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CROP BREEDING, GENETICS & CYTOLOGY

Primitive Accession Derived Germplasm by Cultivar Crosses as Sources for Cotton Improvement

I. Phenotypic Values and Variance Components

^a USDA-ARS, Crop Science Research Laboratory, P.O. Box 5367, Mississippi State, MS 39762
^b Department of Plant and Soil Sciences, Mississippi State University, Mississippi State, MS 39762

^* Corresponding author (JMcCarty{at}msa-msstate.ars.usda.gov).

	ABSTRACT

TOP NOTES ABSTRACT INTRODUCTION MATERIALS AND METHODS RESULTS AND DISCUSSION REFERENCES

 
Cotton, Gossypium hirsutum L., is grown primarily for its spinnable fibers. To meet textile mill requirements and producer demands, both fiber quality and yield must be improved. This study was conducted to compare yield and fiber quality when exotic derived lines were crossed to cultivars. Fourteen germplasm lines with high fiber strength derived from selected primitive accessions were crossed as male parents to each of five cultivars. The F₂ populations and parents were grown at two different field locations in 1998 and 1999. Parents and F₃s were grown at two locations in 2000. Parents, F₂, and F₃ populations were evaluated for yield, yield components, and fiber quality traits. Combination of location and year was considered an environment for data analyses. An additive-dominance, additive x additive (ADAA), and genotype x environment interaction genetic model was used for data analysis. A mixed linear model approach, minimum norm quadratic unbiased estimation (MINQUE) was used to estimate genetic variance components on the basis of the ADAA model. Cultivars had higher yields and lint percentages than germplasm lines. Fiber strength for germplasm lines exceeded cultivars. Lint percentage, boll weight, micronaire reading, elongation, and fiber length were similar between F₂ and F₃ populations and near mid-parent values. Both additive and additive x additive (AA) epistatic effects significantly controlled all agronomic and most fiber traits. Significant dominance effects were detected for all traits except fiber elongation. Additive x environment and dominance x environment interaction effects were detected for most traits. Significant AA x environment interaction effects were detected for agronomic traits but not fiber traits. The additive x environment effect was significant; however, it made a small contribution to the total variance. This study suggest that crosses between day-neutral derived lines from photoperiodic primitive accessions and commercial cultivars may allow both the utilization of heterosis and the use of genetic variation for pure line development.

Abbreviations: A, additive effect • AA, additive x additive effect • ADAA, additive-dominance, additive x additive model • MINQUE, minimum norm quadratic unbiased estimation

	INTRODUCTION

TOP NOTES ABSTRACT INTRODUCTION MATERIALS AND METHODS RESULTS AND DISCUSSION REFERENCES

 
FIBER QUALITY has become increasingly more important to the textile industry in recent years because of technological changes in spinning. Many of the current high yielding, commercial, upland cotton cultivars do not possess the fiber quality desired by the textile industry. Cotton breeders are striving to meet the challenge to improve fiber quality. To improve the efficiency of developing a cultivar with high yield and acceptable fiber quality, it is important to have a better understanding of the genetic information for yield, its components, and fiber traits.

A review of research concerning the genetic properties of agronomic and fiber traits was provided by Meredith (1984). Many recent agronomic and fiber trait studies focused on combining ability by using cultivars as diallel parents (Meredith, 1990; Tang et al., 1993a, 1993b). These studies provided a good understanding of the genetic behavior of fiber traits in cotton. Because of the difficulty producing enough F₁ seeds for experiments across different environments, F₂ populations were often planted in field experiments (Meredith, 1990; Tang et al. (1993a)(1993b); Shoemaker, 2000; Cheatham, 2001). Traditionally, the data of F₂ and or their parents were analyzed by ANOVA methods. However, such analyses of F₂–bulk population means may be biased because of F₂ segregation. Recently, mixed linear model approaches have been used in cotton to estimate genetic variances and covariances and to predict genetic effects (Wu et al., 1995; Tang et al., 1996; McCarty et al., 1998). However, many studies were based on additive–dominance (AD) and its genotype x environment (GxE) genetic models (Meredith, 1990; Tang et al. (1993a)(1993b); Shoemaker, 2000; Cheatham, 2001).

