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Article(id=1276262780437140263, tenantId=1146029695717560320, journalId=1235980609244409860, issueId=1276262756814815737, articleNumber=null, orderNo=null, doi=10.3969/j.issn.1000-2561.2024.08.004, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=null, receivedDate=1689782400000, receivedDateStr=2023-07-20, revisedDate=1693152000000, revisedDateStr=2023-08-28, acceptedDate=null, acceptedDateStr=null, onlineDate=1782214358671, onlineDateStr=2026-06-23, pubDate=1724515200000, pubDateStr=2024-08-25, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1782214358671, onlineIssueDateStr=2026-06-23, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1782214358671, creator=13701087609, updateTime=1782214358671, updator=13701087609, issue=Issue{id=1276262756814815737, tenantId=1146029695717560320, journalId=1235980609244409860, year='2024', volume='45', issue='8', pageStart='1521', pageEnd='1760', issueExtLink='null', onlineDate='null', pubDate='1724515200000', pubDateStr='2024-08-25', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1782214353040, creator='13701087609', updateTime=1782214460949, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext={EN=IssueExt(id=1276263209816420382, tenantId=1146029695717560320, journalId=1235980609244409860, issueId=1276262756814815737, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1276263209816420383, tenantId=1146029695717560320, journalId=1235980609244409860, issueId=1276262756814815737, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null, downloadFileDto=null}, startPage=1552, endPage=1559, ext={EN=ArticleExt(id=1276262780877542185, articleId=1276262780437140263, tenantId=1146029695717560320, journalId=1235980609244409860, language=EN, title=Molecular Cloning and Characterization of CeTIP2;1, an Aquaporin Gene from Tigernut (Cyperus esculentus L.), columnId=1236256430337085821, journalTitle=Chinese Journal of Tropical Crops, columnName=Omics & Biotechnology, runingTitle=null, highlight=null, articleAbstract=

Tigernut (Cyperus esculentus L.), most likely originated from Africa and Mediterranean, represents a novel herbaceous oil crop accumulating high levels of oil in its underground tubers. Water balance is essential for tuber development and metabolism in tigernut. Tonoplast intrinsic proteins (TIPs), which include five phylogenetic groups (i.e. TIP1–5), constitute a subfamily of aquaporin facilitating the fast and passive transport of water across vacuolar membranes. Based on available genome and transcriptome data, one TIP gene named CeTIP2;1 was isolated from tigernut tubers by using the RT-PCR technique. Sequence analysis showed that the full gene length of CeTIP2;1 is 3323 bp, including two introns with a coding sequence of 747 bp; the gene was predicted to encode 248 amino acids with the theoretical molecular weight of 24.73 kDa, the isoelectric point of 5.09, the grand average of hydropathicity of 0.948, the aliphatic index of 114.60, and the instability index of 21.76, implying its stable, acidic, and hydrophobic features, which is consistent with its tonoplast-localization; presence of one conservative MIP domain was observed, which possesses six transmembrane helices, two half helices, and two typical NPA motifs. CeTIP2;1 was shown to exhibit the sequence similarity of 87.20% with AtTIP2;1, which was considerably higher than 46.23% with SoPIP2;1. Further phylogenetic analysis revealed that CeTIP2;1 belongs to the TIP2 group and is a true ortholog of AtTIP2;1. Interestingly, the phylogenetic analysis also supported that TIP2 has diverged into two subgroups sometime before monocot-eudicot divergence. Tissue-specific expression analysis showed that CeTIP2;1 was highly expressed in all tissues examined in this study, i.e., leaf, sheath, root, shoot apex, rhizome, and tuber, with most in tuber/rhizome and lowest in shoot apex. A bell-like expression pattern was observed during tuber development, peaking at the medium stage of swelling and lowest in maturation. Moreover, CeTIP2;1 was also detected in the tuber proteomes, implying its high abundance and key roles. These findings would 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=1276262786133005103, articleId=1276262780437140263, tenantId=1146029695717560320, journalId=1235980609244409860, language=CN, title=油莎豆水通道蛋白基因CeTIP2;1的克隆与分析, columnId=1236256430517440904, journalTitle=热带作物学报, columnName=组学与生物技术, runingTitle=null, highlight=null, articleAbstract=

