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By examining the ecological stoichiometry of soil-microbe-extracellular enzyme systems under nitrogen and phosphorus addition, this research explored the characteristics of soil nutrient limitations and the feedback mechanisms of microbial communities to metabolic constraints. The results showed that fertilization significantly decreased soil carbon to phosphorus ratio, microbial biomass carbon to phosphorus ratio (MBC∶MBP), microbial biomass nitrogen to phosphorus ratio (MBN∶MBP), and enzymatic stoichiometric carbon to nitrogen ratio (EEAC∶N). In contrast, soil nutrient content, microbial biomass, extracellular enzyme activities, and enzymatic stoichiometric nitrogen to phosphorus ratio (EEAN∶P) were significantly increased. Enzyme vector modeling and vector analysis indicated that long-term alfalfa cultivation led to varying degrees of carbon and phosphorus limitation in soil microorganisms. Nitrogen and phosphorus addition alleviated the microbial phosphorus limitation and increased microbial carbon use efficiency (CUE). In conclusion, appropriate supplementation of nitrogen and phosphorus fertilizers is recommended for soils under long-term alfalfa cultivation in the Loess hill region, to mitigate nutrient limitations on soil microorganisms., authors=ZHU Wen-zhi1, LUO Zhu-zhu1,2, LI Liang-liang3, HE Ren-yuan4, LIU Jia-he1, authorsList=ZHU Wen-zhi, LUO Zhu-zhu, LI Liang-liang, HE Ren-yuan, LIU Jia-he, authorCompany=1 College of Resources and Environment, Gansu Agricultural University, Lanzhou, Gansu Province 730070, China; 2 State Key Laboratory of Arid Habitat Crop Science, Lanzhou, Gansu Province 730070, China; 3 College of Pratacultural Science, Gansu Agricultural University, Lanzhou, Gansu Province 730070, China; 4 College of Forestry, Gansu Agricultural University, Lanzhou, Gansu Province 730070, 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=1309575112588878346, articleId=1309575110885990921, tenantId=1146029695717560320, journalId=1308397196074385509, language=CN, title=氮磷添加缓解了紫花苜蓿土壤微生物养分限制, columnId=1309565024058835950, journalTitle=草地学报, columnName=研究论文, runingTitle=null, highlight=null, articleAbstract=本研究以黄土丘陵区紫花苜蓿地为研究对象,设置不施肥(CK)、单施氮肥(T1)、单施磷肥(T2)和氮磷肥配施(T3)4个处理,通过分析氮磷添加下紫花苜蓿土壤-微生物-胞外酶生态化学计量特征,探讨土壤养分限制特征及微生物对代谢限制的反馈机制。结果表明,施肥降低了土壤碳磷比、微生物量碳磷比(MBC∶MBP)、微生物量氮磷比(MBN∶MBP)和酶化学计量碳氮比(EEAC∶N),但增加了土壤养分含量、微生物量、胞外酶活性和酶化学计量氮磷比(EEAN∶P)。酶向量模型和矢量分析结果表明,长期种植苜蓿土壤微生物受到不同程度的碳限制和磷限制,氮磷添加缓解了土壤微生物磷限制,提高了微生物碳利用效率(CUE)。综上,黄土丘陵区长期种植苜蓿土壤应适当补施氮磷肥,以缓解土壤微生物的养分限制。, authors=祝文知1, 罗珠珠1,2, 李亮亮3, 何仁元4, 刘家鹤1, authorsList=祝文知, 罗珠珠, 李亮亮, 何仁元, 刘家鹤, authorCompany=1 甘肃农业大学资源与环境学院, 甘肃 兰州 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detailUrlEn=https://castjournals.cast.org.cn/joweb/cdxb/EN/10.11733/j.issn.1007-0435.2026.06.013, pdfUrlCn=https://castjournals.cast.org.cn/joweb/cdxb/CN/PDF/10.11733/j.issn.1007-0435.2026.06.013, pdfUrlEn=https://castjournals.cast.org.cn/joweb/cdxb/EN/PDF/10.11733/j.issn.1007-0435.2026.06.013, aliStartDate=null, aliEndDate=null, collectionFlag=false, citedCount=null, citedUrl=null, previewStatus=0, delFlag=0, hasFullText=0, orderTime=1781452800000, fullTextJson=null, articleText=null, reference=XIAO X Q,ZHANG H K,FENG Y S,et al. Effects of plant residues on C: N: P of soil,microbial biomass,and extracellular enzyme in an Alpine Mea-dow on the Qinghai-Tibetan Plateau,China[J]. Chinese Journal of Applied Ecology,2023,34(1):58-66 肖向前,张海阔,冯娅斯,等. 