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Article(id=1276175421876081167, tenantId=1146029695717560320, journalId=1235980609244409860, issueId=1276175380184695804, articleNumber=null, orderNo=null, doi=10.3969/j.issn.1000-2561.2024.09.006, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=null, receivedDate=1693152000000, receivedDateStr=2023-08-28, revisedDate=1696003200000, revisedDateStr=2023-09-30, acceptedDate=null, acceptedDateStr=null, onlineDate=1782193530757, onlineDateStr=2026-06-23, pubDate=1727193600000, pubDateStr=2024-09-25, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1782193530757, onlineIssueDateStr=2026-06-23, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1782193530757, creator=13701087609, updateTime=1782193530757, updator=13701087609, issue=Issue{id=1276175380184695804, tenantId=1146029695717560320, journalId=1235980609244409860, year='2024', volume='45', issue='9', pageStart='1761', pageEnd='1997', issueExtLink='null', onlineDate='null', pubDate='1727193600000', pubDateStr='2024-09-25', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1782193520816, creator='13701087609', updateTime=1782193908264, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext={EN=IssueExt(id=1276177005326504448, tenantId=1146029695717560320, journalId=1235980609244409860, issueId=1276175380184695804, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1276177005326504449, tenantId=1146029695717560320, journalId=1235980609244409860, issueId=1276175380184695804, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null, downloadFileDto=null}, startPage=1823, endPage=1831, ext={EN=ArticleExt(id=1276175423058874897, articleId=1276175421876081167, tenantId=1146029695717560320, journalId=1235980609244409860, language=EN, title=Cloning, Subcellular Localization, and Expression Analysis of CeTIP4;1, an Aquaporin Gene Expressed in Cyperus esculentus Tubers, columnId=1236256430337085821, journalTitle=Chinese Journal of Tropical Crops, columnName=Omics & Biotechnology, runingTitle=null, highlight=null, articleAbstract=

Tonoplast intrinsic proteins (TIPs), which comprise a subfamily of aquaporin (AQP) mediating the water balance within plant cells, could be divided into five phylogenetic groups on the basis of sequence similarity, i.e., TIP1–TIP5. Based on the full-length transcriptome previously described in tigernut (Cyperus esculentus), a tuber-abundant TIP gene named CeTIP4;1 was cloned and characterized in this study. Sequence analyses revealed that CeTIP4;1 contained two conservative introns with a complete coding sequence of 753 base pairs (bp), putatively encoding 250 amino acids with the theoretical molecular weight of 25.57 kDa, the isoelectric point (pI) value of 6.02, and the grand average of hydropathicity (GRAVY) value of 0.884, implying its acidic and hydrophobic features. The deduced peptide possessed one conservative major intrinsic protein (MIP) domain and exhibited similar structural features with the structure resolved AtTIP2;1. Classification of CeTIP4;1 into the TIP4 group was supported by the phylogenetic analysis, which showed that CeTIP4;1 clusters with AtTIP4;1, OsTIP4;1, OsTIP4;2 and OsTIP4;3 with sequence similarities of 69.62%–82.94%. After gene cloning, a plant expression vector for subcellular localization, i.e., pNC-Cam1304-CeTIP4;1, was successfully constructed. When transiently overexpressed in tobacco (Nicotiana benthamiana) leaves, confocal laser scanning microscopy imaging showed that CeTIP4;1 was unexpectedly localized to the cell membrane. Further expression analysis indicated that CeTIP4;1 exhibited an apparent tissue-specific expression patter, predominately expressed in tuber, root, rhizome, and shoot apex. Moreover, among the four typical stages of developmental tuber examined in this study, i.e., initiation, medium swelling, late swelling, and maturation, CeTIP4;1 exhibited a bell-like expression pattern, peaking in medium swelling tubers and lowest in maturation tubers. These results lay a solid foundation for further uncovering the mechanism of water balance in tigernut tubers.

