Comparative mapping of QTLs for agronomic traits of rice across environments by using a doubled-haploid population

We report here the RFLP mapping of quantitative trait loci (QTLs) which affect some important agronomic traits in cultivated rice. An anther culture-derived doubled-haploid (DH) population was established from a cross between indica and japonica rice varieties. A molecular linkage map comprising 137 markers was constructed based on this population which covered the rice genome at intervals of 14.8 cM on average. The linkage map was used to locate QTLs for such important agronomic traits as heading date, plant height, number of spikelets per panicle, number of grains per panicle, 1 000-grain weight and the percentage of seed set, by interval mapping. Evidence of genotype-by-environment interaction was found by comparing QTL maps of the same population grown in three diverse environments. A total of 22 QTLs for six agronomic traits was detected which were significant in at least one environment, but only seven were significant in all three environments; seven were significant in two environments and eight could only be detected in a single environment. However, QTLs-by-environment interaction was trait dependent. QTLs for spikelets and grains per panicle were common across environments while traits like heading date and plant height were more sensitive to environment. Received: 22 February 1996 / Accepted: 10 May 1996     

利用水稻重组自交系群体对一些重要农艺性状进行基因定位和互作分析

对水稻（OryzasativaL．）籼（indica）、粳（Japonica）交（窄叶青8号×京系17）通过一粒传获得了一个自交9代的重组自交系（RIL）群体,该群体含有107个稳定纯合的株系,通过构建分子连锁图谱,对水稻播抽期、株高、每穗颖花数、每穗实粒数、200粒重和结实率进行数量性状基因座位（QTL）分析,定位了影响播抽期的2个QTLs、株高的2个QTLs、每穗颖花数的2个QTLs和每穗实粒数的1个QTL。其中,控制播抽期的1个QTL即qHD－8为主效基因,控制株高的1个QTL即qPH－1为主效基因,其余为微效基因。同时分析了影响水稻生产力的数量性状基因座位间的互作。结果表明,影响播抽期、株高、每穗颖花数、每穗实粒数和结实率的互适型互作共有24个,对性状有6．2％～10．9％的贡献率。     

Quantitative trait loci for rice yield-related traits using recombinant inbred lines derived from two diverse cultivars

The thousand-grain weight and spikelets per panicle directly contribute to rice yield. Heading date and plant height also greatly influence the yield. Dissection of genetic bases of yield-related traits would provide tools for yield improvement. In this study, quantitative trait loci (QTL) mapping for spikelets per panicle, thousand-grain weight, heading date and plant height was performed using recombinant inbred lines derived from a cross between two diverse cultivars, Nanyangzhan and Chuan7. In total, 20 QTLs were identified for four traits. They were located to 11 chromosomes except on chromosome 4. Seven and five QTLs were detected for thousand-grain weight and spikelets per panicle, respectively. Four QTLs were identified for both heading date and plant height. About half the QTLs were commonly detected in both years, 2006 and 2007. Six QTLs are being reported for the first time. Two QTL clusters were identified in regions flanked by RM22065 and RM5720 on chromosome 7 and by RM502 and RM264 on chromosome 8, respectively. The parent, Nanyangzhan with heavy thousand-grain weight, carried alleles with increased effects on all seven thousand-grain weight QTL, which explained why there was no transgressive segregation for thousand-grain weight in the population. In contrast, Chuan7 with more spikelets per panicle carried positive alleles at all five spikelets per panicle QTL except qspp5. Further work on distinction between pleiotropic QTL and linked QTL is needed in two yield-related QTL clusters.     

RFLP mapping of major and minor genes for important agronomic characters in rice

A molecular genetic map with 233 RFLP markers which covered about 2070 cM of rice genome was constructed based on a doubled haploid (DH) population derived from anther culture of a cross between an indica variety Gui630 and a japonica variety 02428. Quantitative trait loci (QTLs) for agronomic characters such as number of panides, heading date, plant height, number of spikelets, number of grains, fertility and 1 000-grain weight were analyzed using interval mapping approach. 8 major genes and 29 minor genes were identified associating with these traits. The results also indicated that great phenotypic difference between parents was profitable in detection of major genes.     

