外源油菜素内酯提高黄瓜幼苗低氧耐性的蛋白基础及光合生理
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摘要
无土栽培生产实践中管理不当及土壤水分过高容易导致植物根系供氧不足从而影响植株正常生长发育。黄瓜(Cucumis sativus L.)是世界性的重要蔬菜作物,也是我国设施栽培的主要蔬菜作物之一。然而,由于黄瓜根系细弱、易早衰、喜湿,但对根际氧气需求较高,根际低氧逆境下其生长势较难维持。油菜素内酯是能调节植物生长发育的重要激素之一,在促进植物生长、增强抗逆性等方面起着至关重要的作用。研究表明,外源油菜素内酯对低氧胁迫下黄瓜植株呼吸代谢和抗氧化系统相关酶活性具有一定的调节作用,但关于外源油菜素内酯对低氧胁迫下黄瓜植株蛋白及光合研究报道甚少。
本文以耐低氧性较弱的黄瓜品种‘中农6号’为试材,采用营养液栽培的方法,通过营养液中添加外源24-表油菜素内酯(EBR),研究了低氧胁迫下油菜素内酯对黄瓜幼苗相关蛋白表达、酶活性及光合作用的影响,探讨了油菜素内酯在黄瓜植株适应低氧胁迫中作用的蛋白基础及光合生理机制。主要研究结果如下：
1.外源油菜素内酯对低氧胁迫下黄瓜幼苗生长和可溶性蛋白表达的影响
低氧胁迫下,黄瓜幼苗生长明显受到抑制,根系可溶性蛋白含量降低、而叶片可溶性蛋白含量升高；施用EBR后缓解了低氧胁迫对黄瓜幼苗生长的抑制作用,其干重、根系总长、根尖数较单纯低氧处理显著增加,并且使黄瓜幼苗根系的可溶性蛋白含量提高,而使叶片可溶性蛋白含量降低。SDS-PAGE电泳分析表明,低氧胁迫与对照相比,根系中出现相对分子质量约为63.96×103的新蛋白条带,有11种蛋白表达量增强,施用EBR后根系中这些表达增强的蛋白条带均有所减弱,叶片中则有9种蛋白条带发生明显变化,这些蛋白的表达可能与外源EBR缓解低氧胁迫伤害作用密切相关,外源EBR可通过调节黄瓜幼苗中可溶性蛋白含量及其表达,从而缓解低氧胁迫对幼苗生长的抑制作用,减轻低氧胁迫对植株的伤害。
2.外源油菜素内酯对低氧胁迫下黄瓜幼苗根系呼吸酶、抗氧化酶及其相关基因表达的影响
低氧胁迫增强了黄瓜幼苗根系丙酮酸脱羧酶(PDC)、乙醇脱氢酶(ADH)、乳酸脱氢酶(LDH)同工酶的表达,低氧胁迫下施用外源EBR的3d时PDC、ADH同工酶的表达量分别比单纯低氧处理提高了18.8%、28.8%,而6、9d时PDC、ADH、LDH同工酶的表达又分别降低19.5%、25.6%、53.4%及26.4%、26.0%、28.4%。低氧胁迫下,异柠檬酸脱氢酶(IDH)、琥珀酸脱氢酶(SDH)、苹果酸脱氢酶(MDH)及苹果酸酶(ME)同工酶表达量明显的加强,低氧下施用外源EBR处理后6、9d明显增强了IDH、MDH同工酶的表达,而减弱了SDH同工酶的表达。低氧胁迫增强了黄瓜幼苗根系UDPG焦磷酸转化酶(UGPase)及醛缩酶(ALD)的表达,低氧胁迫3、6d UGPase的表达量分别提高了20.50%、30.80%,ALD的表达量分别提高了47.81%、56.17%,与单纯低氧处理相比,施用外源EBR处理3dALD的表达减少了11.20%,6d UGPase减少了22.15%。与对照相比,单纯低氧胁迫处理的黄瓜幼苗根系3、6、9d超氧化物歧化酶(SOD)、过氧化物酶(POD)、过氧化氢酶(CAT)同工酶表达量均低于对照,低氧胁迫下营养液增施EBR,其SOD同工酶的表达3、6d比单纯的低氧处理分别提高了5.22%、10.25%,POD同工酶的表达也分别增加了13.81%、9.46%,而CAT表达量无明显差异。油菜素内酯对短期根际低氧胁迫下黄瓜幼苗根系呼吸及抗氧化相关酶基因的表达表明,低氧处理3d黄瓜幼苗根系的ldh、pdc、idh、sdh、 mdh、ald、ugpase、pfk、hk、pod基因表达量上调,cat、sod、fk基因表达量下调,低氧下外源增施油菜素内酯ldh、idh、pfk基因表达量则进一步上调,而pdc、sdh、 mdh、cat、sod、ugpase基因表达量又稍有下降。可见,低氧胁迫下营养液添加EBR可调节黄瓜根系无氧呼吸、有氧呼吸酶和相关保护酶同工酶及基因的表达,从而增强黄瓜植株的低氧耐性,缓解低氧胁迫对黄瓜幼苗根系的伤害。
3.外源油菜素内酯对低氧胁迫下黄瓜幼苗根系细胞超微结构、ATPase活性及其线粒体抗氧化系统的影响
低氧胁迫下,黄瓜幼苗根系线粒体超氧阴离子(O-2)产生速率、过氧化氢(H202)和丙二醛(MDA)含量显著升高,根尖细胞受到伤害,ATPase活性及根系活力下降,处理3d其SOD、POD、CAT酶活性均受到诱导高于对照,随着处理时间延长至6d时又显著低于对照；而低氧胁迫下营养液施加EBR可在较长时间内维持较高的SOD、POD、CAT等抗氧化酶活性,降低O了产生速率、H2O2和MDA含量,提高了根系活力及其ATPase活性,根尖细胞受伤减轻。可见油菜素内酯对缓解低氧胁迫对黄瓜幼苗根系细胞超微结构、ATPase活性及其线粒体抗氧化系统均有良好的效应,可有效缓解低氧胁迫造成的伤害。4.外源油菜素内酯对低氧胁迫下黄瓜幼苗光合作用及其多胺、无机离子含量的影响低氧胁迫下,黄瓜植株的净光合速率(Pn)、表观量子效率(AQY)和羧化效率(CE)显著降低,黄瓜幼苗的PSII有效光化学效率(Fv'/Fm')、光化学淬灭系数(qP)、实际光化学效率(ΦPSII)、相对电子传递速率(rETR)显著下降,而非光化学淬灭系数(qN)和PS Ⅱ反应中心天线耗散能量(Drate)显著升高,细胞超微结构异常,说明低氧胁迫下黄瓜植株受到了光抑制。而低氧胁迫下,外源EBR可修复或缓解低氧胁迫对叶绿体和线粒体的伤害,提高植株的Pn、Fv'/Fm'、qP、rETR、ΦPSII以及光化学反应的能量(Prate),降低天线耗散的能量(Drate),说明外源EBR对缓解低氧胁迫对黄瓜幼苗光合特性和细胞超微结构的影响均有良好的效应,提高光能利用效率,缓解黄瓜植株的光抑制,维持较高的光合性能,从而减轻低氧胁迫对黄瓜幼苗的伤害。
