# Genetic effects of thirteen Gossypium barbadense L. chromosome substitution lines in topcrosses with upland cotton cultivars: I. Yield and yield components.

UPLAND COTTON (Gossypium hirsutum L., 2n = 52) is the most extensively cultivated cotton species and has high lint productivity. It is an allelotetraploid with 13 pairs of chromosomes from subgenome A and 13 from subgenome D. The level of intraspecific polymorphism revealed by assessment at the DNA molecular level is low in G. hirsutum, especially among agriculturally elite types, (Gutierrez et al., 2002; Ulloa and Meredith, 2000; Wendel et al., 1989). Gossypium barbadense L., also a cultivated species, shares the same two subgenomes with G. hirsutum, and has superior fiber properties of length, micronaire, and strength. Interspecific germplasm introgression is particularly attractive as it utilizes natural resources and can be targeted to one or more specific traits or genes. However, crossing these two species usually leads to difficulties such as infertility, cytological abnormalities and distorted segregation in the [F.sub.2] generation and beyond.Chromosome substitution has been an indispensable method for intraspecific germplasm introgression into bread wheat (Triticum aestivum L., 2n = 42) and has been useful for genetic analysis and breeding (Al-Quadhy et al., 1988; Berke et al., 1992a, 1992b; Campbell et al., 2003; Campbell et al., 2004; Kaeppler, 1997; Law, 1966; Mansur et al., 1990; Shah et al., 1999; Yen and Baenziger, 1992; Yen et al., 1997; Zemetra and Morris, 1988, and Zemetra et al., 1986). Methods for development of interspecific chromosome substitution in G. hirsutum were outlined by Endrezzi (1963), and several of the initially discovered G. hirsutum monosomics were used to substitute G. barbadense chromosomes into G. hirsutum (Endrezzi, 1963; Kohel et al., 1977; Ma and Kohel, 1983). We have developed and released 17 disomic, chromosome substitution (CS-B) lines through hypoaneuploid-based backcross chromosome substitution, using as recurrent parent a previously developed monosomic or monotelodisomic near-isogenic backcross derivative of TM-1. In each CS-B line, a pair of chromosomes (or chromosome arms) of G. hirsutum inbred TM-1 was replaced by the respective pair from G. barbadense doubled-haploid 3-79 lines (Stelly et al., 2004a, 2004b, 2005). These substitution lines are near-isogenic to the recurrent parent TM-1 for 25 chromosome pairs, and are near-isogenic to one another, for 24 chromosome pairs. Such a highly uniform genetic background among these CS-B lines provides an opportunity to associate traits of importance with specific chromosomes or chromosome arms, in the TM-1 background using comparative analysis. In our previous research with only data from CS-B lines and TM-1 lines, we could not separate additive and epistatic effects (Saha et al., 2004a, 2004b). Study of several CS-B lines (Kohel et al., 1977; Ma and Kohel 1983) indicated that chromosome 6 was associated with higher lint percentage, finer fiber, and later flowering; whereas, chromosome 17 was associated with short fiber length. QTL for boll size, lint percentage, fiber length, and fiber elongation were mapped to chromosome 16 using 178 families from the cross of a substitution line for chromosome 16 and TM-1 (Ren et al., 2002). Recent analyses of an expanded set of new and resynthesized CS-B lines, per se, showed that chromosomes 16 and 18, from 3-79, were associated with reductions in yield, and chromosome 25 of 3-79 was associated with reduced micronaire and increased fiber length and strength compared with TM-1 (Saha et al., 2004a, 2004b). These studies provide good genetic information relative to genetic association of traits with substituted chromosomes or chromosome arms. However, the merit of these CS-B lines for breeding and genetic improvement of upland cultivars has not been investigated before this study.

We crossed 13 CS-B lines, the recurrent parent TM-1, and the donor parent 3-79 with five elite cultivars. The 75 [F.sub.2] hybrids and 20 parents were evaluated in four environments for yield traits. The objectives were to estimate variance components and genetic effects to determine which CS-B lines can be used as good combiners for improving cotton fiber quality and yield in breeding programs. In addition, we can determine specific chromosomes associated with important traits. In the present manuscript, we focus on yield and yield components.

MATERIALS AND METHODS

Development of Plant Materials

Five elite cultivars, Deltapine 90 (DP90); Sure-Grow 747 (SG 747); Phytogen 355(PSC 355); Stoneville 474 (ST 474), and FiberMax 966(FM 966); representing the major cotton seed breeding companies in the USA, were crossed as females with 13 CS-B lines (Stelly et al., 2004b, 2005), TM-1, and G. barbadense line 3-79 (Table 1). The CS-B lines include five subgenome A and eight subgenome D chromosomes or chromosome arms from 3-79 (G. barbadense) substituted into TM-1.

Field Design and Procedures

The 75 top crosses were made at Mississippi State, MS, in the summer of 2002. The [F.sub.1] seeds were sent to a winter nursery in Tecoman, Mexico, to produce the [F.sub.2]. The resulting 75 F2 hybrids, five cultivars, 13 CS-B lines, TM-1, and 3-79 parents were planted at two locations in 2003 and 2004 at the Plant Science Research Center at Mississippi State, MS (33.4[degrees] N 88.8'W). Soil type for Env. 1 was a Marietta loam (Fine-loamy, siliceous, active, fluvaquentic Eutrudepts) and for Env. 2, 3, and 4 was a Leeper silty clay loam (Fine, smectitic, nonacid, thermic Vertic Epiaquept). Plots were planted in a plant two skip-one row pattern with single 9 m rows in Env. 1 and 12 m rows in Env. 2, 3, and 4, replicated 4 times in randomized complete block arrangements. Rows were 0.97 m apart and plants were spaced approximately 10 cm apart within the row. Planting dates were 28 May 2003 for Env. 1 and 2 and 13 May 2004 for Env. 3 and 4. Harvest dates were 3 November for Env. 1, 31 October for Env. 2, 2 to 9 November for Env. 3, and 29 October for Env. 4. Standard cultural practices were followed in each environment. A 25-boll sample was hand harvested from first position bolls near the middle nodes of plants in each plot. This sample was ginned on a 10-saw laboratory gin, after which, the lint and seed fractions were weighed, and lint percentage was calculated by dividing lint fraction by total weight of seed and lint and multiplying by 100. After the boll sample was collected, all plots were harvested with a commercial cotton picker modified to bag seed cotton from each plot in 2003 while in 2004 the picker was equipped with load cells for weighing seed cotton per plot.

Data Analyses

Analysis of Phenotypic Data

Yield and yield component data were analyzed by analysis of variance using SAS procedures (SAS Institute 2001). Means were separated by Fisher's protected least significant difference (LSD) at the 0.05 level. Parents and [F.sub.2] were analyzed separately. Mean squares for genotype effects were considerably greater than mean squares for the genotype x environment interactions (data not shown), thus we report parent and [F.sub.2] data as means across environments.

Genetic Analysis

An additive-dominance (AD) genetic model, with G x E interaction was also used for data analysis (Tang et al., 1996; Wu et al., 1995; Zhu, 1993, 1994). This genetic model is based on the following two genetic assumptions: (i) normal diploid segregation and (ii) dominance effects (interaction effects between alleles at each locus). The genetic model for parent i at environment h is expressed as follows,

[y.sub.hiik(P)) = [mu] + [E.sub.h] + 2[A.sub.i] + [D.sub.ii] + 2A[E.sub.hi] + D[E.sub.hii] + [B.sub.k(h)] + [e.sub.hiik]

The genetic model for a F2 between parents i and j at environment h is expressed as follows,

[MATHEMATICAL EXPRESSION NOT REPRODUCIBLE IN ASCII.]

where, [mu] is population mean, [E.sub.h] is environmental effect, [A.sub.i] or [A.sub.j] is the additive effect, [D.sub.ii], [D.sub.jj], or [D.sub.ij] is the dominance effect, A[E.sub.hi] or A[E.sub.hj] is the additive x environment interaction effect, D[E.sub.hii], D[E.sub.hjj], or D[E.sub.hij] is the dominance x environment interaction effect, [B.sub.k(h)] is the block effect, and [e.sub.hijk] is the random error.

