Mapping quantitative trait loci for salt tolerance at germination and the seedling stage in barley (Hordeum vulgare L.)

Quantitative trait loci (QTLs) controlling salt tolerance at germination and the seedling stage in barley (Hordeum vulgare L.) were identified by interval mapping analysis using marker information from two doubled haploid (DH) populations derived from the crosses, Steptoe/Morex and Harrington/TR306. Interval mapping analysis revealed that the QTLs for salt tolerance at germination in the DH lines of Steptoe/Morex were located on chromosomes 4 (4H), 6(6H), and 7(5H), and in the DH lines of Harrington/TR306 on chromosomes 5(1H) and 7(5H). In both DH populations, the most effective QTLs were found at different loci on chromosome 7(5H). Genetic linkage between salt tolerance at germination and abscisic acid (ABA) response was found from QTL mapping. The QTLs for the most effective ABA response at germination were located very close to those for salt tolerance on chromosome 7 (5H) in both crosses. The QTLs for salt tolerance at the seedling stage were located on chromosomes 2(2H), 5(1H), 6(6H), and 7(5H) in the DH lines of Steptoe/Morex, and on chromosome 7(5H) in the DH lines of Harrington/TR 306. Their positions were different from those of QTLs controlling salt tolerance at germination, indicating that salt tolerance at germination and at the seedling stage were controlled by different loci. This revised version was published online in July 2006 with corrections to the Cover Date.     

Mapping QTLs related to salinity tolerance of rice at the young seedling stage

Using a population of recombinant inbred lines of the 164 genotypes derived from a cross between ‘Milyang 23’ (indica) and ‘Gihobyeo’ (japonica) in rice (Oryza sativa L.), salt tolerance was evaluated at a young seedling stage in concentrations of 0.5% and 0.7% NaCl. Mapping quantitative trait loci (QTLs) related to salt tolerance was carried out by interval mapping using Qgene 3.0. Two QTLs (qST1 and qST3) conferring salt tolerance at young seedling stage were mapped on chromosome 1 and 3, respectively, and explained 35.5–36.9% of the total phenotypic variation in 0.5% and 0.7% NaCl. The favourable allele of qST1 was contributed by ‘Gihobyeo’, and that of qST3 by ‘Milyang 23’. The results obtained in 0.5% and 0.7% NaCl for 2 years were similar in flanked markers and phenotypic variation.     

Inheritance of tolerance to zinc deficiency in barley

Barley (Hordeum vulgare L.) is often grown on alkaline zinc (Zn)‐deficient soils where reductions in yield and grain quality are frequently reported. Currently, the use of Zn‐based fertilizer along with Zn‐deficiency‐tolerant genotypes is considered the most thorough approach for cropping the Zn‐deficient soils; however, developing or breeding genotypes with higher Zn efficiency requires a good understanding of the inheritance of tolerance to Zn deficiency. This study was conducted to determine genetic control of this trait in barley. Two parental cultivars ('Skiff, moderately tolerant; and ‘Forrest’, sensitive), 185 F₂ plants, and 48 F₂‐derived F₃ families from this cross were screened to determine inheritance of tolerance to Zn deficiency using a visual score of deficiency symptoms. The segregation ratios observed indicated that greater tolerance to Zn deficiency in ‘Skiff compared with ‘Forrest’ at the seedling stage is controlled by a single gene with no dominance. The results also indicate that visual scores are useful for genetic analysis of tolerance to Zn deficiency.     

"汕优63"的产量及其构成因子的数量性状基因位点分析

本文选用优良杂交组合珍汕97／明恢63的F2稻蔸群体为材料,对产量及产量构成因子（有效穗、每穗实粒数和千粒重）的表现分布在两年内进行了考查。通过对537个RFLP标记的54对SSR标记的筛选,共找到了151（133个RFLP,18个SSP）个多型性标记,用这些多型性标记构建了该群体的饱和分子标记遗传连锁图,对产量和产量构成因子进行了QTL定位,在两年内共检测到36个QTLs,其中8个QTLs在两年内都被检测到,其余28个QTLs分别在一年内检测到,通过对这些QTLs的定位、效应大小及作用方向的分析,为分析该组合所表现出的强大杂种优势打下了基础。     

QTL analysis of flooding tolerance in soybean at an early vegetative growth stage

