Diversas pesquisas têm reportado que o aumento da produtividade da cultura do milho se deve muito mais à tolerância a estresses abióticos do que a aumentos per se. O déficit hídrico é um dos mais importantes fatores envolvidos na redução de produtividade nessa cultura, mas devido à dificuldade de seleção direta para tolerância à seca, são empregados caracteres secundários como o caráter stay green ou senescência retardada de folhas e colmos. Todavia, estudos sobre a herança desse caráter são escassos na literatura. Em vista disso, este trabalho teve como objetivo estudar a herança deste caráter em milho tropical, utilizando o delineamento III (Comstock e Robinson, 1952). A partir do cruzamento entre as linhagens endogâmicas L08-05F e L38-05D, foram obtidas 100 progênies F2:3, as quais foram retrocruzadas com cada um dos parentais, resultando em 200 progênies de retrocruzamento. Estas progênies foram avaliadas em dez ambientes, com duas repetições por ambiente, em Piracicaba SP, utilizando o delineamento ?-látice. A avaliação do stay green foi feita em dez plantas competitivas por parcela, 120 dias após a semeadura, utilizando uma escala de notas variando de 1 a 5, em que 1 correspondeu às plantas com todas as folhas acima e pelo menos duas folhas verdes abaixo da espiga e 5 à planta com todas as folhas secas. O caráter florescimento feminino foi avaliado e utilizado como covariável para ajuste das variações de maturação das progênies, e as médias das parcelas foram utilizadas para as análises. A estimativa da variância aditiva foi muito superior à de dominância e a variância da interação aditiva por ambiente apresentou maior magnitude que a variância aditiva. O tipo de ação gênica foi de dominância parcial, mas devido ao desequilíbrio de ligação da população, este pode estar superestimado. O coeficiente de herdabilidade em nível de plantas apresentou baixa magnitude (~25%) enquanto que em nível de médias a magnitude foi elevada (~70%). Estes resultados indicam que os efeitos aditivos são mais importantes que os dominantes no controle do caráter stay green e que os efeitos aditivos não são consistentes nos diversos ambientes de avaliação. Assim, o nível de heterose deve apresentar baixa magnitude, e a seleção fenotípica não pode ser realizada em nível de plantas, mas baseada em médias de experimentos com repetições, conduzidas em diversos ambientes para minimizar os efeitos destes e suas interações com os genótipos. A linhagem L08-05F apresentou maior nível de stay green que a linhagem L38-05D e, portanto, pode ser utilizada como fonte de alelos favoráveis para serem transferidos para outras linhagens visando aumentar o nível deste caráter em linhagens que apresentarem baixos níveis de senescência retardada. / Several researches have reported that the increase in maize productivity could be attributed to the tolerance to abiotic stresses rather than to an increase on a plant productivity per se. Moisture stress is one of the most important stresses that reduce the maize productivity, but because of the difficulties for the direct selection for tolerance to moisture stress secondary traits as delayed senescence, also known as stay green trait, has been employed for this purpose. However, there is limited information reported on the inheritance of this trait. Thus, the objective of this research was to study the inheritance of the stay green trait in maize using the design III (Comstock and Robinson, 1952). One hundred F2:3 progenies were developed from a population derived from the cross between the inbred lines L08-05F and L38-05D; and these progenies were backcrossed to the inbred parents giving rise to 200 backcrossed progenies. These progenies were evaluated in ten environments, with two replications per environment, in Piracicaba City, São Paulo State, using the experimental design ?-lattice. The stay green trait was recorded in ten competitive plants per plot 120 days after sowing, using a scale with scores from 1 to 5, where score 1 was assigned to the plants with all leaves above the ear and at least two leaves below the ear green, and score 5 was assigned to plants with all leaves senescent. The trait days to silking was also recorded and used as covariate to adjust the maturation variation of the backcrossed progenies, and the mean of the plots was used for analysis. The estimate of the additive variance was quite larger than the dominance variance, the additive x environment variance was quite larger than the additive variance, and the average level of dominance was partial dominance, but since the population was in linkage disequilibrium the level of dominance may be overestimated. The heritability coefficient on a plant level was of low magnitude (~25%) while that one on a mean level presented high magnitude (~70%). These results indicated that the additive effects were more important than the dominance effects for the control of the stay green trait, and that the additive effects are not consistent across the environments. Therefore, the level of heterosis should present low magnitude, and the phenotypic selection should not be conducted on a plant level; instead selection should be based on the means of replicated experiments assessed in several environments to minimize their effects and the genotype by environment interactions. The inbred line L08-05F presented higher level of stay green than the inbred L38-05D and thus the former inbred could be used as a source of favorable alleles to be transferred to others inbreds to increase the level of this trait in inbreds that have low levels of delayed senescence.

