Identification of Related QTLs at Late Developmental Stage in Rice (Oryza sativa L.) Under Two Nitrogen Levels

QTL underlying related traits at the late developmental stage under two different nitrogen levels were investigated in rice using a population of chromosome segment substitution lines (CSSL) derived from a cross between Teqing and Lemont. A total of 31 QTLs referring 5 traits, that is, plant height (PH), panicle number per plant (PN), chlorophyll content (CC), shoot dry weight (SDW) and grain yield per plant (YD), were detected. Under normal N level, 3 QTLs were detected for each trait, while under low N level, 5,4, 5 and 2 QTLs were detected for PH, PN, CC and SDW respectively. Most of the QTLs were located on chromosome 2, 3, 7, 11 and 12. QTLs controlling different traits or the same trait under different N levels were mapped on the same or adjacent intervals, forming several clusters in rice chromosomes. More than two traits were controlled by QTLs on one of four intervals (RM30-RM439, RM18-RM478, RM309-RM270, and RM235-RM17), suggesting that there were some pleiotropic effects. It was supposed that some QTLs only detected at low N level might be associated with the ability to tolerate the low N stress in rice.     

两种供氮水平下水稻生长后期相关性状的QTL定位

以特青为母本与Lemont杂交,然后用特青为轮回亲本回交,建立特青背景下的染色体片段置换系（CSSL）群体。在正常和低氮条件下分别在生长后期对株高（PH）、单株穗数（PN）、叶绿素含量（CC）、地上部干物重（SDW）和单株籽粒产量（YD）等性状进行了QTL分析,共检测到31个QTL。其中在正常供氮水平下控制PH、PN、CC、SDW和YD的QTL数目均为3个;在低氮水平下检测到5、4、5和2个影响PH、PN、CC和SDW的QTL,在低氮水平下没有检测到控制YD的位点。大部分QTL集中在第2、3、7、11和12染色体上,影响不同性状或在两种供氮水平下影响同一性状的QTL在染色体上成串或成簇分布。其中RM30-RM439、RM18-RM478、RM309-RM270、RM235-RM17等区域同时检测到控制两个以上性状的QTL,表现出明显的一因多效现象。推测仅在低氮水平下检测到的QTL可能跟水稻对低氮胁迫耐性有一定的关联。     

水稻耐亚铁毒QTLs的定位

亚铁毒是潜育性水稻土中限制水稻产量的主要因子。利用龙杂8503／IR64的F2和等价的F3群体,在营养液中培养来定位耐亚铁毒的QTLs。通过构建101SSR标记的遗传连锁图谱来确定耐亚铁毒QTLs的位置和特性。借助叶片棕色斑点指数、株高和最大根长3个性状,利用营养液在水稻苗期来评价F2单株、F3群体和亲本龙杂8503、IR64,共检测到叶片棕色斑点指数、株高和最大根长的QTLs20个,分布在水稻的10条染色体上,表明这些性状受多基因控制。控制叶片棕色斑点指数的QTLs分别定位在第1染色体的RM315-RM212、第2染色体的RM6-RM240和第4染色体的RM252-RM451之间。与前人的研究结果比较发现：1）位于第4染色体RM252-RM451之间的控制叶片棕色斑点指数的QTL与水稻功能图谱上控制叶绿素含量减少的QTL的位置一致。另一个位于第1染色体的RM315-RM212之间的控制叶片棕色斑点指数的QTL与水稻功能图谱上位于C178-R2635之间控制叶绿素含量的QTL连锁。2）位于第2染色体RM6-RM240之间的第3个控制叶片棕色斑点指数的QTL与位于RZ58-CD0686的控制钾吸收的QTL连锁。     