Soybean cultivars are sensitive to flooding stress and their seed yields are substantially reduced in response to the stress. This study was conducted to investigate the genetic basis of flooding tolerance at an early vegetative growth stage. Sixty recombinant inbred lines derived from a cross between a relatively tolerant cv. ‘Misuzudaizu’ and a sensitive cv. ‘Moshidou Gong 503’ were grown in pots in a vinyl plastic greenhouse in 2002 and 2003. At the two‐leaf stage, half of the pots were waterlogged by water placed in plastic containers and adjusted to 5 cm above the soil surface. After 3 weeks of treatment, the pots were returned to the greenhouse and grown until maturity. Flooding tolerance was evaluated by dividing the seed weight of the treated plants by that of the control plants. Quantitative trait loci (QTL) analysis using 360 genetic markers revealed three QTLs for flooding tolerance, ft1 to ft3 in 2002. The ft1 (molecular linkage group C2) was reproducible and an additional four QTLs, ft4 to ft7, were found in 2003. The ft1 had a high LOD score in both years (15.41 and 7.57) and accounted for 49.2% and 30.5% of the total variance, respectively. A large QTL for days to flowering was consistently observed across treatments and years at a similar position to ft1. Comparing the relative location with markers, the maturity gene probably corresponds to E1. Late maturity may have conferred a longer growth period for recovery from flooding stress.     

Detection of QTLs for salt tolerance in Asian barley (Hordeum vulgare L.) by association analysis with SNP markers

Two hundred ninety-six Asian barley (Hordeum vulgare L.) accessions were assessed to detect QTLs underlying salt tolerance by association analysis using a 384 single nucleotide polymorphism (SNP) marker system. The experiment was laid out at the seedling stage in a hydroponic solution under control and 250 mM NaCl solution with three replications of four plants each. Salt tolerance was assessed by leaf injury score (LIS) and salt tolerance indices (STIs) of the number of leaves (NL), shoot length (SL), root length (RL), shoot dry weight (SDW) and root dry weight (RDW). LIS was scored from 1 to 5 according to the severity of necrosis and chlorosis observed on leaves. There was a wide variation in salt tolerance among Asian barley accessions. LIS and STI (SDW) were the most suitable traits for screening salt tolerance. Association was estimated between markers and traits to detect QTLs for LIS and STI (SDW). Seven significant QTLs were located on chromosomes 1H (2 QTLs), 2H (2 QTLs), 3H (1 QTL), 4H (1 QTL) and 5H (1 QTL). Five QTLs were associated with LIS and 2 QTLs with STI (SDW). Two QTLs associated with LIS were newly identified on chromosomes 3H and 4H.     

Detection of quantitative trait loci for phosphorus deficiency tolerance at soybean seedling stage

Phosphorus deficiency is a primary constraint to soybean productivity in acid and calcareous soils. Our aim was to map quantitative trait loci (QTL) controlling phosphorus deficiency tolerance using 152 recombinant inbred lines derived from a cross between the P stress tolerant variety Nannong94-156 and the P stress sensitive variety Bogao. Five traits were used as parameters to evaluate phosphorus deficiency tolerance at seedling stage under different phosphorus levels in experiments 2005 and 2006. As a result, thirty-four additive QTLs were detected on nine linkage groups, with corresponding contribution ratios of 6.6–19.3%. There were three clusters of QTL found in genomic regions S506-Satt534 (on linkage group B2-1), Sat_183-Satt274 (on linkage group D1b + W), and Sat_185-Satt012 (on linkage group G). The locus flanked by Sat_183-Satt274 on linkage group D1b + W was coincident with four previously discovered QTLs with phosphorus efficiency. Another interesting locus flanked by Sat_185-Satt012 on linkage group G was detected across years. The identified QTL will be useful to improve the stress resistance of soybean against a complex nutritional disorder caused by phosphorus deficiency. In addition, more QTLs were detected under low phosphorus condition and some QTLs were detected that specifically expressed under different phosphorus levels. These particular QTLs could help provide greater understanding of the genetic basis of phosphorus efficiency in soybean.     