油莎豆起源于非洲和地中海沿岸，是一种在块茎中高水平积累油脂的新型草本油料作物。水分平衡对于块茎的发育与代谢至关重要。液泡膜内在蛋白（TIP）是一类液泡膜定位并具有高效水分转运活性的水通道蛋白，包含TIP1~5等5个亚类。本研究基于油莎豆的基因组和转录组数据，采用RT-PCR技术对1个块茎高水平表达的TIP基因CeTIP2；1进行克隆。序列分析表明：CeTIP2；1的基因全长3323 bp，含有2个内含子，编码区长747 bp，编码248个氨基酸，理论分子量为24.73 kDa，等电点为5.09，总平均疏水指数为0.948，脂肪族指数为114.60，不稳定系数为21.76，属于稳定的酸性疏水型蛋白，与其液泡膜定位一致；该蛋白含有保守的MIP结构域，其中包括6个跨膜螺旋、2个半螺旋以及2个典型的NPA基序。CeTIP2；1与AtTIP2；1的序列相似性高达87.20%，远高于与SoPIP2；1的46.23%。进化分析进一步证实，CeTIP2；1隶属于TIP2亚类，是AtTIP2；1的直系同源基因。进化分析同时显示TIP2亚类早在单、双子叶植物分化之前就已经进化出2个小组。表达分析显示：CeTIP2；1在叶片、叶鞘、根、芽尖、匍匐茎、块茎等主要组织中均有较高水平的表达，丰度最高的是块茎和匍匐茎，最低的为芽尖；在块茎的发育过程中，呈现先升后降的钟形趋势，峰值出现在膨大中期，最低的为成熟期。此外，CeTIP2；1还在块茎的蛋白组中被检测到，表明其功能的重要性。这些结果为进一步揭示油莎豆的块茎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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The Plant Journal, 2022, 112(2): 518-534., articleTitle=A seed-like proteome in oil-rich tubers, refAbstract=null)], funds=[Fund(id=1276463042003075420, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276262780437140263, awardId=ZDYF2024XDNY171, language=CN, fundingSource=海南省重点研发计划项目(ZDYF2024XDNY171), fundOrder=null, country=null), Fund(id=1276463042065989981, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276262780437140263, awardId=SCKJ-JYRC-2022-66, language=CN, fundingSource=“崖州湾”菁英人才科技专项(SCKJ-JYRC-2022-66), fundOrder=null, country=null), Fund(id=1276463042124710238, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276262780437140263, awardId=31971688, language=CN, fundingSource=国家自然科学基金项目(31971688), fundOrder=null, country=null)], companyList=[AuthorCompany(id=1276463035061502271, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276262780437140263, xref=null, ext=[AuthorCompanyExt(id=1276463035069890880, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276262780437140263, companyId=1276463035061502271, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=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), AuthorCompanyExt(id=1276463035082473793, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276262780437140263, companyId=1276463035061502271, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=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)])], figs=[ArticleFig(id=1276463041139048790, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276262780437140263, language=EN, label=Fig. 1, caption=Bioinformatics analysis of CeTIP2;1

A: Gene structure of CeTIP2;1; B: Identified conservative domain of CeTIP2;1; C: Sequence alignment of CeTIP2;1, AtTIP2;1, and SoPIP2;1, where six transmembrane helices and two half helices are shaded and conserved residues are marked in different colors; D: Predicted 3D structure for CeTIP2;1; E: Phylogenetic analysis of CeTIP2;1 and TIPs in O. sativa and A. thaliana.

