嫁接于3个枣园病砧的抗病接穗枣疯植原体检测及分子变异

doi:10.11707/j.1001-7488.20211105

摘要/Abstract

摘要：

目的: 检测3个枣园不同抗性枣树材料嫁接传毒后携带植原体状况，分析不同植原体株系传播流行、遗传变异和系统发育及其与枣树品种抗性之间的关系，揭示枣疯植原体群体遗传变异特性，为抗病品系合理利用与病害防控提供依据。方法: 4月采集不同抗性枣树材料嫁接到北京玉泉山枣树资源圃、北京昌平王家园和河北唐县军城镇三地枣园的发病砧木上，当年8-10月分别提取接穗和砧木组织样品总DNA，利用植原体16S rDNA、16S-23S间区（SR）、secA、tuf、fusA-tuf间区、himA及其下游假定蛋白基因等片段进行PCR和LAMP检测，并对PCR产物进行测序，分析遗传变异和系统进化。结果: 抗病品系的接穗材料症状严重度大都低于感病砧木；表现无症接穗T30通过直接和巢式PCR与LAMP均未检测到植原体；3个枣园枣树样品中的植原体16S rDNA、SR、tuf和secA基因序列同源性为100%；根据所测38份样品中himA及其下游假定蛋白基因序列在117、154和190 bp位点的碱基变异，军城镇样品序列聚于T-A型，玉泉山样品和河北唐县梁家沟枣疯序列聚于C-A型，而昌平样品和泗洪、泰安、通州枣疯序列聚于C-C型；根据所测29份样品中fusA-tuf基因间区序列在第39 bp位置有碱基A插入或缺失，将玉泉山和军城镇样品归于TJ2型，而国内其他地区各株系和昌平样品为TJ1型；himA及其下游假定蛋白基因序列及fusA-tuf基因间区序列在3个枣园各有优势菌株序列，序列分型对应且互相佐证。结论: 本研究首次测定枣树‘骏优2号’（Z18）优良品系具有高抗病性，并首次检测到3个枣园内枣疯植原体株系的himA及其下游假定蛋白基因与fusA-tuf基因间区存在明显地域差异，且与样品抗病性强弱无关联；初步排除玉泉山枣园植原体来源于江苏泗洪大枣移栽苗传病的假设。

关键词: 嫁接, 抗病性, 枣疯病, 植原体, 16S rDNA, himA及其下游假定蛋白

Abstract:

Objective: This study aimed to detect the jujube witches'-broom(JWB) phytoplasma status of jujubes cultivated in three jujube orchards after inoculating the trees by graft, and to analyze the transmission, epidemic, heritable variation, and phylogeny of different phytoplasma strains; and reveal the relationship between different phytoplasma strains and different resistance jujube cultivars. Method: In April 2017, different resistant jujube scions were collected and grafted on the phytoplasma-infected rootstocks cultivated in three jujube orchards including Yuquanshan and Changping orchards in Beijing and Juncheng orchard in Tang county, Hebei province. The total DNA of scion and rootstock tissue samples were extracted from August to October of that year, to be used as templates for PCR amplification. Specific PCR products of 16S rDNA, 16S-23S intergenic region(SR), secA, tuf, fusA-tuf intergenic region, himA and its downstream putative protein gene were sequenced. Genetic variation and phylogenetic evolution were analyzed with obtained sequence data. Result: The symptom severity of resistant scions was generally lower than that of susceptible rootstocks. No phytoplasma was detected in all of T30 scions without symptoms of witches'-broom by direct and nested PCR and LAMP. The homology of 16S rDNA, SR, tuf and secA gene sequences of JWB phytoplasma isolates from three orchards was 100%. According to the base variation of himA and its downstream putative protein gene at 117, 154 and 190 bp sites in 38 samples, JWB strains from Junchengzhen were clustered in the T-A type, JWB strains from Yuquanshan and Liangjiagou were clustered in the C-A type, while JWB strains from Changping were clustered in the C-C type. According to the insertion or deletion of base A at 39 bp in the sequence of the fusA-tuf intergenic region in 29 samples, JWB strains from Yuquanshan and Juncheng were clustered in the TJ2(A+) type, while others were clustered in the TJ1(A-) type. There were dominant strains in each orchard that cause the sequence types of both regions(himA and its downstream putative protein gene, intergenic region of fusA-tuf genes) were mutually corroborated. Conclusion: In this study 'Junyou No.2' (Z18) jujube is for the first time identified as the high resistant cultivar to JWB. The variations of different JWB strains from two orchards have no significant correlation with the disease resistance of cultivars, while with the regional difference. The gene sequences of JWB strains from the two regions provide the evidence that JWB strains of Yuquanshan are not originated from Sihong County, Jiangsu Province. The results are helpful to reveal the genetic variation characteristics of JWB phytoplasma population and provide a theory basis for rational utilization of resistant cultivars and JWB disease control.

