Article(id=1266470629487301058, tenantId=1146029695717560320, journalId=1266358857061122103, issueId=1266470523241382909, articleNumber=null, orderNo=null, doi=10.13802/j.cnki.zwbhxb.2026.2025103, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1753286400000, receivedDateStr=2025-07-24, revisedDate=null, revisedDateStr=null, acceptedDate=null, acceptedDateStr=null, onlineDate=1779879727956, onlineDateStr=2026-05-27, pubDate=1777478400000, pubDateStr=2026-04-30, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1779879727956, onlineIssueDateStr=2026-05-27, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1779879727956, creator=13701087609, updateTime=1779879727956, updator=13701087609, issue=Issue{id=1266470523241382909, tenantId=1146029695717560320, journalId=1266358857061122103, year='2026', volume='53', issue='2', pageStart='301', pageEnd='586', issueExtLink='null', onlineDate='null', pubDate='1777478400000', pubDateStr='2026-04-30', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=1, specialIssue=null, createTime=1779879702622, creator='13701087609', updateTime=1782266106964, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext={EN=IssueExt(id=1276479828593349443, tenantId=1146029695717560320, journalId=1266358857061122103, issueId=1266470523241382909, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1276479828593349444, tenantId=1146029695717560320, journalId=1266358857061122103, issueId=1266470523241382909, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null, downloadFileDto=null}, startPage=449, endPage=458, ext={EN=ArticleExt(id=1266470630393270724, articleId=1266470629487301058, tenantId=1146029695717560320, journalId=1266358857061122103, language=EN, title=Genetic analysis of Asiatic rice borer Chilo suppressalis populations in Ningbo City based on whole-genome resequencing, columnId=1266470561661206635, journalTitle=Journal of Plant Protection, columnName=Research reports, runingTitle=null, highlight=null, articleAbstract=

To investigate the population structure and phylogenetic relationships among different geographic populations of Asiatic rice borer Chilo suppressalis in Ningbo City, China,whole-genome resequencing was performed on 37 individuals from eight populations (six from Ningbo, one from Beijing, and one from Hangzhou). Genomic variations were identified, and population genetic analyses including population structure, selective sweeps, genetic diversity, and historical effective population size were conducted. The results showed that a total of 42 149 215 single nucleotide polymorphisms and 11 912 812 insertions/deletions were identified across the 37 genomes, with most variations located in intergenic and intronic regions. Among the six Ningbo populations, Yinzhou, Yuyao and Haishu populations were closely related, whereas the Xiangshan population showed the greatest genetic distance from the other Ningbo populations and exhibited moderate genetic differentiation. Genes within the selective sweep regions of the Xiangshan population was significantly enriched in seven signaling pathways, including melanogenesis and circadian entrainment. Among the Ningbo populations, nucleotide diversity was relatively higher in the Haishu and Yinzhou populations, but lower in the Xiangshan and Fenghua populations. The Ningbo populations experienced a rapid contraction approximately 3 000—1 000 years ago, followed by a rapid population expansion.

, authors=null, authorsList=Shan Xiao, Weigang Chai, Jian Liu, Jianghua Chen, authorCompany=null, correspAuthors=null, authorNote=null, correspAuthorsNote=null, copyrightStatement=null, copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=null, magXml=null, pdfUrl=null, pdf=null, pdfFileSize=null, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=null, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=null, mapNumber=null, fund=null), CN=ArticleExt(id=1266470634977645015, articleId=1266470629487301058, tenantId=1146029695717560320, journalId=1266358857061122103, language=CN, title=基于全基因组重测序的宁波市二化螟种群遗传分析, columnId=1266470561837367405, journalTitle=植物保护学报, columnName=研究论文, runingTitle=null, highlight=null, articleAbstract=

为解析宁波市二化螟Chilo suppressalis不同地理种群的遗传结构以及种群间亲缘关系,对8个二化螟种群(6个宁波市地理种群、1个北京种群和1个杭州种群)的37个样品进行全基因组重测序,鉴定基因组间遗传变异,并进行种群遗传结构、基因组选择性清除、遗传多样性、历史有效种群大小等种群遗传学分析。结果显示:在37个二化螟个体基因组中共鉴定到42 149 215个单核苷酸多态性和11 912 812个插入缺失,这些变异均主要分布在基因间和基因内含子区域。在宁波市6个二化螟种群中,鄞州、余姚、海曙种群的亲缘关系较近,象山种群与宁波市其他5个种群的亲缘关系最远,并表现出中度的遗传分化。二化螟象山种群基因组选择性清除区域内的基因显著富集在黑色素生成、昼夜节律夹带等7个信号通路。在宁波市各种群中,海曙种群和鄞州种群的核苷酸多样性较高,象山种群和奉化种群的核苷酸多样性较低。宁波市二化螟种群在3 000年前—1 000年前左右经历了快速收缩,随后种群规模快速增加。

