氮磷添加对杉木根际土壤丛枝菌根真菌和易提取球囊霉素的影响

王婷; 沈益康; 汪鹞雄; 甘秉平; 李全; 刘美华; 宋新章

doi:10.12356/j.2096-8884.2021-0018

氮磷添加对杉木根际土壤丛枝菌根真菌和易提取球囊霉素的影响

doi: 10.12356/j.2096-8884.2021-0018

浙江农林大学，省部共建亚热带森林培育国家重点实验室，浙江杭州 311300

基金项目: 国家“十三五”重点研发计划课题“人工林生产力形成的关键生理生态与环境控制机制”（2016YFD0600201）和浙江省“万人计划”科技创新领军人才项目（2018R52027）

详细信息

作者简介:
王婷：E-mail: wangting716@126.com

通讯作者:
E-mail: xzsong@126.com

中图分类号: S791.27
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出版历程
- 收稿日期: 2021-09-23
- 录用日期: 2021-12-16
- 网络出版日期: 2022-01-08
- 刊出日期: 2022-01-17

Effects of Nitrogen and Phosphorus Addition on Arbuscular Mycorrhizal Fungi and Easily Extracted Glomalin-related Soil Protein in the Chinese Fir Plantation

State Key Laboratory of Subtropical Silviculture, Zhejiang A&F University, Hangzhou 311300, Zhejiang, China

