Mapping QTLs with epistatic effects and QTL×environment interactions for plant height using a doubled haploid population in cultivated wheat

Quantitative trait loci （QTLs） for plant height in wheat （Triticum aestivum L.） were studied using a set of 168 doubled haploid （DH） lines, which were derived from the cross Huapei 3/Yumai 57. A genetic linkage map was constructed using 283 SSR and 22 EST-SSR markers. The DH population and the parents were evaluated for wheat plant height in 2005 and 2006 in Tai＇an and 2006 in Suzhou. QTL analyses were performed using the software of QTLNetwork version 2.0 based on the mixed linear model. Four additive QTLs and five pairs of epistatic effects were detected, which were distributed on chromosomes 3A, 4B, 4D, 5A, 6A, 7B, and 7D. Among them, three additive QTLs and three pairs of epistatic QTLs showed QTL×environment interactions （QEs）. Two major QTLs, Qph4B and Qph4D, which accounted for 14.51% and 20.22% of the phenotypic variation, were located similar to the reported locations of the dwarfing genes Rhtl and Rht2, respectively. The Qph3A-2 with additive effect was not reported in previous linkage mapping studies. The total QTL effects detected for the plant height explained 85.04% of the phenotypic variation, with additive effects 46.07%, epistatic effects 19.89%, and QEs 19.09%. The results showed that both additive effects and epistatic effects were important genetic bases of wheat plant height, which were subjected to environmental modifications, and caused dramatic changes in phenotypic effects. The information obtained in this study will be useful for manipulating the QTLs for wheat plant height by molecular marker-assisted selection （MAS）.

Phenotypic Expression of Whitebacked Planthopper Resistance in the Newly Established japonica/indica Doubled Haploid Rice Population

A new doubled haploid (DH) rice population was established from a cross between WBPH-resistant japonica Chunjiang 06 (CJ-06) and susceptible indica TN1. Sucking inhibitory and ovicidal resistance of the DH rice lines were evaluated on the basis of non-preference response of WBPH immigrants and honeydew excretion by WBPH females, and appearance of watery lesions in the necrotic discoloration of leaf sheaths ovipositied by WBPH,respectively. Both the major gene resistance to WBPH, sucking inhibitory and ovicidal resistance, showed 1 (resistant): 1 (susceptible) segregation ratio in the DH population. Relative density of WBPH populations and damage scores in the DH population indicated combined functions of both the major resistance genes as well as QTLs affecting the host plant response to WBPH infestations. Thus, the newly developed CJ-06/TN1 DH population could be a useful material to analyze major genes and QTLs for WBPH resistance in japonica rice.

Differential Expression of Whitebacked Planthopper Resistance in the japonica/indica Doubled Haploid Rice Population under Field Evaluation and Seedbox Screening Test

Whitebacked planthopper (WBPH) -resistance in a japonica / indica doubled haploid (DH) rice population established from a cross between WBPH-resistant japonica Chun]iang 06 and susceptible indica TN1, was comparatively evaluated through a field experiment based on the WBPH immigrant density and standardized seedbox screening test (SSST). All the susceptible DH lines in the field experiment behaved accordingly in SSST. However, 35 of resistant 66 lines (53%) in the field, were categorized to susceptible groups in SSST. Likewise, there were no significant differences in WBPH immigrant densities among 70 DH lines that were highly resistant to susceptible in SSST. The results revealed that SSST could not evaluate properly WBPH resistance in the DH lines. Four QTLs for WBPH-resistance phenotyped by the immigrant density were detected on chromosomes 2, 3, 4, and 11. Of them, the QTL on chromosome 4 was the most effective (LOD 21.8, variance 78%). Five QTLs associated with seedling mortality were mapped on chromosomes 2, 3, 4, 5 and 6. In addition to the QTL (LOD 10.5, variance 68%) on chromosome 4, there was another major QTL (LOD 12.7, variance 71%) located on chromosome 5, which was SSST-specific but might be irrespective of the WBPH resistance traits.