, figureFileSmall=pNQo08HotLzujfFoB6kjMw==, figureFileBig=6u122Cp4rk1wVltykCbmGg==, tableContent=null), ArticleFig(id=1276463041503953239, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276262780437140263, language=CN, label=图1, caption=CeTIP2;1的生物信息学分析

A：CeTIP2;1的基因结构；B：CeTIP2;1的保守结构域；C：CeTIP2;1与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：CeTIP2;1的3D结构预测；E：CeTIP2;1与水稻和拟南芥TIP蛋白的进化分析。

, figureFileSmall=pNQo08HotLzujfFoB6kjMw==, figureFileBig=6u122Cp4rk1wVltykCbmGg==, tableContent=null), ArticleFig(id=1276463041608810840, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276262780437140263, language=EN, label=Fig. 2, caption=Expression profiles of CeTIP2;1

A: Tissue-specific expression profiles of CeTIP2;1; B: Expression profiles of CeTIP2;1 in different stages of developmental tubers; C: CeTIP2;1 protein abundance in different tissues. Lowercase and uppercase letters indicate significant difference (P<0.05) and extremely significant difference (P<0.01), respectively.

, figureFileSmall=euRTqKgSK6szyfLzYCLrWQ==, figureFileBig=8jgJHtidZFAl3NwilxbV9Q==, tableContent=null), ArticleFig(id=1276463041680114009, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276262780437140263, language=CN, label=图2, caption=CeTIP2;1的表达模式

A：CeTIP2;1的组织特异性分析；B：CeTIP2;1在不同发育时期块茎中的表达模式；C：CeTIP2;1在不同组织中的蛋白丰度。小写字母和大写字母分别表示差异显著（P<0.05）和差异极显著（P<0.01）。

, figureFileSmall=euRTqKgSK6szyfLzYCLrWQ==, figureFileBig=8jgJHtidZFAl3NwilxbV9Q==, tableContent=null), ArticleFig(id=1276463041738834266, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276262780437140263, language=EN, label=Tab. 1, caption=

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

, figureFileSmall=null, figureFileBig=null, tableContent=
蛋白名称Protein name氨基酸数量Numbers of aimo acid分子量MW/kDapIGRAVYAIIIMIPα-螺旋Alpha helix/%延伸链Extended strand/%β-转角Beta turn/%无规则卷曲Random coil/%
CeTIP2;124824.735.090.948114.6021.7614..23136.2920.974.8437.90
AtTIP2;125025.035.300.974112.4826.7314..23233.2020.805.6040.40
SoPIP2;128129.909.030.579102.8127.8131..26036.6518.152.1443.06
), ArticleFig(id=1276463041860469083, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276262780437140263, language=CN, label=表1, caption=