In this study, some coefficients for genetic effects were fractions rather than 0 and 1, thus, ANOVA (analysis of variance) and GLM (general linear model) methods were not appropriate for the genetic analyses. The purposes of our study were to calculate the genetic variances and genetic effects for each genetic component. The phenotypic variance was partitioned into components for additive ([V.sub.A]), dominance ([V.sub.D]), additive x environment ([V.sub.AE]), dominance x environment ([V.sub.DE]), and residual ([V.sub.e]) and expressed as proportions of the total phenotypic variance (Tang et al., 1996) Thus, we considered [mu] and [E.sub.h] as fixed and the remaining effects as random. A mixed linear model approach, minimum norm quadratic unbiased estimation with an initial value of 1.0 called MINQUE1, was used to estimate the variance components (Zhu, 1998). Genetic effects were predicted by the adjusted unbiased prediction (AUP) approach (Zhu, 1993). Standard errors of variance components and genetic effects were estimated by jackknife resampling over one replication within each environment. An approximate one-tailed t test (df = 15) was used to detect the significance of variance components and a two-tailed t test was used to detect the significance of genetic effects (Miller, 1974).

By this method, the predicted genetic effects were deviations from the respective population grand mean [mu], not from TM-1. A t test was utilized to detect the significance of genetic effects from zero. For the CS-B parents these are measures of the additive or homozygous dominance effects of the entire genome of the CS-B parent (25 chromosomes from TM-1 and one chromosome from 3-79). In addition, a significant difference for additive effects or homozygous dominance effects, for a quantitative trait, between a specific CS-B line and TM-1 was considered as a significant additive or homozygous dominance chromosome effect because of the specific substituted chromosome or chromosome arm from 3-79.

Among the [F.sub.2] hybrids, the deviation of the heterozygous dominance effect for a CS-B line and a cultivar from the heterozygous dominance for TM-1 line and the same cultivar can be considered as due to the allelic interactions between the substituted chromosome or the chromosome arm from 3-79 in the CS-B line and the homologous chromosome or arm in the respective cultivar.

Mathematically, if we define [D.sub.i1] as the heterozygous dominance effects between the female parent i and the male parent TM-1, and [D.sub.ij] as the heterozygous dominant effect between the female parent i and a CS-B line j, then the value of [D.sub.ij] minus [D.sub.i1] can be considered as the allelic interaction effect (heterozygous dominance effect) due to the specific substituted chromosome or arm from 3-79 and the homologous chromosome or arm in female cultivar parent i. This is essentially a probe of the specific combination between the specific chromosome in the cultivar and the homologous chromosome from 3-79 in the CS-B line.

The significance of the difference between the genetics effects for a CS-B line and TM-1 was detected by the method of Paterson (1939) using the standard error of the difference between two means.

RESULTS AND DISCUSSION

Mean Comparisons among Parents

On the average, the five cultivar parents had higher lint percentage, seed cotton yield, and lint yield than any CS-B line, TM-1, or 3-79 (Table 1). This was an expected result because the TM-1 and 3-79 parents are from obsolete cultivars. ST 474 had the highest lint percentage (43.95%) among parents. CS-B16, CS-B18, CS-B05sh, CS-B22sh, CS-B22Lo, and 3-79 had higher lint percentage than TM-1, while CS-B17, CS-B25, CS-B14sh, and CS-B15sh had lower lint percentage than TM-1 (Table 1). FM 966 had the heaviest bolls (6.04 g) among the parents. TM-1 bolls (5.72 g) were heavier than four cultivars and 3-79 but were lighter than FM 966. No CS-B lines produced heavier bolls than TM-1; however, CS-B07, CS-B17, CS-B18, CS-B25, CS-B05sh, CS-B14sh, CS-B22sh, CS-B22Lo, and 3-79 bolls were lighter than TM-1; whereas, CS-B02, CS-B04, CS-B06, CS-B16, and CS-B15sh were not different in boll weight than TM-1. No CS-B lines yielded more seed cotton than TM-1 while CS-B16, CS-B18, CS-B14sh, and 3-79 yielded less seed cotton than TM-1. Among CS-B lines only CS-B22Lo had higher lint yield than TM-1; whereas, CS-B16, CS-B17, CS-B18, and CS-B14sh had lower lint yields than TM-1, (Table 1).

Mean Comparisons among [F.sub.2] Hybrids

The means of hybrids across cultivars for each CS-B parent had higher lint percentage, heavier bolls, and greater seedcotton and lint yields than most of the CS-B parents, indicating that dominance effects are important for controlling each of these traits (Tables 1-5). Ranges among CS-B hybrids over four environments were 35.98 to 41.16% for lint percentage, 4.75 to 6.39 g for bolls, 2369 to 4055 kg [ha.sub.-1] for seed cotton, and 895 to 1570 kg for lint [ha.sub.-1] (Tables 2-5).

The mean of hybrids with FM 966 had higher lint percentage, heavier bolls, and greater cotton yield than the mean of hybrids with the other four cultivars (Tables 2-5). The mean of hybrids with DP 90 had lower lint percentage (37.45%) than the mean of hybrids with each of the other four cultivars (greater than 38%). The mean lint percentage of nine CS-B hybrids averaged across cultivars was greater than TM-1 hybrids across cultivars. The mean lint percentage of hybrids with CS-B22sh and CS-B22Lo was greater than the mean of all other CS-B hybrids. There were 31 individual CS-B hybrids with lint percentage significantly greater than 38%, 14 significantly greater than 39% and three significantly greater than 40% (Table 2).

The mean boll weight of hybrids with each cultivar ranged from 5.23 to 5.89 g [boll.sup.-1] (Table 3). The hybrids with FM 966 had the heaviest bolls. When averaged across cultivars, there were six CS-B hybrids with smaller bolls and two with larger bolls than hybrids with TM-1. All hybrids with 3-79 had boll weight less than or equal to 4.10 g; whereas, only hybrid ST 474 x CS-B14sh had boll weight less than 5 g. There were 61 hybrids with boll weight significantly greater than 5 g, and 4 hybrids with FM 966 with boll weight significantly greater than 6 g (Table 3).

Mean seed cotton yields of hybrids with FM 966 were significantly greater than hybrids with the other cultivars. This cultivar was also the highest in seed cotton yield. Among individual hybrid yields, the hybrid FM 966 x CS-B02, produced more seed cotton than any hybrid except FM 966 x CS-B22Lo. No CS-B hybrid mean yield across cultivars was significantly greater than the TM-1 hybrid mean across cultivars and three were less than TM-1 hybrids. The hybrids with 3-79 yielded the least seed cotton among all hybrids (Table 4).

The mean lint yields of hybrids with each cultivar ranged from 1112 to 1315 kg [ha.sup.-1]. The average yield of hybrids with DP 90 and ST 474 was significantly lower than hybrids with the other three cultivars. The mean lint yield of hybrids with FM 966 was the highest. Considering the means of hybrids across the five cultivars, hybrids with CS-B22Lo had lint yields significantly greater than TM-1 hybrids; whereas, hybrids with CSB16, CS-B18, and CS-B14sh had lint yields significantly less than hybrids with TM-1. All 3-79 hybrids produced less than 600 kg [ha.sup.-1] of lint. There were 57 of the 75 hybrids with lint yield significantly greater than 1000 and nine hybrids yielded significantly greater than 1200 kg [ha.sup.-1] (Table 5). Thus, hybrids of elite cultivars with specific CS-B lines offer a better opportunity to use genes from 3-79, G. barbadense to improve lint yields than crosses of elite cultivars with the entire genome of 3-79.

