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1.

Key message

Using newly developed euchromatin-derived genomic SSR markers and a flexible Bayesian mapping method, 13 significant agricultural QTLs were identified in a segregating population derived from a four-way cross of tomato.

Abstract

So far, many QTL mapping studies in tomato have been performed for progeny obtained from crosses between two genetically distant parents, e.g., domesticated tomatoes and wild relatives. However, QTL information of quantitative traits related to yield (e.g., flower or fruit number, and total or average weight of fruits) in such intercross populations would be of limited use for breeding commercial tomato cultivars because individuals in the populations have specific genetic backgrounds underlying extremely different phenotypes between the parents such as large fruit in domesticated tomatoes and small fruit in wild relatives, which may not be reflective of the genetic variation in tomato breeding populations. In this study, we constructed F2 population derived from a cross between two commercial F1 cultivars in tomato to extract QTL information practical for tomato breeding. This cross corresponded to a four-way cross, because the four parental lines of the two F1 cultivars were considered to be the founders. We developed 2510 new expressed sequence tag (EST)-based (euchromatin-derived) genomic SSR markers and selected 262 markers from these new SSR markers and publicly available SSR markers to construct a linkage map. QTL analysis for ten agricultural traits of tomato was performed based on the phenotypes and marker genotypes of F2 plants using a flexible Bayesian method. As results, 13 QTL regions were detected for six traits by the Bayesian method developed in this study.
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2.
Crepieux S  Lebreton C  Servin B  Charmet G 《Genetics》2004,168(3):1737-1749
Mapping quantitative trait loci in plants is usually conducted using a population derived from a cross between two inbred lines. The power of such QTL detection and the parameter estimates depend largely on the choice of the two parental lines. Thus, the QTL detected in such populations represent only a small part of the genetic architecture of the trait. In addition, the effects of only two alleles are characterized, which is of limited interest to the breeder, while common pedigree breeding material remains unexploited for QTL mapping. In this study, we extend QTL mapping methodology to a generalized framework, based on a two-step IBD variance component approach, applicable to any type of breeding population obtained from inbred parents. We then investigate with simulated data mimicking conventional breeding programs the influence of different estimates of the IBD values on the power of QTL detection. The proposed method would provide an alternative to the development of specifically designed recombinant populations, by utilizing the genetic variation actually managed by plant breeders. The use of these detected QTL in assisting breeding would thus be facilitated.  相似文献   

3.
Four-way cross (4WC) involving four different inbred lines frequently appears in the cotton breeding programs. However, linkage analysis and quantitative trait loci (QTL) mapping with molecular markers in cotton has largely been applied to populations derived from a cross between two inbred lines, and few results of QTL dissection were conducted in a 4WC population. In this study, an attempt was made to construct a linkage map and identify QTL for yield and fiber quality traits in 4WC derived from four different inbred lines in Gossypium hirsutum L. A linkage map was constructed with 285 SSR loci and one morphological locus, covering 2113.3 cM, approximately 42% of the total recombination length of the cotton genome. A total of 31 QTL with 5.1–25.8% of the total phenotypic variance explained were detected. Twenty-four common QTL across environments showed high stability, and six QTL were environment-specific. Several genomic segments affecting multiple traits were identified. The advantage of QTL mapping using a 4WC were discussed. This study presents the first example of QTL mapping using a 4WC population in upland cotton. The results presented here will enhance the understanding of the genetic basis of yield and fiber quality traits and enable further marker-assisted selection in cultivar populations in upland cotton.  相似文献   

4.
 We report results from a breeding strategy designed to accumulate favorable QTL alleles for grain yield identified in the SteptoeבMorex’ (SM) barley germplasm. Two map lines (SM73 and SM145) from the original mapping population were selected based on their marker genotype and QTL structure. When crossed, these lines would be expected to produce progeny with most favorable QTL alleles. One hundred doubled haploid (DH) lines from the F1 hybrid of this cross were genotyped with ten RFLP markers and one morphological marker defining grain yield to monitor QTL segregation. A subset of 24 lines representing various combinations of putatively favorable and unfavorable QTL alleles, together with Steptoe, ‘Morex’, SM73, and SM145, were phenotyped for grain yield in five environments. Multiple regression procedures were used to explore phenotype and genotype relationships. Most target QTLs showed significant effects. However, significance and magnitude of QTL effects and favorable QTL allele phase varied across environments. All target QTLs showed significant QTL-by-environment interaction (QTL×E), and the QTL on chromosome 2 expressed alternative favorable QTL alleles in different environments. Digenic epistatic effects were also detected between some QTL loci. For traits such as grain yield, marker-assisted selection efforts may be better targeted at determining optimum combinations of QTL alleles rather than pyramiding alleles detected in a reference mapping population. Received: 2 June 1998 / Accepted: 17 September 1998  相似文献   