植物残体对青藏高原高寒草甸土壤、微生物和胞外酶C∶N∶P化学计量特征的影响[J]. 应用生态学报,2023,34(1):58-66 JIA Y H,HU W Y,DENG J,et al. Effects of nitrogen addition on soil microbial nutrient limitation characteristics in grassland in the Loess Hilly Region[J]. Acta Prataculturae Sinica,2025,34(2):221-232 贾蕴欢,胡雯颖,邓健,等. 氮添加对黄土丘陵区草地土壤微生物养分限制特征的影响[J]. 草业学报,2025,34(2):221-232 SINSABAUGH R L,HILL B H,FOLLSTAD SHAH J J. Ecoenzymatic stoichiometry of microbial organic nutrient acquisition in soil and sediment[J]. Nature,2009,462(7274):795-798 YANG R Z,YANG Z,YANG S L,et al. Nitrogen inhibitors improve soil ecosystem multifunctionality by enhancing soil quality and alleviating microbial nitrogen limitation[J]. Science of the Total Environment,2023,880:163238 YIN Q J,JIANG J W,YIN H Q,et al. Effects of organic inputs on soil nutrients and microbial metabolism in newly reclaimed farmlands[J]. Chinese Journal of Applied Ecology,2025,36(4):969-983 殷启杰,姜建武,殷汉琴,等. 有机物料投入对新垦耕地土壤养分和微生物代谢的影响[J]. 应用生态学报,2025,36(4):969-983 ZHU X M,LIU M,KOU Y P,et al. Differential effects of N addition on the stoichiometry of microbes and extracellular enzymes in the rhizosphere and bulk soils of an Alpine shrubland[J]. Plant and Soil,2020,449(1):285-301 TIAN L X,ZHANG Y C,GAO X L,et al. Microbial carbon and nitrogen limitations with mulching of proso millet fields on the Loess Plateau: evidence from soil ecoenzymatic stoichiometry[J]. Land Degradation & Development,2023,34(17):5224-5237 SONG J S,ZHANG H Y,CHANG F D,et al. Humic acid plus manure increases the soil carbon pool by inhibiting salinity and alleviating the microbial resource limitation in saline soils[J]. Catena,2023,233:107527 WIDDIG M,SCHLEUSS P M,BIEDERMAN L A,et al. Microbial carbon use efficiency in grassland soils subjected to nitrogen and phosphorus additions[J]. Soil Biology and Biochemistry,2020,146:107815 YU Z K,JING Y D,LIU L,et al. Changes of soil,microorganisms,enzymes and stoichiometry during the uncultivature of alfalfa farmland[J]. Acta Agrestia Sinica,2023,31(4):1115-1124 于宗恺,景煜都,刘励,等. 苜蓿地撂荒过程中土壤-微生物-胞外酶及化学计量的变化[J]. 草地学报,2023,31(4):1115-1124 WANG S Q,LUO Z Z,NIU Y N,et al. Soil quality evaluation of alfalfa fields in semi-arid areas of the Loess Plateau based on minimum data set[J]. Chinese Journal of Grassland,2023,45(7):81-90 王淑琴,罗珠珠,牛伊宁,等. 基于最小数据集的黄土高原半干旱区苜蓿地土壤质量评价[J]. 中国草地学报,2023,45(7):81-90 BAO S D. Soil Agricultural Chemistry Analysis [M]. 3rd Edition. Beijing: China Agriculture Press, 2000: 25-97 鲍士旦. 土壤农化分析[M]. 3版. 北京:中国农业出版社,2000:25-97 SUN Y J,ZHANG D N,SHEN Y Y,et al. Effects of simulated nitrogen deposition on soil microbial community structure and enzyme activities in Eucalyptus plantations in mid-subtropical region[J]. Scientia Silvae Sinicae,2025,61(5):46-60 孙英杰,张德楠,沈育伊,等. 