Use of component analysis in QTL mapping of complex crop traits: a case study on yield in barley

Genes contributing to the quantitative variation of a complex crop trait can be numerous. However, using existing approaches, the number of quantitative trait loci (QTL) detected for a trait is limited. Therefore, rather than looking for QTL for a complex trait itself, determining QTL for underlying component traits might give more information. In this study the potential of component analysis in QTL mapping of complex traits was examined using grain yield in spring barley as an example. Grain yield was divided into three components: number of spikes/m², number of kernels/spike, and 1000‐kernel weight. These traits were measured for individuals of a recombinant inbred‐line population in field trials conducted over 2 years. By the use of an approximate multiple QTL model, one to eight QTL were detected for each trait in a year. Some QTL were mapped to similar positions in both years. Almost all QTL for yield were found at the position of or in close proximity to QTL for its component traits. A number of QTL for component traits were not detected when yield itself was subjected to QTL analysis. However, relative to the QTL for yield itself, all component‐trait QTL did not explain the variation in yield better. The results in relation to the potential of using component analysis in studying complex crop traits are discussed.     

Identification of QTLs for BYDV tolerance in bread wheat

We searched for QTLs involved in tolerance to barley yellow dwarf (BYD), a serious viral disease of small grain cereals in two wheat populations, Opata × Synthetic (ITMI)and Frontana × INIA66 (F × I), for which marker data had previously been generated. The populations were evaluated in replicated field trials under artificial inoculation with a BYDV-PAV-Mex isolate and under disease-free conditions. Disease symptoms (yellowing, dwarfism and biomass reduction) were visually recorded and agronomic traits (number of tillers,height, biomass, yield and thousand-kernel weight) were measured on five plants per plot. Phenotypic data on all evaluated traits showed normal distribution with high correlation between visual estimates and measured values. Heritabilities were mostly moderate to high in the 114 lines of the ITMI population, and from low to moderate in the 117 lines of the F × I population. QTL analyses were based on genetic maps containing 443 loci for the ITMI population and 317 loci for the F × I population. Using composite interval mapping, 22 QTLs in the ITMI population and seven in the F × I population were detected, explaining9.8–43.3% of total phenotypic variation (σ² _P)per agronomic trait in the first population, and 4.1–13.7% in the second. Individual QTLs explained less than 15.8%of σ² _P. In the F × I population a minor QTL explaining 7% of σ² _P for yellowing was detected on the short arm of 7D, linked to leaf tip necrosis, a morphological marker for linked genes Bdv1, Yr18 andLr34. A QTL consistently detected for several traits on 2D in the ITMI population and on the short arm of group 6 chromosome(6S) in F × I explained 10–15% of σ² _P. The large number of QTLs having mostly small effects and the continuous distribution of all evaluated traits confirmed the polygenic nature and complexity of BYD tolerance in wheat. This revised version was published online in August 2006 with corrections to the Cover Date.     

A major QTL associated with cold tolerance at seedling stage in rice (Oryza sativa L.)

Chilling injury is one of the most important limiting factors affecting rice production in temperate and high-elevation areas. In this study, 146 microsatellite markers were employed to identify quantitative trait loci (QTL) conferring cold tolerance at seedling stage (CTS) .The mapping population consisted of 193 doubled haploid (DH) lines, which derived from a cross between a cold-tolerant japonica variety (AAV002863) and a cold-sensitive indica cultivar (Zhenshan97B). Tolerance to cold was assessed by the survival percentage of seedlings after cold treatment. In a climate chamber, after treatment at 6°C/10°C for 7 d, the measurement was taken on the sixth day of the recovery stage at room temperature. The phenotypic distribution of the DH population approximately fitted normality with skewness and kurtosis less than 0.3, and the difference among the three repetitions was not significant. Five main effect QTLs were identified with LOD > 4.0 on chromosomes 1, 2, 8 using a composite interval mapping approach. The accumulated contribution of the five QTLs was 62.28%, and a major QTL (LOD = 15.09) was identified on chromosome 2 flanked by RM561 and RM341, which explained 27.42% of the total phenotypic variation. Four significant epistatic interactions were also detected with a total contribution of 20.14%. Liang Chen and Qiaojun Lou had made the equal contribution for the research.     