Detection of QTLs with Additive Effects, Epistatic Effects, and QTL × Environment Interactions for Zeleny Sedimentation Value Using a Doubled Haploid Population in Cultivated Wheat

In order to understand the genetic basis for Zeleny sedimentation value （ZSV） of wheat, a doubled haploid （DH） population Huapei 3 × Yumai 57 （Yumai 57 is superior to Huapei 3 for ZSV）, and a linkage map consisting of 323 marker loci were used to search QTLs for ZSV. This program was based on mixed linear models and allowed simultaneous mapping of additive effect QTLs, epistatic QTLs, and QTL x environment interactions （QEs）. The DH population and the parents were evaluated for ZSV in three field trials. Mapping analysis produced a total of 8 QTLs and 2 QEs for ZSV with a single QTL explaining 0.64-14.39% of phenotypic variations. Four additive QTLs, 4 pairs of epistatic QTLs, and two QEs collectively explained 46.11% of the phenotypic variation （PVE）. This study provided a precise location of ZSV gene within the Xwmc 93 and GluD1 interval, which was designated as Qzsv-1D. The information obtained in this study should be useful for manipulating the QTLs for ZSV by marker assisted selection （MAS） in wheat breeding programs.

Mapping QTLs for Drought Tolerance at Seedling Stage in Rice Using Doubled Haploid Population

QTLs for drought tolerance at the rice seedling stage were analyzed using a doubled haploid （DH） population consisted of 251 lines from the cross between a japonica parent Maybelle and an indica parent Baiyeqiu. A genetic linkage map with 226 SSR marker loci was constructed. Single-locus analysis following composite interval mapping （CIM） detected a total of five QTLs located on five different chromosomes of rice. Four QTLs were also detected following two-locus analysis, resolving two pairs of epistatic QTLs （E-QTI_s） with positive and additive genetic effects. The results indicated that the alleles from the parent Baiyeqiu contributed DH population to improve drought tolerance at the seedling stage.

Gametic selection in a doubled haploid population derived from anther culture of indica/japonica hybrid of Oryza Sativa L.

It is one of the key problems for application ofanther culture in hybrid breeding, geneticanalysis, and molecular mapping whether thedoubled haploid (DH) population derived fromanther culture of rice crosses represents a ran-dom array of the microspore population, i.e.whether gametic selection occurs in androgene-sis. A DH population including 132 lines de-

Yield Stability and Adaptability of Elite Doubled Haploid (DH) Rice Genotypes in Different Locations in Central Clay Plains of the Sudan

Stability analysis of grain yield is an efficient tool for the selection of varieties adapted to fairly wide cultivation zone. A study aims to determine adaptability and stability for grain yield and agronomic performance of 10 elite Doubled Haploid (DH) rice genotypes generated through another culture technique, along with local cultivar Umgar as a check under irrigated lowland conditions in three different environments in the central clay plains of the Sudan;during two cropping seasons 2019 and 2020. The trials were conducted in a randomized complete block design with three replications. Grain yield stability was studied, using the Additive Main effect and Multiplicative Interaction (AMMI) model. Bi-plots were developed following GGE bi-plot methodology over the six environments. The combined analysis of variance revealed significant to highly significant (P < 0.001) effects of genotypes, environments, and genotype by environment interaction. Moreover, the analysis of variance based on AMMI indicated significant genotypes, environments, and GE interaction with a total variation of 43.16%, 44.26% and 12.58% respectively. A biplot-AMMI analysis and yield stability index incorporating the AMMI stability value and yield in a single non-parametric index were used to identify the genotypes with the highest and stable yield. The overall mean for grain yield was 5.69 t/ha, with a range of 5.77 to 4.12 t/ha. The genotypes k150147, k150870, k150032, k150912, and k150307 out yielded the local check Umgar by 40%, 38%, 34%, 34% and 33% respectively. The most genotypes were mid-early maturities which were harvested at 107, 113, 108, 109 and 112 days after sowing, respectively. These five genotypes also showed grain yield stability along with their high mean yield performance according to the AMMI analysis and widely adaptable to the tested locations. Therefore, the five out yielding genotypes will be considered for cultivation under irrigated system condition in central clay in the Sudan.

