红暗柳橙脂类相关质体蛋白基因的克隆与功能初步分析
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摘要
‘红暗柳’甜橙(Citrus sinensis[L.]Osbeck)是一个特异性积累番茄红素的普通甜橙红肉突变体。‘红暗柳’与‘暗柳’的比较蛋白组学研究发现一个脂类相关质体蛋白(PAP)的表达伴随果实发育与色素累积显著上升,且在‘红暗柳’中的累积量高出暗柳两倍以上。质体相关蛋白是一大类古老而保守的蛋白质家族,参与脂蛋白结构如质体球蛋白和原纤维的合成,与质体分化和类胡萝卜素形成和储存密切相关。本研究旨在分离编码该蛋白的基因CSPAP并通过遗传转化的方法对其功能验证作初步的探索。主要工作和结果如下：
1、在‘红暗柳’中克隆到CsRAP的cDNA全长为972 bp的片段,与已报道编码PAP蛋白的基因有很高的同源性；构建了CsPAP基因原核表达载体并在大肠杆菌中诱导了预测大小约为29 kD的特异蛋白的表达。
2、构建了CsPAP基因的植物组成型表达载体,并以农杆菌介导的方法对国庆1号(Citrus unshiu)愈伤组织、山金柑(Fortunella hindsii)上胚轴以及番茄(Lycopersicon esculentum)子叶进行了遗传转化。通过筛选培养获得了国庆1号的抗性愈伤组织和番茄的再生抗性苗,经PCR检测为转基因阳性。
3、Real-time PCR检测到一个抗性愈伤系中CsPAP的表达量比未转基因对照高出50倍以上。另外,ABA处理和逆境处理(NaCl.H202和脱水处理)能影响伏令夏橙愈伤组织中CsPAP基因的表达。
'Hong anliu'sweet orange (Citrus sinensis [L.] Osbeck) is a red-fleshed mutant dominantly accumulating lycopene. In the comparative proteomics study of'Hong anliu' (MT, mutant) and'Anliu'(WT, wild type), a plastid lipid-associated protein (PAP) was detected to increase in parallel to the accumulation of pigments during the development of fruits and its expression in MT was over two-fold of that in WT. PAP is a large group of protein family involving in the formation of plastoglobules and fibrils, plastid differentiation and carotenoid sequestration. The research aimed at cloning the gene encoding CsPAP and primarily analyzing its function by genetic transformation.
The isolated cDNA was 972 bp in length and had high identity rate with PAP genes reported in other species. The prokaryotic expression vector carrying cDNA fragment was constructed and transformed into E. coli to induce protein with a predicted molecular weight of 29 kD.
The eukaryotic constitutive expression vector was also constructed to over-express CsPAP gene in callus (Citrus unshiu), epicotyls (Fortunella hindsii) and tomato cotyledon (Lycopersicon esculentum) by Agrobacterium-mediated transformation. Kanamycin-resistant citrus callus and regenerated tomato plants were obtained by selective culture and verified to be transgenic by PCR.
Real-time PCR analysis revealed that CsPAP gene expressed over 50-fold in one line of transgenic citrus callus than in CK. Meanwhile, ABA treatment and certain stress conditions (NaCl, H2O2, and dehydration treatment) could bring out effects on the expression of CsPAP in Valencia callus (Citrus sinensis).
1、在‘红暗柳’中克隆到CsRAP的cDNA全长为972 bp的片段,与已报道编码PAP蛋白的基因有很高的同源性；构建了CsPAP基因原核表达载体并在大肠杆菌中诱导了预测大小约为29 kD的特异蛋白的表达。
2、构建了CsPAP基因的植物组成型表达载体,并以农杆菌介导的方法对国庆1号(Citrus unshiu)愈伤组织、山金柑(Fortunella hindsii)上胚轴以及番茄(Lycopersicon esculentum)子叶进行了遗传转化。通过筛选培养获得了国庆1号的抗性愈伤组织和番茄的再生抗性苗,经PCR检测为转基因阳性。
3、Real-time PCR检测到一个抗性愈伤系中CsPAP的表达量比未转基因对照高出50倍以上。另外,ABA处理和逆境处理(NaCl.H202和脱水处理)能影响伏令夏橙愈伤组织中CsPAP基因的表达。
'Hong anliu'sweet orange (Citrus sinensis [L.] Osbeck) is a red-fleshed mutant dominantly accumulating lycopene. In the comparative proteomics study of'Hong anliu' (MT, mutant) and'Anliu'(WT, wild type), a plastid lipid-associated protein (PAP) was detected to increase in parallel to the accumulation of pigments during the development of fruits and its expression in MT was over two-fold of that in WT. PAP is a large group of protein family involving in the formation of plastoglobules and fibrils, plastid differentiation and carotenoid sequestration. The research aimed at cloning the gene encoding CsPAP and primarily analyzing its function by genetic transformation.
