渤海湾表层海水中浮游细菌群落随离岸距离的分布特征及其影响因素

赵维; 王敬敬; 徐松; 李晴晴; 杨榕; 张小霞; 黄志勇

doi:10.3969/j.issn.0253-4193.2019.12.015

渤海湾表层海水中浮游细菌群落随离岸距离的分布特征及其影响因素

doi: 10.3969/j.issn.0253-4193.2019.12.015

赵维^1,,
王敬敬¹,
徐松¹,
李晴晴¹,
杨榕¹,
张小霞¹,
黄志勇^1, ,

中国科学院天津工业生物技术研究所天津市工业生物系统与过程工程重点实验室，天津 300308

基金项目: 天津市科技计划项目（18ZXSZSF00100,18YFZCNC01180,16YFXTSY00580）；国家自然科学基金项目（3150020230）；中国科学院战略生物资源计划（KFJ-BRP-009）。

详细信息

作者简介:
赵维（1988—），女，河北省沧州市人，助理研究员，主要从事海洋环境微生物多样性研究。E-mail：zhao_w@tib.cas.cn

通讯作者:
黄志勇，研究员，主要从事环境微生物生态学研究。E-mail：huang_zy@tib.cas.cn

中图分类号: P722.4; Q938
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出版历程
- 收稿日期: 2018-12-12
- 修回日期: 2019-06-21
- 网络出版日期: 2021-04-21
- 刊出日期: 2019-12-25

Distribution characteristics and influencing factors of bacterioplankton community with offshore distance variation in the surface seawater of Bohai Bay

Tianjin Key Laboratory for Industrial Biological Systems and Bioprocessing Engineering, Tianjin Institute of Industrial Biotechnology, Chinese Academy of Sciences, Tianjin 300308, China

摘要

摘要: 为研究渤海湾近岸污染对远近岸海域微生态的影响，利用高通量测序技术，对渤海湾不同离岸距离的6个站位采集表层海水，进行浮游细菌群落结构分析，结合环境、空间因素探究影响其变化的主要因素。结果表明：研究区域存在环境因子的梯度变化，如氮营养盐在近岸高于远岸；细菌α-多样性在不同站位间差异不显著，但仍显示在近岸相对较高；细菌群落结构随离岸距离变化显著，γ-变形菌和拟杆菌在近岸显著富集，且与氮营养盐的含量有关；蓝细菌在远岸显著富集，且与氨氮、透明度、电导率有关；邻体矩阵主坐标单独解释部分对群落结构变异的贡献率最大(38.1%)，说明可能存在尚未测量但具有空间结构的环境变量影响群落空间分布；结合功能预测的结果推测近岸区域的富营养与烃类污染等可能影响群落变化。本文从环境和空间影响两方面探讨了渤海湾不同离岸距离的海域浮游细菌群落结构变化，为研究渤海湾海洋生态及环境保护提供一定的参考。
- 渤海湾 /
- 浮游细菌群落结构 /
- 地理分布 /
- 功能预测 /
- 邻体矩阵主坐标
Abstract: In order to study the impact of coastal pollution in Bohai Bay on the microecology of sea areas with different distances from the shore, the bacterioplankton community compositions (BCCs) in surface seawater samples from 6 stations with different offshore distances along the coastal region of Bohai Bay were analyzed through high throughput sequencing technology, and the main factors affecting the variation of BCCs were explored by combining environmental and spatial factors in this region. The results showed that there was gradient change of environmental factor in the studied region, such as the contents of nitrogen nutrients were higher in the nearshore station than those in the offshore station. Although there was no significant difference tested for the alpha diversity among different sites, the diversity indexes were still relatively higher in the nearshore stations. The bacterioplankton community compositions were significantly varied with the change of offshore distances. Members of Gammaproteobacteria and Bacteroidetes were mainly enriched in nearshore stations which were closely related with the contents of nitrogen nutrients, members of Cyanobacteria were mainly enriched in offshore stations, which were closely related to ammonia nitrogen, transparency and conductivity. The variance partitioning analysis showed that PCNM variables purely contributed most (38.1%) to the variation of community structure, indicating that there may be environmental variables with spatial structure within the research scope that had not yet been measured may affect the spatial distribution of the bacterioplankton community. Meanwhile, the results of functional prediction indicated that the eutrophication, hydrocarbon pollution and other environmental conditions in the nearshore station may contribute to the change of BCCs. This study explored the variation of offshore-distance-varied BCC in the coastal region of Bohai Bay from environmental and spatial impact, which may provide reference for the study and protection of marine environment in Bohai Bay.
- Bohai Bay /
- bacterioplankton community composition /
- geographic distribution /
- function prediction /
- PCNM

