菊花（Chrysanthemum morifolium Ramat.）是世界上最重要的观赏植物之一，具有重要的经济价值。花期是菊花重要的性状之一，与其市场价值紧密相关。众所周知，大多数菊花都为短日秋菊品种，花期较为集中。因此为了保持菊花的商品价值，对其进行花期调控尤为重要。目前在菊花生产上主要通过人工控光和控温以达到周年生产的目的，但这两种手段耗费大量的人力和物力，使生产成本大大提高。乙烯利是一种溶于水之后能释放乙烯的一种化学试剂，价格低廉，对环境友好，并且抑制菊花花芽分化的效果明显，目前也被用于菊花花期调控。迄今为止，虽然已经明确乙烯抑制菊花开花的功能，但分子调控机制仍未被揭示。在本研究中，我们从乙烯处理菊花‘神马’的转录组中查找到一个乙烯信号核心转录因子CmEIN3，揭示了乙烯介导菊花成花抑制的转录级联调控模式，主要研究结果如下：
1. 从菊花‘神马’中克隆获得CmEIN3基因，其开放阅读框（ORF）长为1659 bp，编码552个氨基酸，在N端含有一个EIN3保守结构域；组织表达模式分析表明其在营养生长时期的叶片中高表达；在乙烯处理前期会受到诱导上调表达；酵母转录激活实验表明其具有较强的转录激活活性，编码一个核蛋白。为了表征CmEIN3在菊花中的功能，构建pORE-R4-35AA-CmEIN3超表达载体和pDEST-SRDX-CmEIN3融合抑制子基因沉默载体并遗传转化菊花。分别选取两个超表达株系（CmEIN3-OX-2和CmEIN3-OX-5）和融合抑制子基因沉默株系（CmEIN3-SR1和CmEIN3-SR3）进行表型观察。在正常条件下，两个超表达株系相较于野生型分别晚花4 d和3 d，融合抑制子基因沉默株系分别早花6 d和6.5 d；而在外源乙烯利处理下，两个超表达株系相对于野生型分别晚花18.1 d和12.8 d，两个融合抑制子基因沉默株系分别早花14.1 d和15.1 d，说明CmEIN3参与乙烯介导的菊花成花抑制。将CmEIN3-OX-2和野生型材料的叶片送测转录组，数据分析发现一个注释为拟南芥TEM1的同源基因，其在超表达材料中出现差异上调表达。因TEM1是一个典型的成花抑制因子，且存在于乙烯信号下游，故将其作为CmEIN3的候选靶基因进行下一步验证。 
2. 根据已获得的转录组序列，克隆得到CmTEM1基因，ORF长为1023 bp，编码340个氨基酸，含有一个AP2和B3结构域，并且在C末端存在一个BRD抑制基序。组织定量分析表明其在营养生长的各个时期均有表达，但在叶片中的表达最高，且受乙烯诱导上调表达。酵母转录激活和拟南芥原生质体实验表明其不具有转录激活活性；烟草亚细胞定位实验表明其定位于细胞核中。克隆获得CmTEM1的启动子，进一步通过Y1H、ChIP-qPCR及双荧光素酶报告基因实验证明CmEIN3在体内和体外均能结合CmTEM1的启动子，并激活其表达。为进一步探究CmTEM1的功能，构建pORE-R4-35AA-CmTEM1超表达载体和pORE-R4-35AA-amiR-CmTEM1干扰载体并遗传转化菊花，分别获得两个超表达株系（CmTEM1-OX-1和CmTEM1-OX-2）和两个干扰株系株系（amiR-CmTEM1-1和amiR-CmTEM1-2）。表型观察发现，在正常条件下，两个超表达株系相较于野生型分别晚花4 d和8.2 d，干扰株系分别早花1.6 d和5.1 d；而在外源乙烯利处理下，两个超表达株系比野生型分别晚花16.8 d和25.2 d，两个干扰株系分别早花10.6 d和11 d，说明CmTEM1参与乙烯介导的菊花开花。将CmTEM1-OX-2和野生型叶片送测转录组，分析发现菊花开花整合因子CsAFL1的同源基因在超表达株系中显著下调表达，且该基因在乙烯处理菊花‘神马’的转录组中也显著下调表达，猜测其参与乙烯抑制菊花开花。
3. 克隆获得CmAFL1基因，其ORF长为714 bp，编码237个氨基酸，保守结构域分析表明其含有一个MADS-box和K-box结构域，属于MADS-box转录因子家族中的AP1/FUL亚家族成员。组织定量分析表明其在营养生长的各个组织均有表达，但在茎尖分生组织表达最高，受乙烯诱导下调表达，编码一个核定位蛋白。克隆获得CmAFL1基因的启动子，进一步通过Y1H、EMSA、ChIP-qPCR以及双荧光素酶报告基因实验表明CmTEM1能直接结合CmAFL1的启动子，并抑制其表达。分别在野生型（WT）和amiR-CmTEM1背景下沉默CmAFL1，发现CaLCuV-CmAFL1/WT和CaLCuV-CmAFL1/amiR-CmTEM1株系分别比转化空载CaLCuV/WT和CaLCuV/amiR-CmTEM1的株系平均晚花4.0 d和6.3 d，且CaLCuV-CmAFL1/amiR-CmTEM1开花时间早于CaLCuV-CmAFL1/WT，说明CmAFL1在菊花中具有促进开花的功能，且CmTEM1抑制菊花开花部分依赖CmAFL1。实验还表明CmAFL1促进开花功能在拟南芥中保守。

Chrysanthemum (Chrysanthemum morifolium Ramat.) is considered as one of the most important ornamental plants with a significant economic value in the world. Flowering time is one of the important traits of chrysanthemum, which is closely related to its economic value. As is widely known, the majority of chrysanthemums are short-day autumn varieties with a concentrated flowering period. Therefore, it is essential to regulate their flowering time to maintain their commercial value. At present, people mainly use artificial light and temperature control to achieve the purpose of annual production of