Introgression of qPE9-1 allele, conferring the panicle erectness, leads to the decrease of grain yield per plant in japonica rice （Oryza sativa L.）

Panicle architecture is closely related to yield formation. The qPE9-1 gene has been proved to be widely used in high-yield rice cultivar developments, conferring erect panicle character in japonica rice. Recently, qPE9-1 has been successfully cloned; however, the genetic effect on grain yield per plant of the erect panicle allele qPE9-1 is controversial yet. In the present study, a drooping panicle parent Nongken 57, carrying qpe9-1 allele, was used as recurrent parent to successively backcross to a typical erect panicle line from the double haploid (DH) population (Wuyunjing 8/Nongken 57), which was previously shown to carry qPE9-1 allele. Thus a pair of near-isogenic lines (NILs) was developed. The comparison of agronomic traits between the NILs showed that, when qpe9-1 was replaced by qPE9-1, the panicle architecture was changed from drooping to erect; moreover, the panicle length, plant height, 1000-grain weight and the tillers were significantly decreased, consequently resulting in the dramatic decrease of grain yield per plant by 30%. Therefore, we concluded that the qPE9-1 was a major factor controlling panicle architecture, and qPE9-1 had pleiotropic nature, with negative effects on grain yield per plant. This result strongly suggests that the erect panicle allele qPE9-1 should be used together with other favorable genes in the high-yield breeding practice. In addition, the effect of qPE9-1 on eating and cooking quality was also discussed in the present study.     

籼稻花粉无性系变异的研究

对７个籼稻品种通过花药培养获得的１６１个花粉植株进行了花粉无性系变异的研究。结果表明：（１）所有花粉无性系在遗传上都是纯合的,其整齐度与起始亲本相近或超过起始亲本;（２）花粉无性系的变异主要在数量性状上有一定变幅,变异方向有负向也有正向的。同一来源的无性系间的变异系数略高于起始亲本,仅千粒重达到显著标准。以所考查的性状为基数,变异的频率为１２．５％,同时有３－４个性状发生变异的无性系数占总数的４．３％;（３）少数变异体性状有重大变异,并具有育种价值;（４）根据１０种同工酶和ＲＦＬＰ分析,仅从３个性状发生重大变化的变异体中检测出与起始亲本的差异。     

The extent of parental genotypic divergence determines maximal heterosis by increasing fertility in inter-subspecific hybrids of rice (Oryza sativa L.)

The magnitude of heterosis in F₁ hybrids is related not only to the performance of parents per se but also to the genetic diversity between two parents. The extent of genotypic divergence between hybrid rice parents was investigated at the molecular level, using two subsets of rice materials: a subset of doubled haploid (DH) lines derived from an Indica × Japonica cross (Gui630/02428) and another subset of Indica or Japonica lines representative of a broad spectrum of the Asian cultivated rice gene pool, including landraces, primitive cultivars, historically important cultivars, modern elite cultivars, super rice and parents of superior hybrids. 57 entries deliberately selected from the 81-DH lines (in total) were testcrossed to two widely used rice lines in China, photoperiod-sensitive genic male sterile (PGMS) N422s and thermo-sensitive genic male sterile (TGMS) Peiai64s. Results of the two sets of test-cross F₁ populations showed congruently that parental genotypic divergence has a relatively low impact on heterosis for the two yield components, i.e., panicle number and 1000-grain weight, but it has a great bearing on fertility parameters, i.e., filled grains per plant and seedset. Heterosis for grain yield in the two test-cross populations exhibited a sharp maximum when the proportion of Japonica alleles in the male parent was between 50 and 60%, so was the heterosis for fertility parameters correspondingly. Thus fertility parameters were the most sensitive and important factors which were influenced by the extent of parental genotypic divergence. Moreover, our results showed that parents with moderate extent of genotypic divergence played an important role in the use of inter-subspecific rice heterosis.     

