Article(id=1276616206748619574, tenantId=1146029695717560320, journalId=1235980609244409860, issueId=1276616049617408127, articleNumber=null, orderNo=null, doi=10.3969/j.issn.1000-2561.2025.08.015, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=null, receivedDate=1740672000000, receivedDateStr=2025-02-28, revisedDate=null, revisedDateStr=null, acceptedDate=1746633600000, acceptedDateStr=2025-05-08, onlineDate=1782298622070, onlineDateStr=2026-06-24, pubDate=1756051200000, pubDateStr=2025-08-25, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1782298622070, onlineIssueDateStr=2026-06-24, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1782298622070, creator=13701087609, updateTime=1782298622070, updator=13701087609, issue=Issue{id=1276616049617408127, tenantId=1146029695717560320, journalId=1235980609244409860, year='2025', volume='46', issue='8', pageStart='1785', pageEnd='2029', issueExtLink='null', onlineDate='null', pubDate='1756051200000', pubDateStr='2025-08-25', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1782298584608, creator='13701087609', updateTime=1782298660748, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext={EN=IssueExt(id=1276616369089147039, tenantId=1146029695717560320, journalId=1235980609244409860, issueId=1276616049617408127, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1276616369089147040, tenantId=1146029695717560320, journalId=1235980609244409860, issueId=1276616049617408127, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null, downloadFileDto=null}, startPage=1921, endPage=1930, ext={EN=ArticleExt(id=1276616208443118392, articleId=1276616206748619574, tenantId=1146029695717560320, journalId=1235980609244409860, language=EN, title=Effects of AM Fungi on Wisteria sinensis to Pb Stress and Its Physiological Mechanism, columnId=1236256434120348225, journalTitle=Chinese Journal of Tropical Crops, columnName=Plant Cultivation, Physiology & Biochemistry, runingTitle=null, highlight=null, articleAbstract=

A controlled experiment was conducted using Wisteria sinensis seedlings inoculated with Funneliformis mosseae to investigate the effects and physiological mechanisms of arbuscular mycorrhizal fungi (AMF) on the Pb stress tolerance of W. sinensis seedlings. Pb contamination gradients were established (400, 800, 1200 mg/kg) to measure the growth parameters (growth, biomass, tolerance index) and physiological indicators (chlorophyll content, malondialdehyde content, proline content and related antioxidant enzyme activities). AM fungi inoculation significantly enhanced both aerial and root growth of W. sinensis seedlings while improving the Pb stress tolerance. The translocation factor and shoot bio-concentration factor were lower than those of the control, whereas the root bio-concentration factor exceeded the level of the control groups. This demonstrated that AM fungi effectively reduced Pb accumulation in the aerial parts while significantly enhancing Pb sequestration capacity in the roots. Physiological analysis indicated that AM fungi-treated seedlings exhibited higher chlorophyll content and elevated the activities of antioxidant enzymes (SOD, POD, APX), along with significantly lower MDA accumulation compared to the non-inoculated controls. Principal component analysis combined with cluster heat-map visualization further demonstrated that AM fungi primarily enhanced the Pb tolerance of W. sinensis through three mechanisms: activation of antioxidant enzyme systems, regulation of growth-related processes, and modulation of chlorophyll biosynthesis pathways. The results provide theoretical and practical basis for mycorrhizal plant remediation of heavy metal pollution.

, authors=null, authorsList=Jieming WANG, Hongyun PENG, Ruixue LI, authorCompany=null, correspAuthors=Ruixue 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=1276616209806267206, articleId=1276616206748619574, tenantId=1146029695717560320, journalId=1235980609244409860, language=CN, title=AM真菌对Pb污染土壤中紫藤生长及其生理的影响, columnId=1236256434313286224, journalTitle=热带作物学报, columnName=作物栽培与生理生化, runingTitle=null, highlight=null, articleAbstract=

为探究丛枝菌根(AM)真菌对紫藤(Wisteria sinensis)扦插苗抗铅(Pb)胁迫的影响及其生理机理,以接种摩西斗管囊霉(Funneliformis mosseae)的紫藤扦插苗为材料,Pb污染浓度设置为400、800、1200 mg/kg,通过盆栽模拟试验,测定紫藤幼苗相关生长指标(生长量、生物量、耐性指数)和生理指标(叶绿素含量、丙二醛含量、脯氨酸含量、相关抗氧化酶活性)。结果表明:接种AM真菌能促进紫藤幼苗的地上部和根系生长,并提高紫藤对Pb的耐受性。转移系数以及地上部的富集系数低于CK,根系富集系数均高于CK,表明AM真菌降低了紫藤地上部的Pb含量并显著提高紫藤根系对Pb的富集能力。生理机制分析发现,在AM真菌作用下,紫藤叶片的叶绿素含量、超氧化物歧化酶(SOD)、过氧化物酶(POD)、抗坏血酸过氧化物酶(APX)等抗氧化酶活性均高于CK,丙二醛(MDA)积累量显著低于CK。主成分分析和聚类热图分析也显示,AM真菌主要通过激活抗氧化酶活性,调控相关生长指标及叶绿素合成途径来增强紫藤对Pb胁迫的耐受能力。该研究结果为菌根化植物修复重金属污染提供理论与实践依据。

