蓖麻细胞色素P450基因RNAi植物表达载体构建及遗传转化的研究
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摘要
在植物细胞分裂和胚胎发育过程中，植物激素油菜素内酯（BR）和赤霉素（GA）参与节间的发育过程，从而影响植株的高度。植物体内BR和GA缺失将表现矮化表型。上述两种激素的合成调控受到很多酶的影响，大多数酶属于细胞色素P450家族，该家族的基因对植株茎秆的发育有很大影响。蓖麻细胞色素P450基因和水稻细胞色素P450基因氨基酸序列同源性为60.98%，说明它们属于同一亚家族，推测两个基因的功能相似。为了验证蓖麻细胞色素P450基因的功能，从而揭示该基因与激素合成之间的关系，本研究根据已公布的蓖麻细胞色素P450基因的碱基序列，克隆了该基因的两个片段，成功构建了RNAi植物表达载体。分别以蓖麻品种“通篦5号”的子叶、胚轴和子叶节为外植体，研究了不同种类和浓度的细胞分裂素和生长素对外植体再生不定芽和根的影响，获得了再生频率较高的蓖麻子叶节离体再生体系。通过子叶节和农杆菌共培养得到了转基因阳性植株，为研究蓖麻细胞色素P450基因的功能奠定了基础，主要结果如下：
1．根据Genbank（XM_002525863.1）公布的蓖麻细胞色素P450基因的碱基序列设计了两对引物，PCR扩增得到P450基因的两个片段，分别命名为P450s和P450a，与克隆载体成功连接后测序并进行序列比对，结果显示：P450s全长523bp，与原序列一致性为98.28%；P450a全长411bp，与原序列一致性为99.51%，说明克隆的片段是蓖麻细胞色素P450基因片段。
2．利用SacI和KpnI对重组质粒pMD-P450s和中间载体pHANNIBAL双酶切后，将P450s定向连接到线性中间载体pHANNIBAL上，构建了载体pHAN-P450s，酶切和PCR检测正确；然后利用XbaI和HandⅢ双酶切重组质粒pMD-P450a和载体pHAN-P450s，将P450a定向连接到线性载体pHAN-P450s上，构建了载体pHAN-P450s-P450a，酶切和PCR检测正确；最后利用SacI和XbaI双酶切载体pHAN-P450s-P450a和植物表达载体pBI121，将pHAN-P450s-P450a定向连接到线性表达载体pBI121上，构建了RNAi植物表达载体pBI-P450-RNAi，酶切和PCR检测正确。利用DNA冻融法将植物表达载体pBI-P450-RNAi成功转入农杆菌LBA4404，酶切和PCR检测正确，成功制备了用于侵染的农杆菌工程菌液。
3．建立了蓖麻品种“通篦5号”体外再生体系：通过对子叶、胚轴和子叶节三种外植体再生不定芽、再生根和小植株移栽成活等情况综合分析后认为：子叶节为蓖麻品种“通篦5号”离体再生的最佳外植体，子叶节最佳切取时间为子叶未从胚乳中抽出前，胚根和胚轴总长约为3-5cm时。去掉约2/3子叶和全部胚根的呈“”状的子叶节一分为二竖直插入培养基中，筛选的最佳不定芽诱导培养基为1/8MS+8.0mg/L ZT，外植体再生芽频率为75.6%，每个外植体平均不定芽数为2.6个；最佳根分化培养基为1/8MS+2.0mg/LNAA，再生根频率为86.7%，且分化多条较粗壮的主根，每条主根上须根数量也较多，移栽后成活率较高。
4．建立了蓖麻子叶节遗传转化体系：按离体培养时的方式切取“通篦5号”子叶节，在1/8MS+20mg/L AS+8.0mg/L ZT的培养基上预培养2-3d，然后在菌液浓度为OD600=0.8的农杆菌工程菌液LBA4404中侵染10min，侵染后的子叶节放在无菌滤纸上吸取残余的菌液后再接回预培养基上共培养3d后，转接到1/8MS+250mg/L Kan+300mg/L Cef抗性筛选培养基上，大约40d左右完成了不定芽的分化和根的再生。再生小植株移栽成活后通过PCR和Southern blot检测证明为转基因阳性植株。
5．经Northern blot鉴定，RNA干扰的转基因蓖麻P450基因被抑制。转基因蓖麻叶片的过氧化物酶和酯酶同工酶比非转基因蓖麻少了1条谱带。转基因蓖麻表型变化特征为：株高明显降低，叶片变小，叶色加深。初步证明P450基因与蓖麻株高发育密切相关。Western blot蛋白分析结果表明，转基因蓖麻P450蛋白表达量比对照明显减少。
In process of plant cell division and embryonic development, plant hormonebrassinosteroid (BR) and gibberellin (GA) involved in the development of internodesinfluence the plant height. Deletion of BR and GA will show the dwarf phenotype. TheSynthetic and Regulation of hormone BR and GA were affected by many enzymes, most ofwhich belong to the family of cytochrome P450, whose genes have a significant impact on thedevelopment of plant stem. Homology of amino acid sequence was60.98%compared withCytochrome P450from castor and rice, which showed two genes belong to the samesubfamily, and we can speculate that the function of two genes was similar. Two genefragments have been cloned and RNAi expression vector have been constructed successfullyaccording to the nucleotide sequence of cytochrome P450genes from castor which have beenpublished in order to verify functions of cytochrome P450gene of castor and reveal therelationship between the gene and hormone synthesis. Effects of different types andconcentrations cytokinin and auxin on regeneration shoot and root of explants were studiedusing cotyledons, hypocotyls and cotyledonary node of “Tongbi5” as explants, and higherfrequent regeneration vitro system of cotyledonary node of castor was obtained. Transgeniccastor plant was gotten by co-culture of cotyledonary node and Agrobacterium, which hasprovided the foundation for study functions of Cytochrome P450of Castor. The main resultswere as the follows:
