灌丛化对黄土高原草地土壤有机碳组分与稳定性的影响及其微生物调控机制

doi:10.11686/cyxb2025034

草业学报 ›› 2025, Vol. 34 ›› Issue (12): 1-15.DOI: 10.11686/cyxb2025034

• 研究论文 •

灌丛化对黄土高原草地土壤有机碳组分与稳定性的影响及其微生物调控机制

景煜都¹^,²^,³(), 刘小伟⁴, 梁可⁴, 封俊豪⁴, 于强⁵, 郭梁⁵^,⁶()

^1.中国科学院教育部水土保持与生态环境研究中心，陕西杨凌 712100
^2.中国科学院水利部水土保持研究所，陕西杨凌712100
^3.中国科学院大学，北京 100049
^4.西北农林科技大学草业与草原学院，陕西杨凌 712100
^5.西北农林科技大学水土保持与荒漠化整治全国重点实验室，陕西杨凌 712100
^6.宁夏云雾山国家级自然保护区管理局，宁夏固原 756000

收稿日期:2025-02-02 修回日期:2025-04-03 出版日期:2025-12-20 发布日期:2025-10-20
通讯作者: 郭梁
作者简介:E-mail： guoliang2014@nwafu.edu.cn
景煜都（1998-），男，山西运城人，在读硕士。E-mail： jingyudu22@mails.ucas.ac.cn
基金资助:
国家自然科学基金(42377471);宁夏回族自治区自然科学基金(2024A AC05099)

Impacts of shrub encroachment on the fraction and stability of soil organic carbon of grassland on the Loess Plateau， and the underlying microbial mechanisms

Yu-du JING¹^,²^,³(), Xiao-wei LIU⁴, Ke LIANG⁴, Jun-hao FENG⁴, Qiang YU⁵, Liang GUO⁵^,⁶()

^1.The Research Center for Soil and Water Conservation and Ecological Environment，Chinese Academy of Sciences and Ministry of Education，Yangling 712100，China
^2.Institute of Soil and Water Conservation，Chinese Academy of Sciences and Ministry of Water Resources，Yangling 712100，China
^3.University of Chinese Academy of Sciences，Beijing 100049，China
^4.College of Grassland Agriculture，Northwest A&F University，Yangling 712100，China
^5.State Key Laboratory of Soil and Water Conservation and Desertification Control，Northwest A&F University，Yangling 712100，China
^6.Ningxia Yunwushan National Nature Reserve Administration，Guyuan 756000，China

Received:2025-02-02 Revised:2025-04-03 Online:2025-12-20 Published:2025-10-20
Contact: Liang GUO

摘要/Abstract

摘要：

灌丛化显著影响全球草地生态系统的碳循环，但其对土壤有机碳组分及稳定性的影响，尤其是微生物在其中的调控机制，仍然缺乏深入研究。本研究以黄土高原半干旱草原为对象，分析了不同类型灌丛化对土壤理化与生物性质、有机碳组分与占比、微生物群落特性的影响，以及其中的微生物调控机制。结果表明，半灌木（白莲蒿）和灌木（矮脚锦鸡儿）扩张的生态影响差异不显著，但两种类型的灌丛化显著提高了土壤含水量（SWC）、土壤总有机碳（SOC）含量、可溶性有机碳（DOC）含量与全量养分，促进了碳循环相关酶活性上升与微生物残体碳积累，并显著提高了矿物结合态有机碳（MAOC）、颗粒态有机碳（POC）的绝对含量与MAOC的相对占比，显著降低了土壤pH与POC的相对占比。此外，灌丛化显著改变了土壤细菌与真菌群落组成，提高了土壤细菌的硝化作用、硫化物氧化与铁呼吸功能，同时增强了真菌的凋落物腐生与木本腐生营养型的丰度。土壤微生物残体碳及细菌的功能均与POC和MAOC的含量和占比显著相关。偏最小二乘路径分析进一步表明，与碳循环相关的胞外酶活性和微生物残体碳分别是调控土壤有机碳组分含量与其相对占比的直接微生物因素，而微生物群落组成通过间接途径影响这些过程。总之，灌丛化不仅显著增加了草地土壤碳库的总量及各有机碳组分的含量，还提高了土壤碳库的稳定性，而土壤微生物分泌的胞外酶与微生物残体碳在该过程中起到重要调控作用。本研究为深入理解灌丛化对草地土壤碳组分的微生物调控机制，并为预测黄土高原灌丛化草地土壤碳库未来变化提供了理论支持。

