四川小麦转基因技术体系构建及应用研究
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摘要
1992年，Vasil等首次报道了以胚性愈伤为受体，通过基因枪法将gus／bar基因转入小麦品种“Pavon”，标志着转基因小麦研究获得突破性进展。一些改良品质、抗病、抗虫、抗逆和抗除草剂的基因已先后被转移到小麦基因组中，转化技术从单纯依赖基因枪发展到农杆菌介导和花粉管通道等多种途径，受体基因型己从“模式材料”扩展到推广品种。四川小麦转基因研究还处于初级阶段，研究基础薄弱。多年育种实践表明，外省育成品种难以适应四川特有的生态环境，因此急需充分利用我省自育的小麦品种，建立一套切实可行的转基因技术体系。本研究通过评价一些四川主要的小麦推广品种和农艺性状优良的育成品系的组培再生能力，从中筛选出优良的基因型供转基因研究使用，对农杆菌介导和基因枪法的一些影响因素进行研究优化，成功地利用基因枪法将抗除草剂bar基因导入小麦新品种川农16。本研究获得的主要研究结果如下：
(1) 从四川麦区育成的8个小麦品种(系)中筛选出具有较强组培愈伤诱导能力和植株再生能力的国家审定(2003年)新品种川农16作为转基因研究的受体基因型。其幼胚在供试的6个培养基中平均愈伤诱导率达92.9％，显著高于其它基因型，在个别愈伤诱导培养基的诱导率可达99.25％，且平均出芽率仅1.13％，愈伤质量较好。所诱导的愈伤在MS_(10)培养基上的绿芽分化率达77.78％。结果表明，我们选育的川农16适合作为四川小麦转基因研究的受体基因型。
(2) 从供试的6种愈伤诱导培养基和2种再分化培养基中筛选到具有较强愈伤
诱导能力的MSZ培养基和较高绿芽分化率的MS:。培养基。MSZ对8个供试
材料幼胚的平均愈伤诱导率为94.0%，高于其它愈伤诱导培养基的诱导能力
其平均出芽率为7.8%。分析表明MS大量元素加倍的愈伤诱导培养基MSZ
比较适合于在转基因操作初期用来诱导愈伤。各基因型在供试的2个再分
化培养基Mslo和MS:。.，上的平均绿芽分化率分别为64.9%和40.2%，仅有
1个基因型(异源2号)在MSJO-，培养基上的绿芽分化率高于Msl。，分别
为87.5%和75.0%。据此认为，在小麦组织培养中利用MS:培养基进行愈
伤诱导，并用MS:。培养基进行绿芽分化培养是一套适宜的组培再生体系。
(3)分别采用不同浓度种类的筛选剂对川农16和川麦犯以及川育16的幼胚愈
伤组织筛选发现，Kan不能抑制幼胚愈伤的生长，平均出愈率达87%，愈
伤组织生长良好。川农16的幼胚在含不同G418剂量的MSZ中生长受抑制，
但出愈率不随浓度变化而变化，说明G418不能很好地抑制幼胚生长，未能
筛选出合适的选择浓度。川农16和力i麦32的幼胚在含不同L一PPT剂量的
Ms:培养基中，生长受到明显抑制，L一PPT浓度为3m创L时就能完全抑制
幼胚的生长，因此在小麦遗传转化诱导愈伤和继代阶段L一PPT合适的筛选
浓度是2一3mg/L。用根癌农杆菌菌株(EHA 1 01)对这3个基因型的幼胚进
行遗传转化，统计抗性愈伤获得率结果表明，侵染时间】hr对幼胚的生存和
转化较为有利:侵染前对幼胚进行高渗处理和乙酞丁香酮(AS)处理后，
出愈率与未经处理相比并未表现出明显的增加趋势，但基因型对不同处理
的存在差异。
(4)通过6种不同轰击压与轰击距离的组合，在显微镜下观察发现1100PsiX
12cm，1350PsiX12cm两种参数的轰击效果较好，金粉比较均匀地分布于
培养皿中心直径Zcm的区域，边缘部分与中心部分的分散程度较一致。而
1100Psix6cm这个参数的分散状态最差，中心部分金粉集中，并且结块严
重，边缘部分金粉较少。结果说明在利用基因枪进行轰击时，为使金粉尽
可能地轰击到较多的受体，宜采用lloopsiX12em，135opsiX12em两种轰
击参数。轰击7天后随机挑选了112块愈伤检测g卫绍基因活性，其中有6
块出现了明显蓝斑，占5.4%。并且这6块愈伤中有4块来自必0.6林mX n00psi
X 12em的轰击参数处理，另2块分别来自必0.6林mXll00psiXgem，必0.6林m
X 135OPsiXgcm的处理。结果表明，直径0.6林m金粉用于轰击效果相对最
好。
(5)利用上述基因枪轰击参数将含有抗除草剂bar基因的质粒载体pDM803用
于转化6个小麦基因型，通过在2-smg几PPT筛选后获得了27株再生小麦
植株。利用bar基因上的一段特异序列而设计的引物进行PCR检测表明，
其中有2株来源于川农16的抗性再生植株能扩增出一条约3lobP的特征片
段，与质粒载体pDM803的扩增片段大小一致。对扩增片段回收纯化，并
进行序列测定表明，扩增片段碱基序列与从NCBI数据库下载的2个bar
基因全序列(X05822，X17220)的一段完全吻合，仅在3’端的286位上
有一个碱基是G，而在X05822，X 1 7220上是C，虽然它们与一个人工
修饰后的ba:基因序列(AY028212)有较大的差异，但5个DNA序列
编码的氨基酸序列完全一致。据此认为，抗除草剂bar基因已经整合
到小麦品种川农16的基因组上，目前这2株转基因植株已经正常结实。
(6)对40个通过基因工程获得的组培单株收获的种子利用A队GE进行醇溶蛋
白位点检测，从中发现了一些材料与对照相比，部分位点发生了细微的变
化。分析表明，98一
First successful generation of transgenic wheat (Triticum aetuvivum L.) plants was reported by Vasil et al. (1992) through particle bombardment of plasmid vector pBARGUS into cells of type-C, long-term regenerable embryogenic callus derived form c.v. Pavan. It is a symboliz milestone in the history of wheat transformation. Although wheat transformation has lagged behind other important cereals such as rice and maize due to many reasons, in recent