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Genomes of the cosmopolitan fruit pest Bactrocera dorsalis (Diptera: Tephritidae) reveal its global invasion history and thermal adaptation. Journal of Advanced Research, 53: 61-74, articleTitle=Genomes of the cosmopolitan fruit pest Bactrocera dorsalis (Diptera: Tephritidae) reveal its global invasion history and thermal adaptation, refAbstract=null), Reference(id=1266746491617042887, tenantId=1146029695717560320, journalId=1266358857061122103, articleId=1266470629487301058, doi=null, pmid=null, pmcid=null, year=2019, volume=20, issue=5, pageStart=1064, pageEnd=null, url=null, language=null, rfNumber=null, rfOrder=35, authorNames=Zhao J, Xu L, Sun Y, Song PP, Han ZJ, journalName=International Journal of Molecular Sciences, refType=null, unstructuredReference=Zhao J, Xu L, Sun Y, Song PP, Han ZJ. 2019. UDP-glycosyltransferase genes in the striped rice stem borer, Chilo suppressalis (Walker), and their contribution to chlorantraniliprole resistance. International Journal of Molecular Sciences, 20(5): 1064, articleTitle=UDP-glycosyltransferase genes in the striped rice stem borer, Chilo suppressalis (Walker), and their contribution to chlorantraniliprole resistance, refAbstract=null), Reference(id=1266746491675763144, tenantId=1146029695717560320, journalId=1266358857061122103, articleId=1266470629487301058, doi=null, pmid=null, pmcid=null, year=2019, volume=36, issue=7, pageStart=969, pageEnd=978, url=null, language=null, rfNumber=null, rfOrder=36, authorNames=Zhu L, Feng S, Gao Q, Liu W, Ma WH, Wang XP, journalName=Chronobiology International, refType=null, unstructuredReference=Zhu L, Feng S, Gao Q, Liu W, Ma WH, Wang XP. 2019. Host population related variations in circadian clock gene sequences and expression patterns in Chilo suppressalis . Chronobiology International, 36(7): 969-978, articleTitle=Host population related variations in circadian clock gene sequences and expression patterns in Chilo suppressalis, refAbstract=null), Reference(id=1266746491763843529, tenantId=1146029695717560320, journalId=1266358857061122103, articleId=1266470629487301058, doi=null, pmid=null, pmcid=null, year=2024, volume=40, issue=5, pageStart=1207, pageEnd=1213, url=null, language=null, rfNumber=null, rfOrder=37, authorNames=Zhuo FY, Chen XX, Xia YX, Fu Q, Wang S, Xu HX, Hu F, Zhang J, journalName=Chinese Journal of Biological Control, refType=null, unstructuredReference=Zhuo FY, Chen XX, Xia YX, Fu Q, Wang S, Xu HX, Hu F, Zhang J. 2024. The occurrence characteristics of rice diseases and insect pests and the integration of green control technology in China from 2013 to 2022. Chinese Journal of Biological Control, 40(5): 1207-1213 (in Chinese), articleTitle=null, refAbstract=null), Reference(id=1266746491860312522, tenantId=1146029695717560320, journalId=1266358857061122103, articleId=1266470629487301058, doi=null, pmid=null, pmcid=null, year=2024, volume=40, issue=5, pageStart=1207, pageEnd=1213, url=null, language=null, rfNumber=null, rfOrder=38, authorNames=卓富彦, 陈学新, 夏玉先, 傅强, 王甦, 徐红星, 胡飞, 张杰, journalName=中国生物防治学报, refType=null, unstructuredReference=卓富彦, 陈学新, 夏玉先, 傅强, 王甦, 徐红星, 胡飞, 张杰. 2024. 2013—2022年我国水稻病虫害发生特点与绿色防控技术集成. 中国生物防治学报, 40(5): 1207-1213, articleTitle=2013—2022年我国水稻病虫害发生特点与绿色防控技术集成, refAbstract=null)], funds=[Fund(id=1266746487879917986, tenantId=1146029695717560320, journalId=1266358857061122103, articleId=1266470629487301058, awardId=2023S014, language=CN, fundingSource=宁波市公益性科技计划项目(2023S014), fundOrder=null, country=null), Fund(id=1266746487993164195, tenantId=1146029695717560320, journalId=1266358857061122103, articleId=1266470629487301058, awardId=2024NKYP006, language=CN, fundingSource=宁波市农业科学研究院院长基金项目(2024NKYP006), fundOrder=null, country=null)], companyList=[AuthorCompany(id=1266746478941856105, tenantId=1146029695717560320, journalId=1266358857061122103, articleId=1266470629487301058, xref=null, ext=[AuthorCompanyExt(id=1266746478950244714, tenantId=1146029695717560320, journalId=1266358857061122103, articleId=1266470629487301058, companyId=1266746478941856105, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=Ningbo Key Laboratory of Testing and Control for Characteristic Agro-Product Quality and Safety, Ningbo Academy ofAgricultural Sciences, Ningbo 315040, Zhejiang Province, China), AuthorCompanyExt(id=1266746478958633323, tenantId=1146029695717560320, journalId=1266358857061122103, articleId=1266470629487301058, companyId=1266746478941856105, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=宁波市农业科学研究院,宁波市特色农产品质量安全检测与控制重点实验室,浙江 宁波 315040)])], figs=[ArticleFig(id=1266746485916983692, tenantId=1146029695717560320, journalId=1266358857061122103, articleId=1266470629487301058, language=EN, label=Fig. 1, caption=Density distribution of single nucleotide polymorphisms (A) and insertions/deletions (B)across 29 chromosomes of Chilo suppressalis, figureFileSmall=+iT62VY1d1NRphPYW4mFUw==, figureFileBig=yp8xe9PDK9fBsPlcQnyUqw==, tableContent=null), ArticleFig(id=1266746486026035597, tenantId=1146029695717560320, journalId=1266358857061122103, articleId=1266470629487301058, language=CN, label=图1, caption=二化螟29条染色体上单核苷酸多态性(A)和插入缺失(B)的密度分布图, figureFileSmall=+iT62VY1d1NRphPYW4mFUw==, figureFileBig=yp8xe9PDK9fBsPlcQnyUqw==, tableContent=null), ArticleFig(id=1266746486319636878, tenantId=1146029695717560320, journalId=1266358857061122103, articleId=1266470629487301058, language=EN, label=Fig. 2, caption=Genetic structure analysis of eight Chilo suppressalis populations, figureFileSmall=hAmkRZd+zsxxBEKSWgGD2A==, figureFileBig=riDFl7HgA5C5mVrja7O6HQ==, tableContent=null), ArticleFig(id=1266746486416105871, tenantId=1146029695717560320, journalId=1266358857061122103, articleId=1266470629487301058, language=CN, label=图2, caption=8个二化螟种群的遗传结构分析

A:8个二化螟种群的交叉验证误差;B:祖先个数K为2~4时二化螟的种群结构。BJ:北京种群;YZ:鄞州种群;FH:奉化种群;FY:富阳种群;HS-JSG:海曙集士港种群;HS-GL:海曙古林种群;YY:余姚种群;XS:象山种群。A:Cross-validation errors of eight Chilo suppressalis populations; B: population structure of C. suppressalisp with ancestral population K=2-4. BJ: Beijing population; YZ:Yinzhou population; FH: Fenghua population; FY: Fuyang population; HS-JSG: Haishu-Jishigang population; HS-GL: Haishu-Gulin population; YY: Yuyao population; XS: Xiangshan population.

