生物技术通报 ›› 2026, Vol. 42 ›› Issue (9): 1-13.doi: 10.13560/j.cnki.biotech.bull.1985.2025-1037
• 综述与专论 •
收稿日期:2025-09-26
出版日期:2026-02-09
发布日期:2026-02-09
通讯作者:
陈赢男,女,博士,教授,研究方向 :林木遗传育种;E-mail: chenyingnan@njfu.edu.cn作者简介:秦子璐,女,博士研究生,研究方向 :林木遗传育种;E-mail: qinzilu@njfu.edu.cn
基金资助:
QIN Zi-lu(
), SUN Hai-yan, CHEN Ying-nan(
)
Received:2025-09-26
Published:2026-02-09
Online:2026-02-09
摘要:
植物表皮毛是植物表皮细胞分化形成的特化突起结构,作为植物长期适应环境的重要进化特征,其在抵御生物与非生物胁迫及形成棉纤维、茶叶次生代谢物等产物中具有关键作用。本文系统综述了草本植物(拟南芥、番茄、棉花及水稻)和木本植物(杨树、茶树及桃树)表皮毛发育分子调控机制的研究进展。在拟南芥中,MYB-bHLH-WD40(MBW)转录复合体构成调控表皮毛起始与形态建成的核心保守模块,通过正负反馈环路实现精准调控。番茄中则呈现出以HD-Zip转录因子为主导的、更为复杂的调控网络。在棉纤维发育过程中,R2R3 MYB等转录因子展现出关键作用及功能特异性分化。作为单子叶植物,水稻表皮毛发育形成了区别于双子叶植物的独特调控网络。杨树、茶树等木本植物仍存在部分保守MBW核心调控机制,而桃果实表皮毛发育则演化出不依赖完整MBW复合体的新路径。本文通过比较分析不同物种表皮毛调控网络的保守性与特异性,为揭示植物表皮毛发育的分子进化机制提供了理论依据,并为作物抗逆性与经济性状的分子育种改良提供潜在靶点。
秦子璐, 孙海燕, 陈赢男. 植物表皮毛发育分子调控机制研究进展[J]. 生物技术通报, 2026, 42(9): 1-13.
QIN Zi-lu, SUN Hai-yan, CHEN Ying-nan. Research Progress in the Molecular Regulatory Mechanism of Plant Trichome Development[J]. Biotechnology Bulletin, 2026, 42(9): 1-13.
植物 Plant | 基因家族 Gene family | 基因名称 Gene name | 转基因或突变体材料表型 Phenotype of transgenic or mutant materials | 互作蛋白 Interacting protein | 下游调控基因 Downstream regulatory gene | 参考文献 Reference |
|---|---|---|---|---|---|---|
拟南芥 Arabidopsis thaliana | R2R3 MYB | AtNOK | 野生型中nok突变体的表皮毛分支数量增加,过表达NOK分支数量减少; gl3-sst nok突变体中,过表达AtNOK导致额外的分支形成 | --- | --- | [ |
| R2R3 MYB | AtGL1 | gl1突变体叶片无毛; 过表达AtGL1叶片表皮毛减少 | AtGL3/AtEGL3; AtTTG1 | AtGL2 | [ | |
| R2R3 MYB | AtMYB23 | mby23突变体表皮毛分支减少; 过表达AtMYB23导致下胚轴产生异位表皮毛 | AtGL3/AtEGL3; AtTTG1 | AtGL2 | [ | |
| R2R3 MYB | AtMYB82 | AtGL1启动子驱动AtMYB82可以回补gl1突变体的表皮毛缺陷; 将AtMYB82与显性抑制结构域(SRDX)融合以抑制其功能,导致拟南芥仅叶缘残留少量表皮毛; 过表达N端截短型AtMYB82,导致拟南芥出现无毛叶片 | --- | --- | [ | |
| bHLH | AtGL3 | gl3突变体表皮毛分支减少 | AtGl1;AtTTG1 | AtGL2 | [ | |
