光周期对草原龙胆生长发育的影响及相关基因的功能分析
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摘要
草原龙胆又名草原龙胆,是国际市场上非常流行的切花,矮生品种的培育使其在园林中的应用形式更为广泛。本文对‘德拉迷斯’草原龙胆展开光周期方面的研究,一方面从栽培的角度研究‘德拉迷斯’的光周期和花芽分化特性；另一方面从分子的角度探讨光周期调控‘德拉迷斯’开花的机理。
本研究的主要结果如下：
1.通过7种不同日长的光周期处理,确定了‘德拉迷斯’草原龙胆为量性长日照植物,促进开花的临界日长为14h。光照时间越长,植株生长越高,而真叶数减少。通过比较几种处理下‘德拉迷斯’的开花时间以及开花时的品质,确定14h是生产上能保证得到较好开花品质且比较节能的光周期。
2.采用石蜡切片法对‘德拉迷斯’的花芽分化过程进行显微观察,可以将为期3周的分化过程划分为6个时期：营养生长期、花芽分化初期、萼片原基分化期、花瓣原基分化期、雄蕊原基分化期和雌蕊原基分化期。据此,将花芽分化初期确定为成花转变时期,此时顶端分生组织向上升起,并变得宽阔平坦,出现在萼片原基分化之前。
3.通过观测‘德拉迷斯’花芽分化时期对应的营养生长指标,建立了花芽分化时期(DP)与营养生长(VGTs)之间的相关性。结果显示,株高、冠幅、真叶数和出苗周数都与花芽分化时期存在显著相关性,可以用来预测花芽分化进程。根据回归方程推算,‘德拉迷斯’成花转化时期发生在21片真叶期。因此,建议生产上在21片真叶之前用14h光周期进行处理,以期加速开花。
4.采用同源克隆法从草原龙胆中克隆到光周期途径基因EgGI的gDNA和cDNA全长。通过几种光周期下的实时定量表达(qRT-PCR)分析均表明,EgGI受内源昼夜节律钟的调控,其表达量在光照8-12hr后达到峰值,至暗期下降。另外,分析在长、短日照下成花转变前后EgGI的表达水平,发现LD条件下EgGI的表达水平均明显高于SD条件下,推测EgGI可能参与草原龙胆成花转变之前的调控过程。
5.从草原龙胆中克隆到两个CO基因的同源基因EgCOL1和EgCOL2。通过同源比对和进化树分析,发现EgCOL1划分在CO基因家族的第I组a亚类,与拟南芥AtCOLl和AtCOL2同源性最高；EgCOL2划分在第Ⅰ组c亚类,与拟南芥AtCOL5同源性最高。
6.分析EgCOLl和EgCOL2在几种光周期下的表达节律,结果表明EgCOLl的表达受内源昼夜节律钟调控,而EgCOL2在四种光周期下昼夜节律性不明显。另外,分析EgCOLl和EgCOL2在成花转变前后的表达水平,发现两个基因在LD下的表达水平都要高于SD,且在成花转变时期都有明显的表达峰值,其中EgCOLl的对比更显著。推测这两个基因可能都参与了草原龙胆的成花转变过程。
7.分别构建EgCOLl和EgCOL2基因的超表载体转化烟草,成功获得转基因烟草。表型观测发现,超量表达EgCOLl的烟草与对照在开花时间和外部形态特征上没有明显差别。超量表达EgCOL2的烟草出现了明显的早花现象,qRT-PCR分析发现,强表型植株EgCOL2表达水平明显较高,且上调烟草NtFT和NtSOCl的表达。说明EgCOL2基因具有促进烟草早花的功能。
Lisianthus is one of the most highly ranked cut flowers in international markets, and it is now enjoying increasing popularity as a pot plant in the landscape for the cultivation of dwarf varieties. The photoperiod regulation mechanism of lisianthus has been studied in this paper with the cultivar'Tiramisu'. On the one hand, we studied the photoperiod characteristics and floral transition in the aspect of cultivation, on the other hand, we explore the photoperiodic flowering mechanism from the molecular perspective.
The main results of this study are as follows:
1. The effects of the photoperiod on the flowering traits of Eustoma grandiflorum 'Tiramisu'(TDC) were investigated under7day-length treatments (10h/12h/14h/16h/18h/20h/24h).'Tiramisu'was deduced to be a facultative long-day plant, of which the critical day-length for flowering was14h. Plants tended to be taller and less leaf number under longer days. Compared the flowering time and flower quality in these photoperiods,14h was supposed to be the optimal photoperiod in the commercial production of lisianthus for the high flowering quality and low energy consumption.