5.
Durum wheat has the tendency of accumulating more cadmium (Cd), a biotoxic heavy metal, in seeds than other commonly grown cereals, thus posing a serious food safety/public health concern. This could have serious negative impact on the national pasta industry and the international export market of durum wheat. The phenotyping for selecting low Cd lines is expensive and time consuming. The use of markers could be a more sustainable approach for selecting lines with low Cd levels. Here, a RIL population developed from a cross between Grenora (high Cd) × Haurani (low Cd) and two association mapping panels consisting of advanced breeding lines from the North Dakota durum wheat breeding program were used to identify QTL and associated markers for Cd. A major QTL, with Haurani contributing low Cd uptake allele and explaining 54.3 % phenotypic variation, was detected on chromosome 5BL. Association mapping using 2010 collection validated the results of linkage mapping and identified major QTL on 5BL. The 2009 collection, showed the presence of a major QTL on chromosome 2B. The SNP marker associated with major QTL on 5BL was converted to user friendly KASPar assay. The major QTL and associated KASPar marker were further validated using another RIL population developed from a cross of Strongfield (low Cd) and Alkabo (high Cd). The development of suitable marker assay, associated with major Cd uptake QTL, would help the selection for low Cd accumulating lines, minimizing the costly phenotypic evaluation for this important trait.  相似文献   

6.
为了提高黄淮海麦区小麦育种材料的赤霉病抗性,采用分子标记辅助选择的方法,将来自望水白的4个抗赤霉病主效QTL 3B-QTL、4B-QTL、5A-QTL和6B-QTL导入不同的感病背景中,在后代BC1F3和BC1F4株系中评价它们的抗病效应和农艺性状回复情况。结果表明:(1)导入4个抗病QTL株系的平均病小穗率和病粒率分别为12.2%和6.3%,而受体亲本则分别达到59.1%和44.2%,抗病性显著提高;(2)病小穗数和病粒率与穗长及株高极显著负相关,但与可育小穗数、百粒重、旗叶长和旗叶宽等农艺性状指标没有显著相关性。因此,通过导入抗病主效QTL可以显著改善感病材料的抗性,为进一步选育高产抗病品种提供基础材料。不良农艺性状的紧密连锁阻碍着抗赤霉病主效QTL的高效利用,需要通过继续回交或与其他品种杂交来打破这种遗传连锁关系。  相似文献   

7.
Briggs WH  McMullen MD  Gaut BS  Doebley J 《Genetics》2007,177(3):1915-1928
An ultimate objective of QTL mapping is cloning genes responsible for quantitative traits. However, projects seldom go beyond segments <5 cM without subsequent breeding and genotyping lines to identify additional crossovers in a genomic region of interest. We report on a QTL analysis performed as a preliminary step in the development of a resource for map-based cloning of domestication and improvement genes in corn. A large backcross (BC)1 population derived from a cross between maize (Zea mays ssp. mays) and teosinte (ssp. parviglumis) was grown for the analysis. A total of 1749 progenies were genotyped for 304 markers and measured for 22 morphological traits. The results are in agreement with earlier studies showing a small number of genomic regions having greater impact on the morphological traits distinguishing maize and teosinte. Despite considerable power to detect epistasis, few QTL interactions were identified. To create a permanent resource, seed of BC1 plants was archived and 1000 BC2S6 BC1-derived lines are in development for fine mapping and cloning. The identification of four BC1 progeny with crossovers in a single gene, tb1, indicated that enough derived lines already exist to clone many QTL without the need to generate and identify additional crossovers.  相似文献   

