植物学报 ›› 2021, Vol. 56 ›› Issue (2): 218-231.DOI: 10.11983/CBB20141
宋松泉1,2,*(
), 刘军2,*(), 杨华3, 张文虎2, 张琪2, 高家东2
首页收稿日期:2020-08-03
接受日期:2021-01-05
出版日期:2021-03-01
发布日期:2021-03-17
通讯作者:
宋松泉,刘军
作者简介:liujun@gdaas.cn基金资助:
Songquan Song1,2,*(), Jun Liu2,*(), Hua Yang3, Wenhu Zhang2, Qi Zhang2, Jiadong Gao2
Received:2020-08-03
Accepted:2021-01-05
Online:2021-03-01
Published:2021-03-17
Contact:
Songquan Song,Jun Liu
摘要：
种子萌发是子代植株建立、生长和繁育的重要阶段, 在种子植物生命周期中起重要作用。种子休眠是在发育过程中形成的, 在生理成熟期达到峰值。种子休眠与萌发的植物激素调控可能是种子植物中一种高度保守的机制。细胞分裂素(CK)是植物体内的一种重要信号分子, 调控植物生长发育的许多方面。生物活性CK的水平由其生物合成、活化、失活、再活化和降解之间的平衡所调控, 种子的发育、休眠与萌发受生物活性CK的水平和信号转导途径调控。该文综述了CK的生物合成与代谢、信号转导以及对种子发育、休眠与萌发的调控作用, 提出了在本领域需要进一步研究的科学问题, 旨在为理解CK调控种子发育、休眠与萌发的分子机理提供参考。
宋松泉, 刘军, 杨华, 张文虎, 张琪, 高家东. 细胞分裂素调控种子发育、休眠与萌发的研究进展. 植物学报, 2021, 56(2): 218-231.
Songquan Song, Jun Liu, Hua Yang, Wenhu Zhang, Qi Zhang, Jiadong Gao. Research Progress in Seed Development, Dormancy and Germination Regulated by Cytokinin. Chinese Bulletin of Botany, 2021, 56(2): 218-231.
图1 细胞分裂素(CK)代谢和信号转导模型(改自Kieber and Schaller, 2018; Zubo and Schaller, 2020) Ade: 腺嘌呤; ADP: 腺苷二磷酸; AMP: 腺苷一磷酸; ATP: 腺苷三磷酸; CKX: CK氧化酶/脱氢酶; CYP735A: 细胞色素P450单加氧酶; cZOG: 顺式玉米素-O-葡糖基转移酶; DMAPP: 二甲基丙烯基二磷酸; HK: 组氨酸激酶; HP: 组氨酸磷酸转移蛋白; IPT: 异戊烯基转移酶; LOG: CK核苷5′-单磷酸磷酸核糖水解酶; SC: 侧链; Type-B RRs: 类型-B反应调控因子; Type-A RRs: 类型-A反应调控因子; UGT: UDP糖基转移酶
Figure 1 A model of cytokinin (CK) metabolism and signaling transduction (modified from Kieber and Schaller, 2018; Zubo and Schaller, 2020) Ade: Adenine; ADP: Adenosine diphosphate; AMP: Adenosine monophosphate; ATP: Adenosine triphosphate; CKX: CK oxidase/dehydrogenase; CYP735A: Cytochrome P450 monooxygenase; cZOG: cis-Zeatin-O-glucosyltransferase; DMAPP: Dimethylallyl diphosphate; HK: Histidin (His) kinase; HP: His phosphotransfer protein; IPT: Isopentenyltransferase; LOG: CK nucleoside 5′-monophosphate phosphoribohydrolase; SC: Sidechain; Type-B RRs: Type-B response regulators; Type-A RRs: Type-A response regulators; UGT: UDP glycosyltransferase
图2 细胞分裂素信号转导途径中的分子间相互作用(改自Lomin et al. 2018; Arkhipov et al. 2019) 细胞分裂素信号被膜定位的组氨酸(His)激酶受体感受, 通过His磷酸转移蛋白被传递到细胞核, 从而激活细胞核中的转录因子家族。CK: 细胞分裂素; ATP: 腺苷三磷酸; ADP: 腺苷二磷酸; P: 磷酸; D: 保守的天冬氨酸; H: 保守的His; DI: 感受器模块的二聚体界面结构域; PAS和PAS-like: 感受器模块的CHASE结构域的子结构域; TM1和TM2: 跨膜结构域1和2; HisKA (DHpD): His激酶A结构域(二聚合作用和His磷酸转移结构域); H-ATPase (CAD): 腺苷三磷酸酶结构域(催化与ATP结合结构域); REC-like: 类接收器结构域; REC: 接收器结构域; HPt: 含有His的磷酸转移蛋白(磷酸传递蛋白); RR-B: B型反应调控因子(转录因子)。蛋白-蛋白相互作用(PPI)用红色虚线表示。
Figure 2 Intermolecular interactions in the cytokinin signaling pathway (modified from Lomin et al. 2018; Arkhipov et al. 2019) Cytokinin signaling is perceived by membrane-localized Histidine (His) kinase receptors and is transduced to nucleus through a His phosphotransfer protein to activate a family of transcription factors in the nucleus. CK: Cytokinin; ATP: Adenosine triphosphate; ADP: Adenosine diphosphate; P: Phosphate; D: Conserved Aspartate; H: Conserved His; DI: Dimerization interface domain of the sensor module; PAS and PAS-like: Subdomains of the CHASE domain of the sensor module; TM1 and TM2: Transmembrane domain 1 and 2; HisKA (DHpD): His kinase A domain (dimerization and His phosphotransfer domain); H-ATPase (CAD): Adenosine triphosphatase domain (catalytic and ATP-binding domain); REC-like: Receiver-like domain; REC: Receiver domain; HPt: His-containing phosphotransfer protein (phosphotransmitter); RR-B: Type B response regulator (transcription factor). Protein-protein interactions (PPI) are indicated by a red dotted line.
图3 种子发育和萌发过程中与细胞分裂素(CK)代谢和信号转导组分有关的蛋白(酶)的变化模式图(改自Jameson et al. 2016; Tuan et al. 2019; Nguyen et al. 2020) IPT: 异戊烯基转移酶; LOG: CK核苷5′-单磷酸磷酸核糖水解酶; CKX: CK氧化酶/脱氢酶; HK: 组氨酸激酶; HP: 组氨酸磷酸转移蛋白; Type-B RRs: 类型-B反应调控因子; Type-A RRs: 类型-A反应调控因子
Figure 3 The model of the change of proteins (enzymes) involved in cytokinin (CK) metabolism and signaling components during seed development and germination (modified from Jameson et al. 2016; Tuan et al. 2019; Nguyen et al. 2020) IPT: Isopentenyltransferase; LOG: CK nucleoside 5′-monophosphate phosphoribohydrolase; CKX: CK oxidase/ dehydrogenase; HK: Histidin (His) kinase; HP: His phosphotransfer protein; Type-B RRs: Type-B response regulators; Type-A RRs: Type-A response regulators
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