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.

No description available.

Genetics has been identified as a subject area which many students find difficult to comprehend. The researcher, who is also a lecturer at the University of KwaZulu-Natal, had noted over a number of years that students find the field of quantitative genetics particularly challenging. The aim of this investigation was two-fold. Firstly, during the diagnostic phase of the investigation, to obtain empirical evidence on the nature of difficulties and alternative conceptions that may be experienced by some students in the context of quantitative genetics. Secondly, to develop, implement and assess an intervention during the remediation phase of the study which could address the identified difficulties and alternative conceptions. The research was conducted from a human constructivist perspective using an action research approach. A mixed-method, pragmatic paradigm was employed. The study was conducted at the University of KwaZulu-Natal over four years and involved third-year students studying introductory modules in quantitative genetics. Empirical evidence of students' conceptual frameworks, student difficulties and alternative conceptions was obtained during the diagnostic phase using five research instruments. These included: free-response probes, multiple-choice diagnostic tests, student-generated concept maps, a word association study and student interviews. Data were collected, at the start and completion of the modules, to ascertain the status of students' prior knowledge (prior knowledge concepts), and what they had learnt during the teaching of the module (quantitative genetics concepts). Student-generated concept maps and student interviews were used to determine whether students were able to integrate their knowledge and link key concepts of quantitative genetics. This initial analysis indicated that many students had difficulty integrating their knowledge of variance and heritability, and could not apply their knowledge of quantitative genetics to the solution of practical problems. Multiple-choice diagnostic tests and interviews with selected students were used to gather data on student difficulties and alternative conceptions. The results suggested that students held five primary difficulties or alternative conceptions with respect to prior knowledge concepts: (1) confusion between the terms variation and variance; (2) inappropriate association of heterozygosity with variation in a population; (3) inappropriate association of variation with change; (4) inappropriate association of equilibrium with inbred populations and with values of zero and one; and, (5) difficulty relating descriptive statistics to graphs of a normal distribution. Furthermore, three major difficulties were detected with respect to students understanding of quantitative genetics concepts: (1) students frequently confused individual and population measures such as breeding value and heritability; (2) students confused the terms heritability and inheritance; and, (3) students were not able to link descriptive statistics such as variance and heritability to histograms. Students found the concepts of variance and heritability to be particularly challenging. A synthesis of the results obtained from the diagnostic phase indicated that many of the difficulties and alternative conceptions noted were due to confusion between certain terms and topics and that students had difficulty with the construction and interpretation of histograms. These results were used to develop a model of the possible source of students' difficulties. It was hypothesized and found that the sequence in which concepts are introduced to students at many South African universities could be responsible for difficulties and alternative conceptions identified during the study, particularly the inappropriate association of terms or topics. An intervention was developed to address the identified difficulties and alternative conceptions. This intervention consisted of a series of computer-based tutorials and concept mapping exercises. The intervention was then implemented throughout a third year introductory module in quantitative genetics. The effectiveness of the intervention was assessed using the multiple-choice diagnostic tests and interview protocols developed during the diagnostic phase. The knowledge of the student group who participated in the intervention (test group) was compared against a student group from the previous year that had only been exposed to conventional teaching strategies (control group). t-tests, an analysis of covariance and a regression analysis all indicated that the intervention had been effective. Furthermore, an inductive analysis of the student responses indicted that most students understanding of the concepts of variance, heritability and histograms was greatly improved. The concept maps generated by students during the remediation phase, and data from the student interviews, provided an indication of the nature and extent of the conceptual change which had occurred during the teaching of the module. The results showed that most of the conceptual change could be classified as conceptual development or conceptual capture and not conceptual exchange. Furthermore, it seemed that conceptual change had occurred when considered from an epistemological, ontological and affective perspective, with most students indicating that they felt they had benefited from all aspects of the intervention. The findings of this research strongly suggest an urgent need to redesign quantitative genetics course curricula. Cognisance should be taken of both the sequence and the manner in which key concepts are taught in order to enhance students' understanding of this highly cognitively demanding area of genetics. / Thesis (Ph.D.)-University of KwaZulu-Natal, Pietermaritzburg, 2006.

