SNAC1基因在棉花上的应用及盐胁迫下棉花基因组DNA甲基化分析
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摘要
植物特别是作物对非生物逆境(干旱、冷害和高盐等)的适应性对于作物的产量和品质有着非常重要的影响。干旱和高盐是植物逆境最普遍的形式,在许多地区是农业发展的制约因素,因而抗逆育种一直受到重视。利用基因工程手段,提高作物的耐盐、耐早能力,具有深远的应用前景。植物在逆境胁迫下基因组DNA通常发生着甲基化变化,探明胁迫下植物基因组DNA甲基化的水平及变化模式对研究其抗逆机制有着重要意义。
棉花是一种重要的经济作物,利用基因工程技术对于提高棉花的产量、增强抗逆能力以及改良纤维品质具有重要作用。本研究以最终获得用于抗逆遗传改良的转基因棉花植株为目的,通过导入并超量表达对逆境起调控作用的转录因子基因——SNAC1基因,系统地对这些转基因再生棉花植株进行抗逆性分析和鉴定。初步鉴定出SNAC1基因在抗逆遗传改良中的有效性,并对SNAC1基因作为筛选标记基因用于棉花遗传转化进行了研究。另外,以棉花品系YZ1为材料对其盐胁迫下根组织DNA甲基化的水平及变化模式进行了研究。主要结果如下:
1.SNAC1基因是具有耐旱及耐盐的转录因子基因,本研究利用农杆菌介导的方法把SNAC1基因导入到陆地棉品系YZ1。PCR检测和RT-PCR分析表明,SNAC1基因已经整合到转基因棉花的基因组中且得到表达,共获得9个克隆系的再生植株及后代。田间农艺性状调查表明,转基因S1和S2家系的T_1代棉花植株在产量等农艺性状上与非转基因植株比较没有大的变异。棉花离体叶片失水速率测定表明,田间种植的转基因T_1代棉花植株叶片的失水速率为25%左右,低于非转基因植株叶片失水速率(30.4%)。同时,电镜扫描观察证实了离体叶片失水胁迫后转基因植株叶片的气孔关闭数量比例(71.8%)要高于野生型植株(51.5%)。水培抗盐性鉴定结果表明,在200 mmol/L NaCl胁迫5天后转基因T_2代植株幼苗的叶片相对含水量(RWC)为72%,而非转基因植株叶片的相对含水量为61%,并且非转基因植株叶片萎焉脱落要早于转基因植株。同时,转基因T_2代植株幼苗在高盐胁迫及胁迫后恢复过程中的相对生长速度(RGR)分别为7.9%和109.5%,高于非转基因植株的相对生长速度(分别为-7.4%和53.1%)。另外,在150 mmol/L NaCl胁迫下转基因胚性愈伤组织能够继续生长,而非转基因胚性愈伤组织则失水变白死亡。对转基因阳性和阴性T_1代植株的qRT-PCR分析结果表明,SNAC1基因能够调控棉花部分基因的表达,如G2(NAC familyprotein)与G3(NAM-like protein)基因的表达上升以及G5(GSA-AM2)、G11(SAMS)与AF38(ABF2)基因的表达下降。
2.本文以SNAC1基因作为筛选标记基因,采用农杆菌介导法将SNAC1基因导入棉花细胞,同时研究了NaCl作为筛选剂用于棉花遗传转化的可行性。经过棉花下胚轴的农杆菌菌液浸染、共培养以及NaCl筛选培养,获得了转基因棉花胚性愈伤组织。对获得的胚性愈伤组织进行PCR检测证实外源基因已经整合到棉花基因组中且阳性率达50%以上,同时GUS染色也表明GUS基因得到表达。另外,本文还对NaCl作为筛选剂在农杆菌介导的棉花遗传转化中的应用浓度及方法进行了研究,即:在农杆菌侵染后棉花下胚轴的共培养时间为3～4天,愈伤组织诱导初期的NaCl筛选浓度在1.1%～1.5%(W/V)之间,随着愈伤组织的生长其耐盐性有所增强而逐渐提高NaCl的筛选浓度。与正常培养条件相比,培养基中NaCl的存在不利于胚性愈伤组织的分化,因此经过3～4次愈伤组织继代筛选后要及时去除培养基中的NaCl以促进胚性愈伤组织的分化。
3.本研究以陆地棉品系YZ1为材料,调查不同NaCl浓度处理下棉花幼苗生长及根基因组DNA甲基化的水平和变化模式。结果表明,100 mmol/L NaCl对棉花幼苗的株高和根长生长有促进作用,而200 mmol/L NaCl则显著抑制株高和根长的生长。此外,在100～200 mmol/LNaCl胁迫下棉花幼苗的侧根数量被严重抑制。MSAP分析结果表明,经100、150和200 mmol/L NaCl胁迫处理后棉花幼苗根基因组DNA甲基化水平分别为38.1%、35.2%和34.5%,均低于对照(0 mmol/LNaCl)的甲基化水平(41.2%),同时棉花幼苗根DNA甲基化的水平与NaCl处理浓度呈显著负相关(r=-0.986)。与对照(0 mmol/L NaCl)相比,在100、150和200 mmol/LNaCl胁迫下棉花幼苗根基因组DNA发生甲基化的比率分别为6.4%、7.6%、11.3%,而去甲基化的比率分别为12.7%、11.1%、8.2%.序列分忻结果显示,部分MSAP差异片段与功能基因同源。RT-PCR分析表明,分别与MSAP差异片段M2及M3高度同源的海岛棉gyspy反转座子反转录酶基因及陆地棉cDNA基因在处理(200 mmol/LNaCI)的棉花幼苗根中有表达,但在对照(0 mmol/L NaCl)的根中则没有表达。
Plant's adaptability to abiotic stresses such as drought,cold and salinity plays an important role in plant's yield and quality.Drought and salinity stresses are the most common form of adversity,which are the limiting factors for agricultural development in many areas.Therefore,it is important to improve crop's tolerance to stresses by genetic engineering.Plant genomic DNA was always methylated under stresses and studying the level and pattern of genomic DNA methylation under stresses will help to identify the mechanisms on stress resistance.