, authors=null, authorsList=Zhi ZOU, Yujiao ZHENG, 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=1276175432328286746, articleId=1276175421876081167, tenantId=1146029695717560320, journalId=1235980609244409860, language=CN, title=油莎豆块茎表达CeTIP4;1基因的克隆、亚细胞定位及表达分析, columnId=1236256430517440904, journalTitle=热带作物学报, columnName=组学与生物技术, runingTitle=null, highlight=null, articleAbstract=

液泡膜内在蛋白（tonoplast intrinsic protein，TIP）是调控植物细胞内水分平衡的主要水通道蛋白类型，其根据进化关系可分为TIP1～TIP5等5个亚类。本研究基于前期获得的油莎豆全长转录组对1个块茎高水平表达的TIP基因（命名为CeTIP4;1）进行克隆和鉴定。序列分析显示，该基因含有2个保守的内含子，编码区全长753 bp，预测编码250个氨基酸，理论分子量为25.57 kDa、等电点（pI）为6.02、总平均疏水指数（GRAVY）为0.884，属于典型的酸性疏水型蛋白；该蛋白含有1个保守的MIP（major intrinsic protein）结构域，且具有已结晶AtTIP2;1类似的结构特征。进化分析显示，CeTIP4;1与AtTIP4;1、OsTIP4;1、OsTIP4;2和OsTIP4;3聚在一起，序列相似性为69.62%~82.94%，支持其划归为TIP4亚类。本研究在基因克隆的基础上进一步构建了其亚细胞定位载体pNC-Cam1304-CeTIP4;1。在烟草叶片中的瞬时过表达和共聚焦显微镜观察显示，CeTIP4;1定位在细胞膜。进一步的表达分析显示，CeTIP4;1具有明显的组织特异性，主要在块茎、根、匍匐茎和芽尖中表达；在起始期、膨大中期、膨大晚期和成熟期等4个典型的块茎发育时期中，CeTIP4;1基因的表达呈现先升后降的趋势，峰值出现在膨大中期，最低的是成熟期，与含水量趋势基本一致。本研究结果为深入解析油莎豆块茎的水分平衡机制奠定坚实的基础。

, authors=

邹智（1982—），男，硕士，研究员，研究方向：基因组进化与油脂调控；E-mail：。

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邹智（1982—），男，硕士，研究员，研究方向：基因组进化与油脂调控；E-mail：。

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邹智（1982—），男，硕士，研究员，研究方向：基因组进化与油脂调控；E-mail：。

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A: Gene structure of CeTIP4;1; B: Identified conservative domain of CeTIP4;1; C: Sequence alignment of CeTIP4;1, CeTIP1;1, AtTIP2;1, and SoPIP2;1; D: Predicted 3D structure of CeTIP4;1; E: Phylogenetic analysis of CeTIP4;1 and TIPs in O. sativa and A. thaliana.

, figureFileSmall=7wqB7iLr63OZht+pd1iKTQ==, figureFileBig=uekTLREJh+nt4our98a2vA==, tableContent=null), ArticleFig(id=1276465438594826357, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276175421876081167, language=CN, label=图1, caption=CeTIP4;1的生物信息学分析

A：CeTIP4;1的基因结构；B：CeTIP4;1的保守结构域；C：CeTIP4;1与CeTIP1;1、AtTIP2;1和SoPIP2;1的序列比对，6个跨膜螺旋和2个半螺旋用灰色背景显示，保守残基用不同颜色显示；D: CeTIP4;1的3D结构预测；E：CeTIP4;1与AtTIP和OsTIP的进化分析。

, figureFileSmall=7wqB7iLr63OZht+pd1iKTQ==, figureFileBig=uekTLREJh+nt4our98a2vA==, tableContent=null), ArticleFig(id=1276465438934564982, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276175421876081167, language=EN, label=Fig. 2, caption=Subcellular localization analysis of CeTIP4;1, figureFileSmall=rjpYMZf6CnZvr+tlpfEvzg==, figureFileBig=Dtg/HK47aYHtPE/a5OmkxQ==, tableContent=null), ArticleFig(id=1276465439001673847, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276175421876081167, language=CN, label=图2, caption=CeTIP4;1的亚细胞定位, figureFileSmall=rjpYMZf6CnZvr+tlpfEvzg==, figureFileBig=Dtg/HK47aYHtPE/a5OmkxQ==, tableContent=null), ArticleFig(id=1276465439068782712, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276175421876081167, language=EN, label=Fig. 3, caption=Expression profiles of CeTIP4;1

A: Tissue-specific expression profiles of CeTIP4;1; B: Expression profiles of CeTIP4;1 in different stages of developmental tubers; Different lowercase letters indicate significant difference (P<0.05).

