Article(id=1274057340614631962, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1274057338156769818, articleNumber=null, orderNo=null, doi=10.13343/j.cnki.wsxb.20250530, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1752163200000, receivedDateStr=2025-07-11, revisedDate=null, revisedDateStr=null, acceptedDate=1761753600000, acceptedDateStr=2025-10-30, onlineDate=1781688540843, onlineDateStr=2026-06-17, pubDate=1780502400000, pubDateStr=2026-06-04, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1781688540843, onlineIssueDateStr=2026-06-17, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1781688540843, creator=13701087609, updateTime=1781688540843, updator=13701087609, issue=Issue{id=1274057338156769818, tenantId=1146029695717560320, journalId=1192105938417971205, year='2026', volume='66', issue='6', pageStart='2561', pageEnd='3114', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=1, specialIssue=null, createTime=1781688540257, creator=13701087609, updateTime=1781688602467, updator=13701087609, preIssue=null, nextIssue=null, ext={EN=IssueExt(id=1274057599193486082, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1274057338156769818, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1274057599193486083, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1274057338156769818, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null}, startPage=2580, endPage=2591, ext={EN=ArticleExt(id=1274057341377995292, articleId=1274057340614631962, tenantId=1146029695717560320, journalId=1192105938417971205, language=EN, title=Brassinolide enhance plant stress resistance in acidic soil by regulating rhizosphere microorganisms: mechanisms and prospects, columnId=1192149543727808575, journalTitle=Acta Microbiologica Sinica, columnName=Review, runingTitle=null, highlight=null, articleAbstract=

Acidic soil accounts for approximately 50% of the world’s available arable land. Its highly active aluminum ions and low pH environment not only directly inhibit plant growth but also significantly alter the microbial community structure in the rhizosphere, weaken the functions of beneficial microorganisms, and exacerbate soil-borne diseases. Brassinolide (BR), as a group of important plant signaling molecules, play a core role in enhancing the plant stress resistance in acidic soil by precisely regulating plant-microorganism interactions. BR promote the root secretion of organic acids such as malic acid and oxalic acid by activating BZR1/BES1 and transcription factors. These secretions act as carbon sources and chemotactic signals to specifically recruit beneficial microorganisms such as Paenibacillus azotofixans, Pseudomonas, and ectomycorrhizal fungi, reshaping the microbial community structure in the rhizosphere. The microbial community reassembly induced by BR significantly enhances aluminum ion chelation, nutrient activation, and pathogen inhibition. For instance, nitrogen-fixing bacteria enriched utilize malic acid for metabolic activities and secrete auxin and other substances to promote plant growth in acidic environments. Ectomycorrhizal fungi alleviate aluminum toxicity through oxalic acid secretion. Meanwhile, BR, in collaboration with plant hormones such as auxin and gibberellin, optimizes the root structure, expands the microbial colonization niche, and forms a complex synergistic network for enhancing stress resistance. Future research should focus on the specific regulatory mechanisms of BR on the rhizosphere microbiome, unveil the direct action pathways of BR as microbial signaling molecules, and develop efficient BR-microbial compound preparations in combination with microbial community engineering, providing innovative strategies and application solutions for the regulation of acidic soil microorganisms.

