荒漠豆科灌丛根际土壤氨氧化和反硝化微生物功能基因丰度及群落多样性特征

doi:10.11686/cyxb2023239

摘要/Abstract

摘要：

为探索宁夏荒漠草原豆科灌丛根际土壤中参与氨氧化和反硝化的微生物功能基因丰度和群落组成对植物类型和生长时期的响应特征，本研究以宁夏白芨滩国家级自然保护区内典型的豆科灌丛群落柠条、沙冬青、猫头刺和毛刺不同生长时期（营养期、盛花期和果实期）的根际土壤为材料，通过功能基因实时荧光定量PCR和高通量测序方法，分析4种灌丛根际土壤中氨氧化和反硝化微生物功能基因丰度、群落结构组成特征及其与土壤因子的相关性。结果表明，功能基因丰度及群落结构因灌丛类型和生长时期不同而存在差异。柠条灌丛根际土壤中各功能基因丰度显著高于其他3种灌丛，且在盛花期达到最高。柠条各生长期根际土壤中氨氧化细菌（ammonia-oxidizing bacteria， AOB）群落多样性高于其他灌丛类型，而毛刺根际氨氧化古菌（ammonia-oxidizing archaea， AOA）和nirK型反硝化菌群落多样性最低。各灌丛根际AOA以未分类类群为绝对优势，占79.34%~98.37%，奇古菌门和泉古菌门分别仅占0.28%~20.37%和0.28%~2.35%；AOB在柠条根际以变形菌门为优势，其他3种植物根际则以未分类类群为优势；nirK和nirS型反硝化菌均以变形菌门为优势门，但nirK型反硝化菌的组成受植物类型和生长时期的影响更显著。功能基因的丰度与土壤全氮、速效钾和pH显著正相关。全钾、速效钾、有机质显著影响氨氧化微生物的群落组成，pH是反硝化微生物群落结构的主要影响因子。

关键词: 固沙灌丛, 氨氧化, 反硝化, 功能基因, 群落组成, 多样性

Abstract:

The aim of this study was to explore the patterns of functional gene abundance and community composition of microorganisms involved in ammonia oxidation and denitrification in rhizosphere soil of leguminous shrubs on the Ningxia desert steppe among different plant types and growth periods. We collected rhizosphere soil samples from typical leguminous shrub communities in the Baijitan National Nature Reserve of Ningxia， including Caragana korshinskii， Ammopiptanthus mongolicus， Oxytropis aciphylla， and Caragana tibetica at different growth stages （vegetative， flowering， and fruiting stages）. Using real-time quantitative PCR and high-throughput sequencing methods， the abundance of functional genes and the characteristics of community structure of ammonia-oxidizing and denitrifying microorganisms in rhizosphere soil of four shrubs， as well as their correlations with soil factors， were analyzed. It was found that the abundance of functional genes and community structure varied among different shrub types and growth periods. The abundance of functional genes was significantly higher in the rhizosphere soil of C. korshinskii than in the rhizosphere soil of the other shrubs， and reached the highest level during the flowering period. The diversity of the ammonia-oxidizing bacteria （AOB） community in the rhizosphere soil of C. korshinskii at different growth periods was higher than that of the AOB communities in the rhizosphere soils of the other shrub types， while the diversity of ammonia-oxidizing archaea （AOA） and nirK-type denitrifying bacteria communities was lowest in the rhizosphere soil of C. tibetica. Unclassified taxa were the predominant AOA in rhizosphere soil of each shrub with an extremely high abundance of 79.34%-98.37%， whereas Thaumarchaeota and Crenarchaeota only accounted for 0.28%-20.37% and 0.28%-2.35%， respectively. Proteobacteria was the dominant phylum of AOB in the rhizosphere of C. korshinskii， while Norank_d__Bacteria was the dominant AOB in the rhizosphere soil of the other plants. Both nirK- and nirS- type denitrifying bacteria were dominated by Proteobacteria， but the composition of nirK- type denitrifying bacteria was more significantly influenced by the plant type and growth period. The abundance of functional genes was significantly positively correlated with soil total nitrogen， available potassium， and pH. The soil total potassium， available potassium， and organic matter significantly affected the community composition of ammonia-oxidizing microorganisms， and pH was the main factor affecting the community structure of denitrifying microorganisms.

Key words: sand-fixation shrublands, ammonia oxidation, denitrification, functional genes, community composition, diversity

刘爽, 姚佳妮, 张钧杰, 代金霞. 荒漠豆科灌丛根际土壤氨氧化和反硝化微生物功能基因丰度及群落多样性特征[J]. 草业学报, 2024, 33(5): 115-127.

Shuang LIU, Jia-ni YAO, Jun-jie ZHANG, Jin-xia DAI. Functional gene abundance and community diversity of ammonia-oxidizing and denitrifying microorganisms in the rhizosphere soil of desert leguminous shrubs[J]. Acta Prataculturae Sinica, 2024, 33(5): 115-127.

图/表 8

表1 氨氧化和反硝化功能基因实时荧光定量PCR的引物

Table 1 Primers for real-time fluorescent quantitative PCR of functional genes related to ammonia oxidation and denitrification

功能基因Functional genes	引物Primers	引物序列Primer sequence （5′-3′）	长度Length （bp）	参考文献Reference
amoA	amoAF	STAATGGTCTGGCTTAGACG	600	［14］
amoA	amoAR	GCGGCCATCCATCTGTATGT	600	［14］
bamoA	bamoA1F	GGGGTTTCTACTGGTGGT	491	［14］
bamoA	bamoA2R	CCCCTCKGSAAAGCCTTCTTC	491	［14］
nirK	nirK1aCuF	ATCATGGTSCTGCCGCG	459	［15］
nirK	nirK3aCuR	GCCTCGATCAGRTTGTGGTT	459	［15］
nirS	nirS4F	TTCRTCAAGACSCAYCCGAA	332	［16］
nirS	nirS6R	CGTTGAACTTRCCGGT	332	［16］

图1 4种豆科灌丛根际土壤中氨氧化和反硝化功能基因丰度不同大写字母表示同一生长时期不同灌丛之间差异显著（P<0.05），不同小写字母表示同一灌丛不同生长时期之间差异显著（P<0.05）。Different capital letters represent significant differences among different shrubs at the same growth period （P<0.05）， while different lowercase letters represent significant differences among different growth periods of the same shrub （P<0.05）.

