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Article(id=1277330185972224552, tenantId=1146029695717560320, journalId=1235980609244409860, issueId=1277330185204666919, articleNumber=null, orderNo=null, doi=10.3969/j.issn.1000-2561.2024.03.001, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=null, receivedDate=1671984000000, receivedDateStr=2022-12-26, revisedDate=1673625600000, revisedDateStr=2023-01-14, acceptedDate=null, acceptedDateStr=null, onlineDate=1782468847983, onlineDateStr=2026-06-26, pubDate=1711296000000, pubDateStr=2024-03-25, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1782468847983, onlineIssueDateStr=2026-06-26, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1782468847983, creator=13701087609, updateTime=1782468847983, updator=13701087609, issue=Issue{id=1277330185204666919, tenantId=1146029695717560320, journalId=1235980609244409860, year='2024', volume='45', issue='3', pageStart='443', pageEnd='652', issueExtLink='null', onlineDate='null', pubDate='1711296000000', pubDateStr='2024-03-25', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=0, createTime=1782468847800, creator='13701087609', updateTime=1782468948575, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext={EN=IssueExt(id=1277330607961150151, tenantId=1146029695717560320, journalId=1235980609244409860, issueId=1277330185204666919, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1277330607961150152, tenantId=1146029695717560320, journalId=1235980609244409860, issueId=1277330185204666919, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null, downloadFileDto=null}, startPage=443, endPage=449, ext={EN=ArticleExt(id=1277330186261631530, articleId=1277330185972224552, tenantId=1146029695717560320, journalId=1235980609244409860, language=EN, title=Subcellular Localization and Multimerization Analyses of HbPIP2;3, an Efficient Water Transporter from Hevea brasiliensis, columnId=1236256430337085821, journalTitle=Chinese Journal of Tropical Crops, columnName=Omics & Biotechnology, runingTitle=null, highlight=null, articleAbstract=

Aquaporins (AQPs), assembled in tetramers at the biological membrane, constitute a class of integral membrane proteins facilitating the passive transport of water. Natural rubber, which is specifically synthesized in the laticifer located in the secondary phloem of the rubber tree (Hevea brasiliensis) trunk, is expelled out in the form of latex that represents the cytoplasmic content of laticiferous cells upon tapping. Water, accounting for approximately 70% of the latex upon each tapping, plays a crucial role in the latex production by regulating the latex viscosity and laticifer turgor. Previous studies suggested that the water balance of laticifers is governed by plasma membrane intrinsic proteins (PIPs) especially HbPIP2;3, an efficient and abundant water transporter. To uncover the regulation mechanism of HbPIP2;3 in the laticifer water balance, its 861 bp full-length coding sequence (CDS) was isolated using RT-PCR. Sequence analysis revealed that HbPIP2;3 putatively encoded 286 amino acids with the theoretical molecular weight (MW) of 30.58 kDa, the isoelectric point (pI) of 8.50, the instability index (II) of 31.68, and the grand average of hydropathicity (GRAVY) of 0.449, implying its basic, hydrophobic, and stable features. The protein was shown to contain one conserved MIP (major intrinsic protein) domain, which including six typical transmembrane helices (i.e. TM1-6) as well as two half helices (i.e. HB and HE). 3D prediction via homology modeling suggested that HbPIP2;3 could assemble in homotetramer. In accordance with the bioinformatics prediction, transient over-expression of HbPIP2;3 in Nicotiana benthamiana leaves supported the cell membrane localization, which was also confirmed by bimolecular fluorescence complementation (BiFC). Moreover, BiFC revealed that HbPIP2;3 could interact itself, which was further confirmed by yeast two-hybrid. The results showed that HbPIP2;3 may be involved in the laticifer water balance in the form of homotetramer at the plasma membrane, however, whether it could also function in hereotetramer still needs to be addressed.