CeTIP2;1与AtTIP2;1、SoPIP2;1理化特性和二级结构比较

, figureFileSmall=null, figureFileBig=null, tableContent=
蛋白名称Protein name氨基酸数量Numbers of aimo acid分子量MW/kDapIGRAVYAIIIMIPα-螺旋Alpha helix/%延伸链Extended strand/%β-转角Beta turn/%无规则卷曲Random coil/%
CeTIP2;124824.735.090.948114.6021.7614..23136.2920.974.8437.90
AtTIP2;125025.035.300.974112.4826.7314..23233.2020.805.6040.40
SoPIP2;128129.909.030.579102.8127.8131..26036.6518.152.1443.06
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油莎豆水通道蛋白基因CeTIP2;1的克隆与分析
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邹智 , 郑玉皎
热带作物学报 | 组学与生物技术 2024,45(8): 1552-1559
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热带作物学报 |组学与生物技术 2024 , 45 (8) : 1552 -1559
油莎豆水通道蛋白基因CeTIP2;1的克隆与分析
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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
Molecular Cloning and Characterization of CeTIP2;1, an Aquaporin Gene from Tigernut (Cyperus esculentus L.)
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: 2024-08-25 doi: 10.3969/j.issn.1000-2561.2024.08.004
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油莎豆起源于非洲和地中海沿岸，是一种在块茎中高水平积累油脂的新型草本油料作物。水分平衡对于块茎的发育与代谢至关重要。液泡膜内在蛋白（TIP）是一类液泡膜定位并具有高效水分转运活性的水通道蛋白，包含TIP1~5等5个亚类。本研究基于油莎豆的基因组和转录组数据，采用RT-PCR技术对1个块茎高水平表达的TIP基因CeTIP2；1进行克隆。序列分析表明：CeTIP2；1的基因全长3323 bp，含有2个内含子，编码区长747 bp，编码248个氨基酸，理论分子量为24.73 kDa，等电点为5.09，总平均疏水指数为0.948，脂肪族指数为114.60，不稳定系数为21.76，属于稳定的酸性疏水型蛋白，与其液泡膜定位一致；该蛋白含有保守的MIP结构域，其中包括6个跨膜螺旋、2个半螺旋以及2个典型的NPA基序。CeTIP2；1与AtTIP2；1的序列相似性高达87.20%，远高于与SoPIP2；1的46.23%。进化分析进一步证实，CeTIP2；1隶属于TIP2亚类，是AtTIP2；1的直系同源基因。进化分析同时显示TIP2亚类早在单、双子叶植物分化之前就已经进化出2个小组。表达分析显示：CeTIP2；1在叶片、叶鞘、根、芽尖、匍匐茎、块茎等主要组织中均有较高水平的表达，丰度最高的是块茎和匍匐茎，最低的为芽尖；在块茎的发育过程中，呈现先升后降的钟形趋势，峰值出现在膨大中期，最低的为成熟期。此外，CeTIP2；1还在块茎的蛋白组中被检测到，表明其功能的重要性。这些结果为进一步揭示油莎豆的块茎水分平衡机制奠定坚实的基础。

油料作物  /  块茎  /  液泡膜内在蛋白  /  进化分析  /  表达模式

Tigernut (Cyperus esculentus L.), most likely originated from Africa and Mediterranean, represents a novel herbaceous oil crop accumulating high levels of oil in its underground tubers. Water balance is essential for tuber development and metabolism in tigernut. Tonoplast intrinsic proteins (TIPs), which include five phylogenetic groups (i.e. TIP1–5), constitute a subfamily of aquaporin facilitating the fast and passive transport of water across vacuolar membranes. Based on available genome and transcriptome data, one TIP gene named CeTIP2;1 was isolated from tigernut tubers by using the RT-PCR technique. Sequence analysis showed that the full gene length of CeTIP2;1 is 3323 bp, including two introns with a coding sequence of 747 bp; the gene was predicted to encode 248 amino acids with the theoretical molecular weight of 24.73 kDa, the isoelectric point of 5.09, the grand average of hydropathicity of 0.948, the aliphatic index of 114.60, and the instability index of 21.76, implying its stable, acidic, and hydrophobic features, which is consistent with its tonoplast-localization; presence of one conservative MIP domain was observed, which possesses six transmembrane helices, two half helices, and two typical NPA motifs. CeTIP2;1 was shown to exhibit the sequence similarity of 87.20% with AtTIP2;1, which was considerably higher than 46.23% with SoPIP2;1. Further phylogenetic analysis revealed that CeTIP2;1 belongs to the TIP2 group and is a true ortholog of AtTIP2;1. Interestingly, the phylogenetic analysis also supported that TIP2 has diverged into two subgroups sometime before monocot-eudicot divergence. Tissue-specific expression analysis showed that CeTIP2;1 was highly expressed in all tissues examined in this study, i.e., leaf, sheath, root, shoot apex, rhizome, and tuber, with most in tuber/rhizome and lowest in shoot apex. A bell-like expression pattern was observed during tuber development, peaking at the medium stage of swelling and lowest in maturation. Moreover, CeTIP2;1 was also detected in the tuber proteomes, implying its high abundance and key roles. These findings would lay a solid foundation for further uncovering the mechanism of water balance in tigernut tubers.