两个水稻骨干恢复系重要农艺性状的遗传基础研究

水稻骨干恢复系是指在杂交稻育种中广泛应用的一类恢复系。探明骨干恢复系的遗传基础,发掘其重要农艺性状基因/QTL,对分子标记辅助选择水稻恢复系育种具有重要应用价值。本研究以生产上广泛应用的三系骨干恢复系成恢727和两系骨干恢复系9311为亲本,培育了具有250个系的重组自交系群体。分别在2015年三亚和2016年合肥两个环境下进行了9个重要农艺性状表型和SSR分子标记基因型鉴定,用SAS9.2分析表型数据,用QTL Ici Mapping v4.1进行QTL定位分析。在三亚和合肥两个环境下共检测到39个QTL,三亚检测到16个,分布于第1、2、4、7、8、10、11和12染色体上;合肥检测24个,分布于第1、2、3、7、8、9、10和12染色体上。其中qPH1-1在三亚和合肥两个环境下都能检测到,加性效应分别为-1.75和-2.46。在检测到的39个QTL中,有24个QTL的增效等位基因来自恢复系成恢727,15个QTL的增效等位基因来自9311。共计有26个QTL曾被前人定位,13个属于尚未见文献报道的新QTL。另外,在RM279~RM521、RM336~RM3534、RM25~RM547、RM553~RM160、RM222~RM271区段内检测到5个多效性QTL位点。其中RM25~RM547位点与已经克隆的基因Ghd8位置相近。RM553~RM160位点是一个新的多效性位点,分别控制每穗实粒数、单株产量和结实率,而且效应和表型变异贡献率都较大。其余3个位点在前人的研究中分别有所报道,但其多效性则是在本研究中首次发现。在本研究新发掘到的QTL中,控制穗数的QTL qPN12-1,控制穗长的QTL qPL1-2和qPL10-1,控制总粒数的QTL qSNP2-1和qSNP10-1,控制结实率的QTL qSF3-1,控制千粒重QTL qTGW7-1和控制产量的QTL qGY1-1效应均比较大,解释的表型遗传变异比例也较高。本研究的结果将会为相关性状QTL的精细定位、克隆和育种应用奠定基础。     

水稻幼苗活力性状的低温反应数量性状基因座检测

以籼粳交“密阳23/吉冷1号”的F2：3代200个家系作为作图群体,在12℃冷水胁迫下,进行苗高、苗鲜重和苗干重等水稻幼苗活力性状的低温反应鉴定,并利用由SSR标记构建的分子连锁图谱为基础,对冷水胁迫下苗高、苗鲜重和苗干重以及它们的低温反应指数进行了数量性状基因座（QTLs）检测。研究结果表明,低温胁迫下上述幼苗活力性状在F3家系群中均表现为接近正态的连续分布,表现为由多基因控制的数量性状;在第1、2、7、8和12染色体上,检测到与幼苗活力性状的低温反应相关的QTL共12个,对表型变异的贡献率范围为5．2％-17．9％,其中位于第2染色体RM262-RM263区间和第12染色体RM270-RM17区间的与低温下苗高相关的qCSH2和qCSH12,以及位于第12染色体RM19-RM270区间和第1染色体RM129-RM9区间的分别控制低温下苗干重及其低温反应指数的qSDW12和qCSDW1对表型变异的贡献率较大,分别为16．6％、17．9％、15．9％和16．2％。其增效等位基因均来自吉冷1号,前两者均表现为加性效应,后两者分别表现为显性和超显性。     

Mapping QTLs for Plant Phenology and Production Traits Using Indica Rice (Oryza sativa L.) Lines Adapted to Rainfed Environment

Drought is a major abiotic stress limiting rice production and yield stability in rainfed ecosystems. Identifying quantitative trait loci (QTL) for rice yield and yield components under water limited environments will help to develop drought resilient cultivars using marker assisted breeding (MAB) strategy. A total of 232 recombinant inbred lines of IR62266/Norungan were used to map QTLs for plant phenology and production traits under rainfed condition in target population of environments. A total of 79 QTLs for plant phenology and production traits with phenotypic variation ranging from 4.4 to 72.8% were detected under non-stress and drought stress conditions across two locations. Consistent QTLs for phenology and production traits were detected across experiments and water regimes. The QTL region, RM204-RM197-RM217 on chromosome 6 was linked to days to 50% flowering and grain yield per plant under both rainfed and irrigated conditions. The same genomic region, RM585-RM204-RM197 was also linked to harvest index under rainfed condition with positive alleles from Norungan, a local landrace. QTLs for plant production and drought resistance traits co-located near RM585-RM204-RM197-RM217 region on chromosome 6 in several rice genotypes. Thus with further fine mapping, this region may be useful as a candidate QTL for MAB, map-based cloning of genes and functional genomics studies for rainfed rice improvement.     