Variance Components

We expect that chromosome substitution lines (i.e., CS-B) when crossed with elite cultivars will reduce a trait's phenotypic complexity when compared to interspecific lineages with the same elite cultivars. This reduction in phenotypic complexity will improve variance component estimates.

The variance components were expressed as proportions of the phenotypic variance and are summarized in Table 6. Additive effects contributed the most to yield and yield components (lint percentage 57.6%, boll weight 58.1%, seed cotton yield 37.7%, and lint yield 41.9%). Dominance effects also made significant contributions to the phenotypic variance (lint percentage 30.2%, boll weight 24.8%, seed cotton 15.9%, and lint yield 17.6%). The G x E interaction variance [([V.sub.AE] + [V.sub.DE])/[V.sub.P]] contributed less than 10% to the phenotypic variance for lint percentage and boll weight and accounted for 24 and 22% to the phenotypic variance for seed cotton yield and lint cotton yield, respectively. Thus, the larger proportion of the phenotypic variance attributed to additive effects, indicates that these effects can be utilized in breeding programs designed to produce cultivars. The significant dominance components also indicate that specific CS-B lines may be useful for developing hybrid cottons. In this present study, we bridge across genetic interest to plant breeding interest as we probe the genetic effects of a specific 3-79 chromosome or chromosome arm and the homologous chromosome or arm in the elite cultivars.

Additive Genetic Effects

Additive effects can be considered as measuring the general combining ability (GCA) of parents and we report these as deviations from the population grand mean. We can also calculate the additive effects of only the specific chromosome or chromosome arm from 3-79 that is substituted into the CS-B parent and this is calculated as the difference in the additive effect value for a CS-B parent and the additive effect value for the TM-1 parent.

Lint Percentage

Additive effects for lint percentage were significantly different from TM-1, but negative, for CS-B02, CS-B04, CS-B07, CS-B25, CS-B05sh, CS-B14sh, and CS-B15sh. However, CS-B16, CS-B18, CS-B22sh, and CS-B22Lo had positive additive effects for lint percentage that were significantly greater than TM-1. Thus, alleles on chromosomes 16, 18, 22sh, and 22Lo from 3-79 contribute greater additive effects for lint percentage than alleles on the homologous chromosomes in TM-1 (Table 7).

Boll Weight

FM 966, the cultivar with the heaviest bolls, had the largest additive effects (0.64 g [boll.sup.-1]) for boll weight (Table 7). SG 747 also had significant positive additive effects; whereas, DP 90 and ST 474 had significant negative additive effects on boll weight. Although eight CS-B lines had significantly lower boll weight than TM-1; additive effects for chromosomes 4, 6, and 15sh from 3-79 were significantly greater than additive effects for homologous chromosomes in TM-1; whereas, additive effects for boll weight for chromosomes 18, 25, 05sh, 14sh, 22sh, and 22Lo from 3-79 were less than the homologous chromosomes in TM-1. Additive effects for 3-79 were negative and less than TM-1.

Seed Cotton Yield

Additive effects for seed cotton yield were positive and significantly different than zero for all cultivars and CS-B02; however, no chromosome from 3-79 had additive effects significantly greater than additive effects of the homologous chromosome in TM-1. Additive effects on seed cotton yield for chromosomes 16, 17, 18, and 14sh from 3-79 were negative and less than additive effects for homologous chromosomes from TM-1. A large negative additive effect on yield was found for the whole genome 3-79 (Table 7).

Lint Yield

Additive effects on lint yield for all cultivars were positive and greater than TM-1. Chromosomes 22sh and 22Lo from 3-79 had positive additive effects that were significantly greater than the homologous chromosomes from TM-1; however, additive effects for chromosomes 16, 17, 18, and 14sh from 3-79 were negative and less than the homologous chromosomes from TM-1 (Table 7).

Dominance Genetic Effects

Dominance effects can be considered as measuring the specific combining ability (SCA) of parents in specific hybrid combinations. We estimated two types of dominance effects, homozygous ([D.sub.ii]) and heterozygous ([D.sub.ij]). A negative homozygous dominance effect for a parent will result in inbreeding depression in generations following the use of this parent in a cross. High heterozygous dominance effects between two parents should result in high heterosis in the [F.sub.1] or [F.sub.2] hybrid. The homozygous dominance effects are summarized in Table 8 and the heterozygous dominance effects in Tables 9-12.

Homozygous Dominance Effects

The largest homozygous dominance effect among cultivars for lint percent was ST 474 with positive 3.3% (Table 8). This cultivar also had the highest lint percentage among cultivars (Table 1). All CS-B lines (Table 8) had significant negative homozygous dominance effects for lint percentage except CS-B16 (+0.60) and CS-B22Lo (not different from zero). Homozygous dominance effects for boll weight were negative for all cultivars and CS-B lines (Table 8). Homozygous dominance effects for seed cotton yield were similar to those for lint cotton yield. Homozygous dominance effects for lint yield for all CS-B parents were significant and negative, except for CS-B6, CS-B16, and CS-B25, which were not different from zero. FM 966 had a significant negative homozygous dominance effects whereas, the effect for the other four cultivars were not significantly different from zero (Table 8).

Heterozygous Dominance Effects

Lint Percentage

In hybrids with DP 90, heterozygous dominance effects for lint percentage for substituted chromosomes 17 and 22sh were significantly greater than for the homologous chromosome in TM-1 (Table 9). In hybrids with SG 747, dominance effects for substituted chromosomes 4, 18, and 22sh were significantly greater than homologous chromosomes in TM-1. In hybrids with PSC 355, dominance effects for substituted chromosome 4, 6, 7, 17, 18, 05sh, 14sh, 15sh, and 22sh were significantly greater than homologous chromosomes in TM-1. In hybrids with ST 474, dominance effects for substituted chromosome 17 was significantly greater than chromosome 17 in TM-1; whereas, dominance effects for substituted chromosomes 6, 5sh, and 15sh from 3-79 were significantly less than the homologous chromosomes in TM-1. ST 474 had the highest lint percentage among cultivars and CS-B17 had the lowest. In hybrids with FM 966, substituted chromosomes 6, 16, 22sh, and 22Lo had heterozygous dominance effects for lint percentage that were significantly greater than the homologous chromosomes in TM-1. These data indicate that specific substituted chromosomes from 3-79 interact to provide significant, positive, heterozygous dominance effects greater than the homologous TM-1 chromosome and much better than the negative dominance effects from the whole genome 3-79 crosses. Thus, these effect are essentially the lieterozygous dominance effects of alleles on individual chromosomes from 3-79 with the homologous chromosome in cultivars or they could be viewed as chromosome specific heterozygous dominance effects. Thus, these data show that we can use specific chromosome substitution lines in crosses with cultivars and overcome the negative dominance effects on lint percentage obtained when the whole genome of 3-79 is used in crosses. This should also avoid the hybrid incompatibility of the interspecific genome cross between G. hirsutum and G. barbadense.

Boll Weight

When the entire genome of 3-79 was crossed with each cultivar, heterozygous dominance effects were very negative and less than TM-1 for all hybrids, except with SG747 (Table 10). In hybrids with DP 90, substituted chromosomes 6, 7, 17, 14sh, and 22sh from 3-79 had heterozygous dominance effects significantly greater than the homologous chromosomes from TM-1 (Table 10). No chromosomes showed heterozygous dominance effects different from TM-1 in hybrids with SG 747. In hybrids with PSC 355, substituted chromosomes 7, 25, and 22sh had heterozygous dominance effects that were negative and significantly less than homologous chromosomes from TM-1. In hybrids with ST 474, substituted chromosomes 4 and 15sh had positive heterozygous dominance effects significantly greater than homologous chromosomes from TM-1. In hybrids with FM 966, chromosomes 4, 16, and 22sh from 3-79 had positive heterozygous dominance effects significantly greater than homologous chromosomes from TM-1. Breeders should be able to increase boll weight of hybrids by using specific CS-B lines in crosses with DP 90, ST 474, and FM 966.

