A Review of Microsatellite Markers and Their Applications in Rice Breeding Programs to Improve Blast Disease Resistance

Over the last few decades, the use of molecular markers has played an increasing role in rice breeding and genetics. Of the different types of molecular markers, microsatellites have been utilized most extensively, because they can be readily amplified by PCR and the large amount of allelic variation at each locus. Microsatellites are also known as simple sequence repeats (SSR), and they are typically composed of 1–6 nucleotide repeats. These markers are abundant, distributed throughout the genome and are highly polymorphic compared with other genetic markers, as well as being species-specific and co-dominant. For these reasons, they have become increasingly important genetic markers in rice breeding programs. The evolution of new biotypes of pests and diseases as well as the pressures of climate change pose serious challenges to rice breeders, who would like to increase rice production by introducing resistance to multiple biotic and abiotic stresses. Recent advances in rice genomics have now made it possible to identify and map a number of genes through linkage to existing DNA markers. Among the more noteworthy examples of genes that have been tightly linked to molecular markers in rice are those that confer resistance or tolerance to blast. Therefore, in combination with conventional breeding approaches, marker-assisted selection (MAS) can be used to monitor the presence or lack of these genes in breeding populations. For example, marker-assisted backcross breeding has been used to integrate important genes with significant biological effects into a number of commonly grown rice varieties. The use of cost-effective, finely mapped microsatellite markers and MAS strategies should provide opportunities for breeders to develop high-yield, blast resistance rice cultivars. The aim of this review is to summarize the current knowledge concerning the linkage of microsatellite markers to rice blast resistance genes, as well as to explore the use of MAS in rice breeding programs aimed at improving blast resistance in this species. We also discuss the various advantages, disadvantages and uses of microsatellite markers relative to other molecular marker types.

[1]  R. Varshney,et al.  Molecular markers and their applications in wheat breeding , 1999 .

[2]  V. Shenoy,et al.  Marker-assisted breeding of Pi-1 and Piz-5 genes imparting resistance to rice blast in PRR78, restorer line of Pusa RH-10 Basmati rice hybrid , 2013 .

[3]  P. Cregan,et al.  Characterization of trinucleotide SSR motifs in wheat , 2002, Theoretical and Applied Genetics.

[4]  J. D. Austin,et al.  When technology meets conservation: increased microsatellite marker production using 454 genome sequencing on the endangered Okaloosa Darter (Etheostoma okaloosae). , 2010, The Journal of heredity.

[5]  Y. G. Yu,et al.  Divergence and allelomorphic relationship of a soybean virus resistance gene based on tightly linked DNA microsatellite and RFLP markers , 2004, Theoretical and Applied Genetics.

[6]  D. Mackill,et al.  Molecular Markers and Their Use in Marker-Assisted Selection in Rice , 2008 .

[7]  H. Yoshida,et al.  Development of PCR-based allele-specific and InDel marker sets for nine rice blast resistance genes , 2006, Theoretical and Applied Genetics.

[8]  M. Yano,et al.  Towards the Understanding of Complex Traits in Rice: Substantially or Superficially? , 2009, DNA research : an international journal for rapid publication of reports on genes and genomes.

[9]  Yulin Jia,et al.  Development of Monogenic and Digenic Rice Lines for Blast Resistance Genes Pi‐ta, Pi‐kh/Pi‐ks , 2010 .

[10]  M. Morgante,et al.  PCR-amplified microsatellites as markers in plant genetics. , 1993, The Plant journal : for cell and molecular biology.

[11]  H. Leung,et al.  GENETIC AND MOLECULAR DISSECTION OF BLAST RESISTANCE IN RICE USING RFLP, SIMPLE SEQUENCE REPEATS AND DEFENSE-RELATED CANDIDATE GENE MARKERS , 2003 .

[12]  M. Wingfield,et al.  Microsatellite discovery by deep sequencing of enriched genomic libraries. , 2009, BioTechniques.

[13]  Y. Cho,et al.  A novel gene, Pi40(t), linked to the DNA markers derived from NBS-LRR motifs confers broad spectrum of blast resistance in rice , 2007, Theoretical and Applied Genetics.

[14]  Wei Li,et al.  Identification of SSR markers for a broad-spectrum blast resistance gene Pi20(t) for marker-assisted breeding , 2008, Molecular Breeding.