模拟氮沉降对中亚热带桉树人工林土壤微生物群落结构及酶活性的影响[J]. 林业科学,2025,61(5):46-60 SINSABAUGH R L,TURNER B L,TALBOT J M,et al. Stoichiometry of microbial carbon use efficiency in soils[J]. Ecological Monographs,2016,86(2):172-189 YANG Y,LIU H,YANG X,et al. Plant and soil elemental C: N: P ratios are linked to soil microbial diversity during grassland restoration on the Loess Plateau,China[J]. Science of the Total Environment,2022,806(Pt 1):150557 SINSABAUGH R L,LAUBER C L,WEINTRAUB M N,et al. Stoichiometry of soil enzyme activity at global scale[J]. Ecology Letters,2008,11(11):1252-1264 MEN X X,BAO Y,ZHAI D P,et al. Soil microbial metabolic limitations under subalpine coniferous and broad–leaved forests responding to litter input and removal treatments[J]. Functional Ecology,2025,39(10):2803-2816 LIU S,ZHANG X Y,WANG H M,et al. Phosphorus-transforming microbes enhance phosphatase catalytic efficiency to alleviate phosphorus limitation under nitrogen and phosphorus additions in subtropical forest soil[J]. Soil Biology and Biochemistry,2025,209:109915 LE J J,SU Y,PENG Q W,et al. Effects of nitrogen addition on soil enzyme activities and ecoenzymatic stoichiometry in Alpine grassland of the Tianshan Mountains[J]. Arid Zone Research,2020,37(2):382-389 勒佳佳,苏原,彭庆文,等. 氮添加对天山高寒草原土壤酶活性和酶化学计量特征的影响[J]. 干旱区研究,2020,37(2):382-389 BOOT C M,HALL E K,DENEF K,et al. Long-term reactive nitrogen loading alters soil carbon and microbial community properties in a subalpine forest ecosystem[J]. Soil Biology and Biochemistry,2016,92:211-220 LIAN J L,CHEN J,CUI H,et al. Soil extracellular enzyme stoichiometry reveals the nutrient limitations of soil microbial metabolism under precipitation changes in Ningxia desert steppe of China[J]. European Journal of Soil Biology,2025,127:103774 MA W J,LI J,GAO Y,et al. Responses of soil extracellular enzyme activities and microbial community properties to interaction between nitrogen addition and increased precipitation in a semi-arid grassland ecosystem[J]. Science of the Total Environment,2020,703:134691 A’BEAR A D,JONES T H,KANDELER E,et al. Interactive effects of temperature and soil moisture on fungal-mediated wood decomposition and extracellular enzyme activity[J]. Soil Biology and Biochemistry,2014,70:151-158 YU Q S,NI X F,HAGEDORN F,et al. Field experiments and a meta-analysis reveal a minor influence of nitrogen addition on phosphorus fractions in forests[J]. Global Change Biology,2025,31(4):e70156 FAN Z Z,WANG X,WANG C,et al. Effect of nitrogen and phosphorus addition on soil enzyme activities: A meta-analysis[J]. Chinese Journal of Applied Ecology,2018,29(4):1266-1272 范珍珍,王鑫,王超,等. 整合分析氮磷添加对土壤酶活性的影响[J]. 应用生态学报,2018,29(4):1266-1272 ZHANG J X,LIU X J,HAO F,et al. Effects of nitrogen and phosphorus regulation on alfalfa nitrogen accumulation and the nitrogen and phosphorus nutrients of soil[J]. Acta Agrestia Sinica,2016,24(1):61-68 张进霞,刘晓静,郝凤,等. 氮磷调控对紫花苜蓿氮积累与土壤氮磷营养的影响[J]. 