GENETIC GAIN FROM DIFFERENT SELECTION METHODS IN Eucalyptus macarthurii PROGENIES IN DIFFERENT ENVIRONMENTS

GANHOS GENÉTICOS POR DIFERENTES MÉTODOS DE SELEÇÃO EM PROGÊNIES DE Eucalyptus macarthurii EM DIFERENTES AMBIENTES

ABSTRACT

The aims of this research was to estimate genetic parameters of Eucalyptus macarthurii progenies and to predict genetic gain for different selection methods. In order to evaluate the gains, eleven progenies tests were studied. These tests were installed in 1997, in randomized blocks design, in two sites, with 5 replicates of five plants per plot. The progenies were evaluated, from the first to the eighth year, regarding the variable diameter at breast height (DBH). The genetic parameters were estimated using the mixed model procedure (REML/BLUP). The selection of the progenies for establishing a Clonal Seed Orchard was based on the analysis by location, joint analysis and on the harmonic mean of the relative performance of genotypic values (HMRPGV) with different selection intensities among and within families: three, four, five and six individuals per progeny with the selection of the top 30 individuals, taking into consideration the effective population size (N e ). The best age to carry out the selection was seven years old, however, early selection can be made from age two onwards. Each environment characteristics influenced progenies phenotypic expression, so the higher gains in balance with N e , were estimated with the selection of individuals for each site, and the selection of five or six individuals per progeny.

Keywords:
Genetic parameters; Progeny test; Clonal seed orchard; REML/BLUP; HMRPGV

RESUMO

O objetivo desta pesquisa foi estimar parâmetros genéticos de progênies de Eucalyptus macarthurii e estimar ganhos genéticos por diferentes métodos de seleção. Foram utilizados testes de onze progênies, instalados em 1997 em blocos ao acaso, em dois ambientes, com 5 repetições de 5 plantas por parcela. Os indivíduos foram avaliados desde o primeiro ao oitavo ano, quanto à variável diâmetro à altura do peito. Os parâmetros genéticos foram estimados pela metodologia de modelos mistos (REML/ BLUP). A seleção de progênies para formação de Pomar Clonal de Sementes baseou-se na análise por local, análise conjunta e no método da média harmônica do desempenho relativo dos valores genéticos (MHPRVG) com diferentes intensidades de seleção entre e dentro de famílias: três, quatro, cinco e seis indivíduos por família, com seleção dos 30 melhores indivíduos, considerando-se o tamanho efetivo populacional (N e ). A idade mais indicada para se realizar a seleção foi aos sete anos de idade, porém uma seleção precoce pode ser realizada a partir dos dois anos. As particularidades de cada ambiente influenciaram na expressão fenotípica das progênies, de modo que os maiores ganhos, em equilíbrio com o N e , foram estimados com a seleção dos indivíduos para cada local, sendo a seleção entre cinco e seis indivíduos por progênie.

Palavras chave:
Parâmetros genéticos; Testes de progênies; Pomar clonal de sementes; REML/BLUP; MHPRVG

INTRODUCTION

Some species of the genus Eucalyptus have great genotypic plasticity in Brazil, growing satisfactorily in wide an edaphoclimatic amplitude. However, the species show fundamental differences regarding the responses to environmental stimuli and it is therefore indispensable to run tests evaluating their ability to adapt in each desired location (FERREIRA, 1992). In this context, the selection of superior genotypes for genetic improvement is fundamental to generate more productive forests which are suited to different regions, having greater resistance to adverse environmental conditions (ODA et al., 2007; ROSADO et al., 2012). Eucalyptus macarthurii H. Deane & Maiden is endemic to New South Wales, Australia and this species is found near the swamps and creeks, growing in damp and cold locations (BROOKER; KLEINIG, 1999). In its place of origin, the species can be found from 700 m up (BENSON; MCDOUGALL, 1998; LITTLE; GARDNER, 2003).