低氧胁迫下,黄瓜幼苗叶片及根系内源腐胺(Put)、亚精胺(Spd)、精胺(Spm)、多胺(PAs)总含量和Put/PAs值显著提高,(Spd+Spm)/Put值显著降低,根系内源多胺变化幅度大于叶片,且幼苗叶片的K+、Ca2+、Mg2、Fe3+、Zn2+、Mn2+及根系的K+、Ca2+、Mg2+、Zn2+等离子含量均显著下降；外源EBR显著提高了低氧胁迫下黄瓜幼苗游离态Spm、结合态Spd、Spm及束缚态Put、Spd、Spm的含量,促进PAs进一步积累,降低Put/PAs比值,提高(Spd+Spm)/Put比值,显著增加黄瓜幼苗叶片Ca2+、Mg2+、Fe3+、Zn2+、Mn2+及根系Ca2+、Mg2+、Fe3+、Mn2+的含量。可见,外源EBR可调节黄瓜幼苗内源多胺含量及其形态,促进植株根系的养分吸收,从而缓解低氧胁迫对黄瓜幼苗的伤害。
Plants frequently experience limited oxygen availability, mainly due to mismanagement in production practice of soilless culture and soil flooding. The growth of plant was inhibited under hypoxic stress. Cucumber (Cucumis sativus L.) is one of the important vegetables in world, and major protected cultivation vegetable crops in China. Cucumber roots are weak and senilism, so they require higher rhizosphere oxygen cucumber is highly sensitive to hypoxia. Brassinolide (BR) has been recognized as a new kind of phytohormones and played an essential role in plant development and stress resistance. Previous researches suggested that BRs could regulate respiration, carbon metabolism and antioxidant system activity of cucumber seedlings under hypoxic stress. However, reports about the effects of brassinolide on photosynthesis and protein in cucumber seedling under hypoxic stress were limited.
Cucumber(Cucumis sativus L.)'Zhongnong6' was applied in our study, and exogenous24-epibrassinolide (EBR) was added in nutrient solution. The influence of EBR on protein expression, enzyme activities and photosynthesis in cucumber under hypoxic condition was studied, and the physiological mechanisms of cucumber responds for hypoxia were investigated. The results are as follows:
1. Effects of exogenous brassinolide on the growth and soluble protein expression in cucumber seedlings under hypoxic stress