Comparative study of QTLs for agronomic traits of riceOriza sativa L.) between salt stress and nonstress environment

Genotype-by-environment interactions (GxE) are commonly observed for quantitative traits. In the present study, a doubled haploid (DH) population and its genetic linkage map were used to comparatively study QTLs in salt stress and nonstress environments. A total of 24 QTLs were detected for five agronomic traits, which were distributed on all the chromosomes except 9 and 11. Under the salt stress, nine (37.5%) QTLs were detected, including one for 1 000-grain weight (GW), two for heading date (HD), one for plant height (PH), two for grains per panicle (GPP), and three for effective tillers (ET), while in the nonstress environment, 17 QTLs (70.8%) were detected, including five for GW, six for HD, three for PH, two for GPP, and one for ET. Two QTLs (8.3%) were consistently detected in both environments. One was identified on chromosome 4 for HD and the other on Chr.6 for GPP. Furthermore, three regions carrying multiple QTLs were identified on chromosomes 1, 4 and 8 respectively. For example, on chromosome 8, three QTLs for HD, GW and PH, respectively were identified between RG885-GA408 in nonstress environment, but not in the stress environment. The comparative study of QTLs detected in extremely different (salt stress and nonstress) environments revealed that there existed several QTLs for important agronomic traits on chromosome 8 which were affected significantly by salt stress.     

QTL analysis of agronomic traits in barley based on the doubled haploid progeny of two elite North American varieties representing different germplasm groups

A better understanding of the genetics of complex traits, such as yield, may be achieved by using molecular tools. This study was conducted to estimate the number, genome location, effect and allele phase of QTLs determining agronomic traits in the two North American malting barley (Hordeum vulgare L.) quality variety standards. Using a doubled haploid population of 140 lines from the cross of two-rowed Harrington×six-rowed Morex, agronomic phenotypic data sets from nine environments, and a 107-marker linkage map, we performed QTL analyses using simple interval mapping and simplified composite interval mapping procedures. Thirty-five QTLs were associated, either across environments or in individual environments, with five grain and agronomic traits (yield, kernel plumpness, test weight, heading date, and plant height). Significant QTL×environment interaction was detected for all traits. These interactions resulted from both changes in the magnitude of response and changes in the sign of the allelic effect. QTLs for multiple traits were coincident. The vrs1 locus on chromosome 2 (2H), which determines inflorescence row type, was coincident with the largest-effect QTL determining four traits (yield, kernel plumpness, test weight, and plant height). QTL analyses were also conducted separately for each sub-population (six-rowed and two-rowed). Seven new QTLs were detected in the sub-populations. Positive transgressive segregants were found for all traits, but they were more prevalent in the six-rowed sub-population.QTL analysis should be useful for identifying candidate genes and introgressing favorable alleles between germplasm groups. Received: 18 August 2000 / Accepted: 15 December 2000     

Comparative study of QTLs for agronomic traits of riceOriza sativa L.) between salt stress and nonstress environment

Genotype-by-environment interactions (GxE) are commonly observed for quantitative traits. In the present study, a doubled haploid (DH) population and its genetic linkage map were used to comparatively study QTLs in salt stress and nonstress environments. A total of 24 QTLs were detected for five agronomic traits, which were distributed on all the chromosomes except 9 and 11. Under the salt stress, nine (37.5%) QTLs were detected, including one for 1 000-grain weight (GW), two for heading date (HD), one for plant height (PH), two for grains per panicle (GPP), and three for effective tillers (ET), while in the nonstress environment, 17 QTLs (70.8%) were detected, including five for GW, six for HD, three for PH, two for GPP, and one for ET. Two QTLs (8.3%) were consistently detected in both environments. One was identified on chromosome 4 for HD and the other on Chr.6 for GPP. Furthermore, three regions carrying multiple QTLs were identified on chromosomes 1, 4 and 8 respectively. For example, on chromosome 8, three QTLs for HD, GW and PH, respectively were identified between RG885-GA408 in nonstress environment, but not in the stress environment. The comparative study of QTLs detected in extremely different (salt stress and nonstress) environments revealed that there existed several QTLs for important agronomic traits on chromosome 8 which were affected significantly by salt stress.     

