Article(id=1309574766298755233, tenantId=1146029695717560320, journalId=1308397196074385509, issueId=1309574765791244449, articleNumber=null, orderNo=null, doi=10.11733/j.issn.1007-0435.2026.07.001, pmid=null, cstr=null, oa=null, hot=0, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=null, receivedDate=1770566400000, receivedDateStr=2026-02-09, revisedDate=1774800000000, revisedDateStr=2026-03-30, acceptedDate=null, acceptedDateStr=null, onlineDate=1790156555008, onlineDateStr=2026-09-23, pubDate=null, pubDateStr=null, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1790156555008, onlineIssueDateStr=2026-09-23, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1790156555008, creator=13701087609, updateTime=1790156555008, updator=13701087609, issue=Issue{id=1309574765791244449, tenantId=1146029695717560320, journalId=1308397196074385509, year='2026', volume='34', issue='7', pageStart='2341', pageEnd='2752', issueExtLink='null', onlineDate='null', pubDate='1784044800000', pubDateStr='2026-07-15', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1790156554887, creator='13701087609', updateTime=1790156849638, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext={EN=IssueExt(id=1309576002150424823, tenantId=1146029695717560320, journalId=1308397196074385509, issueId=1309574765791244449, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1309576002154619128, tenantId=1146029695717560320, journalId=1308397196074385509, issueId=1309574765791244449, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null, downloadFileDto=null}, startPage=2341, endPage=2354, ext={EN=ArticleExt(id=1309574766659465379, articleId=1309574766298755233, tenantId=1146029695717560320, journalId=1308397196074385509, language=EN, title=Effects of Grassland Degradation on Soil Carbon Cycle, Microbial Functional Genes and Metabolism, columnId=null, journalTitle=Acta Agrestia Sinica, columnName=null, runingTitle=null, highlight=null, articleAbstract=Under the context of global climate change, the influence mechanism of grassland degradation on soil carbon cycling remain poorly understood. In this study, we investigated 15 paired degraded and non-degraded grassland sites on the Qinghai-Tibet Plateau, integrated microcosm incubation, metagenomics and metabolomics to assess the impacts of degradation on key carbon cycling processes, organic carbon fractions, microbial functional genes, and metabolic intermediates. The results showed that grassland degradation significantly reduced labile carbon pools, with particulate organic carbon, dissolved organic carbon, and microbial biomass carbon decreasing by 48.21%, 30.87%, and 20.21%, respectively. Meanwhile, soil organic carbon mineralization increased by 56.71%, and carbon accumulation efficiency declined by 32.51%. Degradation also led to a decrease in the diversity of carbon cycling functional genes (-2.95%, P<0.05), while the abundance of starch-degrading genes and carbohydrate-related metabolites increased significantly (by 9.39% and 49.6%, respectively). These changes indicated that intensified microbial carbon limitation and a shift in metabolic strategy from synthesis to decoomposition. This study elucidates the key mechanisms underlying soil carbon cycling responses to grassland degradation from a multi-factor coupling perspective, revealing that environmental factors and microbial traits jointly regulated carbon cycling, with microbial functional genes and metabolic intermediates playing a dominat role., authors=DING Cheng-xiang1, LI Xing-fu2, SU De-rong3, LIU Yu4, authorsList=DING Cheng-xiang, LI Xing-fu, SU De-rong, LIU