Research and Breeding Application Progress of the Technique of Producing Double Haploid of Wheat by Wide Hybridization between Wheat and Maize

The technique of producing doublehaploid of wheat by distant hybridization between wheat and maize has characterized with better inducing effect, shorter in- ducing period, easy operation, and so on. At present, it is the most efficient and has great potential of application in breeding of wheat. This article reviewed princi- ple and production process of the technique, research situation of the three key in- dicators of the technology（embryo rate, seedling rate and success rate of doubling）in recent years, and application of the technology in breeding, genetics, germplasm improvement of wheat. At last, both the achievements and the direction of further improvement and development of the technology in our program were discussed.

Resistance Performance to Whitebacked Planthopper in Different Phenotypes of Japonica/Indica Doubled Haploid Rice Lines

Field performance of whitebacked planthopper （WBPH）-resistance of four phenotypes was evaluated in Chunjiang 06 （C J-06） / TN1 DH rice lines, which were expressed by different combinations of sucking inhibitory and ovicidal traits inherited independently from C J-06. WBPH established the highest populations in susceptible DH lines that had neither sucking inhibitorynor ovicidal resistance. Both immigration and subsequent population levels were kept below the damage-causing density in the sucking inhibitory DH lines even under a WBPH outbreak. WBPH could not build up populations in the DH lines having both the sucking inhibitory and ovicidal resistance. Although WBPH immigrated preferentially to non-sucking inhibitory DH lines with ovicidal resistance, subsequent population buildup was significantly suppressed. It was concluded that the differential performance to WBPH-resistance in CJ-06 / TN1 DH lines was primarily due to the sucking inhibitory trait, and complementarity to the ovicidal trait.

普通小麦(浙农林12×CASL7AS)DH系株高的分析

小麦株高与植株倒伏密切相关,影响小麦产量。为挖掘小麦株高QTL位点和候选基因,从遗传和分子水平上解释株高分子机理,本研究将野生二粒小麦染色体臂置换系CASL7AS为母本,课题组自选株系浙农林12(简称ZNL12)为父本,杂交F 1经过花培得到178个DH群体。通过4年2个种植点株高数据,利用55K SNP芯片构建高密度遗传图谱,对小麦株高性状进行QTL分析。该遗传连锁图谱包含3655个SNP标记,长度为4738.45 cM,标记间平均遗传距离为1.30 cM,覆盖了小麦21条染色体。其中,A、B、D染色体组分别含有标记1466、1492和697个。QTL分析共检测出53个QTL位点,分布于1A、3A、5A、7A、1B、3B、4B、5B、7B、2D、4D、6D和7D染色体上,可解释2.80%~38.50%表型变异。其中,稳定主效QTL有2个,分别为QPh.zafu.4B-1和QPh.zafu.4D-1,其贡献率分别为25.18%~38.50%和20.67%。经细胞生物学分析,基因型为(0,2)矮秆植株胚芽鞘的表皮细胞长度显著短于基因型为(2,0)的高秆植株,基因型为(0,0)、(2,2)中间型的植株胚芽鞘表皮细胞长度无显著差异,由此推测小麦株高的变异由细胞长度因素决定。QPh.zafu.4B-1候选基因Rht-B1b基因编码区矮秆株系,第904核苷酸处发生碱基“C-T”的突变,形成终止密码子(CAG-TAG);而QPh.zafu.4D-1候选基因Rht-D1b基因编码区矮秆株系,第781核苷酸处碱基突变“G-T”,编码氨基酸中缬氨酸变为天冬氨酸。结果为小麦株高候选基因的筛选和QTL定位提供了优异的遗传位点和候选基因,未来可用于小麦抗倒伏相关的分子标记辅助育种。