Submitted by Jadson Francisco de Jesus SILVA (jadson@uefs.br) on 2018-02-22T21:50:40Z No. of bitstreams: 1 PREDI??O DE PAR?METROS GEN?TICOS E INCREMENTO DA QUALIDADE DEFINITIVO.pdf: 1627949 bytes, checksum: 31c34b863f924737f87f81fa9f625403 (MD5) / Made available in DSpace on 2018-02-22T21:50:40Z (GMT). No. of bitstreams: 1 PREDI??O DE PAR?METROS GEN?TICOS E INCREMENTO DA QUALIDADE DEFINITIVO.pdf: 1627949 bytes, checksum: 31c34b863f924737f87f81fa9f625403 (MD5) Previous issue date: 2017-03-17 / The study aimed to estimate genetic parameters and genetic variability, as well as to determine the correlations among fruit variables associated with production and pulp quality attributes in pumpkin progenies. In 2013, the parameters and prediction of progeny gains allowed the ranking of the best individuals in 10 progenies from self pollination. In 2015 the divergence was determined, selecting genotypes for qualitative and quantitative characteristics, and studying the correlations among fruit variables associated with production and pulp quality attributes in 11 evaluated genotypes. In 2013, the REML/BLUP analysis allowed the ranking considering all variables evaluated indicating the individuals 10/1/1, 10/2/12 and 6/1/3 as more promising, being the first two of ?piriform? format, and the third of "moranga" format. In 2015, it was found great variability in the progenies evaluated and similarity among progenies C. moschata 1, C. moschata 3, C. moschata 4 and C. moschata 8. The promising progenies for fruit mass and ?-carotene were C. Moschata 4 and C. moschata 7 for advances aiming to increase of production and the nutritional characteristics of the fruit. Estimates of carotenoid contents generated by the Hue angle are only valid for contrasting genotypes due to the grouping of genotypes with carotenoid content four times higher than ?Jacarezinho? cultivar. The variables explained 98% of the variation in the basic variable in the path analysis and four variables can be used for direct selection aiming to increase of fruit mass. / O estudo objetivou obter estimativas de par?metros gen?ticos e de variabilidade gen?tica, bem como determinar as correla??es entre vari?veis do fruto associadas ? produ??o e atributos de qualidade da polpa em prog?nies de ab?bora. Em 2013, os par?metros e a predi??o de ganhos gen?ticos permitiram o ranqueamento dos melhores indiv?duos em 10 prog?nies provenientes de autofecunda??es. Em 2015 determinou-se a diverg?ncia, sendo feita a sele??o de gen?tipos para caracteres qualitativos e quantitativos, al?m de estudar as correla??es entre vari?veis do fruto associadas ? produ??o e atributos de qualidade da polpa em 11 gen?tipos avaliados. Em 2013, a an?lise via REML/BLUP possibilitou o ranqueamento considerando todas as vari?veis avaliadas indicando os indiv?duos 10/1/1, 10/2/12 e 6/1/3 como mais promissores, sendo os dois primeiros de formato ?piriforme? e o terceiro de formato ?moranga?. Em 2015, constatou-se grande variabilidade nas prog?nies avaliadas e similaridade entre as prog?nies C. moschata 1, C. moschata 3, C. moschata 4 e C. moschata 8. As prog?nies promissoras para massa do fruto e ?-caroteno foram C. moschata 4 e C. moschata 7 para avan?os visando aumento da produ??o e das caracter?sticas nutricionais do fruto. As estimativas dos teores de carotenoides geradas pelo ?ngulo Hue somente s?o v?lidas para gen?tipos contrastantes devido ao agrupamento de gen?tipos com teores de carotenoides quatro vezes maiores em rela??o a cultivar Jacarezinho. As vari?veis explicaram 98% da varia??o ocorrida na vari?vel b?sica na an?lise de trilha e quatro vari?veis podem ser utilizadas para a sele??o direta visando aumento da massa do fruto.

