{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,4]],"date-time":"2026-08-04T08:47:16Z","timestamp":1785833236536,"version":"3.56.0"},"reference-count":54,"publisher":"MDPI AG","issue":"8","license":[{"start":{"date-parts":[[2026,8,4]],"date-time":"2026-08-04T00:00:00Z","timestamp":1785801600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"Science and Technology Development Plan Project of Jilin Province, China","award":["YDZJ202501ZYTS403"],"award-info":[{"award-number":["YDZJ202501ZYTS403"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Microorganisms"],"abstract":"<jats:p>Cool-climate maize fields are characterized by low soil temperatures, strong seasonal hydrothermal fluctuations, and peat-influenced soil profiles, which may lead to patterns of nitrogen (N) cycling distinct from those in conventional agricultural soils. During maize growth, soils from three depths were characterized using physicochemical measurements, N-transformation and enzyme-activity assays, metagenomic sequencing, Mantel tests, variation partitioning analysis, and partial least squares path modeling (PLS-PM). Soil environmental factors varied significantly over time and with depth; soil organic matter (SOM) and total nitrogen (TN) increased with depth, while ammonium nitrogen (NH4+-N) predominated early and nitrate nitrogen (NO3\u2212-N) predominated during the middle and late growth stages. The nitrogen fixation rate (NFR), nitrification rate (NitR), and denitrification rate (DNR) all peaked in August and showed a spatial pattern characterized by nitrogen fixation in the deepest layer and denitrification in the upper and middle layers. Bacterial communities varied less spatiotemporally than fungal communities. The genes nifK, hao, nirS\/nirK, NR, nrfC, and hzsA\/hzsC were identified as key nitrogen-cycling functional genes. Mantel tests and PLS-PM further characterized these relationships, with PLS-PM showing that soil physicochemical properties were positively associated with bacterial community composition (\u03b2 = 0.87, p &lt; 0.01), which, in turn, was negatively associated with N-cycling functional genes (\u03b2 = \u22120.97, p &lt; 0.001). Together, these pathways were associated with variation in N-cycling processes. Overall, this study advances an integrated understanding of N-cycling patterns and their potential controls in cool-climate maize fields and provides a scientific basis for optimizing N management strategies.<\/jats:p>","DOI":"10.3390\/microorganisms14081705","type":"journal-article","created":{"date-parts":[[2026,8,4]],"date-time":"2026-08-04T08:02:19Z","timestamp":1785830539000},"page":"1705","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Interaction Mechanisms Among Soil Environmental Factors, Microbial Communities, and Nitrogen-Cycling Functional Genes in Cool-Climate Maize Fields"],"prefix":"10.3390","volume":"14","author":[{"given":"Qingqing","family":"Dai","sequence":"first","affiliation":[{"name":"Agriculture College, Yanbian University, Yanji 133002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yuhang","family":"Wang","sequence":"additional","affiliation":[{"name":"Agriculture College, Yanbian University, Yanji 133002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mingji","family":"Jin","sequence":"additional","affiliation":[{"name":"College of Geography and Ocean Sciences, Yanbian University, Yanji 133002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shuo","family":"Wang","sequence":"additional","affiliation":[{"name":"Agriculture College, Yanbian University, Yanji 133002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mingji","family":"Han","sequence":"additional","affiliation":[{"name":"Agriculture College, Yanbian University, Yanji 133002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,8,4]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","unstructured":"Futa, B., Gmitrowicz-Iwan, J., Skersien\u0117, A., \u0160lepetien\u0117, A., and Para\u0161otas, I. (2024). Innovative Soil Management Strategies for Sustainable Agriculture. Sustainability, 16.","DOI":"10.3390\/su16219481"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"147","DOI":"10.3103\/S0147687420040055","article-title":"Soil Is an Essential Component of the Biosphere and the Global Food System (Critical Assessment of the Situation)","volume":"75","author":"Khomiakov","year":"2020","journal-title":"Mosc. Univ. Soil Sci. Bull."