Comparative study of QTLs for agronomic traits of rice (Oriza sativa L.) between salt stress and nonstress environment

Genotype-by-environment interactions (GxE) are commonly observed for quantitative traits. In the present study, a doubled haploid (DH) population and its genetic linkage map were used to comparatively study QTLs in salt stress and nonstress environments. A total of 24 QTLs were detected for five agronomic traits, which were distributed on all the chromosomes except 9 and 11. Under the salt stress, nine (37.5%) QTLs were detected, including one for 1 000-grain weight (GW), two for heading date (HD), one for plant height (PH), two for grains per panicle (GPP), and three for effective tillers (ET), while in the nonstress environment, 17 QTLs (70.8%) were detected, including five for GW, six for HD, three for PH, two for GPP, and one for ET. Two QTLs (8.3%) were consistently detected in both environments. One was identified on chromosome 4 for HD and the other on Chr.6 for GPP. Furthermore, three regions carrying multiple QTLs were identified on chromosomes 1, 4 and 8 respectively. For example, on chromosome     

水稻穗部性状的QTL与环境互作分析

分别在两年收集珍汕97／明恢63的重组自交系群体的表现数据,运用混合线性模型的QTL定位方法,联合分析穗部5个性状的QTLs7及QTL与环境互作关系。每穗颖花数、每穗实粒数、结实率、穗长和穗着密度分别检测到10、3、6、8和7个QTLs分别解释各性状变异的29．13％、19．2％、29．46％、26．39％和35．76％。对于同一性状,高值亲本和低值亲本中均存在增效和减效QTL。相关性状QTL的位置表现相同或相似,高值亲本和低值亲本中均存在增效和减效QTL。相关性状QTL的位置表现相同或相似,成族分布。1个穗长QTL,2个每穗颖花数QTL3,3个结实率QTLs表现与环境显著互作,QTL与环境互作效应的贡献率比相应的QTL贡献率略大。遗传力稍高的每穗实粒数和穗着粒密度的DQTL与环境不互作。     

Identification of quantitative trait loci and epistatic interactions for plant height and heading date in rice

Appropriate heading date and plant height are prerequisites for attaining the desired yield level in rice breeding programs. In this study, we analyzed the genetic bases of heading date and plant height at both single- locus and two-locus levels, using a population of 240 F_2:3 families derived from a cross between two elite rice lines. Measurements for the traits were obtained over 2 years in replicated field trials. A linkage map was constructed with 151 polymorphic marker loci, based on which interval mapping was performed using Mapmaker/QTL. The analyses detected six QTLs for plant height and six QTLs for heading date; collectively the QTLs for heading date accounted for a much greater amount of phenotypic variation than did the QTLs for plant height. Two-way analyses of variance, with all possible two-locus combinations, detected large numbers (from 101 to 257) of significant digenic interactions in the 2 years for both traits involving markers distributed in the entire genome; 22 and 39 were simultaneously detected in both years for plant height and heading date, respectively. Each of the interactions individually accounted for only a very small portion of the phenotypic variation. The majority of the significant interactions involved marker loci that did not detect significant effects by single-locus analyses, and many of the QTLs detected by single-locus analyses were involved in epistatic interactions. The results clearly demonstrated the importance of epistatic interactions in the genetic bases of heading date and plant height. Received: 5 May 2001 / Accepted: 3 August 2001     

水稻F2不育和抽穗期QTL分析

对台中65（粳稻）/Bhadua（籼稻）杂交F2代群体构建了RFLP连锁图谱,含94个分布较为均匀的标记。对F2小穗不育性状进行单点分析和区间分析的结果基本一致：有两个F2小穗不育QTL座位分别位于染色体1的XNpb113～XNpb346之间和染色体8的G187～XNpb397之间,而且该两个QTL均为新检测出的座位;检测出5个抽穗期TQL,其中3个座位在单点分析和区间分析中的结果一致,分别位于染色体1的XNpb113～XNpb346,染色体4的C891～C335,染色体的8的C166～C1121,另外,染色体6的XNpb27为单点分析结果,染色体10的R716～C405为区间分析结果。由于染色体1上的F2不育QTL和抽穗期QTL重叠,该QTL座位是由于遗传效应所至还是由于环境因素（迟抽穗）所至有待构建近等基因系进一步研究。;位于染色体1和10上的抽穗期QTL座位为新检测的座位。对新检测的F2不育和抽穗期QTL座位正在建立相应的近等基因系以精确定位和克隆上述基因。     