水稻抽穗期数量性状基因的定位及遗传效应分析

本研究利用特早抽穗粳稻品种石狩白毛和籼稻品种明恢63杂交的F2分离群体共116株,构建了含88个共显性分子标记的连锁图谱,对水稻(Oryza sativA L．)抽穗期进行基因定位。利用2种分析软件MAPMAKER／QTL和QTLMapper进行分析,共检测到3个抽穗期的数量性状基因座(QTLs)。2个软件共同发现第7染色体上RM214与A5106标记区间内存在1个主效QTL Hd7a（Hd7c),来自明恢63的这个位点的等位基因可使抽穗期延迟。其余2个QTLs分别位于第7、9染色体上。同时检测到有5对位点间存在上位性作用,但相对贡献率较小,表明上位性效应也是影响抽穗期的遗传基础。     

不同生态环境下玉米产量性状QTL分析

以玉米（Zea mays L.）自交系黄早四和Mo17为亲本得到的191个F₂单株为作图群体,衍生的184个F_2∶3 家系作为性状评价群体,分析了单株穗数、穗行数、行粒数、百粒重和单株籽粒产量在北京和新疆2个生态环境下的表现和数量性状基因位点的定位结果。QTL检测结果表明,2个环境共检测出47个QTL,分布于除第10染色体以外的9条染色体,其中与单株穗数相关的QTL共10个,可解释的表型变异为5.3%~25.6%;与穗行数相关的QTL共13个,可解释的表型变异为4.5%~23.2%;与行粒数相关的QTL有9个,解释的表型变异为5.4%~13.7%;与百粒重相关的QTL达10个,可解释的表型变异为4.9%~13.3%;与单株籽粒产量相关的QTL有5个,可解释的表型变异为6.1%~35.8 %。大部分产量QTL只在单一环境下被检测到,说明产量相关QTL与环境之间存在明显的互作。表型相关显著的产量性状,它们的QTL容易在相同或相邻标记区间检测到。研究还发现了若干个QTL富集区域,可能是发掘通用QTL的候选位点。     

Identification of QTLs controlling rice drought tolerance at seedling stage in hydroponic culture

‘Drought avoidance’ and ‘drought tolerance’ are two mechanisms by which plants adapt under water stress. These mechanisms are difficult to evaluate separately in field experiments. Using hydroponic culture, we studied the genetic control of drought tolerance in rice (Oryza sativa L.) without the effect of drought avoidance. A backcross inbred population of ‘Akihikari’ (lowland cultivar) × ‘IRAT109’ (upland cultivar) with 106 lines was cultured with (stressed condition) and without (non-stressed condition) polyethylene glycol (PEG) at seedling stage. The relative growth rate (RGR), specific water use (SWU), and water use efficiency (WUE) showed significant genotype × environment interactions with or without PEG, indicating that each line responded differently to water stress. A quantitative trait locus (QTL) analysis revealed that these interactions were QTL specific. A total of three QTLs on chromosomes 2, 4, and 7 were detected for RGR. The QTL on chromosome 7 had a constant effect across environments, while the QTL on chromosome 4 had an effect only under non-stressed condition and that on chromosome 2 only under stressed condition. The stress-specific QTL on chromosome 2 was not co-located with any QTLs for root system depth previously reported from the same mapping population. However, this QTL was co-located with a stress-specific QTL for SWU, suggesting that the control of transpiration was relevant to dry matter production under drought. We concluded that PEG-treated hydroponic culture is very effective for use in genetic analyses of drought tolerance at seedling stage.     

QTLs in barley controlling seedling elongation of deep-sown seeds

In drought areas, in which topsoil moisture is low, barley seeds are generally sown in the subsoil (deep-seeding). In order for the shoots of germinating seeds to emerge from the deep soil cover, the coleoptile and the first internode of the seedlings must elongate as an adaptive response to deep-seeding. Here, we have mapped quantitative trait loci (QTLs) for these adaptive characteristics. Elongation of the coleoptile and first internode was investigated using seeds sown under two soil cover conditions: at a depth of 9 cm beneath a soil mixture; and, at a depth of 12 cm beneath vermiculite. We identified multiple alleles for increased coleoptile and first internode elongation using a doubled haploid population of 150 lines generated from a cross between the barley cultivars Harrington and TR306. Composite interval mapping analyses of the data revealed two moderate and eleven small effect QTLs, with at least one QTL on each chromosome. The QTLs on chromosomes 5H and 7H had moderate effects on coleoptile elongation (18.5–27.6% of PVE: phenotypic variance explained; 2.6–3.2 mm of Add: additive effect) and first internode elongation (PVE: 16.6–19.6%; Add: 3.1–3.2 mm). The small effect QTLs showed PVEs of less than 15% and an Add range of 1.2–3.2 mm for both characters. A marker assisted selection approach, using markers linked to the QTLs for seedling elongation at deep-seeding, may eventually enable development of drought tolerant barley hybrids.     

