betA基因的异源表达提高了棉花耐盐和耐低温能力
摘要
甘氨酸甜菜碱(GB)是一种重要的渗透保护物质,在低温、高盐和干旱等逆境胁迫下,许多植物积累甘氨酸甜菜碱作为渗透保护物质以维持细胞正常的渗透压,稳定酶和蛋白复合物的结构和功能,保持膜的完整性。来自大肠杆菌的betA基因编码的胆碱脱氢酶(CDH)兼具有胆碱单氧化物酶和甜菜碱醛脱氢酶的功能,转移单一的betA基因即可赋予植物积累甘氨酸甜菜碱的能力。
生产中低温和盐渍是限制棉花产量和质量的重要因素之一。提高棉花的耐盐、耐低温能力,对于提高棉花产量,拓宽植棉区域具有重要意义。工作以本实验室得到的转betA基因鲁棉研19的T3代棉花为材料,以鲁棉研19为野生型对照,通过对转基因植株后代的耐盐和耐低温特性进行了分析,结果表明转基因棉花的耐盐和耐低温能力都有明显提高。
betA基因的表达提高了转基因棉花叶片中GB的累积量
betA基因的过表达显著提高了转基棉花叶片中GB的含量。利用核磁共振法测定了转基因株系LM1、LM2、LM3、LM4及野生型对照鲁棉研19植株叶片中GB的含量。在正常生长状态下,鲁棉研19植株叶片中GB含量约为99.29μmolg~(-1)DW,转基因株系GB的含量在125.98-214.34μmolg~(-1)DW之间。250mMNaCl处理10天后,棉花植株叶片的GB含量大幅提高,株系LM1、LM2、LM3和LM4植株叶片甜菜碱含量分别为321.06、357.32、355.51和276.09μmolg~(-1)DW,显著高于对照株系176.00μmolg~(-1)DW的GB含量。5℃低温处理24h后,株系LM1、LM2、LM3和LM4植株的GB含量分别为170.78、205.93、248.21和189.46μmolg~(-1)DW,分别是野生型对照109.37μmolg~(-1)DW GB含量的1.56、1.88、2.27和1.73倍,转基因棉花植株叶片的GB含量显著高于野生型对照植株。转基因株系中甜菜碱含量的升高可能是由于高盐、低温胁迫后betA基因的表达水平增强引起的。
转betA基因棉花的耐盐性分析
在250mM NaCl浇灌条件下,转基因株系LM1、LM2、LM3、LM4第9天的种子萌发率分别为52.04%、66.04%、60.44%和42.40%,均显著高于野生型28.28%的萌发率。株系LM2、LM3生物量也显著高于野生型对照。250mM NaCl浇灌21天后,野生型对照植株叶片的相对含水量由80.79%降至48.54%,株系LM1、LM2、LM3、LM4叶片的相对含水量分别为62.90%、64.93%、64.44%和63.53%,转基因植株叶片失水速度比对照缓慢。转基因棉花叶片中渗透保护物质可溶性糖、游离氨基酸含量均高于野生型对照,株系LM1、LM2、LM3、LM4的细胞溶质势较野生型对照分别低8.86%、13.89%、3.50%和8.61%,即转基因棉花细胞中积累了更多的溶质,增强了细胞吸取水分和持水能力,从而有利于植株保持正常形态和细胞膨压。株系LM2、LM3细胞膜离子渗漏率分别比野生型对照植株低23.08%和23.56%,叶片丙二醛含量也比野生型对照植株低38.72%和36.03%,说明转基因植株在盐胁迫条件下细胞膜损伤和膜脂过氧化程度都较轻。转基因株系LM2、LM3的净光合速率、气孔导度和蒸腾速率都明显高于野生型对照,转基因棉花的光合作用系统受到渗透胁迫的影响较野生型对照为轻。在含盐量为0.45%的盐碱地中,转基因株系LM2、LM3的单株籽棉产量远远高于野生型对照,这为培育耐盐棉花新品种提供了优良材料,对于棉花育种具有重要意义。
转betA基因棉花的耐低温性分析
在15℃条件下,转基因株系LM1、LM2、LM3、LM4第13天的种子萌发率分别为67.33%、78.20%、70.11%和74.25%,均显著高于野生型53.24%的萌发率。5℃低温处理24h,野生型对照植株叶片的相对含水量由87.94%降至53.43%,转基因株系LM3的叶片相对含水量为66.28%,显著高于对照株系,转基因植株叶片持水能力比对照强。转基因株系LM1、LM2、LM3、LM4的离子渗漏率分别比对野生型对照低22.59%、37.95%、40.28%、23.11%,其中株系LM2、LM3的细胞膜离子渗漏率显著低于野生型对照植株,叶片丙二醛含量也比野生型对照植株低32.96%和39.65%,说明低温条件对转基因株系棉花叶片细胞膜的损害相对较轻,耐低温能力较强。株系LM2、LM3的细胞保护酶SOD的活性比野生型对照分别高4.41%和6.35%,说明转基因植株在低温胁迫条件下,体内保护酶活性处于较高水平,具有较强清除体内自由基的能力。低温胁迫后转基因株系的净光合速率、气孔导度、胞间CO_2浓度和蒸腾速率都明显高于野生型对照,转基因株系LM2、LM3的净光合速率分别比野生型对照植株高13.81%和31.61%,转基因植株与野生型对照相比,其光合系统受低温的伤害较轻。
因此,本研究表明来自大肠杆菌的betA基因在棉花中的异源表达可以提高转基因棉花植株中甘氨酸甜菜碱的含量,转基因棉花LM2、LM3的抗逆性有不同程度的提高,为棉花抗逆育种提供了新材料。
Glycinebetaine(GB)is important one kind of compatible osmolyte.Some plant accumulated the osmoprotectant GB to maintain the cell normal turgor,the structure and function of enzyme,protein complex,and the integrity of membrane responding the stress such as low temperature,salt and drought.The betA gene from E.coli. encodes chroline dehydrogenase(CDH)which has both the function of Choline Monooxygenase(CMO)and betaine aldehy dedehydrogenase(BADH). Transformation of the single gene betA can endow the transgenic plants the capability of accumulating GB.
Low temperature and soil salinization are important factors affecting the quality and yield of cotton.There is important significance to improve the salt and low temperature tolerance of cotton for increasing the yield and plant area of cotton.The tolerance to salt and low temperature stress of four T3 progeny of transgenic lines from Lumianyanl9 was investigated.The result shows that the tolerance of transgenic plants improved significantly.
