生长素介导环境信号调控植物的生长发育

doi:10.11983/CBB17135

摘要/Abstract

摘要： 植物是一类营固着生活的自养型生物, 如何更好地适应周围环境对植物的生存至关重要。生长素是调控植物生长发育的重要激素之一。近年来的研究发现, 生长素不仅能够响应内在的发育信号, 而且能够介导各种环境信号, 参与植物生长发育和生长反应的调控。该文主要从光信号、温度信号、重力信号、营养元素和金属离子信号等方面重点阐述生长素如何介导上述各种不同的环境信号, 从而调控植物的生长发育。

关键词: 生长素, 环境信号, 生长发育

Abstract: Because plants are sessile and photo-autotrophic, they must adapt to the surrounding environment. Auxin is one of the most important plant hormones essential for plant growth and development. Recently, auxin was found to regulate plant growth by responding to endogenous developmental signals and by mediating various environmental cues. In this review, we focus on how auxin regulates plant growth by mediating various environmental cues such as light, temperature, gravity, nutrient element and metal ion signals.

Key words: auxin, environment cues, growth and development

刘广超, 丁兆军. 生长素介导环境信号调控植物的生长发育. 植物学报, 2018, 53(1): 17-26.

Guangchao Liu , Zhaojun Ding. Auxin Regulates Plant Growth and Development by Mediating Various Environmental Cues. Chinese Bulletin of Botany, 2018, 53(1): 17-26.

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链接本文: https://www.chinbullbotany.com/CN/10.11983/CBB17135

https://www.chinbullbotany.com/CN/Y2018/V53/I1/17

图/表 1

图1 生长素介导的环境信号调控植物生长发育的工作模型(A) 在重力响应过程中, 生长素的非对称分布主要是通过改变外运载体PIN的表达模式来实现的。这一调控过程一方面依赖于PID/WAG对PIN蛋白的磷酸化, 另一方面受转录因子FLP/MYB88直接的转录调节。(B) 在光信号通路过程中, 蓝光受体向光素PHOT通过抑制PID蛋白的活性, 降低对PIN3的磷酸化修饰水平进而参与生长素在下胚轴中的非对称分布。另一蓝光受体隐花色素CRY一方面和蓝光受体向光素PHOT及红光受体PHYB共同抑制ABCB19蛋白表达水平, 另一方面则通过转录因子PIF4直接结合在IAA19的启动子区并抑制下游生长素响应因子ARF7的转录活性参与下胚轴向光的弯曲生长。在植物庇荫反应中, 红光受体PHYB通过PIF家族转录因子直接调控生长素合成相关基因YUCCA的转录水平参与茎的向光弯曲; 而SAV3及SAV4同样参与了植物的庇荫反应。(C) 在响应温度信号通路中, 生长素受体TIR1通过直接与热激蛋白HSP90互作参与植物对高温环境的适应; 另一方面蓝光受体CRY1与PIF4直接互作调控YUC8的转录参与高温对下胚轴的伸长。此外, 高温诱导生长素甲基转移酶IAMT1的表达, 降低植物子房内生长素信号, 进而出现显著的雄性不育表型。(D) 在响应不同离子对植物根系发育的调控网络中, 生长素合成基因TAR2介导了低氮对植物侧根发生的调控, 而硝态氮受体基因NRT1.1通过抑制生长素的极性运输促进了高氮下植物侧根的起始。在低钾处理时, ARF2可被磷酸化, 进而解除对HAK5转录的抑制, 增强植物对钾离子的吸收能力。生长素合成基因TAA1和YUCCAs特异性地在转换区受到铝胁迫的异位诱导表达, 造成此区生长素的积累, 而ARF7可以直接转录调控细胞分裂素合成基因IPT的表达, 最终抑制主根的伸长。镉胁迫则通过维持生长素的内稳态, 进而抑制下游ROS水平, 参与对根伸长的调控。

Figure 1 An overview of plant growth and development in response to environment signal mediated by auxin (A) Nonsymmetrical of auxin in response to gravity due to altering expression pattern of auxin polar transporter PIN, which is regulated by phosphorylation of PIN through PID/WAG kinase or transcriptional regulation of FLP/MYB88. (B) In response to phototropism, blue light receptor PHOT reduced phosphorylation of PIN3 by inhibiting PID activity, thus mediated nonsymmetrical of auxin in hypocotyl. Another blue light receptor CRY inhibited ABCB919 expression together with PHOT and PHYB, and decreased ARF7 expression through binding to IAA19 promoter with PIF4. In shade condition, PHYB participated in phototropism of shoot through PIF to regulate YUCCA expression. SAV3 and SAV4 also mediate plant shade response. (C) In response to temperature signal, auxin receptor TIR1 can interact with HSP90 to participate in high temperature stress. While CRY1, together with PIF4, induced YUC8 expression to increase auxin level in hypocotyl under high temperature condition. In addition, high temperature could induce the expression of auxin transmethylase IAMT1, reduced auxin signal in ovary and led to male sterile. (D) Auxin synthesis gene TAR2 mediated lateral root development under low nitrogen condition, and nitrate nitrogen receptor NRT1.1 induced lateral root initiation by inhibiting auxin polar transport. ARF2 can be phosphorylated in low potassium, thus relieve the inhibition of HAK5 and enhanced the absorbing ability of phosphorus. TAA1 and YUCCA can be specific induced in the root TZ under aluminum stress, which caused excess auxin. While ARF7 directly regulated IPT expression, leading to the inhibition of root growth. Another metal ion cadmium maintained the auxin homeostasis to reduce downstream ROS level to regulate root growth.

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