reports indicated that wheat genetic transformations is becoming a routine technique adopted worldwide in different laboratories. This technique was successfully being used for introduction of agronomically important genes for quality improvement, resistance against fungal pathogens, insects, drought stress and herbicide. The development of methodologies for the delivery of exogenous genes provided the new alternative approaches, such as Agrobacterium-mediated and pollen-tube pathway method for particle bombardment. The several elite culitvar genotypes were used, wh
ich is not depended on 'model' genotypes, such as Bobwhite.
In China, transgenic wheat plants were regenerated in different laboratories. However, in Sichuan province, the transformation of wheat was just carried out in few laboratories. In practice wheat breeding engineering, most foreign cultivars couldn't adapted to the unique entironment in Sichuan. So, it's necessary to establish a genetic transformation system for the elite wheat cultivars planted in Sichuan. In present study, the response in tissue culture of 8 elite varieties or breeding lines with good agronomic characteristics were investigated in order to select the best suitable donor genotypes for genetic transformation. Moreover, the main parameters of
Agrobacterium-mediated and particle bombardment were demonstrated with the aim of optimize the procedures for their genetic transformation. Then two fertile transgenic plants ( c.v chuannongl6) with bar gene were regenerated successfully via microprojectile bombardment method and a transformation system for Sichuan elite cultivars was constructed successfully too. The main results were described as folio wings:
1. The frequencies of induced-callus and regeneration capabilities of 8 elite wheat genotypes with good agronomical traits in Sichuan were investigated by the F-test and shortest significant ranges tests(SSR). The results showed that the immature embryo derived from novel cultivar Chuannongl6, which presented the highest callus induction frequency and better regeneration capabilities, was suitable donor explant for wheat transformation. The average frequency of induced-callus derived from immature embryo of chuannong16 (92.9%) was significant highest among 8 genotypes. In given media(N6o), it was 99.25%, with the lowest frequency of initiated shoot directly from explants(1.13%) and more embryogenic callus. 77.78% calli derived from Chuannong 16 cultured by dedifferentiation media MSio regenerated the leaf-like green spot. Chuannongl6, with stable and super high yield, is the new breakthrough of Sichuan wheat breeding. So, the elite genotype Chuannongl6 was selected for donor genotype in Sichuan wheat transformation.
