The effect of seed-dispersal timing on seedling recruitment is modulated by environmental conditions that vary across altitude in a threatened palm.

BACKGROUND AND AIMS: The timing of seed dispersal determines the environmental conditions that plants face during early life stages. In seasonal environments, selection is expected to favour dispersal timing that is matched to environmental conditions suitable for successful recruitment. Our aim here was to test whether the timing of seed dispersal influences seedling establishment success in two populations of Euterpe edulis that are located at contrasting altitudes, have different seed-dispersal phenologies and are subjected to distinct climatic conditions. METHODS: We sowed E. edulis seeds in contrasting altitudes on different dates, and monitored seed germination, emergence and seedling establishment at each altitude over 4 years. At the high-altitude site, five seed-dispersal cohorts were established during the natural dispersal period. At the low-altitude site, three seed-dispersal cohorts were established during natural dispersal, and two were established either before or after natural dispersal. KEY RESULTS: At the high-altitude site, seed-dispersal timing did not affect seed germination, seedling emergence or seedling establishment success. In contrast, at the low-altitude site, late seed dispersal near the end of the wet season resulted in a lower probability of seedling establishment, possibly due to the exposure of seeds, germinants and seedlings to unfavourable drought conditions. In addition, at the low-altitude site, the natural seed-dispersal period was poorly matched to favourable environmental conditions for seedling establishment. CONCLUSIONS: The greater effect of seed-dispersal timing on seedling establishment at the low-altitude site is probably related to a more seasonal and drought-prone environment that favours a restricted period of seed dispersal. The magnitude of the effect of dispersal timing on seedling establishment success was modulated by environmental conditions that vary across altitude. Furthermore, reproductive phenology appears to be subject to more intense selection at the lower limit of the altitudinal range, due to a more restrictive window of opportunity for successful seedling establishment.

Testing Wallace's intuition: water type, reproductive isolation and divergence in an Amazonian fish.

Alfred Russel Wallace proposed classifying Amazon rivers based on their colour and clarity: white, black and clear water. Wallace also proposed that black waters could mediate diversification and yield distinct fish species. Here, we bring evidence of speciation mediated by water type in the sailfin tetra (Crenuchus spilurus), a fish whose range encompasses rivers of very distinct hydrochemical conditions. Distribution of the two main lineages concords with Wallace's water types: one restricted to the acidic and nutrient-poor waters of the Negro River (herein Rio Negro lineage) and a second widespread throughout the remaining of the species' distribution (herein Amazonas lineage). These lineages occur over a very broad geographical range, suggesting that despite occurring in regions separated by thousands of kilometres, individuals of the distinct lineages fail to occupy each other's habitats, hundreds of metres apart and not separated by physical barrier. Reproductive isolation was assessed in isolated pairs exposed to black-water conditions. All pairs with at least one individual of the lineage not native to black waters showed significantly lower spawning success, suggesting that the water type affected the fitness and contributed to reproductive isolation. Our results endorse Wallace's intuition and highlight the importance of ecological factors in shaping diversity of the Amazon fish fauna.

Arboreality constrains morphological evolution but not species diversification in vipers.

An increase in ecological opportunities, either through changes in the environment or acquisition of new traits, is frequently associated with an increase in species and morphological diversification. However, it is possible that certain ecological settings might prevent lineages from diversifying. Arboreality evolved multiple times in vipers, making them ideal organisms for exploring how potentially new ecological opportunities affect their morphology and speciation regimes. Arboreal snakes are frequently suggested to have a very specialized morphology, and being too large, too small, too heavy, or having short tails might be challenging for them. Using trait-evolution models, we show that arboreal vipers are evolving towards intermediate body sizes, with longer tails and more slender bodies than terrestrial vipers. Arboreality strongly constrains body size and circumference evolution in vipers, while terrestrial lineages are evolving towards a broader range of morphological variants. Trait-dependent diversification models, however, suggest similar speciation rates between microhabitats. Thus, we show that arboreality might constrain morphological evolution but not necessarily affect the rates at which lineages generate new species.

Ecological and evolutionary consequences of tri-trophic interactions: Spatial variation and effects of plant density.