The isolated cDNA was 972 bp in length and had high identity rate with PAP genes reported in other species. The prokaryotic expression vector carrying cDNA fragment was constructed and transformed into E. coli to induce protein with a predicted molecular weight of 29 kD.
The eukaryotic constitutive expression vector was also constructed to over-express CsPAP gene in callus (Citrus unshiu), epicotyls (Fortunella hindsii) and tomato cotyledon (Lycopersicon esculentum) by Agrobacterium-mediated transformation. Kanamycin-resistant citrus callus and regenerated tomato plants were obtained by selective culture and verified to be transgenic by PCR.
Real-time PCR analysis revealed that CsPAP gene expressed over 50-fold in one line of transgenic citrus callus than in CK. Meanwhile, ABA treatment and certain stress conditions (NaCl, H2O2, and dehydration treatment) could bring out effects on the expression of CsPAP in Valencia callus (Citrus sinensis).
引文
1.邓秀新,徐建国.柑橘产业现状.见：中国柑橘学会编著,中国柑橘产业.北京：中国农业出版社,2008,17-24
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3.景士西主编.园艺植物育种学总论.北京：中国农业出版社,2000,116-118
4.刘庆.‘暗柳’甜橙红色突变体性状形成的分子机理研究.[博士学位论文].武汉：华中农业大学图书馆,2008
5.刘永忠.脐橙(Citrus sinensis Osbeck)晚熟芽变性状形成机理研究.[博士学位论文].武汉：华中农业大学图书馆,2006
6.沈德绪主编.果树育种学.北京：中国农业出版社,1992,79-91
7.陶能国.甜橙(Citrus sinensis Osbeck)红肉突变体类胡萝卜素合成相关基因的克隆与特性分析.[博士学位论文].武汉：华中农业大学图书馆,2006
8.徐娟.几个柑橘产区果实色泽评价及红肉脐橙(Citrus sinensis L. cv. Cara cara)果肉呈色机理初探.[博士学位论文].武汉：华中农业大学图书馆,2002
9.杨颖丽,史如霞,魏学玲,范庆,王莱.NaCl胁迫对唐古特白刺愈伤组织的生理效应.西北植物学报,2008,28(11)：2238-2243
10.张建成.红肉脐橙类胡萝卜素合成酶基因(CsPSY, CsLCYb)功能分析与crtB转基因对柑橘类胡萝卜素生物合成的影响.[博士学位论文].武汉,华中农业大学图书馆,2009
11.张敏,邓秀新.柑橘芽变选种以及芽变性状形成机理研究进展.果树学报,2006：23(6)：871-876
12.周文静.红肉脐橙(Citrus sinensis Osbeck)番茄红素β-环化酶基因(LCYb)功能的初步验证和分析.[硕士学位论文].武汉：华中农业大学图书馆,2008
13.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.植物生理与分子生物学学报,2004,30(6)：609-618
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2.段艳欣.根癌农杆菌介导的柑橘转化体系优化与转LFY、 API基因植株的培育.[博士学位论文].武汉：华中农业大学图书馆,2006
3.景士西主编.园艺植物育种学总论.北京：中国农业出版社,2000,116-118
4.刘庆.‘暗柳’甜橙红色突变体性状形成的分子机理研究.[博士学位论文].武汉：华中农业大学图书馆,2008
5.刘永忠.脐橙(Citrus sinensis Osbeck)晚熟芽变性状形成机理研究.[博士学位论文].武汉：华中农业大学图书馆,2006
6.沈德绪主编.果树育种学.北京：中国农业出版社,1992,79-91
7.陶能国.甜橙(Citrus sinensis Osbeck)红肉突变体类胡萝卜素合成相关基因的克隆与特性分析.[博士学位论文].武汉：华中农业大学图书馆,2006
8.徐娟.几个柑橘产区果实色泽评价及红肉脐橙(Citrus sinensis L. cv. Cara cara)果肉呈色机理初探.[博士学位论文].武汉：华中农业大学图书馆,2002
9.杨颖丽,史如霞,魏学玲,范庆,王莱.NaCl胁迫对唐古特白刺愈伤组织的生理效应.西北植物学报,2008,28(11)：2238-2243
10.张建成.红肉脐橙类胡萝卜素合成酶基因(CsPSY, CsLCYb)功能分析与crtB转基因对柑橘类胡萝卜素生物合成的影响.[博士学位论文].武汉,华中农业大学图书馆,2009
11.张敏,邓秀新.柑橘芽变选种以及芽变性状形成机理研究进展.果树学报,2006：23(6)：871-876
12.周文静.红肉脐橙(Citrus sinensis Osbeck)番茄红素β-环化酶基因(LCYb)功能的初步验证和分析.[硕士学位论文].武汉：华中农业大学图书馆,2008
13.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.植物生理与分子生物学学报,2004,30(6)：609-618
14. Alquezar B, Rodrigo M J, Zacarias L. Regulation of carotenoid biosynthesis during fruit maturation in the red-fleshed orange mutant Cara Cara. Phytochem,2008,69: 1997-2007
