豆峻岭;刘文革;

• 1:中国农业科学院郑州果树研究所

摘要(Abstract)：

番茄红素广泛存在于各种植物中,它不仅是植物花、果实等形成红色的主要色素,也是对人体健康有益的一种类胡萝卜素。它在植物体内的合成及分解涉及许多中间物质,并且受到多种酶基因的调控,而这些中间物质也参与植物体内其他物质的合成代谢。目前围绕着番茄红素的生物合成以及合成途径中相关酶基因的研究越来越多,并且通过对关键酶基因的超量表达、RNA沉默已经获得了一些高番茄红素植株。综述了国内外关于番茄红素合成途径相关酶基因的研究、影响其生物合成的一些相关因素以及为提高植物中番茄红素含量而进行的一些基因工程方面的研究,以期为进一步转基因研究以及高番茄红素植物品种的选育提供思路。

关键词(KeyWords)： 番茄红素;生物合成;酶基因;影响因素

Abstract:

Keywords:

基金项目(Foundation): 国家自然科学基金(31171979);; 现代农业产业技术体系建设专项(CARS-26-03);; 中国农业科学院与北京大兴区政府科研合作项目

作者(Author): 豆峻岭;刘文革;

Email:

DOI: 10.13925/j.cnki.gsxb.2013.04.001

参考文献(References)：

• [1]MILLAYDET,BULL.Processing of tomatoes and solanombin extrac-tion[J].Soc Naney,1875,2(1):21-23.
• [2]FORD N A,ERDMAN J W.Are lycopene metabolites metabolically active[J].Acta Biochimica Polonica,2012,59(1):1-4.
• [3]GIOVANUCCI E,ASCHERIO A,RIMM E B.Intake of carotenoids and retinol in relationship to risk of prostate cancer[J].Journal of the National Cancer Institute,1995(87):1767-1776.
• [4]ILAHY R,HDIDER C,LENUCCI M S,TLILI I,DALESSANDRO G.Phytochemical composition and antioxidant activity of high-ly-copene tomato(Solanum lycopersicum L.)cultivars grown in South-ern Italy[J].Scientia Horticulturae,2011,127:255-261.
• [5]BLUM A,MONIR M,WIRSANSKY I,BEN-ARZI S.The benefi-cial effects of tomatoes[J].European Journal of Internal Medicine,2005,16(6):402-404.
• [6]WATZL B,BUB A,BRIVIBA K,RECHKEMMER G.Supplemen-tation of a low-carotenoid diet with tomato or carrotjuice modulates immune functions in healthy men[J].Annals Nutrition Metabolism,2003,47(6):255-261.
• [7]NGUYEN M L.Lycopene:Chemical and biological properties[J].Food Technology,1999,53(2):38-45.
• [8]MINLEE J,JOUNG J G,MCQUINN R,CHUNG M Y,ZHANG J F,DENISE T,GIOVANNONI J.Combined transcriptome,genetic diversity and metabolite profiling in tomato fruit reveals that the ethylene response factor SlERF6plays an important role in ripening and carotenoid accumulation[J].The Plant Journal,2012,70:191-204.
• [9]GALPAZ N,WANG Q,MENDA M,ZAMIR D,HIRSCHBERG J.Abscisic acid deficiency in the tomato mutant high-pigment3lead-ing to increased plastid number and higher fruit lycopene content[J].The Plant Journal,2008,53:717-730.
• [10]SUN Ming-qi,PAN Si-yi,HU Jian-zhong,FU Hong-fei,XU Xi-ao-yun.Study on effect of environment on stability of Citrus peel carotenoid[J].Food Science,2008,29(6):127-129.孙明奇,潘思轶,胡建中,傅虹飞,徐晓云.环境条件对柑橘果皮类胡萝卜素稳定性影响研究[J].食品科学,2008,29(6):127-129.
• [11]HIRSEHBERG J.Carotenoid biosynthesis in flowering plants[J].Cur-rent Opinion in Plant Biology,2001,4:210-218.
• [12]ISAACSON T,RONEN G,ZAMIR D,HIRSCHBERG J.Cloning of tangerine from tomato reveals a carotenoid isomerase essential for the production ofβ-carotene and xanthophylls in plants[J].The Plant Cell,2002,14:333-342.
• [13]NAQVI S,ZHU C F,FARRE G,SANDMANN G,CHIRSTO P.Synergistic metabolism in hybrid corn indicates bottlenecks in the carotenoid pathway and leads to the accumulation of extraordinary levels of the nutritionally important carotenoid zeaxanthin[J].Plant Biotechnology Journal,2011,9:384-393.
