Indica-japonica Cross Research Articles

Two complementary genes in a presence-absence variation contribute to indica-japonica reproductive isolation in rice

Understanding the evolutionary forces in speciation is a central goal in evolutionary biology. Asian cultivated rice has two subspecies, indica and japonica, but the underlying mechanism of the partial reproductive isolation between them remains obscure. Here we show a presence-absence variation (PAV) at the Se locus functions as an indica-japonica reproductive barrier by causing hybrid sterility (HS) in indica-japonica crosses. The locus comprises two adjacent genes: ORF3 encodes a sporophytic pollen killer, whereas ORF4 protects pollen in a gametophytic manner. In F1 of indica-japonica crosses, pollen with the japonica haplotype, which lacks the sequence containing the protective ORF4, is aborted due to the pollen-killing effect of ORF3 from indica. Evolutionary analysis suggests ORF3 is a gene associated with the Asian cultivated rice species complex, and the PAV has contributed to the reproductive isolation between the two subspecies of Asian cultivated rice. Our analyses provide perspectives on rice inter-subspecies post-zygotic isolation, and will promote efforts to overcome reproductive barriers in indica-japonica hybrid rice breeding.

A minimal genome design to maximally guarantee fertile inter-subspecific hybrid rice

Hybrid rice has made considerable contributions to achieve the ambitious goal of food security for the world’s population. Hybrid rice from indica/xian and japonica/geng subspecies shows much higher heterosis and is thereby an important innovation in promoting rice production in the next decade. However, such inter-subspecific hybrid rice has long suffered from serious hybrid sterility, which is a major challenge that needs to be addressed. In this study, we performed a genome design strategy to produce fertile inter-subspecific hybrid by creation of wide compatibility varieties that are able to overcome hybrid sterility. Based on combined genetic analyses in two indica-japonica crosses, we determined that four hybrid sterility loci, S5, f5, pf12 and Sc, are the major QTLs controlling inter-subspecific hybrid sterility and thus the minimal targets that can be manipulated for breeding sub-specific hybrid rice. We then cloned the pf12 locus, one of the most effective loci for hybrid male sterility, by map-based cloning, and showed that artificial disruption of pf12A gene at this locus could successfully rescue hybrid fertility. We further dissected the genetic basis of wide compatibility using three pairwise crosses from a wide-compatibility variety Dular and representative indica and japonica varieties. On this basis, we constructed and assembled different combinations of naturally compatible alleles of four loci, S5, Sc, pf12, and f5, and found that the improved lines could fully recover pollen and embryo sac fertility in test-crossed F1s, thereby completely fulfilling the demands of inter-subspecific hybrid spikelet fertility in agricultural production. This breeding scheme would facilitate redesign of future inter-subspecific hybrid rice with a higher yield potential.

The Pyramiding of Elite Allelic Genes Related to Grain Number Increases Grain Number per Panicle Using the Recombinant Lines Derived from Indica-japonica Cross in Rice.

Indica(xian)-japonica(geng) hybrid rice has many heterosis traits that can improve rice yield. However, the traditional hybrid technology will struggle to meet future needs for the development of higher-yield rice. Available genomics resources can be used to efficiently understand the gene-trait association trait for rice breeding. Based on the previously constructed high-density genetic map of 272 high-generation recombinant inbred lines (RILs) originating from the cross of Luohui 9 (indica, as female) and RPY geng (japonica, as male) and high-quality genomes of parents, here, we further explore the genetic basis for an important complex trait: possible causes of grain number per panicle (GNPP). A total of 20 genes related to grains number per panicle (GNPP) with the differences of protein amino acid between LH9 and RPY were used to analyze genotype combinations, and PCA results showed a combination of PLY1, LAX1, DTH8 and OSH1 from the RPY geng with PYL4, SP1, DST and GNP1 from Luohui 9 increases GNPP. In addition, we also found that the combination of LAX1-T2 and GNP1-T3 had the most significant increase in GNPP. Notably, Molecular Breeding Knowledgebase (MBK) showed a few aggregated rice cultivars, LAX1-T2 and GNP1-T3, which may be a result of the natural geographic isolation between the two gene haplotypes. Therefore, we speculate that the pyramiding of japonica-type LAX-T2 with indica-type GNP1-T3 via hybridization can significantly improve rice yield by increasing GNPP.