, authors=

汪结明(1977—),男,博士,副教授,研究方向:园林植物应用。

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* 李瑞雪(LI Ruixue),E-mail:
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Effects of arbuscular myeorrhizal fungi on growth and eeophysiological characteristies of Imperata cylindrica under heavy metal stress[J]. Journal of Shanxi University (Natural Science Edition), 2023, 46(2): 439-448. 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Physiological and biochemical evaluation of synergistic responses in drought and cold tolerance of Sophora davidii seedlings[J]. Chinese Journal of Tropical Crops, 2023, 44(10): 2051-2059. (in Chinese), articleTitle=Physiological and biochemical evaluation of synergistic responses in drought and cold tolerance of Sophora davidii seedlings, refAbstract=null), Reference(id=1276616219969065885, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276616206748619574, doi=null, pmid=null, pmcid=null, year=2019, volume=164, issue=null, pageStart=10, pageEnd=19, url=null, language=null, rfNumber=[28], rfOrder=45, authorNames=ZHANG X, ZHANG H, LOU X, TANG M, journalName=Environmental and Experimental Botany, refType=null, unstructuredReference=ZHANG X, ZHANG H, LOU X, TANG M. Mycorrhizal and non-mycorrhizal Medicago truncatula roots exhibit differentially regulated NADPH oxidase and antioxidant response under Pb stress[J]. 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Effects of Cd stress on seedling growth and activitiesin antioxidant enzymes of lettuce[J]. Ecology and Environmental Sciences, 2009, 18(2): 494-497. 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*表示接种处理与CK间差异显著(P<0.05);不同小写字母表示不同Pb浓度处理间差异显著(P<0.05)。

, figureFileSmall=SWtGDk2NhUEOP56Q/LAF5A==, figureFileBig=1WBoWU1oo9GLNU5QkR2bFA==, tableContent=null), ArticleFig(id=1276616214294172516, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276616206748619574, language=EN, label=Fig. 2, caption=Pb content in above-ground and root of W. sinensis with different treatments, figureFileSmall=Crb+jnh0iuG59lxxSglU9g==, figureFileBig=ZuM9UPEpYYKHPedA7maXwA==, tableContent=null), ArticleFig(id=1276616214365475685, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276616206748619574, language=CN, label=图2, caption=不同处理紫藤地上部和根系的Pb含量

*表示接种处理与CK间差异显著(P<0.05);不同小写字母表示不同Pb浓度处理间差异显著(P<0.05)。

, figureFileSmall=Crb+jnh0iuG59lxxSglU9g==, figureFileBig=ZuM9UPEpYYKHPedA7maXwA==, tableContent=null), ArticleFig(id=1276616214428390246, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276616206748619574, language=EN, label=Fig. 3, caption=BCF and TF of Pb in W. sinensis with different treatments, figureFileSmall=aSIz3Lh2TTfGnt0uOXnYWg==, figureFileBig=fpIAexG3F4xJ75RSpdN/5Q==, tableContent=null), ArticleFig(id=1276616214487110503, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276616206748619574, language=CN, label=图3, caption=不同处理紫藤的Pb富集系数与转移系数

*表示接种处理与CK间差异显著(P<0.05);不同小写字母表示不同Pb浓度处理间差异显著(P<0.05)。

, figureFileSmall=aSIz3Lh2TTfGnt0uOXnYWg==, figureFileBig=fpIAexG3F4xJ75RSpdN/5Q==, tableContent=null), ArticleFig(id=1276616214558413672, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276616206748619574, language=EN, label=Fig. 4, caption=Chlorophy content of W. sinensis leaves with different treatments, figureFileSmall=lp8maMhnY161Vf+ZqsWSbA==, figureFileBig=XWDIdPhDa1LxZiQ8opdOMA==, tableContent=null), ArticleFig(id=1276616214633911145, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276616206748619574, language=CN, label=图4, caption=不同处理紫藤叶片的叶绿素含量

