植物保护学报

引物名称 Primer name	引物序列（5′→3′） Primer sequence （5′→3′）	用途 Application
HcAtg8-F	GGATCCATGAAATTCCAATACAAAGAAGAACAT	DNA克隆 DNA cloning
HcAtg8 -R	GTCGACGTTAATATCCATAAACATTTTCATCAGA
qHcAtg8-F	CAGAGCCGACATTACTGGT	实时荧光定量PCR qRT-PCR
qHcAtg8-R	AATGGTCTGCGGTAACAAC
HcActin-F	CTACCTCACGCCATTCTC
HcActin-R	AGCTTCTCCTTGATGTCAC

引物名称 Primer name	引物序列（5′→3′） Primer sequence （5′→3′）	用途 Application
HcAtg8-F	GGATCCATGAAATTCCAATACAAAGAAGAACAT	DNA克隆 DNA cloning
HcAtg8 -R	GTCGACGTTAATATCCATAAACATTTTCATCAGA
qHcAtg8-F	CAGAGCCGACATTACTGGT	实时荧光定量PCR qRT-PCR
qHcAtg8-R	AATGGTCTGCGGTAACAAC
HcActin-F	CTACCTCACGCCATTCTC
HcActin-R	AGCTTCTCCTTGATGTCAC

美国白蛾自噬基因 HcAtg8 的鉴定及其对沉默 HcV-ATPase A 的响应

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王薄毓 , 王晓洁 , 韩莹 , 王海 , 赵丹 , 王倩 , 姚亚林 , 陆秀君

植物保护学报 | 研究论文 2026,53(2): 459-466

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植物保护学报 |研究论文 2026 , 53 (2) : 459 -466

美国白蛾自噬基因 HcAtg8 的鉴定及其对沉默 HcV-ATPase A 的响应

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王薄毓, 王晓洁, 韩莹, 王海, 赵丹, 王倩, 姚亚林, 陆秀君

作者信息

河北农业大学植物保护学院，保定 071001

通讯作者:

E-mail：luxiujun@hebau.edu.cn

E-mail：yaoyalin@hebau.edu.cn

Identification of the autophagy gene HcAtg8 in fall webworm Hyphantria cunea and its response to HcV-ATPaseA silencing

Boyu Wang, Xiaojie Wang, Ying Han, Hai Wang, Dan Zhao, Qian Wang, Yalin Yao, Xiujun Lu

Affiliations

College of Plant Protection, Hebei Agricultural University, Baoding 071001, Hebei Province, China

出版时间: 2026-04-30 doi: 10.13802/j.cnki.zwbhxb.2026.2025117

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摘要

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为明确美国白蛾Hyphantria cunea调控细胞能量代谢的V型质子ATP酶催化亚基A（V-type proton ATPase catalytic subunit A，V-ATPase A）基因HcV-ATPase A与自噬相关基因（autophagy-related gene，Atg）HcAtg8的级联效应，采用PCR技术克隆HcAtg8基因，分析该基因的生物学信息及系统进化关系，利用原核表达系统体外表达HcAtg8，并进行Western blot检测，通过实时荧光定量PCR（quantitative real-time PCR， qRT-PCR）技术分析美国白蛾不同龄期及不同肠道部位中HcAtg8基因的表达情况，以及沉默HcV-ATPase A后中肠自噬相关组织的病理学变化及HcAtg8基因的表达情况。结果显示：克隆得到的美国白蛾自噬基因HcAtg8编码区为354 bp，编码117个氨基酸，HcAtg8蛋白和家蚕Bombyx mori BmAtg8聚在一个分支，亲缘关系较近。构建得到pMAL-c2X-HcAtg8重组表达质粒，体外原核表达得到大小为56.4 kD的HcAtg8蛋白，诱导8 h后表达量最高。美国白蛾HcAtg8基因在不同龄期和不同肠道部位均有表达，在蛹中和中肠的表达量最高。沉默HcV-ATPase A后，处理组自噬溶酶体数量为11.0个，显著高于对照组的3.5个，自噬溶酶体数量上调3.1倍，处理组脂滴数量为73.8个，显著高于对照组的12.5个，脂滴数量上调5.9倍；沉默HcV-ATPase A后溶酶体酸性环境改变，导致自噬小体及相关代谢产物积累；qRT-PCR结果显示自噬相关基因HcAtg8在24 h时的表达量为对照的6.1倍，在48 h时表达量最高，为对照的375.2倍；在72 h时该基因表达量降低，为对照的2.5倍，与组织切片的结果相符。表明通过沉默HcV-ATPase A导致了HcAtg8的高表达以及自噬小体和脂滴的积累，破坏了中肠细胞的稳态，导致细胞功能障碍或者死亡。

