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Article(id=1276601551179346059, tenantId=1146029695717560320, journalId=1235980609244409860, issueId=1276601397818814642, articleNumber=null, orderNo=null, doi=10.3969/j.issn.1000-2561.2025.01.002, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=null, receivedDate=1721664000000, receivedDateStr=2024-07-23, revisedDate=1723996800000, revisedDateStr=2024-08-19, acceptedDate=null, acceptedDateStr=null, onlineDate=1782295127910, onlineDateStr=2026-06-24, pubDate=1737734400000, pubDateStr=2025-01-25, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1782295127910, onlineIssueDateStr=2026-06-24, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1782295127910, creator=13701087609, updateTime=1782295127910, updator=13701087609, issue=Issue{id=1276601397818814642, tenantId=1146029695717560320, journalId=1235980609244409860, year='2025', volume='46', issue='1', pageStart='1', pageEnd='245', issueExtLink='null', onlineDate='null', pubDate='1737734400000', pubDateStr='2025-01-25', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1782295091347, creator='13701087609', updateTime=1782295207335, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext={EN=IssueExt(id=1276601884408418422, tenantId=1146029695717560320, journalId=1235980609244409860, issueId=1276601397818814642, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1276601884408418423, tenantId=1146029695717560320, journalId=1235980609244409860, issueId=1276601397818814642, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null, downloadFileDto=null}, startPage=10, endPage=20, ext={EN=ArticleExt(id=1276601551447781517, articleId=1276601551179346059, tenantId=1146029695717560320, journalId=1235980609244409860, language=EN, title=Identification and Expression Profile Analysis of KAS Gene Family in Camellia oleifera, columnId=1236256430337085821, journalTitle=Chinese Journal of Tropical Crops, columnName=Omics & Biotechnology, runingTitle=null, highlight=null, articleAbstract=

Beta-ketoacyl-acyl carrier protein synthase (KAS) is the key enzyme system in plant fatty acid biosynthesis. In order to investigate the function of the KAS gene family in the maturation and oil accumulation of Camellia oleifera, this study used bioinformatics methods and analyzed ther physicochemical properties, chromosomal localization, subcellular localization, secondary structure, phylogenetic trees, conserved motifs, promoter cis-acting elements, and RT-qPCR to analyze the expression patterns in the oil rapid accumulation stage of C. oleifera. The findings indicated that 14 Camellia oleifera KAS family members were distributed on seven chromosomes with relative molecular masses between 1.13 kDa and 5.15 kDa, and most of them were acid-stabilized proteins. Except for CoKAS II-2, which was localized in mitochondria and chloroplasts, all of them were localized in chloroplasts. Phylogenetic analysis revealed that CoKAS was divided into three subclasses, with similarities in structure and conserved motifs among KAS family numbers in the same subfamily. The promoter regions of the CoKAS gene family members were enriched with the cis-acting elements growth and development-related, light response, hormone induction, and adversity response. Expression analysis revealed that most of the CoKAS genes were highly expressed during the oil rapid accumulation stage. In addition, it was found a strong correlation between the members of the CoKAS and CoMYB gene families during fruit ripening in C. oleifera. The findings would provide a theoretical basis for comprehensively analyzing the function of CoKAS genes.

, authors=null, authorsList=Xiaoxia LIU, Zihao YU, Huiyang JING, Kaifeng XIONG, Zuodong QIN, authorCompany=null, correspAuthors=Zuodong QIN, 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=1276601555235238044, articleId=1276601551179346059, tenantId=1146029695717560320, journalId=1235980609244409860, language=CN, title=油茶KAS基因家族的鉴定及表达分析, columnId=1236256430517440904, journalTitle=热带作物学报, columnName=组学与生物技术, runingTitle=null, highlight=null, articleAbstract=

β-酮脂酰ACP合成酶（beta-ketoacyl-acyl carrier protein synthase，KAS）是植物脂肪酸生物合成过程中的关键酶系。为探究KAS在油茶种子成熟和油脂积累中的作用，本研究通过生物信息学方法对CoKAS基因家族成员进行鉴定，分析其理化性质、染色体定位、亚细胞定位、二级结构、进化关系、保守基序、启动子顺式作用元件，并采用RT-qPCR技术分析油茶果实油脂快速积累期的表达模式。结果表明：鉴定出的14个CoKAS家族成员分布在7条染色体上，相对分子质量在1.13~5.15 kDa之间，且大部分为酸性稳定蛋白。除CoKAS II-2定位于线粒体和叶绿体外，其余均定位于叶绿体。系统发育分析显示CoKAS基因被分为3个亚类，同一亚族成员的基因结构和保守基序具有相似性。CoKAS基因家族成员的启动子区域富含与生长发育、光响应、激素诱导和胁迫应答相关的顺式作用元件。通过表达分析发现，大部分CoKAS基因在油茶种子油脂快速积累期的表达量较高。此外，在油茶果实成熟过程中，CoKAS与CoMYB基因家族间存在有较强相关性的成员。本研究结果为深入解析CoKAS基因在油茶油脂生物合成中的调控机制提供理论基础。

, authors=

刘晓霞（1989—），女，硕士，实验师，研究方向：天然产物的开发及其生物合成调控。

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* 覃佐东（QIN Zuodong），E-mail：。
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刘晓霞（1989—），女，硕士，实验师，研究方向：天然产物的开发及其生物合成调控。

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刘晓霞（1989—），女，硕士，实验师，研究方向：天然产物的开发及其生物合成调控。