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E-mail: LIANG Yuting,
ZHAO Yuan,
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酸性土壤约占全球可用耕地的50%,其高活性铝离子和低pH环境不仅直接抑制植物生长,更显著改变根际微生物群落结构,削弱有益微生物功能,进而加剧土传病害。油菜素内酯(brassinolide, BR)作为重要的植物信号分子,通过精细调控“植物-微生物”互作在增强植物对酸性土壤的抗逆性中发挥核心作用。BR通过激活BZR1/BES1和转录因子促进根系分泌苹果酸、草酸等有机酸。这些分泌物作为碳源和趋化信号特异性招募固氮类芽孢杆菌、假单胞菌、外生菌根真菌等有益微生物,重塑根际微生物群落结构。BR诱导的微生物群落重构显著增强了铝离子螯合、养分活化及病原菌抑制功能。例如,富集的固氮菌利用苹果酸进行代谢活动,同时分泌生长素等物质促进植物在酸性环境中生长;外生菌根真菌通过分泌草酸缓解铝毒。同时,BR与生长素、赤霉素等激素协同优化根系构型,扩大微生物定殖生态位,形成复杂的协同抗逆网络。未来研究需聚焦BR对根际微生物组的特异性调控机制,解析BR作为微生物信号分子的直接作用途径,并结合微生物群落工程开发高效的BR-微生物复合制剂,为酸性土壤微生物调控提供创新策略和应用方案。

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作者贡献声明

张若琳:起草文章的初稿、修改和完善,以及最终的文字润色和校对;张理:对文献中的数据进行整理、统计和分析,文字润色和校对;马志远:对综述提出修改意见、框架和研究问题;姜美彤:背景资料的文字润色和校对;丁骥贤:对综述提出修改意见,项目管理与协调;张慧:项目管理与协调;赵远:框架设计、论文审阅;梁玉婷:提出综述的主题、框架和研究问题、文章审阅与修改。

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油菜素内酯通过调控根际微生物增强植物在酸性土壤中的抗逆性:机制与展望
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张若琳 1, 2 , 张理 2, 3 , 马志远 2 , 姜美彤 2 , 丁骥贤 2 , 张慧 2 , 赵远 1 , 梁玉婷 2
微生物学报 | 综述 2026,66(6): 2580-2591
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微生物学报 | 综述 2026, 66(6): 2580-2591
油菜素内酯通过调控根际微生物增强植物在酸性土壤中的抗逆性:机制与展望
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张若琳1, 2, 张理2, 3, 马志远2, 姜美彤2, 丁骥贤2, 张慧2, 赵远1 , 梁玉婷2
作者信息
  • 1.常州大学 环境科学与工程学院,江苏 常州
  • 2.中国科学院南京土壤研究所,江苏 南京
  • 3.中国科学院大学,北京
Brassinolide enhance plant stress resistance in acidic soil by regulating rhizosphere microorganisms: mechanisms and prospects
Ruolin ZHANG1, 2, Li ZHANG2, 3, Zhiyuan MA2, Meitong JIANG2, Jixian DING2, Hui ZHANG2, Yuan ZHAO1 , Yuting LIANG2
Affiliations
  • 1.School of Environmental Science and Engineering, Changzhou University, Changzhou, Jiangsu, China
  • 2.Institute of Soil Science, Chinese Academy of Sciences, Nanjing, Jiangsu, China
  • 3.University of Chinese Academy of Sciences, Beijing, China
出版时间: 2026-06-04 doi: 10.13343/j.cnki.wsxb.20250530
