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Article(id=1256518443751301392, tenantId=1146029695717560320, journalId=1255847803461844995, issueId=1256518442379763982, articleNumber=null, orderNo=null, doi=10.13346/j.mycosystema.250178, pmid=null, cstr=32115.14.j.mycosystema.250178, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1749484800000, receivedDateStr=2025-06-10, revisedDate=null, revisedDateStr=null, acceptedDate=1754496000000, acceptedDateStr=2025-08-07, onlineDate=1777506941974, onlineDateStr=2026-04-30, pubDate=1774108800000, pubDateStr=2026-03-22, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1777506941974, onlineIssueDateStr=2026-04-30, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1777506941974, creator=13701087609, updateTime=1777506941974, updator=13701087609, issue=Issue{id=1256518442379763982, tenantId=1146029695717560320, journalId=1255847803461844995, year='2026', volume='45', issue='3', pageStart='240320', pageEnd='250282', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1777506941647, creator=13701087609, updateTime=1777507117568, updator=13701087609, preIssue=null, nextIssue=null, ext={EN=IssueExt(id=1256519180338213460, tenantId=1146029695717560320, journalId=1255847803461844995, issueId=1256518442379763982, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1256519180338213461, tenantId=1146029695717560320, journalId=1255847803461844995, issueId=1256518442379763982, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null}, startPage=250178, endPage=, ext={EN=ArticleExt(id=1256518447039635744, articleId=1256518443751301392, tenantId=1146029695717560320, journalId=1255847803461844995, language=EN, title=Functions of autophagy protein CfAtg7 in Colletotrichum fructicola, columnId=1256263562373226548, journalTitle=Mycosystema, columnName=Research paper, runingTitle=null, highlight=null, articleAbstract=

Anthracnose is prevalent in major oil-tea production regions across the country, severely compromising yield and quality of oil-tea Camellia oleifera. Colletotrichum fructicola is the dominant pathogenic fungus responsible for this disease, and understanding its pathogenic mechanisms is fundamental for controlling Ca. oleifera anthracnose. In eukaryotes, autophagy is a conserved intracellular degradation pathway and serves as the primary route for degrading macromolecular proteins and damaged organelles. Atg7, a core autophagy protein, plays crucial roles in many plant-pathogenic fungi. However, its biological functions in Co. fructicola remains unclear. In this study, the CfATG7 gene knockout mutant ΔCfatg7 and its complementary strain ΔCfatg7/CfATG7 were constructed, and their phenotype and pathogenicity were assayed. The results demonstrate that CfAtg7 is involved in regulating autophagy, vegetative growth, conidiation, and appressorium formation. It also participates in the responses to endoplasmic reticulum stress, cell wall integrity stress, and oxidative stress, revealing the pleiotropic roles of the autophagy protein CfAtg7 in Co. fructicola. This study elucidated the biological functions of CfAtg7 in Co. fructicola, providing experimental evidence for the development of novel fungicides targeting this protein.

, correspAuthors=Shengpei ZHANG, authorNote=null, correspAuthorsNote=
*
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油茶炭疽病广泛发生于全国各大油茶产区，严重危害油茶产量和品质，果生刺盘孢Colletotrichum fructicola是其优势致病菌，解析该病菌的致病机制是防治油茶炭疽病的基础。真核生物的细胞自噬是一种保守的胞内降解途径，是降解大分子蛋白与各受损细胞器的主要途径。Atg7作为自噬核心蛋白，在很多植物病原真菌中发挥着重要功能。然而，目前暂不清楚Atg7在果生刺盘孢中发挥的生物学功能。本实验构建了CfATG7基因敲除突变体ΔCfatg7及其互补菌株ΔCfatg7/CfATG7，并对其进行了生物学表型及致病力测定。研究结果表明CfAtg7参与调控油茶果生刺盘孢的细胞自噬、营养生长、分生孢子和附着胞形成，也参与对内质网压力胁迫、细胞壁完整性胁迫以及氧化胁迫的应答，揭示了自噬相关蛋白CfAtg7在果生刺盘孢中所发挥的多效作用。本研究阐明了CfAtg7在果生刺盘孢中的生物学功能，为以该蛋白为靶标的新型杀菌剂的开发提供了实验依据。