The expression of betA increased the accumulation of GB in transgenic cotton
The expression of betA increases the level of GB in transgenic cotton leaves significantly.The level of GB in the leaves of transgenic lines LM1,LM2,LM3,LM4 and wild-type Lumianyanl9 are detected by NMR method.The concentration of GB in the leaves of wild-type was 99.29μmolg~(-1)DW,the level of GB in the transgenic lines were between 125.98-214.34μmolg~(-1)DW under normal growing condition.After ten days treated by 250mM NaCl,the concentration of GB increased significantly,the four transgenic lines accumulated 321.06,357.32,355.51 and 276.09μmolg~(-1)DW respectively,significantly higher levels of GB than the wild-type plant with 176.00μmolg~(-1)DW GB.After 24 hours' treatment with 5℃,the GB content in transgenic lines LM1,LM2,LM3 and LM4 reached 170.78,205.93,248.21 and 189.46μmolg~(-1)DW,they were 1.56,1.88,2.27 and 1.73 times compared to that in wild-type plants.It is possible that significant increase of the level of GB in transgenic lines resulted from the enhanced expression of betA under the stress of salt or low temperature.
Analysis of salt tolerance of betA-expressing cotton
The germination rates of the transgenic lines LM1,LM2,LM3 and LM4 treated by 250mM NaCl for 9 days are 52.04%,66.04%,60.44%and 42.40%respectively, each of which is significantly higher than that of the wild-type Lumianyan19.The biomass of the lines LM2,LM3 is significantly higher than that of the wild-type.After 21 days salt stress under 250mMNaCl,the relative water content of the wild-type plants reduced from 80.79%to 48.54%,whereas that of the four transgenic lines are 62.90%,64.93%,64.44%and 63.53%,the water loss rates of the transgenic lines more slowly than that of the wild-type plants.The content of other osmoprotectant such as soluble sugar,free amino acids in the leaves of the transgenic cotton are obvious higher than those in wild-type.The solute potential of the lines LM1,LM2, LM3,LM4 is 8.86%,13.89%,3.50%and 8.61%lower than that of the wild-type.The results indicated that more solutes were accumulated in transgenic plants to enable cells to absorb more water and to maintain cell normal shape and turgor under stress. The cell membrane ion leakage,the content of malondialdehyde of lines LM2,LM3 is 23.08%,23.56%and 38.72%,36.03%lower than that of the wild-type respectively, which suggested less membrane damage and lipid peroxidation occurred in these transgenic lines.After osmotic stress,net photosynthesis,stomatal conductance and transpiration rate of the four transgenic lines were significantly higher than those of wild-type,the effect to the Photosynthesis System of transgenic cottons is less than that of the wild-type.The seed cotton yield per plant of the lines LM2,LM3 is significantly higher than that of the wild-type on the salinized land(salt content 0.45%),which were of great value in cotton breeding and production.
