[1]刘世名,李魏,戴良英.大豆疫霉根腐病抗性研究进展[J].大豆科学,2016,35(02):320-329.[doi:10.11861/j.issn.1000-9841.2016.02.0320]
　LIU Shi-ming,LI Wei,DAI Liang-ying.Progresses in Research on the Resistance of Soybean to Phytophythora Root Rot Caused by Phytophthora sojae[J].Soybean Science,2016,35(02):320-329.[doi:10.11861/j.issn.1000-9841.2016.02.0320]
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大豆疫霉根腐病抗性研究进展

《大豆科学》[ISSN:1000-9841/CN:23-1227/S] 卷: 第35卷 期数: 2016年02期 页码: 320-329 栏目: 出版日期: 2016-03-20
Title:
Progresses in Research on the Resistance of Soybean to Phytophythora Root Rot Caused by Phytophthora sojae
作者:
刘世名1; 2; 李魏1; 戴良英1
1．湖南农业大学 植物保护学院，湖南 长沙 410128；?
2.中国农业科学院 植物保护研究所，北京 100193
Author(s):
LIU Shi-ming1; 2;  LI Wei1;  DAI Liang-ying1
1.College of Plant Protection, Hunan Agricultural University, Changsha 410128, China;?
2.Institute of Plant Protection, Chinese Academy of Agricultural Sciences, Beijing 100193, China
关键词:
大豆; Phytophthora sojae; 疫霉根腐病; 抗病性; 抗性位点基因
Keywords:
Soybean;  Phytophthora sojae;  Phytophthora root rot;  Resistance;  Resistant loci and genes
DOI:
10.11861/j.issn.1000-9841.2016.02.0320
文献标志码:
A
摘要:
大豆疫霉根腐病由卵菌大豆疫霉菌（Phytophthora sojae）侵染引起，是一种大豆生产上危害非常严重的病害。大豆对疫霉菌的抗性表现有2种，一种是由单位点显性基因Rps控制的完全抗性，对大豆疫霉菌小种具有特异性，另一种为多基因控制的部分抗性，由数量性状位点控制，对所有大豆疫霉菌小种具有广谱抗性。迄今已定位、鉴定了26个单位点Rps基因，分别分布在第2（1个）、3（12个）、10（1个）、13（4个）、16（2个）、17（1个）、18（4个）及19（1个）条染色体上。其中，Rps1k提供了最稳定的PRR抗性。尽管在Rps2等位点区域检测到了抗病基因簇，但至今仅从Rps1k位点分离出了功能基因Rps1k-1和Rps1k-2，二者均为CC-NBS-LRR类型基因，且在细胞内可重组形成Rps1k-3。单位点基因的PRR抗性一般能维持8~15年左右，QTL控制的PRR抗性则较稳定、持久。有待不断发现新的PRR抗性资源和鉴定新的抗性基因，以及阐明大豆抗PRR的遗传与分子机制以长期、有效地防治大豆PRR。本文主要针对大豆的PRR抗性研究，尤其是近年在多个新Rps基因的鉴定、相关基因组学（转录组学）、小RNA及蛋白质组学上的研究，以及影响大豆PRR抗性的基因功能鉴定等方面所取得的进展进行了综述。
Abstract:
Abstract: Phytophythora root rot caused by Oomycete Phytophthora sojae, is a devastating pest in production of soybean (Glycine max L Merr).There are two types of soybean resistance to PRR: One is controlled by single gene (locus) with complete resistance to specific races, and the other one is controlled by multigenes, which are quantitative trait loci, with broad but partial resistance to all races of Phytophthora sojae.To date, totally, 26 single genes underlying complete resistance to PRR (resistance to Phytophthora sojae, Rps) have been identified, which distribute on Chromosomes 2 (1 gene), 3 (12 genes), 10 (1 gene), 13 (4 genes), 16 (2 genes), 17 (1 gene), 18 (4 genes) and 19 (1 gene), respectively Among them, Rps1k provides the most stable resistance to PRR. Some clusters of disease-resistant genes were detected around the Rps genes regions, but so far, only Rps1k-1 and Rps1k-2 in Rps1k locus were isolated among all the identified Rps genes loci Moreover, these 2 genes can further form Rps1k-3 through recombination Generally, the resistance of single gene can maintain about 8-15 years, whereas the resistance of QTL can remain much longer as well as stable.To long-term and effectively control PRR, screening more PRR-resistant soybean germplasm resources and identifying more new PRR-resistant genes are required, and finally the genetic and molecular mechanisms of soybean resistance to PRR would be elucidated. Here, this review focuses on the recent progresses in identification of novel Rps genes, research on genomics (transcriptome), small RNA and proteomics related with soybean resistance to PRR, and functional identification of the genes influencing the resistance to PRR.