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酸性土壤约占全球可用耕地的50%,其高活性铝离子和低pH环境不仅直接抑制植物生长,更显著改变根际微生物群落结构,削弱有益微生物功能,进而加剧土传病害。油菜素内酯(brassinolide, BR)作为重要的植物信号分子,通过精细调控“植物-微生物”互作在增强植物对酸性土壤的抗逆性中发挥核心作用。BR通过激活BZR1/BES1和转录因子促进根系分泌苹果酸、草酸等有机酸。这些分泌物作为碳源和趋化信号特异性招募固氮类芽孢杆菌、假单胞菌、外生菌根真菌等有益微生物,重塑根际微生物群落结构。BR诱导的微生物群落重构显著增强了铝离子螯合、养分活化及病原菌抑制功能。例如,富集的固氮菌利用苹果酸进行代谢活动,同时分泌生长素等物质促进植物在酸性环境中生长;外生菌根真菌通过分泌草酸缓解铝毒。同时,BR与生长素、赤霉素等激素协同优化根系构型,扩大微生物定殖生态位,形成复杂的协同抗逆网络。未来研究需聚焦BR对根际微生物组的特异性调控机制,解析BR作为微生物信号分子的直接作用途径,并结合微生物群落工程开发高效的BR-微生物复合制剂,为酸性土壤微生物调控提供创新策略和应用方案。

油菜素内酯  /  酸性土壤  /  根际微生物组  /  微生物群落重构  /  植物-微生物互作  /  微生物信号传导

Acidic soil accounts for approximately 50% of the world’s available arable land. Its highly active aluminum ions and low pH environment not only directly inhibit plant growth but also significantly alter the microbial community structure in the rhizosphere, weaken the functions of beneficial microorganisms, and exacerbate soil-borne diseases. Brassinolide (BR), as a group of important plant signaling molecules, play a core role in enhancing the plant stress resistance in acidic soil by precisely regulating plant-microorganism interactions. BR promote the root secretion of organic acids such as malic acid and oxalic acid by activating BZR1/BES1 and transcription factors. These secretions act as carbon sources and chemotactic signals to specifically recruit beneficial microorganisms such as Paenibacillus azotofixans, Pseudomonas, and ectomycorrhizal fungi, reshaping the microbial community structure in the rhizosphere. The microbial community reassembly induced by BR significantly enhances aluminum ion chelation, nutrient activation, and pathogen inhibition. For instance, nitrogen-fixing bacteria enriched utilize malic acid for metabolic activities and secrete auxin and other substances to promote plant growth in acidic environments. Ectomycorrhizal fungi alleviate aluminum toxicity through oxalic acid secretion. Meanwhile, BR, in collaboration with plant hormones such as auxin and gibberellin, optimizes the root structure, expands the microbial colonization niche, and forms a complex synergistic network for enhancing stress resistance. Future research should focus on the specific