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Article(id=1276204182059221793, tenantId=1146029695717560320, journalId=1235980609244409860, issueId=1276204178091413862, articleNumber=null, orderNo=null, doi=10.3969/j.issn.1000-2561.2024.12.003, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=null, receivedDate=1715875200000, receivedDateStr=2024-05-17, revisedDate=1718121600000, revisedDateStr=2024-06-12, acceptedDate=null, acceptedDateStr=null, onlineDate=1782200387730, onlineDateStr=2026-06-23, pubDate=1735056000000, pubDateStr=2024-12-25, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1782200387730, onlineIssueDateStr=2026-06-23, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1782200387730, creator=13701087609, updateTime=1782200387730, updator=13701087609, issue=Issue{id=1276204178091413862, tenantId=1146029695717560320, journalId=1235980609244409860, year='2024', volume='45', issue='12', pageStart='2487', pageEnd='2737', issueExtLink='null', onlineDate='null', pubDate='1735056000000', pubDateStr='2024-12-25', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1782200386783, creator='13701087609', updateTime=1782200456354, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext={EN=IssueExt(id=1276204470308565242, tenantId=1146029695717560320, journalId=1235980609244409860, issueId=1276204178091413862, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1276204470308565243, tenantId=1146029695717560320, journalId=1235980609244409860, issueId=1276204178091413862, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null, downloadFileDto=null}, startPage=2515, endPage=2523, ext={EN=ArticleExt(id=1276204182923248419, articleId=1276204182059221793, tenantId=1146029695717560320, journalId=1235980609244409860, language=EN, title=Cloning and Expression Analysis of HbLPAT2 Gene from Hevea brasiliensis, columnId=1236256430337085821, journalTitle=Chinese Journal of Tropical Crops, columnName=Omics & Biotechnology, runingTitle=null, highlight=null, articleAbstract=

Lysophosphatidyl acyltransferase (LPAT), a key enzyme which controls the production of lysophosphatidic into phosphatidic, plays an important role in phospholipid biosynthesis in plants. Hevea brasiliensis is an important source for the production of natural rubber, which has an irreplaceable role in the fields of national defense, medical treatment, high-end products, and has high social and economic value. Cloning and analysis of HbLPAT2 in H. brasiliensis would provide scientific basis for further exploration of its phospholipid biosynthesis function and the effect of phospholipid on nature rubber. In this study HbLPAT2 was cloned from the latex of the H. brasiliensis by molecular biology techniques, and the bioinformatics analysis, expression analysis and subcellular localization analysis were performed. In this study, two key genes responsible for phospholipid biosynthesis in the HbLPAT gene family were successfully cloned from the latex of the H. brasiliensis, named HbLPAT2a and HbLPAT2b, the full length coding sequence of HbLPAT2a and HbLPAT2b genes was 1161 bp and 939 bp, encoding 386 and 312 amino acids. Both HbLPAT2a and HbLPAT2b contained a PLN02380 lysopyatidyl acyltransferase functional domain and contained a highly conserved NH(X)4D motif; HbLPAT2a protein had 3 transmembrane domains, and HbLPAT2b protein had 1 transmembrane domain, both of which had no signal peptide, were hydrophilic proteins, and contained multiple phosphorylation sites. The subcellular localization analysis showed that both HbLPAT2a and HbLPAT2b proteins were located in the endoplasmic reticulum. In different organs, the highest expression level of HbLPAT2a and HbLPAT2b was found in seeds, suggesting that they have a lipid biosynthesis function. However, in the latex of H. brasiliensis, the first and second expression level of whole HbLPAT gene family were HbLPAT2a and HbLPA2b. The result is helpful to understanding that HbLPAT2a and HbLPAT2b play an important role in the phospholipid biosynthetic pathway in latex of H. brasiliensis and to providing a scientific basis for genetic improvement of latex quality of H. brasiliensis.

, authors=null, authorsList=Xiaolin WANG, Peng QU, Mingyang GAN, Jian QIU, Ruishen FAN, authorCompany=null, correspAuthors=Ruishen FAN, 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=1276204193773912879, articleId=1276204182059221793, tenantId=1146029695717560320, journalId=1235980609244409860, language=CN, title=橡胶树HbLPAT2基因的克隆与表达分析, columnId=1236256430517440904, journalTitle=热带作物学报, columnName=组学与生物技术, runingTitle=null, highlight=null, articleAbstract=

溶血磷酸酰基转移酶（lysophosphatidyl acyltransferase，LPAT）是控制溶血磷脂酸生成磷脂酸的关键酶，在植物磷脂合成中发挥重要作用。橡胶树（Hevea brasiliensis）是生产天然橡胶的重要来源，其在国防、医疗、高端制品等领域具有不可替代的作用，具有很高的社会价值和经济价值。本研究采用分子生物学技术在橡胶树胶乳中成功克隆出2个HbLPAT基因家族中负责磷脂合成的关键基因，分别命名为HbLPAT2a和HbLPAT2b，其编码序列全长分别为1161 bp和939 bp，分别编码了386个和312个氨基酸。HbLPAT2a和HbLPAT2b均含有1个PLN02380的溶血磷脂酸酰基转移酶功能结构域和高度保守的NH(X)4D基序。HbLPAT2a有3个跨膜结构域，HbLPAT2b有1个跨膜结构域，二者均无信号肽，为亲水性蛋白，且含有多个磷酸化位点。亚细胞定位分析表明，HbLPAT2a和HbLPAT2b蛋白均定位于内质网上。不同组织部位表达分析表明，HbLPAT2a和HbLPAT2b基因均在橡胶树种子中的表达量最高，推测其具有脂质合成功能，而在胶乳中，HbLPAT2a和HbLPAT2b的表达量在HbLPAT基因家族中位居前2位。本研究结果有助于了解HbLPAT2a和HbLPAT2b在橡胶树胶乳磷脂合成过程中的功能，为后期橡胶树胶乳品质的遗传改良提供科学依据。