关键词

美国白蛾 / 自噬相关基因 / 基因克隆表达 / 沉默HcV-ATPase A / 中肠组织变化

Abstract

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To elucidate the cascade effect between the V-type proton ATPase catalytic subunit A (V-ATPase A), a key regulator of cellular energy metabolism, and the autophagy-related gene HcAtg8 in the fall webworm Hyphantria cunea, the HcAtg8 gene was cloned by PCR. The bioinformatic characteristics and phylogenetic relationships were analyzed. The HcAtg8 protein was expressed in vitro using a prokaryotic expression system and detected via Western blot. Quantitative real-time PCR (qRT-PCR) was used to analyze the expression of HcAtg8 in different developmental stages and tissues (foregut, midgut, and hindgut). Additionally, after silencing HcV-ATPase A, histopathological changes in midgut tissues related to autophagy and the expression of HcAtg8 were examined. The results showed that the cloned coding region of HcAtg8 was 354 bp in length, encoding 117 amino acids. Phylogenetic analysis indicated that HcAtg8 clustered closely with BmAtg8 from Bombyx mori, suggesting a close evolutionary relationship. A recombinant expression vector pMAL-c2X-HcAtg8 was constructed, and a 56.4 kD HcAtg8 protein was obtained by prokaryotic expression in vitro, with the highest expression level observed after 8 h of induction. HcAtg8 was expressed across all developmental stages and gut tissues, with the highest expression observed in pupae. Among gut tissues, expression was highest in the midgut. After silencing HcV-ATPase A, the number of autophagosomes in the treated group was 11.0, significantly higher than that in the control group (3.5), representing a 3.1-fold increase. The number of lipid droplets in the treated group was 73.8, significantly higher than that in the control group (12.5), representing a 5.9-fold increase. Silencing HcV-ATPase A disrupted the acidic environment of lysosomes, leading to the accumulation of autophagosomes and related metabolites. qRT-PCR results showed that the expression level of HcAtg8 was 6.1-fold higher than that of the control at 24 h, peaked at 375.2-fold at 48 h, and subsequently decreased to 2.5-fold at 72 h, consistent with the histological observations. These results indicate that silencing HcV-ATPase A leads to the upregulation of HcAtg8 expression and the accumulation of autophagosomes and lipid droplets, thereby disrupting midgut cellular homeostasis, and resulting in cellular dysfunction or death.

Key words

Hyphantria cunea / autophagy-related gene / gene cloning and expression / HcV-ATPase A silencing / midgut tissue changes

引用本文

王薄毓, 王晓洁, 韩莹, 王海, 赵丹, 王倩, 姚亚林, 陆秀君. 美国白蛾自噬基因 HcAtg8 的鉴定及其对沉默 HcV-ATPase A 的响应. 植物保护学报, 2026 , 53 (2) : 459 -466 . DOI: 10.13802/j.cnki.zwbhxb.2026.2025117

