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Article(id=1312009711029936586, tenantId=1146029695717560320, journalId=1311385839672066169, issueId=1312009474789958403, articleNumber=null, orderNo=null, doi=10.13870/j.cnki.stbcxb.2026.04.040, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1763654400000, receivedDateStr=2025-11-21, revisedDate=1773504000000, revisedDateStr=2026-03-15, acceptedDate=1776355200000, acceptedDateStr=2026-04-17, onlineDate=1790737091070, onlineDateStr=2026-09-30, pubDate=1787846400000, pubDateStr=2026-08-28, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1790737091070, onlineIssueDateStr=2026-09-30, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1790737091070, creator=13701087609, updateTime=1790737091070, updator=13701087609, issue=Issue{id=1312009474789958403, tenantId=1146029695717560320, journalId=1311385839672066169, year='2026', volume='40', issue='4', pageStart='1', pageEnd='488', issueExtLink='null', onlineDate='null', pubDate='1787846400000', pubDateStr='2026-08-28', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=1, specialIssue=null, createTime=1790737034746, creator='13701087609', updateTime=1790739471092, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext={EN=IssueExt(id=1312019693662204413, tenantId=1146029695717560320, journalId=1311385839672066169, issueId=1312009474789958403, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1312019693662204414, tenantId=1146029695717560320, journalId=1311385839672066169, issueId=1312009474789958403, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null, downloadFileDto=null}, startPage=205, endPage=215, ext={EN=ArticleExt(id=1312009711256429003, articleId=1312009711029936586, tenantId=1146029695717560320, journalId=1311385839672066169, language=EN, title=Effects of Nitrogen Application on Rhizosphere Bacterial Denitrification Pathways， Root Growth， and Yield of Sunflowers in Salt-Affected Farmland, columnId=1312009477243626245, journalTitle=Journal of Soil and Water Conservation, columnName=Basic Research, runingTitle=null, highlight=null, articleAbstract=

Objective This study aims to investigate the specific mechanisms by which nitrogen application regulates rhizosphere bacterial denitrification pathways in salt-affected farmland， thereby promoting sunflower root growth and yield. Methods Field plot experiments with drip irrigation under plastic mulch were conducted in sunflower farmland from 2023 to 2024. Four nitrogen application levels were set： the local conventional nitrogen application rate as control （CK： 200 kg/hm2）， 25% nitrogen reduction （N1： 150 kg/hm2）， 50% nitrogen reduction （N2： 100 kg/hm2）， and 75% nitrogen reduction （N3： 50 kg/hm2）. Results The two-year experiments demonstrated that moderate nitrogen reduction improved the rhizosphere soil environment in salt-affected farmland and promoted sunflower root growth and yield formation. Compared with CK， both N1 and N2 treatments significantly increased Ca2+ （17.22%， 10.56%） and Mg2+ （34.75%， 25.40%） contents in rhizosphere soil at the budding stage， and reduced Cl- and SO42- contents. Furthermore， N1 treatment significantly promoted oxalic acid and citric acid secretion by root at the budding stage （59.07% and 60.11%）， and N2 treatment significantly promoted formic acid secretion by root at both the budding stage and the maturity stage. Regarding functional genes， N1， N2， and N3 treatments reduced the relative abundance of denitrifying genes narG， nirK， and nirS， whereas N1 treatment increased the abundance of the nosZ gene and altered the composition of nosZ-type bacterial communities， including increasing the relative abundance of nosZ-type Acidobacteria and Proteobacteria. Regarding crop growth and yield formation， N1 and N2 treatments significantly increased root surface area density （RSAD） at the flowering stage and root length density （RLD） at the budding stage. Over two years， N1 treatment significantly increased sunflower 100-seed weight （9.07%）， seed weight per head （9.08%）， and yield （5.05%）. In contrast， N2 and N3 treatments significantly reduced RSAD values （23.66%， 29.46%）， 100-seed weight （21.67%， 29.51%）， and seed weight per head （21.66%， 29.49%） at the maturity stage. Conclusion A 25% nitrogen reduction treatment has a potential inhibitory effect on the initial stage of denitrification （NO3-→NO） and a potential promoting effect on the complete denitrification process （N2O→N2） in rhizosphere soil. This not only suppresses nitrogen loss from salt-affected farmland at the source but also promotes N2 reduction. Meanwhile， it shows significant advantages in promoting plant root growth and yield formation， as well as improving the rhizosphere soil ionic environment. These findings provide a theoretical basis for scientific nitrogen application and nitrogen loss control in the Tumuchuan Plain irrigation district.

, authors=Xueqin Fan1, 2, Weiping Li1, 2, Jiapeng Zhang1, 2, Zhaonan Chi1, 2, Dongliang Zhang1, 2, Lixia Cao1, 2, Meng Zhang1, 2, authorsList=Xueqin Fan, Weiping Li, Jiapeng Zhang, Zhaonan Chi, Dongliang Zhang, Lixia Cao, Meng Zhang, authorCompany=null, correspAuthors=Weiping Li, 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=1312009714037252566, articleId=1312009711029936586, tenantId=1146029695717560320, journalId=1311385839672066169, language=CN, title=施氮对盐渍农田向日葵根际土壤细菌反硝化途径与根系生长及产量的影响, columnId=1312009477407204103, journalTitle=水土保持学报, columnName=基础研究, runingTitle=null, highlight=null, articleAbstract=

目的 为探究施氮调控盐渍农田向日葵根际土壤细菌反硝化途径进而促进根系生长及产量的具体途径。 方法 2023—2024年开展膜下滴灌向日葵田间小区试验，以当地常规施氮量为对照（CK：200 kg/hm2），并设减氮25%（N1：150 kg/hm2），减氮50%（N2：100 kg/hm2），减氮75%（N3：50 kg/hm2）共4个处理。 结果 2 a试验表明，适度减氮有助于改善盐渍农田根际土壤环境并促进向日葵根系与产量形成。与CK相比，N1、N2处理均显著提高现蕾期根际土壤Ca2⁺（17.22%、10.56%）、Mg2⁺（34.75%、25.40%）含量，降低Cl-、SO42-含量，同时N1处理显著促进现蕾期根系分泌草酸与柠檬酸（59.07%和60.11%），N2处理显著促进现蕾期和成熟期根系分泌甲酸；功能基因方面，N1、N2、N3处理降低narG、nirK、nirS反硝化功能基因相对丰度，然而N1处理提高nosZ基因的丰度，并改变nosZ型菌群的组成，提高nosZ型Acidobacteria和Proteobacteria的相对丰度。作物生长与产量形成方面，N1、N2处理显著增加开花期根表面积密度（RSAD）与现蕾期根长密度（RLD）值，2 a N1处理显著促进向日葵百粒重（9.07%）、单盘籽粒重（9.08%）增加向日葵产量（5.05%）；相比之下，N2与N3处理显著降低成熟期RSAD值（23.66%、29.46%）、百粒重（21.67%、29.51%）与单盘籽粒重（21.66%、29.49%）。 结论 减氮25%处理对根际土壤反硝化初始过程（NO3-→NO）有潜在的抑制作用，对完全反硝化过程（N2O→N2）有潜在的促进作用，不仅从源头上阻控盐渍农田土壤氮素损失，还能促进N2的还原。同时在促进植株根系生长与产量形成与改善根际土壤离子环境方面具有显著优势。研究结果可为土默川平原灌区科学施氮与阻控氮素损失提供理论参考。

