QTL detection of seven spike-related traits and their genetic correlations in wheat using two related RIL populations

Spike-related traits contribute greatly to grain yield in wheat. To localize wheat chromosomes for factors affecting the seven spike-related traits??i.e., the spike length (SL), the basal sterile spikelet number (BSSN), the top sterile spikelet number (TSSN), the sterile spikelet number in total (SSN), the spikelet number per spike (SPN), the fertile spikelet number (FSN) and the spike density (SD)??two F_8:9 recombinant inbred line (RIL) populations were generated. They were derived from crosses between Weimai 8 and Jimai 20 (WJ) and between Weimai 8 and Yannong 19 (WY), comprising 485 and 229 lines, respectively. Combining the two new linkage maps and the phenotypic data collected from the four environments, we conducted quantitative trait locus (QTL) detection for the seven spike-related traits and evaluated their genetic correlations. Up to 190 putative additive QTL for the seven spike-related traits were detected in WJ and WY, distributing across all the 21 wheat chromosomes. Of these, at least nine pairwise QTL were common to the two populations. In addition, 38 QTL showed significance in at least two of the four different environments, and 18 of these were major stable QTL. Thus, they will be of great value for marker assisted selection (MAS) in breeding programs. Though co-located QTL were universal, every trait owned its unique QTL and even two closely related traits were not excluded. The two related populations with a large/moderate population size made the results authentic and accurate. This study will enhance the understanding of the genetic basis of spike-related traits.     

利用高密度SNP 遗传图谱定位小麦穗部性状基因

小麦穗部性状之间相关性密切, 其中穗粒数和千粒重是重要的产量构成要素, 挖掘与穗部性状相关联的基因位点对分子标记辅助育种及解释基因效应具有重要意义。本研究以RIL群体(山农01-35×藁城9411) 173个F_8:9株系为材料, 利用90 k小麦SNP基因芯片、DArT芯片技术及传统的分子标记技术构建的高密度遗传图谱, 在5个环境下进行穗部相关性状QTL定位。检测到位于1B、4B、5B、6A染色体上7个控制千粒重的加性QTL, 解释表型变异率6.00%~36.30%, 加性效应均来自大粒母本山农01-35; 检测到8个控制穗长的加性QTL, 解释表型变异率14.34%~25.44%; 3个控制穗粒数的加性QTL; 5个控制可育小穗数的加性QTL; 3个控制不育小穗数的加性QTL, 贡献率为8.70%~37.70%; 4个控制总小穗数的加性QTL; 6个控制小穗密度的加性QTL。通过基因型与环境互作分析, 检测到32个加性QTL, 解释表型变异率0.05%~1.05%。在4B染色体区段EX_C101685–RAC875_C27536检测到控制粒重、穗长、穗粒数、可育小穗数、不育小穗数、总小穗数的一因多效QTL,其贡献率为5.40%~37.70%, 该位点在多个环境中被检测到, 是稳定主效QTL。在6A染色体wPt-0959-TaGw2-CAPS区间上检测到控制粒重、总小穗数的QTL。研究结果为穗部性状的分子标记开发、基因精细定位和功能基因克隆奠定了基础。     

QTL analysis for wheat falling number in a recombinant inbred line population segregated with 1BL/1RS translocation in a rainfed agricultural area of China

Falling number (FN) is an inner quality trait in wheat (Triticum aestivum L.) ultimately determining the end use of wheat kernels. In this 3-year study, 171 recombinant inbred lines derived from Chuannong17 (a 1BL/1RS tranlocation parent) × Mianyang11 were planted in the Sichuan Basin, a rainfed agricultural area in southwestern China. In this climate, we found that FN had significant differences between 1BL/1RS translocation lines and non-1BL/1RS translocation lines in two of the 3 years and the heavy fluctuation of rainfall and temperature resulted in decreasing FN in grain filling period. We used 191 simple sequence repeats markers to construct a genetic linkage map and then detected 11 additive effect FN quantitative trait loci (QTL) on chromosomes 2B, 3D, 4A, 4D, 6B and 7D, explaining 5.48–31.91% of the phenotypic variance. The FN QTL on chromosomes 4A, 4D and 6B were major or stable and detected at least in 2 years, whereas the Qfn.sicau-3D.1 in 2015 year explained the maximum phenotypic variation (about 31.91%). Furthermore, FN QTLs additive and epistatic effects as well as their interactions with environment were estimated by a mixed linear model approach. We found that the additive effect QTLs had no significant additive × environment interaction, while the paired QTLs had significant additive × additive epistatic effects however none had a significant additive × additive epistasis × environment interaction effect, excluding the paired QTLs Qfn.sicau-3B/Qfn.sicau-5B.     