, authors=null, authorsList=Zhi ZOU, Xueying QIAO, Yujiao ZHENG, Jianghua YANG, 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=1277330187486368310, articleId=1277330185972224552, tenantId=1146029695717560320, journalId=1235980609244409860, language=CN, title=橡胶树HbPIP2；3的亚细胞定位与多聚化分析, columnId=1236256430517440904, journalTitle=热带作物学报, columnName=组学与生物技术, runingTitle=null, highlight=null, articleAbstract=

水通道蛋白是一类高效转运水分子的膜内在蛋白，其在生物膜上以四聚体的形式起作用。天然橡胶在橡胶树的乳管细胞中特异合成，并在割胶过程以胶乳的形式被排出。胶乳系乳管细胞的胞质成分，其含水量高达70%，水分通过调节胶乳的粘稠度和乳管膨压进而影响橡胶树的产排胶能力，是决定胶乳产量的关键因素。前期研究显示，橡胶树乳管的水分平衡主要由质膜内在蛋白（plasma membrane intrinsic protein，PIP）特别是HbPIP2；3介导。为揭示HbPIP2；3调控乳管水分平衡的分子机制，本研究采用RT-PCR技术对其861 bp的编码区进行分离。序列分析显示：HbPIP2；3预测编码286个氨基酸，理论分子量为30.58 kDa，等电点为8.50，不稳定系数为31.68，总平均疏水指数为0.449，为稳定的疏水型碱性蛋白；该蛋白含有保守的MIP结构域，其中包括6个典型的跨膜螺旋和2个半螺旋；基于同源建模的3D结构预测显示其可以形成同源四聚体。生物信息学预测和在烟草叶片中的亚细胞定位分析显示，HbPIP2；3定位在细胞膜，这同时也得到了双分子荧光互补（bimolecular fluorescence complementation，BiFC）实验的证实。BiFC实验显示，HbPIP2；3可在细胞膜上形成同源四聚体，这进一步得到酵母双杂交结果的验证。本研究结果表明，HbPIP2；3可通过同源四聚体的方式调控乳管的水分平衡，但是否存在异源互作模式还有待进一步研究。

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邹智（1982—），男，硕士研究员，研究方向：基因组进化与逆境适应机制、油脂代谢与调控；E-mail：。

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邹智（1982—），男，硕士研究员，研究方向：基因组进化与逆境适应机制、油脂代谢与调控；E-mail：。

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邹智（1982—），男，硕士研究员，研究方向：基因组进化与逆境适应机制、油脂代谢与调控；E-mail：。

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articleId=1277330185972224552, language=CN, orderNo=2, keyword=水通道蛋白), Keyword(id=1277330191697449563, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277330185972224552, language=CN, orderNo=3, keyword=亚细胞定位), Keyword(id=1277330191789724252, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277330185972224552, language=CN, orderNo=4, keyword=双分子荧光互补), Keyword(id=1277330192150434397, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277330185972224552, language=CN, orderNo=5, keyword=酵母双杂交)], refs=[Reference(id=1277330194398581356, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277330185972224552, doi=null, pmid=null, pmcid=null, year=2008, volume=8, issue=1, pageStart=45, pageEnd=null, url=null, language=null, rfNumber=[1], rfOrder=0, authorNames=DANIELSON J A, JOHANSON U, journalName=BMC Plant Biology, refType=null, unstructuredReference=DANIELSON J A, JOHANSON U. 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PIP1 aquaporins: Intrinsic water channels or PIP2 aquaporin modulators?[J] FEBS Letters, 2015, 589(23): 3508-3515., articleTitle=PIP1 aquaporins: Intrinsic water channels or PIP2 aquaporin modulators?, refAbstract=null)], funds=[Fund(id=1277330194172088938, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277330185972224552, awardId=31971688; 31700580, language=CN, fundingSource=国家自然科学基金项目(31971688; 31700580), fundOrder=null, country=null), Fund(id=1277330194293723755, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277330185972224552, awardId=320RC705, language=CN, fundingSource=海南省自然科学基金项目(320RC705), fundOrder=null, country=null)], companyList=[AuthorCompany(id=1277330187675111992, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277330185972224552, xref=1., ext=[AuthorCompanyExt(id=1277330187683500601, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277330185972224552, companyId=1277330187675111992, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=1.National Key Laboratory for Tropical Crop Breeding / Hainan Key Laboratory for Biosafety Monitoring and Molecular Breeding in Off-Season Reproduction Regions / Institute of Tropical Biosciences and Biotechnology / Sanya Research Institute, Chinese Academy of Tropical Agricultural Sciences, Haikou, Hainan 571101, China), AuthorCompanyExt(id=1277330187691889210, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277330185972224552, companyId=1277330187675111992, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=1.热带作物生物育种全国重点实验室/海南省南繁生物安全与分子育种重点实验室/中国热带农业科学院热带生物技术研究所/三亚研究院，海南海口　571101)]), AuthorCompany(id=1277330187750609467, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277330185972224552, xref=2., ext=[AuthorCompanyExt(id=1277330187758998076, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277330185972224552, companyId=1277330187750609467, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=2.Rubber Research Institute, Chinese Academy of Tropical Agricultural Sciences, Haikou, Hainan 571101, China), AuthorCompanyExt(id=1277330187767386685, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277330185972224552, companyId=1277330187750609467, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=2.中国热带农业科学院橡胶研究所，海南海口　571101)])], figs=[ArticleFig(id=1277330192582447710, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277330185972224552, language=EN, label=Fig. 1, caption=Bioinformatics analysis of HbPIP2;3, figureFileSmall=xNkvKnX8DR3H4ihcP46l7A==, figureFileBig=LTKDPCddfyKE/Lm9xXwhmg==, tableContent=null), ArticleFig(id=1277330192678916703, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277330185972224552, language=CN, label=图1, caption=HbPIP2；3的生物信息学分析

