ABA介导的PacMYBA调控红肉甜樱桃果实花色苷合成的研究
摘要
植物中的花色苷属于水溶性色素,在果实和蔬菜中主要呈现出蓝色,紫色和红色。植物花色苷合成路径属于苯丙氨酸解氨酶途径中的类黄酮分支。近期研究表明,一类R2R3-MYB转录因子对花色苷合成具有显著地调控作用。本研究在分析了甜樱桃果实发育动态及相关生理指标变化的基础上,克隆了甜樱桃果实花色苷合成路径关键酶基因和R2R3-MYB转录因子,开展了甜樱桃花色苷合成和调控机制的研究,主要研究结果如下：
(1)对甜樱桃果实发育做了果实发育动态及生理指标分析。发现甜樱桃果实发育呈现典型的双“s”动态发育过程。果实可溶性糖包括葡萄糖,果糖和山梨醇,蔗糖未检出。果实叶绿素含量在小绿果时期后呈现下降趋势,而花色苷含量在转色期后迅速增加。
(2)首次利用RACE技术克隆得到甜樱桃花色苷合成路径5个关键酶基因全长序列和部分CHS基因序列,依次命名为：PacCHS, PacCHI, PacF3H, PacDFR, PacANS, PacUFGT。 qRT-PCR分析表明,在甜樱桃果实发育前期PacCHS, PacCHI和PacF3H表达水平逐渐升高,果实转色期后表达量迅速下降,临近成熟又快速提高。而PacDFR, PacANS和PacUFGT则在果实发育前期一直较低的表达水平,在果实转色期后则逐渐上升并且维持在一个较高的表达水平。
(3)克隆得到甜樱R2R3-MYB类转录因子命名为PacMYBA。蛋白序列比对发现PacMYBA与已知的花色苷调控相关的转录因子MdMYB10在R2R3区域相似度达到84%,全蛋白序列相似率达到68%。亚细胞定位发现PacMYBA定位于细胞核中,且没有转录自激活活性。组织特异性表达分析表明PacMYBA在成熟的果皮和果肉中表达量最高。在甜樱桃果实发育进程中PacMYBA的表达与果实花色苷积累呈现显著相关。在PacMYBA启动子中发现了多个参与植物生长发育响应、逆境响应和植物激素响应的顺式作用元件。
(4)在模式植物拟南芥中过表达PacMYBA,转基因植株授粉后10d内荚果果皮变红,而野生型植株荚果依然为绿色,表明PacMYBA具有调控花色苷合成的作用。应用病毒介导的基因沉默手段(VIGS)成功地沉默了发育果实中的PacMYBA,注射含有PacMYBA-TRV2和TRV1的樱桃果实相对于注射空TRV载体果实没有正常地积累花色苷物质,说明PacMYBA直接参与调控了甜樱桃果实花色苷物质的合成。
(5)转色期前外源施用ABA可以显著促进樱桃果实花色苷的合成,ABA生物合成抑制剂NDGA处理则显著抑制了果实花色苷的合成,说明外源施加ABA对果实花色苷的合成具有促进作用。用30μmol/L ABA温育溶液处理甜樱桃果实圆片,PacMYBA表达量迅速升高,处理后1h即显著高于对照；用NDGA处理后PacMYBA表达量始终低于对照和ABA处理,表明PacMYBA的表达可能受ABA调控.用VIGS技术沉默ABA生物合成途径中关键酶基因PacNCEDl可以降低甜樱桃果实内源ABA含量,且显著抑制了甜樱桃果实花色苷的合成。6个花色苷合成路径关键酶基因和PacMYBA的相对表达量在PacNCED1-RNAi果实中显著低于TRV空载体和对照果实。这进一步说明ABA作为一种信号分子参与了甜樱桃果实花色苷的合成。
Anthocyanins are the major water-soluble pigments in higher plants and are responsible for the blue, purple, and red colors of many fruits and vegetables. The anthocyanin biosynthetic pathway is part of the flavonoid pathway, which is a branch of the phenylpropanoid pathway. Recent studies have shown that a kind of R2R3MYB transcription factors have significantly regulation effect on anthocyanin biosynthesis. The present study describes the identification and functional characterization of the sweet cherry R2R3MYB protein, PacMYBA and also first obtained five full-length genes of anthocyanin biosynthesis pathway. The present work also first systematically expounds the role of plant hormone ABA in the fruit development process of sweet cherry. The main results are follows:
(1) In the present study we have studied the physiological and morphological changes during sweet cherry fruit development. The results showed that sweet cherry exhibited a biphasic growth pattern. The soluble sugars of sweet cherry including:glucose, fructose and sorbitol. No sucrose was detected in the sweet cherry. The levels of Chl declined continually after the SG stage, whereas the anthocyanin content increased rapidly after veraison.
