不同光、温对冷敏感植物黄瓜与耐冷植物Rumex K-1 PSⅠ和PSⅡ光化学活性的影响
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摘要
光合系统II(PSII)与光合系统I (PSI)共同组成了光合线性电子传递链,但两系统对温度/光胁迫存在着不同的敏感性。我们以冷敏感植物黄瓜(津春-4号)和起源于北纬45度的耐低温植物杂交酸膜(Rumex K-1(Patientia×R. tianschaious))为实验材料,通过测定光合气体交换、放氧活性、叶绿素a调制式脉冲荧光、820nm光透射与叶绿素a快相荧光同步信号、活性氧(ROS)与活性氧清除酶活性、叶绿体的超微结构等,研究了低温和高温在不同光强条件下对植物叶片光合机构的影响。主要结果如下:
1.在4℃与300μmolm~(-2)s~(-1)PFD胁迫下,黄瓜叶片饱和光下的CO_2同化速率和最大放氧速率出现急剧的下降,PSI的最大光化学效率(△I/Io)显著下降,同时PSII最大光化学效率(Fv/Fm)降低,表明发生了低温光抑制。在黄瓜与Rumex K-1叶片中,我们均发现:PSI光化学活性(△I/Io)的下降明显快于PSII捕获光量子驱动电子到QA-后电子传递链的效率(ΨO)。以上结果表明,对冷敏感植物黄瓜和耐冷植物Rumex K-1来说,PSI均为低温光抑制主要位点。
2.低温光胁迫下,黄瓜叶片PSII开放反应中心的捕光效率(Fv’/Fm’)和光下PSII实际光化学效率(ФPSII)均出现明显的降低,同时,电子传递链还原程度(1-qP)显著增强。我们在黄瓜和Rumex K-1叶片上均看到PSII活化压的增加与PSI光抑制间存在密切的关系。这可能是PSI光抑制限制了PSII的电子向PSI传递,增加了电子受体QA的还原状态。同时,Q_A~-的积累可减少流向PSI的电子,这可以减缓对PSI的进一步的光抑制。
3.低温光下,虽然黄瓜叶片超氧化物歧化酶(SOD)活性升高,但过氧化物酶(CAT),抗坏血酸-谷胱甘肽循环中的关键酶:抗坏血酸过氧化物酶(APX),谷胱甘肽还原酶(GR)和脱氢抗坏血酸还原酶(DHAR)的活性在逐步失活,单脱氢抗坏血酸还原酶(MDHAR)则几乎不变化,同时,叶片H_2O_2,O_2~(·|-)和丙二醛(MDA)含量逐渐积累。以上事实说明虽然黄瓜叶片SOD活性升高,但其他抗氧化酶系统活性的下降,不能有效地清除积累的活性氧,从而导致细胞的氧化破坏。
4.与耐冷植物Rumex K-1相比,在低温光胁迫下,黄瓜叶绿体膜脂组分的变化更为剧烈,其中磷脂酰胆碱(PC)、磷脂酰甘油(PG)以及硫代异鼠李糖基甘油二酯(SQDG)的不饱和度下降幅度更大。这可能是导致黄瓜PSII和PSI光化学活性以及放氧复合体活性对低温比较敏感原因之一。
5.黄瓜叶绿体对低温-光胁迫更加敏感。低温下,黄瓜类囊体逐渐扭曲和膨胀,叶绿体内膜形成小泡,最终,叶绿体被膜瓦解,使得基质与胞质组分出现混合。而Rumex K-1叶绿体能够进行有效的避光运动,其叶绿体的机构几乎未受低温-光胁迫的影响。我们认为,在过剩光能条件下,叶绿体避光运动在减少光能吸收,降低叶绿体的氧化胁迫方面具有重要的生理意义。
6.高温(40℃)黑暗条件下,冷敏感植物黄瓜叶片放氧活性及PSI, PSII光化学活性不受影响,而耐冷植物Rumex K-1的放氧活性显著降低,PSII最大光化学效率下降,同时PSI反应中心色素P700还原明显地变缓。
7. 40℃,照光条件下,黄瓜光合系统活性维持稳定。100μmolm~(-2)s~(-1)光下Rumex K-1的OEC活性和PSII光化学活性几乎不受影响,同时P700+还原速率保持稳定。而300μmolm~(-2)s~(-1)光下,Rumex K-1的PSII,OEC活性受到轻微地抑制,导致P700~+还原速率出现轻微的下降。以上结果表明高温下,冷敏感植物黄瓜具有较高的热稳定性,而耐冷植物RumexK-1的OEC和PSII表现为热敏感性,在高温胁迫下PSII电子供应能力显著下降,限制了P700~+的还原。
The effects of different temperatures coupled with different light intensity on activity of photosynthetic apparatus in leaves of a chilling sensitive plant cucumber (Cucumis sativus L. cvs. Jinchun No4) and a chilling-tolerant plant Rumex K-1(Rumex Patientia×R. tianschaious) were studied by measurements of photosynthetic gas exchange, oxygen- evolution, modulated chlorophyll a fluorescence, 820nm transmission signal paralleling the chlorophyll a fluorescence transient, changes in reactive oxygen species (ROS) and activity of ROS scavenging enzymic system, ultrastructure of chloroplasts and so on. The main results obtained are followed:
1. With treatment of chilling temperature (4℃)coupled wth 300μmolm~(-2)s~(-1) light, the sharp decreases were observed in the light-saturated net CO_2 fixation and maximal oxygen evolution rate of cucumber. The maximal efficiency of PSI (△I/Io) reduced significantly, coupled with the decline of maximal photochemistry (Fv/Fm) in cucumber leaves. However, the redox states of PSI activity (△I/Io) was decreased more severely than the efficiency that a trapped exciton can move an electron into the electron transport chain beyond Q_A~- (ΨO) in both cucmuber and Rumex K-1 intact leaves. These results suggested that PSI should be the primary target of chilling-photoinhibition in the both chilling-sensitve and chilling-tolerant plants.
