Correlated responses of fatty acid composition, grain quality, and agronomic traits to nine cycles of recurrent selection for increased oil content in oat

Increases in the groat-oil content of oat (Avena sativa L.) increase the energy value of the grain and improve the feasibility of extracting oat oil for use as a vegetable oil. Nine cycles of recurrent selection for greater groat-oil content conducted in a genetically broad-based oat population resulted in dramatic increases in groat-oil content. Our objectives were todetermine if selection for greater groat-oil content affected fatty acid composition, grain quality traits (test weight and seed weight), or agronomic traits (straw yield, biomass, harvest index, heading date, and height). We evaluated 100 random lines from the base (C0) population and each of the nine selection cycle populations in three environments in order to estimate means, genetic variances, heritabilities, and genotypic and phenotypic correlations of grain quality and agronomic traits. We also evaluated 20 random lines from each population to estimate changes in fatty acid contents. Oleate and stearate contents increased over cycles of selection, as did the ratio of unsaturated to saturated fatty acids. Palmitate, linoleate, and linolenate contents and all grain quality and agronomic traits except harvest index decreased over cycles of selection. There was no evidence for reduced genetic variance or heritability in C9 for any trait, but the genotypic and phenotypic correlations between agronomic traits and oil content fluctuated over cycles. Selection for increased groat-oil content improved oil quality but reduced grain quality and agronomic performance of the population. This revised version was published online in August 2006 with corrections to the Cover Date.     

Why SR52 is such a great maize hybrid? II. Heritability,correlation and path-coefficient analyses

Although exceptional heterosis for yield in the world’s first commercial single cross maize (Zea mays L.) hybrid SR52 has been confirmed, the role of secondary traits hasn’t been established. Objectives of this study were to establish relationship between grain yield and secondary traits, while assessing heritability and genotypic variation in SR52’s segregating populations. Knowledge on associations between yield and secondary traits would be crucial in extracting productive inbred lines from the population. Traits were subjected to correlation and path-coefficient analyses. Path analysis model accounted for over 70% of variation (R²) in all populations. Consistently high positive correlations with grain yield were observed for ear length and number of kernel rows on ear. Ear girth (0.19), and number of kernels per row (0.48) showed high direct effects on yield in the F₂ population and the trend was consistent in the backcrosses. Positive indirect effects of most traits on grain yield were negligible with the exception of kernel per row via total number of kernels on ear, which was above 0.40 in the F₂ and BCP1. Ear length can be exploited for indirect selection for yield in SR52’s segregating generations. 100-kernel weight and number of kernel rows on ear can, to a lesser extent, be targeted in selecting for grain yield. High to medium phenotypic variability in segregating generations of SR52 for traits such as ear length, number kernel rows on ear and grain yield should give impetus to selection of productive inbred lines from SR52 hybrid because of presence of diversity in the gene pool.     

Identification of QTL underlying the resistance of soybean to pod borer, <Emphasis Type="Italic">Leguminivora glycinivorella</Emphasis> (Mats.) obraztsov,and correlations with plant,pod and seed traits

Soybean (Glycine max L. Merr.) pod borer (Leguminivora glycinivorella (Mats.) Obraztsov) (SPB) results in severe loss in soybean yield and quality in certain regions of the world, especially in Northeastern China, Japan and Russia. The aim here was to evaluate the inheritance of pod borer resistance and to identify quantitative trait loci (QTL) underlying SPB resistance for the acceleration of the control of this pest. Used were the 129 recombinant inbred lines (RILs) of the F_5:6 derived population from ‘Dong Nong 1068’ × ‘Dong Nong 8004’ and 131 SSR markers. Correlations between the percentage of damaged seeds (PDS) by pod borer and plant, pod and seed traits that were potentially related to SPB resistance were analyzed. The results showed highly significant correlations between PDS by pod borer and plant height (PH), maturity date (MA), pod color (PC), pubescence density (PB), 100-seed weight (SW) and protein content existed. Soybeans with dwarf stem, light color of pod coat, small seeds, lower density of pubescence, early maturity and low content of protein seemed to have higher resistance to SPB. The correlated traits had potential to inhibit egg deposition and thereby to decrease the damage by SPB. Three QTL directly associated with the resistance to SPB judged by PDS at harvest were identified. qRspb-1 (Satt541–Satt253) and qRspb-2 (Satt253–Satt314) were both on linkage group (LG) H and qRspb-3 (Satt288–Satt199) on LG G. The three QTL explained 10.96, 9.73 and 11.59% of the phenotypic variation for PDS, respectively. In addition, 12 QTL that underlay 10 of 13 traits potentially related with SPB resistance were found. These QTL detected jointly provide potential for marker assisted selection to improve cultivar resistance to SPB. Guiyun Zhao, Jian Wang, and Yingpeng Han have equal contribution to the paper.     