O delineamento III foi proposto para estimar as variâncias aditivas e de dominância e o grau médio de dominância de caracteres quantitativos. Com o advento dos marcadores moleculares, Cockerham & Zeng (1996) desenvolveram uma metodologia genético-estatística associando o delineamento III com marcadores moleculares. Esta metodologia foi proposta visando estimar, com o uso de quatro contrastes ortogonais, os efeitos aditivos, dominantes e epistáticos dos QTLs ligados a marcadores moleculares. O objetivo desta pesquisa foi usar ambas as metodologias para análise genética da produção de grãos, componentes da produção e número de ramificações do pendão em uma população referência F2 de milho. Duzentos e cinqüenta progênies F2:3 foram retrocruzadas com ambas linhagens genitoras, dando origem a 500 progênies de retrocruzamento. Estas progênies foram alocadas em cinco látices 10x10 e avaliadas em seis ambientes em três estações experimentais próximas a Piracicaba, SP, com duas repetições por ambientes. Estimativas de variância aditiva e de dominância, assim como o grau médio de dominância dos caracteres avaliados, apresentaram magnitudes similares às reportadas em populações de milho temperado para todos os caracteres. Estimativas do grau médio de dominância foram inferiores a um para o diâmetro da espiga, número de fileiras, peso de 500 grãos e número de ramificações do pendão, mostrando que os efeitos aditivos foram mais importantes que os efeitos de dominância para estes caracteres. Para prolificidade, comprimento da espiga e número de grãos por fileira, o grau médio de dominância não diferiu de dominância completa, sugerindo que os efeitos de dominância foram importantes para estes caracteres. Para produção de grãos, o resultado do grau médio de dominância sugeriu sobredominância. Porém, como é conhecido, o desequilíbrio de ligação causa viéses nas estimativas de grau médio de dominância e, conseqüentemente, estas estimativas podem ser menores, sendo que, provavelmente tenha ocorrido pseudo-sobredominância para produção de grãos. A análise do delineamento III com marcadores moleculares mostrou que os QTLs estão distribuídos em todos os cromossomos para todos caracteres. As somas em módulo dos efeitos dos QTLs mostraram que para produção de grãos, prolificidade, comprimento da espiga, e número de grãos por fileira, os efeitos de dominância foram superiores aos efeitos aditivos, e estes superiores aos efeitos epistáticos; para diâmetro da espiga, peso de 500 grãos, número de fileiras, e número ramificações do pendão, os efeitos aditivos foram maiores que os efeitos de dominância, e estes superiores aos efeitos epistáticos, exceto para número de fileiras em que os efeitos epistáticos foram maiores que os efeitos de dominância. Os efeitos epistáticos foram detectados para todos caracteres e contribuíram mais para os componentes da produção que para a produção de grãos per se. A análise clássica do delineamento III e a associada a marcadores moleculares forneceram resultados de grande utilidade, mas o delineamento III com marcadores permitiu a estimação dos efeitos genéticos de regiões específicas do genoma e sugeriu que os efeitos epistáticos foram muito importantes na expressão e na herança dos caracteres analisados. / The Design III was proposed to estimate additive and dominance variances, and the average levels of dominance of quantitative traits. With the advent of the molecular markers, Cockerham & Zeng (1996) developed a genetic-statistical procedure using the Design III with molecular markers. This procedure was designed to estimate additive, dominance and epistatic effects of the QTLs linked to molecular markers from four orthogonal contrasts. The objectives of this research were to use both methodologies for the genetic analysis of grain yield, yield components, and number of tassel branches in an F2 reference maize population. Two-hundred and fifty F2:3 progenies were backcrossed to the two parental inbred lines, which gave rise to 500 backcrossed progenies. These progenies were allocated in five 10x10 lattices design, and evaluated in six environments in three experimental stations near Piracicaba, SP, with two replications per environment. Estimates of additive and dominance variances, as well as the average levels of dominance for the traits evaluated, had magnitudes similar to those already reported for temperate maize populations for all traits. Estimates of the average level of dominance were lower than one for ear diameter, kernel row number, weight of 500 kernels, and tassel branches number, showing that the additive effects were more important than the dominance effects for these traits. For prolificacy, ear length, and kernels per row number, the average levels of dominance did not differ from complete dominance, suggesting that the dominance effects were important for these traits. For grain yield, the result suggested overdominance as the average level of dominance. However, as is well-known, linkage disequilibrium causes biases in the estimates of the average levels of dominance and, therefore, these estimates could be lower, and probably for grain yield a pseudo-overdominance was detected. The analysis of the Design III with molecular markers showed that QTLs were distributed in all chromosomes for all traits. The sum in module of the QTLs effects showed that for grain yield, prolificacy, ear length, and kernels per row number, dominance effect was greater than additive effect, and the latter greater than epistatic effect; whereas for ear diameter, weight of 500 kernels, kernel row number, and tassel branches number, additive effect was greater than dominance effect, and the latter greater than epistatic effect, except for kernel row number where epistatic effect was greater than dominance effect. Epistatic effects were detected for all traits, and had higher contribution for the yield components than for yield per se. Both traditional and QTL analysis of the Design III presented very useful results, but the Design III with molecular markers allowed the estimation of the genetic effects from specific genomic regions, and suggested that the epistatic effects play a very important role for the expression and inheritance of the traits assessed.

