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生物工程学报 Chinese Journal of Biotechnology DOI: 10.13345/j.cjb.

1 生物工程学报 Chinese Journal of Biotechnology DOI: /j.cjb 郑世刚等 / 白藜芦醇合成酶基因在基因工程中的应用及功能研究进展 March 25, 2014, 30(3): Chin J Biotech, All rights reserved 341 综述郑世刚 1,2, 李臻 2, 赵善仓 3, 王庆国 2, 刘炜 ,2,,,.., 2014, 30(3): Zheng SG, Li Z, Zhao SC, et al. Advances of resveratrol synthase gene in the application of genetic engineering and biofunctional investigation. Chin J Biotech, 2014, 30(3): 摘要 : 白藜芦醇合成酶 (Resveratrol synthase, RS) 是白藜芦醇 (Resveratrol, Res) 合成途径中的关键酶以往研究报道,RS 基因已在多种植物和微生物中进行了转化和表达, 并在植物的代谢及调控等方面发挥生物学作用文中主要围绕 RS 基因对植物的转化, 及异源表达后植物体内代谢产物的变化, 转 RS 基因对植物抗病原菌活性抗自由基活性和生长发育的影响, 以及利用 RS 基因在微生物中生产 Res 的相关进展进行了综述并对 RS 基因在生物工程方面的应用前景进行展望 : 白藜芦醇合成酶基因, 白藜芦醇, 代谢, 生物功能, 应用前景 Advances of resveratrol synthase gene in the application of genetic engineering and biofunctional investigation Shigang Zheng 1,2, Zhen Li 2, Shancang Zhao 3, Qingguo Wang 2, and Wei Liu 1,2 1 College of Life Science, Qingdao Agricultural University, Qingdao , Shandong, China 2 Bio-Tech Research Center, Shandong Academy of Agricultural Sciences; Key Laboratory of Genetic Improvement, Ecology and Physiology of Crops, Shandong Province, Jinan , Shandong, China 3 Agricultural Quality Standards and Testing Technology Institute of Shandong Academy of Agricultural Sciences, Jinan , Shandong, China Abstract: Resveratrol synthase (RS) plays a key role in resveratrol (Res) biosynthesis. RS gene has been formerly Received: May 21, 2013; Accepted: July 2, 2013 Supported by: National Transform Science and Technology Program (No. 2009ZX B), Shandong Province Germplasm Innovation and Utilization Project ( ). Corresponding author: Wei Liu. Tel: ; wheiliu@163.com (No. 2009ZX B) ( )

2 342 ISSN CN /Q Chin J Biotech March, 2014 Vol.30 No.3 reported to be transformed into many plant species and microorganisms, and to play certain roles in metabolic and regulation processes. In this paper, the transformations of RS gene in plants, and the related changes of biological properties, such as metabolites, anti-pathogen activities, anti-radical properties, and developmental characters in transgenic plants, as well as the production of resveratrol in microbes by utilizing RS gene were summarized. Moreover, the application prospects of RS gene in bioengineering were also addressed. Keywords: resveratrol synthase gene, resveratrol, metabolism, biological properties, application prospects (Resveratrol, Res) (Ultraviolet, UV) [1] Res [2] Res Res Res (Resveratrol synthase, RS) Res [3] RS (Stilbene synthase, STS) 4- A A Res [3] RS Res Res RS Res Res 1 RS 基因在植物中的转化及相关代谢产物 1.1 RS 基因的克隆及转化 Res RS RS RS 1988 Schröder Arachis hypogaea 1.6 kb RS cdna (Chalcone synthase, CHS) 70% 74% CHS [4] Melchior Vitis vinifera L. RS PSV kb cdna 43 kda A A Res PSV25 RS [5] RS PSV25 RS Vst1 [6] StSy [7] RS AhRS [8] AhRS3 [9] bp RS PNRS1 RS 90% RS [10] Parthenocissus henryana Polygonum cuspidatum Sorghum bicolor PhSTS [11] PcRS [12] SbSTS1 [13] RS 1990 Hain RS AhRS