Boyu Wang, Xiaojie Wang, Ying Han, Hai Wang, Dan Zhao, Qian Wang, Yalin Yao, Xiujun Lu. Identification of the autophagy gene HcAtg8 in fall webworm Hyphantria cunea and its response to HcV-ATPaseA silencing[J]. Journal of Plant Protection, 2026 , 53 (2) : 459 -466 . DOI: 10.13802/j.cnki.zwbhxb.2026.2025117

正文

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美国白蛾Hyphantria cunea属鳞翅目灯蛾科，是世界性检疫害虫，在中国连续多年暴发成灾且为害范围逐年扩大（Ning et al.，2022）。该害虫寄主范围广、食性杂、抗逆性强，以幼虫群集结网为害树叶并取食叶肉，残留表皮枯萎，严重时可造成整颗植株死亡（杨忠岐，2004；刘丹等，2023）。生产上防治美国白蛾多采用化学防治和生物防治相结合的方法（杨忠岐等，2018；Edosa et al.，2019）。鉴定新型靶标基因是生物防治新领域，利用靶标基因防控美国白蛾是目前研究的重点、热点和难点。目前已鉴定得到影响美国白蛾代谢的V型质子ATP酶催化亚基A（V-type protein ATPase catalytic subunit A，V-ATPase A）基因HcV-ATPase A、6-磷酸果糖谷氨酰胺氨基转移酶（glutamine：fructose-6-phosphate aminotransferase，GFAT）基因HcGFAT和蜕皮激素诱导蛋白E75基因HcE75，利用其开发新型核酸纳米农药将为美国白蛾防控提供新途径（Zhang et al.，2022；Zou et al.，2022；Wang et al.，2024）。

自噬是真核生物中普遍存在并高度保守的生理过程，是程序化细胞死亡的一种类型，对于维持细胞稳态和避免细胞衰老至关重要（Mizushima et al.，2008；Shelly et al.，2009；刘勇等，2024）。自噬通过形成具有双层膜的自噬体，将细胞中错误折叠的蛋白质、受损的细胞器运送到溶酶体降解后进行循环利用（Kuo et al.，2018）。自噬相关基因（autophagy-related gene，Atg）所编码的蛋白ATG参与自噬体的形成和选择性自噬的调控（Sheng & Qin，2019），其中ATG8蛋白是参与整个自噬过程的重要标志蛋白，也是自噬体膜形成的重要标志（Mizushima et al.，2011）。溶酶体腔内的酸性环境是其发挥活性的关键，腔内酸性环境主要依靠液泡型ATP酶V-ATPase维持。研究表明自噬活性影响昆虫生长发育，如自噬抑制剂作用于蜜蜂幼虫后，幼虫自噬活性受到显著抑制，进而导致蛹的死亡（陈文凤，2023）；沉默褐飞虱Nilaparvata lugens Atg13基因后，其存活率显著降低，组织ATP含量显著降低（吴建艮等，2022）。

V-ATPase是一种普遍存在的多亚基复合物，由可溶性V1复合物和膜相关V0复合物组成（王梦珂等，2020）。V1复合物的A亚基可以催化ATP水解（Nishi & Forgac，2002；Forgac，2007）。V-ATPase依靠ATP水解释放能量，调控蛋白质转运、受体介导内吞与pH平衡（Breton & Brown，2013）；同时可利用水解ATP产生的动力完成质子跨膜运输，是维持细胞离子平衡的核心功能酶。在拟南芥Arabidopsis thaliana中，V-ATPase特异性抑制剂Concanamycin A能抑制内吞转运、分泌途径以及自噬小体降解，表明V-ATPase在植物自噬过程中起着非常重要的作用（蔺姗，2019）。在人和模型动物脑内发现V-ATPase的部分亚基下调会影响溶酶体酸化功能和自噬溶酶体的降解（Gao et al.，2011；Chen et al.，2018）。抑制V-ATPase与组织蛋白酶D的活性会导致水解酶功能异常并引起自噬阻滞，进而产生神经毒性（韩解，2022）。本实验室前期研究发现，沉默美国白蛾HcV-ATPase A基因后，自噬小体增多（Wang et al.，2024）。基于此，本研究利用美国白蛾转录组数据筛选得到自噬相关基因HcAtg8，通过克隆HcAtg8基因的编码区分析其编码蛋白的序列结构和系统进化关系，分析HcAtg8基因在不同龄期和肠道中的表达水平，以及沉默HcV-ATPase A后中肠细胞自噬相关组织的病理学变化及HcAtg8基因的表达趋势，以期为明确美国白蛾HcV-ATPase A与自噬间的相互作用提供依据。