Key words: graft, disease resistance, jujube witches'-broom disease, phytoplasma, 16S rDNA, himA and its downstream hypothetical protein

中图分类号:

S763.7

张文鑫,于少帅,田国忠,王合,任争光,王圣洁,孔德治,李永,林彩丽. 嫁接于3个枣园病砧的抗病接穗枣疯植原体检测及分子变异[J]. 林业科学, 2021, 57(11): 49-58.

Wenxin Zhang,Shaoshuai Yu,Guozhong Tian,He Wang,Zhengguang Ren,Shengjie Wang,Dezhi Kong,Yong Li,Caili Lin. Detection and Molecular Variation of Jujube Witches' Broom Phytoplasma in Resistant Jujube Scions Grafted on Diseased Root Stocks in Three Jujube Orchards[J]. Scientia Silvae Sinicae, 2021, 57(11): 49-58.

图/表 7

表1

表2

3个枣园不同品种接穗嫁接后症状表现及植原体检测结果①"

样品编号 Sample No.	树号 Tree No.	症状与病级Symptom and degrees				分子检测Molecular detection
样品编号 Sample No.	树号 Tree No.	品种 Cultivar	症状 Symptom	病级 Disease degrees		PCR	Nested PCR	LAMP
YQ-SC-1	Tree 1	T27	NS	0		+	+	+
YQ-SC-2	Tree 1	T27	YL	Ⅲ		+	+	+
YQ-SC-3	Tree 1	T27	P	Ⅰ		+	+	+
YQ-RS-1	Tree 1	T19	LL、WB	Ⅴ		+	+	+
YQ-RS-2	Tree 1	T19	LL、WB	Ⅲ		+	+	+
YQ-SC-4	Tree 1	Z18	YL	i		-	-	-
YQ-RS-3	Tree 1	T19	NS	0		+	+	+
YQ-RS-4	Tree 1	T19	LL、WB	Ⅴ		+	+	+
YQ-RS-5	Tree 1	T19	WB	Ⅲ		+	+	+
YQ-SC-5	Tree 1	50	LL、WB	Ⅴ		+	+	+
YQ-SC-6	Tree 2	T30	NS	0		-	-	-
YQ-Z18	Tree 3	Z18	NS	0		-	-	+
YQ-SC-7	Tree 4	T30	NS	0		-	-	-
YQ-SC-8	Tree 5	T30	NS	0		-	-	-
YQ-RS-6	Tree 5	M13	LL、WB	Ⅴ		+	+	+
YQ-M11	Tree 6	M11	NS	0		-	-	+
YQ-SC-9	Tree 7	M11	P	Ⅱ		+	+	+
YQ-SC-10	Tree 8	T27	LL、WB	Ⅴ		+	+	+
YQ-RS-7	Tree 8	M15	LL、WB	Ⅴ		+	+	+
YQ-SC-11	Tree 9	T30	LL、WB	Ⅴ		-	-	-
YQ-RS-8	Tree 9	M15	LL、WB	Ⅴ		+	+	+
YQ-T27	Tree 10	T27	NS	0		-	-	+
JC-RS-1	Tree 11	Pozao	LL、WB	Ⅴ		+	+	+
JC-SC-1	Tree 11	T27	P	Ⅱ		+	+	+
JC-RS-2	Tree 12	Pozao	LL	Ⅰ		-	-	+
JC-SC-2	Tree 12	T27	PE	Ⅰ		+	-	+
JC-RS-3	Tree 13	Pozao	LL、WB	Ⅴ		+	+	+
JC-SC-3	Tree 13	Z18	NS	0		+	-	+
JC-RS-4	Tree 14	Pozao	LL、WB	Ⅴ		-	-	+
JC-SC-4	Tree 14	Pozao	LL、WB	Ⅴ		+	-	+
JC-ST-1	Tree 15	Pozao	LL、WB	Ⅴ		-	-	+
JC-RS-5	Tree 15	Pozao	LL、WB	Ⅴ		-	-	-
JC-SC-5	Tree 15	Z25	LL	Ⅰ		+	+	+
JC-SC-6	Tree 16	50	LL、WB	Ⅴ		-	+	+
JC-SC-7	Tree 17	87	LL、WB	Ⅴ		+	+	+
JC-SC-8	Tree 18	127	LL、WB	Ⅴ		+	+	+
JC-SC-9	Tree 19	Z25	P	Ⅰ		-	-	+
CP-SC-1	Tree 20	T27	P	Ⅱ		-	-	+
CP-RS-1	Tree 20	Dongzao	NS	0		+	+	+
CP-SC-2	Tree 21	Z18	LL、PE	Ⅰ		+	+	+
CP-RS-2	Tree 21	Dongzao	LL、WB	Ⅴ		-	+	+
CP-SC-3	Tree 22	50	NS	0		+	+	+
CP-RS-3	Tree 22	Dongzao	LL	Ⅰ		-	+	+
CP-SC-4	Tree 23	87	PE	Ⅰ		-	-	-
CP-RS-4	Tree 23	Dongzao	LL	Ⅰ		+	+	+
CP-SC-5	Tree 24	T30	NS	0		-	-	-
CP-RS-5	Tree 24	Dongzao	LL	Ⅰ		-	-	-
CP-SC-6	Tree 25	M11	P	Ⅱ		+	+	+
CP-RS-6	Tree 25	Dongzao	LL	Ⅰ		+	+	+
CP-SC-7	Tree 26	L2	P	Ⅲ		+	+	+
CP-RS-9	Tree 26	Dongzao	LL、WB	Ⅱ		-	-	+
CP-SC-9	Tree 27	T31	WB	Ⅱ		-	-	-
CP-RS-7	Tree 27	Dongzao	NS	0		+	+	+
CAF-SC-1	Tree 28	T27	NS	0		-	-	-
CAF-SC-2	Tree 28	M11	IC	Ⅰ		+	+	+
CAF-SC-3	Tree 28	Jikang	PE、IC	Ⅱ		+	+	+
CAF-RS-1	Tree 28	Unknow	NS	0		-	-	-
JWB-JSSH-QC1	Tree 29	T19	LL、WB	Ⅴ		+	+	+
JWB-JSSH-FJT	Tree 30	T19	LL、WB	Ⅴ		+	+	+
JWB-JSSH-QC2	Tree 31	T19	LL、WB	Ⅴ		+	+	+
JWB-JSSH-QC3	Tree 32	T19	LL、WB	Ⅴ		+	+	+
JWB-SDTA	Tree 33	/	LL、WB	Ⅴ		+	+	+
JWB-HBTX-LJG	Tree 34	Pozao	LL、WB	Ⅴ		+	+	+
JWB-HBTX-SJG	Tree 35	Pozao	LL、WB	Ⅴ		+	+	+
JWB-SDTA-GSZD	Tree 36	Changhongzao	LL、WB	Ⅴ	+	+	+
JWB-BJTZ-RZY	Tree 37	/	LL、WB	Ⅴ	+	+	+

