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Article(id=1276175697936781542, tenantId=1146029695717560320, journalId=1235980609244409860, issueId=1276175380184695804, articleNumber=null, orderNo=null, doi=10.3969/j.issn.1000-2561.2024.09.003, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=null, receivedDate=1704816000000, receivedDateStr=2024-01-10, revisedDate=1705161600000, revisedDateStr=2024-01-14, acceptedDate=null, acceptedDateStr=null, onlineDate=1782193596585, onlineDateStr=2026-06-23, pubDate=1727193600000, pubDateStr=2024-09-25, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1782193596585, onlineIssueDateStr=2026-06-23, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1782193596585, creator=13701087609, updateTime=1782193596585, 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=1791, endPage=1800, ext={EN=ArticleExt(id=1276175699077632232, articleId=1276175697936781542, tenantId=1146029695717560320, journalId=1235980609244409860, language=EN, title=Screening of AcSWEET11 Interacting Proteins Using Yeast Twohybrid System in Pineapple (Ananas comosus L.), columnId=1236256430337085821, journalTitle=Chinese Journal of Tropical Crops, columnName=Omics & Biotechnology, runingTitle=null, highlight=null, articleAbstract=

SWEET (sugars will eventually be exported transporter) plays an important role in pant flowering. However, the mechanism of flower was unclear in pineapple. In this study, the interacting proteins of AcSWEET11 were identified, in order to lay a foundation for analysis of the flowering mechanism of pineapple. In this study, the interacting proteins of AcSWEET11 were screened from the pineapple flowering cDNA membrane library by co-transformed, and then the expression levels of the candidate proteins during flowering were analyzed. pBT3-STE-AcSWEET11+pPR3-N was nontoxicity for NMY51 yeast cells but had autonomous activation. Further study showed that autonomous activation was restrained after adding 3ʹAT in TDO and QDO. A total of 81 positive clones were screened by yeast two-hybrid, and 48 proteins were identified by sequencing which were probably interactive with AcSWEET11. The proteins were contained E3 ubiquitin-protein ligase RING1-like, Trehalose-phosphate synthase 7, Cytochrome P450 90A1-like and Transcription factor LUX. GO and KEGG revealed that 48 proteins may participate in lipid metabolism, amino acid metabolism, carbohydrate metabolism, signal transduction and transport and catabolism. And the expression of Trehalose-phosphate synthase 7 (XP_020105459.1), Protein TIFY 3-like (XP_020082835.1), 40S ribosomal protein S27 (XP_020092770.1) and Heterogeneous nuclear ribonucleoprotein 1-like (XP_020112516.1) were down regulated during flowering induction that consistent with AcSWEET1 while the expression of Dihydrolipoyl dehydrogenase 2 (XP_020113798.1), Putative lipid-transfer protein DIR1 (XP_020086640.1), Clathrin assembly protein At4g32285 (XP_020108161.1) were up regulated. AcSWEET11 might interact with the proteins such as Trehalose-phosphate synthase 7 to participate in pineapple flowering. The protein interaction network of AcSWEET11 was plentiful to provide a foundation for analysis the regulatory mechanism of AcSWEET11 in pineapple flowering.

, authors=null, authorsList=Wenqiu LIN, Shenghui LIU, Xiumei ZHANG, Hongna ZHANG, Yunhe LI, Qingsong WU, authorCompany=null, correspAuthors=Qingsong WU, 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=1276175703133524216, articleId=1276175697936781542, tenantId=1146029695717560320, journalId=1235980609244409860, language=CN, title=利用酵母双杂交筛选菠萝AcSWEET11的互作蛋白, columnId=1236256430517440904, journalTitle=热带作物学报, columnName=组学与生物技术, runingTitle=null, highlight=null, articleAbstract=

SWEET（sugars will eventually be exported transporter）基因在植物开花过程中具有重要的作用，但AcSWEET11在菠萝成花中的作用机制尚不清楚。通过鉴定成花过程中与AcSWEET11的互作蛋白，为菠萝成花机制的解析奠定基础。本研究利用共转化的方法在菠萝成花过程的cDNA膜文库中筛选AcSWEET11的互作蛋白，分析候选蛋白的表达量。结果表明，pBT3-STE-AcSWEET11+pPR3-N对NMY51酵母细胞无毒性，但有自激活活性。进一步研究结果显示，在TDO/3ʹAT培养基和QDO培养基上自激活受到抑制。利用该系统筛选到了81个阳性克隆，经测序鉴定出48个与AcSWEET11互作的候选蛋白，包括E3 ubiquitin-protein ligase RING1-like、Trehalose-phosphate synthase 7、Cytochrome P450、Transcription factor LUX等。GO和KEGG分析结果显示，48个蛋白主要分布在细胞进程、代谢过程、刺激反应和催化活性等生物过程，参与脂类代谢、氨基酸代谢和碳水化合物代谢、信号转导和运输与分解代谢等新陈代谢途径。Trehalose-phosphate synthase 7（XP_020105459.1）、Protein TIFY 3-like（XP_020082835.1）、40S ribosomal protein S27（XP_020092770.1）、Heterogeneous nuclear ribonucleoprotein 1-like（XP_020112516.1）等4个基因与AcSWEET11表达趋势一致，在菠萝成花过程中下调表达；Dihydrolipoyl dehydrogenase 2（XP_020113798.1）、Putative lipid-transfer protein DIR1（XP_020086640.1）、clathrin assembly protein At4g32285（XP_020108161.1）等3个基因在菠萝成花过程中上调表达。这些结果表明，AcSWEET11可能通过与Trehalose-phosphate synthase 7等蛋白发生互作，参与菠萝成花过程。本研究进一步丰富了AcSWEET11的蛋白互作网络，为AcSWEET11在菠萝成花中的调控机制的解析奠定基础。