Cotton is one of the most important economic crops in the world,and genetic engineering played an important role in increasing cotton's yield,enhancing stress resistance and improving cotton fiber quality.The objective of this study was to gain transgenic cotton plants for genetic improvement,SNAC1 gene,a transcriptional factor regulating stress response,which was transformed into cotton and the analysis and identification of stress resistance were carried out on transgenic cotton plants systematically.The availability of SNAC1 gene was indentified preliminarily in genetic improvement and SNAC1 gene as a selection marker gene for genetic transformation of cotton was studied also.In addition,the cotton strain of YZ1 was used to study the level and pattern of DNA methylation in cotton roots under salt stress.The following results were obtained:
1.SNAC1 gene in rice can enhance drought- and salt-tolerance,and then was transformed into Gossypium hirsutum YZ1 by Agrobacterium -mediated genetic transformation in this study.PCR detection and RT-PCR analysis indicated SNAC1 gene had been integrated into the transgenic cotton genome and at the same time regenerated plants of 9 clones were gained totally.The field surveys showed that T_1 generation of transgenic plants did not vary in many agronomic traits such as yield,compared to non-transgenic plants.The determination of rate of water loss of excised-leaves of cotton demonstrated that the RWL(rate of water loss) of T_1 generationof transgenic plants was 25%,and was lower than that(30.4%) of the non-transgenic plants.Then the observation of SEM(scanning electron microscope) also suggested the ratio of stomatal closure in the leaves of transgenic plants was 71.8%and was higher than that(51.5%) of non-transgenic plants.The results of salt-tolerance identification by water culture showed that RWC (Relative water content) of leaves of T_2 generation of transgenic seedlings was 72%under stress of 200 mmol/L NaCl after 5 d and that of non-transgenic cotton's leaves was 61%. What is more,the leaves of non-transgenic plants withered and broken off earlier than that of transgenic plants.In addion,the RGR(Relative growth rate) of transgenic plant seedlings were 7.9%and 109.5%respectively during salt stress and sress revovery,and the RGR of non-transgenic plants were only-7.4%and 53.1%respectively.Under the 150 mmol/L NaCl stess transgenic embryogenic calli can continue to grow,but non-transgenic embryogenic calli lost water and died.The results of qRT-PCR analysis suggested that in T_1 generation of transgenic positive and negative plants SNAC1 gene could regulate expressions of some genes in cotton,for example expressions of G2(NAC family protein) and G3(NAM-like protein) genes were up-regulated and expressions of G5(GSA-AM2), G11(SAMS) and AF38(ABF2) genes were down-regulated.
