Article(id=1226554097655001107, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1226554095926952065, articleNumber=null, orderNo=null, doi=10.13343/j.cnki.wsxb.20240825, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1734537600000, receivedDateStr=2024-12-19, revisedDate=null, revisedDateStr=null, acceptedDate=1745856000000, acceptedDateStr=2025-04-29, onlineDate=1770362885153, onlineDateStr=2026-02-06, pubDate=1751558400000, pubDateStr=2025-07-04, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1770362885153, onlineIssueDateStr=2026-02-06, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1770362885153, creator=13701087609, updateTime=1770362885153, updator=13701087609, issue=Issue{id=1226554095926952065, tenantId=1146029695717560320, journalId=1192105938417971205, year='2025', volume='65', issue='7', pageStart='2771', pageEnd='3233', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1770362884741, creator=13701087609, updateTime=1770363575040, updator=13701087609, preIssue=null, nextIssue=null, ext={EN=IssueExt(id=1226556991309529548, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1226554095926952065, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1226556991309529549, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1226554095926952065, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null}, startPage=2811, endPage=2829, ext={EN=ArticleExt(id=1226554098858766358, articleId=1226554097655001107, tenantId=1146029695717560320, journalId=1192105938417971205, language=EN, title=Advances in fungal apoptosis-like cell death and targeted drugs, columnId=1192149543727808575, journalTitle=Acta Microbiologica Sinica, columnName=Review, runingTitle=null, highlight=null, articleAbstract=

Currently, the drugs available for treating fungal infections show limited number and off-target effects. Thus, there is an urgent need to develop new antifungal drugs. Fungal apoptosis-like cell death (ALCD) is a cell death phenomenon that occurs during the normal development stage of organisms. This article summarizes the characteristics, signaling pathways, and key factors involved in fungal ALCD and introduces the natural and synthetic drugs that can induce fungal apoptosis. The natural drugs include lipid peptides, farnesol, statins, and alkaloids from microorganisms, organic acids and essential oils from plants, and melittins from insects. Furthermore, this article establishes the basic molecular landscape of drug-induced fungal ALCD. This article provides a theoretical basis for formulating a new strategy for resisting pathogenic fungi and developing targeted antifungal drugs.

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目前，治疗真菌感染的药物数量有限且已产生脱靶效应，研发新型抗真菌药物迫在眉睫。真菌凋亡样细胞死亡(apoptosis-like cell death, ALCD)是生物体在正常发育阶段细胞发生的死亡现象。本文概述了真菌ALCD的特征、所涉及的信号通路及关键因子，介绍了可诱导真菌凋亡的天然药物和人工合成药物。其中，天然药物包括来自微生物源的脂肽、法尼醇、他汀类、生物碱，来自植物源的有机酸、精油，以及来自昆虫的蜂毒素，并绘制了药物诱导真菌ALCD的基础分子景观。本文为制定新的抗病原真菌策略和研发靶向抗真菌药物提供了理论依据。

