利用籼粳回交群体分析水稻粒形性状相关QTLs

水稻谷粒的外观性状对稻米外观品质存在重要的影响。该研究利用SSR标记,以回交群体Balilla／NTH∥Balilla为作图群体,构建了水稻12条染色体的连锁图,该遗传图谱包括：108个分子标记,平均图距为11．9cM。以构建的遗传图谱为基础,采用区间作图法对谷粒外观性状,包括粒长、粒宽和粒形进行了数量性状基因(QTL)定位。结果表明,粒长、粒宽和粒形在回交群体中均呈近似的正态分布,表现出典型的数量性状特征。QTL定位结果表明,第12染色体上RM101-RM270区间内存在一个与粒长性状相关的QTL,(qGL-12),加性效应约为0．26mm,贡献率为16．7％。在第2和第3染色体上RM154-RM211和RM257-RM175区问内,分别检测到qGW-2和qGW-3两个位点与粒宽性状有关,加性效应为分别为-0．10mm和-0．12mm,贡献率分别为11．5％和16．6％。对于粒形性状,共检测到3个QTLs,qLW-2、qLW-6和qLW-7,分别位于第2、6和7染色体上。其中qLW-2和qLW-7的加性效应分别约为0．09和0．10,两个QTLs分别可解释表型变异的12．7％和18．3％;而qLW-6的加性效应约为-0．13,可解释粒形变异的11．5％。文中还讨论了粒形和稻米外观品质同时改良的可能性。     

利用水稻重组自交系群体定位谷粒外观性状的数量性状基因

用区间作图和混合线性模型的复合区间作图两种方法,对水稻（Oryza sativa L）珍汕97和明恢63组合的重组自交系群体的谷粒外观性状－粒长,粒宽和粒形进行了数量性状基因（QTL）定位,用区间作图法在LOD≥2．4水平上（近拟于a=0.005),1998年对粒长,粒宽和粒形分别检测到6,2放2个QTLs,1999年对以上3个性状分别检测到3,2和2个QTLs,其中7个QTLs在两年均检测到,位于第3染色体RG393－C1087区间的QTL效应大,同时影响粒长和粒形,两年贡献分别为57．5％,61．4％和26．7％,29．9％,位于第5染色体RG360－C734B区间的QTL效应大,同时影响粒宽和粒形,两年贡献率分别为44．2％,53．2％和32．1％和36．0％,用混合线性模型的复合区间作图法在P＝0．005水平上,对粒长,粒宽和粒形分别检测到8,5和5个QTLs,共解释各自性状变异的58．81％,44．75％和57．47％,只检测到1个QTL与环境之间存在的显互作。     

稻米外观品质性状遗传与分子定位研究进展

稻米外观品质主要是指稻米的粒形、垩白、透明度和籽粒色泽等,它不仅直接影响到人们的喜好,还与其他品质性状诸如蒸煮食用、加工等密切相关。因此,外观品质对稻米的商品价值有着十分重要的影响。本文从经典遗传与现代分子生物学两个方面对稻米主要外观品质的遗传研究进展进行了较全面的综述,包括粒长、粒宽、长宽比、粒厚、垩白、透明度和籽粒色泽等。综合近年来的遗传研究结果发现,大多数稻米外观品质性状都是由数量基因控制的。利用分子标记技术已将控制外观品质的QTL（qualitative trait locus）定位在不同的染色体上,为下一步的稻米外观品质改良提供了有利条件。     

5个籼稻背景的高代回交置换系的置换片段分析

以轮回亲本籼稻品种9311（Oryza sativassp.indica‘Yangdao6’）为对照,选用132个亲本间有多态性的SSR标记,对以粳稻品种日本晴（Oryzasativassp.japonica‘Nipponbare’）为供体的5个高代回交置换系的农艺性状及置换片段进行分析。5个置换系在粒长、粒宽、千粒重、剑叶长、株高及落粒性等方面与籼稻品种9311之间有极显著差异,其余性状与籼稻品种9311间的差异不显著;从置换系中检测出8个置换片段,总长度为236.0cM,平均长度为29.5cM;从置换片段上检出包括2个千粒重、1个粒长、1个粒宽、1个剑叶长、1个株高和1个落粒性共7个QTLs,分别分布在水稻第1、3、5、6和第10染色体上。其中,第1染色体上控制剑叶长的QTL和第6染色体上控制株高的QTL可能是新发现的QTLs。实验结果进一步丰富了置换系群体的数量和质量,也为QTLs的精细定位及分子设计育种奠定了基础。     

