Chinese Bulletin of Botany ›› 2024, Vol. 59 ›› Issue (2): 257-277.DOI: 10.11983/CBB23096
• SPECIAL TOPICS • Previous Articles Next Articles
Wen Chen*,†(
), Yingying Zhou,†, Ping Luo, Yongyi Cui
HomeReceived:2023-07-22
Accepted:2023-11-14
Online:2024-03-01
Published:2023-12-26
Contact:
* E-mail: About author:† These authors contributed equally to this paper
Wen Chen, Yingying Zhou, Ping Luo, Yongyi Cui. Molecular Mechanism of Petal Doubling of Flower in Angiosperm[J]. Chinese Bulletin of Botany, 2024, 59(2): 257-277.
Figure 1 Flower development models of Angiosperm (refer to Soltis et al., 2007; Thomson and Wellmer, 2019) (A) Classic ABC model (represented by plants such as Arabidopsis thaliana, snapdragon and rose); (B) Modified ABC model (represented by plants of Tulipa, Lilium, Ranunculus and Aquilegia); (C) ABCDE model (represented by model plants such as Arabidopsis thaliana and snapdragon); (D) (A)BCD model (represented by plants such as gerbera and rice, the defined (A) combines A and E function genes). Arrows represent the regulation of functional genes on specific floral organs; T-shaped lines represent repression. Se: Sepals; Pt: Petals; St: Stamens; Ca: Carpels; Ov: Ovules
Figure 2 HOT model and perianth code for flower development of orchids (refer to Hsu et al., 2015; Zhang et al., 2022) (A) HOT model for flower development of orchids (PI and four clades of AP3 of B-class genes specify the orchid perianth formation at the late inflorescence and floral bud stages. At the late inflorescence stage, PI and AP3Bs determine the formation of sepals. PI and both AP3A1 and AP3Bs control the lateral petal formation; at the floral bud stage, PI, AP3As and other MADS-box genes control the lip morphogenesis); (B) Perianth code of orchids (in the sepal, petal and lip, PI interacts with different AP3-like and AGL-like proteins to form L complex (OAP3-2/OAGL6-2/OAGR6-2/OPI) or SP complex (OAP3-1/OAGL6-1/OGL6-1/OPI). The presence of L complex results in lip formation, and the presence of SP complex results in sepal/petal formation; the coexistence or co-absence of the SP and L complexes produces sepal/petal and lip intermediate structures). Numbers 1, 2, 3/4 represent flower whorls.
