基于组织培养和授粉后浸蘸花柱的两种棉花遗传转化体系的建立
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摘要
农杆菌介导的遗传转化是目前广泛采用的转基因方法,其操作方法简单,无需昂贵设备,导入外源基因多为单拷贝或低拷贝,并多呈孟德尔式遗传规律,遗传稳定性好,易于研究和育种利用。但农杆菌介导的遗传转化依赖于组织培养,受转化受体材料的基因型限制。棉花成功组织培养的种质材料相对有限,农杆菌介导的遗传转化也受限于基因型。本研究致力于突破棉花遗传转化的基因型限制,开展了两方面相关的研究：拓宽可组织培养再生并适合于转化的棉花基因型；尝试以农杆菌浸蘸棉花花柱的整株活体转化方法进行棉花遗传转化,避开遗传转化依赖于组织培养的限制。
新疆是我国第一大优质产棉区,其棉花产量占全国总产量1/3以上。但诸多因素如干旱气候、病虫害等不利于新疆棉花生产的可持续发展。新疆棉花品种的改良,尤其是针对新疆特定生态条件而开展的耐盐、抗旱、抗病、抗冻、高产、优质、早熟等品种培育,将促进新疆棉花生产发展。本研究以新疆陆地棉4个主栽品种为材料,研究其体细胞胚胎发生能力,以期建立新疆陆地棉有效的体细胞胚胎发生再生体系,扩展可再生棉花基因型,促进新疆转基因棉研究和生产。研究利用多种浓度的激素组合成功地诱导获得了体细胞胚并进一步发育成苗。研究发现,所用4种激素组合均能有效诱导愈伤组织,其中又以0.02mg l-1KT+0.1mg l-12,4-D和0.10mg l-1KT+0.1mg l-12,4-D组合的诱导效果最佳。2个愈伤诱导措施：①愈伤组织诱导培养基中KNO3(?)量加倍,②下胚轴外植体纵切面接触培养基,有利于胚性愈伤组织的诱导和产生。挑选黄绿色、灰绿色或浅绿色的质地疏松的愈伤组织继代于无激素且KNO3含量加倍的培养基中可产生胚性愈伤组织,并在高比例KT/2,4-D (0.05mg l-1KT+0.01mg l-12,4-D或0.10mg1-1KT+0.01mg1-12,4-D)促进下发育成胚。借助在培养基上垫滤纸产生干燥作用,并间隔使用强透气效果的棉塞对培养三角瓶进行透气处理,体细胞胚可成熟发育并产生根系发达的正常再生植株。应用此法,4个实验材料在6-8个月内即可获得大量再生苗。
以胚性愈伤组织为外植体,通过农杆菌介导转化将抗病基因hcm1基因和抗旱基因bcp基因导入新疆陆地棉品种新陆中20。转化的胚性愈伤在100mg1-1卡纳霉素和0.5,1.0,1.5mg1-1Phosphinothricin (PPT)两种不同选择压进行抗性愈伤组织筛选。在筛选过程中尽可能地铺散胚性愈伤使之与选择培养基充分接触,增强选择剂的选择效果,减少非转化子的逃逸。胚性愈伤转化6个月后即分化再生转基因苗,九个培养皿的转化愈伤在8个月内即可获得100个以上独立的再生株系。转化再生植株产生了广泛的体细胞无性系变异,变异频率高达100%。对所获得的转基因植株进行PCR检测,发现在100mg1-1卡纳霉素和1.5mg1-1PPT筛选下获得的再生植株,选择标记nptⅡ基因和bar基因PCR阳性率在90%以上,抗病基因hcml基因和抗旱基因bcp基因PCR阳性率在75%以上。卡那霉素或Basta除草剂抗性鉴定表明nptⅡ基因和bar基因在PCR检测呈阳性的植株中成功表达。Southern杂交结果表明,部分PCR检测呈阳性植株已经整合hcml基因或bcp基因。
在棉花授粉后的特定时间内,以农杆菌-蔗糖溶液浸蘸棉花花柱进行整株活体转化。实验证明,农杆菌在附加0.05%(v/v)表面活性剂Silwet L-77和40mg1-1乙酰丁香酮的10%蔗糖溶液中能通过花粉管通道有效地进行转化并获得转基因种子。研究发现,农杆菌浸蘸花柱的时间和位点对转化成功与否至关重要。授粉当天上午9：00-11：00进行浸蘸柱头处理未能获得转基因植株；授粉当天下午17：00-19：00浸蘸柱头收获4株转基因植株,转化率为0.07-0.17%；授粉当天下午17：00-19：00切除柱头后滴加农杆菌-蔗糖溶液可进一步提高转化率,收获7株转基因植株,转化率为0.46-0.93%；授粉次日上午9：00=11：00去除花柱后进行浸蘸处理收获2株转基因植株,转化率为0.04-0.06%。4个实验均进行7天(次)独立处理,有3个实验的2次或3次独立处理获得了转基因植株,转化率从0.04%到0.93%不等。PCR和Southern杂交验证表明外源基因bar基因已整合至棉花基因组中,T0和T1植株除草剂抗性鉴定亦证明bar基因在转基因植株中成功表达。多次独立实验处理的成功获得转基因植株证明了农杆菌浸蘸棉花花柱的整株活体转化体系的可重复性和可靠性。与传统的依赖于组织培养的转基因方法相比,农杆菌浸蘸棉花花柱的整株活体转化体系更为简便、经济和高效,同时还避免了组织培养过程当中产生的体细胞无性系变异等问题。
