利用单片段代换系研究水稻产量相关性状QTL加性及上位性效应

产量及其相关性状如单株有效穗数、千粒重、穗实粒数、穗总粒数和结实率等是水稻重要的农艺性状,了解产量及其相关性状QTL的加性及上位性效应对以分子标记聚合育种改良水稻产量具有重要意义。本文以16个单片段代换系及15个双片段代换系分析了水稻产量相关性状QTL的加性及上位性效应。共检出影响产量及其相关性状的13个QTL,包括产量性状1个、单株有效穗数1个、千粒重4个、穗实粒数4个、穗总粒数2个和结实率1个,分布于第2、第3、第4、第7和第10染色体上。此外,检出12对双基因互作。结果显示,2个正向(或负向)产量性状QTL聚合,往往会产生负向(或正向)的上位性效应,能否产生更大(或更小)的目标性状,取决于双片段遗传效应(加性效应与上位效应代数和)绝对值与单片段最大加性效应绝对值的差。本研究结果对实施高产分子标记聚合育种方法有重要参考价值。     

Identification of quantitative trait loci underlying seed protein and oil contents of soybean across multi‐genetic backgrounds and environments

Seed protein and oil contents are important quantitative traits in soybean. Previously, quantitative trait loci (QTL) associated with seed protein and oil were mostly identified in single genetic background. The objective of this work was to identify QTL and their epistatic effects underlying seed protein and oil contents in three recombinant inbred line populations (two of them used one common female parent) across eight environments by composite interval mapping. Forty QTL underlying protein content and 35 QTL underlying oil content were identified. Among them, nine were universal QTL underlying protein content and four were universal QTL underlying oil content. Epistatic interactions between QTL underlying seed protein/oil and different genetic backgrounds were detected. Different pairs of epistatic interactions were observed in diverse genetic backgrounds across multi‐environments. Common marker intervals were observed to simultaneously underlie seed protein and oil contents with different epistatic interactions. The results in this study suggested that a specific genotype with high oil content and low protein content might significantly affect the selection of soybean lines for high seed protein.     

烤烟几种化学成分的QTL初步分析

以含137个株系的烤烟DH群体(G-28×NC2326)及其亲本为材料, 在以前作图数据的基础上, 新增23个标记。将这些标记数据合并起来构建了包括11个ISSR标记和158个RAPD标记、由27个连锁群组成的烤烟分子标记遗传连锁图, 覆盖长度2 094.6 cM, 相邻标记间的平均图距为15.95 cM。利用4个环境下的试验数据进行了总糖、烟碱、氧化钾3种烟叶化学成分的QTL初步分析, 共检测到7个加性效应QTL和9对加加上位性效应QTL, 其中3个加性QTL和3对上位性QTL存在QTL与环境互作效应(QE)。表明在烤烟总糖、烟碱、氧化钾的遗传控制中除加性效应外, 上位性效应也具有重要作用。对于烟碱、氧化钾检测到加性QTL与环境互作效应, 对于总糖、氧化钾检测到上位性QTL与环境互作效应, 利用这些与环境具有互作效应的QTL进行标记辅助选择时宜考虑特定的环境条件。     

大豆叶片性状和叶绿素含量QTL间的上位性和环境互作效应

以丰产性好、抗旱力强的栽培大豆晋豆23为母本,山西农家品种半野生大豆灰布支黑豆为父本杂交衍生的447个RIL作为供试群体。将亲本及447个家系分别于2011、2012和2013年采用随机试验种植,按照标准测量叶长、叶宽和叶柄长3个性状,并于2012年8月1日和8月8日和2013年8月2日和8月9日各测量1次叶绿素含量。采用QTLNETwork 2.0混合线性模型分析方法和主基因+多基因混合遗传分离分析法,对大豆叶片性状和叶绿素含量进行遗传分析和QTL间的上位性和环境互作效应研究。结果表明,叶长受2对加性-加性×加性上位性混合主基因控制,叶宽受3对等效主基因控制,叶柄长受4对加性-加性×加性上位性主基因控制,叶绿素含量受4对加性主基因控制;检测到10个与叶长、叶宽、叶柄长和叶绿素含量相关的QTL,分别位于A1、A2、C2、H_1、L和O染色体。其中2个叶长QTL分别位于C2和L染色体,是2对加性×加性上位互作效应及环境互作效应QTL;3个叶宽加性与环境互作QTL分别位于A2、C2和O染色体;2个叶柄长QTL分别位于L和O染色体;3个叶绿素含量QTL分别位于A1、C2和H_1染色体。叶片性状和叶绿素含量的遗传机制较复杂,加性效应、加性×加性上位互作效应及环境互作效应是大豆叶片性状和叶绿素含量的重要遗传基础。建议大豆分子标记辅助育种中,一方面要考虑起主要作用的QTL,另一方面要注重上位性QTL的影响,这对于性状的遗传和稳定表达具有积极的意义。     