regulatory mechanisms of BR on the rhizosphere microbiome, unveil the direct action pathways of BR as microbial signaling molecules, and develop efficient BR-microbial compound preparations in combination with microbial community engineering, providing innovative strategies and application solutions for the regulation of acidic soil microorganisms.

brassinolide  /  acidic soil  /  rhizosphere microbiome  /  microbial community reassembly  /  plant-microorganism interaction  /  microbial signal transduction
张若琳, 张理, 马志远, 姜美彤, 丁骥贤, 张慧, 赵远, 梁玉婷. 油菜素内酯通过调控根际微生物增强植物在酸性土壤中的抗逆性:机制与展望. 微生物学报, 2026 , 66 (6) : 2580 -2591 . DOI: 10.13343/j.cnki.wsxb.20250530
Ruolin ZHANG, Li ZHANG, Zhiyuan MA, Meitong JIANG, Jixian DING, Hui ZHANG, Yuan ZHAO, Yuting LIANG. Brassinolide enhance plant stress resistance in acidic soil by regulating rhizosphere microorganisms: mechanisms and prospects[J]. Acta Microbiologica Sinica, 2026 , 66 (6) : 2580 -2591 . DOI: 10.13343/j.cnki.wsxb.20250530
全球约有25亿hm2的耕地和潜在耕地属于酸性土壤,约占全球可用耕地总面积的50%[1]。土壤酸化不仅由自然因素(如土壤风化、盐基离子淋失)引发,也受人为因素(如氮肥过量使用、大量排放导致酸沉降等)影响[2]。酸化会增强土壤毒性金属的活性,其中铝毒被认为是抑制植物在酸性土壤中生长的最主要限制因子[3]。铝毒通过抑制根系发育、降低光合作用效率,进而抑制作物产量[4]。同时,土壤酸化还会降低氮、磷等养分的有效性,进一步限制作物生产力[5]。此外,土壤酸化进程还伴随着土壤微生物群落结构的明显变化,具体表现为真菌与细菌的比值上升、病原微生物富集,以及拮抗菌群活性被显著抑制,进而增加土传病害的发生概率[6]。因此,从微生物学和微生态学角度寻找有效缓解酸性土壤对作物不利影响的创新策略已成为当前农业可持续发展的重要研究课题。
油菜素内酯(brassinolide, BR)被认为是植物的第六类激素,在促进植物生长、调节营养代谢、强化根系发育及提升作物抗逆性方面具有重要作用。1970年,美国科学家Mitchell首次从油菜花粉中筛选并分离出一种具有植物生长调节作用的生理活性物质,将其命名为油菜素(brassin)[7]。目前,植物界中已发现60余种油菜素内酯类化合物,这类物质被统一称作油菜素甾醇(brassinosteroids, BRs)。其中,同系物中使用范围最广的有3种,分别是芸苔素内酯(brassinolide, BL)、28-homobrassinolide (28-HomoBL/HBL)以及24-epibrassinolide (24-EpiBL/EBR)[8]。BR的研究涵盖多种模式植物和经济作物,包括拟南芥、水稻、小麦、番茄、棉花等。这些研究揭示了BR信号通路促进植物生长发育的核心机制——调控植物表型重塑和营养获取能力[9]。近年研究表明,BR能通过改变根系分泌物组成和质量影响根际微生物的群落结构、功能活性和生态网络[10]。目前,关于BR通过调控根际微生物组增强植物对酸性土壤适应性的分子机制和生态效应尚待系统解析和深入研究。
本文从微生物学角度出发,系统回顾了BR对根际微生物群落的调控作用和机制,重点阐述了BR通过调控根系分泌物-微生物互作增强植物对酸性土壤适应性的生理生态机制,以期为深入挖掘BR的微生物调控功能提供理论依据和研究思路。
