三种石斛的光合电子传递、碳同化动态研究
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摘要
本文以报春石斛、金钗石斛、鼓槌石斛为试验材料,测定了其碳同化、光合作用系统电子传递特征;同步研究了三种石斛PSII荧光动力学和PSI氧化还原信号在不同处理之间的变化规律;检测了三种石斛叶片中苹果酸含量和磷酸烯醇式丙酮酸羧化酶日变化动态;观察了三种石斛气孔形态和开闭特点,探究了碳同化、电子传递和气孔运动三者之间的联动规律。
结果表明,报春石斛为景天科酸代谢植物;正常栽培条件下和轻度干旱环境条件下金钗石斛和鼓槌石斛的碳同化行C3途径,比较严重的干旱胁迫可以诱导金钗石斛和鼓槌石斛行景天科酸代谢途径,而且鼓槌石斛可以昼夜不停地进行碳同化,全天保持二氧化碳交换速率为正值。
冬季温室栽培条件下,三种石斛对光照条件的需求差异很大。报春石斛和鼓槌石斛可以适应1000μmolphotonm-2s-1的光照,而金钗石斛的光饱和点在300μmolphotonm-2s-1。石斛属植物光合作用速率普遍偏低,三种石斛光合速率最高者为鼓槌石斛,可以达到3.92μmolCO2m-2s-1。光合作用日变化显示,报春石斛表现为景天科酸代谢植物特征。
生长季节报春石斛的光饱和点有所降低,为700μmolphotonm-2s-1;光合作用速率提高,高达3.826μmolCO2m-2s-1;光合作用日变化呈现出CAM植物的四阶段特征。鼓槌石斛的光饱和点降低为800-900μmolphotonm-2s-1之间,最大光合作用速率升高到5.912μmolCO2m-2s-1;光合作用日变化呈单峰曲线,最大值出现在中午前后。金钗石斛更适合荫蔽环境,光合作用日变化呈双峰曲线,中午光照最强烈的时段表现为一定程度的光抑制。
报春石斛晚间黑暗条件下PEPC羧化酶活性最强,而在中午光照最充足时,酶的活性相对较弱;金钗石斛的PEPC羧化酶活性日变化特点与报春石斛相似,但是活性比较弱;鼓槌石斛的PEPC羧化酶活性很弱,昼夜变化幅度很小,表现为黑暗状态下比光下更强。
在冬季,报春石斛的最大光量子光化学效率低于0.8,叶绿体光合作用机构处于非正常状态,叶片失绿,说明其进入休眠状态;金钗石斛的最大光量子光化学效率为0.828,但是其实际光量子光化学效率、光化学猝灭系数在冬季普遍处于较低水平,说明其光合作用活性不高,表现出一定程度的休眠;鼓槌石斛能够很好地适应冬季温室环境,整个冬季,最大光量子光化学效率都比较高,即便是在临晨低温的11月中下旬,其Fv/Fm高达0.8,可见其适应低温的能力是最强的。而且,鼓槌石斛天线色素吸收的能量用于光化学反应的份额比较高,PSII反应中心开放的比例高,利于电子的快速传递,迅速形成同化力。因此,冬季温室栽培的鼓槌石斛保持着较高的光合作用能力。
黑暗处理后报春石斛的最大荧光值低于同一光合作用能力的金钗石斛,也低于12h黑暗处理的报春石斛,表明2h黑暗处理的报春石斛的电子传递链存在电子传递,为叶绿体呼吸作用(chlororespiration)提供了证据。
暗适应后照光处理的时间对于光化学反应有影响,对于本试验条件下的C3植物金钗石斛和鼓槌石斛,4h以上的光化光照射有利于其光合作用机构的正常运转;对于CAM途径的报春石斛,暗适应后2h的照光使其光合作用速率达到最高,其光化学反应与二氧化碳供应关系密切。
报春石斛叶片苹果酸含量表现出规律性的昼夜变化,其节律与典型CAM植物相似,呈现四阶段的鲜明特征。
CAM植物报春石斛光下气孔近乎关闭,但是光合作用系统电子传递正常进行。鼓槌石斛、金钗石斛明显存在气孔不均匀开闭现象。三种石斛的气孔结构相似,气孔复合体为不规则型,没有副卫细胞;保卫细胞外侧罩有拱盖。
文章首次在园林植物上根据叶绿素荧光动力学和PSI氧化还原信号同步研究两个光合作用系统电子传递状态;首次对不作任何处理的鲜活叶片样品进行快速显微观察,捕捉到真实的气孔不均匀开闭实时状态;并且,对同步研究碳同化、电子传递、气孔运动三者的联动关系做出创造性探索。
Three dendrobia named Dendrobium primulinum, D. nobile, D.chrysotoxum were employed in the trials. Carbon assimilation, electron transport across the two photosystems were explored; the fluctuation of chlorophyll fluorescence related to PSII and P700 signal pertinent to PSI were investigated synchronously; the diurnal changes of malic acid contents in fresh leaves was measured; the diurnal changes of the activity of phosphoenolpyruvate carboxylase (PEPC) was inspected; stomata morphology and stomata movement were observed; moreover, the experiment design was initiated to realize synchronous study on the linkage between three physiological processes including carbon assimilation, electron transport and stomata movement.