• [14]ZHU Chang-fu,CHEN Xing,WANG Ying-dian.Carotenoid biosyn-thesis in plants and application of its relative genes in gene engi-neering[J].Journal of Plant Physiology and Molecular Biology,2004,30(6):609-615.朱长甫,陈星,王英典.植物类胡萝卜素生物合成及其相关基因在基因工程中的应用[J].植物生理与分子生物学学报,2004,30(6):609-615.
• [15]WANG Yu-ping,LIU Qing-chang,ZHAI Hong.Expression and regulation of genes related to plant carotenoid biosynthesis and their application in plant gene engineering[J].Molecular Plant Breeding,2006,4(1):103-110.王玉萍,刘庆昌,翟红.植物类胡萝卜素生物合成相关基因的表达调控及其在植物基因工程中的应用[J].分子植物育种,2006,4(1):103-110.
• [16]DUCREUX L J,MORRIS W L,HEDLEY P E,SHEPHERD T,DAVIES H V,MILLAM S,TAYLOR M A.Metabolic engineering of high carotenoid potato tubers containing enhanced levels of beta-carotene and lutein[J].Jounal of Experimental Botany,2005,56(409):81-89.
• [17]THORUP T A,TANYOLAC B,LIVINGSTONE K D,POPOVSKY S,PARAN I,JAN M.Candidate gene analysis of organ pigmentation loci in the Solanaceae[J].Proceedings of the National Academy of Sciences of the United States of America,2000,97(21):11192-11197.
• [18]ZHU Hai-sheng,LI Yong-ping,WEN Qing-fang.Cloning and ex-pression analysis of phytoene synthase gene in fragaria ananassa duch.[J].Scientia Agricultura Sinica,2011,44(2):349-357.朱海生,李永平,温庆放.草莓八氢番茄红素合成酶基因的克隆及其表达特性[J].中国农业科学,2011,44(2):349-357.
• [19]TAO Neng-guo.Cloning and characterization of carotenoid biosyn-thetic genes from sweet orange(Citrus Sinesis Osbeck)red-fleshed mutants[D].Wuhan:Huazhong Agricultural University,2006.陶能国.甜橙红肉突变体类胡萝卜素合成相关基因的克隆与相关性分析[D].武汉:华中农业大学,2006.
• [20]ZHAO Wen-en,KANG Bao-shan,HU Guo-qin.Advances in re-search on carotenoids in watermelon flesh[J].Journal of Fruit Sci-ence,2008,25(6):908-915.赵文恩,康保姗,胡国勤.西瓜瓤类胡萝卜素研究进展[J].果树学报,2008,25(6):908-915.
• [21]KANG B,ZHAO W,HOU Y B,TIAN P.Expression of carotenogenic genes during the development and ripening of watermelon fruit[J].Scientia Horticulturae,2010,124:368-375.
• [22]BANG H,KIM S,LESKOVAR D.Differential expression of carotenoid biosynthesis genes among different colored flesh in watermelons[Citrullus lanatus(Thunb)][J].Hort Science,2004,39:869-870.
• [23]OKADA K,SAITO T,NAKAGAWA T,KAWAMUKAI M,KAMIYA Y.Five geranylgeranyl diphosphate synthases expressed in different organs are localized into three subcellular compartments in Ara-bidopsis[J].Plant Physiology,2000,122(4):1045-1056.
• [24]AMENT K,VAN SCHIE C C,BOUWMEESTER H J,HARING M A,SCHUURINK R C.Induction of a leaf specific geranylgeranyl pyrophosphate synthase and emission of(E,E)-4,8,12-trimethyl-trideca-1,3,7,11-tetraene in tomato are dependent on both jas-monic acid and salicylic acid signaling pathways[J].Planta,2006,224(5):1197-1208.
• [25]ZHU C,YAMAMURA S,KOIWA H,NISHIHARA M,SAND-MANN G.cDNA cloning and expression of carotenogenic genes dur-ing flower develo Pmnt in Gentiana lutea[J].Plant Molecular Biolo-gy,2002,8(3):277-285.
• [26]WELSCH R,WUST F,BAR C,AL-BABILI S,BEYER P.A third phytoene synthase is devoted to abiotic stress-induced abscisic acid formation in rice and defines functional diversification of phytoene synthase genes[J].Plant Physiology,2008,147(1):367-380.
