转rol ABC基因枳橙砧木的评价
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摘要
柑橘是我国重要的果树,培育矮化砧木和接穗品种一直是柑橘育种的重要目标之一。柑橘天然矮化砧木种质极少,传统育种因面临漫长的童期、高度杂合,以及珠心胚干扰等问题,目前进展缓慢。应用矮化基因,通过基因工程技术将一个或多个基因转入到柑橘基因组中,可望加快柑橘砧木矮化育种的进程。
rol ABC基因克隆于Agrobactecium rhizogenes菌A4的Ri质粒的TL-DNA区,转入rol基因可诱发植株出现节间变短,分枝增多,顶端优势散失,大量毛根形成,植株显著矮化等性状,rol基因的表达与植物矮化性状形成关系密切,其诱发的大量毛状根在提高植物抗逆性方面引起重视,通过遗传转化技术将rol基因转入林木和果树砧木的研究近年来已有报道。
枳橙[Citrus sinensis (L.) Osb x Poncirus trifoliate (L.) Raf.]是一种重要的柑橘杂交砧木,因具有抗柑橘衰退病和脚腐病特性,在欧美国家已成为新兴的柑橘砧木被大力推广应用,如进一步改良获得矮化、抗性枳橙将有更大的应用价值。
本实验室及合作者通过农杆菌介导法己将rol ABC基因转入Troyer枳橙中,并通过组织培养繁殖B、D、E三个转rol ABC基因枳橙株系,转基因枳橙表现出典型的rol基因性状,根系也产生大量毛状根。在此基础上,本研究系统阐述了转rol ABC基因枳橙的分子基础、形态特征、生理特性、外源基因表达模式变化,并研究其用作基砧、中间砧对柑橘接穗生长发育的影响,探索转rol ABC基因枳橙自身及诱导接穗矮化的机理,并按照农业部转基因安全办要求,对转rol ABC基因枳橙砧木的安全性进行了综合评价。主要研究结论如下：
1、PCR、Southern杂交鉴定确定,rol ABC基因已经转入到Troyer枳橙,B,D系插入一个拷贝,E系中插入两个拷贝,Semi-RT PCR和qRT PCR证实rol A、rol C基因在转基因B、D、E系中均能正常转录表达,其中rol C基因表达量最高,其次是rol A基因,rol B基因转录水平十分微弱。
2、转rol基因枳橙及其实生后代表现出典型的rol基因形态特征,E系生长势较旺盛,植株较高,表现出更好的环境适应性,基部分枝相比B、D系更多。三者均表现良好的嫁接亲和力。B、D、E系转基因枳橙具有狭长而密集的导管结构,单位面积细胞数未有减少；光合速率明显高于野生植株,叶绿素含量增加与光合效率提高有密切关系；SOD、POD、Pro均显著提高,多年栽培表现出良好的抗逆性。
3、建立适合于rol A、rol B、rol C基因实时荧光定量RT-PCR分析方法。2009年6月15至9月15,每隔15天,采取转基因枳橙B、D、E系7个组织(芽、幼茎、嫩叶、功能叶、皮、老叶、根),相对定量法分析外源基因的时空表达与植株生长的关系。发现在B、D、E系转rol基因枳橙中,rol C基因表达最强,其次是rol A, rol B基因表达水平最低。rol A、rol C基因表达在6月30日和7月30日呈现两个表达峰值,7月15日呈现一个明显的低谷,并且证实高温抑制生长与rol基因表达下调有关。rol A基因表达主要在老叶、芽、功能叶中表达,rolB基因主要在根系中表达,rol C基因主要在嫩茎、芽和功能叶中表达。取4个时间点,分析rol基因时空表达与矮化性状的相关性发现：植株生长高度减少率(plant height reduced rate, RPHRR)与rol A(r=0.75, p<0.01), rol C(r=0.74, P＜0.01)基因在韧皮部的转录水平成正相关,除根中rol基因的表达外,rol A、rol C基因在其它6个组织中的表达也直接促进转基因枳橙矮化。
4、转rol基因枳橙既不属于GA缺陷型,也不属于GA不敏感型,喷施GA3能促进其茎伸长生长,但恢复不到野生型水平。幼芽中GA1、GA4和IAA显著降低(P＜0.05), POD酶活性显著提高(P<0.05)。转rol基因枳橙幼芽中GA20oxl基因mRNA水平相比对照显著下调(P<0.01)。在幼芽、嫩茎中,rol C基因与GA20ox1表达负相关。故认为rol基因通过在幼芽中的高表达下调了GA20ox1基因转录表达,进而抑制了活性GAs在幼芽中的合成,顶端分生组织较低量的活性GAs限制植物茎伸长。
5、转rol ABC基因枳橙果实种子变小,且从楔形变成椭圆形,萌发比野生植株晚一周左右。T1代在幼苗期植株就表现出典型的rol基因矮化性状,萌芽率有所下降,99%为珠心苗,生长一致,但主干粗度有所下降。PCR, RT-PCR鉴定确认rol基因在实生后代表达同T0代,认为转rol ABC基因枳橙T1代能稳定遗传rol基因的典型性状。
6、以转rol ABC基因枳橙B、D、E做基砧和中间砧木,嫁接大红甜橙、沙田柚,嫁接4年后,提取接穗DNA进行PCR和qRT PCR分析,确证外源基因不能通过嫁接转移。rol ABC基因枳橙一年生嫁接树光合作用明显提升,四年生嫁接树未见与对照有差别；两个接穗品种均表现12%-30%不同程度的诱导矮化效果；2009年和201O年连续2年果实品质分析发现,E系上嫁接大红甜橙,果实总糖含量分别上升14.2%和19.3%。