The results demonstrated that D.primulinum is a CAM (Crassulacean acid metabolism) species. Under normal conditions and in moderate arid environment, D.nobile and D.chrysotoxum performed C3 photosynthesis. However, serious drought could induce them switching into CAM pathway, furthermore, the carbon assimilation could operate unremittingly round the clock in D.chrysotoxum, and it always kept net absorb of carbon dioxide day and night.
In green house during winter, the requirement for light condition was different significantly from one to another among the three dendrobia. D.primulinum and D.chrysotoxum could be adapt to the highlight as intense as 1000μmolphotonm-2s-1, while the light saturation point was 300μmolphotonm-2s-1. Generally, the photosynthetic rate was low in dendrobia. Among the three dendrobia, the maximum photosynthetic rate was 3.92μmolCO2m-2s-1in D.chrysotoxum during winter. Diurnal change of photosynthesis in D.primulinum exhibited typical feature of CAM with nocturnal carbon assimilation.
In growth season, the light saturation point of D.primulinum declined to 300μmolphotonm-2s-1, and the maximum photosynthetic rate increased to 3.826μmolCO2m-2s-1. The diurnal change of photosynthesis was composed by four distinct phases. The light saturation point of D.chrysotoxum lowered to 800μmolphotonm-2s-1, and maximum photosynthetic rate rose to 5.912μmolCO2m-2s-1, while the diurnal change of photosynthesis showed single summit at noon.
D.nobile preferred to shade environment, whose diurnal change of photosynthesis showed double summits, and there was photo inhibit at noon when illumination was intense.
Activity of PEPC of D.primulinum from dark leaves was strong, whilst it was relatively weak at noon when sun light was bright. The diurnal change of activity of PEPC in D.nobile was resembled to D.primulinum. The activity of PEPC in D.chrysotoxum was very weak, and the magnitude of diurnal change was small; anyway, its activity from dark leaves was stronger than one from illuminated leaves.
The maximal photochemical efficiency of PSII in D.primulinum during winter was lower than 0.8, and the photosynthetic apparatus was in abnormal state that there was chlorophyll deficiency, which indicated that D.primulinum fell into dormancy. The maximal photochemical efficiency in D.nobile was 0.828, but its actual photochemical efficiency and photochemical quenching coefficient were very small, which showed that its photosynthesis was sluggish and was dormant to some certain during winter. D.chrysotoxum could perfectly acclimatize itself to the greenhouse environment in winter, its maximal photochemical efficiency was higher even as high as 0.8 when encountered to the chilling stress. The ratio for photochemistry from the energy absorbed by antenna pigment in D.chrysotoxum was high, and there were more open centers of PSII, which was conducive to electron transport so as to rapidly produce assimilatory power. Hence, the photosynthesis processes were vigorous in D.chrysotoxum during winter.
After 2 hours of darkness treatment, the maximal fluorescence was lower than one of D.nobile which was in the same level based on the photosynthetic rate as D.primulinum, and the Fm was also lower than the value of maximal fluorescence of D.primulinum after 12 hours of darkness treatment. Therefore, there was electron transport in the D.primulinum treated in darkness for 2 hours, which probably related to chlororepiration.
The effect of illumination period on photochemical reaction after darkness adaptation was significant. Longer than 4 hours of actinic light illumination was beneficial to operating efficiently of photosynthesis apparatus for C3 species including D.nobile and D. chrysotoxum. As to D.primulinum, 2 hours of illumination after darkness adaptation resulted in the maximal photosynthetic rate, and there was close relationship between photochemical reaction and carbon dioxide supply.
Malic acid contents showed orderly diurnal fluctuation in the leaf of D.primulinum, and its rhythm demonstrated the characteristics of four phases exclusive to CAM species.
The stomata of D.primulinum were nearly close when exposed to light intenser than actinic light, but its electron transport operated smoothly. It was found that there was unevenly stomata close in D.nobile and D.chrysotoxum. Stomata configuration were similar among the three dendrobia, their stomata apparatus types were anomocytic, without subsidiary cells, and there was a cuticle stomata ledge outside of guard cells.