Fine Mapping and Candidate-Gene Analysis of an open glume multi-pistil 3 (mp3) in Rice (Oryza sativa L.)

The rice mutant mp3 was derived from an indica–japonica cross between Rejing35 and XieqingzaoB, producing an inconstant number of pistils ranging from one to four pistils in a floret at heading stage, which also developed an open-glume with one or two seeds and twin seedlings at mature and seedling stage. Several altered characteristics, including filling grain panicle–1 (62.90), grain-setting rate (60.48%) and grain yield plant–1 (13.42 g), decreased but an increase in 1000-grain weight (36.87 g) was observed. Genetic analysis revealed that the mp3 mutant phenotype was controlled by a single recessive gene. Using a chromosome walking strategy in the F2 population of 02428/mp3, the mp3 gene was fine mapped between L3-135 and RM7576, with a physical distance of 30.617 kb on rice chromosome 3. Four candidate genes were found in this region referred to the rice genome annotations. LOC_Os03g11614/OsMADS1 corresponded with the mutant mp3 phenotype. Sequencing showed no sequence alterations in the coding and promoter sequence of the LOC_Os03g11614/OsMADS1 of mp3. The mp3 gene may be an allelic gene with three previously reported genes but controlled different mutant floral organ phenotypes in rice. Therefore, this mp3 gene provided a novel perspective on the biological function of OsMADS1 in the development of rice floral organ.

QTL Analysis and Heterosis Loci of Effective Tiller Using Three Genetic Populations Derived from Indica-Japonica Crosses in Rice

Effective panicle numbers (PNs) and Tiller numbers (TNs) are important traits affecting rice (Oryza sativa L.) architecture and grain yield. However, the molecular mechanisms underlying PN and TN heterosis remain unknown in rice. In addition, new PN- or TN-related genes need to be detected and discovered. In this study, in order to detect rice quantitative trait loci (QTLs) and the heterosis-related loci of PN or TN in rice, we developed a high generation recombinant inbred line (RIL) population from a cross of two elite cultivars, Luohui9 (Xian/Indica) and RPY geng (Geng/Japonica), and two testcross hybrid populations derived from the crosses of RILs and two cytoplasmic male sterile lines, YTA (Xian/Indica) and Z7A (Geng/Japonica). Finally, nine QTLs of PN across four seasons were identified, and two QTLs of TN in 191HB were mapped. Besides this, six heterosis-related QTLs of PN and five heterosis-related QTLs of TN were located. We found that heterosis-related QTLs of PN or TN covered multiple known genes, such as MOC1, TAC1 and OsETR2. Furthermore, homologous gene analysis identified one candidate gene of PN (LOC_10g25720). Together, these findings uncover multiple heterosis-related loci, and provide a new insight into the heterosis mechanism of PN and TN in rice.

Bin mapping-based QTL analyses using three genetic populations derived from indica-japonica crosses uncover multiple grain shape heterosis-related loci in rice.

Exploitation of heterosis between indica and japonica has important significance in scientific research and agriculture application. However, the molecular mechanism of grain shape heterosis in indica-japonica hybrid remains unknown in rice (Oryza sativa L.). To reveal the genetic mechanism of grain shape in indica--japonica hybrid, we constructed a high-generation recombinant inbred line (RIL) population and two testcross hybrid populations derived from the cross of RILs and two cytoplasmic male sterile material (YTA and Z7A) and then performed a bin mapping-based quantitative trait locus (QTL) mapping of multiple grain shape traits, such as grain length (GL), grain width (GW), and grain length-to-width ratio (GLWR). A total of sixteen QTLs and 30 heterosis-related QTLs of grain shape traits were detected. We found that GS3, GS5, and OsPPKL2 were also correlated with grain shape both in RILs and two testcross hybrid populations. Homologous gene analysis emphasized two candidate grain shape-associated genes (LOC_Os06g14260 and LOC_Os04g51950). Our findings uncover multiple grain shape heterosis-related loci and provides a new insight into heterosis mechanism of grain shape in rice.