, figureFileSmall=OxpADMotrDLaR05XIbBs2w==, figureFileBig=G9hLheUeElOLFSaHCG9gGw==, tableContent=null), ArticleFig(id=1276465439148474489, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276175421876081167, language=CN, label=图3, caption=CeTIP4;1的表达模式

A: CeTIP4;1的组织特异性分析；B: CeTIP4;1在不同发育时期块茎中的表达模式；不同小写字母表示差异显著（P<0.05）。

, figureFileSmall=OxpADMotrDLaR05XIbBs2w==, figureFileBig=G9hLheUeElOLFSaHCG9gGw==, tableContent=null), ArticleFig(id=1276465439328829562, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276175421876081167, language=EN, label=Tab. 1, caption=

Comparison of physicochemical properties and secondary structure of CeTIP4;1, CeTIP1;1, CeTIP2;1, AtTIP2;1, and SoPIP2;1

, figureFileSmall=null, figureFileBig=null, tableContent=
蛋白名称Protein nameaaMW/kDapIGRAVYAIIIMIPα-螺旋Alpha helix/%延伸链Extended strand/%β-转角Beta turn/%无规则卷曲Random coil/%
CeTIP4;125025.576.020.884119.4033.703~23536.0022.804.4036.80
CeTIP1;125125.756.010.831107.7727.811~24933.8621.516.7737.85
CeTIP2;124824.735.090.948114.6021.7614~23136.2920.974.8437.90
AtTIP2;125025.035.300.974112.4826.731~24433.2020.805.6040.40
SoPIP2;128129.909.030.579102.8127.8131~26036.6518.152.1443.06
), ArticleFig(id=1276465439391744123, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276175421876081167, language=CN, label=表1, caption=

CeTIP4;1与CeTIP1;1、CeTIP2;1、AtTIP2;1和SoPIP2;1理化特性和二级结构比较

, figureFileSmall=null, figureFileBig=null, tableContent=
蛋白名称Protein nameaaMW/kDapIGRAVYAIIIMIPα-螺旋Alpha helix/%延伸链Extended strand/%β-转角Beta turn/%无规则卷曲Random coil/%
CeTIP4;125025.576.020.884119.4033.703~23536.0022.804.4036.80
CeTIP1;125125.756.010.831107.7727.811~24933.8621.516.7737.85
CeTIP2;124824.735.090.948114.6021.7614~23136.2920.974.8437.90
AtTIP2;125025.035.300.974112.4826.731~24433.2020.805.6040.40
SoPIP2;128129.909.030.579102.8127.8131~26036.6518.152.1443.06
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油莎豆块茎表达CeTIP4;1基因的克隆、亚细胞定位及表达分析
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邹智 , 郑玉皎
热带作物学报 | 组学与生物技术 2024,45(9): 1823-1831
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热带作物学报 |组学与生物技术 2024 , 45 (9) : 1823 -1831
油莎豆块茎表达CeTIP4;1基因的克隆、亚细胞定位及表达分析
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邹智（1982—），男，硕士，研究员，研究方向：基因组进化与油脂调控；E-mail：。

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邹智（1982—），男，硕士，研究员，研究方向：基因组进化与油脂调控；E-mail：。

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邹智 , 郑玉皎
作者信息
  • 热带作物生物育种全国重点实验室/中国热带农业科学院热带生物技术研究所/中国热带农业科学院三亚研究院，海南海口　571101
Cloning, Subcellular Localization, and Expression Analysis of CeTIP4;1, an Aquaporin Gene Expressed in Cyperus esculentus Tubers
Zhi ZOU , Yujiao ZHENG
Affiliations
  • National Key Laboratory for Tropical Crop Breeding / Institute of Tropical Biosciences and Biotechnology, Chinese Academy of Tropical Agricultural Sciences / Sanya Research Institute, Chinese Academy of Tropical Agricultural Sciences, Haikou, Hainan 571101, China
出版时间: 2024-09-25 doi: 10.3969/j.issn.1000-2561.2024.09.006
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液泡膜内在蛋白（tonoplast intrinsic protein，TIP）是调控植物细胞内水分平衡的主要水通道蛋白类型，其根据进化关系可分为TIP1～TIP5等5个亚类。本研究基于前期获得的油莎豆全长转录组对1个块茎高水平表达的TIP基因（命名为CeTIP4;1）进行克隆和鉴定。序列分析显示，该基因含有2个保守的内含子，编码区全长753 bp，预测编码250个氨基酸，理论分子量为25.57 kDa、等电点（pI）为6.02、总平均疏水指数（GRAVY）为0.884，属于典型的酸性疏水型蛋白；该蛋白含有1个保守的MIP（major intrinsic protein）结构域，且具有已结晶AtTIP2;1类似的结构特征。进化分析显示，CeTIP4;1与AtTIP4;1、OsTIP4;1、OsTIP4;2和OsTIP4;3聚在一起，序列相似性为69.62%~82.94%，支持其划归为TIP4亚类。本研究在基因克隆的基础上进一步构建了其亚细胞定位载体pNC-Cam1304-CeTIP4;1。在烟草叶片中的瞬时过表达和共聚焦显微镜观察显示，CeTIP4;1定位在细胞膜。进一步的表达分析显示，CeTIP4;1具有明显的组织特异性，主要在块茎、根、匍匐茎和芽尖中表达；在起始期、膨大中期、膨大晚期和成熟期等4个典型的块茎发育时期中，CeTIP4;1基因的表达呈现先升后降的趋势，峰值出现在膨大中期，最低的是成熟期，与含水量趋势基本一致。本研究结果为深入解析油莎豆块茎的水分平衡机制奠定坚实的基础。