粳稻蒸煮食味品质相关性状的QTL分析

以沈农265和丽江新团黑谷杂交衍生的重组自交系群体(RILs)为实验材料,对12个粳稻(Oryza sativa subsp.japonica)蒸煮食味品质相关性状进行QTL分析。共检测到29个蒸煮食味品质相关的QTLs,分布于除第8染色体外的11条染色体上,LOD值介于2.50–16.47之间,加性效应值为–132.69–471.85,单个QTL贡献率为10.36%–73.24%。在第6染色体RM508–RM253区域检测到1个蒸煮营养食味品质多效性QTL簇,其中q AC6表型贡献率最大,解释73.24%的表型变异;在第10染色体PM166–RM258区域检测到2个与蒸煮食味品质相关的QTLs,分别是控制口感的q CTS10和综合评分的q CCS10。此外,检测到15个与RVA特征谱相关的QTLs,在第6染色体RM253–RM402区域检测到3个与RVA谱特征值相关的QTLs,表型贡献率均大于12%。这些定位结果将为粳稻蒸煮食味相关品质的分子遗传机理研究奠定基础。     

Validation and dissection of quantitative trait loci for leaf traits in interval RM4923-RM402 on the short arm of rice chromosome 6

Validation and dissection of a QTL region for leaf traits in rice which has been reported in a number of independent studies were conducted. Three sets of near isogenic lines (NILs) were originated from a residual heterozygous line derived the indica cross Zhenshan 97B/Milyang 46. They were overlapping and totally covered a 4.2-Mb heterogenous region extending from RM4923 to RM402 on the short arm of rice chromosome 6. Each NIL set consisted of 10 maternal lines and 10 paternal lines. They were measured for the length, width, perimeter and area of the top three leaves and the number of spikelets per panicle, number of grains per panicle and grain weight per panicle. In NIL sets 6-4 and 6-7, differing in intervals RM4923-RM225 and RM19410-RM6119, respectively, significant variations with the enhancing alleles from the female parent ZS97 were shown for the length, perimeter and area except for the area of the third leaf from top in 6-4, but the effects were lower in 6-4 than in 6-7. No significant effects were detected for the three traits in the remaining NIL set. It was shown that flag leaf length (FLL) is the primary target of the QTLs detected. Two QTLs for FLL linked in repulsion phase were resolved, of which qFLL6.2 located in the 1.19-Mb interval RM3414-RM6917 had a major effect with the enhancing allele from Zhenshan 97B, and qFLL6.1 located in the 946.8-kb interval RM19350-RM19410 had a smaller effect with the enhancing allele from Milyang 46. The two QTLs also exerted pleiotropic effects on the yield traits.     

Genetic analysis of rooting ability of transplanted rice (Oryza sativa L.) under different water conditions

In order to assess the benefits of superior rooting ability of rice (Oryza sativa L.) for growth after transplanting under water-limiting conditions, genetic differences in the rooting ability of rice seedlings 30 d after sowing, with their visible roots either pruned or not pruned, were quantified by several root parameters 4 d after transplanting (DAT), under flooded or non-flooded paddy fields (four treatments in total), together with production traits at maturity. Ninety-eight recombinant inbred lines from the two japonica ecotypes, a lowland variety Otomemochi, and an upland variety Yumenohatamochi, were genotyped with 107 simple sequence repeat (SSR) markers. Otomemochi in general produced more adventitious roots, partitioned a greater proportion of biomass to roots, and had a greater increment of root dry weight (DeltaRW) at 4 DAT than Yumenohatamochi, but these variety differences were less clear under non-flooded conditions without root pruning. Several quantitative trait loci (QTLs) associated with rooting ability were identified mainly in chromosomes 1, 4, and 6 across the four treatments or in combined analysis. On the other hand, QTLs for DeltaRW around RM2357 in chromosome 5 and for maximum new root length in RM215-RM205 in chromosome 9 were found only under root pruning treatments and under non-flooded conditions without root pruning, respectively. Greater DeltaRW was associated with higher head dry weight per hill at maturity only in the non-flooded and root-pruning treatments. This study suggests the importance of rooting ability after transplanting and, possibly, other mechanisms for adaptation to non-flooded conditions.     