, authors=

林文秋（1989—），女，硕士，助理研究员，研究方向：菠萝遗传育种与生物技术。

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* 吴青松（WU Qingsong），E-mail：。
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林文秋（1989—），女，硕士，助理研究员，研究方向：菠萝遗传育种与生物技术。

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林文秋（1989—），女，硕士，助理研究员，研究方向：菠萝遗传育种与生物技术。

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Journal of Advanced Research, 2022, 48: 17-31., articleTitle=The clock component OsLUX regulates rice heading through recruiting OsELF3-1 and OsELF4s to repress Hd1 and Ghd7, refAbstract=null)], funds=[Fund(id=1276465458480021755, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276175697936781542, awardId=321QN302, language=CN, fundingSource=海南省自然科学基金青年基金项目(321QN302), fundOrder=null, country=null), Fund(id=1276465458547130620, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276175697936781542, awardId=32001989, language=CN, fundingSource=国家自然科学基金青年基金项目(32001989), fundOrder=null, country=null)], companyList=[AuthorCompany(id=1276465448145256642, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276175697936781542, xref=null, ext=[AuthorCompanyExt(id=1276465448153645251, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276175697936781542, companyId=1276465448145256642, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=South Subtropical Crop Research Institute, Chinese Academy of Tropical Agricultural Sciences / Laboratory of Tropical Fruit Biology, Ministry of Agriculture and Rural Affairs / Key Laboratory of Hainan Province for Postharvest Physiology and Technology of Tropical Horticultural Products, Zhanjiang, Guangdong 524091, China), AuthorCompanyExt(id=1276465448166228164, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276175697936781542, companyId=1276465448145256642, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=中国热带农业科学院南亚热带作物研究所/农业农村部热带果树生物学重点实验室/海南省热带园艺产品采后生理与保鲜重点实验室，广东湛江　524091)])], figs=[ArticleFig(id=1276465456978460907, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276175697936781542, language=EN, label=Fig. 1, caption=Toxicity and autonomous activation of bait vector

A-C: The situations of pNubG-Fe65+pTSU2-APP yeast cells grows in DDO、TDO/3ʹAT、QDO culture, respectively; D-F: The situations of pPR3-N+pTSU2-APP yeast cells grows in DDO、TDO/3ʹAT、QDO culture, respectively; G-L: The situations of pBT3-STE- AcSWEE11 yeast cells grows in DDO, TDO, 5 mmol/L TDO/3ʹAT, 10 mmol/L TDO/3ʹAT, 15 mmol/L TDO/3ʹAT and QDO culture, respectively.

, figureFileSmall=fNf+CV48/gJcCHICL3XbKA==, figureFileBig=ISMh47bioS/04FYIJcVi6w==, tableContent=null), ArticleFig(id=1276465457053958380, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276175697936781542, language=CN, label=图1, caption=诱饵毒性及自激活检测

A~C：pNubG-Fe65+pTSU2-APP分别在DDO、TDO/3ʹAT、QDO平板的生长情况；D~F：pPR3-N+pTSU2-APP分别在DDO、TDO/3ʹAT、QDO平板的生长情况；G~L：pBT3-STE-AcSWEE11分别在DDO、TDO、5 mmol/L TDO/3ʹAT、10 mmol/L TDO/3ʹAT、15 mmol/L TDO/3ʹAT和QDO平板的生长情况。

, figureFileSmall=fNf+CV48/gJcCHICL3XbKA==, figureFileBig=ISMh47bioS/04FYIJcVi6w==, tableContent=null), ArticleFig(id=1276465457355948269, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276175697936781542, language=EN, label=Fig. 2, caption=Verification of system function for bait vector pBT3-STE-AcSWEET11, figureFileSmall=3hF79XhDhc49xYpnb/6P8Q==, figureFileBig=FLn/Tl6xfmzxc9dRh4Xd7Q==, tableContent=null), ArticleFig(id=1276465457431445742, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276175697936781542, language=CN, label=图2, caption=诱饵载体pBT3-STE-AcSWEET11的系统功能验证, figureFileSmall=3hF79XhDhc49xYpnb/6P8Q==, figureFileBig=FLn/Tl6xfmzxc9dRh4Xd7Q==, tableContent=null), ArticleFig(id=1276465457502748911, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276175697936781542, language=EN, label=Fig. 3, caption=Screened of interacting proteins

A: The situations of the first transformed yeast cells grows in TDO/X/3ʹAT culture; B: The situations of the second transformed yeast cells grows in TDO/X/3’AT culture; C: The situations of the first transformed yeast cells grows in QDO/X culture; D: The situation of the second transformed yeast cells grows in QDO/X culture.

, figureFileSmall=+wMLBBt1SZEn+/I+3Jswkw==, figureFileBig=MS+VpptqTa30+UxPWvib5g==, tableContent=null), ArticleFig(id=1276465457586634992, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276175697936781542, language=CN, label=图3, caption=互作蛋白的筛选

A：第一次转化的酵母细胞在TDO/X/3ʹAT培养基上生长的情况；B：第二次转化的酵母细胞在TDO/X/3ʹAT培养基上生长的情况；C：第一次转化的酵母细胞在QDO/X培养基上生长的情况；D：第二次转化的酵母细胞点种QDO/X培养基上生长的情况。

, figureFileSmall=+wMLBBt1SZEn+/I+3Jswkw==, figureFileBig=MS+VpptqTa30+UxPWvib5g==, tableContent=null), ArticleFig(id=1276465457662132465, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276175697936781542, language=EN, label=Fig. 4, caption=Result of PCR products from partially positive clonal colonies by agarose gel electrophoresis

M: DL 2000 DNA marker; 1-25: The PCR products of colonies.