[15]  Xinping C. Zhao,et al.  Characterization and genetic mapping of a short, highly repeated, interspersed DNA sequence from rice (Oryza sativa L.) , 1992, Molecular and General Genetics MGG.

[16]  S. Fukai,et al.  Molecular Breeding for Rainfed Lowland Rice in the Mekong Region , 2005 .

[17]  A. McClung,et al.  SSR Markers Closely Linked to the Pi-z Locus are Useful for Selection of Blast Resistance in a Broad Array of Rice Germplasm , 2006, Molecular Breeding.

[18]  Ashutosh Kumar Singh,et al.  Incorporation of blast resistance into “PRR78”, an elite Basmati rice restorer line, through marker assisted backcross breeding , 2012 .

[19]  P. Bentzen,et al.  Microsatellites: genetic markers for the future , 1994, Reviews in Fish Biology and Fisheries.

[20]  A. Davierwala,et al.  Potential of (GATA)n microsatellites from rice for inter- and intra-specific variability studies , 2001, BMC Evolutionary Biology.

[21]  .. S.Farooq,et al.  Molecular Markers in Plant Breeding-II. Some Pre-requisites for Use , 2002 .

[22]  D. Mackill,et al.  Molecular mapping and marker-assisted selection for major-gene traits in rice. , 2001 .

[23]  Transferability of microsatellite and sequence tagged site markers in Oryza species. , 2003, Hereditas.

[24]  R. Chauhan,et al.  RAPD and Pathotype Analyses of Magnaporthe grisea Populations from the north-western Himalayan Region of India , 2002 .

[25]  Ashutosh Kumar Singh,et al.  SNPs in stress-responsive rice genes: validation, genotyping, functional relevance and population structure , 2012, BMC Genomics.

[26]  M. Yano,et al.  The Pib gene for rice blast resistance belongs to the nucleotide binding and leucine-rich repeat class of plant disease resistance genes. , 1999, The Plant journal : for cell and molecular biology.

[27]  T. Sharma,et al.  Development and validation of microsatellite markers linked to the rice blast resistance gene Pi-z of Fukunishiki and Zenith , 2008, Euphytica.

[28]  D. Mackill,et al.  Evaluation of Genetic Diversity in Rice Subspecies Using Microsatellite Markers , 2002 .

[29]  P. Cregan,et al.  Automated sizing of fluorescent-labeled simple sequence repeat (SSR) markers to assay genetic variation in soybean , 1997, Theoretical and Applied Genetics.

[30]  M. Stephens,et al.  Inference of population structure using multilocus genotype data: linked loci and correlated allele frequencies. , 2003, Genetics.

[31]  E. D. Earle,et al.  Nuclear DNA content of some important plant species , 1991, Plant Molecular Biology Reporter.

[32]  A. Parco,et al.  Fine mapping and DNA marker-assisted pyramiding of the three major genes for blast resistance in rice , 2000, Theoretical and Applied Genetics.

[33]  Tao Wu,et al.  Genetic improvement of resistance to blast and bacterial blight of the elite maintainer line Rongfeng B in hybrid rice (Oryza sativa L.) by using marker-assisted selection , 2012 .

[34]  G. Lu,et al.  Two broad-spectrum blast resistance genes, Pi9(t) and Pi2(t), are physically linked on rice chromosome 6 , 2002, Molecular Genetics and Genomics.

[35]  R. Rathour,et al.  Virulence structure of theMagnaporthe grisea rice population from the northwestern Himalayas , 2006, Phytoparasitica.

[36]  Jeremy D. Edwards,et al.  Development and evaluation of a high-throughput, low-cost genotyping platform based on oligonucleotide microarrays in rice , 2008, Plant Methods.

[37]  S. Tanksley,et al.  Abundance, variability and chromosomal location of microsatellites in wheat , 1995, Molecular and General Genetics MGG.

[38]  Robert J Henry,et al.  Potential of SSR markers for plant breeding and variety identification in Australian barley germplasm , 2003 .

[39]  Lin Liu,et al.  Comparison of Next-Generation Sequencing Systems , 2012, Journal of biomedicine & biotechnology.

[40]  P. Pulkkinen,et al.  Microsatellite sequences in a conifer, Pinus sylvestris. , 1995, Genome.

[41]  X. C. Liu,et al.  SSR heterogenic patterns of parents for marking and predicting heterosis in rice breeding , 1998, Molecular Breeding.