chrysanthemums, but these two approaches require massive investment of human resources, resulting in increased production costs. Ethephon as an affordable and eco-friendly chemical reagent, dissolved easily in water, induces the release of ethylene and can effectively suppress flower bud initiation in chrysanthemums. Many producers use it to regulate the flowering time of chrysanthemums. So far, although the inhibitory effect of ethylene on chrysanthemum flowering has been established, a comprehensive understanding of the underlying molecular regulatory mechanisms has yet to be elucidated. In this study, we have identified the primary regulators CmEIN3 of ethylene signaling from a transcriptome data of wild type chrysanthemum 'Jinba' with the treatment of ethephon, and revealed a transcriptional regulatory module of ethylene-mediated flowering inhibition in chrysanthemums, the following are the main findings:
1. The transcription factor CmEIN3 was isolated from the chrysanthemum 'Jinba', with an open reading frame (ORF) of 1659 bp, encoding 552 amino acids with a conserved EIN3 domain at its N-terminal. Tissue expression pattern analysis showed that CmEIN3 was highly expressed in leaves at vegetative growth stage and induced by ethylene. Yeast transcriptional activation assay indicated that CmEIN3 had strong transcriptional activation activity, and it was a nuclear localized protein. To elucidate the role of CmEIN3 in chrysanthemum, the pORE-R4-35AA-CmEIN3 (CmEIN3-OX) overexpressing vector and pDEST-SRDX-CmEIN3 (CmEIN3-SRDX) chimeric repressor gene-silencing technology (CRES-T) vector were constructed, then transformed into chrysanthemum respectively. We selected two overexpressing lines (CmEIN3-OX-2 and CmEIN3-OX-5) and two silencing lines (CmEIN3-SR1 and CmEIN3-SR3) for phenotypic observation. Under control conditions, it was found that the two overexpressing lines flowered 4 d and 3 d later, while the gene silenced lines flowered 6 d and 6.5 d earlier than wild type, respectively. With the exogenous ethephon treatment, the two overexpressing lines flowered 18.1 d and 12.8 d later, and the two gene silenced lines flowered 14.1 d and 15.1 d earlier than wild type, respectively, which suggested that CmEIN3 was involved in ethylene-mediated inhibition of flowering in chrysanthemum. To further identified the function of CmEIN3, the leaves of CmEIN3-OX-2 and the wild-type plants were used for RNA-seq, it was discovered that the homolog TEM1 gene CmTEM1, which plays a role in the ethylene signaling pathway and acts as a repressor of flowering, exhibited increased expression level in CmEIN3-OX-2. Therefore, TEM1 was selected as the candidate target gene of CmEIN3 for further validation.