草地学报,2016,24(1):61-68 QIN W K,FENG J G,ZHANG Q F,et al. Nitrogen and phosphorus addition mediate soil priming effects via affecting microbial stoichiometric balance in an Alpine meadow[J]. Science of the Total Environment,2024,908:168350 LI Y N,LI S Y,SHI S B,et al. Study on soil enzyme stoichiometric characteristics of different grazing intensities in desert steppe[J]. Acta Agrestia Sinica,2022,30(9):2239-2248 李雅男,李邵宇,史世斌,等. 荒漠草原不同放牧强度下土壤酶化学计量特征的研究[J]. 草地学报,2022,30(9):2239-2248 WANG C,YANG Q N,ZHANG C,et al. Vegetation restoration of abandoned cropland improves soil ecosystem multifunctionality through alleviating nitrogen-limitation in the China Danxia[J]. Frontiers in Plant Science,2023,14:1116179 ROSINGER C,ROUSK J,SANDÉN H. Can enzymatic stoichiometry be used to determine growth-limiting nutrients for microorganisms?A critical assessment in two subtropical soils[J]. Soil Biology and Biochemistry,2019,128:115-126 YIN H B,HUANG Q Y,SHA G,et al. Response of soil extracellular enzyme activity and stoichiometric characteristics to nitrogen deposition in new volcanic lava platform[J]. Acta Ecologica Sinica,2025,45(13):6292-6303 阴红彬,黄庆阳,沙刚,等. 新期火山熔岩台地土壤胞外酶活性及化学计量特征对氮沉降的响应[J]. 生态学报,2025,45(13):6292-6303 ÅGREN G I,WETTERSTEDT J A M,BILLBERGER M F K. Nutrient limitation on terrestrial plant growth-modeling the interaction between nitrogen and phosphorus[J]. New Phytologist,2012,194(4):953-960 PANG C L,ZHANG Z H,ZHU X M,et al. Divergent microbial metabolic limitations across soil depths after two decades of high nitrogen inputs in a primary tropical forest[J]. Global Change Biology,2025,31(8):e70440 WU M Y,CHEN L,CHEN S G,et al. Soil microbial carbon and nitrogen limitation constraints soil organic carbon stability in arid and semi-arid grasslands[J]. Journal of Environmental Management,2025,373:123675 SUN Y N,LI Q,LI Y K,et al. The effect of nitrogen and phosphorus applications on soil enzyme activities in Qinghai-Tibetan Apine meadows[J]. Acta Prataculturae Sinica,2016,25(2):18-26 孙亚男,李茜,李以康,等. 氮、磷养分添加对高寒草甸土壤酶活性的影响[J]. 草业学报,2016,25(2):18-26 SCHLEUSS P M,WIDDIG M,HEINTZ-USCHART A,et al. Interactions of nitrogen and phosphorus cycling promote P acquisition and explain synergistic plant-growth responses[J]. Ecology,2020,101(5):e03003 YU X C,ZHU H Y. Enacting partner specificity in legume–rhizobia symbioses[J]. aBIOTECH,2025,6(2):311-327 ZHANG J Y,YANG X M,SONG Y H,et al. Revealing the nutrient limitation and cycling for microbes under forest management practices in the Loess Plateau-Ecological stoichiometry[J]. Geoderma,2020,361:114108 XU M P,LI W J,WANG J Y,et al. Soil ecoenzymatic stoichiometry reveals microbial phosphorus limitation after vegetation restoration on the Loess Plateau,China[J]. Science of the Total Environment,2022,815:152918 XIE W J,ZHANG Y P,LI J Y,et al. Straw application coupled with N and P supply enhanced microbial biomass,enzymatic activity,and carbon use efficiency in saline soil[J]. Applied Soil Ecology,2021,168:104128 GU Y J,HAN C L,FAN J W,et al. Alfalfa forage yield,soil water and P availability in