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Journal of Integrative Agriculture, 2018, 17(7): 1631-1640., articleTitle=Light shading improves the yield and quality of seed in oil-seed peony (Paeonia ostii Feng Dan), refAbstract=null), Reference(id=1276601947885015473, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276601551179346059, doi=null, pmid=null, pmcid=null, year=2017, volume=212, issue=null, pageStart=165, pageEnd=174, url=null, language=null, rfNumber=[26], rfOrder=29, authorNames=GAUTHIER M, PELLET D, MONNEY C, HERRERA J M, ROUGIER M, BAUX A, journalName=Field Crops Research, refType=null, unstructuredReference=GAUTHIER M, PELLET D, MONNEY C, HERRERA J M, ROUGIER M, BAUX A. Fatty acids composition of oilseed rape genotypes as affected by solar radiation and temperature[J]. 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Plant Physiology, 2017, 173(2): 1211-1225., articleTitle=MYB89 transcription factor represses seed oil accumulation, refAbstract=null), Reference(id=1276601949738897845, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276601551179346059, doi=null, pmid=null, pmcid=null, year=2017, volume=8, issue=null, pageStart=null, pageEnd=null, url=null, language=null, rfNumber=[30], rfOrder=33, authorNames=DUAN S W, JIN C Y, LI D, GAO C H, QI S H, LIU K G, HAI J B, MA H L, CHEN M G, journalName=Frontiers in Plant Science, refType=null, unstructuredReference=DUAN S W, JIN C Y, LI D, GAO C H, QI S H, LIU K G, HAI J B, MA H L, CHEN M G. MYB76 inhibits seed fatty acid accumulation in Arabidopsis[J]. Frontiers in Plant Science, 2017, 8: 251917., articleTitle=MYB76 inhibits seed fatty acid accumulation in Arabidopsis, refAbstract=null), Reference(id=1276601949873115574, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276601551179346059, doi=null, pmid=null, pmcid=null, year=2014, volume=26, issue=9, pageStart=3519, pageEnd=3537, url=null, language=null, rfNumber=[31], rfOrder=34, authorNames=BARTHOLE G, TO A, MARCHIVE C, BRUNAUD V, SOUBIGOU-TACONNAT L, BERGER N, DUBREUCQ B, LEPINIEC L, BAUD S, journalName=The Plant Cell, refType=null, unstructuredReference=BARTHOLE G, TO A, MARCHIVE C, BRUNAUD V, SOUBIGOU-TACONNAT L, BERGER N, DUBREUCQ B, LEPINIEC L, BAUD S. MYB118 represses endosperm maturation in seeds of Arabidopsis[J]. The Plant Cell, 2014, 26(9): 3519-3537., articleTitle=MYB118 represses endosperm maturation in seeds of Arabidopsis, refAbstract=null)], funds=[Fund(id=1276601945628479889, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276601551179346059, awardId=2024JJ7188, language=CN, fundingSource=湖南省自然科学基金项目(2024JJ7188), fundOrder=null, country=null), Fund(id=1276601945683005842, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276601551179346059, awardId=23C0354, language=CN, fundingSource=湖南省教育厅科学研究项目(23C0354), fundOrder=null, country=null), Fund(id=1276601945737531795, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276601551179346059, awardId=23XKY07, language=CN, fundingSource=湖南科技学院科学研究项目(23XKY07), fundOrder=null, country=null)], companyList=[AuthorCompany(id=1276601936338096476, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276601551179346059, xref=null, ext=[AuthorCompanyExt(id=1276601936346485085, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276601551179346059, companyId=1276601936338096476, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=College of Chemistry and Bioengineering, Hunan University of Science and Engineering / Hunan Engineering Technology Research Center for Comprehensive Development and Utilization of Biomass Resources, Yongzhou, Hunan 425199, China), AuthorCompanyExt(id=1276601936354873694, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276601551179346059, companyId=1276601936338096476, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=湖南科技学院化学与生物工程学院/湖南省生物质资源综合开发利用工程技术研究中心，湖南永州　425199)])], figs=[ArticleFig(id=1276601942751187329, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276601551179346059, language=EN, label=Fig. 1, caption=Phylogenetic trees of KAS family members between C. oleifera and Arabidopsis thaliana

The numbers in the graph indicate that the bootstrap confidence value is greater than 75%.

, figureFileSmall=oj+FSa90Y4WFBst8DrwN6Q==, figureFileBig=WiXzfeunzySkF4mQEfHpLQ==, tableContent=null), ArticleFig(id=1276601942809907586, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276601551179346059, language=CN, label=图1, caption=油茶和拟南芥KAS蛋白系统进化关系

图中数值表示自展支持率大于75%。

, figureFileSmall=oj+FSa90Y4WFBst8DrwN6Q==, figureFileBig=WiXzfeunzySkF4mQEfHpLQ==, tableContent=null), ArticleFig(id=1276601942902182275, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276601551179346059, language=EN, label=Fig. 2, caption=Gene structure, conserved structural domains, and motif analysis of CoKAS family

A: Gene structure analysis of the CoKAS family; B: Conserved structural domains analysis of the CoKAS family; C: Conserved motif analysis of the CoKAS family; D: Conserved motif sequences of the CoKAS family LOGO.

, figureFileSmall=GEF+0O4vSTpgS9Jh8XkdhA==, figureFileBig=iqpqce+ni8M6GVn0IUN5vQ==, tableContent=null), ArticleFig(id=1276601942977679748, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276601551179346059, language=CN, label=图2, caption=COKAS家族基因结构、保守结构域和保守基序分析

