盐生植物根系性状对滨海湿地微生物脱氮过程的影响机制研究进展

doi:10.11707/j.1001-7488.LYKX20260101

摘要/Abstract

摘要：

滨海湿地作为海陆过渡带的重要生态系统，受潮汐作用、盐度梯度以及盐生植被分布等多重环境因子的共同影响，形成了独特的氮（N）循环模式，在缓解近岸水体富营养化和维持生态系统功能方面具有重要作用。滨海湿地土壤脱N过程的驱动机制已受到广泛关注，但现有研究多侧重于环境因子或微生物过程的单一解析，盐生植物对微生物脱N过程的影响机制仍缺乏系统梳理。盐生植物根系性状是调控植被特征与土壤N循环过程耦合关系的重要功能属性。本研究系统梳理了滨海湿地水文与盐度条件以及盐生植物对脱N过程的影响。重点从根系功能性状的角度，整合了根系生理性状、化学性状和形态性状通过碳（C）输入、根系泌氧、调节根际微环境等途径对微生物脱N过程的影响，以期揭示根系介导的微生物群落结构重塑、核心脱N功能类群富集以及C—N、铁（Fe）—N等关键元素耦合脱N过程的调控机制。本研究旨在填补盐生植物根系性状与滨海湿地微生物脱N过程之间系统认识的不足，为未来相关研究的深化开展提供理论参考。

关键词: 滨海湿地, 脱氮过程, 根系性状, 根系?微生物互作, 根际过程

Abstract:

Coastal wetlands are key transitional ecosystems located at the terrestrial-aquatic interface, and influenced by multiple environmental factors such as tidal regimes, salinity gradients, and distribution of halophyte vegetation, thereby forming unique nitrogen (N) cycling patterns. Coastal wetlands play a critical role in alleviating nearshore eutrophication and maintaining ecosystem functions. Although the drivers of soil N removal processes in coastal wetlands have been widely studied, existing research has largely focused on environmental factors or microbial processes separately, whereas the regulating mechanisms of halophytes on microbial denitrification still lacks a systematic review. Halophyte root functional traits are also increasingly considered a key functional attribute in regulating the coupling relationship between vegetation characteristics and soil nitrogen cycling processes. This study systematically reviews current knowledge on the effects of hydrological and salinity gradients, as well as halophytes, on N removal in coastal wetlands, with a particular focus on root traits. We integrate how root chemical, physiological, and morphological traits influence microbial N removal through carbon (C) inputs, radial oxygen loss, and rhizosphere microenvironment regulation. The study further reveals the regulatory mechanisms of root-mediated microbial community restructuring, enrichment of key denitrifying taxa, and the regulation of N removal via coupled elemental cycles, including C–N and iron–N interactions. The aim of this review is to fill the systematic knowledge gap between halophyte root traits to microbial N removal in coastal wetlands, and provide a theoretical basis for advancing future research.

Key words: coastal wetland, nitrogen removal processes, root traits, root-microbial interactions, rhizosphere processes

中图分类号:

S938.1
S316

王绍坤,李晶,崔丽娟. 盐生植物根系性状对滨海湿地微生物脱氮过程的影响机制研究进展[J]. 林业科学, 2026, 62(4): 1-11.

Shaokun Wang,Jing Li,Lijuan Cui. Research Progress on Mechanisms of Halophyte Plant Root Traits Influencing Microbial Nitrogen Removal in Coastal Wetlands[J]. Scientia Silvae Sinicae, 2026, 62(4): 1-11.

图/表 4

图1

表1

盐生植物根系性状调控滨海湿地脱氮过程的微生物机制"