燕大1817/北农6号重组自交系群体穗部性状的QTL定位

小麦穗部性状与单株产量密切相关。本研究以小麦骨干亲本燕大1817与优良品系北农6号衍生的269个重组自交系为材料,通过在北京和河北石家庄的2年田间试验数据,利用本实验室已构建的高密度SNP和SSR遗传连锁图谱进行穗长、穗粒数和穗粒重QTL定位。采用完备复合区间作图法共检测到29个穗部性状加性效应QTL,其中10个穗长QTL分布于1B、2D、3A、3B、4A、5A、5B、6A和7D染色体上,解释的表型变异率为2.96%~9.63%,QSl.cau-4A.2在所有5个环境中均能被检测到,解释的表型变异为5.89%~9.62%,另有7个QTL能在2个或2个以上环境中被检测到;8个穗粒数相关QTL分布于1A、3A、3D、4A和5B染色体上,解释的表型变异为4.06%~11.17%,为单个环境QTL。11个与穗粒重相关QTL分布于1A、1B、2A、2D、3A、4D、5A、5B和6B染色体上,解释的表型变异为2.79%~16.12%,其中QGws.cau-1B、QGws.cau-3A和QGws.cau-6B.2在2个或者2个以上环境中能被检测到。另外,鉴定出6个分布于1A、2D、3A、4A和5B染色体上的QTL富集区段。     

Analysis of contributors to grain yield in wheat at the individual quantitative trait locus level

In wheat, strong genetic correlations have been found between grain yield (GY) and tiller number per plant (TN), fertile spikelet number per spike (FSN), kernel number per spike (KN) and thousand‐kernel weight (TKW). To investigate their genetic relationships at the individual quantitative trait locus (QTL) level, we performed both normal and multivariate conditional QTL analysis based on two recombinant inbred lines (RILs) populations. A total of 79 and 48 normal QTLs were identified in the International Triticeae Mapping Initiative (ITMI)/SHW‐L1 × Chuanmai 32 (SC) populations, respectively, as well as 55 and 35 conditional QTLs. Thirty‐two QTL clusters in the ITMI population and 18 QTL clusters in the SC population explained 0.9%–46.2% of phenotypic variance for two to eight traits. A comparison between the normal and conditional QTL mapping analyses indicated that FSN made the smallest contribution to GY among the four GY components that were considered at the QTL level. The effects of TN, KN and TKW on GY were stronger at the QTL level.     

小麦籽粒产量及穗部相关性状的QTL定位

由小麦品种花培3号和豫麦57杂交获得DH群体168个株系,种植于3个环境中,利用305个SSR标记对籽粒产量和穗部相关性状(穗长、穗粒数、总小穗数、可育小穗数、小穗着生密度、千粒重和粒径)进行了QTL定位。利用基于混合线性模型的QTLNetwork 2.0软件,共检测到27个加性效应和13对上位效应位点,其中 8个加性效应位点具有环境互作效应。相关性高的性状间有一些共同的QTL位点,表现出一因多效或紧密连锁效应。5D染色体区段Xwmc215–Xgdm63,检测到控制籽粒产量、穗粒数、总小穗数、可育小穗数和小穗着生密度5个性状的QTL位点,各位点的遗传贡献率较大且遗传效应方向相同,增效等位基因均来源于豫麦57,适用于分子标记辅助育种和聚合育种。控制千粒重与穗粒数的QTL位于染色体不同区段,有利于实现穗粒数与粒重的遗传重组。     