In the South of Brazil, in higher altitude regions, this species has shown rapid growth and high tolerance to adverse climatic conditions, being a promising species for breeding programs aiming at its potential usage, as timber for sawmills, pulp, biomass source and properties for medicinal oil production (PIRRAGLIA et al., 2012), with an average volume over bark of 182.44 m³/ha and average annual increment of 36.49m³/ha/year for 5 years old plantations in the municipality of Lages, SC, Brazil (FONSECA et al., 1979). However, today, there is a need for studies about its productivity, behavior and quality of wood in Brazil.

To grow eucalyptus forests, which are more productive and adapted to different regions as well as meeting market requirements, it is important to consider the genetic aspects, once the forest is the result of actions and interactions of their genotypes with the environment (MARTINS et al., 2005). Therefore, it is advisable to evaluate its performance in more than one location so that the effect of interaction can be estimated and attenuated (SILVA et al., 2011). Estimates of genetic parameters and prediction of gains from progenies tests in several locations have been a great help in defining strategies to improve with the selection and/or recombination of individuals and progenies for a new selection cycle.

For forest species genetic improvement programs, time is a critical factor, because the exploitation involves long time consuming and expensive cycles (BELTRAME et al., 2012). Thus, the practice of selecting superior genotypes based on the first years of production, also known as early selection, is an advantageous alternative because it aims at decreasing the time needed to complete an improved generation (FONSECA et al., 2010).

In addition to these strategies, which aim at the genetic gain with greater accuracy and more precociously, studies of the parameters of adaptability and phenotypic stability of the genotypes have been a great contribution to this aspect. These provide information about the behavior of each genotype in several environmental conditions with the identification of genotypes which are more responsive to environmental variations (CRUZ et al., 2012). Thus, several methods of selection have been employed in Eucalyptus, with emphasis to the procedure REML/BLUP effects (restricted maximum likelihood/best linear unbiased prediction) (ROCHA et al., 2006, 2007), as being the best procedure for generating estimates of variance components, with the maximization of selective accuracy (FARIAS NETO et al., 2009).

The major objective for the selection of individuals and progenies is to generate populations that focus on producing improved seeds with greater genetic gains. One of the strategies for the production of improved seed is the clonal seeds orchard - CSO, which presents the advantage of producing seeds earlier, allowing high selection intensity and low possibility of inbreeding.

Based on these statements, this study has as objective to indicate the most appropriate age and determine the best method for progenies selection of Eucalyptus macarthurii individuals, as well as to estimate the genetic gains for the formation of a production population (CSO-clonal seed orchard) for Três Barras and Timbó Grande, SC.

MATERIAL AND METHODS

In total, eleven progenies of Eucalyptus macarthurii open-pollinated progenies were used, with seeds from Pietermaritzburg, South Africa provenance, imported from the Institute for Commercial Forestry Research (ICFR). The tests were implemented in October 1997, in two locations in the north of Santa Catarina: Três Barras and Timbó Grande, in randomized blocks design, with 5 replicates and 5 plants per plot, in a spacing of 2.5m x 3.0m. Seeds of the species E. dunnii and E. nitens were added as witnesses. The silvicultural practices were performed according to the characteristics required for each environment.

The geographical location, altitude and edaphoclimatic conditions of the two sites, are presented in Table 1. The climate for the two regions is classified according to Köppen as Cfb, humid mesothermic, with mild summer and uniformly distributed rainfall, without a dry season and with average air temperature in the cooler months between -3 ° C and 18 ° C and average air temperature in the warmest month of about 22 ° C (ALVARES et al., 2013).

TABLE 1
Geographical location, altitude, edaphoclimatic conditions and soil types (according to the Brazilian System of Soil Classification) of the progenies test sites.

The individuals were evaluated every year, from one to eight years old, with measurements of total height (H, m), except in the first year, and diameter at breast height (DBH, cm). The individual volume was estimated by the equation 1, where, VOL = individual volume with bark and ff is the form factor of k 0.45.

[1]

The estimation of genetic parameters and the correlation between the variables of measured growth was performed by location and for each age, using the software SELEGEN (RESENDE, 2007), which uses the best linear unbiased prediction (BLUP effects) and restricted maximum likelihood (REML). We also carried out genetic correlation between the variable DBH for ages two to seven years old, for possible early selection of individuals. The mixed linear model employed was the following (equation 2), where y is the vector of fixed effects data (averages of blocks through the locations, r), individual genetic additive effects (random, a) and errors or residuals (random, p); and X, Z and W are the matrixes of incidence for r, G and P, respectively.