Citrus is one of the most important fruit crops in the world. In the past tens of years, obtaining dwarf characteristics was one of the main goals of citrus genetic improvement. There are no useful dwarfing rootstocks for citrus (Deng 1994; Grosser et al.,1988). Citrus traditional hybridization has made slow progress due to long juvenile period, high heterozygosity, sexual incompatibility and nucellar embryony. Biotechnology methods allow the introduction of one or few genes of dwarf genes into a citrus genome relatively quickly, and offer an alternative way to promote citrus rootstock dwarfing breeding.
The rol A, rol B and rol C genes of Agrobacterium rhizogenes A4 are located on the TL-DN A of the Ri plasmid. rol genes have been introduced into a range of crops, and a wide of morphological and physiological changes occurred, these include dwarfing effects on plant height due to short internodes, wrinkled leaves and delayed flower timing, increased branching associated with loss of apical dominance, and inducing full hairy-root syndrome. Recently, rol genes were used to transform fruit trees for dwarfing breeding, especially for creating dwarfing rootstock with tolerance to environmental stress.
Troyer citrange [Citrus sinensis (L.) Osb.χPoncirus trifoliata (L.) Raf.] is an important hybridization rootstock in citriculture. Which were find widespread use in European and America countries due to resist citrus tristeza virus (CTV) and citrus foot rot. To obtain dwarf and resistance by genetic improvement could enhance the value of Troyer citrange to a certain extent.
Previously, rol ABC genes were transferred into Troyer citrange and three clones were obtained. These transgenic clones were propagated by tissue culture during the past six years. Many years observations showed that transgenic clones had typical dwarfing characters (up to 50% height reduction), and increased rooting ability. In the present study, morphological characteristics of rol ABC transgenic citrange clones were observed, molecular confirmation and expression of transgenes were investigated at different time intervals and in various tissues using qRT-PCR, many aspects of physiological property were determined, and the rootstock-scions interaction were surveyed, with the aim of evaluating the potential of transgenic Troyer citrange and understanding the induction mechanism of dwarfing by rol genes.