参考文献/References:

［1］Grau C R, Dorrance A E, Bond J, et al. Fungal diseases［M］//Boerma H R, Specht J E.Soybeans: Improvement, proAduction and uses 3rd ed. Madison: American Society of Agronomy, 2004:679-763.

［2］Sugimoto T, Watanabe K, Yoshida S, et al. Field application of calcium to reduce Phytophthora stem rot of soybean, and calcium distribution in plants［J］. Plant Disease, 2010, 94: 812-819.
［3］Sugimoto T, Kato M, Yoshida S, et al.Pathogenic diversity of Phytophthora sojae and breeding strategies to develop Phytophthora-resistant soybeans［J］.Breeding Science, 2012, 61: 511-522.
［4］Gordon S G, Berry S A, St Martin S K, et al. Genetic analysis of soybean plant introductions with resistance to Phytophthora sojae［J］. Phytopathology, 2007, 97: 106-112.
［5］Gordon S G, Kowitwanich K, Pipatpongpinyo W, et al.Molecular marker analysis of soybean plant introductions with resistance to Phytophthora sojae［J］. Phytopathology, 2007, 97: 113-118.
［6］Zhang J, Sun S, Wang G, et al. Characterization of Phytophthora resistance in soybean cultivars/lines bred in Henan province［J］. Euphytica, 2014, 196: 375-384.
［7］Lin F, Zhao M, Ping J, et al. Molecular mapping of two genes conferring resistance to Phytophthora sojae in a soybean landrace PI 567139B［J］. Theoretical and Applied Genetics, 2013, 126: 2177-2185.
［8］Sugimoto T, Yoshida S, Kaga A, et al. Genetic analysis and identification of DNA markers linked to a novel Phytophthora sojae resistance gene in the Japanese soybean cultivar Waseshiroge［J］. Euphytica, 2011, 182: 133-145.［9］Sun S, Zhao S L, Wang J M, et al.Characterization and mapping of RpsYu25, a novel resistance gene to Phytophthora sojae［J］. Plant Breeding, 2011, 130: 139-143.
［10］Sun J, Li L H, Zhao J, et al. Genetic analysis and fine mapping of RpsJS, a novel resistancegene to Phytophthora sojae in soybean ［Glycine max(L) Merr］ ［J］.Theoretical and Applied Genetics, 2014, 127: 913-919.
［11］Wu X, Zhang B, Sun S, et al. Identification, genetic analysis and mapping of resistance to Phytophthora sojae of Pm28 in soybean［J］.Agricultura Scientia China, 2011, 10: 1506-1511.
［12］Zhang J, Xia C, Wang X, et al. Genetic characterization and fine mapping of the novel Phytophthora resistance gene in a Chinese soybean cultivar［J］. Theoretical and Applied Genetics, 2013, 126: 1555-1561.
［13］Zhang J, Xia C, Duan C, et al. Identification and candidate gene analysis of a novel Phytophthora resistance gene Rps10 in a Chinese soybean cultivar［J］. PLoS ONE, 2014, 8: e69799.
［14］郭娜, 胡冠军, 赵晋铭, 等. 一对单显性大豆抗疫霉根腐病基因的遗传分析及定位［J］.南京农业大学学报, 2015, 38(4): 532-537.(Guo N, Hu G J, Zhao J M, et al. Genetic analysis and mapping of a single dominant Phytophthora sojae resistance gene in soybean［J］.Journal of Nanjing Agricultural University, 2015, 38(4): 532-537.)
［15］武晓玲,周斌,孙石,等.大豆对大豆疫霉菌株Pm14抗性的遗传分析及基因定位［J］.中国农业科学, 2011, 44(3): 456-460. (Wu X L, Zhou B, Sun S, et al.Genetic analysis and mapping of resistance to Phytophthora sojae of Pm14 in soybean［J］. Scientia Agricultura Sinica, 2011, 44(3): 456-460.)