显性无腺体与隐性无腺体陆地棉品种间杂种二代利用研究

利用４个隐性低酚棉品系为母本，５个显性无腺体品系为父本，按不完全双列杂交（ＮＣⅡ）设计配制２０个组合，利用亲本、Ｆ１和Ｆ２材料采用ＡＤＡＡ模型研究表明，衣分以加性效应为主，其余产量及产量构成因素以显性遗传效应为主。子棉产量和铃重有较低的上位遗传效应。纤维品质性状中，绒长与整齐度以显性遗传效应为主，绒长还存在少量上位效应；比强度与伸长率仅存在显著的加性遗传效应，麦克隆值以加性遗传效应为主。Ｆ１皮棉产量、第一次收花率、单株铃数、铃重和衣分的群体超亲优势平均值分别为：２７．９％、１８．７％、１６．２％、９．７％和１％，Ｆ２分别为：１０．８％、５．１％、４．５％、４．６％和－１．２％。Ｆ１绒长的群体平均优势为２．５％，比强度和伸长率为－１．８和０．８％，麦克隆值和整齐度分别为－０．２％和－２．７％。Ｆ２绒长略呈下降趋势，比强度、伸长率和麦克隆值Ｆ２与Ｆ１相近，整齐度Ｆ２略优于Ｆ１。表明了显性无腺体基因的杂种优势利用前景广阔。     

大豆产量及主要农艺性状QTL的上位性互作和环境互作分析

以栽培大豆晋豆23为母本,半野生大豆灰布支黑豆ZDD2315为父本杂交衍生的F2:15和F2:16的447个RIL家系为遗传群体,绘制SSR遗传图谱,采用混合线性模型方法,对2年大豆小区产量及主要农艺性状进行加性QTL、加性×加性上位互作及环境互作分析。结果检测到9个与小区产量、茎粗、有效分枝、主茎节数、株高、结荚高度相关的QTL,分别位于J_2、I、M连锁群上,其中小区产量、茎粗、株高、有效分枝和主茎节数QTL的加性效应为正值,说明增加这些性状的等位基因来源于母本晋豆23。同时,检测到7对影响小区产量、茎粗、株高和结荚高度的加性×加性上位互作效应及环境互作效应的QTL,共发现14个与环境存在互作的QTL。上位效应和QE互作效应对大豆小区产量及主要农艺性状的遗传影响较大。大豆分子标记辅助育种中,既要考虑起主要作用的QTL,又要注重上位性QTL,才有利于性状的稳定表达和遗传。     

陆地棉重组自交系产量及产量构成因子性状的上位性QTL分析

棉花的产量及产量构成因子性状是以复杂的方式遗传,遗传力较低并易受环境条件影响。经典数量遗传学指出,上位性是复杂性状的遗传基础。本研究以湘杂棉2号F8和F9世代重组自交系为材料,调查了3个环境下的产量及产量构成因子性状,并构建了遗传连锁图。旨在定位产量及产量构成因子性状的上位性QTL并分析QTL与环境的互作效应。所有产量及产量构成因子性状均检测到上位性QTL,共检测到16对加性互作QTL（AA）,涉及的位点中仅4个有单位点效应,这反映了上位性的复杂性及其对产量和产量构成因子性状的重要贡献。共检测到17对QTL加性和环境互作（AE）,以及14对上位性QTL与环境的互作,表明环境因素对产量和产量构成因子性状起重要影响作用。研究结果还表明上位性效应作为湘杂棉2号的遗传基础起着重要作用。对各性状在不同环境的优良基因型进行了预测。综合优良家系（GSL）和特定环境下的优良家系（SL）的性状表现高于两亲本,表明湘杂棉2号重组自交系各性状都有提高的潜力。由于QTL加性和环境互作以及上位性QTL与环境互作的影响,预测的优良家系基因型会随着环境的改变而不同,表明应针对特定环境开展棉花育种。     