8.
Specific traits are an important consideration in plant breeding. In popcorn, inferior agronomic traits could be improved using dent or flint corn backcrossed with popcorn. In this study, we used advanced backcross quantitative trait locus (AB-QTL) analysis to identify trait-improving QTL alleles from a dent maize inbred Dan232, and compared the detection of QTL in the BC2S1 population with QTL results using F2:3 families of the same population. Two hundred and twenty BC2S1 families developed from a cross between Dan232 and an elite popcorn inbred N04 were evaluated for nine plant traits in replicated field trials under two environments. Using composite interval mapping (CIM), a total of 28 significant QTL were detected, and of these, 23 (82.14%) had favorable alleles contributed by the dent corn parent Dan232. Nine QTL (32.14%) detected in the BC2S1 population were also located in or near the same chromosome intervals in the F2:3 population. All of the favorable QTL alleles from Dan232 could be used in marker-assisted selection (MAS) to improve the respective plant traits in popcorn breeding. In addition, their near isogenic lines (QTL-NILs) could be obtained through selfing or another 1–2 backcross with N04. Also, N04 improved for the studied plant traits could be developed from the BC2S1 families used in this study. This study demonstrated that the AB-QTL method can be applied to identify favorable QTL from dent corn inbred in popcorn breeding and, once identified, the alleles could be used in marker-assisted selection to improve the respective plant traits.  相似文献   

9.
Sorghum, a cereal of economic importance ensures food and fodder security for millions of rural families in the semi-arid tropics. The objective of the present study was to identify and validate quantitative trait loci (QTL) for grain yield and other agronomic traits using replicated phenotypic data sets from three post-rainy dry sorghum crop seasons involving a mapping population with 245 F9 recombinant inbred lines derived from a cross of M35-1 × B35. A genetic linkage map was constructed with 237 markers consisting of 174 genomic, 60 genic and 3 morphological markers. The QTL analysis for 11 traits following composite interval mapping identified 91 QTL with 5–12 QTL for each trait. QTL detected in the population individually explained phenotypic variation between 2.5 and 30.3 % for a given trait and six major genomic regions with QTL effect on multiple traits were identified. Stable QTL across seasons were identified. Of the 60 genic markers mapped, 21 were found at QTL peak or tightly linked with QTL. A gene-based marker XnhsbSFCILP67 (Sb03g028240) on SBI-03, encoding indole-3-acetic acid-amido synthetase GH3.5, was found to be involved in QTL for seven traits. The QTL-linked markers identified for 11 agronomic traits may assist in fine mapping, map-based gene isolation and also for improving post-rainy sorghum through marker-assisted breeding.  相似文献   

10.
Utilization of quantitative trait loci (QTL) identified in bi-parental mapping populations has had limited success for improving complex quantitative traits with low to moderate heritability. Association mapping in contemporary breeding germplasm may lead to more effective marker strategies for crop improvement. To test this approach, we conducted association mapping of two complex traits with moderate heritability; Fusarium head blight (FHB) severity and the grain concentration of mycotoxin associated with disease, deoxynivalenol (DON). To map FHB resistance in barley, 768 breeding lines were evaluated in 2006 and 2007 in four locations. All lines were genotyped with 1,536 SNP markers and QTL were mapped using a mixed model that accounts for relatedness among lines. Average linkage disequilibrium within the breeding germplasm extended beyond 4 cM. Four QTL were identified for FHB severity and eight QTL were identified for the DON concentration in two independent sets of breeding lines. The QTL effects were small, explaining 1–3% of the phenotypic variation, as might be expected for complex polygenic traits. We show that using breeding germplasm to map QTL can complement bi-parental mapping studies by providing independent validation, mapping QTL with more precision, resolving questions of linkage and pleiotropy, and identifying genetic markers that can be applied immediately in crop improvement.  相似文献   

11.
While many reports on genetic analysis of Fusarium head blight (FHB) resistance in bread wheat have been published during the past decade, only limited information is available on FHB resistance derived from wheat relatives. In this contribution, we report on the genetic analysis of FHB resistance derived from Triticum macha (Georgian spelt wheat). As the origin of T. macha is in the Caucasian region, it is supposed that its FHB resistance differs from other well-investigated resistance sources. To introduce valuable alleles from the landrace T. macha into a modern genetic background, we adopted an advanced backcross QTL mapping scheme. A backcross-derived recombinant-inbred line population of 321 BC2F3 lines was developed from a cross of T. macha with the Austrian winter wheat cultivar Furore. The population was evaluated for Fusarium resistance in seven field experiments during four seasons using artificial inoculations. A total of 300 lines of the population were genetically fingerprinted using SSR and AFLP markers. The resulting linkage map covered 33 linkage groups with 560 markers. Five novel FHB-resistance QTL, all descending from T. macha, were found on four chromosomes (2A, 2B, 5A, 5B). Several QTL for morphological and developmental traits were mapped in the same population, which partly overlapped with FHB-resistance QTL. Only the 2BL FHB-resistance QTL co-located with a plant height QTL. The largest-effect FHB-resistance QTL in this population mapped at the spelt-type locus on chromosome 5A and was associated with the wild-type allele q, but it is unclear whether q has a pleiotropic effect on FHB resistance or is closely linked to a nearby resistance QTL.  相似文献   