QTL mapping of yield-related traits in the wheat germplasm 3228

The new wheat germplasm 3228, a putative derivative of tetraploid Agropyron cristatum Z559 and the common wheat Fukuhokomugi, has superior features in yield-related traits, particularly in spike morphological traits, such as large spike and superior grain number. To identify favorable alleles of these traits in 3228, 237 F_2:3 families were developed from the cross 3228/Jing 4839. A genetic map was constructed using 179 polymorphic SSR and EST-SSR markers. A total of 76 QTL controlling spike number per plant (SNP), spike length (SL), spikelet number per spike (SNS), floret number per spikelet (FNS), grain number per spike (GNS) and thousand-grain weight (TGW) were detected on 16 chromosomes. Each QTL explained 1.24–27.01% of the phenotypic variation, and 9 QTL (28.95%) were detected in two or all environments. Additive effects of 45 QTL were positive with 3228 alleles increasing the QTL effects, 31 QTL had negative effects indicating positive contributions from Jing 4839. Three important clusters involving all traits were located on chromosomes 5A, 6A and 4B, and several co-located QTL were also found. Most of the QTL detected on the three chromosome regions could contribute to the use of 3228 in breeding for grain yield improvement.     

QTL for yield, yield components and canopy temperature depression in wheat under late sown field conditions

A wheat (Triticum aestivum L.) recombinant inbred line (RIL) population was used to identify quantitative trait loci (QTL) associated with yield, yield components, and canopy temperature depression (CTD) under field conditions. The RIL population, consisting of 118 lines derived from a cross between the stress tolerant cultivar ‘Halberd’ and heat stress sensitive cultivar ‘Karl92’, was grown under optimal and late sown conditions to impose heat stress. Yield and yield components including biomass, spikes m^?2, thousand kernel weight, kernel weight and kernel number per spike, as well as single kernel characteristics were determined. In addition, CTD was measured during both moderate (32–33 °C) and extreme heat stress (36–37 °C) during grain-filling. Yield traits showed moderate to high heritability across environments with a large percentage of the variance explained by genetic effects. Composite interval mapping detected 25 stable QTL for the 15 traits measured, with the amount of phenotypic variation explained by individual QTL ranging from 3.5 to 27.1 %. Two QTL for both yield and CTD were co-localized on chromosomes 3BL and 5DL and were independent of phenological QTL. At both loci, the allele from Halberd was associated with both higher yield and a cooler crop canopy. The QTL on 3BL was also pleiotropic for biomass, spikes m^?2, and heat susceptibility index. This region as well as other QTL identified in this study may serve as potential targets for fine mapping and marker assisted selection for improving yield potential and stress adaptation of wheat.     

新疆冬小麦品种农艺及产量性状遗传多样性分析

Grain yield is one of the most important goals in wheat breeding, and agronomic or yield-related traits can directly reflect the characteristics of varieties. In order to determine the evolution of genetic diversity in agronomic traits of Xinjiang winter wheat varieties and their adaptabilities to different ecological environments, 134 winter wheat landraces and 54 moderns bred varieties from Xinjiang were selected for agronomic trait investigation. They were planted in three different ecological environments (Urumqi and Yining in Xinjiang province, and Tai’an in Shandong province) for two consecutive growth seasons, and nine agronomic and yield-related traits were surveyed and analyzed. The estimated broad sense heritability of nine agronomic and yield traits was in descending order: plant height > grain width > grain length/width ratio > spike length > spikelet number > thousand- kernel weight > grain number per spike > grain length > fertile spikelet number. Correlation analyses of nine agronomic and yield traits showed that these traits were correlated with each other. It was found that the plant height, spike length and grain length/width ratio of landraces were higher than that in modern bred varieties, but the grain number per spike, thousand kernel weight, grain length and grain width in landraces were less than that in modern bred varieties. However, the correlation coefficient of these nine traits was higher in modern bred varieties than that in landraces. These variations reflected the evolution of Xinjiang winter wheat varieties in agronomic traits in recent years. This study may provide important information for breeders to select the breeding parents in different winter wheat regions.     

Mapping quantitative trait loci influencing panicle‐related traits from Chinese common wild rice (Oryza rufipogon) using introgression lines

Panicle‐related traits are important agronomic traits which directly associated with grain yield. In this study, we investigated quantitative trait loci (QTLs) associated with panicle‐related traits using a set of 265 introgression lines (ILs) of common wild rice (Oryza rufipogon Griff.) in the background of Indica high‐yielding cultivar Guichao 2 (O. sativa L.). A total of 39 QTLs associated with panicle‐related traits including panicle length (PL), primary branch number (PBN), secondary branch number (SBN), spikelet number per panicle (SPP) and spikelet density (SD), were detected in the ILs with single‐point analysis. The alleles of 20 QTLs derived from wild rice showed positive effects, and some QTLs, such as, QPl1b for PL, QPbn8 for PBN, QSd4 and QSd11b for SD and QSpp4 for SPP showed larger positive effects, providing good candidates and useful information for marker‐aided improvement of yield potential of rice. Most of the QTLs controlling SPP, SBN and SD were located in cluster or closely linked on chromosomes, and the directions of their additive effects were consistent, which explained the genetic basis of significant correlations between their phenotypic characters.     

