La cléistogamie est un régime de reproduction mixte particulier – contrairement aux régimes mixtes classiques où un seul morphe floral effectue l'auto- et l'allofécondation, les espèces cléistogames produisent des fleurs fermées (cléistogames, CL) qui sont obligatoirement autofécondées et des fleurs ouvertes (chasmogames) qui sont potentiellement allofécondées. D'autres particularités sont associées aux régimes cléistogames : i) les fleurs fermées ne peuvent pas exporter de pollen (pollen discounting total), résultant avec l'absence de l'avantage automatique de l'autofécondation, ii) les couts de production des deux morphes floraux ne sont pas les mêmes, iii) les différences observées entre les descendants CL et CH sont déterminée par le degré de consanguinité mais aussi par le type de fleur dont un descendant est issu. Le taux de fleurs CH d'un individu est souvent plastique, ainsi la cléistogamie peut être un moyen d'ajuster le taux d'allofécondation en fonction des conditions environnementales. Nous avons étudié une espèce annuelle cléistogame, Lamium amplexicaule, qui peut produire une génération de plantes au printemps et une à l'automne et dont le taux CH dépend de la variation des facteurs associés avec la saison. Nous avons récolté des données d'observation en population naturelle, des expériences en conditions semi-naturelles au printemps et à l'automne, effectué des analyses génotypiques avec des marqueurs microsatellites et construit un modèle théorique simple dans le but de i) étudier la variation du taux CH et son caractère plastique ; ii) estimer le taux d'allofécondation des fleurs CH et son lien avec le taux CH ; iii) tester le caractère adaptatif de la cléistogamie plastique iv) tester différents scénarios évolutifs qui pourrait expliquer le maintien de la cléistogamie plastique de L. amplexicaule. Nos résultats montrent que la cléistogamie chez L. amplexicaule est un trait plastique et adapté à la variation entre saisons et aussi que la variation du taux CH se traduit en variation du taux d'allofécondation global. Nous résultats infirment une hypothèse classique souvent utilisée pour expliquer l'évolution de la cléistogamie, à savoir que la production des deux types des fleurs en proportions variables est une adaptation à la disponibilité des ressources, et suggèrent que c'est plutôt la différence de valeur sélective entre les descendants CL et CH ou la variation de l'abondance des pollinisateurs qui expliquent mieux la plasticité adaptative de la cléistogamie chez L. amplexicaule. Dans les perspectives de ce travail, nous proposons d'approfondir les études qui explorent le lien entre l'effet des forces évolutives qui opèrent au sein des régimes mixtes classiques (un seul morphe floral) et l'effet de la spécialisation des structures florales pour différents modes de reproduction. / Cleistogamy differs from classical mixed mating systems, for which species with single floral morph self-fertilize at intermediate rates: cleistogamous plants produce both closed cleistogamous flowers (CL) that are obligately selfed and open chasmogamous flowers (CH) that are potentially outcrossed. Because CL flowers cannot export pollen (total pollen discounting), cleistogamous species do not benefit from the automatic advantage of selfing. Furthermore, costs for producing the two floral types are different, and the two types of progeny they produce (CL and CH) have different properties that go beyond the differences between selfed and outcrossed progeny. The proportion of individual CH flowers is often plastic, suggesting this trait is an adaptation of the outcrossing rate to environmental variation. Here, we studied an annual cleistogamous species, Lamium amplexicaule, that has both spring and autumn generations each year, and whose CH proportion correlates with variation in seasonal cues. We combined data from field surveys, semi-natural experimental studies in spring and autumn, genetic analyses of neutral markers, and some theoretical modeling to i) assess the variation in CH proportion and its plasticity, ii) assess the outcrossing rate of CH flowers and its relation to the CH proportion, iii) test the adaptive character of plastic cleistogamy, and iv) test evolutionary scenarios that could explain the maintenance of plastic cleistogamy in L. amplexicaule. We show that cleistogamy in L. amplexicaule is plastic and adaptive to seasonal variation, and that CH proportion variation translates into variation of the overall outcrossing rate. Classical explanations for cleistogamy evolution relying on resource allocation to CL and CH flowers do not fit our data; we instead propose that the adaptive character of plastic cleistogamy could be due to environmentally dependent variation in fitness of CL and CH progeny and pollinator abundance. More studies of the evolution of cleistogamy need to account for the combined effect of classical evolutionary forces that operate on the reproductive systems of monomorphic flower species and the effect of floral specialization to different mating types.