12.
Linkage analysis of quantitative trait loci in multiple line crosses   总被引:8,自引:0,他引:8  
Yi N  Xu S 《Genetica》2002,114(3):217-230
Simple line crosses, for example, backcross and F2, are commonly used in mapping quantitative trait loci (QTL). However, these simple crosses are rarely used alone in commercial plant breeding; rather, crosses involving multiple inbred lines or several simple crosses but connected by shared inbred lines may be common in plant breeding. Mapping QTL using crosses of multiple lines is more relevant to plant breeding. Unfortunately, current statistical methods and computer programs of QTL mapping are all designed for simple line crosses or multiple line crosses but under a regular mating system. It is not straightforward to extend the existing methods to handle multiple line crosses under irregular and complicated mating designs. The major hurdle comes from irregular inbreeding, multiple generations, and multiple alleles. In this study, we develop a Bayesian method implemented via the Markov chain Monte Carlo (MCMC) algorithm for mapping QTL using complicated multiple line crosses. With the MCMC algorithm, we are able to draw a complete path of the gene flow from founder alleles to their descendents via a recursive process. This has greatly simplified the problem caused by irregular mating and inbreeding in the mapping population. Adopting the reversible jump MCMC algorithm, we are able to simultaneously search for multiple QTL along the genome. We can even infer the posterior distribution of the number of QTL, one of the most important parameters in QTL study. Application of the new MCMC based QTL mapping procedure is demonstrated using two different mating designs. Design I involves two inbred lines and their derived F1, F2, and BC populations. Design II is a half-diallel cross involving three inbred lines. The two designs appear different, but can be handled with the same robust computer program.  相似文献   

13.
Oil content in rapeseed (Brassica napus L.) is generally regarded as a character with high heritability that is negatively correlated with protein content and influenced by plant developmental and yield related traits. To evaluate possible genetic interrelationships between these traits and oil content, QTL for oil content were mapped using data on oil content and on oil content conditioned on the putatively interrelated traits. Phenotypic data were evaluated in a segregating doubled haploid population of 282 lines derived from the F1 of a cross between the old German cultivar Sollux and the Chinese cultivar Gaoyou. The material was tested at four locations, two each in Germany and in China. QTLMapper version 1.0 was used for mapping unconditional and conditional QTL with additive (a) and locus pairs with additive × additive epistatic (aa) effects. Clear evidence was found for a strong genetic relationship between oil and protein content. Six QTL and nine epistatic locus pairs were found, which had pleiotropic effects on both traits. Nevertheless, two QTL were also identified, which control oil content independent from protein content and which could be used in practical breeding programs to increase oil content without affecting seed protein content. In addition, six additional QTL with small effects were only identified in the conditional mapping. Some evidence was apparent for a genetic interrelationship between oil content and the number of seeds per silique but no evidence was found for a genetic relationship between oil content and flowering time, grain filling period or single seed weight. The results indicate that for closely correlated traits conditional QTL mapping can be used to dissect the genetic interrelationship between two traits at the level of individual QTL. Furthermore, conditional QTL mapping can reveal additional QTL with small effects that are undetectable in unconditional mapping.  相似文献   