Comparison and analysis of main effects,epistatic effects,and QTL × environment interactions of QTLs for agronomic traits using DH and RILs populations in rice

Two genetic linkage maps based on doubled haploid (DH) and recombinant inbred lines (RILs) populations, derived from the same indica-japonica cross ‘Samgang × Nagdong’, were constructed to analyze the quantitative trait loci (QTLs) affecting agronomic traits in rice. The segregations of agronomic traits in RILs population showed larger variations than those in DH population. A total of 10 and 12 QTLs were identified on six chromosomes using DH population and seven chromosomes using RILs population, respectively. Three stable QTLs including pl9.1, ph1.1, and gwp11.1 were detected through different years. The percentages of phenotypic variation explained by individual QTLs ranged from 8 to 18% in the DH population and 9 to 33% in the RILs population. Twenty-three epistatic QTLs were identified in the DH population, while 21 epistatic QTLs were detected in the RILs population. Epistatic interactions played an important role in controlling the agronomic traits genetically. Four significant main-effect QTLs were involved in the digenic interactions. Significant interactions between QTLs and environments (QE) were identified in two populations. The QTLs affecting grain weight per panicle (GWP) were more sensitive to the environmental changes. The comparison and QTLs analysis between two populations across different years should help rice breeders to comprehend the genetic mechanisms of quantitative traits and improve breeding programs in marker-assisted selection (MAS).     

Multiple‐line cross QTL mapping for grain yield and thousand kernel weight in triticale

Grain yield and its component trait thousand kernel weight are important traits in triticale breeding programmes. Here, we used a large mapping population of 647 doubled haploid lines derived from four families to dissect the genetic architecture underlying grain yield and thousand kernel weight by multiple‐line cross QTL mapping. We identified 3 QTL for grain yield and 13 for thousand kernel weight which cross‐validated explained 5.2% and 48.2% of the genotypic variance, respectively. Both traits showed a positive phenotypic correlation, and we found two QTL overlapping between them. Full two‐dimensional epistasis scans revealed epistatic QTL for both traits, suggesting that epistatic interactions contribute to their genetic architecture. Based on QTL identified in our results, we conclude that the potential for marker‐assisted selection is limited for grain yield but more promising for thousand kernel weight.     

Molecular genetic analysis of flour color using a doubled haploid population in bread wheat (<Emphasis Type="Italic">Triticum aestivum</Emphasis> L.)

Flour color is an important trait in the assessment of flour quality for the production of many end products. In this study, quantitative trait loci (QTLs) with additive effects, epistatic effects, and QTL × environment (QE) interactions for flour color in bread wheat (Triticum aestivum L.) were studied, using a set of 168 doubled haploid (DH) lines derived from a Huapei 3 × Yumai 57 cross. A genetic map was constructed using 283 simple sequence repeats (SSR) and 22 expressed sequence tags (EST)-SSR markers. The DH and parents were evaluated for flour color in three environments. QTL analyses were performed using QTLNetwork 2.0 software based on a mixed linear model approach. A total of 18 additive QTLs and 24 pairs of epistatic QTLs were detected for flour color, which were distributed on 19 of the 21 chromosomes. One major QTL, qa1B, closely linked to barc372 0.1 cM, could account for 25.64% of the phenotypic variation of a* without any influence from the environments. So qa1B could be used in the molecular marker-assisted selection (MAS) in wheat breeding programs. The results showed that both additive and epistatic effects were important genetic basis for flour color, and were also sometimes subject to environmental modifications. The information obtained in this study should be useful for manipulating the QTLs for flour color by MAS in wheat breeding programs. Kun-Pu Zhang and Guang-Feng Chen contributed equally to this study.     