酸性土壤环境显著影响根际微生物的群落结构组成和功能活性表达。土壤pH值是调控微生物群落组成的关键环境因子,在pH<5.5的酸性条件下细菌多样性通常会下降,群落结构趋于简化,而真菌群落结构则会产生特异性变化,部分耐受类群呈现显著富集态势[11]。土壤酸碱度能显著调控根际微生物群落结构:中性或弱碱性条件有利于维持细菌多样性,而酸性环境则会增强真菌的竞争能力,并抑制拮抗细菌的活性,从而加剧植物病害发生[12]。为应对酸性胁迫增加的病害风险,植物通过根系分泌物主动招募有益微生物[13]。例如,拟南芥可分泌苹果酸以富集枯草芽孢杆菌(Bacillus subtilis)等益生菌,增强抗逆能力[14];耐铝植物根际则富集青霉属(Penicillium)、芽枝霉属(Cladosporium)等真菌,并形成更稳定的共生网络[15]。土豆在致病疫霉(Phytophthora infestans)侵染下也可招募放线菌以增强防御[16]。酸性土壤促进丛枝菌根真菌(arbuscular mycorrhizal fungi, AMF)与植物根系的共生整合,显著提升AMF在根系中的定殖效率,并同步放大植物对菌根途径的生理依赖[17-19]
在微生物功能基因表达和代谢途径方面,酸化显著抑制硝化菌和氨氧化菌的活性,改变氮循环途径和效率,影响氮素利用[20]。具有溶磷功能的微生物曲霉属(Aspergillus)在酸性环境中通过分泌草酸、柠檬酸等有机酸有效活化固定态磷素,提高磷有效性[13,21-22]。固氮菌属(Azotobacter)是广泛使用的促生根细菌,其固氮能力对植物根系生长至关重要[23]。此外,酸性土壤中病原微生物如镰孢(Fusarium)和疫霉(Phytophthora)的竞争能力和致病性增强,而具有生防潜力的拮抗菌如假单胞菌(Pseudomonas)和木霉(Trichoderma)的活性受到抑制,微生态平衡被破坏[24]
植物可借助对自身根系分泌物的化学组成实施精准调控以适配根际微生物群落的结构特征,进而适应特定的酸性胁迫生境。铝胁迫条件下,植物增加有机酸分泌量并改变分泌组分,这些分泌物不仅直接螯合铝离子降低毒性,还作为特异性信号分子招募有益微生物类群[25]。微生物可以分泌生长素(indole-3-acetic acid, IAA)等植物激素,促进植物根系发育并提升其功能[26]。例如,植物根际促生菌(plant growth promoting rhizobacteria, PGPR)如芽孢杆菌属(Bacillus)能够合成IAA、赤霉素(gibberellic acid, GA)等激素,显著促进根系伸长、侧根增多,并在干旱、盐碱等胁迫下提升植株抗氧化酶活性[27-28];还能产生铁载体等代谢产物抑制病原菌生长和繁殖[29]。优化发酵条件后,铜绿假单胞菌(Pseudomonas aeruginosa) Gxun-2产铁载体能力增强,对多种根腐病原菌的抑菌圈直径显著增大,抑菌率均大于70%[30];此外,微生物还能通过有机酸分泌调节根际pH环境[31]。以真菌枯萎病病原菌尖孢镰孢菌(Fusarium oxysporum)为例,其分泌的快速碱化因子肽激素可诱导质外体pH升高,进而促进该真菌在植物体内定殖[32]。最近研究表明,合成微生物群落(synthetic microbial community, SynCom)在提高植物酸性土壤适应性方面具有巨大应用潜力,可通过人工设计优化群落结构和功能[33-34]
BR作为一类高效能激素,其含量的剧烈波动会对植物的生长发育产生重要影响。目前,关于BR信号通路的理解主要源自拟南芥等模式植物的遗传与分子生物学研究。BR的生物合成以甾醇类化合物菜油甾醇(campesterol, CR)为前体,依赖去黄化素2 (de-etiolated 2, DET2)以及多种细胞色素P450单加氧酶(如CYP90家族)的协同催化,最终形成具有最高生物活性的BR[35]。在BR稳态维持的调控网络中,信号转导通路下游的转录因子通过抑制多个参与生物合成的关键酶基因(如DWF4CPD)的表达,实现对生物合成过程的负反馈调控。DWF4基因编码一种C-22羟化酶,该酶属于细胞色素P450家族,在拟南芥中由转移DNA (transfer-DNA, T-DNA)插入引起DWF4功能丧失的突变体,表现出典型的油菜素内酯合成缺陷表型[36]。通过T-DNA插入技术构建的cpd突变体,因其CPD基因(编码C-23羟化酶)功能丧失而造成内源激素缺陷,从而在黑暗条件下呈现黄化抑制表型[37]。此外,CPD基因的表达受BR的负反馈调控作用,转录因子油菜素内酯不敏感1 (brassinosteroid resistant 1, BZR1)可特异性识别并结合CPD的启动子区域,从而抑制该基因的转录活性[38]