, authors=

王晓琳（2002—），女，本科生，研究方向：林木生物技术。

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* 范睿深（FAN Ruishen），E-mail：。
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2.云南农业大学热带作物学院，云南普洱　665099, bio={"content":"

王晓琳（2002—），女，本科生，研究方向：林木生物技术。

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王晓琳（2002—），女，本科生，研究方向：林木生物技术。

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Frontiers in Plant Science, 2019, 10: 1573., articleTitle=Effect of single and combined expression of lysophosphatidic acid acyltransferase, glycerol-3-phosphate acyltransferase, and diacylglycerol acyltransferase on lipid accumulation and composition in Neochloris oleoabundans, refAbstract=null)], funds=[Fund(id=1276204221888332654, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276204182059221793, awardId=322QN407, language=CN, fundingSource=海南省自然科学基金青年项目(322QN407), fundOrder=null, country=null), Fund(id=1276204222286791535, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276204182059221793, awardId=202305AF150125, language=CN, fundingSource=云南省黄华孙专家工作站(202305AF150125), fundOrder=null, country=null)], companyList=[AuthorCompany(id=1276204194658911025, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276204182059221793, xref=1., ext=[AuthorCompanyExt(id=1276204194679882546, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276204182059221793, companyId=1276204194658911025, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=1.Rubber Research Institute, Chinese Academy of Tropical Agricultural Sciences, Haikou, Hainan 571101, China), AuthorCompanyExt(id=1276204195007038259, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276204182059221793, companyId=1276204194658911025, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=1.中国热带农业科学院橡胶研究所，海南海口　571101)]), AuthorCompany(id=1276204195422274356, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276204182059221793, xref=2., ext=[AuthorCompanyExt(id=1276204195443245877, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276204182059221793, companyId=1276204195422274356, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=2.College of Tropical Crops, Yunnan Agricultural University, Pu'er, Yunnan 665099, China), AuthorCompanyExt(id=1276204195468411702, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276204182059221793, companyId=1276204195422274356, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=2.云南农业大学热带作物学院，云南普洱　665099)])], figs=[ArticleFig(id=1276204213923349340, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276204182059221793, language=EN, label=Fig. 1, caption=Phylogenetic analysis of HbLPAT and AtGPAT gene family in H. brasiliensis and A. thaliana, figureFileSmall=c157DN543sZhLER57n2GlA==, figureFileBig=XxODi8plXsJekm3/U2UmDQ==, tableContent=null), ArticleFig(id=1276204214707684189, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276204182059221793, language=CN, label=图1, caption=橡胶树HbLPAT和拟南芥AtGPAT基因家族系统进化分析, figureFileSmall=c157DN543sZhLER57n2GlA==, figureFileBig=XxODi8plXsJekm3/U2UmDQ==, tableContent=null), ArticleFig(id=1276204216402183006, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276204182059221793, language=EN, label=Fig. 2, caption=Relative expression levels of different HbLPAT in latex of H. brasiliensis, figureFileSmall=Fn4QaExWEzbTxw0gBatJwg==, figureFileBig=SwsR97W8pBrGqSKFGhreLg==, tableContent=null), ArticleFig(id=1276204216477680479, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276204182059221793, language=CN, label=图2, caption=橡胶树胶乳中不同HbLPATs的相对表达量

不同小写字母表示差异显著（P<0.05）。

, figureFileSmall=Fn4QaExWEzbTxw0gBatJwg==, figureFileBig=SwsR97W8pBrGqSKFGhreLg==, tableContent=null), ArticleFig(id=1276204216863556448, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276204182059221793, language=EN, label=Fig. 3, caption=Multiple alignment analysis of HbLPAT2 in H. brasiliensis, figureFileSmall=xkkItwiSaFqq8KcyvYD6BQ==, figureFileBig=nE9Z03OILkrU0AQzaMn8Zg==, tableContent=null), ArticleFig(id=1276204217253626721, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276204182059221793, language=CN, label=图3, caption=橡胶树HbLPAT2蛋白序列的对比分析

A：HbLPAT2a的功能结构域；B：HbLPAT2b的功能结构域；C：LPAT2蛋白序列比对，红色虚线框代表保守基序。

, figureFileSmall=xkkItwiSaFqq8KcyvYD6BQ==, figureFileBig=nE9Z03OILkrU0AQzaMn8Zg==, tableContent=null), ArticleFig(id=1276204217329124194, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276204182059221793, language=EN, label=Fig. 4, caption=Relative expression levels of HbLPAT2 genes in different tissues of H. brasiliensis, figureFileSmall=FeZ5h+yNeJo3Cgi+KBXFFA==, figureFileBig=A6ItG3Oy6TDADWR1OMoFFw==, tableContent=null), ArticleFig(id=1276204217702417251, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276204182059221793, language=CN, label=图4, caption=HbLPAT2基因在橡胶树不同组织中的相对表达量

不同小写字母表示差异显著（P<0.05）。

, figureFileSmall=FeZ5h+yNeJo3Cgi+KBXFFA==, figureFileBig=A6ItG3Oy6TDADWR1OMoFFw==, tableContent=null), ArticleFig(id=1276204218088293220, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276204182059221793, language=EN, label=Fig. 5, caption=Subcellular localization of HbLPAT2a and HbLPAT2b, figureFileSmall=CsnrcsCIOATieKQBPthckw==, figureFileBig=d5waa9vZrvCJqKheYimSTg==, tableContent=null), ArticleFig(id=1276204218167984997, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276204182059221793, language=CN, label=图5, caption=HbLPAT2a和HbLPAT2b基因的亚细胞定位