水稻籼粳交DH群体籽粒充实度的遗传分析

籽粒充实度差是限制亚种间杂交稻产量潜力发挥的重要因素。作者对籼粳交（圭630／02428）DH群体的籽粒充实度进行了研究,结果表明,籽粒充实度是受多基因控制的数量性状,在该DH群体中发生分离的基因估计数为5－6对。通过估测三级（偏度）和四级（峰度）统计量,检测到控制籽粒充实度的多基因间存在重叠作用。DH群体中籽粒充实度表现出明显的超亲分离,表明基因重组可实现控制籽粒充实度的增效基因的聚合。籽粒充实度与单穗产量、穗均实粒数和干物质／总库容之比呈显著或极显著正相关,与千粒重、穗均颖花数和穗均总库容相关不显著。     

不同遗传背景下杂交籼稻功能叶与产量构成因素的相关性

为明确水稻功能叶与产量构成因素间的相关性,以不同遗传背景下籼稻的10个不育系和16个恢复系为亲本,按照NCII设计配制两套双列杂交组合,对水稻12个功能叶性状与8个产量性状构成因素进行了相关分析,结果表明：3片功能叶叶长与叶面积、剑叶宽、倒2叶宽等性状之间均存在极显著正相关,功能叶夹角之间也存在极显著正相关,但不同遗传背景对夹角性状与9个形态性状之间的相关性则存在明显差异,在第1套组合中,其相关系数均为负值,且相关均不显著;而第2套组合则相反。8个产量构成因素中,单株穗数与平均穗长、着粒密度、穗实粒数以及穗着粒数之间存在极显著负相关,平均穗长与穗着粒数、结实率与单株产量呈显著或极显著正相关,遗传背景对产量组成上有较大影响,在第1套组合中单株产量主要由结实率、单株穗数以及穗实粒数等性状决定,而在第2套中则主要由穗实粒数和结实率等性状决定。在功能叶与产量构成因素的相关中,叶长、叶面积、剑叶宽、倒2叶宽与着粒密度、穗实粒数以及穗着粒教等3个性状之间存在显著或极显著正相关。12个水稻功能叶性状与8个产量构成因素之间的主成分分析表明,在不同的遗传背景下,产量构成因素均主要受叶面积和叶夹角影响,两种不同遗传背景中其累积贡献率分别为69．8％和84．0％。     

水稻双单倍体群体的分子标记图示基因型分析

利用窄叶青８号（籼稻）／京系１７（粳稻）花培产生的双单倍体群体建立了一个包含１６０个分子标记的遗传连锁图,在此基础上利用ＨＹＰＥＲＧＥＮＥ软件建立了５２个ＤＨ系的图示基因型,并对ＤＨ系的亲本基因组比率和染色体的交换重组频率进行了比较分析。结果表明本实验所用的ＤＨ群体没有显著偏离正态分布,籼粳稻杂交后代中植株的籼粳表现与同工酶、形态指数和基因组比率的分析结果一致,此外还发现ＤＨ群体中出现了大量的交换罕见染色体。利用图示基因型分析发现株高和分子标记ＲＺ９７８和ＲＧ４Ａ相关,生育期和ＲＲＫ０８－１、ＲＧ４７７和ＲＧ５１１相关。本文还就图示基因型分析技术在ＤＨ群体的遗传分析和选择育种中的应用进行了讨论．     

A Pyramid Breeding of Eight Grain-yield Related Quantitative Trait Loci Based on Marker-assistant and Phenotype Selection in Rice （Oryza sativa L.）

1000-Grain weight and spikelet number per panicle are two important components for rice grain yield.In our previous study,eight quantitative trait loci(QTLs)conferring spikelet number per panicle and 1000-grain weight were mapped through sequencing-based genotyping of 150 rice recombinant inbred lines(RILs).In this study,we validated the effects of four QTLs from Nipponbare using chromosome segment substitution lines(CSSLs),and pyramided eight grain yield related QTLs.The new lines containing the eight QTLs with positive effects showed increased panicle and spikelet size as compared with the parent variety 93-11.We further proposed a novel pyramid breeding scheme based on marker-assistant and phenotype selection(MAPS).This scheme allowed pyramiding of as many as 24 QTLs at a single hybridization without massive cross work.This study provided insights into the molecular basis of rice grain yield for direct wealth for high-yielding rice breeding.     