根系性状 Root traits	影响机制 Mechanisms	微生物类群 Microbial taxa	参考文献 References
根系泌氧 Radial oxygen loss	根系泌氧塑造基质中好氧和微好氧区域，影响氮去除过程反应强度 Radial oxygen loss creates aerobic and microaerobic zones within the substrate, thereby influencing the intensity of nitrogen removal processes	拟杆菌纲，地杆菌属，假单胞菌 Petrimonas, Geobacter, Pseudomonas	Hu et al.，2023a；2023b
根系生物量 Root biomass	植物较高的根系生物量提供更多碳源输入，促进反硝化过程 Greater root biomass enhances carbon inputs, thereby facilitating denitrification	nirS, nirK型反硝化微生物 nirS, nirK type denitrifiers	Choudhury et al.，2022
比表面积 Specific root area	高比表面积植物根系，提供更大附着界面，提高功能微生物多样性和活性 Higher root surface area provides greater carrier for microorganisms, enhancing their diversity and activity	放线菌门，节杆菌属Actinobacteria, Arthrobacter	Zhang et al.，2022
根直径，比根长 Root diameter，specific root length	控制分泌速率和细根周转的速率进一步调节碳输入，调控反硝化过程 Regulation of exudation rates and root turnover further modulates carbon inputs, thereby regulating denitrification	固氮螺菌属，慢生根瘤菌属，中华根瘤菌属Azospira， Bradyrhizobium， Sinorhizobium	Gillespie et al.，2021；Wang et al.，2024a
根氮浓度 Root nitrogen content	不仅决定了植物本身的养分吸收率，还减少了土壤中的氮底物，抑制反硝化 This not only determines the nutrient uptake rate of plants themselves, but also reduces nitrogen substrates in the soil, thereby inhibiting denitrification		Cantarel et al.，2015；Moreau et al.，2015
根系次生代谢产物 Root secondary metabolites	脂肪酸甲酯、脂肪酸酰胺、豆甾醇作为信号分析促进反硝化菌氮去除活性 Fatty acid methyl esters, fatty acid amides, and stigmasterol function as signaling molecules, enhancing the nitrogen removal activity of denitrifying bacteria	假单胞菌属Pseudomonas	Sun et al.，2016；Lu et al.，2021
根系初生代谢产物 Root primary metabolites	有机酸、蛋白质、氨基酸、糖类作为碳源促进反硝化过程 Organic acids, proteins, amino acids, and carbohydrates serve as carbon sources to promote denitrification	绿脓杆菌，地杆菌属，梭状芽孢杆菌属Pseudomonas aeruginosa，Geobacter, Clostridium	Yin et al.，2019；Ma et al.，2021
生物反硝化抑制剂 Biological denitrification inhibitors	根系分泌的黄酮和萜类化合物抑制反硝化；类腐殖酸以及氨氧化过程产生的羟胺抑制厌氧氨氧化活性 Flavonoids and terpenoids exuded by roots inhibit denitrification, while humic-like substances and hydroxylamine generated during ammonia oxidation suppress anaerobic ammonium oxidation activity	nirS, nirK, nosZ型反硝化微生物 nirS, nirK, nosZ type denitrifiers	Li et al.，2025；Hu et al.，2023a