水稻垩白性状遗传育种研究进展

水稻(Oryza sativa L.)垩白包括垩白率和垩白度,是重要的外观品质之一,对其他品质性状也有重要影响,阐明水稻垩白性状的遗传机制十分重要。近年来随着水稻功能基因组学和分子标记技术的发展,越来越多的垩白基因获得了克隆。本文综述了水稻垩白的评价指标、与其他品质性状的关系、遗传基础、垩白QTL定位和垩白基因克隆,并提出了借助显性核不育系进行轮回选择改良水稻垩白的分子育种策略,以期为水稻垩白性状的分子改良提供参考和借鉴。     

籼稻稻米碾磨与外观品质性状的QTL定位

文章利用籼籼交组合特青/IRBB衍生的重组自交系群体, 在2个环境下对稻米碾磨品质和外观品质进行QTL定位。共计检测到控制稻米碾磨品质的QTL 12个和控制外观品质的QTL 18个, 包括糙米率8个、精米率2个、整精米率2个、粒长7个、粒宽5个和长宽比6个, 这些QTL分布于除第4和12染色体外的其他10条染色体上。其中, 第3染色体涵盖粒形基因GS3的区域对粒长、长宽比、糙米率和整精米率具有较大效应, 其献率分别为56.71%、42.23%、10.05%和4.91%; 第5染色体涵盖粒宽基因GW5的区域对粒宽、长宽比、糙米率和精米率具有较大效应, 表型变异贡献率分别为59.51%、36.68%、19.51%和4.56%。此外, 第6染色体涵盖直链淀粉含量基因Wx的区域对糙米率和精米率具有较小效应。GS3和GW5对糙米率和粒形具有重要作用。     

5 个籼稻背景的高代回交置换系的置换片段分析

以轮回亲本籼稻品种9311(Oryz a sativa ssp. indica ‘Yangdao 6’)为对照, 选用132个亲本间有多态性的SSR标记, 对以粳稻品种日本晴(Oryz a sativa ssp. japonica 'Nipponbare’)为供体的5个高代回交置换系的农艺性状及置换片段进行分析。5个置换系在粒长、粒宽、千粒重、剑叶长、株高及落粒性等方面与籼稻品种9311之间有极显著差异, 其余性状与籼稻品种9311间的差异不显著; 从置换系中检测出8个置换片段, 总长度为236.0 cM, 平均长度为29.5 cM; 从置换片段上检出包括2个千粒重、1个粒长、1个粒宽、1个剑叶长、1个株高和1个落粒性共7个QTLs , 分别分布在水稻第1、3、5、6和第10染色体上。其中, 第1染色体上控制剑叶长的QTL和第6染色体上控制株高的QTL可能是新发现的QTLs。实验结果进一步丰富了置换系群体的数量和质量, 也为QTLs 的精细定位及分子设计育种奠定了基础。     

几个优良籼稻亲本品质性状的配合力和杂种优势分析

以3个不育系和10个恢复系为材料,采用NCII交配设计研究10个米质性状的配合力和杂种优势。结果表明：①大多数品质性状的量值介于双亲之间,除粒重表现一定的超亲优势、垩白度和粒宽表现一定的正向平均优势外,其他品质性状优势不明显。②杂种稻米的品质性状主要受不育系或恢复系的影响,其中粒长、粒宽和直链淀粉含量3个性状,不育系的影响要高于恢复系;而对于整精米率、粒重、垩白率、垩白度和糊化温度,则恢复系的影响要高于不育系。③就优质育种的利用价值而言,不育系以广占63-4S为好,恢复系以扬稻6号为好,R527、镇恢084次之,用上述亲本选配的杂交组合米质较好;恢复系特青、盐恢559表现为一般配合力效应低,特殊配合力方差小,优质育种利用价值不大。     

水稻穗部性状的QTL与环境互作分析

分别在两年收集珍汕97／明恢63的重组自交系群体的表现数据,运用混合线性模型的QTL定位方法,联合分析穗部5个性状的QTLs7及QTL与环境互作关系。每穗颖花数、每穗实粒数、结实率、穗长和穗着密度分别检测到10、3、6、8和7个QTLs分别解释各性状变异的29．13％、19．2％、29．46％、26．39％和35．76％。对于同一性状,高值亲本和低值亲本中均存在增效和减效QTL。相关性状QTL的位置表现相同或相似,高值亲本和低值亲本中均存在增效和减效QTL。相关性状QTL的位置表现相同或相似,成族分布。1个穗长QTL,2个每穗颖花数QTL3,3个结实率QTLs表现与环境显著互作,QTL与环境互作效应的贡献率比相应的QTL贡献率略大。遗传力稍高的每穗实粒数和穗着粒密度的DQTL与环境不互作。     

