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Article(id=1316714666835951796, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1316714454184739470, articleNumber=null, orderNo=null, doi=10.13343/j.cnki.wsxb.20260259, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1774800000000, receivedDateStr=2026-03-30, revisedDate=null, revisedDateStr=null, acceptedDate=1777046400000, acceptedDateStr=2026-04-25, onlineDate=1791858839949, onlineDateStr=2026-10-13, pubDate=1791043200000, pubDateStr=2026-10-04, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1791858839949, onlineIssueDateStr=2026-10-13, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1791858839949, creator=13701087609, updateTime=1791858839949, updator=13701087609, issue=Issue{id=1316714454184739470, tenantId=1146029695717560320, journalId=1192105938417971205, year='2026', volume='66', issue='10', pageStart='4661', pageEnd='5310', issueExtLink='null', onlineDate='null', pubDate='1791043200000', pubDateStr='2026-10-04', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=1, specialIssue=null, createTime=1791858789249, creator='13701087609', updateTime=1791885400527, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext={EN=IssueExt(id=1316826070066774161, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1316714454184739470, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1316826070066774162, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1316714454184739470, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null, downloadFileDto=null}, startPage=5136, endPage=5148, ext={EN=ArticleExt(id=1316714667137941685, articleId=1316714666835951796, tenantId=1146029695717560320, journalId=1192105938417971205, language=EN, title=Roles of histone variant H2A.Z in the growth, development, and pathogenic mechanism of Fusarium graminearum, columnId=1192149543992045670, journalTitle=Acta Microbiologica Sinica, columnName=Research Article, runingTitle=null, highlight=null, articleAbstract=

[Objective] As a key component of eukaryotic chromatin, histone variant H2A.Z plays an important role in gene expression regulation. This study explored the regulatory effects of H2A.Z on the biological characteristics and pathogenicity of Fusarium graminearum, aiming to decipher its epigenetic regulation mechanism in the pathogenicity of pathogenic fungi. [Methods] The genome of F. graminearum was excavated by bioinformatics methods, and the H2A.Z homologous genes were identified by multiple sequence alignment and phylogenetic analysis. The gene deletion mutants were constructed by homologous recombination combined with PEG-mediated protoplast transformation. The phenotypic differences in morphological development, pathogenicity, and environmental adaptability between the wild type and the mutants were systematically evaluated by morphological observation, pathogenicity determination, and abiotic stress response experiments. [Results] H2A.Z was encoded by FgHTZ1. Phenotypic analysis showed that the deletion of FgHTZ1 resulted in significant retardation of colony growth, abnormal mycelial polar growth and conidial morphogenesis, and significant attenuation in pathogenicity. In addition, the mutants showed significantly altered sensitivity to osmotic stress (NaCl and sorbitol), oxidative stress (H2O2), and cell wall damage (Congo red), exhibiting a complex phenotype of enhanced tolerance to cell wall disruptors but hypersensitivity to osmotic/oxidative stress. [Conclusion] FgHTZ1 is the coding gene of H2A.Z in F. graminearum. As a positive regulator, this gene not only regulates the vegetative growth, morphological development, and pathogenicity of F. graminearum but also participates in mediating osmotic stress and oxidative stress responses, playing a key role in maintaining cell wall integrity. This study provides an important theoretical basis for further understanding the molecular functions of histone variants in the pathogenic mechanisms of plant pathogenic fungi.

, authors=Feifei SHANG, Jianhu LIU, Hongzhe CAO, Jinping ZANG, Kang ZHANG, Jihong XING*, Jingao DONG*, authorsList=Feifei SHANG, Jianhu LIU, Hongzhe CAO, Jinping ZANG, Kang ZHANG, Jihong XING, Jingao DONG, authorCompany=null, correspAuthors=Jihong XING, Jingao DONG, authorNote=

These authors contributed equally to this work.

, correspAuthorsNote=
*E-mail: XING Jihong,
DONG Jingao,
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【目的】 组蛋白变体H2A.Z作为真核生物染色质的关键组分，在基因表达调控中发挥重要作用。本研究旨在探究H2A.Z对禾谷镰孢(Fusarium graminearum)生物学特性及致病性的调控作用，解析其在病原真菌致病过程中的表观遗传调控机制。 【方法】 利用生物信息学手段对禾谷镰孢基因组进行挖掘，通过多序列比对和系统发育分析鉴定H2A.Z同源基因；采用同源重组策略结合聚乙二醇(polyethylene glycol, PEG)介导的原生质体转化体系构建基因缺失突变体；通过形态学观察、致病力测定及非生物胁迫响应实验系统评估野生型与突变体在形态发育、致病性及环境适应性方面的差异。 【结果】 本研究明确了禾谷镰孢H2A.Z由FgHTZ1基因编码。表型分析显示，FgHTZ1缺失导致菌落生长显著迟缓、菌丝极性生长与分生孢子形态建成异常，且致病性大幅减弱。此外，突变体对渗透胁迫(NaCl、山梨醇)、氧化应激(H2O2)及细胞壁损伤(刚果红)的敏感性发生显著改变，表现出对细胞壁干扰剂的耐受性增强但对渗透/氧化胁迫超敏的复杂表型。 【结论】 禾谷镰孢组蛋白变体H2A.Z的编码基因为FgHTZ1，该基因作为正调控因子，不仅调控禾谷镰孢的营养生长、形态发育和致病力，还参与介导菌体对渗透胁迫与氧化胁迫的适应性应答，并对维持细胞壁完整性具有关键作用。本研究为深入理解组蛋白变体在植物病原真菌致病机制中的分子功能提供了重要的理论基础。