The independent evolution of the sexes may often be constrained if male and female homologous traits share a similar genetic architecture. Thus, cross-sex genetic covariance is assumed to play a key role in the evolution of sexual dimorphism (SD) with consequent impacts on sexual selection, population dynamics and the speciation process. I used quantitative genetics tools to assess the importance of sex-specific genetic variance in facilitating the evolution of body mass and horn size SD in wild bighorn sheep from Ram Mountain, Alberta. I also developed a bighorn sheep genetic linkage map composed of 247 microsatellite markers to gain insights about the genetic architecture of trait variation. Finally, I conducted systematic reviews and meta-analyses of published cross-sex genetic correlations (rMF, a standardized estimate of cross-sex genetic covariance) to test basic hypotheses about the importance of sex-specific genetic variance in the evolution of SD and mechanisms responsible for generating such variance. My results demonstrated that sex-specific genetic variance was present in bighorn sheep and that it likely played an important role in alleviating intralocus sexual conflicts. The quantitative trait locus (QTL) mapping analysis resulted in the identification of numerous loci influencing body mass and horn dimensions, some of which had apparent sex-specific effects. An analysis of 553 rMF estimates recovered from 114 published sources allowed demonstrating that 1) the evolution of SD was generally constrained by positive cross-sex genetic covariance, 2) levels of SD were often sub-optimal, and 3) sex-specific genetic variance was an important mechanism allowing the evolution of SD. In addition, I confirmed the long-standing hypothesis of a general decline in rMF with age. Sexual dimorphism is an important evolutionary phenomenon, but our understanding of its evolution is still limited. After decades of speculation, my research has provided clear empirical evidence for the importance of sex-specific genetic variance in allowing its evolution. / Ecology

Darwin's (1859) theory of evolution by natural selection accounts for the adaptations of organisms, but, as Fisher (1930) famously said, 'natural selection is not evolution.' Evolutionary theory has two major components: i) natural selection, which involves the underlying dynamics of populations; and ii) adaptive evolutionary change, which involves the optimisation of phenotypes for fitness maximisation. Many of the traditional theoretical frameworks in evolutionary theory have focussed on studying optimisation processes that generate biological adaptations. In recent years, however, a number of evolutionary theorists have turned to using frameworks such as the 'replicator dynamics' or 'eco-evolutionary dynamics', to explore the dynamics of natural selection. There has, however, been little attempt to explore how these dynamical systems frameworks relate to more traditional frameworks in evolutionary theory or how they incorporate the principles that embody the process of evolution by natural selection, namely, phenotypic variation, differential reproductive success, and heritability. In this thesis, I use these principles to provide the formal foundations of a general framework - a mathematical synthesis - in which the future state of an evolutionary system can be predicted from its present state; what I will call a 'dynamical systems theory of natural selection.' Given the state of an existing biological system, and a set of assumptions about how individuals within the system interact, the job of the dynamical systems theory of natural selection is no less than to predict the future in its entirety.

Orientador: Mário Luiz Teixeira de Moraes / Banca: Marco Eustáquio de Sá / Banca: Miguel Luiz Menezes Freitas / Banca: Edson Seizo Mori / Banca: Ananda Virgínia de Aguiar / Resumo: A aroeira Myracrodruon urundeuva (F.F. & M.F. Allemão) é conhecida pela extraordinária durabilidade e resistência de sua madeira, possui reconhecido valor econômico e apresenta diversas aplicações, entre elas, a confecção de esteios, postes moirões e dormentes, extração de tanino e bálsamo. Seis populações da espécie estão sendo conservadas ex situ em delineamentos experimentais na Fazenda de Ensino, Pesquisa e Extensão da FEIS/UNESP, em Selvíria-MS, onde se pretende ampliar a base genética, recombinar a variabilidade genética e produzir sementes para reflorestamentos. Das seis populações duas são procedentes de área com pouca perturbação antrópica: Estação Ecológica do Instituto Florestal em Paulo de Faria-SP e Estação Ecológica do Seridó em Serra Negra do Norte-RN. As outras quatro são de área com forte perturbação antrópica: Bauru-SP, Itarumã-GO, Petrolina-PE e Selvíria-MS. Dessa forma, foram avaliados nove testes de progênies de M. urundeuva para caracteres quantitativos empregando-se a metodologia REML/BLUP (máxima verossimilhança restrita / melhor predição linear não viciada), assumindo que as progênies são parentes no grau de meios-irmãos e a metodologia MHPRVG para a análise de adaptabilidade e estabilidade nos testes