Yu, authorCompany=1 Qinghai University, Xining, Qinghai Province 810016, China; 2 Institute of Industrial Development Planning, National Forestry and Grassland Administration, Beijing 100010, China; 3 Beijing Forestry University, Beijing 100083, China; 4 Chinese Research Academy of Environmental Sciences, Beijing 100012, China, correspAuthors=null, authorNote=null, correspAuthorsNote=null, copyrightStatement=null, copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=null, magXml=null, pdfUrl=null, pdf=null, pdfFileSize=null, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=null, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=null, mapNumber=null, fund=null), CN=ArticleExt(id=1309574766579773602, articleId=1309574766298755233, tenantId=1146029695717560320, journalId=1308397196074385509, language=CN, title=草地退化对土壤碳循环、微生物功能基因及代谢的影响, columnId=1309565024058835950, journalTitle=草地学报, columnName=研究论文, runingTitle=null, highlight=null, articleAbstract=在全球气候变化背景下,草地退化对土壤碳循环的影响机制仍缺乏系统认知。本研究以青藏高原15对未退化—退化配对样地为对象,结合微宇宙培养、宏基因组与宏代谢组分析,综合评估退化对土壤碳循环关键过程、有机碳组分、微生物功能基因及代谢产物的影响。结果表明:草地退化显著降低活性碳库(颗粒有机碳、溶解性有机碳和微生物生物量碳分别下降48.21%、30.87%和20.21%),同时增强有机碳矿化(提高56.71%),降低碳积累效率(下降32.51%)。退化还导致碳循环功能基因多样性下降(-2.95%,P<0.05),但淀粉分解基因及碳水化合物代谢产物显著增加(分别上升9.39%和49.6%,P<0.05),表明微生物碳限制加剧,代谢策略由合成向分解转变。本研究从多要素耦合视角阐明了草地退化调控土壤碳循环的关键机制,表明环境因子与微生物特征协同驱动碳循环变化,其中微生物功能基因与代谢产物发挥主导作用。, authors=丁成翔1, 李兴福2, 苏德荣3, 刘禹4, authorsList=丁成翔, 李兴福, 苏德荣, 刘禹, authorCompany=1 青海大学, 青海 西宁 810016; 2 国家林业和草原局产业发展规划院, 北京 100010; 3 北京林业大学, 北京 100083; 4 中国环境科学研究院, 北京 100012, correspAuthors=null, authorNote=丁成翔(1980-),男,回族,宁夏中卫人,博士,主要从事草地生态学研究,E-mail:2011990016@qhu.edu.cn, correspAuthorsNote=null, copyrightStatement=null, copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=null, magXml=null, pdfUrl=null, pdf=TxzUECf24jtRK66Mu2ACIQ==, pdfFileSize=10529578, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=null, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=null, mapNumber=null, fund=青海省中央引导地方科技发展资金项目 (2024ZY011); 青海省重点研发与转化计划科技成果转化专项项目 (2024-NK-138); 中央级公益性科研院所基本科研业务费青年探索项目 (2025YSKY-39))}, authors=null, keywords=[Keyword(id=1309574864965566701, tenantId=1146029695717560320, journalId=1308397196074385509, articleId=1309574766298755233, language=CN, orderNo=1, keyword=土壤碳循环), Keyword(id=1309574865326276846, tenantId=1146029695717560320, 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Grassland carbon sink in China and its promotion strategies[J]. Bulletin of National Natural Science Foundation of China, 2023, 37(4):587-602 彭云峰, 常锦峰, 赵霞, 等. 中国草地生态系统固碳能力及其提升途径[J]. 中国科学基金, 2023, 37(4):587-602 BAI Y F, COTRUFO M F. Grassland soil carbon sequestration: current understanding, challenges, and solutions[J]. Science, 2022, 377(6606):603-608 SCHARLEMANN J P, TANNER E V, HIEDERER R, et al. Global soil carbon: understanding and managing the largest terrestrial carbon pool[J]. Carbon Management, 2014, 5(1):81-91 LIU L L, SAYER E J, DENG M F, et al. The grassland carbon cycle: mechanisms, responses to global changes, and potential contribution to carbon neutrality[J]. Fundamental Research, 2023, 3(2):209-218 WU M C, XIE S Y, ZANG J X, et al. Multiple anthropogenic environmental stressors structure soil bacterial diversity and community network[J]. Soil Biology and Biochemistry, 2024, 198:109560 BROWN R W, CHADWICK D R, ZANG H D, et al. Use of metabolomics to quantify changes in soil microbial function in response to fertiliser nitrogen supply and extreme drought[J]. Soil Biology and Biochemistry, 2021, 160:108351 CAO J J, ADAMOWSKI J F, DEO R C, et al. Grassland degradation on the Qinghai-Tibetan Plateau: reevaluation