表1

图2

图3

参考文献 0

	陈飞扬. 2023. 河口潮滩湿地反硝化型甲烷厌氧氧化及其影响机制研究. 上海: 华东师范大学.
	Chen F Y. 2023. Denitrifying anaerobic methane oxidation and its influencing mechanism in estuarine intertidal wetlands. Shanghai: East China Normal University. ［in Chinese］
	陈顺涛. 2023. 河口潮滩湿地氮素转化过程及环境调控机制. 上海: 华东师范大学.
	Chen S T. 2023. Nitrogen transformation processes and its environmental controls in estuarine tidal flat wetland. Shanghai: East China Normal University. ［in Chinese］
	崔丽娟, 康晓明, 张曼胤, 等. 2019. 中国滨海湿地生态系统功能及服务评价. 北京: 中国林业出版社.
	Cui L J, Kang X M, Zhang M Y, et al. 2019. Assessment of ecosystem functions and services of China’s coastal wetlands. Beijing: China Forestry Publishing House. ［in Chinese］
	崔丽娟, 李　伟, 窦志国, 等. 近30年中国滨海滩涂湿地变化及其驱动力. 生态学报, 2022, 42 (18): 7297- 7307. doi: 10.5846/stxb202203080551
	Cui L J, Li W, Dou Z G, et al. Changes and driving forces of the tidal flat wetlands in coastal China during the past 30 years. Acta Ecologica Sinica, 2022, 42 (18): 7297- 7307. doi: 10.5846/stxb202203080551
	李　晶, 雷茵茹, 崔丽娟, 等. 我国滨海滩涂湿地现状及研究进展. 林业资源管理, 2018, (2): 24- 28. doi: 10.13466/j.cnki.lyzygl.2018.02.005
	Li J, Lei Y R, Cui L J, et al. Current status and research progress of coastal tidal flat wetlands in China. Forest Resources Management, 2018, (2): 24- 28. doi: 10.13466/j.cnki.lyzygl.2018.02.005
	李　威, 于珍珍, 何　徽, 等. 碳输入改变对湿地松人工成熟林土壤微生物群落的影响. 林业科学, 2025, 61 (6): 232- 242. doi: 10.11707/j.1001-7488.LYKX20240238
	Li W, Yu Z Z, He H, et al. Effects of carbon input change on soil bacterial and fungal community structure and diversity in a mature Pinus elliottii plantation. Scientia Silvae Sinicae, 2025, 61 (6): 232- 242. doi: 10.11707/j.1001-7488.LYKX20240238
	刘世荣, 陈远其, 聂秀青, 等. 2026. 树种多样性对森林生态系统多功能性和韧性的影响. 林业科学, 62(1): 1−18.
	Liu S R, Chen Y Q, Nie X Q, et al. 2026. Effects of tree species diversity on multifunctionality and resilience of forest ecosystems. Scientia Silvae Sinicae, 62(1): 1−18. ［in Chinese］
	刘晓娟, 马克平. 植物功能性状研究进展. 中国科学(生命科学), 2015, 45 (4): 325- 339.
	Liu X J, Ma K P. Plant functional traits: concepts, applications and future directions. Science in China (Series C), 2015, 45 (4): 325- 339.
	刘志君, 崔丽娟, 李　伟, 等. 互花米草入侵对盐城滨海湿地nirS型反硝化细菌多样性及群落结构的影响. 生态环境学报, 2022, 31 (4): 704- 714. doi: 10.16258/j.cnki.1674-5906.2022.04.008
	Liu Z J, Cui L J, Li W, et al. Effects of Spartina alterniflora invasion on the diversity and community structure of nirS-type denitrifying bacteria in Yancheng coastal wetlands. Ecology and Environment Sciences, 2022, 31 (4): 704- 714. doi: 10.16258/j.cnki.1674-5906.2022.04.008
	王汝苗, 李　晶, 刘魏魏, 等. 微生物代谢可塑性对退化湿地固碳的调控机制及生态恢复启示. 林业科学, 2025, 61 (7): 52- 58.
	Wang R M, Li J, Liu W W, et al. Regulating mechanisms of microbial metabolic plasticity on carbon sequestration in degraded wetlands and its implications for ecological restoration. Scientia Silvae Sinicae, 2025, 61 (7): 52- 58.
	中国海洋生态环境状况公报. 2023. 生态环境部.
	China Marine Ecological Environment Status Bulletin. 2023. Ministry of Ecology and Environment of the People’s Republic of China. ［in Chinese］
	An Z R, Chen F Y, Zheng Y L, et al. Role of n-DAMO in mitigating methane emissions from intertidal wetlands is regulated by saltmarsh vegetations. Environmental Science & Technology, 2024, 58 (2): 1152- 1163. doi: 10.1021/acs.est.3c07882
	Attia S, Russel J, Mortensen M S, et al. Unexpected diversity among small-scale sample replicates of defined plant root compartments. The ISME Journal, 2022, 16 (4): 997- 1003. doi: 10.1038/s41396-021-01094-7
	Bardgett R D, Mommer L, De Vries F T. Going underground: root traits as drivers of ecosystem processes. Trends in Ecology & Evolution, 2014, 29 (12): 692- 699. doi: 10.1016/j.tree.2014.10.006