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图 1 渤海湾取样点

Fig. 1 Sampling sites in the Bohai Bay

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图 2 基于营养盐类含量变化的环境因子聚类分析与各站位群落组成柱状图

Fig. 2 Clustering analysis of environmental factors based on the nutrient contents and histogram of bacterial community composition at each site

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图 3 基于Jaccard距离（a）和Bray-Curtis距离（b）的OTU水平的群落聚类分析

Fig. 3 Principal coordinates analysis (PCoA) plot derived from the Jaccard (a) and Bray-Curtis (b) distance among seawater samples based on the occurrences and abundances of Operational Taxonomic Units (OTUs)

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图 4 基于LDA分析的3组间差异微生物展示图

具有显著差异的分类单元节点被着以不同颜色，该分类单元的最高等级对应的分支区域也着以相同颜色。如果分类单元在组间没有显著差异，则相应节点为黄色

Fig. 4 Least discriminant analysis (LDA) effect size taxonomic cladogram comparing all samples categorized by three bacterial groups

Significantly discriminant taxon nodes are colored and branch areas are shaded according to the highest ranked group for that taxon. If the taxon is not significantly differentially represented among sample groups, the corresponding node is colored yellow

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图 5 差异物种和环境因子的相关性分析

热图代表各差异物种和环境因子的Pearson相关性系数，星号代表显著性：* 0.05，** 0.001

Fig. 5 Heat map demonstrate correlations between highly abundant discriminant taxa of three groups and seawater environmental parameters

Value of the heat map bar represents the Pearson correlation coefficient (r ), and the asterisk represents the significance of the relationship with * 0.05 and ** 0.001

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图 6 群落结构变异因素分析

基于Hellinger转化的OTU数据，通过海水环境因子（左上）、地理因素包括线性趋势（右上）和邻体矩阵主坐标（下）来对群落结构进行变异分解分析。每一个解释部分都经过前向选择选出最适模型的环境因子子集、线性趋势变量子集和邻体矩阵主坐标子集（n=6）

Fig. 6 Variation partitioning of bacterial communities

Hellinger transformed OTU matrix was used as responsible variable by the water environmental factors (Env., top left and the spatial factors including linear trend (Trend, top right) and PCNM variables (bottom). Forward selection procedures were used to select the best subset of environmental, trend and PCNM variables explaining community variation respectively (n = 6)

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图 7 基于FAPROTAX功能预测的结果进行的主坐标分析

Fig. 7 PCoA plot based on function prediction through FAPROTAX

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表 1 取样站位的环境特征与海水理化性质

Tab. 1 Environment and seawater properties of sampling sites

变量	TJ13	TJ14	TJ16	TJ17	TJ23	TJ27
经度	118.093 6°E	118.884 1°E	117.766 9°E	118.488 6°E	117.742 2°E	117.727 3°E
纬度	38.673 9°N	38.659 1°N	38.670 1°N	38.634 8°N	38.667°N	38.671 4°N
离岸距离/km	44.0	113.1	16.0	79.0	14.0	12.0
风向	NE	NE	NW	NE	NW	NW
风速/m·s⁻¹	1.0	3.0	2.7	4.0	2.0	0.8
水深/m	12.8	22.0	5.0	8.5	4.0	3.9
气压/kPa	1 012	1 012	1 012	1 011	1 012	1 012
气温/℃	26.4	24	28.2	23.4	28.3	30.4
水温/℃	26.9	24.0	27.0	23.5	27.0	28.4
湿度/%	63.6	76.9	62.9	68.4	30.4	56.3
透明度/cm	100	300	200	200	200	170
盐度	23.5	29.5	29.6	29.5	29.5	29.4
溶解氧含量/mg·L^–1	7.51	6.70	6.51	6.88	5.79	6.85
pH	8.18	8.10	8.12	8.06	8.12	8.15
电导率/mS·cm^–1	45.1	45.6	45.4	45.6	45.3	45.2
化学需氧量/mg·L ^–1	0.64	1.13	1.05	0.56	1.35	1.60
亚硝酸盐/mg·L ^–1	0.030	0.006	0.134	0.016	0.108	0.106
硝酸盐/mg·L ^–1	0.114	0.127	0.318	0.103	0.402	0.356
氨氮/mg·L ^–1	0.011	0.133	0.009	0.043	0.019	0.064
无机氮/mg·L ^–1	0.155	0.266	0.461	0.162	0.529	0.526
活性硅酸盐/mg·L ^–1	0.768	0.890	0.712	0.712	0.812	1.010
活性磷酸盐/mg·L ^–1	0.004	0.003	0.007	0.004	0.016	0.005