A：COKAS家族基因结构分析；B：COKAS家族保守结构域分析；C：COKAS家族保守基序分析；D：COKAS家族保守基序序列LOGO。

, figureFileSmall=GEF+0O4vSTpgS9Jh8XkdhA==, figureFileBig=iqpqce+ni8M6GVn0IUN5vQ==, tableContent=null), ArticleFig(id=1276601943078343045, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276601551179346059, language=EN, label=Fig. 3, caption=Cis-acting elements in promoters of CoKAS family members, figureFileSmall=Xcp51h+jACNsAUy28CGlig==, figureFileBig=W2MsEczFQh6ygz6ioc4XbQ==, tableContent=null), ArticleFig(id=1276601943158034822, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276601551179346059, language=CN, label=图3, caption=油茶COKAS家族成员启动子顺势作用元件, figureFileSmall=Xcp51h+jACNsAUy28CGlig==, figureFileBig=W2MsEczFQh6ygz6ioc4XbQ==, tableContent=null), ArticleFig(id=1276601943216755079, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276601551179346059, language=EN, label=Fig. 4, caption=Expression patterns of CoKAS genes at five different stages of seed development, figureFileSmall=4GA7cSx7cg6Ii2uqNFV8hA==, figureFileBig=G8b0X3WAa5bb0Z8dHaNvJA==, tableContent=null), ArticleFig(id=1276601943300641160, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276601551179346059, language=CN, label=图4, caption=油茶COKAS基因在油茶果实不同发育时期的表达模式, figureFileSmall=4GA7cSx7cg6Ii2uqNFV8hA==, figureFileBig=G8b0X3WAa5bb0Z8dHaNvJA==, tableContent=null), ArticleFig(id=1276601943380332937, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276601551179346059, language=EN, label=Fig. 5, caption=Analysis of expression patterns of CoKAS gene family members in oil rapid accumulation stage

Different lowercase letters indicate significant difference among treatments (P<0.05).

, figureFileSmall=+LcSigWmind5RSKqhiXG+g==, figureFileBig=268OJR4p20Rho09qk6xJZA==, tableContent=null), ArticleFig(id=1276601945032888714, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276601551179346059, language=CN, label=图5, caption=油脂快速积累期CoKAS基因家族成员表达分析

不同小写字母表示处理间差异显著（P<0.05）。

, figureFileSmall=+LcSigWmind5RSKqhiXG+g==, figureFileBig=268OJR4p20Rho09qk6xJZA==, tableContent=null), ArticleFig(id=1276601945120969099, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276601551179346059, language=EN, label=Fig. 6, caption=Correlation analysis of expression of CoKAS and transcription factors CoMYB

Deeper colors and larger circles indicate a greater correlation between the two.

, figureFileSmall=X4JC6D/y2EtOouc6g0ycxA==, figureFileBig=p660/HYwUBkDR6K/Ki+TlQ==, tableContent=null), ArticleFig(id=1276601945204855180, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276601551179346059, language=CN, label=图6, caption=油茶COKAS与转录因子COMYB表达的相关性分析

颜色越深和圆圈越大表明二者之间的相关性越强。

, figureFileSmall=X4JC6D/y2EtOouc6g0ycxA==, figureFileBig=p660/HYwUBkDR6K/Ki+TlQ==, tableContent=null), ArticleFig(id=1276601945271964045, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276601551179346059, language=EN, label=Tab. 1, caption=

Primers sequence for CoKAS family

, figureFileSmall=null, figureFileBig=null, tableContent=
基因名称Gene name正向引物（5′–3′）Forward primer (5′–3′)反向引物（5′–3′）Reverse primer (5′–3′)
EF1a2CGTTATCGTTGCCTCTGCTCGCGTTATGTTTTCCTGTTTTGTC
CoKAS I-1ATGGGTCTCGTCTCCGTCTTGGGTAGGGAATTTGGACGCA
CoKAS I-3GCATCACTCACATTGACCGCTGCAGTAGCGAAGGCAATCA
CoKAS II-3TGAGAAGAACAATGGCGGCTCACGTCGTTTCCAAGCACTG
CoKAS II-5GGCATAGCTGGTGCTTGTTCGTCGCTTGCTTGCTCTCAAC
CoKAS II-6CTGGGGATTTGGCAGAGGTTATCCACCAGAAGCTCCAAGC
CoKAS III-1CGAGTACCCAGGCTTGTCAGGGATCCCAGTGCGAACAGAT
CoKAS III-2CGAGTACCCAGGCTTGTCAGGGATCCCAGTGCGAACAGAT
CoKAS III-3AAGGCTGGTCTCACTTCGTCAGAAATGACCCGCTCTGGTG
), ArticleFig(id=1276601945343267214, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276601551179346059, language=CN, label=表1, caption=

油茶KAS家族成员的引物序列

, figureFileSmall=null, figureFileBig=null, tableContent=
基因名称Gene name正向引物（5′–3′）Forward primer (5′–3′)反向引物（5′–3′）Reverse primer (5′–3′)
EF1a2CGTTATCGTTGCCTCTGCTCGCGTTATGTTTTCCTGTTTTGTC
CoKAS I-1ATGGGTCTCGTCTCCGTCTTGGGTAGGGAATTTGGACGCA
CoKAS I-3GCATCACTCACATTGACCGCTGCAGTAGCGAAGGCAATCA
CoKAS II-3TGAGAAGAACAATGGCGGCTCACGTCGTTTCCAAGCACTG
CoKAS II-5GGCATAGCTGGTGCTTGTTCGTCGCTTGCTTGCTCTCAAC
CoKAS II-6CTGGGGATTTGGCAGAGGTTATCCACCAGAAGCTCCAAGC
CoKAS III-1CGAGTACCCAGGCTTGTCAGGGATCCCAGTGCGAACAGAT
CoKAS III-2CGAGTACCCAGGCTTGTCAGGGATCCCAGTGCGAACAGAT
CoKAS III-3AAGGCTGGTCTCACTTCGTCAGAAATGACCCGCTCTGGTG
), ArticleFig(id=1276601945414570383, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276601551179346059, language=EN, label=Tab. 2, caption=