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1080\/03650340.2024.2325697","article-title":"Investigating Soil Quality in Cold Highland Agricultural Fields with Different Soil Types","volume":"70","author":"Qiu","year":"2024","journal-title":"Arch. Agron. Soil Sci."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"e20402","DOI":"10.1002\/tpg2.20402","article-title":"Advances and Opportunities in Unraveling Cold-Tolerance Mechanisms in the World\u2019s Primary Staple Food Crops","volume":"17","author":"Jan","year":"2024","journal-title":"Plant Genome"},{"key":"ref_5","first-page":"281","article-title":"Untangling the Increasing Elevation of Cropland in China from 1980 to 2020","volume":"4","author":"Chen","year":"2023","journal-title":"Geogr. Sustain."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"782","DOI":"10.1016\/j.scitotenv.2017.12.309","article-title":"Shifts in Soil Bacterial and Archaeal Communities during Freeze-Thaw Cycles in a Seasonal Frozen Marsh, Northeast China","volume":"625","author":"Ren","year":"2018","journal-title":"Sci. Total Environ."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"115170","DOI":"10.1016\/j.geoderma.2021.115170","article-title":"Linking Soil Water Retention Capacity to Pore Structure Characteristics Based on X-Ray Computed Tomography: Chinese Mollisol under Freeze-Thaw Effect","volume":"401","author":"Liu","year":"2021","journal-title":"Geoderma"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"810","DOI":"10.1038\/s41598-020-79988-3","article-title":"Modelling Adaptation Strategies to Reduce Adverse Impacts of Climate Change on Maize Cropping System in Northeast China","volume":"11","author":"Jiang","year":"2021","journal-title":"Sci. Rep."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"105685","DOI":"10.1016\/j.apsoil.2024.105685","article-title":"Effects of Organic Nitrogen Addition on Soil Microbial Community Assembly Patterns in the Sanjiang Plain Wetlands, Northeastern China","volume":"204","author":"Wang","year":"2024","journal-title":"Appl. Soil Ecol."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"379","DOI":"10.1007\/s11104-019-04391-5","article-title":"Bacteria and Fungi Differentially Contribute to Carbon and Nitrogen Cycles during Biological Soil Crust Succession in Arid Ecosystems","volume":"447","author":"Zhao","year":"2020","journal-title":"Plant Soil"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"263","DOI":"10.1038\/nrmicro.2018.9","article-title":"The Microbial Nitrogen-Cycling Network","volume":"16","author":"Kuypers","year":"2018","journal-title":"Nat. Rev. Microbiol."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"013004","DOI":"10.1088\/1748-9326\/ac417e","article-title":"A Review of the Importance of Mineral Nitrogen Cycling in the Plant-Soil-Microbe System of Permafrost-Affected Soils\u2014Changing the Paradigm","volume":"17","author":"Ramm","year":"2022","journal-title":"Environ. Res. Lett."},{"key":"ref_13","doi-asserted-by":"crossref","unstructured":"Gong, Y., Yang, T., Yan, J., and Yu, X. (2025). The Influence of Seasonal Freeze\u2013Thaw in Northeast China on Greenhouse Gas Emissions and Microbial Community Structure in Peat Soil. Water, 17.","DOI":"10.3390\/w17162395"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"175228","DOI":"10.1016\/j.scitotenv.2024.175228","article-title":"Dynamic Responses of Soil Microbial Communities to Seasonal Freeze-Thaw Cycles in a Temperate Agroecosystem","volume":"950","author":"Deng","year":"2024","journal-title":"Sci. Total Environ."