In vitro selection and field responses of somaclonal variant plants of rice cv PR113 for drought tolerance

Drought is the major environmental stress that limits rice productivity worldwide. In vitro somaclonal variation using different selection agents has been used for crop improvement. Here, rice plants of cv PR113 were selected in vitro on 30, 50 and 70 g L^-1 polyethylene glycol 6,000 (PEG). Callus growth, proliferation, calli volume (first and second culture) and plantlet regeneration (third culture) were found to be decreased upto a certain level to acquire tolerance to PEG-induced drought. From the field data, 30 g L^-1 PEG lines showed higher vegetative growth (plant height, tiller number, leaf number, shoot weight and root growth) as compared with 50 g L^-1 PEG selected somaclone lines under limited irrigation. The yield parameters-panicle length, panicle weight, grains per panicle, 1,000-grain weight, grain yield per plant, harvest index and grain straw ratio were also higher in 30 g L^-1 PEG lines as compared with 50 g L^-1 PEG lines. The results, therefore indicate that 30 g L^-1 PEG selected somaclone lines were more suited than 50 g L^-1 PEG selected somaclone lines under stress as compared with WT. The finding suggests that rice cv PR113 somaclones generated on PEG are found to be drought tolerant under field condition with better yield.     

普通野生稻中增产相关QTL的发掘

普通野生稻(Oryza Rufipogon)是重要的遗传资源,发掘其优良等位基因将对水稻遗传改良产生重要影响。文章从以珍汕97为轮回亲本,普通野生稻为供体的BC2F1群体中选择一个与珍汕97表型明显不同的单株BC2F1-15,经过连续自交获得回交重组自交系BC2F5群体。均匀分布于12条染色体的126个多态性SSR(Simplesequence repeats)标记基因型分析,发现BC2F1-15单株在30%的标记位点为杂合基因型;利用该群体共检测到4个抽穗期、3个株高、4个每穗颖花数、2个千粒重和1个单株产量QTL。在第7染色体RM481-RM2区间,检测到抽穗期、每穗颖花数和产量QTL,野生稻等位基因表现增效作用;其他3个每穗颖花数QTL位点,野生稻等位基因也均具有增效作用。结果表明野生稻携带有增产相关的等位基因,这些有利等位基因无疑是水稻遗传改良可资利用的新资源。     

Over-expression of the rice LRK1 gene improves quantitative yield components

In rice ( Oryza sativa L.), the number of panicles, spikelets per panicle and grain weight are important components of grain yield. These characteristics are controlled by quantitative trait loci (QTLs) and are derived from variation inherent in crops. As a result of the complex genetic basis of these traits, only a few genes involved in their control have been cloned and characterized. We have previously map-cloned a gene cluster including eight leucine-rich repeat receptor-like kinase ( LRK ) genes in Dongxiang wild rice ( Oryza rufipogon Griff.), which increased the grain yield by 16%. In the present study, we characterized the LRK1 gene, which was contained in the donor parent (Dongxiang wild rice) genome and absent from the recurrent parent genome (Guichao2, Oryza sativa L. ssp. indica ). Our data showed that rice LRK1 is a plasma membrane protein expressed constitutively in leaves, young panicles, roots and culms. The over-expression of rice LRK1 results in increased panicles, spikelets per panicle, weight per grain and enhanced cellular proliferation, leading to a 27.09% increase in total grain yield per plant. The increased number of panicles and spikelets per panicle are associated with increased branch number. Our data suggest that rice LRK1 regulates rice branch number by enhancing cellular proliferation. The functional characterization of rice LRK1 facilitates an understanding of the mechanisms involved in cereal crop yield, and may have utility in improving grain yield in cereal crops.     

A comparative screening of hybrid,modern varieties and local rice cultivar for brown leaf spot disease susceptibility and yield performance

Five rice cultivars, one hybrid (WR96), three modern (BR16, BR26, and BRRI Dhan27) and one local (Pari) were screened for reaction to brown leaf spot disease caused by Cochliobolus miyabeanus and performance of yield-related characters. The severity of brown leaf spot varied with growth stages of rice plant as well as different cultivars tested under field condition. Low disease severity was observed at maximum tillering stage compared to moderate to high at dough stage, with hybrid cultivar WR96 showing highest disease, while local cultivar Pari had the lowest. Brown spot disease severity in different cultivars under induced epiphytic condition also followed the similar trend. The results also revealed that most of the yield-contributing characters examined showed wide variations among the cultivars. Modern cultivar BR16 produced the highest panicle length, number of grain per panicle and grain yield per hectare. At the same time as local cultivar Pari generated the lowest number of tiller per plant, panicle length, grain number per panicle and grain yield per hectare. Moreover, hybrid cultivar WR96 produced the highest percentage of spotted grain per panicle and seed yielding C. miyabeanus, and also the lower percentage of seed germination, while the reverse was observed in local cultivar Pari. These findings may allow producers and breeders to select rice cultivar, resistant or tolerant to brown leaf spot disease and to avoid significant reductions in grain yields.     