当BR水平较低时,BR信号通过细胞膜上的受体激酶油菜素内酯不敏感1 (brassinosteroid insensitive 1, BRI1)感知,激酶抑制剂1 (kinase inhibitor 1, BKI1)会占据BRI1的位点,抑制BRI1与BRI1相关受体激酶1 (BRI1-associated receptor kinase 1, BAK1)的相互作用[39]。当BR水平较高时,BRI1和BAK1蛋白磷酸化后会启动后续的磷酸化级联反应,向下游信号传递[40]。BRI1-BAK1复合物通过磷酸化级联反应激活下游信号分子BRI1抑制子1 (BRI1 suppressor 1, BSU1),油菜素内酯不敏感2 (brassinosteroid-insen sitive 2, BIN2)是转录因子BZR1和油菜素内酯不敏感EMS抑制子1 (BR-insensitive-EMS-suppressor 1, BES1)/BZR2磷酸化的重要信号因子,BSU1因子在BR较高水平下产生并使BIN2去磷酸化,进而被降解,最终会抑制BES1和BZR1的磷酸化过程[41]。此外,蛋白磷酸酶2A (PP2A)也会介导BES1和BZR1的去磷酸化和激活,去磷酸化的转录因子BZR1和BES1进入细胞核调控BR响应基因的表达[42]。BR信号转导的核心调控机制如图1A所示。转录因子BZR1与BES1可直接调控参与细胞伸长、分裂、分化及逆境应答相关基因的表达,进而协同调控植物的生长发育进程与胁迫适应能力[43]。如图1B所示,当根系细胞进入快速伸长区,BR信号增强,BZR1的表达处于高峰;然后当根系细胞进入成熟区,BZR1的表达骤然降低[44](圆圈所画范围是BR信号最强部分)。BR信号转导的核心在于通过磷酸化/去磷酸化和泛素化过程精确调控关键组分的活性与稳定性,这一复杂且高度动态的信号系统使植物得以在快速变化的环境中实现生长发育与胁迫响应的高度协调[41]
在转录调控层面,一项早期研究表明,BR处理促进了对水稻稻瘟病菌(Magnaporthe oryzae)的抗性[45]。近年研究揭示了水稻中的GAI-RGA-SCR (GRAS)家族蛋白OsDLA通过与GSK3/SHAGGY样(GSK3/SHAGGY-like, GSK2)激酶和OsWRKY53形成模块,参与油菜素内酯信号途径并正向调控对M. oryzae的抗性[46]。BIN2在水稻中的同源蛋白为GSK2激酶[47]。植物特有的GRAS蛋白家族在植物生长发育进程中行使多种生物学功能[48]OsWRKY53过表达的转基因植株,其叶片张开角度显著增加,且该类植株对BR表现出高度敏感特性[49]OsDLA基因编码的是GRAS蛋白家族中的一个成员,BR处理和M. oryzae侵染可诱导OsDLA上调表达;OsDLA与OsWRKY53转录因子发生互作,抑制了GSK2介导的OsWRKY53磷酸化降解,大量OsWRKY53进入细胞核激活了BR响应基因的表达,进而增强了水稻对稻瘟病的抗性[46]
在根系分泌物层面,BR通过调控根系分泌物的组成和含量实现对铝胁迫的响应及根际微环境的调控。拟南芥铝激活苹果酸转运蛋白(Arabidopsis aluminum-activated malate transporter, AtALMT1)主要在根系中表达,其核心功能是受铝离子激活后向细胞外分泌苹果酸[50]。铝胁迫下,AtALMT1基因调控的根系苹果酸分泌在拟南芥铝胁迫耐性调控中起到关键作用[51]。BZR1与BES1转录因子借助转录水平调控AtALMT1的表达,间接调控根系苹果酸的分泌。其中,BZR1/BES1与STOP1间以拮抗作用为主,与WRKY46则通过协同作用联合调控AtALMT1的表达;与BZR1/BES1的调控途径不同,BIN2及其同源蛋白主要在蛋白水平对AtALMT1实施调控,进而影响根系苹果酸的分泌过程[52]
BR与其他激素共同调控根系分泌物,进而促进有益根际微生物定殖,同时抑制致病菌的活性。例如,番茄蔗糖转运蛋白SlSUT2可以与BR生物合成和信号传导组分发生物理互作以调节丛枝菌根(arbuscular mycorrhiza, AM)共生[53]。BR能够诱导水稻中赤霉素(gibberellin, GA)生物合成途径的关键基因D18/GA3ox-2的表达,进而使水稻幼苗体内具备生物活性的GA含量显著上升[54]
在酸性土壤环境中BR通过精确调控微生物群落结构和代谢活性,显著缓解铝毒害效应。BR促进根系分泌有机酸,这些有机酸不仅直接螯合铝离子,还作为微生物生长的碳源,促进微生物产生柠檬酸特异性铝螯合剂[55]。其中,苹果酸是根系分泌物中的核心有机酸组分,它借助浓度梯度产生化学信号分子,招募具备趋化特性的微生物类群。例如,黄杆菌(Flavobacterium)和固氮类芽孢杆菌(Paenibacillus arsenophilus)可通过感知苹果酸的浓度梯度,依靠鞭毛运动定向迁移至根际区域[56]。苹果酸为微生物提供可利用的碳源,促进其在根际的定殖与增殖。例如,固氮螺菌(Azospirillum humicireducens)通过利用苹果酸进行代谢活动,同时分泌IAA等促生物质,与植物形成互惠关系[57]。此外,苹果酸还通过降低根际土壤pH值创造酸性微环境,筛选出耐酸或嗜酸的微生物类群行使养分活化功能。