de progênies provenientes da Estação Ecológica de Paulo de Faria em quatro sistemas de plantio (ambientes) diferentes. Este trabalho permitiu estudar os caracteres de desenvolvimento silvicultura nos testes de progênies, a variabilidade genética, o ganho de seleção a partir do Índice Multiefeito, a propagação sexuada e assexuada, a sobreposição de geração e a estabilidade e adaptabilidade das progênies de M. urundeuva. Verificou-se que as progênies de M. urundeuva em relação aos caracteres silviculturais analisados: altura total, diâmetro médio da copa (DMC), diâmetro à altura do peito (DAP)... (Resumo completo, clicar acesso eletrônico abaixo) / Abstract: The aroeira Myracrodruon urundeuva (F.F. & M.F. Allemão) has been known for the extraordinary durability and resistance of its wood, it has recognized economic value and presents several uses, as for, posts, and sleepers, tannin extraction, and balm. Six populations of the species has been conserved by ex situ design in Experimental Station of Education, Research, and Extension of FEIS/UNESP, in Selvíria-MS Brazil, where to intend to enlarge the genetic base, mix up the genetic variability and to produce seeds for reforestations. Two out of six populations are coming from sites with little anthopic disturbance: Ecological Station of the Instituto Forestal in Paulo de Faria-SP, Brazil and Ecological Station of Seridó in Serra Negra- RN, Brazil. The others four are from sites with large anthopic disturbance: Bauru-SP, Itarumã-GO, Petrolina-PE and Selvíria-bad. In that way, were evaluated nine progeny trial of M. urundeuva to study the traits using the methodology of REML/BLUP (máxima verossimilhança restrita/melhor predição linear não viciada), stating that the are half site and the MHPRVG methodology for the adaptability and stability analyses by trials from Ecological Station of Instituto Florestal of Paulo de Faria by four different plantation systems (atmospheres). This research to study the traits of development forestation in the progeny trials, the genetic variability, the selection gain starting by Index Multi-effects', the propagation sexual and assexual, Superiority of elite trees in relationship of the progenies and the stability and adaptability of the progenies of M. urundeuva. Verified that the progenies of M. urundeuva in relation to the traits analyzed silviculturais: total height, medium diameter of the cup (DMC), diameter to the height of the chest (DAP) they had a satisfactory development, and the progenies originating from of the Ecological Station... (Complete abstract click electronic access below) / Doutor

Dentro da biologia evolutiva, uma das questões centrais é a compreensão de como os processos evolutivos, em particular a seleção e o acaso (deriva genética) moldaram a diversidade obervada nos organismos. Dentro deste contexto, a utilização de abordagens como a integração morfológica e a genética quantitativa nos fornecem poderosas ferramentas. Enquanto a primeira descreve os padrões de relação entre caracteres e testa hipóteses sobre as relações de desenvolvimento e/ou funções subjacentes, a segunda possui ferramentas para investigar as forças que podem ter gerado os fenótipos atuais. Utilizando como modelo de estudo os gêneros de marsupiais da Ordem Didelphimorphia, analisei a evolução morfológica craniana unindo estas duas linhas de pesquisa. Central a genética quantitativa está a matriz de covariância genética (G) que descreve a porção da variância que é efetivamente herdada (no sentido de transmissão de valor entre gerações) e conseqüentemente serve como substrato à seleção. Apesar de ter sido desenvolvida inicialmente para estudos em escala microevolutiva, o arsenal matemático da genética quantitativa pode ser estendida a escalas macroevolutivas caso a matriz G permaneça relativamente estável. Entretanto, como a estimativa de uma matriz G requer um número grande de espécimes aparentados e com genealogia conhecida, utilizei ao longo deste trabalho sua correspondente fenotípica (P). Desta forma, no primeiro capítulo demonstrei a similaridade das matrizes de covariância e correlação P dos marsupiais Didelphimorphia. Em contrapartida, as magnitudes de integração morfológica, que medem o grau de associação entre os caracteres, se mostraram mais variáveis dentro do grupo. A história evolutiva (filogenia) e as distâncias morfológicas entre os gêneros não parecem ter tido influência sobre os padrões nem as magnitudes, já que não se correlacionaram significativamente a eles. No segundo capítulo, estendi estas análises para os gêneros ao longo do seu desenvolvimento e novamente os padrões se mostraram semelhantes, tanto quando comparei a ontogenia de um único gênero, quando analisei diferentes gêneros para diferentes classes etárias. Em contrapartida, as magnitudes de integração se mostraram mais variáveis, com uma tendência a diminuírem com o aumento da idade. Por conta destes resultados, comparei as matrizes de correlação empíricas com matrizes teóricas que expressam hipóteses de modularidade baseadas no desenvolvimento/função compartilhada pelas regiões do crânio. Testei para