmex., provavelmente refletindo não uma maior exposição a parasitas mas uma estratégia de maior investimento em defesas. As jardineiras e as forrageadoras das espécies A. laevigata e Ac. subterraneus brunneus tiveram bulas maiores em relação às lixeiras, isto mostrando uma menor qualidade (em termos de defesas contra parasitas) das lixeiras. Neste caso, suponhamos que estes indivíduos, sendo de menor qualidade, são alocados a tarefas mais perigosas e morrerão rapidamente. Acromyrmex subterraneus brunneus foi a única que apresentou colônias de bactéria mutualista, dispostas no tórax. Houve uma relação positiva do tamanho desta colônia com o volume da bula da glândula metapleural. Essas variações nesses sistemas de defesas implicam que as formigas podem apresentar diferentes estratégias para defender o ninho contra patógenos e parasitas.

abstract: The most advanced social insects, the eusocial insects, form often large societies in which there is reproductive division of labor, queens and workers, have overlapping generations, and cooperative brood care where daughter workers remain in the nest with their queen mother and care for their siblings. The eusocial insects are composed of representative species of bees and wasps, and all species of ants and termites. Much is known about their organizational structure, but remains to be discovered. The success of social insects is dependent upon cooperative behavior and adaptive strategies shaped by natural selection that respond to internal or external conditions. The objective of my research was to investigate specific mechanisms that have helped shaped the structure of division of labor observed in social insect colonies, including age polyethism and nutrition, and phenomena known to increase colony survival such as egg cannibalism. I developed various Ordinary Differential Equation (ODE) models in which I applied dynamical, bifurcation, and sensitivity analysis to carefully study and visualize biological outcomes in social organisms to answer questions regarding the conditions under which a colony can survive. First, I investigated how the population and evolutionary dynamics of egg cannibalism and division of labor can promote colony survival. I then introduced a model of social conflict behavior to study the inclusion of different response functions that explore the benefits of cannibalistic behavior and how it contributes to age polyethism, the change in behavior of workers as they age, and its biological relevance. Finally, I introduced a model to investigate the importance of pollen nutritional status in a honeybee colony, how it affects population growth and influences division of labor within the worker caste. My results first reveal that both cannibalism and division of labor are adaptive strategies that increase the size of the worker population, and therefore, the persistence of the colony. I show the importance of food collection, consumption, and processing rates to promote good colony nutrition leading to the coexistence of brood and adult workers. Lastly, I show how taking into account seasonality for pollen collection improves the prediction of long term consequences. / Dissertation/Thesis / Doctoral Dissertation Applied Mathematics 2018

What selective forces and molecular mechanisms govern the integration of worker body size and morphology, brain architecture, and behavior in insect societies? Workers of the remarkably polyphenic and socially complex fungus-growing leafcutter ant Atta cephalotes exhibit a striking agricultural division of labor. The number of morphologically distinct and behaviorally differentiated worker groups, adaptive mosaic neural phenotypes, and brain transcriptomes have not been examined and the influences of socioecological challenges on behavioral performance, cognition, and brain evolution are unclear. We quantified worker morphological and behavioral variation to assess the number of worker size classes and characterized their social roles. We discriminated multiple worker size groups using a Gaussian mixture model: mid-sized workers (“medias”) had the most diverse task repertories and serve dominant roles in leaf harvesting, whereas workers of other size classes performed fewer, more specialized behaviors. We used variation among tasks in sensorimotor functions and task performance frequencies to create an estimate of sensory integration and processing demands across worker size groups. This metric predicted that medias require the greatest neural investment due to the high diversity of sensory inputs and motor functions associated with their task set. We quantified the volumes of key neuropils in brains of workers of different sizes and determined their allometries, finding that our estimate corresponded to proportional investment in the mushroom bodies, a brain compartment responsible for learning, memory, and sensory integration, and identifying allometric scaling patterns in other brain centers. Additionally, we measured whole-brain gene expression and identified significant differences in expression levels for numerous genes likely to underpin behavior. Differences were most pronounced between the smallest (fungal gardener “minims”) and largest (defensive “majors”), although not all expression differences were driven by worker size. Overrepresented gene functional categories included those related to sensory processing (enriched in genes upregulated in medias and minims) and metabolism (enriched in genes upregulated in majors). These results identify the nature of selective forces favoring differentiation along morphological, neuroanatomical, behavioral, and molecular axes among A. cephalotes workers and the impact of advanced division of labor on brain evolution. / 2023-03-01T00:00:00Z