1 材料与方法

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1.1 材料

供试昆虫、蛋白：美国白蛾卵购于中国林业科学研究院森林生态环境与保护研究所昆虫病毒研发中心，于河北农业大学害虫生物防治实验室、昆虫病理与分子生物学实验室内人工饲养，饲养至3龄幼虫时供试，饲养条件为温度（26±1） ℃、相对湿度（60±5）%、光周期16 L∶8 D。绿色荧光蛋白（green fluorescent protein，gfp）为本课题组保存并提供。

供试培养基：Luria-Bertani（LB）液体培养基成分为胰蛋白胨10 g、酵母提取物5 g、NaCl 10 g、ddH₂O定容至1 L；加入12 g琼脂即为LB固体培养基。

试剂及仪器：大肠杆菌Escherichia coli DH5α、BL21（DE3）及质粒pMAL-c2X由河北农业大学害虫生物防治实验室保存并提供；磷酸盐缓冲液（phosphate buffered saline，PBS）、PVDF膜、环氧树脂812、限制性内切酶BamH I、Sal I、T4 DNA Ligase、高保真酶Primer STAR^® Max DNA Polymerase、反转录试剂盒cDNA Synthesis Kit、荧光定量酶SYBR^® Premix Ex Taq TM II，宝生物工程（大连）有限公司；异丙基-β-D-硫代半乳糖苷（Isopropyl β-D-1-thiogalactopyranoside，IPTG）、羊抗鼠抗体IgG-HRP、动物组织总RNA提取试剂盒，天根生化科技（北京）有限公司；通用型DNA纯化回收试剂盒、鼠抗Anti-MBP抗体、RIPA裂解液，生工生物工程（上海）股份有限公司；其他试剂均为国产分析纯。Mastercycler Gradient PCR仪，德国Eppendorf公司；DYCP-32B琼脂糖凝胶电泳系统，北京六一仪器厂；Mini protein III蛋白质电泳系统装置，美国Bio-Rad公司；HT7800透射电子显微镜，日立高新技术公司。

1.2 方法

1.2.1 美国白蛾总RNA提取及第一链cDNA的合成

分别收集美国白蛾卵、1~6龄幼虫、蛹及成虫的健康试虫，每10头（粒）为1个重复，共3次重复；同时分别收集美国白蛾3龄幼虫的前肠、中肠、后肠组织，每10头为1个重复，共3次重复，于-80 ℃保存备用。参照动物组织总RNA提取试剂盒说明书提取各发育阶段以及各肠道组织样品的RNA，参照反转录试剂盒cDNA Synthesis Kit说明书反转录获得第一链cDNA备用。

