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Article(id=1190335350317875419, tenantId=1146029695717560320, journalId=1189982191388893191, issueId=1190335347767743264, articleNumber=null, orderNo=null, doi=10.16438/j.0513-4870.2024-1147, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1731945600000, receivedDateStr=2024-11-19, revisedDate=1736870400000, revisedDateStr=2025-01-15, acceptedDate=null, acceptedDateStr=null, onlineDate=1761727662876, onlineDateStr=2025-10-29, pubDate=1744387200000, pubDateStr=2025-04-12, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1761727662876, onlineIssueDateStr=2025-10-29, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1761727662876, creator=13701087609, updateTime=1761727662876, updator=13701087609, issue=Issue{id=1190335347767743264, tenantId=1146029695717560320, journalId=1189982191388893191, year='2025', volume='60', issue='4', pageStart='843', pageEnd='1182', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1761727662269, creator=13701087609, updateTime=1761729313427, updator=13701087609, preIssue=null, nextIssue=null, ext={EN=IssueExt(id=1190342273276678997, tenantId=1146029695717560320, journalId=1189982191388893191, issueId=1190335347767743264, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1190342273276678998, tenantId=1146029695717560320, journalId=1189982191388893191, issueId=1190335347767743264, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null}, startPage=1156, endPage=1165, ext={EN=ArticleExt(id=1190335350573727964, articleId=1190335350317875419, tenantId=1146029695717560320, journalId=1189982191388893191, language=EN, title=Genome-wide identification and functional analysis of the JAZ gene family in Tripterygium wilfordii, columnId=1190335348761793317, journalTitle=Acta Pharmaceutica Sinica, columnName=Original Articles, runingTitle=null, highlight=null, articleAbstract=

JAZ proteins, as inhibitors in the jasmonic acid (JA) signaling pathway, play a crucial role in regulating plant growth and development, as well as the biosynthesis of secondary metabolites in plants. To explore the functions of the JAZ gene family in Tripterygium wilfordii, our study systematically identified the members of the JAZ gene family in T. wilfordii. We analyzed their physicochemical properties, chromosome localization, phylogenetic tree, and protein conserved motifs. Additionally, we investigated the expression patterns of TwJAZs in different tissues, constructed a gene regulatory network map, and ultimately cloned the full-length sequences of 18 JAZ genes. The results showed that the 18 TwJAZs were unevenly distributed across 12 chromosomes, encoding amino acid numbers ranging from 122 to 525, with molecular weights of 13.87 to 54.73 kDa, and isoelectric points ranging from 6.83 to 10.27. Subcellular localization prediction indicated that 13 TwJAZs were localized in the nucleus, while 5 TwJAZs were located in the cytoplasm and chloroplasts. The phylogenetic tree analysis revealed that the JAZ family proteins of T. wilfordii could be divided into five subfamilies, with members of the same subfamily sharing similar conserved motifs. Expression pattern analysis demonstrated that TwJAZs were predominantly expressed in flowers, leaves, and peeled stems, with most TwJAZs showing the highest expression levels after 4 hours of methyl jasmonate (MeJA) induction. The transcription factor regulatory network shows that TwJAZs are strongly correlated with transcription factor families such as AP2/ERF-ERF, NAC, and bHLH. This study comprehensively identified all the JAZ gene family sequences in T. wilfordii, initially clarifying the structural and functional characteristics of the TwJAZs, and laying an important foundation for further research on the functions and regulatory mechanisms of TwJAZs.

, correspAuthors=Li-chan TU, authorNote=null, correspAuthorsNote=null, copyrightStatement=Copyright ©2025 Acta Pharmaceutica Sinica. All rights reserved., 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, authorCompany=null, fund=null, authors=null, authorsList=Ke-yan WANG, Li-mei TANG, Lin-jie WU, Yu-ting WANG, Xia-wei CHEN, Li-chan TU), CN=ArticleExt(id=1190336159805965273, articleId=1190335350317875419, tenantId=1146029695717560320, journalId=1189982191388893191, language=CN, title=药用植物雷公藤JAZ基因家族的全基因组鉴定及功能分析, columnId=1190335348896011050, journalTitle=药学学报, columnName=研究论文, runingTitle=null, highlight=null, articleAbstract=

JAZ蛋白作为茉莉酸途径的抑制因子, 对植物生长发育、植物次级代谢产物合成的调控具有重要作用。为研究雷公藤(Tripterygium wilfordii) JAZ基因家族的功能, 本研究系统鉴定了雷公藤JAZ基因家族成员, 分析其理化性质、基因染色体定位、系统进化树、蛋白质保守基序, 同时通过雷公藤转录组测序数据分析TwJAZs基因在不同部位的表达模式, 构建基因网络调控图谱, 最终基因克隆获得18条TwJAZs基因的全长序列。结果表明, 18个TwJAZs基因不均匀地分布在12条染色体上, 其编码的氨基酸数量为122~525个, 分子质量为13.87~54.73 kDa, 等电点为6.83~10.27。亚细胞预测显示13个TwJAZs蛋白定位在细胞核上, 5个位于细胞质和叶绿体上。系统进化树结果表明雷公藤JAZ家族蛋白可分为5个亚家族, 同一亚家族成员具有相似的保守基序。表达模式分析结果表明TwJAZs基因主要在花、叶、去皮茎中具有较高的表达, 且大部分TwJAZs基因在茉莉酸甲酯(MeJA) 诱导4 h后的表达量最高。转录因子调控网络显示TwJAZs主要与AP2/ERF-ERF、NAC、bHLH等转录因子家族呈现强相关。本研究全面鉴定获得雷公藤中所有的JAZ基因家族序列, 初步明确了TwJAZs基因家族的结构和功能特点, 为深入研究TwJAZs蛋白功能和调控机制奠定重要基础。

