农杆菌介导的枣树遗传转化研究
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摘要
枣树(Ziziphus jujuba Mill.)是我国原产重要果树。枣树由于花小、人工去雄难、坐果率低且胚败育现象严重,使得其杂交育种非常困难。枣疯病是由植原体引起致死性传染病害,几乎分布于国内外所有枣区,绝大多数枣树品种都对其敏感,已成为制约枣产业发展的严重障碍。随着生物技术的不断发展,基因工程已成为现代植物改良育种的有效手段。本研究旨在探索建立农杆菌介导的枣树遗传转化体系,并利用发根农杆菌提高枣树对枣疯病的抗性,为枣树遗传改良和枣疯病防治提供技术支撑。主要研究结果如下:
1.建立了冬枣的花药再生体系。诱导花药愈伤的适宜培养基为1/2MS+1.0mg/L 2,4-D+20g/L麦芽糖+5.5g/L琼脂;花药愈伤的适宜增殖培养基为MS+0.4mg/L TDZ+0.1mg/L NAA +20g/L麦芽糖+5.5g/L琼脂;1.0mg/L TDZ和0.4mg/L NAA的组合是诱导冬枣花药愈伤出芽的适宜生长调节剂组合,诱导出芽率达100%,平均出芽数为3.9个/块;0.2mg/L BA、0.1mg/L IBA和1.5g/L PVA是玻璃苗恢复的适宜组合,玻璃苗恢复率为48.9%;在添加1.5mg/L IAA的生根培养基中,生根率最高(43.3%),根长最长(4.2cm)。
2.首次建立了月光枣的花药再生体系。诱导花药愈伤的适宜培养基为1/2MS+1.0mg/L 2,4-D+20g/L麦芽糖+5.5g/L琼脂;月光花药愈伤的适宜增殖培养基为MS+0.6mg/L TDZ+0.1mg/L NAA+20g/L麦芽糖+5.5g/L琼脂。适宜月光愈伤分化出芽的培养基为:MS+5.5g/L琼脂+20g/L麦芽糖+0.8mg/L TDZ+4.0mg/L AgNO3。其诱导出芽率为100%,平均出芽数为10.4个/块;在培养基MS+5.5g/L琼脂+30g/L蔗糖+0.2mg/L BA+0.1mg/L IBA+1.5g/L PVA中玻璃苗可恢复成正常苗;添加2.0mg/L IBA的生根培养基中月光生根率最高;GA3的添加对于枣愈伤的生长和玻璃苗的恢复均为抑制作用。
3.建立了根癌农杆菌介导的冬枣叶片遗传转化体系。适宜的遗传转化条件为:根癌农杆菌的OD600为0.8,侵染20min后,在共培养基MS+5.5g/L琼脂+20g/L麦芽糖+1.0mg/L TDZ+0.1mg/L IBA+200μM AS(pH5.8)中暗培养3d。首次将双抗虫基因(Bt基因和蛋白抑制剂基因API)转入冬枣中,得到了8个转基因株系。
4.建立了冬枣叶片及冬枣疯组培苗的发根农杆菌介导的遗传转化体系。冬枣叶片诱导毛根的适宜条件是在菌液浓度OD600为0.8时侵染叶片20min,在共培养基1/2MS+4.0mg/L IBA+5.5g/L琼脂+30g/L蔗糖(pH5.8)中暗培养3-5d,然后转接入培养基1/2MS0+CRO100mg/L中,光照条件为光照/黑暗为14/10h。冬枣疯组培苗茎段诱导出毛根的适宜条件为:在菌液中添加50μM的AS预活化1~2h,OD600为0.8侵染5~10min,共培养5d,共培养基为MS+5.5g/L琼脂+20g/L蔗糖+100mg/L CRO,光照条件为光照/黑暗为14/10h。首次通过发根农杆菌侵染使冬枣疯组培苗转化为正常植株,得到了30个抗病性提高的转健苗株系。
Chinese jujube (Ziziphus jujuba Mill.) is an important native fruit tree of China. Conventional croos is very difficult in Chinese jujube for reasons of very low fruit-setting, very high embryo abortion as well as very small flower. With the development of biotechnology, genetic engineering has become an effective means for modern plant breeding improvement.This study is aimed at to establishing an Agrobacterium-mediated genetic transformation system and laying a foundation for genetic engineering in Chinese jujube. The main results of the study are as follows:
1. An optimized regeneration system for anther wall of Z. Jujuba Mill.‘Dongzao’was established. The optimal medium for inducing callus from anther is 1/2MS+1.0mg/L 2,4-D+20g/L maltose+5.5g/L agar. The optimal medium of callus proliferation is MS+0.4mg/L TDZ+0.2mg/L NAA+20g/L maltose+5.5g/L agar. The combination of 1.0 mg/L TDZ and 0.4 mg/L NAA is the best one to shoots induction from callus with the highest shoot induction rate (100%) and a high number of shoots per callus (3.9). The combination of 0.2 mg/L BA, 0.1 mg/L IBA and 1.5 mg/L PVA promoted the highest rate of hyperhydric shoots restoration (48.9%).The highest rooting (43.3%), and highest root length (4.2 cm) was recorded at 1.5 mg /l IAA.
