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Article(id=1215670319793225898, tenantId=1146029695717560320, journalId=1149652044408987649, issueId=1215670311140381365, articleNumber=null, orderNo=null, doi=10.19812/j.cnki.jfsq11-5956/ts.20250318001, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1742227200000, receivedDateStr=2025-03-18, revisedDate=null, revisedDateStr=null, acceptedDate=null, acceptedDateStr=null, onlineDate=1767767990301, onlineDateStr=2026-01-07, pubDate=1753372800000, pubDateStr=2025-07-25, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1767767990301, onlineIssueDateStr=2026-01-07, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1767767990301, creator=13701087609, updateTime=1767767990301, updator=13701087609, issue=Issue{id=1215670311140381365, tenantId=1146029695717560320, journalId=1149652044408987649, year='2025', volume='16', issue='14', pageStart='1', pageEnd='326', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1767767988237, creator=13701087609, updateTime=1767970098618, updator=13701087609, preIssue=null, nextIssue=null, ext={EN=IssueExt(id=1216518023599538606, tenantId=1146029695717560320, journalId=1149652044408987649, issueId=1215670311140381365, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1216518023599538607, tenantId=1146029695717560320, journalId=1149652044408987649, issueId=1215670311140381365, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null}, startPage=120, endPage=127, ext={EN=ArticleExt(id=1215670320195879132, articleId=1215670319793225898, tenantId=1146029695717560320, journalId=1149652044408987649, language=EN, title=Research progress on Fusarium pathogens and food security, columnId=1151895321388347923, journalTitle=Journal of Food Safety & Quality, columnName=Food Analysis and Detection, runingTitle=null, highlight=null, articleAbstract=

Fusarium is a ubiquitous pathogen worldwide that can affect the growth and development of plants and pose a serious threat to global food security and biodiversity. Almost all Fusarium bacteria produce mycotoxins, which are diverse and pathogenic. The most toxic ones include trichothecenes, zearalenone (ZEA) and fumonisins. On the one hand, mycotoxins, as pathogenic agents, can infect a wide range of plants, leading to symptoms such as stunted growth, fruit or seed rot, yellowing of leaves, wilting, ulceration, and root or stem rot. These effects result in significant crop damage and economic losses, making mycotoxin contamination a major concern in agriculture. On the other hand, Fusarium is also considered to be the causative agent of fungal infections in humans and animals, and its toxins contaminate cereals and food, causing related pathologies in humans and animals, and have a wide range of clinical manifestations in immunocompromised patients. Most often, Fusarium is difficult to control because the fungus easily overcomes host resistance to various control methods. This review summarized the latest research on Fusarium, exploring its interaction mechanisms with plants and its impact on food security. Additionally, it provided a detailed overview of various methods for detecting Fusarium, aiming to promote research on the comprehensive management of this inevitable food contaminant to support the sustainable development of global food security.

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镰刀菌是世界范围内一种普遍存在的病原菌, 影响植物的生长发育, 严重威胁全球粮食安全和生物多样性。几乎所有的镰刀菌都会产生真菌毒素, 其毒素种类繁多且致病性强。毒性较大的主要有单端孢霉烯族毒素、玉米赤霉烯酮(zearalenone, ZEA)和伏马菌素等。一方面, 真菌毒素作为致病因子可感染多种植物, 引起生长发育迟缓、果实或种子腐烂、叶子发黄、枯萎、溃烂、根或茎腐烂等症状, 造成严重的作物损失与经济损失, 是农业中备受关注的主题; 另一方面, 镰刀菌也被认为是人类和动物真菌感染的病原体, 其毒素污染粮食和食物, 引起人类和动物的相关病症, 在免疫功能低下的患者中具有广泛的临床表现。大多数情况下, 镰刀菌病很难控制, 因为真菌很容易克服寄主对各种控制方法的抗性。本文综述了镰刀菌的最新研究进展, 探讨了其与植物互作机制及其对粮食安全的影响。同时, 详细介绍了识别镰刀菌的不同方法, 旨在推动对这种不可避免的食品污染物进行综合管理的研究, 以促进全球粮食安全的可持续发展。

, correspAuthors=平文丽, authorNote=null, correspAuthorsNote=
*平文丽(1982—), 女, 博士, 助理研究员, 主要研究方向为烟草分子抗病育种。E-mail:
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张雪珂(1994—), 女, 硕士, 研究实习员, 主要研究方向为烟草分子抗病育种。E-mail:

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张雪珂(1994—), 女, 硕士, 研究实习员, 主要研究方向为烟草分子抗病育种。E-mail:

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张雪珂(1994—), 女, 硕士, 研究实习员, 主要研究方向为烟草分子抗病育种。E-mail:

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Nature Climate Change, 2013, 3: 985-988., articleTitle=Crop pests and pathogens move pole wards in a warming world, refAbstract=null)], funds=[Fund(id=1215670327309419244, tenantId=1146029695717560320, journalId=1149652044408987649, articleId=1215670319793225898, awardId=2024ZC044, language=CN, fundingSource=河南省农业科学院自主创新项目(2024ZC044), fundOrder=null, country=null), Fund(id=1215670327506551550, tenantId=1146029695717560320, journalId=1149652044408987649, articleId=1215670319793225898, awardId=2025ZC100, language=CN, fundingSource=河南省农业科学院自主创新项目(2025ZC100), fundOrder=null, country=null)], companyList=[AuthorCompany(id=1215670322523717987, tenantId=1146029695717560320, journalId=1149652044408987649, articleId=1215670319793225898, xref=null, ext=[AuthorCompanyExt(id=1215670322532106597, tenantId=1146029695717560320, journalId=1149652044408987649, articleId=1215670319793225898, companyId=1215670322523717987, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=Key Laboratory for Green Preservation & Control of Tobacco Diseases and Pests in Huanghuai Growing Area, Tobacco Research Institute, Henan Academy of Agricultural Sciences, Xuchang 461000, China), AuthorCompanyExt(id=1215670322536300901, tenantId=1146029695717560320, journalId=1149652044408987649, articleId=1215670319793225898, companyId=1215670322523717987, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=河南省农业科学院烟草研究所, 烟草行业黄淮烟区烟草病虫害绿色防控重点实验室, 许昌 461000)])], figs=[ArticleFig(id=1215670325048689300, tenantId=1146029695717560320, journalId=1149652044408987649, articleId=1215670319793225898, language=EN, label=Fig.1, caption=Morphological characteristics of Fusarium conidia[13], figureFileSmall=Jn9UMJBqT5hgkIZ3G39ezw==, figureFileBig=Khtp6dH/jCXL2uSFdqPgYQ==, tableContent=null), ArticleFig(id=1215670325199684258, tenantId=1146029695717560320, journalId=1149652044408987649, articleId=1215670319793225898, language=CN, label=图1, caption=镰刀菌分生孢子形态特征[13]

注: A~D. 大分生孢子形态; E~H. 大分生孢子顶端细胞形状; I~L. 大分生孢子基细胞形状; M. 小分生孢子形状; N. 小分生孢子形态; O. 微分生孢子链。

, figureFileSmall=Jn9UMJBqT5hgkIZ3G39ezw==, figureFileBig=Khtp6dH/jCXL2uSFdqPgYQ==, tableContent=null), ArticleFig(id=1215670325627503293, tenantId=1146029695717560320, journalId=1149652044408987649, articleId=1215670319793225898, language=EN, label=Fig.2, caption=Model of interaction between Fusarium mycotoxins and plants, figureFileSmall=Xue4WbxMo7Ab92oyrmZLPA==, figureFileBig=N0rrYTdBflytDUA5Xuo7Dg==, tableContent=null), ArticleFig(id=1215670326881600198, tenantId=1146029695717560320, journalId=1149652044408987649, articleId=1215670319793225898, language=CN, label=图2, caption=镰刀菌属毒素与植物互作的模式图

注: 病原相关分子模式(pathogen associated molecular pattern, PAMP); 病原体相关分子模式触发免疫(PAMP triggered immunity, PTI); 效应子触发免疫反应(effector triggered immunity, ETI); 活性氧(reactive oxygen species, ROS); 程序性细胞死亡(programmed cell death, PCD)。

, figureFileSmall=Xue4WbxMo7Ab92oyrmZLPA==, figureFileBig=N0rrYTdBflytDUA5Xuo7Dg==, tableContent=null), ArticleFig(id=1215670326994846417, tenantId=1146029695717560320, journalId=1149652044408987649, articleId=1215670319793225898, language=EN, label=Table 1, caption=

Fusarium species and their major mycotoxins

, figureFileSmall=null, figureFileBig=null, tableContent=
镰刀菌属 产生的主要霉菌毒素
黄色镰刀菌(Fusarium culmorum) 脱氧雪腐镰刀菌醇, 3-乙酰脱氧雪腐镰刀菌烯醇, 15-乙酰脱氧雪腐镰刀菌烯醇, 雪腐镶刀菌醇, 镰刀菌烯酮, 玉米赤烯
木贼镰刀菌(Fusarium equiseti) 玉米赤烯, 二乙酰氧基马兜铃醇
禾谷镰刀菌(Fusarium graminearium) 脱氧雪腐镰刀菌醇, 15-乙酰脱氧雪腐镰刀菌烯醇, 雪腐镶刀菌醇, 镰刀菌烯酮, 玉米赤烯
尖孢镰刀菌(Fusarium oxysporiuun) 脱氧雪腐镰刀菌醇, 恩镰孢菌素A, 恩镰孢菌素A1, 恩镰孢菌素B1, 串珠镰刀菌素, 白僵菌素, 镰刀菌酸
梨孢镰刀菌(Fusarium poae) T-2毒素, HT-2毒素, 雪腐镶刀菌醇, 二乙酰氧基马兜铃醇, 镰刀菌烯酮
层出镰刀菌(Fusarium proliferatum) 伏马菌素B1, 串珠镰刀菌素, 白僵菌素
茄病镰刀菌(Fusarium solani) 伏马菌素B1, 伏马菌素B2, 串珠镰刀菌素, 镰刀菌酸
拟枝孢镰刀菌(Fusarium sporotrichioides) T-2毒素, HT-2毒素, 新茄病镰刀菌烯醇, 二乙酰氧基马兜铃醇, 镰刀菌烯酮, 玉米赤烯
轮枝样镰刀菌(Fusarium verticiloides) 伏马菌素B1, 伏马菌素B2, 串珠镰刀菌素, 白僵菌素, 镰刀菌酸
接骨木镰刀菌(Fusarium sambucinum) 脱氧雪腐镰刀菌醇, 二乙酰氧基马兜铃醇, T-2毒素
), ArticleFig(id=1215670327145841374, tenantId=1146029695717560320, journalId=1149652044408987649, articleId=1215670319793225898, language=CN, label=表1, caption=

