243份云南普通小麦地方品种抗条锈病鉴定及分子标记检测

doi:10.3864/j.issn.0578-1752.2021.04.002

摘要/Abstract

摘要：

【目的】小麦条锈病由条形柄锈菌小麦专化型（Puccinia striiformis. f. sp. tritici,Pst）引起的气传性真菌病害,是世界范围内最具破坏性的小麦病害之一。发掘抗病资源和培育抗病品种是防治条锈病最经济有效的措施。通过鉴定和评价云南小麦地方品种对中国当前流行的条锈菌生理小种的抗性水平,综合分析可能携带的抗条锈病基因,为小麦抗病育种提供理论依据。【方法】利用当前中国小麦生产上毒性强、流行频率高的条锈菌生理小种CYR32和CYR34对243份云南小麦地方品种进行苗期抗病性鉴定,成株期使用CYR32、CYR33、CYR34和贵农致病类群等混合生理小种进行抗性鉴定,并利用小麦生产上重要抗条锈病基因Yr5、Yr10、Yr15、Yr18、Yr26、Yr28、Yr29、Yr30、Yr36、Yr39、Yr41、Yr48、Yr65、Yr67、Yr80和Yr81的紧密连锁标记对供试材料进行分子检测。【结果】在243份小麦材料中,18份种质对CYR32表现苗期抗性,32份种质对CYR34表现苗期抗性,对CYR32和CYR34均表现苗期抗病的有8份,占3.29%。结合苗期和成株期抗性鉴定,174份地方种质表现为稳定的成株期抗性,占71.6%,其中105份种质高抗条锈病。分子检测结果显示,携带Yr10、Yr18、Yr29、Yr30、Yr65和Yr81抗性基因的材料分别有48、44、4、6、4和101份。同时携带2、3和4个抗性基因的材料各有34、4和1份。所有供试材料均未检测到Yr5、Yr15、Yr26、Yr28、Yr36、Yr39、Yr41、Yr48、Yr67和Yr80。此外,74份地方种质未检测到以上所有抗性基因,其中58份具有成株期抗性,可能携带其他已知或新的条锈病抗性基因。【结论】云南小麦地方品种作为中国特有的小麦种质资源,具有优良的条锈病抗性。243份供试小麦品种对当前条锈菌流行小种抗性水平整体偏高,筛选得到174份具有稳定抗性的地方种质,同时有74份可能携带其他已知或未知的抗性基因,可作为进一步挖掘抗条锈病新基因或QTL的亲本来源。

关键词: 小麦地方品种, 条锈病, 抗病鉴定, 分子标记, Yr

Abstract:

【Objective】As one of the most destructive wheat diseases in the world, stripe rust is an airborne fungal disease caused by fungus Puccinia striiformis. f. sp. Tritici, (Pst). Breeding resistant cultivars by exploiting resistant genetic resources is the most economical and effective strategy to combat the rusts. To provide theoretical basis for the management of wheat stripe rust by identifying and evaluating the resistance level of Yunnan wheat landraces to the current predominant races of the pathogen, and synthetically screened resistance genes that may be carried in the tested materials. 【Method】In this study, 243 wheat landraces originated from Yunnan were inoculated with two highly toxic and prevalent stripe rust races CYR32 and CYR34 at seedling stage. At the adult plant stage of wheat, the mixture of the current predominant races of CYR32, CYR33, CYR34, and Guinong Pathogenic group was used as the inoculums to test the resistance of the tested wheat. The wheat landraces were screened with the molecular markers closely linked to known stripe rust resistance genes Yr5, Yr10, Yr15, Yr18, Yr26, Yr28, Yr29, Yr30, Yr36, Yr39, Yr41, Yr48, Yr65, Yr67, Yr80, and Yr81.【Result】The results showed that the 243 of the tested wheat materials, of which 18 were resistant to CYR32, 32 resistant to CYR34, and only 8 (3.29%) were resistant to both races at the seedling stage. Based on identification results of the seedling stage and the adult plant stage, 174 (71.6%) exhibited adult plant resistance, of which 105 were high resistance to stripe rust. The results of molecular detection indicated that 48, 44, 4, 6, 4, and 101 out of the 243 tested wheat materials carried the resistance genes Yr10, Yr18, Yr29, Yr30, Yr65, and Yr81, respectively. There were 34, 4 and, 1 simultaneously carrying 2, 3, and 4 resistance genes, respectively. Yr5, Yr15, Yr26, Yr28, Yr36, Yr39, Yr41, Yr48, Yr67, and Yr80 were not detectable in all of the wheat landraces. In addition, 74 landraces not detected any of the 16 resistance genes mentioned the above, and 58 exhibited adult plant resistance, which, presumably, may carry other known or new resistance genes to wheat stripe rust.【Conclusion】As a special wheat germplasm resource in China, Yunnan wheat landraces have excellent resistance to stripe rust. This study determined that resistance of the 243 tested wheat landraces to the current predominant races of the pathogen was high on the whole, and 174 wheat landraces with stable resistance were identified. Among them, 74 lines might carry other known or new resistance genes to wheat stripe rust, which could be used as parents’ sources for further exploration of new stripe rust resistance genes or QTL.