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pmcid=null, year=2016, volume=32, issue=10, pageStart=84, pageEnd=96, url=null, language=null, rfNumber=[39], rfOrder=38, authorNames=杨正婷, 刘建祥, journalName=生物技术通报, refType=null, unstructuredReference=杨正婷, 刘建祥, 2016. 植物内质网胁迫应答研究进展. 生物技术通报, 32(10): 84-96, articleTitle=植物内质网胁迫应答研究进展, refAbstract=null), Reference(id=1256518507420836753, tenantId=1146029695717560320, journalId=1255847803461844995, articleId=1256518443751301392, doi=null, pmid=null, pmcid=null, year=2007, volume=27, issue=4, pageStart=358, pageEnd=362, url=null, language=null, rfNumber=[40], rfOrder=39, authorNames=叶青, 郑民华, journalName=国际病理科学与临床杂志, refType=null, unstructuredReference=叶青, 郑民华, 2007. 自噬的分子机制与病理生理意义. 国际病理科学与临床杂志, 27(4): 358-362, articleTitle=自噬的分子机制与病理生理意义, refAbstract=null)], funds=[Fund(id=1256518490492625598, tenantId=1146029695717560320, journalId=1255847803461844995, articleId=1256518443751301392, awardId=2023JJ20100, language=EN, fundingSource=Natural Science Foundation of Hunan Province(2023JJ20100), 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journalId=1255847803461844995, articleId=1256518443751301392, companyId=1256518456309047688, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=中南林业科技大学 森林生物资源与有害生物综合管理湖南省普通高等学校重点实验室 木本油料资源利用全国重点实验室，湖南 长沙 410004)])], figs=[ArticleFig(id=1256518477653860945, tenantId=1146029695717560320, journalId=1255847803461844995, articleId=1256518443751301392, language=EN, label=Fig. 1, caption=Domain prediction and phylogenetic analysis of CfAtg7, and generation of knockout mutants and complemented strains. A: Phylogenetic tree of CfAtg7 with an amino acid substitution rate of 0.10; the data indicate bootstrap value. Species and their corresponding GenBank accession numbers are as follows: Co. gloeosporioides (EQB55638.1, CgAtg7), Fusarium graminearum (XP_011319173.1, FgAtg7), Magnaporthe oryzae (XP_003715525.1, MoAtg7), Aspergillus oryzae (KDE85504.1, AoAtg7), A. nidulans (XP_050467950.1, AnAtg7), Ustilago maydis (XP_011391717.1, UmAtg7), Saccharomyces cerevisiae (NP_012041.1, ScAtg7). B: Domain prediction of CfAtg7; the gray regions indicate the ATG7 and ThiF domains, respectively. C: Schematic diagram of CfATG7 gene knockout strategy. D: Verification of Knockout mutant and complemented strain; M, DNA marker DL2000; 1, ΔCfatg7; 2, ΔCfatg7/CfATG7; Wild-type is positive control; H2O is negative control; Primer pair 1, ATG7BWF/ H855R; Primer pair 2, ATG7NBF/ATG7NBR., figureFileSmall=Xo0Qmne0jb1N1RqtqK3GrQ==, figureFileBig=8Uq6XDIBEWCAR2SbSIX0ig==, tableContent=null), ArticleFig(id=1256518478006182485, tenantId=1146029695717560320, journalId=1255847803461844995, articleId=1256518443751301392, language=CN, label=图1, caption=CfAtg7结构域预测、系统发育分析和敲除突变体及回补菌株的获得 A：CfAtg7的系统发育进化树，氨基酸替代率为0.10，数字代表支持率；物种及其对应的GenBank登录号如下：Co. gloeosporioides (EQB55638.1)、Fusarium graminearum (XP_011319173.1)、Magnaporthe oryzae (XP_003715525.1)、Aspergillus oryzae (KDE85504.1)、A. nidulans (XP_050467950.1)、Ustilago maydis (XP_011391717.1)、Saccharomyces cerevisiae (NP_012041.1)；B：CfAtg7的结构域预测, 灰色区域分别代表ATG7和ThiF结构域；C：CfATG7基因敲除策略图；D：突变菌株和回补菌株的电泳验证；M，DNA marker DL2000；1，ΔCfatg7；2，ΔCfatg7/CfATG7；WT作阳性对照；H2O作阴性对照；引物对1，ATG7BWF/H855R；引物对2，ATG7NBF/ATG7NBR, figureFileSmall=Xo0Qmne0jb1N1RqtqK3GrQ==, figureFileBig=8Uq6XDIBEWCAR2SbSIX0ig==, tableContent=null), ArticleFig(id=1256518480547930724, tenantId=1146029695717560320, journalId=1255847803461844995, articleId=1256518443751301392, language=EN, label=Fig. 2, caption=Mycelial growth and conidiation level in ΔCfatg7 strain. A: Colony of three strains on two culture media; B: Data analysis of colony diameter and conidial production. Error bars represent standard deviation (SD). **P<0.01; ***P<0.001; ****P<0.000 1. The same below., figureFileSmall=8ot8vmlMxuBYEL9edTfFqg==, figureFileBig=2luEJAeHKvKSHk/zlP17rw==, tableContent=null), ArticleFig(id=1256518480912835180, tenantId=1146029695717560320, journalId=1255847803461844995, articleId=1256518443751301392, language=CN, label=图2, caption=ΔCfatg7菌丝生长和产孢量 A：菌株在两种培养基上的菌落；B：菌落直径和分生孢子产量的数据分析；误差线表示标准差，**P<0.01；***P<0.001；****P<0.000 1. 