, authors=樊雪琴1, 2, 李为萍1, 2, 张家鹏1, 2, 池曌男1, 2, 张栋良1, 2, 曹丽霞1, 2, 张萌1, 2, authorsList=樊雪琴, 李为萍, 张家鹏, 池曌男, 张栋良, 曹丽霞, 张萌, authorCompany=null, correspAuthors=李为萍, authorNote=

樊雪琴（2000—），女，硕士研究生，主要从事灌溉排水原理与管理决策研究。E-mail：

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李为萍（1978—），女，博士，教授，博士生导师，主要从事灌溉排水原理与管理决策研究。E-mail：
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樊雪琴（2000—），女，硕士研究生，主要从事灌溉排水原理与管理决策研究。E-mail：

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樊雪琴（2000—），女，硕士研究生，主要从事灌溉排水原理与管理决策研究。E-mail：

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注：图中不同小写字母表示不同施氮处理间差异显著（p<0.05）。下同。

, figureFileSmall=jBjdueZXCsDMhveoziTCuQ==, figureFileBig=LknajHO10nksReacbT10hg==, tableContent=null), ArticleFig(id=1312009719162692118, tenantId=1146029695717560320, journalId=1311385839672066169, articleId=1312009711029936586, language=EN, label=Fig.2, caption=Relative abundance of nirK， narG， nosZ， and nirS denitrifying functional genes in sunflower rhizosphere soil under nitrogen application treatments, figureFileSmall=as86W8+lb5FSEWFdeeA9vw==, figureFileBig=I1b5F0+5lZmw4wBZRKjuCw==, tableContent=null), ArticleFig(id=1312009719242383895, tenantId=1146029695717560320, journalId=1311385839672066169, articleId=1312009711029936586, language=CN, label=图2, caption=施氮处理下向日葵根际土壤反硝化细菌 nirK、narG、nosZ、nirS 功能基因的相对丰度

注：narG（NO3-→NO2-）编码的硝酸盐还原酶、nirK（NO2-→NO）编码含铜离子的亚硝酸盐还原酶、nirS（NO2-→NO）编码含细胞色素cd1亚硝酸盐还原酶、nosZ（N2O→N2）编码N2O还原酶；图中对反硝化功能基因的原始 reads counts 进行测序深度标准化，计算其占所有功能基因 reads counts 的比例，图中蓝、粉、红色分别对应向日葵现蕾期、开花期与成熟期。

, figureFileSmall=as86W8+lb5FSEWFdeeA9vw==, figureFileBig=I1b5F0+5lZmw4wBZRKjuCw==, tableContent=null), ArticleFig(id=1312009719322075672, tenantId=1146029695717560320, journalId=1311385839672066169, articleId=1312009711029936586, language=EN, label=Fig.3, caption=Changes in sunflower root indicators under nitrogen application treatments, figureFileSmall=c6f8dMONS1DrbZ0dE7wm6Q==, figureFileBig=qoyUnpN8BA4/kkQZslyJXg==, tableContent=null), ArticleFig(id=1312009719384990233, tenantId=1146029695717560320, journalId=1311385839672066169, articleId=1312009711029936586, language=CN, label=图3, caption=施氮处理下向日葵根系指标的变化, figureFileSmall=c6f8dMONS1DrbZ0dE7wm6Q==, figureFileBig=qoyUnpN8BA4/kkQZslyJXg==, tableContent=null), ArticleFig(id=1312009719456293402, tenantId=1146029695717560320, journalId=1311385839672066169, articleId=1312009711029936586, language=EN, label=Fig.4, caption=Contents of soluble ion components in sunflower rhizosphere soil under nitrogen application treatments, figureFileSmall=/hooRYRi+YisBSdGJ2whmw==, figureFileBig=LQbiUjFWue7o1wJykt4S1g==, tableContent=null), ArticleFig(id=1312009719523402267, tenantId=1146029695717560320, journalId=1311385839672066169, articleId=1312009711029936586, language=CN, label=图4, caption=施氮处理下向日葵根际土壤可溶性离子组分含量, figureFileSmall=/hooRYRi+YisBSdGJ2whmw==, figureFileBig=LQbiUjFWue7o1wJykt4S1g==, tableContent=null), ArticleFig(id=1312009719594705436, tenantId=1146029695717560320, journalId=1311385839672066169, articleId=1312009711029936586, language=EN, label=Fig.5, caption=Correlation analysis among physicochemical factors of sunflower rhizosphere soil， bacterial denitrifying functional genes， and root indicators under nitrogen application treatments, figureFileSmall=3Hoa7vJhgGJHaZEtUUGSpQ==, figureFileBig=B6Im1N121hBUWI6AC1AcTg==, tableContent=null), ArticleFig(id=1312009719666008605, tenantId=1146029695717560320, journalId=1311385839672066169, articleId=1312009711029936586, language=CN, label=图5, caption=施氮处理下向日葵根际土壤理化因子与细菌反硝化功能基因及根系指标的相关性, figureFileSmall=3Hoa7vJhgGJHaZEtUUGSpQ==, figureFileBig=B6Im1N121hBUWI6AC1AcTg==, tableContent=null), ArticleFig(id=1312009719749894686, tenantId=1146029695717560320, journalId=1311385839672066169, articleId=1312009711029936586, language=EN, label=Table 1, caption=

Experimental design table

, figureFileSmall=null, figureFileBig=null, tableContent=
处理灌水定额/（m3·hm-2）

灌溉定额/

（m3·hm-2）

底肥/

（kg·hm-2）

追肥/

（kg·hm-2）

施氮总量/

（kg·hm-2）

春灌现蕾期开花期灌浆期
CK9004502252251 80050150200
N19004502252251 80050100150
N29004502252251 8005050100
N39004502252251 80050050
), ArticleFig(id=1312009719875723807, tenantId=1146029695717560320, journalId=1311385839672066169, articleId=1312009711029936586, language=CN, label=表1, caption=

试验设计

, figureFileSmall=null, figureFileBig=null, tableContent=
处理灌水定额/（m3·hm-2）

灌溉定额/

（m3·hm-2）

底肥/

（kg·hm-2）

追肥/

（kg·hm-2）

施氮总量/

（kg·hm-2）

春灌现蕾期开花期灌浆期
CK9004502252251 80050150200
N19004502252251 80050100150
N29004502252251 8005050100
N39004502252251 80050050
), ArticleFig(id=1312009720177713696, tenantId=1146029695717560320, journalId=1311385839672066169, articleId=1312009711029936586, language=EN, label=Table 2, caption=

Concentrations of organic acids secreted by roots under nitrogen application treatments