, correspAuthors=屠李婵, authorNote=null, correspAuthorsNote=
屠李婵, E-mail:
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J Fruit Sci (果树学报), 2023, 40: 1294-1311., articleTitle=null, refAbstract=null)], funds=[Fund(id=1190350204827177824, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1190335350317875419, awardId=82204559, language=CN, fundingSource=国家自然科学基金资助项目(82204559), fundOrder=null, country=null), Fund(id=1190350204936229729, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1190335350317875419, awardId=LQ23H280002, language=CN, fundingSource=浙江省自然科学基金资助项目(LQ23H280002), fundOrder=null, country=null)], companyList=[AuthorCompany(id=1190350201404625701, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1190335350317875419, xref=null, ext=[AuthorCompanyExt(id=1190350201413014310, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1190335350317875419, companyId=1190350201404625701, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=School of Medicine, Hangzhou City University, Hangzhou 310015, China), AuthorCompanyExt(id=1190350201421402919, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1190335350317875419, companyId=1190350201404625701, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=浙大城市学院医学院, 浙江 杭州 310015)])], figs=[ArticleFig(id=1190350203623412560, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1190335350317875419, language=EN, label=null, caption=null, figureFileSmall=gHZqhry8w1ijfTJjWAnJvQ==, figureFileBig=4ZbLeh7fGfDc7uPWlO14wg==, tableContent=null), ArticleFig(id=1190350203686327121, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1190335350317875419, language=CN, label=Figure 1, caption=

Chromosome localization of JAZ genes in T. wilfordii. Tw: T. wilfordii

, figureFileSmall=gHZqhry8w1ijfTJjWAnJvQ==, figureFileBig=4ZbLeh7fGfDc7uPWlO14wg==, tableContent=null), ArticleFig(id=1190350203761824594, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1190335350317875419, language=EN, label=null, caption=null, figureFileSmall=bgorouh6bLS+SUprpmFd/w==, figureFileBig=gCcwqbpmqGOWvYxFueo3fQ==, tableContent=null), ArticleFig(id=1190350203816350547, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1190335350317875419, language=CN, label=Figure 2, caption=

Phylogenetic tree of JAZ proteins in T. wilfordii. At: Arabidopsis thaliana; Sm: Salvia miltiorrhiza; Vv: Vitis vinifera; Sl: Solanum lycopersicum

, figureFileSmall=bgorouh6bLS+SUprpmFd/w==, figureFileBig=gCcwqbpmqGOWvYxFueo3fQ==, tableContent=null), ArticleFig(id=1190350203896042324, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1190335350317875419, language=EN, label=null, caption=null, figureFileSmall=L9vbluW10RdJQ3i4/0QH0A==, figureFileBig=dgh3RN4pn2aghXnD4ThErA==, tableContent=null), ArticleFig(id=1190350203958956885, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1190335350317875419, language=CN, label=Figure 3, caption=

Gene structure of JAZ genes in T. wilfordii

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Conserved motifs of JAZ family members in T. wilfordii

, figureFileSmall=m9YRvt38d5GaqLgOI988lg==, figureFileBig=+FRQZB4HJPOYuPBMEFhsLg==, tableContent=null), ArticleFig(id=1190350204147700568, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1190335350317875419, language=EN, label=null, caption=null, figureFileSmall=Fi/m9hnX/FFwdXCiOExClQ==, figureFileBig=7jcMsjRCMC9FrfKrzrAeiA==, tableContent=null), ArticleFig(id=1190350204210615129, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1190335350317875419, language=CN, label=Figure 5, caption=

Expression patterns of TwJAZs. A: Expression patterns of TwJAZs induced by MeJA in suspension cells of T. wilfordii. Tw_0 h means that the suspension cells of T. wilfordii were collected for 0 h without any treatment. CK-4 h, CK-12 h, CK-24 h, CK-48 h, CK-72 h, CK-120 h, CK-240 h means that the T. wilfordii suspension cells were collected after control treatment for 4, 12, 24, 48, 72, 120, 240 h. MJ-4 h, MJ-12 h, MJ-24 h, MJ-48 h, MJ-72 h, MJ-120 h, MJ-240 h means that the T. wilfordii suspension cells were collected after induction treatment with MeJA for 4, 12, 24, 48, 72, 120 and 240 h, respectively. B: Expression patterns of TwJAZs in different tissues of T. wilfordii. Tw_F: Flower; Tw_L: Leaf; Tw_PS: Peeled stem; Tw_SB: Stem bark; Tw_RB: Root bark; Tw_RP: Root phloem; Tw_RX: Root xylem

, figureFileSmall=Fi/m9hnX/FFwdXCiOExClQ==, figureFileBig=7jcMsjRCMC9FrfKrzrAeiA==, tableContent=null), ArticleFig(id=1190350204277723994, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1190335350317875419, language=EN, label=null, caption=null, figureFileSmall=woi6nTC3cRMfb0j8iC02wQ==, figureFileBig=8PrQ+zXGXDGD+I044IMHFQ==, tableContent=null), ArticleFig(id=1190350204332249947, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1190335350317875419, language=CN, label=Figure 6, caption=

Regulatory network of JAZ and other transcription factors family of T. wilfordii. Orange lines represent r > 0.8, and green lines represent r < -0.8. The darker the color, the stronger the correlation

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Gene name Forward primer (5′-3′) Reverse primer (5′-3′)
TwJAZ1 ATGGAGAGAGATTTTCTGGGACTGG CTAATGTGTAGCCGGAAGAGGATAGG
TwJAZ2 ATGTCGAATTTTGCTCAAACC TTAATTCATGGACTCTTGCTTTCTC
TwJAZ3 ATGGAGAGAGATTTTCTGGGACTGG TCAGCATTCCCGTTGATTGCTAATT
TwJAZ4 ATGTCGACCGCCACGGA TCACTGTGGTGATGGAACC
TwJAZ5 ATGGAGAGAGATTTCTTGGGTTTGA TCATGGATTCATGGCCAGCAGTTC
TwJAZ6 ATGTCGAGCTCGACGGAAAGGTC TCACTGTGGTGATGGAGCCATCAAG
TwJAZ7 ATGTCAAGAGCACCTGTGGAGCTCG TTAGGACTGGCAAGCAAACGGCG
TwJAZ8 ATGGCAGGACCTGAAGTGAAG TCAAGCAGCCTGAGGTTCATTAGA
TwJAZ9 ATGTCGACCGGGACGGAAAGG TCACTCCGGTGATAAGCCAAGCCA
TwJAZ10 ATGGATAGAGCACCAGTGGAGCTCG CTAGCAAGCAAAGGGCGACACAGAT
TwJAZ11 ATGGAGATAGATTTCTTGGGTT TCACGGATTCATGGCTAGC
TwJAZ12 ATGATGAGATCAAGGAAGTGC TTAATTAACAATGTATGGGGA
TwJAZ13 ATGAGAGCAATGAAGTGCAATCTTG TTAATTAACAGTGTATGGGGATTTT
TwJAZ14 ATGGACACTGGAGCCACGC TTATTCTCCAGCAGCAAATGAGCAG
TwJAZ15 ATGGACACCGAAGCCATGCCGT CTAAGGTAGATGGCCTTCCAGGCTT
TwJAZ16 ATGAAAAGGAATTGCAATCTTGAAC CTATTGATTGAAATGATGATAGGGG
TwJAZ17 ATGAAAAGGAATTGCAATCTTGAAC CTATTGACTCGGGCGATGATAGG
TwJAZ18 ATGAGTCCAAACTGCAATCTTG CTAGTGATTAAAGGTATATGGAAAT
), ArticleFig(id=1190350204487439197, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1190335350317875419, language=CN, label=Table 1, caption=