similaridade das duas regiões principais (face e neurocrânio), cinco sub-regiões (base e abóboda craniana, oral, nasal e zigomático), uma matriz 198 de integração somática (caracteres neurais vs. faciais) e uma de integração total, concatenando as cinco sub-regiões. Foram encontradas somente correlações significativas para as regiões da face e suas sub-regiões nasal e oral. Apesar das variações nas magnitudes de integração observadas nestes dois primeiros capítulos, as respostas evolutivas obtidas para estes gêneros foram todas muito similares. Este resultado, associado a constatação de que grande parte da variação entre os espécimes é devida ao tamanho, ou variações de forma associadas a ele (alometria), me levaram a analisar as conseqüências evolutivas associadas a estas variações alométricas. Assim, no terceiro capítulo analisei as direções de resposta à seleção simulada antes e após a remoção do tamanho. Variações alométricas exercem forte influência sobre estes crânios tornando-os muito integrados morfologicamente, ou seja, a percepção dos módulos que os compõem é muito difícil (baixa modularidade). Isto se reflete em uma maior variação ao longo da linha de menor resistência evolutiva (que resume o eixo de maior variação disponível entre os espécimes), já que os módulos não podem responder a seleção de forma independente (alta integração morfológica). Em marsupiais, esta linha é está alinhada na direção de variações de tamanho. Desta forma, não importa em que direção a seleção esteja atuando, as respostas evolutivas serão usualmente na direção desta linha de menor resistência e conseqüentemente de tamanho. Uma vez que o tamanho é removido, as magnitudes de integração diminuem e a modularidade do crânio aumenta. Por conseguinte, ele passa a ser capaz de responder à seleção em uma gama maior de direções porque aumenta também a relativa independência destes módulos em relação aos demais. Como estes resultados indicam que variações entre os Didelphimorphia são direcionadas a variações no tamanho, devido justamente às restrições impostas pela linha de menor resistência evolutiva, analisei as trajetórias ontegenéticas de dois gêneros irmãos, Didelphis e Philander, separando os efeitos da forma e tamanho nesta diversificação. Para tanto empreguei análises de morfometria tradicional, morfométrica e análises de coeficientes alométricos que apontaram para uma maior diferenciação no tamanho entre eles. Apesar da similaridade entre as trajetórias ontogenéticas destes dois gêneros, espécimes de Didelphis nascem maiores e crescem por mais tempo, como indicado pela sua trajetória mais longa. / One of the central goals in evolutionary biology is related to how evolutionary processes, mainly natural selection and genetic drift, shaped living organisms. The combined use of Morphologial Integration and Quantitative Genetics give us powerful tools to accomplish this goal. Morphological Integration is concerned of how characters are related, as well as their underlying genetics/developmental relationship, while Quantitative Genetics have methodologies designed to explore the phenotypic forces underlying diversity among organisms. Using Didelphimorphia marsupials as a study group, I combined these two approaches to study its morphologic cranial diversification. In Quantitative Genetics, the genetic additive covariance matrix (G) resume the genetic variation underlying resemblance among relatives, which is the portion of the variance responsive to selection. Initially developed to microevolutionary scale studies, it can be extended to a macroevolutionary scale if it remains relatively similar in that time scale. However, as G matrix estimations require a huge number of related specimens with known genealogy, I used its phenotypic (P) counterpart which was more easily obtained. In the first chapter I showed high similarities among Didelphimorphia marsupials covariance and correlation P matrices. On the other hand, integration magnitudes (which measure the average correlation among traits) vary among taxa. Neither phylogeny nor morphologic distances showed any association with the similarity in patterns and magnitudes of integration. In the second chapter, I did these same analyses, but throughout genera ontogeny. Again, there was a high similarity among taxa in patterns of integration, both when I analyzed the ontogeny for each genus separately or against each other (at different age class). Morphological integration magnitudes showed the same variation obtained for adults, with a tendency to decrease at older ages. Taking these results into account, I compared the phenotypic correlation matrices to theoretical matrices, based on hypotheses of shared developmental and functional units. I searched for modularity in the two main skull regions (face and neurocranium) and five sub regions (cranium base and vault, face, nasal, and oral). I also looked for modularity concerning somatic development (Neurocranium vs. Face) and total modularity, as the 200 summation of the five sub-regions. Only Face and its sub-regions nasal and oral, showed significant correlations to the phenotypic genera