2. The significant effects of medium on callus induction were observed by .F-test and SSR. Among 6 media, the frequencies of callus induction on MS2 and N6o were higher than that of the others. The average frequencies of callus induction on MS2 and N60 were 94.0% and 93.1%, respectively. The lowest frequency (83.6%) of callus induction was observed on N61. When MS2 and N6o were selected for callus induction media, it was observed that the frequency of initiated shoot directly from explants on N60 (60.2%) was higher than that of MS2. It indicated that MS2 was superior to N60 for callus induction. Thus, we chose MS2 as callus induction medium in the following procedures. Comparison of the differentiation frequency for each genotype on two
media MS10 and MS10-1(87.5%, 75.0%), it indicated that MS10 was superior to MS10-1 for green shoot differentiation. So, the call
(1) 从四川麦区育成的8个小麦品种(系)中筛选出具有较强组培愈伤诱导能力和植株再生能力的国家审定(2003年)新品种川农16作为转基因研究的受体基因型。其幼胚在供试的6个培养基中平均愈伤诱导率达92.9％，显著高于其它基因型，在个别愈伤诱导培养基的诱导率可达99.25％，且平均出芽率仅1.13％，愈伤质量较好。所诱导的愈伤在MS_(10)培养基上的绿芽分化率达77.78％。结果表明，我们选育的川农16适合作为四川小麦转基因研究的受体基因型。
(2) 从供试的6种愈伤诱导培养基和2种再分化培养基中筛选到具有较强愈伤
诱导能力的MSZ培养基和较高绿芽分化率的MS:。培养基。MSZ对8个供试
材料幼胚的平均愈伤诱导率为94.0%，高于其它愈伤诱导培养基的诱导能力
其平均出芽率为7.8%。分析表明MS大量元素加倍的愈伤诱导培养基MSZ
比较适合于在转基因操作初期用来诱导愈伤。各基因型在供试的2个再分
化培养基Mslo和MS:。.，上的平均绿芽分化率分别为64.9%和40.2%，仅有
1个基因型(异源2号)在MSJO-，培养基上的绿芽分化率高于Msl。，分别
为87.5%和75.0%。据此认为，在小麦组织培养中利用MS:培养基进行愈
伤诱导，并用MS:。培养基进行绿芽分化培养是一套适宜的组培再生体系。
(3)分别采用不同浓度种类的筛选剂对川农16和川麦犯以及川育16的幼胚愈
伤组织筛选发现，Kan不能抑制幼胚愈伤的生长，平均出愈率达87%，愈
伤组织生长良好。川农16的幼胚在含不同G418剂量的MSZ中生长受抑制，
但出愈率不随浓度变化而变化，说明G418不能很好地抑制幼胚生长，未能
筛选出合适的选择浓度。川农16和力i麦32的幼胚在含不同L一PPT剂量的
Ms:培养基中，生长受到明显抑制，L一PPT浓度为3m创L时就能完全抑制
幼胚的生长，因此在小麦遗传转化诱导愈伤和继代阶段L一PPT合适的筛选
浓度是2一3mg/L。用根癌农杆菌菌株(EHA 1 01)对这3个基因型的幼胚进
行遗传转化，统计抗性愈伤获得率结果表明，侵染时间】hr对幼胚的生存和
转化较为有利:侵染前对幼胚进行高渗处理和乙酞丁香酮(AS)处理后，
出愈率与未经处理相比并未表现出明显的增加趋势，但基因型对不同处理
的存在差异。
(4)通过6种不同轰击压与轰击距离的组合，在显微镜下观察发现1100PsiX
12cm，1350PsiX12cm两种参数的轰击效果较好，金粉比较均匀地分布于
培养皿中心直径Zcm的区域，边缘部分与中心部分的分散程度较一致。而
1100Psix6cm这个参数的分散状态最差，中心部分金粉集中，并且结块严
重，边缘部分金粉较少。结果说明在利用基因枪进行轰击时，为使金粉尽
可能地轰击到较多的受体，宜采用lloopsiX12em，135opsiX12em两种轰
击参数。轰击7天后随机挑选了112块愈伤检测g卫绍基因活性，其中有6
块出现了明显蓝斑，占5.4%。并且这6块愈伤中有4块来自必0.6林mX n00psi
X 12em的轰击参数处理，另2块分别来自必0.6林mXll00psiXgem，必0.6林m
X 135OPsiXgcm的处理。结果表明，直径0.6林m金粉用于轰击效果相对最
好。
(5)利用上述基因枪轰击参数将含有抗除草剂bar基因的质粒载体pDM803用
于转化6个小麦基因型，通过在2-smg几PPT筛选后获得了27株再生小麦
植株。利用bar基因上的一段特异序列而设计的引物进行PCR检测表明，
其中有2株来源于川农16的抗性再生植株能扩增出一条约3lobP的特征片
段，与质粒载体pDM803的扩增片段大小一致。对扩增片段回收纯化，并
进行序列测定表明，扩增片段碱基序列与从NCBI数据库下载的2个bar
基因全序列(X05822，X17220)的一段完全吻合，仅在3’端的286位上
有一个碱基是G，而在X05822，X 1 7220上是C，虽然它们与一个人工
修饰后的ba:基因序列(AY028212)有较大的差异，但5个DNA序列
编码的氨基酸序列完全一致。据此认为，抗除草剂bar基因已经整合
到小麦品种川农16的基因组上，目前这2株转基因植株已经正常结实。
(6)对40个通过基因工程获得的组培单株收获的种子利用A队GE进行醇溶蛋
白位点检测，从中发现了一些材料与对照相比，部分位点发生了细微的变
化。分析表明，98一
First successful generation of transgenic wheat (Triticum aetuvivum L.) plants was reported by Vasil et al. (1992) through particle bombardment of plasmid vector pBARGUS into cells of type-C, long-term regenerable embryogenic callus derived form c.v. Pavan. It is a symboliz milestone in the history of wheat transformation. Although wheat transformation has lagged behind other important cereals such as rice and maize due to many reasons, in recent reports indicated that wheat genetic transformations is becoming a routine technique adopted worldwide in different laboratories. This technique was successfully being used for introduction of agronomically important genes for quality improvement, resistance against fungal pathogens, insects, drought stress and herbicide. The development of methodologies for the delivery of exogenous genes provided the new alternative approaches, such as Agrobacterium-mediated and pollen-tube pathway method for particle bombardment. The several elite culitvar genotypes were used, wh
