上海大金山岛不同植被类型土壤细菌群落的变异

doi:10.17521/cjpe.2023.0172

摘要/Abstract

摘要：

揭示次生演替过程中土壤细菌群落对植被类型变化的响应格局, 有助于深化对生态系统地上地下相互作用机制的理解。该研究以上海大金山岛处于演替前、中和后期的落叶灌丛、落叶阔叶林和常绿阔叶林为对象, 通过测定土壤碳、氮、磷含量以及土壤细菌群落特征, 分析土壤细菌多样性、类群关系网络结构和标志性类群如何随植被演替而更替。结果发现: 土壤养分含量在常绿阔叶林显著高于落叶阔叶林, 但土壤细菌多样性在落叶阔叶林显著高于常绿阔叶林, 土壤养分含量和细菌多样性在落叶灌丛处于中等水平。土壤细菌的相关性网络节点、密度和复杂性在落叶阔叶林最高, 在落叶灌丛中等, 在常绿阔叶林最低。落叶灌丛和落叶阔叶林的优势土壤细菌分别为根瘤菌目(Rhizobiales)和伯克氏菌目(Burkholderiales)等具有潜在固氮功能的类群, 常绿阔叶林的优势土壤细菌为黄单胞菌目(Xanthomonadales)和嗜热芽菌目(Thermogemmatisporales)等具有潜在致病和抗病性功能的类群以及酸杆菌目(Acidobacteriales)等与纤维素降解相关的类群。该结果表明: 海岛植被演替过程中, 植物种类组成和土壤养分供给性的改变会极大地重塑土壤细菌多样性、群落组成、互作网络结构和标志类群。海岛常绿阔叶林显著更低的土壤细菌多样性、趋于松散简化的细菌网络结构以及具有潜在致病和抗病功能的标志类群的出现, 说明了顶极群落地下部分对地上部分退化趋势的响应。

关键词: 次生演替, 岛屿, 灌丛, 森林, 细菌多样性, 16S rDNA

Abstract:

Aims Revealing the response patterns of soil bacterial community to changes in vegetation type during secondary succession can improve our understanding of the mechanisms that structure the above- and below-ground interactions in ecosystems.

Methods To investigate how soil bacterial diversity, taxa network structure and biomarkers shift with vegetation succession, this study measured soil carbon, nitrogen and phosphorus contents, as well as soil bacterial community properties across shrubland, deciduous broadleaf forest and evergreen broadleaf forests, representing the early-, middle- and late-successional stages, respectively, on Dajinshan Island, Shanghai.

Important findings Soil nutrient contents were significantly higher in evergreen broadleaf forest than in deciduous broadleaf forest. However, soil bacterial diversity was significantly higher in deciduous broadleaf forest than in evergreen broadleaf forest, while soil nutrient content and bacterial diversity were medium in deciduous shrubland. The correlation network nodes, density and complexity of soil bacteria were highest in deciduous broadleaf forest, medium in deciduous shrubland, and lowest in evergreen broadleaf forest. The dominant soil bacteria in deciduous shrubland and broadleaf forest was Rhizobiales and Burkholderiales, respectively, which belong to functional group of nitrogen-fixing. The dominant soil bacterial in evergreen broadleaf forest were characterized by functional groups of pathogenicity and resistance such as Xanthomonadales and Thermogemmatisporales, and functional group associated with cellulose degradation such as Acidobacteriales. These results suggest that changes in plant species composition and soil nutrient availability during island vegetation succession can greatly reshape species diversity, community composition, interactive network structure and biomarkers of soil bacteria. In evergreen broadleaf forest, lowered soil bacterial diversity, simplified bacterial network structure, and emerged biomarkers of functional groups of pathogenicity and resistance suggest a response of belowground to the degraded trend of aboveground in the studied climax forest.

Key words: secondary succession, island, shrubland, forest, bacterial diversity, 16S rDNA

杨安娜, 李曾燕, 牟凌, 杨柏钰, 赛碧乐, 张立, 张增可, 王万胜, 杜运才, 由文辉, 阎恩荣. 上海大金山岛不同植被类型土壤细菌群落的变异. 植物生态学报, 2024, 48(3): 377-389. DOI: 10.17521/cjpe.2023.0172

YANG An-Na, LI Zeng-Yan, MOU Ling, YANG Bai-Yu, SAI Bi-Le, ZHANG Li, ZHANG Zeng-Ke, WANG Wan-Sheng, DU Yun-Cai, YOU Wen-Hui, YAN En-Rong. Variation in soil bacterial community across vegetation types in Dajinshan Island, Shanghai. Chinese Journal of Plant Ecology, 2024, 48(3): 377-389. DOI: 10.17521/cjpe.2023.0172