	Bergmann J, Weigelt A, van der Plas F, et al. 2020. The fungal collaboration gradient dominates the root economics space in plants. Science Advances, 6(27): eaba3756.
	Bowen J L, Spivak A C, Bernhard A E, et al. Salt marsh nitrogen cycling: where land meets sea. Trends in Microbiology, 2024, 32 (6): 565- 576. doi: 10.1016/j.tim.2023.09.010
	Cantarel A A M, Pommier T, Desclos-Theveniau M, et al. Using plant traits to explain plant–microbe relationships involved in nitrogen acquisition. Ecology, 2015, 96 (3): 788- 799. doi: 10.1890/13-2107.1
	Chen S T, Gao D Z, Zhang J B, et al. Gross nitrogen transformations in intertidal sediments of the Yangtze Estuary: Distribution patterns and environmental controls. Geoderma, 2023a, 429, 116233. doi: 10.1016/j.geoderma.2022.116233
	Chen S T, Gao D Z, Li X F, et al. Nitrogen loss from anaerobic ammonium oxidation coupled to Iron(III) reduction activity across estuarine and coastal wetlands of China: Spatial variations, controlling factors, and environmental implications. Catena, 2023b, 222, 106805. doi: 10.1016/j.catena.2022.106805
	Chen Y, Wen Y, Zhou Q, et al. Effects of plant biomass on nitrogen transformation in subsurface-batch constructed wetlands: a stable isotope and mass balance assessment. Water Research, 2014, 63, 158- 167. doi: 10.1016/j.watres.2014.06.015
	Cheng S S, Gong X, Xue W F, et al. Evolutionarily conserved core microbiota as an extended trait in nitrogen acquisition strategy of herbaceous species. New Phytologist, 2024, 244 (4): 1570- 1584. doi: 10.1111/nph.20118
	Choudhury M I, Hallin S, Ecke F, et al. Disentangling the roles of plant functional diversity and plant traits in regulating plant nitrogen accumulation and denitrification in freshwaters. Functional Ecology, 2022, 36 (4): 921- 932. doi: 10.1111/1365-2435.14001
	Coskun D, Britto D T, Shi W M, et al. How plant root exudates shape the nitrogen cycle. Trends in Plant Science, 2017, 22 (8): 661- 673. doi: 10.1016/j.tplants.2017.05.004
	Cui H, Yang Y C, Ding Y, et al. A novel pilot-scale tubular bioreactor-enhanced floating treatment wetland for efficient in situ nitrogen removal from urban landscape water: Long-term performance and microbial mechanisms. Water Environment Research, 2019, 91 (11): 1498- 1508. doi: 10.1002/wer.1147
	Deng S H, Peng S, Xie B H, et al. Influence characteristics and mechanism of organic carbon on denitrification, N₂O emission and NO₂−accumulation in the iron [Fe(0)]-oxidizing supported autotrophic denitrification process. Chemical Engineering Journal, 2020, 393, 124736. doi: 10.1016/j.cej.2020.124736
	Di Capua F, Pirozzi F, Lens P N L, et al. Electron donors for autotrophic denitrification. Chemical Engineering Journal, 2019, 362, 922- 937. doi: 10.1016/j.cej.2019.01.069
	Fan K K, Weisenhorn P, Gilbert J A, et al. Wheat rhizosphere harbors a less complex and more stable microbial co-occurrence pattern than bulk soil. Soil Biology and Biochemistry, 2018, 125, 251- 260. doi: 10.1016/j.soilbio.2018.07.022
	Fort F. Grounding trait-based root functional ecology. Functional Ecology, 2023, 37 (8): 2159- 2169.
	Freschet G T, Roumet C, Comas L H, et al. Root traits as drivers of plant and ecosystem functioning: current understanding, pitfalls and future research needs. New Phytologist, 2021, 232 (3): 1123- 1158. doi: 10.1111/nph.17072
	Gao J, Hou L J, Zheng Y L, et al. nirS-Encoding denitrifier community composition, distribution, and abundance along the coastal wetlands of China. Applied Microbiology and Biotechnology, 2016, 100 (19): 8573- 8582. doi: 10.1007/s00253-016-7659-5
	Gao D Z, Liu C, Li X F, et al. High importance of coupled nitrification-denitrification for nitrogen removal in a large periodically low-oxygen estuary. Science of the Total Environment, 2022, 846, 157516. doi: 10.1016/j.scitotenv.2022.157516