, figureFileSmall=hAmkRZd+zsxxBEKSWgGD2A==, figureFileBig=riDFl7HgA5C5mVrja7O6HQ==, tableContent=null), ArticleFig(id=1266746486504186256, tenantId=1146029695717560320, journalId=1266358857061122103, articleId=1266470629487301058, language=EN, label=Fig. 3, caption=Principal component analysis of 37 Chilo suppressalis samples, figureFileSmall=lbDnILO5luCqlXPi3izzig==, figureFileBig=yNTWKfOndFKxWQaGb3MHTg==, tableContent=null), ArticleFig(id=1266746486567100817, tenantId=1146029695717560320, journalId=1266358857061122103, articleId=1266470629487301058, language=CN, label=图3, caption=37个二化螟样品的主成分分析, figureFileSmall=lbDnILO5luCqlXPi3izzig==, figureFileBig=yNTWKfOndFKxWQaGb3MHTg==, tableContent=null), ArticleFig(id=1266746486630015378, tenantId=1146029695717560320, journalId=1266358857061122103, articleId=1266470629487301058, language=EN, label=Fig. 4, caption=Phylogenetic tree of 37 Chilo suppressalis samples, figureFileSmall=nj1Mdpc0YYhIJ6NH6yHvGg==, figureFileBig=PKQZaU+E7YCip7iwzVXGLQ==, tableContent=null), ArticleFig(id=1266746486692929939, tenantId=1146029695717560320, journalId=1266358857061122103, articleId=1266470629487301058, language=CN, label=图4, caption=37个二化螟样品的系统发育树

BJ1~BJ5:北京种群;YZ1~YZ5:鄞州种群;FH1~FH3:奉化种群;FY1~FY5:富阳种群;HS-JSG1~HS-JSG5:海曙集士港种群;HS-GL1~HS-GL5:海曙古林种群;YY1~YY5:余姚种群;XS1~XS4:象山种群。BJ1-BJ5: Beijing population; YZ1-YZ5: Yinzhou population; FH1-FH3: Fenghua population; FY1-FY5: Fuyang population; HS-JSG1-HS-JSG5: Haishu-Jishigang population; HS-GL1-HS-GL5: Haishu-Gulin population; YY1-YY5: Yuyao population; XS1-XS4: Xiangshan population.

, figureFileSmall=nj1Mdpc0YYhIJ6NH6yHvGg==, figureFileBig=PKQZaU+E7YCip7iwzVXGLQ==, tableContent=null), ArticleFig(id=1266746486751650196, tenantId=1146029695717560320, journalId=1266358857061122103, articleId=1266470629487301058, language=EN, label=Fig. 5, caption=Selective sweep analysis of the genome in the Xiangshan population of Chilo suppressalis, figureFileSmall=5oSKkGbFW1xILfdoByiX6g==, figureFileBig=DZLceG7y7gNblx22ldUISQ==, tableContent=null), ArticleFig(id=1266746486831341973, tenantId=1146029695717560320, journalId=1266358857061122103, articleId=1266470629487301058, language=CN, label=图5, caption=二化螟象山种群基因组选择性清除结果

πxs:二化螟象山种群的核苷酸多样性;πother:分别为二化螟鄞州、奉化、富阳、海曙集士港、海曙古林和余姚种群的核苷酸多样性。πxs: Nucleotide diversity of the Xiangshan population of C. suppressalis; πother: nucleotide diversity of the Yinzhou, Fenghua, Fuyang, Haishu-Jishigang, Haishu-Gulin, and Yuyao populations, respectively.

, figureFileSmall=5oSKkGbFW1xILfdoByiX6g==, figureFileBig=DZLceG7y7gNblx22ldUISQ==, tableContent=null), ArticleFig(id=1266746486902645142, tenantId=1146029695717560320, journalId=1266358857061122103, articleId=1266470629487301058, language=EN, label=Fig. 6, caption=KEGG pathway enrichment analysis of selected genes in the Xiangshan population of Chilo suppressalis, figureFileSmall=ggcfS2+vl7H0+jy8T6XpBA==, figureFileBig=ay0P1DOBp0OnG8O6ZRK1rw==, tableContent=null), ArticleFig(id=1266746486965559703, tenantId=1146029695717560320, journalId=1266358857061122103, articleId=1266470629487301058, language=CN, label=图6, caption=二化螟象山种群受选择基因的KEGG通路富集分析, figureFileSmall=ggcfS2+vl7H0+jy8T6XpBA==, figureFileBig=ay0P1DOBp0OnG8O6ZRK1rw==, tableContent=null), ArticleFig(id=1266746487024279960, tenantId=1146029695717560320, journalId=1266358857061122103, articleId=1266470629487301058, language=EN, label=Fig. 7, caption=Historical effective population size of Chilo suppressalis populations in Ningbo City, figureFileSmall=Rbo+RtzkYb/qHSdiDrrqgQ==, figureFileBig=NbmAwyreypiovqgMOmsYGg==, tableContent=null), ArticleFig(id=1266746487095583129, tenantId=1146029695717560320, journalId=1266358857061122103, articleId=1266470629487301058, language=CN, label=图7, caption=宁波市二化螟种群历史有效种群大小, figureFileSmall=Rbo+RtzkYb/qHSdiDrrqgQ==, figureFileBig=NbmAwyreypiovqgMOmsYGg==, tableContent=null), ArticleFig(id=1266746487158497690, tenantId=1146029695717560320, journalId=1266358857061122103, articleId=1266470629487301058, language=EN, label=Table 1, caption=

Sequencing statistics of genome resequencing samples from Chilo suppressalis populations