, figureFileSmall=null, figureFileBig=null, tableContent=
年份生育期处理甲酸苹果酸草酸乙酸柠檬酸
2023现蕾期CK116.71±5.84c627.00±31.35c1 666.14±83.31c10.04±0.50c10.78±0.54c
N1172.07±8.60a994.86±49.74a2 650.37±132.52a36.82±1.84a17.26±1.06a
N2132.50±6.63b774.40±38.72b1 855.01±92.75b34.73±1.74a16.03±0.80b
N3108.71±5.44c502.34±25.12d1 248.53±62.43d18.87±0.94b15.36±0.77b
开花期CK110.63±5.53c938.74±46.94b2 306.83±115.34b11.85±0.59c20.13±1.01b
N1175.26±8.76a1 672.36±83.62a3 521.65±176.08a69.99±3.50a33.91±1.70a
N2131.08±6.55b1 013.10±50.66b2 519.58±125.98b29.66±1.48b21.34±1.07b
N3107.36±5.37c629.20±31.46c879.26±43.96c7.95±0.40d9.93±0.50c
成熟期CK82.91±4.15c934.02±46.70b1 393.70±69.69b2.67±0.13c12.76±0.64c
N1139.12±6.96a1 118.70±55.94a1 785.55±89.28a15.81±0.79a16.03±0.80a
N295.18±4.76b1 007.60±50.38b1 216.37±60.82c9.38±0.47b14.08±0.70b
N363.50±3.18d675.76±33.79c469.84±23.49d9.60±0.48b6.24±0.31d
2024现蕾期CK140.05±7.00c645.81±32.29c1 649.48±82.47c11.04±0.55c16.26±0.81b
N1206.48±10.32a1 023.68±51.18a2 623.87±131.19a40.50±2.03a23.39±1.17a
N2159.00±7.95b797.63±39.88b1 836.46±91.82b38.20±1.91a17.63±0.88b
N3130.45±6.52c517.41±25.87d1 236.04±61.80d20.76±1.04b16.90±0.84b
开花期CK132.76±6.64c965.87±48.29b2 283.76±114.19b13.04±0.65c22.14±1.11b
N1210.31±10.52a1 722.53±86.13a3 486.43±174.32a76.99±3.85a37.30±1.87a
N2157.30±7.86b1 043.49±52.17b2 494.38±124.72b32.63±1.63b23.47±1.17b
N3128.83±6.44c648.08±32.40c870.47±43.52c8.75±0.44d10.92±0.55c
成熟期CK99.49±4.97c962.04±48.10b1 379.76±68.99b2.94±0.15c14.04±0.70c
N1166.94±8.35a1 152.26±57.61a1 767.69±88.38a17.39±0.87a17.63±0.88a
N2114.22±5.71b1 037.83±51.89b1 402.21±70.11b10.32±0.52b15.49±0.77b
N376.20±3.81d696.03±34.80c465.14±23.26c10.56±0.53b6.86±0.34d
), ArticleFig(id=1312009720571978273, tenantId=1146029695717560320, journalId=1311385839672066169, articleId=1312009711029936586, language=CN, label=表2, caption=

施氮处理下根系分泌的有机酸浓度 (μg/g)

, figureFileSmall=null, figureFileBig=null, tableContent=
年份生育期处理甲酸苹果酸草酸乙酸柠檬酸
2023现蕾期CK116.71±5.84c627.00±31.35c1 666.14±83.31c10.04±0.50c10.78±0.54c
N1172.07±8.60a994.86±49.74a2 650.37±132.52a36.82±1.84a17.26±1.06a
N2132.50±6.63b774.40±38.72b1 855.01±92.75b34.73±1.74a16.03±0.80b
N3108.71±5.44c502.34±25.12d1 248.53±62.43d18.87±0.94b15.36±0.77b
开花期CK110.63±5.53c938.74±46.94b2 306.83±115.34b11.85±0.59c20.13±1.01b
N1175.26±8.76a1 672.36±83.62a3 521.65±176.08a69.99±3.50a33.91±1.70a
N2131.08±6.55b1 013.10±50.66b2 519.58±125.98b29.66±1.48b21.34±1.07b
N3107.36±5.37c629.20±31.46c879.26±43.96c7.95±0.40d9.93±0.50c
成熟期CK82.91±4.15c934.02±46.70b1 393.70±69.69b2.67±0.13c12.76±0.64c
N1139.12±6.96a1 118.70±55.94a1 785.55±89.28a15.81±0.79a16.03±0.80a
N295.18±4.76b1 007.60±50.38b1 216.37±60.82c9.38±0.47b14.08±0.70b
N363.50±3.18d675.76±33.79c469.84±23.49d9.60±0.48b6.24±0.31d
2024现蕾期CK140.05±7.00c645.81±32.29c1 649.48±82.47c11.04±0.55c16.26±0.81b
N1206.48±10.32a1 023.68±51.18a2 623.87±131.19a40.50±2.03a23.39±1.17a
N2159.00±7.95b797.63±39.88b1 836.46±91.82b38.20±1.91a17.63±0.88b
N3130.45±6.52c517.41±25.87d1 236.04±61.80d20.76±1.04b16.90±0.84b
开花期CK132.76±6.64c965.87±48.29b2 283.76±114.19b13.04±0.65c22.14±1.11b
N1210.31±10.52a1 722.53±86.13a3 486.43±174.32a76.99±3.85a37.30±1.87a
N2157.30±7.86b1 043.49±52.17b2 494.38±124.72b32.63±1.63b23.47±1.17b
N3128.83±6.44c648.08±32.40c870.47±43.52c8.75±0.44d10.92±0.55c
成熟期CK99.49±4.97c962.04±48.10b1 379.76±68.99b2.94±0.15c14.04±0.70c
N1166.94±8.35a1 152.26±57.61a1 767.69±88.38a17.39±0.87a17.63±0.88a
N2114.22±5.71b1 037.83±51.89b1 402.21±70.11b10.32±0.52b15.49±0.77b
N376.20±3.81d696.03±34.80c465.14±23.26c10.56±0.53b6.86±0.34d
), ArticleFig(id=1312009720681030178, tenantId=1146029695717560320, journalId=1311385839672066169, articleId=1312009711029936586, language=EN, label=Table 3, caption=

Yield indicators of sunflower under nitrogen application treatments

, figureFileSmall=null, figureFileBig=null, tableContent=
年份处理产量/（kg·hm-2）百粒重/g单盘籽粒重/g
2023CK3 622.19±181.11a24.56±1.23b199.14±9.96b
N13 761.92±188.10a26.79±1.34a217.21±10.86a
N23 562.22±178.11a20.32±1.02c147.25±7.36c
N33 027.89±151.40b18.29±0.92d132.53±6.63c
2024CK3 604.29±180.21a24.08±1.20b195.23±9.76b
N13 828.96±191.45b26.26±1.31a212.95±10.65a
N23 221.06±161.05c17.80±0.89c161.54±8.08c
N32 737.90±136.90d16.02±0.80c145.39±7.27d
), ArticleFig(id=1312009721083683364, tenantId=1146029695717560320, journalId=1311385839672066169, articleId=1312009711029936586, language=CN, label=表3, caption=

施氮处理下向日葵产量指标

, figureFileSmall=null, figureFileBig=null, tableContent=
年份处理产量/（kg·hm-2）百粒重/g单盘籽粒重/g
2023CK3 622.19±181.11a24.56±1.23b199.14±9.96b
N13 761.92±188.10a26.79±1.34a217.21±10.86a
N23 562.22±178.11a20.32±1.02c147.25±7.36c
N33 027.89±151.40b18.29±0.92d132.53±6.63c
2024CK3 604.29±180.21a24.08±1.20b195.23±9.76b
N13 828.96±191.45b26.26±1.31a212.95±10.65a
N23 221.06±161.05c17.80±0.89c161.54±8.08c
N32 737.90±136.90d16.02±0.80c145.39±7.27d
), ArticleFig(id=1312009721188540965, tenantId=1146029695717560320, journalId=1311385839672066169, articleId=1312009711029936586, language=EN, label=Table 4, caption=

Physicochemical properties of sunflower rhizosphere soil at different growth stages under nitrogen application treatments in 2023