2. A high efficient regeneration system using anther wall of Z. Jujuba Mill.‘Yueguang’was firstly established. The optimal medium for inducing callus for anther wall was 1/2MS+1.0mg/L 2,4-D+20g/L maltose+5.5g/L agar. The optimal medium of callus proliferation is MS+0.6mg/L TDZ+0.2mg/L NAA+20g/L maltose+5.5g/L agar. The optimal differentiation medium of callus was MS+5.5g/L agar+20g/L maltose+0.8mg/L TDZ+4.0mg/L AgNO3with the highest shoot induction response (100%) and highest number of shoots per callus (10.4). The combination of 0.2 mg/L BA, 0.1 mg/L IBA and 1.5 mg/L PVA promoted the highest rate of hyperhydric shoots restoration. The highest rooting (43.3%), and highest root length (4.2 cm) was recorded at 2.0 mg /L IBA. The addition of GA3 inhabited the growth of callus and hyperhydric shoots restoration.
3.The genetic transgenic system of‘Dongzao’leaf mediated by Agrobacterium tumefacienswas investigated and the optimal conditions were determined as follows: bacterial concentration OD600 0.8, infection time 20 min, co-culture 3 days and the co-culture medium MS+20g/L maltose+5.5g/L agar+1.0mg/L TDZ+0.1mg/L IBA+200μΜAS.
Two insect-resistant genes (Bt gene and protease inhibitor gene, API) were firstly into‘Dongzao’and . 8 transgenic lines have been obtained.
4. The genetic transgenic system of Z. Jujuba Mill.‘Dongzao’and‘Dongzao’with witches’broom disease (JWB) mediated by Agrobacterium rhizogenes were firstly established. For‘Dongzao’leaf, the optimal conditions were determined as follows: bacterial concentration OD600 0.8, infection time 20 min, co-culture 3-5 days, the co-culture medium was 1/2MS+30g/L sugar+5.5g/L agar+4.0mg/L IBA and the medium of inducing hairy root was 1/2MS+30g/L sugar+5.5g/L agar+100mg/L CRO under a 14-h light and 10-h dark cycle. For stems of diseased‘Dongzao’, the optimal conditions were determined as follows: bacterial concentration OD600 0.8 with activation of 50μΜAS for 1-2h before infection, infection time 5-10 min, co-culture 5 days, the co-culture medium was MS+30g/L sugar+5.5g/L agar and the medium of inducing hairy root was MS+30g/L sugar+5.5g/L agar+100mg/L CRO under under a 14-h light and 10-h dark cycle. This is the first time to thansform Chinese jujube with JWB into healthy one via Agrobacterium rhizogenesinfection. 30 recovering plantlet lines have been obtained.
1.建立了冬枣的花药再生体系。诱导花药愈伤的适宜培养基为1/2MS+1.0mg/L 2,4-D+20g/L麦芽糖+5.5g/L琼脂;花药愈伤的适宜增殖培养基为MS+0.4mg/L TDZ+0.1mg/L NAA +20g/L麦芽糖+5.5g/L琼脂;1.0mg/L TDZ和0.4mg/L NAA的组合是诱导冬枣花药愈伤出芽的适宜生长调节剂组合,诱导出芽率达100%,平均出芽数为3.9个/块;0.2mg/L BA、0.1mg/L IBA和1.5g/L PVA是玻璃苗恢复的适宜组合,玻璃苗恢复率为48.9%;在添加1.5mg/L IAA的生根培养基中,生根率最高(43.3%),根长最长(4.2cm)。
2.首次建立了月光枣的花药再生体系。诱导花药愈伤的适宜培养基为1/2MS+1.0mg/L 2,4-D+20g/L麦芽糖+5.5g/L琼脂;月光花药愈伤的适宜增殖培养基为MS+0.6mg/L TDZ+0.1mg/L NAA+20g/L麦芽糖+5.5g/L琼脂。适宜月光愈伤分化出芽的培养基为:MS+5.5g/L琼脂+20g/L麦芽糖+0.8mg/L TDZ+4.0mg/L AgNO3。其诱导出芽率为100%,平均出芽数为10.4个/块;在培养基MS+5.5g/L琼脂+30g/L蔗糖+0.2mg/L BA+0.1mg/L IBA+1.5g/L PVA中玻璃苗可恢复成正常苗;添加2.0mg/L IBA的生根培养基中月光生根率最高;GA3的添加对于枣愈伤的生长和玻璃苗的恢复均为抑制作用。
3.建立了根癌农杆菌介导的冬枣叶片遗传转化体系。适宜的遗传转化条件为:根癌农杆菌的OD600为0.8,侵染20min后,在共培养基MS+5.5g/L琼脂+20g/L麦芽糖+1.0mg/L TDZ+0.1mg/L IBA+200μM AS(pH5.8)中暗培养3d。首次将双抗虫基因(Bt基因和蛋白抑制剂基因API)转入冬枣中,得到了8个转基因株系。
4.建立了冬枣叶片及冬枣疯组培苗的发根农杆菌介导的遗传转化体系。冬枣叶片诱导毛根的适宜条件是在菌液浓度OD600为0.8时侵染叶片20min,在共培养基1/2MS+4.0mg/L IBA+5.5g/L琼脂+30g/L蔗糖(pH5.8)中暗培养3-5d,然后转接入培养基1/2MS0+CRO100mg/L中,光照条件为光照/黑暗为14/10h。冬枣疯组培苗茎段诱导出毛根的适宜条件为:在菌液中添加50μM的AS预活化1~2h,OD600为0.8侵染5~10min,共培养5d,共培养基为MS+5.5g/L琼脂+20g/L蔗糖+100mg/L CRO,光照条件为光照/黑暗为14/10h。首次通过发根农杆菌侵染使冬枣疯组培苗转化为正常植株,得到了30个抗病性提高的转健苗株系。