不同生长环境下水稻结实率数量性状位点的检测

以籼稻密阳23与粳稻吉冷1号配制所获得的F_2:3群体200个家系作为作图群体，在北京、昆明、三亚、公主岭和韩国春川等5个点进行水稻结实率的鉴定，并利用SSR标记对水稻结实率数量性状位点进行检测。结果表明，水稻结实率表型值及其在F₃家系群中的分布以及所检测到的QTL数目因生长环境不同而有较大差异，说明QTL与环境有明显的互作效应。水稻结实率在F₃家系群中呈接近正态或偏态的连续分布，是多个基因所控制的数量性状。共检测到与水稻结实率相关的QTL 14个，分布于第1、2、3、4、6、7、8、10和12染色体上，对表型变异的贡献率为4.9%～15.3%。分别位于第1、2、6和12染色体RM1~RM259、RM263~RM6、RM340~RM30、RM270~RM17区间的qSSR1、qSSR2、qSSR6和qSSR12至少在2种生长环境下均检测到，对表型变异的贡献率分别为4.9%～8.4%、4.8%～7.2%、7.6%～10.7%和7.4%～10.4%。以上多数QTL增效等位基因均来自吉冷1号，基因作用方式主要为部分显性或显性或超显性。     

Inheritance of nitrogen and phosphorus utilization efficiency in spring barley at the vegetative growth stages under high and low nutrition

Inheritance of the nitrogen and phosphorus utilization efficiencies (NUE and PUE, respectively) and that of the tolerance (T) to limited NP nutrition was investigated in spring barley crosses at the vegetative growth stages. Plants were grown in sand-vermiculite cultures under high and low NP nutrition. In a diallel set (Ps and F₂s), both general combining ability (GCA) and specific combining ability (SCA) effects were significant for the variation in NUE and PUE, while the variation in T was mainly associated with GCA effects. The contribution of nonadditive genes for the utilization efficiencies was found to increase under nutrient shortages. Overdominance of genes was detected. The characters exhibited low heritabilities (0.10-0.42). Generation means analysis in two cross-combinations revealed significant effects of epistatic gene interactions. It was assumed that the involvement of both the dominance effects and epistatic interactions would not facilitate selection efforts to improve the characters in spring barley at its vegetative growth. Such selection should be performed among families of later generations. The genotype-nutrition interactions observed suggest that selection under diverse nutrition rates would be necessary for the more precise evaluation of barley efficiency under less favourable soil fertility.     

Identification and mapping of a QTL for rachis internode length associated with cleistogamy in barley

General knowledge of the closed flowering trait, or cleistogamy, of barley is still limited. The relationship between cleistogamy and spike morphology characters was studied and linkage of cleistogamy genes with a highly significant quantitative trait locus (QTL) for rachis internode length on the long arm of chromosome 2H was detected. The mapping populations consisted of 129 doubled haploid lines of ‘Mikamo Golden’ × ‘Harrington’ and 150 F₂ plants of ‘Misato Golden’ × ‘Satsuki Nijo’. The phenotypic variance explained by this QTL accounted for 77.5% and 82.6% of the variance in rachis internode lengt, respectively, in these two populations. The peaks of the QTL coincided with the positions of the cleistogamy gene loci.     

利用重组自交系群体对水稻产量相关性状的QTL分析

利用水稻(Oryza sativa L．)珍汕97和SLG组合的重组自交系群体和177个微卫星标记,采用复合区间作图法对产量相关性状进行了数量性状基因(QTL)定位,共得到38个QTLs。单个QTL解释这些性状6．6％～32．9％的性状变异,各性状QTL的累积表型贡献率达21．3％～79．5％。大多数性状之间具有显著的表型相关性,相关性较高的性状之间常具有较多共同或紧密连锁的QTL。