, figureFileSmall=null, figureFileBig=null, tableContent=
生育期处理pHNO3--N/（mg·kg-1）NH4+-N/（mg·kg-1）SOM/（g·kg-1）
现蕾期CK8.45±0.42ab36.16±1.81a33.58±1.52b9.88±0.49a
N18.02±0.40b33.75±1.69a38.14±1.73a9.81±0.34a
N28.75±0.44ab28.13±1.41b33.56±1.43b7.63±0.38b
N38.99±0.45a26.44±1.32b30.05±1.36c6.95±0.35b
开花期CK8.55±0.43c30.74±1.54a39.06±1.77b11.40±0.57a
N18.22±0.41d32.74±1.64a45.45±2.06a11.80±0.59a
N28.75±0.44b24.55±1.23b37.03±1.68b11.00±0.55a
N39.10±0.46a24.45±1.22b33.48±1.52c8.18±0.41b
成熟期CK8.76±0.44b27.13±1.36a36.02±1.63b13.60±0.68a
N18.30±0.42c30.15±1.51a41.09±1.86a15.60±0.63a
N28.85±0.44b23.10±1.16b35.28±1.51b10.20±0.51b
N39.16±0.46a22.11±1.11b32.47±1.47c8.43±0.42c
), ArticleFig(id=1312009722849485350, tenantId=1146029695717560320, journalId=1311385839672066169, articleId=1312009711029936586, language=CN, label=表4, caption=

施氮处理下2023年向日葵生育期根际土壤理化性质

, figureFileSmall=null, figureFileBig=null, tableContent=
生育期处理pHNO3--N/（mg·kg-1）NH4+-N/（mg·kg-1）SOM/（g·kg-1）
现蕾期CK8.45±0.42ab36.16±1.81a33.58±1.52b9.88±0.49a
N18.02±0.40b33.75±1.69a38.14±1.73a9.81±0.34a
N28.75±0.44ab28.13±1.41b33.56±1.43b7.63±0.38b
N38.99±0.45a26.44±1.32b30.05±1.36c6.95±0.35b
开花期CK8.55±0.43c30.74±1.54a39.06±1.77b11.40±0.57a
N18.22±0.41d32.74±1.64a45.45±2.06a11.80±0.59a
N28.75±0.44b24.55±1.23b37.03±1.68b11.00±0.55a
N39.10±0.46a24.45±1.22b33.48±1.52c8.18±0.41b
成熟期CK8.76±0.44b27.13±1.36a36.02±1.63b13.60±0.68a
N18.30±0.42c30.15±1.51a41.09±1.86a15.60±0.63a
N28.85±0.44b23.10±1.16b35.28±1.51b10.20±0.51b
N39.16±0.46a22.11±1.11b32.47±1.47c8.43±0.42c
), ArticleFig(id=1312009722962731559, tenantId=1146029695717560320, journalId=1311385839672066169, articleId=1312009711029936586, language=EN, label=Table 5, caption=

Physicochemical properties of sunflower rhizosphere soil at different growth stages under nitrogen application treatments in 2024

, figureFileSmall=null, figureFileBig=null, tableContent=
生育期处理pHNO3--N/（mg·kg-1）NH4+-N/（mg·kg-1）SOM/（g·kg-1）
现蕾期CK8.37±0.42ab39.78±1.99a36.94±1.85b11.07±0.55a
N17.94±0.40b37.13±1.86a41.95±2.10a11.34±0.72a
N28.66±0.43ab30.94±1.55b35.72±1.74b8.55±0.43b
N38.90±0.45a29.08±1.45b33.06±1.65c7.78±0.39b
开花期CK8.46±0.42a33.81±1.69a42.97±2.15b12.77±0.64a
N18.14±0.41a36.01±1.80a50.00±2.50a13.22±0.66a
N28.66±0.43a27.01±1.35b40.73±2.04b9.32±0.62b
N39.01±0.45a26.90±1.34b36.83±1.84c9.16±0.46b
成熟期CK8.67±0.43a29.84±1.49b39.62±1.98b15.23±0.76b
N18.22±0.41a33.17±1.66a45.20±2.26a17.47±0.87a
N28.76±0.44a25.41±1.27c38.61±1.83b11.42±0.57c
N39.07±0.45a24.32±1.22c35.72±1.79c9.44±0.47d
), ArticleFig(id=1312009723046617640, tenantId=1146029695717560320, journalId=1311385839672066169, articleId=1312009711029936586, language=CN, label=表5, caption=

施氮处理下2024年向日葵生育期根际土壤理化性质

, figureFileSmall=null, figureFileBig=null, tableContent=
生育期处理pHNO3--N/（mg·kg-1）NH4+-N/（mg·kg-1）SOM/（g·kg-1）
现蕾期CK8.37±0.42ab39.78±1.99a36.94±1.85b11.07±0.55a
N17.94±0.40b37.13±1.86a41.95±2.10a11.34±0.72a
N28.66±0.43ab30.94±1.55b35.72±1.74b8.55±0.43b
N38.90±0.45a29.08±1.45b33.06±1.65c7.78±0.39b
开花期CK8.46±0.42a33.81±1.69a42.97±2.15b12.77±0.64a
N18.14±0.41a36.01±1.80a50.00±2.50a13.22±0.66a
N28.66±0.43a27.01±1.35b40.73±2.04b9.32±0.62b
N39.01±0.45a26.90±1.34b36.83±1.84c9.16±0.46b
成熟期CK8.67±0.43a29.84±1.49b39.62±1.98b15.23±0.76b
N18.22±0.41a33.17±1.66a45.20±2.26a17.47±0.87a
N28.76±0.44a25.41±1.27c38.61±1.83b11.42±0.57c
N39.07±0.45a24.32±1.22c35.72±1.79c9.44±0.47d
), ArticleFig(id=1312009723126309417, tenantId=1146029695717560320, journalId=1311385839672066169, articleId=1312009711029936586, language=EN, label=Table 6, caption=

Principal component scoring coefficient matrix of rhizosphere soil physicochemical properties， bac-terial denitrification functional genes， crop roots， and yield under nitrogen application treatments

, figureFileSmall=null, figureFileBig=null, tableContent=
指标主成分1主成分2
RSAD0.952-0.194
产量0.8520.491
NH4+-N0.940-0.285
柠檬酸0.8960.280
Ca2+0.273-0.857
Mg2+0.8440.293
nosZ0.735-0.611
nirK0.2670.924
特征值4.6232.299
贡献率/%57.79057.790
累积贡献率/%28.74086.530
), ArticleFig(id=1312009723193418282, tenantId=1146029695717560320, journalId=1311385839672066169, articleId=1312009711029936586, language=CN, label=表6, caption=

施氮处理下根际土壤理化性质、细菌反硝化功能基因、作物根系与产量的主成分评分系数矩阵

, figureFileSmall=null, figureFileBig=null, tableContent=
指标主成分1主成分2
RSAD0.952-0.194
产量0.8520.491
NH4+-N0.940-0.285
柠檬酸0.8960.280
Ca2+0.273-0.857
Mg2+0.8440.293
nosZ0.735-0.611
nirK0.2670.924
特征值4.6232.299
贡献率/%57.79057.790
累积贡献率/%28.74086.530
), ArticleFig(id=1312009723260527147, tenantId=1146029695717560320, journalId=1311385839672066169, articleId=1312009711029936586, language=EN, label=Table 7, caption=

Comprehensive scores and rankings among rhi-zosphere soil physicochemical properties， bacte-rial denitrifying functional genes， crop roots， and yield under nitrogen application treatments

, figureFileSmall=null, figureFileBig=null, tableContent=
年份处理隶属函数值（F1）隶属函数值（F2）

综合

得分

排序
2023CK0.470.980.642
N10.690.600.661
N20.360.970.563
N30.090.510.234
2024CK0.630.650.642
N10.890.250.681
N20.520.520.523
N30.230.020.164
), ArticleFig(id=1312009723331830316, tenantId=1146029695717560320, journalId=1311385839672066169, articleId=1312009711029936586, language=CN, label=表7, caption=