镰刀菌种类及其主要真菌毒素

, figureFileSmall=null, figureFileBig=null, tableContent=
镰刀菌属 产生的主要霉菌毒素
黄色镰刀菌(Fusarium culmorum) 脱氧雪腐镰刀菌醇, 3-乙酰脱氧雪腐镰刀菌烯醇, 15-乙酰脱氧雪腐镰刀菌烯醇, 雪腐镶刀菌醇, 镰刀菌烯酮, 玉米赤烯
木贼镰刀菌(Fusarium equiseti) 玉米赤烯, 二乙酰氧基马兜铃醇
禾谷镰刀菌(Fusarium graminearium) 脱氧雪腐镰刀菌醇, 15-乙酰脱氧雪腐镰刀菌烯醇, 雪腐镶刀菌醇, 镰刀菌烯酮, 玉米赤烯
尖孢镰刀菌(Fusarium oxysporiuun) 脱氧雪腐镰刀菌醇, 恩镰孢菌素A, 恩镰孢菌素A1, 恩镰孢菌素B1, 串珠镰刀菌素, 白僵菌素, 镰刀菌酸
梨孢镰刀菌(Fusarium poae) T-2毒素, HT-2毒素, 雪腐镶刀菌醇, 二乙酰氧基马兜铃醇, 镰刀菌烯酮
层出镰刀菌(Fusarium proliferatum) 伏马菌素B1, 串珠镰刀菌素, 白僵菌素
茄病镰刀菌(Fusarium solani) 伏马菌素B1, 伏马菌素B2, 串珠镰刀菌素, 镰刀菌酸
拟枝孢镰刀菌(Fusarium sporotrichioides) T-2毒素, HT-2毒素, 新茄病镰刀菌烯醇, 二乙酰氧基马兜铃醇, 镰刀菌烯酮, 玉米赤烯
轮枝样镰刀菌(Fusarium verticiloides) 伏马菌素B1, 伏马菌素B2, 串珠镰刀菌素, 白僵菌素, 镰刀菌酸
接骨木镰刀菌(Fusarium sambucinum) 脱氧雪腐镰刀菌醇, 二乙酰氧基马兜铃醇, T-2毒素
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镰刀菌病原体与粮食安全研究进展
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张雪珂 , 平文丽 *
食品安全质量检测学报 | 食品分析与检测 2025,16(14): 120-127
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食品安全质量检测学报 | 食品分析与检测 2025, 16(14): 120-127
镰刀菌病原体与粮食安全研究进展
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张雪珂 , 平文丽*
作者信息
  • 河南省农业科学院烟草研究所, 烟草行业黄淮烟区烟草病虫害绿色防控重点实验室, 许昌 461000

    • 张雪珂(1994—), 女, 硕士, 研究实习员, 主要研究方向为烟草分子抗病育种。E-mail:

通讯作者:

*平文丽(1982—), 女, 博士, 助理研究员, 主要研究方向为烟草分子抗病育种。E-mail:
Research progress on Fusarium pathogens and food security
Xue-Ke ZHANG , Wen-Li PING*
Affiliations
  • Key Laboratory for Green Preservation & Control of Tobacco Diseases and Pests in Huanghuai Growing Area, Tobacco Research Institute, Henan Academy of Agricultural Sciences, Xuchang 461000, China
出版时间: 2025-07-25 doi: 10.19812/j.cnki.jfsq11-5956/ts.20250318001
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摘要
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镰刀菌是世界范围内一种普遍存在的病原菌, 影响植物的生长发育, 严重威胁全球粮食安全和生物多样性。几乎所有的镰刀菌都会产生真菌毒素, 其毒素种类繁多且致病性强。毒性较大的主要有单端孢霉烯族毒素、玉米赤霉烯酮(zearalenone, ZEA)和伏马菌素等。一方面, 真菌毒素作为致病因子可感染多种植物, 引起生长发育迟缓、果实或种子腐烂、叶子发黄、枯萎、溃烂、根或茎腐烂等症状, 造成严重的作物损失与经济损失, 是农业中备受关注的主题; 另一方面, 镰刀菌也被认为是人类和动物真菌感染的病原体, 其毒素污染粮食和食物, 引起人类和动物的相关病症, 在免疫功能低下的患者中具有广泛的临床表现。大多数情况下, 镰刀菌病很难控制, 因为真菌很容易克服寄主对各种控制方法的抗性。本文综述了镰刀菌的最新研究进展, 探讨了其与植物互作机制及其对粮食安全的影响。同时, 详细介绍了识别镰刀菌的不同方法, 旨在推动对这种不可避免的食品污染物进行综合管理的研究, 以促进全球粮食安全的可持续发展。