Hypoxic stress significantly decreased the cucumber growth and soluble protein content in roots, but increased in leaves. EBR could alleviate the growth inhibition in cucumber seedling which induced by hypoxic stress with increasing longer roots, dry weight and root tips. Meanwhile, exogenous EBR inhibited the changes of growth and soluble protein content under hypoxic stress. Hypoxic stress induced a new protein (relative molecular weight63.96×103) in cucumber roots and increased the abundant of eleven proteins at least. However, Exogenous EBR could significantly decrease the expression of these proteins and markedly affected nine protein bands in leaves. Therefore, it was possible that these proteins have relations with exogenous EBR alleviating the effect of hypoxic stress in cucumber seedlings. Thus, we considered exogenous EBR could alleviat the inhibition of hypoxic stress on cucumber seedlings by changing the soluble protein content and expression.
2. Effects of exogenous brassinolide on protein expression of respiratory metabolism and antioxidant system in cucumber seedling roots under hypoxic stress
Pyruvate decarboxylase (PDC), alcohol dehydrogenase (ADH), lactate dehydrogenase (LDH) were induced by hypoxic stress. After application of EBR for three days, the expression of PDC and ADH isozymes increased by18.8%and28.8%compared to the hypoxic treatment. However, the abundance of PDC, ADH and LDH isozymes decresed by19.5%,25.6%,53.4%and26.4%,26.0%,28.4%on the6th and9th day, respectively. Hypoxia enhanced the expression of succinate dehydrogenase (SDH), isocitrate dehydrogenase (IDH), malate dehydrogenase (MDH), malic enzyme (ME) isozymes, and after application of EBR at6th and9th, IDH and MDH isozymes were significantly increased but SDH isozymes was decreased. UGPase、ALD were induced by hypoxic stress, the abundance of UGPase and ALD incresed by20.50%,30.80%and47.81%,56.17%on the3th and6th day, respectively. Compared to the hypoxic treatment, the abundance of ALD at3th and UGPase at6th after application of EBR was significantly increased by11.20%and22.15%, respectively. Compared to control, the SOD, POD and CAT isozymes of cucumber seedlings under hypoxic stress treatment were decreased at3th,6th and9th days. However, BR increased the expression of SOD isozymes by5.22%and10.25%on3th day and6th day compared to hypoxic treatment and POD isozymes were also increased13.81%and9.46%, respectively. But CAT isozymes were not changed significantly.