1. Two pairs of primers were designed basing on nucleotide sequence of cytochromeP450of castor which was published by Genbank(XM_002525863.1), and two fragments ofP450gene named as P450s and P450a were gotten by PCR, and sequence and comparisonwas carried out after connecting successfully with the cloning vector.The results showed thatlengths of P450s and P450a were523bp and411bp and had98.28%and99.51%sequencesimilar with the original respectively. These results indicated the cloned fragment wascytochrome P450of castor.
2. Recombinant plasmid pMD-P450s and intermedium vector pHANNIBAL weredouble digest by SacI and KpnI and inserted the P450s into Linear intermediate vectorpHANNIBAL, and vector pHAN-P450s was constructed, which was correct by double digestion and PCR. Then plasmid pMD-P450a and vector pHAN-P450s were recombined bydouble digest by XbaI and HandⅢ and inserted the P450a into linear intermediate vectorpHAN-P450s, and pHAN-P450s-P450a vector was constructed, which was correct byenzymes digestion and PCR. At last, recombinant plasmid pHAN-P450s-P450a and plantexpression vector pBI121were double digest by SacI and XbaI and pHAN-P450s-P450a wasinserted into linear intermediate vector pBI121, and RNAi plant expression vectorpBI-P450-RNAi was constructed, which was correct by enzymes digestion and PCR. Theplant expression vector pBI-P450-RNAi was successfully inserted into the AgrobacteriumLBA4404, which was correct by enzymes digestion and PCR, Agrobacterium culture mediumfor infection was successfully gained.
3. Regeneration system of “Tongbi5” was established: after analyzing regenerations ofshoot, root from cotyledons, hypocotyls and cotyledonary node and transplanting survival rateof regeneration plantlets in general, the result was as follow: the optimum regenerationexplants of “Tongbi5” was cotyledonary node and the optimum cutting time was beforecotyledon head emerging from endosperm when radicle and hypocotyl was3-5cm.Cotyledonary node that showed “” after cutting two-thirds of cotyledonary and all radicle,divided it into two parts and inserted into the medium vertically. And the optimum mediumwas1/8MS+8.0mg/L ZT, which regeneration frequency of explants achieved for75.6%andaverage regeneration buds per explants was2.6. The optimum medium for root differentiationwas1/8MS+2.0mg/L NAA, which root regeneration frequency achieved for86.7%andmultistory stocky taproot with many branch roots could be differentiated and hadhigh-survival rate after transplant.
4. The genetic transformation system of castor cotyledonary node was established:cotyledonary node of “Tongbi.5” was cut by patterns of vitro culture, pre-cultured onmedium(1/8MS+20mg/L AS+8.0mg/L ZT) for2-3days, infested10minute withagrobacterium culture medium LBA4404with the concentration of OD600=0.8, thencotyledonary node was co-cultured for3days after the remnants of bacteria was absorbed bysterile filtered paper, transferred cotyledonary node into the resistant screeningmedium(1/8MS+250mg/L Kan+300mg/LCef), it took40days for shoot differentiation and root regeneration. Regeneration plantlets was transplanted and detected by PCR and Southernblot, and proved it was positive transgenic plants.