, authors=尚飞飞, 柳健虎, 曹宏哲, 藏金萍, 张康, 邢继红*, 董金皋*, authorsList=尚飞飞, 柳健虎, 曹宏哲, 藏金萍, 张康, 邢继红, 董金皋, authorCompany=null, correspAuthors=邢继红, 董金皋, authorNote=

作者贡献声明

尚飞飞：实验操作；柳健虎：数据分析及论文初稿撰写；曹宏哲、藏金萍：审核与编辑写作；张康：项目管理，提供资源；邢继红：项目管理，提供资源，负责论文最终定稿；董金皋：指导监督。

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Trends in Cell Biology, 2014, 24(11): 642-650., articleTitle=Environmental responses mediated by histone variants, refAbstract=null)], funds=[Fund(id=1316829958501159454, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1316714666835951796, awardId=32072369, language=EN, fundingSource=National Natural Science Foundation of China(32072369), fundOrder=null, country=null), Fund(id=1316829958564074015, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1316714666835951796, awardId=32072369, language=CN, fundingSource=国家自然科学基金(32072369), fundOrder=null, country=null), Fund(id=1316829958635377184, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1316714666835951796, awardId=246Z6506G, language=EN, fundingSource=Central Guidance for Local Technology Development Funding(246Z6506G), fundOrder=null, country=null), Fund(id=1316829958698291745, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1316714666835951796, awardId=246Z6506G, language=CN, fundingSource=中央引导地方科技发展资金(246Z6506G), fundOrder=null, country=null)], companyList=[AuthorCompany(id=1316829952822071774, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1316714666835951796, xref=null, ext=[AuthorCompanyExt(id=1316829952826266079, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1316714666835951796, companyId=1316829952822071774, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=Key Laboratory of Hebei Province for Plant Physiology and Molecular Pathology, Hebei Agricultural University, Baoding, Hebei, China), AuthorCompanyExt(id=1316829952834654688, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1316714666835951796, companyId=1316829952822071774, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=河北农业大学，河北省植物生理与分子病理学重点实验室，河北 保定)])], figs=[ArticleFig(id=1316829957062513166, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1316714666835951796, language=EN, label=Figure 1, caption=Phylogenetic and conservative domain analysis of HTZs and HTAs. A: Phylogenetic tree of HTZs and HTAs of multiple species (Fo: Fusarium oxysporum; Fg: Fusariumgraminearum; Tg: Trichoderma gamsii; At: Aspergillus tanneri; Ao: Aspergillus ochraceoroseus; Ca: Candida albicans; Sc: Saccharomyces cerevisiae; Cn: Cryptococcus neoformans); B: Conserved domains of HTZs and HTAs of multiple species., figureFileSmall=pYp78H97f8ko16X6yUl7Xg==, figureFileBig=Mxd9u4/hxgqU8k5Cp5nRpA==, tableContent=null), ArticleFig(id=1316829957150593551, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1316714666835951796, language=CN, label=图1, caption=HTZs 和 HTAs 的系统发育分析与保守结构域分析, figureFileSmall=pYp78H97f8ko16X6yUl7Xg==, figureFileBig=Mxd9u4/hxgqU8k5Cp5nRpA==, tableContent=null), ArticleFig(id=1316829957247062544, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1316714666835951796, language=EN, label=Figure 2, caption=Construction and identification of the FgHTZ1 knockout mutants in Fusarium graminearum. A: Structure diagram of FgHTZ1 knockout mutants (1-3: Three exons of FgHTZ1 gene; FgHTZ1-I: FgHTZ1 homology arm I; FgHTZ1-II: FgHTZ1 homology arm II; Bar: Bialaphos resistance gene); B, C, D: Identification of FgHTZ1 mutants by PCR amplification using primer I-F/B-R, B-F/II-R, and Bar-F/Bar-R (Lane M: DNA marker; Lane 1: Wild type PH-1; Lanes 2, 3, 4: FgHTZ1 mutant Δhtz1-1, Δhtz1-2, and Δhtz1-3); E: The growth status of FgHTZ1 mutants on PDA medium containing 200 μg/mL bialaphos; F: Colony diameters of Figure 2E; G: Relative expression levels of FgHTZ1 gene in PH-1 and the FgHTZ1 mutants. Error bars indicate mean±SD. Asterisks indicate significant difference compared with wild type PH-1 by Student’s t-test (**: P<0.01). The same below., figureFileSmall=nmlqkNinshgDHTl7HyByfg==, figureFileBig=DQxv6DmfohULZ4+0mXocaQ==, tableContent=null), ArticleFig(id=1316829957301588497, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1316714666835951796, language=CN, label=图2, caption=禾谷镰孢 FgHTZ1 