The effects of EBRs on the gene expression of respiration and antioxidant in cucumber seedlings under short-term hypoxic stress suggested that the gene expression of ldh, pdc, idh, sdh, mdh, ald, ugpase, pfk, hk, pod increased and cat, sod and fk decreased at3th day of treatment. After application the EBR, the expression of ldh、idh and pfk were further increased but the pdc, sdh, mdh, cat, sod and ugpase had a slight decreased. Our results suggested that hypoxic stress inhibited the growth of cucumber seedling roots and EBR might regulate the isozyme expressions of anaerobic respiration, aerobic respiration and antioxidant enzymes of cucumber roots thereby alleviated the suppression effects of hypoxic stress.
3. Effects of exogenous brassinolide on the mitochondria antioxidant system, ATPase activity and cellular ultrastructure in cucumber seedling roots under hypoxic stress
Under hypoxic stress, the superoxide anion (O2-) production rate, hydrogen peroxide (H2O2) content and malondialdehyde (MDA) contents were significantly enhanced, and root and ATPase activity were decreased resulting in cell damage. Compared with the control, activities of SOD, POD and CAT were induced and were higher at3d, and significantly lowered at6d by hypoxic stress. However, the treatment of EBR could keep higher activities of SOD, POD and CAT for a longer time, reduce the generation of O2, H2O2and MDA contents and significantly increased the root and ATPase activity, injured of root tip cell was released. The results indicated that exogenous EBR raised the activities of mitochondria antioxidant enzymes, decreased the O2-production rate, H2O2and MDA contents. Therefore, exogenous EBR might improve the mitochondria antioxidant system, ATPase activity and cellular ultrstructure of cucumber seedlings under hypoxic stress thereby alleviating the adverse effects of hypoxic stress.
4. Effects of exogenous brassinolide on photosynthesis, poly amines and inoroganic ions content in cucumber seedlings under hypoxic stress
When exposed to hypoxic stress, net photosynthetic rate (Pn), apparent quanta yield (AQY) and carboxylation efficiency (CE) were decreased significantly, and quantum efficency of open PSⅡ reaction center (Fv'/Fm'), photochemical quenching (qP), PSⅡ photochemisty rate (ΦPSⅡ), however the raletive electron transport rate(rETR)in leaves of cucumber seedlings were significantly decreased, but the allocation of antenna heat dissipation energy (Drate), and non-photochemical quenching(qN) were significantly increased, and the cellular ultrastructure was abnormal, which indicated that hypoxia caused photo-inhibition to cucumber seedlings. However, EBR could alleviate or release the damages in chloroplast and mitochondria under hypoxic stress, and enhanced Pn, Fv'/Fm', qP, rETR, PSⅡ and Prate. The results suggested that EBR could play a positive role in alleviating damage in mitochondria and chloroplast ultrastructure and photosynthetic characteristics, and make the leaf keep a higher photosynthetic performance, and alleviate the damage of hypoxic stress.
Hypoxic stress significantly increased the contents of polyamines including Put, Spd, Spm, total PAs and Put/PAs ratio in leaves and root, and decreased (Spd+Spm)/Put ratio. The change of root was greater than that of leaf. K+, Ca2+, Mg2+, Fe3+, Zn2+and Mn2+content in leaves and K+, Ca2+, Mg2+, Zn2+content in roots significantly decreased, too. However, Exogenous EBR increased the contents of the free Spm, conjugated Spd, Spm and bound Put, Spd, Spm. Meanwhile, EBR further increased the PAs content and (Spd+Spm)/Put ratio and reduced the Put/PAs ratio. Ca2+, Mg2+, Fe3+, Zn2+and Mn2+content in leaves and Ca2+, Mg2+, Fe3+, Mn2+content in roots significantly increased, too. Thus, EBR not only could regulate the formation and contents of polyamines but also promote the absorption of nutrients, make the leaf keep a higher photosynthetic performance, and alleviate the damage of hypoxic stress.