oil crop  /  tuber  /  tonoplast intrinsic protein  /  phylogenetic analysis  /  expression profile
邹智, 郑玉皎. 油莎豆水通道蛋白基因CeTIP2;1的克隆与分析. 热带作物学报, 2024 , 45 (8) : 1552 -1559 . DOI: 10.3969/j.issn.1000-2561.2024.08.004
Zhi ZOU, Yujiao ZHENG. Molecular Cloning and Characterization of CeTIP2;1, an Aquaporin Gene from Tigernut (Cyperus esculentus L.)[J]. Chinese Journal of Tropical Crops, 2024 , 45 (8) : 1552 -1559 . DOI: 10.3969/j.issn.1000-2561.2024.08.004
正文
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水通道蛋白（aquaporin，AQP）是一类广泛存在于生物体内的膜内在蛋白，因其高效的水分转运活性而得名[1-2]。与动物和微生物相比，植物AQP家族出现了明显的扩张，主要包含液泡膜内在蛋白（tonoplast intrinsic protein，TIP）、质膜内在蛋白（plasma membrane intrinsic protein，PIP）、类根瘤26膜内在蛋白（NOD26-like intrinsic protein，NIP）、小分子碱性膜内在蛋白（small basic intrinsic protein，SIP）和未鉴定膜内在蛋白（X intrinsic protein，XIP）等5大类[2-7]。其中，TIP定位在液泡膜，是介导细胞内水分平衡的关键AQP类型[8-12]。根据进化关系，TIP可进一步分为TIP1~5等5个亚类[3-7]。
油莎豆（Cyperus esculentus L.），又名油莎草或虎坚果，是一种起源于非洲和地中海沿岸的新型草本油料作物[13]。系统分析显示，油莎豆隶属于禾本目莎草科，其单套染色体的基因组大小约为225.6 Mb[14-18]。相比于传统油料作物在种子中积累油脂，油莎豆是迄今唯一已知在块茎中高水平积累油脂（24%~35%）的作物，这使其成为研究营养组织油脂代谢与调控的理想模型[14,19-21]。油莎豆具有适应性广、抗逆性强、产量高、适合机械化等特点，这有助于在不挤占现有耕地的情况下充分利用沙化边际土地额外增加我国食用油供给，减少对国外大豆的刚性需求，进而服务国家的战略需求[22]。作为主要的经济器官，积极开展块茎的发育生物学研究具有重要的理论意义和现实价值。本研究报道1个在块茎中高水平表达的TIP基因CeTIP2;1，其中包括基因结构、序列特征、进化关系及表达特性，以期为下一步的功能分析及块茎水分平衡机制的阐释奠定基础。
1　材料与方法
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1.1　材料
1.1.1　植物材料
本研究所用油莎豆品系为热研3号[14]，其中，起始期（起始后1 d）、膨大中期（起始后20 d）、膨大晚期（起始后25 d）和成熟期（起始后35 d）等不同发育时期的块茎采集于中国热带农业科学院热带生物技术研究所文昌试验基地[23]。
1.1.2　菌株、载体及试剂耗材
大肠杆菌（Escherichia coli）DH5α感受态和植物过表达载体pCAMBIA1301由本实验室制备和保存；酶、试剂盒及生化试剂购自相应的试剂公司。
1.2　方法
1.2.1　块茎总RNA的提取和反转录
参照文献[14]，采用天根植物多糖多酚RNA提取试剂盒分别提取不同发育时期块茎的总RNA，并采用赛默飞反转录试剂盒合成cDNA第一链。
1.2.2　 CeTIP2;1基因的克隆与载体构建
