水稻垩白相关基因及其功能的研究进展

水稻是我国最重要的粮食作物之一,高垩白性状严重降低了稻米的食用与加工品质。水稻的垩白性状主要由遗传控制,并受环境因素的影响。降低稻米的垩白含量是水稻育种改良的重要目标。近年来,研究者们揭示了多个与垩白相关的遗传学机制。本文从能量产生、能量物质分配、淀粉合成和淀粉体堆积结构等途径综述了水稻垩白性状相关基因及其遗传特性的最新研究进展,从而为水稻垩白形成的机理研究与稻米品质遗传改良提供参考。     

稻米品质的分子水平研究进展

随着分子生物学和各项技术的发展,稻米品质的研究取得了一些进展。从与稻米品质中的直链淀粉含量、胶稠度、垩白、糊化温度及碾磨品质有关的基因或QTL,ADP葡聚糖焦磷酸化酶、淀粉合成酶、分支酶及脱支酶在淀粉合成中的作用以及胚乳淀粉粒的形成、形态与结构等方面作了综述 。     

粳稻品质性状间及其与植株性状和产量性状间的遗传相关

利用朱军等提出的种子性状遗传模型,采用 3×3 NCⅡ正反交设计的亲本和部分组合F2代种子,分析了品质性状糙米率、垩白粒率、垩白面积和AC间及上述品质性状与株高、穗部性状等的遗传相关,以期为粳稻育种后代选择提供指导。结果表明,精米重与糙米率存在极显著的母体加性相关;虽然控制品质性状的主要遗传效应分量与植株性状相应遗传效应分量遗传协方差不显著,但在其他相应遗传效应分量方面存在着复杂的关系。     

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。     

籼稻稻米蛋白质含量与外观品质性状间不同遗传体系的条件遗传相关分析

选用稻米蛋白质含量等品质差异较大的7个籼型不育系(A)及相应的保持系(B)与5个籼型恢复系(R)杂交,组成7×5不完全双列杂交组合。应用包括胚乳、细胞质和母体植株基因的遗传主效应以及基因型×环境互作效应的数量性状遗传模型及非条件和条件分析方法,研究了籼稻稻米蛋白质含量与外观品质性状间的遗传相关性,进一步揭示了籼稻糙米重、稻米直链淀粉含量对稻米蛋白质含量与外观品质性状间遗传相关性的影响。非条件分析的结果表明,除了与糙米厚的相关性未达到显著水平以外,蛋白质含量与其它稻米外观品质性状间的遗传相关性均达极显著水平,其中与糙米宽间的相关性表现为正值,其余为负向相关。条件分析的结果显示,糙米重和直链淀粉含量对稻米蛋白质含量与外观品质性状间的遗传相关性均可产生较大的影响。糙米重主要通过基因型×环境互作效应影响蛋白质含量与外观品质性状间的遗传相关性,其中对蛋白质含量与糙米长、糙米宽、糙米厚间遗传相关性的影响主要表现为负向作用,而对蛋白质含量与糙米长宽比、糙米长厚比间遗传相关性的影响则表现为正向作用。稻米直链淀粉主要通过细胞质遗传主效应和母体加性效应影响蛋白质含量与糙米长、糙米长宽比、糙米长厚比间的遗传相关性,而对蛋白质含量与糙米宽间的相关性影响主要表现为负向作用。     

云南高原粳稻品质特性分析

采用表型主成分及聚类分析法,对76个云南高原粳稻的11个品质性状进行了分析。结果表明,（1）直链淀粉含量的平均值达食用稻品种品质1级标准,糙米率、透明度、碱消值和胶稠度达2级标准,精米率达3级标准,其余品质性状的平均值均在3级以下。品种间变异最小的品质性状是糙米率和碱消值,变异最大的是垩白度,其次是垩白粒率。（2）稻米品质性状间存在复杂的相关关系。（3）在选出的6个影响云南稻米品质的主因子中,精米率、垩白粒率和垩白度因子对稻米品质的累积贡献率为49.6%。（4）76个云南粳稻聚为4类,其中第一类包括52个品种,表明遗传距离近,遗传基础狭窄。在今后的云南粳稻品质育种中,应降低垩白粒率和垩白度,提高精米率和整精米率,同时拓宽遗传基础。     