1. PCR. southern blot analysis revealed that rol A, rol B and rol C were integrated into the citrange genome, exactly two copies in clones B and D, and one in clone E. Semi RT-PCR and quantitative RT-PCR results indicated that rol A and rol C were over-expressed, with the highest expression for rol C, while rol B had a very weak expression signal.
2. The transgenic citrange clones B, D and E showed typical dwarfing characteristics of rol gene. E clone showed stronger growth vigour and better environmental resistance, also the lateral branches growth thickly compare with B, D clones. Three clones have a narrow and intensive vessel structure, the cell number per cm2 was not decreased, all transgenic citranges reveal good grafting compatibility with Dahong sweet orange and Shatian pumelo, POD, SOD, Pro and chlorophyll content all dramatically increased compare to wild type citrange, and transgenic citrange show an stronger photosynthesis and environmental resistance.
3. Developing suitable qRT-PCR method for analysising the expression of rol A, rol B, rol C genes in transgenic citrange. From 15 June to 15 September in 2009. seven plant tissues including buds, tender leaves, young stems, functional leaves, old leaves, bark and roots were used as materials for q RT PCR analysis, at the same stage, 10 plants per transgenic clone and wild-type citrange were chosen for morphological observations. Try to study the relation between rol genes'expression and plant growth. The result indicated that rol C gene had the highest relative expression in all tested tissues except old leaves, rol B had the lowest expression, and the expression of rol A was intermediate of the three genes. The expression of rol A and rol C showed two peaks on 30 June and 30 July, and one lowest point on 15 July. The temporal changes of rol genes'expression were found to be influenced by environmental condition. On July 15, the highest temperature was supressed rol genes'expression and restrained plant growth activity. The expression of rol A was mainly in old leaves, buds and functional leaves. Expression of rol B was highest in roots, and very low in other tissues. Expression of rol C was primarily in buds, tender stems and functional leaves, The summed expressions of the rol genes at four different sampling times (15 June, 15 July,15 August and 15 September), the relationship between the expression of rol genes in single tissues and RPHRR were showed the expression of rol A and rol C in bark was significantly positively correlated with RPHRR (r=0.75 and r=0.74, respectively; P<0.01). The expression of rol A and rol C in buds, tender stems, tender leaves, functional leaves and old leaves was significantly correlated with their expression in bark. This indicates that expression of rol A and rol C in these tissues was also involved in the induction of dwarfing.
4. Transgenic citrange was neither defective genotype of GA3, nor unsensitive genotype of GA3. However, the spraying of GA3 elongated the internode of transgenic citrange, it was still shorter than wild type citrange. The content of IAA. GA] and GA4 decreased (P<0.05) while peroxidase activity increased markedly in the tender buds of transgenic citrange (P<0.05). The GA20ox1 expression quantity markedly decreased in apical bud (P＜0.01), the expression of GA20oxl in tender leaves was higher than wild type. The expression of rol C gene and GA20oxl gene reveal negative correlation in tender bud, tender stems, we supposed that rol genes' expression could inhibit GAs'synthesis by down regulate the expression of GA20αχl gene in bud, lower bioactive GAs could suppress citrange growth.
5. The seed of transgenic citrange was smaller than CK, the shape was turned into ellipse from wedge-shape. Germination was later than CK about seven days, the germination rate of T1 seed was decreased partly, dwarf character was exhibited in young seedling.99% of the seedlings were originated from nucellus and growing congruously, However, the trunk diameter of transgenic citrange was smaller compare with CK plants, rol genes were steadily inherited to next generation, which was detected by PCR and RT-PCR.