假单胞菌和枯草芽孢杆菌(B. subtilis)等通过调节自身代谢途径适应酸性条件,并与植物的苹果酸分泌协同促进养分活化[58]。BR能够上调植物根系分泌草酸[59]。外生菌根真菌(ectomycorrhizal fungi, ECMF)可以利用草酸和菌丝体对Al3+的吸附提高抗铝性[60]。这是由草酸的结构特点决定的,草酸利用其二元酸特性与金属离子螯合形成五元环稳定结构,瞬时阻断离子毒性,实现高效解毒[61]。苏畅等[59]的研究证实,铝胁迫条件下水稻铝敏感突变体osalr3对外源施加的BR表现出明显应答:相关试验结果显示,经外源BR处理后,osalr3突变体中的草酸含量显著高于对照野生型植株,且可有效减轻铝胁迫造成的毒害效应,植株呈现更优的生长表型特征。此外,草酸的分泌有利于养分的活化和利用,在磷匮乏的酸性土壤中ECMF能够利用草酸溶解难溶性磷提高磷获取效率[62]
除调控有机酸的分泌外,BR还可通过调节根系细胞壁的组成协同根际微生物降低根际铝毒[63]。微生物所分泌的胞外聚合物(extracellular polymeric substance, EPS)含有丰富的多糖、蛋白质及羧基等活性功能基团,可借助物理吸附、离子交换及络合等作用途径高效螯合铝离子,进而降低根际环境中可利用铝的浓度水平[26]。与此同时,部分根际微生物还能将吸附的铝离子转运至细胞内部,通过液泡或其他细胞器进行区室化储存,或与有机酸、酚类等形成复合物,实现内部解毒[64]。在此基础上,BR能促进根系细胞壁中半纤维素的合成并调节果胶代谢,使水溶性果胶含量增加,从而进一步增强细胞壁对金属离子的结合能力[65]。此外,BR处理还显著上调水稻根、茎中超氧化物歧化酶(superoxide dismutase, SOD)、过氧化物酶(peroxidase, POD)和过氧化氢酶(catalase, CAT)的活性,减轻铝诱导的氧化损伤[66-67]。这些细胞壁-微生物协同作用,使得根际微生物在酸性土壤中能够更有效地吸附、隔离铝离子,从而降低铝对植物的毒害。
近期研究证实,SynCom可推动水稻根部BR的生物合成过程,进而提升木葡聚糖内转葡糖基酶(xyloglucan endotransglycosylase, XET)的催化活性,同时重构木聚糖侧链结构,使Al结合位点数量降低了47.5%;与此同时,SynCom还能生成代谢中间体甲羟戊酸(mevalonate, MVA),该物质可进一步诱导根部BR的生物合成,而BR含量的升高会进一步启动XET的功能,触发细胞壁重构过程,进而提升水稻对铝胁迫的耐受能力[68]
BR可借助重构微生物群落结构及互作网络,提升对病害的抑制效能。当病原体侵染植物时,外源施加的BR或植物自身内源BR,能够启动下游免疫相关基因的表达程序,进而强化植物针对病原体的免疫反应[69]。特别是在酸性土壤环境中,植物与微生物互作关系变得更加复杂。土壤酸化显著改变土壤微生物群落结构[70],在这一过程中,细菌群落的变化导致其对土传真菌如黄色镰孢菌(Fusarium culmorum)、疫霉等病原体的抑制能力下降[71]。然而,BR能够有效增强植物对土传真菌的抗性,缓解病原体侵害。例如,BR能够显著提高大麦对多种不同生活方式的真菌病原体的抗性。在大麦种子发芽后,向盆栽土壤中添加20 mL 1 µmol/L EBR进行根部灌溉,在真菌处理后第15天采集基部样本,结果表明2种镰孢菌引起的大麦镰孢菌幼苗疫病症状分别减少了28%和35%[72]
在酸性土壤中,霉菌感染对作物生长构成了严重威胁。例如,甜瓜疫霉(Pythium melonis)是亚洲地区葫芦科作物的主要病原体之一。研究表明,通过对黄瓜植株进行20 mL 1 μmol/L EBR根部灌溉,处理2 d后发现显著增强了黄瓜根系韧皮部和木质部血管细胞壁中木质素的积累,使植株下胚轴更厚实,从而提高对P. melonis的抗性。该机制通过延缓菌丝的扩展和减少感染区域有效预防了疫霉病的发生[73]
此外,土壤pH为5.0时芸薹根肿菌(Plasmodiophora brassicae)引起的十字花科作物根肿病发病率和病情指数最高[74]。BR通过激活转录因子BZR,抑制FLS2SUPPRESSOR OF NPR1-1CONSTITUTIVE 1 (SNC1)等基因,从而直接参与病原体免疫基因的表达[75]
在酸性土壤条件下,BR处理可显著降低土传真菌病害的发病率与病情指数。BR对根系代谢重塑、根际微生物定殖优化及植物抗逆性增强方面具有多维度协同作用:一方面,BR促使根系分泌有机酸,这类物质既通过“趋化效应”吸引功能性微生物向根表富集,并作为碳源与能量来源为微生物提供营养支撑,进而调控根际微生物的选择性定殖过程[56,76];另一方面,BR重塑根细胞壁、减少铝结合位点,增强植物-微生物互作过程[68]。同时,BR能够延缓菌丝的扩展,减少感染区域,并上调根系中SOD、POD等抗氧化酶活性[67],机制如图2所示。