<p>In this thesis, quantitative genetic approaches, common-garden experiments, and field studies were combined to examine patterns of population differentiation and the genetic architecture of characters of putative adaptive significance in the widely distributed perennial herb Lythrum salicaria. In this work, I (1) documented patterns of population differentiation in phenology, life-history, and morphology along latitudinal gradients at different geographical scales, (2) investigated the genetic architecture of phenology, flower morphology, and inflorescence size, and (3) combined estimates of phenotypic selection in the field with information on the genetic variance-covariance matrix (G) to examine potential constraints to adaptive evolution. </p><p>A common-garden experiment demonstrated latitudinal variation in life-history, and phenology of growth and reproduction among L. salicaria populations sampled across Sweden. Flower morphology varied significantly among populations, but was, with the exception of calyx length, not related to latitude of origin. A second experiment, which included two Swedish, two Dutch, and two Spanish populations, indicated that the latitudinal gradient in reproductive and vegetative phenology might extend throughout Europe.</p><p>A quantitative-genetic study of two Swedish populations revealed significant additive genetic variation for all phenological and morphological traits investigated. The G matrices of the populations differed significantly according to common principal component analysis, and genetic correlations within the study populations did not strictly correspond to trait correlations observed among populations.</p><p>In a field study, I detected directional selection through female function for larger inflorescences in two consecutive years. Relative fitness increased disproportionately with inflorescence size in the year when supplemental hand-pollination indicated that pollen limitation was severe. Genetic correlations with inflorescence size considerably influenced predicted response to selection in other characters.</p><p>Taken together, the results suggest that among-population differences in phenology and life-history in L. salicaria have evolved in response to latitudinal variation in length of the growing season. They demonstrate that the evolutionary potential of local populations may be considerable. The genetic covariance structure substantially influences predicted short-term evolutionary trajectories. However, the weak correspondence between genetic correlations documented within populations and trait correlations among populations, suggest that the G matrix has not imposed strict constraints on patterns of among-population differentiation.</p>

This thesis develops and evaluates statistical methods for different types of genetic analyses, including quantitative trait loci (QTL) analysis, genome-wide association study (GWAS), and genomic evaluation. The main contribution of the thesis is to provide novel insights in modeling genetic variance, especially via random effects models. In variance component QTL analysis, a full likelihood model accounting for uncertainty in the identity-by-descent (IBD) matrix was developed. It was found to be able to correctly adjust the bias in genetic variance component estimation and gain power in QTL mapping in terms of precision.  Double hierarchical generalized linear models, and a non-iterative simplified version, were implemented and applied to fit data of an entire genome. These whole genome models were shown to have good performance in both QTL mapping and genomic prediction. A re-analysis of a publicly available GWAS data set identified significant loci in Arabidopsis that control phenotypic variance instead of mean, which validated the idea of variance-controlling genes.  The works in the thesis are accompanied by R packages available online, including a general statistical tool for fitting random effects models (hglm), an efficient generalized ridge regression for high-dimensional data (bigRR), a double-layer mixed model for genomic data analysis (iQTL), a stochastic IBD matrix calculator (MCIBD), a computational interface for QTL mapping (qtl.outbred), and a GWAS analysis tool for mapping variance-controlling loci (vGWAS).