Agrobacterium-mediated transformation is the main method used in the field of biotechnology. It is attractive because of the ease of the protocol coupled with minimal equipment costs. Moreover, transgenic plants developed by this method often contain simple copy insertions. These advantages were a driving force to adapt the method to many different crops including cotton. But cotton as well as many economically important plant species, or elite varieties of particular species, remain highly recalcitrant to Agrobacterium-mediated transformation. In this study, we endeavored to extend the host range and transformation efficiency of cotton by Agrobacterium, to seek a novel mothed of in planta transformation avoiding from the process of tissue culture.
Xinjiang is an excellent cotton growing region in China, producing more than one-third of Chinese cotton production. But worsening conditions of cotton diseases and insect pests as well as the dry condition in Xinjiang had caused a bottleneck in sustainable development of cotton in this region. Cultivars with resistance to abiotic or biotic stresses, such as salt tolerance, cold resistance, and disease resistance, as well as good quality and high yield are needed in Xinjiang. Genetic improvement through genetic transformation can be useful in fulfilling this demand. An efficient somatic embryo production and maturation procedure was developed to regenerate plantlets from hypocotyls of four cotton cultivars grown in Xinjiang. Calli were effectively produced by0.01-0.10mg l-1kinetin (KT) and0.10mg l-12,4-dichlorophenoxyacetic acid (2,4-D), with a better result by0.02mg l-1KT+0.1mg l-12,4-D and0.10mg l-1KT+0.1mg l-12,4-D. Split hypocotyl segments and double amounts of KNO3during induction of calli were beneficial to the emerge of embryogenic calli. Embryogenic calli and globular-stage somatic embryos were effectively initiated with high concentration of KT and low concentration of2,4-D in ECM media (0.05mg l-1KT 0.01mg l-12,4-D and0.10mg l-1KT+0.01mg I-12,4-D). Embryos were further developed into plantlets in MSBF medium under the conditions of dehydration and ventilation, which were achieved using filter paper on medium and cotton tampon sealing flask. Using this protocol, normal plantlets with strong roots were developed from these four cotton cultivars in6-8months.