水稻穗颈维管束及穗部性状的QTL分析

以籼稻(Oryza sativa L. ssp. indica ZYQ8)和粳稻(O. sativa ssp. japonica JX17)的杂交F1代花培加倍的DH群体为材料考察了该群体的穗颈节大小维管束数、一次枝梗数、每穗颖花数、穗颈节顶部直径和穗长,并用该群体构建的分子图谱进行数量性状座位(QTL)分析.检测到控制大维管束的3个QTL (qLVB-1、qLVB-6和qLVB-7)分别位于第1、第6和第7染色体;控制小维管束的2个QTL (qSVB-4和qSVB-6)分别位于第4和第6染色体;控制一次枝梗的4个QTL (qPRB-4a、qPRB-4b、qPRB-6和qPRB-7)分别位于第4 (2个)、第6和第7染色体;每穗颖花数的3个QTL (qSPN-4a、qSPN-4b和 qSPN-6)分别位于第4 (2个)和第6染色体上;穗颈节顶部直径的5个QTL (qPTD-2、qPTD-5、qPTD-6、qPTD-8和qPTD-12)分别位于第2、第5、第6、第8和第12染色体;穗长的3个QTL (qPL-4、qPL-6和qPL-8)分别位于第4、第6、第8染色体上.其中qLVB-6、qSVB-6、qSPN-6、qPTD-6和qPL-6均位于第6染色体的G122-G1314b之间;qPL-8和qPTD-8位于第8染色体的GA408-BP127a之间;qPRB-4a和qSPN-4a位于第4染色体的G177-CT206之间;qPL-4和qSPN-4b位于第4染色体CT404-CT500之间;qSVB-4所在的区间与qPL-4、qSPN-4b和qPRB-4b所在的区间相邻.     

Sensitivities of rice grain yield and other panicle characters to late-stage drought stress revealed by phenotypic correlation and QTL analysis

A lowland rice variety (Zhenshan97B) was crossed with an upland variety (IRAT109) to construct a set of recombinant inbred lines (RILs). The population was evaluated under both well-watered (control) and drought-stress (drought) conditions for 2 years. Panicle water potential (PWP), panicle length (PL), grain number per plant (GNP), primary branch number (PBN), second branch number (SBN), spikelet density (SPD), and dry grain weight per plant (DGW) were measured. Phenotypic correlation and path analysis were used to interpret the ranking of importance of other panicle traits to grain yield. Comparison of such rankings under control and drought conditions showed varied responses of panicle traits to drought stress. It was indicated that GNP was an important contributor to DGW under control, and even more important under drought. Thirty-two quantitative trait loci (QTLs) for panicle traits and dry grain weight were identified, with contribution rates ranging from 3.33% to 22.66%. Eleven epistatic QTLs were detected. Cases of collocated QTLs under control and drought were found for PL, SPD, GNP, PBN, and SBN, but not for DGW and PWP. The effectiveness of selection under normal and stressed conditions is discussed.     

QTLs linked to leaf epicuticular wax,physio-morphological and plant production traits under drought stress in rice (<Emphasis Type="Italic">Oryza sativa</Emphasis> L.)