Fig.1 Abundance of functional genes related to ammonia oxidation and denitrification in rhizosphere soil of four leguminous shrubs

表2 4种豆科灌丛根际土壤氨氧化和反硝化微生物多样性指数

Table 2 Diversity index of ammonia-oxidizing and denitrifying microorganisms in rhizosphere soil of four leguminous shrubs

样品编号 Sample number	氨氧化古菌 Ammonia-oxidizing archaea		氨氧化细菌 Ammonia-oxidizing bacteria		nirK型反硝化菌 nirK-type denitrifying bacteria		nirS型反硝化菌 nirS-type denitrifying bacteria
样品编号 Sample number	Chao 1	Shannon	Chao 1	Shannon	Chao 1	Shannon	Chao 1	Shannon
SDQ₁	23.00	1.4690	22.00	1.7425	274.69	3.9031	105.60	2.0859
SDQ₂	30.00	1.8068	23.00	1.7253	169.50	3.4446	119.43	3.5405
SDQ₃	27.50	1.3555	19.00	1.8469	211.50	3.3924	92.33	2.7693
NT₁	23.00	1.2244	35.00	2.2478	291.13	3.3304	152.20	3.2573
NT₂	18.00	1.2407	30.00	2.3161	271.02	3.4343	155.24	3.1518
NT₃	24.00	1.3009	32.00	2.4139	298.03	4.1252	114.50	3.5996
MTC₁	28.00	1.9160	24.00	1.6897	204.45	3.3538	113.00	2.9958
MTC₂	24.50	1.5782	37.00	1.6630	220.30	3.5430	136.65	3.3694
MTC₃	24.33	1.5728	38.00	2.0907	205.12	3.1738	68.00	2.0021
MC₁	23.33	0.6176	10.00	1.1828	162.00	2.5022	66.00	3.0166
MC₂	20.00	0.7633	16.00	1.7892	177.96	3.1210	100.43	3.0036
MC₃	23.20	0.5309	25.50	1.7542	160.00	2.4473	82.50	3.0530

图2 4种豆科灌丛根际土壤氨氧化微生物在门和属水平的群落组成AOA：氨氧化古菌Ammonia-oxidizing archaea； AOB：氨氧化细菌Ammonia-oxidizing bacteria. SDQ：沙冬青A. mongolicus； NT：柠条C. korshinskii； MTC：猫头刺O. aciphylla； MC：毛刺C. tibetica. 1、2、3分别代表营养期、盛花期和果实期。1， 2， and 3 respectively represent the nutritional period， flowering period， and fruit period. 下同The same below.

Fig.2 Community composition of ammonia-oxidizing microorganisms in rhizosphere soil of four leguminous shrubs at the phylum and genus levels

图3 4种豆科灌丛根际土壤反硝化微生物在门和属水平的群落组成

Fig.3 Community composition of denitrifying microorganisms in rhizosphere soil of four leguminous shrubs at phylum and genus levels

表3 氨氧化和反硝化功能基因丰度与土壤理化性质的Pearson相关性分析

Table 3 Pearson correlation analysis between abundance of functional genes related to ammonia oxidation and denitrification and soil physicochemical properties

基因 Gene	全氮 Total N	全磷 Total P	全钾 Total K	有机质 Soil organic matter （SOM）	硝态氮 Nitrate N	铵态氮 Ammonium N	亚硝酸盐氮Nitrite N	速效磷 Available P	速效钾 Available K	pH
amoA	0.690*	0.600*	0.724**	0.839**	0.207	0.183	0.183	0.483	0.798**	0.685*
bamoA	0.784**	0.689*	0.656*	0.789**	0.451	0.370	0.300	0.697*	0.861**	0.707*
nirK	0.663*	0.531	0.540	0.713**	0.302	0.234	0.051	0.570	0.791**	0.712**
nirS	0.587*	0.448	0.492	0.540	0.456	0.262	0.162	0.618*	0.781**	0.850**

图4 氨氧化和反硝化功能基因丰度与土壤理化性质的冗余分析TN：全氮Total nitrogen； TP：全磷Total phosphorous； TK：全钾Total potassium； SOM：有机质Soil organic matter； NO3--N：硝态氮Nitrate nitrogen； NH4+-N：铵态氮Ammonium nitrogen； NO2--N：亚硝酸盐氮Nitrite nitrogen； AP：速效磷Available phosphorous； AK：速效钾Available potassium. 下同The same below.

Fig.4 Redundancy analysis between abundance of functional genes related to ammonia oxidation and denitrification with soil physicochemical properties

图5 氨氧化和反硝化功能微生物群落与土壤理化因子关联分析A：氨氧化古菌Ammonia-oxidizing archaea； B：氨氧化细菌Ammonia-oxidizing bacteria； C： nirK型反硝化菌nirK-type denitrifying bacteria； D： nirS型反硝化菌nirS-type denitrifying bacteria. *： P<0.05； **： P<0.01； ***： P<0.001.

Fig.5 Correlation analysis between ammonia-oxidizing and denitrifying microbial communities with soil physicochemical properties

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