蓝色：内质网标记CFP-HDEL；绿色：HbLPAT2a-GFP和HbLPAT2b-GFP；白色箭头：HbLPAT2a-GFP内质网共定位处和HbLPAT2b-GFP内质网共定位处。

, figureFileSmall=CsnrcsCIOATieKQBPthckw==, figureFileBig=d5waa9vZrvCJqKheYimSTg==, tableContent=null), ArticleFig(id=1276204218545472358, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276204182059221793, language=EN, label=Tab. 1, caption=

Primers in this study

, figureFileSmall=null, figureFileBig=null, tableContent=
引物名称Primer name引物序列（5ʹ-3ʹ）Primer sequence (5ʹ-3ʹ)用途Usage
HbLPAT2a-FATGGCGATTCCTGCGGCAGT基因
HbLPAT2a-RTTAGTGTTGTTTGTCTTGTCCTGTTTC克隆
HbLPAT2b-FATGGCGCTTCCTGCGGCTGT
HbLPAT2b-RTAATAGAACTGCAAAATCACAAGAGAC
HbLPAT2a-GFagtagatctggtaccgcattcATGGCGATTCCTGCGGCA亚细胞
定位
HbLPAT2a-GRgcccttgctcaccattGTGTTGTTTGTCTTGTCCTGTTTC分析
HbLPAT2b-GFagtagatctggtacctgggggATGGCGCTTCCTGCGGC
HbLPAT2b-GRgcccttgctcaccatTAGAACTGCAAAATCACAAGAGAC
HbLPAT1-QFCATCTACTTCATTGCGGTTCTATG荧光定量检测
HbLPAT1-QRAAGAGCCATTCAAAGCAGAGTT
HbLPAT2a-QFGGACTTCTCTTCTTCTCTTCTGG
HbLPAT2a-QRGCTCTATTGATCGTTCTGTATGTG
HbLPAT2b-QFTGTTGTGGCTGGATCTTGTATG
HbLPAT2b-QRCAATGTCACTTCTGTGGTTGGA
HbLPAT3-QFGCTTGTCTGAAGATTGAAGTGTATG
HbLPAT3-QRAATGTCGCTCCTGTGATTGC
HbLPAT4-QFGTCTGATATGTCTGCTGGTGTT
HbLPAT4-QRTGGCAAGCCGTGTCATTAAG
HbLPAT5-QFACTAAGGATGATAAGAGGTGTTGTG
HbLPAT5-QRCCTGCTCCAATGTATGCTGAA
HbActin-FGATGTGGATATCAGGAAGGA
HbActin-RCATACTGCTTGGAGCAAGA
), ArticleFig(id=1276204220214805351, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276204182059221793, 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（5ʹ-3ʹ）Primer sequence (5ʹ-3ʹ)用途Usage
HbLPAT2a-FATGGCGATTCCTGCGGCAGT基因
HbLPAT2a-RTTAGTGTTGTTTGTCTTGTCCTGTTTC克隆
HbLPAT2b-FATGGCGCTTCCTGCGGCTGT
HbLPAT2b-RTAATAGAACTGCAAAATCACAAGAGAC
HbLPAT2a-GFagtagatctggtaccgcattcATGGCGATTCCTGCGGCA亚细胞
定位
HbLPAT2a-GRgcccttgctcaccattGTGTTGTTTGTCTTGTCCTGTTTC分析
HbLPAT2b-GFagtagatctggtacctgggggATGGCGCTTCCTGCGGC
HbLPAT2b-GRgcccttgctcaccatTAGAACTGCAAAATCACAAGAGAC
HbLPAT1-QFCATCTACTTCATTGCGGTTCTATG荧光定量检测
HbLPAT1-QRAAGAGCCATTCAAAGCAGAGTT
HbLPAT2a-QFGGACTTCTCTTCTTCTCTTCTGG
HbLPAT2a-QRGCTCTATTGATCGTTCTGTATGTG
HbLPAT2b-QFTGTTGTGGCTGGATCTTGTATG
HbLPAT2b-QRCAATGTCACTTCTGTGGTTGGA
HbLPAT3-QFGCTTGTCTGAAGATTGAAGTGTATG
HbLPAT3-QRAATGTCGCTCCTGTGATTGC
HbLPAT4-QFGTCTGATATGTCTGCTGGTGTT
HbLPAT4-QRTGGCAAGCCGTGTCATTAAG
HbLPAT5-QFACTAAGGATGATAAGAGGTGTTGTG
HbLPAT5-QRCCTGCTCCAATGTATGCTGAA
HbActin-FGATGTGGATATCAGGAAGGA
HbActin-RCATACTGCTTGGAGCAAGA
), ArticleFig(id=1276204220659401576, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276204182059221793, language=EN, label=Tab. 2, caption=

Bioinformatics softwares in this study

, figureFileSmall=null, figureFileBig=null, tableContent=
软件Soft网址Website功能Function
ProtParamhttps://web.expasy.org/protparam/蛋白质理化特性分析
Plant-mPLochttp://www.csbio.sjtu.edu.cn/bioinf/plant-multi/亚细胞定位预测
TMHMM 2.0https://services.healthtech.dtu.dk/services/TMHMM-2.0/蛋白质跨膜结构预测
SignalP - 5.0https://services.healthtech.dtu.dk/services/SignalP-5.0//蛋白质信号肽预测
NerPhos 3.1https://services.healthtech.dtu.dk/services/NetPhos-3.1/蛋白质磷酸化位点分析
SOPMAhttps://npsa-pbil.ibcp.fr/cgi-bin/npsa_automat.pl?page=npsa_sopma.html蛋白质二级结构预测
CDShttps://www.ncbi.nlm.nih.gov/Structure/cdd/wrpsb.cgi蛋白质功能结构域分析
), ArticleFig(id=1276204221024306026, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276204182059221793, language=CN, label=表2, caption=