[2]

For the study of the interaction between Genotype versus environment the following model was used equation 3, where: i is the vector of the genotype versus environment interaction effects and T its respective matrix of incidence.

[3]

Therefore, for the selection of the progenies considering simultaneously productivity, stability and adaptability, the harmonic mean relative performance of genetic values (HMRPGV) predicted by BLUP method was used based on the work by Resende (2004; 2007). The stability corresponds to the Harmonic Means of Genotypic Values (HMGV) through the locations; adaptability, could then be said as the relative performance of the genotypic values in relation to the average of each site (RPGV) and the stability and adaptability simultaneously (Productivity, represented by the character DBH) corresponds to the harmonic mean of the relative performance of genotypic values (HMRPGV). In this way, the statistical model used (equation 4) was, where: g is the vector of random individual genotypic effects.

[4]

The genetic parameters between the different ages were compared to check which age is the most indicated to perform the selection, assuring greater genetic gain. This way, the selection of individuals to be recombined into a production population (CSO-clonal seed orchard), was estimated for the best age. For this, individual selection simulations were made using different methods, through different intensities of selection, with the selection of the best 30 individuals: Method 1 - selection of best three individuals within families; Method 2 - Selection of the four best individuals within families; Method 3 - Selection of the best five individuals within families; and Method 4 - selection of six best individuals within families.

The selection genetic gain in percentage (g%) and the effective population size were estimated according to Resende and Bertolucci (1995), where: Kf = average number of individuals selected by progeny; σ² _kf = variance of the number of individuals selected by progeny; N² _f = number of selected progeny; where: SG is the selection gain; is the average of the 30 selected plants; is the average of the evaluated plants; and is the individual heritability in the narrow sense.

[5]

[6]

RESULTS AND DISCUSSION

Both variables (diameter and total height) were highly genetically correlated to volume, over 84%. The diameter, in all analyses and at different ages, presented the greatest genetic correlation to volume (between 90% and 96%). Silva (2008), found similar results in progeny tests of E. benthamii. And a great number of studies of this genetic association in arboreal species have shown high genetic correlation between diameter and volume (SEBBENN, 2009; FREITAS et al., 2008). Rocha et al. (2007) concluded that diameter is the most used variable due to its easy evaluation and accuracy. Thus, this was the variable used to estimate the genetic parameters, as well as the selection gains of the progenies test.

The genetic parameters analyzed for the variable diameter, for each site and for the joint analysis, in every age, are presented in Tables 2, 3 and 4. For the analysis of the progenies in Três Barras (Table 2), the heritability ranged from 32% to 78%, increasing with the age, with a slight decrease at age 8.

TABLE 2
Genetic parameters of variable DBH at each age, from 1 to 8 years old, for E. macarthurii progeny test in the municipality of Três Barras, SC.

• Where: Vg: additive genetic variance; Vparc: environmental variance between plots; Vf: individual phenotypic variance; h2a = h2: individual heritability in the narrow sense, i.e. of additive effects; h2aj: individual heritability in the narrow sense, adjusted for the effects of plots; c2parc = c2: coefficient of determination of the plots effects; h2mp: heritability of average of progenies, assuming full survival; Acprog: Accuracy of selection, assuming full survival; h2ad: additive heritability within the plot; CVgi%: individual additive genetic coefficient of variation; CVgp%: coefficient of genotypic variation between progenies; CVe%: coefficient of variation; CVr = CVgp/CVe = coefficient of variation concerning residuals; ENP: variance of the genotypic values of progenies error of prediction, assuming full survival; SEP: standard deviation of the genotypic value of progenies predicted, assuming full survival.