Chez les animaux sexués, les stratégies d'appariement et de dispersion sont influencées par de nombreux facteurs de l'environnement. Certains facteurs sont communs, ce qui suggère que ces deux stratégies n'évoluent pas indépendamment. Parmi ces facteurs, la structure génétique des populations pourrait jouer un rôle important dans cette évolution conjointe. Par une approche théorique, j'ai étudié l'évolution de la différence de dispersion entre les sexes en lien avec les stratégies d'appariement (monogamie, monoandrie, polyandrie). En monoandrie et polyandrie, stratégies où les deux sexes ont des nombres de partenaires différents, la compétition entre apparentés et la variation spatiale du succès reproducteur plus importante chez les mâles que les femelles, favorisent l'évolution d'une dispersion biaisée en faveur des mâles. Je me suis aussi intéressé à la place des gènes dans le génome et j'ai montré que la localisation de gènes de dispersion sur les chromosomes sexuels peut, à elle-seule, faire évoluer un différentiel de dispersion entre les sexes. J'ai exploré cette dernière possibilité dans le cadre de conflits génomiques potentiels. Parallèlement à cette approche théorique, j'ai effectué une étude expérimentale chez le lézard vivipare (Zootoca vivipara). J'ai recherché le rôle d'un effet maternel, médié par la corticostérone, sur le lien entre la stratégie d'appariement des femelles et la stratégie de dispersion de leurs jeunes. Je me suis aussi intéressé au coût de l'accouplement via la transmission de pathogènes entre partenaires. Mes résultats suggèrent l'absence d'effet direct des accouplements multiples sur la communauté bactérienne cloacale. / In sexually reproducing animals, dispersal and mating strategies are influenced by many environmental factors, some of these factors being common, which suggests that these two strategies could evolve jointly. Among these factors, population genetic structure could play a great role on this joint evolution. Using a theoretical approach, I studied how mating strategies (monogamy, monoandry, polyandry) can influence the evolution of sex-biased dispersal, with a particular focus on processes linked to relatedness. In monoandry and polyandry, i.e., strategies where both sexes have different numbers of mates, kin competition and a larger spatial variation in male than female reproductive success can lead to a male-biased dispersal. I have also investigated the importance of gene position in the genome and showed a link between dispersal genes and sex chromosomes can produce sex-biased dispersal. I have discussed this evolution of dispersal gene on sex chromosomes under a possible genomic conflict. In parallel to these theoretical studies, I have conducted an experimental study in the common lizard (Zootoca vivipara). I have investigated the role of maternal effect, through the influence of corticosterone, in the relationship between the mating strategy of females and the dispersal strategy of their offspring. I also tested experimentally the transmission of bacteria through mating in relation to the possible cost of transmission of pathogens. While I did not found any direct effect of multiple mating on cloacal bacterial community, my results suggest an effect due to a female response, which opens a new perspective in the study of the consequences of mating strategies.