, figureFileSmall=r0husNMN7FQqOR92WcO04Q==, figureFileBig=Oy+OVO2pRl+pw7+/RObIKA==, tableContent=null), ArticleFig(id=1276465457737629938, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276175697936781542, language=CN, label=图4, caption=部分阳性克隆菌落PCR产物琼脂糖凝胶电泳结果

M：DL 2000 DNA marker；1~25：菌落PCR产物。

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Screening of proteins interacting with AcSWEET11 by yeast two-hybrid system

, figureFileSmall=null, figureFileBig=null, tableContent=
编号No.基因注释Gene annotationGenBank登录号GenBank accession number
1Tubby-like F-box proteinXP_020088583.1
2F-box protein PP2-B11OAY76934.1
3Dolichyl-diphosphooligosaccharide-protein glycosyltransferase 48 kDa subunitXP_020096081.1
4Cytochrome P450 90A1-likeXP_020101190.1
5Cytochrome P450 98A1XP_020114040.1
6PLAT domain-containing protein 3-likeXP_020091149.1
7Dihydrolipoyl dehydrogenase 2XP_020113798.1
8Tubulin alpha chainXP_020095799.1
9Hypothetical protein ACMD2_18406OAY83140.1
10Protein SRG1OAY83593.1
11Putative lipid-transfer protein DIR1XP_020086640.1
12T-complex protein 1 subunit zetaXP_020100745.1
13Pentatricopeptide repeat-containing protein At3g62890XP_020086106.1
14Probable clathrin assembly protein At4g32285XP_020108161.1
15AdenosylhomocysteinaseXP_020106819.1
16Lysine-specific demethylase JMJ25XP_020112685.1
17Basic transcription factor 3-likeXP_020094731.1
18Protein TIFY 3-likeXP_020082835.1
19Vacuolar proton pump subunit B2XP_020088591.1
2040S ribosomal protein S27XP_020092770.1
2160S ribosomal protein L18a-2XP_020079935.1
22Ribosomal protein L2YP_009116384.1
23N-alpha-acetyltransferase 16，NatA auxiliary subunitXP_020105507.1
24Heterogeneous nuclear ribonucleoprotein 1-likeXP_020112516.1
25Lipoxygenase 6XP_020110641.1
26DNA polymerase lambda glyoxysomal fatty acidXP_020083041.1
27beta-oxidation multifunctional protein MFP-a-like isoform X2XP_020103271.1
28Serine/threonine-protein kinase EDR1-like isoform X2XP_020098526.1
29Small nuclear ribonucleoprotein Sm D1OAY80757.1
30Polyubiquitin-likeXP_020091802.1
31E3 ubiquitin-protein ligase RING1-likeXP_020112621.1
32Serine/threonine-protein kinase STY46-likeXP_020115320.1
33Photosynthetic NDH subunit of lumenal location 4XP_020089715.1
34Hypothetical protein ACMD2_07758OAY73788.1
3560S acidic ribosomal protein P0XP_020109789.1
36Inhibitor of bruton tyrosine kinaseOAY68292.1
37Acyl-CoA-binding proteinXP_020113385.1
38Transcription factor LUXXP_020097739.1
39Probable alpha，alpha-trehalose-phosphate synthase 7XP_020105459.1
40Proteasome subunit beta type-5-likeXP_020096619.1
41Trithorax group protein osa-likeXP_020113943.1
42Uncharacterized protein LOC109720880XP_020103811.1
43Uncharacterized protein LOC109712697XP_020092020.1
44Uncharacterized protein LOC109707352 isoform X4XP_020084163.1
45Uncharacterized protein LOC109711902 isoform X3XP_020090847.1
46Uncharacterized protein LOC109711741 isoform X2XP_020090525.1
47Uncharacterized protein LOC109707959XP_020085097.1
48Uncharacterized protein LOC109718933 isoform X1XP_020101014.1
), ArticleFig(id=1276465458312249594, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276175697936781542, language=CN, label=表1, caption=