	Gao X Q, Bi Y X, Su L, et al. Unveiling the nitrogen and phosphorus removal potential: Comparative analysis of three coastal wetland plant species in lab-scale constructed wetlands. Journal of Environmental Management, 2024, 351, 119864. doi: 10.1016/j.jenvman.2023.119864
	Gillespie L M, Hättenschwiler S, Milcu A, et al. Tree species mixing affects soil microbial functioning indirectly via root and litter traits and soil parameters in European forests. Functional Ecology, 2021, 35 (10): 2190- 2204. doi: 10.1111/1365-2435.13877
	Glaister B J, Fletcher T D, Cook P L M, et al. Interactions between design, plant growth and the treatment performance of stormwater biofilters. Ecological Engineering, 2017, 105, 21- 31. doi: 10.1016/j.ecoleng.2017.04.030
	Guyonnet J P, Vautrin F, Meiffren G, et al. The effects of plant nutritional strategy on soil microbial denitrification activity through rhizosphere primary metabolites. FEMS Microbiology Ecology, 2017, 93 (4): fix022. doi: 10.1093/femsec/fix022
	Giblin A E, Weston N B, Banta G T, et al. The effects of salinity on nitrogen losses from an oligohaline estuarine sediment. Estuaries and Coasts, 2010, 33 (5): 1054- 1068. doi: 10.1007/s12237-010-9280-7
	Hamonts K, Trivedi P, Garg A, et al. Field study reveals core plant microbiota and relative importance of their drivers. Environmental Microbiology, 2018, 20 (1): 124- 140. doi: 10.1111/1462-2920.14031
	He N P, Li Y, Liu C C, et al. Plant trait networks: improved resolution of the dimensionality of adaptation. Trends in Ecology & Evolution, 2020, 35 (10): 908- 918. doi: 10.1016/j.tree.2020.06.003
	Herms C H, Hennessy R C, Bak F, et al. Back to our roots: exploring the role of root morphology as a mediator of beneficial plant–microbe interactions. Environmental Microbiology, 2022, 24 (8): 3264- 3272. doi: 10.1111/1462-2920.15926
	Hinshaw S E, Tatariw C, Flournoy N, et al. Vegetation loss decreases salt marsh denitrification capacity: implications for marsh erosion. Environmental Science & Technology, 2017, 51 (15): 8245- 8253. doi: 10.1021/acs.est.7b00618
	Hu B L, Shen L D, Xu X Y, et al. Anaerobic ammonium oxidation (anammox) in different natural ecosystems. Biochemical Society Transactions, 2011, 39 (6): 1811- 1816. doi: 10.1042/BST20110711
	Hu J P, He Y Y, Li J H, et al. Planting halophytes increases the rhizosphere ecosystem multifunctionality via reducing soil salinity. Environmental Research, 2024, 261, 119707. doi: 10.1016/j.envres.2024.119707
	Hu S S, Chen Z B, Vosátka M, et al. Arbuscular mycorrhizal fungi colonization and physiological functions toward wetland plants under different water regimes. Science of the Total Environment, 2020, 716, 137040. doi: 10.1016/j.scitotenv.2020.137040
	Hu X J, Xie J X, Xie H J, et al. Towards a better and more complete understanding of microbial nitrogen transformation processes in the rhizosphere of subsurface flow constructed wetlands: effect of plant root activities. Chemical Engineering Journal, 2023a, 463, 142455. doi: 10.1016/j.cej.2023.142455
	Hu X J, Yue J Y, Yao D D, et al. Plant development alters the nitrogen cycle in subsurface flow constructed wetlands: Implications to the strategies for intensified treatment performance. Water Research, 2023b, 246, 120750. doi: 10.1016/j.watres.2023.120750
	Hu Y K, Liu G F, Pan X, et al. Abundance-weighted plant functional trait variation differs between terrestrial and wetland habitats along wide climatic gradients. Science China Life Sciences, 2021, 64 (4): 593- 605. doi: 10.1007/s11427-020-1766-1
	Huang F J, Lin X B, Hu W F, et al. Nitrogen cycling processes in sediments of the Pearl River Estuary: spatial variations, controlling factors, and environmental implications. Catena, 2021, 206, 105545. doi: 10.1016/j.catena.2021.105545