Doctor of Philosophy / Department of Biology / Theodore J. Morgan / Natural populations are highly complex and consist of genetically variable individuals that belong to continuously varying age classes. Genotype and age interact to determine how individuals respond to environmental stress, which ultimately determines the evolutionary trajectories and persistence of populations in variable environments. For small ectothermic species, seasonal and diurnal variation in temperature is an important source of environmental stress that impacts activity patterns and suites of phenotypes directly related to whole organism fitness. I used the genetic and ecological model Drosophila melanogaster to investigate the influence of seasonal and diurnal thermal variability on survival and reproduction in genetically diverse populations. First, I characterized changes in cold tolerance and phenotypic plasticity within a natural population as it responded to seasonal shifts in developmental and short-term acclimation and thermal selection. I found that seasonal variation in cold tolerance was significantly influenced by developmental acclimation that occurred in the field as well as in the lab, where flies that developed under warmer conditions had reduced cold tolerance relative to flies that developed under cooler conditions. Second, I characterized the effect of variation in age on stress response phenotypes in a genetically variable population. I measured genotype- and age-specific responses to multiple environmental stressors, and identified regions of the genome that were associated with age-specific stress tolerance. Genome-wide association mapping revealed that age-specific phenotypes were influenced by distinct sets of polymorphisms and genes, suggesting that the evolution of age-related decline in phenotypes is driven by mutation accumulation within phenotypes, but both mutation accumulation and antagonistic pleiotropy between phenotypes. Next, I characterized the costs and benefits of acclimation for survival and reproduction to understand how physiological and behavioral plasticity interact to determine fitness. I found that phenotypic plasticity and the capacity for acclimation significantly influenced behavioral reproductive success, but the thermal cues that led to adaptive acclimation response in survival also led to decreased reproductive success. However, genotypes with the capacity to acclimate were more likely to survive thermal variation and more likely to reproduce, suggesting that genetic capacity for phenotypic plasticity has important implications for whole organism fitness. Finally, I measured the effect of acclimation on the induction of diapause and ability to survive cold stress in the recently introduced invasive species Drosophila suzukii. D. suzukii is endemic to Asia and was first detected in California in 2008 and in Topeka, KS in 2013. Its recent invasion history thus provides an interesting model to understand the role of plasiticy in the response to a novel and variable environment. I found that diapause was induced through a plastic response to acclimation and short photoperiod, though diapause was more drastically induced by acclimation. Overall, my research provides critical insights into how organisms respond to thermal variation by intergrating quantitative genetics, ecology, evolution, and life history tradeoffs. Collectively, my research demonstrates that the ability of organisms to survive thermal stress is a function of genetic capacity to tolerate stress, genetic capacity for phenotypic plasticity, prior exposure to thermal variation, and the age of the individual.

Orientador: Gonçalo Amarante Guimarães Pereira / Dissertação (mestrado) - Universidade Estadual de Campinas, Instituto de Biologia / Made available in DSpace on 2018-08-26T17:23:36Z (GMT). No. of bitstreams: 1 Almeida_LudimilaDias_M.pdf: 4371531 bytes, checksum: 15deaceac3a09118222822415b70daaf (MD5) Previous issue date: 2015 / Resumo: O ciclo sexual de basidiomicetos é controlado pelo sistema mating type. Este é formado por dois loci multigênicos não ligados A e B, o locus A codifica duas proteínas homeodomínio HD1 e HD2, capazes de heterodimerização, enquanto o locus B apresenta genes para receptores de feromônio e feromônios. Em fungos heterotálicos, o desenvolvimento sexual depende da especificidade entre os quatro alelos, sistema este chamado tetrapolar, e é ativado apenas por interações específicas entre alelos parentais necessariamente diferentes, assegurando que hifas geneticamente iguais sejam incompatíveis. Em contrapartida, a condição na qual hifas geneticamente iguais são compatíveis é denominada homotalismo. Fungos basidiomicetos são tipicamente heterotálicos, no entanto, apesar de pertencer a este filo, o fitopatógeno Moniliophthora perniciosa, causador da doença Vassoura de Bruxa no cacaueiro, é classificado como homotálico primário. Curiosamente, apesar desta classificação, M. perniciosa contém um sistema genético tetrapolar, sendo o primeiro fungo descrito com essa característica. Neste trabalho, foi realizada a caracterização dos loci mating type em M. perniciosa e verificamos o perfil transcricional destes genes com o objetivo de entender os mecanismos moleculares que atuam no seu comportamento homotálico. Primeiramente, foram identificados no genoma um locus A e um locus B, além de genes atuantes