芥蓝游离小孢子培养技术优化及其在育种中的应用

以16个芥蓝品种(F1)为供体材料,比较分析了优化花蕾的消毒、小孢子的游离释放以及过滤等环节对芥蓝游离小孢子培养效果的影响。结果表明,16份参试芥蓝供体材料都具有胚胎发生能力,并全部获得了不同类型的再生植株,实现了较大规模、批量地获得芥蓝双单倍体(DH)。经田间综合评价筛选,应用DH系育成了不同熟性的芥蓝一代杂种20M4芥蓝、21M2芥蓝和22M11芥蓝。

Linkage Map Construction and Quantitative Trait Loci Analysis for Bolting Based on a Double Haploid Population of Brassica rapa

Early bolting of Chinese cabbage （Brassica rapa L.） during spring cultivation often has detrimental effects on the yield and quality of the harvested products. Breeding late bolting varieties is a major objective of Chinese cabbage breeding programs. In order to analyze the genetic basis of bolting traits, a genetic map of B. rapa was constructed based on amplified fragment-length poiymorphism （AFLP）, sequence-related amplified poiymorphism （SRAP）, simple sequence repeat （SSR）, random amplification of polymorphic DNA （RAPD）, and isozyme markers. Marker analysis was carried out on 81 double haploid （DH） lines obtained by microspore culture from F1 progeny of two homozygous parents： B. rapa L. ssp. pekinensis （BY） （an extra-early bolting Chinese cabbage line） and B. rapa L. ssp. rapifera （MM） （an extra-late bolting European turnip line）. A total of 326 markers including 130 AFLPs, 123 SRAPs, 16 SSRs, 43 RAPDs and 14 isozymes were used to construct a linkage map with 10 linkage groups covering 882 cM with an average distance of 2.71 cM between loci. The bolting trait of each DH line was evaluated by the bolting index under controlled conditions. Quantitative trait loci （QTL） analysis was conducted using multiple QTL model mapping with MapQTL5.0 software. Eight QTLs controlling bolting resistance were identified. These QTLs, accounting for 14.1% to 25.2% of the phenotypic variation with positive additive effects, were distributed into three linkage groups. These results provide useful information for molecular marker-assisted selection of late bolting traits in Chinese cabbage breeding programs.

Mapping QTLs for heading synchrony in a doubled haploid population of rice in two environments

Simultaneous heading of plants within the same rice variety, also refer to heading synchrony, is an important factor that affects simultaneous ripening of the variety. Understanding of the genetic basis of heading synchrony may contribute to molecular breeding of rice with simultaneous heading and ripening. In the present study, a doubled haploid （DH） population, derived from a cross between Chunjiang 06 and TN1 was used to analyze quantitative trait locus （QTL） for heading synchrony related traits, i.e., early heading date （EHD）, late heading date （LHD）, heading asynchrony （HAS）, and tiller number （PN）. A total of 19 QTLs for four traits distributed on nine chromosomes were detected in two environments. One QTL, qHAS-8 for HAS, explained 27.7% of the phenotypic variation, co-located with the QTLs for EHD and LHD, but it was only significant under long-day conditions in Hangzhou, China. The other three QTLs, qHAS-6, qHAS-9, and qHAS-10, were identified under short-day conditions in Hainan, China, each of which explained about 11% of the phenotypic variation. Two of them, qHAS-6 and qHAS-9, were co-located with the QTLs for EHD and LHD. Two QTLs, qPN-4 and qPN-5 for PN, were detected in Hangzhou, and qPN-5 was also detected in Hainan. However, none of them was co-located with QTLs for EHD, LHD, and HAS, suggesting that PN and HAS were controlled by different genetic factors. The results of this study can be useful in marker assisted breeding for improvement of heading synchrony.