基因敲除突变体的构建与鉴定, figureFileSmall=nmlqkNinshgDHTl7HyByfg==, figureFileBig=DQxv6DmfohULZ4+0mXocaQ==, tableContent=null), ArticleFig(id=1316829957372891666, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1316714666835951796, language=EN, label=Figure 3, caption=The influence of FgHTZ1 mutation on the growth of Fusarium graminearum. A: Colony morphology of PH-1 and FgHTZ1 mutants; B: Growth rate of PH-1 and FgHTZ1 mutants; C: Mycelium weight of PH-1 and FgHTZ1 mutants (**: P<0.01)., figureFileSmall=JXia2Anxo0UmjkPxVwaegA==, figureFileBig=kRC061pJ9V9yDx227wgZpQ==, tableContent=null), ArticleFig(id=1316829957448389139, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1316714666835951796, language=CN, label=图3, caption=禾谷镰孢 FgHTZ1 基因突变对菌落生长的影响, figureFileSmall=JXia2Anxo0UmjkPxVwaegA==, figureFileBig=kRC061pJ9V9yDx227wgZpQ==, tableContent=null), ArticleFig(id=1316829957528080916, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1316714666835951796, language=EN, label=Figure 4, caption=The influence of FgHTZ1 mutation on hyphal development of Fusarium graminearum. A: Hyphal morphology of PH-1 and FgHTZ1 mutants [CFW: Calcofluor white (Bar=20 μm)]; B: Hyphal length of PH-1 and FgHTZ1 mutants (*: P<0.05; **: P<0.01); C: Hyphal width of PH-1 and FgHTZ1 mutants. Error bars indicate mean±SD. Asterisks indicate significant difference compared with wild type PH-1 by Student’s t-test (**: P<0.01)., figureFileSmall=hBBvtY0qaJwqIYlQgSVKHA==, figureFileBig=HWh9kBs3TxcXs1IGJe+90Q==, tableContent=null), ArticleFig(id=1316829957611966997, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1316714666835951796, language=CN, label=图4, caption=禾谷镰孢 FgHTZ1 基因突变对菌丝发育的影响, figureFileSmall=hBBvtY0qaJwqIYlQgSVKHA==, figureFileBig=HWh9kBs3TxcXs1IGJe+90Q==, tableContent=null), ArticleFig(id=1316829957712630294, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1316714666835951796, language=EN, label=Figure 5, caption=The influence of FgHTZ1 mutation on conidial development of Fusarium graminearum. A: Conidial morphology of PH-1 and FgHTZ1 mutants (Bar=10 μm); B, C: Conidial length and width of PH-1 and FgHTZ1 mutants; D: The number of conidia produced by PH-1 and FgHTZ1 mutants. *: P<0.05; **: P<0.01., figureFileSmall=/lr/y5gXLQSZ62vIgV1R+w==, figureFileBig=DL7Ud6FUvKUebe6p/kVp1A==, tableContent=null), ArticleFig(id=1316829957800710679, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1316714666835951796, language=CN, label=图5, caption=禾谷镰孢 FgHTZ1 基因突变对分生孢子发育的影响, figureFileSmall=/lr/y5gXLQSZ62vIgV1R+w==, figureFileBig=DL7Ud6FUvKUebe6p/kVp1A==, tableContent=null), ArticleFig(id=1316829957897179672, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1316714666835951796, language=EN, label=Figure 6, caption=The pathogenicity of FgHTZ1 mutants. A: Pathogenicity detection of PH-1 and FgHTZ1 mutants (Bar=2 cm); B: Measurement of lesion areas of ‘B73’ inoculated with PH-1 and FgHTZ1 mutants (*: P<0.05; **: P<0.01)., figureFileSmall=O0+NibbgSZaoM+hKbSPfNA==, figureFileBig=edQ+/jfstDhpTVEiFQnH8Q==, tableContent=null), ArticleFig(id=1316829957972677145, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1316714666835951796, language=CN, label=图6, caption=FgHTZ1 突变体的致病力测定, figureFileSmall=O0+NibbgSZaoM+hKbSPfNA==, figureFileBig=edQ+/jfstDhpTVEiFQnH8Q==, tableContent=null), ArticleFig(id=1316829958052368922, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1316714666835951796, language=EN, label=Figure 7, caption=Sensitivity of FgHTZ1 mutants to abiotic stress. A: Sensitivity detection of wild type and mutants to various stresses; B: Promotion rate of NaCl on the growth of PH-1 and FgHTZ1 mutants; C, D, E: Inhibition rate of sorbitol, H2O2, Congo red on the growth of PH-1 and FgHTZ1 mutants. *: P<0.05; **: P<0.01., figureFileSmall=v2hMbMUr75UmicAjLmyYlw==, figureFileBig=ONyk11UIVNqKxXQOX4UVnQ==, tableContent=null), ArticleFig(id=1316829958119477787, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1316714666835951796, language=CN, label=图7, caption=FgHTZ1 突变体对非生物胁迫的敏感性测定, figureFileSmall=v2hMbMUr75UmicAjLmyYlw==, figureFileBig=ONyk11UIVNqKxXQOX4UVnQ==, tableContent=null), ArticleFig(id=1316829958203363868, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1316714666835951796, language=EN, label=Table 1, caption=

