桑轮纹病发生区桑叶表面细菌群落结构在冠层内的分异

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尹诗琳^{1, 2,},
张建华^{1, 3},
李星月⁴,
唐甜¹,
王谢^{1, 3,,}
1.四川省农业科学院土壤肥料研究所成都 610066
2.四川农业大学林学院成都 611130
3.农业部西南山地农业环境重点实验室成都 610066
4.四川省农业科学院植物保护研究所成都 610066
基金项目: 国家现代农业产业技术体系建设专项CARS-18

详细信息

作者简介:尹诗琳, 主要研究方向为桑园土壤微生物多样性与病虫害生物防治。E-mail: yinshilin6793@163.com

通讯作者:王谢, 主要研究方向为桑树的土壤-植物-微生物系统结构与功能。E-mail: wangxiechangde@hotmail.com

中图分类号:S888.71

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出版历程

收稿日期:2020-07-06
录用日期:2020-09-15
刊出日期:2021-03-01

Differentiation in the bacterial community structure of mulberry leaf surfaces in the canopy where mulberry ring rot disease occurs

YIN Shilin^{1, 2,},
ZHANG Jianhua^{1, 3},
LI Xingyue⁴,
TANG Tian¹,
WANG Xie^{1, 3,,}
1. Institute of Soil and Fertilizer, Sichuan Academy of Agricultural Sciences, Chengdu 610066, China
2. College of Forestry, Sichuan Agricultural University, Chengdu 611130, China
3. Key Laboratory of Southwest Mountain Agricultural Environment of the Ministry of Agriculture, Chengdu 610066, China
4. Institute of Plant Protection, Sichuan Academy of Agricultural Sciences, Chengdu 610066, China
Funds: the Special Fund for the Industrial System Construction of Modern Agriculture of ChinaCARS-18

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Corresponding author:WANG Xie, E-mail: wangxiechangde@hotmail.com

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摘要
摘要:探明桑轮纹病发生区冠层内叶表面微生物多样性、结构组成和功能的变化情况，有助于快速筛选控制桑轮纹病病原菌（Gonatophragmium mori）潜在的拮抗微生物。本研究基于高通量测序技术，分析冠层上部和下部叶片上表面和下表面细菌相对丰度的差异。结果发现：1）在多样性指数（Shannon）上，冠层上部叶片是冠层下部叶片的1.26倍，冠层内叶片上表面是下表面的1.49倍。2）在结构上，冠层下部叶片的下表面（LB）和冠层上部叶片的下表面（UB）的优势属均为泛菌属（Pantoea），其相对丰度分别为38.04%和25.31%，而冠层下部叶片的上表面（LS）为沙雷氏菌属（Serratia）、冠层上部叶片的上表面（US）为寡养单胞菌属（Stenotrophomonas），其相对丰度分别为18.0%、23.73%。3）在功能上，冠层下部叶片细菌的碳水化合物和氨基酸的运输和代谢功能比冠层上部叶片强，然而脂质的运输和代谢功能比冠层上部叶片弱；冠层上部叶片上表面细菌的细胞壁生物发生功能比下表面强，而氨基酸的运输和代谢功能弱于下表面；冠层下部叶片上表面细菌的次生代谢物的生物合成、运输和分解代谢和脂质运输和代谢功能强于下表面，而细胞运动比下表面弱。4）冠层内叶片表面芽孢杆菌属（Bacillus）、不粘柄菌属（Asticcacaulis）和苯基杆菌属（Phenylobacterium）的相对丰度与病原菌的相对丰度负相关性显著（P < 0.05），相关性最大的为芽孢杆菌属，系数为-0.87。上述结果表明桑轮纹病发生区冠层内桑叶上下表面细菌群落的多样性、结构组成和功能存在显著差异，对进一步研究桑轮纹病的生物防控具有一定的科学意义。
关键词:桑轮纹病/
冠层部位/
多样性指数/
功能/
高通量测序
Abstract:To identify the antagonistic bacteria of Gonatophragmium, this study investigated variations in the diversity, structure, and function of phylloplane microorganisms in the mulberry ring rot disease areas of mulberry canopy. Using high-throughput sequencing technology, we analyzed the relative abundance of bacteria on the upper and lower surfaces of the upper and lower leaves in the canopy. The results showed that 1) the Shannon index in upper canopy was 1.26 times of that in the lower canopy, and the upper surface of leaves was 1.49 times of that in the lower surface. 2) The dominant genera in the lower surface of a leaf in the lower canopy (LB) and the lower surface of a leaf in the upper canopy (UB) were the genus Pantoea, with relative abundances of 38.04% and 25.31%, respectively. In the upper surface of a leaf in the lower canopy (LS), it was Serratia and the upper surface of a leaf in the upper canopy (US) was Stenotrophomonas, with relative abundances of 18.0% and 23.73%, respectively. 3) The transport and metabolism of carbohydrates and amino acids of bacteria in the lower canopy leaves were stronger than those in the upper canopy leaves; however, lipid metabolism and transport were weaker in the lower canopy leaves. The cell wall biogenesis function of bacteria on the upper surface of leaf in upper canopy was stronger than that on the lower surface, whereas the amino acid transport and metabolism functions were weaker on the upper surface. The biosynthesis, transport, and catabolism of secondary metabolites, lipid transport, and the metabolism of bacteria on the upper surface of lower canopy leaves were stronger than those on the lower surface, but cell movement was weaker on the upper surface. The relative abundances of Bacillus, Asticcacaulis, and Phenylobacterium were significantly negatively related to the relative abundance of pathogens (P < 0.05); the most significant correlation was for Bacillus (-0.87). These results indicate significant differences in the diversity, structure, and function of bacterial communities on the upper and lower surfaces of mulberry leaves in the canopy, which will assist further research on the biological control of mulberry ring rot disease.
Key words:Mulberry ring rot/
Canopy part/
Diversity index/
Function/
High-throughput sequencing