, figureFileSmall=null, figureFileBig=null, tableContent=

种群

Population

Clean

data/Gb

Q30/%

比对率

Mapping rate/%

测序深度

Sequencing depth

覆盖度Coverage/%
10×
鄞州种群Yinzhou population13.8±1.892.9±0.699.0±0.516.3±2.182.3±0.769.6±3.5
海曙古林种群Haishu-Gulin population14.8±1.592.8±0.999.5±0.117.5±1.883.0±0.672.1±3.3
海曙集士港种群Haishu-Jishigang population15.4±2.692.9±0.799.4±0.118.2±3.183.1±0.872.6±4.6
奉化种群Fenghua population14.7±3.397.1±0.186.9±9.415.4±5.181.9±1.665.1±10.0
象山种群Xiangshan population13.3±1.193.5±0.792.9±11.214.5±0.982.2±0.667.8±4.1
余姚种群Yuyao population13.2±1.594.1±0.998.4±2.115.3±1.982.1±0.868.6±4.8
富阳种群Fuyang population12.5±0.797.0±0.299.1±0.114.7±0.881.9±0.466.9±2.4
北京种群Beijing Population14.7±2.697.0±0.199.1±0.317.3±3.282.0±0.573.4±3.4
), ArticleFig(id=1266746487271743899, tenantId=1146029695717560320, journalId=1266358857061122103, articleId=1266470629487301058, language=CN, label=表1, caption=

二化螟种群基因组重测序样本测序信息统计

, figureFileSmall=null, figureFileBig=null, tableContent=

种群

Population

Clean

data/Gb

Q30/%

比对率

Mapping rate/%

测序深度

Sequencing depth

覆盖度Coverage/%
10×
鄞州种群Yinzhou population13.8±1.892.9±0.699.0±0.516.3±2.182.3±0.769.6±3.5
海曙古林种群Haishu-Gulin population14.8±1.592.8±0.999.5±0.117.5±1.883.0±0.672.1±3.3
海曙集士港种群Haishu-Jishigang population15.4±2.692.9±0.799.4±0.118.2±3.183.1±0.872.6±4.6
奉化种群Fenghua population14.7±3.397.1±0.186.9±9.415.4±5.181.9±1.665.1±10.0
象山种群Xiangshan population13.3±1.193.5±0.792.9±11.214.5±0.982.2±0.667.8±4.1
余姚种群Yuyao population13.2±1.594.1±0.998.4±2.115.3±1.982.1±0.868.6±4.8
富阳种群Fuyang population12.5±0.797.0±0.299.1±0.114.7±0.881.9±0.466.9±2.4
北京种群Beijing Population14.7±2.697.0±0.199.1±0.317.3±3.282.0±0.573.4±3.4
), ArticleFig(id=1266746487355629980, tenantId=1146029695717560320, journalId=1266358857061122103, articleId=1266470629487301058, language=EN, label=Table 2, caption=

Functional annotation of single nucleotide polymorphisms and insertions/deletionsin the genome of Chilo suppressalis populations

, figureFileSmall=null, figureFileBig=null, tableContent=

功能类别

Functional category

单核苷酸多态性

Single nucleotide polymorphism

插入缺失

Insertion/deletion

数量Number比例Percent/%数量Number比例Percent/%
基因间Intergenic23 153 11150.4306 681 60350.330
内含子Intronic14 238 77731.0104 152 48931.280
编码序列Coding sequences558 7341.22037 2170.280
5'非翻译区5 untranslated region193 0800.42048 9940.370
3'非翻译区3 untranslated region902 7391.970262 9181.980
同义Synonymous292 4300.640--
起始子丢失Start lost5850.0014690.004
终止子获得Stop gained9 3440.0202 1350.016
终止子丢失Stop lost4 5100.0101 3360.010
终止子延长Stop retained1 6120.004430.001
剪接位点区域Splice site region52 0700.11014 1010.110
移码Frameshift--24 1560.180
), ArticleFig(id=1266746487431127453, tenantId=1146029695717560320, journalId=1266358857061122103, articleId=1266470629487301058, language=CN, label=表2, caption=

二化螟种群基因组单核苷酸多态性和插入缺失的功能注释

, figureFileSmall=null, figureFileBig=null, tableContent=

功能类别

Functional category

单核苷酸多态性

Single nucleotide polymorphism

插入缺失

Insertion/deletion

数量Number比例Percent/%数量Number比例Percent/%
基因间Intergenic23 153 11150.4306 681 60350.330
内含子Intronic14 238 77731.0104 152 48931.280
编码序列Coding sequences558 7341.22037 2170.280
5'非翻译区5 untranslated region193 0800.42048 9940.370
3'非翻译区3 untranslated region902 7391.970262 9181.980
同义Synonymous292 4300.640--
起始子丢失Start lost5850.0014690.004
终止子获得Stop gained9 3440.0202 1350.016
终止子丢失Stop lost4 5100.0101 3360.010
终止子延长Stop retained1 6120.004430.001
剪接位点区域Splice site region52 0700.11014 1010.110
移码Frameshift--24 1560.180
), ArticleFig(id=1266746487510819230, tenantId=1146029695717560320, journalId=1266358857061122103, articleId=1266470629487301058, language=EN, label=Table 3, caption=

Genetic differentiation indices among eight Chilo suppressalis populations

, figureFileSmall=null, figureFileBig=null, tableContent=

种群

Population

北京种群鄞州种群富阳种群余姚种群海曙集士港种群海曙古林种群奉化种群
Beijing population

Yinzhou

population

Fuyang

population

Yuyao

population

Haishu-Jishigang

population

Haishu-Gulin populationFenghua population
鄞州种群Yinzhou population0.272-
富阳种群Fuyang population0.2960.028-
余姚种群Yuyao population0.2840.0150.039-
海曙集士港种群Haishu-Jishigang population0.2720.0030.0310.019-
海曙古林种群Haishu-Gulin population0.2520.0030.0310.019<0.001-
奉化种群Fenghua population0.3490.0250.0600.0450.0250.024-
象山种群Xiangshan population0.3480.0560.0860.0730.0550.0540.090
), ArticleFig(id=1266746487594705311, tenantId=1146029695717560320, journalId=1266358857061122103, articleId=1266470629487301058, language=CN, label=表3, caption=