Key words: wheat landraces, stripe rust, resistance identification, molecular markers, Yr

习玲, 王昱琦, 杨修, 朱微, 陈国跃, 王益, 覃鹏, 周永红, 康厚扬. 243份云南普通小麦地方品种抗条锈病鉴定及分子标记检测[J]. 中国农业科学, 2021, 54(4): 684-695.

XI Ling, WANG YuQi, YANG Xiu, ZHU Wei, CHEN GuoYue, WANG Yi, QIN Peng, ZHOU YongHong, KANG HouYang. Evaluation of Resistance to Stripe Rust and Molecular Detection of Resistance Gene(s) in 243 Common Wheat Landraces from the Yunnan Province[J]. Scientia Agricultura Sinica, 2021, 54(4): 684-695.

0
/ / 推荐

导出引用管理器 EndNote|Reference Manager|ProCite|BibTeX|RefWorks

链接本文: https://www.chinaagrisci.com/CN/10.3864/j.issn.0578-1752.2021.04.002

https://www.chinaagrisci.com/CN/Y2021/V54/I4/684

图/表 3

表1

抗条锈病基因的分子标记及引物序列"

Yr基因 Yr gene	类型 Type	分子标记 Molecular marker	引物序列 Primer sequence (5′-3′)	文献Reference
Yr5	DM	Yr5_B	GGGAACACTTCACGATCA	[31]
		Yr5_B	AATTCCTTCATGCCTTCC
		Yr5_A	CGCTTAATTCCCCTTCCTTC
		Yr5_A	TGGCTCCTTATTCGTTCTCTTTC
	KASP	Yr5F	GAAGGTGACCAAGTTCATGCTGCGCCCCTTTTCGAAAAAATA
		Yr5H	GAAGGTCGGAGTCAACGGATTCTAGCATCAAACAAGCTAAATA
		Yr5R	ATGTCGAAATATTGCATAACATGG
Yr10	AFLP	SC200	CTGCAGAGTGACATCATACA	[32]
	AFLP	SC200	TCGAACTAGTAGATGCTGGC	[32]
	DM	Yr10-5	GGAAATGTGGCGGAGTACCA	[16]
		Yr10-5	CGGAAGGGAGAACCACTGTC
		Yr10-6	CAGCTTGACAAGGGCGAGTA
		Yr10-6	CGCTTGTCGCCAATTCCAAA
Yr15	DM	Y15K1_F2	GGAGATAGAGCACATTACAGAC	[33]
		uhw301R	GGAGATAGAGCACATTACAGAC	[33]
		WJL2F	CCTTGTGTGCTACCAGGGTT	[34]
		WJL2R	GGAACTCAAGCCCTTCTGCT	[34]
		WJL3F	AAAAGAGCTCGCCTCCTACG	[35]
		WJL3R	GCCATGATGAGATCGGGAGG	[35]
Yr18	STS	csLV34	GTTGGTTAAGACTGGTGATGG	[17]
	STS		TGCTTGCTATTGCTGAATAGT
