黄山土壤细菌群落及氮循环功能群的垂向分布格局

doi:10.16258/j.cnki.1674-5906.2023.01.007

生态环境学报 ›› 2023, Vol. 32 ›› Issue (1): 56-69.DOI: 10.16258/j.cnki.1674-5906.2023.01.007

黄山土壤细菌群落及氮循环功能群的垂向分布格局

向兴¹^,²(), 满百膺¹^,²^,^*(), 张俊忠³, 罗洋¹, 毛小涛¹, 张超¹, 孙丙华², 王希²

1.上饶师范学院生命科学学院，江西上饶 334001
2.安徽省黄山生物多样性与短尾猴行为生态学国际联合研究中心，安徽合肥 230039
3.西南林业大学生物多样性保护学院，云南昆明 6502224

收稿日期:2022-06-09 出版日期:2023-01-18 发布日期:2023-04-06
通讯作者: *满百膺（1979年生），男，副教授，博士，研究方向为环境微生物。E-mail: shmilying@126.com
作者简介:向兴（1989年生），男（苗族），讲师，博士，研究方向为微生物生态学。E-mail: xiangxing1989@126.com
基金资助:
国家自然科学基金项目(31960026);安徽省黄山生物多样性与短尾猴行为生态学国际联合研究中心开放课题(KF200013)

Vertical Distribution of Bacterial Community and Functional Groups Mediating Nitrogen Cycling in Mount Huangshan, Anhui, China

XIANG Xing¹^,²(), MAN Baiying¹^,²^,^*(), ZHANG Junzhong³, LUO Yang¹, MAO Xiaotao¹, ZHANG Chao¹, SUN Binghua², WANG Xi²

1. College of Life Science, Shangrao Normal University, Shangrao, 334001, P. R. China
2. International Research Center for Huangshan Biodiversity and Tibetan Macaque Behavioral Ecology, Hefei, 230039, P. R. China
3. College of Biodiversity Conservation, Southwest Forestry University, Kunming, 650224, P. R. China

Received:2022-06-09 Online:2023-01-18 Published:2023-04-06

摘要/Abstract

摘要：

山地生态系统是研究土壤微生物垂向变化规律的天然场所，受采样空间尺度、样本异质性、测序深度、变化的生态因子、环境参数及多样性度量指数等因素的制约，目前对该系统土壤细菌群落及氮循环功能群垂向分异格局的认识仍然薄弱。以植被和土壤垂直分带明显的黄山为研究对象，选取海拔926—1800 m的典型森林土壤样本，采用Illuminia HiSeq高通量测序技术获取土壤细菌群落信息，在生物信息学分析中引入谱系结构指数（MNTD和NTI），结合土壤理化参数及FAPROTAX功能预测手段，探究黄山森林土壤细菌群落多样性和氮循环功能群垂向变化的潜在规律。结果显示，（1）变形菌门（Proteobacteria，44.77%）、酸杆菌门（Acidobacteria，21.85%）和放线菌门（Actinobacteria，14.88%）为优势细菌门类，其垂向分布依次呈现递减式、“U”型式和单峰式。（2）细菌群落的物种多样性指数和谱系多样性指数的空间垂向分布呈单峰模式。（3）谱系结构指数分析揭示细菌群落的系统发育关联度随海拔升高呈单调递增模式，表明环境过滤作用显著影响细菌群落的形成过程。（4）系统发育方法和分类学方法均表明pH和温度（t）是黄山森林土壤细菌群落结构垂向分布的主控因素。（5）4个主要的氮循环功能群中，尿素降解功能群（8.14%）、好氧氨氧化功能群（2.73%）和好氧亚硝酸氧化功能群（0.99%）的相对丰度在垂向分布上呈单峰模式，其优势类群依次为中慢生根瘤菌属（Mesorhizobium）、Ellin6067类群和硝化螺菌属（Nitrospira）；固氮功能群（21.45%）在垂向呈“U”型分布模式，其优势类群是慢生根瘤菌属（Bradyrhizobium）。研究结果揭示黄山森林土壤细菌群落多样性、物种组成及氮循环功能群的垂向分异格局，为认识中亚热带森林生态系统土壤微生物群落结构及功能群对环境变化的响应提供理论依据。

关键词: 黄山, 细菌群落, 谱系结构指数, 土壤pH, 氮循环功能群, 垂向分布格局

Abstract:

Mountain ecosystems are natural laboratories to study the vertical changes in soil microorganisms. However, due to the impact of sampling space scale, heterogeneity of samples, sequencing depth, change of ecological factors, environmental parameters and the selection of diversity index, the vertical patterns of soil bacterial communities and nitrogen cycling functional groups in this system are still poorly understood. Thus, we collected the forest soil samples from 926 to 1800 m in Mount Huangshan, with distinct vertical zones of vegetation and soil types, and applied high-throughput sequencing of bacterial 16S ribosomal gene, together with the phylogenetic structure analysis (MNTD and NTI) of bioinformatics, physicochemical parameters and FAPROTAX functional prediction to analyze the vertical distribution patterns of bacterial communities and nitrogen-cycling functional groups. Results showed that (1) Proteobacteria (44.77%), Acidobacteria (21.85%) and Actinobacteria (14.85%) dominated the bacterial communities at phylum level and showed linear, U-shaped and unimodal decreasing with the altitude increase. (2) Species diversity index and phylogenetic diversity index of bacterial communities displayed a unimodal model with the altitude increase. (3) Community phylogenetic relatedness exhibited monotonic increase trends along elevational gradient based on community phylogenetic analysis, suggesting that bacterial community assembly processes were governed by environmental filtering. (4) Both phylogenetic and taxon-based approaches showed that pH and soil temperature (t) were the most important driving forces for the bacterial community vertical distribution. (5) Among the four main nitrogen cycle functional groups, ureolysis group (8.14%), aerobic ammonia oxidation group (2.73%) and aerobic nitrite oxidation group (0.99%) presented unimodal model with the altitude increase, and their dominant taxa were Mesorhizobium, Ellin6067, and Nitrospira respectively. While the nitrogen-fixing group (21.45%) showed a U-shaped pattern, which dominated by Bradyrhizobium. The results reveal a vertical distribution pattern of soil bacteria and nitrogen cycling functional groups in Mount Huangshan forest ecosystem, which provide insight into the response of soil microbial communities and functional groups to environmental change in subtropical forest ecosystems.

Key words: Mount Huangshan, bacterial communities, phylogenetic structure, soil pH, nitrogen-cycling functional groups, vertical distribution pattern

中图分类号:

S154.3
X172

向兴, 满百膺, 张俊忠, 罗洋, 毛小涛, 张超, 孙丙华, 王希. 黄山土壤细菌群落及氮循环功能群的垂向分布格局[J]. 生态环境学报, 2023, 32(1): 56-69.

XIANG Xing, MAN Baiying, ZHANG Junzhong, LUO Yang, MAO Xiaotao, ZHANG Chao, SUN Binghua, WANG Xi. Vertical Distribution of Bacterial Community and Functional Groups Mediating Nitrogen Cycling in Mount Huangshan, Anhui, China[J]. Ecology and Environment, 2023, 32(1): 56-69.

图/表 7

表1 黄山森林土壤理化特征

Table 1 Physicochemical properties of forest soil samples in Mount Huangshan, east China

海拔分组	t/℃	pH	w_(AN)/(mg·kg^-1)	w_(AP)/(mg·kg^-1)	w_(AK)/(mg·kg^-1)	w_(TN)/(g·kg^-1)	w_(TP)/(g·kg^-1)	w_(TK)/(g·kg^-1)	w_(SOM)/(g·kg^-1)
低海拔 (L, n=3)	17.80±0.10^a	4.25±0.28	844.67±281.76^a	8.83±3.37^ab	162.33±32.15^a	8.94±1.79^a	0.67±0.06^a	25.53±3.33	233.67±70.89^a
中低海拔 (ML, n=6)	15.72±0.78^b	4.89±0.63	444.67±150.14^b	2.10±0.91^b	96.67±18.98^b	5.07±1.05^b	0.45±0.08^b	30.75±5.15	124.67±12.06^b
中海拔 (M, n=6)	15.25±0.43^b	5.18±0.80	539.33±164.18^b	12.42±8.07^a	100.83±40.47^b	5.82±1.27^b	0.69±0.17^a	29.27±3.42	148.40±49.35^b
高海拔 (H, n=9)	12.22±0.63^c	4.38±0.42	624.11±156.81^ab	2.70±2.25^b	95.89±37.72^b	5.37±1.466^b	0.50±0.10^b	27.98±1.69	123.62±24.41^b

图1 黄山森林土壤细菌群落α多样性指数的垂向变化趋势扩增序列变体（a：ASV）、物种多样性指数（b：Shannon；c：ACE）和谱系多样性指数（d：PD）采用二次多项式回归模型；谱系结构指数（e：MNTD；f：NTI）采用对数回归模型；R2：拟合度；P：显著性

Figure 1 Vertical variation of α diversity indices of forest soil bacterial communities in Mount Huangshan, east China

图2 黄山森林土壤细菌群落多样性分析及环境影响因子对其变异度的解释图b中彩色部表示显著影响土壤细菌群落变异的环境因子及其解释度；VE：植被，AL：海拔，Unknown：不能被解释的群落变异

Figure 2 PCoA analysis of diversity for forest soil bacterial communities and variability of diversity explained by environment factors in Mount Huangshan, east China

图3 黄山森林土壤细菌群落门水平组成

Figure 3 Composition of soil bacterial communities at phylum level in Mount Huangshan, east China

图4 黄山不同海拔森林土壤细菌优势属的相对丰度 BCP：伯克霍尔德氏菌属

Figure 4 Relative abundance of dominant genus for soil bacteria along different altitude groups in Mount Huangshan, east China

图5 黄山森林土壤细菌群落与环境因子之间的冗余分析显著性水平：*：0.05；**：0.01

Figure 5 RDA analysis between bacterial communities at phylum level and environmental factors in Mount Huangshan, east China

图6 黄山森林土壤细菌氮循环功能群的分类学组成（a）及垂向变化规律（b—e）图b、c、d和e的回归模型：二次多项式回归模型；r2：拟合度；P：显著性

Figure 6 Taxonomic information (a) and vertical distribution patterns (b-e) of nitrogen cycling functional groups in Mount Huangshan

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黄山土壤细菌群落及氮循环功能群的垂向分布格局

Vertical Distribution of Bacterial Community and Functional Groups Mediating Nitrogen Cycling in Mount Huangshan, Anhui, China

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