水稻粒形性状的遗传及相关基因定位与克隆研究进展

作物育种的首要目标是提高产量。水稻粒形是与水稻产量性状直接相关,与品质性状存在着密切关系的数量性状,其评价指标主要是粒长、粒宽、粒厚、长/宽和长/厚。近年来,水稻粒形的数量遗传研究取得了重要进展,并成功定位克隆了一批控制水稻粒形的基因。文章综述了水稻粒形的经典遗传研究、QTL定位、粒形基因的克隆和功能分析以及在水稻超高产育种中的利用。     

Dissecting the Genetic Basis of Extremely Large Grain Shape in Rice Cultivar ＇JZ1560＇

Rice grain shape,grain length(GL),width(GW),thickness(GT)and length-to-width ratio(LWR),are usually controlled by multiple quantitative trait locus(QTL).To elucidate the genetic basis of extremely large grain shape,QTL analysis was performed using an F2 population derived from a cross between a japonica cultivar ’JZ1560’(extremely large grain)and a contrasting indica cultivar ’FAZ1’(small grain).A total number of 24 QTLs were detected on seven different chromosomes.QTLs for GL,GW,GT and LWR explained 11.6%,95.62%,91.5%and 89.9%of total phenotypic variation,respectively.Many QTLs pleiotropically controlled different grain traits,contributing complex traits correlation.GW2 and qSW5/GW5,which have been cloned previously to control GW,showed similar chromosomal locations with qGW2-I/qGT2-I/qLWR2-2 and qGW5-2/qLWR5-l and should be the right candidate genes.Plants pyramiding GW2 and qSW5/GW5 showed a significant increase in GW compared with those carrying one of the two major QTLs.Furthermore,no significant QTL interaction was observed between GW2 and qSW5/GW5.These results suggested that GW2 and qSW5/GW5 might work in independent pathways to regulate grain traits.’JZ1560’ alleles underlying all QTLs contributed an increase in GW and GT and the accumulation of additive effects generates the extremely large grain shape in ’JZ1560’.     

Dissecting the genetic basis for the effect of rice chalkiness, amylose content, protein content, and rapid viscosity analyzer profile characteristics on the eating quality of cooked rice using the chromosome segment substitution line population across eight environments

Quantitative trait locus (QTL) mapping and stability analysis were carried out for 16 rice (Oryza sativa L.) quality traits across eight environments, by using a set of chromosome segment substitution lines with 'Asominori' as genetic background. The 16 quality traits include percentage of grain with chalkiness (PGWC), area of chalky endosperm (ACE), amylose content (AC), protein content (PC), peak viscosity, hot paste viscosity, cool paste viscosity, breakdown viscosity (BDV), setback viscosity (SBV), consistency viscosity, cooked-rice luster (LT), scent, tenderness (TD), viscosity, elasticity, and the integrated values of organleptic evaluation (IVOE). A total of 132 additive effect QTLs are detected for the 16 quality straits in the eight environments. Among these QTLs, 56 loci were detected repeatedly in at least three environments. Interestingly, several QTL clusters were observed for multiple quality traits. Especially, one QTL cluster near the G1149 marker on chromosome 8 includes nine QTLs: qPGWC-8, qACE-8, qAC-8, qPC-8a, qBDV-8a, qSBV-8b, qLT-8a, qTD-8a, and qIVOE-8a, which control PGWC, ACE, AC, PC, BDV, SBV, LT, TD, and IVOE, respectively. Moreover, this QTL cluster shows high stability and repeatability in all eight environments. In addition, one QTL cluster was located near the C2340 marker on chromosome 1 and another was detected near the XNpb67 marker on chromosome 2; each cluster contained five loci. Near the C563 marker on chromosome 3, one QTL cluster with four loci was found. Also, there were nine QTL clusters that each had two or three loci; however, their repeatability in different environments was relatively lower, and the genetic contribution rate was relatively smaller. Considering the correlations among all of the 16 quality traits with QTL cluster distributions, we can conclude that the stable and major QTL cluster on chromosome 8 is the main genetic basis for the effect of rice chalkiness, AC, PC, and rapid viscosity analyzer profile characteristics on the eating quality of cooked rice. Consequently, this QTL cluster is a novel gene resource for controlling rice high-quality traits and should be of great significance for research on formation mechanism and molecule improvement of rice quality.     