酵母双杂交系统筛选出的与AcSWEET11互作的蛋白

, figureFileSmall=null, figureFileBig=null, tableContent=
编号No.基因注释Gene annotationGenBank登录号GenBank accession number
1Tubby-like F-box proteinXP_020088583.1
2F-box protein PP2-B11OAY76934.1
3Dolichyl-diphosphooligosaccharide-protein glycosyltransferase 48 kDa subunitXP_020096081.1
4Cytochrome P450 90A1-likeXP_020101190.1
5Cytochrome P450 98A1XP_020114040.1
6PLAT domain-containing protein 3-likeXP_020091149.1
7Dihydrolipoyl dehydrogenase 2XP_020113798.1
8Tubulin alpha chainXP_020095799.1
9Hypothetical protein ACMD2_18406OAY83140.1
10Protein SRG1OAY83593.1
11Putative lipid-transfer protein DIR1XP_020086640.1
12T-complex protein 1 subunit zetaXP_020100745.1
13Pentatricopeptide repeat-containing protein At3g62890XP_020086106.1
14Probable clathrin assembly protein At4g32285XP_020108161.1
15AdenosylhomocysteinaseXP_020106819.1
16Lysine-specific demethylase JMJ25XP_020112685.1
17Basic transcription factor 3-likeXP_020094731.1
18Protein TIFY 3-likeXP_020082835.1
19Vacuolar proton pump subunit B2XP_020088591.1
2040S ribosomal protein S27XP_020092770.1
2160S ribosomal protein L18a-2XP_020079935.1
22Ribosomal protein L2YP_009116384.1
23N-alpha-acetyltransferase 16，NatA auxiliary subunitXP_020105507.1
24Heterogeneous nuclear ribonucleoprotein 1-likeXP_020112516.1
25Lipoxygenase 6XP_020110641.1
26DNA polymerase lambda glyoxysomal fatty acidXP_020083041.1
27beta-oxidation multifunctional protein MFP-a-like isoform X2XP_020103271.1
28Serine/threonine-protein kinase EDR1-like isoform X2XP_020098526.1
29Small nuclear ribonucleoprotein Sm D1OAY80757.1
30Polyubiquitin-likeXP_020091802.1
31E3 ubiquitin-protein ligase RING1-likeXP_020112621.1
32Serine/threonine-protein kinase STY46-likeXP_020115320.1
33Photosynthetic NDH subunit of lumenal location 4XP_020089715.1
34Hypothetical protein ACMD2_07758OAY73788.1
3560S acidic ribosomal protein P0XP_020109789.1
36Inhibitor of bruton tyrosine kinaseOAY68292.1
37Acyl-CoA-binding proteinXP_020113385.1
38Transcription factor LUXXP_020097739.1
39Probable alpha，alpha-trehalose-phosphate synthase 7XP_020105459.1
40Proteasome subunit beta type-5-likeXP_020096619.1
41Trithorax group protein osa-likeXP_020113943.1
42Uncharacterized protein LOC109720880XP_020103811.1
43Uncharacterized protein LOC109712697XP_020092020.1
44Uncharacterized protein LOC109707352 isoform X4XP_020084163.1
45Uncharacterized protein LOC109711902 isoform X3XP_020090847.1
46Uncharacterized protein LOC109711741 isoform X2XP_020090525.1
47Uncharacterized protein LOC109707959XP_020085097.1
48Uncharacterized protein LOC109718933 isoform X1XP_020101014.1
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利用酵母双杂交筛选菠萝AcSWEET11的互作蛋白
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林文秋 , 刘胜辉 , 张秀梅 , 张红娜 , 李运合 , 吴青松 *
热带作物学报 | 组学与生物技术 2024,45(9): 1791-1800
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热带作物学报 |组学与生物技术 2024 , 45 (9) : 1791 -1800
利用酵母双杂交筛选菠萝AcSWEET11的互作蛋白
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[Author(id=1276465448233337030, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276175697936781542, orderNo=0, firstName=null, middleName=null, lastName=null, nameCn=null, orcid=null, stid=null, country=null, authorPic=null, dead=0, email=null, emailSecond=null, emailThird=null, correspondingAuthor=0, authorType=1, ext={EN=AuthorExt(id=1276465448325611720, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276175697936781542, authorId=1276465448233337030, language=EN, stringName=Wenqiu LIN, firstName=Wenqiu, middleName=null, lastName=LIN, prefix=null, suffix=null, authorComment=null, nameInitials=null, affiliation=null, department=null, xref=null, address=South Subtropical Crop Research Institute, Chinese Academy of Tropical Agricultural Sciences / Laboratory of Tropical Fruit Biology, Ministry of Agriculture and Rural Affairs / Key Laboratory of Hainan Province for Postharvest Physiology and Technology of Tropical Horticultural Products, Zhanjiang, Guangdong 524091, China, bio=null, bioImg=null, bioContent=null, aboutCorrespAuthor=null), CN=AuthorExt(id=1276465448568881353, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276175697936781542, authorId=1276465448233337030, language=CN, stringName=林文秋, firstName=null, middleName=null, lastName=null, prefix=null, suffix=null, authorComment=null, nameInitials=null, affiliation=null, department=null, xref=null, address=中国热带农业科学院南亚热带作物研究所/农业农村部热带果树生物学重点实验室/海南省热带园艺产品采后生理与保鲜重点实验室，广东湛江　524091, bio={"content":"

林文秋（1989—），女，硕士，助理研究员，研究方向：菠萝遗传育种与生物技术。

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林文秋（1989—），女，硕士，助理研究员，研究方向：菠萝遗传育种与生物技术。

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林文秋, 刘胜辉, 张秀梅, 张红娜, 李运合, 吴青松*
作者信息
  • 中国热带农业科学院南亚热带作物研究所/农业农村部热带果树生物学重点实验室/海南省热带园艺产品采后生理与保鲜重点实验室，广东湛江　524091
通讯作者:
* 吴青松（WU Qingsong），E-mail：。
Screening of AcSWEET11 Interacting Proteins Using Yeast Twohybrid System in Pineapple (Ananas comosus L.)
Wenqiu LIN, Shenghui LIU, Xiumei ZHANG, Hongna ZHANG, Yunhe LI, Qingsong WU*
Affiliations
  • South Subtropical Crop Research Institute, Chinese Academy of Tropical Agricultural Sciences / Laboratory of Tropical Fruit Biology, Ministry of Agriculture and Rural Affairs / Key Laboratory of Hainan Province for Postharvest Physiology and Technology of Tropical Horticultural Products, Zhanjiang, Guangdong 524091, China
出版时间: 2024-09-25 doi: 10.3969/j.issn.1000-2561.2024.09.003
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SWEET（sugars will eventually be exported transporter）基因在植物开花过程中具有重要的作用，但AcSWEET11在菠萝成花中的作用机制尚不清楚。通过鉴定成花过程中与AcSWEET11的互作蛋白，为菠萝成花机制的解析奠定基础。本研究利用共转化的方法在菠萝成花过程的cDNA膜文库中筛选AcSWEET11的互作蛋白，分析候选蛋白的表达量。结果表明，pBT3-STE-AcSWEET11+pPR3-N对NMY51酵母细胞无毒性，但有自激活活性。进一步研究结果显示，在TDO/3ʹAT培养基和QDO培养基上自激活受到抑制。利用该系统筛选到了81个阳性克隆，经测序鉴定出48个与AcSWEET11互作的候选蛋白，包括E3 ubiquitin-protein ligase RING1-like、Trehalose-phosphate synthase 7、Cytochrome P450、Transcription factor LUX等。GO和KEGG分析结果显示，48个蛋白主要分布在细胞进程、代谢过程、刺激反应和催化活性等生物过程，参与脂类代谢、氨基酸代谢和碳水化合物代谢、信号转导和运输与分解代谢等新陈代谢途径。Trehalose-phosphate synthase 7（XP_020105459.1）、Protein TIFY 3-like（XP_020082835.1）、40S ribosomal protein S27（XP_020092770.1）、Heterogeneous nuclear ribonucleoprotein 1-like（XP_020112516.1）等4个基因与AcSWEET11表达趋势一致，在菠萝成花过程中下调表达；Dihydrolipoyl dehydrogenase 2（XP_020113798.1）、Putative lipid-transfer protein DIR1（XP_020086640.1）、clathrin assembly protein At4g32285（XP_020108161.1）等3个基因在菠萝成花过程中上调表达。这些结果表明，AcSWEET11可能通过与Trehalose-phosphate synthase 7等蛋白发生互作，参与菠萝成花过程。本研究进一步丰富了AcSWEET11的蛋白互作网络，为AcSWEET11在菠萝成花中的调控机制的解析奠定基础。