2.In this study,employing SNAC1 gene was selected as a marker gene and NaCl as a selection agent,SNAC1(Stress-induced NAC 1) and GUS genes were introduced into cotton genome via Agrobacterium-mediated transformation,and the feasibility of NaCl as a selection agent for the genetic transformation of cotton was researched.Finally transgenic embryonic calli were gained after the infection with Agrobacterium,co-culture and selection culture of NaCl.Transgenic calli were confirmed by PCR analysis and positive rate of transgenic calli was 58%,and at the same time expression of the GUS gene was showed with GUS staining.Furthermore the concentration of NaCl and method for employing NaCl as a selection agent had been studied.The hypocotyls were co-cultured for 3～4 d after Agrobaterium infection.The reasonable selection concentration of NaCl should be 1.1%～1.5%(W/V) and the start concentration of NaCl in the callus induction medium should be lower.Then the concentration was increased as cotton calli proliferated.Because NaCl is not beneficial for embryo differentiation and development,NaCl should be removed from the culture medium to promote embryonic calli differentiation after calli were subcultured for 3～4 times.
3.In this study the growth of cotton seedlings and the level and patterns of DNA methylation in the roots of cotton were investigated uder salt stress.The results showed that 100 mmol/L NaCl obviously promoted plant height and root length of cotton seedlings,but 200 mmol/L NaCl significantly inhibited the growth of cotton seedlings; 100～200 mmol/L NaCl inhibited the numbers of lateral root considerably.The analysis of MSAP showed that the level of global DNA methylation in the roots of cotton seedlings was 41.2%,38.1%,35.2%and 34.5%respectively under the stresses of 0,100,150,200 mmol/L NaCl;there was a significantly negative correlation(r=-0.986) between NaCl concentrations and the level of DNA methylation in the roots of cotton seedlings. Compared to the control(0 mmol/L NaCl) the methylation ratio of root genomic DNA were 6.4%,7.6%and 11.3%respectively,and the demethylation ratio of root genomic DNA were 12.7%,11.1%and 8.2%respectively under the stresses of 100,150 and 200 mmol/L NaCl.In addition,the analysis of sequences suggested that parts of MSAP fragments were homologous to functional genes.The results of RT-PCR showed that M3 fragment homologous to gypsy retrotransposon reverse transcriptase gene of Gossypium barbadense and M2 fragment homologous to Gossypium hirsutum cDNA GH_TMO were expressed in the roots of cotton seedlings under 200 mmol/L NaCl stress,but were not expressed under 0 mmol/L NaCl stress.