镰刀菌  /  真菌  /  病原体  /  鉴定  /  病原菌与植物互作  /  疾病管理  /  粮食安全

Fusarium is a ubiquitous pathogen worldwide that can affect the growth and development of plants and pose a serious threat to global food security and biodiversity. Almost all Fusarium bacteria produce mycotoxins, which are diverse and pathogenic. The most toxic ones include trichothecenes, zearalenone (ZEA) and fumonisins. On the one hand, mycotoxins, as pathogenic agents, can infect a wide range of plants, leading to symptoms such as stunted growth, fruit or seed rot, yellowing of leaves, wilting, ulceration, and root or stem rot. These effects result in significant crop damage and economic losses, making mycotoxin contamination a major concern in agriculture. On the other hand, Fusarium is also considered to be the causative agent of fungal infections in humans and animals, and its toxins contaminate cereals and food, causing related pathologies in humans and animals, and have a wide range of clinical manifestations in immunocompromised patients. Most often, Fusarium is difficult to control because the fungus easily overcomes host resistance to various control methods. This review summarized the latest research on Fusarium, exploring its interaction mechanisms with plants and its impact on food security. Additionally, it provided a detailed overview of various methods for detecting Fusarium, aiming to promote research on the comprehensive management of this inevitable food contaminant to support the sustainable development of global food security.