• [27]CHARLES A D,SUPINYA D,LEWIS D,PAUL S,ALLAN A C.Metabolic and gene expression analysis of apple(Malus×domestica)carotenogenesis[J].Journal of Experimental Botany,2012,63(12):4497-4511.
• [28]GIULIANO G,BARTLEY G E,SCOLNIK P A.Regulation of carotenoid biosynthesis during tomato development[J].The Plant cell,1993,5(4):379-387.
• [29]RONEN G,COHEN M,ZAMIR D,HIRSCHBERG J.Regulation of carotenoid biosynthesis during tomato fruit development:expression of the gene for lycopene epsilon-cyclase is down-regulated during ripening and is elevated in the mutant Delta[J].The Plant Journal,1999,17(4):341-351.
• [30]KATO M,IKOMA Y,MATSUMOTO H,SUGIURA M,HIROSHI-HYODO,YANO M.Accumulation of carotenoids and expression of carotenoid biosynthetic genes during maturation in citrus fruit[J].Plant Physiology,2004,134:824-837.
• [31]BASSENE J B,FROELICHER Y,DUBOIS C,FERRER R M,NAVARRO L,OLLITRAULT P,ANCILLO G.Non-additive phe-notypic and transcriptomic inheritance in a citrus allotetraploid so-matic hybrid between C.reticulate and C.limon:the case of pulp carotenoid biosynthesis pathway[J].Plant Cell Reports,2009,28:1689 1697.
• [32]CLOTAULT J,PELTIER D,BERRUYER R,THOMAS M,BRI-ARD M,GEOFFRIAU E.Expression of carotenoid biosynthesis genes during carrot root development[J].Journal of Experimental Botany,2008,59(13):3563-3573.
• [33]PECKER I,GABBAY R,FRANCIS X,CUNNINGHAM J,JOSEPH H.Cloning and characterization of the cDNA for lycopeneβ-cyclase from tomato reveals decrease in its expression during fruit ripening[J].Plant Molecular Biology,1996,30(4):807-819.
• [34]YUAN Ping-li.The study of lycopene and key gene expression dur-ing fruit development with different ploidy watermelon[D].Beijing:Chinese Academy of Agricultural Sciences,2012.袁平丽.多倍体西瓜发育过程中番茄红素变化及关键酶基因表达[D].北京:中国农业科学院,2012.
• [35]WAN Xue-shan,LIU Wen-ge,YAN Zhi-hong,ZHAO Sheng-jie,HE Nan,LIU Peng.Changes of the contents of functional substances including lycopene,citrulline and ascorbic acid during watermelon fruits development[J].Scientia Agricultura Sinica,2011,44(13):2738-2747.万学闪,刘文革,阎志红,赵胜杰,何楠,刘鹏.西瓜果实发育过程中番茄红素、瓜氨酸和VC等功能物质含量的变化[J].中国农业科学,2011,44(13):2738-2747.
• [36]YUAN Ping-li,LIU Wen-ge,ZHAO Sheng-jie,LU Xu-qiang,YAN Zhi-hong,HE Nan,ZHU Hong-ju.Lycopene content and ex-pression of phytoene synthase and lycopeneβ-cyclase genes in te-traploid watermelon[C].Proceedings of the Xth EUCARPIA Meeting on Genetics and Breeding of Cucurbitaceae.2012:316-324.
• [37]LIU W,ZHAO S,CHENG Z,WAN X,YAN Z,KING S R.Lycopene and citrulline contents in watermelon(Citrullus lanatus)fruit with different ploidy and changes during fruit development[J].Acta Horti-culturae,2009,871:543-549.
• [38]YUAN Ping-li,LIU Wen-ge,LU Xu-qiang,ZHAO Sheng-jie,LU Jin-sheng,YAN Zhi-hong,HE Nan,ZHU Hong-ju.Change of ly-copene contents during fruit development of different ploidy water-melon[J].Journal of Fruit Science,2012,29(5):890-894.袁平丽,刘文革,路绪强,赵胜杰,芦金生,阎志红,何楠,朱红菊.不同倍性西瓜果实发育过程中番茄红素含量动态[J].果树学报,2012,29(5):890-894.
• [39]ZHANG J,TAO N,XU Q,ZHOU W,CAO H,XU J,DENG X.Functional characterization of citrus PSY gene in Hongkong kumquat(Fortunella hindsii Swingle)[J].Plant Cell Reports,2009,28(11):1737-1746.