Mapping QTL for leaf pigment content at dynamic development stage and analyzing Meta-QTL in rice

Leaf pigment content is a complicated quantitative trait which results from the cooperation of multiple related genes or quantitative trait loci (QTL) in rice. So far, more than 600 QTL for leaf pigment content has been reported. However, most QTL positions could not be directly compared with each other. In this study, our goal was to analyze consistent QTL regions affecting leaf pigment content by applying a genome wide QTL meta-analysis approach. 20 QTL related to leaf pigment content were detected at 5 different growth periods in 134 recombinant inbred lines (RILs) from a typical Indica-Japonica cross derived from PA64s and 9311 (two parents of super hybrid rice Liangyoupei 9). There was a candidate gene GLTP in the region of 3 QTL (qFS_Chla4, qFS_Chla+b4, and qFS_Car4), which related to component of chloroplast thylakoid membranes. After sequencing and qRT-PCR analysis, the gene showed significant difference in parents of the RILs. Meta-analysis was performed on 618 QTL collected from 38 previous studies. Consequently, a total of 64 MQTL (Meta-QTL) were obtained, of which 58.95% confidence intervals (CI) were smaller than the average Cl of the original QTL. In addition, 64 MQTL covered 6 of 20 QTL detected. Among them, four main QTL were included in 3 MQTL, and phenotypic variation of 21 MQTL was greater than 20%. The traits with higher number of QTL had higher number of MQTL, and candidate genes in overlapping MQTL include multiple effects. 7 candidate genes related to leaf pigment content were found in 4 MQTL spanning physical intervals < 0.5 Mb. These findings are not only anticipated to help molecular marker-assisted selection and pyramiding, but also contribute to determine candidate genes that regulate pigment content.

Dissecting combining ability effect in a rice NCII-III population provides insights into heterosis in indica-japonica cross

BackgroundCombining ability is a measure for selecting elite parents that make the highest contributions to hybrid performance. However, the genetic bases of combining ability and how they contributed to heterosis is seldomly known.ResultsWe constructed a both NCII and NCIII population derived from an indica-japonica cross to study the relationship among parental performance, combining ability and hybrid performance of 11 agronomic traits. Among them, specific combining ability is more important to grain yield than parental performance and general combining ability. We performed linkage analyses to phenotypic values and combining ability of all 11 traits in Doubled haploid lines and its two backcross populations and identified 108 QTLs in total. Among these QTLs, four known loci, Sd1, Ghd7, Ghd8 and DEP1 contribute a lot to GCA effects of agronomic traits except grain yield and seed setting rate. Three QTLs, Ghd8, S5 and qS12, contribute a lot to SCA effects of grain yield and present overdominace.ConclusionsOur study provides insights into the genetic bases of combining ability and heterosis and will promote the improvements of indica-japonica hybrid breeding.

QTL analyses of heterosis for grain yield and yield-related traits in indica-japonica crosses of rice (Oryza sativa L.)

Two sets of rice materials, 166 RILs derived from a cross between Milyang 23 (Korean indica-type rice) and Tong 88-7 (japonica Rice), and BC1F1 hybrids derived from crosses between the RILs and the female parent, Milyang 23, were produced to identify QTLs for heterosis of yield and yield-related traits. The QTLs were detected from three different phenotype data sets including the RILs, BC1F1 hybrids, and mid-parental heterosis data set acquired from the definition of mid-parental heterosis. A total of 57 QTLs were identified for nine traits. Of eight QTLs detected for yield heterosis, five overlapped with other heterosis QTLs for yield-related traits such as spikelet number per panicle, days to heading, and spikelet fertility. Four QTLs for yield heterosis, gy1.1, py6, gy10, and py11, were newly identified in this study. We identified a total of 17 EpQTLs for yield heterosis that explain 21.4 ∼ 59.0 % of total phenotypic variation, indicating that epistatic interactions may play an important role in heterosis.