Fusarium  /  fungi  /  pathogens  /  identification  /  pathogen and plant interaction  /  disease management  /  food security
张雪珂, 平文丽. 镰刀菌病原体与粮食安全研究进展. 食品安全质量检测学报, 2025 , 16 (14) : 120 -127 . DOI: 10.19812/j.cnki.jfsq11-5956/ts.20250318001
Xue-Ke ZHANG, Wen-Li PING. Research progress on Fusarium pathogens and food security[J]. Journal of Food Safety & Quality, 2025 , 16 (14) : 120 -127 . DOI: 10.19812/j.cnki.jfsq11-5956/ts.20250318001
正文
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0 引言
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超过70%的植物病害可以追溯到威胁粮食安全的真菌或类真菌病原体[1]。此外, 包括镰刀菌在内的许多真菌病原体也会产生真菌毒素, 进一步威胁人类和牲畜的健康, 摄入真菌毒素会造成急性或慢性中毒, 引起肠道、肝脏、肾脏等器官的毒性反应, 甚至可能具有致癌性[2-3]。因此, 镰刀菌对作物生产中的食品安全具有重要影响。镰刀菌属真菌可感染包括小麦、棉花、水稻、番茄、香蕉、茄子及烟草等在内的100多种植物, 是全球重要的植物病原菌之一[4-5]。镰刀菌适应多种环境, 尤其偏好温暖气候, 通常在收获前的田间产生真菌毒素, 其中最具毒性和传播性的包括单端孢霉烯族毒素、伏马菌素和玉米赤霉烯酮(zearalenone, ZEA)等。镰刀菌可能引起的疾病症状包括根或茎腐烂、溃疡病、维管束枯萎、果实或种子腐烂以及叶片疾病等, 往往造成严重的社会经济损失[6]。它们的寄主范围广、内生感染性强、生存和传播方式多样, 是重要的植物病原体。随着新型真菌病原体的出现, 粮食生产和生态环境面临更大威胁, 因此有效控制真菌病害显得尤为迫切且具有挑战性。本文将综述镰刀菌的分类、形态学与分子鉴定技术、毒素与植物互作机制以及其对粮食安全的影响, 旨在提升社会对粮食安全问题的关注, 并为未来的防控策略、作物育种、农业政策制定及社会意识的提升提供参考和有价值的信息。
1 镰刀菌的概述与分类学
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镰刀菌是一大类丝状真菌, 主要存在于空气和土壤中, 在世界范围内, 目前已知的一些重要的植物致病性真菌物种均是镰刀菌属的成员, 可侵染包括小麦、棉花、水稻、番茄、香蕉、茄子和烟草等在内的100多种植物, 是世界上最重要的植物病原菌之一。此外, 镰刀菌真菌也会侵染免疫力低的人和动物, 增加癌症的发生率和引起人和动物的生长缺陷、发育迟缓等疾病, 严重时可危及生命安全并引起生物个体的死亡[7-8]。
镰刀菌属真菌的分类研究是世界上公认的难题, 最早的分类始于1809年, LINK以粉红镰孢菌为模式种建立了镰孢菌属(Fusarium Link)[9]。1821年, FRIES将它放在半知菌的瘤座菌科(Tuberculariae)[9]。1935年第一个较完整的镰孢菌属分类系统被提出, WOLLENWEBER等[10]将两个玫瑰镰刀菌的模式标本重新分类为接骨木镰孢(Fusarium sambucinum)和禾谷镰刀菌(Fusarium graminearum), 目前认为接骨木镰孢为该属的模式种。虽然镰刀菌的分类学继续发生重大变化, 主要以分子分类为基础, 但WOLLENWEBER和REINKING分类系统依然是物种描述的基础。镰刀菌属成员的特征是具有分离的、透明的、形状弯曲且细长的大分生孢子[11-12], 菌丝体和孢子团通常颜色明亮[12]。镰刀菌属包括约70个公认的物种, 通过使用多种方法鉴定, 推定物种约有300个。根据系统发育的物种概念, 许多假定的物种尚未被正式命名[13]。
2 植物致病性镰刀菌真菌种类鉴定
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2.1 植物致病性镰刀菌种类鉴定的形态学特征
形态特征是用来鉴定任何真菌物种的最传统标准。镰刀菌产生一系列棉质的菌丝, 颜色有白色、粉红色、黄色和紫色。一些物种产生大分生孢子或小分生孢子作为无性生殖结构, 如接骨木镰刀菌和轮枝样镰刀菌(Fusarium verticiloides)等; 有些物种可同时产生大分生孢子和小分生孢子, 如黄色镰刀菌(Fusarium culmorum)和尖孢镰刀菌(Fusarium oxysporiuun)等, 这类镰刀菌通过产生不同类型的孢子, 可以适应不同的生存环境。小分生孢子通常用于在不利条件下的传播, 而大分生孢子则有助于在适宜的环境中扩展和繁殖。两种孢子的共存提高了它们的生存和扩散能力[14]。使用显微镜镜检形态特征是鉴定镰刀菌物种的主要手段, 包括大分生孢子的一般形状和大小、小分生孢子的产生、衣孢子、菌核、有性期和色素沉着等。镰刀属真菌在培养特征上是可变的, 因为它们生长环境的变化会导致培养物和分生孢子的形态变化[11]。
2.1.1 普遍存在的特征
镰刀菌属的分类传统上主要依据其形态学特征, 如分生孢子、分生孢子梗、菌丝结构等。常见的特征包括: (1)分生孢子的形态, 可以是弯曲或直的, 形状通常为镰刀形或细长; (2)分生孢子梗的类型, 分生孢子梗有时是单独的, 有时形成簇生或群体结构; (3)菌丝的形态与结构, 菌丝通常无隔或有隔, 颜色可以是白色、黄色、粉色等。