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Study on andrographolide-induced apoptosis of Candida albicans biofilm dispersion cells[J]. China Journal of Chinese Materia Medica, 2012, 37(3): 362-365 (in Chinese)., articleTitle=null, refAbstract=null), Reference(id=1227681752693535578, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226554097655001107, doi=null, pmid=null, pmcid=null, year=2020, volume=104, issue=15, pageStart=6513, pageEnd=6526, url=null, language=null, rfNumber=[141], rfOrder=156, authorNames=MEMARIANI H, MEMARIANI M, journalName=Applied Microbiology and Biotechnology, refType=null, unstructuredReference=MEMARIANI H, MEMARIANI M. Anti-fungal properties and mechanisms of melittin[J]. 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Insect Molecular Biology, 2014, 23(6): 788-799., articleTitle=Identification of a novel antimicrobial peptide, scolopendin 1, derived from centipede Scolopendra subspinipes mutilans and its antifungal mechanism, refAbstract=null), Reference(id=1227681752802587484, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226554097655001107, doi=null, pmid=null, pmcid=null, year=2015, volume=null, issue=null, pageStart=null, pageEnd=null, url=null, language=null, rfNumber=[143], rfOrder=158, authorNames=崔杨, journalName=null, refType=null, unstructuredReference=崔杨. 壳寡糖诱导交链孢菌细胞凋亡及毒素抑制的研究[D]. 天津: 天津科技大学硕士学位论文, 2015., articleTitle=壳寡糖诱导交链孢菌细胞凋亡及毒素抑制的研究, refAbstract=null), Reference(id=1227681752865502045, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226554097655001107, doi=null, pmid=null, pmcid=null, year=2015, volume=null, issue=null, pageStart=null, pageEnd=null, url=null, language=null, rfNumber=[143], rfOrder=159, authorNames=CUI Y, journalName=null, refType=null, unstructuredReference=CUI Y. The research that oligosaccharide induce the Alternaria and inhibit the toxins[D]. Tianjin: Master’s Thesis of Tianjin University of Science and Technology, 2015 (in Chinese)., articleTitle=null, refAbstract=null)], funds=[Fund(id=1227681727024394289, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226554097655001107, awardId=32172468, language=EN, fundingSource=National Natural Science Foundation of China(32172468), fundOrder=null, country=null), Fund(id=1227681727129251903, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226554097655001107, awardId=32172468, language=CN, fundingSource=国家自然科学基金(32172468), fundOrder=null, country=null)], companyList=[AuthorCompany(id=1227681717859840618, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226554097655001107, xref=null, ext=[AuthorCompanyExt(id=1227681717897589361, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226554097655001107, companyId=1227681717859840618, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=Engineering Research Center of Agricultural Microbiology Technology, Ministry of Education, Heilongjiang Provincial Key Laboratory of Plant Genetic Engineering and Biological Fermentation Engineering for Cold Region, Key Laboratory of Microbiology in Colleges and Universities of Heilongjiang Province, School of Life Science, Heilongjiang University, Harbin, Heilongjiang, China), AuthorCompanyExt(id=1227681717901783666, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226554097655001107, companyId=1227681717859840618, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=黑龙江大学 生命科学学院，农业微生物技术教育部工程研究中心，黑龙江省寒区植物基因与生物发酵重点 实验室，黑龙江省普通高校微生物重点实验室，黑龙江 哈尔滨)])], figs=[ArticleFig(id=1227681722293220248, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226554097655001107, language=EN, label=Figure 1, caption=The differences between C14 family proteases and inhibitors of apoptosis proteins in fungi and animals^[59]. A: C14 family proteases; B: Animal IAPs; C: Fungal IAPs., figureFileSmall=/tnjf7WEwxZECvEsWFjGwA==, figureFileBig=6lle/zcVEGFZxPuNvN2ZLA==, tableContent=null), ArticleFig(id=1227681722406466471, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226554097655001107, language=CN, label=图1, caption=真菌和动物C14家族蛋白酶和凋亡抑制蛋白的区别^[59]。A：C14家族蛋白酶；B：动物IAPs；C：真菌IAPs。, figureFileSmall=/tnjf7WEwxZECvEsWFjGwA==, figureFileBig=6lle/zcVEGFZxPuNvN2ZLA==, tableContent=null), ArticleFig(id=1227681722561655734, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226554097655001107, language=EN, label=Figure 2, caption=The molecular mechanism of fungal apoptosis-like cell death., figureFileSmall=It0mNoOH0rE7lv0VQzFG6w==, figureFileBig=5WAoZJ4gWOXgCBlr/Np/yQ==, tableContent=null), ArticleFig(id=1227681722670707647, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226554097655001107, language=CN, label=图2, caption=真菌凋亡样细胞死亡的分子机制, figureFileSmall=It0mNoOH0rE7lv0VQzFG6w==, figureFileBig=5WAoZJ4gWOXgCBlr/Np/yQ==, tableContent=null), ArticleFig(id=1227681722767176645, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226554097655001107, language=EN, label=Figure 3, caption=Molecular mechanism of apoptosis induced by cinnamaldehyde and monoterpene nerolol in fungi., figureFileSmall=S/hMnpuVQmBoAU6CcRwEHA==, figureFileBig=luchf8lhNwrXEReSpUrs4Q==, tableContent=null), ArticleFig(id=1227681722867839949, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226554097655001107, language=CN, label=图3, caption=肉桂醛(CA)和橙花酚单萜(NEL)诱导真菌ALCD的分子机制, figureFileSmall=S/hMnpuVQmBoAU6CcRwEHA==, figureFileBig=luchf8lhNwrXEReSpUrs4Q==, tableContent=null), ArticleFig(id=1227681722981086171, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226554097655001107, language=EN, label=Table 1, caption=