Use of index selection in recurrent selection programs in maize

Summary Phenotypic and genotypic correlations were examined for four traits in seven populations of maize (Zea mays L.) undergoing recurrent selection. Correlations among grain yield and percentage of grain moisture, root lodging, and stalk lodging were low (|r|<0.3) except for the correlation between grain yield and stalk lodging, which was high and negative. The phenotypic and genotypic correlations agreed well from cycle to cycle within populations. Variation of correlations among populations was not significantly larger than variation among cycles. Heritabilities of these traits generally were high (h²>0.5). Two indices, one that used heritabilities as index weights and one that used relative economic weights (base index) as index weights, were compared with the Smith-Hazel index (optimim index). Relative efficiencies of the two indices, in terms of predicted gains for the individual traits and the composite trait, compared with the Smith-Hazel index, were high. The use of an index in which heritabilities were used as index weights was recommended because:1) the heritabilities were the same as the optimum weights when the traits were uncorrelated, and for the data examined the correlations were low; and 2) heritabilities were computed in routine data analyses and were available at no additional cost.Joint contribution: USDA-SEA-AR and Journal Paper No. 10152 of the Iowa Agriculture and Home Economics Exp. Stn., Ames, Iowa. Project 2194.     

Phenotypic recurrent selection in common bean (Phaseolus vulgaris L.) based on performance of S2 progenies

Summary The effect of a recurrent selection procedure for improving seed yield per plant was evaluated in common bean. Progress was registered after three cycles of recurrent selection with S2 progeny evaluation performed on the grain yield/plant. Gain from selection was estimated by field testing 45 random S2 lines from each of the C0, C1 and C2 populations. Response to selection for seed yield/plant was 6 g from C0 to C1 and 4.4 g from C1 to C2. The seed yield improvement was due principally to an increase in pods/plant; little contribution was provided by the 1000-seed weight and seeds/pod traits. Broad-sense heritability and genotypic variance for grain yield remained high in all cycles which suggests further gain from additional selection cycles. Grain yield/plant is significantly correlated to pods/plant and seeds/pod traits in all cycles, indicating that plants with favourable combinations of these traits could be identified. The proposed recurrent selection procedure appears to be effective in improving the population and extracting superior genotypes for varietal development.     

Estimates of selection parameters associated with harvest index in oat lines derived from a bulk population

Summary Harvest index of grain crops is defined as grain yield divided by total plant yield. We estimated the heritability percentages of harvest index and its components, grain yield and plant weight, the genotypic and phenotypic correlations among these three traits, and the genotypic correlations of harvest index with plant height, 100-seed weight, grain number, and heading date by using a population of 1200 F₉-derived oat lines tested in six environments. Furthermore, we examined the relative selection efficiencies of indirect selection for yield through harvest index and of index selection for yield through yield plus harvest index.Heritability percentages were computed by use of variance-component, standard-unit, and realizedheritability methods. The heritability percentages for harvest index, grain yield, and plant weight were similar, and averaged between 50 and 65 precent across environments. Standard-unit and realized heritability percentages agreed closely and generally were lower than those computed via the variance-component method. Expected heritability percentages for harvest index calculated by using grain and plant yield data agreed almost exactly with actual values. Genotypic correlations were 0.88 between grain yield and plant weight, 0.42 between harvest index and grain yield, and –0.07 between harvest index and plant weight. Genotypic and phenotipic correlations were similar in magnitude. Theoretical and actual genotypic and phenotypic correlations of harvest index with grain yield and plant weight agreed closely. Genotypic correlations, computed via parent-offspring relationships, between harvest index and plant height, 100-seed weight, grain number, and heading date averaged –0.41, 0.43, 0.00, and –0.33 respectively.Indirect selection for grain yield through harvest index was 43 percent as efficient as direct selection, and a selection index that combined harvest index and grain yield was no more efficient than direct selection for yield. Harvest index had little value as a selection criterion for grain yield improvement when unrestricted selection was used. Indirect selection for grain yield through harvest index, however, would be expected to retain lines with a more favorable combination of yield, plant height and heading date than would unrestricted direct selection for yield.Journal Paper No. J-7914 of the Iowa Agriculture and Home Economic Experiment Station, Ames, Iowa 50010, Project No. 1752.     