Primers used in this study

, figureFileSmall=null, figureFileBig=null, tableContent=
Gene namesPrimer namesPrimer sequences (5′→3′)
FgHTZ1 homology arm II-FATCGATGCGAAGACGGCGAGGATGT
I-RGAATTCACGGTGGCGCCGCGTC
FgHTZ1 homology arm IIII-FGGATCCTGTCGATGTAGTCGCGTTAA
II-RTCTAGAGCCGCCACATTTAGCCAC
FgHTZ1, BARB-FATCTCGGTGACGGGCAATCGATGCGAAGACGGCG
B-RACGGTGGCGCCGCGTCATGAGCCCAGAACGACGC
BARBAR-FATGAGCCCAGAACGACGC
BAR-RTCAGATCTCGGTGACGGGCA
FgHTZ1RT-FAGCAGTAGAGCGCGGTTGATGTG
RT-RAAGTTCTGGAGTTGGCAGGCGTAAG
FgActinFgActin-FAACATTGTCATGTCTGGTGGTACC
FgActin-RCACTTGCGGTGAACGATTGA
), ArticleFig(id=1316829958320804381, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1316714666835951796, language=CN, label=表1, caption=

本研究所用引物

, figureFileSmall=null, figureFileBig=null, tableContent=
Gene namesPrimer namesPrimer sequences (5′→3′)
FgHTZ1 homology arm II-FATCGATGCGAAGACGGCGAGGATGT
I-RGAATTCACGGTGGCGCCGCGTC
FgHTZ1 homology arm IIII-FGGATCCTGTCGATGTAGTCGCGTTAA
II-RTCTAGAGCCGCCACATTTAGCCAC
FgHTZ1, BARB-FATCTCGGTGACGGGCAATCGATGCGAAGACGGCG
B-RACGGTGGCGCCGCGTCATGAGCCCAGAACGACGC
BARBAR-FATGAGCCCAGAACGACGC
BAR-RTCAGATCTCGGTGACGGGCA
FgHTZ1RT-FAGCAGTAGAGCGCGGTTGATGTG
RT-RAAGTTCTGGAGTTGGCAGGCGTAAG
FgActinFgActin-FAACATTGTCATGTCTGGTGGTACC
FgActin-RCACTTGCGGTGAACGATTGA
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组蛋白变体H2A.Z在禾谷镰孢生长发育及致病机制中的功能解析
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尚飞飞 , 柳健虎 , 曹宏哲 , 藏金萍 , 张康 , 邢继红 * , 董金皋 *
微生物学报 | 研究报告 2026,66(10): 5136-5148
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微生物学报 |研究报告 2026 , 66 (10) : 5136 -5148
组蛋白变体H2A.Z在禾谷镰孢生长发育及致病机制中的功能解析
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尚飞飞, 柳健虎, 曹宏哲, 藏金萍, 张康, 邢继红* , 董金皋*
作者信息
  • 河北农业大学，河北省植物生理与分子病理学重点实验室，河北 保定
通讯作者:
邢继红, 董金皋
作者简介:

作者贡献声明

尚飞飞：实验操作；柳健虎：数据分析及论文初稿撰写；曹宏哲、藏金萍：审核与编辑写作；张康：项目管理，提供资源；邢继红：项目管理，提供资源，负责论文最终定稿；董金皋：指导监督。

Roles of histone variant H2A.Z in the growth, development, and pathogenic mechanism of Fusarium graminearum
Feifei SHANG, Jianhu LIU, Hongzhe CAO, Jinping ZANG, Kang ZHANG, Jihong XING* , Jingao DONG*
Affiliations
  • Key Laboratory of Hebei Province for Plant Physiology and Molecular Pathology, Hebei Agricultural University, Baoding, Hebei, China
    • Corresponding Author:
    *E-mail: XING Jihong,
    DONG Jingao,
出版时间: 2026-10-04 doi: 10.13343/j.cnki.wsxb.20260259
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摘要
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【目的】 组蛋白变体H2A.Z作为真核生物染色质的关键组分，在基因表达调控中发挥重要作用。本研究旨在探究H2A.Z对禾谷镰孢(Fusarium graminearum)生物学特性及致病性的调控作用，解析其在病原真菌致病过程中的表观遗传调控机制。 【方法】 利用生物信息学手段对禾谷镰孢基因组进行挖掘，通过多序列比对和系统发育分析鉴定H2A.Z同源基因；采用同源重组策略结合聚乙二醇(polyethylene glycol, PEG)介导的原生质体转化体系构建基因缺失突变体；通过形态学观察、致病力测定及非生物胁迫响应实验系统评估野生型与突变体在形态发育、致病性及环境适应性方面的差异。 【结果】 本研究明确了禾谷镰孢H2A.Z由FgHTZ1基因编码。表型分析显示，FgHTZ1缺失导致菌落生长显著迟缓、菌丝极性生长与分生孢子形态建成异常，且致病性大幅减弱。此外，突变体对渗透胁迫(NaCl、山梨醇)、氧化应激(H2O2)及细胞壁损伤(刚果红)的敏感性发生显著改变，表现出对细胞壁干扰剂的耐受性增强但对渗透/氧化胁迫超敏的复杂表型。 【结论】 禾谷镰孢组蛋白变体H2A.Z的编码基因为FgHTZ1，该基因作为正调控因子，不仅调控禾谷镰孢的营养生长、形态发育和致病力，还参与介导菌体对渗透胁迫与氧化胁迫的适应性应答，并对维持细胞壁完整性具有关键作用。本研究为深入理解组蛋白变体在植物病原真菌致病机制中的分子功能提供了重要的理论基础。

禾谷镰孢  /  H2A.Z  /  FgHTZ1  /  基因敲除  /  致病力

[Objective] As a key component of eukaryotic chromatin, histone variant H2A.Z plays an important role in gene expression regulation. This study explored the regulatory effects of H2A.Z on the biological characteristics and pathogenicity of Fusarium graminearum, aiming to decipher its epigenetic regulation mechanism in the pathogenicity of pathogenic fungi. [Methods] The genome of F. graminearum was excavated by bioinformatics methods, and the H2A.Z homologous genes were identified by multiple sequence alignment and phylogenetic analysis. The gene deletion mutants were constructed by homologous recombination combined with PEG-mediated protoplast transformation. The phenotypic differences in morphological development, pathogenicity, and environmental adaptability between the wild type and the mutants were systematically evaluated by morphological observation, pathogenicity determination, and abiotic stress response experiments. [Results] H2A.Z was encoded by FgHTZ1. Phenotypic analysis showed that the deletion of FgHTZ1 resulted in significant retardation of colony growth, abnormal mycelial polar growth and conidial morphogenesis, and significant attenuation in pathogenicity. In addition, the mutants showed significantly altered sensitivity to osmotic stress (NaCl and sorbitol), oxidative stress (H2O2), and cell wall damage (Congo red), exhibiting a complex phenotype of enhanced tolerance to cell wall disruptors but hypersensitivity to osmotic/oxidative stress. [Conclusion] FgHTZ1 is the coding gene of H2A.Z in F. graminearum. As a positive regulator, this gene not only regulates the vegetative growth, morphological development, and pathogenicity of F. graminearum but also participates in mediating osmotic stress and oxidative stress responses, playing a key role in maintaining cell wall integrity. This study provides an important theoretical basis for further understanding the molecular functions of histone variants in the pathogenic mechanisms of plant pathogenic fungi.