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"106694","DOI":"10.1016\/j.still.2025.106694","article-title":"Seasonal Dynamics of Prokaryotic Nitrogen Cycling Genes in Cropland Soils: Effects of Soil Texture, Tillage, and Environmental Factors","volume":"253","author":"Yang","year":"2025","journal-title":"Soil Tillage Res."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"109002","DOI":"10.1016\/j.agee.2024.109002","article-title":"Agricultural Soil Microbiomes Differentiate in Soil Profiles with Fertility Source, Tillage, and Cover Crops","volume":"368","author":"Bier","year":"2024","journal-title":"Agric. Ecosyst. Environ."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"8033","DOI":"10.1073\/pnas.1601070113","article-title":"Global Biogeography of Microbial Nitrogen-Cycling Traits in Soil","volume":"113","author":"Nelson","year":"2016","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_18","doi-asserted-by":"crossref","unstructured":"Gao, T., Li, Y., Zhao, C., Chen, J., Jin, R., and Zhu, W. (2023). Factors Driving Changes in Water Conservation Function from a Geospatial Perspective: Case Study of Jilin Province. Front. Ecol. Evol., 11.","DOI":"10.3389\/fevo.2023.1303957"},{"key":"ref_19","doi-asserted-by":"crossref","unstructured":"Wang, Q., Zou, J., Liu, Y., Li, J., Liu, X., Zhang, H., Jiao, F., Meng, Q., and Guo, Z. (2024). Impacts of Farming Activities on Nitrogen Degradability under a Temperate Continental Monsoon Climate. Agronomy, 14.","DOI":"10.3390\/agronomy14061094"},{"key":"ref_20","doi-asserted-by":"crossref","unstructured":"Qu, J.-F., Hou, Y.-L., Ge, M.-Y., Wang, K., Liu, S., Zhang, S.-L., Li, G., and Chen, F. (2017). Carbon Dynamics of Reclaimed Coal Mine Soil under Agricultural Use: A Chronosequence Study in the Dongtan Mining Area, Shandong Province, China. Sustainability, 9.","DOI":"10.3390\/su9040629"},{"key":"ref_21","doi-asserted-by":"crossref","unstructured":"Liu, J., Cai, H., Chen, S., Pi, J., and Zhao, L. (2023). A Review on Soil Nitrogen Sensing Technologies: Challenges, Progress and Perspectives. Agriculture, 13.","DOI":"10.3390\/agriculture13040743"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"703","DOI":"10.1016\/0038-0717(87)90052-6","article-title":"An Extraction Method for Measuring Soil Microbial Biomass C","volume":"19","author":"Vance","year":"1987","journal-title":"Soil Biol. Biochem."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"1185","DOI":"10.1104\/pp.43.8.1185","article-title":"The Acetylene-Ethylene Assay for n(2) Fixation: Laboratory and Field Evaluation","volume":"43","author":"Hardy","year":"1968","journal-title":"Plant Physiol."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"2176","DOI":"10.1007\/s11368-018-02238-7","article-title":"High Soil Temperatures Alter the Rates of Nitrification, Denitrification and Associated N2O Emissions","volume":"19","author":"Lai","year":"2019","journal-title":"J. Soils Sediments"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"567","DOI":"10.5194\/bg-15-567-2018","article-title":"Seasonal and Interannual Dynamics of Soil Microbial Biomass and Available Nitrogen in an Alpine Meadow in the Eastern Part of Qinghai\u2013Tibet Plateau, China","volume":"15","author":"Xu","year":"2018","journal-title":"Biogeosciences"},{"key":"ref_26","doi-asserted-by":"crossref","unstructured":"Liu, Y., Chen, Q., Wang, Z., Zheng, H., Chen, Y., Chen, X., Wang, L., Li, H., and Zhang, J. (2019). Nitrogen Addition Alleviates Microbial Nitrogen Limitations and Promotes Soil Respiration in a Subalpine Coniferous Forest. Forests, 10.","DOI":"10.3390\/f10111038"},{"key":"ref_27","doi-asserted-by":"crossref","unstructured":"Berg, B., and McClaugherty, C. (2020). Plant Litter: Decomposition, Humus Formation, Carbon Sequestration, Springer International Publishing.","DOI":"10.1007\/978-3-030-59631-6"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"35","DOI":"10.2136\/sssabookser9.c2","article-title":"Ecology of Extracellular Enzyme Activities and Organic Matter Degradation in Soil: A Complex Community-Driven Process","volume":"Volume 9","author":"Dick","year":"2011","journal-title":"SSSA Book Series"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"fiab136","DOI":"10.1093\/femsec\/fiab136","article-title":"Cropping Systems Impact Changes in Soil Fungal, but Not Prokaryote, Alpha-Diversity and Community Composition Stability over a Growing Season in a Long-Term Field Trial","volume":"97","author":"Finn","year":"2021","journal-title":"FEMS Microbiol. Ecol."