Primers for gene cloning

, figureFileSmall=null, figureFileBig=null, tableContent=
Gene name Forward primer (5′-3′) Reverse primer (5′-3′)
TwJAZ1 ATGGAGAGAGATTTTCTGGGACTGG CTAATGTGTAGCCGGAAGAGGATAGG
TwJAZ2 ATGTCGAATTTTGCTCAAACC TTAATTCATGGACTCTTGCTTTCTC
TwJAZ3 ATGGAGAGAGATTTTCTGGGACTGG TCAGCATTCCCGTTGATTGCTAATT
TwJAZ4 ATGTCGACCGCCACGGA TCACTGTGGTGATGGAACC
TwJAZ5 ATGGAGAGAGATTTCTTGGGTTTGA TCATGGATTCATGGCCAGCAGTTC
TwJAZ6 ATGTCGAGCTCGACGGAAAGGTC TCACTGTGGTGATGGAGCCATCAAG
TwJAZ7 ATGTCAAGAGCACCTGTGGAGCTCG TTAGGACTGGCAAGCAAACGGCG
TwJAZ8 ATGGCAGGACCTGAAGTGAAG TCAAGCAGCCTGAGGTTCATTAGA
TwJAZ9 ATGTCGACCGGGACGGAAAGG TCACTCCGGTGATAAGCCAAGCCA
TwJAZ10 ATGGATAGAGCACCAGTGGAGCTCG CTAGCAAGCAAAGGGCGACACAGAT
TwJAZ11 ATGGAGATAGATTTCTTGGGTT TCACGGATTCATGGCTAGC
TwJAZ12 ATGATGAGATCAAGGAAGTGC TTAATTAACAATGTATGGGGA
TwJAZ13 ATGAGAGCAATGAAGTGCAATCTTG TTAATTAACAGTGTATGGGGATTTT
TwJAZ14 ATGGACACTGGAGCCACGC TTATTCTCCAGCAGCAAATGAGCAG
TwJAZ15 ATGGACACCGAAGCCATGCCGT CTAAGGTAGATGGCCTTCCAGGCTT
TwJAZ16 ATGAAAAGGAATTGCAATCTTGAAC CTATTGATTGAAATGATGATAGGGG
TwJAZ17 ATGAAAAGGAATTGCAATCTTGAAC CTATTGACTCGGGCGATGATAGG
TwJAZ18 ATGAGTCCAAACTGCAATCTTG CTAGTGATTAAAGGTATATGGAAAT
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Gene name Gene accession number Subcellular localization Amino acids length Molecular mass/Da pI Instability index GRAVY Aliphatic index
TwJAZ1 XM_038823720.1 Nucleus 475 50 000.36 9.48 59.00 -0.311 71.37
TwJAZ2 XM_038825238.1 Chloroplast 188 20 507.21 9.30 47.77 -0.570 61.81
TwJAZ3 XM_038832473.1 Nucleus 525 54 734.12 9.63 56.77 -0.340 72.76
TwJAZ4 XM_038839784.1 Nucleus 267 29 396.56 8.76 60.22 -0.415 70.56
TwJAZ5 XM_038841851.1 Cytoplasm 372 39 398.01 9.52 46.19 -0.215 76.64
TwJAZ6 XM_038842845.1 Nucleus 243 26 511.32 8.92 60.64 -0.379 67.08
TwJAZ7 XM_038847386.1 Nucleus 199 21 995.15 9.54 69.42 -0.524 72.96
TwJAZ8 XM_038855066.1 Chloroplast 210 22 335.41 8.39 54.65 -0.421 73.90
TwJAZ9 XM_038856651.1 Chloroplast 259 28 336.48 7.55 48.04 -0.479 71.20
TwJAZ10 XM_038856882.1 Nucleus 197 21 779.84 9.52 55.87 -0.538 79.64
TwJAZ11 XM_038862888.1 Cytoplasm 309 32 640.27 7.87 39.04 -0.183 76.47
TwJAZ12 XM_038865522.1 Nucleus 130 15 173.15 9.50 85.68 -0.877 64.46
TwJAZ13 XM_038866766.1 Nucleus 122 13 867.57 9.14 71.03 -0.772 71.89
TwJAZ14 XM_038863286.1 Nucleus 346 37 057.45 8.62 48.86 -0.566 66.24
TwJAZ15 XM_038843897.1 Nucleus 301 32 761.85 8.70 53.08 -0.644 64.78
TwJAZ16 XM_038864546.1 Nucleus 122 13 984.79 8.91 52.35 -0.765 79.02
TwJAZ17 XM_038868010.1 Nucleus 127 14 451.3 10.27 66.92 -0.746 82.13
TwJAZ18 XM_038864545.1 Nucleus 139 15 630.34 6.83 86.73 -0.783 55.40
), ArticleFig(id=1190350204651017055, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1190335350317875419, language=CN, label=Table 2, caption=

Analysis of the physicochemical properties and subcellular localization of JAZ gene family in T. wilfordii