In Timbó Grande (Table 3), the values of heritability (h2a) ranged from 9% to 43.8% of from lower to higher age. These results were considered high in the older ages, according to Simeon et al. (2002) classification for the variable diameter, except for values found in the joint analysis (Table 4), which varied from 5%, for the age of 2 years, to 15.8% for 8 years. Garcia and Nogueira (2005) affirm that the estimates of heritability coefficient, high and positive, show good genetic control in the expression of the characters and show great potential for selection within the experiment, with good prospects for genetic progress. Similar results of high heritability for the variable diameter were found for E. resinífera (Sato et al. 2007), E. pellita (Zanata et al., 2010) and E. grandis (Miranda et al., 2015).

TABLE 3
Genetic parameters of variable DBH at each age, from 1 to 8 years old, for E. macarthurii progeny test in the municipality of Timbó Grande, SC.

• Where: Vg: additive genetic variance; Vparc: environmental variance between plots; Vf: individual phenotypic variance; h2a = h2: individual heritability in the narrow sense, i.e. of additive effects; h2aj: individual heritability in the narrow sense, adjusted for the effects of plots; c2parc = c2: coefficient of determination of the plots effects; h2mp: heritability of average of progenies, assuming full survival; Acprog: Accuracy of selection, assuming full survival; h2ad: additive heritability within the plot; CVgi%: individual additive genetic coefficient of variation; CVgp%: coefficient of genotypic variation between progenies; CVe%: coefficient of variation; CVr = CVgp/ CVe = coefficient of variation concerning residuals; ENP: variance of the genotypic values of progenies error of prediction, assuming full survival; SEP: standard deviation of the genotypic value of progenies predicted, assuming full survival.

TABLE 4
Genetic parameters of variable DBH at ages 1 to 8 years old, E. macarthurii progenies test for the joint analysis of both sites Três Barras and Timbó Grande, SC.

• Where : Vg: additive genetic variance; Vparc: environmental variance between plots; Vf: individual phenotypic variance; h2a = h2: individual heritability in the narrow sense, i.e. of additive effects; h2aj: individual heritability in the narrow sense, adjusted for the effects of plots; c2parc = c2: coefficient of determination of the plots effects; h2mp: heritability of average of progenies, assuming full survival; Acprog: Accuracy of selection, assuming full survival; h2ad: additive heritability within the plot; CVgi%: individual additive genetic coefficient of variation; CVgp%: coefficient of genotypic variation between progenies; CVe%: coefficient of variation; CVr = CVgp/CVe = coefficient of variation concerning residuals; ENP: variance of the genotypic values of progenies error of prediction, assuming full survival; SEP: standard deviation of the genotypic value of progenies predicted, assuming full survival.

The accuracy values increase with the age of the individuals, being considered high (over 70%) from the first year of age onwards, except for the joint analysis. In older ages these values were higher (over 90%). In a genetic variation study of E. camaldulensis, the accuracy found was high for diameter, height and taper with values ranging from 85% to 89% (Moraes, et al., 2007). The accuracy of selection indicates the likelihood and reliability to reach the expected results (Pires et al., 2011).

The genetic correlation between the performance of the progenies in the different sites, as shown in Table 4 for joint analysis, had results below 27%, except in the first year, which indicates low performance of the progenies considering a single population for both locations. Similar results were found in the work by Santos et al. (2015), for Eucalyptus clones in Rio Grande do Sul. According to Resende (2002), only when the genetic correlation value between locations is over 70% it is possible to carry out a single improvement population. The low genotypic performance correlation (27%) indicates complex type genotype x environment interaction. Therefore, the performance of the progenies is differentiated based on the variations of the environment, because when subjected to different environments, the performance of one progeny compared to other varies, so that progeny which are superior in one environment may not be in another (PIRES et al., 2011). This result also indicates the importance of studying the ability of stability and adaptability of the progenies in different places of this research and for each location where genetic testing of the species is warranted.