［16］姚海燕, 王晓鸣, 武小菲, 等.大豆品种早熟18抗疫霉根腐病基因的SSR分子标记［J］. 植物遗传资源学报, 2010, 11(2): 213-217. (Yao H Y, Wang X M, Wu X F, et al. Molecular mapping of Phytophthora resistance gene in soybean cultivar Zaoshu 18［J］. Journal of Plant Genetic Resources, 2010, 11(2): 213-217.)
［17］朱振东,霍云龙,王晓鸣,等.一个抗大豆疫霉根腐病新基因的分子鉴定［J］.作物学报, 2007, 33(1): 154-157.(Zhu Z D, Huo Y L, Wang X M, et al. Molecular identification of a novel Phytophthora resistance gene in soybean［J］. Acta Agronomic Sinica, 2007, 33(1): 154-157.)
［18］Bhattacharyya M K, Narayanan N N, Gao H, et al. Identification of a large cluster of coiled coil-nucleotide binding site-leucine rich repeat-type genes from the Rps1 region containing Phytophthora resistance genes in soybean［J］. Theoretical and Applied Genetics, 2005, 111: 75-86.
［19］Gao H, Narayanan N N, Ellison L, et al. Two classes of highly similar coiled coil-nucleotide binding-leucine rich repeat genes isolated from the Rps1-k. locus encode Phytophthora resistance in soybean［J］. Molecular Plant Microbe and Interaction, 2005, 18: 1035-1045.
［20］Tyler B M.Molecular basis of recognition between Phytophthora pathogens and their hosts［J］. Annual Review of Phytopathology, 2002, 40: 137-167.
［21］Schneider D S. Plant immunity and film noir: What gumshoe detectives can teach us about plant-pathogen interactions［J］. Cell, 2002, 109: 537-540.
［22］MacGregor T, Bhattacharyya M, Tyler B, et al. Genetic and physical mapping of Avr1a in Phytophthora sojae［J］. Genetics, 2002, 160: 949-959.
［23］Shan W, Cao M, Dan L, et al.The Avr1b locus of Phytophthora sojae encodes an elicitor and a regulator required for avirulence on soybean plants carrying resistance gene Rps1b［J］. Molecular Plant Microbe and Interaction, 2004, 17: 394-403.
［24］Song T, Kale S, Arredondo F, et al.Two RxLR avirulence genes in Phytophthora sojae determine soybean Rps1k-mediated disease resistance［J. Molecular Plant Microbe and Interaction, 2013, 26: 711-720.
［25］Gijzen M, F-rster H, Coffey M D, et al. Cosegregation of Avr4 and Avr6 in Phytophthora sojae［J］Canadian Journal of Botany, 1996, 74: 800-802
［26］Whisson S C, Drenth A, Maclean D J, et al.Phytophthora sojae avirulence genes, RAPD and RFLP markers used toconstruct a detailed genetic linkage map［J］. Molecular Plant Microbe and Interaction, 1995, 8: 988-995.
［27］Whisson S C, Basnayake S, Maclean D J, et al.Phytophthora sojae avirulence genes χ2?and Avr6 are locatedin a 24 kb, recombination-rich region of genomic DNA［J］. Fungal Genetics and Biology, 2004, 41: 62-74.
［28］Dou D, Kale S, Liu T, et al. Different domains of Phytophthora sojae effector Avr4/6 are recognized by soybean resistance genes Rps4 and Rps6［J］. Molecular Plant Microbe and Interaction, 2010, 23: 425-435.
［29］Qutob D, Chapman B P, Gijzen M. Transgenerational gene silencing causes gain of virulence in a plant pathogen［J］. Nature Communications, 2013, 4: 1349
［30］Na R, Yu D, Qutob D, et al. Deletion of the Phytophthora sojae avirulence gene Avr1d causes gain of virulence on Rps1d［J］. Molecular Plant Microbe and Interaction, 2013, 26: 969-976.
［31］Bernard R L, Smith P E, Kaufmann M J, et al. Inheritance of resistance to Phytophthora root and stem rot in soybean［J］.Agronomy Journal, 1957, 49: 391.
［32］Buzzell R I, Anderson T R.Inheritance and race reaction of a new soybean Rps1 allele［J］. Plant Disease, 1992, 76: 600-601.
［33］Mueller E H, Athow K L, Laviolette F A. Inheritance of resistance to four physiologic races of Phytophthora megasperma var sojae［J］. Phytopathology, 1978, 68: 1318-1322.