本研究所用生物信息学软件

, figureFileSmall=null, figureFileBig=null, tableContent=
软件Soft网址Website功能Function
ProtParamhttps://web.expasy.org/protparam/蛋白质理化特性分析
Plant-mPLochttp://www.csbio.sjtu.edu.cn/bioinf/plant-multi/亚细胞定位预测
TMHMM 2.0https://services.healthtech.dtu.dk/services/TMHMM-2.0/蛋白质跨膜结构预测
SignalP - 5.0https://services.healthtech.dtu.dk/services/SignalP-5.0//蛋白质信号肽预测
NerPhos 3.1https://services.healthtech.dtu.dk/services/NetPhos-3.1/蛋白质磷酸化位点分析
SOPMAhttps://npsa-pbil.ibcp.fr/cgi-bin/npsa_automat.pl?page=npsa_sopma.html蛋白质二级结构预测
CDShttps://www.ncbi.nlm.nih.gov/Structure/cdd/wrpsb.cgi蛋白质功能结构域分析
), ArticleFig(id=1276204221120775020, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276204182059221793, language=EN, label=Tab. 3, caption=

Bioinformatics analysis of HbLPATs in H. brasiliensis

, figureFileSmall=null, figureFileBig=null, tableContent=
项目ItemHbLPAT2aHbLPAT2b
分子量/kDa43.2335.41
分子式C1997H3155N515O529S12C1637H2555N423O433S10
原子数62085058
等电点（pI）9.535.02
负电荷残基（Asp+Glu）3129
正电荷残基（Arg+Lys）4433
脂肪系数109.38109.68
不稳定系数41.2039.73
亲水性0.2310.201
亚细胞定位Endoplasmic reticulumEndoplasmic reticulum
跨膜结构域31
信号肽00
磷酸化位点S: 20; T: 11; Y: 2S: 16; T: 9; Y: 1
α-螺旋/%44.8247.44
延伸链/%17.3617.95
β-转角/%3.635.13
无规则卷曲/%34.2029.49
), ArticleFig(id=1276204221489873773, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276204182059221793, language=CN, label=表3, caption=

橡胶树HbLPATs生物信息学分析

, figureFileSmall=null, figureFileBig=null, tableContent=
项目ItemHbLPAT2aHbLPAT2b
分子量/kDa43.2335.41
分子式C1997H3155N515O529S12C1637H2555N423O433S10
原子数62085058
等电点（pI）9.535.02
负电荷残基（Asp+Glu）3129
正电荷残基（Arg+Lys）4433
脂肪系数109.38109.68
不稳定系数41.2039.73
亲水性0.2310.201
亚细胞定位Endoplasmic reticulumEndoplasmic reticulum
跨膜结构域31
信号肽00
磷酸化位点S: 20; T: 11; Y: 2S: 16; T: 9; Y: 1
α-螺旋/%44.8247.44
延伸链/%17.3617.95
β-转角/%3.635.13
无规则卷曲/%34.2029.49
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橡胶树HbLPAT2基因的克隆与表达分析
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王晓琳 1, 2 , 曲鹏 2 , 甘明洋 1, 2 , 仇键 1 , 范睿深 1, *
热带作物学报 | 组学与生物技术 2024,45(12): 2515-2523
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热带作物学报 |组学与生物技术 2024 , 45 (12) : 2515 -2523
橡胶树HbLPAT2基因的克隆与表达分析
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王晓琳（2002—），女，本科生，研究方向：林木生物技术。

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王晓琳1, 2, 曲鹏2, 甘明洋1, 2, 仇键1, 范睿深1, *
作者信息
  • 1.中国热带农业科学院橡胶研究所，海南海口　571101
  • 2.云南农业大学热带作物学院，云南普洱　665099
通讯作者:
* 范睿深（FAN Ruishen），E-mail：。
Cloning and Expression Analysis of HbLPAT2 Gene from Hevea brasiliensis
Xiaolin WANG1, 2, Peng QU2, Mingyang GAN1, 2, Jian QIU1, Ruishen FAN1, *
Affiliations
  • 1.Rubber Research Institute, Chinese Academy of Tropical Agricultural Sciences, Haikou, Hainan 571101, China
  • 2.College of Tropical Crops, Yunnan Agricultural University, Pu'er, Yunnan 665099, China
出版时间: 2024-12-25 doi: 10.3969/j.issn.1000-2561.2024.12.003
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摘要
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溶血磷酸酰基转移酶（lysophosphatidyl acyltransferase，LPAT）是控制溶血磷脂酸生成磷脂酸的关键酶，在植物磷脂合成中发挥重要作用。橡胶树（Hevea brasiliensis）是生产天然橡胶的重要来源，其在国防、医疗、高端制品等领域具有不可替代的作用，具有很高的社会价值和经济价值。本研究采用分子生物学技术在橡胶树胶乳中成功克隆出2个HbLPAT基因家族中负责磷脂合成的关键基因，分别命名为HbLPAT2a和HbLPAT2b，其编码序列全长分别为1161 bp和939 bp，分别编码了386个和312个氨基酸。HbLPAT2a和HbLPAT2b均含有1个PLN02380的溶血磷脂酸酰基转移酶功能结构域和高度保守的NH(X)4D基序。HbLPAT2a有3个跨膜结构域，HbLPAT2b有1个跨膜结构域，二者均无信号肽，为亲水性蛋白，且含有多个磷酸化位点。亚细胞定位分析表明，HbLPAT2a和HbLPAT2b蛋白均定位于内质网上。不同组织部位表达分析表明，HbLPAT2a和HbLPAT2b基因均在橡胶树种子中的表达量最高，推测其具有脂质合成功能，而在胶乳中，HbLPAT2a和HbLPAT2b的表达量在HbLPAT基因家族中位居前2位。本研究结果有助于了解HbLPAT2a和HbLPAT2b在橡胶树胶乳磷脂合成过程中的功能，为后期橡胶树胶乳品质的遗传改良提供科学依据。