In this thesis, quantitative genetic approaches, common-garden experiments, and field studies were combined to examine patterns of population differentiation and the genetic architecture of characters of putative adaptive significance in the widely distributed perennial herb Lythrum salicaria. In this work, I (1) documented patterns of population differentiation in phenology, life-history, and morphology along latitudinal gradients at different geographical scales, (2) investigated the genetic architecture of phenology, flower morphology, and inflorescence size, and (3) combined estimates of phenotypic selection in the field with information on the genetic variance-covariance matrix (G) to examine potential constraints to adaptive evolution. A common-garden experiment demonstrated latitudinal variation in life-history, and phenology of growth and reproduction among L. salicaria populations sampled across Sweden. Flower morphology varied significantly among populations, but was, with the exception of calyx length, not related to latitude of origin. A second experiment, which included two Swedish, two Dutch, and two Spanish populations, indicated that the latitudinal gradient in reproductive and vegetative phenology might extend throughout Europe. A quantitative-genetic study of two Swedish populations revealed significant additive genetic variation for all phenological and morphological traits investigated. The G matrices of the populations differed significantly according to common principal component analysis, and genetic correlations within the study populations did not strictly correspond to trait correlations observed among populations. In a field study, I detected directional selection through female function for larger inflorescences in two consecutive years. Relative fitness increased disproportionately with inflorescence size in the year when supplemental hand-pollination indicated that pollen limitation was severe. Genetic correlations with inflorescence size considerably influenced predicted response to selection in other characters. Taken together, the results suggest that among-population differences in phenology and life-history in L. salicaria have evolved in response to latitudinal variation in length of the growing season. They demonstrate that the evolutionary potential of local populations may be considerable. The genetic covariance structure substantially influences predicted short-term evolutionary trajectories. However, the weak correspondence between genetic correlations documented within populations and trait correlations among populations, suggest that the G matrix has not imposed strict constraints on patterns of among-population differentiation.

This thesis develops and evaluates statistical methods for different types of genetic analyses, including quantitative trait loci (QTL) analysis, genome-wide association study (GWAS), and genomic evaluation. The main contribution of the thesis is to provide novel insights in modeling genetic variance, especially via random effects models. In variance component QTL analysis, a full likelihood model accounting for uncertainty in the identity-by-descent (IBD) matrix was developed. It was found to be able to correctly adjust the bias in genetic variance component estimation and gain power in QTL mapping in terms of precision.  Double hierarchical generalized linear models, and a non-iterative simplified version, were implemented and applied to fit data of an entire genome. These whole genome models were shown to have good performance in both QTL mapping and genomic prediction. A re-analysis of a publicly available GWAS data set identified significant loci in Arabidopsis that control phenotypic variance instead of mean, which validated the idea of variance-controlling genes.  The works in the thesis are accompanied by R packages available online, including a general statistical tool for fitting random effects models (hglm), an efficient generalized ridge regression for high-dimensional data (bigRR), a double-layer mixed model for genomic data analysis (iQTL), a stochastic IBD matrix calculator (MCIBD), a computational interface for QTL mapping (qtl.outbred), and a GWAS analysis tool for mapping variance-controlling loci (vGWAS).