*表示接种处理与CK间差异显著(P<0.05);不同小写字母表示不同Pb浓度处理间差异显著(P<0.05)。

, figureFileSmall=lp8maMhnY161Vf+ZqsWSbA==, figureFileBig=XWDIdPhDa1LxZiQ8opdOMA==, tableContent=null), ArticleFig(id=1276616214692631402, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276616206748619574, language=EN, label=Fig. 5, caption=Changes in physiological and biochemical indicators of W. sinensis leaves with different treatments, figureFileSmall=AfCPnXL5mtr7FVvp4HskNw==, figureFileBig=J9/Wn/R3SyZ22cS0OXxwSA==, tableContent=null), ArticleFig(id=1276616214772323179, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276616206748619574, language=CN, label=图5, caption=不同处理紫藤叶片相关生理生化指标的变化

*表示接种处理与CK间差异显著(P<0.05);不同小写字母表示不同Pb浓度处理间差异显著(P<0.05)。

, figureFileSmall=AfCPnXL5mtr7FVvp4HskNw==, figureFileBig=J9/Wn/R3SyZ22cS0OXxwSA==, tableContent=null), ArticleFig(id=1276616214839432044, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276616206748619574, language=EN, label=Fig. 6, caption=PCA and cluster heatmapm of growth and physiological and biochemical indicators of W. sinensis under AM inoculation and different Pb concentrations stress, figureFileSmall=SSV+WgJXRaLT0uXhKP8gFA==, figureFileBig=4oh3pnMliB1DBkGTAU3tcA==, tableContent=null), ArticleFig(id=1276616214898152301, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276616206748619574, language=CN, label=图6, caption=接种AM真菌和不同浓度Pb胁迫下紫藤生长和生理生化指标的PCA和聚类热图

A:PCA荷载图;B:PCA得分图;C:聚类热图。

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AM真菌对Pb污染土壤中紫藤生长及其生理的影响
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汪结明 , 彭虹云 , 李瑞雪 *
热带作物学报 | 作物栽培与生理生化 2025,46(8): 1921-1930
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热带作物学报 |作物栽培与生理生化 2025 , 46 (8) : 1921 -1930
AM真菌对Pb污染土壤中紫藤生长及其生理的影响
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汪结明, 彭虹云, 李瑞雪*
作者信息
  • 湖南科技大学建筑与设计学院,湖南湘潭 411201
通讯作者:
* 李瑞雪(LI Ruixue),E-mail:
Effects of AM Fungi on Wisteria sinensis to Pb Stress and Its Physiological Mechanism
Jieming WANG, Hongyun PENG, Ruixue LI*
Affiliations
  • School of Architecture and Design, Hunan University of Science and Technology, Xiangtan, Hunan 411201, China
出版时间: 2025-08-25 doi: 10.3969/j.issn.1000-2561.2025.08.015
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为探究丛枝菌根(AM)真菌对紫藤(Wisteria sinensis)扦插苗抗铅(Pb)胁迫的影响及其生理机理,以接种摩西斗管囊霉(Funneliformis mosseae)的紫藤扦插苗为材料,Pb污染浓度设置为400、800、1200 mg/kg,通过盆栽模拟试验,测定紫藤幼苗相关生长指标(生长量、生物量、耐性指数)和生理指标(叶绿素含量、丙二醛含量、脯氨酸含量、相关抗氧化酶活性)。结果表明:接种AM真菌能促进紫藤幼苗的地上部和根系生长,并提高紫藤对Pb的耐受性。转移系数以及地上部的富集系数低于CK,根系富集系数均高于CK,表明AM真菌降低了紫藤地上部的Pb含量并显著提高紫藤根系对Pb的富集能力。生理机制分析发现,在AM真菌作用下,紫藤叶片的叶绿素含量、超氧化物歧化酶(SOD)、过氧化物酶(POD)、抗坏血酸过氧化物酶(APX)等抗氧化酶活性均高于CK,丙二醛(MDA)积累量显著低于CK。主成分分析和聚类热图分析也显示,AM真菌主要通过激活抗氧化酶活性,调控相关生长指标及叶绿素合成途径来增强紫藤对Pb胁迫的耐受能力。该研究结果为菌根化植物修复重金属污染提供理论与实践依据。