Characterization of Imcrop, a Mutator-like MITE family in the rice genome

Sequence comparisons of ammonium transporter 1–2 genes (OsAMT1-2) in different rice accessions revealed a MITE insertion in the upstream region of the gene. The 391-bp MITE, classified as a Mutator superfamily member and named Imcrop, included terminal inverted repeat (TIR) and 9-bp target site duplication (TSD) sequences. We identified 151 Imcrop elements dispersed on 12 chromosomes of the japonica reference genome. Of these, 12.6% were found in genic regions and 33.1% were located within 1.5 kb of annotated rice genes. We constructed comparative insertion maps with 111 and 102 intact Imcrop elements in the japonica and indica reference genomes, respectively. The Imcrop elements showed relatively even distribution across all chromosomes although their frequency was higher on chromosomes 1, 3, and 4 in both genomes. Seventy seven Imcrop elements were detected in both subspecies, whereas 34 and 25 insertions were found only in the japonica or indica genome, respectively. We compared insertion polymorphisms of 19 Imcrop elements found inside genes in 48 Korean rice cultivars, consisting of 42 japonica and six Tongil-types (indica-japonica cross). Thirteen insertions were common to all cultivars indicating these elements were present before indica-japonica divergence. The six other elements showed insertion polymorphisms among accessions, showing their recent insertion history or no critical positive effect of their insertion on the rice genome.

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).

Identification and confirmation of three neutral alleles conferring wide compatibility in inter-subspecific hybrids of rice (Oryza sativa L.) using near-isogenic lines

Wide-compatibility varieties (WCVs) are a special class of rice germplasm that is able to produce fertile hybrids when crossed to both indica and japonica subspecies. Previous studies determined 'Dular' and 02428 as two WCVs and identified a number of QTLs as having large effects on fertility of inter-subspecific hybrids. In this study, we developed five near-isogenic lines (NILs) for three of the QTLs, f5, f6 and S5, by backcrossing and marker-assisted selection, using "Dular" and 02428 as the donors and "Zhenshan 97" as the recipient. Three of the NILs each carried one introgressed allele, and two NILs each carried two introgressed alleles in combinations. The NILs were testcrossed to an indica tester "Nanjing 11" and a japonica tester "Balilla". The results showed that the f5 allele from "Dular"(f5-Du) is a neutral allele conferring wide compatibility, with a large effect on both pollen and spikelet fertility, and the f6 allele from "Dular" (f6-Du) is a neutral allele for spikelet fertility with smaller effect. The S5 allele from 02428 (S5-08) was confirmed to be a neutral allele for spikelet fertility. It is likely that f6 and S5 are the same locus as deduced by their genomic locations and effects. The results also showed that even in combination, two neutral alleles of different loci were not able to produce normal fertility hybrids in typical indica-japonica crosses. The implications of the findings in rice breeding programs are discussed.

Utilization of Diverse Rice Ecotypes in Heterosis Breeding.

A classification based on RFLP markers and morphological trait index was conducted in four photoperiod- and/or thermo-sensitive genic male sterile rice (PGMS or TGMS) lines and 49 rice varieties of nine different ecotypic origins. The results indicated that the combination of the two classification methods enabled to classify clearly and similarly the materials into two groups, indica and japonica. The 49 varieties were test-crossed to the four PGMS or TGMS lines used as female parents. Grain yield of the test-crossed F1s of diverse ecotypes exhibited remarkable differences, especially in three parameters, i.e. number of combinations over a check, extent of standard heterosis and heterobeltiosis. In general, the first two showed higher values in indica-japonica crosses than in crosses between two different ecotypes within the same subspecies. For each female parent except for LS2S, there were corresponding optimum ecotypes of male parents. In terms of grain yield advantages and above-mentioned parameters, patterns of heterotic combinations were as follows: Peiai64S in combination with japonica varieties of northeastern China, restorers of japonica rice hybrids, or japonica varieties of North China; N422S in combination with indica varieties of China, indica varieties of Korea; 108S with American new varieties, japonica varieties of northeastern China, and indica varieties of China. LS2S showed a lower general combining ability.

Genetic analysis and gene mapping of a rice few-tillering mutant in early backcross populations (Oryza sativa L.)