[42]  Ling Wang,et al.  Fine Mapping of the Blast Resistance Gene Pi15, Linked to Pii, on Rice Chromosome 9 , 2003 .

[43]  M. Smale,et al.  Molecular tools in plant genetic resources conservation: a guide to the technologies. , 1997 .

[44]  J. Jiang,et al.  Identification of a 118-kb DNA fragment containing the locus of blast resistance gene Pi-2(t) in rice , 2002, Molecular Genetics and Genomics.

[45]  Genetic dissection of rice blast resistance by QTL mapping approach using an F3 population , 2013, Molecular Biology Reports.

[46]  M. Rakoczy-Trojanowska,et al.  Characteristics and a comparison of three classes of microsatellite-based markers and their application in plants. , 2004, Cellular & molecular biology letters.

[47]  L. Lipovich,et al.  Mapping and genome organization of microsatellite sequences in rice (Oryza sativa L.) , 2000, Theoretical and Applied Genetics.

[48]  M. Rai,et al.  Microsatellite markers: an overview of the recent progress in plants , 2011, Euphytica.

[49]  N. Gemmell,et al.  Fast, cost-effective development of species-specific microsatellite markers by genomic sequencing. , 2009, BioTechniques.

[50]  J. Weber.  Informativeness of human (dC-dA)n.(dG-dT)n polymorphisms. , 1990, Genomics.

[51]  F. Lin,et al.  Identification and mapping of Pi41, a major gene conferring resistance to rice blast in the Oryza sativa subsp. indica reference cultivar, 93-11 , 2009, Theoretical and Applied Genetics.

[52]  Shen Chen,et al.  Genetic and physical mapping of Pi37(t), a new gene conferring resistance to rice blast in the famous cultivar St. No. 1 , 2005, Theoretical and Applied Genetics.

[53]  M. Morgante,et al.  Identification and characterization of microsatellites in Norway spruce (Picea abies K.). , 1997, Genome.

[54]  M. Litt,et al.  A hypervariable microsatellite revealed by in vitro amplification of a dinucleotide repeat within the cardiac muscle actin gene. , 1989, American journal of human genetics.

[55]  Huanming Yang,et al.  A Draft Sequence of the Rice Genome (Oryza sativa L. ssp. indica) , 2002, Science.

[56]  G. Eizenga,et al.  Association mapping of yield and its components in rice cultivars , 2007, Molecular Breeding.

[57]  Liu Shi.  Improvement of Resistance to Rice Blast in Zhenshan 97 by Molecular Marker-aided Selection , 2003 .

[58]  H. Matsumura,et al.  Large-scale DNA polymorphism study of Oryza sativa and O. rufipogon reveals the origin and divergence of Asian rice , 2007, Theoretical and Applied Genetics.

[59]  W. Powell,et al.  How much effort is required to isolate nuclear microsatellites from plants? , 2003, Molecular ecology.

[60]  Looking back to move forward - a personal perspective on pig molecular genetics from RFLPs to nextgen sequencing , 2012 .

[61]  M. Bharathi,et al.  Identification of Microsatellite (SSR) and RAPD Markers Linked to Rice Blast Disease Resistance gene in Rice (Oryza sativa L.) , 2011 .

[62]  J. L. Weber,et al.  Survey of plant short tandem DNA repeats , 1994, Theoretical and Applied Genetics.

[63]  Ryan D. Hernandez,et al.  Genome-Wide Patterns of Nucleotide Polymorphism in Domesticated Rice , 2007, PLoS genetics.

[64]  Xinqiong Liu,et al.  The Blast Resistance Gene Pi37 Encodes a Nucleotide Binding Site–Leucine-Rich Repeat Protein and Is a Member of a Resistance Gene Cluster on Rice Chromosome 1 , 2007, Genetics.

[65]  Haofeng Chen,et al.  HighSSR: high-throughput SSR characterization and locus development from next-gen sequencing data , 2012, Bioinform..

[66]  J. Staub,et al.  Genetic Markers, Map Construction, and Their Application in Plant Breeding , 1996 .

[67]  D. Fujita,et al.  Development of pyramided lines with two resistance genes, Pish and Pib, for blast disease (Magnaporthe oryzae B. Couch) in rice (Oryza sativa L.) , 2010 .

[68]  M. Morgante,et al.  Microsatellites are preferentially associated with nonrepetitive DNA in plant genomes , 2002, Nature Genetics.