For the first time, chlorophyll fluorescence kinetics and P700 signal were simultaneously monitored and served as probe to study the electron transport across two photosystems in ornamental plants. For the first time, micro-observation was actualized on fresh leaf free from special treatment which disturbed the stomata movement, snapshooting the actual stomata uneven close. The blueprint tailored for synchronously exploring carbon assimilation, electron transport and stomata movement was initiated, and the primary experiment conducted lately foretold the bright future.
结果表明,报春石斛为景天科酸代谢植物;正常栽培条件下和轻度干旱环境条件下金钗石斛和鼓槌石斛的碳同化行C3途径,比较严重的干旱胁迫可以诱导金钗石斛和鼓槌石斛行景天科酸代谢途径,而且鼓槌石斛可以昼夜不停地进行碳同化,全天保持二氧化碳交换速率为正值。
冬季温室栽培条件下,三种石斛对光照条件的需求差异很大。报春石斛和鼓槌石斛可以适应1000μmolphotonm-2s-1的光照,而金钗石斛的光饱和点在300μmolphotonm-2s-1。石斛属植物光合作用速率普遍偏低,三种石斛光合速率最高者为鼓槌石斛,可以达到3.92μmolCO2m-2s-1。光合作用日变化显示,报春石斛表现为景天科酸代谢植物特征。
生长季节报春石斛的光饱和点有所降低,为700μmolphotonm-2s-1;光合作用速率提高,高达3.826μmolCO2m-2s-1;光合作用日变化呈现出CAM植物的四阶段特征。鼓槌石斛的光饱和点降低为800-900μmolphotonm-2s-1之间,最大光合作用速率升高到5.912μmolCO2m-2s-1;光合作用日变化呈单峰曲线,最大值出现在中午前后。金钗石斛更适合荫蔽环境,光合作用日变化呈双峰曲线,中午光照最强烈的时段表现为一定程度的光抑制。
报春石斛晚间黑暗条件下PEPC羧化酶活性最强,而在中午光照最充足时,酶的活性相对较弱;金钗石斛的PEPC羧化酶活性日变化特点与报春石斛相似,但是活性比较弱;鼓槌石斛的PEPC羧化酶活性很弱,昼夜变化幅度很小,表现为黑暗状态下比光下更强。
在冬季,报春石斛的最大光量子光化学效率低于0.8,叶绿体光合作用机构处于非正常状态,叶片失绿,说明其进入休眠状态;金钗石斛的最大光量子光化学效率为0.828,但是其实际光量子光化学效率、光化学猝灭系数在冬季普遍处于较低水平,说明其光合作用活性不高,表现出一定程度的休眠;鼓槌石斛能够很好地适应冬季温室环境,整个冬季,最大光量子光化学效率都比较高,即便是在临晨低温的11月中下旬,其Fv/Fm高达0.8,可见其适应低温的能力是最强的。而且,鼓槌石斛天线色素吸收的能量用于光化学反应的份额比较高,PSII反应中心开放的比例高,利于电子的快速传递,迅速形成同化力。因此,冬季温室栽培的鼓槌石斛保持着较高的光合作用能力。
黑暗处理后报春石斛的最大荧光值低于同一光合作用能力的金钗石斛,也低于12h黑暗处理的报春石斛,表明2h黑暗处理的报春石斛的电子传递链存在电子传递,为叶绿体呼吸作用(chlororespiration)提供了证据。
暗适应后照光处理的时间对于光化学反应有影响,对于本试验条件下的C3植物金钗石斛和鼓槌石斛,4h以上的光化光照射有利于其光合作用机构的正常运转;对于CAM途径的报春石斛,暗适应后2h的照光使其光合作用速率达到最高,其光化学反应与二氧化碳供应关系密切。
报春石斛叶片苹果酸含量表现出规律性的昼夜变化,其节律与典型CAM植物相似,呈现四阶段的鲜明特征。
CAM植物报春石斛光下气孔近乎关闭,但是光合作用系统电子传递正常进行。鼓槌石斛、金钗石斛明显存在气孔不均匀开闭现象。三种石斛的气孔结构相似,气孔复合体为不规则型,没有副卫细胞;保卫细胞外侧罩有拱盖。
文章首次在园林植物上根据叶绿素荧光动力学和PSI氧化还原信号同步研究两个光合作用系统电子传递状态;首次对不作任何处理的鲜活叶片样品进行快速显微观察,捕捉到真实的气孔不均匀开闭实时状态;并且,对同步研究碳同化、电子传递、气孔运动三者的联动关系做出创造性探索。
Three dendrobia named Dendrobium primulinum, D. nobile, D.chrysotoxum were employed in the trials. Carbon assimilation, electron transport across the two photosystems were explored; the fluctuation of chlorophyll fluorescence related to PSII and P700 signal pertinent to PSI were investigated synchronously; the diurnal changes of malic acid contents in fresh leaves was measured; the diurnal changes of the activity of phosphoenolpyruvate carboxylase (PEPC) was inspected; stomata morphology and stomata movement were observed; moreover, the experiment design was initiated to realize synchronous study on the linkage between three physiological processes including carbon assimilation, electron transport and stomata movement.