, figureFileSmall=null, figureFileBig=null, tableContent=
Gene name Gene accession number Subcellular localization Amino acids length Molecular mass/Da pI Instability index GRAVY Aliphatic index
TwJAZ1 XM_038823720.1 Nucleus 475 50 000.36 9.48 59.00 -0.311 71.37
TwJAZ2 XM_038825238.1 Chloroplast 188 20 507.21 9.30 47.77 -0.570 61.81
TwJAZ3 XM_038832473.1 Nucleus 525 54 734.12 9.63 56.77 -0.340 72.76
TwJAZ4 XM_038839784.1 Nucleus 267 29 396.56 8.76 60.22 -0.415 70.56
TwJAZ5 XM_038841851.1 Cytoplasm 372 39 398.01 9.52 46.19 -0.215 76.64
TwJAZ6 XM_038842845.1 Nucleus 243 26 511.32 8.92 60.64 -0.379 67.08
TwJAZ7 XM_038847386.1 Nucleus 199 21 995.15 9.54 69.42 -0.524 72.96
TwJAZ8 XM_038855066.1 Chloroplast 210 22 335.41 8.39 54.65 -0.421 73.90
TwJAZ9 XM_038856651.1 Chloroplast 259 28 336.48 7.55 48.04 -0.479 71.20
TwJAZ10 XM_038856882.1 Nucleus 197 21 779.84 9.52 55.87 -0.538 79.64
TwJAZ11 XM_038862888.1 Cytoplasm 309 32 640.27 7.87 39.04 -0.183 76.47
TwJAZ12 XM_038865522.1 Nucleus 130 15 173.15 9.50 85.68 -0.877 64.46
TwJAZ13 XM_038866766.1 Nucleus 122 13 867.57 9.14 71.03 -0.772 71.89
TwJAZ14 XM_038863286.1 Nucleus 346 37 057.45 8.62 48.86 -0.566 66.24
TwJAZ15 XM_038843897.1 Nucleus 301 32 761.85 8.70 53.08 -0.644 64.78
TwJAZ16 XM_038864546.1 Nucleus 122 13 984.79 8.91 52.35 -0.765 79.02
TwJAZ17 XM_038868010.1 Nucleus 127 14 451.3 10.27 66.92 -0.746 82.13
TwJAZ18 XM_038864545.1 Nucleus 139 15 630.34 6.83 86.73 -0.783 55.40
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药用植物雷公藤JAZ基因家族的全基因组鉴定及功能分析
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王柯燕 , 唐丽媚 , 吴林捷 , 王雨婷 , 陈夏唯 , 屠李婵 *
药学学报 | 研究论文 2025,60(4): 1156-1165
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药学学报 | 研究论文 2025, 60(4): 1156-1165
药用植物雷公藤JAZ基因家族的全基因组鉴定及功能分析
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王柯燕, 唐丽媚, 吴林捷, 王雨婷, 陈夏唯, 屠李婵*
作者信息
  • 浙大城市学院医学院, 浙江 杭州 310015

通讯作者:

屠李婵, E-mail:
Genome-wide identification and functional analysis of the JAZ gene family in Tripterygium wilfordii
Ke-yan WANG, Li-mei TANG, Lin-jie WU, Yu-ting WANG, Xia-wei CHEN, Li-chan TU*
Affiliations
  • School of Medicine, Hangzhou City University, Hangzhou 310015, China
出版时间: 2025-04-12 doi: 10.16438/j.0513-4870.2024-1147
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摘要
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JAZ蛋白作为茉莉酸途径的抑制因子, 对植物生长发育、植物次级代谢产物合成的调控具有重要作用。为研究雷公藤(Tripterygium wilfordii) JAZ基因家族的功能, 本研究系统鉴定了雷公藤JAZ基因家族成员, 分析其理化性质、基因染色体定位、系统进化树、蛋白质保守基序, 同时通过雷公藤转录组测序数据分析TwJAZs基因在不同部位的表达模式, 构建基因网络调控图谱, 最终基因克隆获得18条TwJAZs基因的全长序列。结果表明, 18个TwJAZs基因不均匀地分布在12条染色体上, 其编码的氨基酸数量为122~525个, 分子质量为13.87~54.73 kDa, 等电点为6.83~10.27。亚细胞预测显示13个TwJAZs蛋白定位在细胞核上, 5个位于细胞质和叶绿体上。系统进化树结果表明雷公藤JAZ家族蛋白可分为5个亚家族, 同一亚家族成员具有相似的保守基序。表达模式分析结果表明TwJAZs基因主要在花、叶、去皮茎中具有较高的表达, 且大部分TwJAZs基因在茉莉酸甲酯(MeJA) 诱导4 h后的表达量最高。转录因子调控网络显示TwJAZs主要与AP2/ERF-ERF、NAC、bHLH等转录因子家族呈现强相关。本研究全面鉴定获得雷公藤中所有的JAZ基因家族序列, 初步明确了TwJAZs基因家族的结构和功能特点, 为深入研究TwJAZs蛋白功能和调控机制奠定重要基础。

雷公藤  /  JAZ  /  生物信息学分析  /  表达模式  /  基因克隆

JAZ proteins, as inhibitors in the jasmonic acid (JA) signaling pathway, play a crucial role in regulating plant growth and development, as well as the biosynthesis of secondary metabolites in plants. To explore the functions of the JAZ gene family in Tripterygium wilfordii, our study systematically identified the members of the JAZ gene family in T. wilfordii. We analyzed their physicochemical properties, chromosome localization, phylogenetic tree, and protein conserved motifs. Additionally, we investigated the expression patterns of TwJAZs in different tissues, constructed a gene regulatory network map, and ultimately cloned the full-length sequences of 18 JAZ genes. The results showed that the 18 TwJAZs were unevenly distributed across 12 chromosomes, encoding amino acid numbers ranging from 122 to 525, with molecular weights of 13.87 to 54.73 kDa, and isoelectric points ranging from 6.83 to 10.27. Subcellular localization prediction indicated that 13 TwJAZs were localized in the nucleus, while 5 TwJAZs were located in the cytoplasm and chloroplasts. The phylogenetic tree analysis revealed that the JAZ family proteins of T. wilfordii could be divided into five subfamilies, with members of the same subfamily sharing similar conserved motifs. Expression pattern analysis demonstrated that TwJAZs were predominantly expressed in flowers, leaves, and peeled stems, with most TwJAZs showing the highest expression levels after 4 hours of methyl jasmonate (MeJA) induction. The transcription factor regulatory network shows that TwJAZs are strongly correlated with transcription factor families such as AP2/ERF-ERF, NAC, and bHLH. This study comprehensively identified all the JAZ gene family sequences in T. wilfordii, initially clarifying the structural and functional characteristics of the TwJAZs, and laying an important foundation for further research on the functions and regulatory mechanisms of TwJAZs.