[69]  Gabriel O. Romero,et al.  Genetic fingerprinting: Advancing the frontiers of crop biology research , 2009 .

[70]  H. Leung,et al.  Association between molecular markers and blast resistance in an advanced backcross population of rice , 2004, Theoretical and Applied Genetics.

[71]  M. Pandey,et al.  Inheritance of Blast Resistance and Associated Microsatellite Markers in Rice Cultivar ‘Laxmi’ , 2007 .

[72]  G. Eizenga,et al.  Molecular diversity and genome-wide linkage disequilibrium patterns in a worldwide collection of Oryza sativa and its wild relatives , 2008, Euphytica.

[73]  M. Wang,et al.  Microsatellite Markers in Plants and Insects. Part I: Applications of Biotechnology , 2009 .

[74]  R. Varshney,et al.  The development and use of microsatellite markers for genetic analysis and plant breeding with emphasis on bread wheat , 2000, Euphytica.

[75]  Y. Cho,et al.  The pi40 gene for durable resistance to rice blast and molecular analysis of pi40-advanced backcross breeding lines. , 2009, Phytopathology.

[76]  Zhonghua Wang,et al.  Rapid survey for presence of a blast resistance gene Pi-ta in rice cultivars using the dominant DNA markers derived from portions of the Pi-ta gene , 2007 .

[77]  M. Ismail,et al.  Blast resistance in rice: a review of conventional breeding to molecular approaches , 2012, Molecular Biology Reports.

[78]  J. Tohme,et al.  Identification of microsatellite markers linked to the blast resistance gene Pi-1(t) in rice , 2008, Euphytica.

[79]  R. Tarchini,et al.  A Single Amino Acid Difference Distinguishes Resistant and Susceptible Alleles of the Rice Blast Resistance Gene Pi-ta , 2000, Plant Cell.

[80]  J. Bohlmann,et al.  Efficient Genetic Mapping of Single Nucleotide Polymorphisms Based Upon DNA Mismatch Digestion , 2005, Molecular Breeding.

[81]  K. Suwabe,et al.  Identification and characterization of microsatellites in eggplant , 2003 .

[82]  A. McClung,et al.  Development of DNA Markers Suitable for Marker Assisted Selection of Three Pi Genes Conferring Resistance to Multiple Pyricularia grisea Pathotypes , 2004 .

[83]  W. Chaoyang,et al.  Improvement of Resistance to Rice Blast in Zhenshan 97 by Molecular Marker-aided Selection , 2003 .

[84]  Jin Xue-hui.  Identification of Rice Blast Resistance Gene Pi-1 in Some Japonica Rice Varieties with SSR Marker , 2006 .

[85]  M. Latif,et al.  Analysis of simple sequence repeat markers linked with blast disease resistance genes in a segregating population of rice (Oryza sativa). , 2011, Genetics and molecular research : GMR.

[86]  N E Morton,et al.  Parameters of the human genome. , 1991, Proceedings of the National Academy of Sciences of the United States of America.

[87]  P. N. Cheremisinoff,et al.  Applications of biotechnology , 1985 .

[88]  P. Jarne,et al.  Microsatellites, from molecules to populations and back. , 1996, Trends in ecology & evolution.

[89]  O. Panaud,et al.  Frequency of microsatellite sequences in rice (Oryza sativa L.). , 1995, Genome.

[90]  Amanda J. Garris,et al.  Genetic Structure and Diversity in Oryza sativa L. , 2005, Genetics.

[91]  A. Prabhu,et al.  Tagging microsatellite marker to a blast resistance gene in the irrigated rice cultivar Cica-8 , 2012 .

[92]  T. Mackay.  The genetic architecture of quantitative traits. , 2001, Annual review of genetics.

[93]  L. Borges,et al.  Recuperación del genoma del padre recurrente en un programa de retrocruzas asistido por marcadores en arroz , 2006 .

[94]  M. Blair,et al.  Microsatellite marker development, mapping and applications in rice genetics and breeding , 1997, Plant Molecular Biology.

[95]  M. Yano,et al.  Saturation mapping with subclones of YACs: DNA marker production targeting the rice blast disease resistance gene, Pi-b , 1997, Theoretical and Applied Genetics.

[96]  R. Varshney,et al.  Cereal Genomics: An Overview , 2004 .

[97]  A. Rossman,et al.  Pyricularia Grisea the Correct Name for the Rice Blast Disease Fungus , 1990 .

