植物学报 ›› 2018, Vol. 53 ›› Issue (5): 726-737.DOI: 10.11983/CBB17078 cstr: 32102.14.CBB17078
• 专题论坛 • 上一篇
收稿日期:2017-04-09
接受日期:2017-08-30
出版日期:2018-09-01
发布日期:2018-11-29
通讯作者:
陈国雄
作者简介: 共同第一作者。
基金资助:
Zhang Jiwei1,2, Zhao Jiecai1, Zhou Qin1,2, Chen Guoxiong1,*(
)
Received:2017-04-09
Accepted:2017-08-30
Online:2018-09-01
Published:2018-11-29
Contact:
Chen Guoxiong
About author: These authors contributed equally to this paper
摘要: 表皮毛是大多数植物地上部分表皮组织所延伸出来的一种特化的毛状结构附属物。表皮毛在植物表皮层和环境间构筑了一道天然的物理屏障, 不但对植物的生长发育具有重要意义, 而且还具有非常高的应用价值和经济价值。近几年, 研究者从不同植物中不断克隆出新的表皮毛发育相关基因, 在揭示植物调控表皮毛生长发育的分子机制方面取得很大进展。该文综述了植物表皮毛的最新研究进展, 并展望了植物表皮毛的研究方向及应用开发价值。
张继伟, 赵杰才, 周琴, 陈国雄. 植物表皮毛研究进展. 植物学报, 2018, 53(5): 726-737.
Zhang Jiwei, Zhao Jiecai, Zhou Qin, Chen Guoxiong. Progress in Research of Plant Trichome. Chinese Bulletin of Botany, 2018, 53(5): 726-737.
| 基因名称 | 物种 | 基因序列号 | 产物 | 功能及突变体表型描述 | 参考文献 |
|---|---|---|---|---|---|
| GL1 | 拟南芥 | AF495524 | R2R3-MYB类型转录因子 | 调控表皮毛的启动, 突变体表现为无表皮毛, 但过表达反而导致表皮毛减少 | |
| TTG1 | 拟南芥 | BT011856 | 含有WD40重复序列蛋白 | 调控表皮毛的启动, 突变体表现为无表皮毛, 且突变体种子的休眠特性消失 | |
| GL3 | 拟南芥 | AF246291 | bHLH类型转录因子 | 调控表皮毛的启动, 突变体表现为表皮毛数量减少 | |
| GL2 | 拟南芥 | AJ457046 | HD2-ZIP结构转录因子 | 调控表皮细胞命运, 突变体表皮毛减少, 且大部分无分枝 | |
| TTG2 | 拟南芥 | AF404862 | WRKY类型转录因子 | 控制表皮毛早期发育, 突变体表现为分枝减少或无分枝 | |
| EGL3 | 拟南芥 | AF251687 | bHLH类型转录因子 | 正调节因子, 功能与GL3冗余, 而且gl3/egl3双突变体表现为无表皮毛 | |
| MYB23 | 拟南芥 | Z68158 | R2R3-MYB类型转录因子 | 异位表达可以促进表皮毛生长 | |
| TRY | 拟南芥 | AY161286 | R3-MYB类型转录因子 | 负调节因子, 突变体表皮毛成簇生长且分枝增加 | |
| CPC | 拟南芥 | AB004871 | R3-MYB类型转录因子 | 负调节因子, 突变体表现为根毛减少, 表皮毛簇生 | |
| WER | 拟南芥 | Z95789 | R2R3-MYB类型转录因子 | 控制根毛发育, 功能缺失导致根毛显著增加 | |
| ETC1 | 拟南芥 | AY519518 | MYB类型转录因子 | 负调节因子 | |
| ETC2 | 拟南芥 | AY234411 | MYB类型转录因子 | 负调节因子 | |
| ETC3 | 拟南芥 | AB264292 | MYB类型转录因子 | 突变体表现为表皮毛分枝增加而根毛密度减少 | |
| KAK | 拟南芥 | AY265959 | 泛素连接酶 | 调节表皮毛核内复制, 突变体表现为表皮毛分枝增加, DNA含量增加 | |
| GIS | 拟南芥 | BT003340 | 含C2H2的转录因子 | 调控表皮细胞和花序的分化, 突变体表皮毛减少 | |
| GIS2 | 拟南芥 | NM_120748 | 含C2H2的转录因子 | 调控表皮毛的分化, 受赤霉素和细胞分裂素的调节 | |
| RHL2 | 拟南芥 | AJ297842 | DNA拓扑异构酶 | 调节表皮毛核内复制, 调控表皮毛形态 | |