大分生孢子是鉴定镰刀菌种培养物的唯一最重要的培养特征。大分生孢子最显著的特征是其形状, 其次是间隔的大小和数量, 最后是顶细胞、基细胞或足细胞的性质[13]。就形状而言可分为3种类型: (1)直型大分生孢子, 相对少见, 通常呈直线形, 例如燕麦镰刀菌(Fusarium avenaceum); (2)弯曲型大分生孢子, 这些孢子呈明显的弯曲状, 类似镰刀形。孢子的宽度在整个长度上保持相对一致, 通常呈背侧曲率, 例如木贼镰刀菌(Fusarium equiseti); (3)带状大分生孢子, 呈弯曲状, 形态上类似于弯月形, 通常具有明显的弯曲性, 并且分生孢子较长, 宽度在整个长度上变化较大, 如禾谷镰刀菌(图1)。大分生孢子的长度可以根据不同种类的镰刀菌有所不同, 例如亚美尼亚镰刀菌(Fusarium armeniacum)产生较长的孢子, 而黄色镰刀菌则产生较短的孢子。然而, 孢子大小通常是一个相对恒定的特征, 其变化通常表明培养条件不适宜。一般来说, 大分生孢子呈现3至5隔, 隔数的确定应根据隔距的范围和每个孢子的平均间隔数来判断。
2.1.2 其他重要特征
上述特征在几乎所有镰刀菌种中都普遍存在。但也有些特征仅限于少数几种, 这些特征也是镰刀菌种类鉴定的重要因素。例如, 中分生孢子远形性、圆形孢子的盘绕菌丝以及菌核样结构的形成等, 都是有助于初级鉴定的相关特征。
色素沉着是另一个关键的次要鉴定特征[13]。不同种类的镰刀菌会产生从黄色到橙色、胭脂红等多种颜色。在马铃薯葡萄糖琼脂培养基(potato dextrose agar, PDA)上可以检测到色素沉着的模式, 通常首选12 h的明暗周期的光照条件。色素可能对光照或pH敏感, 在培养基中呈现扩散状或不扩散状, 通常在培养一周后可观察到色素变化。
生长速度也是鉴定的次要特征之一, 通常以PDA上的菌落直径来衡量。菌落生长速度因种类而异, 某些品种如黄皮梢腐病菌(Fusarium lateritium)生长较慢, 而黄色镰刀菌和禾谷镰刀菌等则生长较快[7]。
尽管形态学鉴定非常常用, 但也存在耗时且很容易导致误认等缺陷, 特别是对于系统发育关系较近的物种。为进一步区分物种, 次生代谢物和真菌毒素也被用作鉴定依据。通过分析不同镰刀菌产生的特定次生代谢物和毒素, 可以快速、准确地识别不同种类的镰刀菌及其致病性特点, 对真菌进行初步分组, 并可进一步分析, 最终将真菌划分为特定物种[15-18]。已知镰刀菌会产生许多毒素, 这些毒素可以有效地用于它们的物种鉴定(表1)。
2.2 基于遗传多样性鉴定植物致病性镰刀菌种的分子工具
虽然单独的形态学鉴定可能存在问题, 但在实践中仍然是有帮助的, 并且经常与分子方法结合使用。不同的分子工具, 有助于根据遗传多样性描述物种之间的差异。目前常见的分子工具有: (1)随机扩增多态性DNA (random amplified polymorphic DNA, RAPD)[19]。RAPD分析已被有效地用于细菌和真菌的基因组分析。此外, RAPD已被用于鉴定镰刀菌物种, 如尖孢镰刀菌、燕麦镰刀菌、梨孢镰刀菌、茄病镰刀菌和串珠镰刀菌(Fusarium moniliforme)。尽管RAPD具有反应灵敏度高的优势, 但由于结果的可重复性较差, 并且需要严格的聚合酶链式反应(polymerase chain reaction, PCR)条件, 因此影响了其在真菌分类中的应用; (2)限制性片段长度多态性(restriction fragment length polymorphism, RFLP)[20]。RFLPs已被广泛用于研究菌根和土壤真菌种群/群落的多样性, 该技术能够检测低拷贝编码序列, 且非常稳定, 但其实验操作烦琐, 检测周期长, 成本高昂, 已经逐渐被其他基于PCR的指纹技术所取代。(3)扩增片段长度多态性(amplified fragment length polymorphism, AFLP)[21]。AFLP技术是RAPD的改良版本, 该技术可以一次扩增50~100个片段, 同时检测不同基因组区域的多种多态性。它的灵敏度和可重复性也非常高。目前可用于区分尖孢镰刀菌非致病菌株与普通镰刀菌的分离株, AFLP图谱也被广泛用于尖孢镰刀菌复合体的系统发育分析, 但该技术对DNA的纯度和内切酶的质量要求很高; (4)基因间间隔区(inter genic spacer, IGS)[22]。核糖体DNA重复单位由高度保守的基因和更多的可变间隔区组成, IGS将核糖体DNA重复单位分开。核糖体DNA重复单位的数量在不同物种之间不同, 这导致IGS的长度和限制位点的差异。IGS-RFLP已被用于分析密切相关的物种或群落内部和之间的遗传变异。通过对IGS区域RFLP分析, 该技术已有效地用于与镰刀菌属亲缘关系密切物种的系统发育分析, 如梨孢镰刀菌与九州镰刀菌(Fusarium kyushuense), 这两个物种之间存在明显的分化; (5) β-微管蛋白(β-tubulin)[23]。β-微管蛋白基因序列已被广泛用于各种真菌的系统发育研究。HATSCH等[24]利用β-微管蛋白基因序列和其他标记基因对Fusarium langsethiae、梨孢镰刀菌和拟枝孢镰刀菌进行了分类研究; (6)翻译延伸因子1A (translation elongation factors 1 alpha, TEF-1α)[25]。TEF-1α在镰刀菌的物种水平上具有很高的信息量, 已被广泛用于镰刀菌分类; (7)内部转录间隔物(internal transcribed spacer, ITS)[26]已被用于真菌菌株的鉴别和诊断, ITS区高度保守的引物位点使得它很容易从真菌物种中扩增出来。通常情况下, 植物病原真菌的分子鉴定是通过对ITS区域进行PCR扩增, 然后进行限制性内切分析或直接测序, 并在GenBank或其他数据库中进行生物大分子序列比对搜索工具(basic local alignment search tool, BLAST)搜索来完成的; (8)基质辅助激光解析电离飞行时间质谱(matrix-assisted laser desorption/ionization time of flight, MALDI-TOF)技术[27]。与基于PCR的方法不同, 该技术是一种先进的工具, 可以使用完整的细胞或细胞提取物获得质谱图、与相关的生物质谱数据库进行比对, 快速准确地鉴定真菌和其他微生物。