A：HbPIP2；3的保守结构域预测；B：HbPIP2；3、HbPIP1；1和SoPIP2；1的序列比对；C：HbPIP2；3的3D结构预测。

, figureFileSmall=xNkvKnX8DR3H4ihcP46l7A==, figureFileBig=LTKDPCddfyKE/Lm9xXwhmg==, tableContent=null), ArticleFig(id=1277330193127707232, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277330185972224552, language=EN, label=Fig. 2, caption=Subcellular localization analysis of HbPIP2;3, figureFileSmall=gYLpwmmWXFHbZcFAgg8hTQ==, figureFileBig=yhYQV/JjUmNxlwHDA4afDw==, tableContent=null), ArticleFig(id=1277330193375171169, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277330185972224552, language=CN, label=图2, caption=HbPIP2；3的亚细胞定位, figureFileSmall=gYLpwmmWXFHbZcFAgg8hTQ==, figureFileBig=yhYQV/JjUmNxlwHDA4afDw==, tableContent=null), ArticleFig(id=1277330193454862946, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277330185972224552, language=EN, label=Fig. 3, caption=Multimerization analysis of HbPIP2;3 on the basis of BiFC, figureFileSmall=h9kpxIndi71xnveEHfQD6g==, figureFileBig=+IvYL7JSrCgP2Iixtx0sbg==, tableContent=null), ArticleFig(id=1277330193530360419, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277330185972224552, language=CN, label=图3, caption=基于BiFC的HbPIP2；3多聚化分析, figureFileSmall=h9kpxIndi71xnveEHfQD6g==, figureFileBig=+IvYL7JSrCgP2Iixtx0sbg==, tableContent=null), ArticleFig(id=1277330193597469284, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277330185972224552, language=EN, label=Fig. 4, caption=Self-activation analysis of HbPIP2;3, figureFileSmall=bqlric0Shl3u0L7yFXQlxg==, figureFileBig=wAdj4yw2yzp5hT2KozVtvA==, tableContent=null), ArticleFig(id=1277330193807184485, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277330185972224552, language=CN, label=图4, caption=HbPIP2；3的自激活检测, figureFileSmall=bqlric0Shl3u0L7yFXQlxg==, figureFileBig=wAdj4yw2yzp5hT2KozVtvA==, tableContent=null), ArticleFig(id=1277330193878487654, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277330185972224552, language=EN, label=Fig. 5, caption=Multimerization analysis of HbPIP2;3 on the basis of yeast two-hybrid, figureFileSmall=Z7CDHLROlx39mjRt/mJyqQ==, figureFileBig=0PwqR2tfgd7At7aI6bI2zQ==, tableContent=null), ArticleFig(id=1277330193933013607, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277330185972224552, language=CN, label=图5, caption=基于酵母双杂交的HbPIP2；3多聚化分析, figureFileSmall=Z7CDHLROlx39mjRt/mJyqQ==, figureFileBig=0PwqR2tfgd7At7aI6bI2zQ==, tableContent=null), ArticleFig(id=1277330194000122472, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277330185972224552, language=EN, label=Tab. 1, caption=