Quantitative trait loci mapping of forage agronomic traits in six mapping populations derived from European elite maize germplasm

Four agronomic traits were analysed including dry matter concentration (DMC) and dry matter yield (DMY) for stover, plant height (PHT) and days from planting to silking (DPS). We mapped quantitative trait loci (QTL) in three populations with doubled haploid lines (DHL), one RIL population and two testcross (TC) populations derived from crosses between two of the four populations mentioned above to elite tester lines, based on field phenotyping at multiple locations and years for each; 146–168 SSRs were used for genotyping of the four mapping populations. Significant high phenotypic and genotypic correlations were found for all traits at two locations, while DMC was negatively correlated with the other traits. A total of 42, 41, 54, and 45 QTL were identified for DMC, DMY, PHT, and DPS, respectively, with 9, 7, 12, and 7 major QTL for each trait. Most detected QTL displayed significant interactions with environment. Major QTL detected in more than two populations will contribute to marker‐assisted breeding and also to fine mapping candidate genes associated with maize agronomic traits.     

Correlated response to recurrent selection for groat-oil content in oats

Summary One hundred random oat (Avena sativa L.) lines from a base (C₀) and each of three populations (C₁, C₂, and C₃) improved for groat (caryopsis) oil content by phenotypic recurrent selection were evaluated for correlated changes in several unselected agronomic traits. In addition, the parents of the base population and four check varieties were evaluated for the same traits. Phenotypic recurrent selection for high groat-oil content resulted in no significant correlated response in mean expression of any trait. Mean grain yield, biomass, groat yield, and harvest index of the improved populations were equal or superior to the mean of the parents and, with the exception of harvest index, equivalent to the mean of the check varieties. Mean test weight and seed weight of all populations were lower than for parents or check varieties. Selection for high groat-oil content caused a decline in genotypic variance for test weight and groat fraction, but reductions in genotypic variance for heading date and plant height may have resulted from culling for good agronomic type. Broad-sense heritability remained moderate to high for all traits except groat fraction. Phenotypic and genotypic correlation coefficients revealed negative, though mostly nonsignificant, relationships between groat-oil content and several traits, which may reflect a purported bioenergetic limitation to increasing groat-oil content in oats. Oil yield, however, was positively correlated with grain and groat yield, groat fraction, biomass, and harvest index. Results suggest that development of high-oil oat cultivars with current levels of production traits via phenotypic recurrent selection is possible.Journal Paper no. J-13038 of the Iowa Agric. and Home Econ. Exp. Stn., Ames, IA 50011. Project 2447.     

Quantitative trait loci mapping of forage stover quality traits in six mapping populations derived from European elite maize germplasm

Four forage maize stover quality traits were analysed including in vitro digestibility of organic matter (IVDOM), neutral detergent fibre (NDF), water‐soluble carbohydrates (WSC) and digestibility of NDF (DNDF). We mapped quantitative trait loci (QTL) in three DH (doubled haploid) populations (totally 250–720 DH lines): one RIL population (358 lines) and two testcross (TC) populations, based on field phenotyping at multiple locations and years for each. High phenotypic and genotypic correlations were found for all traits and significant (p < .01) at two locations, and NDF was negatively correlated with the other traits. QTL analyses were conducted by composite interval mapping. A total of 33, 23, 32 and 25 QTL were identified for IVDOM, NDF, WSC and DNDF, respectively, with three, four, five and two major QTL for each. Few consistent QTL for IVDOM, WSC and DNDF were detected in more than two populations. This study contributed to the identification of key QTL associated with forage maize digestibility traits and is beneficial for marker‐assisted breeding and fine mapping of candidate genes associated with forage maize quality.     

Effects of divergent selection on correlations between morphological traits in white clover (Trifolium repens L.)