为鉴定油莎豆中AtTIP2;1的直系同源基因，首先从CNGBdb（https://db.cngb.org/search/assembly/CNA-0051961/）和NCBI（https://www.ncbi.nlm.nih.gov/bioproject/703731）数据库分别下载油莎豆的基因组和转录组数据； 以AtTIP2;1作为种子， 采用本地TBLASTN程序（E值设为1e–10）搜索油莎豆的基因组，并利用StringTie（v2.2.0）将转录组数据比对到基因组以界定转录区域；根据基因序列设计引物对CeTIP2;1F/R（AAACGGAGACCCAACAAGCAAAAC/CAACCAAACAACATCAGGAAACCA）和CeTIP2;1HF/R（AGTGGTCTCTGT CCAGTCCTATGGCAGGCA TTGCTTTTGG/GGTCTCA GCAGACCACAAGTTTAGTACTCGCTGC TGGACACTGG），分别用于全长cDNA的分离和植物过表达载体的构建，具体的PCR反应和载体构建流程详见文献[15]。
1.2.3　序列分析
采用在线软件GSDS（v2.0）（http://gsds.gao-lab.org/）分析基因结构，并分别用Protparam（https://web.expasy.org/protparam/）、CDD（https://www.ncbi.nlm.nih.gov/cdd）、PlantmPLoc（http://www.csbio.sjtu.edu.cn/bioinf/plantmulti/）、SOPMA（https://npsa-prabi.ibcp.fr/cgi-bin/npsa_automat.pl?page=/NPSA/npsa_sopma.html）和SWISS-MODEL（https://swissmodel.expasy.org/interactive）等在线软件分析蛋白的理化特性、保守结构域、亚细胞定位、二级结构和三级结构。
1.2.4　多序列比对与进化分析
采用MEGA（v6.06）软件中的MUSCLE进行蛋白多序列比对，其中，跨膜螺旋区和保守残基根据与已结晶AtTIP2;1[12]和菠菜（Spinacia oleracea）SoPIP2;1[24]的序列比对进行鉴定；进化树的构建采用MEGA软件中的邻接法以及1000次自举重复。
1.2.5　转录谱分析
转录组读段的比对采用StringTie软件，并用FKPM（fragments per kilobase of exon per million fragments mapped）法均一化基因在不同样本中的相对表达水平。
1.2.6　蛋白谱分析
油莎豆的蛋白组数据下载于ProteomeXchange/PRIDE（https://www.ebi.ac.uk/pride/archive/projects/PXD021894），质谱数据的检索与分析采用Proteome Discoverer（v2.4.1.15）软件。
1.2.7　荧光定量分析
参照文献[17]，以CeUCE2作为内参，应用引物对CeTIP2;1Fq/Rq（TCGTCATATTCAGCCCGGTGC/GGGTGCTTACCGG TAAGGTACTTG）进行荧光定量分析，每样本3次生物学重复。
1.3　数据处理
采用SPSS 20软件进行统计分析，采用邓肯单因素多样本差异显著性进行方差分析。
2　结果与分析
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2.1　 CeTIP2;1基因的克隆与序列分析
通过同源搜索，从油莎豆基因组的Scaffold27鉴定1个AtTIP2;1同源基因，二者在蛋白水平的序列相似性高达87.20%，将其命名为CeTIP2;1。CeTIP2;1基因的全长为3323 bp，包含2个内含子，5'和3' UTR分别为307 bp和1956 bp（图1A）。随后，采用RT-PCR技术对其747 bp的编码区序列进行分离，并成功构建了植物过表达载体pCAMBIA1301-CeTIP2;1，桑格测序未发现碱基差异。进一步的序列分析显示，CeTIP2;1编码区的GC含量为56.76%，编码248个氨基酸，理论分子量为24.73 kDa，等电点为5.09，总平均疏水指数为0.948，脂肪族指数为114.60，不稳定系数为21.76，属于稳定的酸性疏水型蛋白，这与AtTIP2;1类似，但不同于SoPIP2;1的碱性特性（表1）。CDD分析显示，CeTIP2;1的14~231位为保守的MIP结构域（图1B）。