棉花是一种重要的经济作物,利用基因工程技术对于提高棉花的产量、增强抗逆能力以及改良纤维品质具有重要作用。本研究以最终获得用于抗逆遗传改良的转基因棉花植株为目的,通过导入并超量表达对逆境起调控作用的转录因子基因——SNAC1基因,系统地对这些转基因再生棉花植株进行抗逆性分析和鉴定。初步鉴定出SNAC1基因在抗逆遗传改良中的有效性,并对SNAC1基因作为筛选标记基因用于棉花遗传转化进行了研究。另外,以棉花品系YZ1为材料对其盐胁迫下根组织DNA甲基化的水平及变化模式进行了研究。主要结果如下:
1.SNAC1基因是具有耐旱及耐盐的转录因子基因,本研究利用农杆菌介导的方法把SNAC1基因导入到陆地棉品系YZ1。PCR检测和RT-PCR分析表明,SNAC1基因已经整合到转基因棉花的基因组中且得到表达,共获得9个克隆系的再生植株及后代。田间农艺性状调查表明,转基因S1和S2家系的T_1代棉花植株在产量等农艺性状上与非转基因植株比较没有大的变异。棉花离体叶片失水速率测定表明,田间种植的转基因T_1代棉花植株叶片的失水速率为25%左右,低于非转基因植株叶片失水速率(30.4%)。同时,电镜扫描观察证实了离体叶片失水胁迫后转基因植株叶片的气孔关闭数量比例(71.8%)要高于野生型植株(51.5%)。水培抗盐性鉴定结果表明,在200 mmol/L NaCl胁迫5天后转基因T_2代植株幼苗的叶片相对含水量(RWC)为72%,而非转基因植株叶片的相对含水量为61%,并且非转基因植株叶片萎焉脱落要早于转基因植株。同时,转基因T_2代植株幼苗在高盐胁迫及胁迫后恢复过程中的相对生长速度(RGR)分别为7.9%和109.5%,高于非转基因植株的相对生长速度(分别为-7.4%和53.1%)。另外,在150 mmol/L NaCl胁迫下转基因胚性愈伤组织能够继续生长,而非转基因胚性愈伤组织则失水变白死亡。对转基因阳性和阴性T_1代植株的qRT-PCR分析结果表明,SNAC1基因能够调控棉花部分基因的表达,如G2(NAC familyprotein)与G3(NAM-like protein)基因的表达上升以及G5(GSA-AM2)、G11(SAMS)与AF38(ABF2)基因的表达下降。
2.本文以SNAC1基因作为筛选标记基因,采用农杆菌介导法将SNAC1基因导入棉花细胞,同时研究了NaCl作为筛选剂用于棉花遗传转化的可行性。经过棉花下胚轴的农杆菌菌液浸染、共培养以及NaCl筛选培养,获得了转基因棉花胚性愈伤组织。对获得的胚性愈伤组织进行PCR检测证实外源基因已经整合到棉花基因组中且阳性率达50%以上,同时GUS染色也表明GUS基因得到表达。另外,本文还对NaCl作为筛选剂在农杆菌介导的棉花遗传转化中的应用浓度及方法进行了研究,即:在农杆菌侵染后棉花下胚轴的共培养时间为3～4天,愈伤组织诱导初期的NaCl筛选浓度在1.1%～1.5%(W/V)之间,随着愈伤组织的生长其耐盐性有所增强而逐渐提高NaCl的筛选浓度。与正常培养条件相比,培养基中NaCl的存在不利于胚性愈伤组织的分化,因此经过3～4次愈伤组织继代筛选后要及时去除培养基中的NaCl以促进胚性愈伤组织的分化。
3.本研究以陆地棉品系YZ1为材料,调查不同NaCl浓度处理下棉花幼苗生长及根基因组DNA甲基化的水平和变化模式。结果表明,100 mmol/L NaCl对棉花幼苗的株高和根长生长有促进作用,而200 mmol/L NaCl则显著抑制株高和根长的生长。此外,在100～200 mmol/LNaCl胁迫下棉花幼苗的侧根数量被严重抑制。MSAP分析结果表明,经100、150和200 mmol/L NaCl胁迫处理后棉花幼苗根基因组DNA甲基化水平分别为38.1%、35.2%和34.5%,均低于对照(0 mmol/LNaCl)的甲基化水平(41.2%),同时棉花幼苗根DNA甲基化的水平与NaCl处理浓度呈显著负相关(r=-0.986)。与对照(0 mmol/L NaCl)相比,在100、150和200 mmol/LNaCl胁迫下棉花幼苗根基因组DNA发生甲基化的比率分别为6.4%、7.6%、11.3%,而去甲基化的比率分别为12.7%、11.1%、8.2%.序列分忻结果显示,部分MSAP差异片段与功能基因同源。RT-PCR分析表明,分别与MSAP差异片段M2及M3高度同源的海岛棉gyspy反转座子反转录酶基因及陆地棉cDNA基因在处理(200 mmol/LNaCI)的棉花幼苗根中有表达,但在对照(0 mmol/L NaCl)的根中则没有表达。
Plant's adaptability to abiotic stresses such as drought,cold and salinity plays an important role in plant's yield and quality.Drought and salinity stresses are the most common form of adversity,which are the limiting factors for agricultural development in many areas.Therefore,it is important to improve crop's tolerance to stresses by genetic engineering.Plant genomic DNA was always methylated under stresses and studying the level and pattern of genomic DNA methylation under stresses will help to identify the mechanisms on stress resistance.