Embryogenic calli of Xinluzhong20were transformed with Agrobacterium carrying drought resistant gene bcp or disease resistant gene hcml under the selection by Phosphinothricin (PPT) or Kanamycin, respectively. Low density of transformed embryogenic calli in Petri dish provided more effective contact between embryogenic calli and selective agents, and increased the selective efficiency with less escape. Transformed embryogenic calli could regenerate into transgenic plantlets after6months. And the number of transgenic plantlets obtained from9Petri dishes of transformed embryogenic calli may reach as many as100independent transgenic lines in8months, tremendously improving transformation frequency and reducing the overall time period significantly. A frequency of100%somaclonal variation was observed among the regenerated plants. PCR tests of plantlets regenerated under selection by100mg l-1kanamycin or1.5mg l-1PPT showed a positive rate of90%in terms of nptⅡ or bar gene, while75%positive rate in terms of hcml gene or bcp gene. The resistance test with Kanamycin or Basta proofed the expression of the integrated nptll or bar gene. And Southern blot analysis confirmed the integration of hcml gene or bcp gene into the cotton genome of some of PCR positive plants, suggesting that Xinluzhong20was efficiently susceptible to Agrobacterium-mediated transformation.
Simple incubation of Agrobacterium with cotton pistil after pollination could produce transformed seeds, enabling transformed plants to be produced without the necessity of a tissue culture system.10%sucrose inoculation medium containing0.05%(v/v) Silwet L-77and40mg l-1acetosyringone facilitated the entrance of Agrobacterium into interior of germ line and transformation of cotton. The importance of Agrobacterium inoculation time and pistil position (stigma, style and stigma excisions) for inoculation was evaluated in terms of transgenic plant production. No transformants produced from pistil drip during9:00～11:00on the first day of flowering. Pistil drip during17:00～19:00on the first day of flowering resulted in0.07-0.17%Basta resistant plants/number of viable seeds generated, and stigma excision prior to pistil drip during this time period gave rise to a transformation efficiency of0.46-0.93%, in contrast with0.04-0.06%generated from pistil drip during9:00～11:00on the second day of flowering. Among four experiments with seven indepentant treatment days, three experiments produced transformed plants from two or three indepentant treatment days. The transformation efficiency ranged from0.04%to0.93%, with the highest rate of transformation from the treatment time period during the afternoon of flowering. PCR and Southern blot analysis consistently confirmed the integration of bar gene into the TO cotton genome, and herbicide resistance test of TO and T1generations showed bar gene expressed and functioned well in these transgenics. Transgenic cotton plants produced