摘要

摘要: 目的分析氮磷添加对杉木（Cunninghamia lanceolata）根际丛枝菌根真菌（Arbuscular mycorrhizal fungi，AMF）性状和群落结构、易提取球囊霉素（Easily extracted glomalin-related soil protein，EE-GRSP）以及土壤理化性质的影响，为杉木人工林的可持续经营提供参考。方法以杉木人工林中的10 a杉木为研究对象，以未添加氮磷为控制对照组（N0: 0 kg N·hm⁻²·a⁻¹ + P0: 0 mg P·kg⁻¹），设置了不同水平的单独氮添加（N30 + P0: 30 kg N·hm⁻²·a⁻¹ + 0 mg P·kg⁻¹和N60 + P0: 60 kg N·hm⁻²·a⁻¹ + 0 mg P·kg⁻¹）、单独磷添加（N0 + P20: 0 kg N·hm⁻²·a⁻¹ + 20 mg P·kg⁻¹和N0 + P40: 0 kg N·hm⁻²·a⁻¹ + 40 mg P·kg⁻¹）及其复合作用（N30 + P20，N30 + P40，N60 + P20，N60 + P40），共9个处理，测定分析了AMF侵染率、孢子密度和易提取球囊霉素含量，并通过高通量测序技术分析了不同处理下杉木根际土壤AMF群落结构和多样性指数。结果 1）在所有处理下，杉木根际土壤的AMF优势属均为球囊霉属（Glomus，> 95%）。2）与对照相比，单独氮添加显著提高了AMF的侵染率（191.5%—537.7%，P < 0.05）和孢子密度（59.2%—85.7%，P < 0.05），但显著降低了易提取球囊霉素的含量（9.8%—16.8%，P < 0.05）。N30 + P0处理显著提高了AMF丰富度（P < 0.05）。3）单独磷添加显著提高了AMF的侵染率（471.4%—658.3%，P < 0.05），显著降低了易提取球囊霉素的含量（8.9%—31.7%，P < 0.05），对孢子密度无显著影响（P > 0.05）。N0 + P40显著提高了AMF多样性（P < 0.05）。4）N30 + P20显著降低了AMF的侵染率（79.7%）、显著提高了孢子密度（79.3%）和易提取球囊霉素含量（22.5%）（P < 0.05）；在N60处理下，磷添加对侵染率和易提取球囊霉素无显著影响（P > 0.05），N60 + P40显著降低了孢子密度（64.8%，P < 0.05）。5）土壤含水量与孢子密度和易提取球囊霉素含量呈显著正相关（P < 0.05），与AMF侵染率呈极显著负相关（P < 0.01）。孢子密度与有效氮磷比呈显著正相关（P < 0.05）。AMF侵染率与易提取球囊霉素呈极显著负相关（P < 0.01）。AMF多样性与土壤理化因子无显著相关性（P > 0.05）。结论在自然状态下，适当的磷添加（40 mg P·kg⁻¹）或氮添加（30 kg N·hm⁻²·a⁻¹）可以提高AMF的侵染率和多样性，从而促进AMF和杉木的共生关系，提高杉木生产力；在未来氮沉降增加的背景下，磷添加将提高土壤营养元素含量，降低AMF的侵染率，从而减弱AMF和杉木的共生关系。
- 丛枝菌根真菌 /
- 多样性 /
- 氮沉降 /
- 磷添加 /
- 杉木
Abstract: Objective The aim of this study is to analyze the effects of nitrogen (N) and phosphorus (P) addition on the characteristics and community structure of arbuscular mycorrhizal fungi (AMF), easily extracted glomalin-related soil protein (EE-GRSP), and soil physicochemical properties in the rhizosphere of Chinese fir (Cunninghamia lanceolata), further to provide a scientific reference for the sustainable management of Chinese fir plantations. Method The 10-year-old Chinese firs in the plantation were selected, 9 treatments were set up, including the control (without N and P addition), only N addition (N30 + P0: 30 kg N·hm⁻²·a⁻¹ + 0 mg P·kg⁻¹, N60 + P0: 60 kg N·hm⁻²·a⁻¹ + 0 mg P·kg⁻¹), only P addition (N0 + P20: 0 kg N·hm⁻²·a⁻¹ + 20 mg P·kg⁻¹, N0 + P40: 0 kg N·hm⁻²·a⁻¹ + 40 mg P·kg⁻¹), and their combinations (N30 + P20, N30 + P40, N60 + P20, N60 + P40), with three independent replicates for each treatment. AMF colonization rate, spore density and easily extracted GRSP (EE-GRSP) were analyzed, as the high-throughput sequencing technology was used to analyze the community structure and diversity index of AMF by different treatments. Result The dominant genus of AMF in the rhizosphere of Chinese fir was Glomus (> 95%) under all treatments. Compared to the control, N addition significantly increased the AMF colonization rate (191.5%−537.7%, P < 0.05) and the spore density (59.2%−85.7%, P < 0.05), while significantly reduced the EE-GRSP concentration (9.8%−16.8%, P < 0.05). N30 + P0 treatment significantly increased the AMF richness (P < 0.05). Phosphorus addition significantly increased the AMF colonization rate (471.4%−658.3%, P < 0.05), reduced the EE-GRSP concentration (8.9%−31.7%, P < 0.05), however, it showed no significant effect on the spore density (P > 0.05). N0 + P40 significantly increased the AMF diversity (P < 0.05). N30 + P20 significantly reduced the AMF colonization rate (79.7%, P < 0.05), while increased the spore density (79.3%, P < 0.05) and the EE-GRSP concentration (22.5%, P < 0.05). Under N60 treatment, P addition did not significantly influence the colonization rate and EE-GRSP concentration (P > 0.05), except N60 + P40 significantly reduced the spore density (64.8%, P < 0.05). Soil moisture content was significantly positive with spore density and EE-GRSP (P < 0.05), but negative with AMF colonization rate (P < 0.01). There was a significant positive correlation between soil spore density and soil AN∶AP (P < 0.05). Shannon index and Chao1 index had no significant correlation with soil physical and chemical factors (P > 0.05). Conclusion In fact, appropriate P addition (40 mg P·kg⁻¹) or N addition (30 kg N·hm⁻²·a⁻¹) could increase the colonization rate and diversity of AMF, promote the symbiotic relationship between AMF and Chinese fir, and further enhance the Chinese fir productivity. Under the scenario of increasing nitrogen deposition, P addition could increase soil nutrients, reduce the colonization rate of AMF, and weaken the symbiotic relationship between AMF and Chinese fir.
- Arbuscular mycorrhizal fungi /
- diversity /
- nitrogen deposition /
- phosphorus addition /
- Chinese fir

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图 1 AMF群落花瓣图（OTU水平）

Figure 1. Petal diagram of AMF community at the OTU level

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图 2 不同氮磷添加下土壤AMF群落组成

Figure 2. Soil AMF community compositions under different N and P addition treatments in the Chinese fir plantation

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图 3 不同氮磷添加下杉木AMF的Shannon和Chao1指数

Figure 3. Shannon and Chao1 indexes of under different N and P addition treatments in the Chinese fir plantation

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图 4 不同氮磷添加下杉木AMF侵染率（a）和孢子密度（b）

Figure 4. AMF colonization rate (a) and spore density (b) under different N and P addition treatments in the Chinses fir plantation

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图 5 不同氮磷添加下杉木的易提取球囊霉素含量

Figure 5. EE-GRSP under different N and P addition in the Chinese fir plantation

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表 1 不同氮和磷添加下杉木土壤理化性质差异

Table 1. Differences of physical and chemical properties of Chinese fir soil under different nitrogen and Phosphorus Additions