Basic information of CoKAS family

, figureFileSmall=null, figureFileBig=null, tableContent=
基因名称Gene name染色体Chromosome氨基酸数量Number of amino acids分子质量Molecular weight/kDa等电点pI不稳定系数Instability index脂肪族指数Aliphatic index水合平均值GRAVY grand average of hydro-pathicity二级结构Secondary structure/%亚细胞定位Subcellular localization
α-螺旋Alpha helixβ-转角Beta turn无规则卷曲Random coil延伸链Extended strand
CoKAS I-144424.726.3338.0181.70–0.16435.076.5643.4414.93Chloroplast
CoKAS I-244815.158.7032.6777.75–0.17833.066.6544.0716.22Chloroplast
CoKAS I-344765.108.8932.9278.36–0.18034.666.7243.7014.92Chloroplast
CoKAS I-441071.134.5442.3580.37–0.16033.649.3542.9914.02Chloroplast
CoKAS II-194755.086.3541.5781.83–0.05635.375.8944.0014.74Chloroplast
CoKAS II-222412.536.1335.9783.150.01135.275.8144.4014.52Chloroplast Mitochondrion
CoKAS II-324795.128.8838.7685.18–013738.006.2640.7115.03Chloroplast
CoKAS II-4104584.888.0041.0685.04–0.11737.345.6842.2313.76Chloroplast
CoKAS II-5104024.276.9839.4177.24–0.14338.067.2139.5515.17Chloroplast
CoKAS II-6143183.346.2032.0182.89–0.01336.797.5538.9916.67Chloroplast
CoKAS III-1154054.265.5638.2689.560.03728.408.5139.5123.95Chloroplast
CoKAS III-2143944.135.4138.7288.860.02427.168.3840.3624.11Chloroplast
CoKAS III-3144004.246.2443.9691.250.00328.259.0038.7524.00Chloroplast
CoKAS III-4154004.246.5343.9691.970.02226.509.0040.5024.00Chloroplast
), ArticleFig(id=1276601945498456464, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276601551179346059, language=CN, label=表2, caption=

CoKAS家族成员基本信息

, figureFileSmall=null, figureFileBig=null, tableContent=
基因名称Gene name染色体Chromosome氨基酸数量Number of amino acids分子质量Molecular weight/kDa等电点pI不稳定系数Instability index脂肪族指数Aliphatic index水合平均值GRAVY grand average of hydro-pathicity二级结构Secondary structure/%亚细胞定位Subcellular localization
α-螺旋Alpha helixβ-转角Beta turn无规则卷曲Random coil延伸链Extended strand
CoKAS I-144424.726.3338.0181.70–0.16435.076.5643.4414.93Chloroplast
CoKAS I-244815.158.7032.6777.75–0.17833.066.6544.0716.22Chloroplast
CoKAS I-344765.108.8932.9278.36–0.18034.666.7243.7014.92Chloroplast
CoKAS I-441071.134.5442.3580.37–0.16033.649.3542.9914.02Chloroplast
CoKAS II-194755.086.3541.5781.83–0.05635.375.8944.0014.74Chloroplast
CoKAS II-222412.536.1335.9783.150.01135.275.8144.4014.52Chloroplast Mitochondrion
CoKAS II-324795.128.8838.7685.18–013738.006.2640.7115.03Chloroplast
CoKAS II-4104584.888.0041.0685.04–0.11737.345.6842.2313.76Chloroplast
CoKAS II-5104024.276.9839.4177.24–0.14338.067.2139.5515.17Chloroplast
CoKAS II-6143183.346.2032.0182.89–0.01336.797.5538.9916.67Chloroplast
CoKAS III-1154054.265.5638.2689.560.03728.408.5139.5123.95Chloroplast
CoKAS III-2143944.135.4138.7288.860.02427.168.3840.3624.11Chloroplast
CoKAS III-3144004.246.2443.9691.250.00328.259.0038.7524.00Chloroplast
CoKAS III-4154004.246.5343.9691.970.02226.509.0040.5024.00Chloroplast
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油茶KAS基因家族的鉴定及表达分析
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刘晓霞 , 喻子昊 , 敬辉扬 , 熊凯峰 , 覃佐东 *
热带作物学报 | 组学与生物技术 2025,46(1): 10-20
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热带作物学报 |组学与生物技术 2025 , 46 (1) : 10 -20
油茶KAS基因家族的鉴定及表达分析
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刘晓霞, 喻子昊, 敬辉扬, 熊凯峰, 覃佐东*
作者信息
  • 湖南科技学院化学与生物工程学院/湖南省生物质资源综合开发利用工程技术研究中心，湖南永州　425199
通讯作者:
* 覃佐东（QIN Zuodong），E-mail：。
Identification and Expression Profile Analysis of KAS Gene Family in Camellia oleifera
Xiaoxia LIU, Zihao YU, Huiyang JING, Kaifeng XIONG, Zuodong QIN*
Affiliations
  • College of Chemistry and Bioengineering, Hunan University of Science and Engineering / Hunan Engineering Technology Research Center for Comprehensive Development and Utilization of Biomass Resources, Yongzhou, Hunan 425199, China
出版时间: 2025-01-25 doi: 10.3969/j.issn.1000-2561.2025.01.002
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摘要
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β-酮脂酰ACP合成酶（beta-ketoacyl-acyl carrier protein synthase，KAS）是植物脂肪酸生物合成过程中的关键酶系。为探究KAS在油茶种子成熟和油脂积累中的作用，本研究通过生物信息学方法对CoKAS基因家族成员进行鉴定，分析其理化性质、染色体定位、亚细胞定位、二级结构、进化关系、保守基序、启动子顺式作用元件，并采用RT-qPCR技术分析油茶果实油脂快速积累期的表达模式。结果表明：鉴定出的14个CoKAS家族成员分布在7条染色体上，相对分子质量在1.13~5.15 kDa之间，且大部分为酸性稳定蛋白。除CoKAS II-2定位于线粒体和叶绿体外，其余均定位于叶绿体。系统发育分析显示CoKAS基因被分为3个亚类，同一亚族成员的基因结构和保守基序具有相似性。CoKAS基因家族成员的启动子区域富含与生长发育、光响应、激素诱导和胁迫应答相关的顺式作用元件。通过表达分析发现，大部分CoKAS基因在油茶种子油脂快速积累期的表达量较高。此外，在油茶果实成熟过程中，CoKAS与CoMYB基因家族间存在有较强相关性的成员。本研究结果为深入解析CoKAS基因在油茶油脂生物合成中的调控机制提供理论基础。