(2) We first obtained the anthocyanin biosynthesis pathway genes of sweet cherry by using RACE technology and named as PacCHS, PacCHI, PacF3H, PacDFR, PacANS, and PacUFGT. The expression of PacCHS, PacCHI and PacF3H were at high levels during the early stages, declined during veraison (i.e., the onset of ripening; DQ YW, IR), and increased again in the last two stages. In accordance with the pattern of anthocyanin accumulation during ripening, the expression of PacDFR, PacANS and PacUFGT were expressed at low levels during the early developmental stages, and expression rapidly increased during veraison and remained at high levels through to the final developmental stage.
(3) We obtained a single putative R2R3MYB gene product, which we named PacMYBA, using RACE (rapid amplification of cDNA ends) PCR. Compared to apple MdMYB10, PacMYBA shared84%amino acid identity in the R2R3DNA-binding domain and68%identity over the whole protein. The nuclear localization of PacMYBA was consistent with its predicted function as a transcription factor and it is has no self-transcriptional activation activity. Tissue specific expression analysis showed that PacMYBA have the highest expression level in mature skin and flesh. The expression level of PacMYBA during sweet cherry fruit development is positive correlation with the anthocyanin biosynthesis of sweet cherry. We also found several cis-elements which related to plant growth and development response, stress response and plant hormone response in the promoter sequence of PacMYBA.
(4) The immature seeds of transgenic plants displayed red pigmentation<10days after anthesis, whereas the seeds of wild-type plants (ecotype Columbia) had no pigmentation. These data show that PacMYBA plays a role in anthocyanin biosynthesis.Silencing of PacMYBA, using a Tobacco rattle virus-induced gene silencing technique, resulted in sweet cherry fruit that lacked red pigment. Those fruits which were infiltrated with TRV alone pigmentation normally. These results shown that PacMYBA involve in regulating anthocyanin biosynthesis of sweet cherry directly.
(5) Abscisic acid (ABA) treatment significantly induced anthocyanin accumulation, while treatment with the ABA biosynthesis inhibitor nordihydroguaiaretic acid (NDGA) blocked anthocyanin production. The exoression level of PacMYBA increased quickly treated with30μmol/L ABA. After1h treatment with ABA the expression level of PacMYBA was significantly higher than control. This result suggested that PacMYBA might be a ABA-response gene.The uncolored phenotype was also observed in the fruits, silencing of PacNCED1, which encodes a key enzyme in the ABA biosynthesis pathway. The endogenous ABA content as well as the transcript levels of six structural genes and PacMYBA in PacNCED1-RNAi fruit was significantly lower than in the TRV vector control fruit. These results suggest that ABA is a signal molecule that promotes red-color sweet cherry fruit accumulating anthocyanin.
(1)对甜樱桃果实发育做了果实发育动态及生理指标分析。发现甜樱桃果实发育呈现典型的双“s”动态发育过程。果实可溶性糖包括葡萄糖,果糖和山梨醇,蔗糖未检出。果实叶绿素含量在小绿果时期后呈现下降趋势,而花色苷含量在转色期后迅速增加。
(2)首次利用RACE技术克隆得到甜樱桃花色苷合成路径5个关键酶基因全长序列和部分CHS基因序列,依次命名为：PacCHS, PacCHI, PacF3H, PacDFR, PacANS, PacUFGT。 qRT-PCR分析表明,在甜樱桃果实发育前期PacCHS, PacCHI和PacF3H表达水平逐渐升高,果实转色期后表达量迅速下降,临近成熟又快速提高。而PacDFR, PacANS和PacUFGT则在果实发育前期一直较低的表达水平,在果实转色期后则逐渐上升并且维持在一个较高的表达水平。
(3)克隆得到甜樱R2R3-MYB类转录因子命名为PacMYBA。蛋白序列比对发现PacMYBA与已知的花色苷调控相关的转录因子MdMYB10在R2R3区域相似度达到84%,全蛋白序列相似率达到68%。亚细胞定位发现PacMYBA定位于细胞核中,且没有转录自激活活性。组织特异性表达分析表明PacMYBA在成熟的果皮和果肉中表达量最高。在甜樱桃果实发育进程中PacMYBA的表达与果实花色苷积累呈现显著相关。在PacMYBA启动子中发现了多个参与植物生长发育响应、逆境响应和植物激素响应的顺式作用元件。
(4)在模式植物拟南芥中过表达PacMYBA,转基因植株授粉后10d内荚果果皮变红,而野生型植株荚果依然为绿色,表明PacMYBA具有调控花色苷合成的作用。应用病毒介导的基因沉默手段(VIGS)成功地沉默了发育果实中的PacMYBA,注射含有PacMYBA-TRV2和TRV1的樱桃果实相对于注射空TRV载体果实没有正常地积累花色苷物质,说明PacMYBA直接参与调控了甜樱桃果实花色苷物质的合成。
(5)转色期前外源施用ABA可以显著促进樱桃果实花色苷的合成,ABA生物合成抑制剂NDGA处理则显著抑制了果实花色苷的合成,说明外源施加ABA对果实花色苷的合成具有促进作用。用30μmol/L ABA温育溶液处理甜樱桃果实圆片,PacMYBA表达量迅速升高,处理后1h即显著高于对照；用NDGA处理后PacMYBA表达量始终低于对照和ABA处理,表明PacMYBA的表达可能受ABA调控.用VIGS技术沉默ABA生物合成途径中关键酶基因PacNCEDl可以降低甜樱桃果实内源ABA含量,且显著抑制了甜樱桃果实花色苷的合成。6个花色苷合成路径关键酶基因和PacMYBA的相对表达量在PacNCED1-RNAi果实中显著低于TRV空载体和对照果实。这进一步说明ABA作为一种信号分子参与了甜樱桃果实花色苷的合成。