籼稻糙米厚度的发育遗传研究

应用包括３套遗传体系基因效应的数量性状发育遗传模型,分析了１２个籼稻亲本在４个不同稻米发育时期的糙米厚性状。结果表明,三倍体胚乳、二倍体母体植株基因的加性和显性效应以及细胞质效应均可以明显影响各个稻米发育时期的糙米厚度,其中灌浆始期以二倍体母体植株效应为主,灌浆中后期以三倍体胚乳效应为主,成熟期则以细胞质效应为主。在４个不同发育时期中,控制糙米厚的基因加性效应和显性效应交替为主。胚乳显性方差和母体     

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。     

籼稻稻米外观品质的细胞质,母体和胚乳遗传效应分析

利用浙协2号A等9个籼型不育系和T49等5个籼型恢复系进行不完全双列杂交,研究了籼稻稻米外观品质的遗传效应．结果表明,稻米外观品质性状的表现受制于胚乳、母体和细胞质三套遗传体系．糙米长、长宽比和长厚比等性状以母体遗传率为主,而糙米宽和糙米厚则以胚乳直接遗传率为主,糙米长和长宽比等性状的细胞质遗传率亦很重要．结果还发现外观品质性状间存在着较强的遗传相关,其中糙米长与糙米宽、糙米长与糙米厚、糙米宽与糙米厚、糙米宽与长宽比、糙米厚与长厚比以及糙米长宽比与长厚比性状间以胚乳直接加性和母体加性相关为主．而糙米长与长宽比、糙米长与长厚比、糙米宽与长厚比以及糙米厚与长宽比性状间则以胚乳直接显性和母体显性相关为主．就外观品质的总体情况而言,遗传效应预测值表明参试亲本以V20A、作5A和测早2-2较好,其各种遗传效应能够显著改善稻米品质性状。V20A／102和作5A／测早2-2等组合具有较好的稻米外观品质．     

早籼稻米垩白形成中的生理生化变化特性

以早籼多垩白品种泸红早１号、浙７３３和少垩白品种中优早３号、舟优９０３及中籼中引８５为材料,在自然高温条件下对水稻灌浆过程中胚乳内含物质（如淀粉、可溶性糖等）和酶（淀粉磷酸化酶、Ｑ酶）活性的连续变化进行测定。结果表明,不同垩白度品种其胚乳内部物质增长的时间进程和酶活性的变化存在差异,由此造成灌浆动态明显不同,并导致垩白形成的差异。     

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.     

Natural alleles of a uridine 5ʹ-diphospho-glucosyltransferase gene responsible for differential endosperm development between upland rice and paddy rice

Traditional upland rice generally exhibits insufficient grains resulting from abnormal endosperm development compared to paddy rice. However, the underlying molecular mechanism of this trait is poorly understood. Here, we cloned the uridine 5ʹ-diphospho (UDP)-glucosyltransferase gene EDR1 (Endosperm Development in Rice) responsible for differential endosperm development between upland rice and paddy rice by performing quantitative trait loci analysis and map-based cloning. EDR1 was highly expressed in developing seeds during grain filling. Natural variations in EDR1 significantly reduced the UDP-glucosyltransferase activity of EDR1^YZN compared to EDR1^YD1, resulting in abnormal endosperm development in the near-isogenic line, accompanied by insufficient grains and changes in grain quality. By analyzing the distribution of the two alleles EDR1^YD1 and EDR1^YZN among diverse paddy rice and upland rice varieties, we discovered that EDR1 was conserved in upland rice, but segregated in paddy rice. Further analyses of grain chalkiness in the alleles of EDR1^YD1 and EDR1^YZN varieties indicated that rice varieties harboring EDR1^YZN and EDR1^YD1 preferentially showed high chalkiness, and low chalkiness, respectively. Taken together, these results suggest that the UDP-glucosyltransferase gene EDR1 is an important determinant controlling differential endosperm development between upland rice and paddy rice.     