of causative factors[J]. Rangeland Ecology & Management, 2019, 72(6):988-995 LIU S B, ZAMANIAN K, SCHLEUSS P M, et al. Degradation of Tibetan grasslands: consequences for carbon and nutrient cycles[J]. Agriculture, Ecosystems & Environment, 2018, 252:93-104 HU W J, CAI Y, LI X Q, et al. Microbial carbon accumulation efficiency in global soils resolved via13C-glucose amendment experiments[J]. Global Biogeochemical Cycles, 2025, 39(6):e2025GB008553 ABDALLA K, MUTEMA M, CHIVENGE P, et al. Grassland degradation significantly enhances soil CO2 emission[J]. Catena, 2018, 167:284-292 WANG P Y, WANG J S, WANG S, et al. Soil moisture threshold of methane uptake in Alpine grassland ecosystems[J]. Global Change Biology, 2025, 31(2):e70062 ZHOU J C, LIU S, VAN GROENIGEN K JAN, et al. Contrasting responses of particulate and mineral-associated organic carbon stocks to grazing exclusion in an Alpine meadow[J]. Agriculture, Ecosystems & Environment, 2026, 400:110227 HUANG Q, WANG B R, SHEN J K, et al. Shifts in C-degradation genes and microbial metabolic activity with vegetation types affected the surface soil organic carbon pool[J]. Soil Biology and Biochemistry, 2024, 192:109371 WANG N N, LI K X, YUAN F H, et al. Faster cycling but lower efficiency: A microbial metabolic perspective on carbon loss after wetland conversion to cropland[J]. Soil Biology and Biochemistry, 2024, 189:109260 HAO A H, XUE X, PENG F, et al. Different vegetation and soil degradation characteristics of a typical grassland in the Qinghai-Tibetan Plateau[J]. Acta Ecologica Sinica, 2020, 40(3):964-975 郝爱华, 薛娴, 彭飞, 等. 青藏高原典型草地植被退化与土壤退化研究[J]. 生态学报, 2020, 40(3):964-975 ZHANG S N, LI F C, LI J B, et al. The influence of different degrees of degradation on soil dissolved organic carbon and its characteristics in sloping grassland[J]. Acta Agrestia Sinica, 2025, 33(9):2941-2949 张胜男, 李富翠, 李金波, 等. 不同退化程度对坡地草地土壤可溶性有机碳及其特征的影响[J]. 草地学报, 2025, 33(9):2941-2949 BAO S D. Soil and agricultural chemistry analysis [M].3rd Edition.Beijing:China Agriculture Press, 2000:42-80 鲍士旦.土壤农化分析[M]. 3版. 北京:中国农业出版社, 2000:42-80 BRIDGHAM S D, YE R. Organic matter mineralization and decomposition[J]. Methods in biogeochemistry of wetlands. 2013, 10: 385-406 MA L, JIANG X L, LIU G H, et al. Environmental factors and microbial diversity and abundance jointly regulate soil nitrogen and carbon biogeochemical processes in Tibetan wetlands[J]. Environmental Science & Technology, 2020, 54(6):3267-3277 TAŞ N, PRESTAT E, MCFARLAND J W, et al. Impact of fire on active layer and permafrost microbial communities and metagenomes in an upland Alaskan boreal forest[J]. The ISME Journal, 2014, 8(9):1904-1919 WU H B, GUO Z T, PENG C H. Distribution and storage of soil organic carbon in China[J]. Global Biogeochemical Cycles, 2003, 17(2):2001GB001844 WITZGALL K, VIDAL A, SCHUBERT D I, et al. Particulate organic matter as a functional soil component for persistent soil organic carbon[J]. Nature Communications, 2021, 12:4115 BROOKES P C, LANDMAN A, PRUDEN G, et al. Chloroform fumigation and the release of soil nitrogen: A rapid direct extraction method to measure microbial biomass nitrogen in soil[J]. Soil Biology and Biochemistry, 1985, 17(6):837-842 LI D H, LIU C M, LUO R B, et al. MEGAHIT: an ultra-fast single-node solution for large and complex metagenomics assembly via succinct de Bruijn graph[J]. Bioinformatics, 2015, 31(10):1674-1676 SEEMANN T. Prokka: rapid prokaryotic genome annotation[J]. Bioinformatics, 2014, 30(14):2068-2069 HUERTA-CEPAS J, SZKLARCZYK D, HELLER D, et al. eggNOG 5.0: a hierarchical, functionally and phylogenetically annotated