PREMISE OF STUDY: The factors driving variation in species interactions are often unknown, and few studies have made a link between changes in interactions and the strength of selection. METHODS: We report on spatial variation in functional responses by a seed predator (SP) and its parasitic wasps associated with the herb Ruellia nudiflora. We assessed the influence of plant density on consumer responses and determined whether density effects and spatial variation in functional responses altered natural selection by these consumers on the plant. We established common gardens at two sites in Yucatan, Mexico, and planted R. nudiflora at two densities in each garden. We recorded fruit output and SP and parasitoid attack; calculated relative fitness (seed number) under scenarios of three trophic levels (accounting for SP and parasitoid effects), two trophic levels (accounting for SP but not parasitoid effects), and one trophic level (no consumer effects); and compared selection strength on fruit number under these scenarios across sites and densities. KEY RESULTS: There was spatial variation in SP recruitment, whereby the SP functional response was negatively density-dependent at one site but density-independent at the other; parasitoid responses were density-independent and invariant across sites. Site variation in SP attack led, in turn, to differences in SP selection on fruit output, and parasitoids did not alter SP selection. There were no significant effects of density at either site. CONCLUSIONS: Our results provide a link between consumer functional responses and consumer selection on plants, which deepens our understanding of geographic variation in the evolutionary outcomes of multitrophic interactions.

Evolutionary heritage influences Amazon tree ecology.

Lineages tend to retain ecological characteristics of their ancestors through time. However, for some traits, selection during evolutionary history may have also played a role in determining trait values. To address the relative importance of these processes requires large-scale quantification of traits and evolutionary relationships among species. The Amazonian tree flora comprises a high diversity of angiosperm lineages and species with widely differing life-history characteristics, providing an excellent system to investigate the combined influences of evolutionary heritage and selection in determining trait variation. We used trait data related to the major axes of life-history variation among tropical trees (e.g. growth and mortality rates) from 577 inventory plots in closed-canopy forest, mapped onto a phylogenetic hypothesis spanning more than 300 genera including all major angiosperm clades to test for evolutionary constraints on traits. We found significant phylogenetic signal (PS) for all traits, consistent with evolutionarily related genera having more similar characteristics than expected by chance. Although there is also evidence for repeated evolution of pioneer and shade tolerant life-history strategies within independent lineages, the existence of significant PS allows clearer predictions of the links between evolutionary diversity, ecosystem function and the response of tropical forests to global change.

Functional decoupling between flowers and leaves in the Ameroglossum pernambucense complex can facilitate local adaptation across a pollinator and climatic heterogeneous landscape.

Decoupling between floral and leaf traits is expected in plants with specialized pollination systems to assure a precise flower-pollinator fit, irrespective of leaf variation associated with environmental heterogeneity (functional modularity). Nonetheless, developmental interactions among floral traits also decouple flowers from leaves regardless of selection pressures (developmental modularity). We tested functional modularity in the hummingbird-pollinated flowers of the Ameroglossum pernambucense complex while controlling for developmental modularity. Using two functional traits responsible for flower-pollinator fit [floral tube length (TL) and anther-nectary distance (AN)], one floral trait not linked to pollination [sepal length (SL), control for developmental modularity] and one leaf trait [leaf length (LL)], we found evidence of flower functional modularity. Covariation between TL and AN was ca. two-fold higher than the covariation of either of these traits with sepal and leaf lengths, and variations in TL and AN, important for a precise flower-pollinator fit, were smaller than SL and LL variations. Furthermore, we show that previously reported among-population variation of flowers associated with local pollinator phenotypes was independent from SL and LL variations. These results suggest that TL and AN are functionally linked to fit pollinators and sufficiently decoupled from developmentally related floral traits (SL) and vegetative traits (LL). These results support previous evidences of population differentiation due to local adaptation in the A. pernambucense complex and shed light on the role of flower-leaf decoupling for local adaptation in species distributed across biotic and abiotic heterogeneous landscapes.

Genetic variability for carotenoid content of grains in a composite maize population

Local maize (Zea mays L.) varieties are cultivated by small-scale farmers in western Santa Catarina (SC) State, in southern Brazil. These small areas frequently present many problems related to biotic and non-biotic stresses, which have limited the economic output and income of the farmers. Production from local varieties for human consumption would be an alternative way of improving income and stimulating on farm conservation. The genetic variability of the total carotenoid content (TCC) of kernels in a local maize population was evaluated for their economic exploitation potential as biofortified food. Two independent samples of 96 half-sib families (HSF) plus four checks were evaluated in two groups of experiments in western SC and each one was carried out in two environments. They were set out in a 10 × 10 partially balanced lattice with three replications per location; plots consisted of one row, 5.0 m long with 1.0 m between rows. TCC ranged from 11 to 23 µg g-1, averaging 16 µg g-1 in the pooled analysis over the two sets. The local composite population exhibited genetic variability in order to increase the TCC of grains in the second cycle of selection by the convergent-divergent scheme.