Tripterygium wilfordii  /  JAZ  /  bioinformatics analysis  /  expression pattern  /  gene cloning
王柯燕, 唐丽媚, 吴林捷, 王雨婷, 陈夏唯, 屠李婵. 药用植物雷公藤JAZ基因家族的全基因组鉴定及功能分析. 药学学报, 2025 , 60 (4) : 1156 -1165 . DOI: 10.16438/j.0513-4870.2024-1147
Ke-yan WANG, Li-mei TANG, Lin-jie WU, Yu-ting WANG, Xia-wei CHEN, Li-chan TU. Genome-wide identification and functional analysis of the JAZ gene family in Tripterygium wilfordii[J]. Acta Pharmaceutica Sinica, 2025 , 60 (4) : 1156 -1165 . DOI: 10.16438/j.0513-4870.2024-1147
正文
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中药雷公藤作为我国传统中药, 具有祛风除湿、清热解毒、杀虫止痒、舒筋活血通络的功效[1, 2]。其代表性成分雷公藤甲素和雷公藤红素具有抗肿瘤、抗炎、抗风湿等药理活性, 是近年来最有可能开发成为现代药物的天然产物[3]。茉莉酸(jasmonic acid, JA) 及其衍生物是一种在植物生长发育、病害防御以及机械损伤阶段里具有重要调控作用的植物激素[4, 5]。在前期研究中, 已证明通过茉莉酸甲酯(methyl jasmonate, MeJA) 诱导雷公藤悬浮细胞能显著促进雷公藤甲素等萜类活性成分在植物体内的合成积累[6], 但目前JA信号途径如何调控雷公藤甲素等萜类活性成分的生物合成仍有待探索。
JAZ (jasmoate ZIM-domain) 蛋白作为JA途径的抑制因子, 在响应JA信号途径中发挥着重要作用[7]。在低水平JA条件下, JAZ与JA信号途径的转录因子结合来抑制JA应答基因的表达; 而在高水平JA下, JAZ会迅速降解, 从而释放并触发JA信号通路的激活。Zhang等[8]进一步证实JAZ蛋白能够根据JA的存在与否来调整自身构象, 扮演阻抑蛋白和受体蛋白两种角色, 从而控制植物激素途径的抑制和激活。此外, JAZ还能与其他通路调控因子相互作用, 对信号传导以及植物的生长和代谢具有广泛的影响[9, 10]。因此, 深入研究JAZ蛋白对药用植物次级代谢产物合成及积累的调控机制, 在药用植物品种的优化与培育, 以及提升其有效成分含量的实际应用中, 蕴含着巨大的潜在价值。
JAZ蛋白最先从模式植物拟南芥(Arabidopsis thaliana) 中分离并鉴定获得, 具有ZIM、Jas、NT三个保守结构域。目前, JAZ基因家族在拟南芥、玉米、二穗短柄草和水稻等植物上有较多报道, 并在其表达方式、基因特征、蛋白质特征及功能已有相关研究, 而在药用植物上的研究较少。最新研究发现, 月季(Rosa hybrida) 中的RhCBL4-RhCIPK3模块可以调控JAZ5的降解, 来介导乙烯信号通路调节的花瓣衰老过程[11]。在萝卜(Brassica rapa L. subsp. rapa) 中, JAZ基因不仅参与对ABA、SA、MeJA、盐胁迫和低温胁迫的响应, 还可能在萝卜块根的形成中发挥作用[12]。在拟南芥中, JAZ蛋白能与bHLH转录因子(TT8、GL3、EGL3) 及R2R3 MYB转录因子(MYB75、GL1) 相互作用, 来调控毛状体的启动和花青素的积累[13]。而Zheng等[14]对药用植物丹参中的JAZ基因功能也进行了相关研究。该研究发现在JA信号通路中, SmJAZ3蛋白通过与SmERF73转录因子的互作, 从而调控丹参酮类活性成分的生物合成, 这也为JAZ基因家族调控雷公藤萜类活性成分的生物合成提供了重要参考。
至今为止, 关于雷公藤JAZ基因家族成员是否在其生长发育和次级代谢中参与调控萜类活性成分的生物合成, 尚未有相关研究报道。本研究基于前期组装的高质量雷公藤染色体级别的基因组图谱, 首次利用基因组数据开展雷公藤JAZ基因家族的全基因组鉴定, 对雷公藤JAZ基因家族成员的生物信息学和基因表达模式等进行研究, 明确雷公藤JAZ基因家族成员的特征, 为深入研究雷公藤JAZ家族蛋白的功能机制奠定重要基础。
材料与方法
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植物材料    本团队的雷公藤植株由福建省三明市永安市的大田桃源国有林场提供, 该植株由福建省三明市永安市的大田桃源国有林场涂育合老师鉴定为卫矛科雷公藤属植物雷公藤(Tripterygium wilfordii Hook. f.)。
试剂、菌株和载体    GeneJET凝胶回收试剂盒(批号: 91279139)、PhusionTM Plus PCR Master Mix (批号: 01154558)、胰蛋白胨(批号: 3110563)、琼脂(批号: 23136204)、酵母提取物(批号: 4304391-02) 购自美国Thermo Fisher Scientific公司; 氯化钠(批号: CS31213550) 购自北京酷莱搏科技有限公司; pEASY®-Blunt Zero Cloning Kit (批号: #Q40826)、Trans2K® Plus DNA Marker (批号: #Q20307)、6×DNA Loading Buffer (批号: #Q20722)、大肠杆菌Trans1-T1 (批号: #Q4551130) 购自北京全式金生物技术有限公司; 快速质粒小提试剂盒(DP105, 批号: #X0616) 购自天根生化科技(北京) 有限公司; 2×Flash PCR MasterMix (Dye, 批号: 10380/01722) 购自康为世纪生物科技股份有限公司。