matrices. Despite integration magnitude differences, all evolutionary responses produced by taxa were highly similar. These results, combined with a huge size variation (or size related variation - allometry) across taxa, lead me to search for the evolutionary consequences due to size variation. In the third chapter, I compared evolutionary response directions produce by each genera matrices before and after size removal under a random selection simulation. Allometry strongly affect these skulls, turning them into highly integrated structures with lower modularity (skull modules are not easily distinguished). Because of this, modules cannot evolve relatively independent of other modules and evolutionary responses will strongly affect the whole cranium. This is related to the variation along the lines of least evolutionary resistance. This line is the multivariate direction of greatest genetic or phenotypic variation (the combination of a suite of traits that displays the maximum within-population variance). In Didelphimorphia marsupials, this line is aligned with size variation and regardless the selection direction, evolutionary change is usually aligned to this least resistance line. The removal of size variation diminish the magnitude of integration while increases modularity. Consequently, skulls become able to respond to selection in more directions as modules become relatively more independent of each other. In the last chapter I compared size and shape differences between ontogenetic trajectories of two sister genera Didelphis and Philander. Using traditional and geometric morphometric analysis plus allometric coefficient analysis, I could show that bigger differences between them are size related. Despite similarities, ontogenetic trajectory in Didelphis is longer, leading to bigger specimens.

O principal objetivo deste estudo foi investigar os processos responsáveis pela evolução morfológica craniana de primatas da subordem Strepsirrhini, com maior ênfase ao clado endêmico à ilha de Madagascar. Sistemas multidimensionais morfológicos como o crânio descrevem a quantidade de variação disponível para atuação de processos evolutivos. A partir de 27 marcos anatômicos cranianos e 39 distâncias euclidianas estimei matrizes de variância e covariância fenotípicas (Matrizes P) para 40 espécies das sete famílias viventes de Strepsirrhini e duas espécies do gênero Tarsius. Utilizei a abordagem da genética quantitativa para investigar o padrão de variação nas matrizes P dessa linhagem de primatas, conjuntamente com um par de matrizes fenotípica e genética de Saguinus fuscicolis. Para investigar a estabilidade e conservção do padrão da estrutura de variação ao longo da diversidade deste clado de primatas compararei estas matrizes através de dois métodos complementares que me permitiram avaliar a distribuição da variação no morfoespaço (Random Skewers) e o compartilhamento de direções de variação (Krzanowski) das matrizes. Os resultados obtidos sugerem uma estabilidade das matrizes, possivelmente mantida por seleção estabilizadora interna comum ao clado, aos demais primatas e mamíferos em geral. Esses resultados de comparação de matrizes são apresentados no Capítulo 1 e corroboram a conjectura de Cheverud, indicando que as matrizes cumprem o pressuposto de similaridade estrutural entre as matrizes P e a matriz G. Além disso o padrão de variação apresentou um sinal filogenético e algumas das diferenças observadas nas matrizes parecem estar relacionadas com fatores de dieta. No Capítulo 2 investiguei a evolução da divergência morfológica craniana em um conjunto de 70 espécies, onde testei a hipótese nula de deriva genética como explicação suficiente para a divergencia entre as médias das espécies. Esta hipótese foi avaliada dentro das expectativas de modelos da genética quantitativa através de duas metodologias. Tais testes avaliam a proporcionalidade de variação dentro (W) e entre grupos (B) sob perspectivas complementares. Com o teste de regressão investiguei a manutenção da quantidade de variação associada a cada eixo de variação, e com o teste de correlação avaliei o alinhamento entre estes eixos. Rejeitamos a H_0 em diversos pontos da árvore que representam eventos de cladogenese a nível de família e gênero. Nestes casos aceitamos a H_1 de seleção natural e vemos que a maior parte da diversificação da linhagem de primatas de Madagascar foi influenciada por seleção direcional envolvendo tamanho e alterações anatômicas relacionadas a dieta / The main goal of this study was to investigate the evolutionary processes responsible for Strepsirrhini cranial morphological evolution. Multidimensional morphological structures like the cranium can describe the amount of variance available to evolution. We used 27 landmarks and 39 euclidean distances between them to describe the variation in the cranium. We used a broad phylogenetic scaled comparison of cranial P matrices representing 40 species of Strepsirrhini primates combined to a pair of Saguinus fuscicolis G and P to investigate the stability of variance structure along this lineage. Our results show a relative