Doctor of Philosophy / Department of Biology / Susan J. Brown / All arthropods share a segmented body plan. Detailed studies on segmentation mechanisms in the long-germ insect Drosophila melanogaster identified a segmentation hierarchy composed of maternal, gap, pair-rule, and segment polarity genes. In this hierarchy, pair-rule genes play an important role to translate gradients of regional information from maternal and gap genes into segmental expression of segment polarity genes. However, our understanding of the role of pair-rule genes in other short-germ insects and basally branching arthropods is still limited. To gain insights into the role of pair-rule genes in short-germ segmentation, I analyzed genetic interactions as well as expression patterns and functions of homologs of Drosophila pair-rule genes in the short-germ insect Tribolium castaneum. Interestingly, despite the pair-rule like expression patterns of Tribolium homologs of almost all eight canonical Drosophila pair-rule genes, only five have a segmentation function. Knock-down of primary pair-rule genes caused asegmental and truncated phenotypes while knock-down of secondary pair-rule genes caused typical pair-rule phenotypes. Epistatic analysis between the genes revealed that primary pair-rule genes form a gene circuit to prepattern a two-segmental unit, and secondary pair-rule genes are downstream targets of the gene circuit. The typical pair-rule phenotypes observed in secondary pair-rule gene RNAi embryos led to a detailed comparative analysis of the role of paired (prd) and sloppy-paired (slp) between Drosophila and Tribolium. This study revealed that prd is functionally conserved while the functional parasegmental register for Tribolium slp is opposite that of Drosophila slp. The fact that the register of slp function has evolved differently in the lineages leading to Drosophila and Tribolium reveals an unprecedented flexibility in pair-rule patterning. Despite this flexibility in pair-rule patterning between Drosophila and Tribolium, segmental expression of engrailed (en) and wingless (wg) at parasegmental boundaries is conserved in both insects. Analysis of double and triple RNAi for pair-rule genes in Tribolium revealed that the primary pair-rule genes even-skipped and runt are redeployed to directly regulate en and wg with prd or slp at parasegmental boundaries. This redeployment of primary pair-rule genes seem to compensate for the apparently fewer number of functional secondary pair-rule genes in Tribolium segmentation.

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<p>The importance of cryptic female choice, i.e. female post-copulatory influence over male reproductive success, in driving the evolution of male traits remains controversial. The main aim of this thesis was to understand the post-copulatory consequences of sexual interactions and the importance of cryptic female choice in two species of beetle.</p><p>Males of the red flour beetle Tribolium castaneum use their legs to rub the lateral edges of the female elytra during mating. When manipulating female perception of this behaviour, I found that females preferentially use the sperm of males with vigorous leg rubbing when they mate with more than one male. Leg rubbing also appeared to increase female rate of oviposition. Females do not seem to gain any indirect benefits by preferring males with an intense leg rubbing behaviour since this behaviour was found to have very low narrow sense heritability and did not appear to be condition dependent in its expression.</p><p>Males of the bruchid beetle Callosobruchus maculatus have spiny genitalia that harm their mates. Females kick males during copulation and when prevented from kicking, suffered reduced lifetime offspring production as a consequence of more extensive injuries. Males were not able to delay female remating, increase rate of oviposition or increase sperm precedence by inflicting relatively severe injuries to non-kicking females. Hence, the injuries appear to be side effects of male efforts to remain in copula. When copulation duration was manipulated, ejaculate size and female lifetime offspring production increased with the length of copulation. Females reduced their mating rate when they had access to water, suggesting that they obtain water from the large ejaculates and trade-off their need for additional water against the costs of mating. Males may then reduce the benefits of remating by providing their mates with a large amount of water. Females did not increase their remating propensity to avoid inbreeding when they had mated to brothers. Together, these studies reveal the complexity of sexual interactions and the importance of post-copulatory processes for the fitness of both males and females.</p>

The importance of cryptic female choice, i.e. female post-copulatory influence over male reproductive success, in driving the evolution of male traits remains controversial. The main aim of this thesis was to understand the post-copulatory consequences of sexual interactions and the importance of cryptic female choice in two species of beetle. Males of the red flour beetle Tribolium castaneum use their legs to rub the lateral edges of the female elytra during mating. When manipulating female perception of this behaviour, I found that females preferentially use the sperm of males with vigorous leg rubbing when they mate with more than one male. Leg rubbing also appeared to increase female rate of oviposition. Females do not seem to gain any indirect benefits by preferring males with an intense leg rubbing behaviour since this behaviour was found to have very low narrow sense heritability and did not appear to be condition dependent in its expression. Males of the bruchid beetle Callosobruchus maculatus have spiny genitalia that harm their mates. Females kick males during copulation and when prevented from kicking, suffered reduced lifetime offspring production as a consequence of more extensive injuries. Males were not able to delay female remating, increase rate of oviposition or increase sperm precedence by inflicting relatively severe injuries to non-kicking females. Hence, the injuries appear to be side effects of male efforts to remain in copula. When copulation duration was manipulated, ejaculate size and female lifetime offspring production increased with the length of copulation. Females reduced their mating rate when they had access to water, suggesting that they obtain water from the large ejaculates and trade-off their need for additional water against the costs of mating. Males may then reduce the benefits of remating by providing their mates with a large amount of water. Females did not increase their remating propensity to avoid inbreeding when they had mated to brothers. Together, these studies reveal the complexity of sexual interactions and the importance of post-copulatory processes for the fitness of both males and females.