Fusarium graminearum  /  H2A.Z  /  FgHTZ1  /  gene knockout  /  pathogenicity
尚飞飞, 柳健虎, 曹宏哲, 藏金萍, 张康, 邢继红, 董金皋. 组蛋白变体H2A.Z在禾谷镰孢生长发育及致病机制中的功能解析. 微生物学报, 2026 , 66 (10) : 5136 -5148 . DOI: 10.13343/j.cnki.wsxb.20260259
Feifei SHANG, Jianhu LIU, Hongzhe CAO, Jinping ZANG, Kang ZHANG, Jihong XING, Jingao DONG. Roles of histone variant H2A.Z in the growth, development, and pathogenic mechanism of Fusarium graminearum[J]. Acta Microbiologica Sinica, 2026 , 66 (10) : 5136 -5148 . DOI: 10.13343/j.cnki.wsxb.20260259
正文
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禾谷镰孢(Fusarium graminearum)是引发小麦赤霉病和玉米茎腐病的主要病原菌，对全球粮食安全构成严重威胁。该真菌不仅导致农作物大幅减产，其产生的脱氧雪腐镰刀菌烯醇(deoxynivalenol, DON)和玉米赤霉烯酮(zearalenone, ZEA)等次级代谢产物，更因具有强细胞毒性而严重危害人畜健康[1-2]。随着全基因组测序的完成，禾谷镰孢已成为解析植物-病原真菌互作机制的模式生物，其致病相关基因的功能鉴定与调控网络解析取得了显著进展[3]。深入挖掘禾谷镰孢关键基因的生物学功能，不仅有助于揭示其致病分子机理，更为研发新型绿色防控策略提供理论支撑。
在真核生物中，染色质的基本结构单位核小体由146 bp DNA缠绕组蛋白八聚体(含H2A、H2B、H3、H4各2分子)构成。除保守的核心组蛋白外，组蛋白变体在DNA复制、基因表达调控及染色体动力学行为中扮演关键角色[4-6]。H2A.Z是进化上高度保守的组蛋白变体，约占细胞总H2A蛋白的15%。与典型H2A相比，其C端序列的显著差异赋予了其独特的结构与功能特性[7-8]。研究表明，H2A.Z在哺乳动物中是胚胎干细胞分化与基因激活的关键调控因子，其缺失可导致胚胎发育停滞[9-10]；在酿酒酵母中，H2A.Z则是减数分裂期间DNA重组的必需因子[11-12]。值得注意的是，尽管H2A.Z缺失在酵母中不致死，但在高等生物中却表现出致死效应，暗示该蛋白在不同进化层级的生物中存在功能分化[13-14]。从空间分布上看，H2A.Z偏好性地富集于核小体边缘，尤其在转录起始位点(transcription start site, TSS)附近的核小体缺失区及启动子区域高度聚集[15-16]。
近年来，H2A.Z在植物病原真菌中的功能逐渐受到关注。研究表明，H2A.Z在部分植物病原真菌中参与调控菌丝生长、分生孢子发育及致病性等关键生物学过程。例如，在水稻恶苗病菌(Fusarium fujikuroi)中，H2A.Z的缺失显著影响了菌丝的正常生长和分生孢子的形成，进而降低了病原菌的致病力[17]。然而，尽管H2A.Z在其他植物病原真菌中的功能已有初步探索，关于H2A.Z在禾谷镰孢中的具体调控机制，尤其是其在生长发育、致病性及环境适应性中的分子调控网络目前仍知之甚少。这一研究空白不仅限制了我们对禾谷镰孢致病机理的深入理解，也阻碍了基于表观遗传调控的新型防控策略的开发。
尽管H2A.Z在真核生物中的基础功能已被广泛报道，但其在禾谷镰孢生长发育及致病过程中的具体生物学功能与分子调控机制尚不明确。因此本研究拟通过禾谷镰孢基因组信息精准鉴定H2A.Z编码基因，并通过同源重组技术构建基因敲除突变体。在此基础上，系统评估该基因缺失对菌丝形态建成、产孢能力、次级代谢产物合成及致病性等关键表型的影响。本研究旨在揭示H2A.Z在禾谷镰孢中的功能角色，为解析病原真菌表观遗传调控网络提供直接证据。鉴于H2A.Z兼具高度保守性与物种特异性，本研究不仅有助于深化对禾谷镰孢致病机理的理解，也为探索植物-病原互作的进化机制提供重要的理论依据。
1 材料与方法
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1.1 材料
本研究所用的禾谷镰孢菌PH-1菌株、同源重组载体pBS-Bar、‘郑单958’玉米品种及玉米自交系‘B73’均由河北农业大学河北省植物生理与分子病理学重点实验室提供并保存。
1.2 禾谷镰孢 FgHTZ1 基因的生物信息学鉴定与系统进化分析
从Ensembl真菌数据库(http://fungi.ensembl.org/index.html)下载8种代表性真菌的H2A.Z和H2A蛋白序列，包括禾谷镰孢(Fusarium graminearum)、尖孢镰孢(Fusarium oxysporum)、盖姆斯木霉(Trichoderma gamsii)、构巢曲霉(Aspergillus nidulans)、坦纳曲霉(Aspergillus tanneri)、白色念珠菌(Candida albicans)、新型隐球菌(Cryptococcus neoformans)和酿酒酵母(Saccharomyces cerevisiae)。在进化分析方面，运用MEGA 7软件中的邻近法构建系统发育树。通过Pfam (http://pfam.xfam.org/)和SMART (http:// smart.embl-heidelberg.de/)在线工具对各物种H2A.Z蛋白的功能域进行预测分析，并利用IBS软件可视化功能域结构。