1.2.2 目的基因的鉴定、扩增和克隆测序

利用本实验室前期得到的美国白蛾转录组数据进行筛选，利用LTFViewr5u软件解析数据库序列，进一步利用NCBI上已知Atg8序列进行全长序列比对，获得HcAtg8基因序列，利用DNAMAN 9软件对分析所得的HcAtg8基因序列进行全长引物设计（表1）。以3龄幼虫cDNA为模板进行PCR扩增，25 μL PCR扩增体系：cDNA 1.0 μL、10 μmol/L上下游引物各0.5 μL、2×PrimerStar Max Mix 12.5 μL、ddH₂O 10.5 μL。扩增程序：95 ℃预变性3 min；95 ℃变性45 s，55 ℃复性45 s，72 ℃延伸90 s，35个循环；72 ℃延伸10 min。利用通用型DNA纯化回收试剂盒纯化回收PCR产物。回收产物与pMAL-c2X质粒连接，连接转化至大肠杆菌DH5α感受态细胞，次日挑取白色单菌落进行菌液PCR，反应程序同上，之后经0.8%琼脂糖凝胶电泳检测正确后，提取重组质粒pMAL-c2X-HcAtg8，使用BamH I、Sal I进行单双酶切验证，委托生工生物工程（上海）股份有限公司测序。

1.2.3 美国白蛾HcAtg8基因的原核表达

将测序正确的pMAL-c2X-HcAtg8表达质粒转化至大肠杆菌BL21（DE3）感受态细胞，在含有氨苄西林的LB固体培养基平板上培养16 h时挑选单菌落，并进行PCR验证（Wang et al.，2024）。将重组菌接入10 mL的LB液体培养基中过夜培养，次日转接220 r/min继续振荡培养至OD_{600 nm}为0.65时，加入母液浓度为1 mol/L的IPTG 6 μL，于37 ℃分别诱导2、4、6和8 h后取样1 mL，以未经IPTG诱导的菌液作为阴性对照。离心收集菌沉淀，加入1×PBS重悬沉淀，加入5×Loading Buffer混匀，煮沸10 min，以10%十二烷基硫酸钠-聚丙烯酰胺凝胶电泳（sodium dodecyl sulfate-polyacrylamide gel electrophoresis，SDS-PAGE）分离HcAtg8蛋白，将该蛋白转移至PVDF膜，以小鼠来源的Anti-MBP抗体为一抗，碱性磷酸酶标记的羊抗鼠抗体IgG-HRP为二抗，进行Western blot分析。对蛋白进行纯化并送至河北省生物研究所制备特异抗体。取2头美国白蛾蛹，加入RIPA裂解液后置于冰上进行充分匀浆。4 ℃、12 000 r/min离心15 min，吸取上清液即为总蛋白样品，以特异性抗体为一抗，采用Western blot检测目标蛋白的表达情况。

1.2.4 HcAtg8的生物信息学分析

利用DNAMAN V6.0软件对HcAtg8基因序列的开放阅读框、编码的蛋白分子量等进行预测，利用WOLF PSORT在线软件预测其亚细胞定位，利用ExPASy Proteomics Server在线软件预测其编码蛋白的理化性质，利用SignalP 4.1 Server、NetOGlyc 4.1 Server和NetNGlyc1.0 Server等网站分别对其编码蛋白的信号肽、O-糖基化位点和N-糖基化位点等进行分析，利用SOPMA软件对其编码蛋白的二级结构进行分析。将筛选得到的序列在NCBI上进行BLAST比对分析，下载近缘物种的Atg8的氨基酸序列，利用MEGA 7软件采用邻接法构建系统进化树，Bootstrap设1 000次重复检验。

1.2.5 qRT-PCR检测HcAtg8的时空表达

利用DNAMAN软件设计HcAtg8及内参基因HcActin的实时荧光定量PCR（quantitative real-time PCR，qRT-PCR）特异性引物（表1），以1.2.1中获得的cDNA模板，参照荧光定量试剂盒说明书进行qRT-PCR，反应体系以及反应程序参考Wang et al.（2024）方法。每处理3次重复，反应结束后记录Ct值。以参照样品的表达量为标准参量，利用2^-ΔΔ^Ct 法计算该基因在美国白蛾不同龄期以及肠道组织中的相对表达量，计算基因沉默效率（Wang et al.，2024）。基因沉默效率=（1-处理组靶基因相对表达量/对照组靶基因相对表达量）×100%。