Key elements of apoptosis and their roles

, figureFileSmall=null, figureFileBig=null, tableContent=

Key elements	Inhibit/Activate apoptosis	Drug-target suitability	Comparison with animals or humans	References
AIF	Activate	Yes	Different	[45,56]
meCAS	Activate	Yes	Fungi specific	[68]
IAPs	Inhibit	Yes	Fungi specific	[82]
IAP-APs	Activate	Yes	Fungi specific	[89]
GSH	Inhibit	No	Lack of specificity	[91]
Sphingolipids	Activate	Yes	Different	[92]

), ArticleFig(id=1227681723115303911, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226554097655001107, language=CN, label=表1, caption=

凋亡关键元件及其作用

, figureFileSmall=null, figureFileBig=null, tableContent=

Key elements	Inhibit/Activate apoptosis	Drug-target suitability	Comparison with animals or humans	References
AIF	Activate	Yes	Different	[45,56]
meCAS	Activate	Yes	Fungi specific	[68]
IAPs	Inhibit	Yes	Fungi specific	[82]
IAP-APs	Activate	Yes	Fungi specific	[89]
GSH	Inhibit	No	Lack of specificity	[91]
Sphingolipids	Activate	Yes	Different	[92]

), ArticleFig(id=1227681723211772914, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226554097655001107, language=EN, label=Table 2, caption=

Antifungal drugs for clinical application

, figureFileSmall=null, figureFileBig=null, tableContent=

Drug classification	Drug name	Target	Whether it triggers apoptosis	References
Polyenes	Nystatin	Mainly binds ergosterol	Indirect triggering	[99]
	Amphotericin
Echinocandins	Caspofungin	Inhibition of glucan synthesis	Indirect triggering	[100]
	Mycamine
	Anidulafungin
Pyrimidines	5-flucytosine	Interference with DNA synthesis	No report	[101]
	5-fluorouracil
Azole (imidazole)	Ketoconazole	Inhibition of ergosterol synthesis	Some indirectly trigger mitochondrial pathway apoptosis	[102]
	Clotrimazole
Azole (triazoles)	Itraconazole
	Voriconazole
	Posaconazole
Allylamines	Naftifine	Inhibition of ergosterol synthesis	No report	[103]
	Terbinafine
	Butenafine

), ArticleFig(id=1227681723329213440, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226554097655001107, language=CN, label=表2, caption=

临床应用的抗真菌药物

, figureFileSmall=null, figureFileBig=null, tableContent=

Drug classification	Drug name	Target	Whether it triggers apoptosis	References
Polyenes	Nystatin	Mainly binds ergosterol	Indirect triggering	[99]
	Amphotericin
Echinocandins	Caspofungin	Inhibition of glucan synthesis	Indirect triggering	[100]
	Mycamine
	Anidulafungin
Pyrimidines	5-flucytosine	Interference with DNA synthesis	No report	[101]
	5-fluorouracil
Azole (imidazole)	Ketoconazole	Inhibition of ergosterol synthesis	Some indirectly trigger mitochondrial pathway apoptosis	[102]
	Clotrimazole
Azole (triazoles)	Itraconazole
	Voriconazole
	Posaconazole
Allylamines	Naftifine	Inhibition of ergosterol synthesis	No report	[103]
	Terbinafine
	Butenafine

), ArticleFig(id=1227681723421487116, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226554097655001107, language=EN, label=Table 3, caption=