orthology resource based on 5090 organisms and 2502 viruses[J]. Nucleic Acids Research, 2019, 47(D1):D309-D314 KANEHISA M, GOTO S. KEGG Kyoto encyclopedia of genes and genomes[J]. Nucleic Acids Research, 2000, 28(1):27-30 PANG Z Q, ZHOU G Y, EWALD J, et al. Using MetaboAnalyst 5.0 for LC-HRMS spectra processing, multi-omics integration and covariate adjustment of global metabolomics data[J]. Nature Protocols, 2022, 17(8):1735-1761 BOUCHEZ T, BLIEUX A L, DEQUIEDT S, et al. Molecular microbiology methods for environmental diagnosis[J]. Environmental Chemistry Letters, 2016, 14(4):423-441 CSARDI G, NEPUSZ T. The igraph software[J]. Complex syst, 2006, 1695: 1-9 HU P L, LIU S J, YE Y Y, et al. Effects of environmental factors on soil organic carbon under natural or managed vegetation restoration[J]. Land Degradation & Development, 2018, 29(3):387-397 WU J X, LI S Y, HAN G D. Linkage between soil stoichiometric imbalance and microbial carbon use efficiency in desert steppe under different grazing intensities[J]. Chinese Journal of Applied Ecology, 2024, 35(8):2108-2118 吴佳芯, 李邵宇, 韩国栋. 不同放牧强度下荒漠草原土壤化学计量失衡与微生物碳利用效率的联系[J]. 应用生态学报, 2024, 35(8):2108-2118 LI J Y, CHEN P, LI Z G, et al. Soil aggregate-associated organic carbon mineralization and its driving factors in rhizosphere soil[J]. Soil Biology and Biochemistry, 2023, 186:109182 GUO X W, DAI L C, LI Y K, et al. Major greenhouse gas fluxes in different degradation levels of Alpine meadow on the Qinghai-Tibetan Plateau[J]. Research of Soil and Water Conservation, 2019, 26(5):188-194, 209 郭小伟, 戴黎聪, 李以康, 等. 不同退化程度下的高寒草甸主要温室气体通量[J]. 水土保持研究, 2019, 26(5):188-194, 209 PAN T, HOU S, WU S H, et al. Variation of soil hydraulic properties with Alpine grassland degradation in the eastern Tibetan Plateau[J]. Hydrology and Earth System Sciences, 2017, 21(4):2249-2261 GUO M Z, TANG W, ZHAO G Y, et al. Methane concentration dynamics and association with dissolved carbon in different small artificial waterbodies: implications for origin and emission[J]. Water Research, 2026, 288:124659 YANG Y, NIU B, et al. Effects of electron acceptors on CH4 emission in Alpine wetlands[J]. Mires and Peat, 2023, 29:15 ZHANG H, LIU G H, WU J J. The effect of land degradation and restoration on particulate and mineral-associated organic carbon[J]. Applied Soil Ecology, 2024, 196:105322 HU Y, SUN X Y, ZHANG H L, et al. Linking soil organic carbon mineralization to microbial interactions during Alpine grassland degradation[J]. Journal of Environmental Management, 2025, 394:127432 LIU C, LI L, ZHI Y, et al. Molecular insight into the vertical migration and degradation of dissolved organic matter in riparian soil profiles[J]. Environmental Research, 2024, 245:118013 ZHANG R, BAI Y F, ZHANG T, et al. Driving factors that reduce soil carbon, sugar, and microbial biomass in degraded Alpine grasslands[J]. Rangeland Ecology & Management, 2019, 72(2):396-404 LOUCA S, POLZ M F, MAZEL F, et al. Function and functional redundancy in microbial systems[J]. Nature Ecology & Evolution, 2018, 2(6):936-943 HU Y L, FU L C, AO G, et al. Climate, plant and microorganisms jointly influence soil organic matter fractions in temperate grasslands[J]. Science of the Total Environment, 2025, 958:178133 SU N, RONG X M, LIU Y, et al. Achieving sustainable rice production with reduced nitrogen fertilization through unlocking soil organic nitrogen mineralization by alleviating microbial carbon limitation[J]. Field Crops Research, 2026, 340:110382 WU H W, CUI H L, FU C X, et al. Unveiling the crucial role of soil microorganisms in carbon cycling: A review[J]. Science of the Total Environment, 2024, 909:168627 HERNANDEZ D J, DAVID A S, MENGES E S, et al. Environmental stress destabilizes microbial networks[J]. The ISME Journal, 2021, 15(6):1722-1734)
