孙建平,
刘雅辉^,,
左永梅,
韩民利,
张虹伟,
吕晶晶
河北省农林科学院滨海农业研究所/河北省盐碱地绿化工程技术中心/唐山市耐盐植物重点实验室 唐山 063299
基金项目: 河北省农林科学院基本业务费201810101
河北省农林科学院现代农业科技创新工程项目2019-1-6-2

详细信息

作者简介:孙建平, 主要研究方向为滨海盐碱地改良与农业高效利用。E-mail:bhssjp@163.com

通讯作者:刘雅辉, 主要研究方向为盐碱地改良与农业高效利用。E-mail:bhslyh@126.com

中图分类号:S154.36

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出版历程

收稿日期:2020-03-04
录用日期:2020-05-29
刊出日期:2020-10-01

The bacterial community structure and function of Suaeda salsa rhizosphere soil

SUN Jianping,
LIU Yahui^,,
ZUO Yongmei,
HAN Minli,
ZHANG Hongwei,
LYU Jingjing
Institute of Coastal Agricultural, Hebei Academy of Agriculture and Forestry Sciences/Saline and Alkali Land Greening Engineering Technology Center of Heibei Province/Tangshan Key Laboratory of Plant Salt Tolerance Research, Tangshan 063299, China
Funds: This study was supported by the Basic Business Expenses of Hebei Academy of Agricultural and Forestry Sciences201810101
The Agriculture Science and Technology Innovation Project of Hebei Academy of Agricultural and Forestry Sciences2019-1-6-2

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Corresponding author:LIU Yahui, E-mail:bhslyh@126.com

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摘要
摘要:盐地碱蓬作为生物改良盐碱地的理想材料，其根际土壤微生物对土壤改良发挥着重要作用。为了深入探索环渤海滨海盐碱地碱蓬根际土壤细菌群落结构组成及其功能，采用Illumina Misep高通量测序平台对环渤海地区滨海盐碱地盐地碱蓬根际土壤和裸地土壤进行测序。从16个样本中获得有效序列734 792条，4 285个OTUs，归属于41门、100纲、282目、400科、892属、1 577种。盐地碱蓬根际土壤细菌群落由变形菌门（Proteobacteria）、放线菌门（Actinobacteria）、绿弯曲门（Chloroflexi）、拟杆菌门（Bacteroidetes）、芽单胞菌门（Gemmatimonadetes）、酸杆菌门（Acidobacteria）、厚壁菌门（Firmicutes）、蓝藻细菌门（Cyanobacteria）、髌骨细菌门（Patescibacteria、浮霉菌门（Planctomycetes）组成。Alpha多样性计算结果表明，盐地碱蓬根际土壤细菌群落结构多样性高并与裸地土壤间差异显著；LEfSe（LDA Effect Size）分析发现，盐地碱蓬与裸地差异指示种明显不同。PCoA与相关性Heatmap表明，盐地碱蓬、速效氮、速效钾、速效磷、电导率是影响土壤细菌目类水平群落组成的主要因子。PICRUSt（Phylogenetic Investigation of Communities by Reconstruction of Unobserved States）分析表明微生物群落在新陈代谢等40个功能方面盐地碱蓬根际土壤比裸地土壤高。本研究表明盐地碱蓬覆盖能够降低土壤盐分，增加土壤养分，对土壤细菌群落多样性及其功能有积极作用。
Abstract:Suaeda salsa is an ideal agent for the biological enrichment of saline-alkali soil. Microorganisms in the rhizosphere of this plant play an essential role in soil improvement. The Illumina Misep high-throughput sequencing platform was used to explore the structural composition and function of the bacterial community in the rhizosphere soil of S. salsa and bare soil from coastal saline-alkali land in the Bohai Bay Rim area of Hebei, Shandong, and Tianjin, China. In total, 734 792 effective sequences were obtained from 16 samples, of which 4 285 OUTs belonged to 41 phyla, 100 classes, 282 orders, 400 families, 892 genera, and 1 577 species. The bacterial community in the rhizosphere soil of S. salsa contained Proteobacteria, Actinobacteria, Chloroflexi, Bacteroidetes, Gemmatimonadetes, Acidobacteria, Firmicutes, Cyanobacteria, Patescibacteria, and Planctomycetes. These results were consistent with the Alpha diversity analysis results, indicating that the community was highly diversified and significantly different from that of the bare soil. The LEfSe (LDA Effect Size) analysis showed that indicator species differentially occurred in S. salsa and bare soils. In S. salsa soil, Cyanobacteria, Acidobacteria, Alphaproteobacteria, Oxyphotobacteria, Chloroflexi, Rhizobiales, Nostocales, Sphingomonadales, Sphingomonadaceae, and Bacillus were the indicator species. Based on principal coordinates analyses and a correlation heatmap, the main factors affecting the soil bacterial community at order level were the presence of S. salsa, alkali-hydrolyzable nitrogen, available potassium, available phosphorus, and electrical conductivity. Also, Ectothiorhodospira and Balneolaceae could survive in bare soil with poor fertility, high salinity, and a viscous structure. PICRUSt (Phylogenetic Investigation of Communities by Reconstruction of Unobserved States) analysis showed that 304 metabolic pathways at pathway level 3 were active in both soil, of which 41 pathways, especially those involving in metabolism were different between S. salsa soil and bare land soil. These results indicated that S. salsa growth has a positive effect on the diversity and function of soil bacterial community by improving soil structure and increasing nutrients levels. These findings may be applied to improve saline-alkali land, optimize soil environment, and enhance its usefulness and sustainability.