2. The flower bud differentiation process of 'Tiramisu' was observed by paraffin method. The differentiation process lasted about3weeks and was divided into six stages:vegetative growth phase, flower initiation phase, sepal primordia phase, petal primordia phase, stamen primordia phase and pistil primordia phase. The floral transition was distinguished in the flower initiation phase, which was characterized by a distinctly raised and broadened apical meristem before appearance of sepal primordia.
3. By observing the vegetative growth corresponding to flower bud differentiation phase (DP), the correlation between DP and vegetative growth traits (VGTs) was established. The results indicated that, significant linear relationships existed between each growth characteristics of 'Tiramisu' and its flower bud differentiation phase. It could be estimated from the regression equations that the floral transition of TDC occured at21-leaf stage. Therefore, it is recommended that TDC planted under day-length of14h before21-leaf stage to promote flowering.
4. Photoperiod pathway gene EgGI was isolated from lisianthus and the full gDNA and cDNA sequence was obtained with Tail-PCR method. Analysis the expression rhythms of EgGI in different photoperiods indicated that EgGI mRNA is under the control of endogenous circadian clock, and showed diurnal rhythms peaking at8-12hours later when transferred to light period, then decreased in the dark. In addition, analysis of the expression level of EgGI during floral transition in LD and SD condition indicated that, EgGI mRNA under LD condition was always significant higher than under SD condition, suggesting that EgGI possibly have involved in the process before floral transition.
5. Two CO homologous genes were isolated from lisianthus and designated as EgCOL1and EgCOL2. Homologous alignment and phylogenetic tree analysis indicated that EgCOL1gene was divided to I a subclass of CO family, and the closest homologues were AtCOLl and AtCOL2; EgCOL2was divided to I c subclasses, and was most similar to AtCOL5.
6. Circadian expression of EgCOLl and EgCOL2was analyzed under different photoperiods. The results indicated that EgCOLl was under the control of endogenous circadian clock; while EgCOL2had no obvious expression rhythms in four kinds of photoperiod. In addition, the transcriptional levels of EgCOLl and EgCOL2were much higher in LD condition than in SD condition, and the expression levels of the two genes both reached the peak during the floral transition period. There had more significant difference in the expression level of EgCOL1between SD and LD condition. These two genes might be involved in the process of floral transition.
7. Over-expression vectors of EgCOLl and EgCOL2were constructed and transformed into tobacco. By phenotype observation, there were no differences between the tobacco transformed with EgCOLl and the wild type on flowering time and phenotype. The transgenic tobacco with EgCOL2showed a phenomenon of significant early flowering. Through expression analysis of transgenic plants, found that the plants with strong phenotype have significant higher expression levels in EgCOL2, and the expression level of NtFT and NtSOCl were also up regulated. These results suggest that EgCOL2play an important role in promoting early flowering in tobacco.
本研究的主要结果如下：
1.通过7种不同日长的光周期处理,确定了‘德拉迷斯’草原龙胆为量性长日照植物,促进开花的临界日长为14h。光照时间越长,植株生长越高,而真叶数减少。通过比较几种处理下‘德拉迷斯’的开花时间以及开花时的品质,确定14h是生产上能保证得到较好开花品质且比较节能的光周期。
2.采用石蜡切片法对‘德拉迷斯’的花芽分化过程进行显微观察,可以将为期3周的分化过程划分为6个时期：营养生长期、花芽分化初期、萼片原基分化期、花瓣原基分化期、雄蕊原基分化期和雌蕊原基分化期。据此,将花芽分化初期确定为成花转变时期,此时顶端分生组织向上升起,并变得宽阔平坦,出现在萼片原基分化之前。
3.通过观测‘德拉迷斯’花芽分化时期对应的营养生长指标,建立了花芽分化时期(DP)与营养生长(VGTs)之间的相关性。结果显示,株高、冠幅、真叶数和出苗周数都与花芽分化时期存在显著相关性,可以用来预测花芽分化进程。根据回归方程推算,‘德拉迷斯’成花转化时期发生在21片真叶期。因此,建议生产上在21片真叶之前用14h光周期进行处理,以期加速开花。
4.采用同源克隆法从草原龙胆中克隆到光周期途径基因EgGI的gDNA和cDNA全长。通过几种光周期下的实时定量表达(qRT-PCR)分析均表明,EgGI受内源昼夜节律钟的调控,其表达量在光照8-12hr后达到峰值,至暗期下降。另外,分析在长、短日照下成花转变前后EgGI的表达水平,发现LD条件下EgGI的表达水平均明显高于SD条件下,推测EgGI可能参与草原龙胆成花转变之前的调控过程。
5.从草原龙胆中克隆到两个CO基因的同源基因EgCOL1和EgCOL2。通过同源比对和进化树分析,发现EgCOL1划分在CO基因家族的第I组a亚类,与拟南芥AtCOLl和AtCOL2同源性最高；EgCOL2划分在第Ⅰ组c亚类,与拟南芥AtCOL5同源性最高。
6.分析EgCOLl和EgCOL2在几种光周期下的表达节律,结果表明EgCOLl的表达受内源昼夜节律钟调控,而EgCOL2在四种光周期下昼夜节律性不明显。另外,分析EgCOLl和EgCOL2在成花转变前后的表达水平,发现两个基因在LD下的表达水平都要高于SD,且在成花转变时期都有明显的表达峰值,其中EgCOLl的对比更显著。推测这两个基因可能都参与了草原龙胆的成花转变过程。
7.分别构建EgCOLl和EgCOL2基因的超表载体转化烟草,成功获得转基因烟草。表型观测发现,超量表达EgCOLl的烟草与对照在开花时间和外部形态特征上没有明显差别。超量表达EgCOL2的烟草出现了明显的早花现象,qRT-PCR分析发现,强表型植株EgCOL2表达水平明显较高,且上调烟草NtFT和NtSOCl的表达。说明EgCOL2基因具有促进烟草早花的功能。
Lisianthus is one of the most highly ranked cut flowers in international markets, and it is now enjoying increasing popularity as a pot plant in the landscape for the cultivation of dwarf varieties. The photoperiod regulation mechanism of lisianthus has been studied in this paper with the cultivar'Tiramisu'. On the one hand, we studied the photoperiod characteristics and floral transition in the aspect of cultivation, on the other hand, we explore the photoperiodic flowering mechanism from the molecular perspective.