雷公藤基因家族分析    课题组前期组装的雷公藤基因组图谱已上传至NCBI数据库(GenBank: JAAARO000000000.1)[6], 本研究所使用的雷公藤全基因组蛋白序列来源于NCBI数据库基于已上传的雷公藤基因组组装数据进行注释的版本(NCBI RefSeq assembly: GCF_013401445.1)。从Pfam数据库(http://pfam.xfam.org/) 获得JAZ家族蛋白保守结构域TIFY和Jas的隐马尔可夫文件(ID: PF06200、PF09425)[15]。使用TBtools[16]的Simple HMM Search工具对雷公藤基因组所有蛋白序列进行保守序列筛选(E-value < 0.05)。然后从TAIR (https://www.arabidopsis.org/) 数据库中下载13个拟南芥JAZ蛋白序列, 基因编号分别是AT1G19180、AT1G74950、AT3G17860、AT1G48500、AT1G17380、AT1G72450、AT2G34600、AT1G30135、AT1G70700、AT5G13220、AT3G43440、AT5G20900、AT3G22275。利用TBtools[16]的BlastP工具将拟南芥JAZ蛋白序列与雷公藤蛋白序列进行比对, 将E-value设为1e-5。2种方法结果取交集, 再利用NCBI中的Batch CD-search进行保守结构域验证[15, 17], 最终筛选获得TwJAZs蛋白32条, 属于18个TwJAZs基因, 命名为TwJAZ1~TwJAZ18, 并选择TwJAZ基因的最长转录本序列用于基因家族分析与基因全长克隆实验。
雷公藤RNA提取、cDNA合成与JAZ基因克隆    选取新鲜的雷公藤嫩叶, 放入2.0 mL的离心管中, 加入无菌磁珠于液氮速冻冷却后, 使用混合型碾磨仪MM 400在低温环境中研磨2 min。然后按照CTAB-LiCl法提取雷公藤嫩叶的总RNA, 利用1%琼脂糖凝胶电泳检测RNA的完整性和质量, 并用NanoDrop OneC型核酸定量仪测定RNA浓度。最后按照SMARTTM RACE cDNA Amplification Kit试剂盒的说明书进行反转录合成cDNA, 保存于-20 ℃冰箱中备用。使用Primer Premier 5软件设计雷公藤TwJAZs基因的全长序列引物, 引物由北京擎科生物科技有限公司(杭州) 合成(表 1)。利用全长序列引物和PCR反应, 克隆获得TwJAZs基因全长序列。PCR反应体系: 25.0 μL的PhusionTM Plus PCR Master Mix、正反引物各2.5 μL、1.0 μL cDNA和19.0 μL ddH2O。利用Bio-Rad的T100TM PCR仪进行扩增, 反应程序为: 98 ℃预变性30 s; 98 ℃变性10 s、55 ℃退火20 s、72 ℃延伸45 s、循环35次; 72 ℃终延伸8 min, 4 ℃保存。PCR产物经1%琼脂糖凝胶电泳检测, 切胶回收目的条带后将回收产物连接B-zero载体、转化大肠杆菌Trans-T1感受态细胞中, 通过菌液PCR筛选后送测序验证[18]。
雷公藤JAZ蛋白理化性质分析    利用ExPASy (http://web.expasy.org/prot‐param/) 对雷公藤JAZ家族成员的理化性质进行预测与分析, 包括氨基酸数目、分子量、等电点、不稳定指数等指标。用WOLF PSORT (https://wolfpsort.hgc.jp/) 预测雷公藤JAZ蛋白的亚细胞定位[19]。
雷公藤JAZ基因的染色体定位    利用雷公藤基因组注释GFF文件和TBtools[16]软件, 对雷公藤JAZ基因家族成员的染色体分布进行可视化分析[20]。
雷公藤JAZ家族的系统进化树构建    利用Mega X软件[21]构建雷公藤JAZ家族系统进化树。采用的方法为最大似然法(maximum likelihood, ML), 采用的最优模型为JTT+G+F model, 空位选项为use all sites, 检验参数bootstrap method值取500[22]。
雷公藤JAZ基因结构和保守基序分析    通过基因结构显示系统GSDS在线软件(http://gsds.gao-lab.org) 将雷公藤JAZ家族成员的基因结构可视化[23]。利用在线工具MEME (http://meme.nbcr.net/meme/) 对18个TwJAZs蛋白序列进行保守基序分析, 最终获得18个TwJAZs蛋白的保守基序[24]。
雷公藤转录组分析    基于课题组前期自测的66个雷公藤转录组测序数据[6] (SRA数据库编号: SRX7094753、SRX7094754、SRX7094755、SRX7094756、SRX7094757、SRX7094758、SRX7094759、SRX7094760、SRX7094761、SRX7094763、SRX7094764、SRX7094765、SRX7094766、SRX7094767、SRX7094768、SRX7094769、SRX7094770、SRX7094774、SRX7094775、SRX7094777、SRX7094778、SRX7094779、SRX7094780、SRX7094781、SRX7094782、SRX7094783、SRX7094784、SRX7094785、SRX7094786、SRX7094789、SRX7094790、SRX7094791、SRX7094792、SRX7094793、SRX7094794、SRX7094795、SRX7094796、SRX7094797、SRX7094799、SRX7094800、SRX7094801、SRX7094802、SRX7094803、SRX7094804、SRX7094805、SRX7094739、SRX7094740、SRX7094751、SRX7094762、SRX7094776、SRX7094787、SRX7094798、SRX7094771、SRX7094772、SRX7094749、SRX7094750、SRX7094752、SRX7094773、SRX7094741、SRX7094742、SRX7094743、SRX7094744、SRX7094745、SRX7094746、SRX7094747、SRX7094748), 分析TwJAZs基因在根部(周皮、韧皮部、木质部)、茎部(茎皮、去皮茎)、叶部、花部及MeJA诱导下不同时间的基因表达量(fragments per kilobase million, FPKM), 并利用Heatmap工具绘制雷公藤JAZ基因表达聚类热图, 用于进一步筛选差异表达的TwJAZs基因。