| STI | 拟南芥 | AF264023 | 未知 | 调控表皮毛分枝形成, 突变体表现为表皮毛分枝减少 | |
| KLK | 拟南芥 | AY496701 | 未知 | 调控表皮毛分枝形成 | |
| BRICK1 | 拟南芥 | AY085644 | 未知 | 调控表皮毛分枝形成 | |
| DIS1 | 拟南芥 | AF507911 | 未知 | 调控表皮毛分枝形成, 突变体表现为表皮毛细胞扩增, 且形态不正常 | |
| DIS2 | 拟南芥 | AY084256 | 未知 | 调控表皮毛分枝形成 | |
| SPL9 | 拟南芥 | AJ011638 | 未知 | 受miR156调控, 抑制表皮毛的启动 | |
| TEM | 拟南芥 | AY056169 | RAV转录因子 | 光周期响应因子, 参与调控赤霉素合成与转运, 调节花期与表皮毛形成等 | |
| TCS1 | 拟南芥 | AK222122 | 未知 | 突变体表现为表皮毛有缺陷 | |
| 基因名称 | 物种 | 基因序列号 | 产物 | 功能及突变体表型描述 | 参考文献 |
| TRIL | 黄瓜 | 未知 | HD-ZIP结构转录因子 | 调控表皮毛的启动, 突变体表现为无表皮毛 | |
| CsGL1 | 黄瓜 | 未知 | HD-ZIP结构转录因子 | 调控表皮毛的启动, 突变体表现为无表皮毛 | |
| GoPGF | 棉花 | KP072743 | bHLH类型转录因子 | 调控表皮毛的启动, 沉默GoPGF表现为无表皮毛 | |
| GhMYB2A | 棉花 | JQ868556 | MYB类型转录因子 | 调控棉花纤维形成, 受miRNA调节 | |
| GhMYB2D | 棉花 | JQ868560 | MYB类型转录因子 | 调控棉花纤维形成, 受miRNA调节 | |
| AaMYB1 | 黄花蒿 | KC118530 | MYB类型转录因子 | 调控表皮毛发育和萜类代谢 | |
| AaHD1 | 黄花蒿 | KU744599 | HD-ZIP结构转录因子 | 调控表皮毛发育, 过表达导致表皮毛密度增加 | |
| HI | 番茄 | 未知 | SRA1的1个亚基 | 调控肌动蛋白丝凝聚, 突变体表皮毛形态有缺陷 |
表1 植物表皮毛发育相关重要基因
Table 1 The important genes related to trichome development in plants
| 基因名称 | 物种 | 基因序列号 | 产物 | 功能及突变体表型描述 | 参考文献 |
|---|---|---|---|---|---|
| GL1 | 拟南芥 | AF495524 | R2R3-MYB类型转录因子 | 调控表皮毛的启动, 突变体表现为无表皮毛, 但过表达反而导致表皮毛减少 | |
| TTG1 | 拟南芥 | BT011856 | 含有WD40重复序列蛋白 | 调控表皮毛的启动, 突变体表现为无表皮毛, 且突变体种子的休眠特性消失 | |
| GL3 | 拟南芥 | AF246291 | bHLH类型转录因子 | 调控表皮毛的启动, 突变体表现为表皮毛数量减少 | |
| GL2 | 拟南芥 | AJ457046 | HD2-ZIP结构转录因子 | 调控表皮细胞命运, 突变体表皮毛减少, 且大部分无分枝 | |
| TTG2 | 拟南芥 | AF404862 | WRKY类型转录因子 | 控制表皮毛早期发育, 突变体表现为分枝减少或无分枝 | |
| EGL3 | 拟南芥 | AF251687 | bHLH类型转录因子 | 正调节因子, 功能与GL3冗余, 而且gl3/egl3双突变体表现为无表皮毛 | |
| MYB23 | 拟南芥 | Z68158 | R2R3-MYB类型转录因子 | 异位表达可以促进表皮毛生长 | |
| TRY | 拟南芥 | AY161286 | R3-MYB类型转录因子 | 负调节因子, 突变体表皮毛成簇生长且分枝增加 | |
| CPC | 拟南芥 | AB004871 | R3-MYB类型转录因子 | 负调节因子, 突变体表现为根毛减少, 表皮毛簇生 | |
| WER | 拟南芥 | Z95789 | R2R3-MYB类型转录因子 | 控制根毛发育, 功能缺失导致根毛显著增加 | |
| ETC1 | 拟南芥 | AY519518 | MYB类型转录因子 | 负调节因子 | |
| ETC2 | 拟南芥 | AY234411 | MYB类型转录因子 | 负调节因子 | |
| ETC3 | 拟南芥 | AB264292 | MYB类型转录因子 | 突变体表现为表皮毛分枝增加而根毛密度减少 | |
| KAK | 拟南芥 | AY265959 | 泛素连接酶 | 调节表皮毛核内复制, 突变体表现为表皮毛分枝增加, DNA含量增加 | |