| 类型 | 名称 | 功能及表型 | 重瓣起源类型 | 参考文献 |
|---|---|---|---|---|
| A类功能基因 | AP2 | 月季RcAP2表达水平受低温诱导而被高温抑制, 拟南芥异源过表达RcAP2可诱导雄蕊瓣化 | 雌雄蕊起源 | Han et al., |
| B类功能基因(MADS-box) | PI | 拟南芥中异源过表达桃PpPI可诱导雄蕊瓣化; 拟南芥中异源过表达枇杷EjPI可导致萼片瓣化 | 雌雄蕊起源或苞片起源 | Xia et al., |
| AP3 | 夏堇中共表达TfDEF (AP3)和TfGLO (PI)可导致萼片瓣化; 共抑制两者可导致花瓣萼片化 | 苞片起源 | Sasaki et al., | |
| C类功能基因(MADS-box) | AG | 矮牵牛、三花龙胆和月季中沉默AG同源基因导致雄蕊瓣化; 低温诱导的月季重瓣化可能与RhAG表达区域被低温抑制有关 | 雌雄蕊起源 | Noor et al., |
| E类功能基因(MADS-box) | SEP3 | 莲NnSEP3过表达导致花萼和花瓣数量减少, 类似心皮的器官增加 | 雌雄蕊起源 | 林钟员, |
| SEP4 | 万寿菊TeSEP4过表达导致花瓣和雄蕊数量减少, 该蛋白可与多种MADS-box蛋白互作 | 雌雄蕊起源 | Zhang et al., | |
| 干细胞决定基因 | WUS | WUS与CLV可形成负反馈回路调控花分生组织活性的终止。WUS表达受AG和SEP抑制。WUS功能缺失导致干细胞和花器官数量减少 | 其它 | Ito et al., |
| 边界基因 | PTL | PTL可通过维持生长素平衡确保花瓣正常发育。拟南芥PTL缺失可导致花瓣减少。PTL抑制C类功能基因在第1、2轮花器官的表达 | 其它 | Lampugnani et al., |
| HAN | HAN可通过维持边界细胞中细胞分裂素的平衡而调控花器官发育。其突变导致拟南芥萼片融合及花瓣和雄蕊数量减少 | 其它 | Zhao et al., | |
| CUC | CUCs在花器官边界表达并抑制细胞增殖。CUC1和CUC2基因受miRNA164a-c负调控, CUCs双重突变导致萼片融合和花瓣数量减少 | 其它 | Aida et al., | |
| SUP | SUP主要控制第3轮与第4轮花器官之间的边界, 可能影响生长素合成。sup/ag-1双突变体在心皮位置较ag-1具有更多轮次花瓣的重瓣性状 | 其它 | Prunet et al., | |
| RBE | RBE可通过抑制AG表达而控制第2轮与第3轮花器官边界。其突变导致花瓣被雄蕊或雄蕊类似器官替代 | 其它 | Krizek et al., | |
| 对称性基因 | CYC | 向日葵和非洲菊CYC同源基因的异位表达导致管状花向舌状花转变, 并通过调控细胞增殖影响舌状花花瓣的形状和大小 | 花序起源 | Fambrini et al., |
| 其它 | MYB | 月季中沉默RhMYB123导致雄蕊瓣化, RhAG、RhAGL15和RhSHP1及生长素信号转导途径中多个基因的表达量下降 | 雌雄蕊起源 | Li et al., |
| PAN | 拟南芥pan突变体前三轮花器官的数量均增加。其可能通过控制花器官原基起始时间间隔影响花器官的数量和位置 | 其它 | Running and Meyerowitz, | |
| ULT | 拟南芥中ULT1突变导致4轮花器官数量均增多。其可能通过调节WUS-AG反馈环参与干细胞的终止 | 其它 | Fletcher, |
Table 1 Key transcription factor genes and their functions involved in petal doubling of flower
| 类型 | 名称 | 功能及表型 | 重瓣起源类型 | 参考文献 |
|---|---|---|---|---|
| A类功能基因 | AP2 | 月季RcAP2表达水平受低温诱导而被高温抑制, 拟南芥异源过表达RcAP2可诱导雄蕊瓣化 | 雌雄蕊起源 | Han et al., |
| B类功能基因(MADS-box) | PI | 拟南芥中异源过表达桃PpPI可诱导雄蕊瓣化; 拟南芥中异源过表达枇杷EjPI可导致萼片瓣化 | 雌雄蕊起源或苞片起源 | Xia et al., |