The results demonstrated that D.primulinum is a CAM (Crassulacean acid metabolism) species. Under normal conditions and in moderate arid environment, D.nobile and D.chrysotoxum performed C3 photosynthesis. However, serious drought could induce them switching into CAM pathway, furthermore, the carbon assimilation could operate unremittingly round the clock in D.chrysotoxum, and it always kept net absorb of carbon dioxide day and night.
In green house during winter, the requirement for light condition was different significantly from one to another among the three dendrobia. D.primulinum and D.chrysotoxum could be adapt to the highlight as intense as 1000μmolphotonm-2s-1, while the light saturation point was 300μmolphotonm-2s-1. Generally, the photosynthetic rate was low in dendrobia. Among the three dendrobia, the maximum photosynthetic rate was 3.92μmolCO2m-2s-1in D.chrysotoxum during winter. Diurnal change of photosynthesis in D.primulinum exhibited typical feature of CAM with nocturnal carbon assimilation.
In growth season, the light saturation point of D.primulinum declined to 300μmolphotonm-2s-1, and the maximum photosynthetic rate increased to 3.826μmolCO2m-2s-1. The diurnal change of photosynthesis was composed by four distinct phases. The light saturation point of D.chrysotoxum lowered to 800μmolphotonm-2s-1, and maximum photosynthetic rate rose to 5.912μmolCO2m-2s-1, while the diurnal change of photosynthesis showed single summit at noon.
D.nobile preferred to shade environment, whose diurnal change of photosynthesis showed double summits, and there was photo inhibit at noon when illumination was intense.
Activity of PEPC of D.primulinum from dark leaves was strong, whilst it was relatively weak at noon when sun light was bright. The diurnal change of activity of PEPC in D.nobile was resembled to D.primulinum. The activity of PEPC in D.chrysotoxum was very weak, and the magnitude of diurnal change was small; anyway, its activity from dark leaves was stronger than one from illuminated leaves.
The maximal photochemical efficiency of PSII in D.primulinum during winter was lower than 0.8, and the photosynthetic apparatus was in abnormal state that there was chlorophyll deficiency, which indicated that D.primulinum fell into dormancy. The maximal photochemical efficiency in D.nobile was 0.828, but its actual photochemical efficiency and photochemical quenching coefficient were very small, which showed that its photosynthesis was sluggish and was dormant to some certain during winter. D.chrysotoxum could perfectly acclimatize itself to the greenhouse environment in winter, its maximal photochemical efficiency was higher even as high as 0.8 when encountered to the chilling stress. The ratio for photochemistry from the energy absorbed by antenna pigment in D.chrysotoxum was high, and there were more open centers of PSII, which was conducive to electron transport so as to rapidly produce assimilatory power. Hence, the photosynthesis processes were vigorous in D.chrysotoxum during winter.
After 2 hours of darkness treatment, the maximal fluorescence was lower than one of D.nobile which was in the same level based on the photosynthetic rate as D.primulinum, and the Fm was also lower than the value of maximal fluorescence of D.primulinum after 12 hours of darkness treatment. Therefore, there was electron transport in the D.primulinum treated in darkness for 2 hours, which probably related to chlororepiration.
The effect of illumination period on photochemical reaction after darkness adaptation was significant. Longer than 4 hours of actinic light illumination was beneficial to operating efficiently of photosynthesis apparatus for C3 species including D.nobile and D. chrysotoxum. As to D.primulinum, 2 hours of illumination after darkness adaptation resulted in the maximal photosynthetic rate, and there was close relationship between photochemical reaction and carbon dioxide supply.
Malic acid contents showed orderly diurnal fluctuation in the leaf of D.primulinum, and its rhythm demonstrated the characteristics of four phases exclusive to CAM species.
The stomata of D.primulinum were nearly close when exposed to light intenser than actinic light, but its electron transport operated smoothly. It was found that there was unevenly stomata close in D.nobile and D.chrysotoxum. Stomata configuration were similar among the three dendrobia, their stomata apparatus types were anomocytic, without subsidiary cells, and there was a cuticle stomata ledge outside of guard cells.
For the first time, chlorophyll fluorescence kinetics and P700 signal were simultaneously monitored and served as probe to study the electron transport across two photosystems in ornamental plants. For the first time, micro-observation was actualized on fresh leaf free from special treatment which disturbed the stomata movement, snapshooting the actual stomata uneven close. The blueprint tailored for synchronously exploring carbon assimilation, electron transport and stomata movement was initiated, and the primary experiment conducted lately foretold the bright future.
引文
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