Anthocyanins are the major water-soluble pigments in higher plants and are responsible for the blue, purple, and red colors of many fruits and vegetables. The anthocyanin biosynthetic pathway is part of the flavonoid pathway, which is a branch of the phenylpropanoid pathway. Recent studies have shown that a kind of R2R3MYB transcription factors have significantly regulation effect on anthocyanin biosynthesis. The present study describes the identification and functional characterization of the sweet cherry R2R3MYB protein, PacMYBA and also first obtained five full-length genes of anthocyanin biosynthesis pathway. The present work also first systematically expounds the role of plant hormone ABA in the fruit development process of sweet cherry. The main results are follows:
(1) In the present study we have studied the physiological and morphological changes during sweet cherry fruit development. The results showed that sweet cherry exhibited a biphasic growth pattern. The soluble sugars of sweet cherry including:glucose, fructose and sorbitol. No sucrose was detected in the sweet cherry. The levels of Chl declined continually after the SG stage, whereas the anthocyanin content increased rapidly after veraison.
(2) We first obtained the anthocyanin biosynthesis pathway genes of sweet cherry by using RACE technology and named as PacCHS, PacCHI, PacF3H, PacDFR, PacANS, and PacUFGT. The expression of PacCHS, PacCHI and PacF3H were at high levels during the early stages, declined during veraison (i.e., the onset of ripening; DQ YW, IR), and increased again in the last two stages. In accordance with the pattern of anthocyanin accumulation during ripening, the expression of PacDFR, PacANS and PacUFGT were expressed at low levels during the early developmental stages, and expression rapidly increased during veraison and remained at high levels through to the final developmental stage.
(3) We obtained a single putative R2R3MYB gene product, which we named PacMYBA, using RACE (rapid amplification of cDNA ends) PCR. Compared to apple MdMYB10, PacMYBA shared84%amino acid identity in the R2R3DNA-binding domain and68%identity over the whole protein. The nuclear localization of PacMYBA was consistent with its predicted function as a transcription factor and it is has no self-transcriptional activation activity. Tissue specific expression analysis showed that PacMYBA have the highest expression level in mature skin and flesh. The expression level of PacMYBA during sweet cherry fruit development is positive correlation with the anthocyanin biosynthesis of sweet cherry. We also found several cis-elements which related to plant growth and development response, stress response and plant hormone response in the promoter sequence of PacMYBA.
(4) The immature seeds of transgenic plants displayed red pigmentation<10days after anthesis, whereas the seeds of wild-type plants (ecotype Columbia) had no pigmentation. These data show that PacMYBA plays a role in anthocyanin biosynthesis.Silencing of PacMYBA, using a Tobacco rattle virus-induced gene silencing technique, resulted in sweet cherry fruit that lacked red pigment. Those fruits which were infiltrated with TRV alone pigmentation normally. These results shown that PacMYBA involve in regulating anthocyanin biosynthesis of sweet cherry directly.
(5) Abscisic acid (ABA) treatment significantly induced anthocyanin accumulation, while treatment with the ABA biosynthesis inhibitor nordihydroguaiaretic acid (NDGA) blocked anthocyanin production. The exoression level of PacMYBA increased quickly treated with30μmol/L ABA. After1h treatment with ABA the expression level of PacMYBA was significantly higher than control. This result suggested that PacMYBA might be a ABA-response gene.The uncolored phenotype was also observed in the fruits, silencing of PacNCED1, which encodes a key enzyme in the ABA biosynthesis pathway. The endogenous ABA content as well as the transcript levels of six structural genes and PacMYBA in PacNCED1-RNAi fruit was significantly lower than in the TRV vector control fruit. These results suggest that ABA is a signal molecule that promotes red-color sweet cherry fruit accumulating anthocyanin.
引文
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