Submitted by Cláudia Bueno (claudiamoura18@gmail.com) on 2015-10-14T22:00:35Z No. of bitstreams: 1 Dissertação - Poliana Regina Carloni Di Prado - 2013.pdf: 1121268 bytes, checksum: de6b8400cf11e9a39e0a3bb61045e84e (MD5) / Approved for entry into archive by Luciana Ferreira (lucgeral@gmail.com) on 2015-10-16T10:53:58Z (GMT) No. of bitstreams: 1 Dissertação - Poliana Regina Carloni Di Prado - 2013.pdf: 1121268 bytes, checksum: de6b8400cf11e9a39e0a3bb61045e84e (MD5) / Made available in DSpace on 2015-10-16T10:53:58Z (GMT). No. of bitstreams: 1 Dissertação - Poliana Regina Carloni Di Prado - 2013.pdf: 1121268 bytes, checksum: de6b8400cf11e9a39e0a3bb61045e84e (MD5) Previous issue date: 2013-03-13 / The consumption of pepper genus Capsicum is in increasing demand worldwide. However few works have been conducted aiming their utilization in plant breeding research as well as in preserving their genetic patrimonium. The objectives of this work were to estimate the genetic parameters of a pimenta de bode (Capsicum chinense Jacq) population from the Southwest of Goiás as well as to assess gains from selection carried out in that population based on six selection criteria. Twenty seven genotypes and three commercial controls were used and the experiment arranged in a completely randomized bloc design with three replicates and four plants per plot spaced by 1.2 X 0.8 m. The estimated parameters were: genetic variance σ²g, environmental variance σ²e, phenotypic variance σ²g; genetic variance coefficient CVg%, the ratio between the genetic coefficient of variance and the environmental ratio CVg/CVe, and the heritability in broad sense term h2. The criteria used from gains selection were: selection I – shortening the flowering period; II - increase harvested fruit weight in the first harvest; III – increase fruit production; IV – increase 10 fruit weight; V – reduce 10 fruit weight; VI –shorten ripening period The CVg% was above 10% for the majority of the traits tested as well as the CVg/CVe , above 1, indicating the predominance of the genetic traits in relation to the environmental factors; being these traits considered ideal for use in genetic breeding programs. Heritability in broad sense (h2) varied from 54.27 to 99.66%. The selection criteria used presenting gains from selection were I, III, IV and VI, with 10.30, 69.59, 181 and 1.31 % respectively of gains obtained for the selected trait. / O consumo de pimentas, do gênero Capsicum, possui uma crescente demanda em nível mundial. Por outro lado, poucos trabalhos têm sido realizados com a finalidade de preservar o patrimônio genético dessas espécies e também utilização deste em programas de melhoramento genético. Os objetivos deste trabalho foram estimar os parâmetros genéticos de uma população de pimenta de bode (Capsicum chinense Jacq) do sudoeste goiano e os ganhos de seleção realizados nessa população baseados em seis critérios de seleção. Foram utilzados 27 genótipos e 3 testemunhas comerciais, o experimento foi conduzido em blocos casualizados, com três repetições, quatro plantas por parcela e espaçamento de 1.2x0.8m. Os parâmetros estimados foram variâncias genética σ²g, ambiental σ²e, fenotípica σ²f, o coeficiente de variação genético CVg%, a razão entre o coeficiente de variação genético e o ambiental CVg/CVe e a herdabilidade em sentido amplo h2. Os critérios utilizados foram; seleção I: reduzir os dias para florescimento, II: aumentar o peso na primeira colheita, III: aumentar a produção de frutos, IV: aumentar o peso de 10 frutos, V: reduzir o peso de 10 frutos e VI: reduzir os dias para maturação. O CVg% foi acima de 10% para a maioria das características, o CVg/CVe foi acima de 1 para a maioria das características indicando que os fatores genéticos são predominantes em relação aos ambientais. A h2 variou de 54.27 à 99.66%. Os critérios de seleção utilizados que apresentaram ganhos de seleção foram I, III, IV, VI com 10.30, 69.59, 181, 1.31% respectivamente de ganho realizado para característica selecionada.

O delineamento III foi proposto para estimar as variâncias aditivas e de dominância e o grau médio de dominância de caracteres quantitativos. Com o advento dos marcadores moleculares, Cockerham & Zeng (1996) desenvolveram uma metodologia genético-estatística associando o delineamento III com marcadores moleculares. Esta metodologia foi proposta visando estimar, com o uso de quatro contrastes ortogonais, os efeitos aditivos, dominantes e epistáticos dos QTLs ligados a marcadores moleculares. O objetivo desta pesquisa foi usar ambas as metodologias para análise genética da produção de grãos, componentes da produção e número de ramificações do pendão em uma população referência F2 de milho. Duzentos e cinqüenta progênies F2:3 foram retrocruzadas com ambas linhagens genitoras, dando origem a 500 progênies de retrocruzamento. Estas progênies foram alocadas em cinco látices 10x10 e avaliadas em seis ambientes em três estações experimentais próximas a Piracicaba, SP, com duas repetições por ambientes. Estimativas de variância aditiva e de dominância, assim como o grau médio de dominância dos caracteres