虽然CeTIP2;1与SoPIP2;1的序列相似性仅为46.23%，但都拥有如图1C所示的6个跨膜螺旋（TM1~6）、2个半螺旋（HB和HE）以及2个典型的NPA基序。CeTIP2;1的ar/R选择性滤器为H-I-G-R，与AtTIP2;1一致，但不同于SoPIP2;1的F-H-T-R；CeTIP2;1的Froger位点为T-S-A-Y-I，不同于AtTIP2;1的T-S-A-Y-W和SoPIP2;1的M-S-A-F-W。与SoPIP2;1相比，CeTIP2;1和AtTIP2;1具有较短的N端，且仅含有对应于SoPIP2;1 S96的磷酸化位点；CeTIP2;1的连环C含有1个对应于AtTIP2;1 H131的组氨酸，而SoPIP2;1的相应位点为N（图1C）。亚细胞定位预测显示，CeTIP2;1定位在液泡膜。CeTIP2;1的二级结构以无规则卷曲和α-螺旋为主，分别占37.90%和36.29%，这与AtTIP2;1和SoPIP2;1类似（表1）。基于AtTIP2;1的同源建模显示，CeTIP2;1可形成同源四聚体，每个单体含有6个跨膜螺旋（图1D）。
2.2　进化分析
为进一步鉴定CeTIP2;1的进化关系，本研究将其与已报道的10个OsTIP和10个AtTIP构建了无根进化树。如图1E所示，21个TIP被聚为5组，即TIP1~5。TIP1明显聚成2个分支，其中一支包括OsTIP1;1、AtTIP1;1和AtTIP1;2，另一支包括OsTIP1;2和AtTIP1;3；TIP2也聚成2个分支，其中一支包括CeTIP2;1、OsTIP2;2和AtTIP2;1，另一支包括OsTIP2;1、AtTIP2;2和AtTIP2;3，表明TIP1和TIP2早在单、双子叶植物分化之前就已经进化成2个小组。与此相反，TIP3按物种聚在一起，这与AtTIP1;1/1;2、AtTIP2;2/-2;3和OsTIP4-1/4-2/4-3类似，表明这些基因在单双子叶植物分化之后产生。综上，这些结果表明CeTIP2;1属于AtTIP2;1和OsTIP2;2的直系同源基因。
2.3　 CeTIP2;1的组织特异性分析
为揭示CeTIP2;1在不同组织中的表达特性，利用转录组数据分析其在幼嫩叶片、成熟叶片、叶鞘、根、芽尖、匍匐茎、块茎等主要组织中的表达模式。如图2A所示，CeTIP2;1在所有组织中均有较高水平的表达，其中，表达丰度最高的是块茎和匍匐茎，其次是叶鞘、幼嫩叶片、成熟叶片和根，最低的是芽尖。
2.4　 CeTIP2;1在不同发育时期块茎中的表达模式
为揭示基因表达与块茎发育的关系，采用qRT-PCR技术进一步分析CeTIP2;1在起始期、膨大中期、膨大晚期和成熟期等4个典型发育时期块茎中的表达模式。如图2B所示，结果表明基因在膨大中期的表达丰度最高，极显著高于起始期和膨大晚期，而在成熟期的丰度最低。
2.5　 CeTIP2;1在不同组织中的蛋白定量
为揭示CeTIP2;1在不同组织中的蛋白丰度，研究进一步分析了油莎豆的蛋白组数据，即叶片、根、新鲜收获的成熟块茎（RT）、脱水块茎（DT）、吸水48 h块茎（RT）和萌发块茎（ST）。如图2C所示，结果表明CeTIP2;1在根中的丰度最高，显著高于吸水48 h块茎，而在萌发块茎中的丰度最低；蛋白在块茎的脱水、吸水过程中差异不显著，丰度与叶片相当。
3　讨论
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细胞膜定位的PIP和液泡膜定位的TIP是调控植物细胞水分平衡的主要AQP类型[2,8]。相比于PIP，TIP高度分化，其在高等植物中至少存在5个亚类，且其决定底物特异性的ar/R选择性滤器明显呈现多样化[3-7]。根据ar/R选择性滤器的类型及残基特性，BANSAL等[25]将拟南芥、水稻和玉米中的TIP归为三大主要类型，第一类包含TIP1（H-I-A-V），第二类包含TIP2、TIP3和TIP4（H-I/M/V-G/A/S-R），第三类包含TIP5（N-V-G-C）。