| bHLH | AtEGL3 | egl3突变体表皮毛数量和分支减少; gl3 egl3双突变体完全无毛 | AtGl1;AtTTG1 | AtGL2 | [ | |
| WD40 | AtTTG1 | ttg1突变体叶片和茎无毛 | AtGL1; AtGL3/AtEGL3 | AtGL2 | [ | |
| HD-Zip | AtGL2 | gl2突变体表皮毛分支减少或畸形 | --- | --- | [ | |
| R3 MYB | AtCPC | cpc突变体表皮毛增加; 过表达AtCPC导致表皮毛减少 | AtGL3/AtEAtGL3; AtTTG1 | --- | [ | |
| R3 MYB | AtTRY | try突变体表皮毛成簇生长且分支增加; try cpc双突变体中出现更大的叶表皮毛簇 | AtGL3/AtEGL3; AtTTG1 | --- | [ | |
番茄 Solanum lycopersicum | HD-Zip | SlCD2 | 突变体中叶片VI型腺毛密度减少 | --- | --- | [ |
| HD-Zip | SlWo | SlWo的功能获得性突变体LA3186导致叶片I型毛状体密度增加; SlWo-RNAi则使叶片I型毛状体几乎消失; woW106R 显性负突变体的花药边缘连锁表皮毛数量减少; WoP635R 功能增强突变体的花药边缘连锁表皮毛数量显著增加 | SlCycB2 | --- | [ | |
| HD-Zip | SlHZ45 | 过表达SlHZ45增加了番茄叶片Ⅰ、Ⅳ、Ⅵ型腺体毛 | --- | --- | [ | |
| HD-Zip | SlLn | 过表达SlLn使叶片I、III、V型表皮毛密度增加; 敲除SlLn表皮毛密度明显降低 | SlWo; SlH | SlCycB2; SlCycB3 | [ | |
| HD-Zip | SlHD7/SlHD7L | cr-hd7/hd7l双敲除突变体的花药边缘连锁表皮毛几乎完全缺失 | --- | --- | [ | |
| C2H2 | SlH | 敲除SlH抑制叶片I型表皮毛形成 | SlWo;SlLn | --- | [ | |
| R2R3 MYB | SlMX1 | 过表达SlMX1叶表皮毛增加; SlMX1-RNAi株系叶片表皮毛减少 | --- | --- | [ | |
| MYB-like | SlGCR1/SlGCR2 | gcr1中叶片非腺毛转化为腺毛; gcr2无显著表型; gcr1/2双突变体叶片腺毛显著增多 | --- | SlLFS | [ | |
| bHLH | SlMYC1 | 敲低SlMYC1使叶片VI型腺毛密度降低;myc1纯合突变体和CRISPR-Cas9敲除株系,叶片Ⅵ型腺毛完全缺失 | [ | |||
| --- | SlHl | hl突变体叶片表皮毛严重弯曲、缩短 | --- | --- | [ | |
| --- | SlHl-2 | hl-2突变体叶片和茎杆表皮毛畸形; 过表达SlHl-2可恢复hl-2突变体的叶片表皮毛; CRISPR敲除或RNAi沉默SlHl-2导致野生型叶片表皮毛扭曲 | --- | --- | [ | |
| HD-Zip | SlHDZIV8 | 敲低SlHDZIV8使叶片表皮毛出现扭曲、密度降低 | --- | SlHl-2 | [ | |
| B-type cyclin | SlCycB2/SlCycB3 | 过表达SlCycB2会导致叶片几乎所有非腺毛和腺毛消失,仅残留异常Ⅲ型类似毛; SlCycB2-RNAi抑制其表达则促进叶片Ⅲ、Ⅴ型非腺毛显著增加; 同源基因SlCycB3与SlCycB2功能相似 | --- | --- | [ | |
棉 Gossypium | WD40 | GhTTG1;GhTTG3 | GhTTG1和GhTTG3能恢复拟南芥ttg1突变体的表皮毛;在拟南芥中过表达GhTTG1或GhTTG3使叶片表皮毛增加 | --- | --- | [ |
| R2R3 MYB | GhMYB25 | 在烟草中过表达GhMYB25表皮毛数量增加 | --- | --- | [ | |
| R2R3 MYB | GhMYB25-like | GhMYB25-like- RNAi使棉花种子几乎无纤维 | --- | GhMYB25; GhMYB109 | [ | |