The effects of two generations of divergent selection for stolon diameter and stolon amount on the pattern of phenotypic correlations between important agronomic traits in white clover (Trifolium repens L.) were studied. The effects of selection were: the proportions of the variances of traits under direct selection were considerably lower within selection lines than overall and this was associated with a loss of correlation between traits. Unselected traits showed higher proportions of variances remaining within the selection lines and shifts from non-significant to significant correlations from the plants overall to selection lines. The significance of these results and their relevance to strategies of germplasm improvement is discussed. This revised version was published online in July 2006 with corrections to the Cover Date.     

Quantitative trait loci of frost tolerance and physiologically related trait in faba bean (Vicia faba L.)

In faba bean, field based winter-hardiness is a complex trait that is significantly correlated to frost tolerance. Frost tolerance could be used to indirectly select for faba bean winter-hardiness. The aim of this study was to identify putative QTL associated with frost tolerance and auxiliary traits and to quantify the efficiency of marker assisted selection. Thus, 101 recombinant inbred lines derived from the cross between two frost tolerant lines were tested for their hardened and unhardened frost tolerance and for their leaf fatty acid content in both treatments. Significant differences among the RIL were observed for all studied traits. For frost tolerance, five putative QTL were detected; three for unhardened frost tolerance that explained 40.7% (8.6% after cross-validation, CV) of its genotypic variance and two for hardened frost tolerance that explained 21.8% (1.0% after CV). For fatty acid content, three QTL were detected for oleic acid content in unhardened leaves that explained 62.9% (40.6% after CV) of its genotypic variance. This fatty acid was significantly correlated with unhardened frost tolerance. The unbiased genotypic variance explained enabled to draw realistic prospects of MAS for frost tolerance. In this study, combined MAS was more efficient than classical phenotypic selection and was expected to be higher on larger populations at early generations. Moreover, favourable alleles inherited from the exotic line BPL 4628 could be introgressed to European winter-hardy beans for further improvement.     

Genetic analyses for bioactive potential in tetraploid potato (Solanum tuberosum L.)

A total of four traits related to bioactive potential of the tetraploid potato were studied in a North Carolina II cross‐experiment. The objectives of this study were to estimate genetic parameters for target bioactive compounds in a tetraploid potato population, to determine phenotypic and genotypic correlations and to calculate combining ability effects of cultivars and breeding lines included in the crossing scheme. Total monomeric anthocyanins, phenolics, carotenoids and hydrophilic antioxidant capacity were measured by spectrophotometric methods. The sources of variation female (GCA_f), male (GCA_m) and their interaction (SCA) were significant for bioactive compounds and antioxidant capacity. Narrow‐sense heritability estimates (h²) were moderate for all the traits (0.413–0.657). Both phenotypic and genotypic correlations were highly significant for phenolic compounds and antioxidant capacity. As regards combining ability, positive and negative GCA and also SCA effects have been identified for bioactive compounds and antioxidant capacity. These results indicate that hybrid breeding lines can be obtained by choosing parents with significant GCA, whose bioactive potential can also be further enhanced if the crosses have a significant SCA.     

Mass selection for seed width in oat populations

Heterogeneous and segregating oat populations subjected to no selection and to mass selection for seed width for five consecutive generations were evaluated for 100-seed weight, heading date and plant height.The mass-selection technique, which consisted of passing the seeds over a screen with 12.7×2.4 mm slots and propagating the seeds too wide to go through the slots, was repeated for five consecutive generations (F₃ through F₇). Comparable unselected populations were grown each year. Remnant seeds from each check and mass-selected population were stored for future studies.In 1963, 500 random seeds from each of the six unselected populations (F₂ through F₇), from each of the five mass-selected populations (F₃ through F₇) and from a pure line were space-planted in the field. In 1964, the progenies from 75 random strains (a strain was the progeny of one spaced plant) from each of the 12 populations were sown in a randomized-block experiment with three replicates. In 1966, the strains from the last (F₇) generation of the unselected and mass-selected lines of descent (75 strains from each) were tested for yield in a 7-replicate randomized block experiment.The mass-selection technique increased the mean 100-seed weight, whereas in the unselected line of descent, the mean 100-seed weight did not change. The mean heading dates became later and the mean plant heights taller with the mass-selection procedure. The mean yield of the final mass-selection population was nine percent higher than that of the comparable unselected population.In spite of the associated changes between 100-seed weight and plant height and heading date in successive generations; the genotypic correlations among strains within populations indicated that these three traits were inherited independently. In advanced generations (F₆ and F₇), the genotypic variances for heading date and 100-seed weight were reduced in both the unselected and mass-selected lines of descent, whereas, for plant height no significant change occurred in either.Journal No. J-5586 of the Iowa Agricultural and Home Economics Experiment Station, Ames, Iowa, Project 1176. In cooperation with the Crops Research Division, Agricultural Research Service. U.S. Department of Agriculture.Professor of Plant Breeding.     