橡胶树  /  HbLPAT2  /  生物信息学分析  /  表达分析

Lysophosphatidyl acyltransferase (LPAT), a key enzyme which controls the production of lysophosphatidic into phosphatidic, plays an important role in phospholipid biosynthesis in plants. Hevea brasiliensis is an important source for the production of natural rubber, which has an irreplaceable role in the fields of national defense, medical treatment, high-end products, and has high social and economic value. Cloning and analysis of HbLPAT2 in H. brasiliensis would provide scientific basis for further exploration of its phospholipid biosynthesis function and the effect of phospholipid on nature rubber. In this study HbLPAT2 was cloned from the latex of the H. brasiliensis by molecular biology techniques, and the bioinformatics analysis, expression analysis and subcellular localization analysis were performed. In this study, two key genes responsible for phospholipid biosynthesis in the HbLPAT gene family were successfully cloned from the latex of the H. brasiliensis, named HbLPAT2a and HbLPAT2b, the full length coding sequence of HbLPAT2a and HbLPAT2b genes was 1161 bp and 939 bp, encoding 386 and 312 amino acids. Both HbLPAT2a and HbLPAT2b contained a PLN02380 lysopyatidyl acyltransferase functional domain and contained a highly conserved NH(X)4D motif; HbLPAT2a protein had 3 transmembrane domains, and HbLPAT2b protein had 1 transmembrane domain, both of which had no signal peptide, were hydrophilic proteins, and contained multiple phosphorylation sites. The subcellular localization analysis showed that both HbLPAT2a and HbLPAT2b proteins were located in the endoplasmic reticulum. In different organs, the highest expression level of HbLPAT2a and HbLPAT2b was found in seeds, suggesting that they have a lipid biosynthesis function. However, in the latex of H. brasiliensis, the first and second expression level of whole HbLPAT gene family were HbLPAT2a and HbLPA2b. The result is helpful to understanding that HbLPAT2a and HbLPAT2b play an important role in the phospholipid biosynthetic pathway in latex of H. brasiliensis and to providing a scientific basis for genetic improvement of latex quality of H. brasiliensis.