Effects of Grassland Degradation on Soil Carbon Cycle, Microbial Functional Genes and Metabolism
DING Cheng-xiang1, LI Xing-fu2, SU De-rong3, LIU Yu4
Affiliations
1 Qinghai University, Xining, Qinghai Province 810016, China; 2 Institute of Industrial Development Planning, National Forestry and Grassland Administration, Beijing 100010, China; 3 Beijing Forestry University, Beijing 100083, China; 4 Chinese Research Academy of Environmental Sciences, Beijing 100012, China
Under the context of global climate change, the influence mechanism of grassland degradation on soil carbon cycling remain poorly understood. In this study, we investigated 15 paired degraded and non-degraded grassland sites on the Qinghai-Tibet Plateau, integrated microcosm incubation, metagenomics and metabolomics to assess the impacts of degradation on key carbon cycling processes, organic carbon fractions, microbial functional genes, and metabolic intermediates. The results showed that grassland degradation significantly reduced labile carbon pools, with particulate organic carbon, dissolved organic carbon, and microbial biomass carbon decreasing by 48.21%, 30.87%, and 20.21%, respectively. Meanwhile, soil organic carbon mineralization increased by 56.71%, and carbon accumulation efficiency declined by 32.51%. Degradation also led to a decrease in the diversity of carbon cycling functional genes (-2.95%, P<0.05), while the abundance of starch-degrading genes and carbohydrate-related metabolites increased significantly (by 9.39% and 49.6%, respectively). These changes indicated that intensified microbial carbon limitation and a shift in metabolic strategy from synthesis to decoomposition. This study elucidates the key mechanisms underlying soil carbon cycling responses to grassland degradation from a multi-factor coupling perspective, revealing that environmental factors and microbial traits jointly regulated carbon cycling, with microbial functional genes and metabolic intermediates playing a dominat role.
PENG Y F, CHANG J F, ZHAO X, et al. Grassland carbon sink in China and its promotion strategies[J]. Bulletin of National Natural Science Foundation of China, 2023, 37(4):587-602 彭云峰, 常锦峰, 赵霞, 等. 中国草地生态系统固碳能力及其提升途径[J]. 中国科学基金, 2023, 37(4):587-602 BAI Y F, COTRUFO M F. Grassland soil carbon sequestration: current understanding, challenges, and solutions[J]. Science, 2022, 377(6606):603-608 SCHARLEMANN J P, TANNER E V, HIEDERER R, et al. Global soil carbon: understanding and managing the largest terrestrial carbon pool[J]. Carbon Management, 2014, 5(1):81-91 LIU L L, SAYER E J, DENG M F, et al. The grassland carbon cycle: mechanisms, responses to global changes, and potential contribution to carbon neutrality[J]. Fundamental Research, 2023, 3(2):209-218 WU M C, XIE S Y, ZANG J X, et al. Multiple anthropogenic environmental stressors structure soil bacterial diversity and community network[J]. Soil Biology and Biochemistry, 2024, 198:109560 BROWN R W, CHADWICK D R, ZANG H D, et al. Use of metabolomics to quantify changes in soil microbial function in response to fertiliser nitrogen supply and extreme drought[J]. Soil Biology and Biochemistry, 2021, 160:108351 CAO J J, ADAMOWSKI J F, DEO R C, et al. Grassland degradation on the Qinghai-Tibetan Plateau: reevaluation of causative factors[J]. Rangeland Ecology & Management, 2019, 72(6):988-995 LIU S B, ZAMANIAN K, SCHLEUSS P M, et al. Degradation of Tibetan grasslands: consequences