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Submitted by Reginaldo Soares de Freitas (reginaldo.freitas@ufv.br) on 2017-04-05T18:50:23Z No. of bitstreams: 1 texto completo.pdf: 1802151 bytes, checksum: 55ced28e2e4f99399e06e35026253dfd (MD5) / Made available in DSpace on 2017-04-05T18:50:23Z (GMT). No. of bitstreams: 1 texto completo.pdf: 1802151 bytes, checksum: 55ced28e2e4f99399e06e35026253dfd (MD5) Previous issue date: 2005-06-10 / Coordenação de Aperfeiçoamento de Pessoal de Nível Superior / O interesse por métodos não-químicos de controle de pragas em grãos armazenados vem aumentando, uma vez que os consumidores estão menos tolerantes aos resíduos nos alimentos e há sucessivos relatos de resistência a inseticidas em muitas espécies de insetos de produtos armazenados. O controle biológico surge como uma das alternativas e existem vários casos de sucesso desse método, principalmente quando inserido em programas de manejo integrado de pragas. Pesquisas com o ácaro Acarophenax lacunatus (Cross & Krantz) vêm sendo desenvolvidas e demonstram que sua utilização no controle biológico de coleópteros-praga de produtos armazenados é promissora. Assim, esta pesquisa foi dividida em cinco etapas que abordam estudos básicos com este inimigo natural, visando fornecer subsídios para sua futura liberação em programas de controle biológico em ambientes de armazenamento. A primeira etapa está relacionada com a capacidade de A. lacunatus sobreviver à inanição sob diferentes temperaturas. Foram simuladas situações de escassez de alimento, sendo observado que A. lacunatus conseguiu sobreviver na ausência de alimento por até 108 horas, nas temperaturas de 20, 25 e 28 0C. A segunda etapa avaliou se este ácaro dispersa numa massa de grãos e consegue localizar o hospedeiro Rhyzopertha dominica (Fabricius). Constatou-se a habilidade de A. lacunatus em penetrar até 20 cm de profundidade e localizar ovos de R. dominica em diferentes regiões na massa de grãos, colonizando o ambiente de forma ativa. A terceira etapa consistiu no estudo da biologia de A. lacunatus sobre os hospedeiros Tribolium castaneum (Herbst) e Cryptolestes ferrugineus (Stephens) nas temperaturas 20, 25, 28, 30, 32 e 35 0C. A temperatura influenciou os parâmetros biológicos do ácaro, sendo a taxa intrínseca de crescimento (rm), a taxa líquida reprodutiva (Ro) e a razão sexual diretamente proporcionais ao aumento da temperatura. As maiores progênies (19 indivíduos/fêmea do ácaro em T. castaneum e 15 indivíduos/fêmea do ácaro em C. ferrugineus) foram observadas à temperatura de desenvolvimento. 30 A 0 C, demonstrando quarta que essa temperatura favorece seu etapa abordou um estudo de seleção de linhagens de A. lacunatus, que foram mantidas por sucessivas gerações, isoladamente, sobre os hospedeiros T. castaneum, C. ferrugineus e R. dominica. Teve como objetivo testar a preferência do ácaro por ovos desses coleópteros, em testes mensais, durante nove meses, na tentativa de detectar possíveis mudanças nessa preferência que possam indicar a seleção de linhagens desse ácaro com uma melhor performance sobre estes insetos. Houve uma influência significativa do tempo sobre a taxa de parasitismo, sendo R. dominica o hospedeiro mais parasitado em todas as situações, indicando que esta preferência parece estar fixada geneticamente, uma vez que A. lacunatus vem sendo mantido sobre este hospedeiro desde sua introdução no Brasil. Foi calculada a herdabilidade (h2) para T. castaneum e C. ferrugineus, sendo que este valor foi maior para T. castaneum. Isso indica que a seleção de linhagens desse ácaro com um melhor desempenho sobre T. castaneum pode ser realizada. Já a quinta etapa consistiu no estudo do potencial de parasitismo de A. lacunatus sobre o coleóptero T. castaneum, onde observou-se que o aumento da densidade do ácaro implicou no aumento do número de ovos parasitados, com conseqüente redução das populações desse inseto. Assim, novas informações sobre A. lacunatus são acrescentadas e indicam que sua utilização no controle biológico de T. castaneum, C. ferrugineus e R. dominica pode ser viável. / The interest for non-chemical control methods of stored grain pests is increasing since the consumers are less tolerant about insecticide residues on food and there are reports of insecticide resistance in insects of stored products. The biological control appears as an alternative and there are several success cases of the control method, mainly within integrated pest management programs. Studies with the mite Acarophenax lacunatus (Cross & Krantz) have been carried out and show the potential of its use in the biological control of stored grain beetles. Therefore the present study was divided in five phases encompassing basic investigations with this natural enemy aiming to provide support for its future use in biological control programs in storage facilities. The first phase is related with the capacity of A. lacunatus to survive under starvation at different temperatures. This mite species was able to survive for up to 108 hours at the temperatures of 20, 25 and 28 0C. The second phase focused on assessing the dispersal of this mite species in the grain mass and its ability to locate its host Rhyzopertha dominica (Fabricius). The mite was able to penetrate up to 20 cm deep and locate eggs of R. dominica in different regions of the grain mass efectivelly colonizing the environment. The third phase involved the biology of A. lacunatus on the hosts Tribolium castaneum (Herbst) and Cryptolestes ferrugineus (Stephens) at the temperatures 20, 25, 28, 30, 32 and 35 0C. The temperature affected the biological parameters of the mite with a proportional increase in the intrinsic growth rate (rm) net reproductive rate (Ro) and sex ratio with increased temperature. The greates progenies (19 individuals/female in T. castaneum and 15 individuals/female in C. ferrugineus) were observed at 30 0C temperature showing that this temperature favors the mite development. The fourth phase focused the selection of mite strains maintained for several generations in different hosts (T. castaneum, C. ferrugineus and R. dominica). The objective was to test the host egg preference of each selection line in monthly tests trying to detect changes in preference through time. There was a significant influence of time on the parasitism rate with R. dominica always being the preferred host and indicating that this preference was genetically fixed in the mite population. The heritability (h2) of egg preference for the hosts T. castaneum and C. ferrugineus was calculated and higher values were observed for T. castaneum. This indicates that selection for better performance in T. castaneum is possible. In the fifth phase of the present effort, we investigated the parasitism potential of A. lacunatus in the beetle T. castaneum where an increase in mite density led to an increase in the number of parasitized eggs with a consequent reduction of this insect population. Therefore, new information on A. lacunatus was obtained indicating the viability of its use in biological control programs against T. castaneum, C. ferrugineus and R. dominica.

Submitted by Marco Antônio de Ramos Chagas (mchagas@ufv.br) on 2017-06-27T17:46:33Z No. of bitstreams: 1 texto completo.pdf: 647098 bytes, checksum: b268e217f616628b33c7d54bc5890149 (MD5) / Made available in DSpace on 2017-06-27T17:46:33Z (GMT). No. of bitstreams: 1 texto completo.pdf: 647098 bytes, checksum: b268e217f616628b33c7d54bc5890149 (MD5) Previous issue date: 2001-07-24 / Conselho Nacional de Desenvolvimento Científico e Tecnológico / Com a necessidade de buscar alternativas para o uso de agentes químicos em produtos armazenados, devido ao desenvolvimento de resistência em insetos-praga e à possibilidade de contaminação dos alimentos por resíduos, os programas de erradicação de pragas têm dado maior ênfase ao controle biológico. Em razão do exposto, este trabalho foi dividido em três etapas que abordaram o potencial de parasitismo do ácaro Acarophenax lacunatus (Prostigmata: Acarophenacidae) em cinco espécies de insetos de produtos armazenados. Na primeira etapa, avaliou-se a preferência deste ácaro por ovos dos insetos Rhyzopertha dominica (Coleoptera: Bostrichidae), Dinoderus minutus (Coleoptera: Bostrichidae), Tribolium castaneum (Coleoptera: Tenebrionidae) e Oryzaephilus surinamensis (Coleoptera: Silvanidae). A segunda etapa abordou o estudo do potencial de parasitismo do ácaro em D. minutus em mandioca seca armazenada, e a terceira etapa em R. dominica, T. castaneu m, O. surinamensis e Cryptolestes ferrugineus (Coleoptera: Cucujidae) em trigo parcialmente triturado. Todos os testes foram realizados a 28 o C e escotofase de 24 horas, em câmaras climáticas do tipo B.O.D. O estudo da preferência por ovos foi realizado em dois testes: um sem chance de escolha e outro com chance de escolha, utilizando-se placas de Petri (5 cm de diâmetro), as quais foram divididas em quatro quadrantes. No teste sem chance de escolha, foram colocados, em cada quadrante, cinco ovos de apenas um dos insetos