Evaluation of genetic variances, heritabilities, and correlations for yield and fiber traits among cotton F2 hybrid populations

Summary F₂ hybrid cultivars continue to occupy a small portion of the cotton (Gossypium hirsutum L.) production are in the United States, but occupy a larger proportion of the production area in some other countries. Sixty-four F₂ hybrids resulting from crosses of four commercial cultivars and 16 pest-resistant germplasm lines were evaluated for five fiber and four yield traits in four environments at Mississippi State, MS. An additive-dominance genetic model was employed for these traits. The minimum norm quadratic unbiased estimation (MINQUE) method was used with a mixed model approach for estimating genetic variance and covariance components and for predicting genetic correlations. This study investigated genetic variances, heritabilities, and genetic and phenotypic correlations between agronomic and fiber traits among these 64 F₂ hybrid populations and discussed the usefulness of these populations for use as hybrids or for selections for pure lines.Dominance variance accounted for the major proportion of the phenotypic variances for lint yield, lint percentage, and boll size indicating that hybrids should have an advantage for these traits compared to pure lines. A low proportion of additive variance for fiber traits and the significant additive x environment variance components indicated a lack of substantial useful additive genetic variability for fiber traits. This suggests that selections for pure lines within these F₂ populations would have limited success in improving fiber traits. Genetic and phenotypic correlation coefficients were of comparable magnitude for most pairs of characters. Fiber strength showed a positive additive genetic correlation with boll weight. Dominance genetic correlations of fiber strength with elongation and 2.5% span length were also significant and positive; however, the additive genetic correlation of length and strength was zero.Contribution of the USDA-ARS in cooperation with the Mississippi Agric. and Forestry Exp. Stn.     

Maize participatory breeding in Portugal: Comparison of farmer's and breeder's on‐farm selection

“VASO” is a Portuguese participatory maize breeding project (1984), where several maize landraces such as “Pigarro” have been selected both by a farmer's (phenotypic recurrent selection) and a breeder's approach (S2 lines recurrent selection). The objectives of this study were to determine the phenotypic and genotypic responses to participatory selection using these two different approaches, to clarify to which extent both selection methods preserve genetic diversity, and conclude what is the preferred method to apply in sustainable farming systems. The results, obtained via ANOVA, regression analyses and molecular markers, indicate that for both selection methods, genetic diversity was not significantly reduced, even with the most intensive breeder's selection. Although there were some common outputs, such as the determinated versus indeterminated ears, cob and ear weight ratio per ear and rachis 2, specific phenotypic traits evolved in opposite directions between the two selection approaches. Yield increase was only detected during farmer selection, indicating its interest on PPB. Candidate genes were identified for a few of the traits under selection as potential functional markers in participatory plant breeding.     

Phenotypic and genotypic analyses of diversity and breeding progress in European triticale (× Triticosecale Wittmack)

Knowledge of the extent of phenotypic variability, genetic diversity and the realized breeding progress is central for the optimum design of breeding programmes, but little information is available for triticale (× Triticosecale Wittmack). In this study, a collection of 885 diverse European triticale lines was evaluated in multilocation field trials in 2 years. We observed significant genotypic variances and high heritabilities for several agronomic and morphological traits and significant correlations among different traits including grain yield. Based on a subset of the population of 121 varieties registered in Europe between 1983 and 2014, we observed a substantial breeding progress for grain yield with a significant rate of increase of 53 kg/ha or 0.67% per year. All lines were genotyped by a genotyping‐by‐sequencing approach yielding 58 888 polymorphic markers. Our analyses revealed the absence of major population structure but a certain grouping of lines according to their origin. Taken together, our results on triticale germplasm and its breeding history provide important information for breeding programmes and future selection gain in this crop.     

