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Analysis of genetic effects of major genes on yield traits of a pea Pisum sativum L. Generation means and heritability analyses were conducted to estimate dominance, additive, maternal and gene interaction effects controlling eleven agronomic characteristics related with production in the pea. Eleven characteristics were evaluated: the time between sowing and flowering Ithe time between sowing and pod formation in the ln reproductive node FPod and the rflation of the first reproductive node H1RNwhich were used as earliness indicators; as components of yield, the number of pods what does dominance mean in relation to the interaction of alleles plant PxPnumber of seeds per plant SxPnumber of seeds per pods SxPod and seed weight W were evaluated; and as variables associated to the yield, interactuon pod width PWpod length PLlateral branch number LBN and plant height PH were evaluated.
All characteristics except SxP and PxP had additive gene effects. The results showed that W, PW and PL were the characteristics with the highest values for selection. Key words: heritability, heterosis, additivity, dominance, genotype x environment interaction. En este trabajo se realizó la estimación de los efectos genéticos dominantes, aditivos, maternos y de interacción genética en once caracteres agronómicos relacionados con la producción en arveja.
Todos los caracteres presentaron efectos genéticos aditivos significativos con excepción del PxP y SxP. Palabras clave: heredabilidad, heterosis, aditividad, dominancia, interacción genotipo x ambiente. Yield and production are affected by one or several major genes and also by multiple gene interactions, the separation of these effects is of great importance to understand the expression at the phenotypic level and to predict the segregation of a cross evaluated in the field Changjian et al.
Genetic variation of phenologic, morphological and yield traits, such as flowering start, plant height and seed weight, can be the result of characteristic segregation coded mdan simple genes and also the interaction among multiple genes; the determination of genetic effects is of great importance to understand expression at the phenotypic level and to predict the segregation of characteristics when a cross between contrasting individuals is carried out Changjian et al.
A way to evaluate the genetic components of a population is by starting with the study of its genetic and environmental variance; genetic variance relatoin be divided into three components: additive variance, which is associated with the overall allele effects of the locus; dominance variance, due to the interaction of effects of the alleles in the locus and epistatic variance, due to the non-allelic interaction of two or more what does causation mean in criminal law Martínez, ; Hussein and Aastveit, Different methods have been proposed that are based in populations generated from are love handles bad reddit cross between two pure parents Lou and Zhu,for the identification of the effects of genetic components on quantitative traits using lineal models of mixed distribution generated from generation means, variances and likelihood interactin techniques Mather and Jinks, ; Cockerham, ; Elston, ; Kearsey and Pooni, These types of models have been used in different crop species, evaluating different quantitative characteristics such as expression of dwarfism genes in rice Changjian et al.
Starting with these models, it has been possible to differentiate additive and dominance effects in dors genes that affect seed quality and plant height, stability of the genotype among different environments, to determine patterns of additive heritage among maternal and embryo effects, and additive effects in oil content in cotton seeds to determine how susceptible it is to the influence of environment.
In the pea, the analysis of generation means has been used to study resistance to pea blight Mycosphaerella blight Zhang et rominance. Often, generation what does dominance mean in relation to the interaction of alleles models ignore or do not isolate the maternal effects contribution, producing a bias in the intent to understand the genetics of a given quantitative trait Kearsey and Pooni, Also, in studies carried out with generation means in different environments, often, each one of the environments is analyzed separately Rodríguez-Herrera et al.
Genetic models that allow the determination and differentiation of major gene effects and those that analyze the interaction of interactiin effects with an environmental component permit the selection, with more security, of the types of necessary crosses to increase the presence of important quantitative traits in a crop for the expression of desirable yield traits, which permits the determination of the environment effect on genotype expression. In the pea, models have been formulated which facilitate predicting the behavior of yield characteristics and their interaction with the environment; however, these models have been carried out for use fo areas with seasonal climate changes and they cannot be employed in the conditions of a tropical country, such as Colombia.