response to plastic film mulch and P fertilization in a semiarid environment[J]. Field Crops Research,2018,215:94-103 AN R,YU R P,XING Y,et al. Intercropping efficiently utilizes phosphorus resource in soil via different strategies mediated by crop traits and species combination[J]. Plant and Soil,2023,497(1-2):705-725 YANG L Y,CANARINI A,ZHANG W S,et al. Microbial life-history strategies mediate microbial carbon pump efficacy in response to N management depending on stoichiometry of microbial demand[J]. Global Change Biology,2024,30(5):e17311)
Nitrogen and Phosphorus Addition Alleviated Soil Microbial Nutrient Limitation in Alfalfa Fields
ZHU Wen-zhi, LUO Zhu-zhu, LI Liang-liang, HE Ren-yuan, LIU Jia-he
Affiliations
1 College of Resources and Environment, Gansu Agricultural University, Lanzhou, Gansu Province 730070, China; 2 State Key Laboratory of Arid Habitat Crop Science, Lanzhou, Gansu Province 730070, China; 3 College of Pratacultural Science, Gansu Agricultural University, Lanzhou, Gansu Province 730070, China; 4 College of Forestry, Gansu Agricultural University, Lanzhou, Gansu Province 730070, China
This study focued on alfalfa fields in the Loess hilly region as research objects, with four experimental treatments were established: no fertilization (CK), nitrogen fertilizer alone (T1), phosphorus fertilizer alone (T2), and combined nitrogen-phosphorus fertilization (T3). By examining the ecological stoichiometry of soil-microbe-extracellular enzyme systems under nitrogen and phosphorus addition, this research explored the characteristics of soil nutrient limitations and the feedback mechanisms of microbial communities to metabolic constraints. The results showed that fertilization significantly decreased soil carbon to phosphorus ratio, microbial biomass carbon to phosphorus ratio (MBC∶MBP), microbial biomass nitrogen to phosphorus ratio (MBN∶MBP), and enzymatic stoichiometric carbon to nitrogen ratio (EEAC∶N). In contrast, soil nutrient content, microbial biomass, extracellular enzyme activities, and enzymatic stoichiometric nitrogen to phosphorus ratio (EEAN∶P) were significantly increased. Enzyme vector modeling and vector analysis indicated that long-term alfalfa cultivation led to varying degrees of carbon and phosphorus limitation in soil microorganisms. Nitrogen and phosphorus addition alleviated the microbial phosphorus limitation and increased microbial carbon use efficiency (CUE). In conclusion, appropriate supplementation of nitrogen and phosphorus fertilizers is recommended for soils under long-term alfalfa cultivation in the Loess hill region, to mitigate nutrient limitations on soil microorganisms.