for carbon and nutrient cycles[J]. Agriculture, Ecosystems & Environment, 2018, 252:93-104 HU W J, CAI Y, LI X Q, et al. Microbial carbon accumulation efficiency in global soils resolved via13C-glucose amendment experiments[J]. Global Biogeochemical Cycles, 2025, 39(6):e2025GB008553 ABDALLA K, MUTEMA M, CHIVENGE P, et al. Grassland degradation significantly enhances soil CO2 emission[J]. Catena, 2018, 167:284-292 WANG P Y, WANG J S, WANG S, et al. Soil moisture threshold of methane uptake in Alpine grassland ecosystems[J]. Global Change Biology, 2025, 31(2):e70062 ZHOU J C, LIU S, VAN GROENIGEN K JAN, et al. Contrasting responses of particulate and mineral-associated organic carbon stocks to grazing exclusion in an Alpine meadow[J]. Agriculture, Ecosystems & Environment, 2026, 400:110227 HUANG Q, WANG B R, SHEN J K, et al. Shifts in C-degradation genes and microbial metabolic activity with vegetation types affected the surface soil organic carbon pool[J]. Soil Biology and Biochemistry, 2024, 192:109371 WANG N N, LI K X, YUAN F H, et al. Faster cycling but lower efficiency: A microbial metabolic perspective on carbon loss after wetland conversion to cropland[J]. Soil Biology and Biochemistry, 2024, 189:109260 HAO A H, XUE X, PENG F, et al. Different vegetation and soil degradation characteristics of a typical grassland in the Qinghai-Tibetan Plateau[J]. Acta Ecologica Sinica, 2020, 40(3):964-975 郝爱华, 薛娴, 彭飞, 等. 青藏高原典型草地植被退化与土壤退化研究[J]. 生态学报, 2020, 40(3):964-975 ZHANG S N, LI F C, LI J B, et al. The influence of different degrees of degradation on soil dissolved organic carbon and its characteristics in sloping grassland[J]. Acta Agrestia Sinica, 2025, 33(9):2941-2949 张胜男, 李富翠, 李金波, 等. 不同退化程度对坡地草地土壤可溶性有机碳及其特征的影响[J]. 草地学报, 2025, 33(9):2941-2949 BAO S D. Soil and agricultural chemistry analysis [M].3rd Edition.Beijing:China Agriculture Press, 2000:42-80 鲍士旦.土壤农化分析[M]. 3版. 北京:中国农业出版社, 2000:42-80 BRIDGHAM S D, YE R. Organic matter mineralization and decomposition[J]. Methods in biogeochemistry of wetlands. 2013, 10: 385-406 MA L, JIANG X L, LIU G H, et al. Environmental factors and microbial diversity and abundance jointly regulate soil nitrogen and carbon biogeochemical processes in Tibetan wetlands[J]. Environmental Science & Technology, 2020, 54(6):3267-3277 TAŞ N, PRESTAT E, MCFARLAND J W, et al. Impact of fire on active layer and permafrost microbial communities and metagenomes in an upland Alaskan boreal forest[J]. The ISME Journal, 2014, 8(9):1904-1919 WU H B, GUO Z T, PENG C H. Distribution and storage of soil organic carbon in China[J]. Global Biogeochemical Cycles, 2003, 17(2):2001GB001844 WITZGALL K, VIDAL A, SCHUBERT D I, et al. Particulate organic matter as a functional soil component for persistent soil organic carbon[J]. Nature Communications, 2021, 12:4115 BROOKES P C, LANDMAN A, PRUDEN G, et al. Chloroform fumigation and the release of soil nitrogen: A rapid direct extraction method to measure microbial biomass nitrogen in soil[J]. Soil Biology and Biochemistry, 1985, 17(6):837-842 LI D H, LIU C M, LUO R B, et al. MEGAHIT: an ultra-fast single-node solution for large and complex metagenomics assembly via succinct de Bruijn graph[J]. Bioinformatics, 2015, 31(10):1674-1676 SEEMANN T. Prokka: rapid prokaryotic genome annotation[J]. Bioinformatics, 2014, 30(14):2068-2069 HUERTA-CEPAS J, SZKLARCZYK D, HELLER D, et al. eggNOG 5.0: a hierarchical, functionally and phylogenetically annotated orthology resource based on 5090 organisms and 2502 viruses[J]. Nucleic Acids Research, 2019, 47(D1):D309-D314 KANEHISA M, GOTO S. KEGG Kyoto encyclopedia of genes and