菠萝  /  AcSWEET11  /  酵母双杂交  /  互作蛋白

SWEET (sugars will eventually be exported transporter) plays an important role in pant flowering. However, the mechanism of flower was unclear in pineapple. In this study, the interacting proteins of AcSWEET11 were identified, in order to lay a foundation for analysis of the flowering mechanism of pineapple. In this study, the interacting proteins of AcSWEET11 were screened from the pineapple flowering cDNA membrane library by co-transformed, and then the expression levels of the candidate proteins during flowering were analyzed. pBT3-STE-AcSWEET11+pPR3-N was nontoxicity for NMY51 yeast cells but had autonomous activation. Further study showed that autonomous activation was restrained after adding 3ʹAT in TDO and QDO. A total of 81 positive clones were screened by yeast two-hybrid, and 48 proteins were identified by sequencing which were probably interactive with AcSWEET11. The proteins were contained E3 ubiquitin-protein ligase RING1-like, Trehalose-phosphate synthase 7, Cytochrome P450 90A1-like and Transcription factor LUX. GO and KEGG revealed that 48 proteins may participate in lipid metabolism, amino acid metabolism, carbohydrate metabolism, signal transduction and transport and catabolism. And the expression of Trehalose-phosphate synthase 7 (XP_020105459.1), Protein TIFY 3-like (XP_020082835.1), 40S ribosomal protein S27 (XP_020092770.1) and Heterogeneous nuclear ribonucleoprotein 1-like (XP_020112516.1) were down regulated during flowering induction that consistent with AcSWEET1 while the expression of Dihydrolipoyl dehydrogenase 2 (XP_020113798.1), Putative lipid-transfer protein DIR1 (XP_020086640.1), Clathrin assembly protein At4g32285 (XP_020108161.1) were up regulated. AcSWEET11 might interact with the proteins such as Trehalose-phosphate synthase 7 to participate in pineapple flowering. The protein interaction network of AcSWEET11 was plentiful to provide a foundation for analysis the regulatory mechanism of AcSWEET11 in pineapple flowering.