XIAO X Q,ZHANG H K,FENG Y S,et al. Effects of plant residues on C: N: P of soil,microbial biomass,and extracellular enzyme in an Alpine Mea-dow on the Qinghai-Tibetan Plateau,China[J]. Chinese Journal of Applied Ecology,2023,34(1):58-66 肖向前,张海阔,冯娅斯,等. 植物残体对青藏高原高寒草甸土壤、微生物和胞外酶C∶N∶P化学计量特征的影响[J]. 应用生态学报,2023,34(1):58-66 JIA Y H,HU W Y,DENG J,et al. Effects of nitrogen addition on soil microbial nutrient limitation characteristics in grassland in the Loess Hilly Region[J]. Acta Prataculturae Sinica,2025,34(2):221-232 贾蕴欢,胡雯颖,邓健,等. 氮添加对黄土丘陵区草地土壤微生物养分限制特征的影响[J]. 草业学报,2025,34(2):221-232 SINSABAUGH R L,HILL B H,FOLLSTAD SHAH J J. Ecoenzymatic stoichiometry of microbial organic nutrient acquisition in soil and sediment[J]. Nature,2009,462(7274):795-798 YANG R Z,YANG Z,YANG S L,et al. Nitrogen inhibitors improve soil ecosystem multifunctionality by enhancing soil quality and alleviating microbial nitrogen limitation[J]. Science of the Total Environment,2023,880:163238 YIN Q J,JIANG J W,YIN H Q,et al. Effects of organic inputs on soil nutrients and microbial metabolism in newly reclaimed farmlands[J]. Chinese Journal of Applied Ecology,2025,36(4):969-983 殷启杰,姜建武,殷汉琴,等. 有机物料投入对新垦耕地土壤养分和微生物代谢的影响[J]. 应用生态学报,2025,36(4):969-983 ZHU X M,LIU M,KOU Y P,et al. Differential effects of N addition on the stoichiometry of microbes and extracellular enzymes in the rhizosphere and bulk soils of an Alpine shrubland[J]. Plant and Soil,2020,449(1):285-301 TIAN L X,ZHANG Y C,GAO X L,et al. Microbial carbon and nitrogen limitations with mulching of proso millet fields on the Loess Plateau: evidence from soil ecoenzymatic stoichiometry[J]. Land Degradation & Development,2023,34(17):5224-5237 SONG J S,ZHANG H Y,CHANG F D,et al. Humic acid plus manure increases the soil carbon pool by inhibiting salinity and alleviating the microbial resource limitation in saline soils[J]. Catena,2023,233:107527 WIDDIG M,SCHLEUSS P M,BIEDERMAN L A,et al. Microbial carbon use efficiency in grassland soils subjected to nitrogen and phosphorus additions[J]. Soil Biology and Biochemistry,2020,146:107815 YU Z K,JING Y D,LIU L,et al. Changes of soil,microorganisms,enzymes and stoichiometry during the uncultivature of alfalfa farmland[J]. Acta Agrestia Sinica,2023,31(4):1115-1124 于宗恺,景煜都,刘励,等. 苜蓿地撂荒过程中土壤-微生物-胞外酶及化学计量的变化[J]. 草地学报,2023,31(4):1115-1124 WANG S Q,LUO Z Z,NIU Y N,et al. Soil quality evaluation of alfalfa fields in semi-arid areas of the Loess Plateau based on minimum data set[J]. Chinese Journal of Grassland,2023,45(7):81-90 王淑琴,罗珠珠,牛伊宁,等. 基于最小数据集的黄土高原半干旱区苜蓿地土壤质量评价[J]. 中国草地学报,2023,45(7):81-90 BAO S D. Soil Agricultural Chemistry Analysis [M]. 3rd Edition. Beijing: China Agriculture Press, 2000: 25-97 鲍士旦. 土壤农化分析[M]. 3版. 北京:中国农业出版社,2000:25-97 SUN Y J,ZHANG D N,SHEN Y Y,et al. Effects of simulated nitrogen deposition on soil microbial community structure and enzyme activities in Eucalyptus plantations in mid-subtropical region[J]. Scientia Silvae Sinicae,2025,61(5):46-60 孙英杰,张德楠,沈育伊,等. 模拟氮沉降对中亚热带桉树人工林土壤微生物群落结构及酶活性的影响[J]. 