14.
We report here the second advanced backcross quantitative trait locus (AB-QTL) analysis carried out in winter wheat. Seven agronomic traits were studied in a BC2F1population derived from a cross between the German winter wheat variety Flair and the synthetic wheat line XX86 developed in Japan. We selected 111 BC2F1 lines and genotyped these with 197 microsatellite markers. Field data for seven agronomic traits were collected from corresponding BC2F3 families that were grown at up to six locations in Germany. QTL analyses for yield and yield components were performed using single-marker regression and interval mapping. A total of 57 putative QTLs derived from XX86 were detected, of which 24 (42.1%) were found to have a positive effect from the synthetic wheat XX86. These favourable QTLs were mainly associated with thousand-grain weight and grain weight per ear. Many QTLs for correlated traits were mapped in similar chromosomal regions. The AB-QTL data obtained in the present study are discussed and compared with results from previous QTL analyses.  相似文献   

15.
Mathematically-derived traits from two or more component traits, either by addition, subtraction, multiplication, or division, have been frequently used in genetics and breeding. When used in quantitative trait locus (QTL) mapping, derived traits sometimes show discrepancy with QTL identified for the component traits. We used three QTL distributions and three genetic effects models, and an actual maize mapping population, to investigate the efficiency of using derived traits in QTL mapping, and to understand the genetic and biological basis of derived-only QTL, i.e., QTL identified for a derived trait but not for any component trait. Results indicated that the detection power of the four putative QTL was consistently greater than 90% for component traits in simulated populations, each consisting of 200 recombinant inbred lines. Lower detection power and higher false discovery rate (FDR) were observed when derived traits were used. In an actual maize population, simulations were designed based on the observed QTL distributions and effects. When derived traits were used, QTL detected for both component and derived traits had comparable power, but those detected for component traits but not for derived traits had low detection power. The FDR from subtraction and division in the maize population were higher than the FDR from addition and multiplication. The use of derived traits increased the gene number, caused higher-order gene interactions than observed in component traits, and possibly complicated the linkage relationship between QTL as well. The increased complexity of the genetic architecture with derived traits may be responsible for the reduced detection power and the increased FDR. Derived-only QTL identified in practical genetic populations can be explained either as minor QTL that are not significant in QTL mapping of component traits, or as false positives.  相似文献   

16.
Grain yield and associated agronomic traits are important factors in wheat (Triticum aestivum L.) improvement. Knowledge regarding the number, genomic location, and effect of quantitative trait loci (QTL) would facilitate marker-assisted selection and the development of cultivars with desirable characteristics. Our objectives were to identify QTLs directly and indirectly affecting grain yield expression. A population of 132 F12 recombinant inbred lines (RILs) was derived by single-seed descent from a cross between the Chinese facultative wheat Ning7840 and the US soft red winter wheat Clark. Phenotypic data were collected for 15 yield and other agronomic traits in the RILs and parental lines from three locations in Oklahoma from 2001 to 2003. Twenty-nine linkage groups, consisting of 363 AFLP and 47 SSR markers, were identified. Using composite interval mapping (CIM) analysis, 10, 16, 30, and 14 QTLs were detected for yield, yield components, plant adaptation (shattering and lodging resistance, heading date, and plant height), and spike morphology traits, respectively. The QTL effects ranged from 7 to 23%. Marker alleles from Clark were associated with a positive effect for the majority of QTLs for yield and yield components, but gene dispersion was the rule rather than the exception for this RIL population. Often, QTLs were detected in proximal positions for different traits. Consistent, co-localized QTLs were identified in linkage groups 1AL, 1B, 4B, 5A, 6A, and 7A, and less consistent but unique QTLs were identified on 2BL, 2BS, 2DL, and 6B. Results of this study provide a benchmark for future efforts on QTL identification for yield traits.  相似文献   

17.
Starch pasting viscosity is an important quality trait in cassava (Manihot esculenta Crantz) cultivars. The aim here was to identify loci and candidate genes associated with the starch pasting viscosity. Quantitative trait loci (QTL) mapping for seven pasting viscosity parameters was carried out using 100 lines of an F1 mapping population from a cross between two cassava cultivars Huay Bong 60 and Hanatee. Starch samples were obtained from roots of cassava grown in 2008 and 2009 at Rayong, and in 2009 at Lop Buri province, Thailand. The traits showed continuous distribution among the F1 progeny with transgressive variation. Fifteen QTL were identified from mean trait data, with Logarithm of Odds (LOD) values from 2.77–13.01 and phenotype variations explained (PVE) from10.0–48.4%. In addition, 48 QTL were identified in separate environments. The LOD values ranged from 2.55–8.68 and explained 6.6–43.7% of phenotype variation. The loci were located on 19 linkage groups. The most important QTL for pasting temperature (PT) (qPT.1LG1) from mean trait values showed largest effect with highest LOD value (13.01) and PVE (48.4%). The QTL co‐localised with PT and pasting time (PTi) loci that were identified in separate environments. Candidate genes were identified within the QTL peak regions. However, the major genes of interest, encoding the family of glycosyl or glucosyl transferases and hydrolases, were located at the periphery of QTL peaks. The loci identified could be effectively applied in breeding programmes to improve cassava starch quality. Alleles of candidate genes should be further studied in order to better understand their effects on starch quality traits.  相似文献   