pineapple  /  AcSWEET11  /  yeast two-hybrid system  /  interacting proteins
林文秋, 刘胜辉, 张秀梅, 张红娜, 李运合, 吴青松. 利用酵母双杂交筛选菠萝AcSWEET11的互作蛋白. 热带作物学报, 2024 , 45 (9) : 1791 -1800 . DOI: 10.3969/j.issn.1000-2561.2024.09.003
Wenqiu LIN, Shenghui LIU, Xiumei ZHANG, Hongna ZHANG, Yunhe LI, Qingsong WU. Screening of AcSWEET11 Interacting Proteins Using Yeast Twohybrid System in Pineapple (Ananas comosus L.)[J]. Chinese Journal of Tropical Crops, 2024 , 45 (9) : 1791 -1800 . DOI: 10.3969/j.issn.1000-2561.2024.09.003
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菠萝[Ananas comosus（L.）Merr]是世界重要的热带果树之一，也是我国重要的热带经济作物。菠萝是聚花果植物，花芽分化一旦开始，花序和果实的发育就会持续下去，直到果实成熟[1]，成花与否决定了菠萝产量与品质的形成。目前，生产中菠萝的开花途径有2个，一是经过冬季低温后自然成花[2]，二是栽培12~16个月后经人工诱导成花。自然开花会导致果实成熟期不一致，收获期延长，甚至出现部分植株3~4 a才能自然开花的现象，限制了菠萝产业的发展[3]。为了解决这一问题，生产上主要利用乙烯利、电石等乙烯衍生物对菠萝进行人工诱导成花（催花）达到产期调节的效果[4]，但催花效果受到品种、环境条件和菠萝的生长发育时期的影响。如果催花不当，会造成减产、果实品质下降，甚至是绝收的情况，严重影响了经济效益，阻碍了产业的升级[5]。因此，解析菠萝的成花机制，对促进菠萝产期的调节具有重要意义。
SWEET广泛存在于植物体，在植物的许多生理过程中发挥着重要的作用。自2010年CHEN等[6]从拟南芥中第一次鉴定出糖转运蛋白基因SWEET以来，通过对已经发表的基因组序列进行全基因组检索分析，在多种植物中鉴定到了大量的SWEETs[7-9]。SWEET属于MtN3/saliva家族，膜内区域高度保守，具有双向转运糖的功能，包含7个跨膜a-螺旋结构，形成三螺旋束，促进糖的跨膜运输[10]。不同植物的SWEET系统进化分析结果表明，在植物中SWEET基因家族可以分为4个亚类，且每个亚类的主要转运底物不同：Ⅰ亚类主要转运脱氧葡萄糖，拟南芥的AtSWEET1是第一个以糖转运为特点的糖转运蛋白，具有葡萄糖的转运活性，参与花粉对单糖的吸收[11]；Ⅱ亚类主要转运单糖，葡萄VvSWEET4主要负责葡萄糖的转运，水稻OsSWEET5参与半乳糖转运[11-12]；Ⅲ亚类主要转运蔗糖，拟南芥AtSWEET11和AtSWEET12负责将细胞内的蔗糖流出到细胞壁间隙进而装载到韧皮部用于蔗糖的长距离运输[13]；而Ⅳ亚类主要介导了果糖的单向运输，AtSWEET17定位到液泡膜上，外部供应果糖时，其在根延伸区高度表达，促进根从外部吸收果糖，介导液泡对果糖的摄取并储存[14-15]。SWEETs调控植物花器官的发育，在植物开花过程中发挥着重要的调控作用。在水稻中OsSWEET11在圆锥花序和花药中高表达，遗传转化研究结果表明OsSWEET11调控小孢子在花粉母细胞阶段和未成熟花粉的发育。OsSWEET11通过降低花粉发育过程中淀粉的含量导致育性下降[7]。OsSWEET14基因敲除后，突变植株的种子变小，生长延缓，植株的生殖发育延迟[16]。一些研究结果表明SWEETs不仅能够影响植物的育性，在植物的花器官形成过程中也起着重要的作用。通过分析牡丹花花器官形成过程中葡萄糖含量变化和PsSWEET表达量变化发现，PsSWEET8极可能通过调控牡丹花瓣葡萄糖的转运从而调控开花过程[17]。最新的研究表明，在长日照条件下超表达AtSWEET10后拟南芥提前开花，进一步的研究结果表明AtSWEET10位于FT信号途径的下游[18]，这是首次有直接证据证明SWEET参与了植物成花转变的过程。但是AtSWEET10是如何调控开花的机制尚不清楚。因此，推测AtSWEET10可能在FT的诱导下进行转录然后运输糖或者FT蛋白来促进开花[18]。
已有研究表明，菠萝（无刺卡因）成花过程中可溶性糖和蔗糖含量显著提高[19]，且本课题组前期研究结果证实，AcSWEET11能够促进果实糖积累[20]，并能促进拟南芥提早开花（待发表），但其在成花中的作用机制尚不清楚。本研究利用酵母双杂交技术，筛选菠萝花芽分化过程的cDNA文库中与AcSWEET11互作的候选蛋白，为进一步研究AcSWEET11参与成花调控的机制奠定基础。
1　材料与方法
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1.1　材料
酵母粉（yeast extract）和胰蛋白胨（tryptone）购自OXOID公司；缺陷培养基（minimal synthetical defined medium，SD）DDO（SD/-Leu/-Trp）、TDO（SD/-Leu/-Trp/-His）、TDO/X（SD/-Leu/-Trp/-His/X-a-gal）、QDO（SD/-Leu/-Trp/-His/-Ade）和YPDA培养基购自北京索莱宝科技有限公司；X-a-gal购自YEASEN公司；酵母菌株感受态NMY51和pBT3、pNubG-Fe65、pTSU2-APP、pPR3等载体购自武汉金凯瑞生物工程有限公司；菠萝花芽的cDNA文库由本研究中心保存。
1.2　方法
1.2.1　诱饵载体的毒性和自激活检测
毒性和自激活检测方法参照王会勤[21]的方法。以pNubGFe65和pTSU2-APP共转化NMY51酵母菌为阳性对照，pPR3-N和pTSU2-APP共转化酵母菌为阴性对照。将pBT3-STE-AcSWEET11和猎物空载pPR3-N通过LiAc的方法共转化至酵母NMY51中，在DDO、TDO、TDO/AT和QDO培养基上30 ℃培养3~4 d，观察菌落的生长情况，检测pBT3-STE-AcSWEET11诱饵蛋白的毒性和自激活活性。通过对pBT3-STE-AcSWEET11和Post-Nubal共转化NMY51酵母菌，在DDO、TDO/3ʹAT和QDO培养基上培养，观察菌落的生长情况，检测pBT3-STE-AcSWEET11在酵母NMY51中的表达功能。
1.2.2　互作蛋白的筛选