油茶  /  CoKAS基因家族  /  生物信息学  /  表达分析

Beta-ketoacyl-acyl carrier protein synthase (KAS) is the key enzyme system in plant fatty acid biosynthesis. In order to investigate the function of the KAS gene family in the maturation and oil accumulation of Camellia oleifera, this study used bioinformatics methods and analyzed ther physicochemical properties, chromosomal localization, subcellular localization, secondary structure, phylogenetic trees, conserved motifs, promoter cis-acting elements, and RT-qPCR to analyze the expression patterns in the oil rapid accumulation stage of C. oleifera. The findings indicated that 14 Camellia oleifera KAS family members were distributed on seven chromosomes with relative molecular masses between 1.13 kDa and 5.15 kDa, and most of them were acid-stabilized proteins. Except for CoKAS II-2, which was localized in mitochondria and chloroplasts, all of them were localized in chloroplasts. Phylogenetic analysis revealed that CoKAS was divided into three subclasses, with similarities in structure and conserved motifs among KAS family numbers in the same subfamily. The promoter regions of the CoKAS gene family members were enriched with the cis-acting elements growth and development-related, light response, hormone induction, and adversity response. Expression analysis revealed that most of the CoKAS genes were highly expressed during the oil rapid accumulation stage. In addition, it was found a strong correlation between the members of the CoKAS and CoMYB gene families during fruit ripening in C. oleifera. The findings would provide a theoretical basis for comprehensively analyzing the function of CoKAS genes.