| R2R3 MYB | GhMML4_D12 | VIGS沉默GhMML4_D12后长纤维减少 | --- | --- | [ | |
| R2R3 MYB | GhMML3 | 沉默GhMML3会导致短绒纤维缺失,长纤维产量也大幅减少 | --- | --- | [ | |
| R2R3 MYB | GaMYB2 | 在拟南芥中,AtGL1启动子驱动的GaMYB2可回补gl1突变体的表皮毛; 35S启动子驱动的GaMYB2使野生型和gl1突变体拟南芥种子产生表皮毛 | --- | --- | [ | |
| HD-Zip | GaHOX1 | AtGL2启动子驱动下可完全恢复拟南芥gl2-2突变体表皮毛; 35S启动子驱动时抑制野生型表皮毛发育,莲座叶表皮毛密度也降低 | --- | --- | [ | |
| R2R3 MYB | GhMYB212 | 敲除GhMYB212导致棉花纤维长度缩短 | --- | GhSWEET12 | [ | |
| R2R3 MYB | GhMYB109 | 反义抑制GhMYB109使纤维伸长受阻 | --- | GhACO1; GhACO2 | [ | |
| R2R3 MYB | GhMYB102 | 过表达GhMYB102会导致纤维变短、次生壁增厚 | --- | GhIRX10 | [ | |
| R2R3 MYB | GhMYB30 | 过表达GhMUR3会使纤维长度显著缩短,纤维细度增加; GhMUR3-RNAi会使纤维细度下降 | --- | GhMUR3 | [ | |
| HD-Zip | GhHOX3 | GhHOX3共抑制和RNAi株系纤维长度减少,共抑制株系长纤维完全消失,仅残留极短纤维; 过表达株系纤维长度增加 | --- | GhRDL1 | [ | |
水稻 Oryza sativa | WOX | OsWOX3B/LOC_Os05g02730 | 在有毛品种中通过RNAi抑制OsWOX3B表达后,叶片和颖壳均无毛; 将OsWOX3B基因由自身启动子驱动转入HMK后,转基因植株叶片恢复有毛表型; glr1突变体叶片和颖壳光滑无毛 | --- | --- | [ |
| --- | LOC_Os01g70100 | glr2突变体叶片的长毛、微毛和颖壳表皮毛缺失,但腺毛发育正常 | --- | --- | [ | |
| SBP-box | OsGLR3/OsSPL10 | glr3叶片和颖壳光滑无毛 | --- | OsSCR1 | [ | |
| GRAS | OsSCR1/2 | osscr1的长毛和微毛数量显著减少,腺毛发育正常,osscr2无明显变化; osscr1 osscr2双突变体几乎无长毛和微毛,腺毛数量减少; 过表达OsSCR1或OsSCR2,叶片3种表皮毛数量均显著增加; 将由自身启动子启动的OsSCR1或OsSCR2转入osscr1 osscr2双突变体,OsSCR1可完全恢复双突变体的表皮毛缺陷,OsSCR2仅能部分恢复 | --- | OsWOX3B | [ |
表1 草本植物表皮毛发育的分子调控机制
Table 1 Molecular regulatory mechanisms of trichome development in herbaceous plants
植物 Plant | 基因家族 Gene family | 基因名称 Gene name | 转基因或突变体材料表型 Phenotype of transgenic or mutant materials | 互作蛋白 Interacting protein | 下游调控基因 Downstream regulatory gene | 参考文献 Reference |
|---|---|---|---|---|---|---|
拟南芥 Arabidopsis thaliana | R2R3 MYB | AtNOK | 野生型中nok突变体的表皮毛分支数量增加,过表达NOK分支数量减少; gl3-sst nok突变体中,过表达AtNOK导致额外的分支形成 | --- | --- | [ |