stability in the patterns of variance and covariance of the clade and that P matrices can be considered as surrogates to its underling G. We also report a high association between matrix similarity and phylogenetic distance. We investigated particular contributions of each trait to matrix dissimilarity through a evolutionary perspective considering the main dietary shifts observed in the clade. Our results suggest that differences in observed patterns of variation can be attributed to characters with heterogeneity in the degree of stabilizing selection in the adult cranium and to differences in directional selection involved in chewing of specialized feeding behavior. Under the quantitative genetics theory lies an expectation for the evolution of average phenotypes that if populations have diverged by random processes, patterns of within and between-taxon morphological variation should be proportional. In chapter 2 we tested the null hypothesis that genetic drift is a sufficient explanation to observed divergence in cranial multivariate means of 70 Strepsirrhini primates. We detected deviations from neutrality along the whole clade of Strepsirrhini and specially in more inclusive nodes. We argue that this deviations reflect important historical shifts in the evolution of the clade associated with directional selection for size and in anatomical features related with diet

No presente trabalho buscou-se introduzir o efeito da competição entre plantas nas análises dos testes de progênie/procedências em essências florestais, com o fim de identificar os seus efeitos e as distorções devidas à sua não observância. Para tanto, foram utilizados ensaios com níveis de precisão e mortalidades diferentes, de cinco espécies, a saber: Gallesia gorarema Vell. Moq., Eucaliptus grandis Hill ex Maider, Eucaliptus citridora Hook, Pinus elliottii Engl. var. elliottii e Araucaria angustifolia (Bert.) O. Ktze. Obtiveram-se as esperanças dos quadrados médios das fontes de variação da análise de variância nos delineamentos aqui utilizados. Com base nestas derivações, foi demonstrado explicitamente o viés nas estimativas de parâmetros genéticos quantitativos. Este viés está diretamente relacionado com a magnitude do coeficiente de regressão b e com a grandeza relativa das somas de quadrados de diferentes efeitos contidos na análise de variância da variável competição. Caso ignorado o efeito de competição, quando este influencia a variável resposta Y, os ponderadores b, que compõem o índice de seleção terão estimativas viesadas, gerando erro na seleção dos indivíduos superiores. Na análise de dados observou-se que a inclusão da competição, de maneira geral, reduziu as estimativas das componentes de variância, e por conseqüência, outras estimativas de parâmetros que são função destes, quando comparado com as estimativas feitas por via das análises sem o ajuste para a competição. A análise com a variável competição não mostrou diferenças significativas para o efeito de progênies. Isto demostra que a competição comportou-se de forma aleatória, o que corrobora para que seja colocada na análise como uma covariável; caso contrário esta teria que ser considerada uma componente da performance e introduzida numa análise multicaracterística. Utilizando as análises com e sem ajuste para a competição, para estimar os valores genéticos e o ganho com a seleção, observou-se que os indivíduos selecionados não são concordantes. Isto indica que os equívocos na seleção podem ser comuns, haja vista que o fato de se ajustar os dados faz com que o posto dos indivíduos tidos por superiores seja alterado. É recomendável considerar os efeitos da competição na análise de dados em que os indivíduos estão sujeitos a competir uns com os outros, no seu desenvolvimento. / The aim of this work was to introduce competition effects in the model underlying the analysis of forest tree experiments. Results were compared with analyses in which effects were neglected. Progeny trails with different levels of precision and mortality were used, including the following species: Gallesia gorarema Vell. Moq., Eucaliptus grandis Hill ex Maider, Eucaliptus citridora Hook, Pinus elliottii Engl. var. elliottii and Araucaria angustifolia (Bert.) O. Ktze. Mathematical expectation of mean squares values were derived and the bias of estimates was explicitly shown. Competition effects were found significant in all experiments, but were primarily of random nature. Bias was shown to be directly proportional to the magnitude of the regression parameter b and to the relative magnitude of sums of squares of the competition variable. Including the variable in general lead to a reduction of estimates of variance components and to smaller expected progress from selection. The b coefficients of multi-effect selection index are also biased if competition is ignored. Results indicated that different sets of genotypes could be selected if the analyses of data were carried out with or without the competition effects. Including a competition variable in the analysis of trials in which plants are exposed to competing with each other is recommendable.