Genetic variability for carotenoid content of grains in a composite maize population

Local maize (Zea mays L.) varieties are cultivated by small-scale farmers in western Santa Catarina (SC) State, in southern Brazil. These small areas frequently present many problems related to biotic and non-biotic stresses, which have limited the economic output and income of the farmers. Production from local varieties for human consumption would be an alternative way of improving income and stimulating on farm conservation. The genetic variability of the total carotenoid content (TCC) of kernels in a local maize population was evaluated for their economic exploitation potential as biofortified food. Two independent samples of 96 half-sib families (HSF) plus four checks were evaluated in two groups of experiments in western SC and each one was carried out in two environments. They were set out in a 10 × 10 partially balanced lattice with three replications per location; plots consisted of one row, 5.0 m long with 1.0 m between rows. TCC ranged from 11 to 23 µg g-1, averaging 16 µg g-1 in the pooled analysis over the two sets. The local composite population exhibited genetic variability in order to increase the TCC of grains in the second cycle of selection by the convergent-divergent scheme.(AU)

Adaptive evolution of a generalist parasitoid: implications for the effectiveness of biological control agents.

The use of alternative hosts imposes divergent selection pressures on parasitoid populations. In response to selective pressures, these populations may follow different evolutionary trajectories. Divergent natural selection could promote local host adaptation in populations, translating into direct benefits for biological control, thereby increasing their effectiveness on the target host. Alternatively, adaptive phenotypic plasticity could be favored over local adaptation in temporal and spatially heterogeneous environments. We investigated the existence of local host adaptation in Aphidius ervi, an important biological control agent, by examining different traits related to infectivity (preference) and virulence (a proxy of parasitoid fitness) on different aphid-host species. The results showed significant differences in parasitoid infectivity on their natal host compared with the non-natal hosts. However, parasitoids showed a similar high fitness on both natal and non-natal hosts, thus supporting a lack of host adaptation in these introduced parasitoid populations. Our results highlight the role of phenotypic plasticity in fitness-related traits of parasitoids, enabling them to maximize fitness on alternative hosts. This could be used to increase the effectiveness of biological control. In addition, A. ervi females showed significant differences in infectivity and virulence across the tested host range, thus suggesting a possible host phylogeny effect for those traits.

Field comparison between selection methods at the maize seedling stage in relation to aluminum tolerance

Selection and breeding for aluminum tolerance is a useful approach to increase maize (Zea mays L.) grain yield in acid soils. The objective of this work was the comparison between two screening approaches for the development of Al tolerant maize populations. One cycle of divergent selection for aluminum tolerance, using both nutrient solution and pots with acid soil, were completed in the maize population SIKALQ. The four sub-populations selected through both selection procedures were compared with the original population and checks in field trials repeated over locations. Eight environments stratified into four groups varying from non acid to high aluminum saturation were used. Three variables were analyzed for comparisons of methodologies: grain yield, plant height and days to male flowering. The populations performed similarly in the high productivity locations and the differences between tolerant and sensitive ones were more evident for increasing levels of toxic aluminum. In spite of little differences, selection in pots with acid soil was more efficient than nutrient solution in changing the genetic structure of the population toward more tolerant and sensitive levels.

A seleção e o melhoramento visando tolerância ao alumínio é um procedimento valioso para aumentar a produção de milho (Zea mays L.) em solos ácidos. O objetivo deste trabalho foi comparar a eficiência de duas metodologias de seleção de plântulas, no desenvolvimento de populações de milho tolerantes ao alumínio. Foi realizado na população SIKALQ, um ciclo de seleção divergente para tolerância ao alumínio, utilizando solução nutritiva e vasos com solo ácido. As quatro sub-populações selecionadas pelas duas metodologias foram comparadas com a população original e testemunhas em experimentos de campo em vários locais. Foram utilizados oito locais, estratificados em quatro grupos começando com solos não ácidos até com alta saturação com alumínio. As comparações foram realizadas utilizando três variáveis: produção de grãos, altura de planta e dias até florescimento. As populações apresentaram um comportamento semelhante entre elas nos locais de alta produtividade, e as diferenças entre as tolerantes e suscetíveis ficaram mais evidentes na medida em que aumentou a toxidez por alumínio. Os resultados sugerem que a seleção em vasos com solo ácido foi mais eficiente em alterar a estrutura genética da população do que a solução nutritiva.