e, no centro da placa, uma fêmea de A. lacunatus em processo de fisogastria. O experimento foi realizado com cada uma das espécies. No teste com chance de escolha, foram utilizados ovos de todas as espécies, que foram distribuídos de forma que cada quadrante só recebesse ovos de um dos insetos. Da mesma forma, no centro da placa foi liberada uma fêmea fisogástrica do ácaro. Em nenhum dos testes o ácaro A. lacunatus parasitou ovos de O. surinamensis. No teste sem chance de escolha não foi observada preferência de A. lacunatus por ovos de nenhum dos outros hospedeiros. Entretanto, no teste com livre escolha, A. lacunatus parasitou mais ovos de R. dominica e D. minutus que de T. castaneum. Nas outras etapas do trabalho, avaliou-se a população final dos insetos (larvas e adultos), o número de ovos parasitados e de ácaros em processo de fisogastria, a taxa instantânea de crescimento (r i ) de A. lacunatus e dos insetos, e a perda de peso dos produtos (mandioca e trigo). Foi observado que o ácaro apresenta potencial biótico maior que o dos hospedeiros R. dominica, D. minutus e T. castaneum. Desta forma, o número de larvas e adultos destes insetos sofreu reduções em relação à testemunha, o que também ocorreu com as perdas de matéria seca ocasionadas pelo ataque dos insetos. Também não foi observado parasitismo em populações do coleóptero O. surinamensis. Pelo número de ovos parasitados de R. dominica, D. minutus, T. castaneum e C. ferrugineus, conclui-se que o ácaro A. lacunatus é um inimigo natural eficaz destes insetos, não sendo, portanto, um parasita específico de R. dominica , como se imaginava. / With the necessity of searching for alternatives to the use of chemical control agents in stored products due to development of insecticide resistance in insect-pests and the possibility of foodstuff contamination by residue, pest eradication programs have been putting more emphasis on biological control. As a consequence, the present work was divided in three phases which aimed to assess the parasitism potential of the mite Acarophenax lacunatus (Prostigmata: Acarophenacidae) in five species of insects of stored products. In the first phase, the mite preference for eggs of Rhyzopertha dominica (Coleoptera: Bostrichidae), Dinoderus minutus (Coleoptera: Bostrichidae), Tribolium castaneum (Coleoptera: Tenebrionidae) and Oryzaephilus surinamensis (Coleoptera: Silvanidae) was assessed. In the second phase, the parasitism potential of the mote species of D. minutus attacking dried stored cassava was evaluated and in the third phase the parasitism potential of the same parasitic species was assessed in R. dominica, T. castaneum, O. surinamensis e Cryptolestes ferrugineus (Coleoptera: Cucujidae) infesting wheat. All of the experiments were carried out under controlled conditions of 28 o C and 24 hours scotophase in climatic chambers of the B.O.D. type. The study of egg preference was carried out in two tests: one without opportunity of choice and other with it using Petri dishes divided in four areas. On the non-choice test five eggs of only one species was placed in each different area and a physogastric female of A. lacunatus was placed in the centre of the Petri dish (5 cm diameter). On the choice test there was only eggs of a single species in each of the four areas within the Petri dish and the same procedure was used for each potential prey species. The mite A. lacunatus did not attack eggs of O. surinamensis in any of the tests. No preference of A. lacunatus by eggs of any host was observed on the non-choice test. However, A. lacunatus attacked more eggs R. dominica and D. minutus than of T. castaneum. In the other phases of the work the final insect population (adults and larvae), egg parasitism, number of mites in physogastry, the instantaneous rate of increase of the mite and the insect populations (r i ) and weight loss of the stored product (wheat or cassava). It was observed that the mite has higher biotic potential than the hosts R. dominica, D. minutus and T. castaneum. Therefore, the number of larvae and adults of these insect species were lower than in the control treatment, what was also observed with the losses in dried matter caused by insect infestation. Again, no parasitism on eggs of O. surinamensis was observed. Based on the egg parasitism of R. dominica, D. minutus, T. castaneum and C. ferrugineus, the mite A. lacunatus is an efficient natural enemy for all of this species and not only to R. dominica, as initially reported. / Dissertação importada do Alexandria