• [40]DHARMAPURI S,ROSATI C,PALLARA P,AQUILANI R,BOU-VIER F,GIULIANO G.Metabolic engineering of xanthophyll con-tent in tomato fruits[J].Febs Letters,2002,519(1-3):30-34.
• [41]ZHOU Wen-jing.Functional verification of the lycopene beta-cy-clase gene(Lcyb)from’Cara Cara’navel orange(Citrus Sinensis[L.]Osbeck)[D].Wuhan:Huazhong Agricultural University,2008.周文静.红肉脐橙番茄红素β-环化酶基因(Lcyb)功能的初步验证和分析[D].武汉:华中农业大学,2008.
• [42]KIM S H,KIM Y H,AHN Y O,AHN M J,JEONG J C,LEE H S,KWAK S S.Downregulation of the lycopeneε-cyclase gene increas-es carotenoid synthesis via theβ-branch-specific pathway and en-hances salt-stress tolerance in sweetpotato transgenic calli[J].Physi-ologia Plantarum,2013,147:432-442.
• [43]ROSATI C,AQUILANI R,DHARMAPURI S,GIULIANO G.Metabol-ic engineering of beta-carotene and lycopene content in tomato fruit[J].The Plant Journal,2000,24(3):413-420.
• [44]WAN Qun,ZHANG Xing-guo,SONG Ming.Fruit-specific RNAi-mediated LCY gene silencing enhances content of lycopene in toma-toes[J].Chinese Journal of Biotechnology,2007,23(3):429-433.万群,张兴国,宋明.果实特异性RNAi介导的LCY基因沉默来增加番茄中番茄红素的含量[J].生物工程学报,2007,23(3):429-433.
• [45]MA Chao,MA Bing-gang,HAO Qing-nan,HE Juan,LU Xiao-yan,WU Zhi-ping.Construction of RNA Interference vector for lycopene cyclaseβgene(Lyc-β)and its expression identification in lycoper-sicun esculentum[J].Journal of Agricultural Biotechnology,2010,18(1):10-17.马超,马兵钢,郝青南,何娟,鲁晓燕,吴志苹.番茄红素β环化酶基因(Lyc-β)RNAi载体构建及表达鉴定[J].农业生物技术学报,2010,18(1):10-17.
• [46]NAMBARA E,MARION-POLL A.Abscisic acid biosynthesis and catabolism[J].Annual Review Plant Biology,2005(56):165-185.
• [47]YAMAGUCHI S.Gibberellin biosynthesis in Arabidopsis[J].Phyto-chemistry Review,2006(5):39-47.
• [48]SUN L.Fruit-specific RNAi-mediated suppression of SlNCED1in-creases both lycopene and beta-carotene contents in tomato fruit[J].Journal of Experimental Botany,2012,63(8):3097-3108.
• [49]GAO H,ZHU H,SHAO Y,CHEN A,LU C,ZHU B,LUO Y.Ly-copene accumulation affects the biosynthesis of some carotenoid-re-lated volatiles independent of ethylene in tomato[J].Journal of Inte-grative Plant Biology,2008,50(8):991-996.
• [50]WANG Gui-yuan,XIA Ren-xue.Changes in the contents of ly-copene,beta-carotene,sugar and endogenous GA and ABA in flesh during the fruit development of‘Cara Cara’orange[J].Acta Horticul-turae Sinica,2005,32(3):482-485.王贵元,夏仁学.红肉脐橙果实发育过程中番茄红素、β-胡萝卜素、糖、GA、ABA含量的变化[J].园艺学报,2005,32(3):482-485.
• [51]SOREFAN K,BOOKER J,HAUROGNE K,GOUSSOT M,BAIN-BRIDGE K,FOO E,CHATFIELD S,WARD S,BEVERIDGE C,RAMEAU C,LEYSER O.MAX4and RMS1are orthologous dioxy-genase-like genes that regulate shoot branching in Arabidopsis and pea[J].Genes&Development,2003,17:1469-1476.
• [52]DUN E A,BREWER P B,BEVERIDGE C A.Strigolactones:dis-covery of the elusive shoot branching hormone[J].Trends in Plant Science,2009,14(7):364-372.
• [53]BEVERIDGE C A,KYOZUKA J.New genes in the strigolactone-re-lated shoot branching pathway[J].Current Opinion in Plant Biology,2010,13:34-39.