[98]  I. Ashikawa,et al.  Development of PCR-based SNP markers for rice blast resistance genes at the Piz locus , 2004, Theoretical and Applied Genetics.

[99]  N. Shimizu,et al.  Construction of an 800-kb contig in the near-centromeric region of the rice blast resistance gene Pi-ta 2 using a highly representative rice BAC library , 1997, Molecular and General Genetics MGG.

[100]  D. Tautz.  Hypervariability of simple sequences as a general source for polymorphic DNA markers. , 1989, Nucleic acids research.

[101]  Songnian Hu,et al.  A Genome-wide Microsatellite Polymorphism Database for the Indica and Japonica Rice , 2007, DNA research : an international journal for rapid publication of reports on genes and genomes.

[102]  W. Zhai,et al.  A B-lectin receptor kinase gene conferring rice blast resistance. , 2006, The Plant journal : for cell and molecular biology.

[103]  Takuji Sasaki,et al.  The map-based sequence of the rice genome , 2005, Nature.

[104]  A. Liston,et al.  Contributions of PCR-Based Methods to Plant Systematics and Evolutionary Biology , 1998 .

[105]  S. Mccouch,et al.  Molecular mapping of a new gene for resistance to rice blast (Pyricularia grisea Sacc.) , 2000, Euphytica.

[106]  H. Kanamori,et al.  Distinct evolutionary patterns of Oryza glaberrima deciphered by genome sequencing and comparative analysis , 2011, The Plant journal : for cell and molecular biology.

[107]  S. Gaillard,et al.  Identification and fine mapping of Pi33, the rice resistance gene corresponding to the Magnaporthe grisea avirulence gene ACE1 , 2003, Theoretical and Applied Genetics.

[108]  S Rozen,et al.  Primer3 on the WWW for general users and for biologist programmers. , 2000, Methods in molecular biology.

[109]  V. Gupta,et al.  Potential of molecular markers in plant biotechnology. , 2009 .

[110]  J. Todd,et al.  Microsatellites for linkage analysis of genetic traits. , 1992, Trends in genetics : TIG.

[111]  L. Talbert,et al.  Conversion of AFLP markers to sequence-specific PCR markers in barley and wheat , 1999, Theoretical and Applied Genetics.

[112]  T. Sharma,et al.  RESEARCH ARTICLE Open Access Development of genic-SSR markers by deep transcriptome sequencing in pigeonpea [Cajanus cajan (L.) Millspaugh] , 2022 .

[113]  B. S. Ahloowalia,et al.  Molecular techniques in crop improvement , 2002 .

[114]  X. Liu,et al.  Genetic and physical mapping of Pi36(t), a novel rice blast resistance gene located on rice chromosome 8 , 2005, Molecular Genetics and Genomics.

[115]  Xinqiong Liu,et al.  Identification and fine mapping of Pi39(t), a major gene conferring the broad-spectrum resistance to Magnaporthe oryzae , 2007, Molecular Genetics and Genomics.

[116]  A. Inagaki,et al.  Microsatellite DNA markers for rice chromosomes , 1996, Theoretical and Applied Genetics.

[117]  K. Zenbayashi,et al.  Pi35(t), a new gene conferring partial resistance to leaf blast in the rice cultivar Hokkai 188 , 2006, Theoretical and Applied Genetics.

[118]  A. Torres.  Application of Molecular Markers for Breeding Disease Resistant Varieties in Crop Plants , 2010 .

[119]  Åsmund Bjørnstad,et al.  An overview of molecular marker methods for plants , 2006 .

[120]  S. Tanksley,et al.  Molecular mapping of genes for resistance to rice blast (Pyricularia grisea Sacc.) , 1996, Theoretical and Applied Genetics.

[121]  F. C. Victoria,et al.  Combinaciones de genes en arroz para el desarrollo de resistencia durable a Pyricularia grisea en Colombia , 2002 .

[122]  Ling Wang,et al.  Identification and characterization of a new blast resistance gene located on rice chromosome 1 through linkage and differential analyses. , 2004, Phytopathology.

[123]  Kenneth L. McNally,et al.  Rice Molecular Breeding Laboratories in the Genomics Era: Current Status and Future Considerations , 2008, International journal of plant genomics.

[124]  Wei Zhao,et al.  Gramene: a resource for comparative grass genomics , 2002, Nucleic Acids Res..