丛枝菌根真菌  /  紫藤  /  Pb胁迫  /  生理机理

A controlled experiment was conducted using Wisteria sinensis seedlings inoculated with Funneliformis mosseae to investigate the effects and physiological mechanisms of arbuscular mycorrhizal fungi (AMF) on the Pb stress tolerance of W. sinensis seedlings. Pb contamination gradients were established (400, 800, 1200 mg/kg) to measure the growth parameters (growth, biomass, tolerance index) and physiological indicators (chlorophyll content, malondialdehyde content, proline content and related antioxidant enzyme activities). AM fungi inoculation significantly enhanced both aerial and root growth of W. sinensis seedlings while improving the Pb stress tolerance. The translocation factor and shoot bio-concentration factor were lower than those of the control, whereas the root bio-concentration factor exceeded the level of the control groups. This demonstrated that AM fungi effectively reduced Pb accumulation in the aerial parts while significantly enhancing Pb sequestration capacity in the roots. Physiological analysis indicated that AM fungi-treated seedlings exhibited higher chlorophyll content and elevated the activities of antioxidant enzymes (SOD, POD, APX), along with significantly lower MDA accumulation compared to the non-inoculated controls. Principal component analysis combined with cluster heat-map visualization further demonstrated that AM fungi primarily enhanced the Pb tolerance of W. sinensis through three mechanisms: activation of antioxidant enzyme systems, regulation of growth-related processes, and modulation of chlorophyll biosynthesis pathways. The results provide theoretical and practical basis for mycorrhizal plant remediation of heavy metal pollution.