Seed Cotton Yield

All hybrids with 3-79 exhibited large, significant, negative heterozygous dominance effects for seed cotton yield (Table 11). Crosses of ST 474 x CS-B17 and FM 966 x CS-B02 had positive heterozygous dominance for seed cotton yield which were greater than the homologous chromosomes from TM-1, even though the cross of the entire genome of 3-79 with cultivars has a large negative effect on seed cotton yield (Table 11).

Lint Cotton Yield

All hybrids with 3-79 exhibited large and negative heterozygous dominance effects for lint yield (Table 12). Hybrids of DP 90 x CS-B15sh, ST 474 x CS-B17, and FM 966 x CS-B02 exhibited large and positive dominance effects for lint yield which were greater than corresponding hybrids with TM-1. The final product is lint yield and in these three hybrids, chromosomes, 2, 15sh and 17, from 3-79 contributed alleles that produced significantly greater heterozygous dominance effects for lint yield than alleles from the homologous chromosomes in TM-1.

CONCLUSIONS

We can consider additive effects as representing GCA effects and heterozygous dominance effects as SCA effects. The five cultivars, as expected, had larger additive effects for lint percentage and lint cotton yield than each of the CS-B lines, indicating that these cultivars were good combiners for improving lint percentage and lint cotton yield compared to these CS-B lines. All whole genome [F.sub.2] hybrid lines involving 3-79 displayed "hybrid breakdown" typical for [F.sub.2] populations between these two species. This hybrid breakdown is accompanied by partial sterility, poor seed viability, and other disruptions. The incompatibility between these two species may not necessarily affect specific yield related genes, but could be the result of disruption of regulatory genes determining fertility, seed viability, etc. The value of the CS-B lines is that they allow chromosome specific genetic effects to be dissected from the 3-79 genome and utilized for improvements in yield components and lint yield.

Chromosomes 22sh and 22Lo from 3-79 are good examples of small but significant increases of 54 and 60 kg lint [ha.sup.-1] because of additive effects. Three specific crosses involving chromosomes 2, 17, and 15sh from 3-79 are examples of significant heterozygous dominance effects for lint yield of 396, 273, and 274 kg lint [ha.sup.-1], respectively. These chromosome specific positive effects indicate that the CS-B lines can provide access to useful QTL alleles in G. barbadense 3-79 that are not easily detectable or useful at the whole genome cross. Chromosomes 16, 18, 22sh, and 22Lo showed significant, positive additive effects for lint percentage, 22sh and 22Lo showed significant and positive additive effects for lint yield, and chromosomes 4, 6, and 15sh had significant and positive additive effects for boll weight that were each greater than the respective additive effects of TM-1. These are specific examples of alleles from specific chromosomes or arms of 3-79 that are uncovered in the CS-B lines and that were better than alleles in TM-1. Considering that 3-79 has a smaller boll and produces less lint yield than TM-1 this illustrates the utility of using CS-B lines to uncover useful genetic alleles.

The five cultivars used in these crosses represent diverse commercial breeding programs; therefore, the results from these studies should offer guidance to commercial use of these CS-B germplasm in breeding programs.

Jenkins et al. (2004) reported that chromosome 25 of 3-79 had greater additive effects on fiber length and fiber strength and chromosome arms 22sh and 22Lo had greater additive effects for lint percentage and lint yield than the homologous chromosomes in TM-1 and that most traits showed predominantly additive effects, using data from two environments, for the same top crosses used in this present study. In the current study, we expanded the environments to four and thus obtained a better estimate of these effects.

Plant breeding improvements from whole genome crosses between G. barbadense and G. hirsutum have not been very successful because of incompatibility and other genetic problems. However, use of these CS-B lines allows alleles from single G. barbadense chromosomes to be introgressed into G. hirsutum while avoiding the major problems common to interspecific crosses between these species. These CS-B lines provide a better way to use the favorable alleles from G. barbadense. CS-B lines allow the effects of an individual chromosomes or chromosome arms to be studied. Recombinant substituted (RS) lines, which are recombinant inbred lines for specific chromosomes, can be used to more precisely identify and map gene(s) controlling agronomic traits, fiber traits, and to map molecular markers (Campbell et al., 2003, 2004; Kaeppler, 1997; Shah et al., 1999). RS lines are superior to recombinant inbred lines for identifying genes or QTL of quantitative traits because RS lines increase statistical power and have a more uniform genetic background with only one divergent chromosome or segment. We are developing RS populations in selected CS-B lines to be used for high-resolution dissection and mapping of the QTL governing cotton yield and fiber quality.

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Abbreviations: CS-B, Chromosome substitution line from G. barbadense; GCA, general combining ability; SCA, specific combining ability; RS, Recombinant substituted lines.

Johnie N. Jenkins,* Jixiang Wu, Jack C. McCarty, Sukumar Saha, Osman Gutierrez, Russell Hayes, and David M. Stelly

Johnie N. Jenkins, Jack C. McCarty, Sukumar Saha, Osman Gutierrez, and Russell Hayes, Crop Science Research Laboratory, USDA-ARS, Mississippi State, MS 39762; Jixiang Wu, Department of Plant and Soil Sciences, Mississippi State University, Mississippi State, MS 39762; David M. Stelly, Department of Soil and Crop Sciences, Texas A&M University, College Station, TX 77843-2474. Mention of trade names or commercial products in this manuscript does not imply recommendation or endorsement by the U. S. Department of Agriculture. Joint Contribution of USDA-ARS and Mississippi State University. Journal paper No. J-10780 of Mississippi Agricultural and Forestry Experiment Station. Received 22 Aug. 2005. * Corresponding author (jnjenkins@ars.usda.gov).