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图1滨海盐地碱蓬根际土壤及裸地土壤细菌的稀释曲线
Figure1.Rarefaction curves of bacteria in bare soil and rhizosphere soil of Suaeda salsa in the coastal saline-alkali land


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图2滨海盐碱地盐地碱蓬根际土壤与裸地土壤细菌群落组成
Figure2.Composition of the bacterial community in rhizosphere soil of Suaeda salsa and bare soil in the coastal saline-alkali land


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图3滨海盐碱地盐地碱蓬根际土壤(J)与裸地土壤(D)优势物种组成比例及其在样本中的分布比例
左半圈表示样本中物种组成情况, 外层彩带颜色代表来自哪一分组, 内层彩带颜色代表物种, 长度代表该物种在对应样本中的相对丰度; 右半圈表示该分类学水平下物种在不同样本中的分布比例情况, 外层彩带颜色代表物种, 内层彩带颜色代表不同分组, 长度代表该样本在某一物种中的分布比例。
Figure3.Composition proportion of dominant species in rhizosphere soil of Suaeda salsa (J) and bare soil (D) and their distribution proportions in all samples of the coastal saline-alkali land
The left half circle shows the species composition in the sample, the color of the outer ribbon represents the group from which it comes, the color of the inner ribbon represents the species, and the length represents the relative abundance of the species in the corresponding sample. The right half circle shows the distribution proportion of the species in different samples under the taxonomic level, the outer ribbon represents the species, and the color of the inner ribbon represents different groups, the length represents the distribution proportion of the sample in a certain species.


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图4滨海盐碱地盐地碱蓬根际土壤与裸地土壤组间比较分析
D:裸地土壤; J:盐地碱蓬根际土壤。
Figure4.Group comparison analyses between rhizosphere soil of Suaeda salsa and bare soil in the coastal saline-alkali land
D: bare land soil; J: rhizosphere soil of Suaeda salsa.


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图5滨海盐碱地盐地碱蓬根际土壤与裸地土壤中的特殊群落及影响力
D:裸地土壤; J:盐地碱蓬根际土壤。不同颜色节点表示在对应组别中显著富集, 且对组间差异存在显著影响的微生物类群; 淡黄色节点表示在不同分组中均无显著差异, 或对组间差异无显著影响的微生物类群; 小写字母代表差异指示种, 其中p代表门, c代表纲, o代表目, f代表科, g代表属。
Figure5.Special communities and their effects in rhizosphere soil of Suaeda salsa and bare soil in in the coastal saline-alkali land
D: bare land soil; J: rhizosphere soil of Suaeda salsa. The nodes in different colors represent the microbial groups which are significantly enriched in the corresponding groups and have a substantial impact on between-group variance. The yellowish nodes denote the microbial groups which are not significantly different between groups and or have no significant effect on between-group variance. The lowercase letters represent difference indicator species, of which "p" means phylum, "c" means class, "o" means order, "f" means family and "g" means genus.