Camellia oleifera  /  CoKAS gene family  /  bioinformatics  /  expression analysis
刘晓霞, 喻子昊, 敬辉扬, 熊凯峰, 覃佐东. 油茶KAS基因家族的鉴定及表达分析. 热带作物学报, 2025 , 46 (1) : 10 -20 . DOI: 10.3969/j.issn.1000-2561.2025.01.002
Xiaoxia LIU, Zihao YU, Huiyang JING, Kaifeng XIONG, Zuodong QIN. Identification and Expression Profile Analysis of KAS Gene Family in Camellia oleifera[J]. Chinese Journal of Tropical Crops, 2025 , 46 (1) : 10 -20 . DOI: 10.3969/j.issn.1000-2561.2025.01.002
正文
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β-酮脂酰ACP合成酶（beta-ketoacyl-acyl carrier protein synthase，KAS）是植物脂肪酸从头合成过程中碳链延伸的关键酶系，催化脂酰基-CoA或脂酰基-ACP分子之间C-C键形成[1]。该酶由N端和C端蛋白结构域组成，活性位点位于这2个结构域之间。N端结构域包含参与二聚体形成的结构和活性位点半胱氨酸残基，与C端结构域的氨基酸残基协同作用，实现底物的识别、催化反应及产物的释放[2]。在植物中，已鉴定出KAS I、KAS II和KAS III三种KAS亚型，它们各司其职，共同协作完成脂肪酸链的延伸。KAS III是脂肪酸合成起始阶段的限速酶，以乙酰CoA和丙二酰CoA为底物，合成乙酰乙酰-ACP[3-4]。KAS II则主要负责将棕榈酸（C16:0-ACP）延伸为硬脂酸（C18:0-ACP），调控C16与C18脂肪酸的比例[5]。KAS I能特异性地识别C4至C16长度的酰基-ACP，并促进其碳链的延伸[6]。利用RNA干扰（RNA interference，RNAi）技术下调棉花种子中ghKAS2基因的表达，可以显著提高棉籽中棕榈酸（C16:0）的含量，并使其性状稳定遗传给后代，同时不影响种子的发芽率[5]。水稻OsKASI-2基因的敲除导致细胞膜脂肪酸的不饱和度降低，进而增强了水稻对低温的敏感性，影响其抗寒能力[6]。在拟南芥中，过表芝麻SiKASI可导致绒毡层细胞中油脂的异常积累，并与一种腺苷三磷酸结合盒转运蛋白（ABC transporter）相互作用，从而调控花粉的发育[7]。此外，将麻风树JcKASIII基因过表达于野生型以及KASI或KASII基因敲低的拟南芥突变体中，构建转基因植株，结果表明其种子油组成发生显著性变化[8]。综上表明，KAS基因家族成员在植物生长发育和胁迫应答中起着重要的作用。
油茶（Camellia oleifera）是中国特有的优质木本油料树种，广泛分布于长江流域至华南地区的山地丘陵，具有重要的经济和生态价值[9]。油茶籽油含高达80%以上的不饱和脂肪酸（主要为油酸和亚油酸）[10]，营养丰富，已被联合国粮农组织列为重点推广的健康型高级食用油。然而，现有油茶林产率较低，导致茶油价格居高不下，严重限制了油茶产业的健康发展。油脂积累是油茶籽品质形成的关键环节，揭示其分子机制对于培育高产优质油茶品种具有重要意义。本研究基于油茶果实不同发育时期的转录组数据，鉴定CoKAS基因家族成员，并对其理化性质、系统进化关系、共线性、启动子序列、基因结构和表达模式等进行系统分析，为进一步研究油茶KAS基因家族的功能研究奠定基础。
1　材料与方法
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1.1　油茶CoKAS家族成员鉴定
从NCBI数据库中下载油茶种子发育不同时期段的转录组数据（SRA登录号：PRJNA668531），油茶全基因组序列信息从GitHub（https://github.com/Hengfu-Yin/CON_genome_data）数据库中获取，利用Hisat2和StringTie软件获得转录本信息。从NCBI数据库（https://www.ncbi.nlm.nih.gov/protein）中下载拟南芥KAS蛋白序列，采用blast-2.15.0+搜索其在油茶中的同源序列，阈值为1e-10。从Pfam（http://pfam-legacy.xfam.org/）数据库中下载KAS结构域PF00109（N端）和PF02801（C端）的hmmr文件；使用hmmer 3.4软件对油茶KAS家族成员进行初步筛选，参数设置默认。选取上述2组结果共有的序列，采用NCBI数据库中的ORF finder（https://www.ncbi.nlm.nih.gov/orffinder/）在线工具预测候选序列的开放阅读框（open reading frame，ORF）。通过NCBI CD-Search（https://www.ncbi.nlm.nih.gov/Structure/bwrpsb/bwrpsb.cgi）、SMART（https://smart.embl.de/）和InterPro（https://www.ebi.ac.uk/interpro/）在线软件确定CoKAS家族成员，根据染色体定位信息进行命名。
利用ExPASy ProtParam（https://web.expasy.org/protparam/）在线软件对油茶CoKAS的基本理化性质进行分析。利用SOPMA（https://npsapbil.ibcp.fr/cgi-bin/npsa_automat.pl?page=npsa_sopma.html）在线软件预测CoKAS的二级结构。利用CELLO v.2.5（http://cello.life.nctu.edu.tw/）在线软件分析CoKAS的亚细胞定位。
1.2　CoKAS系统发育树构建
利用MEGA-X软件中Muscle算法对CoKAS和AtKAS蛋白序列进行多序列比对，使用邻近法（neighbor-joining method，NJ）构建系统进化树，bootstrap值设置为1000。
1.3　 CoKAS基因结构分析
利用MEME（https://meme-suite.org/meme/tools/meme）在线软件分析蛋白的保守基序（motif），设定基数为10，其他为默认参数。根据CoKAS基因家族的注释文件，利用GSDS 2.0（https://gsds.gao-lab.org/）在线软件进行基因结构的可视化分析。
1.4　 CoKAS启动子序列分析
利用TBtools软件提取CoKAS基因转录起始位点（ATG）上游2000 bp的启动子序列，利用PlantCARE（https://bioinformatics.psb.ugent.be/webtools/plantcare/html/）在线软件预测其顺式作用元件，利用R软件包对数据进行可视化分析。
1.5　 CoKAS基因在果实发育不同时期的表达模式分析
CoKAS基因家族成员的表达量以每千个碱基的转录每百万映射读取的fragments（fragments per kilobase of exon model per million mapped fragments，FPKM）计算。根据LI等[11]鉴定的CoMYB（Camellia oleifera myeloblastosis）家族成员筛选出亚细胞定位于细胞核的MYB蛋白，采用R软件包进行CoKAS和CoMYB相关性分析及数据可视化分析。
1.6　RT-qPCR分析