Table 1. Mean yield and yield component values for parents. Lint Boll Seed cotton Lint Parent percentage weight yield yield % g kg [ha.sup.-1] DP 90 39.68 4.97 3530 1396 SG 747 42.16 5.39 3512 1471 PSC 355 41.16 5.04 3726 1531 ST 474 43.95 5.01 3549 1541 FM 966 42.10 6.04 3855 1622 TM-1 34.45 5.72 2531 869 CS-B02 34.68 5.70 2687 926 CS-B04 34.44 5.75 2646 906 CS-B06 34.46 5.72 2833 965 CS-B07 34.17 5.31 2753 931 CS-B16 37.69 5.69 1472 559 CS-B17 30.98 5.32 2340 726 CS-B18 35.32 5.32 2049 723 CS-B25 32.30 5.12 2428 783 CS-B05sh 35.10 5.01 2715 937 CS-B14sh 32.74 4.40 2110 686 CS-B15sh 32.97 5.65 2410 794 CS-B22sh 37.35 5.23 2433 896 CS-B22Lo 38.27 4.53 2719 1033 3-79 35.56 3.72 1601 567 LSD 0.05 0.51 0.18 372 140 Table 2. Lint percentage of [F.sub.2] hybrid from crosses of five cultivars with CS-B lines, TM-1, and 3-79. Cultivar Male parent ([dagger]) DP90 SG747 PSC355 ST474 FM966 Mean TM-1 37.29 38.10 37.61 38.80 38.55 38.07 CS-B02 37.46 38.23 37.92 38.94 38.83 38.28 CS-C04 37.31 39.08 39.13 38.85 39.03 38.68 CS-B06 37.94 37.91 38.54 37.62 39.37 38.28 CS-B07 37.60 38.54 39.32 39.22 38.38 38.61 CS-B16 38.21 39.76 38.60 39.84 40.49 39.38 CS-B17 37.07 37.66 37.52 38.99 37.97 37.84 CS-B18 38.16 39.69 39.42 39.73 39.17 39.23 CS-B25 36.90 37.65 36.88 37.69 38.15 37.45 CS-B05sh 37.80 38.80 39.76 38.16 39.56 38.82 CS-B14sh 35.98 37.31 37.44 38.34 37.95 37.40 CS-B15sh 37.05 37.87 37.61 37.36 37.87 37.55 CS-B22sh 39.94 40.51 40.22 40.44 41.16 40.45 CS-B22Lo 39.54 39.74 39.26 40.64 40.88 40.01 3-79 33.55 36.52 33.10 33.83 34.45 34.29 Mean 37.45 38.49 38.15 38.56 38.79 ([dagger]) LSD 0.05 for comparing males averaged across cultivars is 0.30; LSD 0.05 for comparing individual hybrids is 0.57; and LSD 0.05 for comparing female means averaged across CS-B lines, TM-1, and 3-79 individual hybrids is 0.15. Table 3. Boll weight (grams seed cotton [boll.sup.-1] of [F.sub.2] hybrid from crosses of five cultivars with CS-B lines, TM-1, and 3-79. Cultivars Male parent ([dagger]) DP90 SG747 PSC355 ST474 FM966 Mean TM-1 5.42 5.70 5.67 5.42 6.09 5.66 CS-B02 5.38 5.62 5.71 5.61 6.25 5.71 CS-B04 5.36 5.66 5.70 5.76 6.32 5.76 CS-B06 5.68 5.72 5.60 5.61 6.19 5.76 CS-B07 5.59 5.61 5.35 5.43 5.94 5.58 CS-B16 5.32 5.62 5.67 5.29 6.39 5.66 CS-B17 5.51 5.68 5.57 5.42 6.11 5.66 CS-B18 5.37 5.56 5.44 5.27 5.94 5.52 CS-B25 5.27 5.42 5.04 5.33 5.89 5.39 CS-B05sh 5.13 5.40 5.21 5.24 5.90 5.38 CS-B14sh 5.05 5.17 5.09 4.75 5.69 5.15 CS-B15sh 5.57 5.72 5.63 5.69 6.27 5.78 CS-B22sh 5.44 5.34 5.19 5.35 6.11 5.49 CS-B22Lo 4.91 5.17 5.29 5.04 5.60 5.20 3-79 3.50 4.10 3.44 3.38 3.62 3.61 Mean 5.23 5.43 5.31 5.24 5.89 ([dagger]) LSD 0.05 for comparing males averaged across cultivars is 0.09; LSD 0.05 for comparing individual hybrids is 0.19; and LSD 0.05 for comparing female means averaged across CS-B lines, TM-1, and 3-79 individual hybrids is 0.05. Table 4. Seed cotton yields (kg [ha.sup.-1]) of [F.sub.] hybrid from crosses of five cultivars with CS-B lines, TM-1, and 3-79 Cultivar Male parent ([dagger]) DP90 SG747 PSC355 ST474 FM966 Mean TM-1 3154 3495 3556 3093 3558 3371 CS-B02 3300 3419 3334 3336 4055 3489 CS-B04 2982 3255 3437 3300 3481 3291 CS-B06 3228 3487 3525 3049 3609 3380 CS-B07 3236 3472 3590 3110 3639 3409 CS-B16 2369 3053 2438 2419 3039 2663 CS-B17 3090 3150 3272 3290 3343 3229 CS-B18 2953 3114 3097 2797 3257 3043 CS-B25 3055 3338 3334 3250 3388 3273 CS-B05sh 3285 3345 3538 3259 3730 3432 CS-B14sh 2687 2989 3461 3048 3605 3158 CS-B15sh 3445 3391 3432 3313 3507 3417 CS-B22sh 3309 3041 3419 3306 3584 3332 CS-B22Lo 3293 3440 3395 3127 3808 3413 3-79 1231 1468 1666 1366 1156 1378 Mean 2974 3164 3233 3004 3384 ([dagger]) LSD 0.05 for comparing males averaged across cultivars is 183; LSD 0.05 for comparing individual hybrids is 410; and LSD 0.05 for comparing female means averaged across CS-B lines, TM-1, and 3-79 individual hybrids is 105. Table 5. Lint yield (kg [ha.sup.-1]) of [F.sub.2] hybrid from crosses of five cultivars with CS-B lines, TM-1, and 3-79. Cultivar Male parent ([dagger]) DP90 SG747 PSC355 ST474 FM966 Mean TM-1 1167 1318 1333 1187 1367 1275 CS-B02 1233 1293 1265 1289 1570 1330 CS-B04 1104 1264 1337 1272 1354 1266 CS-B06 1217 1304 1351 1124 1415 1282 CS-B07 1211 1331 1406 1199 1391 1308 CS-B16 895 1206 935 952 1229 1043 CS-B17 1134 1176 1225 1274 1266 1215 CS-B18 1115 1227 1219 1096 1273 1187 CS-B25 1123 1250 1219 1212 1286 1218 CS-B05sh 1237 1283 1405 1228 1472 1325 CS-B14sh 954 1109 1287 1156 1364 1174 CS-B15sh 1276 1271 1284 1224 1325 1276 CS-B22sh 1313 1222 1372 1321 1473 1340 CS-B22Lo 1294 1348 1323 1256 1545 1353 3-79 413 535 549 462 397 471 Mean 1112 1209 1234 1150 1315 ([dagger]) LSD 0.05 for comparing males averaged across cultivars is 70; LSD 0.05 for comparing individual hybrids is 157; and LSD 0.05 for comparing female means averaged across CS-B lines, TM-1, and 3-79 individual hybrids is 41. Table 6. Variance Components and standard errors expressed as proportions of the phenotypic variances for yield traits. Lint % [V.sub.A]/[V.sub.P] ([dagger]) 0.576 [+ or -] 0.009 [V.sub.D]/[V.sub.P] 