3 镰刀菌与植物免疫反应的相互作用及其致病机制
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镰刀菌是引起多种植物疾病的主要病原, 能够侵染植物的各个部位, 并诱导细胞死亡。在寄主植物上生长时, 镰刀菌会产生一系列具有植物毒性和真菌毒性的毒素, 这些毒素通过调节植物的生理过程, 促进镰刀菌的定植和繁殖[28]。这些毒素留在受侵染的农产品上并进入食物链, 最终可能引起动物、牲畜和人类真菌中毒。
镰刀菌的基因组可分为核心基因组和适应性基因组, 其中核心基因组负责初级代谢和繁殖, 而适应性基因组则与病原体毒力、寄主特化及其他功能相关。镰刀菌的致病性与其染色体上的一些特定基因密切相关。例如, 镰刀菌通过分泌毒素(如呕吐毒素和三烯醇毒素等)来致病, 这些毒素合成基因常集中在特定染色体区域, 能够抑制植物的免疫反应并导致植物组织的坏死。在某些寄主植物(如番茄)中, 参与毒力和寄主特化的基因位于致病菌株内的致病性染色体上[29]。基因组比较研究表明, 镰刀菌致病性染色体的水平转移能够将非致病菌转变为致病菌。因此, 水平转移可以解释镰刀菌宿主特异性的多系起源[30]。
植物和真菌之间的相互作用是多样而复杂的, 可以改变双方的生理和形态。在病原菌与植物寄主的相互作用中, 毒素一方面可加速病原菌对植物的侵染, 增加其致病力; 另一方面可能会使寄主在受病原菌侵染过程中识别病原菌, 从而激活寄主的防御反应[31-32]。图2为镰刀菌属毒素与植物互作的模式图, 为抵御病原菌, 植物进化出强大的免疫系统, 可分为两部分, 一部分是镰刀菌识别植物细胞后, 分泌细胞壁降解酶, 加速病原菌的侵染, 这些物质可以作为PAMP, 激发植物的PTI, PTI可应对大多数病原菌的入侵。另一部分病原菌已经进化出逃避PTI的方法, 通过向寄主植物细胞注入毒性因子(virulence factors)或效应子(effector molecules)来抑制植物的PTI反应, 部分效应子能够被植物细胞内的核苷酸结合寡聚化结构域(nucleotide binding domain and leucine-rich repeats, NLRs)识别, 从而触发比PTI更强的免疫反应, 即ETI。此外, 病原菌通过分泌致病毒素, 抑制DNA复制, 导致细胞循环受阻; 也可以诱导植物抗氧化酶和防御酶的活性, 激活植物自身的防御系统, 大量的毒素可能会诱导ROS的产生, 从而引发PCD。为应对真菌毒素的危害, 植物通过产生解毒酶来降解毒素、利用转运蛋白将毒素运输至细胞外、或通过分泌次生代谢物抑制毒素的产生, 而这些次生代谢物也可以参与ROS的解毒过程。
目前, 关于病原菌与植物互作中免疫反应的研究大都集中在由模式识别受体(pattern recognition receptors, PRRs)介导的PTI反应以及由病原菌分泌效应蛋白所引起的ETI反应, 然而, 真菌毒素在植物免疫反应中的具体作用仍未被完全解析。因此, 真菌毒素在病原菌侵染过程中诱导的植物免疫反应可能是今后解析植物和病原菌互作机制的研究重点。
4 植物致病性镰刀菌的控制与管理
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镰刀菌是农作物种植过程中一种难以控制的病害, 传统的物理、化学、栽培等防治方法不仅效果不佳, 且成本高昂。镰刀菌广泛分布的一些原因包括: (1)它们可以在多种基质上生长繁殖; (2)它们的孢子传播途径非常高效[29]。镰刀菌是一种土壤传播的植物病原体, 可以在土壤和植物残骸中存活多年。受感染的土壤通过田间使用的工具和动物传播到其他地区, 从而导致新疫情的爆发。这种真菌可以在多种环境条件下生存。新的研究表明, 甲虫可以作为茄病镰刀菌的有效传播媒介[33]。田间真菌接种量、分布情况以及植物生长阶段对农作物减产的程度具有重要影响[34]。大多数干燥和温暖的生长条件有利于镰刀菌的生长。这对于某些地区的作物种植造成了显著影响[35]。
镰刀菌在农业中难以消灭的原因之一是其繁殖体在活体植物的各个器官上持续存在[36]。植物镰刀菌感染的一个主要风险在于, 尽管可能没有明显的外部症状, 但植物的根、茎、叶、花、果等部位均可能已经被病原感染、携带着病原体和真菌毒素[37]。镰刀菌引起的枯萎病通常被认为是单循环病害, 即该病害在同一季节内不会从一株植物传播到另一株[38], 因为能够将病害传播到其他植物引起继发感染的繁殖体只在作物生长季节的后期形成。因此, 控制镰刀菌的扩散和数量的增长是一项非常艰巨的任务。植物在暖湿季节和开花期最容易受到镰刀菌感染和产孢, 但感染的可能性则贯穿整个生长季节[39]。
由于镰刀菌病难以控制, 且真菌容易对各种防治方法产生抗性, 生物防治作为防治手段之一日益受到重视。生物防治不仅符合自然和绿色环保的生态治理趋势, 而且是更加可持续的防控方法。针对镰刀菌的生物防治方法主要包括: 应用有益微生物, 如链霉菌能分泌对镰刀菌具有抑制作用的抗生素、白假丝酵母等真菌通过竞争抑制作用抑制镰刀菌的生长、枯草芽孢杆菌可以通过产生抗菌物质或占据生态位等方式抑制镰刀菌的繁殖; 也有一些植物提取物被发现具有抑制镰刀菌生长的作用。例如, 大蒜、姜等植物提取物中含有天然抗菌物质, SHAH等[40]研究表明生姜素可以有效抑制镰刀菌的侵染。