Dynamic analyses of rice blast resistance for the assessment of genetic and environmental effects

A doubled haploid population was employed to characterize the dynamic changes of the genetic components involved in rice blast resistance, including main-effect quantitative trait loci (QTLs), epistatic QTLs and QTL-by-environment interactions. The study was carried out at three different developmental stages of rice, using natural infection tests over 2 years. The number of main-effect QTLs, epistatic QTLs and their environmental interactions differed across the various measuring stages. One QTL ( d12 ) on chromosome 12 was detected at all stages, whereas most QTLs were active only at one or two stages in the population. These findings suggest that the unstable expression of most QTLs identified for blast resistance was influenced by the developmental status of the plants, epistatic effects between different loci and the environments in which they were grown. These findings demonstrate the complexity of expression of rice blast resistance and have important implications for durable resistance-breeding and map-based cloning of quantitative traits.     

甘蓝型油菜产量及其构成因素的QTL定位与分析

产量性状是复杂的数量性状, 对种子的单株产量及其构成因素(全株总有效角果数、每角粒数、千粒重)进行QTL定位和上位性分析,确定其在染色体上的位置及其遗传效应,可以探讨油菜杂种优势产生原因,提高育种中对产量性状优良基因型选择的效率,达到提高油菜产量的目的。在双低油菜细胞质雄性不育保持系1141B和双高恢复系垦C₁构建的F₂作图群体中,运用SRAP、AFLP和SSR三种标记技术构建了一个甘蓝型油菜(Brassica napus L.)的分子标记遗传连锁图谱。共包含244个标记,分布于20个主要连锁群、1个三联体上,图谱总长度为2 769.5 cM。采用Windows QTL Cartographer Version 2.0统计软件及复合区间作图法,对油菜单株产量及其3大构成因素进行QTL定位,共检测到QTLs 16个分布在9个连锁群上,其中第6和13连锁群最多,均有3个。单个QTL解释性状表型变异的0.38%～73.34%。对于同一性状,等位基因的增效作用既来自母本,亦源自父本;采用双向方差分析法对位点间互作及其上位性进行分析,检测到26对影响产量构成性状的上位性互作效应QTL,说明油菜基因组中存在大量控制产量的互作位点,油菜产量性状的上位性存在着多效性,上位性互作包括QTL与非QTL位点,其中以非QTL位点较多。一般互作位点的独立效应值较小,而互作的效应值显著增大,且一般超过两位点独立效应值之和。反映了控制产量性状基因的复杂性。上位性是甘蓝型油菜产量性状杂种优势的重要遗传基础。     

Inheritance of isoflavone contents in soybean seeds

Summary We studied the genetic basis of isoflavone content inheritance in soybean seeds. The progenitors BARC-8 (low isoflavone content), IAC-100 (high isoflavone content), the F₁ and F₂ populations derived from reciprocal crosses, and backcross populations were analyzed for isoflavone content and composition. Six isoflavones were detected: daidzin (DZ), genistin (GT), glycitin (GC), malonyldaidzin (MDZ), malonylgenistin (MGT) and malonylglycitin (MGC). DZ, GT, MDZ and MGT contents were influenced by the cytoplasm and the nuclear genes of the maternal parent. For this reason, a genetic model was considered that included the cytoplasmic effect and epistasis between nuclear and cytoplasmic genes. Except for GT, the additive effect was the most important one. For GT content the cytoplasmic effect was the most important. Except for MDZ, the epistatic effects were significant for all the isoflavone forms. Our data indicate that genetic improvement for these traits should explore the additive genetic variances in superior lines or the cytoplasmic effect and the epistatic interactions between cytoplasmic and nuclear genes to obtain the largest selection gains.     