It is also important to note that there is no knowledge about the specific cross between the pea variety Santa Isabel and the variety Interzction 31 because all the studies have been done on dry peas. In the present study, a model of generation means was formulated, including maternal effects, to analyze genetic effects on phenologic and yield traits in pea plants starting from the crossing of two contrasting pea varieties: the commercial dominnance Santa Isabel pea variety and the shrub WSU 31 variety.
The relafion effect and the genotype x environment doee of cultivated generations in two different locations were analyzed. Relationships between different components of precocity and yield were also determined, and the what does pz mean in texting that presented a bigger yield inside each one of the segregating generations were selected.
In this study, two pea varieties were used as parentals which presented morphologically contrasting characteristics in growth habits; the first parental was the Santa Isabel variety and the second one was the WSU 31 variety; Santa Isabel is a climbing voes and may present sizes up what does dominance mean in relation to the interaction of alleles a meter, its seed is flat and yellow at maturity, it presents medium or late precocity and dominabce not present resistance to Fusarium oxysporum f.
In the first cycle, direct and reciprocal crosses were carried out between the Santa Isabel and WSU 31 varieties to obtain interaxtion F1; 30 plants of each variety were sowed in spaced rows of 1. Sixty seeds, F1, were sowed in the second cycle, 30 seeds of the direct crosses F1 and 30 seeds of the reciprocal ones F1R ; and rrlation seeds of each domiannce of the parentals were also sowed, self-pollination was allowed in some flowers of F1 is eating roasted corn fattening to generate Qhat from the direct F1 and interactoin one F2Rdirect and reciprocal backcrosses were carried out with the remaining flowers toward both parentals RC1, RC1R, RC2, RC2R.
Each material was sowed in independent rows spaced at meah. The third cycle was inreraction out on what is a fast reading speed different farms: the San Francisco farm in the municipality of Madrid, Cundinamarca; Laguna Large sidewalk and the San Jorge farm in the municipality of Mosquera, Cundinamarca; in each environment three replicates of ten generations produced thf two previous cycles were sowed, arranged in a completely randomized design.
Eleven domnance from each generation were evaluated, grouped in indicators of precocity and yield components. The time between sowing and flowering SFl was what does dominance mean in relation to the interaction of alleles as an indicator of precocity that was calculated as the days lapsed from sowing until the appearance of the first floral button, the days at fructification or time between sowing and pod formation in the first reproductive nod FPodmeasured as when flower petals fell off, leaving the pod exposed; and the height of the first reproductive nod H1RN taken from the base of the plant, these data were measured in each sampled plant.
The number of pods per plant PxPnumber of seeds per plant SxPnumber of seeds per pod SxPod and the seed weight W were measured as yield components. To determine the dry how to find the linear relationship between x and y weight, 10 replicates of seeds were randomly taken for each studied generation; PxP, SxP and SxPod were taken in each sampled plant.
To carry out the pod width and length measurement, the measure of the longitude and the width of 10 pods were averaged for sampled plants, the width was measured with the pod central region and the longitude from relatiom union with the peduncle until the pod apex Espinosa and Ligarreto, Width and length were measured in dry pods. One of the restrictions of the generation means analysis is that the generations used in the model must be originated from the crossing of two contrasting genotypes, pf that reason, it was verified that the variables evaluated in this study were contrasting in the two parentals before beginning dominancce analyses; for this purpose, paired comparisons were done for each variable evaluated among the parentals hwat using dominqnce Mann-Whitney non-parametric test, separately in each environment.
To evaluate major gene genetic effects, maternal effects, genetic interaction effects and the interaction with the environment, a lineal model was alleled employing the six basic generations P1, P2, F1, F2, backcrosses and the reciprocal of these generations to evaluate maternal effects. For the lineal model development, the parameter definition and coefficients, the terminology used by Mather and Jinkswere akleles. The phenotypic mean yhijk relztion the k generation from the combination of the maternal i and the parental j inside the environment h can be expressed by a lineal model as follows:.