油料作物  /  虎坚果  /  块茎  /  液泡膜内在蛋白  /  亚细胞定位  /  表达模式

Tonoplast intrinsic proteins (TIPs), which comprise a subfamily of aquaporin (AQP) mediating the water balance within plant cells, could be divided into five phylogenetic groups on the basis of sequence similarity, i.e., TIP1–TIP5. Based on the full-length transcriptome previously described in tigernut (Cyperus esculentus), a tuber-abundant TIP gene named CeTIP4;1 was cloned and characterized in this study. Sequence analyses revealed that CeTIP4;1 contained two conservative introns with a complete coding sequence of 753 base pairs (bp), putatively encoding 250 amino acids with the theoretical molecular weight of 25.57 kDa, the isoelectric point (pI) value of 6.02, and the grand average of hydropathicity (GRAVY) value of 0.884, implying its acidic and hydrophobic features. The deduced peptide possessed one conservative major intrinsic protein (MIP) domain and exhibited similar structural features with the structure resolved AtTIP2;1. Classification of CeTIP4;1 into the TIP4 group was supported by the phylogenetic analysis, which showed that CeTIP4;1 clusters with AtTIP4;1, OsTIP4;1, OsTIP4;2 and OsTIP4;3 with sequence similarities of 69.62%–82.94%. After gene cloning, a plant expression vector for subcellular localization, i.e., pNC-Cam1304-CeTIP4;1, was successfully constructed. When transiently overexpressed in tobacco (Nicotiana benthamiana) leaves, confocal laser scanning microscopy imaging showed that CeTIP4;1 was unexpectedly localized to the cell membrane. Further expression analysis indicated that CeTIP4;1 exhibited an apparent tissue-specific expression patter, predominately expressed in tuber, root, rhizome, and shoot apex. Moreover, among the four typical stages of developmental tuber examined in this study, i.e., initiation, medium swelling, late swelling, and maturation, CeTIP4;1 exhibited a bell-like expression pattern, peaking in medium swelling tubers and lowest in maturation tubers. These results lay a solid foundation for further uncovering the mechanism of water balance in tigernut tubers.