1.3 禾谷镰孢 FgHTZ1 基因敲除突变体的构建
基于FgHTZ1基因与pBS-Bar载体多克隆位点的匹配性，设计包含Cla I、EcoR I、BamH I和Xba I 4种限制性内切酶位点的特异性引物(表1)，用于扩增FgHTZ1基因的2个同源重组片段I和II。将PCR产物与pBS-Bar载体进行连接，构建FgHTZ1基因敲除载体pBS+II+I。采用PEG介导的原生质体转化方法，将该重组载体导入禾谷镰孢野生型菌株PH-1中。经过抗性培养基筛选和单孢分离纯化后获得转化子。为验证转化效果，根据Bar抗性基因和FgHTZ1基因同源臂序列设计特异性引物进行PCR检测。同时，以FgActin基因为内参，通过RT-qPCR技术定量分析转化子中FgHTZ1基因的表达水平变化。
1.4 突变体的菌落形态与生长速率测定
系统比较禾谷镰孢野生型PH-1及其突变体的菌落形态与生长特性。在固体培养条件下，将菌株接种于PDA平板上，置于25 ℃黑暗培养。每日定时测量并记录菌落直径，以绘制生长曲线并分析生长速率差异；同时仔细观察并记录菌落的形态特征(如边缘形状、气生菌丝密度)及色素产生情况。实验过程中使用直径5 mm的无菌打孔器从菌落边缘切取菌饼，转接至新鲜PDA平板以进行继代培养。在液体培养条件下，将野生型和突变体的分生孢子悬浮液(浓度需保持一致)接种于PDB液体培养基中，25 ℃、200 r/min培养72 h。培养结束后，收集菌丝体并置于37 ℃烘箱中烘干至恒重，测定生物量(干重)。所有实验均设置3次生物学重复，以确保数据的可靠性与统计学意义。
1.5 突变体的菌丝形态观察
将野生型PH-1及突变体菌丝接种于PDA平板上，每皿斜插一张无菌盖玻片以利于菌丝附着，25 ℃暗培养。待菌丝覆盖盖玻片后，用灭菌镊子取出盖玻片置于载玻片上，采用植物细胞壁特异性荧光染料钙荧光白(calcofluor white, CFW)于30 ℃避光染色30 min。随后在暗室条件下利用荧光显微镜观察菌丝形态，重点统计菌丝分隔间距、菌丝宽度等参数。实验设3次生物学重复。
1.6 突变体的分生孢子形态观察与产孢量测定
取预培养5 d (25 ℃)的野生型及突变体菌落，用5 mm打孔器从边缘切取3个菌块，转接至装有100 mL 羧甲基纤维素钠(CMC-Na)液体培养基的三角瓶中，200 r/min培养5 d以诱导产孢。培养结束后，经4层无菌纱布过滤去除菌丝碎片，滤液用于后续分析。利用光学显微镜观察分生孢子的形态特征，并采用血球计数板进行显微计数以测定产孢量。为确保数据准确性，每个处理设置3次生物学重复。
1.7 突变体的致病力测定
选用玉米自交系‘B73’作为寄主材料，采用茎部注射法进行致病力评估。于玉米10叶期，在第二茎节间注射100 μL浓度为1×106个/mL的分生孢子悬浮液，同时设置羧甲基纤维素钠(CMC-Na)液体培养基和无菌水作为阴性对照。注射后用无菌湿纱布包扎伤口以维持湿度，每处理设置3次重复。分别于接种后3、7、14 d进行表型调查，通过纵向剖茎观察病斑扩展情况并测量病斑长度以此量化评价突变体的致病能力。
1.8 突变体对非生物胁迫的敏感性测定
采用生长速率法评估突变体对非生物胁迫的耐受性。将野生型PH-1及突变体菌饼(直径5 mm)接种于含不同胁迫剂的PDA平板上，胁迫条件包括：NaCl (0.8 mol/L)、山梨醇(1.0 mol/L)、H2O2 (10.0 mmol/L)和刚果红(2 mg/mL)。25 ℃黑暗培养72 h后，采用十字交叉法测量菌落直径，并详细记录菌落形态及色素分泌情况。每组设3次生物学重复。以对照组菌落直径为基准，按公式(1)计算抑制率。
抑制率=[(对照菌落直径-处理菌落直径)/对照菌落直径]×100%
1.9 数据统计
采用Graphpad Prism 10.1.2软件进行数据统计分析，具体运用单因素方差分析(one-way analysis of variance, one-way ANOVA)来评估不同处理组之间的差异显著性。在确定存在显著差异后，进一步采用Tukey法进行多重比较，以精确识别各组间的具体差异。为确保实验结果的可靠性和重复性，每个试验均设置3次独立的生物学重复。
2 结果与分析
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2.1 禾谷镰孢 FgHTZ1 基因的生物信息学鉴定与进化分析
本研究利用生物信息学手段，对8种真菌的H2A.Z及H2A蛋白序列进行了系统性进化分析。在基因鉴定方面，通过BLAST搜索禾谷镰孢PH-1基因组数据库，确定FGSG_01057为编码H2A.Z蛋白的基因，命名为FgHTZ1，且蛋白同源性分析表明其与稻瘟病菌、粗糙脉孢菌中的H2A.Z同源蛋白一致性超过75%。经全基因组检索、保守结构域注释与系统发育分析验证，禾谷镰孢基因组中仅FgHTZ1一个基因编码组蛋白变体H2A.Z，不存在其他旁系同源基因。系统发育树构建结果显示，编码H2A.Z的HTZ基因与编码H2A的HTA基因分别形成了独立的单系群(图1A)，表明二者在进化过程中发生了明显分化。其中，禾谷镰孢FgHTZ1与尖孢镰孢FoHTZ1的聚类关系最近，进化距离最短。进一步通过保守结构域预测发现，这些基因均包含典型的H2A核心结构域(图1B)。上述结果从分子进化和结构生物学层面确证了组蛋白变体H2A.Z在真菌界具有高度的保守性特征。
2.2 禾谷镰孢 FgHTZ1 基因敲除突变体的构建与鉴定