Primers used in this study

, figureFileSmall=null, figureFileBig=null, tableContent=
引物名称Primer name引物序列（5′-3′）Primer sequence (5′-3′)用途Usage
HbPIP2;3FATGGCTAAGGACGTGGAAGTT基因克隆
HbPIP2;3RTTAAGCATTGCTCCTGAAGG
pCam1304-HbPIP2;3FAGTGGTCTCTGTCCAGTCCTATGGCTAAGGACGTGGAAGTT亚细胞定位
pCam1304-HbPIP2;3FGGTCTCAGCAGACCACAAGTAGCATTGCTCCTGAAGG
pNC-BiFC-HbPIP2;3FAGTGGTCTCTGTCCAGTCCTATGGCTAAGGACGTGGAAGTT双分子荧光互补
pNC-BiFC-HbPIP2;3RGGTCTCAGCAGACCACAAGTAGCATTGCTCCTGAAGG
pGADT7-HbPIP2;3FTACGACGTACCAGATTACGCTATGGCTAAGGACGTGGAAGTT酵母双杂交
pGADT7-HbPIP2;3RCAGTATCTACGATTCATCTGCTTAAGCATTGCTCCTGAAGG
pGBKT7-HbPIP2;3FATCTCAGAGGAGGACCTGATGGCTAAGGACGTGGAAGTT
pGBKT7-HbPIP2;3RTGCGGCCGCTGCAGGTCGTTAAGCATTGCTCCTGAAGG
), ArticleFig(id=1277330194067231337, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277330185972224552, language=CN, label=表1, caption=

本研究所用引物

, figureFileSmall=null, figureFileBig=null, tableContent=
引物名称Primer name引物序列（5′-3′）Primer sequence (5′-3′)用途Usage
HbPIP2;3FATGGCTAAGGACGTGGAAGTT基因克隆
HbPIP2;3RTTAAGCATTGCTCCTGAAGG
pCam1304-HbPIP2;3FAGTGGTCTCTGTCCAGTCCTATGGCTAAGGACGTGGAAGTT亚细胞定位
pCam1304-HbPIP2;3FGGTCTCAGCAGACCACAAGTAGCATTGCTCCTGAAGG
pNC-BiFC-HbPIP2;3FAGTGGTCTCTGTCCAGTCCTATGGCTAAGGACGTGGAAGTT双分子荧光互补
pNC-BiFC-HbPIP2;3RGGTCTCAGCAGACCACAAGTAGCATTGCTCCTGAAGG
pGADT7-HbPIP2;3FTACGACGTACCAGATTACGCTATGGCTAAGGACGTGGAAGTT酵母双杂交
pGADT7-HbPIP2;3RCAGTATCTACGATTCATCTGCTTAAGCATTGCTCCTGAAGG
pGBKT7-HbPIP2;3FATCTCAGAGGAGGACCTGATGGCTAAGGACGTGGAAGTT
pGBKT7-HbPIP2;3RTGCGGCCGCTGCAGGTCGTTAAGCATTGCTCCTGAAGG
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橡胶树HbPIP2；3的亚细胞定位与多聚化分析
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邹智 1 , 乔雪莹 1 , 郑玉皎 1 , 阳江华 2
热带作物学报 | 组学与生物技术 2024,45(3): 443-449
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热带作物学报 |组学与生物技术 2024 , 45 (3) : 443 -449
橡胶树HbPIP2；3的亚细胞定位与多聚化分析
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邹智（1982—），男，硕士研究员，研究方向：基因组进化与逆境适应机制、油脂代谢与调控；E-mail：。