oil crop  /  tigernut  /  tuber  /  tonoplast intrinsic protein  /  subcellular localization  /  expression profile
邹智, 郑玉皎. 油莎豆块茎表达CeTIP4;1基因的克隆、亚细胞定位及表达分析. 热带作物学报, 2024 , 45 (9) : 1823 -1831 . DOI: 10.3969/j.issn.1000-2561.2024.09.006
Zhi ZOU, Yujiao ZHENG. Cloning, Subcellular Localization, and Expression Analysis of CeTIP4;1, an Aquaporin Gene Expressed in Cyperus esculentus Tubers[J]. Chinese Journal of Tropical Crops, 2024 , 45 (9) : 1823 -1831 . DOI: 10.3969/j.issn.1000-2561.2024.09.006
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液泡膜内在蛋白（tonoplast intrinsic protein，TIP）隶属于水通道蛋白（aquaporin，AQP）家族，因其定位在液泡膜而得名[1]。TIP最先在模式植物拟南芥中被鉴定，拟南芥γ-TIP1（AtTIP1;1）及其同源基因在蟾蜍卵母细胞和酵母中异源表达通常表现出高效的水分转运活性，高于或与细胞膜定位的质膜内在蛋白（plasma membrane intrinsic protein，PIP）相当[2-3]。在体外，有些TIP还被证实可以转运甘油、尿素、硼酸、NH3、H2O2等中性小分子[4-10]。拟南芥（Arabidopsis thaliana）基因组含有10个TIP基因，并根据进化关系分为TIP1~TIP5等5个亚类，其中，除TIP4和TIP5仅有1个成员外，其他亚类含有2~3个成员不等[11]。晶体结构显示，与PIP类似，TIP含有6个由5个亲水环（LA–E）连接的跨膜α螺旋（TM1–6）、2个由LB和LE折向膜内形成的半螺旋以及位于半螺旋顶端的NPA基序[10,12]，2个NPA基序与位于TM2、TM5和LE上的4个保守残基（即ar/R选择性滤器：H2、H5、LE1和LE2）共同决定了孔道的大小和底物的特异性[13-14]；此外，拟南芥AtTIP2;1位于LC的H131为NH3的高效转运所必须[10]。与PIP相比，拟南芥TIP的ar/R选择性滤器变异较大，TIP1为H-I-A-V，TIP2、TIP3和TIP4为H-I-G/A-R，TIP5为N-V-G-C，孔径介于0.67~1.56 Å[15]。研究表明，不同TIP亚类定位于不同类型的液泡，如TIP1主要定位于裂解型液泡，TIP2位于蛋白储藏型液泡，TIP3定位于种子的蛋白储藏型液泡，并在种子的萌发过程中逐步被TIP1所替代[16]。TIP4也在裂解型液泡被发现，主要参与水分、尿素和甘油的转运[5,9]。
油莎豆（Cyperus esculentus L.），又名虎坚果，是一种隶属于禾本目莎草科莎草属的新型草本油料作物[17-20]。不同于传统油料作物在种子中积累油脂，油莎豆不开花或开花不结实，但它可在地下块茎中积累高达35%的油脂，这使其成为研究营养组织油脂合成与调控的理想模型[21-23]。油莎豆块茎起源于地下匍匐茎，其发育过程主要分为起始、膨大和成熟等3个阶段[21,24]。通过前期研究，团队发现海南地区块茎的发育非常迅速，自起始后35 d即可成熟，成熟块茎的含水量约为45%，而在此之前一般维持在85%左右[24]，表明水分平衡在块茎发育和代谢中的重要作用。前期研究显示，在新鲜成熟块茎的蛋白组中鉴定到多个AQP基因，如CePIP1;1、CePIP2;1、CeTIP1;1和CeTIP2;1[19,25-28]。其中，CeTIP1;1隶属于TIP1亚类，在块茎、叶片、叶鞘、根、匍匐茎和芽尖等主要油莎豆组织中均高水平表达，且其编码蛋白定位在烟草叶片的液泡[27]。本研究报道另一个块茎表达的TIP基因CeTIP4;1，该基因源于团队先前构建的油莎豆全长转录组[29]，其与AtTIP4;1同源。研究显示，CeTIP4;1的表达模式及其编码蛋白的亚细胞定位与CeTIP1;1明显不同，这些研究结果将有助于深入解析油莎豆块茎的水分平衡机制。
1　材料与方法
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1.1　材料