Analysis of low temperature tolerance of betA-expressing cotton
The germination rates of the lines LM1,LM2,LM3 and LM4 at the 13~(th)day after seeding at 15℃condition are 67.33%,78.20%,70.11%and 74.25%,each of which is significantly higher than 53.24%which is the germination rate of the wild-type Lumianyanl9.After 24h low temperature treatment at 5℃,the relative water content of the wild-type plants reduced from 87.94%to 53.43%,whereas that of the transgenic line LM3 is 66.28%,which is significantly higher than that of the wild-type plants,the water-retaining capability of the transgenic lines leaves is better than that of the wild-type plants.The cell membrane ion leakage of the four transgenic lines LM1,LM2,LM3,LM4 is 22.59%,37.95%,40.28%and 23.11%respectively, which is lower than that of the wild-type,the content of malondialdehyde of the transgenic lines LM2,LM3 is 32.96%and 39.65%,which is less than that of the wild-type.The results suggested that less membrane damage occurred in these transgenic lines which have stronger low temperature tolerance.The SOD activity in these transgenic lines is also higher than that of wild-type plants under stress.The results implied that the transgenic plant has strong activity of cell protective enzyme, powerful ability to scavenging free radicals under low temperature stress.After low temperature stress,net photosynthesis,stomatal conductance,intercellular CO_2 concentration and transpiration rate of the four transgenic lines were significantly higher than those of wild-type plants,the net photosynthesis of the LM2 and LM3 is 13.81%and 31.61%higher than that of the wild-type plants,the damage to the Photosynthesis System of transgenic cottons by low temperature stress is less than that of the wild-type.
The above physiological analysis of cotton plants transformed with betA gene demonstrated that a proportion of the transgenic lines(LM2,LM3)accumulated higher levels of glycinebetaine and acquired significantly higher tolerance to salt and low temperature stresses.In addition,the transgenic lines with improved stress tolerance provided valuable cotton materials for cotton stress tolerance breeding.
生产中低温和盐渍是限制棉花产量和质量的重要因素之一。提高棉花的耐盐、耐低温能力,对于提高棉花产量,拓宽植棉区域具有重要意义。工作以本实验室得到的转betA基因鲁棉研19的T3代棉花为材料,以鲁棉研19为野生型对照,通过对转基因植株后代的耐盐和耐低温特性进行了分析,结果表明转基因棉花的耐盐和耐低温能力都有明显提高。
betA基因的表达提高了转基因棉花叶片中GB的累积量