下同, figureFileSmall=8ot8vmlMxuBYEL9edTfFqg==, figureFileBig=2luEJAeHKvKSHk/zlP17rw==, tableContent=null), ArticleFig(id=1256518482099823220, tenantId=1146029695717560320, journalId=1255847803461844995, articleId=1256518443751301392, language=EN, label=Fig. 3, caption=Pathogenicity of the three strains on wounded/unwounded Camellia oleifera leaves and apples. A: Pathogenic symptoms on Ca. oleifera leaves; B: Statistical analysis (One-way ANOVA) of pathogenicity on Ca. oleifera leave; C: Pathogenic symptoms on apples caused by the three strains at 5 days post-inoculation (dpi); D: Statistical analysis (One-way ANOVA) of pathogenicity on apples., figureFileSmall=XssXvRKS45fzSggnMhaq+g==, figureFileBig=Vp9YSzs+x5lOvqCnDo7V8w==, tableContent=null), ArticleFig(id=1256518482619916921, tenantId=1146029695717560320, journalId=1255847803461844995, articleId=1256518443751301392, language=CN, label=图3, caption=菌株在无伤和有伤油茶叶片以及苹果上的致病情况 A：菌株接种2 d后，在油茶叶片上的致病情况；B：菌株在油茶叶片上致病情况的单因素方差分析；C：菌株接种5 d后，在苹果上的致病情况；D：菌株在苹果上致病情况的单因素方差分析, figureFileSmall=XssXvRKS45fzSggnMhaq+g==, figureFileBig=Vp9YSzs+x5lOvqCnDo7V8w==, tableContent=null), ArticleFig(id=1256518483395863164, tenantId=1146029695717560320, journalId=1255847803461844995, articleId=1256518443751301392, language=EN, label=Fig. 4, caption=Appressorial formation. *Appressoria; Bars=10 μm., figureFileSmall=UqLeRgZw8Kxs1PoYV/43qg==, figureFileBig=NJD+hDTrRnKoRZ0IlOTpkQ==, tableContent=null), ArticleFig(id=1256518485023253129, tenantId=1146029695717560320, journalId=1255847803461844995, articleId=1256518443751301392, language=CN, label=图4, caption=附着胞形成 *附着胞，标尺=10 μm, figureFileSmall=UqLeRgZw8Kxs1PoYV/43qg==, figureFileBig=NJD+hDTrRnKoRZ0IlOTpkQ==, tableContent=null), ArticleFig(id=1256518485488820880, tenantId=1146029695717560320, journalId=1255847803461844995, articleId=1256518443751301392, language=EN, label=Fig. 5, caption=CfAtg7 is essential for autophagy. A: Colony morphology of WT, ΔCfatg7, and ΔCfatg7/CfATG7 on CM containing either DMSO or rapamycin. rapa. indicates rapamycin. B: Statistical analysis (one-way ANOVA) of rapamycin inhibition rates for each strain. C: Autophagy levels analyzed by fluorescence microscopy. Arrows indicate vacuoles. Scale bars=5 μm. D: Quantification of vacuoles containing GFP signals. E: Autophagy levels analyzed by Western blot. The numbers underneath the blot indicate GFP/(GFP+GFP-CfAtg8); • Target protein., figureFileSmall=t0LFHunI8nsPP4bGquVaYw==, figureFileBig=W13Q30w2DPJ8zzzI+uHO4A==, tableContent=null), ArticleFig(id=1256518485958582937, tenantId=1146029695717560320, journalId=1255847803461844995, articleId=1256518443751301392, language=CN, label=图5, caption=CfAtg7是自噬所必需的 A：WT、ΔCfatg7及回补菌株在含DMSO或雷帕霉素的CM培养基上的菌落形态, rapa.代表雷帕霉素；B：雷帕霉素对各菌株抑制率的单因素方差分析；C：通过荧光显微镜检测自噬水平，箭头指示液泡位置，比例尺=5 μm；D：含GFP信号液泡的占比统计；E：Western blot检测自噬水平，泳道下方的数值为GFP/(GFP+GFP-CfAtg8)，•标记目的蛋白, figureFileSmall=t0LFHunI8nsPP4bGquVaYw==, figureFileBig=W13Q30w2DPJ8zzzI+uHO4A==, tableContent=null), ArticleFig(id=1256518486717751970, tenantId=1146029695717560320, journalId=1255847803461844995, articleId=1256518443751301392, language=EN, label=Fig. 6, caption=Response of ΔCfatg7 to environmental stresses (cell wall stressors: CR, SDS; ER stress: DTT; oxidative stress: H2O2). A: Mycelial growth of the three strains on CM medium supplemented with different stress agents. B: Statistical analysis of growth inhibition rates in ΔCfatg7., figureFileSmall=RuddFzYLyUjJE3tGBSF/RA==, figureFileBig=XrUPJz36VfJkh6EeGOoqag==, tableContent=null), ArticleFig(id=1256518487158153896, tenantId=1146029695717560320, journalId=1255847803461844995, articleId=1256518443751301392, language=CN, label=图6, caption=ΔCfatg7对环境胁迫(细胞壁CR、SDS；内质网DTT；氧化压力H2O2)的应答 A：3个菌株分别接种于CM和添加各种胁迫剂的培养基上菌丝生长情况；B：ΔCfatg7生长抑制率的数据分析, figureFileSmall=RuddFzYLyUjJE3tGBSF/RA==, figureFileBig=XrUPJz36VfJkh6EeGOoqag==, tableContent=null), ArticleFig(id=1256518487686636207, tenantId=1146029695717560320, journalId=1255847803461844995, articleId=1256518443751301392, language=EN, label=Table 1, caption=

Primers used in this study

, figureFileSmall=null, figureFileBig=null, tableContent=
引物
Primer		 引物序列
Sequence (5ʹ→3ʹ)
ATG7KOUF TGGTGCTCGACAAGGTACTGC
ATG7KOUR TTGACCTCCACTAGCTCCAGCCAAGCCCTTGGGCTTTTCGAAAGGTT
ATG7KODF CAAAGGAATAGAGTAGATGCCGACCGGCTCAGGTTTAGAGTCAAAG
ATG7KODR AGACGGCGGACAAAGGTGC
Hyg-F GGCTTGGCTCCAGCTAGTGGAGGT
Hyg-R CTCTATTCCTTTGCCCTCG