from two or three independent treatments of these experiments demonstrated the reliability and reproducibility of this simple transformation procedure by drop dip of Agrobacterium-sucrose onto cotton pistil. Compared with traditional tissue culture based transformation procedures, this Agrobacterium-mediated in planta transformation is technically easier and generally faster to obtain transgenic plants, and it overcomes the problems of somaclonal variation caused by tissue culture.
新疆是我国第一大优质产棉区,其棉花产量占全国总产量1/3以上。但诸多因素如干旱气候、病虫害等不利于新疆棉花生产的可持续发展。新疆棉花品种的改良,尤其是针对新疆特定生态条件而开展的耐盐、抗旱、抗病、抗冻、高产、优质、早熟等品种培育,将促进新疆棉花生产发展。本研究以新疆陆地棉4个主栽品种为材料,研究其体细胞胚胎发生能力,以期建立新疆陆地棉有效的体细胞胚胎发生再生体系,扩展可再生棉花基因型,促进新疆转基因棉研究和生产。研究利用多种浓度的激素组合成功地诱导获得了体细胞胚并进一步发育成苗。研究发现,所用4种激素组合均能有效诱导愈伤组织,其中又以0.02mg l-1KT+0.1mg l-12,4-D和0.10mg l-1KT+0.1mg l-12,4-D组合的诱导效果最佳。2个愈伤诱导措施：①愈伤组织诱导培养基中KNO3(?)量加倍,②下胚轴外植体纵切面接触培养基,有利于胚性愈伤组织的诱导和产生。挑选黄绿色、灰绿色或浅绿色的质地疏松的愈伤组织继代于无激素且KNO3含量加倍的培养基中可产生胚性愈伤组织,并在高比例KT/2,4-D (0.05mg l-1KT+0.01mg l-12,4-D或0.10mg1-1KT+0.01mg1-12,4-D)促进下发育成胚。借助在培养基上垫滤纸产生干燥作用,并间隔使用强透气效果的棉塞对培养三角瓶进行透气处理,体细胞胚可成熟发育并产生根系发达的正常再生植株。应用此法,4个实验材料在6-8个月内即可获得大量再生苗。
以胚性愈伤组织为外植体,通过农杆菌介导转化将抗病基因hcm1基因和抗旱基因bcp基因导入新疆陆地棉品种新陆中20。转化的胚性愈伤在100mg1-1卡纳霉素和0.5,1.0,1.5mg1-1Phosphinothricin (PPT)两种不同选择压进行抗性愈伤组织筛选。在筛选过程中尽可能地铺散胚性愈伤使之与选择培养基充分接触,增强选择剂的选择效果,减少非转化子的逃逸。胚性愈伤转化6个月后即分化再生转基因苗,九个培养皿的转化愈伤在8个月内即可获得100个以上独立的再生株系。转化再生植株产生了广泛的体细胞无性系变异,变异频率高达100%。对所获得的转基因植株进行PCR检测,发现在100mg1-1卡纳霉素和1.5mg1-1PPT筛选下获得的再生植株,选择标记nptⅡ基因和bar基因PCR阳性率在90%以上,抗病基因hcml基因和抗旱基因bcp基因PCR阳性率在75%以上。卡那霉素或Basta除草剂抗性鉴定表明nptⅡ基因和bar基因在PCR检测呈阳性的植株中成功表达。Southern杂交结果表明,部分PCR检测呈阳性植株已经整合hcml基因或bcp基因。
在棉花授粉后的特定时间内,以农杆菌-蔗糖溶液浸蘸棉花花柱进行整株活体转化。实验证明,农杆菌在附加0.05%(v/v)表面活性剂Silwet L-77和40mg1-1乙酰丁香酮的10%蔗糖溶液中能通过花粉管通道有效地进行转化并获得转基因种子。研究发现,农杆菌浸蘸花柱的时间和位点对转化成功与否至关重要。授粉当天上午9：00-11：00进行浸蘸柱头处理未能获得转基因植株；授粉当天下午17：00-19：00浸蘸柱头收获4株转基因植株,转化率为0.07-0.17%；授粉当天下午17：00-19：00切除柱头后滴加农杆菌-蔗糖溶液可进一步提高转化率,收获7株转基因植株,转化率为0.46-0.93%；授粉次日上午9：00=11：00去除花柱后进行浸蘸处理收获2株转基因植株,转化率为0.04-0.06%。4个实验均进行7天(次)独立处理,有3个实验的2次或3次独立处理获得了转基因植株,转化率从0.04%到0.93%不等。PCR和Southern杂交验证表明外源基因bar基因已整合至棉花基因组中,T0和T1植株除草剂抗性鉴定亦证明bar基因在转基因植株中成功表达。多次独立实验处理的成功获得转基因植株证明了农杆菌浸蘸棉花花柱的整株活体转化体系的可重复性和可靠性。与传统的依赖于组织培养的转基因方法相比,农杆菌浸蘸棉花花柱的整株活体转化体系更为简便、经济和高效,同时还避免了组织培养过程当中产生的体细胞无性系变异等问题。
Agrobacterium-mediated transformation is the main method used in the field of biotechnology. It is attractive because of the ease of the protocol coupled with minimal equipment costs. Moreover, transgenic plants developed by this method often contain simple copy insertions. These advantages were a driving force to adapt the method to many different crops including cotton. But cotton as well as many economically important plant species, or elite varieties of particular species, remain highly recalcitrant to Agrobacterium-mediated transformation. In this study, we endeavored to extend the host range and transformation efficiency of cotton by Agrobacterium, to seek a novel mothed of in planta transformation avoiding from the process of tissue culture.