本研究所用油莎豆品系为热研3号，植株种植于中国热带农业科学院文昌实验基地，不同发育时期块茎的采集详见文献[24]；本氏烟草（Nicotiana benthamiana）种植于中国热带农业科学院热带生物技术研究所温室，具体生长条件详见文献[30]；大肠杆菌DH5α和根癌农杆菌GV3101（内含pSoup-P19）感受态、亚细胞定位载体pNC-Cam1304-SubN和pNC-HbPIP2;3-RFP[31]均由本实验室保存；各类酶、试剂盒及生化试剂详见文献[17]。
油莎豆的基因组和转录组数据分别下载于CNGBdb（https://db.cngb.org/search/assembly/CNA-0051961/）和NCBI（https://www.ncbi.nlm.nih.gov/bioproject/703731）数据库；拟南芥和水稻（Oryza sativa）的TIP蛋白下载于TAIR11（https://www.arabidopsis.org/）和RGAP7（http://rice.plantbiology.msu.edu/）数据库；菠菜（Spinacia oleracea）的SoPIP2;1下载于NCBI（https://www.ncbi.nlm.nih.gov/Structure/pdb/1Z98）数据库。
1.2　方法
1.2.1　总RNA的提取和反转录
油莎豆块茎总RNA的提取及反转录方法参照文献[17]，即用天根植物多糖多酚RNA提取试剂盒抽提总RNA，并用宝生物反转录试剂盒合成cDNA第一链。
1.2.2　基因克隆与载体构建
本课题组前期通过对油莎豆的全长转录组[29]进行功能注释，发现1条与AtTIP4;1同源的转录本，序列全长1098 bp，包含1个753 bp的开放读码框。为克隆该基因，采用Primer Premier 5.0软件设计巢式PCR引物对CeTIP4;1F/R（AAACAAAGCAACCAAGACAG/CTTGCATCTCATGATCCAAA）和CeTIP4;1HF/R（AGTGGTCTCTGTCCAGTCCTATGGCAAAGA TTGCATTAGG/GGTCTCAGCAGACCACAAGTT CAAAATTCTTCACCATCTC）。PCR扩增和载体构建参照文献[17]：首先利用外侧引物CeTIP4;1F/R分离基因的全长转录本，随后采用内侧引物CeTIP4;1HF/R扩增基因的编码区序列，最后运用NC克隆试剂盒将目的片段克隆到亚细胞定位载体pNC-Cam1304-SubN。
1.2.3　序列的生物信息学分析
利用本地BLASTN程序将上述CeTIP4;1的转录本比对到油莎豆的基因组，提取相应的基因序列后用GSDS 2.0（http://gsds.gao-lab.org/）在线软件分析其基因结构；蛋白的理化特性和保守结构域分析分别使用Protparam（https://web.expasy.org/protparam/）和CDD（https://www.ncbi.nlm.nih.gov/cdd）在线软件；亚细胞定位、二级和3D结构的预测分别使用Plant-mPLoc（http://www.csbio.sjtu.edu.cn/bioinf/plant-multi/）、SOPMA（https://npsaprabi.ibcp.fr/cgi-bin/npsa_automat.pl?page=/NPSA/npsa_sopma.html）和SWISS-MODEL（https://swissmodel.expasy.org/interactive）在线软件；跨膜螺旋区及保守残基的鉴定基于与已结晶AtTIP2;1[10]和SoPIP2;1[12]的序列比对。
1.2.4　多序列比对与进化树的构建
蛋白的多序列比对采用MUSCLE软件，进化树的构建采用MEGA（v6.06）软件，所用参数为最大似然法、1000次自举重复。
1.2.5　亚细胞定位分析
以前期报道的pNCHbPIP2;3-RFP[31]作为细胞膜定位的阳性标记，参照文献[30]将重组质粒pNC-Cam1304-CeTIP4;1导入农杆菌，挑选单克隆，用含Kan和Rif的LB培养基摇菌，随后与含阳性标记的农杆菌工程菌等量混合，制备浸染液，转化受体，选取4周龄的烟草叶片，荧光观察采用共聚焦显微镜Zeiss LMS880。
1.2.6　基因表达分析
转录谱分析采用StringTie（v2.2.0）软件，即将转录组数据比对到基因组，然后用FKPM（fragments per kilobase of exon per million fragments mapped）法均一化基因的相对表达水平。qRT-PCR分析参照文献[20]，以CeUCE2（Fq: ATCATCAAGGAGACCCAGCG，Rq: CTTAGGGGCAGCCATAGGA）作为内参基因，所用基因特异性引物为CeTIP4;1Fq/Rq（GTCCTCACTTTCTCCCTCCTCT/CACTAACCCGACAAGCAA CG）。
2　结果与分析
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2.1　 CeTIP4;1的克隆与序列分析