基于CSSL的高密度物理图谱定位水稻分蘖角度QTL

对以籼稻9311为遗传背景携带粳稻日本晴基因组的染色体片段置换系（CSSL）的遗传图谱进行分子标记加密,构建了含250个多态标记的高密度物理图谱。以119个CSSLs为材料,P≤0.001为阈值,筛选到分蘖角度与受体亲本9311差异极显著的10个系。结合物理图谱和代换作图方法,共鉴定出5个分蘖角度QTL,其中qTA11的加性效应表现为增效作用,来源于9311的等位基因;其余4个QTL的加性效应为减效作用,均来源于日本晴的等位基因。qTA6-1和qTA6-2分别被定位于第6染色体RM253–RM527之间的3.55Mb区段和RM3139–RM494的1.65Mb区间;qTA9被定位于第9染色体RM257–RM189之间的3.40Mb区段;qTA10被定位在第10染色体RM222–S10-1之间的2.10Mb区段;qTA11被定位于第11染色体RM1761–RM4504之间的3.30Mb区间。以上研究结果为水稻分蘖角度QTL的精细定位和株型育种提供了依据。     

Identification for quantitative trait loci controlling grain shattering in rice

Seed shattering is an important factor causing loss of grain yield before and during rice harvest. In the present study, the quantitative trait loci regarding shattering scale, breaking tensile strength (BTS) and abscission layer (AL), the parameters evaluating seed shattering habit by hand gripping, a digital force gauge and observation on AL, respectively, were identified by using an doubled haploid line (DHL) population from a cross between a loose-shattering type Tongil variety, ‘Samgang’, and a moderately difficult shattering japonica variety, ‘Nagdong’. Eight QTLs consisted in four QTLs for shattering scale, two QTLs for AL, each one QTL for pulling and bending strength were detected on six chromosomes, respectively. Among them, Qss1 with flanking markers RM6696 and RM476 explained 31% of phenotype variation in shattering scale. Furthermore, two new QTLs controlling shattering habit, Qss5-2 and Qal5-1, were located on chromosome 5 at the interval 5028–5037 and 5021-RM289. They explained 10% and 12% of phenotype variations, respectively. A total of eleven digenic epistatic loci were identified for four parameters. The identification of QTLs affecting seed shattering habits is favorable to thoroughly dissect the genetic mechanism of the shattering habit and to apply for marker-assisted selection in rice breeding system of specific regions.     

水稻粒长QTL定位与主效基因的遗传分析

该研究利用短粒普通野生稻矮杆突变体和长粒栽培稻品种KJ01组配杂交组合F_1,构建分离群体F_2;并对该群体粒长进行性状遗传分析,利用平均分布于水稻的12条染色体上的132对多态分子标记对该群体进行QTL定位及主效QTLs遗传分析,为进一步克隆新的主效粒长基因奠定基础,并为水稻粒形育种提供理论依据。结果表明:(1)所构建的水稻杂交组合分离群体F_2的粒长性状为多基因控制的数量性状。(2)对543株F_2分离群体进行QTL连锁分析,构建了控制水稻粒长的连锁遗传图谱,总长为1 713.94 cM,共检测出24个QTLs,只有3个表现为加性遗传效应,其余位点均表现为遗传负效应。(3)检测到的3个主效QTLs分别位于3号染色体的分子标记PSM379～RID24455、RID24455～RM15689和RM571～RM16238之间,且三者对表型的贡献率分别为54.85%、31.02%和7.62%。(4)在标记PSM379～RID24455之间已克隆到的粒长基因为该研究新发现的主效QTL位点。     

水稻低温发芽势的遗传及数量性状基因座分析

利用籼粳交“密阳23／吉冷1号”的F2：3代200个家系作为作图群体,在14℃条件下鉴定萌发7d、11d、14d和17d时低温发芽势,并利用由SSR标记构建的分子连锁图谱为基础,对不同萌发阶段的低温发芽势进行了数量性状基因座（QTLs）检测,同时进行了低温发芽势与其他耐冷性状间的相关分析。结果表明,萌发7d时低温发芽势及其低温反应指数呈现向低发芽势和低的低温反应指数的偏态分布,而萌发11d、14d和17d时低温发芽势及其低温反应指数均呈现接近正态的单峰连续分布。萌发14d时低温发芽势与其他耐冷性的相关性较萌发7d、11d和17d时低温发芽势更为密切,与芽期耐冷性、幼苗期耐冷性、低温下幼苗生长能力和孕穗期耐冷性表现为显著或极显著的正相关。位于第2染色体RM29-RM262区间的qLVG2和第7染色体RM336-RM118区间的qLVG7-2、qCIVG7-2在萌发11d、14d和17d时均检测到;位于RM29-RM262区间的qCIVG2在萌发11d和14d时均检测到,并对表型变异的贡献率随着萌发进程而逐渐增加。与低温发芽势相关的qLVG2贡献率从6．9％增加到14.2％,qLVG7-2贡献率从9．9％增加到11．2％,而与发芽势的低温反应指数相关的qCIVG2贡献率从6.3％增加到9．0％,qCIVG7-2贡献率从8．3％增加12．9％。这些QTL的增效等位基因均来自强耐冷亲本吉冷1号,基因作用方式主要为部分显性。     