| AP3 | 夏堇中共表达TfDEF (AP3)和TfGLO (PI)可导致萼片瓣化; 共抑制两者可导致花瓣萼片化 | 苞片起源 | Sasaki et al., | |
| C类功能基因(MADS-box) | AG | 矮牵牛、三花龙胆和月季中沉默AG同源基因导致雄蕊瓣化; 低温诱导的月季重瓣化可能与RhAG表达区域被低温抑制有关 | 雌雄蕊起源 | Noor et al., |
| E类功能基因(MADS-box) | SEP3 | 莲NnSEP3过表达导致花萼和花瓣数量减少, 类似心皮的器官增加 | 雌雄蕊起源 | 林钟员, |
| SEP4 | 万寿菊TeSEP4过表达导致花瓣和雄蕊数量减少, 该蛋白可与多种MADS-box蛋白互作 | 雌雄蕊起源 | Zhang et al., | |
| 干细胞决定基因 | WUS | WUS与CLV可形成负反馈回路调控花分生组织活性的终止。WUS表达受AG和SEP抑制。WUS功能缺失导致干细胞和花器官数量减少 | 其它 | Ito et al., |
| 边界基因 | PTL | PTL可通过维持生长素平衡确保花瓣正常发育。拟南芥PTL缺失可导致花瓣减少。PTL抑制C类功能基因在第1、2轮花器官的表达 | 其它 | Lampugnani et al., |
| HAN | HAN可通过维持边界细胞中细胞分裂素的平衡而调控花器官发育。其突变导致拟南芥萼片融合及花瓣和雄蕊数量减少 | 其它 | Zhao et al., | |
| CUC | CUCs在花器官边界表达并抑制细胞增殖。CUC1和CUC2基因受miRNA164a-c负调控, CUCs双重突变导致萼片融合和花瓣数量减少 | 其它 | Aida et al., | |
| SUP | SUP主要控制第3轮与第4轮花器官之间的边界, 可能影响生长素合成。sup/ag-1双突变体在心皮位置较ag-1具有更多轮次花瓣的重瓣性状 | 其它 | Prunet et al., | |
| RBE | RBE可通过抑制AG表达而控制第2轮与第3轮花器官边界。其突变导致花瓣被雄蕊或雄蕊类似器官替代 | 其它 | Krizek et al., | |
| 对称性基因 | CYC | 向日葵和非洲菊CYC同源基因的异位表达导致管状花向舌状花转变, 并通过调控细胞增殖影响舌状花花瓣的形状和大小 | 花序起源 | Fambrini et al., |
| 其它 | MYB | 月季中沉默RhMYB123导致雄蕊瓣化, RhAG、RhAGL15和RhSHP1及生长素信号转导途径中多个基因的表达量下降 | 雌雄蕊起源 | Li et al., |
| PAN | 拟南芥pan突变体前三轮花器官的数量均增加。其可能通过控制花器官原基起始时间间隔影响花器官的数量和位置 | 其它 | Running and Meyerowitz, | |
| ULT | 拟南芥中ULT1突变导致4轮花器官数量均增多。其可能通过调节WUS-AG反馈环参与干细胞的终止 | 其它 | Fletcher, |
| 类型 | 作用机制 | 相关基因及其在重瓣化进程中的功能 | 参考文献 |
|---|---|---|---|
| DNA甲基化 | 常发生在CG、CHG和CHH (H包括A、T或C)的胞嘧啶C上, 影响染色质结构、DNA构象及基因表达 | 拟南芥甲基转移酶MET1缺失导致基因组DNA总体甲基化水平下降, 引起花器官同源异型转化及数量变化 | Finnegan et al., |
| 低温可通过诱导月季RhAG启动子中CHH位点甲基化水平升高而抑制RhAG表达, 导致月季雄蕊瓣化 | Ma et al., | ||
| 组蛋白甲基化 | 在转录调节和异染色质形成中发挥重要作用。通常将H3K9和H3K27甲基化作为抑制标记, 而H3K4以及H3K36甲基化则作为激活标记 | 水稻去甲基酶JMJ706功能丧失导致H3K9甲基化程度增强, 影响花的形态和器官数量 | Sun and Zhou, |
| 拟南芥AG蛋白能够通过调控WUS或KNU基因位点的H3K27甲基化修饰状态抑制WUS表达, 进而调控花分生组织的发育 | Liu et al., | ||
| TrxG蛋白家族可调控H3K4甲基化, 其家族成员的突变导致拟南芥雄蕊和花瓣间的同源异型转变 | Grini et al., | ||
| 组蛋白乙酰化 | 受组蛋白乙酰转移酶和脱乙酰酶调控。以赖氨酸乙酰化较常见, 其通常具有激活靶基因的作用 | AP2能够招募组蛋白脱乙酰化酶复合物TPL-HDA19而抑制AG的表达 | Krogan et al., |
| 月季生长素响应因子RhARF18能招募组蛋白脱乙酰酶RhHDA6到RhAG启动子上, 降低H3K9/K14乙酰化水平进而抑制其转录, 引起雄蕊瓣化 | Chen et al., | ||