ATG7BWF CGGGCGAGAAGCCAAAAAT
H855R GCTGATCTGACCAGTTGC
ATG7NBF CTCGCTTCTGTCCTCGTTTG
ATG7NBR CAAGTCCAGGTTGGGTGAAA
ATG7NGFP1F ACTCACTATAGGGCGAATTGGGTACTCAAATTGGTTCCATTGACGGTATTGATGGTA
ATG7NGFP1R CCTCGCCCTTGCTCACCATCTTGGGCTTTTCGAAAGGTT
ATG7NGFP3F GCATGGACGAGCTGTACAAGATGATGGCTGCAATCCAATACGC
ATG7NGFP3R CACCACCCCGGTGAACAGCTCCTCGCCCTTGCTCACTTAAAGCAGCTCTCCGTCTCC
NGFPF ATGGTGAGCAAGGGCGAGG
NGFPR CTTGTACAGCTCGTCCATGC
), ArticleFig(id=1256518489364357809, tenantId=1146029695717560320, journalId=1255847803461844995, articleId=1256518443751301392, language=CN, label=表1, caption=

引物名称及序列

, figureFileSmall=null, figureFileBig=null, tableContent=
引物
Primer		 引物序列
Sequence (5ʹ→3ʹ)
ATG7KOUF TGGTGCTCGACAAGGTACTGC
ATG7KOUR TTGACCTCCACTAGCTCCAGCCAAGCCCTTGGGCTTTTCGAAAGGTT
ATG7KODF CAAAGGAATAGAGTAGATGCCGACCGGCTCAGGTTTAGAGTCAAAG
ATG7KODR AGACGGCGGACAAAGGTGC
Hyg-F GGCTTGGCTCCAGCTAGTGGAGGT
Hyg-R CTCTATTCCTTTGCCCTCG
ATG7BWF CGGGCGAGAAGCCAAAAAT
H855R GCTGATCTGACCAGTTGC
ATG7NBF CTCGCTTCTGTCCTCGTTTG
ATG7NBR CAAGTCCAGGTTGGGTGAAA
ATG7NGFP1F ACTCACTATAGGGCGAATTGGGTACTCAAATTGGTTCCATTGACGGTATTGATGGTA
ATG7NGFP1R CCTCGCCCTTGCTCACCATCTTGGGCTTTTCGAAAGGTT
ATG7NGFP3F GCATGGACGAGCTGTACAAGATGATGGCTGCAATCCAATACGC
ATG7NGFP3R CACCACCCCGGTGAACAGCTCCTCGCCCTTGCTCACTTAAAGCAGCTCTCCGTCTCC
NGFPF ATGGTGAGCAAGGGCGAGG
NGFPR CTTGTACAGCTCGTCCATGC
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自噬蛋白CfAtg7在果生刺盘孢中的功能
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陈妍 , 王伊玲 , 罗晶 , 张盛培 *
菌物学报 | 研究论文 2026,45(3): 250178
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菌物学报 | 研究论文 2026, 45(3): 250178
自噬蛋白CfAtg7在果生刺盘孢中的功能
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陈妍, 王伊玲, 罗晶, 张盛培*
作者信息
  • 中南林业科技大学 森林生物资源与有害生物综合管理湖南省普通高等学校重点实验室 木本油料资源利用全国重点实验室，湖南 长沙 410004
Functions of autophagy protein CfAtg7 in Colletotrichum fructicola
Yan CHEN, Yiling WANG, Jing LUO, Shengpei ZHANG*
Affiliations
  • Key Laboratory of Forest Bio-Resources and Integrated Pest Management for Higher Education in Hunan Province, State Key Laboratory of Utilization of Woody Oil Resource, Central South University of Forestry and Technology, Changsha 410004, Hunan, China
出版时间: 2026-03-22 doi: 10.13346/j.mycosystema.250178
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摘要
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油茶炭疽病广泛发生于全国各大油茶产区，严重危害油茶产量和品质，果生刺盘孢Colletotrichum fructicola是其优势致病菌，解析该病菌的致病机制是防治油茶炭疽病的基础。真核生物的细胞自噬是一种保守的胞内降解途径，是降解大分子蛋白与各受损细胞器的主要途径。Atg7作为自噬核心蛋白，在很多植物病原真菌中发挥着重要功能。然而，目前暂不清楚Atg7在果生刺盘孢中发挥的生物学功能。本实验构建了CfATG7基因敲除突变体ΔCfatg7及其互补菌株ΔCfatg7/CfATG7，并对其进行了生物学表型及致病力测定。研究结果表明CfAtg7参与调控油茶果生刺盘孢的细胞自噬、营养生长、分生孢子和附着胞形成，也参与对内质网压力胁迫、细胞壁完整性胁迫以及氧化胁迫的应答，揭示了自噬相关蛋白CfAtg7在果生刺盘孢中所发挥的多效作用。本研究阐明了CfAtg7在果生刺盘孢中的生物学功能，为以该蛋白为靶标的新型杀菌剂的开发提供了实验依据。

油茶  /  果生刺盘孢  /  自噬相关蛋白  /  CfAtg7

Anthracnose is prevalent in major oil-tea production regions across the country, severely compromising yield and quality of oil-tea Camellia oleifera. Colletotrichum fructicola is the dominant pathogenic fungus responsible for this disease, and understanding its pathogenic mechanisms is fundamental for controlling Ca. oleifera anthracnose. In eukaryotes, autophagy is a conserved intracellular degradation pathway and serves as the primary route for degrading macromolecular proteins and damaged organelles. Atg7, a core autophagy protein, plays crucial roles in many plant-pathogenic fungi. However, its biological functions in Co. fructicola remains unclear. In this study, the CfATG7 gene knockout mutant ΔCfatg7 and its complementary strain ΔCfatg7/CfATG7 were constructed, and their phenotype and pathogenicity were assayed. The results demonstrate that CfAtg7 is involved in regulating autophagy, vegetative growth, conidiation, and appressorium formation. It also participates in the responses to endoplasmic reticulum stress, cell wall integrity stress, and oxidative stress, revealing the pleiotropic roles of the autophagy protein CfAtg7 in Co. fructicola. This study elucidated the biological functions of CfAtg7 in Co. fructicola, providing experimental evidence for the development of novel fungicides targeting this protein.