6. Taking transgenic citrange as inter-rootstock, we grafted Shatian pummel (Citrus grandis [L.] Osbeck cv. Shatian) and Dahong sweet orange on it, rol gene weren't detected in the leaves of scions by PCR and qRT PCR after 4 years, the result indicated rol gene didn't transfer into scions through grafting. The photosynthesis (Pn) ability of Shatian Pummelo scions enhanced compared to control rootstocks after grafting for one year, but the Pn difference of scions wasn't observed after 4 years. The dwarfing rate of scions were from 12%-30% after grafted on transgenic rootstock or inter-rootstock. The fruit quality improved dramatically, the total sugar of E-Dahong was increased 14.2% and 19.3% compare to CK-Dahong in 2009 and in 2010, respectively.
rol ABC基因克隆于Agrobactecium rhizogenes菌A4的Ri质粒的TL-DNA区,转入rol基因可诱发植株出现节间变短,分枝增多,顶端优势散失,大量毛根形成,植株显著矮化等性状,rol基因的表达与植物矮化性状形成关系密切,其诱发的大量毛状根在提高植物抗逆性方面引起重视,通过遗传转化技术将rol基因转入林木和果树砧木的研究近年来已有报道。
枳橙[Citrus sinensis (L.) Osb x Poncirus trifoliate (L.) Raf.]是一种重要的柑橘杂交砧木,因具有抗柑橘衰退病和脚腐病特性,在欧美国家已成为新兴的柑橘砧木被大力推广应用,如进一步改良获得矮化、抗性枳橙将有更大的应用价值。
本实验室及合作者通过农杆菌介导法己将rol ABC基因转入Troyer枳橙中,并通过组织培养繁殖B、D、E三个转rol ABC基因枳橙株系,转基因枳橙表现出典型的rol基因性状,根系也产生大量毛状根。在此基础上,本研究系统阐述了转rol ABC基因枳橙的分子基础、形态特征、生理特性、外源基因表达模式变化,并研究其用作基砧、中间砧对柑橘接穗生长发育的影响,探索转rol ABC基因枳橙自身及诱导接穗矮化的机理,并按照农业部转基因安全办要求,对转rol ABC基因枳橙砧木的安全性进行了综合评价。主要研究结论如下：
1、PCR、Southern杂交鉴定确定,rol ABC基因已经转入到Troyer枳橙,B,D系插入一个拷贝,E系中插入两个拷贝,Semi-RT PCR和qRT PCR证实rol A、rol C基因在转基因B、D、E系中均能正常转录表达,其中rol C基因表达量最高,其次是rol A基因,rol B基因转录水平十分微弱。
2、转rol基因枳橙及其实生后代表现出典型的rol基因形态特征,E系生长势较旺盛,植株较高,表现出更好的环境适应性,基部分枝相比B、D系更多。三者均表现良好的嫁接亲和力。B、D、E系转基因枳橙具有狭长而密集的导管结构,单位面积细胞数未有减少；光合速率明显高于野生植株,叶绿素含量增加与光合效率提高有密切关系；SOD、POD、Pro均显著提高,多年栽培表现出良好的抗逆性。
3、建立适合于rol A、rol B、rol C基因实时荧光定量RT-PCR分析方法。2009年6月15至9月15,每隔15天,采取转基因枳橙B、D、E系7个组织(芽、幼茎、嫩叶、功能叶、皮、老叶、根),相对定量法分析外源基因的时空表达与植株生长的关系。发现在B、D、E系转rol基因枳橙中,rol C基因表达最强,其次是rol A, rol B基因表达水平最低。rol A、rol C基因表达在6月30日和7月30日呈现两个表达峰值,7月15日呈现一个明显的低谷,并且证实高温抑制生长与rol基因表达下调有关。rol A基因表达主要在老叶、芽、功能叶中表达,rolB基因主要在根系中表达,rol C基因主要在嫩茎、芽和功能叶中表达。取4个时间点,分析rol基因时空表达与矮化性状的相关性发现：植株生长高度减少率(plant height reduced rate, RPHRR)与rol A(r=0.75, p<0.01), rol C(r=0.74, P＜0.01)基因在韧皮部的转录水平成正相关,除根中rol基因的表达外,rol A、rol C基因在其它6个组织中的表达也直接促进转基因枳橙矮化。
4、转rol基因枳橙既不属于GA缺陷型,也不属于GA不敏感型,喷施GA3能促进其茎伸长生长,但恢复不到野生型水平。幼芽中GA1、GA4和IAA显著降低(P＜0.05), POD酶活性显著提高(P<0.05)。转rol基因枳橙幼芽中GA20oxl基因mRNA水平相比对照显著下调(P<0.01)。在幼芽、嫩茎中,rol C基因与GA20ox1表达负相关。故认为rol基因通过在幼芽中的高表达下调了GA20ox1基因转录表达,进而抑制了活性GAs在幼芽中的合成,顶端分生组织较低量的活性GAs限制植物茎伸长。
5、转rol ABC基因枳橙果实种子变小,且从楔形变成椭圆形,萌发比野生植株晚一周左右。T1代在幼苗期植株就表现出典型的rol基因矮化性状,萌芽率有所下降,99%为珠心苗,生长一致,但主干粗度有所下降。PCR, RT-PCR鉴定确认rol基因在实生后代表达同T0代,认为转rol ABC基因枳橙T1代能稳定遗传rol基因的典型性状。
6、以转rol ABC基因枳橙B、D、E做基砧和中间砧木,嫁接大红甜橙、沙田柚,嫁接4年后,提取接穗DNA进行PCR和qRT PCR分析,确证外源基因不能通过嫁接转移。rol ABC基因枳橙一年生嫁接树光合作用明显提升,四年生嫁接树未见与对照有差别；两个接穗品种均表现12%-30%不同程度的诱导矮化效果；2009年和201O年连续2年果实品质分析发现,E系上嫁接大红甜橙,果实总糖含量分别上升14.2%和19.3%。