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2023.0172

https://www.plant-ecology.com/CN/Y2024/V48/I3/377

图/表 10

表1 大金山岛9个植物群落类型样地特征

Table 1 Characteristics of the studied plots across 9 plant communities in Dajinshan Island, Shanghai

表2 大金山岛不同植被类型的土壤化学性质(平均值±标准差)

Table 2 Soil chemical properties across vegetation types in Dajinshan Island, Shanghai (mean ± SD)

植被类型 Vegetation type	总氮含量 Total nitrogen content (g·kg^-1)	总磷含量 Total phosphorus content (g·kg^-1)	有机碳含量 Organic matter content (g·kg^-1)	速效氮含量 Available nitrogen content (mg·kg^-1)	速效磷含量 Available phosphorus content (mg·kg^-1)	速效钾含量 Available kalium content (mg·kg^-1)
落叶灌丛 Deciduous shrubland	0.61 ± 0.10^ab	0.11 ± 0.01^b	5.99 ± 1.37^ab	453.70 ± 154.30^ab	4.03 ± 1.00^ab	232.17 ± 29.60^a
落叶阔叶林 Deciduous broadleaf forest	0.49 ± 0.09^b	0.09 ± 0.01^b	4.94 ± 1.09^b	323.80 ± 69.20^b	4.12 ± 1.50^a	234.00 ± 75.30^a
常绿阔叶林 Evergreen broadleaf forest	0.66 ± 0.14^a	0.17 ± 0.04^a	7.86 ± 1.97^a	480.31 ± 132.08^a	14.10 ± 5.80^a	199.41 ± 45.20^a

图1 大金山岛3种植被类型土壤细菌的可操作分类单元(OTU)丰富度(A)、Chao1指数(B)、Shannon-Wiener指数(C)和Simpson指数(D)的差异。DF, 落叶阔叶林; DS, 落叶灌丛; EF, 常绿阔叶林。箱体间不同小写字母表示差异显著(p < 0.05)。

Fig. 1 Differences in operational taxonomic units (OTU) richness (A), Chao1 (B), Shannon-Wiener (C) and Simpson (D) indices of soil bacterial community across three vegetation types in Dajinshan Island, Shanghai. DF, deciduous broadleaf forest; DS, deciduous shrubland; EF, evergreen broadleaf forest. Different lowercase letters among boxplots indicate significant differences (p < 0.05).

图2 大金山岛3种植被类型土壤细菌群落结构的差异。AK, 速效钾含量; AN, 速效氮含量; AP, 速效磷含量; TC, 有机碳含量; TN, 总氮含量; TP, 总磷含量。DF, 落叶阔叶林; DS, 落叶灌丛; EF, 常绿阔叶林。NMDS, 非度量多维尺度。

Fig. 2 Difference in soil bacterial community structure among three vegetation types in Dajinshan Island, Shanghai. AK, available kalium content; AN, available nitrogen content; AP, available phosphorus content; TC, organic matter content; TN, total nitrogen content; TP, total phosphorus content. DF, deciduous broadleaf forest; DS, deciduous shrubland; EF, evergreen broadleaf forest. NMDS, non-metric multidimensional scaling.

图3 大金山岛3种植被类型土壤细菌类群相关性网络。任意两点连线说明可操作分类单元(OTU)之间相关性显著(p < 0.05), 红色代表显著正相关, 蓝色代表显著负相关。DF, 落叶阔叶林; DS, 落叶灌丛; EF, 常绿阔叶林。d, 网络直径; D, 密度; l, 路径长度; n, 节点数。

Fig. 3 Correlation networks of soil bacteria across three vegetation types in Dajinshan Island, Shanghai. The connection between any two points indicates that the correlation between operational taxonomic units (OTUs) is significant (p < 0.05). Red represents positive correlation, while blue represents negative correlation. DF, deciduous broadleaf forest; DS, deciduous shrubland; EF, evergreen broadleaf forest. d, network diameter; D, density; l, path length; n, number of nodes.

表3 大金山岛3种植被类型土壤细菌的相关性网络核心节点

Table 3 Core nodes in correlation networks of soil bacterial across three vegetation types in Dajinshan Island, Shanghai

植被类型 Vegetation type	OTU编号 OTU code	门 Phylum	纲 Class	目 Order	节点度 Nodes degree
落叶灌丛 Deciduous shrubland	OTU137	拟杆菌门 Bacteroidetes	腐螺旋菌纲 Saprospirae	腐螺旋菌目 Saprospirales	19
	OTU34	酸杆菌门 Acidobacteria	DA052	Ellin6513	19
	OTU52	酸杆菌门 Acidobacteria	DA052	Ellin6513	19
落叶阔叶林 Deciduous broadleaf forest	OTU98	变形菌门 Proteobacteria	γ变形菌纲 Gamma proteobacteria	黄单胞菌 Xanthomonadales	54
常绿阔叶林 Evergreen broadleaf forest	OTU104	酸杆菌门 Acidobacteria	DA052	Ellin6513	6
常绿阔叶林 Evergreen broadleaf forest	OTU47	WPS-2	-	-	6