no processamento e sinalização em resposta aos feromônios. O estudo do perfil transcricional destes genes revelou que um receptor tem um perfil de expressão condizente com a fase do ciclo de vida do fungo na qual ocorre o processo de dicariotização. A análise funcional dos receptores foi realizada em um sistema expressão heteróloga, promissor para o estudo de GPCRs (G coupled proteins receptors), porém não permitiu confirmar a presença de alelos compatíveis de receptores e precursores de feromônios no genoma de M. perniciosa como uma possível explicação ao comportamento homotálico. Tendo em vista o locus A, este é formado por um par MpHD1 e MpHD2, o que difere de outros basidiomicetos devido a inserção de uma sequência (11,958kb) interrompendo seus promotores. A hipótese neste cenário é que o transposon encontrado no locus A poderia ter permitido um crossover desigual que trariam genes compatíveis para o mesmo alelo, sendo responsável pelo homotalismo na espécie. Contrariando essa hipótese, os dados obtidos neste projeto indicam que uma possível transição prévia ao homotalismo resultou em uma pressão seletiva relaxada sobre os loci mating type, cuja consequência foi a degeneração nos genes destes loci. Neste contexto, os genes do mating type poderiam não estar mais envolvidos na dicariotização. Este trabalho, portanto, fornece importantes dados para o entendimento da biologia sexual deste fungo, o que futuramente poderá ser correlacionado a sua fitopatogenicidade / Abstract: The basidiomycetes¿ sexual cycle is controlled by the mating type system. The structure of this system comprises two unlinked multigenic loci, A and B. The A locus codes for homeodomain proteins, HD1 e HD2 which form a heterodimer, and B locus presents pheromone receptors and pheromones. In outcrossing (heterothallic) fungi, sexual development depends on the compatibility of four genes in two different allelic versions in a so-called tetrapolar system, and is strictly activated by specific interactions between different parental alleles, ensuring that genetically identical hyphae are incompatible. The phytopathogen Moniliophthora perniciosa causes Witches¿ broom disease in cacao plants, and it is a typical basidiomycete fungi. However, it completes its sexual development through the crossing of genetically identical hyphae, and is the first described homothallic fungi with a complete tetrapolar genetic system. Here we show the characterization of the mating type loci of M. perniciosa and the transcriptional profile of these genes, to uncover the mechanisms underpinning its homothallic behavior. First, we identified an A locus, a B locus and a set of genes that participates in pheromone processing and signalization. Considering the transcriptional profile of these genes, one receptor shows an expression profile consistent with an involvement in dikaryotization. The functional evaluation of the receptors was performed in a heterologous expression system, a promising tool for GPCR (G coupled proteins receptors) proteins study. This system did not allow the confirmation if M. perniciosa contains compatible alleles for receptors and pheromones, one possible explanation for homothallism. Considering A locus, it codes for a pair MpHD1 and MpHD2, which has a sequence insertion (11,958kb) interrupting their promoters, differing from others basidiomycetes. The hypothesis in this scenario is that the insertion of a transposon could have allowed an unequal crossover that brought together compatible genes in the same allele, causing the homothallism in this species. Interestingly, in an opposite direction, our data indicates that a previous transition for homothallism could have resulted in a relaxed selective pressure on mating type loci, with consequences such as the presence of degenerated genes on these loci. In this context, the mating type genes could not necessarily play a role in dikaryotization process. This work provides valuable data for understanding the sexual biology of M. perniciosa, which hereafter could be correlated with its phytopathogenicity / Mestrado / Genetica de Microorganismos / Mestra em Genética e Biologia Molecular

abstract: The present study tested the hypothesis that women dress modestly to signal to other women that they pose no mate poaching threat and are sexually restricted, and that this is especially true for religious women. Participants were 392 Muslim women living in the United States. They read two passages describing fictional situations in which they met with a potential female friend and then indicated what kind of outfit they would wear in both situations. In one situation, the participant obtained a reputation for promiscuity; in the other situation, reputation was not mentioned. I predicted that participants would choose more modest outfits for the promiscuous reputation passage, because if women dress modestly to signal sexual restrictedness, then they should dress more modestly around women with whom they have a reputation for promiscuity—to counteract such a reputation, women may wish to send a strong signal that they are not promiscuous. The hypothesis was partially supported: Less religious women chose more modest outfits for the promiscuous reputation situation than they did for the no reputation situation. This suggests that some women dress modestly to signal sexual restrictedness to other women, but that this is especially true for women who are less religious, not more. More religious women dress more modestly than less religious women, but they may not dress modestly to signal sexual restrictedness. Two important goals for this area of research are to determine the proximate reasons that more religious women dress modestly and to investigate modest dress among non-Muslim religious women. / Dissertation/Thesis / Doctoral Dissertation Psychology 2020