The main results of this study are as follows:
1. The effects of the photoperiod on the flowering traits of Eustoma grandiflorum 'Tiramisu'(TDC) were investigated under7day-length treatments (10h/12h/14h/16h/18h/20h/24h).'Tiramisu'was deduced to be a facultative long-day plant, of which the critical day-length for flowering was14h. Plants tended to be taller and less leaf number under longer days. Compared the flowering time and flower quality in these photoperiods,14h was supposed to be the optimal photoperiod in the commercial production of lisianthus for the high flowering quality and low energy consumption.
2. The flower bud differentiation process of 'Tiramisu' was observed by paraffin method. The differentiation process lasted about3weeks and was divided into six stages:vegetative growth phase, flower initiation phase, sepal primordia phase, petal primordia phase, stamen primordia phase and pistil primordia phase. The floral transition was distinguished in the flower initiation phase, which was characterized by a distinctly raised and broadened apical meristem before appearance of sepal primordia.
3. By observing the vegetative growth corresponding to flower bud differentiation phase (DP), the correlation between DP and vegetative growth traits (VGTs) was established. The results indicated that, significant linear relationships existed between each growth characteristics of 'Tiramisu' and its flower bud differentiation phase. It could be estimated from the regression equations that the floral transition of TDC occured at21-leaf stage. Therefore, it is recommended that TDC planted under day-length of14h before21-leaf stage to promote flowering.
4. Photoperiod pathway gene EgGI was isolated from lisianthus and the full gDNA and cDNA sequence was obtained with Tail-PCR method. Analysis the expression rhythms of EgGI in different photoperiods indicated that EgGI mRNA is under the control of endogenous circadian clock, and showed diurnal rhythms peaking at8-12hours later when transferred to light period, then decreased in the dark. In addition, analysis of the expression level of EgGI during floral transition in LD and SD condition indicated that, EgGI mRNA under LD condition was always significant higher than under SD condition, suggesting that EgGI possibly have involved in the process before floral transition.
5. Two CO homologous genes were isolated from lisianthus and designated as EgCOL1and EgCOL2. Homologous alignment and phylogenetic tree analysis indicated that EgCOL1gene was divided to I a subclass of CO family, and the closest homologues were AtCOLl and AtCOL2; EgCOL2was divided to I c subclasses, and was most similar to AtCOL5.
6. Circadian expression of EgCOLl and EgCOL2was analyzed under different photoperiods. The results indicated that EgCOLl was under the control of endogenous circadian clock; while EgCOL2had no obvious expression rhythms in four kinds of photoperiod. In addition, the transcriptional levels of EgCOLl and EgCOL2were much higher in LD condition than in SD condition, and the expression levels of the two genes both reached the peak during the floral transition period. There had more significant difference in the expression level of EgCOL1between SD and LD condition. These two genes might be involved in the process of floral transition.
7. Over-expression vectors of EgCOLl and EgCOL2were constructed and transformed into tobacco. By phenotype observation, there were no differences between the tobacco transformed with EgCOLl and the wild type on flowering time and phenotype. The transgenic tobacco with EgCOL2showed a phenomenon of significant early flowering. Through expression analysis of transgenic plants, found that the plants with strong phenotype have significant higher expression levels in EgCOL2, and the expression level of NtFT and NtSOCl were also up regulated. These results suggest that EgCOL2play an important role in promoting early flowering in tobacco.
引文
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