林业科学,2025,61(5):46-60 SINSABAUGH R L,TURNER B L,TALBOT J M,et al. Stoichiometry of microbial carbon use efficiency in soils[J]. Ecological Monographs,2016,86(2):172-189 YANG Y,LIU H,YANG X,et al. Plant and soil elemental C: N: P ratios are linked to soil microbial diversity during grassland restoration on the Loess Plateau,China[J]. Science of the Total Environment,2022,806(Pt 1):150557 SINSABAUGH R L,LAUBER C L,WEINTRAUB M N,et al. Stoichiometry of soil enzyme activity at global scale[J]. Ecology Letters,2008,11(11):1252-1264 MEN X X,BAO Y,ZHAI D P,et al. Soil microbial metabolic limitations under subalpine coniferous and broad–leaved forests responding to litter input and removal treatments[J]. Functional Ecology,2025,39(10):2803-2816 LIU S,ZHANG X Y,WANG H M,et al. Phosphorus-transforming microbes enhance phosphatase catalytic efficiency to alleviate phosphorus limitation under nitrogen and phosphorus additions in subtropical forest soil[J]. Soil Biology and Biochemistry,2025,209:109915 LE J J,SU Y,PENG Q W,et al. Effects of nitrogen addition on soil enzyme activities and ecoenzymatic stoichiometry in Alpine grassland of the Tianshan Mountains[J]. Arid Zone Research,2020,37(2):382-389 勒佳佳,苏原,彭庆文,等. 氮添加对天山高寒草原土壤酶活性和酶化学计量特征的影响[J]. 干旱区研究,2020,37(2):382-389 BOOT C M,HALL E K,DENEF K,et al. Long-term reactive nitrogen loading alters soil carbon and microbial community properties in a subalpine forest ecosystem[J]. Soil Biology and Biochemistry,2016,92:211-220 LIAN J L,CHEN J,CUI H,et al. Soil extracellular enzyme stoichiometry reveals the nutrient limitations of soil microbial metabolism under precipitation changes in Ningxia desert steppe of China[J]. European Journal of Soil Biology,2025,127:103774 MA W J,LI J,GAO Y,et al. Responses of soil extracellular enzyme activities and microbial community properties to interaction between nitrogen addition and increased precipitation in a semi-arid grassland ecosystem[J]. Science of the Total Environment,2020,703:134691 A’BEAR A D,JONES T H,KANDELER E,et al. Interactive effects of temperature and soil moisture on fungal-mediated wood decomposition and extracellular enzyme activity[J]. Soil Biology and Biochemistry,2014,70:151-158 YU Q S,NI X F,HAGEDORN F,et al. Field experiments and a meta-analysis reveal a minor influence of nitrogen addition on phosphorus fractions in forests[J]. Global Change Biology,2025,31(4):e70156 FAN Z Z,WANG X,WANG C,et al. Effect of nitrogen and phosphorus addition on soil enzyme activities: A meta-analysis[J]. Chinese Journal of Applied Ecology,2018,29(4):1266-1272 范珍珍,王鑫,王超,等. 整合分析氮磷添加对土壤酶活性的影响[J]. 应用生态学报,2018,29(4):1266-1272 ZHANG J X,LIU X J,HAO F,et al. Effects of nitrogen and phosphorus regulation on alfalfa nitrogen accumulation and the nitrogen and phosphorus nutrients of soil[J]. Acta Agrestia Sinica,2016,24(1):61-68 张进霞,刘晓静,郝凤,等. 氮磷调控对紫花苜蓿氮积累与土壤氮磷营养的影响[J]. 草地学报,2016,24(1):61-68 QIN W K,FENG J G,ZHANG Q F,et al. Nitrogen and phosphorus addition mediate soil priming effects via affecting microbial stoichiometric balance in an Alpine meadow[J]. Science of the Total Environment,2024,908:168350 LI Y N,LI S Y,SHI S B,et al. Study on soil enzyme stoichiometric characteristics of different grazing intensities in desert steppe[J]. Acta Agrestia Sinica,2022,30(9):2239-2248 李雅男,李邵宇,史世斌,等. 荒漠草原不同放牧强度下土壤酶化学计量特征的研究[J]. 