40份CIMMYT小麦品系苗期及成株抗叶锈病基因鉴定

doi:10.7668/hbnxb.20192637

摘要/Abstract

摘要：

为了对40份来自国际玉米小麦改良中心(CIMMYT) 的小麦材料进行抗叶锈病基因鉴定,试验结合系谱分析、基因推导和分子标记检测等方法在苗期对36个已知抗病基因载体品种和供试的40份小麦材料接种17个具有毒性差异的叶锈菌生理小种,通过对比供试小麦材料与已知单基因载体品种侵染型,推导出供试小麦材料中可能携带的已知抗叶锈病基因,同时利用12个与已知抗病基因紧密连锁的标记对供试材料进行标记检测,检测结果与基因推导相互验证。进一步将40份供试品系分别种植于河北保定和河南周口试验田,接种叶锈菌混合小种进行田间成株期抗叶锈性鉴定。结果表明,7个供试小麦材料含有Lr1,携带Lr10的有9个品系,携带Lr11和Lr34的分别有10个品系,含有Lr14a、Lr15和Lr26的分别有2,4,3个小麦品系;另外,经标记检测成株抗叶锈病基因Lr37和Lr46分别存在于22,39个小麦品系中,经田间鉴定有22个小麦品系表现成株抗性。

关键词: 小麦叶锈病, 抗叶锈病基因, 基因推导, 成株抗叶锈性, 分子标记检测

Abstract:

To identify leaf rust resistance genes in 40 wheat materials from the International Maize and Wheat Improvement Center (CIMMYT), combined with pedigree analysis, gene derivation and molecular marker detection, 36 known disease resistance gene vector varieties and 40 wheat materials were vaccinated with 17 physiological species of leaf rust bacteria with different toxicity.By comparing the infected types of the tested wheat lines and the known rust resistance genes,the possible known resistance genes in the tested wheat materials could be postulated.At the same time,12 markers closely linked with known resistance genes were used to detect the 40 CIMMYT lines,and the results could be verified with the gene postulation.Furthermore,40 tested lines were planted in Baoding,Hebei Province and Zhoukou,Henan Province,respectively,and inoculated with mixed Pt races for leaf rust evaluation at adult plant stage in the field.The results showed that 7 wheat lines contained Lr1,Lr10 was found in 9 lines,and each of Lr11 and Lr34 was present in 10 lines.Each of Lr14a,Lr15 and Lr26 was postulated to present in 2,4 and 3 lines,respectively.Each of Lr37 and Lr46 was identified in 22,39 wheat lines,respectively,by using molecular marker detection.In the field test 22 wheat lines showed adult plant resistance to leaf rust.

Key words: Wheat leaf rust, Leaf rust resistance gene, Gene postulation, Adult plant leaf rust resistance, Molecular marker detection

王佳荣, 董瑞, 张梦宇, 高璞, 张培培, 李在峰, 刘大群. 40份CIMMYT小麦品系苗期及成株抗叶锈病基因鉴定[J]. 华北农学报, 2022, 37(2): 201-210. doi: 10.7668/hbnxb.20192637.

WANG Jiarong, DONG Rui, ZHANG Mengyu, GAO Pu, ZHANG Peipei, LI Zaifeng, LIU Daqun. Identification of Leaf Rust Resistance Genes in 40 CIMMYT Wheat Lines at Seedling Stage and Adult Plant[J]. Acta Agriculturae Boreali-Sinica, 2022, 37(2): 201-210. doi: 10.7668/hbnxb.20192637.

http://www.hbnxb.net/CN/Y2022/V37/I2/201

图/表 7

表1 40个小麦品系的系谱及来源

Tab.1 The pedigrees and origins of the 40 wheat lines

序号 No.	品系 Lines	系谱 Pedigree	来源 Origin
1	XYZ 461	WAXWING/7/TNMU/6/CEP80111/CEP81165/5/IAC5/4/YKT406/3/AG/ASN//ATR/8/ATTILA/3*BCN//BAV92/3/TILHI/4/SHA7/VEE#5//ARIV92	CIMMYT
2	XYZ 462	ATTILA2/HUITES//FINSI/3/ATTILA 2/PBW65/4/TRCH/SRTU//KACHU	CIMMYT
3	XYZ 463	WBLL1* 2/4/YACO/PBW65/3/KAUZ* 2/TRAP//KAUZ/5/KACHU #1/6/TRCH/SRTU//KACHU	CIMMYT
4	XYZ 464	TIMBA/KACHU/4/MARCHOUCH* 4/SAADA/3/2* FRET2/KUKUNA//FRET2	CIMMYT
5	XYZ 465	SAUAL/3/KAUZ/PASTOR//PBW343/4/TRCH/SRTU//KACHU	CIMMYT
6	XYZ 466	TRCH/SRTU//KACHU/3/TRCH/SRTU//KACHU	CIMMYT
7	XYZ 467	TILHI/SOKOLL* 2//KINGBIRD #1	CIMMYT
8	XYZ 468	TRCH/SRTU//KACHU* 2/3/PVN	CIMMYT
9	XYZ 469	TRCH/SRTU//KACHU* 2/3/PVN	CIMMYT
10	XYZ 470	TRCH/SRTU//KACHU* 2/3/PVN	CIMMYT
11	XYZ 471	TRCH/SRTU//KACHU* 2/3/KINGBIRD #1	CIMMYT
12	XYZ 472	TRCH/SRTU//KACHU* 2/3/KINGBIRD #1	CIMMYT
13	XYZ 473	WBLL1/KUKUNA//TACUPETO F2001/3/BAJ #1* 2/4/KINGBIRD #1	CIMMYT
14	XYZ 474	KIRITATI/2* WBLL1/5/FRET2/KUKUNA//FRET2/3/YANAC/4/FRET2/KIRITATI/6/BECARD #1	CIMMYT
15	XYZ 475	ROLF07/YANAC//TACUPETO F2001/BRAMBLING/4/WBLL1/KUKUNA//TACUPETO F2001/3/BAJ #1/5/WBLL1/KUKUNA//TACUPETO F2001/3/BAJ #1	CIMMYT
16	XYZ 476	FRET2/KUKUNA//FRET2/3/YANAC/4/FRET2/KIRITATI* 2/5/WHEAR//2* PRL/2* PASTOR	CIMMYT
17	XYZ 477	TRCH/5/BAV92//IRENA/KAUZ/3/HUITES/4/DOLL/6/TRCH/SRTU//KACHU	CIMMYT
18	XYZ 478	WHEAR/KUKUNA/3/C80.1/3* BATAVIA//2* WBLL1* 2/6/WBLL1* 2/4/YACO/PBW65/3/KAUZ* 2/TRAP//KAUZ/5/KACHU #1	CIMMYT
19	XYZ 479	PBW343* 2/KUKUNA//PBW343* 2/KUKUNA* 2/5/PFAU/SERI.1B//AMAD/3/INQALAB 91* 2/KURUKU/4/INQALAB 91* 2/KUKUNA	CIMMYT
20	XYZ 480	PBW343* 2/KUKUNA//PBW343* 2/KUKUNA* 2/6/WBLL1* 2/4/SNI/TRAP#1/3/KAUZ* 2/TRAP//KAUZ/5/KACHU	CIMMYT
21	XYZ 481	PBW343* 2/KUKUNA//PBW343* 2/KUKUNA* 2/6/C80.1/3* BATAVIA//2* WBLL1/5/REH/HARE//2* BCN/3/CROC_1/AE.SQUARROSA (213)//PGO/4/HUITES	CIMMYT
22	XYZ 482	MELON//FILIN/MILAN/3/FILIN/4/TRCH/SRTU//KACHU/5/TRCH/SRTU//KACHU	CIMMYT
23	XYZ 483	PAURAQ/2* NELOKI	CIMMYT
24	XYZ 484	BECARD/KINGBIRD #1//BECARD	CIMMYT
25	XYZ 485	SOKOLL/3/PASTOR//HXL7573/2* BAU/4/2* PASTOR//HXL7573/2* BAU/3/SOKOLL/WBLL1	CIMMYT
26	XYZ 486	SOKOLL/3/PASTOR//HXL7573/2* BAU* 2/4/PAURAQ	CIMMYT
27	XYZ 487	SOKOLL/3/PASTOR//HXL7573/2* BAU* 2/4/EGA BONNIE ROCK	CIMMYT
28	XYZ 488	SOKOLL/3/PASTOR//HXL7573/2* BAU* 2/4/EGA BONNIE ROCK	CIMMYT
29	XYZ 489	SOKOLL/3/PASTOR//HXL7573/2* BAU* 2/4/EGA BONNIE ROCK	CIMMYT
30	XYZ 490	SOKOLL/3/PASTOR//HXL7573/2* BAU* 2/4/GLADIUS	CIMMYT
31	XYZ 491	SOKOLL/3/PASTOR//HXL7573/2* BAU* 2/4/GLADIUS	CIMMYT
32	XYZ 492	WBLL1* 2/BRAMBLING//ND643/2* WBLL1	CIMMYT
33	XYZ 493	ND643/2* WBLL1/4/HUW234+LR34/PRINIA//PBW343* 2/KUKUNA/3/ROLF07	CIMMYT
34	XYZ 494	MERCATO/4/FRAME//MILAN/KAUZ/3/PASTOR/5/WHEAR/SOKOLL	CIMMYT
35	XYZ 495	BABAX/LR42//BABAX/3/ER2000/11/CROC_1/AE.SQUARROSA(213)//PGO/10/ATTILA* 2/9/KT/BAGE//FN/U/3/BZA/4/TRM/5/ALDAN/6/SERI/7/VEE#10/8/OPATA/12/BAVIS	CIMMYT
36	XYZ 496	ATTILA* 2/PBW65//BERKUT/3/WHEAR/SOKOLL	CIMMYT
37	XYZ 497	ND643/2* WBLL1* 2//KINGBIRD #1	CIMMYT
38	XYZ 498	PFAU/MILAN/3/BABAX/LR42//BABAX* 2/4/NIINI #1	CIMMYT
39	XYZ 499	PFAU/MILAN/3/BABAX/LR42//BABAX* 2/4/NIINI #1	CIMMYT
40	XYZ 500	PBW343* 2/KUKUNA//JUCHI/3/ATTILA* 2/PBW65//TNMU	CIMMYT