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邹智1 , 乔雪莹1, 郑玉皎1, 阳江华2
作者信息
  • 1.热带作物生物育种全国重点实验室/海南省南繁生物安全与分子育种重点实验室/中国热带农业科学院热带生物技术研究所/三亚研究院，海南海口　571101
  • 2.中国热带农业科学院橡胶研究所，海南海口　571101
Subcellular Localization and Multimerization Analyses of HbPIP2;3, an Efficient Water Transporter from Hevea brasiliensis
Zhi ZOU1 , Xueying QIAO1, Yujiao ZHENG1, Jianghua YANG2
Affiliations
  • 1.National Key Laboratory for Tropical Crop Breeding / Hainan Key Laboratory for Biosafety Monitoring and Molecular Breeding in Off-Season Reproduction Regions / Institute of Tropical Biosciences and Biotechnology / Sanya Research Institute, Chinese Academy of Tropical Agricultural Sciences, Haikou, Hainan 571101, China
  • 2.Rubber Research Institute, Chinese Academy of Tropical Agricultural Sciences, Haikou, Hainan 571101, China
出版时间: 2024-03-25 doi: 10.3969/j.issn.1000-2561.2024.03.001
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水通道蛋白是一类高效转运水分子的膜内在蛋白，其在生物膜上以四聚体的形式起作用。天然橡胶在橡胶树的乳管细胞中特异合成，并在割胶过程以胶乳的形式被排出。胶乳系乳管细胞的胞质成分，其含水量高达70%，水分通过调节胶乳的粘稠度和乳管膨压进而影响橡胶树的产排胶能力，是决定胶乳产量的关键因素。前期研究显示，橡胶树乳管的水分平衡主要由质膜内在蛋白（plasma membrane intrinsic protein，PIP）特别是HbPIP2；3介导。为揭示HbPIP2；3调控乳管水分平衡的分子机制，本研究采用RT-PCR技术对其861 bp的编码区进行分离。序列分析显示：HbPIP2；3预测编码286个氨基酸，理论分子量为30.58 kDa，等电点为8.50，不稳定系数为31.68，总平均疏水指数为0.449，为稳定的疏水型碱性蛋白；该蛋白含有保守的MIP结构域，其中包括6个典型的跨膜螺旋和2个半螺旋；基于同源建模的3D结构预测显示其可以形成同源四聚体。生物信息学预测和在烟草叶片中的亚细胞定位分析显示，HbPIP2；3定位在细胞膜，这同时也得到了双分子荧光互补（bimolecular fluorescence complementation，BiFC）实验的证实。BiFC实验显示，HbPIP2；3可在细胞膜上形成同源四聚体，这进一步得到酵母双杂交结果的验证。本研究结果表明，HbPIP2；3可通过同源四聚体的方式调控乳管的水分平衡，但是否存在异源互作模式还有待进一步研究。

乳管  /  水通道蛋白  /  亚细胞定位  /  双分子荧光互补  /  酵母双杂交

Aquaporins (AQPs), assembled in tetramers at the biological membrane, constitute a class of integral membrane proteins facilitating the passive transport of water. Natural rubber, which is specifically synthesized in the laticifer located in the secondary phloem of the rubber tree (Hevea brasiliensis) trunk, is expelled out in the form of latex that represents the cytoplasmic content of laticiferous cells upon tapping. Water, accounting for approximately 70% of the latex upon each tapping, plays a crucial role in the latex production by regulating the latex viscosity and laticifer turgor. Previous studies suggested that the water balance of laticifers is governed by plasma membrane intrinsic proteins (PIPs) especially HbPIP2;3, an efficient and abundant water transporter. To uncover the regulation mechanism of HbPIP2;3 in the laticifer water balance, its 861 bp full-length coding sequence (CDS) was isolated using RT-PCR. Sequence analysis revealed that HbPIP2;3 putatively encoded 286 amino acids with the theoretical molecular weight (MW) of 30.58 kDa, the isoelectric point (pI) of 8.50, the instability index (II) of 31.68, and the grand average of hydropathicity (GRAVY) of 0.449, implying its basic, hydrophobic, and stable features. The protein was shown to contain one conserved MIP (major intrinsic protein) domain, which including six typical transmembrane helices (i.e. TM1-6) as well as two half helices (i.e. HB and HE). 3D prediction via homology modeling suggested that HbPIP2;3 could assemble in homotetramer. In accordance with the bioinformatics prediction, transient over-expression of HbPIP2;3 in Nicotiana benthamiana leaves supported the cell membrane localization, which was also confirmed by bimolecular fluorescence complementation (BiFC). Moreover, BiFC revealed that HbPIP2;3 could interact itself, which was further confirmed by yeast two-hybrid. The results showed that HbPIP2;3 may be involved in the laticifer water balance in the form of homotetramer at the plasma membrane, however, whether it could also function in hereotetramer still needs to be addressed.