［34］Ploper L D, Athow K L, Laviolette F A. A new allele at Rps3 locus for resistance to Phytophthora megasperma f. sp glycinea in soybean［J］. Phytopathology, 1985, 75: 690-694.
［35］Weng C, Yu K, Anderson T R, et al.Mapping genes conferring resistance to Phytophthora root rot of soybean, Rps1a and Rps7［J］.Journal of Heredity, 2001, 92: 442-446.［36］Anderson T R, Buzzell R I Inheritance and linkage of the Rps7 gene for resistance to Phytophthora rot of soybean［J］ Plant Disease, 1992, 76: 958-959.
［37］Gordon S G, St. Martin S K, Dorrance A E.Rps8 maps to aresistance gene rich region on soybean molecular linkage group［J］. Food Crop Science, 2006, 46: 168-173.
［38］Sandhu D, Schallock K G, Rivera-Velez N, et al. Soybean Phytophthora resistance gene Rps8 maps closely to the Rps3 region［J］.Journal of Heredity, 2005, 96: 536-541.
［39］Kilen T C, Hartwig E E, Keeling B L. Inheritance of a secondmajor gene for resistance to Phytophthora root rot in soybeans［J］. Crop Science, 1974, 14: 260-262.［40］Graham M A, Marek L F, Shoemaker R C. PCR sampling of disease resistance-like sequences from a disease resistance gene cluster in soybean［J］.Therotical and Applied Genetics, 2002, 105: 50-57.
［41］Athow K L, Laviolette F A, Mueller E H, et al.A new major gene for resistance to Phytophthora megasperma var-sojae in soybean［J］. Phytopathology, 1980, 70: 977-980.
［42］Buzzell R I, Anderson T R. Another major gene for resistance to Phytophthora megasperma var sojae in soybean［J］. Soybean Genetics Newsletter, 1981, 18: 30-33.
［43］Athow K L, Laviolette F A.Rps6, a major gene for resistance to Phytophthora megasperma f sp glycinea in soybean［J］. Phytopathology, 1982, 72: 1564-1567.
［44］Zhang Z, Hao J, Yuan J, et al.Phytophthora-root rot resistancein soybean E00003［J］. Crop Science, 2014, 54: 492-499.
［45］Mideros S, Nita M, Dorrance A E. Characterization of components of partial resistance, Rps2, and root resistance to Phytophthora sojae in soybean［J］.Phytopathology, 2007, 97: 655-662.
［46］Burnham K D, Dorrance A E, van Toai T T, et al. Quantitative trait loci for partial resistance to Phytophthora sojae-in soybean［J］. Crop Science, 2003, 43: 1610-1617.
［47］Han Y, Teng W, Yu K, et al. Mapping QTL tolerance to Phytophthora-root rot in soybean using microsatellite and RAPD/SCAR derived markers［J］. Euphytica, 2008, 162: 231-239.
［48］Weng C, Yu K, Anderson T R, et al. A quantitative trait locus influencing tolerance to Phytophthora- root rot in the soybean cultivar Conrad［J］.Euphytica, 2007, 158: 81-86.
［49］Li X, Han Y, Teng W, et al.Pyramided QTL underlying tolerance to Phytophthora-root rot in mega-environments from soybean cultivars ‘Conrad’ and ‘Hefeng 25’［J］.Theoretical and Applied Genetics, 2010, 121: 651-658.
［50］Wang H, Waller L, Tripathy S, et al. Analysis of genes underlying soybean quantitative trait loci conferring partial resistance to Phytophthora sojae［J］. Plant Genome, 2010, 3: 23-40.
［51］Wang H, Wijeratne A, Wijeratne S, et al.Dissection of two soybean QTL conferring partial resistance to Phytophthora sojae through sequence and gene expression analysis［J］.BMC Genomics, 2012, 13: 428.
［52］Tucker D M, Saghai Maroof M A, Mideros S, et al. Mapping quantitative trait loci for partial resistance to Phytophthora sojae in a soybean interspecific cross［J］. Crop Science, 2010, 50: 628-635.
［53］Wu X, Zhou B, Zhao J, et al. Identification of quantitative trait loci for partial resistance to Phytophthora sojae in soybean［J］. Plant Breeding, 2011, 130: 144-149.
［54］Lee S, Mian M A R, McHale L K, et al. Identification of quantitative trait loci conditioning partial resistanceto Phytophthora sojae in soybean PI 407861A［J］. Crop Science, 2013, 53: 1022-1031
［55］Lee S, Mian M A, McHale L K, et al. Novel quantitative trait loci for partial resistance to Phytophthora sojae in soybean PI 398841［J］.Theoretical and Applied Genetics, 2013, 126: 1121-1132.