表1 40个小麦品系的系谱及来源

Tab.1 The pedigrees and origins of the 40 wheat lines

序号 No.	品系 Lines	系谱 Pedigree	来源 Origin
1	XYZ 461	WAXWING/7/TNMU/6/CEP80111/CEP81165/5/IAC5/4/YKT406/3/AG/ASN//ATR/8/ATTILA/3*BCN//BAV92/3/TILHI/4/SHA7/VEE#5//ARIV92	CIMMYT
2	XYZ 462	ATTILA2/HUITES//FINSI/3/ATTILA 2/PBW65/4/TRCH/SRTU//KACHU	CIMMYT
3	XYZ 463	WBLL1* 2/4/YACO/PBW65/3/KAUZ* 2/TRAP//KAUZ/5/KACHU #1/6/TRCH/SRTU//KACHU	CIMMYT
4	XYZ 464	TIMBA/KACHU/4/MARCHOUCH* 4/SAADA/3/2* FRET2/KUKUNA//FRET2	CIMMYT
5	XYZ 465	SAUAL/3/KAUZ/PASTOR//PBW343/4/TRCH/SRTU//KACHU	CIMMYT
6	XYZ 466	TRCH/SRTU//KACHU/3/TRCH/SRTU//KACHU	CIMMYT
7	XYZ 467	TILHI/SOKOLL* 2//KINGBIRD #1	CIMMYT
8	XYZ 468	TRCH/SRTU//KACHU* 2/3/PVN	CIMMYT
9	XYZ 469	TRCH/SRTU//KACHU* 2/3/PVN	CIMMYT
10	XYZ 470	TRCH/SRTU//KACHU* 2/3/PVN	CIMMYT
11	XYZ 471	TRCH/SRTU//KACHU* 2/3/KINGBIRD #1	CIMMYT
12	XYZ 472	TRCH/SRTU//KACHU* 2/3/KINGBIRD #1	CIMMYT
13	XYZ 473	WBLL1/KUKUNA//TACUPETO F2001/3/BAJ #1* 2/4/KINGBIRD #1	CIMMYT
14	XYZ 474	KIRITATI/2* WBLL1/5/FRET2/KUKUNA//FRET2/3/YANAC/4/FRET2/KIRITATI/6/BECARD #1	CIMMYT
15	XYZ 475	ROLF07/YANAC//TACUPETO F2001/BRAMBLING/4/WBLL1/KUKUNA//TACUPETO F2001/3/BAJ #1/5/WBLL1/KUKUNA//TACUPETO F2001/3/BAJ #1	CIMMYT
16	XYZ 476	FRET2/KUKUNA//FRET2/3/YANAC/4/FRET2/KIRITATI* 2/5/WHEAR//2* PRL/2* PASTOR	CIMMYT
17	XYZ 477	TRCH/5/BAV92//IRENA/KAUZ/3/HUITES/4/DOLL/6/TRCH/SRTU//KACHU	CIMMYT
18	XYZ 478	WHEAR/KUKUNA/3/C80.1/3* BATAVIA//2* WBLL1* 2/6/WBLL1* 2/4/YACO/PBW65/3/KAUZ* 2/TRAP//KAUZ/5/KACHU #1	CIMMYT
19	XYZ 479	PBW343* 2/KUKUNA//PBW343* 2/KUKUNA* 2/5/PFAU/SERI.1B//AMAD/3/INQALAB 91* 2/KURUKU/4/INQALAB 91* 2/KUKUNA	CIMMYT
20	XYZ 480	PBW343* 2/KUKUNA//PBW343* 2/KUKUNA* 2/6/WBLL1* 2/4/SNI/TRAP#1/3/KAUZ* 2/TRAP//KAUZ/5/KACHU	CIMMYT
21	XYZ 481	PBW343* 2/KUKUNA//PBW343* 2/KUKUNA* 2/6/C80.1/3* BATAVIA//2* WBLL1/5/REH/HARE//2* BCN/3/CROC_1/AE.SQUARROSA (213)//PGO/4/HUITES	CIMMYT
22	XYZ 482	MELON//FILIN/MILAN/3/FILIN/4/TRCH/SRTU//KACHU/5/TRCH/SRTU//KACHU	CIMMYT
23	XYZ 483	PAURAQ/2* NELOKI	CIMMYT
24	XYZ 484	BECARD/KINGBIRD #1//BECARD	CIMMYT
25	XYZ 485	SOKOLL/3/PASTOR//HXL7573/2* BAU/4/2* PASTOR//HXL7573/2* BAU/3/SOKOLL/WBLL1	CIMMYT
26	XYZ 486	SOKOLL/3/PASTOR//HXL7573/2* BAU* 2/4/PAURAQ	CIMMYT
27	XYZ 487	SOKOLL/3/PASTOR//HXL7573/2* BAU* 2/4/EGA BONNIE ROCK	CIMMYT
28	XYZ 488	SOKOLL/3/PASTOR//HXL7573/2* BAU* 2/4/EGA BONNIE ROCK	CIMMYT
29	XYZ 489	SOKOLL/3/PASTOR//HXL7573/2* BAU* 2/4/EGA BONNIE ROCK	CIMMYT
30	XYZ 490	SOKOLL/3/PASTOR//HXL7573/2* BAU* 2/4/GLADIUS	CIMMYT
31	XYZ 491	SOKOLL/3/PASTOR//HXL7573/2* BAU* 2/4/GLADIUS	CIMMYT
32	XYZ 492	WBLL1* 2/BRAMBLING//ND643/2* WBLL1	CIMMYT
33	XYZ 493	ND643/2* WBLL1/4/HUW234+LR34/PRINIA//PBW343* 2/KUKUNA/3/ROLF07	CIMMYT
34	XYZ 494	MERCATO/4/FRAME//MILAN/KAUZ/3/PASTOR/5/WHEAR/SOKOLL	CIMMYT
35	XYZ 495	BABAX/LR42//BABAX/3/ER2000/11/CROC_1/AE.SQUARROSA(213)//PGO/10/ATTILA* 2/9/KT/BAGE//FN/U/3/BZA/4/TRM/5/ALDAN/6/SERI/7/VEE#10/8/OPATA/12/BAVIS	CIMMYT
36	XYZ 496	ATTILA* 2/PBW65//BERKUT/3/WHEAR/SOKOLL	CIMMYT
37	XYZ 497	ND643/2* WBLL1* 2//KINGBIRD #1	CIMMYT
38	XYZ 498	PFAU/MILAN/3/BABAX/LR42//BABAX* 2/4/NIINI #1	CIMMYT
39	XYZ 499	PFAU/MILAN/3/BABAX/LR42//BABAX* 2/4/NIINI #1	CIMMYT
40	XYZ 500	PBW343* 2/KUKUNA//JUCHI/3/ATTILA* 2/PBW65//TNMU	CIMMYT