5. P450gene of transgenic castor interfered by RNA was inhibited through Northernblot. One band of peroxidase and esterase isozyme in transgenic castor leaves was less thanthat of non-transgenic castor. Phenotype characteristic of the transgenic castor were follow:plant height was dwarfed, leaves were smaller, leaf color was dark green. These entirepreliminary proved that the gene of P450was closely related to plant height of castor.Western blot analysis showed that the expression of P450protein of transgenic castordecreased significantly compared with the control
1．根据Genbank（XM_002525863.1）公布的蓖麻细胞色素P450基因的碱基序列设计了两对引物，PCR扩增得到P450基因的两个片段，分别命名为P450s和P450a，与克隆载体成功连接后测序并进行序列比对，结果显示：P450s全长523bp，与原序列一致性为98.28%；P450a全长411bp，与原序列一致性为99.51%，说明克隆的片段是蓖麻细胞色素P450基因片段。
2．利用SacI和KpnI对重组质粒pMD-P450s和中间载体pHANNIBAL双酶切后，将P450s定向连接到线性中间载体pHANNIBAL上，构建了载体pHAN-P450s，酶切和PCR检测正确；然后利用XbaI和HandⅢ双酶切重组质粒pMD-P450a和载体pHAN-P450s，将P450a定向连接到线性载体pHAN-P450s上，构建了载体pHAN-P450s-P450a，酶切和PCR检测正确；最后利用SacI和XbaI双酶切载体pHAN-P450s-P450a和植物表达载体pBI121，将pHAN-P450s-P450a定向连接到线性表达载体pBI121上，构建了RNAi植物表达载体pBI-P450-RNAi，酶切和PCR检测正确。利用DNA冻融法将植物表达载体pBI-P450-RNAi成功转入农杆菌LBA4404，酶切和PCR检测正确，成功制备了用于侵染的农杆菌工程菌液。
3．建立了蓖麻品种“通篦5号”体外再生体系：通过对子叶、胚轴和子叶节三种外植体再生不定芽、再生根和小植株移栽成活等情况综合分析后认为：子叶节为蓖麻品种“通篦5号”离体再生的最佳外植体，子叶节最佳切取时间为子叶未从胚乳中抽出前，胚根和胚轴总长约为3-5cm时。去掉约2/3子叶和全部胚根的呈“”状的子叶节一分为二竖直插入培养基中，筛选的最佳不定芽诱导培养基为1/8MS+8.0mg/L ZT，外植体再生芽频率为75.6%，每个外植体平均不定芽数为2.6个；最佳根分化培养基为1/8MS+2.0mg/LNAA，再生根频率为86.7%，且分化多条较粗壮的主根，每条主根上须根数量也较多，移栽后成活率较高。
4．建立了蓖麻子叶节遗传转化体系：按离体培养时的方式切取“通篦5号”子叶节，在1/8MS+20mg/L AS+8.0mg/L ZT的培养基上预培养2-3d，然后在菌液浓度为OD600=0.8的农杆菌工程菌液LBA4404中侵染10min，侵染后的子叶节放在无菌滤纸上吸取残余的菌液后再接回预培养基上共培养3d后，转接到1/8MS+250mg/L Kan+300mg/L Cef抗性筛选培养基上，大约40d左右完成了不定芽的分化和根的再生。再生小植株移栽成活后通过PCR和Southern blot检测证明为转基因阳性植株。
5．经Northern blot鉴定，RNA干扰的转基因蓖麻P450基因被抑制。转基因蓖麻叶片的过氧化物酶和酯酶同工酶比非转基因蓖麻少了1条谱带。转基因蓖麻表型变化特征为：株高明显降低，叶片变小，叶色加深。初步证明P450基因与蓖麻株高发育密切相关。Western blot蛋白分析结果表明，转基因蓖麻P450蛋白表达量比对照明显减少。
In process of plant cell division and embryonic development, plant hormonebrassinosteroid (BR) and gibberellin (GA) involved in the development of internodesinfluence the plant height. Deletion of BR and GA will show the dwarf phenotype. TheSynthetic and Regulation of hormone BR and GA were affected by many enzymes, most ofwhich belong to the family of cytochrome P450, whose genes have a significant impact on thedevelopment of plant stem. Homology of amino acid sequence was60.98%compared withCytochrome P450from castor and rice, which showed two genes belong to the samesubfamily, and we can speculate that the function of two genes was similar. Two genefragments have been cloned and RNAi expression vector have been constructed successfullyaccording to the nucleotide sequence of cytochrome P450genes from castor which have beenpublished in order to verify functions of cytochrome P450gene of castor and reveal therelationship between the gene and hormone synthesis. Effects of different types andconcentrations cytokinin and auxin on regeneration shoot and root of explants were studiedusing cotyledons, hypocotyls and cotyledonary node of “Tongbi5” as explants, and higherfrequent regeneration vitro system of cotyledonary node of castor was obtained. Transgeniccastor plant was gotten by co-culture of cotyledonary node and Agrobacterium, which hasprovided the foundation for study functions of Cytochrome P450of Castor. The main resultswere as the follows:
1. Two pairs of primers were designed basing on nucleotide sequence of cytochromeP450of castor which was published by Genbank(XM_002525863.1), and two fragments ofP450gene named as P450s and P450a were gotten by PCR, and sequence and comparisonwas carried out after connecting successfully with the cloning vector.The results showed thatlengths of P450s and P450a were523bp and411bp and had98.28%and99.51%sequencesimilar with the original respectively. These results indicated the cloned fragment wascytochrome P450of castor.