本研究报道了在油莎豆块茎中高水平表达的TIP基因CeTIP2;1，其编码蛋白包含AQP家族特有的MIP结构域，理化特性与AtTIP2;1高度相似，属于稳定的酸性疏水型蛋白。CeTIP2;1可归为TIP2亚类，主要依据如下：首先，CeTIP2;1与AtTIP2;1和OsTIP2;2的序列相似性在87%以上；其次，3个蛋白在进化分析中聚在一起，并独立于OsTIP2;1、AtTIP2;2和AtTIP2;3。这些证据表明CeTIP2;1是AtTIP2;1的直系同源基因，而非AtTIP2;2和AtTIP2;3。根据先前的比较基因组学分析，与AtTIP3;1和AtTIP3;2一样，AtTIP2;2和AtTIP2;3起源于芸薹属特有的α全基因组重复[5,26]。因为AtTIP2;1和AtTIP2;2/-2;3在水稻中都存在直系同源基因，暗示基因分化发生在单、双子叶植物分化之前，这与TIP1类似[27]。也就是说，在目前鉴定的5个TIP亚类中，TIP1和TIP2早在单、双子叶植物分化之前就发生了进一步的分化，因此，从进化角度讲，单、双子叶植物中的TIP分为7个亚类更为适合。基因结构分析显示，CeTIP2;1含有2个内含子，其内含子数量和位置与AtTIP2;1和OsTIP2;2完全一致[3-4]，表明进化的保守性。CeTIP2;1含有2个典型的NPA基序，ar/R选择性滤器为H-I-G-R，且其连环C含有1个保守的H残基，这与AtTIP2;1和OsTIP2;2完全一致，表明其具有类似的生物学功能。在蟾蜍卵母细胞和酵母体系中功能分析显示，AtTIP2;1和OsTIP2;2可转运水分、尿素和氨[10,12,28-29]。CeTIP2;1的重要性还体现在其泛组织、高水平表达。由于现有蛋白组技术的灵敏性，一般只有较高丰度的蛋白才能检测到。先前研究表明，NIEMEYER等[30]采用nanoLC-MS/MS技术对45个样本进行了蛋白组分析，合计仅检测到2257个蛋白。本研究的定量分析显示，CeTIP2;1在叶片、根和4个时期的块茎中均被检测到，表明其在这些组织中的高水平表达。事实上，组织特异性分析显示CeTIP2;1是1个泛组织表达基因，其在块茎中的高水平表达与块茎的高含水量及活跃的生理代谢是相适应的。据前期研究结果表明，油莎豆块茎发育早期的含水量一致维持在85%左右，在膨大晚期和成熟期逐渐降低到75%和45%[23]。CeTIP2;1在块茎发育过程中的表达模式与水分含量趋势基本一致，峰值出现在膨大中期，而在膨大晚期和成熟期逐渐降低。CeTIP2;1在成熟期的显著下调可能与其蛋白组中丰度低于根组织有关。
综上，本研究报道了1个油莎豆TIP2基因CeTIP2;1，该基因含有2个内含子，属于AtTIP2;1的直系同源基因。该基因在不同组织中均高水平表达，与这些组织特别是块茎的水分平衡密切相关，这为下一步的功能分析及油莎豆水分平衡机制的阐释奠定基础。
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  • 海南省重点研发计划项目(ZDYF2024XDNY171)
  • “崖州湾”菁英人才科技专项(SCKJ-JYRC-2022-66)
  • 国家自然科学基金项目(31971688)
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2024年第45卷第8期
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doi: 10.3969/j.issn.1000-2561.2024.08.004
  • 接收时间：2023-07-20
  • 首发时间：2026-06-23
  • 出版时间：2024-08-25
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  • 收稿日期：2023-07-20
  • 修回日期：2023-08-28
基金
海南省重点研发计划项目(ZDYF2024XDNY171)
“崖州湾”菁英人才科技专项(SCKJ-JYRC-2022-66)
国家自然科学基金项目(31971688)
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
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2种不同金属材料的力学参数
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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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