除了传统的生物防治方法, 一些新型干预技术也为镰刀菌的控制提供了创新的途径。基因编辑技术, 如CRISPR/Cas9, 可以用于修改植物基因, 使其具备更强的抗病性。例如, 编辑植物基因提高其对镰刀菌的抗性, 或是通过转基因技术使植物能够产生对镰刀菌有抑制作用的物质, RADICE等[41]通过CRISPR/Cas9基因编辑技术, 对番茄植物的基因组进行改造, 增强其对镰刀菌的抗性; DEBBARMA等[42]研究了4种富含酚类物质的植物提取物及其混合物在离体和番茄中防治尖孢镰刀菌的效果, 结果表明, 植物提取物的体外抗真菌活性与其总酚含量密切相关, 利用富含多酚的植物粗提取物是一种有效、经济、实用的作物病原菌大规模治理方法。
镰刀菌的生物防治和新型干预技术为应对这一顽固病害提供了多样化解决手段。生物防治利用有益微生物、植物提取物等天然资源, 是一种环保且可持续的方法。与此同时, 基因编辑、纳米技术、微生物代谢产物等新型干预技术正在为未来的镰刀菌防治提供更为创新和高效的途径[43]。随着科技的发展, 这些技术可能为农业生产带来更加智能、绿色的防治方案。
5 植物致病性镰刀菌与全球粮食安全
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镰刀菌是世界大多数地区对农作物最具破坏性的病原真菌, 特别在亚洲、非洲、美洲(美国)、欧洲和大洋洲(澳大利亚), 它对寄主植物造成严重损害, 影响植物发育, 导致作物质量和产量下降。随着全球气候变化的加剧、农业生产方式的转型以及国际贸易的便利化, 镰刀菌的传播和危害更加广泛, 现已成为全球粮食安全的最大威胁[44]。
镰刀菌病原体对世界上许多主要作物构成了严峻的生物威胁。不仅破坏主要提供热量的主粮作物, 如水稻、小麦、玉米和大豆[45-48], 还影响经济作物, 如香蕉、咖啡和大麦和烟草[49-50]。除了直接引起作物病害, 镰刀菌还会在粮食作物中产生多种霉菌毒素, 如脱氧雪腐镰刀菌烯醇、黄曲霉毒素等, 这些毒素能够对人体健康产生重大危害, 尤其是对消化系统、免疫系统和神经系统等造成威胁。脱氧雪腐镰刀菌烯醇被广泛认为是致癌物质, 它不仅影响食品安全, 还可能通过食物对人类健康造成威胁。因此, 镰刀菌的侵染不仅减少了粮食作物的产量, 还可能导致食品的国际贸易受到限制, 从而加剧全球粮食供应的不稳定性。这对一些依赖粮食作物进行国际贸易的国家来说更是严峻挑战[50]。例如, 在小麦生产中, 镰刀菌常常引发小麦镰刀菌病, 该病害不仅降低了小麦的产量, 还严重影响了小麦的质量。小麦中霉菌毒素的超标, 使得这些粮食在市场上无法出售, 甚至可能被销毁。在20世纪90年代, 北美因赤霉病及其毒素造成的经济损失超过30亿美元。在过去10年中, 镰刀菌毒素依然对美国多个州构成严重威胁。以2008年为例, 堪萨斯州的部分农场受到赤霉病的严重影响, 其中小麦中脱氧雪腐镰刀菌醇毒素的浓度高达18000 μg/kg。由于毒素污染过重, 一些田块的作物不得不弃收或焚毁。2008年, 堪萨斯州因此造成的小麦损失高达710万蒲式耳(约42.72万t左右), 价值约5700万美元[51]。2010年和2012年, 中国的小麦主产区, 包括江苏、安徽、湖北、山东、河南、河北、陕西和四川等省, 均出现了赤霉病的大规模爆发, 其中2012年发病面积达927万公顷[52]。
现代农业单一作物种植的模式加速了致命镰刀菌菌株的出现[53], 气候变化也加剧了这一问题, 随着病原体向温暖地区扩展, 疾病的蔓延趋势也随之加剧[54-55]。此外, 植物和植物产品的全球贸易与运输, 助长了植物病原体的扩散, 使其蔓延到原来未受影响的地区[48]。尽管如此, 现代集约化农业仍是全球粮食安全的基石, 能够满足日益增长的人口需求。面对这种变化, 全球面临着真菌病原和新型作物病害的挑战。为了保障未来粮食安全, 加强病原监测、建立预警体系和采用创新的防治策略变得尤为迫切。
6 结束语
收起
镰刀菌是农业、医学和兽医科学领域中最重要的真菌类群之一。在过去的几十年里, 镰刀菌已成为多种植物病害、人类和动物机会性病原及真菌毒素的主要来源。世界上主要粮食作物普遍容易受到镰刀菌等植物病原真菌的侵害, 此外, 镰刀菌引发的真菌病害导致经济作物损失, 严重影响依赖粮食出口的许多发展中国家的经济稳定。由于该真菌对多种防控方法容易产生抗性, 使得有效的病害管理面临挑战。因此, 生物防治作为一种替代方法, 已显示出显著的潜力, 有助于提高全球作物产量。为实现更好的病害控制, 保护未来的收成, 确保持续的全球粮食安全, 亟需加强病害监测、提升预测预报精度, 并开发新的干预措施。
基金
收起
  • 河南省农业科学院自主创新项目(2024ZC044)
  • 河南省农业科学院自主创新项目(2025ZC100)
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2025年第16卷第14期
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doi: 10.19812/j.cnki.jfsq11-5956/ts.20250318001
  • 接收时间：2025-03-18
  • 首发时间：2026-01-07
  • 出版时间：2025-07-25
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  • 收稿日期：2025-03-18
基金
河南省农业科学院自主创新项目(2024ZC044)
河南省农业科学院自主创新项目(2025ZC100)
作者信息
    河南省农业科学院烟草研究所, 烟草行业黄淮烟区烟草病虫害绿色防控重点实验室, 许昌 461000

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*平文丽(1982—), 女, 博士, 助理研究员, 主要研究方向为烟草分子抗病育种。E-mail:
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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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