Genetic diversity for agronomic and fiber traits in day-neutral accessions derived from primitive cotton germplasm

Upland cotton (Gossypium hirsutumL) is an important crop that is cultivated in warm climates through-out the world. Agronomic performance and fiber quality must continually be improved if cotton is to maintain economic viability. Primitive ancestors of cotton contain diversity for trait improvement; however, many of these accessions have a short-day flowering response (photoperiodic) and are not readily useable in breeding programs. In this study, 114 day-neutral derived primitive germplasm lines were evaluated in field trials for two years. Agronomic and fiber trait data were collected and analyzed. Variance components, genotypic values, and genotypic correlations were calculated. Genotypic effects for all traits studied made significant contributions to the phenotypic variation indicating genetic diversity among these lines. The predicted genotypic values showed a wide range of variation for agronomic and fiber traits. Weak genotypic correlations were found between yield and 2.5% span length and fiber strength, two important fiber traits. Although these day-neutral derived accessions had lower lint percentage, they had improved fiber length, strength, micronaire, and comparable yields with two commercial cultivars. Thus, these day-neutral derived accessions are sources of genetic variation that when used in breeding programs offer the potential to improve important traits and expand genetic diversity.Contribution of the USDA-ARS in cooperation with the Mississippi Agric. and Forestry Exp. Stn. Mention of trademark, proprietary product, or vendor does not constitute a guarantee or warranty of the product by USDA, ARS and does not imply its approval to the exclusion of other products or vendors that may also be suitable.     

Evaluation of strategies to optimize training populations for genomic prediction in oat (Avena sativa)

Genomic selection is a promising breeding methodology that could increase selection accuracy and intensity and reduce generation interval. As the cost of genotyping decreases, it will be important to optimize training populations for costly phenotypic experiments for many complex traits. The aim of this research was to evaluate different optimization strategies, by using historical data from the Norwegian oat breeding programme at Graminor. In this paper, we focus on the optimization criteria: genetic diversity, phenotypic variance and genetic similarity between the training and testing populations. The four training population strategies—prediction core, diversity core, phenotypic selection and random selection—were applied to an oat candidate population of 1124 lines. An independent testing population was used to calculate the mean prediction abilities for the traits days to heading and plant height. Moreover, the strategies were tested in three independent wheat populations. The results showed that prediction core was the most promising strategy to select training populations with high genetic similarity to the testing set, high genetic diversity, and high phenotypic variance, which resulted in higher prediction ability across population sizes and traits.     

Inheritance of fruit yield in two watermelon populations in North Carolina

Elite cultivars of watermelon (Citrullus lanatus (Thunb.) Matsum. & Nakai) are high in fruit quality but may not be the highest yielders. The objective of this study was to estimate the heritability of, and genotypic and phenotypic correlations among, yield traits in two watermelon populations developed from crosses between obsolete cultivars with high yield and elite modern cultivars. Field trials were conducted at two locations in North Carolina (Clinton and Kinston). The data were analyzed by regressing S_0:1 progeny data on S₀ parent data to estimate narrow-sense heritability. Narrow-sense heritability estimates were low for all traits measured [total fruit weight (0.04–0.12), marketable fruit weight (0.06–0.15), total fruit number (0.04–0.16), fruit size (0.18–0.19), and percent culls (0.02–0.09) in North Carolina Watermelon 1 (NCWP1) and North Carolina Watermelon 2 (NCWP2) populations, respectively]. Estimates of broad-sense heritability were higher than estimates of narrow-sense heritability. Total fruit weight and marketable fruit weight were highly correlated (r _g = 0.97–1.00). Marketable fruit weight and fruit size used as single selection criteria in NCWP1 and total fruit number in NCWP2, were predicted to give the best correlated response for total fruit weight. Narrow-sense heritability was low for fruit yield; therefore, watermelon breeders should select based on replicated progeny rows in multiple environments to maximize gain.     

籼型黑米稻粒形性状与其中某些矿质元素含量的遗传相关性

采用禾谷类作物种子数量性状遗传模型, 分析籼型黑米稻品种双列杂交F1和F2种子粒形性状与其中矿质元素Fe、 Zn、 Mn和P含量间的多种遗传相关性. 结果表明, 籼型黑米粒重、粒长、粒宽、粒长/粒宽与其中Fe、 Zn、 Mn和P含量除存在明显的表现型相关和基因型相关外, 尚存在较明显的种子直接加性相关、直接显性相关、细胞质相关、