本文以耐低氧性较弱的黄瓜品种‘中农6号’为试材,采用营养液栽培的方法,通过营养液中添加外源24-表油菜素内酯(EBR),研究了低氧胁迫下油菜素内酯对黄瓜幼苗相关蛋白表达、酶活性及光合作用的影响,探讨了油菜素内酯在黄瓜植株适应低氧胁迫中作用的蛋白基础及光合生理机制。主要研究结果如下：
1.外源油菜素内酯对低氧胁迫下黄瓜幼苗生长和可溶性蛋白表达的影响
低氧胁迫下,黄瓜幼苗生长明显受到抑制,根系可溶性蛋白含量降低、而叶片可溶性蛋白含量升高；施用EBR后缓解了低氧胁迫对黄瓜幼苗生长的抑制作用,其干重、根系总长、根尖数较单纯低氧处理显著增加,并且使黄瓜幼苗根系的可溶性蛋白含量提高,而使叶片可溶性蛋白含量降低。SDS-PAGE电泳分析表明,低氧胁迫与对照相比,根系中出现相对分子质量约为63.96×103的新蛋白条带,有11种蛋白表达量增强,施用EBR后根系中这些表达增强的蛋白条带均有所减弱,叶片中则有9种蛋白条带发生明显变化,这些蛋白的表达可能与外源EBR缓解低氧胁迫伤害作用密切相关,外源EBR可通过调节黄瓜幼苗中可溶性蛋白含量及其表达,从而缓解低氧胁迫对幼苗生长的抑制作用,减轻低氧胁迫对植株的伤害。
2.外源油菜素内酯对低氧胁迫下黄瓜幼苗根系呼吸酶、抗氧化酶及其相关基因表达的影响
低氧胁迫增强了黄瓜幼苗根系丙酮酸脱羧酶(PDC)、乙醇脱氢酶(ADH)、乳酸脱氢酶(LDH)同工酶的表达,低氧胁迫下施用外源EBR的3d时PDC、ADH同工酶的表达量分别比单纯低氧处理提高了18.8%、28.8%,而6、9d时PDC、ADH、LDH同工酶的表达又分别降低19.5%、25.6%、53.4%及26.4%、26.0%、28.4%。低氧胁迫下,异柠檬酸脱氢酶(IDH)、琥珀酸脱氢酶(SDH)、苹果酸脱氢酶(MDH)及苹果酸酶(ME)同工酶表达量明显的加强,低氧下施用外源EBR处理后6、9d明显增强了IDH、MDH同工酶的表达,而减弱了SDH同工酶的表达。低氧胁迫增强了黄瓜幼苗根系UDPG焦磷酸转化酶(UGPase)及醛缩酶(ALD)的表达,低氧胁迫3、6d UGPase的表达量分别提高了20.50%、30.80%,ALD的表达量分别提高了47.81%、56.17%,与单纯低氧处理相比,施用外源EBR处理3dALD的表达减少了11.20%,6d UGPase减少了22.15%。与对照相比,单纯低氧胁迫处理的黄瓜幼苗根系3、6、9d超氧化物歧化酶(SOD)、过氧化物酶(POD)、过氧化氢酶(CAT)同工酶表达量均低于对照,低氧胁迫下营养液增施EBR,其SOD同工酶的表达3、6d比单纯的低氧处理分别提高了5.22%、10.25%,POD同工酶的表达也分别增加了13.81%、9.46%,而CAT表达量无明显差异。油菜素内酯对短期根际低氧胁迫下黄瓜幼苗根系呼吸及抗氧化相关酶基因的表达表明,低氧处理3d黄瓜幼苗根系的ldh、pdc、idh、sdh、 mdh、ald、ugpase、pfk、hk、pod基因表达量上调,cat、sod、fk基因表达量下调,低氧下外源增施油菜素内酯ldh、idh、pfk基因表达量则进一步上调,而pdc、sdh、 mdh、cat、sod、ugpase基因表达量又稍有下降。可见,低氧胁迫下营养液添加EBR可调节黄瓜根系无氧呼吸、有氧呼吸酶和相关保护酶同工酶及基因的表达,从而增强黄瓜植株的低氧耐性,缓解低氧胁迫对黄瓜幼苗根系的伤害。
3.外源油菜素内酯对低氧胁迫下黄瓜幼苗根系细胞超微结构、ATPase活性及其线粒体抗氧化系统的影响