AM fungi  /  Wisteria sinensis  /  Pb stress  /  physiological mechanism
汪结明, 彭虹云, 李瑞雪. AM真菌对Pb污染土壤中紫藤生长及其生理的影响. 热带作物学报, 2025 , 46 (8) : 1921 -1930 . DOI: 10.3969/j.issn.1000-2561.2025.08.015
Jieming WANG, Hongyun PENG, Ruixue LI. Effects of AM Fungi on Wisteria sinensis to Pb Stress and Its Physiological Mechanism[J]. Chinese Journal of Tropical Crops, 2025 , 46 (8) : 1921 -1930 . DOI: 10.3969/j.issn.1000-2561.2025.08.015
废弃矿山中残留的铅(Pb)在土壤中具有较强的生物累积性,能够通过植物根系吸收进入植物体内,进而影响植物的正常生长和发育[1]。铅可通过生物放大效应沿食物链传递,最终在人体形成剂量依赖性蓄积,危害人体健康[2]。研究表明,铅胁迫还会抑制植物的光合作用,降低叶绿素含量,影响植物的光能转化效率,在高浓度铅胁迫下,植物的生长受到显著抑制,表现为植株矮小、叶片发黄甚至枯萎[3]。因此,铅污染对生态系统和人类健康造成了严重威胁。传统的重金属尾矿污染土壤治理方法工作量大,成本高、效率低[4]。随着生态园林城市建设的加快推进,利用兼具观赏性的超富集园林植物修复重金属污染土壤,是一种绿色高效的生态治理方法。该方法安全高效、成本低且不产生二次污染,既能满足城市生态景观建设的需求,又能达到减轻重金属对环境的危害的目的,已成为研究热点[5]
丛枝菌根(arbuscular mycorrhizal,AM)真菌是一类广泛存在于自然生态系统中与植物根系形成共生关系的微生物,能够显著增强植物对重金属胁迫的耐受性[6]。有助于宿主植物扩大根系,刺激菌根分泌抗氧化物质来抵御重金属污染的胁迫,同时促进植物吸收营养元素[7]。已有研究表明,部分真菌不仅对植物具有促生效应,还能通过分泌活性物质来优化植物的生理机制以提高其修复污染的效率[8]。研究发现,王族海棠(Malus‘Royalty’)在接种AM真菌后,显著减少铅胁迫对植株生长的抑制,并且光合作用增强[9]。金盏菊(Calendula officinalis L.)在接种AM真菌后,富集重金属的能力也有所增强[10]。然而利用AM真菌联合植物进行土壤重金属修复,主要集中在Cd[11]、Mn[12]、Zn[13]等方面,而较少有关联合观赏藤本植物进行Pb污染修复的研究报道。紫藤(Wisteria sinensis)是豆科木质藤本植物,具有生长迅速、生物量大、寿命长、观赏性强、适应能力强等优点。周敏等[14]研究发现,紫藤对Pb有较强的富集能力,在Pb污染土壤修复方面具有较好的应用潜能。因此,利用丛枝菌根真菌与紫藤的联合修复技术为重金属污染土壤的修复提供了新的可能性。本研究以紫藤为研究对象,在接种AM真菌后进行不同Pb浓度处理,以探究AM真菌对紫藤幼苗Pb富集和转运的影响及其生理机制,为菌根化植物修复重金属污染土壤提供理论与实践依据。
供试材料为紫藤扦插苗,取自湖南湘潭仁芳园艺场。
供试土壤为红壤,取自湖南科技大学生物园植物繁育基地园土,土壤基本理化性质:pH为5.59,Pb含量为39.54 mg/kg,全氮含量为1.72 g/kg,速效氮含量为97.30 mg/kg,速效磷含量为20.50 mg/kg,速效钾含量为81.30 mg/kg。
供试菌株为丛枝菌根真菌摩西斗管囊霉(Funneliformis mosseae),来自亚热带丛枝菌根真菌资源保藏中心,由湖南科技大学生态实验室采用白三叶草(Trifolium repens L.)作为宿主植物扩繁。接种物为菌丝、孢子和菌根,活性孢子含量为每克15个,菌丝侵染率为61%。
湘潭重金属矿区土壤中的Pb平均浓度为401.15 mg/kg[15],综合污染指数达5级,为重度污染[16]。采用双因素完全随机区组设计。采用模拟Pb污染土壤培养,将繁育基地园土粉碎风干过5 mm筛,以液态形式加入Pb(NO32分析纯,以不加Pb(NO32的园土基质作为空白对照(T0)。在前期预试验的基础上,设置3个Pb污染土壤浓度:400、800、1200 mg/kg,编号依次为T1、T2、T3。将不同污染浓度的灭菌园土基质装入花盆中,在土壤深度为10 cm处平铺50 g丛枝菌根真菌菌剂(AM),选择同期扦插的生长健壮、长势一致的紫藤扦插苗,分别移栽至盆中,再覆园土,各浓度处理均设置不加菌剂为对照(CK),即共8个处理:CK-T0、AM-T0、CK-T1、AM-T1、CK-T2、AM-T2、CK-T3、AM-T3。利用原子吸收分光光度计(AA-70000,Shimadzu,日本)检测到T1、T2、T3土壤中的实测Pb浓度分别为397.87、795.96、1193.65 mg/kg。移栽后定期浇水,保持土壤湿润。为了减少Pb流失,花盆底部设置托盘并将渗出的水再回补至盆中土壤内。每个处理5盆,每个生物学重复3次,处理120 d后进行指标测定。