Several genetic mapping studies of agriculturally important taxa (Doebley et al., 1995; Lark et al., 1995; Li et al., 1997; Eshed and Zamir, 1996; Cao et al., 2001; Liao et al., 2001; Lee et al., 2001; Wu, 2003) have provided evidence suggesting that epistasis may be an important genetic factor underlying complex traits. Most of these studies were with taxa other than cotton. The detection of epistasis effects may provide more information that will further our understanding of gene expression and interactions. The detection of epistasis is difficult because most studies lack the genetic structure to resolve genetic variances into sources other than additive and dominance effects. Zhu (1994) developed an ADAA model for detecting additive, dominance, and additive x additive epistasis effects. This method requires three generations, parent, F₁, and F₂. The flexibility of the mixed linear model approach not only allows the extensions of the use to other generations such as F₃ or F₄, but also allows unbalanced genetic designs in different environments. The utilization of such models should aid breeders in the development of improved cultivars.

Both cotton yield and fiber quality have been on a plateau in recent years and the introduction of genes from exotic sources is one approach to increase these important traits. A number of day-neutral lines derived from photoperiodic primitive accessions have been selected at Mississippi State, MS (McCarty et al., 2000, 2003). An understanding of the genetic properties associated with yield and fiber traits of these lines will be helpful in their utilization in breeding programs. To assess their breeding merit, 14 lines were used as male parents and crossed to each of five high-yielding cultivars. The objectives of this research were to determine agronomic and fiber property values and predict the genetic effects associated with 19 parental genotypes and 70 F₂ and F₃ populations. A companion paper (McCarty et al., 2004) provides prediction performances using these same crosses. The genetic information will help maximize efficiency of simultaneous selection for hybrids or pure lines with high fiber quality and yield.

	MATERIALS AND METHODS

TOP NOTES ABSTRACT INTRODUCTION MATERIALS AND METHODS RESULTS AND DISCUSSION REFERENCES

 
Plant Material and Experimental Design
Five cultivars used as female parents were crossed to each of 14 males in 1997. Female parents were 1. ‘Deltapine 50’ (DPL50), 2. ‘DES119’, 3. ‘Stoneville 474’ (ST474), 4. ‘Deltapine Acala 90’ (DPL90), and 5. ‘Sure-Grow 125’(SG125). The male parents were designated 6 through 19 and are as follows: 6. M75-1, 7. M1388-1, 8. M1388-2, 9. M1388-3, 10. M239-1, 11. M239-2, 12. M239-3, 13. M239-4, 14. M239-5, 15. M239-6, 16. M239-7, 17. M237-1, 18. M237-2, and 19. M237-3. Parent 6 was developed from a cross between ‘Deltapine 16’ (DPL16) and accession T75 (PI 549138), where day-neutral flowering habit was selected in the F₂. This selection was then backcrossed to T75. A single high strength, day-neutral plant was selected in the BC₁F₂, BC₁F₃, and BC₁F₄ generation. The BC₁F₄ plant was then advanced via bulk increase to the BC₁F₆. Parents 7 through 9 were developed from a cross between ‘DES56’ and the accession T1388 (PI 415112). A single high strength, day neutral plant was selected in the F₂ and F₃. Three high strength plants were selected in the F₄ and each was then advanced to the F₆. Parents 10 through 16 were from a cross between DPL16 and T239 (PI 163693), where day-neutral flowering habit was selected in the F₂. This day-neutral selection was then backcrossed three times to T239, each time selecting for the day-neutral flowering habit in the F₂. A single high strength, day-neutral plant was selected in the BC₃F₂. In the BC₃F₃, three high strength plants were selected and from these plants seven selections were made in the BC₃F₄. These selections were then advanced to the BC₃F₆. Parents 17 through 19 were from a cross between DPL16 and T237 (PI 163657), where day-neutral flowering habit was selected in the F₂. This selection was then backcrossed to T237. Three BC₁F₂ day-neutral plants were selected for length and strength. A single plant was selected from progeny of these three plants in the BC₁F₃ and BC₁F₄ generation. These three selections were then advanced to the BC₃F₆. Parents 7 through 9, 10 through 16, and 17 through 19 were sister lines selected from F2 individuals of the same cross (McCarty et al., 2003). Since some male parents were derived from the same primitive accession they may influence the genetic components. One way to check this relationship is to analyze the predicted genetic effects (for example, additive effects). This was done and the results showed that there were significant differences among male parents derived from the same accession.

The five female cultivars or their transgenic counter-parts are common cultivar types grown in the Mid South. These five cultivars, in general, have high yield and low fiber quality, with the exception of DPL 90 which has good fiber quality. In contrast, the 14 males were chosen to broaden the genetic base and use primitive lines that generally had excellent fiber strength.