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表 2 海水样品的微生物多样性指数分布

Tab. 2 Analysis of microbial diversity index of the six sea water samples

样品号	OTU 丰度	谱系多样性值	Shannon值
TJ13	628.80 ± 13.30	63.19 ± 2.23	4.83 ± 0.11
TJ14	603.05 ± 27.05	64.57 ± 1.49	4.92 ± 0.05
TJ16	718.30 ± 60.28	69.92 ± 3.73	5.09 ± 0.02
TJ17	827.85 ± 92.55	72.22 ± 2.34	5.43 ± 0.43
TJ23	836.73 ± 125.36	79.48 ± 8.22	5.78 ± 0.28
TJ27	639.23 ± 46.51	63.89 ± 4.82	5.06 ± 0.09
注：表中数据为平均值±标准差; 加粗字体表示相应数值在近岸站位相对高于远岸站位。

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表 3 群落功能在不同组别站位间的相对丰度差异分析

Tab. 3 Difference analysis of relative functional group abundance between different groups of stations

功能组	第1组/%	第2组/%	第3组/%
甲烷营养	0.01±0.02	0.01±0.01	0±0
甲醇氧化	0.55±0.13	0.61±0.06	0.51±0.18
甲基营养	0.57±0.12	0.62±0.06	0.51±0.18
好氧氨氧化	0±0	0.01±0.01	0.01±0.02
有氧亚硝酸盐氧化	0.04±0.02^a	0.09±0.03^b	0.09±0.04^ab
硝化作用	0.05±0.02^a	0.1±0.02^b	0.1±0.06^ab
硫酸盐呼吸	0.01±0.01	0.01±0.01	0.01±0.01
硫化物呼吸	0.01±0.01	0.01±0.01	0.01±0.01
硝化反硝化作用	0.14±0.06	0.22±0.09	0.1±0
亚硝酸盐脱氮作用	0.14±0.06	0.22±0.09	0.1±0
一氧化二氮反硝化作用	0.14±0.06	0.22±0.09	0.1±0
反硝化作用	0.14±0.06	0.22±0.09	0.1±0
Chitinolysis	0.02±0.01	0.01±0.01	0±0
暗氢氧化反应	0.02±0.01^a	0±0^b	0±0^b
固氮作用	0.01±0.01	0.01±0.01	0±0
亚硝酸盐呼吸	0.14±0.06	0.22±0.09	0.1±0
溶纤作用	0.01±0.01	0.01±0.01	0±0
暗硫氧化反应	0.02±0.01	0.29±0.34	0.03±0.01
暗硫化合物氧化反应	0.16±0.16	0.35±0.41	0.03±0.01
锰氧化作用	0.01±0	0.01±0.01	0.01±0.01
发酵作用	2.97±2.23	0.91±0.56	1.46±0.19
有氧化能异养	29.71±4.4^a	17.26±3.3^ab	10.88±2.97^b
人类病原体	0.04±0.02	0.06±0.05	0.05±0.01
人类肠道	0±0	0±0	0.01±0.01
哺乳动物肠道	0±0	0±0	0.01±0.01
动物寄生虫或共生体	0.45±0.15	0.41±0.44	0.27±0.05
芳烃降解	0.76±0.35	0.6±0.65	0.14±0.08
芳香族化合物降解	0.84±0.36	0.68±0.62	0.22±0.07
烃降解	0.83±0.38	0.63±0.68	0.14±0.09
铁呼吸	0.03±0.02	0.04±0.05	0.03±0.01
硝酸盐呼吸	0.15±0.06	0.23±0.09	0.11±0
硝酸盐还原作用	2.92±2.16	1.52±0.18	1.34±0.34
氮呼吸	0.15±0.06	0.23±0.09	0.11±0
细胞内寄生虫	0.85±0.27	2.01±1.16	1.52±0.3
捕食和寄生	0.62±0.37^a	0.09±0.05^b	0.1±0^ab
蓝藻细菌	1.77±0.47^ab	0.3±0.29^b	29.29±10.27^a
非产氧光自养型硫氧化	0.14±0.06	0.22±0.08	0.11±0.01
非产氧光能自养	0.14±0.06	0.22±0.08	0.11±0.01
产氧光能自养	1.77±0.47^ab	0.3±0.29^b	29.29±10.27^a
光能自养	1.9±0.51^ab	0.53±0.34^b	29.4±10.26^a
光能异养	0.14±0.06	0.22±0.09	0.1±0
光营养	1.91±0.51^ab	0.53±0.35^b	29.4±10.26^a
尿素分解	0.1±0.06	0.14±0.11	0.26±0.17
化能异养	30.37±4.33^a	17.91±3.36^ab	11.43±3.2^b
注：表中数据为平均值 ± 标准差；同行不同小写字母表示该功能的相对丰度在不同组别之间依据Kruskal-Wallis检验，在0.05水平存在显著差异（第1组: n=6; 第2组: n= 4; 第3组: n=2）; 加粗字体表示相应数值在近岸站位相对高于远岸站位。

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