| 染色质重塑 | 包括核小体位置改变、核小体去组装及组蛋白修饰等, 需要ATP依赖的染色质重塑复合物参与 | 拟南芥染色质重塑SWI/SNF复合物成员SYD和BRM基因突变均导致第2、3轮花器官的同源异型转化。SYD和BRM均能诱导AP3和AG基因表达从而启动花器官身份决定 | Hurtado et al., |
| miRNA调控 | 通过切割靶基因mRNA、抑制其翻译以及介导DNA甲基化等方式调控基因的表达 | 在桃、香石竹、矮牵牛、月季和玫瑰等植物中发现miRNA172能够抑制AP2类基因的表达进而影响花器官形态建成。该调控机制在真双子叶植物中保守 | Gattolin et al., |
| 拟南芥miR164能够通过调控边界基因CUCs的表达影响花器官数量。草莓中miRNA164-CUC也参与花型调控 | Laufs et al., | ||
| 金鱼草和矮牵牛miR169通过对NF-YA的转录后抑制降低C类基因的活性。miR169突变导致C类基因异位表达, 进而产生雄蕊化的花瓣 | Cartolano et al., | ||
| 山茶miRNA156可负调控CjSPL1/2表达, 使其仅在雄蕊花丝而不在花药中表达, 进而引起雄蕊瓣化 | Li et al., |
Table 2 Epigenetic regulations and mechanisms involved in petal doubling of flower
| 类型 | 作用机制 | 相关基因及其在重瓣化进程中的功能 | 参考文献 |
|---|---|---|---|
| DNA甲基化 | 常发生在CG、CHG和CHH (H包括A、T或C)的胞嘧啶C上, 影响染色质结构、DNA构象及基因表达 | 拟南芥甲基转移酶MET1缺失导致基因组DNA总体甲基化水平下降, 引起花器官同源异型转化及数量变化 | Finnegan et al., |
| 低温可通过诱导月季RhAG启动子中CHH位点甲基化水平升高而抑制RhAG表达, 导致月季雄蕊瓣化 | Ma et al., | ||
| 组蛋白甲基化 | 在转录调节和异染色质形成中发挥重要作用。通常将H3K9和H3K27甲基化作为抑制标记, 而H3K4以及H3K36甲基化则作为激活标记 | 水稻去甲基酶JMJ706功能丧失导致H3K9甲基化程度增强, 影响花的形态和器官数量 | Sun and Zhou, |
| 拟南芥AG蛋白能够通过调控WUS或KNU基因位点的H3K27甲基化修饰状态抑制WUS表达, 进而调控花分生组织的发育 | Liu et al., | ||
| TrxG蛋白家族可调控H3K4甲基化, 其家族成员的突变导致拟南芥雄蕊和花瓣间的同源异型转变 | Grini et al., | ||
| 组蛋白乙酰化 | 受组蛋白乙酰转移酶和脱乙酰酶调控。以赖氨酸乙酰化较常见, 其通常具有激活靶基因的作用 | AP2能够招募组蛋白脱乙酰化酶复合物TPL-HDA19而抑制AG的表达 | Krogan et al., |
| 月季生长素响应因子RhARF18能招募组蛋白脱乙酰酶RhHDA6到RhAG启动子上, 降低H3K9/K14乙酰化水平进而抑制其转录, 引起雄蕊瓣化 | Chen et al., | ||
| 染色质重塑 | 包括核小体位置改变、核小体去组装及组蛋白修饰等, 需要ATP依赖的染色质重塑复合物参与 | 拟南芥染色质重塑SWI/SNF复合物成员SYD和BRM基因突变均导致第2、3轮花器官的同源异型转化。SYD和BRM均能诱导AP3和AG基因表达从而启动花器官身份决定 | Hurtado et al., |
| miRNA调控 | 通过切割靶基因mRNA、抑制其翻译以及介导DNA甲基化等方式调控基因的表达 | 在桃、香石竹、矮牵牛、月季和玫瑰等植物中发现miRNA172能够抑制AP2类基因的表达进而影响花器官形态建成。该调控机制在真双子叶植物中保守 | Gattolin et al., |
| 拟南芥miR164能够通过调控边界基因CUCs的表达影响花器官数量。草莓中miRNA164-CUC也参与花型调控 | Laufs et al., | ||
| 金鱼草和矮牵牛miR169通过对NF-YA的转录后抑制降低C类基因的活性。miR169突变导致C类基因异位表达, 进而产生雄蕊化的花瓣 | Cartolano et al., | ||
| 山茶miRNA156可负调控CjSPL1/2表达, 使其仅在雄蕊花丝而不在花药中表达, 进而引起雄蕊瓣化 | Li et al., |
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