TwJAZs网络调控图谱构建    利用MATLAB R2019b软件进行雷公藤JAZ家族与转录因子家族的皮尔森相关系数(Pearson correlation coefficient) 的计算, 筛选与雷公藤JAZ家族强相关的转录因子家族(r > 0.8, P < 0.05), 并通过Cytoscape软件(version 3.9.1)[25]构建JAZ家族-转录因子家族的调控网络图谱。
结果与分析
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1 雷公藤JAZ基因家族成员的鉴定与克隆
为了获得雷公藤中的TwJAZs基因家族成员, 本研究利用BLAST比对和结构域筛选的方法, 在雷公藤基因组中共鉴定获得18条TwJAZs基因, 并设计18对特异性基因全长引物(表 1)。以雷公藤嫩叶的cDNA作为模板, PCR扩增获得18条基因条带, 经测序验证正确, 命名为TwJAZ1~TwJAZ18。
2 雷公藤JAZ蛋白理化性质分析
ExPASy分析结果显示(表 2), 雷公藤JAZ家族成员蛋白质的氨基酸数目为122~525个, 分子质量为13.87~54.73 kDa, 平均亲水性在-0.877~-0.183, 均为亲水性蛋白。大部分JAZ蛋白等电点在9左右, 为碱性蛋白, 而TwJAZ18等电点为6.83, 为酸性蛋白, 结果说明雷公藤JAZ家族成员大多是碱性蛋白。18个TwJAZs蛋白中, 仅有TwJAZ11为稳定蛋白(instability index < 40)。通过WOLF PSORT对雷公藤JAZ家族蛋白进行亚细胞定位预测, 结果显示绝大部分成员定位在细胞核上(表 2), TwJAZ2、TwJAZ8和TwJAZ9定位在叶绿体上, TwJAZ5和TwJAZ11定位在细胞质上。
3 雷公藤JAZ基因的染色体分布
在雷公藤基因组23条染色体上, TwJAZs基因仅不均匀地分布在12条染色体上, 分别为Chr1、Chr3、Chr4、Chr6、Chr9、Chr10、Chr13、Chr14、Chr16、Chr17、Chr20和Chr23 (图 1)。其中, 在Chr13染色体上分布最多, 共5个TwJAZs基因定位在该条染色体上。此外, 各有2个TwJAZs基因定位在Chr9和Chr14染色体上, 其余9条染色体都只有1个TwJAZ基因。
4 雷公藤JAZ家族的系统进化树
利用18个TwJAZs蛋白与拟南芥[26]、葡萄[27]、番茄[28]和丹参[29]中的JAZ蛋白构建系统发育进化树(图 2)。结果发现, 拟南芥、葡萄、番茄、丹参和雷公藤的JAZ蛋白根据聚类关系可分为五个亚族(group Ⅰ~Ⅴ), 雷公藤JAZ蛋白成员在第Ⅳ亚族分布最多, 在第Ⅲ亚族分布最少。其中, TwJAZ2、TwJAZ4、TwJAZ6和TwJAZ9属于第Ⅰ亚族; TwJAZ8、TwJAZ14和TwJAZ15属于第Ⅱ亚族; TwJAZ7和TwJAZ10属于第Ⅲ亚族; TwJAZ12、TwJAZ13、TwJAZ16、TwJAZ17和TwJAZ18属于第Ⅳ亚族; TwJAZ1、TwJAZ3、TwJAZ5和TwJAZ11属于第Ⅴ亚族。此外, 从图 2进化树分析结果可以看出, 雷公藤的JAZ家族成员与葡萄的JAZ家族成员进化关系相对较近。
5 雷公藤JAZ基因结构分析
雷公藤JAZ基因结构分析结果显示, 不同成员之间基因结构的差异较大, 各成员中的CDS数量为2~8个(图 3)。它们的外显子-内含子结构表明, TwJAZ12、TwJAZ13、TwJAZ16、TwJAZ17和TwJAZ18的基因结构最简单, 只包含2个CDS、2个UTR和1个Intron; TwJAZ7~10均具有5个CDS、2个UTR和4个intron; 而TwJAZ14的CDS数量最多, 具有8个CDS。
6 雷公藤JAZ蛋白的保守基序分析
对雷公藤JAZ家族成员的保守基序进行分析可进一步验证JAZ家族进化关系的正确性。保守基序分析结果表明各成员均含有motif 1、motif 3, 是该家族最保守的两个蛋白基序(图 4)。不同亚族成员之间的motif个数和排列方式都相近, 与系统进化树的结果大致吻合。其中, motif 1、motif 2和motif 3在各个亚家族中均存在, motif 8仅存在于第Ⅰ亚族, motif 7和motif 10仅存在于第Ⅴ亚族, 而motif 9存在于第Ⅰ、Ⅱ、Ⅳ、Ⅴ亚族, 但不存在于Ⅲ亚族。
7 雷公藤JAZ基因家族表达模式分析
为了揭示JAZ基因家族在MeJA处理下和不同组织的表达情况, 本研究分析了66个雷公藤转录组数据。首先, 通过45个雷公藤悬浮细胞的转录组测序数据的分析, 获得TwJAZs基因在对照组和MeJA诱导下的表达量并绘制热图。如图 5A所示, TwJAZs在对照组和MeJA诱导下的表达模式不同, 大部分TwJAZs基因在MeJA诱导后表达呈现上调趋势, 且在MeJA诱导4 h后的表达量最高。
为了进一步研究雷公藤JAZ基因家族在雷公藤不同组织中的表达模式, 本研究分析了雷公藤21个组织转录组测序数据, 获得TwJAZs基因在7个不同组织部位的表达量并绘制热图。组织转录组分析结果表明(图 5B), 大部分TwJAZs在雷公藤的花部(F)、叶部(L) 和去皮茎部(PS) 中的表达量较高。而TwJAZ1、TwJAZ5和TwJAZ8在根的周皮(RB) 中的表达量最高, 其中TwJAZ5基因在根的韧皮部(RP) 中表达量也较高。
8 雷公藤JAZ家族与不同转录因子家族的调控网络
雷公藤JAZ家族与不同转录因子家族的调控网络图谱中(图 6), TwJAZ8、TwJAZ15和TwJAZ18同任何转录因子家族均不呈现强相关。相比之下, 其余雷公藤JAZ家族成员与AP2/ERF-ERF、NAC、bHLH、Tify家族成员呈现出强相关的数量最多, 其次是与GARP-G2-like、MYB、WRKY家族成员的强相关, 而与Alfin-like、AP2/ERF-RAV、B3-ARF、BES1、LOB、MADS-M-type、NF-YA等家族成员的强相关则相对较少, 仅有少数TwJAZs与之存在显著的强相关。已知bHLH、ERF、MYB、C2H2、WRKY和bZIP等家族已陆续被证明在调控植物萜类生物合成中起着关键作用, TwJAZ4、TwJAZ7、TwJAZ10、TwJAZ13、TwJAZ17与上述转录因子家族成员呈现较多强相关, 推测TwJAZ4、TwJAZ7、TwJAZ10、TwJAZ13、TwJAZ17蛋白可能是调控雷公藤甲素等重要萜类活性成分生物合成的关键因子。