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"102","DOI":"10.1038\/nrmicro1341","article-title":"Microbial Biogeography: Putting Microorganisms on the Map","volume":"4","author":"Martiny","year":"2006","journal-title":"Nat. Rev. Microbiol."},{"key":"ref_31","doi-asserted-by":"crossref","unstructured":"Shade, A., Peter, H., Allison, S.D., Baho, D.L., Berga, M., B\u00fcrgmann, H., Huber, D.H., Langenheder, S., Lennon, J.T., and Martiny, J.B.H. (2012). Fundamentals of Microbial Community Resistance and Resilience. Front. Microbiol., 3.","DOI":"10.3389\/fmicb.2012.00417"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"e2304663120","DOI":"10.1073\/pnas.2304663120","article-title":"Enumerating Soil Biodiversity","volume":"120","author":"Anthony","year":"2023","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"107933","DOI":"10.1016\/j.soilbio.2020.107933","article-title":"Mycorrhizal Association of Common European Tree Species Shapes Biomass and Metabolic Activity of Bacterial and Fungal Communities in Soil","volume":"149","author":"Nilsson","year":"2020","journal-title":"Soil Biol. Biochem."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"243","DOI":"10.1128\/MMBR.00001-15","article-title":"Fungal Traits That Drive Ecosystem Dynamics on Land","volume":"79","author":"Treseder","year":"2015","journal-title":"Microbiol. Mol. Biol. Rev."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"5137","DOI":"10.1111\/1462-2920.13512","article-title":"Fungal Community Composition in Soils Subjected to Long-term Chemical Fertilization Is Most Influenced by the Type of Organic Matter","volume":"18","author":"Sun","year":"2016","journal-title":"Environ. Microbiol."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"e02222-17","DOI":"10.1128\/AEM.02222-17","article-title":"Methanogens Are Major Contributors to Nitrogen Fixation in Soils of the Florida Everglades","volume":"84","author":"Bae","year":"2018","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"591","DOI":"10.1890\/03-8002","article-title":"Nitrogen Mineralization: Challenges of a Changing Paradigm","volume":"85","author":"Schimel","year":"2004","journal-title":"Ecology"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"8217","DOI":"10.1073\/pnas.1704504114","article-title":"Nitric Oxide Is an Obligate Bacterial Nitrification Intermediate Produced by Hydroxylamine Oxidoreductase","volume":"114","author":"Caranto","year":"2017","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"699","DOI":"10.1016\/j.tim.2016.05.004","article-title":"A New Perspective on Microbes Formerly Known as Nitrite-Oxidizing Bacteria","volume":"24","author":"Daims","year":"2016","journal-title":"Trends Microbiol."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"6573","DOI":"10.1128\/JB.181.21.6573-6584.1999","article-title":"Prokaryotic Nitrate Reduction: Molecular Properties and Functional Distinction among Bacterial Nitrate Reductases","volume":"181","author":"Cabello","year":"1999","journal-title":"J. Bacteriol."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"1779","DOI":"10.5194\/bg-8-1779-2011","article-title":"Assessment of the Importance of Dissimilatory Nitrate Reduction to Ammonium for the Terrestrial Nitrogen Cycle","volume":"8","author":"Boeckx","year":"2011","journal-title":"Biogeosciences"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"124","DOI":"10.5670\/oceanog.2013.54","article-title":"The Importance of Dissimilatory Nitrate Reduction to Ammonium (DNRA) in the Nitrogen Cycle of Coastal Ecosystems","volume":"26","author":"Giblin","year":"2013","journal-title":"Oceanography"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"104","DOI":"10.1016\/j.jbiotec.2010.12.025","article-title":"Microbial Nitrate Respiration\u2014Genes, Enzymes and Environmental Distribution","volume":"155","author":"Kraft","year":"2011","journal-title":"J. Biotechnol."