光信号与激素调控种子休眠和萌发研究进展

doi:10.11983/CBB19038

摘要/Abstract

摘要： 休眠是种子植物在长期进化过程中产生的适应性性状, 通过抑制种子在不适宜的环境中萌发进而保证植物能够在逆境中生存。此外, 休眠有助于种子的长距离运输和扩散, 因此休眠对种子延续和物种保存具有重要意义。种子由休眠向萌发的发育转变不仅关系到物种的繁衍, 而且对保证农业生产中作物的产量和品质也具有重要作用。种子的休眠和萌发受到内源激素和外源光信号的共同调控。其中, 外源光信号主要通过调控内源ABA和GA的生物合成及信号转导进而调控种子休眠和萌发。该文系统综述了外源光信号和内源激素调控种子休眠和萌发的作用通路以及两类信号通路之间的交互作用, 旨在为农业生产中利用光和激素调控种子休眠与萌发提供参考。

关键词: 种子休眠, 种子萌发, 光信号, 激素, 交互作用

Abstract: Plants have evolved to maintain the dormancy of freshly harvested seeds, which ensures that seeds do not germinate until environmental conditions are optimal. Therefore, dormancy helps seeds spread over long distances to ensure the survival of species. The transition from dormancy to germination is crucial to plant survival and for promoting yield and quality in agricultural production. Seed dormancy and germination are precisely regulated by diverse endogenous hormones and light signals. Light cues regulate seed dormancy and germination by affecting abscisic acid/gibberellic acid biosynthesis and signals. In this review, we summarize the key roles of the hormone pathway and light signal transduction pathways in regulating seed dormancy and germination. We also discuss the interactions (crosstalk) between phytohormone signals and light signals in seed dormancy and germination, in order to apply reference for regulating seed dormancy and germination by using light and hormones in agricultural production.

Key words: seed dormancy, seed germination, light signal, hormones, crosstalk

杨立文,刘双荣,林荣呈. 光信号与激素调控种子休眠和萌发研究进展. 植物学报, 2019, 54(5): 569-581.

Liwen Yang,Shuangrong Liu,Rongcheng Lin. Advances in Light and Hormones in Regulating Seed Dormancy and Germination. Chinese Bulletin of Botany, 2019, 54(5): 569-581.

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链接本文: https://www.chinbullbotany.com/CN/10.11983/CBB19038

https://www.chinbullbotany.com/CN/Y2019/V54/I5/569

图/表 1

图1 光信号通过调控内源脱落酸(ABA)和赤霉素(GA)的生物合成及信号转导调控种子休眠与萌发 (A) 光信号通过调控ABA和GA通路调控种子萌发。PHYB能够介导红光促进PIF1发生泛素化降解, 从而促进种子萌发。PIF1能够通过直接激活DAG1和SOM的转录进而间接调控GA生物合成相关基因的表达, 或者直接诱导DELLA蛋白编码基因RGA和GAI的转录, 最终抑制种子萌发。同样地, PIF1也能通过调控ABA的生物合成和信号转导调控种子萌发。PIF1通过依赖于SOM的途径促进ABA生物合成, 进而抑制种子萌发; 抑或直接诱导ABI3和ABI5的转录进而促进ABA信号转导, 抑制种子萌发。除PIF1之外, PHYB还能调控RVE1的转录间接促进GA的生物合成, 最终促进种子萌发。SPT和CSN蛋白复合体通过依赖于ABI5途径调控种子萌发。SPT通过抑制ABI5的转录抑制ABA信号转导, 促进种子萌发。CSN1通过促进RGL2的泛素化降解进而抑制ABI5的蛋白稳定性, 最终促进种子萌发; 而CSN5a能够直接抑制ABI5蛋白的积累进而促进种子萌发。JAZ3通过抑制ABI5对ABA响应基因EM1的转录激活功能进而促进种子萌发。(B) 光信号通过调控ABA和GA通路调控种子休眠。PHYB能够介导红光抑制RVE1转录, 进而促进下游GA3ox2的转录, 最终抑制种子休眠。在不同生态型拟南芥背景下, SPT调控种子休眠的功能不同。其中, 在Col背景下, SPT通过促进RGL3和ABI5的转录进而促进种子休眠(绿色标识线); 在Ler背景下, SPT通过抑制RGA和ABI4的转录进而抑制种子休眠(红色标识线)。此外, PIF6也参与调控种子休眠。

Figure 1 Light signal regulates seed germination and dormancy via endogenous abscisic acid (ABA) and gibberellin (GA) biosynthesis pathway (A) Light signal regulates seed germination via ABA and GA pathway. PHYB regulates seed germination through promoting the degradation of PIF1 protein. The accumulation of PIF1 in nucleus activates the transcription of DAG1 and SOM, which indirectly regulates the expression of GA biosynthesis gene or directly induces RGA and GAI (DELLA protein encoding genes) transcription and leads to repressing seed germination. Similarly, PIF1 stimulates ABA biosynthesis and ABA signaling pathway to suppress seed germination. PIF1 induces ABA biosynthesis via SOM-dependent pathway to repress seed germination; or it induces the transcription of ABI3 and ABI5 in order to stimulate ABA signaling. Expect for PIF1, PHYB also promotes seed germination via inhibiting RVE1 transcription which indirectly promote GA biosynthesis. SPT and CSN complex could regulate seed germination in an ABI5-dependent manner. SPT suppresses ABI5 transcript to destroy ABA pathway. CSN1 stimulates RGL2 degradation to inhibit ABI5 activity, while CSN5a directly decreases the accumulation of ABI5 in order to provoke seed germination. JAZ3 promotes seed germination by repressing the transcriptional activity of ABI5 which activates ABA-responsing gene EM1 expression. (B) Light signal controls seed dormancy via ABA and GA pathway. PHYB mediates red light to repress the transcription level of GA3ox2, inhibiting seed dormancy. SPT plays different roles in regulating seed dormancy under Col and Ler background of Arabidopsis. SPT promotes seed dormancy through activating the expression of RGL3 and ABI5 under Col background (green line), however, SPT suppresses seed dormancy via inhibiting RGA and ABI4 transcript under Ler background (red line). In addition, PIF6 is also involved in controlling seed dormancy.

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