Identification of QTL for oil content,seed yield,and flowering time in oilseed rape (<Emphasis Type="Italic">Brassica napus</Emphasis>)

A genetic map was constructed with 353 sequence-related amplified polymorphism and 34 simple sequence repeat markers in oilseed rape (Brassica napus L.). The map consists of 19 linkage groups and covers 1,868 cM of the rapeseed genome. A recombinant doubled haploid (DH) population consisting of 150 lines segregating for oil content and other agronomic traits was produced using standard microspore culture techniques. The DH lines were phenotyped for days to flowering, oil content in the seed, and seed yield at three locations for 3 years, generating nine environments. Data from each of the environments were analyzed separately to detect quantitative trait loci (QTL) for these three phenotypic traits. For oil content, 27 QTL were identified on 14 linkage groups; individual QTL for oil content explained 4.20–30.20% of the total phenotypic variance. For seed yield, 18 QTL on 11 linkage groups were identified, and the phenotypic variance for seed yield, as explained by a single locus, ranged from 4.61 to 24.44%. Twenty-two QTL were also detected for days to flowering, and individual loci explained 4.41–48.28% of the total phenotypic variance.     

Identification of coincident QTL for days to heading,spike length and spikelets per spike in <Emphasis Type="Italic">Lolium perenne</Emphasis> L.

Flowering time is a trait which has a major influence on the quality of forage. In addition, flowering and subsequent seed yields are important traits for seed production by grass breeders. In this study, we have identified quantitative trait loci (QTL) for flowering time and morphological traits of the flowering head in an F₁ mapping population in Lolium perenne L (perennial ryegrass), a number of which have not previously been identified in L. perenne mapping studies. QTL for days to heading (DTH) were mapped in both outdoor and glasshouse experiments, revealing three and five QTL for DTH which explained 53% and 42% of the total phenotypic variation observed, respectively. Two QTL for DTH were detected in both environments, although they had contrasting relative magnitudes in each environment. One QTL for spike length and three QTL for spikelets per spike were also identified explaining, a total of 32 and 33% of the phenotypic variance, respectively. Furthermore, the QTL for spike length and spikelets per spike generally coincided with QTL for days to heading, implying co-ordinate regulation by underlying genes. Of particular interest was a region harbouring overlapping QTL for days to heading, spike length and spikelets per spike on the top of linkage group 4, containing the major QTL for spike length identified in this population.     

Detection of quantitative trait loci (QTLs) for seedling traits and drought tolerance in wheat using three related recombinant inbred line (RIL) populations

In order to detect quantitative trait loci (QTLs) for drought tolerance in wheat during seed germination conditional and unconditional QTL analyses of eight seedling traits were conducted under two water regimes using three related F₉ recombinant inbred line populations with a common female parent. A total of 87 QTLs for the eight seedlings traits and 34 specific QTLs related to drought tolerance were detected. Seventy-one of these QTLs were major QTLs with contributions to phenotypic variance of >10 %. Of the 34 QTLs related to drought tolerance only eight were also detected by unconditional analysis of seedling traits under osmotic stress conditions indicating that most of the QTLs related to drought tolerance could not be detected by unconditional QTL analysis. Therefore, conditional QTL analysis of stress-tolerance traits such as drought tolerance was feasible and effective. Of 11 important QTL clusters located on chromosomes 1BL, 1D, 2A, 2B, 2D, 4A, 6B, and 7B, nine were detected in multiple populations and eight were detected by both unconditional and conditional analyses.     