	Cssfr1	L34DINT9F	TTGATGAAACCAGTTTTTTTTCTA
	Cssfr1	L34PLUSR	GCCATTTAACATAATCATGATGGA
Yr26	STS	Xwe173	GGGACAAGGGGAGTTGAAGC	[36]
Yr26	STS	Xwe173	GAGAGTTCCAAGCAGAACAC	[36]
Yr28	InDel	ZCZp343F	TGTGTCATGTTTGGTCGATAGG	[37]
	InDel	ZCZp342R	TCCTCCCTTGTAGCTTCACG
	DM	P175	GCACCGTCCTTCATCTCAGT
	DM	P176	TGCTTTTCCCCGTATCCCTT
Yr29	CAPS	csLV46	CGAGACGTCGTCTTCTCTAAC	[38]
Yr29	CAPS	csLV46	GTGTATGTGTTGATTCTCCTCG	[38]
Yr30	SSR	Xgwm533	GTTGCTTTAGGGGAAAAGCC	[39]
Yr30	SSR	Xgwm533	AAGGCGAATCAAACGGAATA	[39]
Yr36	DM	WKS1_150F	ATGGAGCTCCCACGAAACAAAC	[40]
		WKS1_620R	ACCTCCATGTTGCTCGCATTTGCT
		WKS1_182F	TAGCTCTTCGTGGCCAAAGG
		WKS1_150R	TGGCCACGAAGAGCTAAAGG
Yr39	SSR	Xgwm131	AATCCCCACCGATTCTTCTC	[41]
Yr39	SSR	Xgwm131	AGTTCGTGGGTCTCTGATGG	[41]
Yr41	SSR	Xgwm410	GCTTGAGACCGGCACAGT	[42]
	SSR	Xgwm410	CGAGACCTTGAGGGTCTAGA
	SSR	Xgwm374	ATAGTGTGTTGCATGCTGTGTG
	SSR	Xgwm374	TCTAATTAGCGTTGGCTGCC
Yr48	SSR	Xwmc727	CATAATCAGGACAGCCGCAC	[43]
	SSR	Xwmc727	TAGTGGCCTGATGTATCTAGTTGG
	SSR	Xwms291	CATCCCTACGCCACTCTGC
	SSR	Xwms291	AATGGTATCTATTCCGACCCG
Yr65	SSR	Xgwm18	GGTTGCTGAAGAACCTTATTTAGG	[19]
Yr65	SSR	Xgwm18	TGGCGCCATGATTGCATTATCTTC	[19]
Yr67	SSR	Xbarc182	CCATGGCCAACAGCTCAAGGTCTC	[44]
	SSR	Xbarc182	CGCAAAACCGCATCAGGGAAGCACCAAT
	SSR	Xcfa2040	TCAAATGATTTCAGGTAACCACTA
	SSR	Xcfa2040	TTCCTGATCCCACCAAACAT
Yr80	KASP	KASP_53113	GAAGGTGACCAAGTTCATGCTTGTACAATGACTCCTCGACTAACA	[20]
			GAAGGTCGGAGTCAACGGATTTGTACAATGACTCCTCGACTAACG
			GCCACGCAATATCACCATCG
Yr81	KASP	KASP_3077	GAAGGTGACCAAGTTCATGCTATTCCAAAGTAATTGGCAACAGGTTCA	[21]
			GAAGGTCGGAGTCAACGGATTCCAAAGTAATTGGCAACAGGTTCG
			TGTGGAGCGTGACAATGAGGAAGTT