3 郑世刚等 / 白藜芦醇合成酶基因在基因工程中的应用及功能研究进展 343 Nicotiana tabacum L. Phytophthora megasperma AhRS mrna Res Res UV 8 h 600 ng/g Res UV 24 h 50 ng/g [8] RS Res Arabidopsis thaliana L. [12] Oryza sativa L. [14] Triticum aestivum L. [15] Medicago sativa [16] Actinidia deliciosa [17] Malus domestica Borkh. [18] Carica papaya L. [19] Populus alba L. [20] Lycopersicon esculentum Mill. [7] Brassica napus L. [21] Rehmannia glutinosa L. [9] Lactuca sativa L. [22] Pisum sativum L. [23] Humulus lupulus L. [24] Fragaria ananassa [25] Ipomoea batatas [26] Brassica campestris [27] Daucus carota [28] Res Res RS1 RS PNRS1 [29-30] PNRS1 UV-C Res 4 μg/ml Res 117 μg/g [29] 1.2 转 RS 基因植物中的相关代谢产物 RS RS Res (Piceid; trans-resveratrol-3-o-b-d-glucopyranoside) Res Nicoletti (RP-HPLC) (PDA) (MS) RS 4 ( Res) Res Res [7] Lo - (LC/MS/MS) RS Res Res Res [33] RS Hanhineva CaMV 35S fil1 RS NS-Vitis3 Res [25] Hammerbacher Picea abies RS PaSTS Res RS Res O- O- cjb@im.ac.cn

4 344 ISSN CN /Q Chin J Biotech March, 2014 Vol.30 No.3 表 1 转 RS 基因植物中 Res 及白藜芦醇糖苷的积累 Table 1 Accumulation of Res and piceid in RS transgenic plants RS gene Plant origin Promoter Host plant Metabolites Quantity (μg/g FW) Vst1 CaMV 35S Tobacco (Nicotiana tabacum L.) Res [31] Vst1 Seed-specific p-nap Oilseed rape (Brassica napus L.) Piceid 616 [21] Vst1 Res and CaMV 35S Hop (Humulus lupulus L.) Piceid 560 [24] Vst1 Vst1 Pea (Pisum sativum L.) Piceid [23] Vst1 Vst1 Papaya (Carica papaya L.) Piceid 54 [19] Vst1 Vst1 Apple (Malus domestica Borkh.) Piceid 3 7 [18] StSy CaMV 35S White poplar (Populus alba L.) Piceid [20] StSy Tomato (Lycopersicon Res and CaMV35S esculentum Mill.) Piceid 53 [7] PSV25 CaMV 35S Kiwi (Actinidia deliciosa) Piceid 182 [17] VpSTS Grapevine (Chinese wild Vitis CaMV 35S Res [32] pseudoreticulata) AhRS Groundnut (Arachis hypogaea) AhRS Tobacco (Nicotiana tabacum L.) Res 50 ng/g [8] AhRS Groundnut (Arachis hypogaea) CaMV 35S Alfalfa (Medicago sativa L.) Piceid [16] AhRS3 Groundnut (Arachis hypogaea) CaMV 35S Rehmannia glutinosa L. Piceid 116 [9] AhRS Groundnut (Arachis Purple sweet potato (Ipomoea CaMV 35S hypogaea) batatas) Piceid 340 (DW) [26] SbSTS1 Sorghum bicolor CaMV 35S Arabidopsis tt4 Piceid 580 [13] PhSTS Parthenocissus henryana CaMV 35S Lettuce (Lactuca sativa L.) Res 56.4 [22] PcRS Polygonum cuspidatum CaMV 35S Arabidopsis thaliana L. Piceid 136 [12] PNRS1 Groundnut (Arachis hypogaea) CaMV 35S Tobacco (Nicotiana tabacum L.) Res [29] [34] Res Res (Pterostilbene) Res Res 2011 Xu Res (Resveratrol O-methyltransferase, ROMT) ROMT RS VST1 [35] Rimando