表2 17个小麦叶锈菌生理小种与36个抗叶锈病基因载体品种苗期互作侵染型

Tab.2 Seedling infection types of 36 lines with known leaf rust resistance genes inoculated with 17 Puccinia triticina (Pt)races

序号 No.	Lr 基因 Lr genes	叶锈菌生理小种侵染型Infection types to Pt pathotypes
序号 No.	Lr 基因 Lr genes	FGJQ	FHJQ①	FHJT	FHJR	TGPS	FHJQ②	THDB	PHTS	FHJQ③	FHSQ	FHGS①	FHDS	FHGS②	FHDQ	FHJS	FGJQ	FGDQ
1	RL6003(Lr1)	;1	;1	;1	2	3+	;	3+	4	;1	;1	0	0	;	;1	1	1	;1
2	RL6016(Lr2a)	;1	;	2	1	3	1	3+	;1	;1	1	;	0	;	;1	2	1	;1
3	RL6047(Lr2C)	3+	3+	3+	3+	3	3	3	3+	3	3+	3	3	3	3	3	4	3
4	RL6002(Lr3)	3+	3+	3+	3+	3	3+	3	3+	3	3	3	3	3	3	3	3	3
5	RL6010(Lr9)	0	0	0	0	0	0	0	0	0	0	0	0	0	0	0	0	0
6	RL6005(Lr16)	3+	3+	3+	3+	3+	3+	3+	3	3+	3	3	4	3+	4	4	3+	4
7	RL6064(Lr24)	;1	;	;1	;	;	;	;1	;1	;1	;	;	;1	;	0	1	;	;1
8	RL6078(Lr26)	1	3+	3	3	2+	3+	3	3	3	3	3	3	3	3	3	1	1
9	RL6007(Lr3ka)	2	2	2	2	3+	1	2	3	1	3	;	1	2	1	1	2	1
10	RL6053(Lr11)	3+	3	3+	3+	2+	3	2	3+	3+	3+	3	2	3	2	3	3	2
11	RL6008(Lr17)	3	3	3+	3+	3+	3+	3+	3	3	3+	2	3	2+	3	3	3	3
12	RL6049(Lr30)	1+	1+	1	2	3	1	2	3+	1	2	1	2	2	2	2	1	2
13	RL6051(LrB)	3C	3+	3+	3+	3C	3	3	3+	3+	3	3	3	3+	4	4	4	3+
14	RL6004(Lr10)	3C	3	3+	4	3+	3+	1	3	3+	3	3	3	3+	4	3	3+	4
15	RL6013(Lr14a)	1	2	3	X	3+	2	1+	3	2	X	3	3+	3	2+	3	2	1+
16	RL6009(Lr18)	1	2	3	3	2	2	2	2	2	1	0	0	0	0	2	2	2+
17	RL6019(Lr2b)	2	2+	2	2	3+	2	3+	2	3	3	;1	3	3	2	2	2	3
18	RL6042(Lr3bg)	3	3	3+	3+	4	3+	3+	3+	3+	3	3	3	3+	3+	3	4	4
19	RL4031(Lr13)	3	3	3+	3	3+	3	3	3	3	3	3C	3	3+	4	4	4	3+
20	RL6006(Lr14b)	3+	3+	3+	4	3	3+	3+	4	4	3+	4	3+	3+	4	3	4	4
21	RL605(Lr15)	;	0	0	3+	3+	3	3+	3	0	0	0	1	1	0	0	0;	0
22	RL604(Lr19)	;	;	0	0	;	0	0	0	0	0	0	;1	0	;	0	0;	0
23	RL609(Lr20)	;	;	3+	3	3	1	3+	3+	;	3	;	1	3+	4	4	;	2
24	RL604(Lr21)	1+	2	2	3	2	2	1+	1	1	2	2	3	2	2	2	2+	2
25	RL601(Lr23)	3	3+	3+	3+	3+	3+	3+	3	3+	3	3	3+	3+	4	3	3+	4
26	RL607(Lr28)	0	;	0	0	0	0	0	0	0	0	;	;1	0	;	1	;	0
27	RL608(Lr29)	;	2	2	1	2	1	1+	1	1	1	2	;	1	0	2	1	2
28	RL605(Lr33)	3C	3	3+	3	3	3+	3+	3+	3+	3	3	3+	3+	3	3	3	3
29	E8401(Lr36)	1+	1+	2	1	1	1	1	3	2	1	1	1	2	2	2	2	2
30	KS86NGRC02(Lr39)	3	3+	3	3	3	3+	3+	3+	3+	3	3	3	3+	4	3	3+	3
31	KS91WGRC11(Lr42)	;	1+	1	2	1	1	1	12	1	2	1	1	2	;	1	;	1
32	RL614(Lr44)	3	3	3+	2	3	2	3	4	3+	3C	3	2	3	3	3	3	3
33	RL614(Lr45)	3	3+	3	3+	4	4	3C	2	3+	3	3+	3	3+	2	4	4	3+
34	C98.00(Lr47)	0	0	0	0	0	0	;	0	0	;	;	;1	0	0	0	0	;
35	C78.5(Lr51)	;	;	;1	;	;	;	;	;	;	;	;	;1	;	1	;1	0;	;
36	98M71(Lr53)	1	;	;	;1	1	1	;	;	;	;	;	;1	;	;	;	;	;