O mapeamento de QTL's difere dos demais tipos de pesquisas conduzida em genética. Por se tratar basicamente de um procedimento de testes múltiplos, surge, neste contexto, um problema que se refere ao nível de significância conjunto da análise, e consequentemente, seu poder. Deste modo, avaliou-se, via simulação computacional de dados, o poder de detecção de QTL's da análise de marcas simples, realizada por meio de regressão linear múltipla, utilizando o procedimento stepwise" para seleção das marcas e procedimentos baseados em testes individuais, utilizando os critérios FDR e de Bonferroni para determinação nível de significância conjunto. Os resultados mostraram que o procedimento baseado em regressão múltipla, utilizando o procedimento stepwise" foi mais poderoso em identificar as marcas associadas a QTL's e, mesmo nos casos em que este procedimento apresentou poder ligeiramente inferior aos demais, verificou-se que o mesmo tem como grande vantagem selecionar apenas as marcas mais fortemente ligadas aos QTL's. Dentre os critérios FDR e de Bonferroni, o primeiro mostrou-se, em geral, mais poderoso, devendo ser adotado nos procedimentos de mapeamento por intervalo. Outro problema encontrado na análise de QTL's refere-se µa abordagem da interação QTL's x ambientes. Neste contexto, apresentou-se uma partição da variância da interação genótipos x ambientes em efeitos explicados pelos marcadores e desvios, a partir da qual obtiveram-se os estimadores da proporção da variância genética (pm), e da variância da interação genótipos x ambientes (pms), explicadas pelos marcadores moleculares. Estes estimadores independem de desvios das frequências alélicas dos marcadores em relação µ as esperadas (1:2:1 em uma geração F2, 1:1 em um retrocruzamento, etc.), porém, apresentam uma alta probabilidade de obtenção de estimativas fora do intervalo paramétrico, principalmente para valores elevados destas proporções. Contudo, estas probabilidades podem ser reduzidas com o aumento do número de repetições e/ou de ambientes nos quais as progênies são avaliadas. A partir de um conjunto de dados de produtividade de grãos, referentes µ a avaliação de 68 progênies de milho, genotipadas para 77 marcadores moleculares codominantes e avaliadas em quatro ambientes, verificou-se que as metodologias apresentadas permitiram estimar as proporções pm e pms, bem como classificar as marcas associadas a QTL's, conforme seu nível de interação. O procedimento permitiu ainda a identificação de regiões cromossômicas envolvidas no controle genético do caractere sob estudo conforme sua maior ou menor estabilidade ao longo dos ambientes. / In general terms, QTL mapping di®ers from other research ac-tivities in genetics. Being basically a multiple test procedure, problems arise which are related to the joint level of signi¯cance of the analysis, and consequently, to its power. Using computational simulation of data, the power of simple marker analysis, carried out through multiple linear regression, using stepwise procedures to select the markers was obtained. Procedures based on single tests, using both the FDR and the Bonferroni criteria to determinate the joint level of signi¯cance were also used. Results showed that the procedure based on multiple regression, using the stepwise technique, was the most powerful in identifying markers associated to QTL's. However, in cases where its power was smaller, its advantage was the ability to detect only markers strongly associates with QTL's. In comparision with the Bonferroni method, the FDR criterion was in general more powerful, and should be adopted in the interval mapping procedures. Additional problems found in the QTL analysis refer to the QTL x environment interaction. We consider this aspect by par-titioning the genotype x environment interaction variance in components explained by the molecular markers and deviations. This alowed estimating the proportion of the genetic variance (pm), and genotype x environment variance (pms), explained by the markers. These estimators are not a®ected by deviations of allelic frequencies of the markers in relation to the expected values (1:2:1 in a F2 generation, 1:1 in a backcross , etc). However, there is a high probability of obtaining estimates out of the parametric range, specially for high values of this proportion. Nevertheless, these probabilities can be reduced by increasing the number of replications and/or environments where the progenies are evaluated. Based on a set of grain yield data, obtained from the evaluation of 68 maize progenies genotyped for 77 codominant molecular markers, and evaluated as top crosses in four environments, the presented methodologies allowed estimating proportions pm and pms as well the classification of markers associated to QTL's, with respect to its level of genotype x environment interaction. The procedure also allowed the identification of chromosomic regions, involved in the genetical control of the considered trait, according to its stability, in relation to the observed environmental variation.