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图1不同冠层位置桑叶叶片上下表面细菌OTU丰度的样本韦恩图和聚类树图
LB: 冠层下部叶片的下表面; LS: 冠层下部叶片的上表面; UB: 冠层上部叶片的下表面; US: 冠层上部叶片的上表面。
Figure1.Venn diagram and cluster tree of bacteria based on OTU number of upper and lower surfaces of leaves from different parts of canopy of mulberry
LB: lower surface of leaf from lower canopy; LS: upper surface of leaf from lower canopy; UB: lower surface of leaf from upper canopy; US: upper surface of leaf from upper canopy.

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表1高通量测序使用的引物
Table1.High throughput sequencing primers

引物Primer	序列5'-3' Sequence 5'-3'
341F	CCCTACACGACGCTCTTCCGATCTG (barcode) CCTACGGGNGGCWGCAG
805R	GACTGGAGTTCCTTGGCACCCGAGAATTCCAGACTACHVGGGTATCTAATCC
ITS1	CCCTACACGACGCTCTTCCGATCTN (barcode) CTTGGTCATTTAGAGGAAGTAA
ITS2-Rev	GTGACTGGAGTTCCTTGGCACCCGAGAATTCCAGCTGCGTTCTTCATCGATGC

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表2不同冠层位置桑叶叶片上下表面细菌序列长度统计表
Table2.Sequence length statistics of bicteria in upper and lower surfaces of leaves from different parts of canopy of mulberry

样本 Sample	原始序列 Raw_num	优化序列 Clean_num	高质量序列 Filtered_num	序列长度 Mean_len
冠层下部叶片的下表面Lower surface of leaf from lower canopy (LB)	254 376	234 265	227 294	423.20
冠层下部叶片的上表面Upper surface of leaf form lower canopy (LS)	227 695	209 246	203 185	419.27
冠层上部叶片的下表面Lower surface of leaf from upper canopy (UB)	253 532	232 206	228 482	420.94
冠层上部叶片的上表面Upper surface of leaf from upper canopy (US)	212 680	195 515	194 092	418.25

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表3不同冠层位置桑叶叶片上下表面细菌群落丰富度和多样性
Table3.Bacterial community richness and diversity on upper and lower surfaces of leaves from different parts of canopy of mulberry

样本 Sample	Chao1指数 Chao1 index	ACE指数 ACE index	香农指数 Shannon index	辛普森指数 Simpson index
LB	1184.53±95.24a	2079.80±288.12a	1.44±0.47b	0.46±0.18a
LS	1058.43±215.96a	1610.93±406.28ab	2.70±0.64a	0.20±0.13ab
UB	1307.57±523.09a	1790.37±498.46ab	2.32±0.16a	0.25±0.03a
US	1007.92±121.93a	1464.64±209.73b	2.88±0.46a	0.14±0.06b
LB: 冠层下部叶片的下表面; LS: 冠层下部叶片的上表面; UB: 冠层上部叶片的下表面; US: 冠层上部叶片的上表面。表中数值为平均值±标准偏差, 同列不同小写字母表示处理间差异显著(P < 0.05)。LB: lower surface of leaf from lower canopy; LS: upper surface of leaf from lower canopy; UB: lower surface of leaf from upper canopy; US: upper surface of leaf from upper canopy. The values in the table are mean±SD. Different lowercase letters in a column indicate significant differences at P < 0.05 level.