| R2R3 MYB | AtGL1 | gl1突变体叶片无毛; 过表达AtGL1叶片表皮毛减少 | AtGL3/AtEGL3; AtTTG1 | AtGL2 | [ | |
| R2R3 MYB | AtMYB23 | mby23突变体表皮毛分支减少; 过表达AtMYB23导致下胚轴产生异位表皮毛 | AtGL3/AtEGL3; AtTTG1 | AtGL2 | [ | |
| R2R3 MYB | AtMYB82 | AtGL1启动子驱动AtMYB82可以回补gl1突变体的表皮毛缺陷; 将AtMYB82与显性抑制结构域(SRDX)融合以抑制其功能,导致拟南芥仅叶缘残留少量表皮毛; 过表达N端截短型AtMYB82,导致拟南芥出现无毛叶片 | --- | --- | [ | |
| bHLH | AtGL3 | gl3突变体表皮毛分支减少 | AtGl1;AtTTG1 | AtGL2 | [ | |
| bHLH | AtEGL3 | egl3突变体表皮毛数量和分支减少; gl3 egl3双突变体完全无毛 | AtGl1;AtTTG1 | AtGL2 | [ | |
| WD40 | AtTTG1 | ttg1突变体叶片和茎无毛 | AtGL1; AtGL3/AtEGL3 | AtGL2 | [ | |
| HD-Zip | AtGL2 | gl2突变体表皮毛分支减少或畸形 | --- | --- | [ | |
| R3 MYB | AtCPC | cpc突变体表皮毛增加; 过表达AtCPC导致表皮毛减少 | AtGL3/AtEAtGL3; AtTTG1 | --- | [ | |
| R3 MYB | AtTRY | try突变体表皮毛成簇生长且分支增加; try cpc双突变体中出现更大的叶表皮毛簇 | AtGL3/AtEGL3; AtTTG1 | --- | [ | |
番茄 Solanum lycopersicum | HD-Zip | SlCD2 | 突变体中叶片VI型腺毛密度减少 | --- | --- | [ |
| HD-Zip | SlWo | SlWo的功能获得性突变体LA3186导致叶片I型毛状体密度增加; SlWo-RNAi则使叶片I型毛状体几乎消失; woW106R 显性负突变体的花药边缘连锁表皮毛数量减少; WoP635R 功能增强突变体的花药边缘连锁表皮毛数量显著增加 | SlCycB2 | --- | [ | |
| HD-Zip | SlHZ45 | 过表达SlHZ45增加了番茄叶片Ⅰ、Ⅳ、Ⅵ型腺体毛 | --- | --- | [ | |
| HD-Zip | SlLn | 过表达SlLn使叶片I、III、V型表皮毛密度增加; 敲除SlLn表皮毛密度明显降低 | SlWo; SlH | SlCycB2; SlCycB3 | [ | |
| HD-Zip | SlHD7/SlHD7L | cr-hd7/hd7l双敲除突变体的花药边缘连锁表皮毛几乎完全缺失 | --- | --- | [ | |
| C2H2 | SlH | 敲除SlH抑制叶片I型表皮毛形成 | SlWo;SlLn | --- | [ | |
| R2R3 MYB | SlMX1 | 过表达SlMX1叶表皮毛增加; SlMX1-RNAi株系叶片表皮毛减少 | --- | --- | [ | |
| MYB-like | SlGCR1/SlGCR2 | gcr1中叶片非腺毛转化为腺毛; gcr2无显著表型; gcr1/2双突变体叶片腺毛显著增多 | --- | SlLFS | [ | |
| bHLH | SlMYC1 | 敲低SlMYC1使叶片VI型腺毛密度降低;myc1纯合突变体和CRISPR-Cas9敲除株系,叶片Ⅵ型腺毛完全缺失 | [ | |||
| --- | SlHl | hl突变体叶片表皮毛严重弯曲、缩短 | --- | --- | [ | |
| --- | SlHl-2 | hl-2突变体叶片和茎杆表皮毛畸形; 过表达SlHl-2可恢复hl-2突变体的叶片表皮毛; CRISPR敲除或RNAi沉默SlHl-2导致野生型叶片表皮毛扭曲 | --- | --- | [ | |
| HD-Zip | SlHDZIV8 | 敲低SlHDZIV8使叶片表皮毛出现扭曲、密度降低 | --- | SlHl-2 | [ | |