Selection methods for maize seedlings in greenhouse as related to aluminum tolerance

Genetic improvement is a useful approach to increase aluminum tolerance in maize. This experiment was carried out to compare two screening techniques under greenhouse conditions and estimate the association between the results obtained for both bioassays with grain yield. Nutrient solution (NS) and pots with acid soil (AS) were utilized as screening methodologies to perform one cycle of divergent selection for aluminum tolerance in the tropical maize population SIKALQ. The four sub-populations obtained by both approaches were compared with the original and checks in a greenhouse and in multilocation field trials. Evaluation under the conditions of NS and AS showed that the populations with the best performances were those selected for aluminum tolerance under the same condition of evaluation. The variables measured in greenhouse showed good correlation and the most closely related were fresh root weight (FRW) and dry rot weight (DRW) (0.79). All of these variables showed low correlation with yield in non acid conditions ( 0.30). The correlation was more important (~ 0.50) as soil aluminum saturation increased. The best correlation was observed for visual scoring (VS) (0.68), FRW (0.47) and net root growth (NRG) (0.52).

O melhoramento genético tem se mostrado como estratégia eficiente para melhorar a tolerância ao alumínio em milho. O objetivo deste trabalho foi comparar dois metodos de seleção em casa de vegetação e estimar a associação entre os resultados obtidos em cada um desses ensaios com a produção de grãos no campo. Solução nutritiva (NS) e vasos com solo ácido (AS) foram utilizadas nas metodologias de seleção para realizar um ciclo de seleção divergente para tolerância ao alumínio na população de milho tropical SIKALQ. As quatro subpopulações obtidas foram comparadas com a original e as testemunhas em experimentos de casa de vegetação e de campo. A avaliação sob as condições de NS e AS mostraram que as populações com melhor comportamento foram aquelas selecionadas para tolerância ao alumínio nas mesmas condições da avaliação. As variáveis medidas na casa de vegetação apresentaram boa correlação e as mais fortemente relacionadas foram peso fresco da raiz (FRW) e peso seco da raiz (DRW) (0.79). Todas as variáveis apresentaram baixa correlação com a produção de grãos em solos sem acidez ( 0.30). A correlação foi mais importante (~ 0.50) na medida que a saturação com alumínio do solo foi aumentando. Os melhores valores de correlação foram observados para avaliação visual (VS) (0.68), FRW (0.47) e crescimento líquido da raiz (NRG) (0.52).

Selection methods for maize seedlings in greenhouse as related to aluminum tolerance

Genetic improvement is a useful approach to increase aluminum tolerance in maize. This experiment was carried out to compare two screening techniques under greenhouse conditions and estimate the association between the results obtained for both bioassays with grain yield. Nutrient solution (NS) and pots with acid soil (AS) were utilized as screening methodologies to perform one cycle of divergent selection for aluminum tolerance in the tropical maize population SIKALQ. The four sub-populations obtained by both approaches were compared with the original and checks in a greenhouse and in multilocation field trials. Evaluation under the conditions of NS and AS showed that the populations with the best performances were those selected for aluminum tolerance under the same condition of evaluation. The variables measured in greenhouse showed good correlation and the most closely related were fresh root weight (FRW) and dry rot weight (DRW) (0.79). All of these variables showed low correlation with yield in non acid conditions ( 0.30). The correlation was more important (~ 0.50) as soil aluminum saturation increased. The best correlation was observed for visual scoring (VS) (0.68), FRW (0.47) and net root growth (NRG) (0.52).