表1

图1

图2

参考文献 62

[1]	GUPTA P K, MIR R R, MOHAN A, KUMAR J. Wheat genomics: Present status and future prospects. International Journal of Plant Genomics, 2008: 1-36.
[2]	WELLINGS C R. Global status of stripe rust: A review of historical and current threats. Euphytica, 2011,179(1):129-141.
[3]	LINE R F, CHEN X M. Successes in breeding for and managing durable resistance to wheat rusts. Plant Disease, 1995,79(12):1254-1255.
[4]	王琪琳, 张立, 魏国荣, 曾庆东, 赵杰, 王晓杰, 黄丽丽, 康振生. “西北-华北-长江中下游”条锈病流行区系当前小麦品种(系)抗条锈病性评价. 中国农业科学, 2010,43(14):2889-2896.
	WANG Q L, ZHANG L, WEI G R, ZENG Q D, ZHAO J, WANG X J, HUANG L L, KANG Z S. Evaluation of resistance of current wheat cultivars to stripe rust in Northwest China, North China and the Middle and Lower Reaches of Changjiang River Epidemic Area. Scientia Agricultura Sinica, 2010,43(14):2889-2896. (in Chinese)
[5]	LI J B, DUNDAS I, DONG C M, LI G R, TRETHOWAN R, YANG Z J, HOXHA S, ZHANG P. Identification and characterization of a new stripe rust resistance gene Yr83 on rye chromosome 6R in wheat. Theoretical and Applied Genetics, 2020,133(4):1095-1107.
[6]	KUMAR S, GOYAL A, MOHAN A, BALYAN H S, GUPTA P K. An integrated physical map of simple sequence repeats in bread wheat. Australian Journal of Crop Science, 2013,7:460-468.
[7]	LU Y M, LAN C X, LIANG S S, ZHOU X C, LIU D, ZHOU G, LU Q L, JING J X, WANG M N, XIA X C, HE Z H. QTL mapping for adult-plant resistance to stripe rust in Italian common wheat cultivars Libellula and Strampelli. Theoretical and Applied Genetics, 2009,119(8):1349-1359.
[8]	PAYNE P I, NIGHTINGALE M A, KRATTIGER A F, HOLT L M. The relationship between HMW glutenin subunit composition and the bread-making quality of British-grown wheat varieties. Journal of the Science of Food and Agriculture, 1987,40(1):51-65.
[9]	金善宝. 中国小麦品种及其系谱. 北京: 中国农业出版社, 1983: 366-381.
	JIN S B. Chinese Wheat Varieties and Their Genealogy.Beijing: China Agriculture Press, 1983: 366-381. (in Chinese)
[10]	韩德俊, 张培禹, 王琪琳, 曾庆东, 吴建辉, 周新力, 王晓杰, 黄丽丽, 康振生. 1980份小麦地方品种和国外种质抗条锈性鉴定与评价. 中国农业科学, 2012,45(24):5013-5023.
	HAN D J, ZHANG P Y, WANG Q L, ZENG Q D, WU J H, ZHOU X L, WANG X J, HUANG L L, KANG Z S. Identification and evaluation of resistance to stripe rust in 1980 wheat landraces and abroad germplasm. Scientia Agricultura Sinica, 2012,45(24):5013-5023. (in Chinese)
[11]	张二喜, 李金昌, 吕莉莉, 王伟. 小麦地方品种资源对条锈病的抗性鉴定及评价. 甘肃农业科技, 2006(7):9-12.
	ZHANG E X, LI J C, LÜ L L, WANG W. Evaluation of resistance to strike rust landrace wheat germplasm resources. Gansu Agricultural Science and Technology, 2006(7):9-12. (in Chinese)
[12]	蔺瑞明, 郄彦敏, 冯晶, 徐世昌. 中国小麦农家品种抗条锈病的鉴定与评价. 沈阳农业大学学报, 2010,41(5):535-539.
	LIN R M, QIE Y M, FENG J, XU S C. Identification of the yellow rust resistance gene-carrying wheat landraces in China. Journal of Shenyang Agricultural University, 2010,41(5):535-539. (in Chinese)
[13]	ZHENG S G, LI Y F, LU L, LIU Z H, ZHANG C H, AO D H, LI L R, ZHANG C Y, LIU R, LUO C P, WU Y, ZHANG L. Evaluating the contribution of Yr genes to stripe rust resistance breeding through marker-assisted detection in wheat. Euphytica, 2017,213(2):1-16.
[14]	YE X L, LI J, CHENG Y K, YAO F J, LONG L, YU C, WANG Y Q, WU Y, LI J, WANG J R, JIANG Q T, LI W, MA J, WEI Y M, ZHENG Y L, CHEN G Y. Genome-wide association study of resistance to stripe rust (Puccinia striiformis f. sp. tritici) in Sichuan wheat. BioMed Central Plant Biology, 2019,19(1):147.