低氧胁迫下,黄瓜幼苗根系线粒体超氧阴离子(O-2)产生速率、过氧化氢(H202)和丙二醛(MDA)含量显著升高,根尖细胞受到伤害,ATPase活性及根系活力下降,处理3d其SOD、POD、CAT酶活性均受到诱导高于对照,随着处理时间延长至6d时又显著低于对照；而低氧胁迫下营养液施加EBR可在较长时间内维持较高的SOD、POD、CAT等抗氧化酶活性,降低O了产生速率、H2O2和MDA含量,提高了根系活力及其ATPase活性,根尖细胞受伤减轻。可见油菜素内酯对缓解低氧胁迫对黄瓜幼苗根系细胞超微结构、ATPase活性及其线粒体抗氧化系统均有良好的效应,可有效缓解低氧胁迫造成的伤害。4.外源油菜素内酯对低氧胁迫下黄瓜幼苗光合作用及其多胺、无机离子含量的影响低氧胁迫下,黄瓜植株的净光合速率(Pn)、表观量子效率(AQY)和羧化效率(CE)显著降低,黄瓜幼苗的PSII有效光化学效率(Fv'/Fm')、光化学淬灭系数(qP)、实际光化学效率(ΦPSII)、相对电子传递速率(rETR)显著下降,而非光化学淬灭系数(qN)和PS Ⅱ反应中心天线耗散能量(Drate)显著升高,细胞超微结构异常,说明低氧胁迫下黄瓜植株受到了光抑制。而低氧胁迫下,外源EBR可修复或缓解低氧胁迫对叶绿体和线粒体的伤害,提高植株的Pn、Fv'/Fm'、qP、rETR、ΦPSII以及光化学反应的能量(Prate),降低天线耗散的能量(Drate),说明外源EBR对缓解低氧胁迫对黄瓜幼苗光合特性和细胞超微结构的影响均有良好的效应,提高光能利用效率,缓解黄瓜植株的光抑制,维持较高的光合性能,从而减轻低氧胁迫对黄瓜幼苗的伤害。
低氧胁迫下,黄瓜幼苗叶片及根系内源腐胺(Put)、亚精胺(Spd)、精胺(Spm)、多胺(PAs)总含量和Put/PAs值显著提高,(Spd+Spm)/Put值显著降低,根系内源多胺变化幅度大于叶片,且幼苗叶片的K+、Ca2+、Mg2、Fe3+、Zn2+、Mn2+及根系的K+、Ca2+、Mg2+、Zn2+等离子含量均显著下降；外源EBR显著提高了低氧胁迫下黄瓜幼苗游离态Spm、结合态Spd、Spm及束缚态Put、Spd、Spm的含量,促进PAs进一步积累,降低Put/PAs比值,提高(Spd+Spm)/Put比值,显著增加黄瓜幼苗叶片Ca2+、Mg2+、Fe3+、Zn2+、Mn2+及根系Ca2+、Mg2+、Fe3+、Mn2+的含量。可见,外源EBR可调节黄瓜幼苗内源多胺含量及其形态,促进植株根系的养分吸收,从而缓解低氧胁迫对黄瓜幼苗的伤害。
Plants frequently experience limited oxygen availability, mainly due to mismanagement in production practice of soilless culture and soil flooding. The growth of plant was inhibited under hypoxic stress. Cucumber (Cucumis sativus L.) is one of the important vegetables in world, and major protected cultivation vegetable crops in China. Cucumber roots are weak and senilism, so they require higher rhizosphere oxygen cucumber is highly sensitive to hypoxia. Brassinolide (BR) has been recognized as a new kind of phytohormones and played an essential role in plant development and stress resistance. Previous researches suggested that BRs could regulate respiration, carbon metabolism and antioxidant system activity of cucumber seedlings under hypoxic stress. However, reports about the effects of brassinolide on photosynthesis and protein in cucumber seedling under hypoxic stress were limited.