密穗型水稻品种籽粒垩白性状改良研究

采用籽粒长宽比较大、穗部着粒密的散穗型材料(EG23)改良粳稻密穗型品种的籽粒垩白性状.结果表明,经改良后得到的密穗型品系EA6,与原亲本浙粳20比较,其穗部长度缩短,每穗总粒数增加,着粒密度增大,而籽粒垩白特性得到明显的改善,表明在穗部长度和着粒结构未得到改良的情况下,调节籽粒长宽比对改善密穗型品种籽粒垩白性状具有可能性.穗部不同粒位籽粒垩白性状改良的效果不同,穗顶部和穗中部的改良效果明显优于穗基部.设计的4个不同杂交配组方式中,以反回交配组方式(浙粳20/ EG23//浙粳20)选育效果最好.EA6具有较好的农艺性状,既可作为优异种质资源利用,也可直接应用于生产.这一结果从育种实践上较好地协调了密穗型品种高产与优质的矛盾,对于培育既有密穗型的高产株型又有优良籽粒外观品质的水稻品种具有重要意义.     

水稻外观品质的数量性状基因位点分析

利用由98个家系组成的Nipponbare(粳)／Kasalath(秒)∥Nipponbare回交重组自交系(backcross inbred lines,BILs)群体(BC1F9)及其分子连锁图谱,采用复合区间作图的方法,在2个不同年份对粒长、粒宽、粒形、垩白率、垩白大小、垩白度和透明度等7个稻米外观品质性状的数量性状基因位点(Quantiative trait loci,QTL)进行了定位分析。共定位到33个四QTLs,单个性状QTL数目在4-7个之间,以垩白率最多,为7个;粒长和垩白大小次之,为5个;其他性状均为4个,表明该组合外观品质是由多基因控制的数量性状。单个QTL对性状变异解释率粒长为6．2％-15．2％,粒宽为8．3％-32．5％,长宽比为6．8％-19．8％,垩白率为6．4％-28．5％,垩白大小为6．1％-16．9％,垩白度为9．3％-17．2％,透明度为5．6％-25．2％．QTL在染色体上成集中分布的特点,第3染色体C1488-C563、第5染色体R830-R3166和R1436-R2289、第6染色体R2147-R2171均有3个以上的QTLs分布。比较2年的检测结果表明,外观品质性状的QTL定位都受环境影响,但不同性状受影响的程度差异很大。粒长和粒形的QTL定位受环境影响很小,垩白率、垩白大小和垩白度的QTL定位受环境影响很大。     

Du1, encoding a novel Prp1 protein, regulates starch biosynthesis through affecting the splicing of Wxb pre-mRNAs in rice (Oryza sativa L.)

Starch is the major component of cereal grains. In rice, starch properties determine the eating and cooking quality. The dull endosperm of rice grains is a classical morphological and agronomical trait that has long been exploited for breeding and genetics study. To understand the molecular mechanism that regulates the starch biosynthesis in rice grains, we characterized a classic rice mutant dull endosperm1 (du1) and isolated Du1 through a map-based cloning approach. Du1, encoding a member of pre-mRNA processing (Prp1) family, is expressed mainly in panicles. Du1 specifically affects the splicing efficiency of Wx(b) and regulates starch biosynthesis by mediating the expression of starch biosynthesis genes. Analysis of du1wx shows that Du1 acts upstream of Wx(b). These results strongly suggest that Du1 may function as a regulator of the starch biosynthesis by affecting the splicing of Wx(b) and the expression of other genes involved in the rice starch biosynthetic pathways.     