本研究利用同源重组技术构建了FgHTZ1基因敲除载体pBS+II+I (图2A)，并通过PEG介导的原生质体转化法筛选获得3个具有草铵膦(bialaphos)抗性的转化子，分别命名为Δhtz1-1、Δhtz1-2和Δhtz1-3。PCR验证结果显示，使用外侧引物组合I-F/B-R和B-F/II-R在突变体中均扩增出约1 500 bp的特异性条带，而内侧引物B-F/B-R仅扩增出约500 bp的短片段(图2B-2D)，这一结果确证了外源片段已通过同源重组准确插入目标位点。在抗性培养基上，野生型PH-1的生长受到显著抑制(菌落直径3.3 cm)，而突变体菌株则表现出良好的抗性且生长旺盛(菌落直径达8.0-8.5 cm) (图2E-2F)。RT-qPCR检测显示，突变体中FgHTZ1基因的转录水平较野生型显著下调(图2G)。综合分子鉴定、抗性表型及转录分析结果，确认FgHTZ1基因敲除突变体构建成功。
2.3 禾谷镰孢 FgHTZ1 基因突变对菌落生长及生物量积累的影响
菌落形态观察显示，FgHTZ1突变体与野生型PH-1在菌落形态上无显著差异，但色素合成能力明显减弱(图3A)，表明FgHTZ1可能参与调控禾谷镰孢的色素代谢途径。生长速率分析表明，Δhtz1-1和Δhtz1-2突变株的生长速率与野生型基本一致，而Δhtz1-3突变株的菌落扩展受到显著抑制。接种后第1天起，Δhtz1-3突变株的菌落直径即显著小于野生型PH-1，且该差异在第2天和第3天持续存在(图3B)。值得注意的是，尽管Δhtz1-3的生长速率减慢，其菌丝干重(1.9 g)却显著高于野生型(1.4 g)；而其余2株突变体的生物量(1.2-1.4 g)与野生型无显著差别(图3C)。上述结果揭示了FgHTZ1基因缺失对病原真菌营养生长和生物量形成的复杂调控效应，暗示该基因可能在不同遗传背景或突变株中存在功能分化。
2.4 禾谷镰孢 FgHTZ1 基因突变对菌丝发育的影响
利用光学显微镜与荧光显微镜对野生型及突变体的菌丝形态进行对比观察，结果显示二者在菌丝外观上未呈现显著差异(图4A)。然而，形态学定量分析结果显示，突变体菌丝的隔膜间距及菌丝直径均较野生型明显减小(图4B、4C)。上述微观结构的改变表明FgHTZ1基因的缺失对菌丝的极性生长及细胞形态建成具有关键调控作用。
2.5 禾谷镰孢 FgHTZ1 基因突变对分生孢子发育的影响
显微形态学观察显示，FgHTZ1突变体的分生孢子仍保持典型的镰刀状形态，且具有3-5个隔膜，在整体形态特征上与野生型无显著差异(图5A)。然而，形态学定量分析揭示了细微但显著的表型变化：突变体分生孢子的长度(0.17-0.19 mm)较野生型(0.28 mm)显著缩短，而孢子宽度则未受明显影响(图5B、5C)。更为关键的是，突变体的产孢能力表现出显著缺陷，其分生孢子产量[(1.0-2.5)×105个/mL]显著低于野生型(3.0×105个/mL) (图5D)。上述结果表明，FgHTZ1基因虽不决定孢子的基本形态建成，但对孢子的尺寸调控及产量形成具有不可或缺的关键作用。
2.6 禾谷镰孢 FgHTZ1 基因突变对致病力的影响
致病力测定结果显示，野生型菌株接种玉米后，在接种3 d即可观察到明显的变色病斑，且随着病程推进病斑逐渐扩展并颜色加深。相比之下，FgHTZ1突变体菌株接种后表现出显著减弱的致病表型，病斑扩展受限且症状较轻(图6A)。为进一步量化致病力差异，本研究对病斑面积进行了统计分析。结果表明，在接种后3、7、14 d，突变体的病斑扩展面积均显著低于野生型(图6B)。上述表型与定量数据共同证实，FgHTZ1基因的缺失显著削弱了禾谷镰孢的致病能力。
2.7 禾谷镰孢 FgHTZ1 基因突变对非生物胁迫响应的影响
为阐明FgHTZ1基因在胁迫应答中的生物学功能，本研究系统分析了突变体对渗透胁迫(NaCl、山梨醇)、氧化胁迫(H2O2)及细胞壁干扰剂(刚果红)的响应特性。在渗透胁迫下，NaCl处理对野生型和突变体均表现出一定的生长促进效应，但突变体的促进幅度显著弱于野生型(图7A、7B)；而在山梨醇胁迫下，野生型生长仅受到轻微抑制，Δhtz1-1和Δhtz1-2突变株的生长受抑制程度则显著加重(图7A、7C)。在氧化胁迫下，Δhtz1-2和Δhtz1-3对H2O2的敏感性显著增强(图7A、7D)。尤为值得注意的是，在细胞壁干扰剂刚果红处理组中，野生型表现出明显的生长抑制，而3个突变株却表现出显著增强的耐受性(图7A、7E)。上述结果证实，FgHTZ1不仅参与调控渗透胁迫和氧化胁迫响应途径，还在维持细胞壁结构完整性方面发挥着关键作用。
3 讨论
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在真核生物中，DNA复制完成后核心组蛋白H2A、H2B、H3和H4被精确地组装到染色质中，这些组蛋白在转录调控、DNA复制以及染色体分离等核心生命过程中发挥着不可替代的作用[18]。值得注意的是，H2A.Z作为H2A的一个重要变体，在进化过程中表现出了高度的保守性，这种保守性不仅体现在其氨基酸序列上，更重要的是体现在其功能上。凭借其独特的理化性质，H2A.Z能够显著改变核小体的稳定性，进而重塑染色质的高级结构，实现对基因表达的精细调控。在模式植物拟南芥中H2A.Z已被证实参与调控开花时间、免疫应答以及温度信号传导等多个关键生理过程[19-21]；而在酿酒酵母中H2A.Z则是确保减数分裂过程精确性的决定性因素[22]。