Hevea brasiliensis  /  HbLPAT2  /  bioinformatics analysis  /  expression analysis
王晓琳, 曲鹏, 甘明洋, 仇键, 范睿深. 橡胶树HbLPAT2基因的克隆与表达分析. 热带作物学报, 2024 , 45 (12) : 2515 -2523 . DOI: 10.3969/j.issn.1000-2561.2024.12.003
Xiaolin WANG, Peng QU, Mingyang GAN, Jian QIU, Ruishen FAN. Cloning and Expression Analysis of HbLPAT2 Gene from Hevea brasiliensis[J]. Chinese Journal of Tropical Crops, 2024 , 45 (12) : 2515 -2523 . DOI: 10.3969/j.issn.1000-2561.2024.12.003
正文
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橡胶树（Hevea brasiliensisaa）属大戟科（Euphorbiaceae）三叶橡胶树属（Hevea）植物，迄今为止已有100多年的栽培历史[1]。橡胶树原产于巴西，于1904年引入我国云南，现主产区为海南和云南[2-3]。橡胶树是商用天然橡胶（nature rubber，NR）的主要来源，NR作为一种重要的工业材料，综合性能优异，具有不可替代的作用，广泛应用于航空航天、工农业、交通运输、医疗卫生等领域[4]。NR由橡胶树乳管细胞中排出的胶乳加工而来，胶乳的实质是乳管的细胞质，其主要成分为橡胶烃，并含有少量的脂质和蛋白质等非胶物质。NR中的脂质（主要是磷脂）和蛋白质作为支点与橡胶烃构成网状结构，其含量不仅直接影响NR的品质，也进一步影响其工艺操作性能与产品使用性能，是NR优于合成橡胶的根本原因[5-7]。目前，提高胶乳品质是我国橡胶树育种的重要方向之一。
脂质是一类高溶于非极性溶剂而低溶于水的有机生物分子，其化学本质是醇类和脂肪酸形成的酯类物质及其衍生物[8]。脂肪酸的酯化最终合成油脂是在内质网中通过Kennedy途径实现[9]，参与该途径的酶主要有3-磷酸甘油酰基转移酶（glycerol-3-phosphate acyltransferase，GPAT）、溶血磷酸酰基转移酶（lysophosphatidyl acyltransferase，LPAT）和二酰甘油酰基转移酶（diacylglycerol acyltransferase，DGAT）[10]，其中LPAT在磷脂合成中起到关键作用。甘油三酯合成的第一步反应是GPAT催化脂酰-CoA与3-磷酸甘油在sn-1位发生酯化反应生成溶血磷脂酸酶（lysophosphatidic acid，LPA）[11-12]，然后由LPAT在sn-2位发生酯化反应生成磷脂酸（phosphatidic acid，PA）[10]，再由磷脂酸磷酸化酶（phosphatidic acid phosphatase, PAP）脱去磷酸基团反应生成二酰甘油酯（diacylgycerol，DAG），最后DAG在sn-3发生催化脱酰基生成三酰基甘油酯（triacylglycerol，TAG）[12]。
LPAT是控制溶血磷脂酸生成磷脂酸的关键酶[13]，根据LPAT基因家族的亚细胞定位及进化起源，可将该基因家族分为2类：质体型LPATs和微粒体型LPATs[14]。质体型LPATs对软脂酸底物有强烈的偏好性，具有典型的原核酶类特征[10,15]。微粒体型LPATs种类较多，主要参与真核途径的膜脂及油脂合成[16]。前人在拟南芥中已对LPAT基因家族进行了较为系统的研究，AtLPAT基因家族成员有5个，其中，AtLPAT1定位在叶绿体，属于质体型；AtLPAT2定位于内质网中，属于微粒体型；AtLPAT3亚细胞定位目前不清楚，但是该基因对花可以特异性表达，其功能暂时不清楚；AtLPAT4、AtLPAT5的亚细胞定位和功能目前均尚不明确[10]。已有研究表明，拟南芥（Arabidopsis thaliana）定位在内质网上的AtLPAT2参与了磷脂的从头合成途径，在拟南芥生长发育各个阶段起着重要的作用，而且它能通过刺激磷脂的合成而参与拟南芥对低磷胁迫的响应[17-19]；在花生（Arachis hypogaca）中，超表达AhLPAT2能显著提高种子的含油量[20]；在蓖麻（Ricinus communis）中，超表达RcLPAT2不仅能提高其总含油量，也能提高油脂（TAG）中的不饱和脂肪酸的含量[21]。
提高胶乳品质是当前我国橡胶树育种研究的重要方向之一，增加磷脂含量可提升天然橡胶的加工性能[2]，这使得通过脂质含量的改变来提高胶乳品质成为可能。目前有关橡胶树油脂合成路径的研究较少，脂质积累路径不清，严重影响橡胶树脂质和磷脂的遗传改良，成为提高胶乳品质的主要障碍。借助已知的油脂合成途径以及油料植物改良思路，本研究选择橡胶树磷脂积累和油脂合成路径上的关键基因HbLPAT2为研究对象，对其进行序列、结构和系统发育等生物信息学分析，并通过基因表达分析以及亚细胞定位验证了解橡胶树HbLPAT2的基因结构及分子特征，为进一步研究橡胶树HbLPAT2基因在胶乳磷脂合成过程中的功能提供理论依据，对橡胶树高品质胶乳品种的选育及遗传改良具有一定的参考价值。
1　材料与方法
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1.1　材料
供试材料为10年生的热研73397橡胶树组培苗植株，种植于中国热带农业科学院橡胶研究所儋州试验场三队（19°31′10″N，109°29′41″E）。橡胶树割制为S/2 d4（1/2树围、每4 d割1刀，不涂乙烯利）。于2023年8月连续采集3刀的胶乳样品，收集5~10 min流出的胶乳，割胶当天同时采集其他组织。胶乳用液氮速冻后，放于离心管，其他组织使用锡纸包好，立即放入液氮速冻后于–80 ℃冰箱保存，用于RNA提取。
1.2　方法
1.2.1　橡胶树RNA的提取及cDNA的合成
取50~100 mg组织样品，液氮磨碎后，按照RNA提取试剂盒（TranGen, EasyPure Plant RNA Kit）说明书提取总RNA，使用NanoDrop 2000（Thermo Scientific）检测RNA的纯度和浓度。取检测合格的RNA进行反转录，使用反转录试剂盒（Thermo Scientific, RevertAid First Strand cDNA Synthesis Ki）合成cDNA。
1.2.2　 HbLPAT2的克隆与鉴定