2. The efficiency of excitation capture by open PSII centers (Fv’/Fm’), and actual photochemical efficiency of PSII (ФPSII) decreased considerably, at the same time, the reduction of electron transfer chain (1-qP) increased greatly in cucumber leaves. The close correlation of excitation pressure on PSII and PSI photoinhibition were observed in both the cucumber and Rumex K-1 leaves, indicating that the reduction state of QA is dependent of the extent of PSI photoinhibition. The accumulation of QA- could lessen electron flow to PSI,this might be important for protection against the photoinhibition of PSI.
3. The content of H2O_2, O 2 and MDA in leaves increased largely. The activity of catelase (CAT), ascorbate peroxidase (APX), glutathione reducase (GR) and dehydroascorbate reductase (DHAR) were decreased in the leaves of cucumber treated with chilling temperature under moderate light, while activity of monodyhydroascorbate reductase (MDHAR) remained unchanged, and the activity of superoxide dismulate (SOD) increased. These data suggested that although the activity of SOD was increased, the activitity of antioxidant system was generally decreased in chilling-sensitive plant cucumber treated with chilling temperatures under moderate light, so that they were not able to scavenge the accumulated reactive oxygen species (ROS), resulting in cellular oxidative damage in cucumber leaves.
4. Compared with the chilling–tolerant plant Rumex K-1, the membrane lipid composition of cucumber changed more noticeably and the polyunsaturated fatty acids index of PC, PG and SQDG decreased more remarkable in cucumber treated with chilling temperature combined with moderate light, which might be one of the important reseaons resulting in chilling-sensitivity of photochemical activity of PSI, PSII and OEC in cucumber leaves.
5.Compared with the chilling–tolerant plant Rumex K-1, chloroplasts of cucumber were more vulnerable to chilling-light stress. After chilling treatment under moderate light, the thylakoid progressivly became distorted, swelled, and small vesicles were formed from inner membrane of chloroplasts. The chloroplast envelopes were eventually disintegrated, as a result, stromal contents mixed with the cytoplasm. While a striking chloroplasts reoriention was observed in Rumex K-1 leaves. The chloroplasts of Rumex K-1 were not affected obviously under chilling-light treatment. This observations supported the view that chloroplast avoidance movement play an important role in decreasing the light absorbation by photosystems, reducing oxidantive pressure on chloroplasts under excess light .
6. Photosynthetic oxygen evolution, PSI and PSII photochemistry was maintained steady in cucumber leaves exposed to high temperature (40℃) in the dark , while OEC activity and PSII photochemistry as well as the rate of P700~+ reduction were markedly reduced in Rumex K-1 leave exposed to high temperature (40℃) in the dark.