Chinese jujube (Ziziphus jujuba Mill.) is an important native fruit tree of China. Conventional croos is very difficult in Chinese jujube for reasons of very low fruit-setting, very high embryo abortion as well as very small flower. With the development of biotechnology, genetic engineering has become an effective means for modern plant breeding improvement.This study is aimed at to establishing an Agrobacterium-mediated genetic transformation system and laying a foundation for genetic engineering in Chinese jujube. The main results of the study are as follows:
1. An optimized regeneration system for anther wall of Z. Jujuba Mill.‘Dongzao’was established. The optimal medium for inducing callus from anther is 1/2MS+1.0mg/L 2,4-D+20g/L maltose+5.5g/L agar. The optimal medium of callus proliferation is MS+0.4mg/L TDZ+0.2mg/L NAA+20g/L maltose+5.5g/L agar. The combination of 1.0 mg/L TDZ and 0.4 mg/L NAA is the best one to shoots induction from callus with the highest shoot induction rate (100%) and a high number of shoots per callus (3.9). The combination of 0.2 mg/L BA, 0.1 mg/L IBA and 1.5 mg/L PVA promoted the highest rate of hyperhydric shoots restoration (48.9%).The highest rooting (43.3%), and highest root length (4.2 cm) was recorded at 1.5 mg /l IAA.
2. A high efficient regeneration system using anther wall of Z. Jujuba Mill.‘Yueguang’was firstly established. The optimal medium for inducing callus for anther wall was 1/2MS+1.0mg/L 2,4-D+20g/L maltose+5.5g/L agar. The optimal medium of callus proliferation is MS+0.6mg/L TDZ+0.2mg/L NAA+20g/L maltose+5.5g/L agar. The optimal differentiation medium of callus was MS+5.5g/L agar+20g/L maltose+0.8mg/L TDZ+4.0mg/L AgNO3with the highest shoot induction response (100%) and highest number of shoots per callus (10.4). The combination of 0.2 mg/L BA, 0.1 mg/L IBA and 1.5 mg/L PVA promoted the highest rate of hyperhydric shoots restoration. The highest rooting (43.3%), and highest root length (4.2 cm) was recorded at 2.0 mg /L IBA. The addition of GA3 inhabited the growth of callus and hyperhydric shoots restoration.
3.The genetic transgenic system of‘Dongzao’leaf mediated by Agrobacterium tumefacienswas investigated and the optimal conditions were determined as follows: bacterial concentration OD600 0.8, infection time 20 min, co-culture 3 days and the co-culture medium MS+20g/L maltose+5.5g/L agar+1.0mg/L TDZ+0.1mg/L IBA+200μΜAS.
Two insect-resistant genes (Bt gene and protease inhibitor gene, API) were firstly into‘Dongzao’and . 8 transgenic lines have been obtained.
4. The genetic transgenic system of Z. Jujuba Mill.‘Dongzao’and‘Dongzao’with witches’broom disease (JWB) mediated by Agrobacterium rhizogenes were firstly established. For‘Dongzao’leaf, the optimal conditions were determined as follows: bacterial concentration OD600 0.8, infection time 20 min, co-culture 3-5 days, the co-culture medium was 1/2MS+30g/L sugar+5.5g/L agar+4.0mg/L IBA and the medium of inducing hairy root was 1/2MS+30g/L sugar+5.5g/L agar+100mg/L CRO under a 14-h light and 10-h dark cycle. For stems of diseased‘Dongzao’, the optimal conditions were determined as follows: bacterial concentration OD600 0.8 with activation of 50μΜAS for 1-2h before infection, infection time 5-10 min, co-culture 5 days, the co-culture medium was MS+30g/L sugar+5.5g/L agar and the medium of inducing hairy root was MS+30g/L sugar+5.5g/L agar+100mg/L CRO under under a 14-h light and 10-h dark cycle. This is the first time to thansform Chinese jujube with JWB into healthy one via Agrobacterium rhizogenesinfection. 30 recovering plantlet lines have been obtained.
引文
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