[15]	管方念, 龙黎, 姚方杰, 王昱琦, 江千涛, 康厚扬, 蒋云峰, 李伟, 邓梅, 李豪, 陈国跃. 152份黄淮海麦区小麦农家品种抗条锈性评价及重要条锈病抗性基因的分子检测. 中国农业科学, 2020,53(18):3629-3637.
	GUAN F N, LONG L, YAO F J, WANG Y Q, JIANG Q T, KANG H Y, JIANG Y F, LI W, DENG M, LI H, CHEN G Y. Evaluation of resistance to stripe rust and molecular detection of important known Yr gene(s) of 152 Chinese Wheat Landraces from the Yellow and Huai River Valleys. Scientia Agricultura Sinica, 2020,53(18):3629-3637. (in Chinese)
[16]	LIU W, FRICK M, HUEL R, NYKIFORUK C L, WANG X M, GAUDET D A, EUDES F, CONNER R L, KUZYK A, CHEN Q, KANG Z S, LAROCHE A. The stripe rust resistance gene Yr10 encodes an evolutionary-conserved and unique CC-NBS-LRR sequence in wheat. Molecular Plant, 2014,7(12):1740-1755.
[17]	LAGUDAH E S, KRATTINGER S G, HERRERA-FOESSEL S, SINGH R P, HUERTA-ESPINO J, SPIELMEYER W, BROWN- GUEDIRA G, SELTER L L, KELLER B. Gene-specific markers for the wheat gene Lr34/Yr18/Pm38 which confers resistance to multiple fungal pathogens. Theoretical and Applied Genetics, 2009,119(5):889-898.
[18]	MOORE J W, HERRERA-FOESSEL S, LAN C, SCHNIPPENKOETTER W, AYLIFFE M, HUERTA-ESPINO J, LILLEMO M, VICCARS L, MILNE R, PERIYANNAN S, KONG X Y, SPIELMEYER W, TALBOT M, BARIANA H, PATRICK J W, DODDS P, SINGH R, LAGUDAH E. A recently evolved hexose transporter variant confers resistance to multiple pathogens in wheat. Nature Genetics, 2015,47(12):1494-1498.
[19]	CHENG P, XU S, WANG M N, SEE D R, CHEN X M. Molecular mapping of genes Yr64 and Yr65 for stripe rust resistance in hexaploid derivatives of durum wheat accessions PI 331260 and PI 480016. Theoretical and Applied Genetics, 2014,127(10):2267-2277.
[20]	NSABIYERA V, BARIANA H S, QURESHI N, WONG D, HAYDEN M J, BANSAL U K. Characterization and mapping of adult plant stripe rust resistance in wheat accession Aus27284. Theoretical and Applied Genetics, 2018,131(7):1459-1467.
[21]	GESSESE M, BARIANA H, WONG D, HAYDEN M, BANSAL U. Molecular mapping of stripe rust resistance gene Yr81 in a common wheat landrace Aus27430. Plant Disease, 2019,103(6):1166-1171.
[22]	曾学琦, 恩在诚, 伍绍云. 云南省小麦品种资源的特点及分布. 云南农业科技, 1989(5):3-6.
	ZENG X Q, EN Z C, WU S Y. Characteristics and distribution of wheat variety resources in Yunnan province. Yunnan Agricultural Science and Technology, 1989(5):3-6. (in Chinese)
[23]	李明菊. 云南省小麦条锈病流行体系的研究现状. 植物保护, 2004(3):30-33.
	LI M J. Current research situation on epidemic system of wheat stripe rust in Yunnan province. Plant Protection, 2004(3):30-33. (in Chinese)
[24]	QUAN W, HOU G L, CHEN J, DU Z Y, LIN F, GUO Y, LIU S, ZHANG Z J. Mapping of QTL lengthening the latent period of Puccinia striiformis in winter wheat at the tillering growth stage. European Journal of Plant Pathology, 2013,136(4):715-727.
[25]	LIU T G, PENG Y L, CHEN W Q, ZHANG Z Y. First detection of virulence in Puccinia striiformis f. sp. tritici in China to resistance genes Yr24 (= Yr26) present in wheat cultivar Chuanmai 42. Plant Disease, 2010,94(9):1163.
[26]	WU J H, WANG Q L, CHEN X M, WANG M J, MU J M, LV X N, HUANG L L, HAN D J, KANG Z S. Stripe rust resistance in wheat breeding lines developed for Central Shaanxi, an overwintering region for Puccinia striiformis f. sp. tritici in China. Canadian Journal of Plant Pathology, 2016,38(3):317-324.