7. The PSI and PSII photochemistry were hardly affected in cucumber leaves treated with 40℃under low and moderate light intensities. The OEC activity, PSII photochemistry and the rate of P700+ reduction were not obviously chaged in Rumex K-1 leaves eoxposed to 40℃under low light. Whereas, under 300μmol m~(-2) s~(-1) PFD combined with 40℃, photochemistry of PSII, activity of OEC and P700+ reduction were slightly inhibited in Rumex K-1. These results indicated that the photosynthetic apparatus of chilling-sensitive cucumber was thermotolerant, however, the OEC complex and PSII photochemisty of Rumex K-1 were heat-sensitive, the severely impaired electron donation from PSII limited the P700+ reduction in Rumex K-1.
1.在4℃与300μmolm~(-2)s~(-1)PFD胁迫下,黄瓜叶片饱和光下的CO_2同化速率和最大放氧速率出现急剧的下降,PSI的最大光化学效率(△I/Io)显著下降,同时PSII最大光化学效率(Fv/Fm)降低,表明发生了低温光抑制。在黄瓜与Rumex K-1叶片中,我们均发现:PSI光化学活性(△I/Io)的下降明显快于PSII捕获光量子驱动电子到QA-后电子传递链的效率(ΨO)。以上结果表明,对冷敏感植物黄瓜和耐冷植物Rumex K-1来说,PSI均为低温光抑制主要位点。
2.低温光胁迫下,黄瓜叶片PSII开放反应中心的捕光效率(Fv’/Fm’)和光下PSII实际光化学效率(ФPSII)均出现明显的降低,同时,电子传递链还原程度(1-qP)显著增强。我们在黄瓜和Rumex K-1叶片上均看到PSII活化压的增加与PSI光抑制间存在密切的关系。这可能是PSI光抑制限制了PSII的电子向PSI传递,增加了电子受体QA的还原状态。同时,Q_A~-的积累可减少流向PSI的电子,这可以减缓对PSI的进一步的光抑制。
3.低温光下,虽然黄瓜叶片超氧化物歧化酶(SOD)活性升高,但过氧化物酶(CAT),抗坏血酸-谷胱甘肽循环中的关键酶:抗坏血酸过氧化物酶(APX),谷胱甘肽还原酶(GR)和脱氢抗坏血酸还原酶(DHAR)的活性在逐步失活,单脱氢抗坏血酸还原酶(MDHAR)则几乎不变化,同时,叶片H_2O_2,O_2~(·|-)和丙二醛(MDA)含量逐渐积累。以上事实说明虽然黄瓜叶片SOD活性升高,但其他抗氧化酶系统活性的下降,不能有效地清除积累的活性氧,从而导致细胞的氧化破坏。
4.与耐冷植物Rumex K-1相比,在低温光胁迫下,黄瓜叶绿体膜脂组分的变化更为剧烈,其中磷脂酰胆碱(PC)、磷脂酰甘油(PG)以及硫代异鼠李糖基甘油二酯(SQDG)的不饱和度下降幅度更大。这可能是导致黄瓜PSII和PSI光化学活性以及放氧复合体活性对低温比较敏感原因之一。
5.黄瓜叶绿体对低温-光胁迫更加敏感。低温下,黄瓜类囊体逐渐扭曲和膨胀,叶绿体内膜形成小泡,最终,叶绿体被膜瓦解,使得基质与胞质组分出现混合。而Rumex K-1叶绿体能够进行有效的避光运动,其叶绿体的机构几乎未受低温-光胁迫的影响。我们认为,在过剩光能条件下,叶绿体避光运动在减少光能吸收,降低叶绿体的氧化胁迫方面具有重要的生理意义。
6.高温(40℃)黑暗条件下,冷敏感植物黄瓜叶片放氧活性及PSI, PSII光化学活性不受影响,而耐冷植物Rumex K-1的放氧活性显著降低,PSII最大光化学效率下降,同时PSI反应中心色素P700还原明显地变缓。
7. 40℃,照光条件下,黄瓜光合系统活性维持稳定。100μmolm~(-2)s~(-1)光下Rumex K-1的OEC活性和PSII光化学活性几乎不受影响,同时P700+还原速率保持稳定。而300μmolm~(-2)s~(-1)光下,Rumex K-1的PSII,OEC活性受到轻微地抑制,导致P700~+还原速率出现轻微的下降。以上结果表明高温下,冷敏感植物黄瓜具有较高的热稳定性,而耐冷植物RumexK-1的OEC和PSII表现为热敏感性,在高温胁迫下PSII电子供应能力显著下降,限制了P700~+的还原。
The effects of different temperatures coupled with different light intensity on activity of photosynthetic apparatus in leaves of a chilling sensitive plant cucumber (Cucumis sativus L. cvs. Jinchun No4) and a chilling-tolerant plant Rumex K-1(Rumex Patientia×R. tianschaious) were studied by measurements of photosynthetic gas exchange, oxygen- evolution, modulated chlorophyll a fluorescence, 820nm transmission signal paralleling the chlorophyll a fluorescence transient, changes in reactive oxygen species (ROS) and activity of ROS scavenging enzymic system, ultrastructure of chloroplasts and so on. The main results obtained are followed:
1. With treatment of chilling temperature (4℃)coupled wth 300μmolm~(-2)s~(-1) light, the sharp decreases were observed in the light-saturated net CO_2 fixation and maximal oxygen evolution rate of cucumber. The maximal efficiency of PSI (△I/Io) reduced significantly, coupled with the decline of maximal photochemistry (Fv/Fm) in cucumber leaves. However, the redox states of PSI activity (△I/Io) was decreased more severely than the efficiency that a trapped exciton can move an electron into the electron transport chain beyond Q_A~- (ΨO) in both cucmuber and Rumex K-1 intact leaves. These results suggested that PSI should be the primary target of chilling-photoinhibition in the both chilling-sensitve and chilling-tolerant plants.