O melhoramento genético tem se mostrado como estratégia eficiente para melhorar a tolerância ao alumínio em milho. O objetivo deste trabalho foi comparar dois metodos de seleção em casa de vegetação e estimar a associação entre os resultados obtidos em cada um desses ensaios com a produção de grãos no campo. Solução nutritiva (NS) e vasos com solo ácido (AS) foram utilizadas nas metodologias de seleção para realizar um ciclo de seleção divergente para tolerância ao alumínio na população de milho tropical SIKALQ. As quatro subpopulações obtidas foram comparadas com a original e as testemunhas em experimentos de casa de vegetação e de campo. A avaliação sob as condições de NS e AS mostraram que as populações com melhor comportamento foram aquelas selecionadas para tolerância ao alumínio nas mesmas condições da avaliação. As variáveis medidas na casa de vegetação apresentaram boa correlação e as mais fortemente relacionadas foram peso fresco da raiz (FRW) e peso seco da raiz (DRW) (0.79). Todas as variáveis apresentaram baixa correlação com a produção de grãos em solos sem acidez ( 0.30). A correlação foi mais importante (~ 0.50) na medida que a saturação com alumínio do solo foi aumentando. Os melhores valores de correlação foram observados para avaliação visual (VS) (0.68), FRW (0.47) e crescimento líquido da raiz (NRG) (0.52).

Field comparison between selection methods at the maize seedling stage in relation to aluminum tolerance

Selection and breeding for aluminum tolerance is a useful approach to increase maize (Zea mays L.) grain yield in acid soils. The objective of this work was the comparison between two screening approaches for the development of Al tolerant maize populations. One cycle of divergent selection for aluminum tolerance, using both nutrient solution and pots with acid soil, were completed in the maize population SIKALQ. The four sub-populations selected through both selection procedures were compared with the original population and checks in field trials repeated over locations. Eight environments stratified into four groups varying from non acid to high aluminum saturation were used. Three variables were analyzed for comparisons of methodologies: grain yield, plant height and days to male flowering. The populations performed similarly in the high productivity locations and the differences between tolerant and sensitive ones were more evident for increasing levels of toxic aluminum. In spite of little differences, selection in pots with acid soil was more efficient than nutrient solution in changing the genetic structure of the population toward more tolerant and sensitive levels.

A seleção e o melhoramento visando tolerância ao alumínio é um procedimento valioso para aumentar a produção de milho (Zea mays L.) em solos ácidos. O objetivo deste trabalho foi comparar a eficiência de duas metodologias de seleção de plântulas, no desenvolvimento de populações de milho tolerantes ao alumínio. Foi realizado na população SIKALQ, um ciclo de seleção divergente para tolerância ao alumínio, utilizando solução nutritiva e vasos com solo ácido. As quatro sub-populações selecionadas pelas duas metodologias foram comparadas com a população original e testemunhas em experimentos de campo em vários locais. Foram utilizados oito locais, estratificados em quatro grupos começando com solos não ácidos até com alta saturação com alumínio. As comparações foram realizadas utilizando três variáveis: produção de grãos, altura de planta e dias até florescimento. As populações apresentaram um comportamento semelhante entre elas nos locais de alta produtividade, e as diferenças entre as tolerantes e suscetíveis ficaram mais evidentes na medida em que aumentou a toxidez por alumínio. Os resultados sugerem que a seleção em vasos com solo ácido foi mais eficiente em alterar a estrutura genética da população do que a solução nutritiva.

Inbreeding in two maize subpopulations selected for tassel size

The present work was based on the evaluation of full sib and S1 progenies from the maize (Zea mays L.) population ESALQ-PB1 after six cycles of divergent selection for tassel size. The following traits were analyzed: plant height , ear height, tassel branch number, tassel length, and tassel weight. For all traits, the experimental units were means of three plants per plot. Progenies were evaluated in eleven experiments (completely randomized blocks) with three replications in Piracicaba -- SP, Brazil. Means of inbred (m1) and non-inbred (m0) progenies were used to estimate inbreeding depression (I = m1 -- m0) and the contribution of homozygotes (mu0 + a*) and heterozygotes (d*) to the population mean. Considering the five traits under study, inbreeding depression varied from 1.9 to 15.9% but significance for that effect was detected only for plant height. Plant traits exhibited higher depression than tassel traits; and tassel branch number seems to be more sensitive to inbreeding than the other two tassel traits. Except for plant height, it was apparent that inbreeding depression was higher in the subpopulation negatively selected for tassel size (T--). Estimates of A = mu0 + a* and d* indicate a smaller effect of dominance deviations for all traits, as compared to the contribution of homozygotes. Significant variability among progenies was detected in most instances.