BR作为一类重要植物激素,在植物响应多种逆境胁迫中发挥着核心调控作用,特别是在酸性土壤环境下,BR通过多层次机制促进植物适应能力的提升,具体表现为:(1) 精确调控根系分泌物组成和质量,定向招募有益微生物类群;(2) 重构微生物群落结构,抑制土传病害发生和传播;(3) 优化微生物-植物信号对话,增强植物的系统抗性。这些机制共同构成了复杂的BR-微生物协同抗逆网络,提高植物环境适应性。
尽管BR在缓解生物与非生物胁迫方面已有较多应用,但仍需进一步深入研究以全面揭示其在缓解植物酸铝胁迫的机制和潜在应用价值。从分子生物学层面,应深入研究BR如何在基因和信号转导途径调控微生物对酸性胁迫的响应,并探讨BR是否通过与微生物表面受体结合,激活其固氮或有机酸代谢通路;从微生物生态学层面,则需要探讨BR对根际微生物群落结构和功能的具体影响,进一步研究BR介导的根系分泌物变化对微生物群落的定向筛选,验证BR处理是否能增加根际有益微生物丰度。
未来可以开展长期田间实验,监测除产量外的生理指标与土壤理化指标数据,并对不同作物与区域进行研究,验证BR在粮食作物和经济作物抗酸铝胁迫的效果。基于BR与功能微生物的协同效应,开发BR缓释剂与微生物菌剂的复合制剂,对比单一BR或单一菌剂的应用优势,验证其在酸性红壤中的应用效果;同时,选择3-4个典型红壤类型的田块,结合气候与种植制度,制定BR施用规程并明确最佳窗口期,将BR嵌入种前、中期、收获前环节;最后,强化土壤环境保障,在试点建观测站设4个处理并进行5-10年监测,跟踪土壤微生物、重金属及酶活性,并建立预警模型预判风险,以此实现BR在农业生产中的有效且可持续应用。
综上所述,BR在调控酸性土壤植物-微生物互作中具有显著的调节作用。为了充分挖掘其潜力及应用价值,未来需要进行更加系统、全面地研究,包括机制解析、长期田间试验以及应用潜力的探索。这不仅有助于提升农业生产效率,还能促进生态环境的可持续发展。
  • 国家重点研发计划(2021YFD1900400)
  • 国家自然科学基金(42425703)
  • 国家自然科学基金(42377121)
  • 江苏省研究生实践创新计划(SJCX24_1654)
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2026年第66卷第6期
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doi: 10.13343/j.cnki.wsxb.20250530
  • 接收时间:2025-07-11
  • 首发时间:2026-06-17
  • 出版时间:2026-06-04
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  • 收稿日期:2025-07-11
  • 录用日期:2025-10-30
基金
the National Key Research and Development Program of China(2021YFD1900400)
国家重点研发计划(2021YFD1900400)
the National Natural Science Foundation of China(42425703)
国家自然科学基金(42425703)
the National Natural Science Foundation of China(42377121)
国家自然科学基金(42377121)
the Postgraduate Research & Practice Innovation Program of Jiangsu Province(SJCX24_1654)
江苏省研究生实践创新计划(SJCX24_1654)
作者信息
    1.常州大学 环境科学与工程学院,江苏 常州
    2.中国科学院南京土壤研究所,江苏 南京
    3.中国科学院大学,北京
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https://castjournals.cast.org.cn/joweb/wswxb/CN/10.13343/j.cnki.wsxb.20250530
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2种不同金属材料的力学参数

Family
属数
Number of
genus
种数
Number of
species
占总种数比例
Percentage of
total species (%)

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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