5 郑世刚等 / 白藜芦醇合成酶基因在基因工程中的应用及功能研究进展 345 ROMT RS [36] 1 RS Res μg/g Res μg/g [18,23] μg/g [21,24] Res Res 34 μg/g [29] HPLC Res ( ) A A CHS RS ɑ- β- [18,24] Hüsken Vst1 UDP 4- Res [21] RS [7] 2 转 RS 基因植物的表型和功能 2.1 抗病原菌活性 Botrytis cinerea RS mrna Res Res [2] RS RS Hain RS RS mrna RS [8] Hain RS Vst1 RS [6] 2005 Serazetdinova RS Vst1 Vst2 Puccinia recondita f. sp. tritici Septoria nodorum Berk. [15] Zhu Vst1 RS mrna Phytophthora palmivora [19] Vst1 Vst1 mrna Pyricularia oryzae Vst1 mrna [37] Vst1 Xanthomonas oryzae [14] RS cjb@im.ac.cn

6 346 ISSN CN /Q Chin J Biotech March, 2014 Vol.30 No.3 [38] RS RS Hipskind CaMV 35S RS Phoma medicaginis [16] CaMV 35S 2005 Lim RS AhRS3 Fusarium oxysporum [9] 2011 Liu RS PcRS Colletotrichum higginsianum [12] Thévenot RS PR10 Vst1 Res Res [39] RS 2000 Kobayashi CaMV 35S RS PSV25 [17] 2004 Giorcelli CaMV 35S RS StSy Melampsora pulcherrima Res e- Res Res [20] 2009 Hanhineva RS NS-Vitis3 Res [25]

7 郑世刚等 / 白藜芦醇合成酶基因在基因工程中的应用及功能研究进展抗自由基活性 RS Res H 2 O 2 O 2 (ROS) 2005 Giovanna RS StSy Res Res [7] Morelli (EPR) RS Res [40] 2009 Annalisa TomLoxB RS StSy Res [41] RS [40] RS (Phorbol ester) Res [41] Ureňa 0.16 mmol/l Res (5 s) 75 d Res RS [42] [43] [44] Res RS UV 2010 Tang UV-C Res RS mrna UV-C Res UV-C SOD (GSR) (APX) (DHAR) Res 3,4- Res UV-C Res UV-C [45] RS Res RS Res RS [29] cjb@im.ac.cn

8 348 ISSN CN /Q Chin J Biotech March, 2014 Vol.30 No.3 [46] Res ROS RS RS PNRS1 13 [30] RS Res UV [2] RS Res UV Res 2.3 花青素着色改变和雄性不育现象 RS RS 1997 Fischer RS Vst1 RS CHS [31] 2011 Liu RS PcRS CHS RS CHS [12] 2012 Rimando O- SbOMT3 RS AhRS3 Res RS [36] RS RS CHS RS RS CHS RS RS [9,24] RS A RS Fischer RS Vst1 Vst1 Antirrhinum majus tap1 RS CHS RS CHS [31] 2006

9 郑世刚等 / 白藜芦醇合成酶基因在基因工程中的应用及功能研究进展 349 Höfig PrMALE1 RS 70% [47] 2011 Ingrosso RS StSy [48] [48] RS 3 RS 基因在微生物中的转化及应用 RS Res Res Res RS Res Res RS A p- A RS p- A Res Res Res Becker Hybrid poplar A 4CL216 RS Vst1 Saccharomyces cerevisiae FY23 p- p- A Res Res [49] Watts RS 4- A 4CL1 Escherichia coli BW mg/l Res [50] Res 2 Res Res Zhang 4CL RS 4- Res Rhodobacter sphaeroides 4CL RS Res 0.65 mg/l 5.25 mg/l 4CL RS cjb@im.ac.cn

