植物学报 ›› 2016, Vol. 51 ›› Issue (4): 565-574.DOI: 10.11983/CBB15137
• 专题论坛 • 上一篇
收稿日期:
2015-08-03
接受日期:
2015-10-13
出版日期:
2016-07-01
发布日期:
2016-08-05
通讯作者:
袁政
作者简介:
# 共同第一作者
基金资助:
Yanjie Yu, Dabing Zhang, Zheng Yuan*
Received:
2015-08-03
Accepted:
2015-10-13
Online:
2016-07-01
Published:
2016-08-05
Contact:
Yuan Zheng
About author:
# Co-first authors
摘要: WOX蛋白家族是植物特有的一类转录因子家族, 是植物胚胎建成、干细胞维持和器官发生等发育过程中的重要调控因子。越来越多的研究表明, 作为干细胞维持的关键因子之一, WOX蛋白家族通过相似或特异的调控网络参与植物初生分生组织(茎尖和根尖分生组织)和次生分生组织(维管分生组织)等各级干细胞的维持和分化。该文综述了近年来WOX蛋白家族调控干细胞发育分子机制的研究进展, 并对其在单、双子叶植物中功能的保守性进行了比较和分析。
于燕杰, 张大兵, 袁政. WOX蛋白家族调控干细胞发育分子机制的研究进展. 植物学报, 2016, 51(4): 565-574.
Yanjie Yu, Dabing Zhang, Zheng Yuan. The Updated Functional Study of WOX Protein Family in Regulating Stem Cell Development. Chinese Bulletin of Botany, 2016, 51(4): 565-574.
图1 WOX家族基因参与调控拟南芥和水稻茎尖及根尖分生组织发育的分子网络示意图(改自Gaillochet et al., 2015) (A) 植物茎尖分生组织结构; (B) 拟南芥茎尖分生组织调控网络: WUS在组织中心特异表达, WUS蛋白经过胞间连丝结构运输到中央区直接与CLV3启动子或者调控元件结合促进CLV3的表达; CLV3在中央区特异表达, CLV3小肽主要通过与受体激酶CLV1和BAM1结合传递信号, 抑制WUS表达, CLV3小肽也可以通过与受体激酶RPK2结合传递信号, 抑制WUS表达; (C) 水稻茎尖分生组织调控网络: FCP1和OsWOX4在茎尖分生组织中表达区域重合, FCP1分泌的小肽可能通过未知的受体传递信号, 抑制OsWOX4的表达; (D) 植物根尖分生组织结构; (E) 拟南芥根尖分生组织调控网络(左): 在远端中柱细胞特异表达的CLE40分泌的小肽通过与在远端中柱细胞起始细胞(根尖干细胞)特异表达的受体激酶ACR4结合传递信号, 抑制静止中心标志基因WOX5的表达; 拟南芥静止中心调控(右): ROW1可直接与WOX5启动子结合抑制其表达, WOX5通过直接与细胞分裂蛋白CYCDs的启动子结合, 抑制细胞分裂蛋白活性, 来维持静止中心较低的分裂速率; (F) 水稻FCP2在根尖表达, FCP2分泌的小肽可能通过一种未知的受体传递信号, 抑制静止中心标志基因OsWOX9的表达
Figure 1 Regulatory networks of WOX family in shoot and root apical meristems of Arabidopsis and rice (modified from Gaillochet et al., 2015) (A) The structure of shoot apical meristem (SAM); (B) The WUSCHEL-CLAVATA feedback loop in Arabidopsis: WUS expresses in the organizing center, and WUS protein moves symplastically through plasmodesmata to the overlying domain to determinate stem cell fate, WUS directly promotes the expression of CLV3, which encodes a small secreted peptide that signals through intracellular signal transduction pathways, inducing signaling transduction of CLV1, BAM1 and RPK2 to restrict the expression of WUS; (C) FCP1 and OsWOX4 express in the SAM, and FCP1 represses the expression of OsWOX4 in an unknown pathway; (D) The structure of root apical meristem (RAM); (E) CLE40 expresses in the columella, then this small secreted peptide combines with ACR4 in the columella to transfer signal and repress the expression of WOX5, which expresses specifically in the quiescent center (QC) (left); and involves in the maintenance of the QC (right): in QC, ROW1 specifically binds in the promoter of WOX5 to repress its transcription, and WOX5 suppresses QC divisions that are otherwise promoted by CYCDs, in part by represses CYCD’s expression directly; (F) OsWOX9 expresses specifically in the QC and its expression is specially repressed by FCP2.