A rice mutant, G069, characteristic of few tiller numbers, was found in anther culture progeny from the F(1) hybrid between an indica-japonica cross, Gui630x02428. The mutant has another two major features: delayed tillering development and yellowing apex and margin on the mature leaves. As a donor parent, G069 was further backcrossed with the recurrent parent,02428, for two turns to develop aBC (2)F(2) population. Genetic analysis in the BC (2)F(2) population showed that the traits of few-tillering and yellowing apex and margin on the mature leaves were controlled by one recessive gene. A pool of equally mixed genomic DNA, from few-tillering individual plants in BC (2)F(2), was constructed to screen polymorphism with simple sequence repeat (SSR) markers in comparison with the 02428 genome. One SSR marker and three restriction fragment length polymorphism (RFLP) markers were found possibly linked with the recessive gene. By using these markers, the gene of few-tillering was mapped on chromosome 2 between RFLP marker C424 and S13984 with a genetic distance of 2.4 cM and 0.6 cM, respectively. The gene is designated ft1.

Abundant Polymorphism in the Flanking Regions of Two Loci for Basic PR-1 Proteins as Markers for indica-aponica Differentiation in Rice (Oryza sativa L.).

In an attempt to use new sources of disease resistance, genomic sequences of two genes Rpr1-1(t) and Rpr1-2(t) of PR-1 proteins, which are one of the major pathogenesis-related (PR) proteins in plants, were obtained by TAIL-PCR on the basis of a registered genomic and from expressed sequence tag (EST) data, respectively. The genetic loci of Rpr1-1(t) and Rpr1-2(t) were identified on chromosome 1 and 5, respectively, using recombinant inbred lines (RILs) from and an indica-japonica cross. Abundant sequence polymorphism between alleles of indica and japonica cultivars were found in the two genes. Polymorphic insertions including the Tnr1 (transposable element in rice #1) sequence in indica cultivars and a minor insertion in japonica cultivars were detected in the ORF flanking regions of Rpr1-1(t). In the ORF flanking regions of Rpr1-2(t), a unique insertion of 827 dp was found in japonica cultivars, while two insertions were found in an indica cultivar, Milyang 23. Despite such diversity in the flanking regions, only a few amino-acid residues were different between the alleles. Based on a test for polymorophism among some cultivars, it was found that variation of inserted/deleted sequences may provide a useful molecular tool for investigating indica-japonica differentiation in rice.

Gamete Abortion Locus Detected by Segregation Distortion of Isozyme Locus Pgi1 in indica-japonica Hybrids of Rice (Oryza sativa L.).

In an attempt to stabilize the pollen fertility of indicajaponica hybrids of rice by the incorporation of segregation-distortion-neutral alleles, a locus causing distorted segregation of male gamete in indica-japonica hybrids was identified with an isozyme marker gene Pgi1 on chromosome 3. In seven F2 populations of indica-japonica crosses and 25 F3 lines from IR 36/Kamairazu, the japonica-type allele Pgi1-2 was transmitted via male gamete with a ratio of 0.29 to 0.38 instead of the normal one of 0.5 due to the presence of a gametophyte gene ga14. The recombination value between the locus ga14 and the marker gene Pgi1 was estimated to be 0.29 by using 57 F3 lines of IR 36/Kamairazu. The male gamete transmission rate(k) of ga14-j was estimated to be 0.13∼0.22 against the ga14-i instead of the normal one of 0.5 based on the data from the F2 and F3 lines. The locus ga14 which reduced the frequency of male gametes harboring ga14-j in some indica-japonica hybrids was located in the order of Pgi1∼ga14∼bc1 on chromosome 3.

Epistatic QTLs affecting hybrid breakdown in recombinant inbred populations derived from indica-japonica crosses

Epistatic QTLs affecting hybrid breakdown in recombinant inbred populations derived from indica-japonica crosses

Genetic effects and genotype × environment interactions for cooking quality traits in Indica-japonica crosses of rice (Oryza sativa L.)

Indica-japonica hybridization is an important approach for developing superior performing hybrids in rice (Oryza sativa L.). In view of the scanty information available on cooking quality characters in indica-japonica crosses, an investigation was undertaken to estimate genetic and genotype × environment variance and covariance components of amylose content, gel consistency and alkali digestion value, and to determine the relative importance of direct genetic effects, maternal genetic effects and cytoplasmic effects in the genetic variations of the three quality characters. Two indica photo-sensitive genic male sterile (PGMS) lines and four japonica varieties were used as parents to make crosses. Genetic model with genotype × environment interactions for triploid endosperm was used for genetic studies of the three cooking quality characters. Variance component analysis revealed that genetic variations of the three characters were mainly attributable to direct additive and maternal additive effects, and the three traits had significant direct and maternal heritabilities. Genotype × environment interactions were mainly dominance × environment (including direct dominance × environment and maternal dominance × environment) and cytoplasm × environment interactions. Environment factors could only affect the expression extent of dominant genes, without changing their directions. Predicted values of genetic effects indicated that the parental lines, ‘VI-70’ and ‘H9304-1’, appeared to be best for amylose content, ‘T 1950’ and ‘Suxuan’ appeared to be best for gel consistency and alkali digestion value.