[27]	刘博, 刘太国, 章振羽, 贾秋珍, 王保通, 高利, 彭云良, 金社林, 陈万权. 中国小麦条锈菌条中34号的发现及其致病特性. 植物病理学报, 2017,47(5):681-687.
	LIU B, LIU T G, ZHANG Z Y, JIA Q Z, WANG B T, GAO L, PENG Y L, JIN S L, CHEN W Q. Discovery and pathogenicity of CYR34, a new race of Puccinia striiformis f. sp. tritici in China. Acta Phytopathologica Sinica, 2017,47(5):681-687. (in Chinese)
[28]	LINE R F, QAYOUM A, Virulence, aggressiveness, Evolution and distribution of races of Puccinia striiformis(the cause of stripe of wheat) in North America, 1968-1987. Technical Bulletin, 1992: 1-54.
[29]	陈国跃, 姚琦馥, 刘亚西, 邓梅, 吴文雄, 何员江, 余马. 四川地方小麦品种条锈病抗性研究. 四川农业大学学报, 2013,31(1):1-8.
	CHEN G Y, YAO Q F, LIU Y X, DENG M, WU W X, HE Y J, YU M. Studies on yellow rust resistance of Sichuan wheat landraces. Journal of Sichuan Agricultural University, 2013,31(1):1-8. (in Chinese)
[30]	HILL-AMBROZ K L, BROWN-GUEDIRA G L, FELLERS J P. Modified rapid DNA extraction protocol for high throughput microsatellite analysis in wheat. Crop Science, 2002,42(6):2088-2091.
[31]	MARCHAL C, ZHANG J P, ZHANG P, FENWICK P, STEUEMAGEL B, ADAMSKI N M, BOYD L, MCINTOSH R, WULFF B B H , BERRY S , LAGUDAH E , UAUY C . BED-domain-containing immune receptors confer diverse resistance spectra to yellow rust. Nature Plants, 2018,4(9):662-668.
[32]	邵映田, 牛永春, 朱立煌, 翟文学, 徐世昌, 吴立人. 小麦抗条锈病基因Yr10的AFLP标记. 科学通报, 2001(8):669-672.
	SHAO Y T, NIU Y C, ZHU L H, ZHAI W X, XU S C, WU L R. AFLP markers of wheat stripe rust resistance gene Yr10. Chinese Science Bulletin, 2001(8):669-672. (in Chinese)
[33]	KLYMIUK V, YANIV E, HUANG L, RAATS D, FATIUKHA A, CHEN S S, FENG L H, FRENKEL Z, KRUGMAN T, LIDZBARSKY G, CHANG W, JÄÄSKELÄINEN M J , SCHUDOMA C , PAULIN L , LAINE P , BARIANA H , SELA H , SALEEM K , SØRENSEN C K , HOVMØLLER M S , DISTELFELD A , CHALHOUB B , DUBCOVSKY J , KOROL A B , SCHULMAN A H , FAHIMA . Cloning of the wheat Yr15 resistance gene sheds light on the plant tandem kinase- pseudokinase family. Nature Communications, 2018,9(1):178-202.
[34]	HE Y, FENG L H, JIANG Y, ZHANG L Q, YAN J, ZHAO G, WANG J R, CHEN G Y, WU B H, LIU D C, HUANG L, FAHIMA T. Distribution and nucleotide diversity of Yr15 in wild emmer populations and Chinese wheat germplasm. Pathogenetics, 2020,9(3):212.
[35]	HUANG L, FENG L H, HE Y, TANG Z Z, HE J S, SELA H, KRUGMAN T, FAHIMA T, LIU D C, WU B H. Variation in stripe rust resistance and morphological traits in wild emmer wheat populations. Agronomy, 2019,9(2):44.
[36]	WANG C M, ZHANG Y P, HAN D J, KANG Z S, LI G P, CAO A Z, CHEN P D. SSR and STS markers for wheat stripe rust resistance gene Yr26. Euphytica, 2008,159(3):359-366.
[37]	ZHANG C Z, HUANG L, ZHANG H F, HAO Q Q, LYU B, WANG M N, EPSTEIN L, LIU M, KOU C L, QI J, CHEN F J, LI M K, GAO G, NI F, ZHANG L Q, HAO M, WANG J R, CHEN X M, LUO M C, ZHENG Y L, WU J J, LIU D C, FU D L. An ancestral NB-LRR with duplicated 3'UTRs confers stripe rust resistance in wheat and barley. Nature Communications, 2019,10(1):4023.
[38]	REN Y, SINGH R P, BASNET B R, LAN C X, HUERTA-ESPINO J, LAGUDAH E S, PONCE-MOLINA L J, LAN C X. Identifcation and mapping of adult plant resistance loci to leaf rust and stripe rust in common wheat cultivar Kundan. Plant Disease, 2017,101:456-463.