开花后不同光周期条件下大豆农艺性状和品质性状的QTL分析

以开花期相近的181个大豆重组自交系(RIL)为材料,研究开花后不同光照长度对大豆主要农艺性状的影响,并在利用SSR标记构建大豆遗传图谱的基础上,分别在长日(16 h)和短日(12 h)条件下检测与主要农艺性状及其光周期敏感度(PS)相关的QTL。结果表明,开花后光照处理对大豆农艺性状和品质性状有较大影响,不同性状的光周期敏感度差异明显,株高＞主茎节数＞蛋白质含量、脂肪含量＞百粒重＞单株荚数＞蛋白质和脂肪总量。利用复合区间作图法检测到12个与株高、主茎节数、单株荚数、百粒重、蛋白质和脂肪总量等性状及各性状对开花后光周期处理的敏感度相关的QTL,分别定位于A1、A2、B1、B2、C1、D1a、F、L等8个连锁群上。其中,在短日条件下检测到4个QTL,可解释的遗传变异范围在11.37%~26.63%之间;在长日条件下检测到3个QTL,可解释的遗传变异范围在11.84%~27.85%之间;检测到5个与不同性状光周期敏感度有关的QTL,可解释相对应性状表型变异的范围在6.15%~21.44%之间。针对同一性状,未检测到在长日和短日条件下均起作用的主效QTL, 说明开花后光周期对大豆产量和品质性状相关基因的表达有较大影响。     

普通小麦对仓储害虫玉米象的抗性QTL分析

以普通小麦品种川农16与人工合成小麦衍生品种川麦42为亲本构建了含有127个株系的重组自交系(RIL)群体,将2008和2009年收获的种子用玉米象(Sitophilus zeamais)虫卵进行接种,并于2009年11月开始储藏,次年8月和10月进行虫害调查。利用复合区间作图法(CIM)对RIL群体的抗虫性进行QTL定位分析。利用2008年收获群体检测到位于3B、2D和3D染色体的3个QTL,贡献率依次为10.2%、8.5%和8.3%;利用2009年收获群体检测到位于3D和4B的2个QTL,贡献率分别为8.5%和11.3%。位于3D染色体Xbarc6–Xgwm112标记区间内的QTL在年度间重复检测到,贡献率为8.3%~8.5%,抗性位点来自川农16。     

小麦收获指数与主要农艺性状的相关性探析?

小麦收获指数是指籽粒产量占地上部生物产量的百分率。研究小麦品种主要农艺性状和经济性状间与其产量的相互关系,采用相关性分析与回归方法,对影响收获指数的10个主要农艺性状及因素进行了研究分析。结果表明：供试小麦HI具有较大的改良空间;HI与穗颈长、株高、穗长、小穗数、主茎生物产量有显著负相关,与单穗质量、产量呈正相关,与生物产量没有显著相关性;主成分分析结果显示,3个主成分累积贡献率达 87.427%,表明 3 个主成分已覆盖所有性状的主要信息;通过回归分析,小麦HI与单穗籽粒产量、生物产量、千粒质量有显著回归关系。结论: 通过对10份小麦材料农艺性状的统计分析,得出如下结论。HI受单穗质量、生物产量、千粒重影响较为明显有显著回归关系,与单穗质量、产量和穗粒数有明显正效应,生物产量对其有负效应。小麦HI提高可通过选育生物产量不宜过大,而穗粒数较高、单穗重较大的小麦品种。     

Evaluation of the effects of five QTL regions on Fusarium head blight resistance and agronomic traits in spring wheat (Triticum aestivum L.)

Fusarium head blight (FHB) is an important disease of wheat (Triticum aestivum L.). The aim of this study was to determine the effects of quantitative trait locus (QTL) regions for resistance to FHB and estimate their effects on reducing FHB damage to wheat in Hokkaido, northern Japan. We examined 233 F₁-derived doubled-haploid (DH) lines from a cross between ‘Kukeiharu 14’ and ‘Sumai 3’ to determine their reaction to FHB during two seasons under field conditions. The DH lines were genotyped at five known FHB-resistance QTL regions (on chromosomes 3BS, 5AS, 6BS, 2DL and 4BS) by using SSR markers. ‘Sumai 3’ alleles at the QTLs at 3BS and 5AS effectively reduced FHB damage in the environment of Hokkaido, indicating that these QTLs will be useful for breeding spring wheat cultivars suitable for Hokkaido. Some of the QTL regions influenced agronomic traits: ‘Sumai 3’ alleles at the 4BS and 5AS QTLs significantly increased stem length and spike length, that at the 2DL QTL significantly decreased grain weight, and that at the 6BS QTL significantly delayed heading, indicating pleiotropic or linkage effects between these agronomic traits and FHB resistance.     