表2 17个小麦叶锈菌生理小种与36个抗叶锈病基因载体品种苗期互作侵染型

Tab.2 Seedling infection types of 36 lines with known leaf rust resistance genes inoculated with 17 Puccinia triticina (Pt)races

序号 No.	Lr 基因 Lr genes	叶锈菌生理小种侵染型Infection types to Pt pathotypes
序号 No.	Lr 基因 Lr genes	FGJQ	FHJQ①	FHJT	FHJR	TGPS	FHJQ②	THDB	PHTS	FHJQ③	FHSQ	FHGS①	FHDS	FHGS②	FHDQ	FHJS	FGJQ	FGDQ
1	RL6003(Lr1)	;1	;1	;1	2	3+	;	3+	4	;1	;1	0	0	;	;1	1	1	;1
2	RL6016(Lr2a)	;1	;	2	1	3	1	3+	;1	;1	1	;	0	;	;1	2	1	;1
3	RL6047(Lr2C)	3+	3+	3+	3+	3	3	3	3+	3	3+	3	3	3	3	3	4	3
4	RL6002(Lr3)	3+	3+	3+	3+	3	3+	3	3+	3	3	3	3	3	3	3	3	3
5	RL6010(Lr9)	0	0	0	0	0	0	0	0	0	0	0	0	0	0	0	0	0
6	RL6005(Lr16)	3+	3+	3+	3+	3+	3+	3+	3	3+	3	3	4	3+	4	4	3+	4
7	RL6064(Lr24)	;1	;	;1	;	;	;	;1	;1	;1	;	;	;1	;	0	1	;	;1
8	RL6078(Lr26)	1	3+	3	3	2+	3+	3	3	3	3	3	3	3	3	3	1	1
9	RL6007(Lr3ka)	2	2	2	2	3+	1	2	3	1	3	;	1	2	1	1	2	1
10	RL6053(Lr11)	3+	3	3+	3+	2+	3	2	3+	3+	3+	3	2	3	2	3	3	2
11	RL6008(Lr17)	3	3	3+	3+	3+	3+	3+	3	3	3+	2	3	2+	3	3	3	3
12	RL6049(Lr30)	1+	1+	1	2	3	1	2	3+	1	2	1	2	2	2	2	1	2
13	RL6051(LrB)	3C	3+	3+	3+	3C	3	3	3+	3+	3	3	3	3+	4	4	4	3+
14	RL6004(Lr10)	3C	3	3+	4	3+	3+	1	3	3+	3	3	3	3+	4	3	3+	4
15	RL6013(Lr14a)	1	2	3	X	3+	2	1+	3	2	X	3	3+	3	2+	3	2	1+
16	RL6009(Lr18)	1	2	3	3	2	2	2	2	2	1	0	0	0	0	2	2	2+
17	RL6019(Lr2b)	2	2+	2	2	3+	2	3+	2	3	3	;1	3	3	2	2	2	3
18	RL6042(Lr3bg)	3	3	3+	3+	4	3+	3+	3+	3+	3	3	3	3+	3+	3	4	4
19	RL4031(Lr13)	3	3	3+	3	3+	3	3	3	3	3	3C	3	3+	4	4	4	3+
20	RL6006(Lr14b)	3+	3+	3+	4	3	3+	3+	4	4	3+	4	3+	3+	4	3	4	4
21	RL605(Lr15)	;	0	0	3+	3+	3	3+	3	0	0	0	1	1	0	0	0;	0
22	RL604(Lr19)	;	;	0	0	;	0	0	0	0	0	0	;1	0	;	0	0;	0
23	RL609(Lr20)	;	;	3+	3	3	1	3+	3+	;	3	;	1	3+	4	4	;	2
24	RL604(Lr21)	1+	2	2	3	2	2	1+	1	1	2	2	3	2	2	2	2+	2
25	RL601(Lr23)	3	3+	3+	3+	3+	3+	3+	3	3+	3	3	3+	3+	4	3	3+	4
26	RL607(Lr28)	0	;	0	0	0	0	0	0	0	0	;	;1	0	;	1	;	0
27	RL608(Lr29)	;	2	2	1	2	1	1+	1	1	1	2	;	1	0	2	1	2
28	RL605(Lr33)	3C	3	3+	3	3	3+	3+	3+	3+	3	3	3+	3+	3	3	3	3
29	E8401(Lr36)	1+	1+	2	1	1	1	1	3	2	1	1	1	2	2	2	2	2
30	KS86NGRC02(Lr39)	3	3+	3	3	3	3+	3+	3+	3+	3	3	3	3+	4	3	3+	3
31	KS91WGRC11(Lr42)	;	1+	1	2	1	1	1	12	1	2	1	1	2	;	1	;	1
32	RL614(Lr44)	3	3	3+	2	3	2	3	4	3+	3C	3	2	3	3	3	3	3
33	RL614(Lr45)	3	3+	3	3+	4	4	3C	2	3+	3	3+	3	3+	2	4	4	3+
34	C98.00(Lr47)	0	0	0	0	0	0	;	0	0	;	;	;1	0	0	0	0	;
35	C78.5(Lr51)	;	;	;1	;	;	;	;	;	;	;	;	;1	;	1	;1	0;	;
36	98M71(Lr53)	1	;	;	;1	1	1	;	;	;	;	;	;1	;	;	;	;	;

表3 17个小麦叶锈菌生理小种与40个小麦品系的苗期互作侵染型

Tab.3 Seedling infection types of the 40 wheat lines with 17 Puccinia triticina (Pt) races