Genetic Analysis of Developmental Traits in Chickpea

Chickpea (Cicer arietinum L.) is an important legume crop in India. The present study was conducted to investigate the inheritance of several developmental traits in three crosses of chickpea, viz., WFWG III’בT20’, ‘T88’בBold Seeded’, and ‘NP34’בP1528-1-1’, each having seven generations. The seven generations were P₁, P₂, F₁, B₁, B₂, F₂, and F₃. The experimental lay-out was randomized complete block design with three replications. Data were acquired on days to flowering (DF), days to maturity (DM), plant height in cm (PH), number of primary branches (PB), and number of secondary branches (SB). Generation mean analysis was used to estimate the genetic components; narrow sense heritability was estimated using variance components; and correlation analysis to estimate correlation coefficients among different traits. Genetic differences were found in all three crosses for all traits studied. Additive, dominance, and epistatic effects were found for many traits'. Duplicate epistasis was observed for all traits except number of PB. Higher order interactions and/or linkage were detected for DM and SB. For many traits the relative magnitudes of the genetic effects differed among crosses, thus the extrapolation to other crosses may be difficult. The inheritance becomes more complex as the fate of the character is decided at a later stage in the life cycle. Positive heterosis was observed for some traits, but the exploitation of this component may not Feasible since stable male sterile lines are not available. Early maturity and high yield ‘may be selected independently because of the absence of any significant correlation between these two traits.     

利用BC2F2高代回交群体定位水稻籽粒大小和形状QTL

以我国优良籼稻恢复系蜀恢527为轮回亲本, 以来自菲律宾的Milagrosa为供体亲本, 培育了样本容量为199株的BC₂F₂高代回交群体。选取85个均匀分布在12条染色体上的多态性SSR标记进行基因型分析, 同时对粒长、粒宽、长宽比和千粒重4种性状进行了表型鉴定。采用性状－标记间的单向和双向方差分析对上述性状进行了QTL定位。单向方差分析(P<0.01)共检测到了10个控制粒长、粒宽、长宽比和千粒重的QTL, 其中有3个具有多效性。由于粒长和长宽比的高度相关性, 控制长宽比的2个QTL均能在粒长QTL中检测到。位于第3染色体着丝粒区域的qgl3b是一个控制粒长、长宽比和千粒重的主效QTL, 它可以分别解释粒长、长宽比和千粒重表型变异的29.37%、26.15%和17.15%。该QTL对于粒长、长宽比和千粒重均表现较大的加性效应(来自蜀恢527的等位基因为增效)和负向超显性。位于第8染色体的qgw8位点是一个控制粒宽的主效QTL, 同时也是控制千粒重的微效QTL, 能解释粒宽表型变异的21.47%和千粒重表型变异的5.16%。该QTL对粒宽和千粒重均具有较大的加性效应(来自蜀恢527的等位基因为增效)和正向部分显性。双向方差分析(P<0.005)共检测到61对显著的上位性互作, 涉及54个QTL, 其中23个是能同时影响2~4个性状的多效位点, 且有8个位点与单向方差分析检测到的相同。控制长宽比的13对上位性互作位点中, 与控制粒长的上位性互作位点完全相同的有8对。以上结果为进一步开展水稻籽粒大小和形状有利基因的精细定位、克隆和分子设计育种奠定了基础。     

利用BC2F2高代回交群体定位水稻籽粒大小和形状QTL

以我国优良籼稻恢复系蜀恢527为轮回亲本, 以来自菲律宾的Milagrosa为供体亲本, 培育了样本容量为199株的BC₂F₂高代回交群体。选取85个均匀分布在12条染色体上的多态性SSR标记进行基因型分析, 同时对粒长、粒宽、长宽比和千粒重4种性状进行了表型鉴定。采用性状－标记间的单向和双向方差分析对上述性状进行了QTL定位。单向方差分析(P<0.01)共检测到了10个控制粒长、粒宽、长宽比和千粒重的QTL, 其中有3个具有多效性。由于粒长和长宽比的高度相关性, 控制长宽比的2个QTL均能在粒长QTL中检测到。位于第3染色体着丝粒区域的qgl3b是一个控制粒长、长宽比和千粒重的主效QTL, 它可以分别解释粒长、长宽比和千粒重表型变异的29.37%、26.15%和17.15%。该QTL对于粒长、长宽比和千粒重均表现较大的加性效应(来自蜀恢527的等位基因为增效)和负向超显性。位于第8染色体的qgw8位点是一个控制粒宽的主效QTL, 同时也是控制千粒重的微效QTL, 能解释粒宽表型变异的21.47%和千粒重表型变异的5.16%。该QTL对粒宽和千粒重均具有较大的加性效应(来自蜀恢527的等位基因为增效)和正向部分显性。双向方差分析(P<0.005)共检测到61对显著的上位性互作, 涉及54个QTL, 其中23个是能同时影响2~4个性状的多效位点, 且有8个位点与单向方差分析检测到的相同。控制长宽比的13对上位性互作位点中, 与控制粒长的上位性互作位点完全相同的有8对。以上结果为进一步开展水稻籽粒大小和形状有利基因的精细定位、克隆和分子设计育种奠定了基础。     