0.302 [+ or -] 0.011 [V.sub.AE]/[V.sub.P] 0.007 [+ or -] 0.002 [V.sub.DE]/[V.sub.P] 0.061 [+ or -] 0.009 [V.sub.e]/[V.sub.P] 0.054 [+ or -] 0.005 Boll weight [V.sub.A]/[V.sub.P] ([dagger]) 0.581 [+ or -] 0.013 [V.sub.D]/[V.sub.P] 0.248 [+ or -] 0.016 [V.sub.AE]/[V.sub.P] 0.000 [+ or -] 0.000 [V.sub.DE]/[V.sub.P] 0.079 [+ or -] 0.016 [V.sub.e]/[V.sub.P] 0.092 [+ or -] 0.005 Seed cotton yield [V.sub.A]/[V.sub.P] ([dagger]) 0.377 [+ or -] 0.017 [V.sub.D]/[V.sub.P] 0.159 [+ or -] 0.021 [V.sub.AE]/[V.sub.P] 0.105 [+ or -] 0.010 [V.sub.DE]/[V.sub.P] 0.135 [+ or -] 0.024 [V.sub.e]/[V.sub.P] 0.224 [+ or -] 0.022 Lint yield [V.sub.A]/[V.sub.P] ([dagger]) 0.419 [+ or -] 0.017 [V.sub.D]/[V.sub.P] 0.176 [+ or -] 0.021 [V.sub.AE]/[V.sub.P] 0.097 [+ or -] 0.009 [V.sub.DE]/[V.sub.P] 0.124 [+ or -] 0.022 [V.sub.e]/[V.sub.P] 0.184 [+ or -] 0.021 ([dagger]) [V.sub.A] = additive variance; [V.sub.D] = dominance variance; [V.sub.AE] = additive x environment variance; [V.sub.DE] = dominance x environment variance; [V.sub.e] = error variance; [V.sub.P] = phenotypic variance. Table 7. Additive effects and standard errors for yield and yield components expressed as deviations from population grand mean. Male parents Lint percentage ([dagger]) ([doouble dagger]) % g DP 90 1.64 [+ or -] 0.12 ** * SG 747 2.56 [+ or -] 0.09 ** * PSC 355 2.31 [+ or -] 0.10 ** * ST 474 2.18 [+ or -] 0.10 ** * PM 966 3.03 [+ or -] 0.12 ** * TM-1 -1.00 [+ or -] 0.08 ** CS-B02 -0.78 [+ or -] 0.07 ** * CS-B04 -0.35 [+ or -] 0.08 ** * CS-B06 -0.78 [+ or -] 0.09 ** CS-B07 -0.42 [+ or -] 0.11 ** * CS-B16 0.20 [+ or -] 0.11 * CS-B17 -1.07 [+ or -] 0.08 ** CS-B18 0.17 [+ or -] 0.10 * CS-B25 -1.53 [+ or -] 0.12 ** * CS-B05sh -0.26 [+ or -] 0.08 ** * CS-B14sh -1.61 [+ or -] 0.09 ** * CS-B15sh -1.47 [+ or -] 0.11 ** * CS-B22sh 1.33 [+ or -] 0.11 ** * CS-B22Lo 0.82 [+ or -] 0.11 ** * 3-79 -4.97 [+ or -] 0.17 ** * Male parents Boll weight ([dagger]) ([doouble dagger]) g DP 90 -0.09 [+ or -] 0.03 ** * SG 747 0.11 [+ or -] 0.03 ** PSC 355 0.01 [+ or -] 0.02 * ST 474 -0.09 [+ or -] 0.03 * * PM 966 0.64 [+ or -] 0.04 ** * TM-1 0.19 [+ or -] 0.03 ** CS-B02 0.25 [+ or -] 0.03 ** CS-B04 0.29 [+ or -] 0.03 ** * CS-B06 0.29 [+ or -] 0.03 ** * CS-B07 0.13 [+ or -] 0.03 ** CS-B16 0.19 [+ or -] 0.04 ** CS-B17 0.20 [+ or -] 0.03 ** CS-B18 0.06 [+ or -] 0.03 * CS-B25 -0.07 [+ or -] 0.02 ** * CS-B05sh -0.07 [+ or -] 0.03 * * CS-B14sh -0.28 [+ or -] 0.03 ** * CS-B15sh 0.31 [+ or -] 0.03 ** * CS-B22sh 0.03 [+ or -] 0.03 * CS-B22Lo -0.23 [+ or -] 0.02 ** 3-79 -1.86 [+ or -] 0.03 ** * Seed cotton Male parents yield ([dagger]) ([doouble dagger]) kg [ha.sup.-1] DP 90 236 [+ or -] 59 ** * SG 747 529 [+ or -] 67 ** * PSC 355 575 [+ or -] 78 ** * ST 474 277 [+ or -] 65 ** * PM 966 769 [+ or -] 68 ** * TM-1 56 [+ or -] 38 CS-B02 170 [+ or -] 62 * CS-B04 -29 [+ or -] 58 CS-B06 51 [+ or -] 75 CS-B07 85 [+ or -] 66 CS-B16 -612 [+ or -] 64 ** * CS-B17 -79 [+ or -] 54 * CS-B18 253 [+ or -] 55 ** * CS-B25 -38 [+ or -] 82 CS-B05sh 108 [+ or -] 80 CS-B14sh -140 [+ or -] 56 * * CS-B15sh 110 [+ or -] 58 CS-B22sh 21 [+ or -] 59 CS-B22Lo 89 [+ or -] 69 3-79 -1922 [+ or -] 64 ** Lint cotton Male parents yield ([dagger]) ([doouble dagger]) kg [ha.sup.-1] DP 90 135 [+ or -] 22 ** * SG 747 262 [+ or -] 26 ** * PSC 355 283 [+ or -] 31 ** * ST 474 153 [+ or -] 26 ** * PM 966 382 [+ or -] 28 ** * TM-1 -12 [+ or -] 14 CS-B02 43 [+ or -] 25 CS-B04 -21 [+ or -] 23 CS-B06 -8 [+ or -] 28 CS-B07 19 [+ or -] 25 CS-B16 -233 [+ or -] 26 ** * CS-B17 -65 [+ or -] 20 ** * CS-B18 -94 [+ or -] 22 ** * CS-B25 -65 [+ or -] 31 CS-B05sh 36 [+ or -] 31 CS-B14sh -106 [+ or -] 22 ** * CS-B15sh -6 [+ or -] 23 CS-B22sh 54 [+ or -] 24 * * CS-B22Lo 60 [+ or -] 27 * * 3-79 -816 [+ or -] 24 ** * * Significantly different at 0.05 probability level. ** Sinificantly different at 0.01 probability level. ([dagger]) Difference from zero. ([dagger]) Difference between CS-B line and TM-1. Table 8. Homozygous dominance effects and standard errors for yield and yield components expressed as deviations from population grand mean. Male parents Lint percentage ([dagger]) ([double dagger]) % DP 90 -0.26 [+ or -] 0.31 SG 747 0.53 [+ or -] 0.39 * PSC 355 -0.05 [+ or -] 0.37 ST 474 3.31 [+ or -] 0.47 ** * FM 966 -0.54 [+ or -] 0.43 TM-1 -0.41 [+ or -] 0.23 CS-B02 -0.58 [+ or -] 0.23 * CS-B04 -1.74 [+ or -] 0.28 ** * CS-B06 -0.85 [+ or -] 0.27 ** CS-B07 -1.93 [+ or -] 0.37 ** * CS-B16 0.60 [+ or -] 0.26 * * CS-B17 -4.05 [+ or -] 0.29 ** * CS-B18 -1.91 [+ or -] 0.30 ** * CS-B25 -1.65 [+ or -] 0.40 ** * CS-B05sh -1.28 [+ or -] 0.31 ** * CS-B14sh -1.00 [+ or -] 0.28 ** CS-B15sh -1.06 [+ or -] 0.34 ** CS-B22sh -2.13 [+ or -] 0.45 ** * CS-B22Lo -0.08 [+ or -] 0.21 3-79 9.10 [+ or -] 0.26 ** * Male parents Boll weight ([dagger]) ([double dagger]) g DP 90 -0.35 [+ or -] 0.12 ** * SG 747 -0.30 [+ or -] 0.10 ** PSC 355 -0.46 [+ or -] 0.09 ** * ST 474 -0.30 [+ or -] 0.11 * FM 966 -0.67 [+ or -] 0.10 ** * TM-1 -0.10 [+ or -] 0.12 CS-B02 -0.23 [+ or -] 0.13 CS-B04 -0.27 [+ or -] 0.10 * CS-B06 -0.31 [+ or -] 0.12 * CS-B07 -0.42 [+ or -] 0.13 ** CS-B16 -0.12 [+ or -] 0.11 CS-B17 -0.56 [+ or -] 0.09 ** * CS-B18 -0.25 [+ or -] 0.14 CS-B25 -0.22 [+ or -] 0.10 * CS-B05sh -0.33 [+ or -] 0.08 ** CS-B14sh -0.56 [+ or -] 0.09 ** * CS-B15sh -0.43 [+ or -] 0.09 ** * CS-B22sh -0.31 [+ or -] 0.12 * CS-B22Lo -0.51 [+ or -] 0.10 ** * 3-79 1.94 [+ or -] 0.06 ** * Seed cotton Male parents yield ([dagger]) ([double dagger]) kg [ha.sup.