序号 No.	品系 Lines	品种所含基因 Genes	叶锈菌生理小种侵染型Infection types to Pt pathotypes
序号 No.	品系 Lines	品种所含基因 Genes	FGJQ	FHJQ①	FHJT	FHJR	TGPS	FHJQ②	THDB	PHTS	FHJQ③	FHSQ	FHGS①	FHDS	FHGS②	FHDQ	FHJS	FGJQ	FGDQ
1	XYZ 461	$Lr 11 d 、 Lr 26 c 、 Lr 46 e$	;	1	;1	2	1	3	12	3	;1	;1	3	1	1	1	1	1	1
2	XYZ 462	$Lr 11 d 、 Lr 37 e 、 Lr 46 e$	2+	12	3	12	1	3	12	2	12	2+	2	1	1	1	1	1	1
3	XYZ 463	$Lr 37 e 、 Lr 46 e$	3	12	3	2+	3	3	12	3	2	2+	2	1	1	1	2	1	1
4	XYZ 464	$Lr 10 e 、 Lr 14 ac 、 Lr 34 e 、 Lr 37 e 、 Lr 46 e$	1	12	3	2	3	;1	1	3	1	2	1	1	1	1	1	1	1
5	XYZ 465	$Lr 26 c 、 Lr 37 e 、 Lr 46 e$	;	12	3	2+	2	3	1	3C	2	2+	2	1	1	1	1	;1	1
6	XYZ 466	$Lr 37 e 、 Lr 46 e$	3	2	3C	2+	3	2	1	3C	2	3C	2+	1	12	1	2	1	2
7	XYZ 467	$Lr 10 e 、 Lr 34 e 、 Lr 46 e$	3	;1	;1	;	1	;1	;	3	1	1	12	1	;	;	1	;	1
8	XYZ 468	$Lr 37 e 、 Lr 46 e$	3	1	3	3	3	3	3	3	3C	3	3	3	1	2	3	1	3
9	XYZ 469	$Lr 11 d 、 Lr 37 e 、 Lr 46 e$	;	0	;1	3	2	1	1	2+	;1	;1	3+	1	;	1	1	1	1
10	XYZ 470	$Lr 37 e 、 Lr 46 e$	;	3	3	3	3	2	1+	3C	;	1	3	1	;	1	1	;	1
11	XYZ 471	$Lr 10 e 、 Lr 37 e 、 Lr 46 e$	3	3C	3	2	3	3	1	3C	2+	3	2	2	1	1	1	2	1
12	XYZ 472	$Lr 10 e 、 Lr 37 e 、 Lr 46 e$	3C	2	3	1	3	2	;1	2+	2+	2	1	1	1	1	1	1	1
13	XYZ 473	$Lr 1 e 、 Lr 10 e 、 Lr 34 e 、 Lr 46 e$	;	;	;1	1	12	;1	1	2	;1	;1	1	1	;1	1	1	1	1
14	XYZ 474	$Lr 1 d 、 Lr 10 e 、 Lr 11 d 、 Lr 46 e$	1	2	1	2	1	2	12	3	2	2+	2	1	1	1	2	1	1
15	XYZ 475	$Lr 11 d 、 Lr 34 e 、 Lr 46 e$	3	1	1	1	;1	1	12	3-	;1	2	2+	2	1	2	1	1	1
16	XYZ 476	$Lr 34 e$	;	3	3+	2	;1	3	12	3C	3	;1	3	3	1	2	3	1	3
17	XYZ 477	$Lr 10 e 、 Lr 37 e 、 Lr 46 e$	2	1	2	2	;1	2	1+	2	;1	12	3	1	1	2	3	1	3
18	XYZ 478	$Lr 1 e 、 Lr 10 e 、 Lr 34 e 、 Lr 37 e 、 Lr 46 e$	;	;	;	1	;1	;1	1	;1	;1	;1	;	1	;1	1	1	1	1
19	XYZ 479	$Lr 1 e 、 Lr 34 e 、 Lr 46 e$	;1	;1	1	1	;	;1	;1	12	;1	;1	12	1	;1	1	1	1	1
20	XYZ 480	$Lr 1 e 、 Lr 34 e 、 Lr 37 e 、 Lr 46 e$	1	1	1	1	;	;1	12	2	12	;1	0	2	;	1	1	0	1
21	XYZ 481	$Lr 1 e 、 Lr 34 e 、 Lr 46 e$	1	1+	1	12	;	;1	12	;1	;1	;1	0	1	0	1	1	;	1
22	XYZ 482	$Lr 1 e 、 Lr 11 d 、 Lr 37 e 、 Lr 46 e$	1	1	2	2	;	1	1	2	1	1	1	2	1 2	1	1	;	1
23	XYZ 483	$Lr 11 d 、 Lr 46 e$	;	;1	1	1	;1	;1	1+	3	;1	1	1	1 2	1	1	1	1	1
24	XYZ 484	$Lr 11 d 、 Lr 37 e 、 Lr 46 e$	3	3	3	1	;1	;1	;	2	1	12	12	1	1	1	1	;	1
25	XYZ 485	$Lr 46 e$	3	3+	3+	2+	3+	3+	3	2+	3	3+	1	1	1	1	1	;	1
26	XYZ 486	$Lr 46 e$	3	3	3+	2	3+	3	1	2+	1+	12	1	1	1	1	1	1	1
27	XYZ 487	$Lr 46 e$	3+	3+	3	3	3	3	3+	3	3	1	1	2	3	0;	4	3+	1
28	XYZ 488	$Lr 15 d 、 Lr 46 e$	;	;	;	1	3	;1	3	;1	;1	1	1	1	;	1	0	;	1
29	XYZ 489	$Lr 15 d 、 Lr 46 e$	;	;	;	;1	3	3	3	;1	;1	1	1	1	;	1	0;	;	1
30	XYZ 490	$Lr 15 d 、 Lr 46 e$	;	;	;	2	4	;1	3+	;1	;	1	1	1	0	1	0	;	1
31	XYZ 491	$Lr 37 e 、 Lr 46 e$	4	3+	3+	3+	4	3+	3+	3+	4	1	3	3	4	4	4	4	1
32	XYZ 492	$Lr 46 e$	;1	3	3	1+	;1	1	3	;1	2	1	1	1	2	1	1	1	1
33	XYZ 493	$Lr 11 d 、 Lr 46 e$	1	1+	3	;	;1	3+	1	;1	;1	1	1	;	2	1	1	;	1
34	XYZ 494	$Lr 37 e 、 Lr 46 e$	3	2	3	3C	3+	3+	12	3	2+	2+	2+	2	3C	1	2	;	1
35	XYZ 495	$Lr 11 d 、 Lr 14 ac 、 Lr 26 c 、 Lr 37 e 、 Lr 46 e$	;1	;	;	;1	;1	;1	1	;1	1	12	1	;	1	1	3	;	1
36	XYZ 496	$Lr 37 e 、 Lr 46 e$	;	;	;	;1	;1	;1	;	;1	;	;	1	;	1	1	;	1	1
37	XYZ 497	$Lr 15 d 、 Lr 34 e 、 Lr 37 e 、 Lr 46 e$	1	;	;	3	3+	;1	1	3-	;1	1	1	;	1	1	;	1	1
38	XYZ 498	$Lr 37 e 、 Lr 46 e$	;	;	;	;	;	;	;1	;	;1	;	1	;	1	1	1	;	1
39	XYZ 499	$Lr 37 e 、 Lr 46 e$	;	;	;	1	;	;	;1	;	1	;	;1	;	1	1	1	;	1
40	XYZ 500	$Lr 10 e 、 Lr 15 d 、 Lr 46 e$	;	;	;	;	;	;	;1	;1	;	;1	;1	;	1	1	1	;	1

表3 17个小麦叶锈菌生理小种与40个小麦品系的苗期互作侵染型

Tab.3 Seedling infection types of the 40 wheat lines with 17 Puccinia triticina (Pt) races