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表4不同冠层位置桑叶叶片上下表面细菌各分类水平上数量排名前10物种相对丰度在各样本中所占比例
Table4.Proportions of relative abundances of top ten items at each taxonomic level of bacteria on upper and lower surfaces ofleaves from different parts of canopy of mulberry ?%

分类Taxonomic	名称Name	LB	LS	UB	US
门Phylum	变形菌门Proteobacteria	99.29	88.04	91.22	78.52
	拟杆菌门Bacteroidetes	0.23	10.45	5.98	19.16
	放线菌门Actinobacteria	0.15	1.13	0.97	1.21
	厚壁菌门Firmicutes	0.22	0.18	0.80	0.64
	广古生菌门Euryarchaeota	0.02	0.08	0.48	0.23
	酸杆菌门Acidobacteria	0.00	0.03	0.16	0.03
	未分类菌门Unclassified	0.03	0.01	0.09	0.02
	浮霉菌门Planctomycetes	0.04	0.03	0.05	0.02
	Candidatus Saccharibacteria	0.01	0.01	0.04	0.01
	奇古菌门Thaumarchaeota	0.00	0.00	0.04	0.02
纲Class	γ-变形菌纲Gammaproteobacteria	94.99	67.53	77.77	52.82
	α-变形菌纲Alphaproteobacteria	4.06	16.09	10.47	18.98
	黄杆菌纲Flavobacteriia	0.08	8.31	5.04	10.30
	β-变形菌纲Betaproteobacteria	0.23	4.42	2.79	6.68
	鞘脂杆菌纲Sphingobacteria	0.08	1.86	0.45	8.21
	放线菌纲Actinobacteria	0.15	1.13	0.97	1.21
	芽胞杆菌纲Bacilli	0.19	0.12	0.47	0.43
	纤维粘网菌纲Cytophagia	0.06	0.23	0.12	0.46
	拟杆菌纲Bacteroidia	0.01	0.04	0.35	0.19
	梭菌纲Clostridia	0.02	0.05	0.30	0.18
目Order	肠杆菌目Enterobacteriales	84.10	43.80	48.65	11.72
	假单胞菌目Pseudomonadales	7.31	10.47	19.31	16.44
	黄色单胞杆菌目Xanthomonadales	3.58	13.26	9.77	24.62
	黄杆菌目Flavobacteriales	0.08	8.31	5.05	10.31
	根瘤菌目Rhizobiales	2.44	7.12	4.94	4.99
	鞘脂单胞菌目Sphingomonadales	0.84	4.39	2.22	9.01
	伯克氏菌目Burkholderiales	0.23	4.41	2.73	6.60
	鞘脂杆菌目Sphingobacteriales	0.08	1.86	0.45	8.22
	柄杆菌目Caulobacterales	0.19	1.74	2.78	4.59
	红螺菌目Rhodospirales	0.47	0.27	0.22	0.15
科Family	肠杆菌科Enterobacteriaceae	84.09	43.80	48.71	11.72
	黄色单胞菌科Xanthomonadaceae	3.58	13.26	9.77	24.62
	莫拉式菌科Moraxellaceae	2.49	2.86	14.68	12.59
	黄杆菌科Flavobacteriaceae	0.08	8.31	5.05	10.31
科Family	假单胞菌科Pseudomonadaceae	4.82	7.60	4.64	3.85
	鞘脂菌科Sphingomonadaceae	0.83	4.32	2.05	8.90
	根瘤菌科Rhizobiaceae	1.42	5.06	2.46	2.96
	鞘脂杆菌科Sphingobacteriaceae	0.05	1.85	0.41	8.18
	柄杆菌科Caulobacteraceae	0.19	1.74	2.78	4.59
	丛毛单胞菌科Comamonadaceae	0.14	3.85	2.39	0.76
属Genus	泛菌属Pantoea	38.04	13.86	25.31	7.69
	未分类菌属Unclassified	20.99	7.40	23.29	3.75
	寡养单胞菌属Stenotrophomonas	3.35	12.17	7.40	23.73
	沙雷氏菌属Serratia	17.74	18.00	0.37	0.00
	不动杆菌属Acinetobacter	0.28	1.72	14.62	12.48
	假单胞菌属Pseudomonas	4.81	7.59	4.63	3.83
	金黄杆菌属Chryseobacterium	0.00	6.45	3.51	9.39
	鞘氨醇单胞菌属Sphingomonas	0.77	3.79	1.88	8.70
	克吕沃尔菌属Kluyvera	6.95	5.09	0.00	0.40
	短波单胞菌属Brevundimonas	0.19	1.67	2.50	4.46
LB: 冠层下部叶片的下表面; LS: 冠层下部叶片的上表面; UB: 冠层上部叶片的下表面; US: 冠层上部叶片的上表面。LB: lower surface of leaf from lower canopy; LS: upper surface of leaf from lower canopy; UB: lower surface of leaf from upper canopy; US: upper surface of leaf from upper canopy.