| B-type cyclin | SlCycB2/SlCycB3 | 过表达SlCycB2会导致叶片几乎所有非腺毛和腺毛消失,仅残留异常Ⅲ型类似毛; SlCycB2-RNAi抑制其表达则促进叶片Ⅲ、Ⅴ型非腺毛显著增加; 同源基因SlCycB3与SlCycB2功能相似 | --- | --- | [ | |
棉 Gossypium | WD40 | GhTTG1;GhTTG3 | GhTTG1和GhTTG3能恢复拟南芥ttg1突变体的表皮毛;在拟南芥中过表达GhTTG1或GhTTG3使叶片表皮毛增加 | --- | --- | [ |
| R2R3 MYB | GhMYB25 | 在烟草中过表达GhMYB25表皮毛数量增加 | --- | --- | [ | |
| R2R3 MYB | GhMYB25-like | GhMYB25-like- RNAi使棉花种子几乎无纤维 | --- | GhMYB25; GhMYB109 | [ | |
| R2R3 MYB | GhMML4_D12 | VIGS沉默GhMML4_D12后长纤维减少 | --- | --- | [ | |
| R2R3 MYB | GhMML3 | 沉默GhMML3会导致短绒纤维缺失,长纤维产量也大幅减少 | --- | --- | [ | |
| R2R3 MYB | GaMYB2 | 在拟南芥中,AtGL1启动子驱动的GaMYB2可回补gl1突变体的表皮毛; 35S启动子驱动的GaMYB2使野生型和gl1突变体拟南芥种子产生表皮毛 | --- | --- | [ | |
| HD-Zip | GaHOX1 | AtGL2启动子驱动下可完全恢复拟南芥gl2-2突变体表皮毛; 35S启动子驱动时抑制野生型表皮毛发育,莲座叶表皮毛密度也降低 | --- | --- | [ | |
| R2R3 MYB | GhMYB212 | 敲除GhMYB212导致棉花纤维长度缩短 | --- | GhSWEET12 | [ | |
| R2R3 MYB | GhMYB109 | 反义抑制GhMYB109使纤维伸长受阻 | --- | GhACO1; GhACO2 | [ | |
| R2R3 MYB | GhMYB102 | 过表达GhMYB102会导致纤维变短、次生壁增厚 | --- | GhIRX10 | [ | |
| R2R3 MYB | GhMYB30 | 过表达GhMUR3会使纤维长度显著缩短,纤维细度增加; GhMUR3-RNAi会使纤维细度下降 | --- | GhMUR3 | [ | |
| HD-Zip | GhHOX3 | GhHOX3共抑制和RNAi株系纤维长度减少,共抑制株系长纤维完全消失,仅残留极短纤维; 过表达株系纤维长度增加 | --- | GhRDL1 | [ | |
水稻 Oryza sativa | WOX | OsWOX3B/LOC_Os05g02730 | 在有毛品种中通过RNAi抑制OsWOX3B表达后,叶片和颖壳均无毛; 将OsWOX3B基因由自身启动子驱动转入HMK后,转基因植株叶片恢复有毛表型; glr1突变体叶片和颖壳光滑无毛 | --- | --- | [ |
| --- | LOC_Os01g70100 | glr2突变体叶片的长毛、微毛和颖壳表皮毛缺失,但腺毛发育正常 | --- | --- | [ | |
| SBP-box | OsGLR3/OsSPL10 | glr3叶片和颖壳光滑无毛 | --- | OsSCR1 | [ | |
| GRAS | OsSCR1/2 | osscr1的长毛和微毛数量显著减少,腺毛发育正常,osscr2无明显变化; osscr1 osscr2双突变体几乎无长毛和微毛,腺毛数量减少; 过表达OsSCR1或OsSCR2,叶片3种表皮毛数量均显著增加; 将由自身启动子启动的OsSCR1或OsSCR2转入osscr1 osscr2双突变体,OsSCR1可完全恢复双突变体的表皮毛缺陷,OsSCR2仅能部分恢复 | --- | OsWOX3B | [ |
植物 Plant | 基因家族 Gene family | 基因名称 Gene name | 转基因或突变体材料表型 Phenotype of transgenic or mutant materials | 互作蛋白 Interacting protein | 下游调控基因Downstream regulatory gene | 参考文献 Reference |