Cucumber(Cucumis sativus L.)'Zhongnong6' was applied in our study, and exogenous24-epibrassinolide (EBR) was added in nutrient solution. The influence of EBR on protein expression, enzyme activities and photosynthesis in cucumber under hypoxic condition was studied, and the physiological mechanisms of cucumber responds for hypoxia were investigated. The results are as follows:
1. Effects of exogenous brassinolide on the growth and soluble protein expression in cucumber seedlings under hypoxic stress
Hypoxic stress significantly decreased the cucumber growth and soluble protein content in roots, but increased in leaves. EBR could alleviate the growth inhibition in cucumber seedling which induced by hypoxic stress with increasing longer roots, dry weight and root tips. Meanwhile, exogenous EBR inhibited the changes of growth and soluble protein content under hypoxic stress. Hypoxic stress induced a new protein (relative molecular weight63.96×103) in cucumber roots and increased the abundant of eleven proteins at least. However, Exogenous EBR could significantly decrease the expression of these proteins and markedly affected nine protein bands in leaves. Therefore, it was possible that these proteins have relations with exogenous EBR alleviating the effect of hypoxic stress in cucumber seedlings. Thus, we considered exogenous EBR could alleviat the inhibition of hypoxic stress on cucumber seedlings by changing the soluble protein content and expression.
2. Effects of exogenous brassinolide on protein expression of respiratory metabolism and antioxidant system in cucumber seedling roots under hypoxic stress
Pyruvate decarboxylase (PDC), alcohol dehydrogenase (ADH), lactate dehydrogenase (LDH) were induced by hypoxic stress. After application of EBR for three days, the expression of PDC and ADH isozymes increased by18.8%and28.8%compared to the hypoxic treatment. However, the abundance of PDC, ADH and LDH isozymes decresed by19.5%,25.6%,53.4%and26.4%,26.0%,28.4%on the6th and9th day, respectively. Hypoxia enhanced the expression of succinate dehydrogenase (SDH), isocitrate dehydrogenase (IDH), malate dehydrogenase (MDH), malic enzyme (ME) isozymes, and after application of EBR at6th and9th, IDH and MDH isozymes were significantly increased but SDH isozymes was decreased. UGPase、ALD were induced by hypoxic stress, the abundance of UGPase and ALD incresed by20.50%,30.80%and47.81%,56.17%on the3th and6th day, respectively. Compared to the hypoxic treatment, the abundance of ALD at3th and UGPase at6th after application of EBR was significantly increased by11.20%and22.15%, respectively. Compared to control, the SOD, POD and CAT isozymes of cucumber seedlings under hypoxic stress treatment were decreased at3th,6th and9th days. However, BR increased the expression of SOD isozymes by5.22%and10.25%on3th day and6th day compared to hypoxic treatment and POD isozymes were also increased13.81%and9.46%, respectively. But CAT isozymes were not changed significantly.