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"2784","DOI":"10.1111\/1462-2920.13134","article-title":"Ecology and Physiology of Anaerobic Ammonium Oxidizing Bacteria","volume":"18","author":"Oshiki","year":"2016","journal-title":"Environ. Microbiol."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"752","DOI":"10.1128\/AEM.07113-11","article-title":"Hydrazine Synthase, a Unique Phylomarker with Which To Study the Presence and Biodiversity of Anammox Bacteria","volume":"78","author":"Harhangi","year":"2012","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"e02230-21","DOI":"10.1128\/spectrum.02230-21","article-title":"Comparison of the Responses of Soil Fungal Community to Straw, Inorganic Fertilizer, and Compost in a Farmland in the Loess Plateau","volume":"10","author":"Yin","year":"2022","journal-title":"Microbiol. Spectr."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"489","DOI":"10.1146\/annurev-ecolsys-102710-145034","article-title":"Functional Ecology of Free-Living Nitrogen Fixation: A Contemporary Perspective","volume":"42","author":"Reed","year":"2011","journal-title":"Annu. Rev. Ecol. Evol. Syst."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"e02546-18","DOI":"10.1128\/AEM.02546-18","article-title":"To Fix or Not To Fix: Controls on Free-Living Nitrogen Fixation in the Rhizosphere","volume":"85","author":"Smercina","year":"2019","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"285","DOI":"10.1111\/j.1574-6976.2002.tb00616.x","article-title":"Enzymology and Bioenergetics of Respiratory Nitrite Ammonification","volume":"26","author":"Simon","year":"2002","journal-title":"FEMS Microbiol. Rev."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"1771","DOI":"10.1038\/s41467-017-01406-6","article-title":"Anaerobic Microsites Have an Unaccounted Role in Soil Carbon Stabilization","volume":"8","author":"Keiluweit","year":"2017","journal-title":"Nat. Commun."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"fix006","DOI":"10.1093\/femsec\/fix006","article-title":"Revisiting Life Strategy Concepts in Environmental Microbial Ecology","volume":"93","author":"Ho","year":"2017","journal-title":"FEMS Microbiol. Ecol."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"1354","DOI":"10.1890\/05-1839","article-title":"Toward an Ecological Classification of Soil Bacteria","volume":"88","author":"Fierer","year":"2007","journal-title":"Ecology"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"16160","DOI":"10.1038\/nmicrobiol.2016.160","article-title":"Exploiting rRNA Operon Copy Number to Investigate Bacterial Reproductive Strategies","volume":"1","author":"Roller","year":"2016","journal-title":"Nat. Microbiol."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"108111","DOI":"10.1016\/j.soilbio.2020.108111","article-title":"How Microbes Can, and Cannot, Be Used to Assess Soil Health","volume":"153","author":"Fierer","year":"2021","journal-title":"Soil Biol. Biochem."}],"container-title":["Microorganisms"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2076-2607\/14\/8\/1705\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,4]],"date-time":"2026-08-04T08:29:24Z","timestamp":1785832164000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2076-2607\/14\/8\/1705"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,4]]},"references-count":54,"journal-issue":{"issue":"8","published-online":{"date-parts":[[2026,8]]}},"alternative-id":["microorganisms14081705"],"URL":"https:\/\/doi.org\/10.3390\/microorganisms14081705","relation":{},"ISSN":["2076-2607"],"issn-type":[{"value":"2076-2607","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,4]]}}}