甘菊成花相关基因表达模式和功能的研究
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摘要
花期不仅是影响菊花(Chrysanthemumxmorifolium Ramat.)观赏性状的重要因素之一,也是重要的栽培性状通过有效操控成花的关键基因,培育出自然花期适合产业化生产的菊花新品种,具有重要的理论和实践意义。由于菊花遗传背景复杂,对其成花的分子机理进行研究十分困难。为此,本文作者以甘菊[Chrysanthemum lavandulifolum (Fisch. ex Trautv.) Makino]作为菊花成花机理的研究模式,首先利用高通量测序技术获得了甘菊混合样本的转录组信息,并对其进行了生物信息学分析。然后利用升级版的DGE (Up-graded Digital Gene Expression)技术分析了甘菊幼苗期和现蕾时的基因表达特征。由于花分生组织特征基囚在植物成花过程中所起的关键性作用,本文作者利用模式杭物拟南芥验证了甘菊开花基因DFL和DenFUL的功能。获得了以下主要成果：
1.高通量测序总计获得了4GB的原始数据,从头组装后获得了108,737条Unigene.其平均长度为349bp。根据蛋白同源性比对,58,093条Unigene被注释(E-value<10~5)通过GO、 COG和pathway等生物信息学方法分析证明该转录组文库为甘菊成花及其他基础生物学研究提供了一个非常有价值的基因资源库。
2.确定了甘菊成花受到光周期途径、春化途径、GA途径、自主途径和FRI依赖途径的调控,鉴定了可以用于菊花花期改良的56个重要候选基因。筛选到6,204条编码转录因子的Unigene,这些基因分别属于57个不同的转录因子家族
3.通过升级版DGE技术筛选到在甘菊现蕾时特异性表达的基因类群。甘菊光周期途径、赤霉素途径、自主途径和春化途径中绝大多数促进开花的基因在甘菊现蕾时表达量上升。此外,基础代谢途径中的糖类和淀粉代谢途径、氮素代谢和新陈代谢途径中基因,以及MIKC、SBP、YABBY、 ARF、 Dof和MYB等转录因子家族的基因调控甘菊成花。
4.在拟南芥中过表达甘菊DFL基因能互补拟南芥lfy突变体,并使野生型拟南芥花期提前14天左右。使用LFY启动子驱动DFL在野生型拟南芥表达,可以使花期提前7天左右。此外,本研究还发现PLFY::DFL:EGFP转化野生型拟南芥出现了共抑制现象。
5.过表达DenFUL使拟南芥成花时间大幅度缩短；项部总状花变成本次生枝变成侧花。此外，DenFUL转基因拟南芥莲座叶变小,角果上的花柱变细、伸长,但是没有影响果实的开裂性。
6.通过转基因植物活体检测、洋葱表皮和拟南芥原生质体瞬时表达分别证明DFL和DenFUL均定位于细胞核中。
7.迪过分析转基因拟南芥内源基因的表达模式证明,DFL能够激活拟南芥中AP1,AP3和AG花器官特征基因表达,抑制TFL1基因表达。而在DenFUL转基因拟南芥中LFY和FT的表达量升高,TFL1和FLC的表达量降低。
综合上述研究结果得出了本研究的主要结论：甘菊成花除了受到光周期途径、春化途径、赤毒素途径和自主途径4条遗传途径的控制外,糖类和淀粉等基础代谢途径中的基因以及转录因子也起到了重要作用,通过互补试验证实了DFL基因在甘菊成花过程中起到了关键性的作用；DenFUL具有调控甘菊开花的作用,对果实和叶片的发育也有一定影响。这些研究初步揭示了甘菊成花调控的分子机理,为菊花花期改良奠定了基础。
The flowering period of Chrysanthemum is important horticultural and cultivated characters. It is highly desirable to effectively manipulate the key genes involved in flowering and breed the new variety of Chrysanthemum with the natural florescence suitable for industrial production, while maintaining normal growth and without affecting their resistance to diseases and pests. However, research at the molecular level is limited by a lack of sequence data. Due to the complex genetic background of Chrysanthemum, an analysis of a Chrysanthemum relative species-Chrysanthemum lavandulifolium(Fisch. ex Trautv.) transcriptome aimed to obtain transcript sequence data will be helpful to elucidate the molecular mechanism of flowering control in Chrysanthemum. In this study, the transcriptome of C. lavandlifolium under different developmental stages and stress treatments was sequenced by Illumina/solexa technology, and then bioinformatically analyzed. The genes spatio-temporal expression profiles between seeding stage and squaring period were performed by up-graded version of the DGE. We perform the function analysis of DFL and DenFUL in Arabidopsis by transgenic technology. The major results and conclusions are described as follows:
1. RNA-seq generated4GB raw data, which was then de novo assembled into108,738Unigenes with a mean length of349bp. A total of58,093Unigenes were obtained non-redundant annotation(E-value<10-5). GO, COG and pathway bioinformatically analysis showed that this transcriptome library provide a valuable resource for studying the C.lavandulifolium flowering mechanism and others basic biology.