Multi-Environmental Genetic Analysis of Grain Size Traits Based on Chromosome Segment Substitution Line in Rice (<i>Oryza sativa</i> L.)

Grain size traits are critical agronomic traits which directly determine grain yield, but the genetic bases of these traits are still not well understood. In this study, a total of 154 chromosome segment substitution lines (CSSLs) population derived from a cross between a japonica variety Koshihikari and an indica variety Nona Bokra was used to investigate grain length (GL), grain width (GW), length-width ratio (LWR), grain perimeter (GP), grain area (GA), and thousand grain weight (TGW) under four environments. QTL mapping analysis of six grain size traits was performed by QTL IciMapping 4.2 with an inclusive composite interval mapping (ICIM) model. A total of 64 QTLs were identified for these traits, which mapped to chromosomes 1, 2, 3, 4, 6, 7, 8, 10, 11, and 12 and accounted for 1.6%–27.1% of the total phenotypic variations. Among these QTLs, thirty-six loci were novel and seven QTLs were identified under four environments. One locus containing the known grain size gene, qGL3/GL3.1/OsPPKL1, also have been found. Moreover, five pairs of digenic epistatic interactions were identified except for GL and GP. These findings will facilitate fine mapping of the candidate gene and QTL pyramiding to genetically improve grain yield in rice.     

A haplotype inference algorithm for trios based on deterministic sampling

Background

The three-dimensional shape of grain, measured as grain length, width, and thickness (GL, GW, and GT), is one of the most important components of grain appearance in rice. Determining the genetic basis of variations in grain shape could facilitate efficient improvements in grain appearance. In this study, an F_7:8 recombinant inbred line population (RIL) derived from a cross between indica and japonica cultivars (Nanyangzhan and Chuan7) contrasting in grain size was used for quantitative trait locus (QTL) mapping. A genetic linkage map was constructed with 164 simple sequence repeat (SSR) markers. The major aim of this study was to detect a QTL for grain shape and to fine map a minor QTL, qGL7.

Results

Four QTLs for GL were detected on chromosomes 3 and 7, and 10 QTLs for GW and 9 QTLs for GT were identified on chromosomes 2, 3, 5, 7, 9 and 10, respectively. A total of 28 QTLs were identified, of which several are reported for the first time; four major QTLs and six minor QTLs for grain shape were also commonly detected in both years. The minor QTL, qGL7, exhibited pleiotropic effects on GL, GW, GT, 1000-grain weight (TGW), and spikelets per panicle (SPP) and was further validated in a near isogenic F₂ population (NIL-F₂). Finally, qGL7 was narrowed down to an interval between InDel marker RID711 and SSR marker RM6389, covering a 258-kb region in the Nipponbare genome, and cosegregated with InDel markers RID710 and RID76.

Conclusion

Materials with very different phenotypes were used to develop mapping populations to detect QTLs because of their complex genetic background. Progeny tests proved that the minor QTL, qGL7, could display a single mendelian characteristic. Therefore, we suggested that minor QTLs for traits with high heritability could be isolated using a map-based cloning strategy in a large NIL-F₂ population. In addition, combinations of different QTLs produced diverse grain shapes, which provide the ability to breed more varieties of rice to satisfy consumer preferences.     

Development and high-throughput genotyping of substitution lines carring the chromosome segments of indica 9311 in the background of japonica Nipponbare

Chromosome segment substitution lines (CSSLs) are useful for the precise mapping of quartitative trait loci (QTLs) and dissection of the genetic basis of complex traits.In this study,two whole-genome sequenced rice cultivars,the japonica Nipponbare and indica 9311 were used as recipient and dtonor,respectively.A population with 57 CSSLs was developed after crossing and back-crossing assisted by mo lecular rnarkers,and genotypes were identified using a high-throughput resequencing strategy,Detailed graphical genotypes of 38 lines were constructed based on resequencing data.These CSSLs had a total of 95 substituted segments derived from indica 9311,with an average of about 2.5 segments pet CSSL and eight segments per chromosome,and covered about 87.4％ of the rice whole genome.A multiple linear regression QTL analysis mapped four QTLs for 1000-grain weight.The largest-effect QTL was located in a region on chromosome 5 that contained a cloned major QTL GW5/qSW5 for grain size in rice.These CSSLs with a background of Nipponbare may provide powerful tools for future whole-genome discovery and functional study of essential genes/QTLs in rice,and offer ideal materials and foundations for japonica breeding.     