普通野生稻中增产相关QTL的发掘

普通野生稻(Oryza Rufipogon)是重要的遗传资源,发掘其优良等位基因将对水稻遗传改良产生重要影响。文章从以珍汕97为轮回亲本,普通野生稻为供体的BC2F1群体中选择一个与珍汕97表型明显不同的单株BC2F1-15,经过连续自交获得回交重组自交系BC2F5群体。均匀分布于12条染色体的126个多态性SSR(Simplesequence repeats)标记基因型分析,发现BC2F1-15单株在30%的标记位点为杂合基因型;利用该群体共检测到4个抽穗期、3个株高、4个每穗颖花数、2个千粒重和1个单株产量QTL。在第7染色体RM481-RM2区间,检测到抽穗期、每穗颖花数和产量QTL,野生稻等位基因表现增效作用;其他3个每穗颖花数QTL位点,野生稻等位基因也均具有增效作用。结果表明野生稻携带有增产相关的等位基因,这些有利等位基因无疑是水稻遗传改良可资利用的新资源。     

Identification of Quantitative Trait Loci for Bacterial Blight Resistance Derived from Oryza meyeriana and Agronomic Traits in Recombinant Inbred Lines of Oryza sativa

Bacterial blight (BB) is one of the major diseases that affect rice productivity. In previous studies, BB resistance was transferred to cultivated rice Oryza sativa from wild rice Oryza meyeriana using asymmetric somatic hybridization. One of the resistant hybrid progenies (Y73) has also been shown to possess novel resistance gene(s) different from any of those previously associated with BB resistance. We have mapped quantitative trait loci (QTLs) for BB resistance in a recombinant inbred line (RIL) population derived from a cross between Y73 and a BB‐susceptible cv. IR24. Five QTLs were detected where Y73 alleles contributed to increased BB resistance. Three minor QTLs were identified on chromosomes 3, 10 and 11, and two major QTLs on chromosomes 1 and 5, respectively. QTL on chromosome 5, designated qBBR5, had the strongest effect on BB resistance, explaining approximately 37% of the phenotypic variance. Using the same RIL population, we also mapped QTLs for agronomic traits including plant height (PH), heading date (HD), plant yield (PYD) and PYD component traits. A total of 21 QTLs were identified, of which four were detected for PH, six for HD, three for panicle number per plant (PNPP), one for spikelets per panicle (SPP), six for 1000‐grain weight (TGW) and one for PYD. qPH1 (a QTL for PH) was found in the same interval as qBBR1 for BB resistance, and qHD11 for HD and qBBR11 for BB resistance also shared a similar interval. Additionally, BB resistance was significantly correlated with PH or HD in the RIL population. This suggests that the resistance genes may have pleiotropic effects on, or close linkage to, genes controlling PH or HD. These results will help deduce the resistance mechanisms of the novel resistance gene(s) and provide the basis for cloning them and using them in marker‐assisted breeding.     

QTLs for agronomic traits in the Mediterranean environment identified in recombinant inbred lines of the cross 'Arta' × <Emphasis Type="Italic">H. spontaneum</Emphasis> 41-1

A genetic linkage map has been developed for recombinant inbred lines (RILs) of the cross 'Arta' × Hordeum spontaneum 41-1. One hundred and ninety four RILs, randomly chosen from a population of 494 RILs, were mapped with 189 markers including one morphological trait (btr = brittle rachis locus). The linkage map extended to 890 cM. Agronomic traits such as grain yield, biological yield, days to heading, plant height, cold tolerance and others were evaluated at the ICARDA research stations Tel Hadya and Breda during the years 1996–97 and 1997–98. QTLs for agronomic traits related to drought resistance were localized. For the most-important character 'plant height under drought stress', QTLs on 2H, 3H and 7H were detected. The 'plant height' QTLs, specially the one on 3H, showed pleiotropic effects on traits such as days to heading, grain yield and biological yield. QTLs were also identified for other traits associated with adaptation to the Mediterranean environment such as cold tolerance, days to heading and tiller number. The identification of QTLs for agronomic traits is a first step to analyze and to dissect complex characters such as adaptation to drought tolerance.Comunicated by R. Hagemann     