It was found that the variable diameter at ages two, four, five and six years is highly correlated with the diameter at 7 years old (over 0.70), for both locations, except for the age two years in Timbó Grande (Table 5). Thus, it is possible to perform early selection from two to three years of age for E. macarthurii. In a study conducted by Sato et al. (2010), with progenies of Corymbia maculata, correlations between growth traits at ages 4 and 21 years were also significant, positive and high, suggesting that early selection at age 4 could be made. This is possible because of the variables DBH, height and volume present significant correlations among themselves, in different ages, which enables the efficiency in the early selection of genetic material (MASSARO et al., 2010). Therefore, for each location analyzed separately, the best age to perform the selection of progenies test would be at seven years old, for presenting high values of heritability and accuracy of selection, however with the possibility of early selection from two years of age.

TABLE 5
Genetic correlations of the variable DBH between the age of seven years and other ages, from two to six years old, for both locations, Timbó Grande and Três Barras, SC.

The different selection strategies of progenies were carried out based on the performance analysis of the progenies in two separate locations and joint analysis, considering the results of the genetic gain with the yield, stability and adaptability of progenies. The average performance in both locations selected the best seven families with genotypic values free of interaction (Table 6).

TABLE 6
Estimates of genetic gain predicted for the DBH in E. macarthurii, progenies at age 7 years, considering the average performance in both location Três Barras and Timbó Grande, SC.

• Where: g: genotypic effect; u + g: genotypic predicted value; u + g + gem: average genotypic value in the environments.

There was coincidence of 100% for the ordering of the seven most productive progenies by criteria of simultaneous adaptability and stability (HMRPGV) and average productivity (Table 6 and 7). The value of HMRPGV x general average value is the genotypic progenies average in the studied locations, considering the instability as a negative factor and the adaptability as a positive one (CARBONELL et al., 2007, MAIA et al., 2009). Therefore, these seven progenies might be recommended for planting in both areas, following the interaction pattern of the experimental network, contemplating three attributes: production, stability and adaptability.

TABLE 7
Stability and adaptability (MHRPGV and MHRPGV*MG) of genotypic values of E. macarthurii families for the variable DBH at 7 years old.

• In that: HMRPGV: harmonic means of the relative performance of the geno typic values; HMRPGV*MG: refers to the general average HMRPGV multi plied in all environments.

This way, it is necessary to compare the prediction of genetic gains between analysis for each site, by joint analysis and by yield, stability and adaptability (HMRPGV) (Table 8). The genetic gain, with the average of the harmonic mean relative performance of genetic values (HMRPGV), was larger than only by joint analysis. However, the genetic gain obtained with the selection was even greater in the analysis by location, between 36.2% and 43.9% for Três Barras and 18.8% to 24.4% for Timbó Grande.

TABLE 8
The genetic gain predicted for the variable diameter and effective population size by different selection methods, analysis by location, joint analysis and HMRPGV interaction analysis for E. macarthurii, at 7 years old.

• where: G%: genetic gain in percentage; Nf: number of families selected; Kf: average number of individuals selected per family; Ne: effective population size.

The genetic gain increases with the reduction of the selection intensity within progenies, consequently reducing the number of families selected. It is important to determine the number of individuals for recombination of the production population, the appropriate effective population size to ensure safety in the selection considering the inbreeding in the generation of planting. Therefore, the intensity of selection that ensures a balance between the genetic gain and the effective population size (Ne) is between five and six individuals per family in the selection of 30 individuals with higher genetic value in the population, with Ne ranging from 15.9 to 17.8 for the two sites. In the studies by Rocha et al. (2006a), with E. urophylla progeny test, similar results were found, twenty one progenies were selected with genetic gain of 21.22% and a Ne of 18.3.

CONCLUSION

There is high genetic correlation between the variables diameter at breast height total height and volume over bark. The ideal age for performing the selection in E. macarthurii stands is at seven years old, with higher value of heritability and genetic gain, being possible however to perform early selection starting at two years of age, assuring high selection reliability.

The genetic correlation between the performance of the progenies in the different environments was low, thus, the genetic gain obtained with the selection is greater when raising a production population (CSO) in each location.

The intensity of selection for the formation of a CSO, which ensures balance between the genetic gain and the Ne, lies between five and six individuals per family in the selection of the 30 individuals with higher genetic value in the population.

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Publication Dates

• Publication in this collection
  Jul-Sep 2016

History

• Received
  05 July 2016
• Accepted
  29 Aug 2016