序号 No.	品系 Lines	品种所含基因 Genes	叶锈菌生理小种侵染型Infection types to Pt pathotypes
序号 No.	品系 Lines	品种所含基因 Genes	FGJQ	FHJQ①	FHJT	FHJR	TGPS	FHJQ②	THDB	PHTS	FHJQ③	FHSQ	FHGS①	FHDS	FHGS②	FHDQ	FHJS	FGJQ	FGDQ
1	XYZ 461	$Lr 11 d 、 Lr 26 c 、 Lr 46 e$	;	1	;1	2	1	3	12	3	;1	;1	3	1	1	1	1	1	1
2	XYZ 462	$Lr 11 d 、 Lr 37 e 、 Lr 46 e$	2+	12	3	12	1	3	12	2	12	2+	2	1	1	1	1	1	1
3	XYZ 463	$Lr 37 e 、 Lr 46 e$	3	12	3	2+	3	3	12	3	2	2+	2	1	1	1	2	1	1
4	XYZ 464	$Lr 10 e 、 Lr 14 ac 、 Lr 34 e 、 Lr 37 e 、 Lr 46 e$	1	12	3	2	3	;1	1	3	1	2	1	1	1	1	1	1	1
5	XYZ 465	$Lr 26 c 、 Lr 37 e 、 Lr 46 e$	;	12	3	2+	2	3	1	3C	2	2+	2	1	1	1	1	;1	1
6	XYZ 466	$Lr 37 e 、 Lr 46 e$	3	2	3C	2+	3	2	1	3C	2	3C	2+	1	12	1	2	1	2
7	XYZ 467	$Lr 10 e 、 Lr 34 e 、 Lr 46 e$	3	;1	;1	;	1	;1	;	3	1	1	12	1	;	;	1	;	1
8	XYZ 468	$Lr 37 e 、 Lr 46 e$	3	1	3	3	3	3	3	3	3C	3	3	3	1	2	3	1	3
9	XYZ 469	$Lr 11 d 、 Lr 37 e 、 Lr 46 e$	;	0	;1	3	2	1	1	2+	;1	;1	3+	1	;	1	1	1	1
10	XYZ 470	$Lr 37 e 、 Lr 46 e$	;	3	3	3	3	2	1+	3C	;	1	3	1	;	1	1	;	1
11	XYZ 471	$Lr 10 e 、 Lr 37 e 、 Lr 46 e$	3	3C	3	2	3	3	1	3C	2+	3	2	2	1	1	1	2	1
12	XYZ 472	$Lr 10 e 、 Lr 37 e 、 Lr 46 e$	3C	2	3	1	3	2	;1	2+	2+	2	1	1	1	1	1	1	1
13	XYZ 473	$Lr 1 e 、 Lr 10 e 、 Lr 34 e 、 Lr 46 e$	;	;	;1	1	12	;1	1	2	;1	;1	1	1	;1	1	1	1	1
14	XYZ 474	$Lr 1 d 、 Lr 10 e 、 Lr 11 d 、 Lr 46 e$	1	2	1	2	1	2	12	3	2	2+	2	1	1	1	2	1	1
15	XYZ 475	$Lr 11 d 、 Lr 34 e 、 Lr 46 e$	3	1	1	1	;1	1	12	3-	;1	2	2+	2	1	2	1	1	1
16	XYZ 476	$Lr 34 e$	;	3	3+	2	;1	3	12	3C	3	;1	3	3	1	2	3	1	3
17	XYZ 477	$Lr 10 e 、 Lr 37 e 、 Lr 46 e$	2	1	2	2	;1	2	1+	2	;1	12	3	1	1	2	3	1	3
18	XYZ 478	$Lr 1 e 、 Lr 10 e 、 Lr 34 e 、 Lr 37 e 、 Lr 46 e$	;	;	;	1	;1	;1	1	;1	;1	;1	;	1	;1	1	1	1	1
19	XYZ 479	$Lr 1 e 、 Lr 34 e 、 Lr 46 e$	;1	;1	1	1	;	;1	;1	12	;1	;1	12	1	;1	1	1	1	1
20	XYZ 480	$Lr 1 e 、 Lr 34 e 、 Lr 37 e 、 Lr 46 e$	1	1	1	1	;	;1	12	2	12	;1	0	2	;	1	1	0	1
21	XYZ 481	$Lr 1 e 、 Lr 34 e 、 Lr 46 e$	1	1+	1	12	;	;1	12	;1	;1	;1	0	1	0	1	1	;	1
22	XYZ 482	$Lr 1 e 、 Lr 11 d 、 Lr 37 e 、 Lr 46 e$	1	1	2	2	;	1	1	2	1	1	1	2	1 2	1	1	;	1
23	XYZ 483	$Lr 11 d 、 Lr 46 e$	;	;1	1	1	;1	;1	1+	3	;1	1	1	1 2	1	1	1	1	1
24	XYZ 484	$Lr 11 d 、 Lr 37 e 、 Lr 46 e$	3	3	3	1	;1	;1	;	2	1	12	12	1	1	1	1	;	1
25	XYZ 485	$Lr 46 e$	3	3+	3+	2+	3+	3+	3	2+	3	3+	1	1	1	1	1	;	1
26	XYZ 486	$Lr 46 e$	3	3	3+	2	3+	3	1	2+	1+	12	1	1	1	1	1	1	1
27	XYZ 487	$Lr 46 e$	3+	3+	3	3	3	3	3+	3	3	1	1	2	3	0;	4	3+	1
28	XYZ 488	$Lr 15 d 、 Lr 46 e$	;	;	;	1	3	;1	3	;1	;1	1	1	1	;	1	0	;	1
29	XYZ 489	$Lr 15 d 、 Lr 46 e$	;	;	;	;1	3	3	3	;1	;1	1	1	1	;	1	0;	;	1
30	XYZ 490	$Lr 15 d 、 Lr 46 e$	;	;	;	2	4	;1	3+	;1	;	1	1	1	0	1	0	;	1
31	XYZ 491	$Lr 37 e 、 Lr 46 e$	4	3+	3+	3+	4	3+	3+	3+	4	1	3	3	4	4	4	4	1
32	XYZ 492	$Lr 46 e$	;1	3	3	1+	;1	1	3	;1	2	1	1	1	2	1	1	1	1
33	XYZ 493	$Lr 11 d 、 Lr 46 e$	1	1+	3	;	;1	3+	1	;1	;1	1	1	;	2	1	1	;	1
34	XYZ 494	$Lr 37 e 、 Lr 46 e$	3	2	3	3C	3+	3+	12	3	2+	2+	2+	2	3C	1	2	;	1
35	XYZ 495	$Lr 11 d 、 Lr 14 ac 、 Lr 26 c 、 Lr 37 e 、 Lr 46 e$	;1	;	;	;1	;1	;1	1	;1	1	12	1	;	1	1	3	;	1
36	XYZ 496	$Lr 37 e 、 Lr 46 e$	;	;	;	;1	;1	;1	;	;1	;	;	1	;	1	1	;	1	1
37	XYZ 497	$Lr 15 d 、 Lr 34 e 、 Lr 37 e 、 Lr 46 e$	1	;	;	3	3+	;1	1	3-	;1	1	1	;	1	1	;	1	1
38	XYZ 498	$Lr 37 e 、 Lr 46 e$	;	;	;	;	;	;	;1	;	;1	;	1	;	1	1	1	;	1
39	XYZ 499	$Lr 37 e 、 Lr 46 e$	;	;	;	1	;	;	;1	;	1	;	;1	;	1	1	1	;	1
40	XYZ 500	$Lr 10 e 、 Lr 15 d 、 Lr 46 e$	;	;	;	;	;	;	;1	;1	;	;1	;1	;	1	1	1	;	1

图1 13个分子标记对40份国外小麦品种(系)的部分PCR检测
M.DL2000;Z.郑州5389;1.XYZ461;2.XYZ462;3.XYZ463;4.XYZ464;5.XYZ465;6.XYZ466;7.XYZ467;8.XYZ468;9.XYZ469;10.XYZ470;11.XYZ471。Lr19和Lr26右上角的+、-表示检测所使用的标记不同。

Fig.1 Detection of partial PCR results amplified by 13 molecular markers in 40 foreign wheat varieties (lines)
M.DL2000;Z.Zhengzhou 5389;1.XYZ461;2.XYZ462;3.XYZ463;4.XYZ464;5.XYZ465;6.XYZ466;7.XYZ467;8.XYZ468;9.XYZ469;10.XYZ470;11.XYZ471.The+,-in the upper right corner of the Lr19 and Lr26 indicate that the marks used for the detection are different.