ich is not depended on 'model' genotypes, such as Bobwhite.
In China, transgenic wheat plants were regenerated in different laboratories. However, in Sichuan province, the transformation of wheat was just carried out in few laboratories. In practice wheat breeding engineering, most foreign cultivars couldn't adapted to the unique entironment in Sichuan. So, it's necessary to establish a genetic transformation system for the elite wheat cultivars planted in Sichuan. In present study, the response in tissue culture of 8 elite varieties or breeding lines with good agronomic characteristics were investigated in order to select the best suitable donor genotypes for genetic transformation. Moreover, the main parameters of
Agrobacterium-mediated and particle bombardment were demonstrated with the aim of optimize the procedures for their genetic transformation. Then two fertile transgenic plants ( c.v chuannongl6) with bar gene were regenerated successfully via microprojectile bombardment method and a transformation system for Sichuan elite cultivars was constructed successfully too. The main results were described as folio wings:
1. The frequencies of induced-callus and regeneration capabilities of 8 elite wheat genotypes with good agronomical traits in Sichuan were investigated by the F-test and shortest significant ranges tests(SSR). The results showed that the immature embryo derived from novel cultivar Chuannongl6, which presented the highest callus induction frequency and better regeneration capabilities, was suitable donor explant for wheat transformation. The average frequency of induced-callus derived from immature embryo of chuannong16 (92.9%) was significant highest among 8 genotypes. In given media(N6o), it was 99.25%, with the lowest frequency of initiated shoot directly from explants(1.13%) and more embryogenic callus. 77.78% calli derived from Chuannong 16 cultured by dedifferentiation media MSio regenerated the leaf-like green spot. Chuannongl6, with stable and super high yield, is the new breakthrough of Sichuan wheat breeding. So, the elite genotype Chuannongl6 was selected for donor genotype in Sichuan wheat transformation.
2. The significant effects of medium on callus induction were observed by .F-test and SSR. Among 6 media, the frequencies of callus induction on MS2 and N6o were higher than that of the others. The average frequencies of callus induction on MS2 and N60 were 94.0% and 93.1%, respectively. The lowest frequency (83.6%) of callus induction was observed on N61. When MS2 and N6o were selected for callus induction media, it was observed that the frequency of initiated shoot directly from explants on N60 (60.2%) was higher than that of MS2. It indicated that MS2 was superior to N60 for callus induction. Thus, we chose MS2 as callus induction medium in the following procedures. Comparison of the differentiation frequency for each genotype on two
media MS10 and MS10-1(87.5%, 75.0%), it indicated that MS10 was superior to MS10-1 for green shoot differentiation. So, the call
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