10 350 ISSN CN /Q Chin J Biotech March, 2014 Vol.30 No.3 表 2 通过基因工程手段在微生物中进行 Res 的生产 Table 2 Production of Res in microbes by genetic engineering RS gene Co-transformed Res quantity Microorganism Medium Precursor genes (mg/l) Vst1 4CL216 Saccharomyces cerevisiae FY23 SCDL p-coumaric acid 1.45 [49] StSy 4CL2 Saccharomyces cerevisiae YNB p-coumaric acid 6 [54] CEN.PK113-3b Vst1 TAL, 4CL1 Saccharomyces cerevisiae WAT11 SD 4-coumaric acid 14.4 [51] Vst1 4CL1 Saccharomyces cerevisiae YEPD p-coumaric acid 262/391 [52] CEN.PK2-1 / Barra Grande AhRS 4CL1, PAL, C4H, Saccharomyces cerevisiae W303-1A YP Tyrosine 5.8 [55] ACC1 Vst1 4CL, PAL, C4H, Saccharomyces cerevisiae RESV11 CM Phenylalanine 0.31 [56] CPR, AhRS 4CL1 Escherichia coli BW27784 M9 4-coumaric acid 100 [50] AhRS LE4CL Escherichia coli JM109 M9 p-coumaric acid 171 [53] StSy 4CL2 Escherichia coli BL21 2 YT p-coumaric acid 16 [54] Vst1 4CL1 Escherichia coli BW27784 YM9 p-coumaric acid [53] Res Res 14.4 mg/l [51] 2010 Sydor 4CL RS YEPD Res SD YEPD Saccharomyces cerevisiae CEN.PK2-1 Res 262 mg/l S. cerevisiae Barra Grande Res 391 mg/l Res [52] 2011 Lim RS Res 4CL RS puc (BW27784) Res 1.4 g/l 0.05 mmol/l (Cerulenin) Res 2.3 g/l [53] Res Res 2009 Trantas PAL P450 CPR 4- C4H 4CL RS Res [56] 2012 Shin PAL 4- C4H 4CL RS 2% 2.6 mg/l p- 3.3 mg/l Res A ACC1 A Res 4.3 mg/l Res 5.8 mg/l [55] Res

11 郑世刚等 / 白藜芦醇合成酶基因在基因工程中的应用及功能研究进展研究展望 RS Res RS ROS RS RS RS Res RS RS Res Res REFERENCES [1] Delaunois B, Cordelier S, Conreux A, et al. Molecular engineering of resveratrol in plants. Plant Biotechnol J, 2009, 7(1): [2] Kovacic P, Somanathan R. Multifaceted approach to resveratrol bioactivity: focus on antioxidant action, cell signaling and safety. Oxid Med Cell Longev, 2010, 3(2): [3] Han JJ, Liu W, Bi YP. Advances in resveratrol studies. Chin J Biotech, 2008, 24(11): (in Chinese).,,.., 2008, 24(11): [4] Schröder G, Brown JW, Schröder J. Molecular analysis of resveratrol synthase: cdna, genomic clones and relationship with chalcone synthae. Eur J Biochem, 1988, 172(1): [5] Melchior F, Kindl H. Grapevine stilbene synthase cdna only slightly differing from chalcone synthase cdna is expressed in Escherichia coli into a catalytically active enzyme. FEBS Lett, 1990, 268(1): [6] Hain R, Reif HJ, Krause E, et al. Disease resistance results from foreign phytoalexin expression in a novel plant. Nature, 1993, 361(6408): [7] Nicoletti I, De Rossi A, Giovinazzo G, et al. Identification and quantification of stilbenes in fruits of transgenic tomato plants (Lycopersicon esculentum Mill.) by reversed phase HPLC with photodiode array and mass spectrometry detection. J Agric Food Chem, 2007, 55(9): [8] Hain R, Bieseler B, Kindl H, et al. Expression of a stilbene synthase gene in Nicotiana tabacum results in synthesis of the phytoalexin resveratrol. Plant Mol Biol, 1990, 15(2): [9] Lim JD, Yun SJ, Chung IM, et al. Resveratrol synthase transgene expression and accumulation of resveratrol glycoside in Rehmannia glutinosa. Mol Breeding, 2005, 16(3): [10] Han JJ, Liu W, Bi YP. Molecular cloning of peanut resveratrol synthetic enzyme 1 (PNRS1) and its expression in prokaryote. Acta Agron Sin, 2010, 36(2): (in Chinese).,,. PNRS1., 2010, 36(2): [11] Zhong J, Liu SJ, Ma SS, et al. Effect of matrix attachment regions on resveratrol production in tobacco with transgene of stilbene synthase from Parthenocissus henryana. Acta Botan Sin, 2004, 46(8): (in Chinese).,,,.., 2004, 46(8): [12] Liu ZY, Zhuang CX, Sheng SJ, et al. Overexpression of a resveratrol synthase gene cjb@im.ac.cn