图4 大金山岛3种植被类型土壤优势细菌门的相对丰度。DF, 落叶阔叶林; DS, 落叶灌丛; EF, 常绿阔叶林。

Fig. 4 Relative abundance distribution of dominant soil bacterial phyla across three vegetation types in Dajinshan Island, Shanghai. DF, deciduous broadleaf forest; DS, deciduous shrubland; EF, evergreen broadleaf forest.

表4 大金山岛3种植被类型土壤细菌优势门相对多度差异的显著性检验结果

Table 4 Results of significance test for the difference in relative abundance of dominant soil bacterial phylum among three vegetation types in Dajinshan Island, Shanghai

门 Phylum	落叶灌丛-落叶阔叶林 Deciduous shrubland - deciduous broadleaf forest		落叶阔叶林-常绿阔叶林 Deciduous broadleaf forest - evergreen broadleaf forest		落叶灌丛-常绿阔叶林 Deciduous shrubland - evergreen broadleaf forest
门 Phylum	t	p	t	p	t	p
变形菌门 Proteobacteria	-0.65	0.79	0.63	0.81	-0.09	0.99
酸杆菌门 Acidobacteria	-0.50	0.87	-0.81	0.70	-1.15	0.49
放线菌门 Actinobacteria	2.71	0.03	-1.01	0.58	1.73	0.21
WPS-2	-0.89	0.65	-5.04	0.00	-5.06	0.00
绿弯菌门 Chloroflexi	-0.76	0.73	-1.29	0.41	-1.80	0.19
浮霉菌门 Planctomycetes	2.31	0.07	-0.03	0.99	-2.17	0.10
厚壁菌门 Firmicutes	-1.84	0.18	1.03	0.56	-0.89	0.65
AD3	-1.35	0.38	-0.30	0.95	-1.53	0.29
疣微菌门 Verrucomicrobia	3.45	0.01	0.69	0.77	3.85	0.00

图5 大金山岛3种植被类型土壤细菌相对丰度显著性标志类群。由内至外辐射的圆圈代表由门至目的分类级别, 不同分类级别上的每一个小圆圈代表该水平下的一个分类, 无显著差异的物种统一着色为黄色, 差异显著的物种跟随组别着色, 未能在图中显示(以字母表示)的标志物对应的物种名展示在右侧。DF, 落叶阔叶林; DS, 落叶灌丛; EF, 常绿阔叶林。

Fig. 5 The most markable abundant taxa (biomarkers) of soil bacterial among three vegetation types on Dajinshan Island, Shanghai. The circle from inside to outside represents the taxonomy level from phylum to order, and each small circle at a given classification level represents a subordinate class within that category. The species with none of significant difference are colored by yellow, the species with significant difference are accompanied by the group color, and the species that cannot show in circle (represented by letters) and are corresponding to the markers display on the right side of the figure. DF, deciduous broadleaf forest; DS, deciduous shrubland; EF, evergreen broadleaf forest.

表5 大金山岛土壤细菌优势标志类群相对丰度与土壤养分含量的相关性

Table 5 Pearson correlation coefficients between relative abundance of main biomarker groups of soil bacterial and soil nutrients contents in Dajinshan Island, Shanghai

目的类别 Oder level	总氮含量 Total nitrogen content	总磷含量 Total phosphorus content	有机碳含量 Organic carbon content	速效氮含量 Available nitrogen content	速效磷含量 Available phosphorus content	速效钾含量 Available kalium content
肠杆菌目 Enterobacteriales	-0.17	-0.13	-0.09	-0.23	-0.14	0.12
酸杆菌目 Acidobacteriales	-0.27	-0.01	-0.10	-0.26	0.15	-0.22
伯克氏菌目 Burkholderiales	-0.04	-0.13	-0.09	-0.25	-0.14	0.13
黄单胞菌目 Xanthomonadales	0.24	0.45^*	0.33	0.24	0.39	-0.27
嗜热芽菌目 Thermogemmatisporales	-0.04	0.30	0.08	-0.00	0.37	-0.25
红螺菌目 Rhodospirillales	0.15	0.48^*	0.26	0.27	0.60	-0.39
根瘤菌目 Rhizobiales	0.19	0.28	0.18	0.42^*	0.24	-0.40
土壤红杆菌目 Solirubrobacterales	0.33	0.36	0.31	0.38	0.18	-0.33
Gaiellales	0.20	-0.03	0.07	0.31	-0.17	-0.13

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