(1)生长指标。处理120 d后,测定各处理紫藤幼苗植株的主蔓长(AGL,cm)、地上部生物量(AGB,g)、根长(GL,cm)、根生物量(RW,g)。利用耐性指数(tolerance index,Ti)分析AM真菌作用下紫藤对Pb污染的耐受能力,耐性指数反映了植物对Pb耐受能力的大小。计算公式:Ti=处理组植物生长参数/对照组植物生长参数,耐性指数为各处理的Ti均值。
(2)Pb含量指标。利用原子吸收分光光度计检测紫藤不同部位的Pb含量及土壤中的Pb含量。利用检测到的Pb浓度,分别计算其Pb转运系数(translocation factors,TF)及生物富集系数(bio-concentration factors,BCF),分析其修复效率,探讨其转运机制。计算公式:TF=地上部平均Pb浓度/根部平均Pb浓度;BCF=地上(地下)部平均Pb浓度/土壤平均Pb浓度。
(3)生理生化指标。使用乙醇-丙酮混合溶液浸泡法测定叶绿素含量[17],包括叶绿素a(chl.a)、叶绿素b(chl.b)、总叶绿素含量(chl.a+b);采用氮蓝四唑光化还原法测定超氧化物歧化酶(SOD)活性;采用硫代巴比妥酸法测定丙二醛(MDA)含量[18];采用愈创木酚法测定过氧化物酶(POD)活性;采用抗坏血酸比色法测定抗坏血酸过氧化物酶(APX)活性[19];采用试剂盒(Solarbio,BC0295)测定脯氨酸(Pro)含量。
使用Excel 2020、SPSS 8.0、Origin 2022等软件整理、分析数据;利用Origin 2022软件绘制图表。
图1A图1B所示,随Pb处理浓度的升高,紫藤扦插苗的主蔓长和根长呈持续下降的趋势,表明Pb对紫藤的生长造成了一定伤害,且Pb浓度越高伤害越大。T2、T3处理中,接种AM真菌的主蔓长和根长均显著高于CK,表明接种AM真菌对Pb胁迫下的紫藤生长具有显著促进作用。在AM真菌作用下各处理植株的主蔓长和根长的下降幅度均小于CK,T2与T1处理相比,CK的主蔓长下降21.1%,而AM真菌处理仅下降3.8%。说明接种AM真菌提高了紫藤对Pb胁迫的耐受性。随着土壤Pb污染浓度升高至1200 mg/kg(T3),植株主蔓长和根长均降至最低,且下降幅度高于T2处理,但AM真菌处理后的下降幅度小于CK,说明T3处理时Pb污染已对紫藤的生长造成了显著伤害,但AM真菌能缓解Pb污染对植株的伤害,但有一定阈值范围。
图1C图1D可看出,除T0处理的紫藤地上部生物量外,其他不同Pb浓度处理接种AM真菌的紫藤地上部生物量和根生物量均高于CK。随着Pb处理浓度的增加,紫藤地上部生物量和和根生物量均持续下降。T2处理中接种AM真菌的紫藤地上部生物量和根生物量均显著下降,并且CK的地上部生物量和根生物量均显著低于AM真菌处理,表明T2处理的Pb污染未对接种AM真菌的紫藤造成显著胁迫,与CK相比,AM真菌促进了紫藤地上部和根的生长,分别提高了26.7%和32.0%,说明AM真菌提高了紫藤对Pb胁迫的耐受性。T3处理时,地上部和根生物量均降至最低,与T2相比,其下降幅度均达到显著水平,说明T3处理的Pb污染浓度已对接种AM真菌的紫藤造成了显著伤害,同时也表明AM真菌对提高紫藤的抗Pb胁迫的能力是有阈值范围的。
图1E可知,紫藤的耐性指数(Ti)随着Pb处理浓度的增加持续下降,但AM真菌作用下各处理Ti的下降幅度均小于CK。T2处理中,CK的耐性指数仅为78.0%,而接种AM真菌的耐性指数却达到了84.2%,表明与CK相比,接种AM真菌提高了紫藤对Pb污染的耐受性。T3处理中,CK与AM真菌处理的耐性指数均降至最低,分别为41.7%和63.1%,说明1200 mg/kg(T3)的Pb污染浓度对接种AM真菌的紫藤造成了显著伤害,表明AM真菌提高了紫藤对Pb胁迫的耐受能力,但耐受Pb胁迫浓度已达最高值。
图2可看出,随着Pb处理浓度的升高,紫藤地上部和根系的Pb含量均呈先升后降的变化趋势,表明植株中Pb富集量随着Pb处理浓度的升高而增加,但在超过阈值后不会再富集Pb,导致T3处理的植株中Pb富集量反而降低。T2处理中,AM真菌处理的植株地上部和根系的Pb含量均显著高于CK,分别比CK提高了18.3%和31.0%,说明AM真菌提高了紫藤对Pb的富集能力,且根系的富集提高幅度更高。与T2处理相比,T3处理地上部和根系的Pb含量大幅下降,说明AM真菌对提高紫藤对Pb的富集能力有一定阈值,在T2处理时已基本达到了富集上限。
图3A图3B可知,在各Pb浓度处理中,接种AM真菌的紫藤扦插苗地上部和根系富集Pb的能力均显著高于CK,T1、T2处理的提升最大,分别提高了19.5%和20.0%。随着Pb胁迫浓度的增加,地上部及根系的Pb富集能力逐渐下降,在T3时降至最低,表明AM真菌能显著提高紫藤地上部及根系对Pb的富集能力。综上,Pb污染下AM真菌处理能够通过增强紫藤地上部及根系的富集能力来减少Pb对植株的伤害,但其富集能力是有阈值范围的。