图2 Lr46分子标记检测
M.DNA Marker;1.TcLr46;2.郑州5389;3.XYZ461;4.XYZ462;5.XYZ463;6.XYZ464;7.XYZ465;8.XYZ466;9.XYZ467;10.XYZ468;11.XYZ469;12.XYZ470;13.XYZ471;14.XYZ472;15.XYZ473;16.XYZ474;17.XYZ475;18.XYZ476;19.XYZ477;20.XYZ478;21.XYZ479;22.XYZ480;23.XYZ481;24.XYZ482。

Fig.2 Molecular marker detection for Lr46 of wheat
M.DNA Marker;1.TcLr46;2.Zhengzhou 5389;3.XYZ461;4.XYZ462;5.XYZ463;6.XYZ464;7.XYZ465;8.XYZ466;9.XYZ467;10.XYZ468;11.XYZ469;12.XYZ470;13.XYZ471;14.XYZ472;15.XYZ473;16.XYZ474;17.XYZ475;18.XYZ476;19.XYZ477;20.XYZ478;21.XYZ479;22. XYZ480;23.XYZ481;24.XYZ482.

表4 22个成株抗性品系与小麦叶锈菌混合生理小种的苗期互作侵染型及成株期田间最终严重度(FDS)

Tab.4 Seedling infection types and final disease severity(FDS)in field with mixed Puccinia triticina(Pt)races of 22 wheat lines with slow-rusting resistance

序号 No.	品系 Lines	成株期抗病基因 APR genes	苗期混合生理小种侵染型 Seedling ITs to mixed Pt isolates	2018—2019年周口FDS/% 2018—2019 Zhoukou FDS	2019—2020年保定FDS/% 2019—2020 Baoding FDS
1	XYZ461	Lr46	3	5	10
3	XYZ463	Lr37、Lr46	3	5	10
4	XYZ464	Lr37、Lr46	3	1	5
5	XYZ465	Lr34、Lr37、Lr46	3	10	10
6	XYZ466	Lr37、Lr46	3	5	5
7	XYZ467	Lr46	3	5	10
8	XYZ468	Lr34、Lr37、Lr46	3	5	5
9	XYZ469	Lr37、Lr46	3+	1	5
10	XYZ470	Lr37、Lr46	3	10	10
11	XYZ471	Lr37、Lr46	3	5	10
12	XYZ472	Lr37、Lr46	3	5	5
16	XYZ476	Lr34、Lr46	3+	10	10
17	XYZ477	Lr34、Lr37、Lr46	3	1	1
22	XYZ482	Lr34、Lr37、Lr46	3	10	10
23	XYZ483	Lr46	3	1	10
24	XYZ484	Lr46	3	10	10
26	XYZ486	Lr46	3+	1	10
27	XYZ487	Lr46	4	5	5
29	XYZ489	Lr46	3	10	10
30	XYZ490	Lr46	4	10	5
35	XYZ495	Lr37、Lr46	3	1	1
37	XYZ497	Lr37、Lr46	3+	5	1
SAAR			3+	1	5
郑州5389	Zhengzhou 5389		3+	80	90
LSD				11.4

表5 2018—2019年,2019—2020年40个小麦品系以及慢锈对照SAAR和感病对照郑州5389的最终严重度数据方差分析

Tab.5 Analysis of variance for the final disease severity of the 40 wheat lines,slow rusting check SAAR and susceptible check Zhengzhou 5389 in 2018—2019 and 2019—2020

差异源 Source of variation	自由度 df	均方差 MS	F值 F value	P
品系 Lines	59	3 842.052 5	59.18^*	0.000 1
环境 Environment	1	1 118.016 7	17.22^*	0.000 1
重复 Repetition	1	220.416 7	3.39	0.770 3
品系×环境 Lines×Environment	59	113.465 8	1.17^*	0.006 5
品系×重复 Lines×Repetition	59	49.272 6	0.76	0.854 7
误差 Error	60	64.925 0