通过将CeTIP4;1的转录本比对到基因组，发现该基因位于Scaffold40反向链的5 229 867~5 231 478位，基因全长1612 bp，包含2个内含子，5ʹ和3ʹ UTR分别为154、191 bp（图1A）。重组质粒pNC-Cam1304-CeTIP4;1的桑格测序显示，克隆到的序列与基因的对应区域（外显子）完全一致。进一步的序列分析显示，CeTIP4;1编码区的GC含量为58.03%，编码250个氨基酸（aa），理论分子量（MW）为25.57 kDa，等电点（pI）为6.02，总平均疏水指数（GRAVY）为0.884，脂肪族指数（AI）为119.40，不稳定系数（II）为33.70，属于稳定的酸性疏水型蛋白，这与CeTIP1;1、CeTIP2;1和AtTIP2;1类似，但不同于SoPIP2;1的碱性特性（表1）。CDD分析显示，CeTIP4;1的第3~235位残基为保守的MIP结构域（图1B）。CeTIP4;1与CeTIP1;1、CeTIP2;1、AtTIP2;1和SoPIP2;1的序列相似性分别为67.19%、70.92%、73.81%和44.08%，拥有6个相对保守的跨膜螺旋（TM1–6）、2个半螺旋（HB和HE）以及2个典型的NPA基序（图1C）。CeTIP4;1的ar/R选择性滤器为H-I-A-R，不同于CeTIP1;1的H-I-A-V、CeTIP2;1和AtTIP2;1的H-I-G-R以及SoPIP2;1的F-H-T-R；CeTIP4;1的Froger位点为T-S-A-YW，与CeTIP1;1和AtTIP2;1一致，但不同于CeTIP2;1的T-S-A-Y-I和SoPIP2;1的M-S-A-F-W。与SoPIP2;1相比，CeTIP4;1、CeTIP1;1和AtTIP2;1具有较短的N端，且仅含有对应于SoPIP2;1的S96磷酸化位点；CeTIP4;1的LC区含有一个对应于AtTIP2;1的H131组氨酸，而CeTIP1;1和SoPIP2;1的相应位点分别为F和N（图1C）。与CeTIP1;1、CeTIP2;1AtTIP2;1和SoPIP2;1类似，CeTIP4;1的二级结构以无规则卷曲和α-螺旋为主，分别占36.80%和36.00%（表1）。基于AtTIP2;1的同源建模显示，CeTIP4;1含有6个跨膜螺旋，且可形成同源四聚体（图1D）。
2.2　CeTIP4;1进化分析
为确认CeTIP4;1的进化关系，本研究将其与已报道的AtTIP[11]和OsTIP[32]构建如图1E所示的无根进化树。结果显示，这些蛋白被聚为5组，分别对应于前期报道的TIP1~TIP5[11]，其中，CeTIP4;1与拟南芥和水稻中的TIP4亚类聚在一起，其与AtTIP4;1、OsTIP4;1、OsTIP4;2和OsTIP4;3的序列相似性分别为78.57%、71.71%、69.62%和82.94%。以上结果证实CeTIP4;1属于AtTIP4;1的直系同源基因，而水稻中的3个成员可能在其与油莎豆分化之后产生。
2.3　CeTIP4;1亚细胞定位分析
为揭示CeTIP4;1所起的功能部位，利用含pNC-Cam1304-CeTIP4;1的农杆菌工程菌对烟草叶片进行瞬时转化，共聚焦显微镜观察结果如图2所示。虽然Plant-mPLoc预测CeTIP4;1定位在液泡膜，但本研究结果显示其定位在细胞膜，并与细胞膜标记HbPIP2;3-RFP的红色荧光信号高度重合。
2.4　 CeTIP4;1基因表达分析
如图3A所示，转录谱分析显示CeTIP4;1主要在块茎、根、匍匐茎和芽尖中表达，而在幼嫩叶片、叶鞘和成熟叶片中的表达量极低，表现出明显的组织特异性。为进一步揭示基因在块茎发育过程中的表达模式，本研究选取起始期、膨大中期、膨大晚期和成熟期等4个典型的时期进行qRT-PCR分析，结果显示，CeTIP4;1在4个典型时期的表达量呈先升后降的趋势，峰值出现在膨大中期，最低的是成熟期（图3B）。
3　讨论
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在植物中，细胞的水分平衡主要由细胞膜定位的PIP和液泡膜定位的TIP介导[1,33]。PIP高度保守，仅存在2个进化小组，相比而言，TIP高度分化，其在高等植物中至少存在5个进化小组[3,11,32,34-37]。本研究报道的CeTIP4;1隶属于TIP4亚组，其与AtTIP4;1、OsTIP4;1、OsTIP4;2和OsTIP4;3在蛋白水平的相似性介于69.62%~82.94%，且在进化分析中被聚在一起，推测它们可能具有类似的生物学功能。CeTIP4;1含有2个内含子，这与AtTIP4;1和OsTIP4;1一样，但不同于仅含有1个内含子的OsTIP4;2和OsTIP4;3[11,32]。蓖麻（Ricinus communis）、麻疯树（Jatropha curcas）、木薯（Manihot esculenta）、橡胶树（Hevea brasiliensis）和杨树（Populus trichocarpa）等植物中的TIP4成员均含有2个内含子，且内含子位置高度保守[11,34-37]，这极可能是OsTIP4;2和OsTIP4;3的第2内含子发生了丢失，而非CeTIP4;1和OsTIP4;1获得了新的内含子。