avaliados, apresentaram magnitudes similares às reportadas em populações de milho temperado para todos os caracteres. Estimativas do grau médio de dominância foram inferiores a um para o diâmetro da espiga, número de fileiras, peso de 500 grãos e número de ramificações do pendão, mostrando que os efeitos aditivos foram mais importantes que os efeitos de dominância para estes caracteres. Para prolificidade, comprimento da espiga e número de grãos por fileira, o grau médio de dominância não diferiu de dominância completa, sugerindo que os efeitos de dominância foram importantes para estes caracteres. Para produção de grãos, o resultado do grau médio de dominância sugeriu sobredominância. Porém, como é conhecido, o desequilíbrio de ligação causa viéses nas estimativas de grau médio de dominância e, conseqüentemente, estas estimativas podem ser menores, sendo que, provavelmente tenha ocorrido pseudo-sobredominância para produção de grãos. A análise do delineamento III com marcadores moleculares mostrou que os QTLs estão distribuídos em todos os cromossomos para todos caracteres. As somas em módulo dos efeitos dos QTLs mostraram que para produção de grãos, prolificidade, comprimento da espiga, e número de grãos por fileira, os efeitos de dominância foram superiores aos efeitos aditivos, e estes superiores aos efeitos epistáticos; para diâmetro da espiga, peso de 500 grãos, número de fileiras, e número ramificações do pendão, os efeitos aditivos foram maiores que os efeitos de dominância, e estes superiores aos efeitos epistáticos, exceto para número de fileiras em que os efeitos epistáticos foram maiores que os efeitos de dominância. Os efeitos epistáticos foram detectados para todos caracteres e contribuíram mais para os componentes da produção que para a produção de grãos per se. A análise clássica do delineamento III e a associada a marcadores moleculares forneceram resultados de grande utilidade, mas o delineamento III com marcadores permitiu a estimação dos efeitos genéticos de regiões específicas do genoma e sugeriu que os efeitos epistáticos foram muito importantes na expressão e na herança dos caracteres analisados. / The Design III was proposed to estimate additive and dominance variances, and the average levels of dominance of quantitative traits. With the advent of the molecular markers, Cockerham & Zeng (1996) developed a genetic-statistical procedure using the Design III with molecular markers. This procedure was designed to estimate additive, dominance and epistatic effects of the QTLs linked to molecular markers from four orthogonal contrasts. The objectives of this research were to use both methodologies for the genetic analysis of grain yield, yield components, and number of tassel branches in an F2 reference maize population. Two-hundred and fifty F2:3 progenies were backcrossed to the two parental inbred lines, which gave rise to 500 backcrossed progenies. These progenies were allocated in five 10x10 lattices design, and evaluated in six environments in three experimental stations near Piracicaba, SP, with two replications per environment. Estimates of additive and dominance variances, as well as the average levels of dominance for the traits evaluated, had magnitudes similar to those already reported for temperate maize populations for all traits. Estimates of the average level of dominance were lower than one for ear diameter, kernel row number, weight of 500 kernels, and tassel branches number, showing that the additive effects were more important than the dominance effects for these traits. For prolificacy, ear length, and kernels per row number, the average levels of dominance did not differ from complete dominance, suggesting that the dominance effects were important for these traits. For grain yield, the result suggested overdominance as the average level of dominance. However, as is well-known, linkage disequilibrium causes biases in the estimates of the average levels of dominance and, therefore, these estimates could be lower, and probably for grain yield a pseudo-overdominance was detected. The analysis of the Design III with molecular markers showed that QTLs were distributed in all chromosomes for all traits. The sum in module of the QTLs effects showed that for grain yield, prolificacy, ear length, and kernels per row number, dominance effect was greater than additive effect, and the latter greater than epistatic effect; whereas for ear diameter, weight of 500 kernels, kernel row number, and tassel branches number, additive effect was greater than dominance effect, and the latter greater than epistatic effect, except for kernel row number where epistatic effect was greater than dominance effect. Epistatic effects were detected for all traits, and had higher contribution for the yield components than for yield per se. Both traditional and QTL analysis of the Design III presented very useful results, but the Design III with molecular markers allowed the estimation of the genetic effects from specific genomic regions, and suggested that the epistatic effects play a very important role for the expression and inheritance of the traits assessed.

