干旱胁迫下植物根源化学信号研究进展
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国家自然科学基金项目(31070372和30970447);教育部新世纪优秀人才支持计划项目(NCET-07-0396);兰州大学人才引进专项(582449);国家公益性行业(气象)科研专项子课题(GYHY200806021-06和GYHY201106029)


Current progress in eco-physiology of root-sourced chemical signal in plant under drought stress
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    摘要:

    土壤干旱胁迫诱导植物根系产生根源化学信号,经运输系统长距离传输到地上部分,降低气孔导度,抑制蒸腾作用,从而提高植物的水分利用效率。根源化学信号包括脱落酸(ABA)、细胞分裂素(CTK)、生长素、木质部pH值和钙离子(Ca2+)等,其中以ABA为主的植物根源信号通路研究得最为广泛和深入。总结了几种主要的化学根源信号物质的基本性质、主要功能和调节机制,重点对这些信号参与气孔行为、差别基因表达和生长发育方面的研究进展进行了综述。由于干旱条件下植物根源信号反应涉及到从分子到群体的一系列复杂过程,各种信号的生理效应呈现交互作用、耦合发生的特点,今后的热点领域将集中在研究交互网络中合成的的关键物质和揭示这些物质在分子及生理水平上的作用机理上。根源化学信号研究正朝向"以分子和生理研究为基础、不同尺度的结构和功能耦合"的方向发展。

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    Since Blackman and Davies found root-sourced chemical signals formed when soil was drying, many subsequent experiments have elucidated how root-shoot communication might operate. Chemical root-sourced signals are transmitted long-distance to aboveground parts via the transpiration stream, lowering the stomatal conductance,and inhibiting the transpiration rate so as to improve the water use efficiency of plants. It has been shown that the chemical signals act as an "early-warning" response to soil drying in plants. These root-sourced chemical signals include abscisic acid (ABA), cytokinins (CTK), auxin, xylem pH, Ca2+, of which the ABA-based root signal pathway has been the most extensively and thoroughly studied in plants. This paper summarizes the basic properties, main functions and regulatory mechanisms of several major root-sourced chemical agents. Most content is focused on current research progress on stomatal behavior regulation, differential gene expression, and plant growth and development processes under the operation of these signals. Over the last two decades, most research regarding the eco-physiology of chemical root-sourced signals has been accumulated on the crops such as wheat, maize and legumes. According to Xiong and Fan, the soil water content interval (SWCI) from "switching on" to "turning off" root signals was wider in modern wheat varieties than in in older varieties. There is a significant correlation between SWCI and other parameters such as lethal leaf water potential, maintenance rate of grain yield and water use efficiency. The results support the influence of root signals on regulatory mechanisms of reactive oxygen species, osmotica and antioxidant enzyme system in wheat and legume crops. On the other hand, the molecular mechanism of ABA regulating plant drought adaptation as a major chemical root-sourced signal is one of the hottest issues in this field. Important progress was achieved by Miyazonol et al. who showed that the PYR/PYL/RCAR family of START proteins received ABA to inhibit the phosphatase activity of the group-A protein phosphatases 2C (PP2Cs), which acted as major negative regulators in ABA signalling. The crystal structures of the ABA receptor PYL1 were bound with (+)-ABA, and produced a critical complex by the further binding of (+)-ABA-bound PYL1 with the PP2C protein ABI1. This result provided a breakthrough understanding on the structural basis of the mechanism of (+)-ABA-dependent inhibition of ABI1 by PYL1 in ABA signalling. In addition, other types of chemical root-sourced signals have been found to act in a different regulatory role. As a whole, the physiological effects of the various signals mutually interact and are involved with a range of complex processes from the molecular to the population level. Future research will investigate the key substances produced from the network of crosstalk and explore the regulatory mechanism of these substances at the molecular and physiological level. The eco-physiology for root-sourced chemical signals is following a trend of "revealing the coupling structural and functional relationship at different scales under the basis of molecular and physiological mechanisms".

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李冀南,李朴芳,孔海燕,熊俊兰,王绍明,熊友才.干旱胁迫下植物根源化学信号研究进展.生态学报,2011,31(9):2610~2620

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