8个二化螟种群间的遗传分化指数

, figureFileSmall=null, figureFileBig=null, tableContent=

种群

Population

北京种群鄞州种群富阳种群余姚种群海曙集士港种群海曙古林种群奉化种群
Beijing population

Yinzhou

population

Fuyang

population

Yuyao

population

Haishu-Jishigang

population

Haishu-Gulin populationFenghua population
鄞州种群Yinzhou population0.272-
富阳种群Fuyang population0.2960.028-
余姚种群Yuyao population0.2840.0150.039-
海曙集士港种群Haishu-Jishigang population0.2720.0030.0310.019-
海曙古林种群Haishu-Gulin population0.2520.0030.0310.019<0.001-
奉化种群Fenghua population0.3490.0250.0600.0450.0250.024-
象山种群Xiangshan population0.3480.0560.0860.0730.0550.0540.090
), ArticleFig(id=1266746487670202784, tenantId=1146029695717560320, journalId=1266358857061122103, articleId=1266470629487301058, language=EN, label=Table 4, caption=

Nucleotide diversity and Tajima’s D values ofeight Chilo suppressalis populations

, figureFileSmall=null, figureFileBig=null, tableContent=

种群

Population

核苷酸多样性

Nucleotide diversity

Tajima’s D
鄞州种群Yinzhou population0.003 310.691

海曙古林种群

Haishu-Gulin population

0.003 330.666

海曙集士港种群

Haishu-Jishigang population

0.003 310.683
奉化种群Fenghua population0.002 870.427
象山种群Xiangshan population0.002 980.642
余姚种群Yuyao population0.003 190.632
富阳种群Fuyang population0.003 150.743
北京种群Beijing population0.001 360.878
), ArticleFig(id=1266746487745700257, tenantId=1146029695717560320, journalId=1266358857061122103, articleId=1266470629487301058, language=CN, label=表4, caption=

8个二化螟种群的核苷酸多样性和Tajimas D

, figureFileSmall=null, figureFileBig=null, tableContent=

种群

Population

核苷酸多样性

Nucleotide diversity

Tajima’s D
鄞州种群Yinzhou population0.003 310.691

海曙古林种群

Haishu-Gulin population

0.003 330.666

海曙集士港种群

Haishu-Jishigang population

0.003 310.683
奉化种群Fenghua population0.002 870.427
象山种群Xiangshan population0.002 980.642
余姚种群Yuyao population0.003 190.632
富阳种群Fuyang population0.003 150.743
北京种群Beijing population0.001 360.878
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基于全基因组重测序的宁波市二化螟种群遗传分析
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肖山 , 柴伟纲 , 刘健 , 谌江华
植物保护学报 | 研究论文 2026,53(2): 449-458
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植物保护学报 |研究论文 2026 , 53 (2) : 449 -458
基于全基因组重测序的宁波市二化螟种群遗传分析
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肖山, 柴伟纲, 刘健, 谌江华
作者信息
  • 宁波市农业科学研究院,宁波市特色农产品质量安全检测与控制重点实验室,浙江 宁波 315040
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Genetic analysis of Asiatic rice borer Chilo suppressalis populations in Ningbo City based on whole-genome resequencing
Shan Xiao, Weigang Chai, Jian Liu, Jianghua Chen
Affiliations
  • Ningbo Key Laboratory of Testing and Control for Characteristic Agro-Product Quality and Safety, Ningbo Academy ofAgricultural Sciences, Ningbo 315040, Zhejiang Province, China
出版时间: 2026-04-30 doi: 10.13802/j.cnki.zwbhxb.2026.2025103
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为解析宁波市二化螟Chilo suppressalis不同地理种群的遗传结构以及种群间亲缘关系,对8个二化螟种群(6个宁波市地理种群、1个北京种群和1个杭州种群)的37个样品进行全基因组重测序,鉴定基因组间遗传变异,并进行种群遗传结构、基因组选择性清除、遗传多样性、历史有效种群大小等种群遗传学分析。结果显示:在37个二化螟个体基因组中共鉴定到42 149 215个单核苷酸多态性和11 912 812个插入缺失,这些变异均主要分布在基因间和基因内含子区域。在宁波市6个二化螟种群中,鄞州、余姚、海曙种群的亲缘关系较近,象山种群与宁波市其他5个种群的亲缘关系最远,并表现出中度的遗传分化。二化螟象山种群基因组选择性清除区域内的基因显著富集在黑色素生成、昼夜节律夹带等7个信号通路。在宁波市各种群中,海曙种群和鄞州种群的核苷酸多样性较高,象山种群和奉化种群的核苷酸多样性较低。宁波市二化螟种群在3 000年前—1 000年前左右经历了快速收缩,随后种群规模快速增加。

二化螟  /  全基因组重测序  /  种群遗传结构  /  遗传多样性  /  遗传变异  /  遗传分化

To investigate the population structure and phylogenetic relationships among different geographic populations of Asiatic rice borer Chilo suppressalis in Ningbo City, China,whole-genome resequencing was performed on 37 individuals from eight populations (six from Ningbo, one from Beijing, and one from Hangzhou). Genomic variations were identified, and population genetic analyses including population structure, selective sweeps, genetic diversity, and historical effective population size were conducted. The results showed that a total of 42 149 215 single nucleotide polymorphisms and 11 912 812 insertions/deletions were identified across the 37 genomes, with most variations located in intergenic and intronic regions. Among the six Ningbo populations, Yinzhou, Yuyao and Haishu populations were closely related, whereas the Xiangshan population showed the greatest genetic distance from the other Ningbo populations and exhibited moderate genetic differentiation. Genes within the selective sweep regions of the Xiangshan population was significantly enriched in seven signaling pathways, including melanogenesis and circadian entrainment. Among the Ningbo populations, nucleotide diversity was relatively higher in the Haishu and Yinzhou populations, but lower in the Xiangshan and Fenghua populations. The Ningbo populations experienced a rapid contraction approximately 3 000—1 000 years ago, followed by a rapid population expansion.