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.     

Fine mapping of the grain chalkiness QTL qPGWC-7 in rice (Oryza sativa L.)

Chalkiness of rice grain is an important quality component of rice, as it has a profound influence on eating and milling qualities. We has determined the inheritance of percentage of grain with chalkiness (PGWC) using a set of chromosome segment substitution lines, made from a cross between cv. PA64s and cv. 9311. Two loci controlling PGWC, designated as qPGWC-6 and qPGWC-7, were located on, respectively, chromosomes 6 and 7. Comparisons were made between C-51 (a CSSL harbouring qPGWC-7 and having a chalky endosperm) and the recurrent parent 9311 (translucent endosperm) to characterize the physical and chemical differences between translucent and chalky endosperm. Unlike the translucent endosperm, the chalky endosperm contains loosely packed starch granules, and there were significant difference between C-51 and 9311 for amylopectin structure and degree of crystallinity, but not for either amylose content or starch viscosity. Segregation analysis of the F₂ population from the cross between C-51 and 9311 showed PGWC is a semi-dominant trait, controlled by single nuclear gene. A large F₂ population was constructed from the cross C51 × 9311, and used for the fine mapping of qPGWC-7, which was located to a 44-kb DNA fragment, containing thirteen predicted genes. This result provides a springboard for the map-based cloning of qPGWC-7 and allowed for marker-assisted selection for endosperm texture.     

Climate Change: Implications for the Yield of Edible Rice

Global warming affects not only rice yield but also grain quality. A better understanding of the effects of climate factors on rice quality provides information for new breeding strategies to develop varieties of rice adapted to a changing world. Chalkiness is a key trait of physical quality, and along with head rice yield, is used to determine the price of rice in all markets. In the present study, we show that for every ∼1% decrease in chalkiness, an increase of ∼1% in head rice yield follows, illustrating the dual impact of chalk on amount of marketable rice and its value. Previous studies in controlled growing conditions report that chalkiness is associated with high temperature. From 1980–2009 at IRRI, Los Baños, the Philippines, annual minimum and mean temperatures, and diurnal variation changed significantly. The objective of this study was to determine how climate impacts chalkiness in field conditions over four wet and dry seasons. We show that low relative humidity and a high vapour pressure deficit in the dry season associate with low chalk and high head rice yield in spite of higher maximum temperature, but in the opposite conditions of the wet season, chalk is high and head rice yield is low. The data therefore suggest that transpirational cooling is a key factor affecting chalkiness and head rice yield, and global warming per se might not be the major factor that decreases the amount and quality of rice, but other climate factors in combination, that enable the crop to maintain a cool canopy.     

Fine mapping and candidate gene analysis of the floury endosperm gene, FLO(a), in rice

In addition to its role as an energy source for plants, animals and humans, starch is also an environmentally friendly alternative to fossil fuels. In rice, the eating and cooking quality of the grain is determined by its starch properties. The floury endosperm of rice has been explored as an agronomical trait in breeding and genetics studies. In the present study, we characterized a floury endosperm mutant, flo(a), derived from treatment of Oryza sativa ssp. japonica cultivar Hwacheong with MNU. The innermost endosperm of the flo(a) mutant exhibited floury characteristics while the outer layer of the endosperm appeared normal. Starch granules in the flo(a) mutant formed a loosely-packed crystalline structure and X-ray diffraction revealed that the overall crystallinity of the starch was decreased compared to wild-type. The FLO(a) gene was isolated via a map-based cloning approach and predicted to encode the tetratricopeptide repeat domaincontaining protein, OsTPR. Three mutant alleles contain a nucleotide substitution that generated one stop codon or one splice site, respectively, which presumably disrupts the interaction of the functionally conserved TPR motifs. Taken together, our map-based cloning approach pinpointed an OsTPR as a strong candidate of FLO(a), and the proteins that contain TPR motifs might play a significant role in rice starch biosynthetic pathways.