表2

表3

图1

表4

图2

图3

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抗性Resistance	健康接穗Scion	砧木/根蘖Rootstock/Stool shoot
高抗High resistance(HR)	骏枣Z. jujuba ‘Junzao’(T27)、壶瓶枣Z. jujuba ‘Hupingzao’(Z17)
普抗General resistance(R)	洪赵小枣Z. jujuba ‘Hongzhaoxiaozao’(T30)、蜂蜜罐枣Z. jujuba Fengmiguan(M11)、北京古树87号Beijing ancient tree No.87(87)、北京古树127号Beijing ancient tree No.127(127)
中抗Medium resistance(MR)	北京古树50号Beijing ancient tree No.50(50)
感病Susceptibility(S)		泗洪大枣Z. jujuba ‘Sihongdazao’(T19)、酸枣Z. spinosa (Suanzao)、婆枣Z. jujuba ‘Pozao’(Pozao)
高感High susceptibility(HS)	濮阳冬枣Z. jujuba ‘puyangdongzao’(L2)、冀抗Z. jujuba ‘Jikang No.1’(JK)	襄汾圆枣Z. jujuba Xiangfenyuanzao(M15)、晋枣Z. jujuba ‘Jinzao’(M13)、冬枣Z. jujuba ‘Dongzao’(Dongzao)
未知Unknown	石西大酸枣Z. spinosa ‘Shixi’(Z25)、骏优2号Z. jujuba ‘Junyou No.2’(Z18)

基因Genes	扩出数(条) Amplification number	扩出率 Amplification rate(%)	变异位点 Variation site	变异率 Variation rate(%)
16Sr	42	63.6	0	0
SR	42	63.6	0	0
SecA	31	47.0	0	0
Tuf	32	48.5	0	0
himA及其下游假定蛋白基因 himA and its downstream hypothetical protein gene	38	57.6	3	9.6
fusA-tuf基因间区 fusA-tuf intergenic region	29	43.9	1	18.9

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