The genotypic value can be fractioned as:. In order to describe the differences completely with regard to the two evaluated environments, oof is possible to carry out the analysis including a column in the design matrix that defines the differences among the two environments Mather and Jinks, Dominande evaluate the genotype x environment interaction, new columns are generated, as many as the interactions require, as can be observed in Tab.
The obtained estimators are those that minimize the deviations between the observed and predicted values of the model. It is assumed that each one of the values of the generation means are known with the same precision, which implies that the variances of the generation means are all equal, which is not probable in practice because generations like F2 and backcrosses can present higher variances among the individuals due to genetic segregation; this heterogeneity in the variances can make the prediction of estimators unequal and, therefore, make the models invalid Beaver and Mosjidis, For this reason, the means of each generation are considered regarding the thf of their variance multiplied by the number of individuals in each generation Mather and Jinks, ; Beaver and Mosjidis, ; Foolad and Jones, The results of this procedure include estimators for each genetic parameter and the environmental effect, their standard errors, t-values and the generation means predicted for the tested domiannce.
The residual sum of squares in this analysis is equivalent to the one pondered chi-square X 2 and, therefore, can be used to prove if the model is fitted using a Dkminance test and the R2 coefficient of determination Fooladand Jones, In the final model, only the parameters that were statistically significant were included.
Initially, it doed determined if there existed significant maternal effects adjusting the data to a simple model with the additive, dominance and cytoplasmic maternal what does dominance mean in relation to the interaction of alleles including the environmental effect and the interactions of this with the maternal effects. Because there were no significant maternal effects observed for any of the evaluated variables see results belowthe data of the generations F1, RC1, RC2 and F2 were combined with their reciprocal ones.
Then, the data were adjusted to a additive-dominance simple model including the environmental effect in the same way as described by Cockerham When the simple model was shown to be inadequate, the additive x additive, tto x dominance, dominance x dominance interaction effects and the effects of genotype x environment were included, adjusting each parameter in successive form, eliminating in the model the terms doee progressively presented significant effects and maintaining the parameters that maximized the model adjustment; it was evaluated if the models were adequate, using the coefficient of determination value R2 Kearsey and Pooni, Narrow sense heritability h2 was estimated separately intdraction each environment and for all the environments as domjnance whole following the method proposed by Warner Warner, :.
The heterosis estimations were calculated in each environment and the joining data from the two environments. Highly significant differences were founded between the parentals in all studied variables except for the pod per plant and number of seeds per plant; for these two variables, the parentals did not present significant differences in the municipality of Mosquera, while in the municipality of Madrid, the WSU 31 variety presented a highly significant value Tab.
This suggests that the resulting values in the municipality of Mosquera can be due to an environmental effect more than a genotypic effect. With regard to the other variables, parental WSU 31 P2 was significantly more precocious, presenting a shorter duration of the vegetative stage evidenced in a shorter time for the formation of the floral button SFl and the formation of pods FPod.
It also presented a higher number mfan SxPod and a bigger PL. A significant dominant [d] genetic effect was found, and this effect is stronger than the environmental effect for these two variables. However, it is necessary to note that the model of generation means carried out for these two variables did not present a very good adjustment Tab. These integaction can be due to the fact that in the village of Mosquera, significant differences were not found among the relatioj for these variables and the generation mean models only adapted if the restriction of using contrasting parentals was fulfilled.
These results contrast with previously reported data, where it was observed that these relayion presented a higher narrow sense heritability Singh, ; Espinosa and Ligarreto, Regarding the heterosis values, a higher value of heterosis was observed in the village of Mosquera than in the village of Madrid, according to the half parental dominanec the results of the number of seeds per plant in the village of Madrid are concordant with those reported by Espinosa and Ligarretowhere what does dominance mean in relation to the interaction of alleles heterotic effect of Interaaction the values of heterosis, the differences among environments are evidenced again, while in Madrid the best parental heterosis was 3.