讨论
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雷公藤系卫矛科雷公藤属的藤本灌木植物, 临床中多使用其根或根木质部入药, 相关制剂作为疗效确切的免疫抑制剂被广泛用于治疗类风湿性关节炎[30, 31]、狼疮性肾炎[32]、皮肤病[33]等。雷公藤的主要活性成分为雷公藤甲素和雷公藤红素。值得一提的是, 对雷公藤甲素修饰后的化合物能有效抑制胰腺肿瘤的生长和扩散, 具有治疗胰腺癌的潜力[34]。然而, 雷公藤资源现状令人堪忧, 其野生资源面临枯竭、栽培周期长且有效成分含量低, 加之遗传改良研究空白, 限制了雷公藤资源的培育与开发。由此可见, 对雷公藤进行基因组层面的研究显得尤为重要。
JAZ家族蛋白作为JA信号途径的抑制因子, 在植物的生长、发育和应激反应中起关键作用[35]。因此, 深入研究雷公藤JAZ蛋白调控雷公藤生长发育和代谢的功能、对提高其药效成分及优化培育品种具有潜在价值。当前, 许多药用植物都已进行了基因组水平的JAZ基因家族鉴定, 如在木薯基因组中鉴定了16个MeJAZs基因[36]; 在丹参基因组中鉴定了9个SmJAZs基因[37]; 在香椿基因组中鉴定了13个TciJAZs基因[38]。但遗憾的是, 在极具经济和药用价值的传统药用植物雷公藤中, 该家族的研究尚属空白。本研究利用生物信息学的方法, 从雷公藤基因组中鉴定了18个TwJAZs基因, 并将它们分为5个亚族, 它们不均匀地分布在12条染色体上, 推测TwJAZs基因在各自位置发挥不同的作用和功能。此外, 绝大部分TwJAZs蛋白定位在细胞核上, 这与目前绝大多数植物的JAZ蛋白定位相同。
最新研究指出, JAZ蛋白在多种植物中与抗寒性、病害抗性等生理过程紧密相关。例如, 在苹果(apple) 中, MIEL1和JAZ蛋白通过BBX37-ICE1-CBF模块共同调控JA介导的抗寒性[39]; 在黄花苜蓿(Medicago falcata L.) 中, 其15个MfJAZs在调节黄花苜蓿低温耐受性的JA信号中起着重要作用[40]; 在番茄(Solanum lycopersicum) 中, SlJAZ25可能与JA和SA激素调节密切相关, 在番茄抗灰斑病中起到负调控作用[41]; 在甘蔗(Saccharum spp.) 中, ScJAZ22、ScJAZ24、ScJAZ28、ScJAZ30和ScJAZ31在甘蔗应答叶枯病菌侵染过程中可能发挥积极的调控作用[42]; 在葡萄中, VvJAZ9基因受低温诱导表达最为明显[43]。而本研究中的系统进化树分析结果显示, 雷公藤与葡萄进化关系相近, 推测与VvJAZ9相近的TwJAZ4、TwJAZ6、TwJAZ9蛋白可能具有相同的抗寒功能。
雷公藤萜类活性成分是雷公藤中最具成药潜力的活性成分。在转录组分析结果中, 本研究发现18个TwJAZs在雷公藤不同组织中表达差异较大, 猜测TwJAZs基因可能在相关器官的发育中发挥不同功能。相较于其他基因, TwJAZ1、TwJAZ5和TwJAZ8在根的周皮中高表达, 而雷公藤甲素和雷公藤红素在根的周皮中含量最高, 推测这3个基因可能参与调控雷公藤关键萜类活性成分的生物合成与积累。此外, 在JAZ与转录因子的调控网络中, TwJAZ4、TwJAZ7、TwJAZ10、TwJAZ13、TwJAZ17与bHLH、ERF、MYB、C2H2、WRKY和bZIP等已被证明参与调控植物萜类生物合成的家族具有显著的相关性。因此, 也推测TwJAZ4、TwJAZ7、TwJAZ10、TwJAZ13、TwJAZ17可能是调控雷公藤重要萜类活性成分生物合成的关键蛋白。
综上所述, 本研究系统鉴定了18个TwJAZs基因, 并初步探索了雷公藤JAZ家族成员的功能, 为雷公藤的生长发育与重要次级代谢提供了多个关键的候选因子, 为进一步深入解析其家族功能和明确各个JAZ蛋白参与调控的具体路径和机制研究奠定了重要基础。
基金
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  • 国家自然科学基金资助项目(82204559)
  • 浙江省自然科学基金资助项目(LQ23H280002)
文献
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2025年第60卷第4期
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doi: 10.16438/j.0513-4870.2024-1147
  • 接收时间：2024-11-19
  • 首发时间：2025-10-29
  • 出版时间：2025-04-12
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  • 收稿日期：2024-11-19
  • 修回日期：2025-01-15
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国家自然科学基金资助项目(82204559)
浙江省自然科学基金资助项目(LQ23H280002)
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    浙大城市学院医学院, 浙江 杭州 310015

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屠李婵, E-mail:
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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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