Drought stress is the major constraint to rice (Oryza sativa L.) production and yield stability in rainfed ecosystems. Identifying genomic regions contributing to drought resistance will help to develop rice cultivars suitable for rainfed regions through marker-assisted breeding. Quantitative trait loci (QTLs) linked to leaf epicuticular wax, physio-morphological and plant production traits under water stress and irrigated conditions were mapped in a doubled haploid (DH) line population from the cross CT9993-5-10-1-M/IR62266-42-6-2. The DH lines were subjected to water stress during anthesis. The DH lines showed significant variation for epicuticular wax (EW), physio-morphological and plant production traits under stress and irrigated conditions. A total of 19 QTLs were identified for the various traits under drought stress and irrigated conditions in the field, which individually explained 9.6%–65.6% of the phenotypic variation. A region EM15_10-ME8_4-R1394A-G2132 on chromosome 8 was identified for leaf EW and rate of water loss i.e., time taken to reach 70% RWC from excised leaves in rice lines subjected to drought stress. A large effect QTL (65.6%) was detected on chromosome 2 for harvest index under stress. QTLs identified for EW, rate of water loss from excised leaves and harvest index under stress in this study co-located with QTLs linked to shoot and root-related drought resistance traits in these rice lines and might be useful for rainfed rice improvement.     

Studies on some genetic parameters of rice (Oryza sativa L.)

Summary In order to utilize the available useful variation in breeding rice to improve yield and quality of grain, the performance of a wide range of rice germ plasm comprising 30 varieties was evaluated in northern India, a major rice growing belt. Plant performance revealed high genetic divergence and phenotypic variability in the crop, with the maximum range of variation being for grain number per panicle and the minimum for grain dimensions. There were also significant differences among varietal means for ten phenotypic traits. Genotypic and phenotypic variance contributed profoundly to the variance of the phenotypic traits studied, but, since genetic variability in the traits related to yield was considerable, there is scope for further improvement in yielding ability. Grain number per panicle, number of effective tillers per plant and culm length exhibit high heritability, and genotypic coefficient of variation and therefore a high genetic advance. Thus, selection for these traits would be effective in crop improvement. Moreover, grain number per panicle shows a significant positive correlation with yield, and this trait could profitably form a reliable index for the yielding capacity of this crop.     

Development of doubled haploids from an elite <Emphasis Type="Italic">indica</Emphasis> rice hybrid (BS6444G) using anther culture

A total of 200 doubled haploids (DHs) were generated from an elite rice hybrid, ‘BS6444G’ for which an androgenic method was developed by manipulating the physical and chemical factors. The spike pretreated at 10?°C for 7–8 days was effective for callusing and green plant regeneration. The maximum callus frequency was achieved when the anthers cultured in N6 medium supplemented with 2.0 mg L^?1 2,4-diclorophenoxyacetic acid, 0.5 mg L^?1 6-benzylaminopurine and 3% maltose. Calli induced in N6 media also showed significant green shoot regeneration in MS medium supplemented with 0.5 mg L^?1 1-napthalene acetic acid, 0.5 mg L^?1 kinetin, 1.5 mg L^?1 benzylaminopurine and 3% sucrose producing 210 green plants. Assessment of the ploidy status showed 95.71% fertile diploids and 4.2% polyploids; no haploids were observed. A total of 38 sequence-tagged microsatellite (STMS) markers proved able to discriminate a heterozygote from all the 200 DHs. The DHs grown in the field showed significant variation for their agronomic traits. Comparison of traits with control indicates homogeneity within each DH line and significant variance of traits between DH lines. Nine DH lines produce higher grain yield than the hybrid parent which suggests the possibility of exploiting hybrid vigor in indica rice through the development of DH lines of high yielding hybrids.     

Identification of QTL for growth- and grain yield-related traits in rice across nine locations of Asia

Rice double-haploid (DH) lines of an indica and japonica cross were grown at nine different locations across four countries in Asia. Genotype-by-environment (G x E) interaction analysis for 11 growth- and grain yield-related traits in nine locations was estimated by AMMI analysis. Maximum G x E interaction was exhibited for fertility percentage number of spikelets and grain yield. Plant height was least affected by environment, and the AMMI model explained a total of 76.2% of the interaction effect. Mean environment was computed by averaging the nine environments and subsequently analyzed with other environments to map quantitative trait loci (QTL). QTL controlling the 11 traits were detected by interval analysis using mapmaker/qtl. A threshold LOD of >/=3.20 was used to identify significant QTL. A total of 126 QTL were identified for the 11 traits across nine locations. Thirty-four QTL common in more than one environment were identified on ten chromosomes. A maximum of 44 QTL were detected for panicle length, and the maximum number of common QTL were detected for days to heading detected. A single locus for plant height (RZ730-RG810) had QTL common in all ten environments, confirming AMMI results that QTL for plant height were affected the least by environment, indicating the stability of the trait. Two QTL were detected for grain yield and 19 for thousand-grain weight in all DH lines. The number of QTL per trait per location ranged from zero to four. Clustering of the QTL for different traits at the same marker intervals was observed for plant height, panicle number, panicle length and spikelet number suggesting that pleiotropism and or tight linkage of different traits could be the possible reason for the congruence of several QTL. The many QTL detected by the same marker interval across environments indicate that QTL for most traits are stable and not essentially affected by environmental factors.     