12 352 ISSN CN /Q Chin J Biotech March, 2014 Vol.30 No.3 (PcRS) from Polygonum cuspidatum in transgenic Arabidopsis causes the accumulation of trans-piceid with antifungal activity. Plant Cell Rep, 2011, 30(11): [13] Yu CK, Lam CN, Springob K, et al. Constitutive accumulation of cis-piceid in transgenic Arabidopsis overexpressing a sorghum stilbene synthase gene. Plant Cell Physiol, 2006, 47(7): [14] Tian WZ, Ding L, Cao SY, et al. Rice transformation with a phytoalexin gene and bioassay of the transgenic plants. J Integr Plant Biol, 1998, 40(9): (in Chinese).,,,.. :, 1998, 40(9): [15] Serazetdinova L, Oldach KH, Lörz H. Expression of transgenic stilbene synthases in wheat causes the accumulation of unknown stilbene derivatives with antifungal activity. J Plant Physiol, 2005, 162(9): [16] Hipskind JD, Paiva NL. Constitutive accumulation of a resveratrol-glucoside in transgenic alfalfa increases resistance to Phoma medicaginis. Mol Plant Microbe Interact, 2000, 13(5): [17] Kobayashi S, Ding CK, Nakamura Y, et al. Kiwifruits transformed with a vitis stilbene synthase gene produce piceid (resveratrol-glucoside). Plant Cell Rep, 2000, 19(9): [18] Szankowski I, Briviba K, Fleschhut J, et al. Transformation of apple (Malus domestica Borkh.) with the stilbene synthase gene from grapevine (Vitis and a PGIP gene from kiwi (Actinidia deliciosa). Plant Cell Rep, 2003, 22(2): [19] Zhu YJ, Agbayani R, Jackson MC, et al. Expression of the grapevine stilbene synthase gene VST1 in papaya provides increased resistance against diseases caused by Phytophthora palmicora. Planta, 2004, 220(2): [20] Giorcelli A, Sparvoli F, Mattivi F, et al. Expression of the stilbene synthase (StSy) gene from grapevine in transgenic white poplar results in high accumulation of the antioxidant resveratrol glucosides. Transgenic Res, 2004, 13(3): [21] Hüsken A, Baumert A, Milkowski C, et al. Resveratrol glucoside (Piceid) synthesis in seeds of transgenic oilseed rape (Brassica napus L.). Theor Appl Genet, 2005, 111(8): [22] Liu S, Hu Y, Wang X, et al. High content of resveratrol in lettuce transformed with a stilbene synthase gene of Parthenocissus henryana. J Agric Food Chem, 2006, 54 (21): [23] Richter A, Jacobsen HJ, de Kathen A, et al. Transgenic peas (Pisum sativum) expressing polygalacturonase inhibiting protein from raspberry (Rubus idaeus) and stilbene synthase from grape (Vitis vinifera). Plant Cell Rep, 2006, 25(11): [24] Schwekendiek A, Spring O, Heyerick A, et al. Constitutive expression of a grapevine stilbene synthase gene in transgenic hop (Humulus lupulus L.) yields resveratrol and its derivatives in substantial quantities. J Agric Food Chem, 2007, 55(17): [25] Hanhineva K, Kokko H, Siljanen H, et al. Stilbene synthase gene transfer caused alterations in the phenylpropanoid metabolism of transgenic strawberry (Fragaria ananassa). J Exp Bot, 2009, 60(7): [26] Pan LP, Yu SL, Chen CJ, et al. Cloning a peanut resveratrol synthase gene and its expression in purple sweet potato. Plant Cell Rep, 2012, 31(1): [27] Hu XP, Shang WJ, Cai WQ, et al. Transgenic plants of Brassica campestris with resveratrol synthase gene. J Agr Biotechnol, 2006, 14(2): (in Chinese).,,,.., 2006, 14(2): [28] Lin M, Cao L, Guo SH, et al. Transformation of resveratrol synthase (RS) gene of grape into carrot and screening of its transgenic plants with high content of resveratral. J Agr Biotechnol, 2007, 15(6): (in Chinese).