施氮处理下根际土壤理化性质、细菌反硝化功能基因、作物根系与产量间的综合评分及排序

, figureFileSmall=null, figureFileBig=null, tableContent=
年份处理隶属函数值（F1）隶属函数值（F2）

综合

得分

排序
2023CK0.470.980.642
N10.690.600.661
N20.360.970.563
N30.090.510.234
2024CK0.630.650.642
N10.890.250.681
N20.520.520.523
N30.230.020.164
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施氮对盐渍农田向日葵根际土壤细菌反硝化途径与根系生长及产量的影响
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樊雪琴 1, 2 , 李为萍 1, 2 , 张家鹏 1, 2 , 池曌男 1, 2 , 张栋良 1, 2 , 曹丽霞 1, 2 , 张萌 1, 2
水土保持学报 | 基础研究 2026,40(4): 205-215
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水土保持学报 |基础研究 2026 , 40 (4) : 205 -215
施氮对盐渍农田向日葵根际土壤细菌反硝化途径与根系生长及产量的影响
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樊雪琴1, 2 , 李为萍1, 2 , 张家鹏1, 2, 池曌男1, 2, 张栋良1, 2, 曹丽霞1, 2, 张萌1, 2
作者信息
  • 1.内蒙古农业大学旱区水工程生态环境全国重点实验室，呼和浩特 010018
  • 2.内蒙古农业大学水利与土木建筑工程学院，呼和浩特 010018
通讯作者:
李为萍（1978—），女，博士，教授，博士生导师，主要从事灌溉排水原理与管理决策研究。E-mail：
作者简介:

樊雪琴（2000—），女，硕士研究生，主要从事灌溉排水原理与管理决策研究。E-mail：

Effects of Nitrogen Application on Rhizosphere Bacterial Denitrification Pathways， Root Growth， and Yield of Sunflowers in Salt-Affected Farmland
Xueqin Fan1, 2 , Weiping Li1, 2 , Jiapeng Zhang1, 2, Zhaonan Chi1, 2, Dongliang Zhang1, 2, Lixia Cao1, 2, Meng Zhang1, 2
Affiliations
  • 1.State Key Laboratory of Water Engineering Ecology and Environment in Arid Area，Inner Mongolia;Agricultural University，Hohhot 010018，China
  • 2.College of Water Resources and Civil;Engineering，Inner Mongolia Agricultural University，Hohhot 010018，China
出版时间: 2026-08-28 doi: 10.13870/j.cnki.stbcxb.2026.04.040
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摘要
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目的 为探究施氮调控盐渍农田向日葵根际土壤细菌反硝化途径进而促进根系生长及产量的具体途径。 方法 2023—2024年开展膜下滴灌向日葵田间小区试验，以当地常规施氮量为对照（CK：200 kg/hm2），并设减氮25%（N1：150 kg/hm2），减氮50%（N2：100 kg/hm2），减氮75%（N3：50 kg/hm2）共4个处理。 结果 2 a试验表明，适度减氮有助于改善盐渍农田根际土壤环境并促进向日葵根系与产量形成。与CK相比，N1、N2处理均显著提高现蕾期根际土壤Ca2⁺（17.22%、10.56%）、Mg2⁺（34.75%、25.40%）含量，降低Cl-、SO42-含量，同时N1处理显著促进现蕾期根系分泌草酸与柠檬酸（59.07%和60.11%），N2处理显著促进现蕾期和成熟期根系分泌甲酸；功能基因方面，N1、N2、N3处理降低narG、nirK、nirS反硝化功能基因相对丰度，然而N1处理提高nosZ基因的丰度，并改变nosZ型菌群的组成，提高nosZ型Acidobacteria和Proteobacteria的相对丰度。作物生长与产量形成方面，N1、N2处理显著增加开花期根表面积密度（RSAD）与现蕾期根长密度（RLD）值，2 a N1处理显著促进向日葵百粒重（9.07%）、单盘籽粒重（9.08%）增加向日葵产量（5.05%）；相比之下，N2与N3处理显著降低成熟期RSAD值（23.66%、29.46%）、百粒重（21.67%、29.51%）与单盘籽粒重（21.66%、29.49%）。 结论 减氮25%处理对根际土壤反硝化初始过程（NO3-→NO）有潜在的抑制作用，对完全反硝化过程（N2O→N2）有潜在的促进作用，不仅从源头上阻控盐渍农田土壤氮素损失，还能促进N2的还原。同时在促进植株根系生长与产量形成与改善根际土壤离子环境方面具有显著优势。研究结果可为土默川平原灌区科学施氮与阻控氮素损失提供理论参考。

根际细菌  /  盐渍土  /  向日葵  /  反硝化途径  /  施氮  /  宏基因组测序

Objective This study aims to investigate the specific mechanisms by which nitrogen application regulates rhizosphere bacterial denitrification pathways in salt-affected farmland， thereby promoting sunflower root growth and yield. Methods Field plot experiments with drip irrigation under plastic mulch were conducted in sunflower farmland from 2023 to 2024. Four nitrogen application levels were set： the local conventional nitrogen application rate as control （CK： 200 kg/hm2）， 25% nitrogen reduction （N1： 150 kg/hm2）， 50% nitrogen reduction （N2： 100 kg/hm2）， and 75% nitrogen reduction （N3： 50 kg/hm2）. Results The two-year experiments demonstrated that moderate nitrogen reduction improved the rhizosphere soil environment in salt-affected farmland and promoted sunflower root growth and yield formation. Compared with CK， both N1 and N2 treatments significantly increased Ca2+ （17.22%， 10.56%） and Mg2+ （34.75%， 25.40%） contents in rhizosphere soil at the budding stage， and reduced Cl- and SO42- contents. Furthermore， N1 treatment significantly promoted oxalic acid and citric acid secretion by root at the budding stage （59.07% and 60.11%）， and N2 treatment significantly promoted formic acid secretion by root at both the budding stage and the maturity stage. Regarding functional genes， N1， N2， and N3 treatments reduced the relative abundance of denitrifying genes narG， nirK， and nirS， whereas N1 treatment increased the abundance of the nosZ gene and altered the composition of nosZ-type bacterial communities， including increasing the relative abundance of nosZ-type Acidobacteria and Proteobacteria. Regarding crop growth and yield formation， N1 and N2 treatments significantly increased root surface area density （RSAD） at the flowering stage and root length density （RLD） at the budding stage. Over two years， N1 treatment significantly increased sunflower 100-seed weight （9.07%）， seed weight per head （9.08%）， and yield （5.05%）. In contrast， N2 and N3 treatments significantly reduced RSAD values （23.66%， 29.46%）， 100-seed weight （21.67%， 29.51%）， and seed weight per head （21.66%， 29.49%） at the maturity stage. Conclusion A 25% nitrogen reduction treatment has a potential inhibitory effect on the initial stage of denitrification （NO3-→NO） and a potential promoting effect on the complete denitrification process （N2O→N2） in rhizosphere soil. This not only suppresses nitrogen loss from salt-affected farmland at the source but also promotes N2 reduction. Meanwhile， it shows significant advantages in promoting plant root growth and yield formation， as well as improving the rhizosphere soil ionic environment. These findings provide a theoretical basis for scientific nitrogen application and nitrogen loss control in the Tumuchuan Plain irrigation district.