Chilo suppressalis  /  whole genome resequencing  /  population genetic structure  /  genetic diversity  /  genetic variation  /  genetic differentiation
肖山, 柴伟纲, 刘健, 谌江华. 基于全基因组重测序的宁波市二化螟种群遗传分析. 植物保护学报, 2026 , 53 (2) : 449 -458 . DOI: 10.13802/j.cnki.zwbhxb.2026.2025103
Shan Xiao, Weigang Chai, Jian Liu, Jianghua Chen. Genetic analysis of Asiatic rice borer Chilo suppressalis populations in Ningbo City based on whole-genome resequencing[J]. Journal of Plant Protection, 2026 , 53 (2) : 449 -458 . DOI: 10.13802/j.cnki.zwbhxb.2026.2025103
二化螟Chilo suppressalis属鳞翅目草螟科,是中国水稻上最主要的害虫之一,其幼虫钻蛀水稻茎秆,造成水稻枯鞘、枯心、枯孕穗、白穗,甚至倒伏,严重影响水稻的产量和品质。近年来,由于气候变暖、单双季混栽面积增加、抗药性增强等原因,二化螟在中国大部分稻区为害呈加重趋势(叶恭银等,2023)。据统计,2013—2022年二化螟每年对中国水稻造成的产量损失达55.1万t(卓富彦等,2024)。因此,构建科学高效的二化螟防控技术体系已成为保障水稻安全生产的重要课题。
目前,化学防治仍是控制二化螟的主要手段,但长期大量使用杀虫剂导致其抗药性问题日益突出。在宁波市,二化螟已对氯虫苯甲酰胺、阿维菌素、三唑磷、乙基多杀菌素等多种主流杀虫剂产生了明显抗性(任宗杰等,2025)。例如,2019年余姚种群、象山种群对氯虫苯甲酰胺的抗性倍数分别为702.6倍和2 087.5倍,2022年余姚种群对阿维菌素、三唑磷的抗性已达中等水平,居全国前列(Huang et al.,2021Wang et al.,2024)。宁波市作为二化螟抗药性严重的典型区域,揭示其种群抗性演化的分子机制对于制订科学精准的防控策略具有重要意义。在杀虫剂及环境因子的长期选择压力下,二化螟等害虫通过积累适应性遗传变异来不断提升自身对环境的适合度,进而增加防治难度(Zhu et al.,2019范文敏等,2025Peng et al.,2025)。近年来,高通量测序成本持续下降,全基因组重测序技术凭借其高效鉴定全基因组遗传变异的优势被广泛用于动植物种质资源评价、进化起源追溯、环境适应性解析、分子育种和病虫害防控等多个领域(Bourgeois & Warren,2021Song et al.,2023)。例如,Pélissié et al.(2022)对6个地理种群的85个马铃薯甲虫Leptinotarsa decemlineata吡虫啉抗性及敏感个体进行重测序分析,发现不同种群中抗药性相关基因均受到强烈选择,但涉及的基因在不同种群间存在差异,呈现出多样化的抗药性分子机制;Zhang et al.(2023)基于全球橘小实蝇Bactrocera dorsalis地理种群的基因组重测序鉴定出一个关键的热适应相关基因Cyp6a9,并系统解析了橘小实蝇在全球的入侵与扩散路径;Hu et al.(2024)基于全球褐飞虱Nilaparvata lugens地理种群的基因组重测序,系统解析了其在全球的入侵与扩散路径;Peng et al.(2025)对中国24个二化螟地理种群的群体结构与种群演化历史进行了分析,并鉴定到一个耐寒性适应的关键基因goosecoid,该基因的表达水平在低纬度和高纬度种群中呈现显著差异。因此,本研究利用种群基因组学分析解析二化螟演化历史与环境适应性机制。
为解析宁波市二化螟的遗传结构以及种群间亲缘关系,本研究对8个二化螟种群(6个宁波市地理种群、1个北京种群和1个杭州种群)的37个样品进行全基因组重测序,并进行基因组间遗传变异、种群遗传结构、基因组选择性清除、遗传多样性以及历史有效种群大小等种群遗传学分析,以期为二化螟抗药性等环境适应性机制研究提供遗传学基础,并为制订区域性的精准防控策略提供科学依据。
供试种群:2024年6月自宁波市鄞州区、海曙区、奉化区、象山县和余姚市水稻田共采集27头二化螟3~5龄幼虫,即27个样品,其中鄞州区东钱湖镇采集5头,海曙区古林镇和集士港镇各采集5头,奉化区江口街道采集3头,象山县西周镇采集4头,余姚市阳明街道采集5头,分别命名为YZ、HS-GL、HS-JSG、FH、XS和YY种群。2024年5月自杭州市富阳区中国水稻研究所田间采集5头二化螟幼虫,2024年5月自北京绽诺思特生物科技有限公司购买5头二化螟幼虫,为室内长期饲养种群,分别命名为FY种群和BJ种群,作为外群。
仪器:Agilent 2100 Bioanalyzer型核酸分析仪,美国安捷伦科技有限公司;SUB Cell GT型水平电泳槽和PowerPac Basic型电泳仪,美国伯乐公司;MGISEQ-T7型高通量测序仪,深圳华大智造科技股份有限公司。
对8个种群的37头二化螟进行单头建库测序。将37头试虫依次经75%酒精、纯水表面清洗后,置于-80 ℃保存备用。采用CTAB法提取每个二化螟样品的全基因组DNA,用核酸分析仪和电泳仪检测基因组DNA的质量,选取高质量的样品建库。检测合格文库即可在测序仪上进行测序。建库及测序均委托武汉希望组生物科技有限公司完成。使用fastp 0.24.0软件对测序获得的原始数据进行质控,对含接头序列、10%以上为未知序列、质量值小于20的位点总数超过reads长度50%的reads进行过滤。自InsectBase(http://v2.insect-genome.com/)下载二化螟基因组。使用BWA-MEM2 2.2.1软件将质控后的测序数据比对到二化螟参考基因组上,软件参数默认。利用bamdst(https://github.com/shiquan/bamdst)软件对各样本比对率、测序深度及覆盖度进行统计。取基因组比对率大于70%及测序深度大于10倍的样品进行下一步分析。