根据课题组已有的橡胶树胶基因组和胶乳转录组数据[22]，共获得6条注释为LPAT的Unigene序列，通过NCBI的Blast功能进行对比，找到HbLPAT2基因，分别命名为HbLPAT2a和HbLPAT2b。用Premier 6.0软件设计特异引物（表1），以稀释后的胶乳cDNA为模板，使用高保真酶（Vazyme，2×Phanta Flash Master Mix）进行扩增，扩增体系及PCR反应程序同说明书。PCR扩增产物利用1.5%琼脂糖凝胶电泳进行检测，使用胶回收试剂盒（Agarose Gel DNA Extraction Kit Ver.4.0，Takara）将正确条带进行胶回收，加A尾连接T载体后（TA-cloning Kit，TaKaRa）转换大肠杆菌DH5-α感受态细胞，涂板过夜培养后通过菌落PCR筛选阳性克隆，送至北京擎科生物科技有限公司广州分公司测序。
1.2.3　HbLPAT2a和HbLPAT2b的生物信息学分析
利用生物信息学分析相关软件（表2）对HbLPAT2a和HbLPAT2b进行生物信息学分析。
1.2.4　 HbLPAT2a和HbLPAT2b的表达分析
使用Premier 6.0软件设计qRT-PCR检测的引物（表1）。以HbActin为内参基因，按照荧光定量试剂盒（TB GREEN® Premix Ex TaqTM II. ROX plus，TaKaRa）说明书进行定量反应。实验结果采用2-ΔΔCt方法计算相对表达量，每组实验重复3次。
1.2.5　HbLPAT2a和HbLPAT2b的亚细胞定位验证
以pCAM-1300-GFP为表达载体，选择BamH I为单酶切位点，设计HbLPAT2a和HbLPAT2b的同源臂，使用同源重组克隆酶（ClonExpress Ultra One Step Cloning Kit V2，Vazyme）将HbLPAT2a和HbLPAT2b连接到表达载体上，构成HbLPAT2-GFP重组质粒。将重组质粒转入大肠杆菌，测序成功后（方法同1.2.2）使用质粒提取试剂盒（Plasmid Mini Kit I，Omega）提取质粒，将质粒转入农杆菌GV3101（p19）感受态，筛选阳性菌株培养。将HbLPAT2-GFP与内质网maker CFPHDEL按1∶1比例注射烟草表面，同时注射空载体作为对照，培养2~3 d。用烟草叶片制作临时切片，在共聚焦显微镜中观察，低倍镜下找到绿色荧光，20倍镜观察并拍照。
2　结果与分析
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2.1　HbLPAT基因家族的鉴定及表达分析
以橡胶树胶乳转录组数据为基础，将拟南芥LPAT基因家族蛋白序列与橡胶树蛋白序列进行对比，共鉴定出6个橡胶树LPAT基因家族成员。使用MEGA软件构建系统进化树，根据序列比对和聚类分析结果（图1）将6个LPAT基因家族成员分别命名为HbLPAT1、HbLPAT2a、HbLPAT2b、HbLPAT3、HbLPAT4、HbLPAT5。在橡胶树胶乳中对HbLPAT基因家族成员进行表达分析发现，有4个基因在胶乳中表达，其中HbLPAT2b相对表达量最高，显著高于其他3个基因；其次为HbLPAT2a，显著高于HbLPAT1和HbLPAT5；未检测到HbLPAT3和HbLPAT4（图2）。
2.2　 HbLPAT2的克隆及蛋白序列分析
以橡胶树胶乳cDNA为模板，进行PCR扩增，克隆出2个长度分别为2397 bp和1719 bp的基因片段，经胶回收和阳性克隆测序，获得1161 bp的HbLPAT2a和939 bp的HbLPAT2b的蛋白质编码序列（coding sequence, CDS）。CDD预测结果表明，HbLPAT2a和HbLPAT2b蛋白均具有1个完整的溶血磷脂酸酰基转移酶功能结构域（图3A，图3B）。目前LPAT2基因已在多种植物中被成功克隆并公布了序列，酰基转移酶功能在部分植物中也得到了验证，如拟南芥、耧斗菜（Aquilegia coerulea）、茶树（Camellia sinensis）、蜜柑（Citrus unshiu）、古柯（Erythroxylum novogranatense）、麻风树（Jatropha curcas）、白羽扇豆（Lupinus albus）、木薯（Manihot esculenta）、银白杨（Populus alba）、美洲黑杨（Populus deltoides）、大叶栎（Quercus lobata）、欧洲栓皮栎（Quercus suber）、时钟花（Turnera subulata）、油桐（Vernicia fordii）等。将上述植物的LPAT2与橡胶树LPAT2的蛋白序列比对发现，在HbLPAT2中含有5个LPAT2蛋白保守基序（图3C），分别是NH（X）4D（Motif I）、LPVIGW（Motif II）、VLALFVEGTRF（Motif III）、N（X）LIPRIKGFV（Motif IV）和MRSFVPA（X）4T（X）（Motif V）。Motif I的组氨酸（H）及天冬氨酸（D）残基是酰基转移酶催化活性的重要功能位点；Motif III中则含有EGT-Box，它不但是酰基转移酶的功能位点，还含有酰基转移酶作用底物溶血磷脂酸的结合位点。此外，不同物种间的LPAT2序列在C端部分的差异性略大于N端。
2.3　橡胶树HbLPAT2蛋白的生物信息学分析
蛋白质理化特性分析结果如表3所示，橡胶树HbLPAT2a和HbLPAT2b的相对分子质量分别为43.23 kDa和35.41 kDa，原子总数分别为6208和5058，理论等电位点分别为9.53和5.02；脂肪系数分别为109.38和109.68，不稳定系数分别为41.20和39.73，均大于39，推测其属于不稳定蛋白；亲水性系数分别为0.231和0.200，均属于亲水性蛋白。亚细胞定位、跨膜结构域和信号肽预测表明，HbLPAT2a和HbLPAT2b均定位于内质网，HbLPAT2a有3个跨膜结构域，HbLPAT2b有1个跨膜结构域，二者均无信号肽，属于非分泌型蛋白。HbLPAT2a具有33个潜在的磷酸化位点，HbLPAT2b具有26个潜在的磷酸化位点。二级结构域预测表明，HbLPAT2a和HbLPAT2b的α-螺旋（α-helix）分别为44.82%和47.44%，无规则卷曲（random coil）分别为34.20%和29.49%，是构成HbLPAT蛋白的主要结构元件，而延伸链（extended strand）和β-转角（β-turn）分散于整个蛋白中，分别占17.36%~17.95%和3.63%~5.13%。
2.4　橡胶树HbLPAT2s的表达分析
实时荧光定量分析结果表明，HbLPAT2a和HbLPAT2b在橡胶树的根、叶片、种子、花、树皮和胶乳组织部位中均有表达，二者在种子中的相对表达最高，显著高于其他组织；其次为花，也显著高于另外4个组织部位；在剩余4个组织中的表达量由高到低依次为胶乳、树皮、树根和树叶，差异不显著（图4）。
2.5　橡胶树HbLPAT2的亚细胞定位
