生长素与植物根尖干细胞巢的维持

doi:10.11983/CBB17130

摘要/Abstract

摘要： 干细胞巢的维持与后代细胞的分化是多细胞高等生物个体发育的基础。生长素对植物茎尖和根尖分生组织的形态建成, 尤其是对位于植物这2个末端的分生组织中心的干细胞巢的活性维持起着至关重要的作用。该文综述了近几年在植物根尖干细胞发育领域的研究进展, 主要阐释了PLT蛋白途径、SCR-SHR蛋白途径以及环境因子多信号调控模块维持植物根尖分生组织中干细胞巢稳定的机制, 揭示了生长素可以通过就近合成、极性运输以及信号转导3种方式参与这些信号模块的调控, 从而维持生长素在根尖静止中心细胞附近干细胞巢的浓度梯度, 精确地平衡植物干细胞巢中细胞的增殖与分化。

关键词: 根尖分生组织, 干细胞巢, 生长素, 增殖与分化

Abstract: The coordination between the maintenance of stem-cell niches and differentiation of their progeny sets the developmental basis of higher multiple-cellular organisms to ensure coherent organ growth. The plant hormone auxin plays important roles in the morphological establishment of shoot and root apical meristem mediated by the maintenance of shoot and root stem-cell niches. Here we review multiple signaling modules discovered in recent years, including the PLETHORA pathway, SCARECROW-SHORT ROOT pathway and environmental factors, for their mechanisms in maintaining root stem cell niches. Auxin participates in these mechanisms by regulating local synthesis, polar transport and signal transduction. By these means, auxin concentration gradient can be maintained accurately in stem-cell niches around the root apical quiescent center, for balancing the proliferation and differentiation of stem-cell niches.

Key words: root apical meristem, stem cell niche, auxin, proliferation and differentiation

张倩倩, 郑童, 予茜, 葛磊. 生长素与植物根尖干细胞巢的维持. 植物学报, 2018, 53(1): 126-138.

Qianqian Zhang, Tong Zheng, Qian Yu, Lei Ge. Auxin and the Maintenance of Root Stem Cell Niches in Plants. Chinese Bulletin of Botany, 2018, 53(1): 126-138.

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链接本文: https://www.chinbullbotany.com/CN/10.11983/CBB17130

https://www.chinbullbotany.com/CN/Y2018/V53/I1/126

图/表 2

图1 拟南芥根尖及干细胞巢结构组成根尖分生组织(RAM)放射状细胞层排列方式, 从内到外分别是中柱(STELE)、内皮层(EN)、皮层(CO)、表皮(EP)、侧根冠(LRC)、静止中心(QC)、小柱细胞(Col)、小柱干细胞(CSC)、皮层/内皮层起始子细胞(CEI/CEID)、表皮/侧根冠(EP/LRC)、表皮/侧根冠起始细胞和中柱起始细胞。白色加粗轮廓为干细胞巢区, 分别为静止中心细胞(绿色)、小柱起始细胞(淡蓝色)、皮层/内皮层起始细胞(淡紫色)、表皮和侧根冠起始细胞(玫红色)以及中柱起始细胞(暗绿色)。

Figure 1 The root apical and the stem cell niche structure of Arabidopsis thalianaRadial pattern of cell layers in root apical meristem (RAM) from central to outsides: stele (STELE), endodermis (EN), cortex (CO), epidermis (EP), lateral root cap (LRC), quiescent center (QC), columella (Col), columella stem cell (CSC), cortex/endodermal initial/daughter cell (CEI/CEID), epidermis/later root cap (EP/LRC), EP/LRC initiation cell and stele initiation cells. Thick white line surrounds root stem cells, QC (green), CSC (light blue), CEI/CEID (light purple), EP/LRC initiation cell (rose red) and stele initiation cell (dark green).

图2 根尖干细胞巢维持的调控网络模式图根尖干细胞巢维持途径主要有2条: PLT途径和SHR-SCR途径。PLTs (红色三角的密度)蛋白梯度和生长素的浓度梯度(紫色圆圈的密度)维持干细胞巢区的活性; BIRDs蛋白参与限制SHR转移到皮层, 从而与SCR蛋白一起作用界定内皮层与皮层细胞的界限。WOX5既可以维持QC的稳定性, 也可在CSC细胞抑制CDF4活性, 进而维持干细胞活性。CLE40通过与ACR4结合, 以及ARF10和ARF16的表达, 也可将WOX5的表达限制在QC内。橘黄色代表静止中心细胞, 浅黄色代表皮层细胞, 淡红色代表内皮层细胞; 黑色箭头表示促进作用; ⊥表示抑制作用;红色虚线箭头表示蛋白在细胞间移动。

Figure 2 The regulation network for maintenance of the root stem cell nicheThe two major pathways for root stem cell niche maintenance: PLT pathway and SHR-SCR pathway. PLTs protein gradient (density of red triangles) and auxin concentration gradient (density of purple circles) maintain the activity of stem cell niche; BIRDs proteins are involved in repressing the transfer of SHR from endodermis to cortex cells, and work with SCR protein to define the boundary between endodermis and cortex. WOX5 not only maintains the stability of QC, but also repress the activity of CDF4 in CSC cells to maintain the acti- vity of stem cells. CLE40 restricts the expression of WOX5 in QC through binding to ACR4.And the expression of ARF10 and ARF16 has the same effect on WOX5. QC cells are co- lored in orange, cortex cells are colored in light yellow, and endodermis cells are colored in light red; black arrow represents promoted function; ⊥ represents inhibited function; red dotted arrow represents protein movement between cells.

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