Camellia oleifera  /  Colletotrichum fructicola  /  autophagy-related protein  /  CfAtg7
陈妍, 王伊玲, 罗晶, 张盛培. 自噬蛋白CfAtg7在果生刺盘孢中的功能. 菌物学报, 2026 , 45 (3) : 250178 - . DOI: 10.13346/j.mycosystema.250178
Yan CHEN, Yiling WANG, Jing LUO, Shengpei ZHANG. Functions of autophagy protein CfAtg7 in Colletotrichum fructicola[J]. Mycosystema, 2026 , 45 (3) : 250178 - . DOI: 10.13346/j.mycosystema.250178
正文
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油茶Camellia oleifera是我国优质的经济木本食用油料树种，其果实油茶籽经压榨可制成营养价值高且耐贮藏的茶油，具有巨大的生产潜力(谭晓风 2023)。油茶产业的不断扩大为我国粮食安全、乡村振兴和国际竞争提供保障，是国家政策重点扶持的产业之一(邓海艳和陶丽萍 2023)。
油茶炭疽病是油茶最主要的病害之一，导致油茶严重减产，造成巨大经济损失。该病害的主要病原菌有果生刺盘孢Colletotrichum fructicola Prihast., L. Cai & K.D. Hyde、盘长孢状刺盘孢Colletotrichum gloeosporioides (Penz.) Penz. & Sacc.、暹罗刺盘孢Colletotrichum siamense Prihast., L. Cai & K.D. Hyde、山茶刺盘孢Colletotrichum camelliae Massee和卡斯特刺盘孢Colletotrichum karsti You L. Yang, Zuo Y. Liu, K.D. Hyde & L. Cai等，其中果生刺盘孢是其优势致病菌(李河等2016)。课题组前期对果生刺盘孢的致病机制开展了持续研究，但其致病分子机制仍不十分清楚。阐明该病菌致病的分子机理，可为研发绿色新型杀菌剂提供更多新的药物靶标。
果生刺盘孢与寄主接触初期处于营养匮乏状态，丝状病原真菌在营养匮乏阶段可以通过细胞自噬(autophagy)回收利用被降解的营养物质以维持能量来源，平衡细胞稳态，从而使自身新陈代谢处于正常水平(叶青和郑民华 2007；Mizushima & Komatsu 2011)。细胞自噬是真核生物的一种高度保守的胞内降解途径，具有维持胞内生理平衡及抵抗逆境的重要作用(Naqvi et al. 2012)。细胞自噬过程复杂，其中自噬小体的形成尤为重要。Atg8-磷脂酰乙醇胺(phosphatidylethanolamine, PE)功能组和Atg5-Atg12复合体功能组是自噬小体形成的2个主要偶联系统(Kuma et al. 2007；Liu et al. 2010)。自噬关键蛋白Atg7属于一种E1类泛素激活酶，负责类泛素化分子的激活与转移，可同时促进上述2个偶联系统的形成，从而启动细胞自噬(Collier et al. 2021；Yao et al. 2023)。
作为自噬关键蛋白，Atg7在许多常见植物病原真菌中发挥重要作用。例如，Atg7调控灰葡萄孢和禾谷镰孢菌的生长，Atg7介导的细胞自噬调控稻巨座壳Magnaporthe oryzae的致病过程(刘小红等 2008；Ren et al. 2018；吕务云 2019)。近年来，本课题组解析了部分自噬相关蛋白调控果生刺盘孢生长、产孢及致病力的机制(郭源等2021；Chen et al. 2023；Guo & Zhang 2024)。但在果生刺盘孢中尚未报道自噬核心蛋白Atg7的研究。本试验拟研究Atg7的生物学功能，为以该基因及其调控网络为靶标开发新型杀菌剂提供依据。
1 材料与方法
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1.1 供试菌株
野生型菌株(WT) CFLH16分离自感病的油茶叶片，保存在中南林业科技大学经济林培育与保护教育部重点实验室。酵母转化所需的酵母感受态细胞XK125由本实验室保存。大肠杆菌转化所需的感受态大肠杆菌Trelief™5α菌株采购自擎科生物公司。
1.2 CfAtg7系统发育分析
依照酿酒酵母Saccharomyces cerevisiae ScAtg7的氨基酸序列在果生刺盘孢的基因库中鉴定到其同源蛋白CfAtg7。在SMART网站上传CfAtg7的氨基酸序列预测其结构域。同时，使用Mega7.0软件中的邻接法(Neighbor-Joining Algorithm)构建CfAtg7的系统发育树。
1.3 CfATG7基因敲除突变体的获得