为了进一步研究HbLPAT2a和HbLPAT2b的功能及细胞内的表达部位，本研究构建了HbLPAT2-GFP的融合蛋白，利用农杆菌侵染的方法在烟草叶表皮细胞共表达HbLPAT2-GFP和细胞内质网定位的CFP-HDEL。亚细胞定位分析结果表明，空载的GFP信号在细胞中分散表达，HbLPAT2a和HbLPAT2b的GFP信号呈点状分布并与内质网定位的CFP信号重合（图5），说明HbLPAT2a和HbLPAT2b定位于细胞的内质网上，与之前的蛋白亚细胞定位预测结果相同。
3　讨论
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天然橡胶具有弹性强、可塑性高、回弹力强、抗冲击力强以及散热性良好等诸多性能，可用于生产包括航空轮胎和医用手套在内的50多万种橡胶制品，对运输、医药和国防等行业至关重要。天然橡胶之所以具有高弹性是源于其具有较大的分子量和交互网络结构，双亲性的磷脂与橡胶链α端相连有利于天然橡胶形成分支结构。此外，磷脂作为天然橡胶中最重要的非胶物质之一还显著影响胶乳的产量和稳定性，并且对生胶的硫化特性和抗氧老化性能也有一定影响[2]。鉴定参与橡胶树磷脂生物合成的酰基转移酶，对于理解这些基因在胶乳代谢和胶乳品质调控中的功能具有重要的意义。随着分子生物学的发展，大量研究表明，LPAT参与调控植物磷脂合成积累过程，介导酰基-CoA连接到甘油骨架sn-2位上的酰化反应，催化LPA生成PA，是磷脂磷脂生物合成的一个限速酶。目前，LPAT基因已在多种植物中被克隆出来，并对其基因功能进行了初步验证，如紫苏（Perilla frutescens）、花生、水稻（Oryza sativa）等[23-25]，但在橡胶树中未见报道。
本研究克隆获得的2个橡胶树HbLPAT2基因，序列分析表明HbLPAT2共有5个保守的Motifs，具有高度保守性。其中Motif I（NHXXXXD）为膜结合的O-酰基转移酶（MBOAT）家族，其组氨酸（H）及天冬氨酸（D）是催化酰基转移酶活性的重要位点，Motif III（VLALFVEGTRF）上的EGTBox不但是酰基转移酶的功能位点，还是与酰基转移酶作用底物溶血磷脂酸的结合位点[26-27]，基因结构的高度保守意味着它们在植物磷脂生物合成过程中发挥着相似的功能。同时发现这2个蛋白均具有完整的PLN02380（1~374，1~312）溶血磷脂酸酰基转移酶功能结构域，推测HbLPAT2很可能参与橡胶树胶乳的磷脂生物合成和代谢调控。生物信息学分析发现HbLPAT2a和HbLPAT2b分别有3个跨膜结构域和1个跨膜结构域，均无信号肽，为亲水性蛋白，且含有多个磷酸化位点。HbLPAT2的跨膜结构域均位于N端，这些跨膜区有助于HbLPAT2锚定于膜上。磷酸位点作为修饰位点，可激活或抑制染色质构型，调控基因座的转录。HbLPAT2a和HbLPAT2b蛋白分别具有33个和26个潜在的磷酸化位点（数量最多的为丝氨酸），说明可逆磷酸化调控在实现HbLPAT2蛋白的功能上具有一定作用，在以后的分子育种中可以对它加以利用。如AtWRI1受蔗糖非发酵相关蛋白激酶（SNF1-related protein kinase）KIN10介导降解，其70位点的苏氨酸和166位点的丝氨酸发挥了重要作用，2个位点突变后能够增强AtWRI1蛋白的稳定性[28]。
亚细胞定位预测和验证均表明HbLPAT2a和HbLPAT2b定位于内质网上，这与大部分调控磷脂合成基因的定位相同。胶乳中HbLPAT家族成员表达量分析结果表明，HbLPAT2的表达量显著高于另外4个HbLPATs，在拟南芥的AtLPAT家族成员中，AtLPAT2定位于内质网且参与了磷脂的合成。在底物选择上更偏好18:1-CoA[10]，橡胶树胶乳中的脂肪酸主要为棕榈酸（C16:0）、硬脂酸（C18:0）、油酸（C18:1）和亚油酸（C18:2），这4个主要脂肪酸中3个为C18脂肪酸，这可能是橡胶树胶乳的HbLPATs中HbLPAT2表达量高的原因。HbLPAT2a和HbLPAT2b在橡胶树不同组织中的表达分析结果表明，二者均在种子中的相对表达最高，显著高于其他组织，其次为花。种子作为植物合成油脂的主要场所，绝大多数负责油脂合成相关的基因均在种子中高表达，花作为生殖器官，在开花过程中也需要大量脂质的合成[29]，这也解释了HbLPAT2为何会在橡胶树种子和花中的表达量显著高于其他组织。而在其他组织部位中，胶乳的HbLPAT2表达量最高，考虑到与其他组织部位相比胶乳中丰富的磷脂含量，再结合之前的研究结果，说明HbLPAT2是橡胶树胶乳中主要负责磷脂合成的基因，但是否为唯一负责磷脂合成的基因，还需要后续实验进行验证。
4　结论
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本研究通过分子生物学手段首次克隆出了橡胶树HbLPAT2a和HbLPAT2b两个基因并完成了生物信息学分析和表达分析，结果初步表明HbLPAT2a和HbLPAT2b参与了橡胶树胶乳磷脂的生物合成。后续将进一步探究HbLPAT2a和HbLPAT2b通过参与磷脂合成过程对胶乳品质的影响，为橡胶树胶乳品质的遗传改良提供科学基础和分子靶标。
基金
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  • 海南省自然科学基金青年项目(322QN407)
  • 云南省黄华孙专家工作站(202305AF150125)
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2024年第45卷第12期
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doi: 10.3969/j.issn.1000-2561.2024.12.003
  • 接收时间：2024-05-17
  • 首发时间：2026-06-23
  • 出版时间：2024-12-25
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  • 收稿日期：2024-05-17
  • 修回日期：2024-06-12
基金
海南省自然科学基金青年项目(322QN407)
云南省黄华孙专家工作站(202305AF150125)
作者信息
    1.中国热带农业科学院橡胶研究所，海南海口　571101
    2.云南农业大学热带作物学院，云南普洱　665099

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* 范睿深（FAN Ruishen），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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