从SD/-Leu平板挑取生长状态良好、含有pBT3-STE-AcSWEET11的单克隆接种至50 mL的SD/-Leu液体培养基，30 ℃培养至OD600为0.2后，转接至YPDA液体培养基中，震荡培养，使OD600为0.6~0.8（12~16 h）；室温离心，收集菌体，用无菌水重悬，离心后弃上清；加入LiAc重悬菌体，混匀，置冰上，制备含有诱饵载体的NMY51感受态。将菠萝cDNA文库质粒转化含有诱饵载体的NMY51感受态后，均匀涂布于TDO/X/3ʹAT 5 mmol/L平板上，30 ℃培养3~4 d。挑选TDO/X/3ʹAT板上变蓝的单菌落，转接于新的QDO/X平板上，30 ℃培养3 d，统计2种培养基上变蓝的单菌落，即为候选阳性克隆。
1.2.3　候选互作蛋白的测序鉴定
提取阳性克隆质粒，利用通用引物（Up: CTTTCCTTATACATTAGGACC，Dn: GGGACCTAGACTTCAGGTTG）进行PCR扩增。选取条带单一，大于500 bp的有效克隆片段，送至生工生物工程（上海）股份有限公司进行测序验证。测序结果在NCBI（https://www.ncbi.nlm.nih.gov）数据库上进行BLAST比对分析。
1.2.4　生物信息学分析
利用在线网站UniProt（https://www.uniprot.org）注释候选蛋白的功能。将获得的候选蛋白进行Gene Ontology（GO）和Kyoto Encyclopedia of Genes and Genomes（KEGG）通路分析。
1.2.5　候选蛋白基因的表达分析
对乙烯利诱导菠萝成花过程（0、8、16、32 d）进行转录组测序，获得花芽分化过程中基因表达量的数据，利用HemI软件计算各时期基因表达量的FPKM（fragments per kilobase of exon model per million mapped fragments）值，并绘制候选蛋白基因在乙烯利诱导菠萝成花过程中的表达热图。
2　结果与分析
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2.1　诱饵载体的毒性和自激活检测
以pNubG-Fe65和pTSU2-APP共转化NMY51酵母菌为阳性对照，pPR3-N和pTSU2-APP共转化酵母菌为阴性对照。将pBT3-STEAcSWEET11诱饵重组质粒和猎物空载pPR3-N共转化NMY51酵母感受态。结果表明，pBT3-STEAcSWEET11+pPR3-N在DDO培养基上的酵母菌生长状态良好，说明pBT3-STE-AcSWEET11和pPR3-N已成功转入NMY51酵母菌中，且对NMY51酵母细胞无毒性。pNubG-Fe65+pTSU2-APP在DDO、TDO/3ʹAT、QDO培养基上生长良好，pPR3-N+pTSU2-APP在DDO培养基上正常生长，在TDO/3ʹAT、QDO无菌落生长，诱饵重组质粒pBT3-STE-AcSWEET11+pPR3-N的酵母菌在TDO培养基能够生长，表明pBT3-STE- Ac-SWEET11+pPR3-N有自激活现象，激活HIS3的表达（图1A~图1H）。为了抑制酵母菌中HIS3的表达，在TDO培养基中添加不同浓度的3ʹAT。结果显示，在添加3ʹAT后酵母菌没有生长，HIS3的表达受到抑制（图1I~图1K）。进一步分析其在QDO培养基上的生长情况，结果显示，转化pBT3-STE-AcSWEET11+pPR3-N的酵母菌不能生长（图1L）。以上结果表明，pBT3-STE-AcSWEET11+pPR3-N在TDO/3ʹAT培养基和QDO培养基上自激活受到抑制，后续可在TDO/3ʹAT和QDO培养基上进行互作蛋白的筛选。
将Post-Nubal和pBT3-STE-AcSWEET11共转化NMY51酵母菌，分别在DDO、TDO/3ʹAT和QDO培养基上培养。结果表明，在DDO培养基上有菌落生长，说明共转化成功；在TDO/3ʹAT和QDO培养基上菌落生长良好，表明pBT3-STEAcSWEET11在NMY51系统能正确表达，可进行下一步文库筛选（图2）。
2.2　候选互作蛋白的筛选及鉴定
利用菠萝花芽分化的cDNA膜文库筛选与AcSWEET11互作的蛋白。以pTSU2-APP和pNubGFe65共转化菌液为阳性对照（CK+），pTSU2-APP和pPR3-N共转化菌液为阴性对照（CK–），经过2次筛选，共获得81个在TDO/X/3ʹAT培养基中生长良好的蓝色菌落（图3A、图3B）。进一步加压培养，将58个大小均一的单克隆点种在含有QDO/X培养基上进行再次筛选。结果显示，58个蓝色菌斑均能正常生长（图3C、图3D），初步筛选出58个阳性克隆。将这58个阳性克隆进行菌落PCR反应（pPR3-N为阳性对照，CK+；H2O为阴性对照，CK–），电泳检测结果表明，插入片段条带单一，大小约为500~2000 bp（图4）。分别提取阳性克隆质粒并进行测序分析，合并重复序列，测序结果在NCBI进行比对，筛选到48个可能与AcSWEET11相互作用的蛋白（表1）。主要包括E3泛素连接酶RING1-like、海藻糖磷酸合成酶、细胞色素P450、LUX转录因子等。
2.3　候选蛋白的生物信息学分析
对获得的48个候选互作蛋白进行GO和KEGG分类分析。GO分析结果表明，48个蛋白主要富集在细胞进程（cellular process）、代谢过程（metabolic process）、刺激反应（response tostimulus）、细胞解剖实体（cellular anatomical entity）、结合（binding）和催化活性（catalytic activity）等生物过程（图5）。KEGG分析结果显示，48个蛋白主要的KEGG途径包括脂类代谢（lipid metabolism）、氨基酸代谢（amino acid metabolism）、其他次生代谢物的生物合成（biosynthesis of other secondary metabolites）和碳水化合物代谢（carbohydrate metabolism）等新陈代谢途径，翻译（translation）和折叠、分类和降解（folding，sorting and degradation）等遗传信息途径，以及信号转导（signal transduction）与运输和分解代谢（transport and catabolism）等通路（图6）。
2.4　候选蛋白的基因表达分析