选取湘林210#油茶油脂快速积累期的种子，分别在授粉后256 d（S1）、275 d（S2）、303 d（S3）、330 d（S4）采集样品，每组3个生物学重复。根据候选CoKAS基因的CDS序列，通过NCBI的Primer3（https://www.ncbi.nlm.nih.gov/tools/primer-blast/index.cgi?LINK_LOC=BlastHome）在线软件设计引物（表1）。采用TRNzol法提取总RNA，使用FastKing cDNA第一链合成试剂盒（KR116）进行反转录，合成cDNA。以EF1a2为内参，利用SuperReal PreMix Plus（SYBR Green）进行RT-qPCR，CoKAS基因的相对表达量采用2-ΔΔCt法计算，分别采用SPSS 25.0和origin 2020软件进行数据处理和制图。
2　结果与分析
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2.1　CoKAS家族成员的特征分析
通过结合hmmer和blast方法，在油茶果实的转录组数据中共鉴定到14个CoKAS基因家族成员，包括4个KAS I、6个CoKAS II和4个CoKAS III（表2），它们不均匀地分布在油茶的7条染色体上，其中KAS I的4个家族成员均分布在4号染色体上。理化性质分析揭示，这些蛋白家族成员中氨基酸数量超过400个的成员占78.57%，其中相对分子质量最小的为1.13 kDa，最大的为5.15 kDa。理论等电点在4.54~8.89之间，大部分为酸性蛋白。不稳定系数在32.01~43.96之间，大部分为稳定蛋白。所有CoKAS蛋白的脂溶性指数均未超过100，其中有9个蛋白亲水指数小于0，说明64.29%是脂溶性亲水蛋白。在CoKAS蛋白的二级结构中，占比最高的是无规则卷曲，为44.07%（CoKAS I-2）~38.75%（CoKAS III-3）；占比最低的是β转角，以CoKAS I-4的9.35%为最高。亚细胞定位分析显示，除CoKAS II-2定位于线粒体和叶绿体外，其余均定位于叶绿体。
2.2　CoKAS蛋白系统进化关系分析
采用NJ法构建油茶和拟南芥KAS基因家族成员氨基酸序列系统发育树，结果表明，相对于KAS蛋白亚家族Ⅲ，家族Ⅰ和Ⅱ之间的亲缘关系更为接近（图1）。在第Ⅱ进化支中，除了AtKAS II与CoKAS II-1/2处于同一进化亚支外，CoKAS II-3/4/5/6在该分支中被独立分离出来。而在第Ⅰ和Ⅲ进化支中，CoKAS与AtKAS家族成员分别属于不同的进化亚支。综上所述，油茶中存在与拟南芥KAS进化关系较近的同源KAS蛋白，同时2个物种间也存在明显分化的KAS蛋白。
2.3　CoKAS家族基因结构、保守结构域和保守基序分析
为探究CoKAS家族成员基因的结构特征，对其外显子与内含子的组成进行分析。结果揭示CoKAS家族成员间基因结构的多样性，内含子数量在1~13之间，尤其CoKAS II-1/2的内含子数量高达13。在同一亚族内，成员之间的外显子-内含子结构呈现出较高的相似性，如CoKAS III亚族所有成员所含7个内含子的位置大体一致（图2A）。这些发现表明，亚家族间基因结构的差异化可能与新功能的演化紧密相关。通过保守结构域分析发现，所有成员均展现出KAS蛋白的典型特征，包含有cond_enzymes结构域，而PLN02326是CoKAS III亚家族特有的保守结构域（图2B）。CoKAS家族成员的motif从2到10个不等，其中motif 1/2/6基序组成了KAS基因家族典型的保守区域（图2C，图2D）。motif 1具备还原酶活性FabB区域，存在于所有的CoKAS蛋白中；除CoKAS I-4外，CoKAS家族所有成员均含有motif 2基序；而motif 6基序仅在CoKAS I/II亚家族中被发现。CoKAS I和II两个亚家族成员间motif的空间分布、类型和数量具有较高的一致性，但与CoKAS III家族成员存在一定的差异。然而，在CoKAS I和II两个亚家族中，也有个别成员如CoKAS I-4和CoKAS II-2的motif数量相对缺失，这可能与基因串联重复过程中碱基的丢失有关。
2.4　 CoKAS启动子顺式作用元件分析
通过对CoKAS家族基因启动子上游2000 bp序列的顺式作用元件进行分析，鉴定到大量与生长发育、激素、光及逆境响应相关的顺式作用元件（图3）。在所有的生长发育响应元件中，数量最多的是所有成员都具有的MYB和MYC转录因子结合位点。CoKAS启动子序列含有3~22个光调控和响应元件，其中85.71%的成员具有Box 4元件。CoKAS启动子序列含有7~18个胁迫响应元件，除CoKAS I-3外，其他成员都含有响应厌氧诱导的ARE元件。CoKAS家族成员主要对赤霉素、乙烯、茉莉酸甲酯、水杨酸和生长素产生响应，其中71.43%成员对赤霉素有响应，78.57%成员对乙烯有响应。综上所述，CoKAS在油茶生长发育、激素诱导、光诱导和逆境胁迫响应等生物学过程中可能发挥着重要的作用。
2.5　油茶果实发育过程中CoKAS基因的表达模式
基因的表达对其功能的发挥起着至关重要的作用。为研究CoKAS基因在油茶油脂合成中的功能，构建了CoKAS基因家族成员在果实不同发育时期的表达谱（图4）。除CoKAS III亚族外，第I和II亚族均有成员在油茶果实整个发育阶段的表达量较低，其中第II亚族的比例最大，占全部基因的66.67%。随着油茶果实的成熟，有6个CoKAS基因家族成员的表达趋势与种子油含量逐渐增加趋势相同[12]，但CoKAS II-3的表达模式却呈现出相反的趋势。另外，有2个基因在T3阶段高表达，CoKAS I-3的表达量先上升后下降，而CoKAS III-4的表达量先上升后保持不变。这些结果表明，CoKAS基因家族在油茶果实油脂积累过程中可能发挥着重要的作用。为深入探讨CoKAS基因在油茶种子油脂累积过程中的功能，本研究通过RT-qPCR技术，对8个CoKAS基因在湘林210#油茶种子油脂快速累积期的表达水平进行了检测。结果显示，其中5个CoKAS基因的表达与转录组测序数据基本一致（图5）。此外，除CoKAS III-1基因表现出相反的表达趋势外，CoKAS II-5和CoKAS II-6基因在早期阶段呈现出与转录组分析结果不一致的表达模式，但在后期逐渐趋于一致。这种差异可能是由于湘林210#与华硕2个油茶品种间基因组变异或发育调控机制的不同所致。
由图3可知，CoKAS启动子中含有4~20个转录因子MYB的结合位点，这表明该家族成员可能更容易受到MYB的调控。为探索CoMYB转录因子和CoKAS家族成员的结合情况，本研究对定位于细胞核的124个CoMYB与14个CoKAS进行了关联分析，并计算了Pearson相关系数（图6）。结果显示，有7个CoKAS家族成员可能受到21个转录因子CoMYB的调控，其中CoKAS II-6的转录因子CoMYB最多，高达13个。CoMYB46能调控除CoKAS I-4外的其他6个成员基因的表达，而CoMYB85仅调控CoKAS II-3的表达。此外，还发现不同CoKAS基因家族成员可能同时受多种CoMYB的调控，例如CoMYB18、CoMYB46、CoMYB68、CoMYB83与CoKAS I-1、CoKAS II-3、CoKAS II-5、CoKAS III-1、CoKAS III-2都具有较强的相关性（|r|≥0.85），除CoKAS II-3外，其他成员为负相关。以上结果表明，CoMYB可能通过多样的调控网络调节CoKAS基因的表达。
3　讨论