2. Based on the annotations, we screened56important candidate genes that can be used in genetically modified Chrysanthemum flowering. Those genes respectively belong to photoperiod pathway, vernalization pathway, GA pathway, autonomous pathway and FRI-dependent pathway, which indicated these flowering genetic regulatory pathways coexistence in C.lavandulifolium. In addition, we identified 6,204Unigenes that encode transcription factors and belong to57transcription factor families, of which8transcription factor families were related to flowering.
3. Using up-graded version of DGE technology, we identified gene groups that specifically expressed during C.lavandulifolium squaring period. Most genes promoting flowering in the photoperiod pathway, GA pathway, autonomous pathway and vernalization pathways were up-regulated during C.lavandulifolium squaring period. In addition, the basalmetabolic pathway, such as sugar and starch metabolic pathways, nitrogen metabolism and metabolic pathway genes, as well as MIKC, SBP, YABBY, ARF, Dof and MYB transcription factor families were involved in C.lavandulifolium flowering.
4. Heterologous overexpression of DFL extremely promoted Arabidopsis wildtype early flowering, the florescence was advanced about14days; and fully complemented Ify mutant. Using LFY promoter to drive DFL expressed in Arabidopsis can promote flowering about7days. We also found heterologous expression DFL in Arabidopsis wildtype drived by LFY promoter caused co-suppression.
5. DenFUL transgenic Arabidopsis plants were significant early flowering, which advanced about14days than wild-type under long-day conditions and more than40days under short day conditions. In addition, DenFUL transgenic Arabidopsis producing terminal flowers and lateral flowers, the rosette leaves much small than WT and the style of silique was thin and elongate; but did not affect the fruit cracking.
6. Transgenic plants vivo detection, transient expression using agrobacterium-mediated in onion epidermal cells and transient expression using PEG-mediated in Arabidopsis proplast were employed to investigate the subcellular location of DFL and DenFUL, respectively. All the analysis reached to the same patterns that DFL and DenFUL were localized in nuclear as Arabidopsis LFY and API did.
7. Analysis on endogenous genes expression patterns in transgenic Arabidopsis, we found DFL widely activated the expression of B-and C-class floral organ identity genes, inhibited TFL1expression. Overexpression of DenFUL conferred the expression of endogenesis LfYand FT increased while TFL1and FLC decreased.
Comprehensive results in this study, we gain the main conclusions:the flowering of C. lavandulifolium is not only controlled by photoperiod pathway, vernalization pathway, GA pathway, autonomous pathway and FRI-dependent pathway, but also MIKC, SBP, YABBY, AFR transcription factor family and basal metabolism such as sucrose and starch metabolism play important roles. These genes were excellent candidate genes to modify Chrysanthemum flowering. DFL has LFY function and might plays key roles in C.lavandulifolium transformation flowering. DenFUL might play key roles in flower development and also involved in fruit and leaves development. These studies preliminarily reveal molecular mechanism of flowering in C.lavandulifolium, laying foundations on genetic modification of Chrysanthemum flowering.
1.高通量测序总计获得了4GB的原始数据,从头组装后获得了108,737条Unigene.其平均长度为349bp。根据蛋白同源性比对,58,093条Unigene被注释(E-value<10~5)通过GO、 COG和pathway等生物信息学方法分析证明该转录组文库为甘菊成花及其他基础生物学研究提供了一个非常有价值的基因资源库。
2.确定了甘菊成花受到光周期途径、春化途径、GA途径、自主途径和FRI依赖途径的调控,鉴定了可以用于菊花花期改良的56个重要候选基因。筛选到6,204条编码转录因子的Unigene,这些基因分别属于57个不同的转录因子家族
3.通过升级版DGE技术筛选到在甘菊现蕾时特异性表达的基因类群。甘菊光周期途径、赤霉素途径、自主途径和春化途径中绝大多数促进开花的基因在甘菊现蕾时表达量上升。此外,基础代谢途径中的糖类和淀粉代谢途径、氮素代谢和新陈代谢途径中基因,以及MIKC、SBP、YABBY、 ARF、 Dof和MYB等转录因子家族的基因调控甘菊成花。
4.在拟南芥中过表达甘菊DFL基因能互补拟南芥lfy突变体,并使野生型拟南芥花期提前14天左右。使用LFY启动子驱动DFL在野生型拟南芥表达,可以使花期提前7天左右。此外,本研究还发现PLFY::DFL:EGFP转化野生型拟南芥出现了共抑制现象。
5.过表达DenFUL使拟南芥成花时间大幅度缩短；项部总状花变成本次生枝变成侧花。此外，DenFUL转基因拟南芥莲座叶变小,角果上的花柱变细、伸长,但是没有影响果实的开裂性。
6.通过转基因植物活体检测、洋葱表皮和拟南芥原生质体瞬时表达分别证明DFL和DenFUL均定位于细胞核中。
7.迪过分析转基因拟南芥内源基因的表达模式证明,DFL能够激活拟南芥中AP1,AP3和AG花器官特征基因表达,抑制TFL1基因表达。而在DenFUL转基因拟南芥中LFY和FT的表达量升高,TFL1和FLC的表达量降低。
综合上述研究结果得出了本研究的主要结论：甘菊成花除了受到光周期途径、春化途径、赤毒素途径和自主途径4条遗传途径的控制外,糖类和淀粉等基础代谢途径中的基因以及转录因子也起到了重要作用,通过互补试验证实了DFL基因在甘菊成花过程中起到了关键性的作用；DenFUL具有调控甘菊开花的作用,对果实和叶片的发育也有一定影响。这些研究初步揭示了甘菊成花调控的分子机理,为菊花花期改良奠定了基础。