使用GATK 4.6.1.0软件内MarkDuplicates模块去除测序数据中因PCR扩增而产生的重复序列;使用HaplotypeCaller模块检测每个样品的变异,利用CombineGVCFs模块对所有样品变异信息进行合并,采用SelectVariants模块鉴定各样品的单核苷酸多态性和插入缺失;使用VariantFiltration模块过滤低质量的变异。使用SnpEff 5.2软件对获得的每个样品的基因组遗传变异信息进行功能注释。
采用种群结构分析、主成分分析和系统发育分析3种方法对测序的8个二化螟种群进行种群遗传结构分析。使用vcftools 0.1.16软件提取单核苷酸多态性中双等位基因、最小等位基因频率大于0.02、缺失率小于0.2的位点;采用PLINK 1.90b6.21软件在窗口10 bp、步长10 bp下进一步过滤连锁程度r2>0.2的单核苷酸多态性;使用Admixture 1.3.0软件对过滤后的独立单核苷酸多态性进行种群结构分析,种群的祖先个数K设定为1~5,计算每个K值对应的交叉验证误差,根据最小交叉验证误差确定最接近种群真实情况的祖先个数。采用PLINK 1.90b6.21软件对去连锁的单核苷酸多态性数据集进行主成分分析并构建遗传距离矩阵。根据遗传距离矩阵,使用MEGA 11.0软件采用邻接法构建系统发育树,重复1 000次,使用iTOL(https://itol.embl.de/)在线网站对进化树进行美化。采用PLINK 1.90b6.21软件计算每两个种群间的遗传分化指数Fst。当0≤Fst≤0.05时表示种群间遗传分化程度很小,当0.05<Fst≤0.15时表示种群间遗传分化程度中等,当0.15<Fst≤0.25时表示种群间遗传分化程度较大,而Fst>0.25表示种群间遗传分化程度很大(Wright,1965)。
根据1.2.3结果,二化螟象山种群与浙江省其他6个种群表现中度分化。因此对二化螟象山种群和浙江省其他6个种群进行基因组选择性清除分析。在滑动窗口10 kb下,以5 kb为步长使用vcftools 0.1.16软件分别计算鄞州、奉化、富阳、海曙集士港、海曙古林和余姚这6个浙江省二化螟种群的核苷酸多样性πother与象山种群的核苷酸多样性πXS的比值和遗传分化指数,使用R语言将πother/πXS取对数,即log2πother/πXS);使用R语言计算遗传分化指数的标准分数,即Z(Fst),取这两个值前5%的区域为二化螟象山种群基因组的受选择区域。根据基因组注释文件,利用R包GenomicRanges 1.58.0获得受选择区域内的基因信息。利用基迪奥在线网站(https://www.omicshare.com/)对受选择基因进行KEGG(Kyoto encyclopedia of genes and genomes)通路富集分析。
为分析种群的遗传多样性,使用vcftools 0.1.16软件分别计算8个二化螟种群的核苷酸多样性和Tajima’s D值。核苷酸多样性计算时窗口大小设置为10 kb,步长为5 kb。Tajima’s D值计算时窗口大小设置为10 kb。当Tajima’s D值为正时,表明种群经历了瓶颈效应或平衡选择;当Tajima’s D值接近0时,表明种群处于中性进化状态,没有显著的选择压力或种群历史变化;当Tajima’s D值为负时,表明种群可能经历了扩张或者近期正选择(Tajima,1989)。
将奉化、海曙古林镇、海曙集士港镇、象山、余姚和鄞州6个宁波市地理种群所有个体的单核苷酸多态性作为SMC++ 1.15.4软件的输入文件,平均世代时间设为0.4年,突变速率设为8.4×10-9Haag-Liautard et al.,2007Peng et al.,2025),使用SMC++ 1.15.4软件分析这6个种群的历史有效种群大小。
二化螟8个种群测序数据经质量过滤后的clean data为12.5~15.4 Gb,测序质量Q30为92.8%~97.1%,表明各样品测序质量很高(表1)。与二化螟参考基因组相比,8个种群基因组的比对率为86.9%~99.5%;8个种群的测序深度为14.5倍~18.2倍,4倍覆盖度为81.9%~83.1%,10倍覆盖度为65.1%~73.4%,表明样品的测序深度以及覆盖度足够,可满足后续基因组重测序分析。
在37个二化螟个体的基因组中共鉴定到42 149 215个单核苷酸多态性以及11 912 812个插入缺失。单核苷酸多态性主要分布在基因间区域,占总单核苷酸多态性数的50.430%,其次是位于内含子区域,占比为31.010%;558 734个单核苷酸多态性位于基因质编码区,其中292 430个为同义突变,而非同义突变中造成起始子丢失、终止子获得、终止子丢失以及终止子延长的单核苷酸多态性分别为585、9 344、4 510和1 612个(表2)。插入缺失的分布与单核苷酸多态性类似,主要分布在基因间及内含子区域,占比分别为50.330%和31.280%;37 217个插入缺失位于基因编码区,占总插入缺失数的0.280%,其中24 156个插入缺失会造成移码(表2)。基因组中平均每kb含有48.7个单核苷酸多态性和13.8个插入缺失,且单核苷酸多态性和插入缺失均主要分布在染色体两端(图1)。
当祖先个数为2时,交叉验证误差最小,所以8个二化螟种群祖先个数为2最可信(图2-A)。当祖先个数为2时,北京种群与其他7个二化螟种群分离,当祖先个数为3时,象山种群的3个个体与浙江省其他6个二化螟种群分离,而当祖先个数为4时,富阳种群的部分二化螟进一步分离(图2-B)。
在第1主成分上,二化螟北京种群与浙江省7个种群明显分开;而在第2主成分上,二化螟象山种群、富阳种群与浙江省其他5个种群分开,且象山种群偏离程度更大,与种群结构分析结果一致(图3)。
二化螟富阳、余姚、鄞州种群聚为一簇,二化螟海曙集士港、海曙古林种群聚为一簇,而剩下的奉化种群、象山种群以及北京种群各聚为一簇,与种群结构分析以及主成分分析结果一致(图4)。
8个二化螟种群的遗传分化指数介于0~0.349之间,其中北京种群与其他7个种群的遗传分化指数较大,均大于0.250;象山种群与浙江省其他6个种群以及奉化种群与富阳种群的遗传分化指数均大于0.050,为中度分化;其他种群之间的遗传分化指数均很小(表3)。
log2πother/πXS)>2.082且Z(Fst)>1.81的区段为二化螟象山种群基因组受选择区段(图5)。对相邻区段进行合并后共鉴定到747个区段,总长度为9.705 Mb,共有461个基因位于选择区段内。KEGG通路富集分析结果显示,这461个基因主要富集在黑色素生成、昼夜节律夹带等7个通路(图6)。