	King W L, Yates C F, Guo J, et al. The hierarchy of root branching order determines bacterial composition, microbial carrying capacity and microbial filtering. Communications Biology, 2021, 4, 483. doi: 10.1038/s42003-021-01988-4
	Kuypers M M M, Marchant H K, Kartal B. The microbial nitrogen-cycling network. Nature Reviews Microbiology, 2018, 16 (5): 263- 276. doi: 10.1038/nrmicro.2018.9
	Li J, Cui L J, Delgado-Baquerizo M, et al. Plant species and associated root nutritional traits influence soil dominant bacteria in coastal wetlands across China. New Phytologist, 2024, 244 (2): 683- 693. doi: 10.1111/nph.20047
	Li J C, Bol R, Jones D L, et al. Exploiter no more: root metabolites driving denitrification inhibition from diverse plants. Soil Biology and Biochemistry, 2025, 209, 109898. doi: 10.1016/j.soilbio.2025.109898
	Li N, Li B, Nie M, et al. Effects of exotic Spartina alterniflora on saltmarsh nitrogen removal in the Yangtze River Estuary, China. Journal of Cleaner Production, 2020a, 271, 122557. doi: 10.1016/j.jclepro.2020.122557
	Li X F, Hou L J, Liu M, et al. Biogeochemical controls on nitrogen transformations in subtropical estuarine wetlands. Environmental Pollution, 2020b, 263, 114379. doi: 10.1016/j.envpol.2020.114379
	Ling N, Wang T T, Kuzyakov Y. Rhizosphere bacteriome structure and functions. Nature Communications, 2022, 13, 836. doi: 10.1038/s41467-022-28448-9
	Liu S B, He F K, Kuzyakov Y, et al. Nutrients in the rhizosphere: a meta-analysis of content, availability, and influencing factors. Science of the Total Environment, 2022, 826, 153908. doi: 10.1016/j.scitotenv.2022.153908
	Lopes L D, Schachtman D P. Rhizosphere and bulk soil bacterial community succession is influenced more by changes in soil properties than in rhizosphere metabolites across a maize growing season. Applied Soil Ecology, 2023, 189, 104960. doi: 10.1016/j.apsoil.2023.104960
	Lu Y F, Kronzucker H J, Shi W M. Stigmasterol root exudation arising from Pseudomonas inoculation of the duckweed rhizosphere enhances nitrogen removal from polluted waters. Environmental Pollution, 2021, 287, 117587. doi: 10.1016/j.envpol.2021.117587
	Lu Y F, Kronzucker H J, Yu M, et al. Nitrogen-loss and carbon-footprint reduction by plant-rhizosphere exudates. Trends in Plant Science, 2024, 29 (4): 469- 481. doi: 10.1016/j.tplants.2023.09.007
	Ma L, Yang L L, Liu W, et al. Effects of root exudates on rhizosphere bacteria and nutrient removal in pond-ditch circulation systems (PDCSs) for rural wastewater treatment. Science of the Total Environment, 2021, 785, 147282. doi: 10.1016/j.scitotenv.2021.147282
	Ma Y D, Yue K, Heděnec P, et al. Global patterns of rhizosphere effects on soil carbon and nitrogen biogeochemical processes. Catena, 2023, 220, 106661. doi: 10.1016/j.catena.2022.106661
	Ma Z J, Melville D S, Liu J G, et al. Rethinking China’s new great wall. Science, 2014, 346 (6212): 912- 914. doi: 10.1126/science.1257258
	Maisch M, Lueder U, Kappler A, et al. Iron lung: how rice roots induce iron redox changes in the rhizosphere and create niches for microaerophilic Fe(II)-oxidizing bacteria. Environmental Science & Technology Letters, 2019, 6 (10): 600- 605. doi: 10.1021/acs.estlett.9b00403
	Moreau D, Pivato B, Bru D, et al. Plant traits related to nitrogen uptake influence plant-microbe competition. Ecology, 2015, 96 (8): 2300- 2310. doi: 10.1890/14-1761.1
	Navarro-Torre S, Garcia-Caparrós P, Nogales A, et al. Sustainable agricultural management of saline soils in arid and semi-arid Mediterranean regions through halophytes, microbial and soil-based technologies. Environmental and Experimental Botany, 2023, 212, 105397. doi: 10.1016/j.envexpbot.2023.105397