1.2.6 沉默HcV-ATPase A对HcAtg8表达及自噬的影响

利用Wang et al.（2024）方法合成dsV-ATPase A和dsgfp，采用注射法沉默美国白蛾3龄幼虫HcV-ATPase A基因获得试虫，以注射dsgfp作对照，参考1.2.1方法获得沉默HcV-ATPase A基因0、24、48和72 h后虫体的cDNA，以cDNA为模板进行qPT-PCR测定HcAtg8表达水平，反应体系及反应程序同1.2.5。同时，收集沉默HcV-ATPase A 72 h后的美国白蛾幼虫3头，于冰上解剖分离中肠组织，经过固定液固定、脱色、包埋、切片、染色后，将样本在包埋剂中包埋2 h，使用超微切片机切成60~80 nm厚的切片，并用透射电子显微镜观察脂滴与自噬溶酶体的数量变化，每头幼虫观察3张切片，每个切片5个视野，共3个重复。

1.3 数据分析

试验数据利用SPSS 20.0软件进行统计分析，采用单因素方差分析，用Tukey氏多重比较法进行差异显著性检验。

2 结果与分析

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2.1 HcAtg8 的克隆及其序列分析

HcAtg8序列全长为354 bp（图1），编码117个氨基酸，蛋白质分子量为13.98 kD，等电点为9.21，表明该蛋白质为碱性蛋白质。经亚细胞定位分析，发现该蛋白可能定位于细胞核。通过氨基酸序列对比筛选并构建系统发育树，结果表明美国白蛾HcAtg8序列与家蚕BmAtg8亲缘关系最近（图2）。

HcAtg8蛋白化学分子式为C₄₉₉H₇₇₃N₁₄₉O₁₅₁S₆，总原子数为1 578个，进一步分析发现其不稳定系数为59.27，表明该蛋白为不稳定蛋白；其平均亲水系数为-0.508，表明该蛋白是亲水性蛋白；无糖基化位点，无信号肽结构，该蛋白二级结构中α螺旋占10%，β折叠占21%，无规则卷曲占69%。

2.2 HcAtg8 的原核表达

重组质粒pMAL-c2X-HcAtg8经BamH I单酶切后产生大小为7 000 bp的目的条带，经BamH I和Sal l双酶切后，产生6 646 bp的载体条带和354 bp的目的基因条带（图3-A），说明重组载体pMAL-c2X-HcAtg8构建成功。含有pMAL-c2X-HcAtg8的菌株BL21（DE3）经IPTG诱导后表达大小为56.4 kD的重组蛋白，且在8 h时蛋白表达量最高（图3-B~C）。Western blot检测目标蛋白特异性抗体与目的条带发生特异性结合，表明该蛋白在虫体中存在表达（图3-D）。

2.3 美国白蛾 HcAtg8 的时空定位

美国白蛾HcAtg8在各个龄期及肠道中均有表达，在蛹中表达量最高，其次是成虫，在卵及1~3龄幼虫中表达稳定，整体呈现稳定表达、急剧减少、逐步升高趋势（图4-A）；在中肠中表达量最高（图4-B，P<0.05）。

2.4 沉默 HcV-ATPase A 后对自噬的影响

2.4.1 沉默 HcV-ATPase A后自噬超微结构组织变化

沉默HcV-ATPase A后处理组自噬溶酶体数量为11.0个，显著高于对照组的3.5个，自噬溶酶体上调3.1倍；处理组脂滴数量为73.8个，显著高于对照组的12.5个，脂滴上调5.9倍（P<0.05，图5）。

2.4.2 沉默HcV-ATPase A后不同时间HcAtg8的表达水平

沉默HcV-ATPase A 72 h内美国白蛾体内HcAtg8基因的表达量均比对照高，且差异显著，整体呈现表达量先升高后降低的趋势，在24 h表达量为对照的6.1倍，在48 h时表达量最高，为对照的375.2倍，在72 h时该基因表达量降低，为对照的2.5倍（图6）。