[125]  S. Tanksley,et al.  Abundance, polymorphism and genetic mapping of microsatellites in rice , 1993, Molecular and General Genetics MGG.

[126]  S. Fukuoka,et al.  Mapping of a blast field resistance gene Pi39(t) of elite rice strain Chubu 111 , 2008 .

[127]  Li-li Chen,et al.  A Receptor Kinase-Like Protein Encoded by the Rice Disease Resistance Gene, Xa21 , 1995, Science.

[128]  W. Zhai,et al.  Identification of Two Blast Resistance Genes in a Rice Variety, Digu , 2004 .

[129]  L. Andersson,et al.  The abundance of various polymorphic microsatellite motifs differs between plants and vertebrates. , 1993, Nucleic acids research.

[130]  H. Corke,et al.  Genetic diversity and population structure of a diverse set of rice germplasm for association mapping , 2010, Theoretical and Applied Genetics.

[131]  S. Tanksley,et al.  Tagging genes for blast resistance in rice via linkage to RFLP markers , 1991, Theoretical and Applied Genetics.

[132]  M. Dolan,et al.  The eight amino-acid differences within three leucine-rich repeats between Pi2 and Piz-t resistance proteins determine the resistance specificity to Magnaporthe grisea. , 2006, Molecular plant-microbe interactions : MPMI.

[133]  Ling Jiang,et al.  Fine mapping of qSTV11KAS, a major QTL for rice stripe disease resistance , 2011, Theoretical and Applied Genetics.

[134]  J. Enjalbert,et al.  Molecular markers to study genetic drift and selection in wheat populations , 1999 .

[135]  P. Sharp,et al.  A PCR‐based DNA marker for detection of mutant and normal alleles of the Wx‐D1 gene of wheat , 2001 .

[136]  M. Gowda,et al.  Identification of a major blast resistance gene in the rice cultivar ‘Tetep’ , 2004 .

[137]  S. Tanksley.  Molecular markers in plant breeding , 1983, Plant Molecular Biology Reporter.

[138]  A. McClung,et al.  Molecular markers linked to the blast resistance gene Pi-z in rice for use in marker-assisted selection , 2003, Theoretical and Applied Genetics.

[139]  B. Chattoo,et al.  Molecular mapping of a novel blast resistance gene Pi38 in rice using SSLP and AFLP markers , 2006 .

[140]  N. Young,et al.  DNA-Based Markers in Plants , 2001, Advances in Cellular and Molecular Biology of Plants.

[141]  M. Purugganan,et al.  The Extent of Linkage Disequilibrium in Rice (Oryza sativa L.) , 2007, Genetics.

[142]  B. Han,et al.  The Broad-Spectrum Blast Resistance Gene Pi9 Encodes a Nucleotide-Binding Site–Leucine-Rich Repeat Protein and Is a Member of a Multigene Family in Rice , 2006, Genetics.

[143]  M. Lorieux,et al.  Identification of five new blast resistance genes in the highly blast-resistant rice variety IR64 using a QTL mapping strategy , 2003, Theoretical and Applied Genetics.

[144]  D. Mackill.  Applications of Genomics to Rice Breeding , 2003 .

[145]  Zhonghua Wang,et al.  Rice Pi-ta gene Confers Resistance to the Major Pathotypes of the Rice Blast Fungus in the United States. , 2004, Phytopathology.

[146]  T. Sharma,et al.  High-resolution mapping, cloning and molecular characterization of the Pi-kh gene of rice, which confers resistance to Magnaporthe grisea , 2005, Molecular Genetics and Genomics.

[147]  Z. Ye,et al.  Comparative localization of three classes of cell wall proteins. , 1991, The Plant journal : for cell and molecular biology.

[148]  K. Sax,et al.  The Association of Size Differences with Seed-Coat Pattern and Pigmentation in PHASEOLUS VULGARIS. , 1923, Genetics.

[149]  L. Stein,et al.  Development and mapping of 2240 new SSR markers for rice (Oryza sativa L.). , 2002, DNA research : an international journal for rapid publication of reports on genes and genomes.

[150]  Zhonghua Wang,et al.  Development of Dominant Rice Blast Pi‐ta Resistance Gene Markers , 2002, Crop Science.

[151]  Ashutosh Kumar Singh,et al.  Genic non-coding microsatellites in the rice genome: characterization, marker design and use in assessing genetic and evolutionary relationships among domesticated groups , 2009, BMC Genomics.