The flowering period of Chrysanthemum is important horticultural and cultivated characters. It is highly desirable to effectively manipulate the key genes involved in flowering and breed the new variety of Chrysanthemum with the natural florescence suitable for industrial production, while maintaining normal growth and without affecting their resistance to diseases and pests. However, research at the molecular level is limited by a lack of sequence data. Due to the complex genetic background of Chrysanthemum, an analysis of a Chrysanthemum relative species-Chrysanthemum lavandulifolium(Fisch. ex Trautv.) transcriptome aimed to obtain transcript sequence data will be helpful to elucidate the molecular mechanism of flowering control in Chrysanthemum. In this study, the transcriptome of C. lavandlifolium under different developmental stages and stress treatments was sequenced by Illumina/solexa technology, and then bioinformatically analyzed. The genes spatio-temporal expression profiles between seeding stage and squaring period were performed by up-graded version of the DGE. We perform the function analysis of DFL and DenFUL in Arabidopsis by transgenic technology. The major results and conclusions are described as follows:
1. RNA-seq generated4GB raw data, which was then de novo assembled into108,738Unigenes with a mean length of349bp. A total of58,093Unigenes were obtained non-redundant annotation(E-value<10-5). GO, COG and pathway bioinformatically analysis showed that this transcriptome library provide a valuable resource for studying the C.lavandulifolium flowering mechanism and others basic biology.
2. Based on the annotations, we screened56important candidate genes that can be used in genetically modified Chrysanthemum flowering. Those genes respectively belong to photoperiod pathway, vernalization pathway, GA pathway, autonomous pathway and FRI-dependent pathway, which indicated these flowering genetic regulatory pathways coexistence in C.lavandulifolium. In addition, we identified 6,204Unigenes that encode transcription factors and belong to57transcription factor families, of which8transcription factor families were related to flowering.
3. Using up-graded version of DGE technology, we identified gene groups that specifically expressed during C.lavandulifolium squaring period. Most genes promoting flowering in the photoperiod pathway, GA pathway, autonomous pathway and vernalization pathways were up-regulated during C.lavandulifolium squaring period. In addition, the basalmetabolic pathway, such as sugar and starch metabolic pathways, nitrogen metabolism and metabolic pathway genes, as well as MIKC, SBP, YABBY, ARF, Dof and MYB transcription factor families were involved in C.lavandulifolium flowering.
4. Heterologous overexpression of DFL extremely promoted Arabidopsis wildtype early flowering, the florescence was advanced about14days; and fully complemented Ify mutant. Using LFY promoter to drive DFL expressed in Arabidopsis can promote flowering about7days. We also found heterologous expression DFL in Arabidopsis wildtype drived by LFY promoter caused co-suppression.
5. DenFUL transgenic Arabidopsis plants were significant early flowering, which advanced about14days than wild-type under long-day conditions and more than40days under short day conditions. In addition, DenFUL transgenic Arabidopsis producing terminal flowers and lateral flowers, the rosette leaves much small than WT and the style of silique was thin and elongate; but did not affect the fruit cracking.
6. Transgenic plants vivo detection, transient expression using agrobacterium-mediated in onion epidermal cells and transient expression using PEG-mediated in Arabidopsis proplast were employed to investigate the subcellular location of DFL and DenFUL, respectively. All the analysis reached to the same patterns that DFL and DenFUL were localized in nuclear as Arabidopsis LFY and API did.
7. Analysis on endogenous genes expression patterns in transgenic Arabidopsis, we found DFL widely activated the expression of B-and C-class floral organ identity genes, inhibited TFL1expression. Overexpression of DenFUL conferred the expression of endogenesis LfYand FT increased while TFL1and FLC decreased.
Comprehensive results in this study, we gain the main conclusions:the flowering of C. lavandulifolium is not only controlled by photoperiod pathway, vernalization pathway, GA pathway, autonomous pathway and FRI-dependent pathway, but also MIKC, SBP, YABBY, AFR transcription factor family and basal metabolism such as sucrose and starch metabolism play important roles. These genes were excellent candidate genes to modify Chrysanthemum flowering. DFL has LFY function and might plays key roles in C.lavandulifolium transformation flowering. DenFUL might play key roles in flower development and also involved in fruit and leaves development. These studies preliminarily reveal molecular mechanism of flowering in C.lavandulifolium, laying foundations on genetic modification of Chrysanthemum flowering.
引文
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