-1] DP 90 151 [+ or -] 236 * SG 747 -443 [+ or -] 261 PSC 355 -318 [+ or -] 312 ST 474 89 [+ or -] 265 FM 966 -579 [+ or -] 163 ** TM-1 -507 [+ or -] 172 ** CS-B02 -571 [+ or -] 188 ** CS-B04 -219 [+ or -] 117 CS-B06 -193 [+ or -] 259 CS-B07 -336 [+ or -] 185 CS-B16 -256 [+ or -] 121 CS-B17 -431 [+ or -] 151 * CS-B18 -380 [+ or -] 162 * CS-B25 -422 [+ or -] 228 CS-B05sh -423 [+ or -] 181 * CS-B14sh -543 [+ or -] 163 ** CS-B15sh -732 [+ or -] 147 ** CS-B22sh -536 [+ or -] 211 * CS-B22Lo -380 [+ or -] 182 3-79 2461 [+ or -] 261 ** * Lint cotton Male parents yield ([dagger]) ([double dagger]) kg [ha.sup.-1] DP 90 72 [+ or -] 97 * SG 747 -106 [+ or -] 109 PSC 355 -87 [+ or -] 31 ST 474 185 [+ or -] 123 * FM 966 -195 [+ or -] 73 * TM-1 -179 [+ or -] 62 * CS-B02 -229 [+ or -] 73 ** CS-B04 -121 [+ or -] 48 * CS-B06 -87 [+ or -] 97 CS-B07 -177 [+ or -] 71 * CS-B16 -55 [+ or -] 47 CS-B17 -218 [+ or -] 52 ** CS-B18 -162 [+ or -] 63 * CS-B25 -160 [+ or -] 83 CS-B05sh -205 [+ or -] 71 * CS-B14sh -178 [+ or -] 59 ** CS-B15sh -266 [+ or -] 52 ** CS-B22sh -283 [+ or -] 87 ** CS-B22Lo -156 [+ or -] 72 * 3-79 1127 [+ or -] 100 ** * * Significantly different at 0.05 probability level. ** Significantly different at 0.01 probability level. ([dagger]) Difference from zero. ([double dagger]) Difference between CS-B line and TM-1. Table 9. Heterozygous dominance effects and standard errors for lint percentage expressed as deviations from population grand mean. Cultivar Male parents DP90 SG747 TM-1 0.15 [+ or -] 0.44 -0.39 [+ or -] 0.34 CS-B02 0.19 [+ or -] 0.30 -0.43 [+ or -] 0.43 CS-B04 -0.47 [+ or -] 0.33 1.08 [+ or -] 0.40 * CS-B06 1.33 [+ or -] 0.49 -1.02 [+ or -] 0.35 CS-B07 0.38 [+ or -] 0.33 0.11 [+ or -] 0.34 CS-B16 -0.85 [+ or -] 0.41 0.21 [+ or -] 0.47 CS-B17 1.62 [+ or -] 0.30 * 0.59 [+ or -] 0.40 CS-B18 0.35 [+ or -] 0.39 1.38 [+ or -] 0.42 * CS-B25 1.03 [+ or -] 0.39 0.36 [+ or -] 0.47 CS-B05sh 0.14 [+ or -] 0.21 0.01 [+ or -] 0.33 CS-B14sh -1.14 [+ or -] 0.47 -0.54 [+ or -] 0.36 CS-B15sh 0.92 [+ or -] 0.35 0.41 [+ or -] 0.38 CS-B22sh 1.89 [+ or -] 0.47 * 0.83 [+ or -] 0.42 * CS-B22Lo 1.05 [+ or -] 0.40 -0.80 [+ or -] 0.36 3-79 -4.4 [+ or -] 0.62 * -0.25 [+ or -] 0.73 Cultivar Male parents PSC355 ST474 TM-1 -0.66 [+ or -] 0.23 0.57 [+ or -] 0.50 CS-B02 -0.32 [+ or -] 0.44 0.54 [+ or -] 0.26 CS-B04 1.98 [+ or -] 0.47 * 0.04 [+ or -] 0.34 CS-B06 1.12 [+ or -] 0.44 * -2.20 [+ or -] 0.43 * CS-B07 2.59 [+ or -] 0.42 * 1.05 [+ or -] 0.52 CS-B16 -1.52 [+ or -] 0.45 -0.17 [+ or -] 0.42 CS-B17 1.09 [+ or -] 0.45 * 2.93 [+ or -] 0.36 * CS-B18 1.58 [+ or -] 0.33 * 0.91 [+ or -] 0.34 CS-B25 -0.51 [+ or -] 0.30 -0.11 [+ or -] 0.50 CS-B05sh 2.88 [+ or -] 0.33 * -1.93 [+ or -] 0.28 * CS-B14sh 0.52 [+ or -] 0.44 * 1.14 [+ or -] 0.35 CS-B15sh 0.62 [+ or -] 0.41 * -1.27 [+ or -] 0.37 * CS-B22sh 1.01 [+ or -] 0.43 * 0.13 [+ or -] 0.44 CS-B22Lo -1.04 [+ or -] 0.47 0.59 [+ or -] 0.38 3-79 -6.87 [+ or -] 0.56 * -6.63 [+ or -] 0.49 * Cultivar Male parents FM966 TM-1 0.12 [+ or -] 0.36 CS-B02 0.40 [+ or -] 0.35 CS-B04 0.51 [+ or -] 0.40 CS-B06 1.68 [+ or -] 0.35 * CS-B07 -0.69 [+ or -] 0.34 CS-B16 1.33 [+ or -] 0.36 * CS-B17 0.80 [+ or -] 0.42 CS-B18 -0.22 [+ or -] 0.34 CS-B25 0.97 [+ or -] 0.42 CS-B05sh 1.19 [+ or -] 0.41 CS-B14sh 0.38 [+ or -] 0.42 CS-B15sh -0.06 [+ or -] 0.39 CS-B22sh 1.78 [+ or -] 0.37 * CS-B22Lo 1.21 [+ or -] 0.32 * 3-79 -5.2 [+ or -] 0.41 * * Significantly different from TM-1 at 0.05 probability level. Table 10. Heterozygous dominance effects and standard errors for boll weight expressed as deviations from population grand mean. Cultivar Male Parents DP90 SG747 TM-1 -0.05 [+ or -] 0.10 0.12 [+ or -] 0.14 CS-B02 -0.19 [+ or -] 0.11 -0.11 [+ or -] 0.13 CS-B04 -0.32 [+ or -] 0.13 -0.09 [+ or -] 0.10 CS-B06 0.39 [+ or -] 0.15 * 0.06 [+ or -] 0.11 CS-B07 0.57 [+ or -] 0.14 * 0.21 [+ or -] 0.16 CS-B16 -0.25 [+ or -] 0.15 -0.05 [+ or -] 0.13 CS-B17 0.33 [+ or -] 0.06 * 0.26 [+ or -] 0.11 CS-B18 0.17 [+ or -] 0.17 0.17 [+ or -] 0.19 CS-B25 0.19 [+ or -] 0.14 0.09 [+ or -] 0.10 CS-B05sh -0.03 [+ or -] 0.17 0.13 [+ or -] 0.13 CS-B14sh 0.33 [+ or -] 0.11 * 0.16 [+ or -] 0.11 CS-B15sh 0.17 [+ or -] 0.19 0.07 [+ or -] 0.10 CS-B22sh 0.40 [+ or -] 0.15 * -0.22 [+ or -] 0.16 CS-B22Lo -0.09 [+ or -] 0.10 0.05 [+ or -] 0.13 3-79 -1.01 [+ or -] 0.16 * -0.15 [+ or -] 0.18 Cultivar Male Parents PSC355 ST474 TM-1 0.35 [+ or -] 0.16 -0.08 [+ or -] 0.12 CS-B02 0.38 [+ or -] 0.14 0.28 [+ or -] 0.10 CS-B04 0.28 [+ or -] 0.09 0.52 [+ or -] 0.08 * CS-B06 0.07 [+ or -] 0.12 0.22 [+ or -] 0.13 CS-B07 -0.07 [+ or -] 0.12 * 0.23 [+ or -] 0.13 CS-B16 0.36 [+ or -] 0.11 -0.35 [+ or -] 0.13 CS-B17 0.32 [+ or -] 0.11 0.12 [+ or -] 0.13 CS-B18 