Comparative mapping of QTLs for agronomic traits of rice across environments using a doubled haploid population

We report here the RFLP mapping of quantitative triat loci (QTLs) that affect some important agronomic traits in cultivated rice. An anther culturederived doubled haploid (DH) population was established from a cross between an indica and a japonica rice variety. On the basis of this population a molecular linkage map comprising 137 markers was constructed that covered the rice genome at intervals of 14.8cM on average. Interval mapping of the linkage map was used to locate QTLs for such important agronomic traits as heading date, plant height, number of spikelets per panicle, number of grains Per panicle, 1000-grain weight and percentage of seed set. Evidence of genotype-byenvironment interaction was found by comparing QTL maps of the same population grown in three diverse environments. A total of 22 QTLs for six agronomic traits were detected that were significant in at least one environment, but only 7 were significant in all three environments, 7 were significant in two environments and 8 could only be detected in a single environment. However, QTL-by-environment interaction was traitdependent. QTLs for spikelets and grains per panicle were common across environments, while traits like heading date and plant height were more sensitive to environment.     

Analysis of quantitative trait loci which contribute to anther culturability in rice (Oryza sativa L.)

Anther culturability of rice is significantly different between indica and japonica varieties. A doubled haploid (DH) population was established via anther culture of an indica/japonica hybrid on SK₃ medium, which had been shown particularly suitable for anther culture of indica/japonica hybrids. For analyzing the quantitative trait loci (QTLs) responsible for anther culturability, anthers of the DH lines were again cultured with SK₃ medium and parameters for four traits representing the anther culturability were surveyed and analyzed with the molecular map constructed from the same DH population. The parameters for four major traits were as follows: callus induction frequency (CI), green plantlet differentiation frequency (GPD), albino plantlet differentiation frequency (APD), and green plantlet yield frequency (GPY). All four traits displayed continuous distributions among the DH lines. The correlation coefficients between these traits were also tested and showed that there was no relationship between callus induction and green plantlet differentiation frequencies, but both showed strong positive correlation with the frequency of green plantlet yield. For callus induction frequency, five QTLs were identified on chromosomes 6, 7, 8, 10 and 12. Two QTLs for green plantlet differentiation frequency were located on chromosomes 1 and 9. There was a major QTL for albino plantlet differentiation frequency on chromosome 9. No independent QTL was found for green plantlet yield frequency. The results may be useful in the selection of parents with high response to anther culture for rice haploid breeding and in the establishment of permanent DH populations for molecular mapping.     

利用最大似然法进行水稻产量性状基因的分子作图

本研究根据对估计标记－数量性状基因座位（ＱＴＬ）之间重组率的两种分析方法（矩量法和最大似然法）、两种方差模型（ＱＴＬ基因型之间的方差同质和异质模型）的分析,揭示了ＬＯＤ值在标记－ＱＴＬ连锁检测上所得结果的相关性高于重组率估计值的相关性。采用最大似然法和异质方差模型,估计了水稻产量构成有关的ＱＴＬ与分布于１１对染色体上的５１个限制性片段长度多态性（ＲＦＬＰ）标记之间的重组率,并对似然比（以ＬＯＤ值表示）进行Ｘ￣２检验,发现７个存在显著连锁关系的标记－性状组合,其平均重组率为１０．０％。这些标记分布于第１、５、６、８和１１等５对染色体上,涉及７个ＲＦＬＰ标记和３个产量构成性状,即每穗颖花数（ＲＧ５７３、ＲＺ６１７、ＲＧ１０３）、单株穗数（ＲＧ６４Ｂ）和每穗实粒数（ＲＧ１０１、ＲＧ２４４、ＲＧ６５３）。