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14 354 ISSN CN /Q Chin J Biotech March, 2014 Vol.30 No.3 25(22): [45] Tang K, Zhan JC, Yang HR, et al. Changes of resveratrol and antioxidant enzymes during UV-induced plant defense response in peanut seedlings. J Plant Physiol, 2010, 167(2): [46] Liang QX, Cao GQ, Su MJ, et al. Research progress on plant leaf senescence. Chin Agric Sci Bull, 2006, 8(22): (in Chinese).,,,.., 2006, 22(8): [47] Höfig KP, Möller R, Donaldson L, et al. Towards male sterility in Pinus radiata a stilbene synthase approach to genetically engineer nuclear male sterility. Plant Biotechnol J, 2006, 4(3): [48] Ingrosso I, Bonsegna S, De Domenico S, et al. Over-expression of a grape stilbene synthase gene in tomato induces parthenocarpy and causes abnormal pollen development. Plant Physiol Biochem, 2011, 49(10): [49] Becker JV, Armstrong GO, van der Merwe MJ, et al. Metabolic engineering of Saccharomyces cerevisiae for the synthesis of the wine-related antioxidant resveratrol. FEMS Yeast Res, 2003, 4(1): [50] Watts KT, Lee PC, Schmidt-Dannert C. Biosynthesis of plant-specific stilbene polyketides in metabolically engineered Escherichia coli. BMC Biotechnol, 2006, 6: [51] Jeandet P, Delaunois B, Aziz A, et al. Metabolic engineering of yeast and plants for the production of the biologically active hydroxystilbene, resveratrol. J Biomed Biotechnol, 2012, 2012: [52] Sydor T, Schaffer S, Boles E. Considerable increase in resveratrol production by recombinant industrial yeast strains with use of rich medium. Appl Environ Microbiol, 2010, 76(10): [53] Lim CG, Fowler ZL, Hueller T, et al. High-yield resveratrol production in engineered Escherichia coli. Appl Environ Microbiol, 2011, 77(10): [54] Beekwilder J, Wolswinkel R, Jonker H, et al. Production of resveratrol in recombinant microorganisms. Appl Environ Microbiol, 2006, 72(8): [55] Shin SY, Jung SM, Kim MD, et al. Production of resveratrol from tyrosine in metabolically engineered Saccharomyces cerevisiae. Enzyme Microb Technol, 2012, 51(4): [56] Trantas E, Panopoulos N, Ververidis F. Metabolic engineering of the complete pathway leading to heterologous biosynthesis of varius flavonoids and stilbenoids in Saccharomyces cerevisiae. Metab Eng, 2009, 11(6): ( 本文责编陈宏宇 )

Microsoft Word - 793-797 tb20150504赵宏宇s-高校教改纵横.doc

Microsoft Word - 793-797 tb20150504赵宏宇s-高校教改纵横.doc 微生物学通报 Microbiology China tongbao@im.ac.cn Apr. 20, 2016, 43(4): 793 797 http://journals.im.ac.cn/wswxtbcn DOI: 10.13344/j.microbiol.china.150504 高校教改纵横生物工程专业发酵课程群建设探索 * 赵宏宇