［56］Lee S, Mian M A R, Sneller C H, et al.Joint linkage QTL analyses for partial resistance to Phytophthora sojae in soybean using six nested inbred populations with heterogeneous conditions［J］.Theoretical and Applied Genetics, 2014, 127: 429-444.
［57］Sun J, Guo N, Lei J, et al. Association mapping for partial resistance to Phytophthora sojae in soybean (Glycine max［L］Merr) ［J］.Journal of Genetics, 2014, 932: 355-63.
［58］Hoffman T, Schmidt J S, Zheng X Y, et al. Isolation of ethylene-insensitive soybean mutants that are altered in pathogen susceptibility and geneforgene disease resistance［J］. Plant Physiology, 1999, 119: 935-949.
［59］Yu Y G, Buss G R, Maroof M A S. Isolation of a superfamily of candidate disease-resistance genes in soybean based on a conserved nucleotidebinding site［J］. Proceeding of the National Academy of Science, USA, 1996, 93: 11751-11756
［60］Sandhu D, Gao H, Cianzio S, et al. Deletion of a disease resistance nucleotide-binding-site leucine-rich- repeat-like sequence is associated with the loss of the Phytophthora resistance gene Rps4 in soybean［J］. Genetics, 2004, 168: 2157-2167.
［61］Gajendran K, Gonzales M D, Farmer A, et al.Phytophthora- functional genomics database (PFGD): Functional genomics of phytophthora-plant interactions［J］. Nucleic Acids Research, 2006, 34: 465-470.
［62］Narayanan N N, Grosic S, Tasma I M, et al.Identification of candidate signaling genes including regulators of chromosome condensation 1 protein family differentially expressed in the soybean-Phytophthora sojae-interaction［J］. Theoretical and Applied Genetics, 2009, 118: 399-412.
［63］Tyler B M, Jiang R H Y, Zhou L C, et al. Functional genomics and bioinformatics of the Phytophthora sojae-soybean interaction［M］//Gustafson J P, Taylor J, Stacey G. Genomics of disease. New York:Springer, 2007: 67-78.
［64］Zhou L, Mideros S X, Bao L, et al. Infection and genotype remodel the entire soybean transcriptome［J］. BMC Genomics, 2009, 10: 49.
［65］Lin F,Zhao M, Baumann D D, et al. Molecular response to the pathogen Phytophthora sojae among ten soybean near isogenic lines revealed by comparative transcriptomics［J］.BMC Genomics, 2014, 15: 18.
［66］Wong J, Gao L, Yang Y, et al.Roles of small RNAs in soybean defense against Phytophthora sojae infection［J］.Plant Journal, 2014, 79: 928-940.
［67］Guo N, Ye W, Wu X, et al. Microarray profiling reveals microRNAs involving soybean resistance to Phytophthora sojae［J］. Genome, 2011, 54: 954-958.
［68］Li X, Wang X, Zhang S, et al. Identification of soybean microRNAs involved in soybean cyst nematode infection by deep sequencing［J］. PLoS One, 2012, 7: E39650
［69］Zhao J, Zhang Y, Bian X, et al. A comparative proteomics analysis of soybean leaves under biotic and abiotic treatments［J］. Molecular Biology Reports, 2013, 40: 1553-1562.
［70］Parniske M, Ahlborn B, Werner D. Isoflavonoid-inducible resistance to the phytoalexin glyceollin in soybean rhizobia［J］.Journal of Bacteriology, 1991, 173: 3432-3439.
［71］Graham T L, Graham M Y, Subramanian S, et al. RNAi silencing of genes for elicitation or biosynthesis of 5-deoxyisoflavonoids suppresses race-specific resistance and hypersensitive cell death in Phytophthora sojae infected tissues［J］. Plant Physiology, 2007, 144: 728-740.
［72］Yan Q, Cui X, Su L, et al. GmSGT1 is differently required for soybean Rps genes-mediated and basal resistance to Phytophthora sojae［J］. Plant Cell Reports, 2014, 33: 1275-1288.