Identification of Quantitative Trait Loci for Grain Appearance and Milling Quality Using a Doubled-Haploid Rice Population

Improving grain quality, which is composed primarily of the appearance of the grain and its cooking and milling attributes, is a major objective of many rice-producing areas in China. In the present study, we conducted a marker-based genetic analysis of the appearance and milling quality of rice （Oryza sativa L.） grains using a doubled-haploid population derived from a cross between the indica inbred Zhenshan 97 strain and the japonica inbred Wuyujing 2 strain. Quantitative trait locus （QTL） analysis using a mixed linear model approach revealed that the traits investigated were affected by one to seven QTLs that individually explained 4.0%-30.7% of the phenotypic variation. Cumulatively, the QTL for each trait explained from 12.9% to 61.4% of the phenotypic variation. Some QTLs tended to have a pleiotropic or location-linked association as a cause of the observed phenotypic correlations between different traits. Improvement of the characteristics of grain appearance and grain weight, as well as an improvement in the milling quality of rice grains, would be expected by a recombination of different QTLs using marker-assisted selection.

利用DH群体动态检测水稻抗褐飞虱数量性状基因位点

利用籼粳交珍汕97/武育粳2号F1花培获得的190个双单倍体群体(doubled-haploid population,DH系)及其构建的179个SSR分子标记遗传图谱,通过对DH系群体苗期重复接虫试验和2个不同时期对褐飞虱危害程度进行动态调查,并应用Mapmaker/exp Version 3.0和Windows QTL Cartographer V 2.0对水稻抗褐飞虱数量性状基因位点(quantativetraitlocus,QTL)进行动态检测和遗传效应分析.结果表明,在苗期对褐飞虱抗性的检测中,共检测到6个抗性QTL,分别位于第2、3、4、8和10染色体上,各QTL的LOD值分别为2.22～4.64,贡献率为5.04%～13.73%,第3染色体和第4染色体上各有1个QTL的加性效应为正值,表明来自于亲本武育粳2号的这2个位点的等位基因可以提高水稻对褐飞虱的抗性,其余4个QTL的加性效应均为负值,表明来自于亲本珍汕97的这些位点的等位基因可以提高水稻对褐飞虱的抗性.

望水白和苏麦3号构建的DH群体赤霉病抗性比较

利用抗病品种望水白和苏麦3号分别与感病品种Alondra’s杂交,F1花药培养诱导单倍体,经染色体加倍构建了2个DH群体。2001-2003年连续3年在赤霉病常发重病区福建省建阳市进行了赤霉病抗性鉴定。通过聚类分析把2个DH群体分成稳定的抗病DH系、稳定的感病DH系、稳定的中抗DH系以及抗性不稳定的DH系4类,4种类型在2个DH群体中的比例相似。望水白/Alondra’sDH群体赤霉病抗性比苏麦3号/Alondra’sDH群体强,但其抗性不如苏麦3号DH群体稳定。遗传模型分析表明在望水白/Alondra’s和苏麦3号/Alondra’s2组合各自至少存在2对主基因差异。主基因可以解释70%以上表型抗性,多基因可以解释15%左右的表型抗性。由于3年的发病条件有差异,基因间表现出相互独立、加性上位、等加性和累加作用等不同互作模式。

小麦DH群体数量性状的遗传分析

为了研究小麦数量性状的遗传规律,利用郑州8761×川育35050杂交组合的F1花药培养获得的DH群体对小麦株高、穗下节长、穗长等性状进行了遗传分析。结果表明,株高、穗下节长、穗长、单株穗数、结实小穗数、不育小穗数、总小穗数、穗粒数这8个性状的遗传力分别为92%、83%、79%、30%、48%、48%、53%和47%;控制各性状的最少基因对数分别约为5、6、10、12、11、11、11和9对。各性状的偏度系数和峰度系数的估算结果表明,控制株高、穗下节长、穗长、结实小穗数、总小穗数和穗粒数的基因间无互作,控制单株穗数和不育小穗数的基因间存在互补作用。