2. CmTEM1 was cloned with an ORF length of 1023 bp, encoding 340 amino acids with an AP2 and a B3 domain, and also a BRD repressor motif at its C-terminal. Tissue expression pattern analysis suggested that it expressed at all tissues of the vegetative growth stage, but its expression was highest in leaves and induced by ethylene. Transcriptional activity analysis  in yeast cells and Arabidopsis protoplasts both exhibited that it didn’t have transcriptional activation, and subcellular localization assays in tobacco showed that it localized in the nucleus. The promoter of CmTEM1 was further cloned, then Y1H, ChIP-qPCR and dual luciferase assays showed that CmEIN3 could bind to the CmTEM1 promoter in vivo and in vitro, which directly activated its expression. In addition, we further investigated the function of CmTEM1 by constructing overexpressing vector pORE-R4-35AA-CmTEM1 and artificial microRNA vector pORE-R4-35AA-amiR-CmTEM1, and then transformed them into chrysanthemum to obtain two overexpressing lines (CmTEM1-OX-1 and CmTEM1-OX-2) and knock-down lines (amiR-CmTEM1-1 and amiR-CmTEM1-2), respectively. Under normal conditions, the two overexpressing lines flowered 4 d and 8.2 d later and the knock-down lines flowered 1.6 d and 5.1 d earlier than the wild type, respectively. While with exogenous ethephon treatment, the two overexpressing lines flowered 16.8 d and 25.2 d later and the two knock-down lines flowered 10.6 d and 11 d earlier than the wild type, respectively, which indicated that CmTEM1 was involved in ethylene-mediated chrysanthemum flowering. Subsequently, CmTEM1-OX-2 and wild-type plants were used for RNA-seq, and the results revealed that a homolog of the chrysanthemum flowering integrator CsAFL1 was significantly down-regulated in the CmTEM1-OX-2, and it was also down-regulated in the transcriptome of wild type plant with ethephon treatment. It was speculated that CsAFL1 might be involved in the biological process of ethylene inhibition of chrysanthemum flowering. 
3. The CmAFL1 was cloned, with an ORF length of 714 bp, encoding 237 amino acids with a MADS-box and K-box domain. It is a member of the AP1/FUL subfamily of the MADS family. Tissue expression pattern analysis showed that CmAFL1 expressed at all tissues of the vegetative growth stage, abundantly in the apical meristem, and was repressed by ethylene, it encoded a nuclear-localized protein. The promoter of the CmAFL1 gene was cloned. Y1H, EMSA, ChIP-qPCR and dual luciferase assays suggested that CmTEM1 could directly bind to the promoter of CmAFL1 and repressed its expression. Furthermore, the silencing CmAFL1 in both wild-type plants (CaLCuV-CmAFL1/WT) and amiR-CmTEM1 plants (CaLCuV-CmAFL1/amiR-CmTEM1) showed late flowering 4.0 d and 6.3 d comparing with CaLCuV/WT and CaLCuV/amiR-CmTEM1 plants under short-day conditions, respectively, but the flowering time of CaLCuV-CmAFL1/amiR-CmTEM1 plants was earlier than that of CaLCuV-CmAFL1/WT plants, which suggested that CmTEM1 as a direct upstream regulator suppresses flowering by targeting the flowering positive regulator CmAFL1 partially. The heterologous expression of CmAFL1 in Arabidopsis also accelerated flowering.

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