Identification of Genomic Regions and the Isoamylase Gene for Reduced Grain Chalkiness in Rice

Grain chalkiness is an important grain quality related to starch granules in the endosperm. A high percentage of grain chalkiness is a major problem because it diminishes grain quality in rice. Here, we report quantitative trait loci identification for grain chalkiness using high-throughput single nucleotide polymorphism genotyping of a chromosomal segment substitution line population in which each line carried one or a few introduced japonica cultivar Nipponbare segments in the genetic background of the indica cultivar ZS97. Ten quantitative trait loci regions were commonly identified for the percentage of grain chalkiness and the degree of endosperm chalkiness. The allelic effects at nine of these quantitative trait loci reduced grain chalkiness. Furthermore, a quantitative trait locus (qPGC8-2) on chromosome 8 was validated in a chromosomal segment substitution line–derived segregation population, and had a stable effect on chalkiness in a multiple-environment evaluation of the near-isogenic lines. Residing on the qPGC8-2 region, the isoamylase gene (ISA1) was preferentially expressed in the endosperm and revealed some nucleotide polymorphisms between two varieties, Nipponbare and ZS97. Transgenic lines with suppression of ISA1 by RNA interference produced grains with 20% more chalkiness than the control. The results support that the gene may underlie qPGC8-2 for grain chalkiness. The multiple-environment trials of the near-isogenic lines also show that combination of the favorable alleles such as the ISA1 gene for low chalkiness and the GS3 gene for long grains considerably improved grain quality of ZS97, which proves useful for grain quality improvement in rice breeding programs.     

QTL analysis of percentage of grains with chalkiness in Japonica rice (Oryza sativa)

Appearance quality of rice grains is a major problem for rice production in many areas of the world. We conducted a molecular marker-based genetic analysis of percentage of grains with chalkiness (PGWC), which is a determining factor for appearance quality; it strongly affects milling, eating and cooking quality. An F(8) recombinant inbred line population, which consists of 261 lines derived from a cross between Koshihikari (Japonica) and C602 (Japonica), was used for QTL mapping. Three QTLs related to PGWC were detected on chromosomes 5, 8 and 10, together explaining 50.8% of the genetic variation. The 'Koshihikari' alleles qJPGC-5, qJPGC-8 and the 'C602' alleles at qJPGC-10 were associated with reduced PGWC. The QTL contributions to phenotypic variance were 18.2, 9.6 and 25%, respectively. These QTL markers for PGWC could be used for developing improved varieties.     

与稻米高垩白率相关的qPGWC-9的生理功能

利用携带qPGWC-9目标区段而其他置换片段上不带有与垩白率相关QTL(quantitative trait locus)的高垩白染色体片段置换系(chromosomal segment substitution line,CSSL)AIS82,以其轮回亲本Asominori(低垩白)为对照,从源库关系角度探讨qPGWC-9的生理功能。结果发现,籽粒灌浆期AIS82与Asominori的剑叶净光合速率没有显著差异,说明光合作用强弱不是AIS82高垩白率产生的直接原因。灌浆前期AIS82的淀粉合成关键酶活性显著高于Asominori,但中后期AIS82酶活下降快,整个灌浆期变化幅度相对较大,推测qPGWC-9主要影响了淀粉合成关键酶活性的动态变化,从而决定了高垩白表型。     

Identification of a stable quantitative trait locus for percentage grains with white chalkiness in rice (Oryza sativa)

High chalkiness is a major problem in many rice-producing areas of the world, especially in hybrid rice (Oryza sativa L.) in China. We previously showed a major quantitative trait locus for the percentage of grains with white chalkiness (QTLqPGWC-8) in the interval G1149-R727 on chromosome 8 using a chromosome segment substitution line (CSSL). Here, we selected the line-CSSL50 harboring the QTLqPGWC-8 allele from the CSSLs derived from a cross between Asominori (as a recurrent parent) and IR24 (as a donor parent), which had higher percentage chalkiness, markedly different from that of Asominori. There were also significant differences in starch granules, appearance of amylose content (AAC) and milling qualities between Asominori and CSSL50, but not in grain size or thousand grain weight (TGW). The BC(4) F(2) and BC(4) F(3) populations from a cross between CSSL50 and Asominori were used for fine mapping of qPGWC-8. We narrowed down the location of this QTL to a 142 kb region between Indel markers 8G-7 and 8G-9. QTLqPGWC-8 accounted for 50.9% of the difference in PGWC between the parents. The markers tightly linked to qPGWC-8 should facilitate cloning of the gene underlying this QTL and will be of value for marker-assisted selection in breeding rice varieties with better grain quality.