关键词: 黄土高原半干旱草地, 灌丛化, 有机碳组分, 土壤微生物

Abstract:

Shrub encroachment is significantly affecting carbon cycling in grassland ecosystems worldwide， yet its impact on the components and stability of grassland soil organic carbon， and particularly the underlying microbial regulatory mechanisms， remains insufficiently studied. In this study， we focused on the semi-arid grassland of the central Loess Plateau， and analyzed the effects of shrub encroachment on various soil physicochemical and biological properties， organic carbon components and their proportions， microbial community structure， functions， and trophic types， as well as the underlying microbial regulatory mechanisms. The results show that shrub encroachment significantly increased the total organic carbon， dissolved organic carbon， mineral-associated organic carbon （MAOC）， and particulate organic carbon （POC） contents in soil， and the relative proportion of MAOC， but significantly decreased the relative proportion of POC. Furthermore， shrub encroachment significantly altered the bacterial and fungal community composition by affecting soil physicochemical properties， with different impacts on the relative abundance of amplicon sequence variants of different phyla. Shrub encroachment enhanced soil bacterial nitrification， sulfur oxidation， and iron respiration functions， as well as the abundance of fungal saprotrophic and woody saprotrophic type functions， but reduced the abundance of bacterial methylotrophy and aromatic compound degradation functions. The soil microbial residue carbon and bacterial functions were significantly correlated with the contents and proportions of POC and MAOC. A partial least squares path analysis showed that extracellular enzyme activity and microbial residue carbon， related to carbon cycling， were the microbial factors directly regulating the contents and relative proportions of soil organic carbon components， whereas microbial community composition influenced these processes indirectly. These results indicate that shrub encroachment not only increases the total amount of soil carbon and the contents of soil organic carbon components in grassland soils， but also significantly enhances the stability of the soil carbon pool， with extracellular enzymes secreted by soil microbes and microbial residue carbon playing crucial regulatory roles. The findings of this study provide new information about the mechanisms by which microbes regulate organic carbon components in grassland soils under shrub encroachment. These results provide theoretical support for predicting future changes in the soil carbon pool of shrub-encroached grassland on the Loess Plateau.

Key words: semi-arid grasslands on the Loess Plateau, shrub encroachment, organic carbon fractions, soil microorganisms

景煜都, 刘小伟, 梁可, 封俊豪, 于强, 郭梁. 灌丛化对黄土高原草地土壤有机碳组分与稳定性的影响及其微生物调控机制[J]. 草业学报, 2025, 34(12): 1-15.

Yu-du JING, Xiao-wei LIU, Ke LIANG, Jun-hao FENG, Qiang YU, Liang GUO. Impacts of shrub encroachment on the fraction and stability of soil organic carbon of grassland on the Loess Plateau， and the underlying microbial mechanisms[J]. Acta Prataculturae Sinica, 2025, 34(12): 1-15.

图/表 7

表1 灌丛化对土壤理化与生物指标的影响

Table 1 Impacts of shrub encroachment on soil physicochemical and biological properties

土壤指标 Soil property	无灌木扩张 NSE	半灌木扩张 SSE	灌木扩张 SE
SWC （%）	14.80±1.01b	20.41±0.80a	21.26±1.18a
pH	8.18±0.01a	8.05±0.01b	8.06±0.02b
SOC （g·kg^-1） DOC （mg·kg^-1）	16.74±0.87c 51.99±2.06b	25.77±0.87b 72.12±4.16a	28.41±0.69a 74.88±2.70a
TN （g·kg^-1）	1.60±0.04b	2.94±0.17a	3.14±0.17a
TP （g·kg^-1）	0.58±0.01c	0.67±0.01b	0.77±0.02a
CA （g·kg^-1）	8.07±0.03b	8.16±0.10b	8.56±0.05a
BG （nmol·g^-1·h^-1）	15.22±0.91c	36.32±3.35a	29.43±1.28b
CBH （nmol·g^-1·h^-1）	9.69±0.60a	9.48±1.51a	10.38±0.17a

图1 灌丛化对土壤有机碳组分与相对占比及微生物残体碳的影响POC： Particulate organic carbon； MAOC： Mineral associated organic carbon； BNC： Bacterial necromass carbon； FNC： Fungal necromass carbon； MNC： Microbial necromass carbon；误差棒代表均值±标准误Error bars represent means±standard errors. 下同The same below.