rhizosphere bacteria  /  salt-affected soil  /  sunflower  /  denitrification pathway  /  nitrogen application  /  metagenomic sequencing
樊雪琴, 李为萍, 张家鹏, 池曌男, 张栋良, 曹丽霞, 张萌. 施氮对盐渍农田向日葵根际土壤细菌反硝化途径与根系生长及产量的影响. 水土保持学报, 2026 , 40 (4) : 205 -215 . DOI: 10.13870/j.cnki.stbcxb.2026.04.040
Xueqin Fan, Weiping Li, Jiapeng Zhang, Zhaonan Chi, Dongliang Zhang, Lixia Cao, Meng Zhang. Effects of Nitrogen Application on Rhizosphere Bacterial Denitrification Pathways， Root Growth， and Yield of Sunflowers in Salt-Affected Farmland[J]. Journal of Soil and Water Conservation, 2026 , 40 (4) : 205 -215 . DOI: 10.13870/j.cnki.stbcxb.2026.04.040
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反硝化作用是土壤氮素流失的关键途径之一，该过程由多种微生物协同完成，涉及一系列关键功能基因［1］，不仅加剧全球气候变暖，还对臭氧层具有显著的破坏作用［2］。盐渍农田是我国重要的后备耕地资源，已有研究［3-5］多集中于土壤改良，而对其中关键氮循环过程-反硝化作用的研究仍较为薄弱［6］，盐渍农田中氮肥利用率普遍低于常规农田［7］，高氮输入，一方面抑制固氮微生物功能基因nifH表达［8］；另一方面，为反硝化过程提供过量底物（NO3-），从而加速N2O排放［9］，导致氮素气态损失大幅增加。此外，高施氮肥常因离子拮抗，降低植物体内GS/GOGAT途径对NH4+的同化速率［10］，导致土壤中NH4+累积；累积的NH4+经硝化作用生成大量NO3-，而NO3-不易被土壤吸附，易随水淋失，进而导致地下水或地表水的面源污染。白芳芳等［11］研究表明，减氮处理可显著降低农田土壤中反硝化微生物的功能基因丰度，并显著下调编码亚硝酸盐还原酶的关键功能基因（如nirK和nirS）及编码氧化亚氮还原酶的基因（nosZ）的丰度［12］。适度减氮不仅抑制微生物反硝化作用，还可激活植物的抗逆性，促进根系分泌柠檬酸、苹果酸等有机酸［13］。该类有机酸既可螯合Na⁺离子缓解盐胁迫下的渗透胁迫，也能释放被碳酸钙固定的Ca2⁺，提高土壤中钙的有效性［14-15］。已有研究［16-18］多聚焦于湿地、高原湖泊、河口沉积物等典型生态系统中的反硝化作用，然而在盐渍逆境作用下，根际土壤细菌反硝化途径与根系生长及产量形成对不同施氮量的响应尚不明确。为此，本研究以轻中度盐渍向日葵根际土壤为研究对象，基于田间小区定位试验与宏基因组测序技术，探究不同施氮量对盐渍农田根际土壤细菌反硝化途径、根系生长及产量的影响，以期为盐渍农田向日葵氮肥合理施用提供理论依据。
1 材料与方法
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1.1 试验区概况
2023年和2024年5—9月于内蒙古农业大学海流图科技园区农业水土工程试验基地（40°41'30''N，111°02'30''E）开展田间小区试验，试验基地位于土默川平原灌区，属典型温带大陆性季风气候，多年平均降水量为421.6 mm，多年平均气温6.3 ℃，无霜期135 d。2023年向日葵生育期内（5—9月）平均气温21.2 ℃，年日照时间为2 861.3 h，生育期降水量314.6 mm，蒸发量为1 868.1 mm。2024年向日葵生育期平均气温22.9 ℃，年日照时间为2 746.2 h，生育期降水量589.9 mm，蒸发量为1 842.2 mm。试验地0~40 cm土层土壤质地为砂质壤土，体积质量（BD）为1.73 g/cm3，土壤平均全盐量为3.22 g/kg，属轻中度盐渍土。土壤表层基础养分有机质（SOM）为16.79 g/kg，全氮（TN）为990.3 mg/kg，水解性氮（HN）为63.51 mg/kg，有效磷（AP）为43.66 mg/kg，速效钾（AK）为216.38 mg/kg，向日葵整个生育期地下水埋深为3~5 m。
1.2 试验设计
供试作物为食用向日葵“HZ2 399”，播种前对盐渍农田进行春灌压盐（灌水定额900 m3/hm2），春灌采用畦灌，向日葵生育期采用膜下滴灌，底肥施用磷酸二胺（含N 18%），底肥折合纯氮50 kg/hm2，现蕾期追施尿素（含N 46%）。以当地施肥量为对照CK（追施纯氮150 kg/hm2），另设3个减氮梯度，减氮25%（N1∶追氮100 kg/hm2）、减氮50%（N2∶追氮50 kg/hm2）、减氮75%（N3∶0 kg/hm2），该设计采用养分管理研究中的经典梯度设置方法，即通过设定基于常规施肥量的百分比减量梯度，旨在系统探究盐渍农田中作物-土壤系统对氮输入减少的非线性响应关系。试验采用单因素完全随机区组设计，小区规格为10 m×16 m，每个处理设置3次重复，共计15个小区，小区四周砌筑水泥挡板，以防止水、氮、盐的交互影响。每个小区铺设6膜，每膜种植2行，行距50 cm，株距50 cm，种植密度21 000株/hm2，各小区种植密度一致，具体试验设计见表1。
1.3 测定项目与方法
1.3.1 样品采集
试验采用五点取样法，灌水后分别于向日葵关键生育期使用根钻法取出根系并抖落根系附着土壤。5个点土样混和均匀并装入无菌自封袋并分为3份：1份置于-80 ℃下保存用于DNA提取和宏基因组测序；1份室内自然风干，研磨后过2 mm筛，用于土壤理化性质测试分析；1份用于根系分泌物的测定。使用pH计（梅特勒-托利多仪器，上海）测定土水质量比为1∶2.5的土壤酸碱度（pH），土壤铵态氮使用KCl（2 mol/L）浸提靛酚蓝比色法，土壤硝态氮按水土比10∶1加入50 mL KCl（1 mg/L）溶液，采用紫外分光光度计测定。土壤有机质（SOM）含量测定参照NY/T 1121.6—2006。根际有机酸获取采用乙醇浸提法，使用GC-MS联用仪（HP6890/5975C，美国）进行分析。
1.3.2 宏基因组测序及生物信息
称取500 mg glass beads和0.2~0.5 g的土壤，放入2 mL离心管并添加Buffer SLX Mlus进行振荡，加入80 μL Buffer DS孵育10 min，再离心5 min。转移600 μL上清液至2 mL离心管中，加入200 μL Buffer SP 2，振荡后加入100 μL HTR Reagent，冰浴5 min后离心5 min，移上清液400 μL至离心管中，加入40 μL的磁珠，混合均匀加入450 μL Binding Buffe，室温放置2 min。使用Qubit 4荧光计检测DNA的浓度，并用1%的琼脂糖凝胶电泳检测DNA的质量。调整DNA溶液浓度。多样品DNA文库均一化至10 nM后等体积混合，将混合好的文库（10 nM）稀释至合适的浓度后进行上机测序。
1.3.3 向日葵根系指标与产量测定