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KAS作为脂肪酸合酶的关键组成部分，是一种高度保守的酶，广泛存在于地球上的几乎所有生命体中。其家族成员已在多种植物中被鉴定出来，并在植物的发育转变和环境响应中发挥着重要的作用[5,7-8]。然而，关于油茶KAS基因家族成员的鉴定及其功能的研究却鲜有报道。本研究采用同源性比对的方法，成功鉴定到14个CoKAS家族成员，它们定位在油茶的7条染色体上。CoKAS蛋白同一亚族内的理化性质和二级结构相近，这可能与其功能的保守性有关。亚细胞定位结果显示，14个CoKAS中有13个定位于叶绿体，这与叶绿体是脂肪酸合成的主要场所相符[13]。而只有CoKAS II-2定位于线粒体和叶绿体，这可能与其作为双重定位酶的特性有关，它参与线粒体中的脂质合成，尽管这些脂质只占总脂质的一小部分，但它们在调节植物生长和发育方面发挥了重要作用[14-15]。在进化关系上，油茶与拟南芥的KAS同被分为3个亚类，不同分支行使的功能各异。通过对CoKAS家族成员的基因结构进行分析发现，同一亚族成员的外显子-内含子数量及分布、结构域和保守基序特征相似，这可能与同一亚族CoKAS成员的生物学功能相似有关。基因的进化过程中会出现内含子增加或缺失现象[16]，这可能是导致个别基因（CoKAS I-1、CoKAS II-1、CoKAS II-2和CoKAS I-4）外显子-内含子结构发生变异的原因。
启动子区的顺式作用元件对基因表达具有至关重要的调控作用[17]。油茶CoKAS家族基因的启动子区富集大量生长发育相关、光响应、激素诱导和胁迫应答元件，表明CoKAS基因功能多样，能参与不同的生物学过程。乙烯是调控植物生长与发育的关键激素，一方面乙烯可作为内源调节剂刺激植物分生组织生长和胚乳分裂[18]；另一方面，过量的乙烯则抑制分生组织扩张和胚乳细胞分裂速率[19]。研究发现，超长链脂肪酸（very-long-chain fatty acids，VLCFAs）能通过增强乙烯生物合成促进植物分生组织的伸长[20]。过表达OskasI也能使拟南芥根伸长，而缺乏KAS I时会显著降低种子中脂肪酸的水平[21]。此外，KAS I在植物的种子、根、花和幼苗中均有表达[22]。在根中，KAS I主要在分生组织、伸长区和成熟区的维管束中表达。在子叶胚发育的早期阶段，胚乳细胞迅速增殖，且乙烯生物合成的关键酶1-氨基环丙烷-1-羧酸合成酶（ACC合成酶）和KAS I基因表现出较高的表达水平。然而，在子叶胚发育的后期，高浓度的乙烯会促进胚乳细胞程序性死亡，与此同时，KAS I的表达水平有所下降[22-23]。本研究观察到，CoKAS I-2和CoKAS I-3在油茶种子发育过程中的表达量呈现先上升后下降的趋势，其中CoKAS I-2的表达量相对较低。A-box和GCN4_motif这2个参与分生组织（CoKAS I-2）和胚乳（CoKAS I-3）特异性表达的元件，仅在CoKAS I亚家族成员的启动子区域被发现，且这些基因包含乙烯响应元素。据此推测，CoKAS I-2和CoKAS I-3可能通过调节乙烯信号途径，分别参与油茶根系的生长和种子胚乳的发育，而乙烯对CoKAS I基因家族的表达具有负向调控作用。
在植物种子的灌浆中期至末期，光照强度的增加导致种子中亚油酸、棕榈酸和亚麻酸含量上升，而油酸含量则相应降低。然而，在灌浆末期至成熟期，随着光照强度的逐步减弱，油酸含量开始逐渐增加，与此同时，亚油酸、棕榈酸和亚麻酸含量则呈现出下降趋势[24]。研究揭示，在植物种子的灌浆阶段，高强度光照可能对种子质量和产量产生负面影响。相比之下，适宜的遮光处理能有效提高种子重量，并增加不饱和脂肪酸含量，特别是油酸[25-26]。本研究进一步发现，油茶CoKAS基因家族成员的启动子区域含有众多光响应元件，并在灌浆末期表达水平显著变化的成员较多，推测CoKAS基因家族在光信号调控油茶种子油脂积累和脂肪酸组成中起到关键作用。MYB在植物生长发育及胁迫响应中扮演着至关重要的角色。大量的证据证实，MYB转录因子在植物体内油脂积累的过程中起着关键作用。例如，麻风树中的JcMYB1能够直接促进脂肪酸合成的关键酶二酰基甘油酰基转移酶1（diacylglycerol acyltransferase1，DGAT1）的表达，从而增加油脂含量[27]。油桐中，VfMYB36的过量表达能够显著增强拟南芥中与油脂合成相关的基因AtWRI、AtENO1、AtBCCP1、AtKAS1、AtKCS11和AtPAL2的活性，导致种子中亚麻酸和总油量的提升[28]。然而，在脂质代谢中，MYB转录因子更常见的角色是作为抑制因子。在拟南芥种子的脂质生物合成过程中，MYB89通过直接结合启动子位点抑制WRI1和KASI的表达，并间接抑制L1L等关键基因的活性，从而负向调控油脂的合成[29]；MYB76则通过增强脂肪酸降解酶相关基因的表达来实现对脂肪酸的负向调控[30]；而MYB118则通过调控成熟相关基因的活性来抑制胚乳中油脂的合成[31]。在本项研究中，CoKAS家族成员基因的启动子区域含有多个MYB结合位点，通过基因表达关联性分析，发现CoKAS与CoMYB家族成员间存在高度相关性，其中大部分呈现负相关性。上述结果表明，CoMYB可能通过与CoKAS基因启动子区的顺式作用元件结合，从而调控油茶种子中油脂积累和脂肪酸组成。
4　结论
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本研究基于油茶果实不同发育阶段的转录组数据，系统鉴定了14个CoKAS基因家族成员，这些基因分布于7条染色体上。通过系统进化分析，CoKAS基因被分为3个亚类，同一亚类成员的基因结构和保守基序高度保守。在CoKAS基因家族成员的启动子区域，鉴定到大量与生长发育、光响应、激素信号转导和胁迫响应相关的顺式作用元件。此外，表达模式分析显示，大部分CoKAS基因家族成员在油茶果实快速积累油脂期表达量显著上调。进一步的共表达网络分析表明，CoKAS I-1、CoKAS II-3、CoKAS II-5、CoKAS III-1和CoKAS III-2与CoMYB转录因子家族成员存在显著的相关性。这些结果为深入解析CoKAS基因在油茶籽油生物合成中的调控机制提供理论基础。
基金
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  • 湖南省自然科学基金项目(2024JJ7188)
  • 湖南省教育厅科学研究项目(23C0354)
  • 湖南科技学院科学研究项目(23XKY07)
文献
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2025年第46卷第1期
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doi: 10.3969/j.issn.1000-2561.2025.01.002
  • 接收时间：2024-07-23
  • 首发时间：2026-06-24
  • 出版时间：2025-01-25
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  • 收稿日期：2024-07-23
  • 修回日期：2024-08-19
基金
湖南省自然科学基金项目(2024JJ7188)
湖南省教育厅科学研究项目(23C0354)
湖南科技学院科学研究项目(23XKY07)
作者信息
    湖南科技学院化学与生物工程学院/湖南省生物质资源综合开发利用工程技术研究中心，湖南永州　425199

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* 覃佐东（QIN Zuodong），E-mail：。
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https://castjournals.cast.org.cn/joweb/rdzwxb/CN/10.3969/j.issn.1000-2561.2025.01.002
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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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