	Ooi S K, Barry A, Lawrence B A, et al. Vegetation zones as indicators of denitrification potential in salt marshes. Ecological Applications, 2022, 32 (6): e2630. doi: 10.1002/eap.2630
	Peng Y Y, Gu X S, Zhang M P, et al. Simultaneously enhanced autotrophic–heterotrophic denitrification in iron-based ecological floating bed by plant biomass: Metagenomics insights into microbial communities, functional genes and nitrogen metabolic pathways. Water Research, 2024, 248, 120868. doi: 10.1016/j.watres.2023.120868
	Pervaiz Z H, Contreras J, Hupp B M, et al. Root microbiome changes with root branching order and root chemistry in peach rhizosphere soil. Rhizosphere, 2020, 16, 100249. doi: 10.1016/j.rhisph.2020.100249
	Phillips R P, Fahey T J. Tree species and mycorrhizal associations influence the magnitude of rhizosphere effects. Ecology, 2006, 87 (5): 1302- 1313. doi: 10.1890/0012-9658(2006)87[1302:TSAMAI]2.0.CO;2
	Pold G, Saghaï A, Jones C M, et al. Denitrification is a community trait with partial pathways dominating across microbial genomes and biomes. Nature Communications, 2025, 16, 9495. doi: 10.1038/s41467-025-65319-5
	Qi M T, Li Y, Fu Y X, et al. Enhanced coupling of Fe(II) oxidation and nitrate reduction benefits chemoautotrophic carbon fixation in estuarine and coastal sediments. Environmental Science & Technology, 2025, 59 (39): 21189- 21201. doi: 10.1021/acs.est.5c05446
	Ramesh R, Chen Z, Cummins V, et al. Land–ocean interactions in the coastal zone: past, present and future. Anthropocene, 2015, 12, 85- 98. doi: 10.1016/j.ancene.2016.01.005
	Rooks C, Schmid M C, Mehsana W, et al. The depth–specific significance and relative abundance of anaerobic ammonium–oxidizing bacteria in estuarine sediments (Medway Estuary, UK). FEMS Microbiology Ecology, 2012, 80 (1): 19- 29. doi: 10.1111/j.1574-6941.2011.01266.x
	Schulte-Uebbing L F, Beusen A H W, Bouwman A F, et al. From planetary to regional boundaries for agricultural nitrogen pollution. Nature, 2022, 610 (7932): 507- 512. doi: 10.1038/s41586-022-05158-2
	Shi F X, Song C C, Zhang X H, et al. Plant zonation patterns reflected by the differences in plant growth, biomass partitioning and root traits along a water level gradient among four common vascular plants in freshwater marshes of the Sanjiang Plain, Northeast China. Ecological Engineering, 2015, 81, 158- 164. doi: 10.1016/j.ecoleng.2015.04.054
	Steffen W, Richardson K, Rockström J, et al. Planetary boundaries: Guiding human development on a changing planet. Science, 2015, 347 (6223): 1259855. doi: 10.1126/science.1259855
	Stopnisek N, Shade A. Persistent microbiome members in the common bean rhizosphere: an integrated analysis of space, time, and plant genotype. The ISME Journal, 2021, 15 (9): 2708- 2722. doi: 10.1038/s41396-021-00955-5
	Sun L, Lu Y F, Kronzucker H J, et al. Quantification and enzyme targets of fatty acid amides from duckweed root exudates involved in the stimulation of denitrification. Journal of Plant Physiology, 2016, 198, 81- 88. doi: 10.1016/j.jplph.2016.04.010
	Sharp S J, Elgersma K J, Martina J P, et al. Hydrologic flushing rates drive nitrogen cycling and plant invasion in a freshwater coastal wetland model. Ecological Applications, 2021, 31 (2): e02233. doi: 10.1002/eap.2233
	Tang S Y, Liao Y H, Xu Y C, et al. Microbial coupling mechanisms of nitrogen removal in constructed wetlands: a review. Bioresource Technology, 2020, 314, 123759. doi: 10.1016/j.biortech.2020.123759
	Vieira S, Sikorski J, Dietz S, et al. Drivers of the composition of active rhizosphere bacterial communities in temperate grasslands. The ISME Journal, 2020, 14 (2): 463- 475. doi: 10.1038/s41396-019-0543-4