黄淮麦区小麦品种(系)产量性状与分子标记的关联分析

为了获得与小麦产量性状关联的分子标记,筛选相关标记的等位变异,以128份黄淮麦区小麦品种(系)为材料,在4个环境下鉴定产量性状,并选用在小麦全基因组21条染色体上的64个SSR标记、27个EST-SSR标记和47个功能标记检测所有材料的基因型。91个SSR和EST-SSR标记共检测到315个等位变异,单个引物检测到2~7个等位变异,平均3.5个;47个功能标记共检测到107个等位变异,单个引物检测到2~5个等位变异,平均2.3个。关联分析表明,49个位点与4个环境的产量性状及其均值显著关联(P≤0.005),其中38个位点在2个或以上环境或均值下被重复验证,16个位点与2个或以上性状相关联。对相对稳定的等位变异作进一步分析,发掘了一批与产量性状相关的优异等位变异,如降低株高的等位变异Ax2*-null和UMN19*-A362,增加穗长的等位变异barc21-A220,增加可育小穗数的等位变异gpw2111-A156,增加总小穗数的等位变异swes65-A120,增加穗数的等位变异VRN-A1*-A1068,增加穗粒数的等位变异cfd5-A215和增加千粒重的等位变异wmc626-A170。研究结果对利用分子标记辅助选择进行小麦产量性状的遗传改良具有一定的指导意义。     

Genetic loci linking improved heat tolerance in wheat (<Emphasis Type="Italic">Triticum aestivum</Emphasis> L.) to lower leaf and spike temperatures under controlled conditions

Heat stress adversely affects wheat production in many regions of the world and is particularly detrimental during reproductive development. The objective of this study was to identify novel quantitative trait loci (QTL) associated with improved heat tolerance in wheat (Triticum aestivum L.) and to confirm previous QTL results. To accomplish this, a recombinant inbred line (RIL) population was subjected to a three-day 38°C daytime heat stress treatment during early grain-filling. At maturity, a heat susceptibility index (HSI) was calculated from the reduction of three main spike yield components; kernel number, total kernel weight, and single kernel weight. The HSI, as well as temperature depression (TD) of the main spike and main flag leaf during heat stress were used as phenotypic measures of heat tolerance. QTL analysis identified 14 QTL for HSI, with individual QTL explaining from 4.5 to 19.3% of the phenotypic variance. Seven of these QTL co-localized for both TD and HSI. At all seven loci, the allele for a cooler flag leaf or spike temperature (up to 0.81°C) was associated with greater heat tolerance, indicated by a lower HSI. In a comparison to previous QTL results in a RIL population utilizing the same source of heat tolerance, seven genome regions for heat tolerance were consistently detected across populations. The genetic effect of combining three of these QTL, located on chromosomes 1B, 5A, and 6D, demonstrate the potential benefit of selecting for multiple heat tolerance alleles simultaneously. The genome regions identified in this study serve as potential target regions for fine-mapping and development of molecular markers for more rapid development of heat tolerant germplasm.     

QTL for flag leaf size and their influence on yield-related traits in wheat

Flag leaf-related traits (FLRTs) are determinant traits affecting plant architecture and yield potential in wheat (Triticum aestivum L.). In this study, three related recombinant inbred line (RIL) populations with a common female parent were developed to identify quantitative trait loci (QTL) for flag leaf width (FLW), length (FLL), and area (FLA) in four environments. A total of 31 QTL were detected in four environments. Two QTL for FLL on chromosomes 3B and 4A (QFll-3B and QFll-4A) and one for FLW on chromosome 2A (QFlw-2A) were major stable QTL. Ten QTL clusters (C1–C10) simultaneously controlling FLRTs and yield-related traits (YRTs) were identified. To investigate the genetic relationship between FLRTs and YRTs, correlation analysis was conducted. FLRTs were found to be positively correlated with YRTs especially with kernel weight per spike and kernel number per spike in all the three RIL populations and negatively correlated with spike number per plant. Appropriate flag leaf size could benefit the formation of high yield potential. This study laid a genetic foundation for improving yield potential in wheat molecular breeding programs.