图2 WOX4参与拟南芥维管分生组织的调控(改自Hirakawa et al., 2010)
Figure 2 WOX4 is involved in the regulation of vascular meristem in Arabidopsis (modified from Hirakawa et al., 2010)
基因 | 编号 | 分支 | 在已发表文章中的命名(别名) | 其它物种中的 同源基因 | 表达区域 | 主要功能 | 参考文献 |
---|---|---|---|---|---|---|---|
OsWOX1 | LOC_Os04g56780 | 进化支 | MOC3 | 拟南芥WUS | 主要在地上部分、愈伤组织和幼穗 中表达, 在成熟 器官中表达较低 | 对腋芽的形成是必需的, 并与细胞分裂素有着复杂的关系 | Lu et al., 2015 |
OsWOX2 | LOC_Os05g02730 | 进化支 | WOX3; GLR1; OsWOX3B; NUDA/GL | 玉米NS1 (narrow sheath 1)、NS2和拟南芥PRS (PRESSED FLOWER) | 叶原基、幼叶和 生殖器官 | 叶毛发育 | Dai et al., 2007; Li et al., 2012; Zhang et al., 2012 |
OsWOX3 | LOC_Os11g01130 LOC_Os12g01120 | 进化支 | NAL2/3; OsWOX3A | 在叶片和叶鞘的 大小维管束鞘(尤其是韧皮组织)、 茎基部以及根的 维管柱中表达 | 叶片的横轴生长和维管束形成, 小穗的内外稃形态建成以及分蘖和侧根发育及根毛形成 | Cho et al., 2013; Yoo et al., 2013 | |
OsWOX4 | LOC_Os04g55590 | 进化支 | 无 | 拟南芥WOX4 | 在地上各分生组 织中广泛表达 | 营养生长阶段茎尖分生组织的维持; 与细胞分裂素作用相关 | Ohmori et al., 2013 |
OsWOX5 | LOC_Os01g62310 | 进化支 | 无 | 未知 | 未知 | 未知 | |
OsWOX6 | LOC_Os03g20910 | 中间支 | 无 | 未知 | 未知 | 未知 | |
OsWOX7 | LOC_Os01g47710 | 中间支 | DWT1 | 未知 | 在幼穗中强表达 | 参与分蘖发育; 与穂型相关 | Wang et al., 2013, 2014 |
OsWOX8 | LOC_Os01g60270 | 古老支 | 无 | 未知 | 未知 | 未知 | |
OsWOX9 | LOC_Os01g63510 | 进化支 | QHB | 拟南芥WOX5 | 特异地在根尖组 织中心表达 | 根尖分生组织的维持 | Kamiya et al., 2003; Chu et al., 2013 |
OsWOX9C | LOC_Os05g48990 | 中间支 | 无 | 未知 | 未知 | 未知 | |
OsWOX10 | LOC_Os08g14400 | 中间支 | 无 | 未知 | 未知 | 未知 | |
OsWOX11 | LOC_Os07g48560 | 中间支 | 无 | 未知 | 冠根萌发时表达; 成熟根的分裂区 | 在冠根发育时期直接抑制RR2, 以调节细胞增殖 | Zhao et al., 2009 |
表1 水稻中WOX家族基因的功能总结
Table 1 The summary of WOX gene family function in rice
基因 | 编号 | 分支 | 在已发表文章中的命名(别名) | 其它物种中的 同源基因 | 表达区域 | 主要功能 | 参考文献 |
---|---|---|---|---|---|---|---|
OsWOX1 | LOC_Os04g56780 | 进化支 | MOC3 | 拟南芥WUS | 主要在地上部分、愈伤组织和幼穗 中表达, 在成熟 器官中表达较低 | 对腋芽的形成是必需的, 并与细胞分裂素有着复杂的关系 | Lu et al., 2015 |
OsWOX2 | LOC_Os05g02730 | 进化支 | WOX3; GLR1; OsWOX3B; NUDA/GL | 玉米NS1 (narrow sheath 1)、NS2和拟南芥PRS (PRESSED FLOWER) | 叶原基、幼叶和 生殖器官 | 叶毛发育 | Dai et al., 2007; Li et al., 2012; Zhang et al., 2012 |
OsWOX3 | LOC_Os11g01130 LOC_Os12g01120 | 进化支 | NAL2/3; OsWOX3A | 在叶片和叶鞘的 大小维管束鞘(尤其是韧皮组织)、 茎基部以及根的 维管柱中表达 | 叶片的横轴生长和维管束形成, 小穗的内外稃形态建成以及分蘖和侧根发育及根毛形成 | Cho et al., 2013; Yoo et al., 2013 | |
OsWOX4 | LOC_Os04g55590 | 进化支 | 无 | 拟南芥WOX4 | 在地上各分生组 织中广泛表达 | 营养生长阶段茎尖分生组织的维持; 与细胞分裂素作用相关 | Ohmori et al., 2013 |
OsWOX5 | LOC_Os01g62310 | 进化支 | 无 | 未知 | 未知 | 未知 | |
OsWOX6 | LOC_Os03g20910 | 中间支 | 无 | 未知 | 未知 | 未知 | |
OsWOX7 | LOC_Os01g47710 | 中间支 | DWT1 | 未知 | 在幼穗中强表达 | 参与分蘖发育; 与穂型相关 | Wang et al., 2013, 2014 |
OsWOX8 | LOC_Os01g60270 | 古老支 | 无 | 未知 | 未知 | 未知 | |
OsWOX9 | LOC_Os01g63510 | 进化支 | QHB | 拟南芥WOX5 | 特异地在根尖组 织中心表达 | 根尖分生组织的维持 | Kamiya et al., 2003; Chu et al., 2013 |
OsWOX9C | LOC_Os05g48990 | 中间支 | 无 | 未知 | 未知 | 未知 | |
OsWOX10 | LOC_Os08g14400 | 中间支 | 无 | 未知 | 未知 | 未知 | |
OsWOX11 | LOC_Os07g48560 | 中间支 | 无 | 未知 | 冠根萌发时表达; 成熟根的分裂区 | 在冠根发育时期直接抑制RR2, 以调节细胞增殖 | Zhao et al., 2009 |
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