测序水稻品种SSR遗传连锁图谱的构建及其农艺性状基因座分析

利用测序水稻品种"Nipponbare（粳）/广陆矮4号（籼）"杂交F2群体90个单株为作图群体,构建含148个SSR标记的水稻遗传连锁图谱,覆盖基因组全长1737.81cM,标记间平均距11.90cM。利用该图谱及Excel2000和Mapmaker/QTL1.1b软件对分蘖数、穗数、穗长、主穗长、株高、剑叶长等六个农艺性状间的相互关系和基因位点进行分析,结果在LOD>2.2和P<0.005的条件下共检测到28个QTLs,它们分布在水稻所有染色体上,单个QTL对性状的分子贡献率11.1%-42.9%,其中大于20%有10个,并对选用已测序材料为亲本构建图谱来探讨水稻农艺性状的分子基础及其育种意义进行了讨论。     

云南元江普通野生稻穗颈维管束和穗部性状的QTL分析

以云南元江普通野生稻为供体亲本,籼稻品种特青为轮回亲本构建高代回交群体,用SSR标记构建连锁图谱,在第1、2、3、4、7和10染色体上定位到7个控制穗颈大维管束数的QTL,在第1、2、3、4和8染色体上定位到5个控制穗颈小维管束数的QTL,在第11和12以外的10条染色体上,共定位到15个控制穗一、二次枝梗数和穗颖花数QTL。来自野生稻的等位基因大多表现负效,能显著减少群体的穗颈维管束数、枝梗数和颖花数,说明从野生稻演化成栽培稻的过程中,可能淘汰了一些对产量不利的QTL,保留了有利的QTL。相当一部分控制穗颈维管束数、枝梗数及颖花数的QTL在染色体上成簇分布或紧密连锁,且加性效应的方向一致,从理论上解释了这些性状表型显著相关的遗传基础,同时也说明在人工选择或自然选择下,这些性状可能存在平行进化或协同进化的关系。     

QTLs for heading date and plant height under multiple environments in rice

Both heading date and plant height are important traits related to grain yield in rice. In this study, a recombinant inbred lines (RILs) population was used to map quantitative trait loci (QTLs) for both traits under 3 long-day (LD) environments and 1 short-day (SD) environment. A total of eight QTLs for heading date and three QTLs for plant height were detected by composite interval mapping under LD conditions. Additional one QTL for heading date and three QTLs for plant height were identified by Two-QTL model under LD conditions. Among them, major QTLs qHd7.1, qHd7.2 and qHd8 for heading date, and qPh1 and qPh7.1 for plant height were commonly detected. qHd7.1 and qHd7.2 were mapped to small regions of less than 1 cM. Genome position comparison of previously cloned genes with QTLs detected in this study revealed that qHd5 and qPh3.1 were two novel QTLs. The alleles of these QTLs increasing trait values were dispersed in both parents, which well explained the transgressive segregation observed in this population. In addition, the interaction between qHd7.1 and qHd8 was detected under all LD conditions. Multiple-QTL model analysis revealed that all QTLs and their interactions explained over 80% of heading date variation and 50% of plant height variation. Two heading date QTLs were detected under SD condition. Of them, qHd10 were commonly identified under LD condition. The difference in QTL detection between LD and SD conditions indicated most heading date QTLs are sensitive to photoperiod. These findings will benefit breeding design for heading date and plant height in rice.     

