CO通过NO信号上调抗氧化防护和维持离子稳态来提高小麦幼苗耐盐性
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摘要
CO在哺乳动物研究中被认为是一种调控一系列生理过程的重要内源气体信号分子,并且这些调控过程往往是通过NO信号来介导,但CO在植物逆境响应方面研究还比较少。
本研究发现150 mM NaCl胁迫以时间进程方式诱导了小麦幼苗根部CO合成酶-血红素加氧酶(HO EC 1.14.99.3)活性上升并导致内源CO含量增加。继而用外源CO处理来模拟内源CO的变化,我们发现外源50%饱和溶液与150 mM NaCl共处理(S+50%CO)不仅能提高小麦幼苗根部CO含量,还能明显缓解由胁迫导致其生长减缓和相对含水量的下降。一氧化氮供体硝普钠(SNP,S+0.1SNP)和脂多糖乙二胺四乙酸二钠/一氧化氮(DETA/NO,S+DETA/NO)处理能得到与上述CO处理相类似的生物学表型,而SNP降解产物NO_2~-/NO_3~-和SNP类似物KFe_3(CN)_6KFe_4(CN)_6与盐共处理(S+NO_2~-/NO_3~-;S+KFe_3(CN)_6/KFe_4(CN)_6)则更加抑制小麦幼苗的生长。此外,CO与SNP还能上调小麦幼苗根部质膜ATP酶(PM H~+-ATPase)、液泡膜ATP酶(V H~+-ATPase)和液泡膜焦磷酸酶(V H~+-PPase)等与维持离子稳态相关的酶类及抗坏血酸过氧化物酶(APX)、谷胱甘肽还原酶(GR)、超氧化物歧化酶(SOD)、单脱氢抗坏血酸还原酶(MDHAR)和双脱氢抗坏血酸还原酶(DHAR)等抗氧化防护酶的活性并增加相关酶编码基因的转录本,从而提高幼苗根部的K/Na和缓解盐胁迫造成的氧化损伤。进一步的研究发现,CO的这种生物学功能很可能与NO有关,因为结合采用CO/NO清除剂血红蛋白处理(Hb,S+50%CO+Hb)、NO清除剂亚甲基蓝处理(MB,S+50%CO+MB;S+50%CO+0.1SNP+MB)或动物一氧化氮合酶(NOS)抑制剂N~G-硝基-L-精氨酸甲酯处理(L-NAME,S+50%CO+NAME)都不同程度地逆转了CO的各种缓解效应。更重要的是,CO能通过模拟SNP的作用来诱导小麦幼苗根尖大量释放NO,而组合L-NAME(S+50%CO+NAME)和2-(4-羧基苯)-4,4,5,5-四甲基咪唑-1-氧-3-氧化物(cPTIO,S+50%CO+cPTIO)处理则猝灭CO诱导的NO产生。
上述研究表明,CO通过NO信号上调小麦幼苗根部抗氧化防护和维持离子稳态来提高耐盐性。
In mammalian cells, CO was proven as an endogenous gas molecular in regulating a serial of physiological processes, which were usually mediated by NO. However, little information has been investigated in plant stress responses.
In present study, we evaluated that salt stress induced an increase in endogenous carbon monoxide (CO) production and the activity of the CO synthetic enzyme heme oxygenase in wheat seedling roots. In addition, a 50% CO aqueous solution, applied daily, not only resulted in the enhancement of CO release, but led to a significant reversal in dry weight and water loss caused by 150 mM NaCl treatment, which was mimicked by the application of two nitric oxide (NO) donors sodium nitroprusside (SNP) and diethylenetriamine nitric oxide adduct (DETA/NO). While, the degradation product of SNP (NO_2~-/NO_3~-) and SNP analogue KFe_3(CN)_6/KFe_4(CN)_6 produced negative effects. Further analyses showed that CO as well as SNP, apparently up-regulated H~+-pump and antioxidant enzymes activities or related transcripts, thus resulting in the increase of K/Na ratio and the alleviation of oxidative damage in the root tissue. The above ameliorating effects of CO were specific and related to NO, because the CO/NO scavenger hemoglobin (Hb), NO scavenger methylene blue (MB), or nitric oxide synthase (NOS) inhibitor N~G-nitro-L-arginine methyl ester hydrochloride (L-NAME) differentially blocked the above effects. Additionally, CO was able to mimic the effect of SNP by strongly increasing NO release in the root tips after daily treatment, whereas the CO-induced NO signal was quenched by the addition of L-NAME or the specific scavenger of NO 4,4,5,5-tetramethylimidazoline-1-oxyl-3-oxide potassium salt (cPTIO).