laticifer  /  aquaporin  /  subcellular localization  /  BiFC  /  yeast two-hybrid
邹智, 乔雪莹, 郑玉皎, 阳江华. 橡胶树HbPIP2；3的亚细胞定位与多聚化分析. 热带作物学报, 2024 , 45 (3) : 443 -449 . DOI: 10.3969/j.issn.1000-2561.2024.03.001
Zhi ZOU, Xueying QIAO, Yujiao ZHENG, Jianghua YANG. Subcellular Localization and Multimerization Analyses of HbPIP2;3, an Efficient Water Transporter from Hevea brasiliensis[J]. Chinese Journal of Tropical Crops, 2024 , 45 (3) : 443 -449 . DOI: 10.3969/j.issn.1000-2561.2024.03.001
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质膜内在蛋白（plasma membrane intrinsic protein，PIP）是一类原始而高度保守的水通道蛋白（aquaporin，AQP）[1]。根据进化关系和序列特征，PIP可分为PIP1和PIP2两个亚类，前者具有较长的N端，而后者具有较长的C端[2-7]。在蟾蜍卵母细胞中的体外分析显示，PIP2具有高效的水分转运活性，而PIP1无活性或活性极低，究其原因主要是因PIP1不能有效定位到细胞膜所致[8-14]。在玉米原生质中亚细胞定位分析显示，隶属于PIP2亚类的ZmPIP2;1和ZmPIP2;5定位在细胞膜，而属PIP1亚类的ZmPIP1;1、ZmPIP1;2和ZmPIP1;6均定位在内质网[9]。虽然PIP单体本身就具有水分转运活性，但其在生物膜上以四聚体的形式起作用[15]。PIP不仅可形成同源四聚体，同时也可以形成异源四聚体[15-18]。
橡胶树（Hevea brasiliensis Muell. Arg.）隶属于大戟科，是天然橡胶的主要商业来源[19-21]。生产上，橡胶通过用胶刀周期性地切割树皮收集胶乳而获得。胶乳系乳管细胞的胞质成分，水分含量60%~ 70%，水分通过调节胶乳的粘稠度和乳管膨压进而影响橡胶树的产排胶能力，是决定胶乳产量的关键因素[4,11,14,22]。通过前期研究，本团队鉴定到1个在乳管中高丰度表达的AQP基因，即HbPIP2;3，发现该基因的表达水平与品系的胶乳产量和水分含量呈正相关，且该基因在蟾蜍中过表达可显著提高卵母细胞的水分透性[4,14]。为揭示HbPIP2;3调控乳管水分平衡的分子机制，本研究在前期工作基础上对其编码蛋白的亚细胞定位和多聚化特征进行分析，以增进对乳管水分平衡的分子认知，并为利用生物技术手段提高胶乳产量提供理论依据。
1　材料与方法
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1.1　材料
1.1.1　植物材料
基因克隆所用橡胶树品系为热研7-33-97；亚细胞定位和双分子荧光互补（BiFC）所用材料为本氏烟草。
1.1.2　菌株及载体
大肠杆菌DH5α、根癌农杆菌GV3101、pNC-Cam1304-SubN、pNC-BiFC-ENN和pNC-BiFC-ECN由本实验室保存；Y2HGold购自天根生化科技（北京）有限公司；酵母双杂交载体购自宝日医生物技术（大连）有限公司。
1.1.3　主要试剂
各类酶、试剂盒及生化试剂详见文献[23-24]。
1.2　方法
1.2.1　RNA提取和反转录
胶乳总RNA的提取及cDNA第一链的合成参照文献[4]。
1.2.2　基因克隆与载体构建
根据本地热研7-33-97的基因组序列设计引物（表1），并以胶乳来源的cDNA作为模板进行PCR扩增。参照文献[24]，利用同源重组法将HbPIP2;3构建到pNC-Cam1304-SubN、pNC-BiFC-ENN、pNC-BiFC-ECN、pGADT7和pGBKT7。
1.2.3　生物信息学分析