草地学报,2022,30(9):2239-2248 WANG C,YANG Q N,ZHANG C,et al. Vegetation restoration of abandoned cropland improves soil ecosystem multifunctionality through alleviating nitrogen-limitation in the China Danxia[J]. Frontiers in Plant Science,2023,14:1116179 ROSINGER C,ROUSK J,SANDÉN H. Can enzymatic stoichiometry be used to determine growth-limiting nutrients for microorganisms?A critical assessment in two subtropical soils[J]. Soil Biology and Biochemistry,2019,128:115-126 YIN H B,HUANG Q Y,SHA G,et al. Response of soil extracellular enzyme activity and stoichiometric characteristics to nitrogen deposition in new volcanic lava platform[J]. Acta Ecologica Sinica,2025,45(13):6292-6303 阴红彬,黄庆阳,沙刚,等. 新期火山熔岩台地土壤胞外酶活性及化学计量特征对氮沉降的响应[J]. 生态学报,2025,45(13):6292-6303 ÅGREN G I,WETTERSTEDT J A M,BILLBERGER M F K. Nutrient limitation on terrestrial plant growth-modeling the interaction between nitrogen and phosphorus[J]. New Phytologist,2012,194(4):953-960 PANG C L,ZHANG Z H,ZHU X M,et al. Divergent microbial metabolic limitations across soil depths after two decades of high nitrogen inputs in a primary tropical forest[J]. Global Change Biology,2025,31(8):e70440 WU M Y,CHEN L,CHEN S G,et al. Soil microbial carbon and nitrogen limitation constraints soil organic carbon stability in arid and semi-arid grasslands[J]. Journal of Environmental Management,2025,373:123675 SUN Y N,LI Q,LI Y K,et al. The effect of nitrogen and phosphorus applications on soil enzyme activities in Qinghai-Tibetan Apine meadows[J]. Acta Prataculturae Sinica,2016,25(2):18-26 孙亚男,李茜,李以康,等. 氮、磷养分添加对高寒草甸土壤酶活性的影响[J]. 草业学报,2016,25(2):18-26 SCHLEUSS P M,WIDDIG M,HEINTZ-USCHART A,et al. Interactions of nitrogen and phosphorus cycling promote P acquisition and explain synergistic plant-growth responses[J]. Ecology,2020,101(5):e03003 YU X C,ZHU H Y. Enacting partner specificity in legume–rhizobia symbioses[J]. aBIOTECH,2025,6(2):311-327 ZHANG J Y,YANG X M,SONG Y H,et al. Revealing the nutrient limitation and cycling for microbes under forest management practices in the Loess Plateau-Ecological stoichiometry[J]. Geoderma,2020,361:114108 XU M P,LI W J,WANG J Y,et al. Soil ecoenzymatic stoichiometry reveals microbial phosphorus limitation after vegetation restoration on the Loess Plateau,China[J]. Science of the Total Environment,2022,815:152918 XIE W J,ZHANG Y P,LI J Y,et al. Straw application coupled with N and P supply enhanced microbial biomass,enzymatic activity,and carbon use efficiency in saline soil[J]. Applied Soil Ecology,2021,168:104128 GU Y J,HAN C L,FAN J W,et al. Alfalfa forage yield,soil water and P availability in response to plastic film mulch and P fertilization in a semiarid environment[J]. Field Crops Research,2018,215:94-103 AN R,YU R P,XING Y,et al. Intercropping efficiently utilizes phosphorus resource in soil via different strategies mediated by crop traits and species combination[J]. Plant and Soil,2023,497(1-2):705-725 YANG L Y,CANARINI A,ZHANG W S,et al. Microbial life-history strategies mediate microbial carbon pump efficacy in response to N management depending on stoichiometry of microbial demand[J]. Global Change Biology,2024,30(5):e17311