O dimorfismo sexual nas espécies pode surgir a partir da seleção decorrente dos diferentes sistemas reprodutivos. Estudos comportamentais de espécies neotropicais são raros e pouco se sabe sobe as espécies brasileiras. Neste estudo, foram descritos o comportamento de três espécies neotropicais que ocorrem no Cerrado brasileiro: Acanthagrion truncatum, Argia reclusa (Coenagrionidae) e Heaterina rosea (Calopterygidae). Também foi evidenciado o dimorfismo sexual nestas espécies e investigou-se a partir de observações comportamentais, como o dimorfismo se desenvolve em espécies com diferentes táticas reprodutivas. Com os resultados obtidos, vemos que em espécies territoriais os machos são maiores do que as fêmeas, enquanto em espécies não-territoriais as fêmeas são maiores do que os machos. Sugere-se que, diferentemente de outros estudos, em Zygoptera o tipo de sistema reprodutivo pode determinar o dimorfismo sexual. / Sexual size dimorphism (SSD) can in some species result from the selection acting through different mating systems. Behavioral studies of neotropical species are rare, and few is known about the brazilian species. In this study, we described the behavior of three neotropical species that occur in the brazilian neotropical savannah: Acanthagrion truncatum, Argia reclusa (Coenagrionidae) and Heaterina rosea (Calopterygidae). We show the SSD in these species and investigates through behavioral observations, how SSD develops in species with different mating tactics. With our results, we can see that in territorial species the males are larger than females, while in non-territorial species the females are larger than males. We suggest that, unlike other studies, in Zygoptera the kind of mating system adopted by males may determinate the SSD in a species.

Among multi-male, multi-female primate groups, males engage in direct contest competition for access to mates. The priority-of-access model (PoA model) generally predicts that male reproductive success increases with male dominance rank, but the strength of this relationship is expected to decrease with increasing female reproductive synchrony, particularly in seasonally breeding primates. Genetic paternity studies support the model’s predictions, having found a positive relationship between male dominance rank and reproductive success. However, in addition to dominance status and female reproductive synchrony, a number of proximate factors also impact males’ ability to sire offspring, which have not been considered in studies of male reproductive strategies.By integrating behavioral, genetic, morphological, and hormonal analysis as more direct measures of reproductive success in individual males, this dissertation investigated the relationship between male dominance rank and reproductive success and including the proximate factors affecting this relationship in wild northern pig-tailed macaques (Macaca leonina). The main objectives of this study are: 1) to identify the relationships between male dominance rank, male mating success, and male reproductive success, and assess to what extent female synchrony affects these relationships; 2) to identify the proximate factors that may reduce the reproductive success of the top-ranking male and assess variation in male mating tactics related to dominance rank and migration status (i.e., resident males vs. extra-group males); and 3) to evaluate the function of males’ red ornaments that may be used to signal male dominance status (male-male competition) to attract females (female mate choice) or both. The study was conducted at Khao Yai National Park, northeastern Thailand. Systematic data collection on CH group occurred from September 2015-June 2017. The group composition was recorded daily as well as births, deaths, individual emigrations and immigrations, females’ parity status, and the presence of extra-group males (EGMs). Sociosexual data and male-female interactions (i.e., copulations, ejaculatory copulations, consortships, grooming, female proceptive behaviors and receptive behaviors) were recorded during females’ receptive periods. To assess male reproductive success, genetic paternity analyses were conducted on fecal DNA samples collected from 18 adult and subadult males, 22 adult females, and 25 juveniles and infants. To measure red skin coloration of males, hindquarter images were collected non-invasively for seven adult males. From those images, skin color and luminance were computationally quantified to assess variation in male anogenital reflectance. Lastly, fecal samples were collected from nine adult males to assess monthly levels of fecal testosterone by microtitreplate enzyme immunoassay. The distribution of births and matings suggested that northern pig-tailed macaques, at least in this group, are best categorized as moderate seasonal breeders. Indeed, 33-67% of births occurred within a three-month period. Copulation data revealed