下载: 导出CSV
表5不同冠层位置桑叶叶片上下表面两两间细菌丰度存在显著差异(P < 0.05)的属
Table5.Genus significantly different (P < 0.05) among upper and lower surfaces of leaves from different parts of canopy of mulberry

菌属名称Genus	丰度Abundance		丰度差值Effect size	菌属名称Genus	丰度Abundance		丰度差值Effect size
菌属名称Genus	LB	LS	丰度差值Effect size	菌属名称Genus	LB	UB	丰度差值Effect size
勒克氏菌属Leclercia	10	1	0.00	马赛菌属Massilia	37	204	?0.08
薄层菌属Hymenobacter	7	98	?0.05	微杆菌属Microbacterium	26	205	?0.09
金黄杆菌属Chryseobacterium	95	11 843	?6.49	大理石雕菌属Marmoricola	1	37	?0.02
地嗜皮菌属Geodermatophilus	10	152	?0.08	副球菌属Marmoricola	6	75	?0.04
地嗜皮菌属Geodermatophilus	10	152	?0.08	鞘氨醇单胞菌属Sphingomonas	1583	3866	?1.23
	UB	US			LS	US
鞘氨醇单胞菌属Sphingomonas	3866	15 237	?6.70	克雷伯氏菌属Klebsiella	6	0	0.00
异常球菌属Deinococcus	9	41	?0.02	巨球型菌属Megasphaera	0	6	0.00
勒克氏菌属Leclercia	9	0	0.00	异常球菌属Deinococcus	6	41	?0.02
				沉积物杆状菌属Sediminibacterium	0	15	?0.01
LB: 冠层下部叶片的下表面; LS: 冠层下部叶片的上表面; UB: 冠层上部叶片的下表面; US: 冠层上部叶片的上表面。LB: lower surface of leaf from lower canopy; LS: upper surface of leaf from lower canopy; UB: lower surface of leaf from upper canopy; US: upper surface of leaf from upper canopy.

下载: 导出CSV
表6不同冠层位置桑叶叶片上下表面两两间丰度存在显著差异(P < 0.05)的细菌功能分类
Table6.Functional classification with significant differences (P < 0.05) among upper and lower surfaces of leaves from different parts of canopy of mulberry

样本 Sample	功能预测 Function	均值之差 Difference between means
LB-LS	次生代谢物的生物合成、运输和分解代谢 Secondary metabolites biosynthesis, transport, and catabolism	?0.45
	脂质运输和代谢 Lipid transport and metabolism	?0.71
	细胞运动Cell motility	0.35
UB-US	氨基酸的运输和代谢 Amino acid transport and metabolism	0.85
	细胞壁生物发生 Cell wall/membrane/envelope biogenesis	?0.60
LB-UB	脂质运输和代谢 Lipid transport and metabolism	?0.87
	碳水化合物运输和代谢 Carbohydrate transport and metabolism	1.82
LS-US	氨基酸的运输和代谢 Amino acid transport and metabolism	0.92
LB: 冠层下部叶片的下表面; LS: 冠层下部叶片的上表面; UB: 冠层上部叶片的下表面; US: 冠层上部叶片的上表面。LB: lower surface of leaf from lower canopy; LS: upper surface of leaf from lower canopy; UB: lower surface of leaf from upper canopy; US: upper surface of leaf from upper canopy.

下载: 导出CSV

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