Natural drugs that induce apoptosis-like cell death in fungal cells

, figureFileSmall=null, figureFileBig=null, tableContent=

Natural drugs	Source	Target fungi	Reference
Microbial products
C17 fengycin B	Bacillus subtilis	Fusarium oxysporum	[16]
Farnesol	Various fungi	Candida albicans	[54]
Lipopeptide AF (3)	Bacillus subtilis	Candida albicans	[112]
Iturins	Bacillus amyloliquefaciens	Aspergillus niger	[113]
Surfactin	Bacillus velezensis	Fusariumconcentricum	[114]
Lipid 7 peptide mixture	Neobacillus drentensis	Candida albicans	[115]
2-chloro-1,3-dimethoxy-5-methylbenzene	Hericium erinaceus	Candida albicans	[116]
Statins	Aspergillus terreus	Rhizopus oryzae	[117]
Botanical products
Monoterpene nerol	Flores aurantii	Ceratocystis fimbriata	[13]
Anacardic acid	Cashew nut	Magnaporthe oryzae	[19]
Silymarin	Silybum marianum	Candida albicans	[97]
Coptisine	Coptis chinensis	Cryptococcus neoformans	[125]
α-tomatine	Lycopersico esculentum	Fusarium oxysporum	[126]
Chlorogenic acid	Various plants	Candida albicans	[128]
Lauric acid	Galla chinensis	Rhizoctonia solani	[129]
Cinnamaldehyde	Cinnamon	Candida albicans	[130]
Thyme essential oil	Thymus mongolicus	Aspergillus flavus	[131]
Eugenol	Clove	Candida albicans	[134]
Artemisia argyi essential oil	Artemisia argyi	Candida albicans	[137]
Antimycotic peptide	Satureja khuzistanica	Aspergillus fumigatus	[138]
Purpurin	Madder root	Candida dubliniensis	[139]
Andrographolide	Andrographis paniculata	Candida albicans	[140]
Animal products
Melittin	European bee venom	Aspergillus, Botrytis, Candida	[141]
Scopoletin	Centipede	Fungi	[142]

), ArticleFig(id=1227681726726598685, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226554097655001107, language=CN, label=表3, caption=

诱导真菌细胞凋亡样细胞死亡的天然药物

, figureFileSmall=null, figureFileBig=null, tableContent=

Natural drugs	Source	Target fungi	Reference
Microbial products
C17 fengycin B	Bacillus subtilis	Fusarium oxysporum	[16]
Farnesol	Various fungi	Candida albicans	[54]
Lipopeptide AF (3)	Bacillus subtilis	Candida albicans	[112]
Iturins	Bacillus amyloliquefaciens	Aspergillus niger	[113]
Surfactin	Bacillus velezensis	Fusariumconcentricum	[114]
Lipid 7 peptide mixture	Neobacillus drentensis	Candida albicans	[115]
2-chloro-1,3-dimethoxy-5-methylbenzene	Hericium erinaceus	Candida albicans	[116]
Statins	Aspergillus terreus	Rhizopus oryzae	[117]
Botanical products
Monoterpene nerol	Flores aurantii	Ceratocystis fimbriata	[13]
Anacardic acid	Cashew nut	Magnaporthe oryzae	[19]
Silymarin	Silybum marianum	Candida albicans	[97]
Coptisine	Coptis chinensis	Cryptococcus neoformans	[125]
α-tomatine	Lycopersico esculentum	Fusarium oxysporum	[126]
Chlorogenic acid	Various plants	Candida albicans	[128]
Lauric acid	Galla chinensis	Rhizoctonia solani	[129]
Cinnamaldehyde	Cinnamon	Candida albicans	[130]
Thyme essential oil	Thymus mongolicus	Aspergillus flavus	[131]
Eugenol	Clove	Candida albicans	[134]
Artemisia argyi essential oil	Artemisia argyi	Candida albicans	[137]
Antimycotic peptide	Satureja khuzistanica	Aspergillus fumigatus	[138]
Purpurin	Madder root	Candida dubliniensis	[139]
Andrographolide	Andrographis paniculata	Candida albicans	[140]
Animal products
Melittin	European bee venom	Aspergillus, Botrytis, Candida	[141]
Scopoletin	Centipede	Fungi	[142]

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