|---|---|---|---|---|---|---|
杨树 Populus | R2R3 MYB | PtaMYB186; PtaMYB138;PtaMYB38 | 过表达PtaMYB186表皮毛增加; 同时敲除PtaMYB186、PtaMYB138及PtaMYB38,产生无毛突变体 | --- | --- | [ |
| TCP | PtoTCP19 | miR319a通过靶向抑制PtoTCP19表达,增加叶片表皮毛密度 | PtoRGA; PtoGL1 | [ | ||
| MTA | PtrMTA | 过表达PtrMTA使杨树叶片表皮毛密度增加 | --- | --- | [ | |
| R2R3 MYB | PdeMIXTA04 | 在拟南芥gl1突变体中过表达PdeMIXTA04能回补表型; 在野生型中过表达表皮毛密度提高 | PdeMYC; PdeWD40 | --- | [ | |
| MADS-box | PtrAG; PtrSTK | PtrAG-RNAi使杨树雌花花器官形态异常; 当PtrAG和PtrSTK被RNAi共抑制时,种子和种毛完全缺失 | --- | --- | [ | |
茶树 Camellia sinensis | R2R3 MYB | CsMYB1 | 过表达CsMYB1可以回补拟南芥gl1突变体的表皮毛 | CsGL3; CsWD40 | CsGL2 | [ |
桃树 Prunus persica | R2R3 MYB | PpMYB25 | 在拟南芥中过表达PpMYB25叶片表皮毛增加 | --- | PpMYB26; PpHDG11 | [ |
表2 木本植物表皮毛发育的分子调控机制
Table 2 Molecular regulatory mechanisms of trichome development in woody plants
植物 Plant | 基因家族 Gene family | 基因名称 Gene name | 转基因或突变体材料表型 Phenotype of transgenic or mutant materials | 互作蛋白 Interacting protein | 下游调控基因Downstream regulatory gene | 参考文献 Reference |
|---|---|---|---|---|---|---|
杨树 Populus | R2R3 MYB | PtaMYB186; PtaMYB138;PtaMYB38 | 过表达PtaMYB186表皮毛增加; 同时敲除PtaMYB186、PtaMYB138及PtaMYB38,产生无毛突变体 | --- | --- | [ |
| TCP | PtoTCP19 | miR319a通过靶向抑制PtoTCP19表达,增加叶片表皮毛密度 | PtoRGA; PtoGL1 | [ | ||
| MTA | PtrMTA | 过表达PtrMTA使杨树叶片表皮毛密度增加 | --- | --- | [ | |
| R2R3 MYB | PdeMIXTA04 | 在拟南芥gl1突变体中过表达PdeMIXTA04能回补表型; 在野生型中过表达表皮毛密度提高 | PdeMYC; PdeWD40 | --- | [ | |
| MADS-box | PtrAG; PtrSTK | PtrAG-RNAi使杨树雌花花器官形态异常; 当PtrAG和PtrSTK被RNAi共抑制时,种子和种毛完全缺失 | --- | --- | [ | |
茶树 Camellia sinensis | R2R3 MYB | CsMYB1 | 过表达CsMYB1可以回补拟南芥gl1突变体的表皮毛 | CsGL3; CsWD40 | CsGL2 | [ |
桃树 Prunus persica | R2R3 MYB | PpMYB25 | 在拟南芥中过表达PpMYB25叶片表皮毛增加 | --- | PpMYB26; PpHDG11 | [ |
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