18.
小麦苗期水分利用效率及其相关性状的QTL分析   总被引:13,自引:0,他引:13  
以小麦DH群体(旱选10号×鲁麦14)为研究材料,采用复合区间作图法,对小麦幼苗在水分胁迫及非胁迫条件下的水分利用效率(WUE)及其相关性状的QTL进行定位,并对比分析QTL的加性效应.两种水分条件下共检测到14个具显著加性效应的QTL,分布在2A、3A、4A、5A、6A、7A、1B、3B、3D染色体上,可解释表型变异的范围在6.36%~19.73%.其中,非胁迫(对照)条件下检测到10个QTL,包括2个单株WUE的QTL,5个地上部WUE的QTL,1个根系WUE的QTL及2个总耗水量的QTL;水分胁迫条件下上述性状各检测到1个QTL.对于同一性状没有检测到在两种水分条件下均位于同一标记区间的QTL,表明不同水分环境条件下同一性状的QTL表达模式是不同的.论文也讨论了可能用于标记辅助选择的QTL及其分子标记.  相似文献   

19.
Detection of QTL in multiple segregating populations is of high interest as it includes more alleles than mapping in a single biparental population. In addition, such populations are routinely generated in applied plant breeding programs and can thus be used to identify QTL which are of direct relevance for a marker-assisted improvement of elite germplasm. Multiple-line cross QTL mapping and joint linkage association mapping were used for QTL detection. We empirically compared these two different biometrical approaches with regard to QTL detection for important agronomic traits in nine segregating populations of elite rapeseed lines. The plants were intensively phenotyped in multi-location field trials and genotyped with 253 SNP markers. Both approaches detected several additive QTL for diverse traits, including flowering time, plant height, protein content, oil content, glucosinolate content, and grain yield. In addition, we identified one epistatic QTL for flowering time. Consequently, both approaches appear suited for QTL detection in multiple segregating populations.  相似文献   

20.
Plant breeders simultaneously select for qualitative traits controlled by one or a small number of major genes, as well as for polygenic traits controlled by multiple genes that may be detected as quantitative trait loci (QTL). In this study, we applied computer simulation to investigate simultaneous selection for alleles at both major and minor gene (as QTL) loci in breeding populations of two wheat parental lines, HM14BS and Sunstate. Loci targeted for selection included six major genes affecting plant height, disease resistance, and grain quality, plus 6 known and 11 “unidentified” QTL affecting coleoptile length (CL). Parental line HM14BS contributed the target alleles at two of the major gene loci, while parental line Sunstate contributed target alleles at four loci. The parents have similar plant height, but HM14BS has a longer coleoptile, a desirable attribute for deep sowing in rainfed environments. Including the wild-type allele at the major reduced-height locus Rht-D1, HM14BS was assumed to have 13 QTL for increased CL, and Sunstate four; these assumptions being derived from mapping studies and empirical data from an actual HM14BS/Sunstate population. Simulation indicated that compared to backcross populations, a single biparental F1 cross produced the highest frequency of target genotypes (six desired alleles at major genes plus desired QTL alleles for long CL). From 1,000 simulation runs, an average of 2.4 individuals with the target genotype were present in unselected F1-derived doubled haploid (DH) or recombinant inbred line (RIL) populations of size 200. A selection scheme for the six major genes increased the number of target individuals to 19.1, and additional marker-assisted selection (MAS) for CL increased the number to 23.0. Phenotypic selection (PS) of CL outperformed MAS in this study due to the high heritability of CL, incompletely linked markers for known QTL, and the existence of unidentified QTL. However, a selection scheme combining MAS and PS was equally as efficient as PS and would result in net savings in production and time to delivery of long coleoptile wheats containing the six favorable alleles.  相似文献   

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