0.19 [+ or -] 0.15 -0.05 [+ or -] 0.16 CS-B25 -0.43 [+ or -] 0.14 * 0.30 [+ or -] 0.14 CS-B05sh 0.01 [+ or -] 0.14 0.18 [+ or -] 0.11 CS-B14sh 0.28 [+ or -] 0.15 -0.33 [+ or -] 0.13 CS-B15sh 0.16 [+ or -] 0.13 0.40 [+ or -] 0.17 * CS-B22sh -0.27 [+ or -] 0.10 * 0.20 [+ or -] 0.13 CS-B22Lo 0.59 [+ or -] 0.12 0.16 [+ or -] 0.08 3-79 -1.27 [+ or -] 0.16 * -1.28 [+ or -] 0.15 * Cultivar Male Parents FM966 TM-1 0.05 [+ or -] 0.10 CS-B02 0.35 [+ or -] 0.14 CS-B04 0.43 [+ or -] 0.11 * CS-B06 0.17 [+ or -] 0.17 CS-B07 0.02 [+ or -] 0.16 CS-B16 0.72 [+ or -] 0.11 * CS-B17 0.31 [+ or -] 0.10 CS-B18 0.09 [+ or -] 0.17 CS-B25 0.22 [+ or -] 0.16 CS-B05sh 0.30 [+ or -] 0.10 CS-B14sh 0.39 [+ or -] 0.17 CS-B15sh 0.37 [+ or -] 0.17 CS-B22sh 0.54 [+ or -] 0.14 * CS-B22Lo 0.08 [+ or -] 0.12 3-79 -2.06 [+ or -] 0.19 * * Significantly different from TM-1 at 0.05 probability level. Table 11. Heterozygous dominance effects and standard errors for seed cotton yield in kg [ha.sup.-1] expressed as deviations from population grand mean. Cultivar Male parents DP90 SG747 TM-1 72 [+ or -] 216 470 [+ or -] 257 CS-B02 170 [+ or -] 282 136 [+ or -] 272 CS-B04 -247 [+ or -] 301 16 [+ or -] 223 CS-B06 76 [+ or -] 242 312 [+ or -] 213 CS-B07 99 [+ or -] 259 288 [+ or -] 267 CS-B16 -318 [+ or -] 173 771 [+ or -] 231 CS-B17 165 [+ or -] 245 7 [+ or -] 251 CS-B18 208 [+ or -] 245 249 [+ or -] 250 CS-B25 12 [+ or -] 193 299 [+ or -] 301 CS-B05sh 186 [+ or -] 199 22 [+ or -] 222 CS-B14sh -462 [+ or -] 217 -141 [+ or -] 164 * CS-B15sh 661 [+ or -] 221 271 [+ or -] 233 CS-B22sh 458 [+ or -] 362 -351 [+ or -] 215 * CS-B22Lo 223 [+ or -] 163 237 [+ or -] 245 3-79 -1371 [+ or -] 131 * -1174 [+ or -] 193 * Cultivar Male parents PSC355 ST474 TM-1 441 [+ or -] 353 -101 [+ or -] 190 CS-B02 -193 [+ or -] 281 199 [+ or -] 301 CS-B04 236 [+ or -] 206 341 [+ or -] 196 CS-B06 238 [+ or -] 209 -335 [+ or -] 208 CS-B07 371 [+ or -] 182 -210 [+ or -] 225 CS-B16 -616 [+ or -] 326 * -269 [+ or -] 198 CS-B17 99 [+ or -] 278 524 [+ or -] 152 * CS-B18 65 [+ or -] 296 -151 [+ or -] 201 CS-B25 141 [+ or -] 333 357 [+ or -] 181 CS-B05sh 268 [+ or -] 282 87 [+ or -] 338 CS-B14sh 649 [+ or -] 327 213 [+ or -] 243 CS-B15sh 202 [+ or -] 257 346 [+ or -] 196 CS-B22sh 246 [+ or -] 274 410 [+ or -] 216 CS-B22Lo -11 [+ or -] 273 -164 [+ or -] 310 3-79 -929 [+ or -] 181 * -1147 [+ or -] 195 * Cultivar Male parents FM966 TM-1 187 [+ or -] 164 CS-B02 999 [+ or -] 288 * CS-B04 645 [+ or -] 184 CS-B06 147 [+ or -] 353 CS-B07 211 [+ or -] 255 CS-B16 337 [+ or -] 201 CS-B17 -9 [+ or -] 213 CS-B18 13 [+ or -] 235 CS-B25 -3 [+ or -] 283 CS-B05sh 392 [+ or -] 384 CS-B14sh 689 [+ or -] 261 CS-B15sh 92 [+ or -] 255 CS-B22sh 328 [+ or -] 204 CS-B22Lo 565 [+ or -] 273 3-79 -2211 [+ or -] 229 * * Significantly different from TM-1 at 0.05 probability level. Table 12. Heterozygous dominance effects and standard errors for lint cotton yield in kg [ha.sup.1] expressed as deviations from population grand mean. Cultivar Male parents DP90 SG747 TM-1 18 [+ or -] 85 161 [+ or -] 100 CS-B02 67 [+ or -] 112 28 [+ or -] 109 CS-B04 -121 [+ or -] 116 41 [+ or -] 91 CS-B06 67 [+ or -] 93 79 [+ or -] 81 CS-B07 47 [+ or -] 102 123 [+ or -] 110 CS-B16 -160 [+ or -] 69 309 [+ or -] 95 CS-B17 75 [+ or -] 93 -4 [+ or -] 102 CS-B18 67 [+ or -] 96 134 [+ or -] 103 CS-B25 24 [+ or -] 75 116 [+ or -] 119 CS-B05sh 76 [+ or -] 78 40 [+ or -] 90 CS-B14sh -232 [+ or -] 84 * -82 [+ or -] 67 * CS-B15sh 274 [+ or -] 87 * 96 [+ or -] 91 CS-B22sh 235 [+ or -] 151 -115 [+ or -] 88 * CS-B22Lo 121 [+ or -] 68 66 [+ or -] 99 3-79 -567 [+ or -] 42 * -482 [+ or -] 77 * Cultivar Male parents PSC355 ST474 TM-1 140 [+ or -] 141 -35 [+ or -] 79 CS-B02 -84 [+ or -] 112 95 [+ or -] 124 CS-B04 140 [+ or -] 83 132 [+ or -] 80 CS-B06 124 [+ or -] 84 -215 [+ or -] 79 CS-B07 227 [+ or -] 78 -71 [+ or -] 92 CS-B16 -298 [+ or -] 130 * -136 [+ or -] 84 CS-B17 44 [+ or -] 109 273 [+ or -] 62 * CS-B18 62 [+ or -] 123 -62 [+ or -] 82 CS-B25 2 [+ or -] 127 115 [+ or -] 70 CS-B05sh 205 [+ or -] 116 -33 [+ or -] 133 CS-B14sh 231 [+ or -] 127 90 [+ or -] 97 CS-B15sh 72 [+ or -] 100 75 [+ or -] 75 CS-B22sh 137 [+ or -] 115 162 [+ or -] 92 CS-B22Lo -39 [+ or -] 112 -49 [+ or -] 133 3-79 -506 [+ or -] 70 * -558 [+ or -] 71 * Cultivar Male parents FM966 TM-1 62 [+ or -] 67 CS-B02 396 [+ or -] 118 * CS-B04 28 [+ or -] 76 CS-B06 109 [+ or -] 145 CS-B07 49 [+ or -] 103 CS-B16 160 [+ or -] 85 CS-B17 -16 [+ or -] 87 CS-B18 29 [+ or -] 98 CS-B25 -4 [+ or -] 112 CS-B05sh 196 [+ or -] 159 CS-B14sh 245 [+ or -] 103 CS-B15sh 10 [+ or -] 102 CS-B22sh 200 [+ or -] 88 CS-B22Lo 273 [+ or -] 116 3-79 -964 [+ or -] 87 * * Significantly different from TM-1 at probability level.

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Author: | Jenkins, Johnie N.; Wu, Jixiang; McCarty, Jack C.; Saha, Sukumar; Gutierrez, Osman; Hayes, Russell; |
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Publication: | Crop Science |

Geographic Code: | 1USA |

Date: | May 1, 2006 |

Words: | 10027 |

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