	Wan X H, Chen X L, Huang Z Q, et al. Contribution of root traits to variations in soil microbial biomass and community composition. Plant and Soil, 2021, 460 (1): 483- 495. doi: 10.1007/s11104-020-04788-7
	Wang R M, Cui L J, Li J, et al. Factors driving the halophyte rhizosphere bacterial communities in coastal salt marshes. Frontiers in Microbiology, 2023a, 14, 1127958. doi: 10.3389/fmicb.2023.1127958
	Wang F F, Lu Z Y, Tobias C R, et al. Salt marsh expansion into estuarine mangrove mudflats reduces nitrogen removal capacity. Catena, 2023b, 232, 107459. doi: 10.1016/j.catena.2023.107459
	Wang S K, Li J, Cui L J, et al. Latitudinal patterns and phosphorus-driven regulation of abundant and rare fungal communities in coastal wetlands. Applied Soil Ecology, 2025, 206, 105855. doi: 10.1016/j.apsoil.2024.105855
	Wang S K, Li J, Wang R M, et al. Specific root length regulated the rhizosphere effect on denitrification across distinct macrophytes. Geoderma, 2024a, 449, 117002. doi: 10.1016/j.geoderma.2024.117002
	Wang W Q, Guo Y P, Yang L, et al. Methanogen–methanotroph community has a more consistent and integrated structure in rice rhizosphere than in bulk soil and rhizoplane. Molecular Ecology, 2024b, 33 (13): e17416. doi: 10.1111/mec.17416
	Wang J, Zhang Z X, Liang F, et al. Decreased soil pH weakens the positive rhizosphere effect on denitrification capacity. Pedosphere, 2024c, 34 (5): 905- 915. doi: 10.1016/j.pedsph.2023.07.011
	Wen Z H, Yu P, Shen J B, et al. 2025. Do rhizosphere microbiomes match root functional traits? Trends in Ecology & Evolution, 40(9): 885−899.
	Wright I J, Reich P B, Westoby M, et al. The worldwide leaf economics spectrum. Nature, 2004, 428, 821- 827. doi: 10.1038/nature02403
	Wu H L, Wang X Z, He X J, et al. Effects of root exudates on denitrifier gene abundance, community structure and activity in a micro-polluted constructed wetland. Science of the Total Environment, 2017, 598, 697- 703. doi: 10.1016/j.scitotenv.2017.04.150
	Xia Z G, Wang Q, She Z L, et al. Nitrogen removal pathway and dynamics of microbial community with the increase of salinity in simultaneous nitrification and denitrification process. Science of the Total Environment, 2019, 697, 134047. doi: 10.1016/j.scitotenv.2019.134047
	Xiao S Y, Luo M, Liu Y X, et al. Rhizosphere effect and its associated soil-microbe interactions drive iron fraction dynamics in tidal wetland soils. Science of the Total Environment, 2021, 756, 144056. doi: 10.1016/j.scitotenv.2020.144056
	Xiong C, Zhu Y G, Wang J T, et al. Host selection shapes crop microbiome assembly and network complexity. New Phytologist, 2021, 229 (2): 1091- 1104. doi: 10.1111/nph.16890
	Xun W B, Liu Y P, Ma A Y, et al. Dissection of rhizosphere microbiome and exploiting strategies for sustainable agriculture. New Phytologist, 2024, 242 (6): 2401- 2410. doi: 10.1111/nph.19697
	Yin X, Lu J, Wang Y, et al. The abundance of nirS–type denitrifiers and anammox bacteria in rhizospheres was affected by the organic acids secreted from roots of submerged macrophytes. Chemosphere, 2019, 240, 124903.
	Zhang B, Chen L, Guo Q Z, et al. Characteristics of nitrogen distribution and its response to microecosystem changes in green infrastructure with different woody plants. Chemosphere, 2022, 313, 137371. doi: 10.1016/j.chemosphere.2022.137371
	Zheng Y L, Hou L J, Liu M, et al. Community composition and activity of anaerobic ammonium oxidation bacteria in the rhizosphere of salt-marsh grass Spartina alterniflora. Applied Microbiology and Biotechnology, 2016, 100 (18): 8203- 8212. doi: 10.1007/s00253-016-7625-2
	Zhou X, Li B L, Guo Z Y, et al. Niche separation of ammonia oxidizers in mudflat and agricultural soils along the Yangtze River, China. Frontiers in Microbiology, 2018, 9, 3122. doi: 10.3389/fmicb.2018.03122