以野生型DNA为模板，利用引物ATG7UF/ ATG7UR和ATG7DF/ATG7DR扩增目的基因前后各1 000 bp。将其与利用引物Hyg-F/Hyg-R扩增的HPH (hygromycin B，潮霉素B)抗性基因连接成融合片段，根据PEG介导的原生质体转化法将融合片段转入WT的原生质体中，原生质体制备参照Zhang et al. (2022)的方法。利用基因内引物ATG7NBF/ATG7NBR和臂外引物ATG7BWF/ H855R对转化子DNA进行PCR扩增，通过琼脂糖电泳验证CfATG7基因敲除突变体ΔCfatg7。本文涉及引物见表1。
1.4 ΔCfatg7/CfATG7回补菌株的获得
回补载体构建方法参照高亚兰等(2020)所述方法。用引物ATG7NGFP1F/ATG7NGFP1和ATG7NGFP3F/ATG7NGFP3R以WT的DNA为模板分别扩增得到ATG7NGFP1和ATG7NGFP3片段。将其与引物NGFPF/NGFPR扩增得到GFP片段融合后与pYF11线性化质粒共转入酵母XK125感受态细胞中，在SD-Trp培养基上进行筛选得到ATG7NGFP⁚⁚CfATG7，并用引物ATG7NBF/ATG7NGFP3R进行PCR鉴定阳性克隆。提取其质粒转入Trelief™5α中，在LB培养基筛选。鉴定阳性克隆后测序。随后，将测序无误的质粒转入ΔCfatg7原生质体中，在添加Bleomycin的培养基上培养2-3 d，初步筛选出候选ΔCfatg7/CfATG7回补菌株，挑选菌丝在荧光显微镜下观测其荧光强度。
1.5 生物学表型测定
1.5.1 菌落生长和产孢量测定
切取WT、ΔCfatg7和ΔCfatg7/CfATG7的菌块分别接种于CM和MM培养基表面，倒置于28 ℃培养箱中黑暗培养3 d后用十字法测量生长直径并对所得数据进行统计分析，试验重复3次。将以上相同大小的菌块放入50 mL液体CM中，并于28 ℃、180 r/min摇床上培养。滴加10 μL培养2-3 d后的菌液于血球计数板上，分别统计分生孢子数，试验重复3次，每个样本至少计数3个视野。
1.5.2 突变体致病力测定
将以上菌块分别接种于无伤和有伤的离体油茶叶片边缘上以及同一苹果上，在28 ℃培养箱中密封保湿且黑暗培养一段时间后观察发病情况，试验重复3次。拍照记录并用ImageJ软件测量病斑大小，制作出柱形统计图。
1.5.3 附着胞形成率测定
富集培养3 d后的分生孢子并调整其浓度至1×105个/mL，在疏水玻片上于28 ℃培养箱保湿放置12 h，利用光学显微镜观察附着胞的形成情况，并拍摄附着胞照片。
1.5.4 自噬水平的测定
将自噬标记蛋白GFP-CfAtg8分别转入野生型和突变体原生质体中标记自噬小体。菌株在液体CM中180 r/min振荡培养36 h，用ddH₂O洗涤后转移至MM-N培养基中28 ℃、60-80 r/min诱导2-5 h。GFP-CfAtg8是观察细胞自噬的标记蛋白，当启动自噬时，GFP-CfAtg8会从细胞质中逐渐转移至液泡内。因此，可通过荧光显微镜观察液泡中的绿色荧光，评价自噬水平。同时，进入液泡中的GFP-CfAtg8会在蛋白酶的作用下发生降解，由于降解产生的GFP蛋白相对耐蛋白酶水解，因此，也可通过免疫印迹检测GFP的相对含量从而评价自噬水平。
1.5.5 突变体应对外界胁迫敏感性的测定
从WT、ΔCfatg7及ΔCfatg7/CfATG7菌落边缘分别接种至含25 μmol/L雷帕霉素(rapamycin)、10 mmol/L H2O2氧化胁迫剂、0.1%十二烷基磺酸钠(sodium dodecyl sulfate, SDS)和400 μg/mL刚果红(congo red, CR)细胞壁完整性胁迫剂、7.5 mmol/L二硫苏糖醇(DL-dithiothreitol, DTT)内质网胁迫剂的CM培养基表面，于28 ℃倒置黑暗培养3 d。测量菌落直径并统计分析。
1.6 蛋白质提取和免疫印迹分析
使用含蛋白酶抑制剂(0.1 g/mL)的1 mL RIPA裂解液裂解细胞，涡旋振荡混匀后，将裂解液置于4 ℃孵育10 min，重复3次，随后12 000 r/min离心10 min去除细胞碎片。收集上清裂解液作为总蛋白样本，经SDS-PAGE分离后，采用特异性抗体进行Western blot检测。
1.7 数据分析
数据以均值±标准差表示，采用GraphPad9.0软件对试验数据进行单因素方差分析(one-way ANOVA)。显著性水平设定为**P<0.01，***P<0.001，****P<0.000 1，差异极显著。所有实验均包含3次生物学重复，且每次生物学重复至少含3次技术重复。
2 结果与分析
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2.1 CfAtg7系统发育树及结构域分析
构建CfAtg7的系统发育树和结构域图，发现CfAtg7由705个氨基酸组成，其氨基酸序列和盘长孢状刺盘孢Co. gloeosporioides的序列一致性为98.72%，同源性较高，但与酿酒酵母的一致性相对较低(图1A)。SMART蛋白质结构域预测显示，CfAtg7包括ATG7_N结构域、ThiF结构域和一个低复杂度结构域(图1B)。这说明CfAtg7在丝状真菌中高度保守，并有保守的ATG7结构域与泛素化结构域。
2.2 ΔCfatg7及ΔCfatg7/CfATG7的获得