蛋白的理化特性、保守结构域、亚细胞定位和二级结构预测分别采用Protparam（https://web.expasy.org/protparam/）、CDD（https://www.ncbi.nlm.nih.gov/cdd）、WoLF PSORT（https://www.genscript.com/wolf-psort.html）和SOP-MA（https://npsa-prabi.ibcp.fr/cgi-bin/npsa_automat.pl?page=/NPSA/npsa_sopma.html）等在线工具分析。根据与菠菜SoPIP2;1[15]进行序列比对界定相应的跨膜螺旋区，并利用其晶体结构作为模板通过SWISS-MODEL（https://swissmodel.expasy.org/int-eractive）在线软件进行3D建模。
1.2.4　亚细胞定位和双分子荧光互补
含重组质粒农杆菌浸染液的制备及烟草叶片的瞬时转化参照文献[24]，其中，烟草幼苗约为5周龄，转化48 h后用共聚焦显微镜进行荧光观察。
1.2.5　酵母双杂交
诱饵载体pGBKT7-HbPIP2;3的自激活检测及点对点的酵母双杂交参照宝日医酵母双杂交试剂盒说明书。
2　结果与分析
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2.1　基因克隆与载体构建
以HbPIP2;3F/R为引物的首轮PCR成功扩增到1条约860 bp的目的条带。随后，以第一轮PCR产物作为模板，分别以pCam1304-HbPIP2;3F/R、pNC-BiFC-HbPIP2;3F/R、pGADT7-HbPIP2;3F/R或pGBKT7-HbPIP2;3F/R为引物进行第二轮PCR扩增，目的条带切胶回收后分别构建pCam1304-HbPIP2;3、pNC-BiFC-ECN-HbPIP2;3、pNC-BiFC-ENN-HbPIP2;3、pGADT7-HbPIP2;3和pGBKT7-HbPIP2;3等载体。
2.2　序列的生物信息学分析
测序及序列分析表明，HbPIP2;3的CDS全长为861 bp，预测编码286个氨基酸，理论分子量为30.58 kDa，等电点为8.50，脂肪族指数为100.38，总平均疏水指数为0.449，不稳定系数为31.68。CDD分析显示，该蛋白含有1个高度保守的MIP结构域，即32~267位，共包含236个残基（图1A）。序列比对显示，HbPIP2;3与HbPIP1;1、SoPIP2;1的序列相似性分别为72.1%和85.1%，拥有6个典型的跨膜螺旋（即TM1~6）和2个半螺旋（即HB和HE），其中，位于2个半螺旋上的NPA基序，以及位于TM2、TM5和LE上的ar/R选择性滤器完全一致，均为F-H-T-R；在5个Froger位点（即P1~5）中，仅P1存在变异，分别为Q-S-A-F-W、E-S-A-F-W和M-S-A-F-W；此外，对应于SoPIP2;1 S115的磷酸化位点和L197门控位点也完全一致，而对应于SoPIP2;1 S274的磷酸化位点则只存在HbPIP2;3和SoPIP2;1中（图1B）。SOPMA分析显示，HbPIP2;3的α-螺旋占36.36%，延伸链结构占17.48%，β-转角占3.50%，无规则卷曲占42.66%；同源建模进一步证实该蛋白含有6个跨膜螺旋，同时也显示其可以形成同源四聚体（图1C）。亚细胞定位预测显示，HbPIP2;3定位在细胞膜。
2.3　亚细胞定位分析
以空载pNC-Cam1304-SubN为对照，将含pCam1304-HbPIP2;3的农杆菌工程菌微量注射烟草叶片后进行荧光观察，结果如图2所示，转空载的荧光信号散布于细胞的各个区域，包括细胞核，而实验组仅见于细胞膜，这表明HbPIP2;3定位在细胞膜。
2.4　BiFC分析
以转pCam1304-HbPIP2;3的为阳性对照、单转pNC-BiFC-ENN-HbPIP2;3或pNC-BiFC-ECN-HbPIP2;3的为阴性对照，将分别含有pNC-BiFC-ENN-HbPIP2;3和pNC-BiFC-ECN-HbPIP2;3的农杆菌工程菌等量混合后转化烟草，荧光观察结果如图3所示，2个阴性对照均为未见信号，而实验组和阳性对照在细胞膜均发现强烈的荧光信号，这表明HbPIP2;3可在细胞膜上形成多聚体。
2.5　酵母双杂交实验
自激活检测显示（图4），转pGADT7/pGBKT-7-HbPIP2;3的阳性对照在二缺（SD-TL）、三缺（SD-TLH）和四缺（SD-TLHA）营养平板上生长旺盛，而实验组与阴性对照类似，在四缺板上均无生长，这表明pGBKT7-HbPIP2;3无自激活活性。进一步的酵母双杂交实验显示（图5），实验组和阳性对照类似，在四缺板上生长旺盛，而阴性对照无生长，这表明HbPIP2;3在体外也可形成多聚体。