• [54]LIU L,WEI J,ZHANG M,ZHANG L,LI C,WANG Q.Ethylene independent induction of lycopene biosynthesis in tomato fruits by jasmonates[J].Journal of Experimental Botany,2012,3:1-12.
• [55]SHENG Ji-ping,LUO Yun-bo.The research tomato fat oxygen syn-thase in ethylene biosynthesis approach[J].Journal of China Agricul-tural University,1999,4(5):50.生吉萍,罗云波.番茄脂氧合酶参与乙烯生物合成途径研究[J].中国农业大学学报,1999,4(5):50.
• [56]ZHUANG Xue-ping,ZHANG Wen,ZHNEG Cai-xia,DU Wei-min.Effect of glucose,sodium glutamate and ethephon on synthetic quantity of lycopene in tomato fruits[J].Journal of Anhui Agricultural Sciences,2007,35(19):5664-5665,5669.庄学平,张雯,郑彩霞,杜为民.3种物质对番茄果实中番茄红素合成量的影响[J].安徽农业科学,2007,35(19):5664-5665,5669.
• [57]WENG Qian,ZHOU Bao-li,YU Yang,FU Ya-wen.Effects of ex-ogenous ABA,BR,ETH on changes of lycopene’s contents in fruit of tomato[J].Journal of Shenyang Agricultural University,2007,38(6):784-787.翁倩,周宝利,于洋,付亚文.外源ABA、BR和ETH对番茄果实番茄红素含量的影响[J].沈阳农业大学学报,2007,38(6):784-787.
• [58]WANG Gui-yuan,XIA Ren-xue,ZHOU Kai-bing.Effects on changes of dominating pigment contents in peel of cara cara(Citrus sinensis Osbesk)and fruit colouring after exogenous ABA and GA 3 treatments[J].Journal of Wuhan Botanical Research,2004,22(3):273-276.王贵元,夏仁学,周开兵.外源ABA和GA 3对红肉脐橙果皮主要色素含量变化和果实着色的影响[J].武汉植物学研究,2004,22(3):273-276.
• [59]TAO Jun,ZHANG Shang-long.Metabolism of carotenoid and its regulation in garden crop[J].Journal of Zhejiang University:Agri-culture&Life Sciences,2003,9(5):585-590.陶俊,张上隆.园艺植物类胡萝卜素的代谢及其调节[J].浙江大学学报:农业与生命科学版,2003,9(5):585-590.
• [60]ALBAR,PRATTM,LEEH.Fruit-localized phytochromes regulate lycopene accumulation independently of ethylene production in tomato[J].Plant Physiology,2000,123:363-370.
• [61]ROBERTSONL G H,MAHONEY N E,GOODMAN N,PAVLATH A E.Regulation of lycopene formation in cell suspension culture of VFNT tomato(Lycopersicon esculentum)by CPTA,growth regula-tors,sucrose,and temperature[J].Journal of Experimental Botany,1995,46(6):667-673.
• [62]SUN Ming-qi.Study on extraction and identification of carotenoids in citrus peel and transformation regulation during maturation and preservation[D].Wuhan:Huazhong Agricultural University,2007.孙明奇.柑橘皮类胡萝卜素提取鉴定及其在果皮成熟储藏中的变化规律[D].武汉:华中农业大学,2007.
• [63]NIU Xiao-li,ZHOU Zhen-jiang,LI Rui,HU Tian-tian.Effects of water and fertilizer supply on lycopene content in tomato fruit[J].Ac-ta Horticulturae Sinica,2011,38(11):2111-2120.牛晓丽,周振江,李瑞,胡田田.水肥供应对番茄中番茄红素含量的影响[J].园艺学报,2011,38(11):2111-2120.
• [64]HE Nan,ZHAO Sheng-jie,LIU Wen-ge,YAN Zhi-hong,WAN Xue-shan.Effect of potassium fertilizer on lycopene of mature wa-termelon[J].Journal of Changjiang Vegetables,2008,12b:21-22.何楠,赵胜杰,刘文革,阎志红,万学闪.施用钾肥对西瓜番茄红素的影响研究[J].长江蔬菜,2008,12b:21-22.
• [65]HE Chun-mei,LIN Ying,LI Hong-mei.Cultural environment and storage period of tomato fruit in the influence of the content of ly-copene[J].Journal of Henan Agricultural Sciences,2010(5):93-96.何春玫,林莹,李红梅.栽培环境及贮存期对番茄果实内番茄红素含量的影响[J].河南农业科学,2010(5):93-96.
• [66]LI Ji-suo.The primary studies of the influence factors and heredity on the lycopene content of tomato(Lycopersicum Esculentum Mill.)[D].China Agricultural University,2003.李纪锁.番茄中番茄红素含量影响因素及遗传的初步研究[D].中国农业大学,2003.