图3C中可知,各处理的转运系数均小于1,说明各处理对Pb的转运能力均尚未达到超富集水平。由2.3可知,T1处理中,AM处理的植株Pb含量显著上升,并且根系中的上升幅度高于地上部,故转移系数均小于CK,但差异未达显著水平。说明随着Pb处理浓度的升高,接种AM真菌提高了紫藤扦插苗的Pb富集能力,对根的影响更明显。
图4可以看出,叶绿素a、叶绿素b和总叶绿素含量均随着Pb处理浓度的升高总体呈下降趋势,说明Pb胁迫抑制了紫藤叶绿素的合成。接种AM真菌后,各Pb浓度处理的叶绿素a、叶绿素b、总叶绿素含量均显著高于CK,其中,T2处理的总叶绿素含量比CK提高了21.4%,并且T2处理的叶绿素a与叶绿素b含量的比值也明显升高。说明AM真菌能显著缓解Pb胁迫对紫藤叶绿素合成的抑制。
超氧化物歧化酶、过氧化物酶、抗坏血酸过氧化物酶是抗氧化防御系统中重要的保护酶,可以清除植物在生理活动中产生的过多的自由基和过氧化物,从而减轻植物受到的毒害,这些酶的活性水平与植物抗胁迫能力具有一定相关性。如图5A~图5C所示,随着Pb处理浓度的升高,紫藤的3种抗氧化酶活性均呈先升后降的趋势,并且这3种酶的活性在T2处理中均达最大值。接种AM真菌后,各Pb浓度处理的酶活性均高于CK,表明AM真菌处理显著提高了抗氧化酶防御系统的酶活性,从而显著提高了紫藤抗Pb胁迫的能力。随着Pb处理浓度的继续提高,T3处理的酶活性却降低,这表明AM真菌促进紫藤对土壤Pb污染的修复效应具有一定阈值。
在Pb胁迫下,紫藤扦插苗的脯氨酸含量总体呈先升后降的趋势(图5D)。接种AM真菌后,与CK比,各处理的Pro含量分别提升了1.3%、11.5%、14.0%、10.4%。随着Pb处理浓度的提高,紫藤扦插苗的丙二醛含量持续升高,并在T3处理中CK的MDA含量达到最大值(图5E),表明Pb浓度越高对紫藤胁迫越强。接种AM真菌后,各处理的MDA含量均低于CK,且随Pb处理浓度的增加,接种AM真菌的各处理呈先升后降的趋势,而CK却显著增加。这表明接种AM真菌降低了MDA的累积,减轻了氧化应激,从而显著提高了紫藤抗Pb胁迫的能力。
采用主成分分析法(PCA)对所有检测指标进行综合分析,结果发现,前2个主成分PC1和PC2表达了总方差的60.5%,其中PC1包括叶绿素a含量、叶绿素b含量、总叶绿素含量、主蔓长、根长、地上部生物量、根生物量;PC2包括MDA含量、SOD活性、POD活性、AXP活性、Pro含量。如图6A图6B所示,在不同Pb浓度处理下,接种AM真菌处理与CK明显地分为两类。这一结果初步表明,接种AM真菌对提高紫藤幼苗抗Pb胁迫是有效的。
以皮尔逊相关性为聚类内核进行聚类分析,如图6C所示,各检测指标被分为3簇。a簇包括总叶绿素含量、叶绿素a含量、叶绿素b含量等指标,b簇包括Pro含量、APX活性、POD活性、SOD活性、MAD含量等指标,c簇包括根生物量、地上部生物量、根长、主蔓长等指标。聚类热图中,颜色的深浅直观反映了不同处理条件下各指标的变化趋势。随着Pb胁迫浓度的升高,b簇的各指标整体呈上升趋势,表现为颜色由浅到深,表明紫藤通过提高相关酶活性以缓解Pb胁迫的危害。相对地,a簇和c簇中接种AM的各项指标的颜色整体上比CK深,表明AM真菌能有效提高各项指标从而增强紫藤抗Pb胁迫的能力。
ZANGANEH等[20]研究发现,Pb污染会抑制株高发育并破坏其抗氧化系统,从而干扰植物的正常生理过程而引发胁迫反应。本研究也发现在3种不同浓度的Pb胁迫下,与不加Pb的对照组相比,紫藤幼苗的生长受到显著抑制,且随Pb处理浓度的提高其生长指标整体呈下降趋势。
HUSSAIN等[21]研究发现,AM真菌通过促进植物根系生长帮助植物获取土壤中更多的养分,从而对植株生物量产生有益影响。本研究也发现,接种AM真菌能促进紫藤生长,但AM真菌在不同浓度Pb胁迫中的促生效果存在差异,在T2、T3处理中的促生效果更明显。表明AM真菌在一定程度上能够缓解Pb胁迫对紫藤幼苗生长的抑制作用。该促生效果可能与AM真菌促进了紫藤对土壤营养元素的吸收有关,但具体促进了哪种元素有待后续进一步研究。也可能是植物的一种自我保护机制,即紫藤在受到Pb胁迫后,在与AM真菌作用下通过提高自身生长量和生物量,以降低Pb在植株内的相对浓度,从而达到保护自身的目的。
AM真菌对紫藤的Pb富集效应产生了重要影响。已有研究表明,AM真菌可通过降低西芹(Apium graveolens Linn.)地上部Cd含量以及富集系数,增加根部Cd含量,从而降低转移系数来缓解Cd污染带来的损伤,而且可降低Cd污染土壤中植株地上部的Cd含量[7]。TAN等[22]研究发现,植株接种AM真菌后,向地上部的重金属转移系数降低,使重金属固定在植物根系中。本研究也发现,AM真菌处理后地上部及根系的Pb含量、根系富集系数均高于CK,转移系数稍低于CK,表明AM真菌能显著提高紫藤对Pb的富集能力,且根部富集能力提高的幅度高于地上部。WANG等[23]发现Cd污染下接种AM真菌能够提高龙葵根系中的Cd浓度,降低地上部的Cd浓度。而邱丹[24]则研究发现砷(As)污染下接种AM真菌后,使As主要累积在蜈蚣草的地上部,降低了根系中的As浓度。这可能是由于AM真菌对不同植物、不同重金属的转运、富集作用存在差异。