Fig.1 Impacts of shrub encroachment on the content and relative proportion of soil organic carbon fractions and microbial necromass carbon

图2 灌丛化对土壤细菌与真菌门水平群落结构的影响NMDS1和NMDS2分别为非线性多维度量分析第一轴与第二轴NMDS1 and NMDS2 represent the first and second axis of non-metric multidimensional scaling analysis， respectively.

Fig.2 Impacts of shrub encroachment on phylum-level community structures of soil bacteria and fungi

图3 土壤细菌与真菌ASVs的系统发育树及与土壤基本理化性质的相关性系统发育树中，不同颜色代表ASVs属于不同的细菌或真菌门。相关性热图基于Spearman相关性，其中不同颜色代表土壤微生物ASVs与土壤基础理化性质相关性不同，图中只保留具有统计显著性（P<0.05）的相关关系，红色代表负相关，蓝色代表正相关。In the phylogenetic trees， different colors represent ASVs that belong to different bacterial or fungal phylum. In the correlation heatmaps based on Spearman’s correlation， different colors indicate different correlations between soil microbial ASVs and soil basic physicochemical properties. Only the correlations with statistical significance （P<0.05） were retained in the heatmaps， where red indicates negative correlations， while blue indicates positive correlations.

Fig.3 The phylogenetic trees of soil bacterial and fungal ASVs and the correlation between the ASVs and soil basic physicochemical properties

图4 灌丛化对土壤细菌功能与真菌营养型的影响**： P<0.01； *： P<0.05； NS： P>0.05.

Fig.4 Impacts of shrub encroachment on soil bacterial functions and fungal trophic types

图5 土壤有机碳组分影响因素及土壤有机碳组分与土壤微生物性质的相关分析（a）土壤理化因子与微生物群落组成对土壤有机碳组分影响的冗余分析，红色箭头所指为对土壤有机碳组分影响显著的解释变量，灰色箭头所指为对其影响不显著的解释变量。NMDS1：土壤细菌群落组成，以非度量多维尺度分析（NMDS）第一轴坐标表征； F-NMDS1：土壤真菌群落组成，以NMDS第一轴坐标表征。（b）各个解释变量对土壤有机碳组分总体变化的贡献。（c）土壤有机碳组分及占比与土壤微生物残体碳、细菌功能与真菌营养型的Pearson相关性热图，红色代表负相关，蓝色代表正相关；星号代表相关的显著性。（a） Redundance analysis for the effects of soil physicochemical properties and microbial community composition on soil carbon fraction. Red arrows point to properties that significantly affect soil organic carbon fraction， while gray arrows point to properties with no significant effect. NMDS1： Community composition of soil bacteria indicated with the first axis of non-metric multidimensional scaling； F-NMDS1： Community composition of soil fungi indicated with the first axis of non-metric multidimensional scaling. （b） Contribution of individual properties on the changes of soil organic carbon fraction. （c） Heatmap based on Pearson’s correlation between soil organic carbon fraction as well as the ratio of each fraction to total organic carbon and soil microbial properties including necromass carbon， bacterial functions and fungal trophic types. Red represents negative correlations， while red represents positive correlations. Star marks indicate the significance of correlations.

Fig.5 Analysis of the potential drivers of soil organic carbon fraction and the correlation between soil organic carbon fraction and soil microbial properties

图6 灌丛化对土壤有机碳组分及其占比调控机制的偏最小二乘路径模型（a）：灌丛化对土壤有机碳组分及矿物结合态有机碳占比调控机制的偏最小二乘路径模型，每个方框代表不同的显变量，方框中括号内的因子为用于计算该显变量的隐变量，箭头代表模型中的路径，其侧数字为该路径标准化路径系数，星号代表路径的统计显著性The partial-least square path modeling for the regulatory mechanisms of shrub encroachment on soil organic carbon fraction and the ratio of soil mineral-associated organic carbon to total organic carbon. Each square represents each of the different observable variables， the properties bracketed represent hidden variables used to calculate the observable variable. Arrows represent paths in the model， the numbers aside represent the standardized pathway index， the star marks indicate statistical significance of the paths； ***： P<0.001； **： P<0.01； *： P<0.05.

Fig.6 The partial-least square path modeling for the regulatory mechanisms of shrub encroachment on soil organic carbon fractions and their ratio to soil total organic carbon

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灌丛化对黄土高原草地土壤有机碳组分与稳定性的影响及其微生物调控机制

Impacts of shrub encroachment on the fraction and stability of soil organic carbon of grassland on the Loess Plateau， and the underlying microbial mechanisms

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