betA基因的过表达显著提高了转基棉花叶片中GB的含量。利用核磁共振法测定了转基因株系LM1、LM2、LM3、LM4及野生型对照鲁棉研19植株叶片中GB的含量。在正常生长状态下,鲁棉研19植株叶片中GB含量约为99.29μmolg~(-1)DW,转基因株系GB的含量在125.98-214.34μmolg~(-1)DW之间。250mMNaCl处理10天后,棉花植株叶片的GB含量大幅提高,株系LM1、LM2、LM3和LM4植株叶片甜菜碱含量分别为321.06、357.32、355.51和276.09μmolg~(-1)DW,显著高于对照株系176.00μmolg~(-1)DW的GB含量。5℃低温处理24h后,株系LM1、LM2、LM3和LM4植株的GB含量分别为170.78、205.93、248.21和189.46μmolg~(-1)DW,分别是野生型对照109.37μmolg~(-1)DW GB含量的1.56、1.88、2.27和1.73倍,转基因棉花植株叶片的GB含量显著高于野生型对照植株。转基因株系中甜菜碱含量的升高可能是由于高盐、低温胁迫后betA基因的表达水平增强引起的。
转betA基因棉花的耐盐性分析
在250mM NaCl浇灌条件下,转基因株系LM1、LM2、LM3、LM4第9天的种子萌发率分别为52.04%、66.04%、60.44%和42.40%,均显著高于野生型28.28%的萌发率。株系LM2、LM3生物量也显著高于野生型对照。250mM NaCl浇灌21天后,野生型对照植株叶片的相对含水量由80.79%降至48.54%,株系LM1、LM2、LM3、LM4叶片的相对含水量分别为62.90%、64.93%、64.44%和63.53%,转基因植株叶片失水速度比对照缓慢。转基因棉花叶片中渗透保护物质可溶性糖、游离氨基酸含量均高于野生型对照,株系LM1、LM2、LM3、LM4的细胞溶质势较野生型对照分别低8.86%、13.89%、3.50%和8.61%,即转基因棉花细胞中积累了更多的溶质,增强了细胞吸取水分和持水能力,从而有利于植株保持正常形态和细胞膨压。株系LM2、LM3细胞膜离子渗漏率分别比野生型对照植株低23.08%和23.56%,叶片丙二醛含量也比野生型对照植株低38.72%和36.03%,说明转基因植株在盐胁迫条件下细胞膜损伤和膜脂过氧化程度都较轻。转基因株系LM2、LM3的净光合速率、气孔导度和蒸腾速率都明显高于野生型对照,转基因棉花的光合作用系统受到渗透胁迫的影响较野生型对照为轻。在含盐量为0.45%的盐碱地中,转基因株系LM2、LM3的单株籽棉产量远远高于野生型对照,这为培育耐盐棉花新品种提供了优良材料,对于棉花育种具有重要意义。
转betA基因棉花的耐低温性分析
在15℃条件下,转基因株系LM1、LM2、LM3、LM4第13天的种子萌发率分别为67.33%、78.20%、70.11%和74.25%,均显著高于野生型53.24%的萌发率。5℃低温处理24h,野生型对照植株叶片的相对含水量由87.94%降至53.43%,转基因株系LM3的叶片相对含水量为66.28%,显著高于对照株系,转基因植株叶片持水能力比对照强。转基因株系LM1、LM2、LM3、LM4的离子渗漏率分别比对野生型对照低22.59%、37.95%、40.28%、23.11%,其中株系LM2、LM3的细胞膜离子渗漏率显著低于野生型对照植株,叶片丙二醛含量也比野生型对照植株低32.96%和39.65%,说明低温条件对转基因株系棉花叶片细胞膜的损害相对较轻,耐低温能力较强。株系LM2、LM3的细胞保护酶SOD的活性比野生型对照分别高4.41%和6.35%,说明转基因植株在低温胁迫条件下,体内保护酶活性处于较高水平,具有较强清除体内自由基的能力。低温胁迫后转基因株系的净光合速率、气孔导度、胞间CO_2浓度和蒸腾速率都明显高于野生型对照,转基因株系LM2、LM3的净光合速率分别比野生型对照植株高13.81%和31.61%,转基因植株与野生型对照相比,其光合系统受低温的伤害较轻。
因此,本研究表明来自大肠杆菌的betA基因在棉花中的异源表达可以提高转基因棉花植株中甘氨酸甜菜碱的含量,转基因棉花LM2、LM3的抗逆性有不同程度的提高,为棉花抗逆育种提供了新材料。
Glycinebetaine(GB)is important one kind of compatible osmolyte.Some plant accumulated the osmoprotectant GB to maintain the cell normal turgor,the structure and function of enzyme,protein complex,and the integrity of membrane responding the stress such as low temperature,salt and drought.The betA gene from E.coli. encodes chroline dehydrogenase(CDH)which has both the function of Choline Monooxygenase(CMO)and betaine aldehy dedehydrogenase(BADH). Transformation of the single gene betA can endow the transgenic plants the capability of accumulating GB.
Low temperature and soil salinization are important factors affecting the quality and yield of cotton.There is important significance to improve the salt and low temperature tolerance of cotton for increasing the yield and plant area of cotton.The tolerance to salt and low temperature stress of four T3 progeny of transgenic lines from Lumianyanl9 was investigated.The result shows that the tolerance of transgenic plants improved significantly.