使用根钻采用五点取样法进行根系样本采集，将取得的根系样本用清水清洗干净后，采用根系分析系统（WinRHIZO），精准获取根系长度、根系表面积及根系体积等关键数据，进一步通过运算推导出根长密度（RLD）、根体积密度（RVD）及根表面积密度（RSAD）等参数。在向日葵成熟期时，在每个小区随机选取10株长势较为一致的向日葵，各小区分别测产。并对各小区定株的向日葵花盘上，选取最外侧3~4圈的位置摘取籽粒，测百粒重。
1.4 数据分析
采用Excel 2016软件进行数据的处理，使用SPSS.27软件进行单因素方差分析及差异显著性水平（p<0.05）分析，用最小显著差异法进行显著性检验。用Origin 2023软件绘制反硝化功能基因相对丰度图、门水平群落相对丰度图与根系指标图。文中图表数据均为2 a试验结果的平均值，用以体现其共性规律；2 a试验结果不一致之处则单独说明。
2 结果与分析
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2.1 施氮对根际土壤反硝化细菌群落组成及功能基因的影响
施氮显著改变向日葵根际土壤反硝化细菌的群落组成（图1）。与CK处理相比，N2与N3处理显著降低根际土壤中norB、nirK型Proteobacteria的相对丰度，同时显著降低narG、nirS型Acidobacteria相对丰度。值得关注的是，N1与N2处理在调控微生物群落结构方面表现出积极效应，2023年N1处理显著提高nosZ型Proteobacteria（30.03%），2024年N1与N2处理显著提高nosZ型Proteobacteria（24.13%、47.20%）与nosZ型Acidobacteria（40.32%、11.16%）相对丰度，该群落结构得到优化，尤其是完全反硝化细菌群落的富集，有助于推动根际土壤细菌介导的完全反硝化途径（N2O→N2），从而降低N2O的排放潜力。
由图2可知，施氮处理下向日葵根际土壤反硝化功能基因相对丰度。随着施氮量的减少，功能基因nirK、narG、nirS等基因的相对丰度呈逐步下降趋势。然而施氮处理下编码氧化亚氮还原酶的nosZ基因相对丰度由大到小排序为N1>CK>N2>N3，表明减氮25%（N1处理）在抑制根际土壤细菌不完全反硝化功能基因（narG、nirK、nirS）表达的同时，促进完全反硝化功能基因（nosZ）的富集，说明减氮25%处理不仅从源头阻控根际土壤氮素损失，还促进N2O向N2还原，降低盐渍农田氮素损失的同时减少温室气体N2O的排放潜力。
2.2 施氮对向日葵根系生长与产量形成的影响
2 a试验表明，适量减氮对根系生长与产量形成具有积极促进作用（图3、表3）。与CK处理相比，N1、N2处理在开花期显著提高RSAD值，增幅分别为13.40%、10.70%，N1与N2处理还显著增加现蕾期与成熟期根系RLD值；从产量构成来看，N1处理增加向日葵产量（5.05%），并显著提升向日葵百粒重（9.07%）、单盘籽粒重（9.08%）；相比之下，N2与N3处理则显著降低成熟期RSAD值（23.66%、29.46%），N2、N3处理还降低向日葵产量（6.14%、20.22%），并显著降低百粒重（21.67%、29.51%）、单盘籽粒重（21.66%、29.49%）。综上得出，减氮25%处理（N1）促进关键生育期根系发育，最终有利于产量形成。根系分泌物分析表明，各处理下根系分泌的优势有机酸为草酸、甲酸和苹果酸，也检测到乙酸和柠檬酸（表2）。N1处理显著促进现蕾期根系分泌草酸与柠檬酸（59.07%、60.11%）；N2处理显著促进现蕾期和成熟期根系分泌甲酸（14.17%）。然而，当氮素水平低至N3处理（50 kg/hm2）时，成熟期甲酸分泌受到抑制，且现蕾期、开花期与成熟期根系分泌苹果酸与草酸分泌量均显著降低。总体而言，适度减氮（尤其是N1处理）能够激发根系有机酸的分泌，可能与其在促进根系生长和产量形成中发挥的积极作用有关。
2.3 施氮条件下根际土壤理化性质与细菌反硝化途径及根系产量间的综合分析
由表4、表5可知，施氮处理下向日葵生育期根际土壤的理化性质。根际土壤中NO3--N与NH4+-N含量随生育进程推进呈逐渐下降趋势。与CK处理相比，N2和N3处理的现蕾期根际土壤NO3--N含量均显著降低（22.21%、26.89%）。N1处理显著提高现蕾期根际土壤NH4+-N含量（13.57%），而N3处理显著降低现蕾期与成熟期NH4+-N含量。N1与CK处理的现蕾期、开花期与成熟期根际土壤SOM含量无显著差异，然而N2、N3处理显著降低现蕾期与成熟期根际土壤SOM含量，说明过量减氮可能影响土壤养分的维持能力。
由图4可知，施氮处理的根际土壤可溶性离子含量。阴离子方面，N1、N2、N3处理显著降低现蕾期、开花期和成熟期的根际土壤Cl-和SO42-含量；阳离子方面，N1和N2处理均显著提高现蕾期根际土壤Ca2+（17.22%、10.56%）和Mg2+（34.75%、25.40%）含量。N2与N3处理显著降低现蕾期、开花期与成熟期K++Na+含量；N1处理的现蕾期根际土壤K⁺+Na⁺的含量较CK处理无显著差异。综上，适量减氮（N1、N2）处理有助于降低现蕾期根际土壤中Cl-、SO42-含量，同时促进Ca2+、Mg2+等有益阳离子的累积，可能有助于改善根际离子环境，为根系生长发育提供更为均衡的矿质营养条件。
由图5可知，根际土壤理化因子与向日葵根系生长指标皮尔逊相关性分析结果。现蕾期反硝化功能基因narG与SOM、NO3--N和NH4+-N均呈极显著正相关，与HCO3-、Ca2+、TN及甲酸含量呈显著正相关。nosZ与柠檬酸、甲酸及NH4+-N呈极显著正相关，与Mg2+、Ca2+和苹果酸呈显著正相关，而与pH和Cl-呈显著负相关，表明现蕾期反硝化功能基因与根际环境因子关系密切，可能与该阶段根系分泌物活跃、植株氮素需求较高有关。开花期功能基因narG和nirK与甲酸呈显著正相关，与Cl-呈显著负相关，开花期反硝化功能基因与土壤环境因子的相关性较现蕾期有所减弱。成熟期基因narG、nirS和nosZ均与Ca2+和乙酸呈显著正相关。在根系形态方面，现蕾期与开花期的根系表面积密度（RSAD）和根系体积密度（RVD）均与K++Na+和SO42-呈极显著正相关，至成熟期，RSAD、RVD及根长密度（RLD）与HCO3-、K++Na+、SO42-呈显著正相关，表明根际土壤中的盐分离子（尤其是K++Na+和SO42-）可能是调控向日葵根系生长的关键环境因子。
采用主成分分析对2 a施氮处理下成熟期根际土壤理化性质、细菌反硝化功能基因、作物根系与产量指标进行综合评价（表6、表7），前2个主成分的特征值>1，累计贡献率达到86.53%，数据具有可信度。因此，选择2个主成分作为综合评价指标并计算各处理的综合得分。结果显示，不同处理组综合得分排序为N1>CK>N2>N3，说明减氮25%（N1）在改善根际土壤环境、优化反硝化功能基因组成、促进根系生长及提高产量方面的综合效果最佳，而过度减氮（N3）则表现最差。
3 讨 论
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3.1 施氮处理对根际土壤养分与向日葵产量及根系生长和产量的影响
适度减氮有利于提高根长密度（RLD）与根表面积密度（RSAD）［19］，本研究减氮25%与50%处理显著增加现蕾期RLD与开花期RSAD值，可能与现蕾期追施尿素水解后土壤中铵态氮含量增加有关［20］，过量施氮促使NH4+大量同化，进而抑制根系发育［21］。本试验减氮25%处理较常规施氮处理减弱根际土壤NH4+的同化，显著提高现蕾期根际土壤NH4+-N含量（13.57%），从而为根系生长提供更充足的氮源。此外，胡柯鑫等［22］研究也表明，合理减氮能够改善土壤理化性质，增加速效养分含量，促进根系生长发育［23］。本研究减氮25%处理显著提高向日葵百粒重和单盘籽粒重，并最终促进产量的形成，说明在适宜减氮条件下，能够实现籽粒物质的有效积累，从而获得更高的产量，与石丽等［24］研究结果一致。