Two new loci for hybrid sterility in cultivated rice (Oryza sativa L.)

Female gamete abortion in Indica-Japonica crosses of rice was earlier identified to be due to an allelic interaction at the S-5 locus on chromosome 6. Recently, in other crosses of rice, similar allelic interactions were found at loci designated as S-7 and S-8, located on chromosomes 7 and 6 respectively. All of them are independent of each other. At the S-5 locus, Indica and Japonica rice have S-5 (i) and S-5 (j) alleles respectively and Javanicas, such as Ketan Nangka, have a neutral allele S-5 (n) .The S-5 (i) /S-5 (j) genotype is semi-sterile due to partial abortion of female gametes carrying S-5 (j) , but both the S-5 (n) /S-5 (i) and S-5 (n) /S-5 (j) genotypes are fertile. The S-5 (n) allele is thus a "wide-compatibility gene" (WCG), and parents homozygous for this allele are called wide-compatible varieties (WCV). Such parents when crossed with Indica or Japonica varieties do not show F1 hybrid sterility. Wide-compatible parents have been used to overcome sterility barriers in crosses between Indica and Japonica rice. However, a Javanica variety, Ketan Nangka (WCV), showed typical hybrid sterility when crossed to the Indian varieties N22 and Jaya. Further, Dular, another WCV from India, showed typical hybrid sterility when crossed to an IRRI line, IR2061-628-1-6-4-3(IR2061-628). By genetic analyses using isozyme markers, a new locus causing hybrid sterility in crosses between Ketan Nangka and the Indicas was located near isozyme loci Est-1 and Mal-1 on chromosome 4, and was designated as S-9. Another new locus for hybrid sterility in the crosses between Dular and the IR2061-628 was identified and was found linked to four isozyme loci, Sdh-1, Pox-2, Acp-1 and Acp-2, on chromosome 12. It was designated as S-15. On the basis of allelic interactions causing female-gamete abortion, two alleles were found at S-9, S-9 (kn) in Ketan Nangka and S-9 (i) in N22 and Jaya. In the heterozygote, S-9 (kn) /S-9 (i) , which was semisterile, female gametes carrying S-9 (kn) were aborted. The hybrid of Dular and IR2061-628, with a genetic constitution of S-15 (Du) /S-15 (i) , was semi-sterile and the female gametes carrying S-15 (Du) were aborted. A Japonica tester variety, Akihikari, and an Indica variety, IR36, were found to have neutral alleles, S-9 nand S-15 n, at these loci, in addition to S-7 nand at S-7. The accumulation of three neutral alleles into a breeding line should help solve the hybrid sterility problem in wide crosses of rice.

Identification of A New Locus S-16 Causing Hybrid Sterility in Native Rice Varieties (Oryza sativa L.) from Tai-hu Lake Region and Yunnan Province, China.

The presence of semi-sterile panicles in Indica-Japonica crosses of rice is due to an allelic interaction at a gamete abortion locus, S-5 on ( hromosome 6 The S 5 n allcle s referred to as wide compatibility gene" (WCG), since one of the parents carrying this allele does not show hybrid sterility in the Fl' However. Ketan Nangka, which is a donor of S-5 n, sholved typical h.vbrid sterility when it was crossed to local varieties from Taihu Lake, such as Fengjingdao and Feilaifeng, as weil as to those from Yun-nan Provinco, sueh as Dabai, Luanjiaolong and Xiaobaigu. A new locus for the sterilitv in the hvbrid between Ketan Nangka and the local varieties was detected near the Est-5 locus on Chromosome 1, and designated as S-16 following our recent report.s on S-7, S-8 . S-9 and S-15 .