尽管H2A.Z在多种生物中的功能已有广泛研究，但其在禾谷镰孢这一重要植物病原菌中的具体生物学功能却尚未被系统解析。本研究通过生物信息学手段精准鉴定了FgHTZ1基因编码禾谷镰孢H2A.Z，并利用同源重组技术成功构建了该基因的敲除突变体。表型分析揭示，FgHTZ1的缺失对禾谷镰孢产生了多效性影响，这些影响不仅体现在菌株的营养生长和形态建成上，更显著地削弱了其致病能力。具体而言，FgHTZ1缺失突变体在菌落生长速率、菌丝极性生长、分生孢子形态建成及产孢能力等方面均表现出显著缺陷，这些缺陷直接导致其致病力的显著下降。
进一步地分析发现，FgHTZ1的缺失还显著影响了禾谷镰孢对渗透压和氧化胁迫的响应能力。突变体对NaCl和山梨醇等渗透胁迫剂以及H2O2等氧化胁迫剂的敏感性显著增强，这表明FgHTZ1在禾谷镰孢的胁迫应答机制中扮演着至关重要的角色。尽管突变体对细胞壁干扰剂刚果红的耐受性有所增强，但这种耐受性可能源于细胞壁组分或通透性的适应性改变，这些改变在赋予突变体抵抗刚果红能力的同时，也可能削弱细胞壁的整体稳定性，从而使其对渗透和氧化胁迫更为敏感。上述结果暗示FgHTZ1可能参与调控细胞壁合成或修复相关通路，具体机制有待进一步探究。
在深入探讨FgHTZ1的作用机制时，本研究发现其调控作用可能具有剂量依赖性。在3个独立的突变体中，FgHTZ1表达量最低的Δhtz1-3突变体表现出生长迟缓最为显著，这表明FgHTZ1的表达水平对禾谷镰孢的正常生长发育至关重要。此外，本研究还发现FgHTZ1可能通过影响多个生物学过程来调控禾谷镰孢的致病性，这些过程包括但不限于菌丝生长、分生孢子发育、胁迫应答及细胞壁完整性维持等，显示了H2A.Z在病原菌生物学中的复杂而核心的作用。
为了更全面地理解H2A.Z在植物病原菌中的功能，本研究进一步对比了H2A.Z在其他植物病原菌中的调控作用。例如，在水稻恶苗病菌(Fusarium fujikuroi)和粗糙脉孢菌(Neurospora crassa)中，H2A.Z缺失突变体同样表现出显著减缓的菌丝生长速率和降低的致病力[17,23]。本研究在禾谷镰孢中的研究结果与前人研究结论高度一致，进一步支持了H2A.Z在植物病原菌生长发育及致病过程中的核心调控作用。
在分子机制层面，SWR1染色质重塑复合物被证实是介导经典组蛋白H2A被H2A.Z取代的核心调控因子[24]。酵母遗传学实验显示，SWR1复合物功能缺陷会导致H2A.Z无法发挥其生物学功能，且Δswr1 Δhtz1双突变体的生长障碍比Δhtz1单突变体更为严重，这揭示了SWR1与H2A.Z之间紧密的功能关联性[25]。在真核生物中，SWR1复合物介导的H2A.Z整合是基因表达调控的关键环节，它通过改变染色质构象来影响下游基因的转录活性[26-28]。此外，H2A.Z在有丝分裂异染色质中的特异性定位也使其成为发育调控和环境适应过程中的关键转录调控因子[29]。
基于以上研究背景，我们推测在禾谷镰孢中SWR1复合物可能同样通过介导H2A.Z的整合来调控相关基因的表达，进而影响菌株的生长发育及致病性。因此，深入探究禾谷镰孢中SWR1复合物与FgHTZ1的互作机制将有助于从表观遗传调控层面全面阐明H2A.Z调控病原菌生长发育及致病性的分子基础。这不仅有助于更深入地理解禾谷镰孢的致病机理，也为开发基于表观遗传调控的新型防控策略提供了重要的理论依据和潜在的靶点。
4 结论
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本研究鉴定了禾谷镰孢组蛋白变体H2A.Z的编码基因FgHTZ1，并揭示了其作为核心正调控因子的生物学功能。FgHTZ1不仅正向调控禾谷镰孢的营养生长、形态建成和致病力，还参与介导了菌株对渗透胁迫和氧化胁迫的应答反应，并对维持细胞壁完整性具有关键作用。这些发现为深入阐明H2A.Z组蛋白变体在禾谷镰孢生物学特性调控中的分子机理提供了重要的理论依据。
基金
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  • 国家自然科学基金(32072369)
  • 中央引导地方科技发展资金(246Z6506G)
文献
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2026年第66卷第10期
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doi: 10.13343/j.cnki.wsxb.20260259
  • 接收时间：2026-03-30
  • 首发时间：2026-10-13
  • 出版时间：2026-10-04
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  • 收稿日期：2026-03-30
  • 录用日期：2026-04-25
基金
National Natural Science Foundation of China(32072369)
国家自然科学基金(32072369)
Central Guidance for Local Technology Development Funding(246Z6506G)
中央引导地方科技发展资金(246Z6506G)
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    河北农业大学，河北省植物生理与分子病理学重点实验室，河北 保定

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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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