Genetic and QTL Analysis for Low-Temperature Vigor of Germination in Rice

The quantitative trait loci (QTLs) for low-temperature vigor of germination (LVG) with a germination period of 7 d, 11 d, 14 d, and 17 d at 14 °C was identified using F_2:3 population, which included 200 individuals and lines derived from a cross of indica and japonica “Milyang 23/Jileng 1” with microsatellite markers. The correlation coefficient between LVG and other cold tolerance traits was analyzed. LVG and the cold response index for vigor of germination (CIVG) detected when the germination period was 7 d showed a continuous distribution, which was partial to lower LVG and lower CIVG in F₃ lines. LVG and CIVG detected when the germination periods were 11 d, 14 d, and 17 d showed a continuous distribution near normal, which were quantitative traits controlled by multiple genes. LVG detected when the germination period was 14 d was more correlated with other cold tolerance traits than LVG detected when the germination periods were 7 d, 11 d, and 17 d, which was significantly associated with cold tolerance during the bud bursting period, the seedling stage, the booting stage, and the growing ability under cold conditions. qLVG2 located in RM29-RM262 on chromosome 2, qLVG7-2 and qCIVG7-2 located in RM336-RM118 on chromosome 7 were detected when the germination periods were 11 d, 14 d, and 17 d. qCIVG2 located in RM29-RM262 on chromosome 2 was detected when the germination periods were 11 d and 14 d. The variation is due to the observed phenotypic variation by the above QTLs, which was increased following the germination. The variation of qLVG2 related to LVG was increased from 6.9% to 14.2%. The variation of qLVG7-2 associated with LVG was increased from 9.9% to 11.2%. The variation of qCIVG2 correlated with CIVG was increased from 6.3% to 9.0%. The variation of qCIVG7-2 associated with CIVG was increased from 8.3% to 12.9%. These QTL alleles were obtained from the tolerant parent Jileng 1, and the gene action was most likely to be partially dominant.     

水稻糙米重金属Cd含量的QTL分析

镉(Cd²⁺)是一种分布较广泛、毒性较强的一种重金属, 文章利用韭菜青×IR26杂交衍生的一个重组自交系群体(Recombinant inbred lines, RIL)及构建的SSR分子遗传图谱, 对控制糙米中Cd²⁺含量的QTL进行分析, 为选育籽粒中Cd²⁺低吸收或低积累的水稻品种提供参考。结果表明, 在Cd²⁺胁迫(5 mg/kg)处理条件下, 共检测到2个与糙米Cd²⁺含量有关的QTLs, 分别位于水稻第11染色体上的标记RM6288-RM6544和RM167-RM5704之间, 其中qCCBR-11a对表型贡献率为11.17%, 加性效应0.089; qCCBR-11b对表型变异贡献率为7.66%, 加性效应0.075。相关分析显示, 糙米Cd²⁺含量与株高、每穗总粒数、每穗实粒数、结实率和千粒重等产量性状的相关性均不显著, 糙米中Cd²⁺含量是一个相对独立、由基因控制的遗传性状。     

Correlation and Quantitative Trait Loci Analyses of Total Chlorophyll Content and Photosynthetic Rate of Rice (Oryza sativa) under Water Stress and Well-watered Conditions