a positive relationship between male dominance rank and mating success, supporting the predictions of the PoA model. However, the distribution of male reproductive success indicated that: 1) high-ranking males controlled a proportion of paternity much lower than predicted by the PoA model; 2) middle-ranking males controlled a proportion of paternity higher than predicted by the model; and 3) EGMs, not considered in the PoA model, controlled a surprisingly large proportion of paternity despite a low observed mating success. When females were simultaneously receptive, lower-ranking and subadult males engaged in opportunistic and surreptitious copulation and avoided direct competition with higher-ranking males, and most females approached and mated with EGMs out of the resident adult males’ sight. However, one EGM also mated in full sight of resident adult males. This is the first study to report mating and successful paternity by EGMs in a moderately seasonally breeding species.Four male mating tactics were identified: 1) the top-ranking resident male tactic, in which the male competes for the alpha male position to control priority of access to receptive females through long consortships and copulations; 2) the lower-ranking resident male tactic, in which the male copulates opportunistically and surreptitiously out of sight of higher-ranking males mostly during the mating peak; 3) the subordinate EGM tactic, in which the male lives semi-solitarily and copulates opportunistically and surreptitiously, mostly during the mating peaks; and 4) the super-dominant EGM tactic, in which the EGM copulates irrespective of the presence of other males and in full sight of even the highest-ranking resident male. In addition, I found support for female mate choice. Darker and redder males had more mating partners, received more female proceptive behaviors, and were engaged in more consortships and grooming with receptive females. Furthermore, males became redder and darker as female reproductive synchrony increased. Together, these results suggest that male red ornaments exhibited in the male’s anogenital area is attractive to females. Furthermore, behavioral evidence of female mate choice towards EGMs was found. This dissertation provides a comprehensive picture of the complex male mating tactics of northern pig-tailed macaques. To achieve reproductive success, males engage in a diversity of mating tactics, strongly influenced by male dominance rank and the degree of female reproductive synchrony. However, mate-guarding costs, surreptitious copulations by lower-ranking males and EGMs, and female mate choice, need to be included in an extended version of the PoA model to provide stronger predictions of the distribution of male reproductive success in primates.

No description available.

Mixed-ploidy populations, consisting of multiple cytotypes, are an optimal system for studying genome doubling consequences in plants. The role of frequency dependent selection, known as minority cytotype exclusion principle, is very important in them, but there are many factors limiting this selection. In my thesis, I evaluate the changes in cytotype frequencies and pattern in permanent plots in natural mixed-ploidy populations of three plant species - Butomus umbellatus, Knautia serpentinicola and Tripleurospermum inodorum and I also experimentally analyze partial aspects of frequency dependent selection towards minority cytotype in artificially established mixed-ploidy arrays of two plant species - Arabidopsis arenosa and Tripleurospermum inodorum. Varied changes in frequencies of minority cytotype ware revealed in permanent plots. Usually there was decrease, but in two plots there was increase in minority cytotype frequency and in one case it became dominant. Observed changes depend on initial frequencies of minority cytotype in populations, biological properties of species and particular environment of permanent plot. Relative indexes describing the frequency and the strength of disturbances and also the amount of soil nutrients was used to compare the environment of permanent plots across...

To understand animal behaviour, it is important to consider the environment in which it occurs. The environment, consisting of both abiotic factors and social context, is usually highly variable and leads to variation in individual’s and group’s behaviour. To better understand the environmental influences on behaviour of pygmy halfbeaks (Dermogenys collettei), a small live-bearing fish, I viewed videos of shoals of wild halfbeaks in Singapore. I investigated effects of environmental variation (water depth, canopy cover and water vegetation) and social environment (group size and male to female sex ratio) on halfbeaks’ aggression and mating behaviours. I found that environment had little effect and most variation between studied shoals was probably due to social factors. I found some evidence for aggression increase in larger shoals, primarily in males. Sex ratio had different relation with aggression for individual sexes and mating behaviours decreased when sex ratio became more male biased. This study shows that halfbeaks probably modify their behaviour in relation to social environment. I also show that these changes are similar to those described in other species, which strengthens the validity of using halfbeaks to study social interactions.