植物学报 ›› 2022, Vol. 57 ›› Issue (1): 1-11.DOI: 10.11983/CBB21183
• 特邀综述 • 下一篇
收稿日期:
2021-10-25
接受日期:
2022-01-09
出版日期:
2022-01-01
发布日期:
2022-01-17
通讯作者:
熊延
作者简介:
* E-mail: yanxiong@fafu.edu.cn†共同第一作者。
基金资助:
Yanyan Meng1,2†, Nan Zhang1,2†, Yan Xiong1,2,*()
Received:
2021-10-25
Accepted:
2022-01-09
Online:
2022-01-01
Published:
2022-01-17
Contact:
Yan Xiong
About author:
First author contact:†These authors contributed equally to this paper.
摘要:
雷帕霉素靶蛋白(TOR)是真核生物中高度保守的丝氨酸/苏氨酸蛋白激酶, 能整合营养、能量、生长因子及环境信号, 协调细胞增殖、生长和代谢等过程, 是真核生物生长发育的核心调控因子。近年来, 随着相关研究系统的建立, 植物TOR的功能和机制研究取得了众多突破, 发现其进化上保守的生物学功能及植物中特有的信号通路。该文概述了TOR蛋白复合体的构成, 以及植物TOR响应糖、营养元素(氮、磷和硫)、激素及逆境胁迫信号来调控下游基因转录、蛋白翻译、代谢、细胞自噬和胁迫应答等生物学过程的分子机制, 并提出了植物TOR领域一些亟待解决的科学问题, 以期为全面揭示植物TOR的生物学功能提供参考。
孟彦彦, 张楠, 熊延. 植物TOR激酶响应上游信号的研究进展. 植物学报, 2022, 57(1): 1-11.
Yanyan Meng, Nan Zhang, Yan Xiong. Novel Links in the Plant Target of Rapamycin Signaling Networks. Chinese Bulletin of Botany, 2022, 57(1): 1-11.
图1 动、植物中TOR蛋白的结构域和复合体组分 (A) 动、植物中TOR复合体1 (TORC1)。TORC1主要包含TOR、RAPTOR及LST8三个组分。动、植物中的TOR蛋白高度保守, 均由HEAT重复域、FAT、FRB、激酶域及FATC五个结构域组成。其中, HEAT重复域与RAPTOR结合; 激酶域与LST8结合; FRB结构域可被FKBP12介导与雷帕霉素结合。(B) 动物中TOR复合体2 (TORC2)。动物TORC2主要包含TOR、RICTOR、SIN1及LST8四个组分。其中RICTOR与SIN1互作并与HEAT重复域结合, LST8同样与激酶域结合。而尚未在植物中鉴定到RICTOR和SIN1的同源基因, 暗示TORC2在植物中可能不保守。
Figure 1 The domain structures and components of TOR complexes in plants and animals (A) TOR complex 1 (TORC1) in plants and animals. TORC1 mainly contains TOR, RAPTOR and LST8. TOR protein is highly conserved among plants and animals, composed of HEAT repeats, FAT, FRB, kinase and FATC domains. RAPTOR binds to the HEAT repeats, LST8 binds to the kinase domain, and FKBP12 mediates interact between the FRB domain and rapamycin. (B) TOR complex 2 (TORC2) in animals. TORC2 mainly contains TOR, RICTOR, SIN1 and LST8 in animal. RICTOR interacts with SIN1 and binds to HEAT repeats, and LST8 binds to the kinase domain. Either RICTOR or SIN1 has no homologous genes in plants, which indicates that TORC2 is not conserved.
图2 植物TOR复合体1的上游调控因子 植物TOR复合体1感知多重上游信号, 并可能以二聚体的形式发挥作用。其中糖-能量信号、氮/磷/硫营养信号、生长素-ROP2信号以及赤霉素信号是TOR上游的激活信号; 脱落酸-SnRK2信号和冷胁迫信号是TOR上游的抑制信号; 不同病毒对TOR的作用不同, 既有促进也有抑制。FKBP12-雷帕霉素和ATP竞争性抑制剂Torin1/2、AZD8055、WYE132/354以及KU-63794均可特异且高效地抑制TOR激酶活性。红色箭头代表激活作用, 绿色T形线条代表抑制作用, 黑色线条代表兼具激活和抑制作用。
Figure 2 The upstream regulators of TOR complex 1 (TORC1) in plants TORC1 in plants senses multiple upstream signals, and probably functions as a dimerization form. Glucose-energy, nitrogen/phosphate/sulfur, auxin-ROP2, and GA signals are positive upstream regulators of TOR; ABA-SnRK2 and cold stress are negative upstream regulators of TOR; viruses play both positive and negative roles on TOR activity. FKBP12- rapamycin and ATP competitive inhibitors Torin1/2, AZD8055, WYE132/354 and KU-63794 inhibit plant TOR kinase activity specifically and efficiently. The red arrows represent activation, the green T shape lines mean repression, and the black line represents both activation and repression exist.
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