Citrus is one of the most important fruit crops in the world. In the past tens of years, obtaining dwarf characteristics was one of the main goals of citrus genetic improvement. There are no useful dwarfing rootstocks for citrus (Deng 1994; Grosser et al.,1988). Citrus traditional hybridization has made slow progress due to long juvenile period, high heterozygosity, sexual incompatibility and nucellar embryony. Biotechnology methods allow the introduction of one or few genes of dwarf genes into a citrus genome relatively quickly, and offer an alternative way to promote citrus rootstock dwarfing breeding.
The rol A, rol B and rol C genes of Agrobacterium rhizogenes A4 are located on the TL-DN A of the Ri plasmid. rol genes have been introduced into a range of crops, and a wide of morphological and physiological changes occurred, these include dwarfing effects on plant height due to short internodes, wrinkled leaves and delayed flower timing, increased branching associated with loss of apical dominance, and inducing full hairy-root syndrome. Recently, rol genes were used to transform fruit trees for dwarfing breeding, especially for creating dwarfing rootstock with tolerance to environmental stress.
Troyer citrange [Citrus sinensis (L.) Osb.χPoncirus trifoliata (L.) Raf.] is an important hybridization rootstock in citriculture. Which were find widespread use in European and America countries due to resist citrus tristeza virus (CTV) and citrus foot rot. To obtain dwarf and resistance by genetic improvement could enhance the value of Troyer citrange to a certain extent.
Previously, rol ABC genes were transferred into Troyer citrange and three clones were obtained. These transgenic clones were propagated by tissue culture during the past six years. Many years observations showed that transgenic clones had typical dwarfing characters (up to 50% height reduction), and increased rooting ability. In the present study, morphological characteristics of rol ABC transgenic citrange clones were observed, molecular confirmation and expression of transgenes were investigated at different time intervals and in various tissues using qRT-PCR, many aspects of physiological property were determined, and the rootstock-scions interaction were surveyed, with the aim of evaluating the potential of transgenic Troyer citrange and understanding the induction mechanism of dwarfing by rol genes.
1. PCR. southern blot analysis revealed that rol A, rol B and rol C were integrated into the citrange genome, exactly two copies in clones B and D, and one in clone E. Semi RT-PCR and quantitative RT-PCR results indicated that rol A and rol C were over-expressed, with the highest expression for rol C, while rol B had a very weak expression signal.