O presente trabalho foi baseado na avaliação de progênies de irmãos germanos e S1 da população de milho (Zea mays L.) ESALQ-PB1 depois de seis ciclos de seleção divergente para tamanho do pendão. Os seguintes caracteres foram analisados: altura da planta, altura da espiga, número de ramificações do pendão, comprimento do pendão e peso do pendão. Para todos os caracteres, as unidades experimentais foram médias de três plantas por parcela. As progênies foram avaliadas em onze experimentos (blocos completamente casualizados) com três repetições em Piracicaba -- SP, Brasil. As médias das progênies endogâmicas (m1) e não endogâmicas (m0) foram usadas para estimar a depressão por endogamia (I = m1 -- m0) e a contribuição de homozigotos (mi0 + a*) e heterozigotos (d*) para a média populacional. Considerando os cinco caracteres sob estudo, a depressão por endogamia variou de 1,9 to 15,9% mas somente para altura da planta foi detectada significância para aquele efeito. Os caracteres da planta exibiram maior depressão do que os caracteres do pendão; e o número de ramificações do pendão parece ser mais sensível à endogamia do que os outros caracteres do pendão. Com exceção de altura da planta, foi evidente que a depressão por endogamia foi maior na subpopulação selecionada negativamente para tamanho do pendão (T--). As estimativas de A = mi0 + a* e d* indicam um menor efeito dos desvios de dominância para todos os caracteres, quando comparados com a contribuição dos homozigotos. Foi detectada variabilidade significativa entre progênies na maioria dos casos.

Inbreeding in two maize subpopulations selected for tassel size

The present work was based on the evaluation of full sib and S1 progenies from the maize (Zea mays L.) population ESALQ-PB1 after six cycles of divergent selection for tassel size. The following traits were analyzed: plant height , ear height, tassel branch number, tassel length, and tassel weight. For all traits, the experimental units were means of three plants per plot. Progenies were evaluated in eleven experiments (completely randomized blocks) with three replications in Piracicaba -- SP, Brazil. Means of inbred (m1) and non-inbred (m0) progenies were used to estimate inbreeding depression (I = m1 -- m0) and the contribution of homozygotes (mu0 + a*) and heterozygotes (d*) to the population mean. Considering the five traits under study, inbreeding depression varied from 1.9 to 15.9% but significance for that effect was detected only for plant height. Plant traits exhibited higher depression than tassel traits; and tassel branch number seems to be more sensitive to inbreeding than the other two tassel traits. Except for plant height, it was apparent that inbreeding depression was higher in the subpopulation negatively selected for tassel size (T--). Estimates of A = mu0 + a* and d* indicate a smaller effect of dominance deviations for all traits, as compared to the contribution of homozygotes. Significant variability among progenies was detected in most instances.

O presente trabalho foi baseado na avaliação de progênies de irmãos germanos e S1 da população de milho (Zea mays L.) ESALQ-PB1 depois de seis ciclos de seleção divergente para tamanho do pendão. Os seguintes caracteres foram analisados: altura da planta, altura da espiga, número de ramificações do pendão, comprimento do pendão e peso do pendão. Para todos os caracteres, as unidades experimentais foram médias de três plantas por parcela. As progênies foram avaliadas em onze experimentos (blocos completamente casualizados) com três repetições em Piracicaba -- SP, Brasil. As médias das progênies endogâmicas (m1) e não endogâmicas (m0) foram usadas para estimar a depressão por endogamia (I = m1 -- m0) e a contribuição de homozigotos (mi0 + a*) e heterozigotos (d*) para a média populacional. Considerando os cinco caracteres sob estudo, a depressão por endogamia variou de 1,9 to 15,9% mas somente para altura da planta foi detectada significância para aquele efeito. Os caracteres da planta exibiram maior depressão do que os caracteres do pendão; e o número de ramificações do pendão parece ser mais sensível à endogamia do que os outros caracteres do pendão. Com exceção de altura da planta, foi evidente que a depressão por endogamia foi maior na subpopulação selecionada negativamente para tamanho do pendão (T--). As estimativas de A = mi0 + a* e d* indicam um menor efeito dos desvios de dominância para todos os caracteres, quando comparados com a contribuição dos homozigotos. Foi detectada variabilidade significativa entre progênies na maioria dos casos.