The effects of EBRs on the gene expression of respiration and antioxidant in cucumber seedlings under short-term hypoxic stress suggested that the gene expression of ldh, pdc, idh, sdh, mdh, ald, ugpase, pfk, hk, pod increased and cat, sod and fk decreased at3th day of treatment. After application the EBR, the expression of ldh、idh and pfk were further increased but the pdc, sdh, mdh, cat, sod and ugpase had a slight decreased. Our results suggested that hypoxic stress inhibited the growth of cucumber seedling roots and EBR might regulate the isozyme expressions of anaerobic respiration, aerobic respiration and antioxidant enzymes of cucumber roots thereby alleviated the suppression effects of hypoxic stress.
3. Effects of exogenous brassinolide on the mitochondria antioxidant system, ATPase activity and cellular ultrastructure in cucumber seedling roots under hypoxic stress
Under hypoxic stress, the superoxide anion (O2-) production rate, hydrogen peroxide (H2O2) content and malondialdehyde (MDA) contents were significantly enhanced, and root and ATPase activity were decreased resulting in cell damage. Compared with the control, activities of SOD, POD and CAT were induced and were higher at3d, and significantly lowered at6d by hypoxic stress. However, the treatment of EBR could keep higher activities of SOD, POD and CAT for a longer time, reduce the generation of O2, H2O2and MDA contents and significantly increased the root and ATPase activity, injured of root tip cell was released. The results indicated that exogenous EBR raised the activities of mitochondria antioxidant enzymes, decreased the O2-production rate, H2O2and MDA contents. Therefore, exogenous EBR might improve the mitochondria antioxidant system, ATPase activity and cellular ultrstructure of cucumber seedlings under hypoxic stress thereby alleviating the adverse effects of hypoxic stress.
4. Effects of exogenous brassinolide on photosynthesis, poly amines and inoroganic ions content in cucumber seedlings under hypoxic stress
When exposed to hypoxic stress, net photosynthetic rate (Pn), apparent quanta yield (AQY) and carboxylation efficiency (CE) were decreased significantly, and quantum efficency of open PSⅡ reaction center (Fv'/Fm'), photochemical quenching (qP), PSⅡ photochemisty rate (ΦPSⅡ), however the raletive electron transport rate(rETR)in leaves of cucumber seedlings were significantly decreased, but the allocation of antenna heat dissipation energy (Drate), and non-photochemical quenching(qN) were significantly increased, and the cellular ultrastructure was abnormal, which indicated that hypoxia caused photo-inhibition to cucumber seedlings. However, EBR could alleviate or release the damages in chloroplast and mitochondria under hypoxic stress, and enhanced Pn, Fv'/Fm', qP, rETR, PSⅡ and Prate. The results suggested that EBR could play a positive role in alleviating damage in mitochondria and chloroplast ultrastructure and photosynthetic characteristics, and make the leaf keep a higher photosynthetic performance, and alleviate the damage of hypoxic stress.
Hypoxic stress significantly increased the contents of polyamines including Put, Spd, Spm, total PAs and Put/PAs ratio in leaves and root, and decreased (Spd+Spm)/Put ratio. The change of root was greater than that of leaf. K+, Ca2+, Mg2+, Fe3+, Zn2+and Mn2+content in leaves and K+, Ca2+, Mg2+, Zn2+content in roots significantly decreased, too. However, Exogenous EBR increased the contents of the free Spm, conjugated Spd, Spm and bound Put, Spd, Spm. Meanwhile, EBR further increased the PAs content and (Spd+Spm)/Put ratio and reduced the Put/PAs ratio. Ca2+, Mg2+, Fe3+, Zn2+and Mn2+content in leaves and Ca2+, Mg2+, Fe3+, Mn2+content in roots significantly increased, too. Thus, EBR not only could regulate the formation and contents of polyamines but also promote the absorption of nutrients, make the leaf keep a higher photosynthetic performance, and alleviate the damage of hypoxic stress.
引文
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