2. The efficiency of excitation capture by open PSII centers (Fv’/Fm’), and actual photochemical efficiency of PSII (ФPSII) decreased considerably, at the same time, the reduction of electron transfer chain (1-qP) increased greatly in cucumber leaves. The close correlation of excitation pressure on PSII and PSI photoinhibition were observed in both the cucumber and Rumex K-1 leaves, indicating that the reduction state of QA is dependent of the extent of PSI photoinhibition. The accumulation of QA- could lessen electron flow to PSI,this might be important for protection against the photoinhibition of PSI.
3. The content of H2O_2, O 2 and MDA in leaves increased largely. The activity of catelase (CAT), ascorbate peroxidase (APX), glutathione reducase (GR) and dehydroascorbate reductase (DHAR) were decreased in the leaves of cucumber treated with chilling temperature under moderate light, while activity of monodyhydroascorbate reductase (MDHAR) remained unchanged, and the activity of superoxide dismulate (SOD) increased. These data suggested that although the activity of SOD was increased, the activitity of antioxidant system was generally decreased in chilling-sensitive plant cucumber treated with chilling temperatures under moderate light, so that they were not able to scavenge the accumulated reactive oxygen species (ROS), resulting in cellular oxidative damage in cucumber leaves.
4. Compared with the chilling–tolerant plant Rumex K-1, the membrane lipid composition of cucumber changed more noticeably and the polyunsaturated fatty acids index of PC, PG and SQDG decreased more remarkable in cucumber treated with chilling temperature combined with moderate light, which might be one of the important reseaons resulting in chilling-sensitivity of photochemical activity of PSI, PSII and OEC in cucumber leaves.
5.Compared with the chilling–tolerant plant Rumex K-1, chloroplasts of cucumber were more vulnerable to chilling-light stress. After chilling treatment under moderate light, the thylakoid progressivly became distorted, swelled, and small vesicles were formed from inner membrane of chloroplasts. The chloroplast envelopes were eventually disintegrated, as a result, stromal contents mixed with the cytoplasm. While a striking chloroplasts reoriention was observed in Rumex K-1 leaves. The chloroplasts of Rumex K-1 were not affected obviously under chilling-light treatment. This observations supported the view that chloroplast avoidance movement play an important role in decreasing the light absorbation by photosystems, reducing oxidantive pressure on chloroplasts under excess light .
6. Photosynthetic oxygen evolution, PSI and PSII photochemistry was maintained steady in cucumber leaves exposed to high temperature (40℃) in the dark , while OEC activity and PSII photochemistry as well as the rate of P700~+ reduction were markedly reduced in Rumex K-1 leave exposed to high temperature (40℃) in the dark.
7. The PSI and PSII photochemistry were hardly affected in cucumber leaves treated with 40℃under low and moderate light intensities. The OEC activity, PSII photochemistry and the rate of P700+ reduction were not obviously chaged in Rumex K-1 leaves eoxposed to 40℃under low light. Whereas, under 300μmol m~(-2) s~(-1) PFD combined with 40℃, photochemistry of PSII, activity of OEC and P700+ reduction were slightly inhibited in Rumex K-1. These results indicated that the photosynthetic apparatus of chilling-sensitive cucumber was thermotolerant, however, the OEC complex and PSII photochemisty of Rumex K-1 were heat-sensitive, the severely impaired electron donation from PSII limited the P700+ reduction in Rumex K-1.
引文
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