[39]	HAYDEN M J, KUCHEL H, CHALMERS K J. Sequence tagged microsatellites for the Xgwm533 locus provide new diagnostic markers to select for the presence of stem rust resistance gene Sr2 in bread wheat (Triticum aestivum L.). Theoretical and Applied Genetics, 2004,109(8):1641-1647.
[40]	HUANG L, SELA H, FENG L H, CHEN Q J, KRUGMAN T, YAN J, DUBCOVSKY J, FAHIMA T. Distribution and haplotype diversity of WKS resistance genes in wild emmer wheat natural populations. Theoretical and Applied Genetics, 2016,129(5):921-934.
[41]	LIN F, CHEN X M. Genetics and molecular mapping of genes for race-specific all-stage resistance and non-race-specific high-temperature adult-plant resistance to stripe rust in spring wheat cultivar Alpowa. Theoretical and Applied Genetics, 2007,114(7):1277-1287.
[42]	LUO P G, HU X Y, REN Z L, ZHANG H Y, SHU K, YANG Z J. Allelic analysis of stripe rust resistance genes on wheat chromosome 2BS. Genome, 2008,51(11):922-927.
[43]	LOWE I, JANKULOSKI L, CHAO S, CHEN X M, SEE D, DUBCOVSKY J. Mapping and validation of QTL which confer partial resistance to broadly virulent post-2000 North American races of stripe rust in hexaploid wheat. Theoretical and Applied Genetics, 2011,123(1):143-157.
[44]	XU H X, ZHANG J, ZHANG P, QIE Y M, NIU Y C, LI H J, MA P T, XU Y F, AN D G. Development and validation of molecular markers closely linked to the wheat stripe rust resistance gene YrC591 for marker-assisted selection. Euphytica, 2014,198(3):317-323.
[45]	XU Q, WANG J F, ZHAO J R, XU J H, SUN S T, ZHANG H F, WU J J, TANG C L, KANG Z S, WANG X J. A polysaccharide deacetylase from Puccinia striiformis f. sp. tritici is an important pathogenicity gene that suppresses plant immunity. Plant Biotechnology Journal, 2020: 1-13.
[46]	李明菊, 杨子林, 杨莲, 顾中量, 吉勇, 韦丽莉, 张翼亮, 张培花, 宋文宏, 张庆, 赵剑锋, 刘太国. 部分小麦抗条锈基因及生产品种在云南的抗性表现. 植物保护, 2016,42(4):161-168.
	LI M J, YANG Z L, YANG L, GU Z L, JI Y, WEI L L, ZHANG Y L, ZHANG P H, SONG W H, ZHANG Q, ZHAO J F, LIU T G. Resistance of Yr genes and commercial wheat cultivars to yellow rust in Yunnan province. Plant Protection, 2016,42(4):161-168. (in Chinese)
[47]	李明菊, 伍少云. 云南小麦品质改良亲缘材料成株期抗条锈性评价. 麦类作物学报, 2006(1):113-116.
	LI M J, WU S Y. Assessment to adult-plant resistance to stripe rust of parent materials for quality improvement of wheat germplasms in Yunnan. Journal of Triticeae Crops, 2006(1):113-116. (in Chinese)
[48]	李菁, 姚方杰, 龙黎, 王昱琦, 叶雪玲, 邓梅, 蒋云峰, 李伟, 江千涛, 康厚扬, 陈国跃. 中国特有的3个普通小麦亚种抗条锈病评价及其抗性基因分子检测. 植物病理学报, 2020,50(4):426-441.
	LI J, YAO F J, LONG L, WANG Y Q, YE X L, DENG M, JIANG Y F, LI W, JIANG Q T, KANG H Y, CHEN G Y. Evaluation and molecular detection of stripe rust resistance in three subspecies of Chinese endemic wheat. Acta Phytopathologica Sinica, 2020,50(4):426-441. (in Chinese)
[49]	代君丽, 刘珂, 牛永春, 李洪连. 中国小麦地方品种抗条锈病基因推导. 河南农业科学, 2010(12):83-87.
	DAI J L, LIU K, NIU Y C, LI H L. Postulation of resistance genes to stripe rust in local wheat varieties from China. Journal of Henan Agricultural Sciences, 2010(12):83-87. (in Chinese)
[50]	王吐虹, 郭青云, 蔺瑞明, 姚强, 冯晶, 王凤涛, 陈万权, 徐世昌. 中国40个小麦农家品种和甘肃南部40个生产品种抗条锈病基因推导. 中国农业科学, 2015,48(19):3834-3847.
	WANG T H, GUO Q Y, LIN R M, YAO Q, FENG J, WANG F T, CHEN W Q, XU S C. Postulation of stripe rust resistance genes in Chinese 40 wheat landraces and 40 commercial cultivars in the southern region of Gansu province. Scientia Agricultura Sinica, 2015,48(19):3834-3847. (in Chinese)