Crosses and subsequent evaluations were conducted at the Plant Science Research Center, Mississippi State, MS (33.4 N, 88.8 W). F₁ and male parent seed were sent to a winter nursery to produce the F₂ and provide for seed increase. Seed from the 70 F₂ populations and the 19 parents (5 female cultivars and 14 males) were grown at two locations each year in 1998 and 1999. Seed was harvested from the 1999 test (F₂ bulks) and the resulting F₃–bulk populations and parents were grown at two locations in 2000. Generally less than 20% out crossing occurs in the area where the tests were grown, and thus should not have enhanced heterozygosity and potentially heterosis expressed in the F₃ populations.

The experimental design was a randomized complete block with four replicates at each location in each year. The combination of year and location (Loc) was considered as environments (Env) for the purpose of statistical analyses. The environments were as follows: Env 1 = 1998, Loc 1; Env 2 = 1998, Loc 2; Env 3 = 1999, Loc 1; Env 4 = 1999, Loc 2; Env 5 = 2000, Loc 1; Env 6 = 2000, Loc 2. Plot size for Environments 1, 2, 4, and 6 was a single row 12 m in length with row spacing of 0.97 m. Plot size for Environments 3 and 5 was a single row 9 m in length with row spacing of 0.97 m. The planting for Environment 1 was a two planted-one skip row pattern; whereas, other environments were planted in a solid row pattern. The stand density for all environments was one plant spaced approximately 10 cm apart. Environment 1 soil type was a Leeper silty clay loam (Fine, smectitic, nonacid, thermic Vertic Epiaquepts). Environments 2, 4, and 6 soil type were a Marietta silty clay loam (Fine-loamy, siliceous, active, fluvaquentic Eutrudepts). Environments 3 and 5 were Marietta loam (Fine-loamy, siliceous, active, fluvaquentic Eutrudepts). Standard production practices were followed at all environments.

A 25-boll, hand-harvested sample was collected from each plot before machine harvest. These samples were weighed and ginned on a laboratory 10-saw gin to determine boll weight, lint percentage and provide lint samples for fiber analysis. Lint samples were sent to STARLAB Inc., Knoxville, TN, for determination of micronaire reading, elongation (E1), fiber strength (T1), 2.5% span length (2.5% SL), 50% span length (50% SL). The plots were harvested with a mechanical picker, then the seed cotton was weighed and this data was used to calculate yields. Environment 1 in 1998 was not machine harvested because of extreme late season weather conditions. Yield data was collected for parents and F₂ populations from Environments 2, 3, and 4, and parents and F₃s from Environments 5 and 6. Fiber data for parents and F₂s was from Environments 1through 4, and parents and F₃s from Environment 5.

Genetic Models and Analysis Methods
Additive–dominance additive x additive (ADAA) and genotype x environment interaction genetic model was employed for data analysis (Zhu, 1994).

The mixed linear models were as follows:

Parents:

F2:

F3:

where A_i (or A_j) is additive effect from parent i(or j), A_i or A_j N; D_ii, D_jj, or D_ij is the dominance effect, D_ii, D_jj, or D_ij N; AA_ii, AA_jj, or AA_ij is the additive x additive (AA) epistasis effect, AA_ii, AA_jj, or AA_ij N AE_hi (or AE_hj) is additive x environment interaction effect, AE_hi N; DE_hii, DE_hjj, or DE_hij is the dominance x environment interaction effect, DE_hii, DE_hjj or DE_hij N; AAE_hii, AAE_hjj, or AAE_hij is the AA x environment interaction effect, AAE_hii, AAE_hjj, or AAE_hij N B_k(h) is the block effect with B_k N; e_hijk is the random error with e_hijk N.

A mixed linear model approach, minimum norm quadratic unbiased estimation (MINQUE) was used to estimate genetic variance components based on the ADAA model. The Jackknifing over blocks within environments was used to estimate standard errors of variance, and the predicted effects (Miller, 1974). The degrees of freedom were 19 and t tests were used for each parameter. Narrow-sense heritability across environments was defined as h²_N = /V_p, while broad-sense heritability across environments was defined as h²_B = /V_p; the narrow-sense heritability for genotype by environment interaction as h²_NE = /V_p, while broad-sense heritability for genotype by environment interaction was defined as h²_BE = /V_p (Zhu, 1998). The data set was analyzed by a computer program written in C++.