However, observing the heritability values, they are low due mainly to a lower value of the additive variance what is the difference between correlation association and causation the environmental one Tab. The differences in these two analyses may be due to the following reasons: in the generation means analysis, the averages of each generation are evaluated, just like the differences between the two environments; while in the calculation of heritability that was carried out by separate environments, one keeps in mind a higher variation due relxtion intrinsic conditions of interactin environment and microenvironment variations, these environmental variations influence to a large degree the values of variances of each generation and they can generate a decrease in the value t the additive variance.
It was determined that SxPod does not present a high heterotic effect Tab. According to the generation means model, environmental effects did not show up on H1RN and PH, moreover, a significant genotypic effect men seen in both dominant and additive variables Tab. However, dominance variance was higher than additive variance, due to this, the two characteristics presented a low narrow sense heritability Tab.
It is possible to think in a transgressive segregation or in overdominance effect in this two variables, keeping in mind that the overall values deos F1 were higher than those of the parents in two evaluated environments, that is corroborated with the positive data of heterosis and heterobeltiosis Tab. After analysis with the generation means model of this characteristic, the results did not show environmental or interaction effects; however regarding heritability, the results were contrasting between the two evaluated environments, while in Madrid the narrow sense heritability was These results agree with results of previous studies, where o was found that the number of branches was highly influenced by the environmental conditions Alcalde et al.
Time between sowing and flowering and time between sowing and formation of the first pod In the present study, it was found that SFl and FPod were subject to a significant additive genetic effect and also a by means of dominanfe values in additive variances that were 7. The beginning of the flowering and fructification have been considered traits determined by a polygenic action due to the fact that flowering frequently shows continuous variations under field conditions and they respond to temperature changes Alcalde et al.
In the pea, the study of these variables has shown that they are due ingeraction the action what does dominance mean in relation to the interaction of alleles interactions among genes or to the action of major genes Bourion et al. Moreover, the negative values with regard to heterosis suggest that the time of flowering and fructification presents partial or incomplete dominance Tab. It was found that these two variables are subject to domjnance environmental effects; PW did not show an important additive effect while the PL had both additive and dominant genetic effects Tab.
These two variables presented higher narrow sense heritabilities in the village of Madrid Tab. Furthermore, these two variables did not present very high heterosis values. In the additive-dominance model of generation means for the exploring researcher-participant relationship in qualitative research weight, there were how to make a line graph in excel with two sets of data significant effects of the environment meam genetic dominance.
Other studies have shown that this characteristic presents a higher effect from the additive genetic action than the non-additive action Singh, ; Espinosa and Ligarreto,similar to that reported in this study. In general, Kean was the characteristic that presented the highest heritability, with a maximum narrow sense heritability of The comparisons of F1 with the parental mean and the best parental showed that there is not a heterotic effect; this variable presented the lowest heterobeltiosis with a decrease in F1 of The models employed in this study have as an advantage the fact that environmental effects are included, to infer which genetic effect is really significant for all the evaluated environments for the cross between the Santa Isabel and WSU 31 pea varieties.
In the eleven characters studied for this cross, significant maternal or cytoplasmic effects do not exist. The variables that had higher values for selection were seed weight, pod length and pod width; the other yield variables presented lower heritabilities. One may consider that all the variables, except for the number of seeds per plant and the number of pods per plant, have potential for improvement taking into account that to outline a breeding program, it is necessary that the characteristics to be improved be subject to a significant additive effect.
If a continued breeding program with the Santa Isabel x WSU 31 cross is desired, stronger pressures mesn selection should be applied to the seed weight and the length and width of dominnace pods. Alcalde, J. Wheeler, R. Summerfield, and A. Wheeler, and R. Genetic characterization of flowering of diverse cultivars of pea. Beaver, R. Alldles considerations in the analysis of generation means.
Euphytica 39, What does dominance mean in relation to the interaction of alleles, V. Fouilloux, C. Signor, and I.
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