2. The transgenic citrange clones B, D and E showed typical dwarfing characteristics of rol gene. E clone showed stronger growth vigour and better environmental resistance, also the lateral branches growth thickly compare with B, D clones. Three clones have a narrow and intensive vessel structure, the cell number per cm2 was not decreased, all transgenic citranges reveal good grafting compatibility with Dahong sweet orange and Shatian pumelo, POD, SOD, Pro and chlorophyll content all dramatically increased compare to wild type citrange, and transgenic citrange show an stronger photosynthesis and environmental resistance.
3. Developing suitable qRT-PCR method for analysising the expression of rol A, rol B, rol C genes in transgenic citrange. From 15 June to 15 September in 2009. seven plant tissues including buds, tender leaves, young stems, functional leaves, old leaves, bark and roots were used as materials for q RT PCR analysis, at the same stage, 10 plants per transgenic clone and wild-type citrange were chosen for morphological observations. Try to study the relation between rol genes'expression and plant growth. The result indicated that rol C gene had the highest relative expression in all tested tissues except old leaves, rol B had the lowest expression, and the expression of rol A was intermediate of the three genes. The expression of rol A and rol C showed two peaks on 30 June and 30 July, and one lowest point on 15 July. The temporal changes of rol genes'expression were found to be influenced by environmental condition. On July 15, the highest temperature was supressed rol genes'expression and restrained plant growth activity. The expression of rol A was mainly in old leaves, buds and functional leaves. Expression of rol B was highest in roots, and very low in other tissues. Expression of rol C was primarily in buds, tender stems and functional leaves, The summed expressions of the rol genes at four different sampling times (15 June, 15 July,15 August and 15 September), the relationship between the expression of rol genes in single tissues and RPHRR were showed the expression of rol A and rol C in bark was significantly positively correlated with RPHRR (r=0.75 and r=0.74, respectively; P<0.01). The expression of rol A and rol C in buds, tender stems, tender leaves, functional leaves and old leaves was significantly correlated with their expression in bark. This indicates that expression of rol A and rol C in these tissues was also involved in the induction of dwarfing.
4. Transgenic citrange was neither defective genotype of GA3, nor unsensitive genotype of GA3. However, the spraying of GA3 elongated the internode of transgenic citrange, it was still shorter than wild type citrange. The content of IAA. GA] and GA4 decreased (P<0.05) while peroxidase activity increased markedly in the tender buds of transgenic citrange (P<0.05). The GA20ox1 expression quantity markedly decreased in apical bud (P＜0.01), the expression of GA20oxl in tender leaves was higher than wild type. The expression of rol C gene and GA20oxl gene reveal negative correlation in tender bud, tender stems, we supposed that rol genes' expression could inhibit GAs'synthesis by down regulate the expression of GA20αχl gene in bud, lower bioactive GAs could suppress citrange growth.
5. The seed of transgenic citrange was smaller than CK, the shape was turned into ellipse from wedge-shape. Germination was later than CK about seven days, the germination rate of T1 seed was decreased partly, dwarf character was exhibited in young seedling.99% of the seedlings were originated from nucellus and growing congruously, However, the trunk diameter of transgenic citrange was smaller compare with CK plants, rol genes were steadily inherited to next generation, which was detected by PCR and RT-PCR.
6. Taking transgenic citrange as inter-rootstock, we grafted Shatian pummel (Citrus grandis [L.] Osbeck cv. Shatian) and Dahong sweet orange on it, rol gene weren't detected in the leaves of scions by PCR and qRT PCR after 4 years, the result indicated rol gene didn't transfer into scions through grafting. The photosynthesis (Pn) ability of Shatian Pummelo scions enhanced compared to control rootstocks after grafting for one year, but the Pn difference of scions wasn't observed after 4 years. The dwarfing rate of scions were from 12%-30% after grafted on transgenic rootstock or inter-rootstock. The fruit quality improved dramatically, the total sugar of E-Dahong was increased 14.2% and 19.3% compare to CK-Dahong in 2009 and in 2010, respectively.
引文
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