Xinjiang is an excellent cotton growing region in China, producing more than one-third of Chinese cotton production. But worsening conditions of cotton diseases and insect pests as well as the dry condition in Xinjiang had caused a bottleneck in sustainable development of cotton in this region. Cultivars with resistance to abiotic or biotic stresses, such as salt tolerance, cold resistance, and disease resistance, as well as good quality and high yield are needed in Xinjiang. Genetic improvement through genetic transformation can be useful in fulfilling this demand. An efficient somatic embryo production and maturation procedure was developed to regenerate plantlets from hypocotyls of four cotton cultivars grown in Xinjiang. Calli were effectively produced by0.01-0.10mg l-1kinetin (KT) and0.10mg l-12,4-dichlorophenoxyacetic acid (2,4-D), with a better result by0.02mg l-1KT+0.1mg l-12,4-D and0.10mg l-1KT+0.1mg l-12,4-D. Split hypocotyl segments and double amounts of KNO3during induction of calli were beneficial to the emerge of embryogenic calli. Embryogenic calli and globular-stage somatic embryos were effectively initiated with high concentration of KT and low concentration of2,4-D in ECM media (0.05mg l-1KT 0.01mg l-12,4-D and0.10mg l-1KT+0.01mg I-12,4-D). Embryos were further developed into plantlets in MSBF medium under the conditions of dehydration and ventilation, which were achieved using filter paper on medium and cotton tampon sealing flask. Using this protocol, normal plantlets with strong roots were developed from these four cotton cultivars in6-8months.
Embryogenic calli of Xinluzhong20were transformed with Agrobacterium carrying drought resistant gene bcp or disease resistant gene hcml under the selection by Phosphinothricin (PPT) or Kanamycin, respectively. Low density of transformed embryogenic calli in Petri dish provided more effective contact between embryogenic calli and selective agents, and increased the selective efficiency with less escape. Transformed embryogenic calli could regenerate into transgenic plantlets after6months. And the number of transgenic plantlets obtained from9Petri dishes of transformed embryogenic calli may reach as many as100independent transgenic lines in8months, tremendously improving transformation frequency and reducing the overall time period significantly. A frequency of100%somaclonal variation was observed among the regenerated plants. PCR tests of plantlets regenerated under selection by100mg l-1kanamycin or1.5mg l-1PPT showed a positive rate of90%in terms of nptⅡ or bar gene, while75%positive rate in terms of hcml gene or bcp gene. The resistance test with Kanamycin or Basta proofed the expression of the integrated nptll or bar gene. And Southern blot analysis confirmed the integration of hcml gene or bcp gene into the cotton genome of some of PCR positive plants, suggesting that Xinluzhong20was efficiently susceptible to Agrobacterium-mediated transformation.
Simple incubation of Agrobacterium with cotton pistil after pollination could produce transformed seeds, enabling transformed plants to be produced without the necessity of a tissue culture system.10%sucrose inoculation medium containing0.05%(v/v) Silwet L-77and40mg l-1acetosyringone facilitated the entrance of Agrobacterium into interior of germ line and transformation of cotton. The importance of Agrobacterium inoculation time and pistil position (stigma, style and stigma excisions) for inoculation was evaluated in terms of transgenic plant production. No transformants produced from pistil drip during9:00～11:00on the first day of flowering. Pistil drip during17:00～19:00on the first day of flowering resulted in0.07-0.17%Basta resistant plants/number of viable seeds generated, and stigma excision prior to pistil drip during this time period gave rise to a transformation efficiency of0.46-0.93%, in contrast with0.04-0.06%generated from pistil drip during9:00～11:00on the second day of flowering. Among four experiments with seven indepentant treatment days, three experiments produced transformed plants from two or three indepentant treatment days. The transformation efficiency ranged from0.04%to0.93%, with the highest rate of transformation from the treatment time period during the afternoon of flowering. PCR and Southern blot analysis consistently confirmed the integration of bar gene into the TO cotton genome, and herbicide resistance test of TO and T1generations showed bar gene expressed and functioned well in these transgenics. Transgenic cotton plants produced from two or three independent treatments of these experiments demonstrated the reliability and reproducibility of this simple transformation procedure by drop dip of Agrobacterium-sucrose onto cotton pistil. Compared with traditional tissue culture based transformation procedures, this Agrobacterium-mediated in planta transformation is technically easier and generally faster to obtain transgenic plants, and it overcomes the problems of somaclonal variation caused by tissue culture.
引文
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