	RESULTS AND DISCUSSION

TOP NOTES ABSTRACT INTRODUCTION MATERIALS AND METHODS RESULTS AND DISCUSSION REFERENCES

 
Phenotypic Data Analysis
The female parents (cultivars) yielded more, had higher lint percentage but weaker fibers than the male parents (Table 1). Lint percentage for females was significantly higher than that for F₂ and F₃ populations, while mean lint percentage for F₂ and F₃ was significantly higher than that for male parents (Table 1). Since lint percentage is a major contributor to yield the performance of the females was as expected. The data for individual female, male and F₂–bulks by environment were reported by McCarty et al. (2003). Mean lint yield for females was greater than that for males in all environments, except Environment 6 (McCarty et al., 2003). The mean lint yield for the F₂ populations was between female and males, but it was greater than that of the mid-parent.

Variance Components Analysis
Estimated proportions of variance components to phenotypic variance for agronomic and fiber traits are summarized in Table 2. Both genotypic effects (A, D, and AA) and GxE interaction effects (AE, DE, and AAE) equally controlled lint yield; genotypic effects were more important than GxE interaction effects, based on total proportion of variance for lint percentage, boll weight, and fiber length GxE interaction effects were more important than major genetic effects for controlling micronaire reading, elongation, and fiber strength. The results suggested that lint percentage, boll weight, and fiber length were more stable than yield and other fiber traits across the different environments.

Dominance and dominance x environment effects were more important than other effects for lint cotton yield. Additive effects were more important than other effects for lint percentage. Both dominance and additive effects were more important than other genetic effects for boll weight, 50% span length, and 2.5% span length. Additive and dominance x environment effects mainly controlled micronaire reading and elongation, but dominance x environment effects appeared to be more important. Additive, dominance, and dominance x environment effects mainly controlled fiber strength, but dominance x environment effects appeared more important. Additive x environment interaction effects were detected for all agronomic traits but the magnitude was small relative to the total phenotypic variance. Therefore, neither additive effects nor AA effects were sensitive to different environments, while dominance effects were highly dependent on environmental conditions for lint yield, micronaire reading, elongation, and fiber strength. Generally, the proportion of the residual variance component was larger for fiber traits than for agronomic traits (Table 2). Tang et al. (1996) reported similar results. The small portion of lint sample that is used for fiber property determination may result in a large genetic sampling error since F₂ and F₃ populations are segregating.

Narrow-sense heritability across environments for lint yield and 50% span length was smaller than 10%, which was less than that for all other traits (Table 2). Lint percentage had relatively high narrow-sense heritability across environments. Lint percentage, boll weight, and 2.5% fiber span length had similar high broad-sense heritability estimates across environments, which were greater than 50%. Lint yield, fiber strength, and 50% span length gave similar broad-sense heritability across environments, which ranged from 35 to 40%. All traits except boll weight and lint percentage expressed very low narrow-sense heritability for genotype by environment interaction. All traits except boll weight and fiber span length gave large broad-sense heritability for genotype by environment interaction (30% or above). The term h²_N + h²_NE is defined as narrow-sense heritability for a specific environment, while h²_B + h²_BE is defined as broad-sense heritability for a specific environment. On the basis of the low narrow sense heritabilities estimated for a single environment, selection for lint yield, micronaire reading, elongation, fiber strength, and 50% span length would not be effective in early generations. However, the higher single environment narrow-sense heritability obtained for lint percentage indicated selection can be started at an early generation.

Prediction of genetic effects and genotypic values of germplasm derived from primitive lines may help cotton breeders decide the potential of their use for pure line selection. If the cumulative AA epistatic effects between parents are higher than within parents, then the average of genotypic values of advanced generations should be higher than the mid-parent genotypic values, and this heterosis can be fixed through selection. The detailed information on predicted effects and genotype values for different generations is addressed in a McCarty et al. (2004).

	NOTES

TOP NOTES ABSTRACT INTRODUCTION MATERIALS AND METHODS RESULTS AND DISCUSSION REFERENCES

 
Contribution of the USDA-ARS in cooperation with the Mississippi Agric. and Forestry Exp. Stn.

Received for publication July 14, 2003.

	REFERENCES

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This Article Abstract Full Text (PDF) Free Alert me when this article is cited Alert me if a correction is posted Services Related articles in Crop Science Similar articles in this journal Similar articles in ISI Web of Science Alert me to new issues of the journal Download to citation manager Citing Articles Citing Articles via HighWire Citing Articles via ISI Web of Science (9) Citing Articles via Google Scholar Google Scholar Articles by McCarty, J. C. Articles by Wu, J. Search for Related Content PubMed Articles by McCarty, J. C. Articles by Wu, J. Agricola Articles by McCarty, J. C. Articles by Wu, J. Related Collections Crop Genetics Cotton