8个二化螟种群的核苷酸多样性介于1.36×10-3~3.33×10-3之间,其中北京种群的核苷酸多样性最低,这可能是因为北京种群为室内种群,长期的地理隔离及人工饲养驯化导致其遗传多样性降低;在其他7个种群中,奉化种群和象山种群的核苷酸多样性相对较低,分别为2.87×10-3和2.98×10-3,而海曙古林、海曙集士港以及鄞州种群的核苷酸多样性较高,表明其遗传多样性相对较高(表4)。8个二化螟种群的Tajima’s D值介于0.427~0.878之间,其中北京种群的Tajima’s D值最高,而奉化种群的Tajima’s D值最低;8个二化螟种群的Tajima’s D值均大于0,表明各种群可能经历了瓶颈效应或者平衡选择。
宁波市6个二化螟种群最早可追溯到40万年前,各二化螟种群历史较一致。在40万年前—1 000年前左右,各种群规模整体呈下降趋势,尤其是3 000年前—1 000年前左右期间种群经历了快速收缩;而1 000年前左右—400年前,各二化螟种群规模增加。在宁波市6个二化螟种群中,象山种群的历史有效种群最早降到最低值,且远低于其他5个种群,但在近期其快速扩张,在400年前象山种群的历史有效种群大于其他5个种群;在宁波市6个二化螟种群中,奉化种群的历史有效种群最晚降到最低值,且在400年前的历史有效种群最小(图7)。
昆虫种群遗传学研究种群内基因组成、遗传变异及其时空动态变化规律,为解析昆虫的起源演化、迁移扩散以及环境适应机制提供理论支撑(Hopper,2023)。已有二化螟种群遗传学研究主要基于线粒体基因、核糖体基因和微卫星等分子标记,然而这类标记数量有限,基因组覆盖度低(Meng et al.,2008袁龙宇等,2025)。本研究采用全基因组重测序技术在二化螟基因组中鉴定到数千万个单核苷酸多态性,避免了传统分子标记信息量有限、统计效力不足的局限。
二化螟种群遗传结构具有尺度依赖性。例如,Meng et al.(2008)Peng et al.(2025)研究结果显示在全国范围内二化螟种群可划分为5个遗传种群,种群间遗传分化明显,遗传距离与地理距离呈显著正相关;于文娟等(2021)研究结果显示在四川省范围内二化螟种群间遗传分化较小,地理隔离效应不明显;鲁艳辉等(2022)研究结果显示在浙江省范围内二化螟种群间遗传分化较小,地理隔离效应不明显。本研究的系统发育分析结果显示二化螟象山种群单独聚为一支,并与宁波市其他二化螟种群发生了中等程度的分化,究其原因可能是在地理位置上象山县与宁波市鄞州区、奉化区等隔港相望,存在一定的地理隔离,种群间基因交流较少。
本研究通过选择性清除分析发现,在演化过程中二化螟象山种群的黑色素生成、昼夜节律夹带等通路基因可能经历了强烈的定向选择(Stephan,2019)。宁波市象山县三面环海,受海洋水体的影响,其气温、光照、昼夜温差等与宁波市其他区域存在一定差异,而黑色素生成和昼夜节律夹带等通路基因在紫外线防护、温度调节、免疫调节、光照感知与适应、生理节律调节和行为调节等方面发挥着重要作用,有助于种群更好地适应当地环境,从而提高适合度(D’Mello et al.,2016Sugumaran & Barek,2016Dubowy & Sehgal,2017)。Peng et al.(2025)研究结果显示不同纬度二化螟种群的昼夜节律通路基因受到了定向选择,与本研究结果一致。
二化螟对杀虫剂的抗性机制主要是解毒酶介导的代谢抗性以及靶标突变引起的靶标抗性(范文敏等,2025)。目前,关于代谢抗性相关基因的挖掘主要通过酶活性检测、实时荧光定量PCR检测、转录组分析等方法,而靶标抗性的研究则主要是对靶标受体基因的PCR克隆(Zhao et al.,2019Wang et al.,2025)。但基因组中拷贝数增加、启动子区等非编码区域的遗传突变也可导致基因表达量变化,进而介导抗药性的增强或产生(Heckel,2022Hu et al.,2025)。近年来,越来越多的研究通过全基因组重测序系统鉴定基因组内与抗药性等相关的遗传变异。例如,在褐飞虱烟碱型乙酰胆碱受体基因nAChR-7-like及细胞色素P450 CYP4C61基因中鉴定到多个单核苷酸多态性,这些遗传变异影响褐飞虱对吡虫啉及抗褐飞虱水稻IR36的抗性,其中3个单核苷酸多态性可用于区分抗性与敏感基因型(Pang et al.,2024)。目前主要依靠线粒体基因分析二化螟种群的抗性水平与遗传多样性间的关系,尚未有在全基因组层面鉴定抗药性相关遗传变异的研究(鲁艳辉等,2022黄芊等,2025)。
本研究通过全基因组重测序方法分析了宁波市6个二化螟种群基因组的遗传变异,未对各种群进行抗药性测定,后续可通过全基因组关联分析等方法分析与多种杀虫剂相关的遗传变异,为揭示宁波市二化螟抗药性等环境适应性分子进化机制以及防治策略制订提供参考。
  • 宁波市公益性科技计划项目(2023S014)
  • 宁波市农业科学研究院院长基金项目(2024NKYP006)
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doi: 10.13802/j.cnki.zwbhxb.2026.2025103
  • 接收时间:2025-07-24
  • 首发时间:2026-05-27
  • 出版时间:2026-04-30
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  • 收稿日期:2025-07-24
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宁波市公益性科技计划项目(2023S014)
宁波市农业科学研究院院长基金项目(2024NKYP006)
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    宁波市农业科学研究院,宁波市特色农产品质量安全检测与控制重点实验室,浙江 宁波 315040

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2种不同金属材料的力学参数

Family
属数
Number of
genus
种数
Number of
species
占总种数比例
Percentage of
total species (%)

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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