减氮还可能触发植物的胁迫抗逆响应，促进根系分泌有机酸（如柠檬酸、草酸等）［25］，根系分泌物是土壤微生物的重要碳源。本研究减氮25%处理显著促进现蕾期根系分泌草酸与柠檬酸（59.07%和60.11%），减氮50%处理显著促进现蕾期和成熟期根系分泌甲酸（14.17%）。草酸和柠檬酸不仅能增强作物的耐盐性，还可通过螯合作用促进Mg2+、Ca2+等阳离子的释放，提高养分有效性［26］，也为本研究减氮25%与50%处理显著提高现蕾期根际土壤Ca2⁺（17.22%、10.56%）与Mg2⁺（34.75%、25.40%）含量提供合理解释。值得注意的是，减氮50%处理在增加现蕾期根际土壤Ca2+与Mg2⁺含量的同时，显著降低现蕾期K++Na+含量。可能是因为Ca2+和Mg2+可置换土壤胶体吸附的Na+，促进胶体絮凝，改善土壤结构，增强土壤对有益阳离子的吸附能力，从而优化根际离子环境，缓解盐渍化风险。
3.2 施氮处理对根际土壤反硝化细菌群落及其介导的反硝化途径的影响
施氮显著影响土壤中反硝化微生物丰度与群落结构［27-28］，放线菌门与变形菌门在反硝化作用中发挥重要作用［29］，放线菌门作为反硝化途径中的优势菌门，自身就具有抵抗病害，保护植物健康的能力［30］，本研究现蕾期减氮25%处理显著提高放线菌门相对丰度促进向日葵根系生长并维持健康的根际土壤环境。变形菌门（Proteobacteria）中的反硝化细菌依赖高氮环境，可通过降低硝态氮等底物供应来抑制其活性，从而达到减少氮素淋失风险的目的，本研究减氮处理显著降低narG、nirK、nirS型Proteobacteria相对丰度，可能直接使根际土壤反硝化潜势下降，有助于将更多的氮素以硝酸盐（NO3-）的形式留存于土壤提高氮肥利用率。
Ouyang等［31］通过Meta分析发现，农业上施用氮肥可显著提高反硝化基因（nirK、nirS）的丰度，本研究通过减氮处理降低nirS、nirK、narG基因相对丰度，此结论与李志龙等［32］与宋亚娜等［33］研究相同，减量施氮减少反硝化细菌的底物抑制其生长繁殖。Wang等［34］、Gao等［35］研究表明，施氮显著影响土壤N2O排放及nosZ型反硝化细菌的数量和群落结构，黄莹等［36］通过对强酸性茶园土壤的培养发现，施氮显著降低土壤细菌nosZ基因的丰度，然而本研究减氮25%处理（150 kg/hm2）却特异性地提升根际土壤细菌nosZ基因的丰度，可能与减氮促进根系分泌有机酸密切相关；本研究分析得出，现蕾期根际细菌nosZ基因与柠檬酸、甲酸、苹果酸等呈显著正相关，成熟期根际细菌nosZ基因与乙酸呈显著正相关，进一步说明，减氮处理下细菌功能基因nosZ的富集与根系有机酸的分泌密切相关。有机碳含量是影响nosZ基因丰度的重要因子，且介导N2O还原的反硝化细菌（nosZ）对碳源尤为敏感，故本研究适度减施氮肥可能促使根系分泌有机酸，特异性地为nosZ型微生物提供碳源，创造其竞争优势。
4 结 论
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1）减氮处理（N1、N2、N3）显著抑制根际细菌反硝化功能基因narG、nirK、nirS的表达；减氮25%处理（N1）在抑制成熟期根际细菌narG、nirK、nirS表达的同时，促进nosZ的表达，不仅从源头上阻控盐渍农田根际土壤氮素损失，还能减小盐渍向日葵农田土壤中N2O的排放潜力。
2）N1处理显著促进现蕾期根系分泌草酸（59.07%）与柠檬酸（60.11%），并增加根际土壤Ca2⁺（17.22%）与Mg2⁺（34.75%）并降低Cl-、SO42-含量，有效改善盐渍农田根际土壤环境，并显著增加开花期RSAD值（13.40%）、百粒重（9.07%）与单盘籽粒重（9.08%），促进产量形成。
3）减氮50%（N2）处理显著提高现蕾期根际土壤Ca2⁺（10.56%）与Mg2⁺（25.40%），却显著降低成熟期根系RSAD值、百粒重与单盘籽粒重；N2与N3分别导致向日葵减产6.14%和20.22%。基于主成分分析综合评价成熟期根际土壤养分、反硝化功能基因、根系生长及产量指标，减氮25%（150 kg/hm2）处理表现最佳。
基金
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  • 国家自然科学基金项目(U2443210)
  • 国家自然科学基金项目(52469012)
  • 国家自然科学基金项目(52069022)
  • 内蒙古自治区自然科学基金项目(2024LHMS05022)
  • 内蒙古自治区科技重大专项(2023JBGS0003)
文献
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2026年第40卷第4期
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doi: 10.13870/j.cnki.stbcxb.2026.04.040
  • 接收时间：2025-11-21
  • 首发时间：2026-09-30
  • 出版时间：2026-08-28
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  • 收稿日期：2025-11-21
  • 修回日期：2026-03-15
  • 录用日期：2026-04-17
基金
国家自然科学基金项目(U2443210)
国家自然科学基金项目(52469012)
国家自然科学基金项目(52069022)
内蒙古自治区自然科学基金项目(2024LHMS05022)
内蒙古自治区科技重大专项(2023JBGS0003)
作者信息
    1.内蒙古农业大学旱区水工程生态环境全国重点实验室，呼和浩特 010018
    2.内蒙古农业大学水利与土木建筑工程学院，呼和浩特 010018

通讯作者:

李为萍（1978—），女，博士，教授，博士生导师，主要从事灌溉排水原理与管理决策研究。E-mail：
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https://castjournals.cast.org.cn/joweb/stbcxb/CN/10.13870/j.cnki.stbcxb.2026.04.040
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2种不同金属材料的力学参数
科
Family		 属数
Number of
genus		 种数
Number of
species		 占总种数比例
Percentage of
total species (%)		 属
Genus		 种数
Number of
species		 占总种数比例
Percentage of total
species (%)
鹅膏菌科Amanitaceae 2 11 5.26 鹅膏菌属 Amanita 10 4.78
小菇科 Mycenaceae 2 12 5.74 丝盖伞属 Inocybe 5 2.39
多孔菌科 Polyporaceae 8 14 6.70 蜡蘑属 Laccaria 5 2.39
红菇科 Russulaceae 3 23 11.00 小皮伞属 Marasmius 6 2.87
小菇属 Mycena 11 5.26
光柄菇属 Pluteus 5 2.39
红菇属 Russula 17 8.13
栓菌属 Trametes 5 2.39
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