[51]	尉法刚, 王光浩, 王长有, 张宏, 刘新伦, 田增荣, 朱建峰, 陈春环, 吉万全, 王亚娟. 400份小麦品种(系)条锈病成株期抗性鉴定与评价. 植物遗传资源学报, 2020,21(4):846-854.
	WEI F G, WANG G H, WANG C Y, ZHANG H, LIU X L, TIAN Z R, ZHU J F, CHEN C H, JI W Q, WANG Y J. Evaluation and identification of adult resistance to stripe rust from 400 wheat varieties (lines). Journal of Plant Genetic Resources, 2020,21(4):846-854. (in Chinese)
[52]	李敏州, 李强, 巢凯翔, 申雪雪, 樊玉, 王阳, 王保通. 陕西省115个小麦品种(系)抗条锈病基因的分子检测. 植物病理学报, 2015,45(6):632-640.
	LI M Z, LI Q, CHAO K X, SHEN X X, FAN Y, WANG Y, WANG B T. Molecular detection of stripe rust resistance genes in 115 wheat varieties (lines) from Shaanxi province. Acta Phytopathologica Sinica, 2015,45(6):632-640. (in Chinese)
[53]	KRATTINGER S G, LAGUDAH E S, SPIELMEYER W, SINGH R P, HUERTAESPINO J, MCFADDEN H G, BOSSOLINI E, SELTER L L, KELLER B. A putative ABC transporter confers durable resistance to multiple fungal pathogens in wheat. Science, 2009,323(5919):1360-1363.
[54]	杨文雄, 杨芳萍, 梁丹, 何中虎, 尚勋武, 夏先春. 中国小麦育成品种和农家种中慢锈基因Lr34/Yr18的分子检测. 作物学报, 2008,34(7):1109-1113.
	YANG W X, YANG F P, LIANG D, HE Z H, SHANG X W, XIA X C. Molecular characterization of slow-rusting genes Lr34/Yr18 in Chinese wheat cultivars. Acta Agronomica Sinica, 2008,34(7):1109-1113. (in Chinese)
[55]	ZENG Q D, HAN D J, WANG Q L, YUAN F P, WU J H, ZHANG L, WANG X J, HUANG L L, CHEN X M, KANG Z S. Stripe rust resistance and genes in Chinese wheat cultivars and breeding lines. Euphytica, 2014,196(2):271-284.
[56]	张玉薇, 刘博, 刘太国, 高利, 陈万权. 小麦品种抗条锈病基因Yr10, Yr18及1BL/1RS易位的分子检测. 植物保护, 2014,40(1):54-59.
	ZHANG Y W, LIU B, LIU T G, GAO L, CHEN W Q. Molecular detection of Yr10 and Yr18 genes and 1BL/1RS translocation in wheat cultivars. Plant Protection, 2014,40(1):54-59. (in Chinese)
[57]	李北, 徐琪, 杨宇衡, 王琪琳, 曾庆东, 吴建辉, 穆京妹, 黄丽丽, 康振生, 韩德俊. 重庆麦区小麦品种(系)抗条锈性评价与基因分析. 中国农业科学, 2017,50(3):413-425.
	LI B, XU Q, YANG Y H, WANG Q L, ZENG Q D, WU J H, MU J M, HUANG L L, KANG Z S, HAN D J. Stripe rust resistance and genes in Chongqing wheat cultivars and lines. Scientia Agricultura Sinica, 2017,50(3):413-425. (in Chinese)
[58]	何中虎, 夏先春, 陈新民, 庄巧生. 中国小麦育种进展与展望. 作物学报, 2011,37(2):202-215.
	HE Z H, XIA X C, CHEN X M, ZHUANG Q S. Progress of wheat breeding in China and the future perspective. Acta Agronomica Sinica, 2011,37(2):202-215. (in Chinese)
[59]	ZHANG H, ZHANG L, WANG C Y, WANG Y J, ZHOU X L, LV S, LIU X L, KANG Z S, JI W Q. Molecular mapping and marker development for the Triticum dicoccoides-derived stripe rust resistance gene YrSM139-1B in bread wheat cv. Shaanmai 139. Theoretical and Applied Genetics, 2016,129(2):369-376.
[60]	穆京妹. 基于连锁分析和关联分析的小麦抗条锈病基因挖掘及Yr64与Yr65聚合选育[D]. 杨凌: 西北农林科技大学, 2019.
	MU J M. Gene mining of wheat resistance to stripe rust based on linkage analysis and association analysis and aggregation breeding of Yr64 and Yr65[D]. Yangling: Northwest Agriculture and Forestry University, 2019. (in Chinese)
[61]	韩德俊, 康振生. 中国小麦品种抗条锈病现状及存在问题与对策. 植物保护, 2018,44(5):1-12.
	HAN D J, KANG Z S. Current status and future strategy in breeding wheat for resistance to stripe rust in China. Plant Protection, 2018,44(5):1-12. (in Chinese)
[62]	LIU R, LU J, ZHOU M, ZHENG S G, LIU Z H, ZHANG C H, DU M, WANG M X, LI Y F, WU Y, ZHANG L. Developing stripe rust resistant wheat (Triticum aestivum L. lines with gene pyramiding strategy and marker-assisted selection. Genetic Resources and Crop Evolution, 2020,67(2):381-391.