CeTIP4;1含有1个保守的MIP结构域，其中包含6个跨膜螺旋、2个半螺旋以及2个典型的NPA基序，且具有与AtTIP4;1和OsTIP4;3完全一样的ar/R选择性滤器（即H-I-A-R），其孔径大小与AtTIP1;1相当（0.72 Å vs 0.67 Å）[15]，推测CeTIP4;1可能具有AtTIP1;1类似的水分转运活性。事实上，AtTIP4;1和OsTIP4;1均已被证实具有高效的水分转运活性[5,9]。相比于SoPIP2;1[12]，CeTIP4;1与AtTIP2;1[10]的结构比较接近，且其LC区含有1个与NH3转运所必须的H残基[10]，推测CeTIP4;1可能具有高效的NH3转运活性。此外，研究还显示AtTIP4;1可以转运尿素[5]，OsTIP4;1可以转运甘油[9]。
在拟南芥原生质中的亚细胞定位分析显示AtTIP4;1主要定位在液泡[5]，而本研究结果显示CeTIP4;1定位在烟草叶片的细胞膜，其中的原由还有待进一步研究。可能的原因是：（1）TIP4的亚细胞定位与组织类型有关，其中包括液泡的类型与数量，前人用的是原生质[5]，而本研究用的是幼嫩的烟草叶片；（2）TIP4原本定位在液泡，但可以与PIP互作从而定位到细胞膜，因为有研究显示AtTIP1;2、AtTIP2;1和AtTIP3;1均可与AtPIP2;1互作[38]；（3）CeTIP4;1本身就定位在细胞膜，但这与其pI<7的酸性特性不太相符。
作为主要块茎表达的TIP1基因，CeTIP4;1同时被发现在根、匍匐茎、芽尖中也有较高水平的表达，却很少在光合组织叶片和叶鞘中表达，这与CeTIP1;1和CeTIP2;1的组成型高水平表达不同[27-28]。在块茎发育过程中，CeTIP4;1与CeTIP1;1和CeTIP2;1均表现出钟型趋势，但峰值出现不同，CeTIP4;1和CeTIP2;1较早，出现在膨大中期，而CeTIP1;1出现在膨大晚期；此外，CeTIP4;1和CeTIP2;1在成熟期的表达量显著低于起始期，而CeTIP1;1在起始期和成熟期的表达量差异不显著[27-28]。这表明CeTIP4;1与CeTIP2;1类似，主要在块茎发育前期起作用，同时也解释了CeTIP4;1未能在成熟块茎的蛋白组中被鉴定到的原因[19]。与CeTIP1;1相比，CeTIP4;1在块茎发育过程中的表达模式与含水量趋势更为吻合，因为在本研究分析的4个时期中，起始期和膨大中期的含水量约为85%，膨大晚期为75%，成熟期为45%[24]。
综上，本研究首次报道了1个油莎豆TIP4基因CeTIP4;1，该基因含有2个保守的内含子，为AtTIP4;1的直系同源基因；CeTIP4;1定位在细胞膜，具有AtTIP2;1类似的理化特性和结构特征；CeTIP4;1的表达具有明显的组织特异性，且其在块茎发育过程中的表达模式与含水量趋势基本吻合。本研究结果为深入解析油莎豆块茎的水分平衡机制及遗传改良奠定坚实的基础。
基金
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  • 海南省重点研发计划项目(ZDYF2024XDNY171)
  • 国家自然科学基金项目(32460342; 31971688)
文献
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2024年第45卷第9期
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doi: 10.3969/j.issn.1000-2561.2024.09.006
  • 接收时间：2023-08-28
  • 首发时间：2026-06-23
  • 出版时间：2024-09-25
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  • 收稿日期：2023-08-28
  • 修回日期：2023-09-30
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海南省重点研发计划项目(ZDYF2024XDNY171)
国家自然科学基金项目(32460342; 31971688)
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    热带作物生物育种全国重点实验室/中国热带农业科学院热带生物技术研究所/中国热带农业科学院三亚研究院，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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