The expression of betA increased the accumulation of GB in transgenic cotton
The expression of betA increases the level of GB in transgenic cotton leaves significantly.The level of GB in the leaves of transgenic lines LM1,LM2,LM3,LM4 and wild-type Lumianyanl9 are detected by NMR method.The concentration of GB in the leaves of wild-type was 99.29μmolg~(-1)DW,the level of GB in the transgenic lines were between 125.98-214.34μmolg~(-1)DW under normal growing condition.After ten days treated by 250mM NaCl,the concentration of GB increased significantly,the four transgenic lines accumulated 321.06,357.32,355.51 and 276.09μmolg~(-1)DW respectively,significantly higher levels of GB than the wild-type plant with 176.00μmolg~(-1)DW GB.After 24 hours' treatment with 5℃,the GB content in transgenic lines LM1,LM2,LM3 and LM4 reached 170.78,205.93,248.21 and 189.46μmolg~(-1)DW,they were 1.56,1.88,2.27 and 1.73 times compared to that in wild-type plants.It is possible that significant increase of the level of GB in transgenic lines resulted from the enhanced expression of betA under the stress of salt or low temperature.
Analysis of salt tolerance of betA-expressing cotton
The germination rates of the transgenic lines LM1,LM2,LM3 and LM4 treated by 250mM NaCl for 9 days are 52.04%,66.04%,60.44%and 42.40%respectively, each of which is significantly higher than that of the wild-type Lumianyan19.The biomass of the lines LM2,LM3 is significantly higher than that of the wild-type.After 21 days salt stress under 250mMNaCl,the relative water content of the wild-type plants reduced from 80.79%to 48.54%,whereas that of the four transgenic lines are 62.90%,64.93%,64.44%and 63.53%,the water loss rates of the transgenic lines more slowly than that of the wild-type plants.The content of other osmoprotectant such as soluble sugar,free amino acids in the leaves of the transgenic cotton are obvious higher than those in wild-type.The solute potential of the lines LM1,LM2, LM3,LM4 is 8.86%,13.89%,3.50%and 8.61%lower than that of the wild-type.The results indicated that more solutes were accumulated in transgenic plants to enable cells to absorb more water and to maintain cell normal shape and turgor under stress. The cell membrane ion leakage,the content of malondialdehyde of lines LM2,LM3 is 23.08%,23.56%and 38.72%,36.03%lower than that of the wild-type respectively, which suggested less membrane damage and lipid peroxidation occurred in these transgenic lines.After osmotic stress,net photosynthesis,stomatal conductance and transpiration rate of the four transgenic lines were significantly higher than those of wild-type,the effect to the Photosynthesis System of transgenic cottons is less than that of the wild-type.The seed cotton yield per plant of the lines LM2,LM3 is significantly higher than that of the wild-type on the salinized land(salt content 0.45%),which were of great value in cotton breeding and production.
Analysis of low temperature tolerance of betA-expressing cotton
The germination rates of the lines LM1,LM2,LM3 and LM4 at the 13~(th)day after seeding at 15℃condition are 67.33%,78.20%,70.11%and 74.25%,each of which is significantly higher than 53.24%which is the germination rate of the wild-type Lumianyanl9.After 24h low temperature treatment at 5℃,the relative water content of the wild-type plants reduced from 87.94%to 53.43%,whereas that of the transgenic line LM3 is 66.28%,which is significantly higher than that of the wild-type plants,the water-retaining capability of the transgenic lines leaves is better than that of the wild-type plants.The cell membrane ion leakage of the four transgenic lines LM1,LM2,LM3,LM4 is 22.59%,37.95%,40.28%and 23.11%respectively, which is lower than that of the wild-type,the content of malondialdehyde of the transgenic lines LM2,LM3 is 32.96%and 39.65%,which is less than that of the wild-type.The results suggested that less membrane damage occurred in these transgenic lines which have stronger low temperature tolerance.The SOD activity in these transgenic lines is also higher than that of wild-type plants under stress.The results implied that the transgenic plant has strong activity of cell protective enzyme, powerful ability to scavenging free radicals under low temperature stress.After low temperature stress,net photosynthesis,stomatal conductance,intercellular CO_2 concentration and transpiration rate of the four transgenic lines were significantly higher than those of wild-type plants,the net photosynthesis of the LM2 and LM3 is 13.81%and 31.61%higher than that of the wild-type plants,the damage to the Photosynthesis System of transgenic cottons by low temperature stress is less than that of the wild-type.
The above physiological analysis of cotton plants transformed with betA gene demonstrated that a proportion of the transgenic lines(LM2,LM3)accumulated higher levels of glycinebetaine and acquired significantly higher tolerance to salt and low temperature stresses.In addition,the transgenic lines with improved stress tolerance provided valuable cotton materials for cotton stress tolerance breeding.
引文
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