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图6滨海盐碱地盐地碱蓬根际土壤(J)与裸地土壤(D)的系统发生进化关系
Figure6.Evolutionary relationship in rhizosphere soil of Suaeda salsa (J) and bare soil (D) in the coastal saline-alkali land


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图7滨海盐碱地土壤细菌目类水平群落与环境因子的相关分析
x轴和y轴分别为环境因子和物种, 通过计算获得相关性R值和P值。R值在图中以不同颜色展示, 右侧图例是不同R值的颜色区间; P值则用*标出, * 0.01 < P≤0.05, ** 0.001 < P≤0.01, *** P≤0.001。
Figure7.Analysis of correlation between bacterial community at the order level and environmental factors in the coastal saline-alkali land
The x-axis and y-axis are environmental factors and species respectively. The correlation R value and P value are obtained by calculation. R values are shown in different colors, and the legend on the right is the color range of different R values. P values are marked with * for 0.01 < P≤0.05, ** for 0.001 < P≤0.01, *** for P≤0.001.


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图8滨海盐碱地盐地碱蓬根际土壤(J)与裸地土壤(D)的PICRUSt功能预测
Figure8.Prediction of PICRUst function in rhizosphere soil of Suaeda salsa (J) and bare soil (D) in the coastal saline-alkali land


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表1样地基本概况
Table1.Basic information of the soil samples

样地位置 Sample site location	采样点 Sample plot	地理位置 Geographical position	海拔 Altitude (m)	主要植物 Main plant
天津北大港 Tianjin Beidagang	D1, J4, J5, J6	38°44′25.24″N, 117°14′13.36″E	2	盐地碱蓬、芦苇、金叶榆树 Suaeda salsa, Phragmites communis, Ulmus pumila
沧州黄骅市南排河 Nanpaihe Town, Huanghua City, Cangzhou City, Hebei Province	D2, J7, J8, J9	38°25′42.89″N, 117°35′14.58″E	2	盐地碱蓬、芦苇、知风草 S. salsa, P. communis, Eragrostis ferruginea
山东东营市东营区 Dongying District, Dongying City, Shandong Province	D2, J10, J11	37°24′12.31″N, 118°41′10.17″E	0	盐地碱蓬、柽柳、白刺 S. salsa, Tamarix chinensis, Nitraria schoberi
河北唐山市曹妃甸区 Caofeidian District, Tangshan City, Hebei Province	D4, D5, J1, J2, J3	39°12′48.44″N, 118°27′46.43″E	0	盐地碱蓬、芦苇、柽柳 S. salsa, P. communis, T. chinensis
D:裸地土壤; J:碱蓬根际土壤。D: bare land soil; J: rhizosphere soil of Suaeda salsa.

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表2滨海盐碱地盐地碱蓬根际土壤及裸地土壤的化学性质
Table2.Chemical properties of rhizosphere soil of Suaeda salsa and bare soil in the coastal saline-alkali land

样本 Sample	pH	电导率 Electrical conductivity (dS·m-1)	碱解氮 Alkali-hydrolyzable N (mg·kg-1)	速效磷 Available P (mg·kg-1)	速效钾 Available K (mg·kg-1)	有机质 Organic matter (g·kg-1)
裸地土壤Bare land soil	7.46±0.18a	6.47±5.10a	26.80±26.15b	14.91±2.41b	292.90±30.38b	9.45±1.63b
根际土壤Rhizosphere soil	7.48±0.22a	1.30±1.54b	90.71±48.58a	20.45±3.31a	586.14±110.63a	16.00±9.27a
同列不同小写字母表示差异显著(P≤0.05)。Different lowercase letters in the same column indicate significant differences at P≤0.05 level.

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表3滨海盐碱地盐地碱蓬根际土壤及裸地土壤细菌的生物多样性指数
Table3.Biodiversity indexes of bacteria in rhizosphere soil of Suaeda salsa and bare soil in the coastal saline-alkali land

多样性指数 Diversity index	裸地土壤 Bare land soil	根际土壤 Rhizosphere soil
Sobs指数Sobs index	660.80±169.95	705.36±110.52
香农指数Shannon index	4.75±0.38	5.05±0.40
辛普森指数Simpson index	0.024±0.009	0.019±0.014*
Ace指数Ace index	802.58±173.58	800.87±129.08
Chao指数Chao index	838.30±179.28	817.32±129.10
覆盖度指数Coverage index	0.994±0.001	0.995±0.001
*表示裸地土壤和盐地碱蓬根际土壤在P≤0.05水平差异显著。* means significant differences between bare land soil and rhizosphere soil of S. glauca at P≤0.05.

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