Pb胁迫对光系统Ⅰ产生影响,光合作用的产物由于光合电子传递速率下降无法正常分解运转,直接影响光捕获能力[9]。在对景天三七(Sedum aizoon L.)的研究中发现,随着铅离子浓度的增加,景天三七的叶绿素含量总体呈下降趋势[25]。白茅[lmperata cylindrica(L.)P. Beauv.]在Pb胁迫下总叶绿素含量显著降低,接种AM真菌后总叶绿素含量显著提高[26]。本研究也发现,随着Pb处理浓度的提高,叶绿素含量逐渐降低,T2处理的叶绿素总量相较于CK显著增加。由此可知,Pb胁迫抑制了紫藤幼苗叶绿素的合成,接种AM真菌缓解了Pb对叶绿素合成的抑制,但缓解机制有待后续研究。
当植物遭受重金属胁迫时,由SOD和POD共同组成的活性氧清除系统被激活,能够有效抑制氧自由基的积累,以避免自由基对植物细胞造成潜在的负面效应[27]。研究发现,Pb胁迫增加了蒺藜(Tribulus terrestris L.)中超氧负离子及MDA含量,而接种AM真菌后,SOD、POD、APX活性增高[28]。Pro是植物体内重要的渗透调节物质和营养物质,其含量越高说明植物体的代谢活动越强[29]。本研究也发现,紫藤的SOD、POD、APX活性及Pro含量均呈先升后降的趋势。表明AM真菌在低浓度Pb处理时能够通过调节抗氧化酶系统活性以提高植株对Pb的耐受性,但在较高浓度Pb胁迫下,由于植株受胁迫程度过大,可能导致抗氧化酶系统调节效率下降。紫藤的MDA含量也随Pb浓度的升高而升高,与CK相比,接种AM真菌后MDA含量显著降低,表明在Pb胁迫下,AM真菌能够降低植物的MDA含量,以缓解Pb的毒性,进一步证实了AM真菌对紫藤幼苗抗氧化能力的调控作用,且这种调控作用因Pb浓度的不同而有所差异。
当前铅锌矿区等重金属污染严重的区域,土壤肥力低下、植被覆盖率低,传统的修复方法往往难以奏效。紫藤对Pb具有一定的富集能力,与AM真菌联合,紫藤可作为修复Pb污染的先锋植物,可增强生态修复的效果。AM真菌联合紫藤修复技术可以改善Pb污染的土壤质量,促进植被恢复,加速土壤生态系统的恢复,同时也可以发挥紫藤的观赏作用让矿区景观化。因此,该技术在农业土壤生态修复、矿山公园植物营建、观赏植物菌根化工厂化育苗等方面的应用前景广阔。
尽管AM真菌与紫藤的联合修复技术在Pb污染土壤的治理中展现出良好的应用前景,但其实际应用仍存在一定局限性。AM真菌与紫藤的修复效果易受环境因素的影响,并且当土壤中的Pb浓度超过一定限度时,Pb会对AM真菌和紫藤产生严重的毒害作用,导致AM-紫藤共生体的生长受阻,从而无法达到更高的修复目标,因此,为了AM真菌与紫藤的联合修复技术的高效应用仍需开展后续深度研究。
Pd污染下紫藤植株地上部及其根生长量、生物量呈下降趋势,叶绿素合成等受到抑制,抗氧化系统被打破。接种AM真菌后紫藤幼苗在Pb污染土壤中的生长均高于对照,表现出良好的促生效果,显著提升Pb污染土壤中紫藤的地上部及其根生长量、生物量,并提高了紫藤对Pb污染的耐受性。接种AM真菌增强了紫藤根系对Pb的富集能力。AM真菌主要通过增强抗氧化酶活性和Pro含量,缓解Pb对叶绿素合成的胁迫影响等途径,有效增强了紫藤幼苗抗Pb胁迫的能力,紫藤接种AM真菌的菌-苗联合模式可作为修复重金属Pb污染的有效途径。
  • 国家自然科学基金项目(31960327)
  • 教育部人文社会科学研究规划基金项目(24YJAZH144)
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doi: 10.3969/j.issn.1000-2561.2025.08.015
  • 接收时间:2025-02-28
  • 首发时间:2026-06-24
  • 出版时间:2025-08-25
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  • 收稿日期:2025-02-28
  • 录用日期:2025-05-08
基金
国家自然科学基金项目(31960327)
教育部人文社会科学研究规划基金项目(24YJAZH144)
作者信息
    湖南科技大学建筑与设计学院,湖南湘潭 411201

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* 李瑞雪(LI Ruixue),E-mail:
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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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