处理 Treatment	含水率 Soil moisture/%	pH值 pH-value	有效氮 Available N/(mg·kg⁻¹)	有效磷 Available P/(mg·kg⁻¹)	有效氮磷比 AN∶AP	全氮 Total N/(g·kg⁻¹)	全磷 Total P/(g·kg⁻¹)
N0+P0	29.52±2.60Aa	4.59±0.09Ab	31.36±5.82Bb	3.39±1.38Aa	9.99±2.72Cb	0.88±0.31Bb	0.12±0.04Bb
N0+P20	24.49±0.94b	4.78±0.13a	61.97±4.66a	3.35±0.38a	18.54±1.06a	1.62±0.36a	0.23±0.05a
N0+P40	27.79±2.36a	4.80±0.03a	37.33±3.42b	3.70±0.25a	10.09±0.31b	0.87±0.10b	0.17±0.02a
N30+P0	31.65±3.73Aa	4.76±0.15Aa	34.72±8.08Ba	1.74±0.55Ab	20.41±2.02Ba	0.79±0.20Ba	0.11±0.01Bb
N30+P20	36.14±2.78a	4.72±0.07a	50.40±11.64a	2.32±1.10b	23.84±8.83a	1.14±0.22a	0.13±0.01ab
N30+P40	32.80±2.21a	4.67±0.12a	38.08±4.88a	4.79±1.09a	8.09±1.13b	0.97±0.22a	0.15±0.02a
N60+P0	32.59±3.17Aa	4.32±0.01Bb	69.44±14.56Aa	2.32±0.38Ab	30.20±6.97Aa	1.38±0.23Aa	0.17±0.02Aa
N60+P20	31.04±1.32a	4.79±0.15a	46.29±11.98a	3.36±1.15b	14.09±2.19b	1.12±0.39a	0.14±0.03ba
N60+P40	30.84±5.15a	4.60±0.10a	54.13±11.27a	5.60±1.23a	9.74±1.13b	1.30±0.34a	0.18±0.03a
注：对照（N0 + P0）。不同大写字母表示在P0下不同氮添加处理间差异显著(P < 0.05)，不同小写字母表示在相同模拟氮沉降梯度下不同磷添加处理间差异显著(P < 0.05)，下同。 The control(N0 + P0). Capital letters indicate significant differences among N addition treatments under the P0 condition (P < 0.05). Lowercase letters indicate significant differences among P addition treatments at the identical N addition treatments (P < 0.05). The same below.

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表 2 氮磷添加对杉木AMF多样性、侵染率、孢子密度和易提取球囊霉素双因素方差分析

Table 2. Two-way ANOVA of effects of N and P addition on AMF diversity, colonization rate, spore density, and EE-GRSP in the Chinese fir plantation

因素 Factors	氮沉降 Nitrogen deposition		磷添加 Phosphorus addition		交互作用 Interaction
因素 Factors	F值 F-Value	显著性 P Value	F值 F-Value	显著性 P Value	F值 F-Value	显著性 P Value
Shannon指数 Shannon index	0.995	0.389	1.627	0.224	2.395	0.089
Chao1指数 Chao1 index	1.197	0.325	1.029	0.377	2.794	0.058
侵染率 Colonization rate	129.082	0.000	16.642	0.000	309.338	0.000
孢子密度 Spore density	166.913	0.000	95.375	0.000	39.741	0.000
易提取球囊霉素 EE-GRSP	1.830	0.189	1.140	0.342	0.864	0.504

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表 3 AMF多样性、侵染率、孢子密度和易提取球囊霉素与土壤理化性质的相关性分析

Table 3. Spearman's correlation coefficients (r) for relationships between AMF diversity, colonization rate, spore density, EE-GRSP and soil properties

因素 Factors	土壤理化性质 Soil properties
因素 Factors	含水率 Soil moisture	pH值 pH-value	有效氮 Available N	有效磷 Available P	有效氮磷比AN∶AP	全氮 Total N	全磷 Total P	Shannon指数 Shannon index	Chao1指数 Chao1 index	侵染率 Colonization rate	孢子密度 Spore density	易提取球囊霉素 EE-GRSP
Shannon指数 Shannon index	−0.037	0.159	−0.031	0.112	−0.086	−0.081	0.196	1	0.462*	0.078	−0.200	0.338
Chao1指数 Chao1 index	−0.016	0.318	−0.227	−0.103	−0.016	−0.298	−0.243	0.462*	1	0.036	−0.005	0.285
侵染率 Colonization rate	−0.506**	0.428*	0.046	−0.137	0.193	−0.027	0.252	0.078	0.036	1	−0.281	−0.525**
孢子密度 Spore density	0.662**	0.055	0.140	−0.318	0.385*	0.156	−0.295	−0.200	−0.005	−0.281	1	0.034
易提取球囊霉素 EE-GRSP	0.450*	−0.077	0.366	−0.046	0.234	0.324	−0.032	0.338	0.285	−0.525**	0.034	1
注：* P < 0.05；**P < 0.01。

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