为了研究CfAtg7是否影响果生刺盘孢生物学功能的正常发挥，利用反向遗传学方法和同源重组原理，参照1.3的方法，将潮霉素B抗性基因HPH替换CfATG7编码区，获得突变体ΔCfatg7，基因敲除策略见图1C。参照1.4的方法将具有博莱霉素抗性且包含基因自身启动子区域的目的片段回补到ΔCfatg7中，通过绿色荧光筛选获得回补菌株ΔCfatg7/CfATG7，并对所得的基因敲除突变体和回补菌株进行电泳验证(图1)。
2.3 CfAtg7参与调控果生刺盘孢生长和分生孢子形成过程
为了研究CfAtg7是否参与调控果生刺盘孢菌丝的营养生长过程，将菌块接种于CM和MM培养基。在CM培养基上，ΔCfatg7菌落直径仅为(4.68±0.07) cm，显著小于野生型的菌落直径(5.50±0.05) cm。在MM培养基上，WT的菌落直径(4.70±0.09) cm，ΔCfatg7的菌落直径则只有(3.92±0.06) cm，两者具有极显著差异(图2)。上述结果表明CfAtg7参与调控果生刺盘孢菌丝的营养生长过程。
进一步研究CfAtg7是否参与调控果生刺盘孢分生孢子的形成。培养2 d，ΔCfatg7的产孢量为(16±2)×104个/mL，显著小于野生型(78±5)×104个/mL的产孢量(图2B)。培养3 d，ΔCfatg7的产孢量为(717±15)×104个/mL，也显著小于野生型(953±87)× 104个/mL的产孢量。这表明CfAtg7参与调控果生刺盘孢形成无性分生孢子。
2.4 CfAtg7参与调控果生刺盘孢的致病力
将上述菌株分别接种到表面无明显伤口的油茶叶片上和利用针孔处理后有伤口的油茶叶片上，置于28 ℃培养箱中黑暗保湿培养2-3 d，拍摄同一画面中存在叶片和标尺的图片，利用软件ImageJ来测量病斑面积，并用GraphPad9.0软件对试验数据进行单因素方差分析(one-way ANOVA)。在无伤叶片上，野生型和回补菌株形成的病斑面积为(44.3±10.5) mm2和(44.9±7.6) mm2，ΔCfatg7则只形成(1.5±1.2) mm2的病斑，显著减小。在有伤叶片上ΔCfatg7形成的病斑面积为(10.10±6.58) mm2，而WT和ΔCfatg7/CfATG7则产生较ΔCfatg7显著增大的炭疽病病斑(图3A, 3B)。同时，我们将上述菌株接种于已消毒且无明显病斑的苹果上。结果表明，ΔCfatg7在苹果上形成的病斑直径为(1.67±0.58) cm，显著小于WT和ΔCfatg7/CfATG7的病斑直径(图3C, 3D)。上述实验结果说明CfAtg7蛋白参与调控果生刺盘孢的致病力。
2.5 CfAtg7参与调控果生刺盘孢附着胞的形成
附着胞是植物病原菌一种高度特化的感染细胞，对寄主侵染起关键作用(林福呈 2001)。在疏水玻片上滴加分生孢子悬浮液进行保湿培养，在显微镜下观察附着胞形成情况(图4)。28 ℃黑暗培养12 h后，WT孢子萌发率为(59±2)%，ΔCfatg7的孢子萌发率为(61±4)%，两者无显著差异。但ΔCfatg7的附着胞的形成率仅为(2±2)%，极显著低于WT的(37±6)%。这表明CfAtg7蛋白参与调控果生刺盘孢附着胞的形成。
2.6 CfAtg7调控果生刺盘孢的自噬过程
在真核生物中，雷帕霉素靶蛋白(target of rapamycin, TOR)信号通路调节细胞的生长代谢。TOR通路和细胞自噬之间的功能关系复杂，雷帕霉素通常作为一种自噬诱导剂使用(陈淳媛 2007)。CfAtg7作为自噬蛋白，我们首先关注其对雷帕霉素的响应。结果显示该胁迫剂对ΔCfatg7突变体的抑制率显著降低(图5A, 5B)。这说明CfAtg7参与调控果生刺盘孢对自噬诱导剂雷帕霉素的响应。
随后，我们在WT和ΔCfatg7突变体中转入自噬标记蛋白GFP-CfAtg8。在营养匮乏条件MM-N诱导自噬时，GFP-Atg8会转运至液泡降解以实现营养循环(Liu et al. 2015)。未诱导时(0 h)，WT菌株液泡中几乎无绿色荧光；诱导2 h后，ΔCfatg7菌株液泡完全无绿色荧光，WT的GFP阳性液泡比例显著高于ΔCfatg7，直至5 h，ΔCfatg7菌株液泡仍完全无绿色荧光，WT中绿色荧光几乎全部进入菌株液泡(图5C, 5D)。由于从GFP-Atg8上切割的完整GFP片段不会被液泡蛋白酶解，可通过观察游离GFP评估自噬活性。通过免疫印迹分析GFP-CfAtg8的降解情况发现，ΔCfatg7无游离GFP (图5E)，表明其丧失自噬水平。以上结果证实CfAtg7是自噬所必需的。
2.7 CfAtg7参与调控果生刺盘孢对环境胁迫的耐受性
病原真菌在侵染寄主植物的过程中，会引起植物的一系列生理生化反应来抵抗其侵染。真菌细胞壁作为真菌和周围环境的分界，是一层坚硬的屏障(东曼等2022)。在刚果红处理下，胁迫剂对ΔCfatg7的抑制率达(30±3)%，显著高于其对WT的抑制率(8%)；在SDS处理下，胁迫剂对ΔCfatg7的抑制率为65%，对WT的抑制率为(59±1.7)%，统计学差异极显著，表明ΔCfatg7对细胞壁完整性胁迫剂敏感性更高(图6)，说明CfAtg7参与果生刺盘孢应对细胞壁完整性胁迫应答的过程。
内质网是胞内蛋白质的修饰场所，蛋白质的正常修饰是细胞发挥生物学功能的基础(杨正婷和刘建祥 2016)。在内质网胁迫剂DTT处理下，其对野生型抑制率为(31.7±2.3)%，对ΔCfatg7的抑制率为(24.7±0.6)%，显著低于WT和ΔCfatg7/ CfATG7 [抑制率为(31.3±1.5)%] (图6)，表明CfAtg7参与调控果生刺盘孢应对寄主植物的内质网压力。