2. Recombinant plasmid pMD-P450s and intermedium vector pHANNIBAL weredouble digest by SacI and KpnI and inserted the P450s into Linear intermediate vectorpHANNIBAL, and vector pHAN-P450s was constructed, which was correct by double digestion and PCR. Then plasmid pMD-P450a and vector pHAN-P450s were recombined bydouble digest by XbaI and HandⅢ and inserted the P450a into linear intermediate vectorpHAN-P450s, and pHAN-P450s-P450a vector was constructed, which was correct byenzymes digestion and PCR. At last, recombinant plasmid pHAN-P450s-P450a and plantexpression vector pBI121were double digest by SacI and XbaI and pHAN-P450s-P450a wasinserted into linear intermediate vector pBI121, and RNAi plant expression vectorpBI-P450-RNAi was constructed, which was correct by enzymes digestion and PCR. Theplant expression vector pBI-P450-RNAi was successfully inserted into the AgrobacteriumLBA4404, which was correct by enzymes digestion and PCR, Agrobacterium culture mediumfor infection was successfully gained.
3. Regeneration system of “Tongbi5” was established: after analyzing regenerations ofshoot, root from cotyledons, hypocotyls and cotyledonary node and transplanting survival rateof regeneration plantlets in general, the result was as follow: the optimum regenerationexplants of “Tongbi5” was cotyledonary node and the optimum cutting time was beforecotyledon head emerging from endosperm when radicle and hypocotyl was3-5cm.Cotyledonary node that showed “” after cutting two-thirds of cotyledonary and all radicle,divided it into two parts and inserted into the medium vertically. And the optimum mediumwas1/8MS+8.0mg/L ZT, which regeneration frequency of explants achieved for75.6%andaverage regeneration buds per explants was2.6. The optimum medium for root differentiationwas1/8MS+2.0mg/L NAA, which root regeneration frequency achieved for86.7%andmultistory stocky taproot with many branch roots could be differentiated and hadhigh-survival rate after transplant.
4. The genetic transformation system of castor cotyledonary node was established:cotyledonary node of “Tongbi.5” was cut by patterns of vitro culture, pre-cultured onmedium(1/8MS+20mg/L AS+8.0mg/L ZT) for2-3days, infested10minute withagrobacterium culture medium LBA4404with the concentration of OD600=0.8, thencotyledonary node was co-cultured for3days after the remnants of bacteria was absorbed bysterile filtered paper, transferred cotyledonary node into the resistant screeningmedium(1/8MS+250mg/L Kan+300mg/LCef), it took40days for shoot differentiation and root regeneration. Regeneration plantlets was transplanted and detected by PCR and Southernblot, and proved it was positive transgenic plants.
5. P450gene of transgenic castor interfered by RNA was inhibited through Northernblot. One band of peroxidase and esterase isozyme in transgenic castor leaves was less thanthat of non-transgenic castor. Phenotype characteristic of the transgenic castor were follow:plant height was dwarfed, leaves were smaller, leaf color was dark green. These entirepreliminary proved that the gene of P450was closely related to plant height of castor.Western blot analysis showed that the expression of P450protein of transgenic castordecreased significantly compared with the control
引文
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