3 讨论

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自噬是一种细胞内的防御和应激调控机制，能快速清除细胞内受损的细胞器、错误折叠的蛋白质、入侵的病原体，并将降解产物循环再利用，以此维持细胞内环境稳态并应对营养匮乏、氧化应激等不利生存条件。ATG8是自噬研究中监测自噬小体和自噬活性的重要标志物（Qiao et al.，2018）。何珊等（2023）研究发现，白纹伊蚊Aedes albopictus ATG8氨基酸序列同家蚕的相似性为94.2%，扩增片段为357 bp。本研究利用美国白蛾转录组数据筛选并克隆鉴定到美国白蛾HcAtg8基因，系统进化树发育分析结果也显示HcAtg8与家蚕的BmAtg8亲缘关系最近。

不同昆虫的Atg基因具有物种特异性，白纹伊蚊AaAtg8在各发育阶段均有表达，在蛹期表达量最高，这表明AaAtg8可能在发育中起重要作用（Qiao et al.，2018）。在本研究中，HcAtg8在美国白蛾所有发育阶段均有表达，在蛹中表达量最高，与 Qiao et al.（2018）研究的白纹伊蚊Atg8在蛹期表达量最高的结果一致。在黄粉虫Tenebrio molitor的整个发育过程中，TmATG8的相对表达量在中肠中较高（Tettamanti et al.，2007）；在大蜡螟Galleria mellonella中，GmATG8在中肠、卵巢、马氏小管、脂肪体和丝腺中均检测到表达，并且发现表达量最高的组织是在幼虫中肠（Khoa & Takeda，2012）；本研究也获得类似结果，组织特异性结果显示，HcAtg8的表达量在美国白蛾幼虫的中肠最高。

溶酶体的质子泵V-ATP酶负责建立酶活化和货物降解所必需的内腔低pH，V-ATP酶参与溶酶体酸化、溶酶体酶活化和自噬特异性物质降解（Mauvezin & Neufeld，2012）。ATP6V0A1/a1亚基的敲低会抑制细胞的溶酶体酸化（Lee et al.，2015）。ATP6V0A1/a1亚基错义变异体可损害Neuro-2a细胞系的溶酶体酸化和自噬体积累（Aoto et al.，2021）。本研究沉默HcV-ATPase A后，随着溶酶体酸性环境改变，溶酶体降解自噬小体活性下降，推测可能自噬体-溶酶体降解受阻，导致自噬小体及相关代谢产物积累，影响昆虫稳态，进而发挥杀虫作用。研究结果将为进一步探究HcV-ATPase A与美国白蛾细胞自噬反应的相互作用机理提供支撑，也将为阐明HcAtg8基因的功能及作用机制奠定基础。

基金

收起

河北省中央引导地方项目(246Z6507G)
河北省现代农业产业技术体系项目(HBCT2024190208)
河北省现代农业产业技术体系项目(HBCT2024130204)
石家庄市驻冀高校重点研发专项(241490102A)

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2026年第53卷第2期

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doi: 10.13802/j.cnki.zwbhxb.2026.2025117

接收时间：2025-08-15
首发时间：2026-05-27
出版时间：2026-04-30

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收稿日期：2025-08-15

基金

河北省中央引导地方项目(246Z6507G)

河北省现代农业产业技术体系项目(HBCT2024190208)

河北省现代农业产业技术体系项目(HBCT2024130204)

石家庄市驻冀高校重点研发专项(241490102A)

作者信息

河北农业大学植物保护学院，保定 071001

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E-mail：luxiujun@hebau.edu.cn

E-mail：yaoyalin@hebau.edu.cn

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https://castjournals.cast.org.cn/joweb/zwbhxb/CN/10.13802/j.cnki.zwbhxb.2026.2025117

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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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