3　讨论
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作为天然橡胶合成和贮藏的场所，橡胶树乳管是一类高度分化的单一细胞类型组织[19]。与成熟叶片等组织不同，乳管无中央大液泡，取而代之的是成百上千的分散性小液泡，即所谓的黄色体[4,19]。在成熟叶片中，水分的平衡由PIP和液泡定位的液泡内在蛋白（tonoplast intrinsic protein，TIP）共同决定，而乳管的水分平衡则主要由PIP决定[7]，因此，分离和鉴定乳管表达的PIP具有重要理论意义和应用价值[4,11-14]。在3个乳管高丰度表达的PIP中，HbPIP1;4和HbPIP2;7在蟾蜍卵母细胞中异源表达均无水分转运活性[13]，而HbPIP2;3的活性与HbPIP2;1相当[11,14]。由于HbPIP2;1在乳管中的表达丰度很低，因此，HbPIP2;3被认为是调控乳管水分平衡的关键基因[4,14]。
本研究结果显示，HbPIP2;3含有一个AQP家族特有的MIP结构域（Pfam登录号为PF00230），其中包含6个典型的跨膜螺旋和2个半螺旋，等电点>7.0，总平均疏水指数>0，不稳定系数为<40，属于稳定的疏水型碱性蛋白。与隶属于PIP1亚类的HbPIP1;1相比，HbPIP2;3与SoPIP2;1的序列相似性更高，具有较短的N端和较长的C端，且C端含有1个PIP2亚类特有的磷酸化位点[15]。与生物信息学预测结果一致，在烟草叶片中的亚细胞定位分析显示HbPIP2;3定位在细胞膜。与基于同源建模的3D结构预测一致，BiFC和点对点的酵母双杂交实验均表明HbPIP2;3可形成同源四聚体。HbPIP1;1虽然在烟草中也定位在细胞膜[24]，但与HbPIP2;3和SoPIP2;1不同的是，其在蟾蜍中并无水分转运活性[12]。虽然本研究试图通过序列比较寻求答案，但HbPIP1;1具有HbPIP2;3和SoPIP2;1类似的ar/R选择性滤器（F-H-T-R）以及典型的NPA基序，理论上HbPIP1;1应具有高效的水分转运活性，因此，相关的决定因子还有待进一步研究。另外一种可能是，HbPIP1;1自身原本不能定位到细胞膜，但可以跟烟草中的PIP2蛋白互作进而有效定位。事实上，PIP1和PIP2之间的异源互作广泛存在于不同植物中，如玉米、拟南芥、烟草、葡萄、甜菜、草莓、棉花、大麦，甚至还包括卷柏[8-9,16-18,25-27]。因此，进一步筛选和鉴定与HbPIP1;1互作的PIP2蛋白有助于阐明复杂的调控机制。
综上，本研究证实HbPIP2;3在细胞膜上可通过同源四聚体的方式调控橡胶树乳管的水分平衡，但是否存在异源互作模式还有待进一步研究。
基金
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  • 国家自然科学基金项目(31971688; 31700580)
  • 海南省自然科学基金项目(320RC705)
文献
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2024年第45卷第3期
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doi: 10.3969/j.issn.1000-2561.2024.03.001
  • 接收时间：2022-12-26
  • 首发时间：2026-06-26
  • 出版时间：2024-03-25
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  • 收稿日期：2022-12-26
  • 修回日期：2023-01-14
基金
国家自然科学基金项目(31971688; 31700580)
海南省自然科学基金项目(320RC705)
作者信息
    1.热带作物生物育种全国重点实验室/海南省南繁生物安全与分子育种重点实验室/中国热带农业科学院热带生物技术研究所/三亚研究院，海南海口　571101
    2.中国热带农业科学院橡胶研究所，海南海口　571101
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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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