小麦DH群体穗下节间直径、茎壁厚及茎壁面积的QTL定位

由小麦品种花培3号和豫麦57杂交获得168个DH株系,连续两年在山东泰安种植,利用324个SSR标记构建遗传连锁图谱,并基于混合线性模型对控制穗下节间直径、茎壁厚及茎壁面积的QTL遗传效应和环境互作效应进行分析。共检测到10个加性效应位点和6对上位效应位点,其中3个加性位点参与环境互作效应。检测到位于染色体2D、3D和5D(2个)控制穗下节间直径的4个加性QTLs,与控制茎壁厚的3个加性位点相同或相邻,表现出一因多效或紧密连锁效应。两个位于染色体2D和5D控制茎壁厚和茎壁面积QTL有较大遗传贡献率,分别为11.37%和10.98%,适用于分子标记辅助育种和聚合育种。6对上位性效应遗传贡献率较小、无环境互作效应。     

陆地棉产量性状的遗传分析

采用加性-显性与环境互作的遗传模型,分析陆地棉8个杂交亲本和F128个组合的7个产量性状的两年试验资料,估算各项遗传方并非分量和F1、F2的杂种优势.结果表明,产量性状受加性效应和显性效应共同控制,加性×环境各产量性状均极显著,铃重、衣分、籽指还受显性×环境的极显著影响,7项产量性状的广义和狭义遗传率均达到极显著水准.     

Quantitative trait loci mapping and epistatic analysis for grain yield and yield components using molecular markers with an elite maize hybrid

Summary The aim of this investigation was to map quantitative trait loci (QTL) associated with grain yield and yield components in maize and to analyze the role of epistasis in controlling these traits. An F_2:3 population from an elite hybrid (Zong3 × 87-1) was used to evaluate grain yield and yield components in two locations (Wuhan and Xiangfan, China) using a randomized complete-block design. The mapping population included 266 F_2:3 family lines. A genetic linkage map containing 150 simple sequence repeats and 24 restriction fragment length polymorphism markers was constructed, spanning a total of 2531.6 cM with an average interval of 14.5 cM. A logarithm-of-odds threshold of 2.8 was used as the criterion to confirm the presence of one QTL after 1000 permutations. Twenty-nine QTL were detected for four yield traits, with 11 of them detected simultaneously in both locations. Single QTL contribution to phenotypic variations ranged from 3.7% to 16.8%. Additive, partial dominance, dominance, and overdominance effects were all identified for investigated traits. A greater proportion of overdominance effects was always observed for traits that exhibited higher levels of heterosis. At the P ≤ 0.005 level with 1000 random permutations, 175 and 315 significant digenic interactions were detected in two locations for four yield traits using all possible locus pairs of molecular markers. Twenty-four significant digenic interactions were simultaneously detected for four yield traits at both locations. All three possible digenic interaction types were observed for investigated traits. Each of the interactions accounted for only a small proportion of the phenotypic variation, with an average of 4.0% for single interaction. Most interactions (74.9%) occurred among marker loci, in which significant effects were not detected by single-locus analysis. Some QTL (52.2%) detected by single-locus analysis were involved in epistatic interactions. These results demonstrate that digenic interactions at the two-locus level might play an important role in the genetic basis of maize heterosis.