［73］Xu P, Jiang L, Wu J, et al. Isolation and characterization of a pathogenesis-related protein 10 gene (GmPR10) with induced expression in soybean (Glycine max) during infection with Phytophthora sojae［J］. Molecular Biology Reports, 2014, 41: 4899-4909
［74］Jiang L, Wu J, Fan S, et al. Isolation and characterization of a novel pathogenesis-related protein gene (GmPRP) with induced expression in soybean (Glycine max) during infection with Phytophthora sojae［J］. PLoS One, 2015, 10: e0129932.［75］Dong L, Cheng Y, Wu J, et al.Overexpression of GmERF5, a new member of the soybean EAR motif-containing ERF transcription factor, enhances resistance to Phytophthora sojae in soybean［J］.Journal of Experimental Botany, 2015, 66: 2635-2647.?
［76］Borkowska M, Krzymowska M, Talarczyk A, et al. Transgenic potato plants expressing soybean beta-1,3endoglucanase gene exhibit an increased resistance to Phytophthora infestans［J］. Zeitschrift fur Naturforschung C, 1998, 53: 1012-1016.
［77］Guo P, Wang Y, Zhou X, et al.Expression of soybean lectin in transgenic tobacco results in enhanced resistance to pathogens and pests［J］. Plant Science, 2013, 211: 17-22.
［78］Liu T, Ye W, Ru Y, et al.Two host cytoplasmic effectors are required for pathogens is of Phytophthora sojae by suppression of host defenses［J］.Plant Physiology, 2011, 155: 490-501.
［79］Chai C, Lin Y, Shen D, et al.Identification and functional characterization of the soybean GmaPPO12 promoter conferring Phytophthora sojae induced expression［J］.PLoS One, 2013, 8: e67670.
［80］Sumit R, Sahu B B, Xu M, et al.Arabidopsis nonhost resistance gene PSS1 confers immunity against an oomycete and a fungal pathogen but not a bacterial pathogen that cause diseases in soybean［J］.BMC Plant Biology, 2012, 12: 87
［81］Zhang H, Wu Q, Cao S, et al.A novel protein elicitor (SsCut) from Sclerotinia sclerotiorum induces multiple defense responses in plants［J］.Plant Molecular Biology, 2014, 86: 495-511.
［82］Juvale P S, Hewezi T, Zhang C, et al. Temporal and spatial bean pod mottle virus-induced gene silencing in soybean［J］. Molecular Plant Pathology, 2012, 13: 1140-1148.
［83］Kandoth P K, Heinz R, Yeckel G, et al.A virus-induced gene silencing method to study soybean cyst nematode parasitism in Glycine max［J］. BMC Research Notes, 2013, 6: 255.
［84］Liu S, Kandoth P K, Warren S D, et al. A soybean cyst nematode resistance gene points to a new mechanism of plant resistance to pathogens［J］. Nature, 2012, 492: 256-260.
［85］Liu J, Graham M A, Pedley K F, et al. Gaining insight into soybean defense responses using functional genomics approaches［J］. Briefings Functional Genomics, 2015, 14: 283-290.
［86］Jacobs T B, LaFayette P R, Schmitz R J, et al. Targeted genome modifications in soybean with CRISPR/Cas9［J］.BMC Biotechnology, 2015, 15: 16.
［87］Sun X, Hu Z, Chen R, et al. Targeted mutagenesis in soybean using the CRISPR-Cas9 system［J］. Scientific Reports, 2015, 5: 10342
［88］Cook D E, Lee T G, Guo X, et al.Copy number variation of multiple genes at Rhg1 mediates nematode resistance in soybean［J］. Science, 2012, 338: 1206-1209.［89］Schmutz J, Cannon S B, Schlueter J, et al. Genome sequence of the palaeopolyploid soybean［J］. Nature, 2010, 462: 178-183.［90］Song Q, Hyten D L, Jia G, et al. Development and evaluation of SoySNP50K, a high density genotyping array for soybean［J］. PLoS One, 2013, 8: e54985.［91］Han Y, Zhao X, Cao G, et al. Genetic characteristics of soybean resistance to HG type 0 and HG type 1-2-3-5-7 of the cyst nematode analyzed by genome-wide association mapping［J］.BMC Genomics, 2015, 16: 598.

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备注/Memo

基金项目：湖南农业大学“神农学者”计划; 湖南农业大学青年科学基金（14QN19）。

第一作者简介：刘世名（1970-），男，博士，教授，主要从事大豆抗病基因克隆与基因功能研究。E-mail：smliuhn@yahoo.com。

更新日期/Last Update: 2016-04-04