摘要
研究证实过氧化氢(H_2O_2)、一氧化氮(NO)和一氧化碳(CO)均介导脱落酸(ABA)诱导的气孔关闭,也有资料显示壳梭孢素(fusicoccin,FC)有促进气孔开放的效应,但时止今日尚未清楚ABA诱导气孔关闭中CO与H_2O_2的关系,FC抑制ABA诱导的气孔关闭与保卫细胞H_2O_2、NO变化的关系也未见研究。本文以蚕豆为材料,借助表皮条试验和激光共聚焦扫描显微镜(LSCM)技术对上述问题进行了探索。所得结果主要如下:
1.ABA、CO合成底物亚铁血红素(Ht)、CO水溶液和H_2O_2均表现促进气孔关闭的效应,CO产生酶亚铁血红素加氧酶-1(HO-1)专一性抑制剂锌原卟啉(ZnPPIX)、H_2O_2清除剂抗坏血酸(Vc)、过氧化氢酶(CAT)和H_2O_2产生酶NADPH氧化酶抑制剂二苯基碘(DPI)均抑制ABA诱导气孔关闭,Ht诱导的气孔关闭也被Vc、CAT和DPI抑制,这些结果暗示ABA可能通过诱导保卫细胞CO合成增加H_2O_2产生,进而促进气孔关闭。内源H_2O_2检测结果表明,ABA和Ht确有诱导保卫细胞H_2O_2产生的作用,而且ABA促进保卫细胞H_2O_2产生的作用可被ZnPPIX明显抑制,此与前述表皮条试验的结果完全一致。
2.与H_2O_2清除剂Vc、CAT和H_2O_2产生酶NADPH氧化酶抑制剂DPI一样,FC抑制ABA诱导气孔关闭并降低保卫细胞内源H_2O_2水平,表明FC通过降低保卫细胞H_2O_2水平抑制ABA诱导气孔关闭;与H_2O_2清除剂Vc、CAT类似,FC不仅阻止外源H_2O_2诱导气孔关闭、降低保卫细胞H_2O_2荧光,而且促进ABA诱导下已关闭气孔重新开放、降低ABA诱导下已产生的内源H_2O_2,但H_2O_2产生酶NADPH氧化酶抑制剂DPI却无上述效应,这些结果说明FC可能通过清除而非抑制合成降低保卫细胞H_2O_2水平,进而抑制ABA诱导气孔关闭。
3.与NO专一性清除剂c-PTIO、NO合酶(NOS)抑制剂L-NAME一样,FC抑制ABA诱导气孔关闭并降低保卫细胞内源NO水平,表明FC通过降低保卫细胞NO水平抑制ABA诱导气孔关闭;与NO专一性清除剂c-PTIO类似,也能通过降低保卫细胞NO水平抑制ABA诱导的气孔关闭。与NO专一性清除剂c-PTIO相类似,FC不仅阻止外源NO供体硝普钠(SNP)诱导气孔关闭、降低保卫细胞NO荧光,而且促进ABA诱导下已关闭气孔重新开放、降低ABA诱导下已产生的内源NO,但NOS抑制剂L-NAME却无上述效应,这些结果说明FC可能通过清除而非抑制合成降低保卫细胞NO水平,进而抑制ABA诱导气孔关闭。
Previous studies showed that CO, H_2O_2 and NO are involve in ABA-induced stomatal closure, and stomatal opening is induced by fusicoccin (FC). However, the role of CO-mediated H_2O_2 in ABA-induced stomatal closure remains unclear. Also, the effect and action mechanism of FC on ABA-induced stomatal closure are unkown. In the present study, by means of the epidermal strip bioassay and the laser scanning confocal microscopy(LSCM), it was investigated that the relationship of CO and H_2O_2 in ABA-induced stomatal closure and the effects and action mechanism of FC on stomatal closure induced by ABA. The main results are as followed:
1. Stomatal closure were induced by ABA, CO synthesis substrate Hematin (Ht), CO aqueous solution and H_2O_2, ABA-induced stomatal closure was restrained by ZnPPIX (CO specific synthetic inhibitor), CAT(catalase), Vc(H_2O_2 scavenger) and DPI(an inhibitor of H_2O_2 generating enzyme NADPH oxidase), Ht-induced stomatal closure was also prevented by CAT, Vc and DPI. The results suggest that CO Synthesis may mediate ABA-induced H_2O_2 production and stomatal closure. The examination results of endogenous H_2O_2 showed that ABA and Ht induce increase of H_2O_2. However, ABA-induced the production of endogenous H_2O_2 was significantly prevented by ZnPPIX. These results were in accordance with the epidermal strip bioassay.
2. Like CAT. Vc and DPI, FC inhibited ABA-induced stomatal closure and reduced the level of endogenous H_2O_2 in guard cells, which indicated that FC inhibited ABA-induced stomatal closure by reducing endogenous H_2O_2 level. Like CAT and Vc, FC not only prevented stomatal closure induced by exogenous H_2O_2, reduced H_2O_2 fluorescence of guard cells treated by exogenous H_2O_2, but also promoted the closed stoma induced by ABA to reopen, abolished H_2O_2 that had been generated by ABA. However, DPI had no the effects mentioned above. Taken together, our results indicate that FC probably reduce the levels of H_2O_2 in guard cells by scavenging, not restraining H_2O_2 generation, and then prevent ABA-induced stomatal closure.
3. Like c-PTIO(NO scavenger) and L-NAME(NO synthase inhibitor), FC prevented ABA-induced stomatal closure and reduce the level of endogenous NO in guard cells, which indicated that FC prevented ABA-induced stomatal closure by reducing the level of endogenous NO. Like c-PTIO, FC not only prevented SNP-induced stomatal closure, reduced NO fluorescence of guard cells treated by SNP, but also promote the closed stoma induced by ABA to reopen, abolished NO that had been generated by ABA. However, L-NAME had no the effects mentioned above. Hence, it is concluded that FC reduce probably the levels of NO in guard cells via scavenging, not preventing NO generation, and then prevent ABA-induced stomatal closure.
引文
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