| GIS | 拟南芥 | BT003340 | 含C2H2的转录因子 | 调控表皮细胞和花序的分化, 突变体表皮毛减少 | |
| GIS2 | 拟南芥 | NM_120748 | 含C2H2的转录因子 | 调控表皮毛的分化, 受赤霉素和细胞分裂素的调节 | |
| RHL2 | 拟南芥 | AJ297842 | DNA拓扑异构酶 | 调节表皮毛核内复制, 调控表皮毛形态 | |
| STI | 拟南芥 | AF264023 | 未知 | 调控表皮毛分枝形成, 突变体表现为表皮毛分枝减少 | |
| KLK | 拟南芥 | AY496701 | 未知 | 调控表皮毛分枝形成 | |
| BRICK1 | 拟南芥 | AY085644 | 未知 | 调控表皮毛分枝形成 | |
| DIS1 | 拟南芥 | AF507911 | 未知 | 调控表皮毛分枝形成, 突变体表现为表皮毛细胞扩增, 且形态不正常 | |
| DIS2 | 拟南芥 | AY084256 | 未知 | 调控表皮毛分枝形成 | |
| SPL9 | 拟南芥 | AJ011638 | 未知 | 受miR156调控, 抑制表皮毛的启动 | |
| TEM | 拟南芥 | AY056169 | RAV转录因子 | 光周期响应因子, 参与调控赤霉素合成与转运, 调节花期与表皮毛形成等 | |
| TCS1 | 拟南芥 | AK222122 | 未知 | 突变体表现为表皮毛有缺陷 | |
| 基因名称 | 物种 | 基因序列号 | 产物 | 功能及突变体表型描述 | 参考文献 |
| TRIL | 黄瓜 | 未知 | HD-ZIP结构转录因子 | 调控表皮毛的启动, 突变体表现为无表皮毛 | |
| CsGL1 | 黄瓜 | 未知 | HD-ZIP结构转录因子 | 调控表皮毛的启动, 突变体表现为无表皮毛 | |
| GoPGF | 棉花 | KP072743 | bHLH类型转录因子 | 调控表皮毛的启动, 沉默GoPGF表现为无表皮毛 | |
| GhMYB2A | 棉花 | JQ868556 | MYB类型转录因子 | 调控棉花纤维形成, 受miRNA调节 | |
| GhMYB2D | 棉花 | JQ868560 | MYB类型转录因子 | 调控棉花纤维形成, 受miRNA调节 | |
| AaMYB1 | 黄花蒿 | KC118530 | MYB类型转录因子 | 调控表皮毛发育和萜类代谢 | |
| AaHD1 | 黄花蒿 | KU744599 | HD-ZIP结构转录因子 | 调控表皮毛发育, 过表达导致表皮毛密度增加 | |
| HI | 番茄 | 未知 | SRA1的1个亚基 | 调控肌动蛋白丝凝聚, 突变体表皮毛形态有缺陷 |
图1 植物表皮毛分化调控模式该模式显示, 赤霉素在叶肉细胞中的合成及转运是由TEM负责调控, 并在表皮毛前体细胞下面的叶肉细胞中积累和向上转运。赤霉素通过DELLA和GIS调控GL1-GL3-TTG1复合体, 茉莉酸通过JAZ也调控该复合体, 赤霉素和细胞分裂素调节GIS2, 然后, GL1-GL3-TTG1复合体和GIS2进一步调节GL2, 指导表皮毛发育。TRY等负调节因子移动至邻近表皮细胞, 与GL1竞争性结合到GL3上, 阻碍GL2的表达, 从而抑制表皮毛形成。
Figure 1 The regulate pattern of plant trichome differentiationThe diagram shows the synthesis and transport of GA in mesophyll are regulated by TEM, which also accumulate in mesophyll underneath pre-trichome and transfer to trichome. GA regulates the GL1-GL3-TTG1 complex through DELLA and GIS, JA also regulates the complex through JAZ, GA and CK regulate GIS2, then the complex and GIS2 guide trichome development by regulating GL2. The negative regulator TRY moves to neighboring epidermis where it competes with GL1 for binding to GL3, which suppresses the expression of GL2 and inhibit the trichome formation.
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