15. Altman A, Goren R. Promotion of callus formation by abscisic acid in citrus bud cultures. Plant Physiol,1971,47(6):844-846
16. Bramley P M. Regulation of carotenoid formation during tomato fruit ripening and development. JExp Bot,2002,377:2107-2113
17. Brehelin C, Kessler F, Van Wijk K J. Plastoglobules:versatile lipoprotein particles in plastids. Trends Plant Sci,2007,12(6):260-266
18. Burkhardt P K, Beyer P, Wunn J, Klbti A, Armstrong G A, Schledz M, von Lintig J, Potrykus I. Transgenic rice (Oryza sativa) endosperm expressing daffodil (Narcissus pseudonarcissus) phytoene synthase accumulates phytoene biosynthesis. Plant J, 1997,11(5):1071-1078
19. Cazzonelli C I, Pogson B J. Source to sink:regulation of carotenoid biosynthesis in plants. Trends Plant Sci,2010,15(5):266-274
20. Cheng Y J, Guo W W, Yi H L, Pang X M, and Deng X X. An efficient protocol for genomic DNA extraction from Citrus species. Plant Mol Biol Rep,2003,21: 177a-177g
21. Cunningham F X, Gantt E. Genes and enzymes of carotenoid biosynthesis in plants. Annu Rev Plant Phys,1998,49:557-583
22. Cuttriss A J, Chubb A C, Alawady A, Grimm B, Pogson B J. Regulation of lutein biosynthesis and prolamellar body formation in Arabidopsis. Funct Plant Biol,2007, 34(8):663-672
23. Deruere J, Romer S, d'Harlingue A, Backhaus R A, Kuntz M, Camara B. Fibril assembly and carotenoid overaccumulation in chromoplasts:a model for supramolecular lipoprotein structures. Plant Cell,1994,6(1):119-133
24. Ducreux L J M, 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 p-carotene and lutein. JExp Bot,2005,56(409):81-89
25. Fraser P D, Bramley P M. The biosynthesis and nutritional uses of carotenoids. Prog Lipid Res,2004,43:228-265
26. Galpaz N, Wang Q, Menda N, Zamir D, Hirschberg J. Abscisic acid deficiency in the tomato mutant high-pigment 3 leading to increased plastid number and higher fruit lycopene content. Plant J,2007,53(5):717-730
27. Gillet B, Beyly A, Peltier G, Rey P. Molecular characterization of CDSP 34, a chloroplastic protein induced by water deficit in Solanum tuberosum L. plants, and regulation of CDSP34 expression by ABA and high illumination. Plant J,2001, 16(2):257-262
28. Giuliano G, Tavazza R, Diretto G, Beyer P, Taylor M A. Metabolic engineering of carotenoid biosynthesis in plants. Trends Biotechnol,2008,26(3):139-145
29. Joyard J, Ferro M, Masselon C, Seigneurin-Berny D, Salvi D, Garin J, Rolland N. Chloroplast proteomics and the compartmentation of plastidial isoprenoid biosynthetic pathways. Mol Plant,2009,2(6):1154-1180
30. Kato M, Ikoma Y, Matsumoto H, Sugiura M, Hyodo H, Yano M. Accumulation of carotenoids and expression of carotenoid biosynthetic genes during maturation in citrus fruit. Plant Physiol,2004,134(2):824-837
31. Kobayashi S, Goto-Yamamoto N, Hirochika H. Retrotransposon-induced mutations in grape skin color. Science,2004,304:982-982
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