In order to explore the relevant molecular genetic mechanisms of photosynthetic rate (PR) and chlorophyll content (CC) in rice ( Oryza sativa L.), we conducted a series of related experiments using a population of recombinant inbred lines (Zhenshan97B × IRAT109). We found a significant correlation between CC and PR ( R = 0.19**) in well-watered conditions, but no significant correlation during water stress ( r = 0.08). We detected 13 main quantitative trait loci (QTLs) located on chromosomes 1, 2, 3, 4, 5, 6, and 10, which were associated with CC, including six QTLs located on chromosomes 1, 2, 3, 4, and 5 during water stress, and seven QTLs located on chromosomes 2, 3, 4, 6, and 10 in well-watered conditions. These QTLs explained 47.39% of phenotypic variation during water stress and 56.19% in well-watered conditions. We detected four main QTLs associated with PR; three of them ( qPR2 , qPR10 , qPR11 ) were located on chromosomes 2, 10, and 11 during water stress, and one ( qPR10 ) was located on chromosome 10 in well-watered conditions. These QTLs explained 34.37% and 18.41% of the phenotypic variation in water stress and well-watered conditions, respectively. In total, CC was largely controlled by main QTLs, and PR was mainly controlled by epistatic QTL pairs.     

Fine mapping a QTL qCTB7 for cold tolerance at the booting stage on rice chromosome 7 using a near-isogenic line

Low temperature at the booting stage is a serious abiotic stress in rice, and cold tolerance is a complex trait controlled by many quantitative trait loci (QTL). A QTL for cold tolerance at the booting stage in cold-tolerant near-isogenic rice line ZL1929-4 was analyzed. A total of 647 simple sequence repeat (SSR) markers distributed across 12 chromosomes were used to survey for polymorphisms between ZL1929-4 and the cold-sensitive japonica cultivar Towada, and nine were polymorphic. Single marker analysis revealed that markers on chromosome 7 were associated with cold tolerance. By interval mapping using an F₂ population from ZL1929-4 × Towada, a QTL for cold tolerance was detected on the long arm of chromosome 7. The QTL explained 9 and 21% of the phenotypic variances in the F₂ and F₃ generations, respectively. Recombinant plants were screened for two flanking markers, RM182 and RM1132, in an F₂ population with 2,810 plants. Two-step substitution mapping suggested that the QTL was located in a 92-kb interval between markers RI02905 and RM21862. This interval was present in BAC clone AP003804. We designated the QTL as qCTB7 (quantitative trait locus for cold tolerance at the booting stage on chromosome 7), and identified 12 putative candidate genes.     

水稻抗褐飞虱种质Swarnalata抽穗期和休眠性相关QTL检测

为有效利用抗褐飞虱水稻Swarnalata,对2013年南京种植的Swarnalata/02428 F2分离群体进行抽穗期和种子休眠性考察,利用172个分子标记构建了Swarnalata/02428 F2的分子遗传连锁图谱,图谱全长为3311.4c M,标记间平均图距为19.22c M。利用Windows QTL Cartographer V2.5软件对该分离群体进行抽穗期和种子休眠性相关QTL检测,共检测到7个抽穗期相关QTL,分别位于第2、3、6、11染色体,其中位于第11染色体的q HD-11-1贡献率最高,为28.85%;检测到3个种子休眠性相关QTL,分别位于第3、6、9染色体,其中位于第9染色体的q Sd-9贡献率最高,为22.11%。分析表明,本研究检测到的抽穗期QTL与种子休眠QTL所在位置不同,说明该群体中种子休眠与抽穗期没有直接关系,它们分别由不同基因控制。本研究不仅为水稻休眠基因的精细定位及克隆奠定基础,也为更有效利用Swarnalata中的抗褐飞虱基因提供基础和一些优良的中间材料。     

水稻产量库相关穗部性状的遗传分析

收人李源于珍汕97/明恢63的重组系群体中与产量库容有关的10个穗部性状的表现型数据。总体上,每穗颖花数与每穗二次枝梗数、每个二次枝梗上的颖花数、颖花密度有更大的相关性。对所研究的10个性状,两年间共检测到53个QTLs。约43.4%的QTLs能在两年同时检测到。5个染色体区域（第1染色体上G359-RG532和C567-C86-RG236,第2染色体上R712-RM29,第6染色体上P-RG424,第10染色体上C148-RM258）分别对多个穗部性状表现出效应。结果显示相关性状的QTLs大致定位在相似的染色体区域,这表明基因的多效性或紧密连锁是穗部性状间相关的遗传基因。在检测到的大量2位点互作对中,约18.2%在两年都能被检测到。不同性状的共同互作对的比例为8.7%～32.6%。在两年都能检测到2位点组合中,约26.7%的组合同时影响着多个性状,表现出多效效应。结果表明每个性状都由数个QTL、基因型与环境互作、大量的上位性互作所控制。