Mapping quantitative trait loci for yield,yield components and morphological traits in an advanced backcross population between <Emphasis Type="Italic">Oryza rufipogon</Emphasis> and the <Emphasis Type="Italic">Oryza sativa</Emphasis> cultivar Jefferson

An advanced backcross population between an accession of Oryza rufipogon (IRGC 105491) and the U.S. cultivar Jefferson (Oryza sativa ssp. japonica) was developed to identify quantitative trait loci (QTLs) for yield, yield components and morphological traits. The genetic linkage map generated for this population consisted of 153 SSR and RFLP markers with an average interval size of 10.3 cM. Thirteen traits were examined, nine of which were measured in multiple environments. Seventy-six QTLs above an experiment-wise significance threshold of P<0.01 (corresponding to an interval mapping LOD>3.6 or a composite interval mapping LOD>3.9) were identified. For the traits measured in multiple environments, 47% of the QTLs were detected in at least two environments. The O. rufipogon allele was favorable for 53% of the yield and yield component QTLs, including loci for yield, grains per panicle, panicle length, and grain weight. Morphological traits related to the domestication process and/or weedy characteristics, including plant height, shattering, tiller type and awns, were found clustered on chromosomes 1 and 4. Comparisons to previous studies involving wild x cultivated crosses revealed O. rufipogon alleles with stable effects in multiple genetic backgrounds and environments, several of which have not been detected in studies between Oryza sativa cultivars, indicating potentially novel alleles from O. rufipogon. Some O. rufipogon-derived QTLs, however, were in similar regions as previously reported QTLs from Oryza sativa cultivars, providing evidence for conservation of these QTLs across the Oryza genus. In addition, several QTLs for grain weight, plant height, and flowering time were localized to putative homeologous regions in maize where QTLs for these traits have been previously reported, supporting the hypothesis of functional conservation of QTLs across the grasses.     

Quantitative Trait Loci for Panicle Layer Uniformity Identified in Doubled Haploid Lines of Rice in Two Environments

Uniformity of stem height in rice directly affects crop yield potential and appearance, and has become a vital index for rice improvement. In the present study, a doubled haploid (DH) population, derived from a cross between japonica rice Chunjiang 06 and indica rice TN1 was used to analyze the quantitative trait locus (QTL) for three related traits of panicle-layer-uniformity; that is, the tallest panicle height, the lowest panicle height and panicle layer disuniformity in two locations: Hangzhou (HZ) and Hainan (HN). A total of 16 QTLs for three traits distributed on eight chromosomes were detected in two different environments. Two QTLs, qTPH -4 and qTPH -8 were co-located with the QTLs for qLPH -4 and qLPH -8, which were only significant in the HZ environment, whereas the qTPH -6 and qLPH -6 located at the same interval were only significant in the HN environment. Two QTLs, qPLD -10-1 and qPLD -10-2, were closely linked to qTPH-10 , and they might have been at the same locus. One QTL, qPLD -3, was detected in both environments, explaining more than 23% of the phenotypic variations. The CJ06 allele of qPLD -3 could increase the panicle layer disuniformity by 9.23 and 4.74 cm in the HZ and HN environments. Except for qPLD -3, almost all other QTLs for the same trait were detected only in one environment, indicating that these three traits were dramatically affected by environmental factors. The results may be useful for elucidation of the molecular mechanism of panicle-layer-uniformity and marker assisted breeding for super-rice.     

Additive and additive × additive interaction make important contributions to spikelets per panicle in rice near isogenic (Oryza sativa L.) lines

Epistasis plays an important role in the genetic basis of rice yield traits. Taking interactions into account in breeding programs will help the development of high-yielding rice varieties. In this study, three sets of near isogenic lines (NILs) targeting three QTLs for spikelets per panicle (SPP), namely qSPP1, qSPP2 and qSPP7, which share the same Zhenshan 97 genetic background, were used to produce an F 2 population in which the three QTLs segregated simultaneously. The genotypes of the individual F 2 plants at the three QTLs were replaced with three markers that are closely linked to the corresponding QTLs. These QTLs were validated in the F 2 and F 3 popula- tions at the single marker level. qSPP7 exhibited major pleiotropic effects on SPP, plant height and heading date. Multifactor analysis of variance was performed for the F 2 population and its progeny. Additive × additive interaction between qSPP2 and qSPP7 had significant effects on SPP in both the F 2 population and its progeny. Both additive and additive × additive interactions could explain about 73% of the total SPP phenotypic variance. The SPP performance of 27 three-locus combinations was ranked and favorable combinations were rec- ommended for rice breeding in different ecosystems.