大麦DH群体若干数量性状的遗传分析

以大麦品种Y erong与F rank lin进行杂交,对其F1代通过花药培养构建DH群体,利用该群体估测株高(PH)、穗长(SL)、穗下节间长(TF IL)、单株穗数(SPP)、主穗粒数(GPS)、单株秆重(SW PP)、单株粒重(GW PP)、单株粒数(GPP)和千粒重(W TG)9个数量性状的遗传力和基因数目,并通过估算各性状的偏度和峰度系数分析影响各性状的基因间的作用方式。结果表明:①上述各性状的遗传力分别为0.79、0.67、0.54、0.31、0.90、0.30、0.20、0.41和0.77;控制各性状的基因数目分别为8.76、9.30、5.30、24.67、3.63、14.59、14.90、12.36和6.81对。遗传力(Y)依基因对数(X)线性回归方程为Y=-0.029 6X+0.878 5(r=-0.780 0*),回归系数-0.029 6与0的差异显著(t=-3.30*,d f=7),表明多基因性状易受环境影响,同时也佐证了多基因遗传学说。②控制SL、TF IL、GW PP和W TG的多基因间存在互补作用,控制PH、SW PP和SPP各性状的多基因间存在重叠作用,而控制GPS和GPP的多基因间无互作效应。

同一基础材料的玉米双单倍体(DH)系配合力的分析

利用高频玉米单倍体诱导系,对选系材料M35/F35的F1代进行诱导获得大量单倍体籽粒,经自然加倍获得一批纯合双单倍体(DH)系,用测验系京24组配杂交组合分析DH系的配合力表现。结果表明,来源于同一基础材料的不同DH系之间,配合力差异较大,部分DH系的配合力比亲本增加,说明采用单倍体育种可以选育到高配合力的优良玉米自交系,组配出优良品种,从而加快育种进程,应用前景广阔。

小麦DH群体茎秆可溶性碳水化合物含量相关数量性状的遗传分析

以小麦DH群体（早选10号×鲁麦14）的150个株系及其亲本为研究材料，分析雨养和正常灌溉条件下与茎秆可溶性碳水化合物含量相关的8个性状的相关性、遗传力、控制性状的基因数日及基因作用方式。结果表明，在两种水分条件下，DH群体各性状的表型值多数介于双亲之间，且出现超亲分离，变异系数在6．96％～65．72％之间。DH群体及其亲本各性状表型值普遍表现为雨养条件下的高于正常灌溉的。初花期、花后14d和成熟期的茎秆可溶性碳水化合物含量（SSCf、SSCf14和SSCm）之间相关性较低，而SSCf14与其他多数性状在两种水分条件下分别表现极显著或显著的正相关。DH群体各性状的遗传力和调控性状的基因数目在雨养和灌溉条件下有较大差异。在雨养条件下的遗传力，SSCf最高，为0．49，而籽粒灌浆效率（GFE）最低，为0．11，其他6个性状介于二者之间；而灌溉条件下的遗传力，GFE最高，为0．65，SSCf最低，为0．51。在雨养条件下，控制GFE、可溶性碳水化合物的积累效率（AESSC）、SSCfl4和SSCm的基因数日较多，分别为27、26、22和20对，控制可溶性碳水化合物转运效率（RESSC）的基因数目最少，只有9对；而灌溉条件下，控制SSCfl4的基因数目最多（19对），控制RESSC的基因数目最少（4对）。两种条件下控制SSCm和灌溉条件下控制SSCf14的基因间存在互补作用，其他性状的多基因间未检测到互作效应；两种条件下的RESSC和GFE，以及灌溉条件下的AESSC基因间存在重叠作用。说明小麦茎秆可溶性碳水化合物含量相关性状的遗传基础是复杂的，在不同发育时期、不同水分条件下控制这些性状的基因可能具有不同的表达模式。