在侵染过程中，真菌需要和活性氧做斗争，过量的活性氧会破坏生物膜系统，影响细胞多种生理功能，造成细胞损伤(刘欢等 2019)。氧化胁迫剂H2O2对ΔCfatg7的抑制率为(56±1.7)%，对ΔCfatg7/CfATG7和野生型抑制率约为(18± 1.0)%，ΔCfatg7敏感性显著高于WT和回补菌株(图6)，表明CfAtg7蛋白参与调控果生刺盘孢对氧化胁迫的应答。
3 讨论
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本研究在果生刺盘孢Co. fructicola中鉴定了一个自噬关键蛋白CfAtg7，并分析了其生物学功能。我们发现CfAtg7对自噬过程至关重要，并参与果生炭疽菌的生长、分生孢子形成、附着胞发育、胁迫响应及致病性。
CfATG7基因缺失导致菌株生长速率下降，表明CfAtg7对生长具有正向调控作用，这与Atg7蛋白调控灰葡萄孢和禾谷镰孢菌的营养生长结果一致(Ren et al. 2018；吕务云 2019)。上述研究暗示Atg7调控生长的过程在物种间具有保守性。
自噬是真核生物对胞内物质进行重复利用所特有的一种保守过程。我们发现ΔCfatg7在氮饥饿诱导条件下自噬过程被阻断，这与已证明的植物病原菌的研究结果相符。该结果进一步印证了Atg7在自噬过程中的保守性。
近年来，越来越多的研究表明细胞自噬参与植物病原真菌的致病过程。稻巨座壳中细胞自噬可分别协同细胞壁完整性通路和脂质代谢从而控制致病过程(Zhu et al. 2023；Guo et al. 2024)。禾谷镰孢菌中去泛素化介导的细胞自噬调控病菌的致病力(Chen et al. 2024)。我们前期的研究也初步揭示了细胞自噬在果生炭疽菌致病过程中的作用，ΔCfatg7突变体的自噬缺陷可能预示其致病力受损。通过实验，我们发现ΔCfatg7在油茶叶片上的致病力显著减弱，这也与Atg7在稻巨座壳中的研究结果(刘小红等 2008)一致。
我们通过实验证明CfATG7基因缺失仍形成极少数附着胞，但形成率显著降低。推测附着胞形成率的降低是突变体致病力下降的主要原因。同时，ΔCfatg7对细胞壁完整性胁迫和氧化胁迫更为敏感。考虑到细胞壁是真菌与寄主间的第一道屏障，且病原菌必须抵御寄主迸发的ROS才能成功侵染寄主植物，我们认为ΔCfatg7对这两种胁迫的响应缺陷也可能导致其致病力下降。
综上所述，本研究不仅揭示了CfAtg7在自噬中的作用，还阐明了其在果生炭疽菌生长、产孢、附着胞形成、胁迫响应及致病性中的多效性功能。
作者贡献
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陈妍：实验操作、数据分析、论文撰写；王伊玲：实验操作、数据分析；罗晶：实验操作；张盛培：实验设计、论文修改。
利益冲突声明
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该研究不存在任何潜在利益冲突的商业或财务关系。
基金
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  • 湖南省自然科学基金(2023JJ20100)
  • 国家自然科学基金(32001317)
  • 国家重点研发计划(2023YFD1401301)
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2026年第45卷第3期
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doi: 10.13346/j.mycosystema.250178
  • 接收时间：2025-06-10
  • 首发时间：2026-04-30
  • 出版时间：2026-03-22
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  • 收稿日期：2025-06-10
  • 录用日期：2025-08-07
基金
Natural Science Foundation of Hunan Province(2023JJ20100)
湖南省自然科学基金(2023JJ20100)
National Natural Science Foundation of China(32001317)
国家自然科学基金(32001317)
National Key Research & Development Program of China(2023YFD1401301)
国家重点研发计划(2023YFD1401301)
作者信息
    中南林业科技大学 森林生物资源与有害生物综合管理湖南省普通高等学校重点实验室 木本油料资源利用全国重点实验室，湖南 长沙 410004

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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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