参考文献 34

[1]	Khan M H,Bukhari A,Dar Z A,Rizvi S M.Status and strategies in breeding for rust resistance in wheat[J].Agricultural Sciences,2013,4(6):292-301. doi: 10.4236/as.2013.46042.
[2]	Huerta-Espino J,Singh R P,Germán S,McCallum B D,Park R F,Chen W Q,Bhardwaj S C,Goyeau H.Global status of wheat leaf rust caused by Puccinia triticina[J].Euphytica,2011,179(1):143-160.doi:10.1007/s10681-011-0361-x.
[3]	李振岐,曾士迈.中国小麦锈病[M].北京:中国农业出版社,2002:362.
	Li Z Q,Zeng S M.Wheat rust in China[M].Beijing:China Agriculture Press, 2002:362.
[4]	Kumar S,Bhardwaj S C,Gangwar O P,Sharma A,Qureshi N,Kumaran V V,Khan H,Prasad P,Miah H,Singh G P,Sharma K,Verma H,Forrest K L,Trethowan R M,Bariana H S,Bansal U K.Lr80:A new and widely effective source of leaf rust resistance of wheat for enhancing diversity of resistance among modern cultivars[J].Theoretical and Applied Genetics,2021,134(3):849-858.doi:10.1007/s00122-020-03735-5.
[5]	Dyck P L.The association of a gene for leaf rust resistance with the chromosome 7D suppressor of stem rust resistance in common wheat[J].Genome,1987,29(3):467-469.doi:10.1094/PHYTO.1998.88.9.890.
[6]	Singh R P,Mujeeb-Kazi A,Huerta-Espino J.Lr46:a gene conferring slow-rusting resistance to leaf rust in wheat[J].Phytopathology,1998,88(9):890-894.doi:10.1094/PHYTO.1998.88.9.890.
[7]	Herrera-Foessel S A,Lagudah E S,Huerta-Espino J,Hayden M J,Bariana H S,Singh D,Singh R P.New slow-rusting leaf rust and stripe rust resistance genes Lr67 and Yr46 in wheat are pleiotropic or closely linked[J].Theoretical and Applied Genetics,2011,122(1):239-249.doi:10.1007/s00122-010-1439-x.
[8]	Herrera-Foessel S A,Singh R P,Huerta-Espino J,Rosewarne G M,Periyannan S K,Viccars L,Calvo-Salazar V,Lan C X,Lagudah E S.Lr68 :a new gene conferring slow rusting resistance to leaf rust in wheat[J].Theoretical and Applied Genetics,2012,124(8):1475-1486.doi:10.1007/s00122-012-1802-1.
[9]	Hong M.Expression of adult resistance to stripe rust at different growth stages of wheat[J].Plant Disease,1996,80(4):375.doi:10.1094/pd-80-0375.
[10]	Li Z F,Xia X C,He Z H,Li X,Zhang L J,Wang H Y,Meng Q F,Yang W X,Li G Q,Liu D Q.Seedling and slow rusting resistance to leaf rust in Chinese wheat cultivars[J].Plant Disease,2010,94(1):45-53.doi:10.1094/PDIS-94-1-0045.
[11]	王思曼,张静,张培培,Gebrewahid Takele Weldu,闫晓翠,刘大群,李在峰.69份国外小麦品种(系)抗叶锈性鉴定及基因分析[J].麦类作物学报,2020,40(10):1166-1174.doi:10.7606/j.issn.1009-1041.2020.10.03.
	Wang S M,Zhang J,Zhang P P,Weldu G T,Yan X C,Liu D Q,Li Z F.Leaf rust resistance identification and gene analysis of 69 foreign wheat varieties(lines)[J].Journal of Triticeae Crops,2020,40(10):1166-1174.
[12]	赵丽娜,任晓娣,胡亚亚,张涛,张娜,杨文香,刘大群.23 份中国小麦微核心种质抗叶锈性评价[J].中国农业科学,2013,46(3):441-450.doi:10.3864/j.issn.0578-1752.2013.03.001.
	Zhao L N,Ren X D,Hu Y Y,Zhang T,Zhang N,Yang W X,Liu D Q.Evaluation of wheat leaf rust resistance of 23 Chinese wheat mini-core collections[J]. Scientia Agricultura Sinica,2013,46(3):441-450.
[13]	焦悦,王思曼,赵喜兰,张培培,郑慧敏,李在峰,刘大群.50个国外小麦品种(系)抗叶锈性鉴定[J].河南农业科学,2019,48(12):79-88.doi:10.15933/j.cnki.1004-3268.2019.12.012.
	Jiao Y,Wang S M,Zhao X L,Zhang P P,Zheng H M,Li Z F,Liu D Q.Plant resistance to leaf rust in 50 foreign wheat cultivars[J].Journal of Henan Agricultural Sciences,2019,48(12):79-88.
[14]	Gao P,Zhou Y,Gebrewahid T W,Zhang P P,Yan X C,Li X,Yao Z J,Li Z F,Liu D Q.Identification of known leaf rust resistance genes in common wheat cultivars from Sichuan Province in China[J].Crop Protection,2019,115:122-129.doi:10.1016/j.cropro.2018.09.012.
[15]	Zhang P P,Gebrewahid T W,Zhou Y,Li Q L,Li Z F,Liu D Q.Seedling and adult plant resistance to leaf rust in 46 Chinese bread wheat landraces and 39 wheat lines with known Lr genes[J].Journal of Integrative Agriculture,2019,18(5):1014-1023.doi:10.1016/s2095.3119(19)62575-X.
[16]	杨文香. 21个小麦品种(系)抗叶锈性基因推导[J].河北农业大学学报,2000,23(3):69-72.doi:10.3969/j.issn.1000-1573.2000.03.016.
	Yang W X.The postulated genes for resistance to leaf rust in 21 wheat cultivars used in Hebei Province[J].Journal of Hebei Agricultural University,2000,23(3):69-72.
[17]	Long D L,Kolmer J A.A North American system of nomenclature for Puccinia recondita f.sp.tritici[J].Phyt opathology,1989,79(5):525-529.doi:10.1094/Phyto-79-525.
[18]	Roelfs A,Singh R P,Saari E.Rust diseases of wheat:concepts and methods of disease management[M].Mexico:CIMMYT,1992.
[19]	Dubin H J,Torres E.Causes and consequences of the 1976—1977 wheat leaf rust epidemic in northwest Mexico[J].Annual Review of Phytopathology,1981,19:41-49.doi:10.1146/annurev.py.19.090181.000353.
[20]	Sharp P J,Kreis M,Shewry P R,Gale M D.Location of β-amylase sequences in wheat and its relatives[J].Theoretical and Applied Genetics,1988,75(2):286-290.doi:10.1007/BF00303966.
[21]	Feuillet C,Messmer M,Schachermayr G,Keller B.Genetic and physical characterization of the LR1 leaf rust resistance locus in wheat(Triticum aestivum L.)[J].Molecular and General Genetics,1995,248(5):553-562.doi:10.1007/BF02423451.
[22]	Schachermayr G,Siedler H,Gale M D,Winzeler H,Winzeler M,Keller B.Identification and localization of molecular markers linked to the Lr9 leaf rust resistance gene of wheat[J].Theoretical and Applied Genetics,1994,88(1):110-115.doi:10.1007/BF00222402.
[23]	Neu C,Stein N,Keller B.Genetic mapping of the Lr20-Pm1 resistance locus reveals suppressed recombination on chromosome arm 7AL in hexaploid wheat[J].Genome,2002,45(4):737-744.doi:10.1139/g02-040.
[24]	Schachermayr G M,Messmer M M,Feuillet C,Winzeler H,Winzeler M,Keller B.Identification of molecular markers linked to the Agropyron elongatum-derived leaf rust resistance gene Lr24 in wheat[J].Theoretical and Applied Genetics,1995,90(7/8):982-990.doi:10.1007/BF00222911.
[25]	Shan X,Blake T K,Talbert L E.Conversion of AFLP markers to sequence-specific PCR markers in barley and wheat[J].Theoretical and Applied Genetics,1999,98(6/7):1072-1078.doi:10.1007/s001220051169.
[26]	William H M,Hoisington D,Singh R P,González-de-león D.Detection of quantitative trait loci associated with leaf rust resistance in bread wheat[J].Genome,1997,40(2):253-260.doi:10.1139/g97-036.
[27]	Seyfarth R,Feuillet C,Schachermayr G,Winzeler M,Keller B.Development of a molecular marker for the adult plant leaf rust resistance gene Lr35 in wheat[J].Theoretical and Applied Genetics,1999,99(3/4):554-560.doi:10.1007/s001220051268.
[28]	William M,Singh R P,Huerta-Espino J,Islas S O,Hoisington D.Molecular marker mapping of leaf rust resistance gene lr46 and its association with stripe rust resistance gene yr29 in wheat[J].Phytopathology,2003,93(2):153-159.doi:10.1094/PHYTO.2003.93.2.153.
[29]	Helguera M,Khan I A,Dubcovsky J.Development of PCR markers for the wheat leaf rust resistance gene Lr47[J].Theoretical and Applied Genetics,2000,100(7):625-631.doi:10.1007/s001220051397.
[30]	Helguera M,Vanzetti L,Soria M,Khan I A,Kolmer J,Dubcovsky J.PCR markers for Triticum Speltoides leaf rust resistance gene Lr51 and their use to develop isogenic hard red spring wheat lines[J].Crop Science,2005,45(2):728-734.doi:10.2135/cropscis2005.0728.
[31]	Cloutier S,McCallum B D,Loutre C,Banks T W,Wicker T,Feuillet C,Keller B,Jordan M C.Leaf rust resistance gene Lr1,isolated from bread wheat(Triticum aestivum L.)is a member of the large psr567 gene family[J].Plant Molecular Biology,2007,65(1/2):93-106.doi:10.1007/s11103-007-9201-8.
[32]	Singh R P,Rajaram S.Genetics of adult-plant resistance of leaf rust in Frontana and three CIMMYT wheats[J].Genome,1992,35(1):24-31.doi:10.1139/g92-004.
[33]	张小村,李斯深,赵新华,范玉顶,李瑞军.小麦纹枯病抗性的QTL分析和抗病基因的分子标记[J].植物遗传资源学报,2005,6(3):276-279.doi:10.3969/j.issn.1672-1810.2005.03.007.
	Zhang X C,Li S S,Zhao X H,Fan Y D,Li R J.QTL and molecular markers for resistance gene of wheat sharp eyespot[J].Journal of Plant Genetic Resources,2005,6(3):276-279.
[34]	韩烨,何中虎,夏先春,李星,李在峰,刘大群.CIMMYT小麦材料的苗期和成株抗叶锈病鉴定[J].作物学报,2011,37(7):1125-1133.doi:10.3724/SP.J.1006.2011.01125.
	Han Y,He Z H,Xia X C,Li X,Li Z F,Liu D Q.Seedling and slow rusting resistances to leaf rust in CIMMYT wheat lines[J].Acta Agronomica Sinica,2011,37(7):1125-1133.

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