Diversas pesquisas têm reportado que o aumento da produtividade da cultura do milho se deve muito mais à tolerância a estresses abióticos do que a aumentos per se. O déficit hídrico é um dos mais importantes fatores envolvidos na redução de produtividade nessa cultura, mas devido à dificuldade de seleção direta para tolerância à seca, são empregados caracteres secundários como o caráter stay green ou senescência retardada de folhas e colmos. Todavia, estudos sobre a herança desse caráter são escassos na literatura. Em vista disso, este trabalho teve como objetivo estudar a herança deste caráter em milho tropical, utilizando o delineamento III (Comstock e Robinson, 1952). A partir do cruzamento entre as linhagens endogâmicas L08-05F e L38-05D, foram obtidas 100 progênies F2:3, as quais foram retrocruzadas com cada um dos parentais, resultando em 200 progênies de retrocruzamento. Estas progênies foram avaliadas em dez ambientes, com duas repetições por ambiente, em Piracicaba SP, utilizando o delineamento ?-látice. A avaliação do stay green foi feita em dez plantas competitivas por parcela, 120 dias após a semeadura, utilizando uma escala de notas variando de 1 a 5, em que 1 correspondeu às plantas com todas as folhas acima e pelo menos duas folhas verdes abaixo da espiga e 5 à planta com todas as folhas secas. O caráter florescimento feminino foi avaliado e utilizado como covariável para ajuste das variações de maturação das progênies, e as médias das parcelas foram utilizadas para as análises. A estimativa da variância aditiva foi muito superior à de dominância e a variância da interação aditiva por ambiente apresentou maior magnitude que a variância aditiva. O tipo de ação gênica foi de dominância parcial, mas devido ao desequilíbrio de ligação da população, este pode estar superestimado. O coeficiente de herdabilidade em nível de plantas apresentou baixa magnitude (~25%) enquanto que em nível de médias a magnitude foi elevada (~70%). Estes resultados indicam que os efeitos aditivos são mais importantes que os dominantes no controle do caráter stay green e que os efeitos aditivos não são consistentes nos diversos ambientes de avaliação. Assim, o nível de heterose deve apresentar baixa magnitude, e a seleção fenotípica não pode ser realizada em nível de plantas, mas baseada em médias de experimentos com repetições, conduzidas em diversos ambientes para minimizar os efeitos destes e suas interações com os genótipos. A linhagem L08-05F apresentou maior nível de stay green que a linhagem L38-05D e, portanto, pode ser utilizada como fonte de alelos favoráveis para serem transferidos para outras linhagens visando aumentar o nível deste caráter em linhagens que apresentarem baixos níveis de senescência retardada. / Several researches have reported that the increase in maize productivity could be attributed to the tolerance to abiotic stresses rather than to an increase on a plant productivity per se. Moisture stress is one of the most important stresses that reduce the maize productivity, but because of the difficulties for the direct selection for tolerance to moisture stress secondary traits as delayed senescence, also known as stay green trait, has been employed for this purpose. However, there is limited information reported on the inheritance of this trait. Thus, the objective of this research was to study the inheritance of the stay green trait in maize using the design III (Comstock and Robinson, 1952). One hundred F2:3 progenies were developed from a population derived from the cross between the inbred lines L08-05F and L38-05D; and these progenies were backcrossed to the inbred parents giving rise to 200 backcrossed progenies. These progenies were evaluated in ten environments, with two replications per environment, in Piracicaba City, São Paulo State, using the experimental design ?-lattice. The stay green trait was recorded in ten competitive plants per plot 120 days after sowing, using a scale with scores from 1 to 5, where score 1 was assigned to the plants with all leaves above the ear and at least two leaves below the ear green, and score 5 was assigned to plants with all leaves senescent. The trait days to silking was also recorded and used as covariate to adjust the maturation variation of the backcrossed progenies, and the mean of the plots was used for analysis. The estimate of the additive variance was quite larger than the dominance variance, the additive x environment variance was quite larger than the additive variance, and the average level of dominance was partial dominance, but since the population was in linkage disequilibrium the level of dominance may be overestimated. The heritability coefficient on a plant level was of low magnitude (~25%) while that one on a mean level presented high magnitude (~70%). These results indicated that the additive effects were more important than the dominance effects for the control of the stay green trait, and that the additive effects are not consistent across the environments. Therefore, the level of heterosis should present low magnitude, and the phenotypic selection should not be conducted on a plant level; instead selection should be based on the means of replicated experiments assessed in several environments to minimize their effects and the genotype by environment interactions. The inbred line L08-05F presented higher level of stay green than the inbred L38-05D and thus the former inbred could be used as a source of favorable alleles to be transferred to others inbreds to increase the level of this trait in inbreds that have low levels of delayed senescence.