Taken together, above results suggest that CO might confer an increased tolerance to salinity stress by maintaining ion homeostasis and enhancing antioxidant system parameters in wheat seedling roots, both of which were partially mediated by NO signal.
本研究发现150 mM NaCl胁迫以时间进程方式诱导了小麦幼苗根部CO合成酶-血红素加氧酶(HO EC 1.14.99.3)活性上升并导致内源CO含量增加。继而用外源CO处理来模拟内源CO的变化,我们发现外源50%饱和溶液与150 mM NaCl共处理(S+50%CO)不仅能提高小麦幼苗根部CO含量,还能明显缓解由胁迫导致其生长减缓和相对含水量的下降。一氧化氮供体硝普钠(SNP,S+0.1SNP)和脂多糖乙二胺四乙酸二钠/一氧化氮(DETA/NO,S+DETA/NO)处理能得到与上述CO处理相类似的生物学表型,而SNP降解产物NO_2~-/NO_3~-和SNP类似物KFe_3(CN)_6KFe_4(CN)_6与盐共处理(S+NO_2~-/NO_3~-;S+KFe_3(CN)_6/KFe_4(CN)_6)则更加抑制小麦幼苗的生长。此外,CO与SNP还能上调小麦幼苗根部质膜ATP酶(PM H~+-ATPase)、液泡膜ATP酶(V H~+-ATPase)和液泡膜焦磷酸酶(V H~+-PPase)等与维持离子稳态相关的酶类及抗坏血酸过氧化物酶(APX)、谷胱甘肽还原酶(GR)、超氧化物歧化酶(SOD)、单脱氢抗坏血酸还原酶(MDHAR)和双脱氢抗坏血酸还原酶(DHAR)等抗氧化防护酶的活性并增加相关酶编码基因的转录本,从而提高幼苗根部的K/Na和缓解盐胁迫造成的氧化损伤。进一步的研究发现,CO的这种生物学功能很可能与NO有关,因为结合采用CO/NO清除剂血红蛋白处理(Hb,S+50%CO+Hb)、NO清除剂亚甲基蓝处理(MB,S+50%CO+MB;S+50%CO+0.1SNP+MB)或动物一氧化氮合酶(NOS)抑制剂N~G-硝基-L-精氨酸甲酯处理(L-NAME,S+50%CO+NAME)都不同程度地逆转了CO的各种缓解效应。更重要的是,CO能通过模拟SNP的作用来诱导小麦幼苗根尖大量释放NO,而组合L-NAME(S+50%CO+NAME)和2-(4-羧基苯)-4,4,5,5-四甲基咪唑-1-氧-3-氧化物(cPTIO,S+50%CO+cPTIO)处理则猝灭CO诱导的NO产生。
上述研究表明,CO通过NO信号上调小麦幼苗根部抗氧化防护和维持离子稳态来提高耐盐性。
In mammalian cells, CO was proven as an endogenous gas molecular in regulating a serial of physiological processes, which were usually mediated by NO. However, little information has been investigated in plant stress responses.
In present study, we evaluated that salt stress induced an increase in endogenous carbon monoxide (CO) production and the activity of the CO synthetic enzyme heme oxygenase in wheat seedling roots. In addition, a 50% CO aqueous solution, applied daily, not only resulted in the enhancement of CO release, but led to a significant reversal in dry weight and water loss caused by 150 mM NaCl treatment, which was mimicked by the application of two nitric oxide (NO) donors sodium nitroprusside (SNP) and diethylenetriamine nitric oxide adduct (DETA/NO). While, the degradation product of SNP (NO_2~-/NO_3~-) and SNP analogue KFe_3(CN)_6/KFe_4(CN)_6 produced negative effects. Further analyses showed that CO as well as SNP, apparently up-regulated H~+-pump and antioxidant enzymes activities or related transcripts, thus resulting in the increase of K/Na ratio and the alleviation of oxidative damage in the root tissue. The above ameliorating effects of CO were specific and related to NO, because the CO/NO scavenger hemoglobin (Hb), NO scavenger methylene blue (MB), or nitric oxide synthase (NOS) inhibitor N~G-nitro-L-arginine methyl ester hydrochloride (L-NAME) differentially blocked the above effects. Additionally, CO was able to mimic the effect of SNP by strongly increasing NO release in the root tips after daily treatment, whereas the CO-induced NO signal was quenched by the addition of L-NAME or the specific scavenger of NO 4,4,5,5-tetramethylimidazoline-1-oxyl-3-oxide potassium salt (cPTIO).
Taken together, above results suggest that CO might confer an increased tolerance to salinity stress by maintaining ion homeostasis and enhancing antioxidant system parameters in wheat seedling roots, both of which were partially mediated by NO signal.
引文
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