Cotton is one of the most important economic crops in the world,and genetic engineering played an important role in increasing cotton's yield,enhancing stress resistance and improving cotton fiber quality.The objective of this study was to gain transgenic cotton plants for genetic improvement,SNAC1 gene,a transcriptional factor regulating stress response,which was transformed into cotton and the analysis and identification of stress resistance were carried out on transgenic cotton plants systematically.The availability of SNAC1 gene was indentified preliminarily in genetic improvement and SNAC1 gene as a selection marker gene for genetic transformation of cotton was studied also.In addition,the cotton strain of YZ1 was used to study the level and pattern of DNA methylation in cotton roots under salt stress.The following results were obtained:
1.SNAC1 gene in rice can enhance drought- and salt-tolerance,and then was transformed into Gossypium hirsutum YZ1 by Agrobacterium -mediated genetic transformation in this study.PCR detection and RT-PCR analysis indicated SNAC1 gene had been integrated into the transgenic cotton genome and at the same time regenerated plants of 9 clones were gained totally.The field surveys showed that T_1 generation of transgenic plants did not vary in many agronomic traits such as yield,compared to non-transgenic plants.The determination of rate of water loss of excised-leaves of cotton demonstrated that the RWL(rate of water loss) of T_1 generationof transgenic plants was 25%,and was lower than that(30.4%) of the non-transgenic plants.Then the observation of SEM(scanning electron microscope) also suggested the ratio of stomatal closure in the leaves of transgenic plants was 71.8%and was higher than that(51.5%) of non-transgenic plants.The results of salt-tolerance identification by water culture showed that RWC (Relative water content) of leaves of T_2 generation of transgenic seedlings was 72%under stress of 200 mmol/L NaCl after 5 d and that of non-transgenic cotton's leaves was 61%. What is more,the leaves of non-transgenic plants withered and broken off earlier than that of transgenic plants.In addion,the RGR(Relative growth rate) of transgenic plant seedlings were 7.9%and 109.5%respectively during salt stress and sress revovery,and the RGR of non-transgenic plants were only-7.4%and 53.1%respectively.Under the 150 mmol/L NaCl stess transgenic embryogenic calli can continue to grow,but non-transgenic embryogenic calli lost water and died.The results of qRT-PCR analysis suggested that in T_1 generation of transgenic positive and negative plants SNAC1 gene could regulate expressions of some genes in cotton,for example expressions of G2(NAC family protein) and G3(NAM-like protein) genes were up-regulated and expressions of G5(GSA-AM2), G11(SAMS) and AF38(ABF2) genes were down-regulated.
2.In this study,employing SNAC1 gene was selected as a marker gene and NaCl as a selection agent,SNAC1(Stress-induced NAC 1) and GUS genes were introduced into cotton genome via Agrobacterium-mediated transformation,and the feasibility of NaCl as a selection agent for the genetic transformation of cotton was researched.Finally transgenic embryonic calli were gained after the infection with Agrobacterium,co-culture and selection culture of NaCl.Transgenic calli were confirmed by PCR analysis and positive rate of transgenic calli was 58%,and at the same time expression of the GUS gene was showed with GUS staining.Furthermore the concentration of NaCl and method for employing NaCl as a selection agent had been studied.The hypocotyls were co-cultured for 3～4 d after Agrobaterium infection.The reasonable selection concentration of NaCl should be 1.1%～1.5%(W/V) and the start concentration of NaCl in the callus induction medium should be lower.Then the concentration was increased as cotton calli proliferated.Because NaCl is not beneficial for embryo differentiation and development,NaCl should be removed from the culture medium to promote embryonic calli differentiation after calli were subcultured for 3～4 times.
3.In this study the growth of cotton seedlings and the level and patterns of DNA methylation in the roots of cotton were investigated uder salt stress.The results showed that 100 mmol/L NaCl obviously promoted plant height and root length of cotton seedlings,but 200 mmol/L NaCl significantly inhibited the growth of cotton seedlings; 100～200 mmol/L NaCl inhibited the numbers of lateral root considerably.The analysis of MSAP showed that the level of global DNA methylation in the roots of cotton seedlings was 41.2%,38.1%,35.2%and 34.5%respectively under the stresses of 0,100,150,200 mmol/L NaCl;there was a significantly negative correlation(r=-0.986) between NaCl concentrations and the level of DNA methylation in the roots of cotton seedlings. Compared to the control(0 mmol/L NaCl) the methylation ratio of root genomic DNA were 6.4%,7.6%and 11.3%respectively,and the demethylation ratio of root genomic DNA were 12.7%,11.1%and 8.2%respectively under the stresses of 100,150 and 200 mmol/L NaCl.In addition,the analysis of sequences suggested that parts of MSAP fragments were homologous to functional genes.The results of RT-PCR showed that M3 fragment homologous to gypsy retrotransposon reverse transcriptase gene of Gossypium barbadense and M2 fragment homologous to Gossypium hirsutum cDNA GH_TMO were expressed in the roots of cotton seedlings under 200 mmol/L NaCl stress,but were not expressed under 0 mmol/L NaCl stress.
引文
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