为了确定候选互作蛋白基因的表达情况，对前期获得的菠萝花芽分化过程中的转录组数据进行分析，结果表明，共有30个候选蛋白基因检测到表达量，18个候选蛋白基因在乙烯利诱导菠萝成花中未检测到。进一步分析结果表明，Trehalosephosphate synthase 7（XP_020105459.1）、Protein TIFY 3-like（XP_020082835.1）、40S ribosomal protein S27（XP_020092770.1）、Heterogeneous nuclear ribonucleoprotein 1-like（XP_020112516.1）等4个基因与AcSWEET11（XP_020107765.1）表达一致，在菠萝成花过程中下调表达；Dihydrolipoyl dehydrogenase 2（XP_020113798.1）、Putative lipid-transfer protein DIR1（XP_020086640.1）、Clathrin assembly protein At4g32285（XP_020108161.1）等3个基因在菠萝成花过程中上调表达（图7）。表明Trehalose-phosphate synthase 7等7个候选互作蛋白在菠萝成花过程中具有重要作用。
3　讨论
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酵母双杂交是一种筛选互作蛋白的常用方法，广泛应用于未知互作蛋白的筛选和已知蛋白间相互作用的验证。但该系统存在假阳性率和假阴性率高等局限性[22]。自激活和毒性是导致假阳性率和假阴性率高的主要因素。当诱饵蛋白和猎物蛋白单独或结合后，出现毒性，抑制酵母细胞的正常生长，从而出现假阴性；而当诱饵蛋白存在自激活活性，猎物蛋白与诱饵蛋白结合后可激活报告基因的表达，从而出现假阳性[23]。为了抑制诱饵蛋白的自激活活性，不同缺陷培养基以及在缺陷培养基中添加不同浓度3ʹAT被广泛应用于酵母双杂交的过程中。谢瑞莹等[23]通过在TDO培养基中添加10 mmol/L的3ʹAT，抑制了pGBKT7-NtMYB4a+pGADT7的自激活活性；在葡萄中，将pGBKT7-VvJAZ9+pGADT7在QDO/X/A的培养基上培养，其自激活活性受到抑制[24]。本研究结果显示，pBT3-STE-AcSWEET11+pPR3-N存在自激活现象，通过在TDO培养基中添加不同浓度的3ʹAT后或在QDO培养基上培养，酵母菌不能生长，表明3ʹAT和QDO培养基能够抑制pBT3-STE-AcSWEET11+pPR3-N的自激活活性。
菠萝花芽分化是一个复杂的生物学过程，受到自身因素和外界环境条件的影响，涉及糖、氨基酸、植物激素等生理生化的变化[19,25-26]。本课题组的前期研究结果表明，AcSWEET11可促进可溶性糖的积累[20]，但其在成花中的作用机制尚不清楚。本研究利用酵母双杂交技术，筛选了48个与AcSWEET11互作的蛋白，这些蛋白主要富集在氨基酸、碳水化合物以及信号转导和运输、分解等代谢通路上，推测AcSWEET11可能通过氨基酸、碳水化合物和信号转导等途径参与菠萝的成花过程。
拟南芥AtSWEET10能够促进拟南芥提早开花，且处于FT的下游[18]。在本研究中，没有筛选到FT等成花相关的基因，推测AcSWEET11可能不是通过直接与FT等成花相关基因相互作用参与菠萝的成花过程。Trehalose-phosphate synthase（TPS）基因在拟南芥成花启动中具有重要作用，其主要通过调控SPL等基因的表达参与拟南芥的成花过程，是糖作为信号物质参与成花的关键途径[27]。本研究结果表明，AcSWEET11和AcTPS7能够产生互作，且其在菠萝成花过程中的表达模式一致，推测AcSWEET11可能通过TPS信号途径参与菠萝成花。乙烯是诱导菠萝成花的关键激素，外源乙烯通过促进内源乙烯的合成诱导菠萝成花[4]。已有报道表明E3泛素连接酶（E3 ubiquitin-protein ligase RING1）能直接与ACS合成酶（1-aminocyclopropane-1-carboxylate synthase）相互作用来调节乙烯的合成[28]。本研究结果显示，AcSWEET11和E3泛素连接酶存在互作，推测AcSWEET11可能通过与E3泛素连接酶互作来调控乙烯的合成参与菠萝的成花过程。此外，本研究还筛选到了1个LUX转录因子。在水稻中，OsLUX通过调控OsELF3-1和OsELF4s蛋白抑制Hd1/Ghd7的表达参与成花[29]。而在菠萝中，AcSWEET11是否与水稻相似，能够通过与LUX相互作用，调控ELF等蛋白的表达来参与菠萝的成花过程，需要进一步研究。
4　结论
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本研究通过对pBT3-STE-AcSWEET11+pPR3-N的毒性和自激活活性的检测，建立AcSWEET11的酵母双杂交系统。利用该系统筛选了48个与AcSWEET11互作的候选蛋白。对48个蛋白进行GO和KEGG分析，其主要分布在细胞进程、代谢过程、刺激反应和催化活性等生物过程，参与氨基酸代谢、碳水化合物代谢和信号转导和运输等新陈代谢途径，推测AcSWEET11可能通过氨基酸代谢、碳水化合物代谢或者信号转导等途径参与菠萝的成花过程。
基金
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  • 海南省自然科学基金青年基金项目(321QN302)
  • 国家自然科学基金青年基金项目(32001989)
文献
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2024年第45卷第9期
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doi: 10.3969/j.issn.1000-2561.2024.09.003
  • 接收时间：2024-01-10
  • 首发时间：2026-06-23
  • 出版时间：2024-09-25
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  • 收稿日期：2024-01-10
  • 修回日期：2024-01-14
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海南省自然科学基金青年基金项目(321QN302)
国家自然科学基金青年基金项目(32001989)
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    中国热带农业科学院南亚热带作物研究所/农业农村部热带果树生物学重点实验室/海南省热带园艺产品采后生理与保鲜重点实验室，广东湛江　524091

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* 吴青松（WU Qingsong），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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