genomes[J]. Nucleic Acids Research, 2000, 28(1):27-30 PANG Z Q, ZHOU G Y, EWALD J, et al. Using MetaboAnalyst 5.0 for LC-HRMS spectra processing, multi-omics integration and covariate adjustment of global metabolomics data[J]. Nature Protocols, 2022, 17(8):1735-1761 BOUCHEZ T, BLIEUX A L, DEQUIEDT S, et al. Molecular microbiology methods for environmental diagnosis[J]. Environmental Chemistry Letters, 2016, 14(4):423-441 CSARDI G, NEPUSZ T. The igraph software[J]. Complex syst, 2006, 1695: 1-9 HU P L, LIU S J, YE Y Y, et al. Effects of environmental factors on soil organic carbon under natural or managed vegetation restoration[J]. Land Degradation & Development, 2018, 29(3):387-397 WU J X, LI S Y, HAN G D. Linkage between soil stoichiometric imbalance and microbial carbon use efficiency in desert steppe under different grazing intensities[J]. Chinese Journal of Applied Ecology, 2024, 35(8):2108-2118 吴佳芯, 李邵宇, 韩国栋. 不同放牧强度下荒漠草原土壤化学计量失衡与微生物碳利用效率的联系[J]. 应用生态学报, 2024, 35(8):2108-2118 LI J Y, CHEN P, LI Z G, et al. Soil aggregate-associated organic carbon mineralization and its driving factors in rhizosphere soil[J]. Soil Biology and Biochemistry, 2023, 186:109182 GUO X W, DAI L C, LI Y K, et al. Major greenhouse gas fluxes in different degradation levels of Alpine meadow on the Qinghai-Tibetan Plateau[J]. Research of Soil and Water Conservation, 2019, 26(5):188-194, 209 郭小伟, 戴黎聪, 李以康, 等. 不同退化程度下的高寒草甸主要温室气体通量[J]. 水土保持研究, 2019, 26(5):188-194, 209 PAN T, HOU S, WU S H, et al. Variation of soil hydraulic properties with Alpine grassland degradation in the eastern Tibetan Plateau[J]. Hydrology and Earth System Sciences, 2017, 21(4):2249-2261 GUO M Z, TANG W, ZHAO G Y, et al. Methane concentration dynamics and association with dissolved carbon in different small artificial waterbodies: implications for origin and emission[J]. Water Research, 2026, 288:124659 YANG Y, NIU B, et al. Effects of electron acceptors on CH4 emission in Alpine wetlands[J]. Mires and Peat, 2023, 29:15 ZHANG H, LIU G H, WU J J. The effect of land degradation and restoration on particulate and mineral-associated organic carbon[J]. Applied Soil Ecology, 2024, 196:105322 HU Y, SUN X Y, ZHANG H L, et al. Linking soil organic carbon mineralization to microbial interactions during Alpine grassland degradation[J]. Journal of Environmental Management, 2025, 394:127432 LIU C, LI L, ZHI Y, et al. Molecular insight into the vertical migration and degradation of dissolved organic matter in riparian soil profiles[J]. Environmental Research, 2024, 245:118013 ZHANG R, BAI Y F, ZHANG T, et al. Driving factors that reduce soil carbon, sugar, and microbial biomass in degraded Alpine grasslands[J]. Rangeland Ecology & Management, 2019, 72(2):396-404 LOUCA S, POLZ M F, MAZEL F, et al. Function and functional redundancy in microbial systems[J]. Nature Ecology & Evolution, 2018, 2(6):936-943 HU Y L, FU L C, AO G, et al. Climate, plant and microorganisms jointly influence soil organic matter fractions in temperate grasslands[J]. Science of the Total Environment, 2025, 958:178133 SU N, RONG X M, LIU Y, et al. Achieving sustainable rice production with reduced nitrogen fertilization through unlocking soil organic nitrogen mineralization by alleviating microbial carbon limitation[J]. Field Crops Research, 2026, 340:110382 WU H W, CUI H L, FU C X, et al. Unveiling the crucial role of soil microorganisms in carbon cycling: A review[J]. Science of the Total Environment, 2024, 909:168627 HERNANDEZ D J, DAVID A S, MENGES E S, et al. Environmental stress destabilizes microbial networks[J]. The ISME Journal, 2021, 15(6):1722-1734