盐碱土上马蔺的渗透调节和光合适应性研究
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摘要
本文研究了生长在盐碱土上马蔺的生长、渗透调节和内源激素ABA的变化,盐碱土生长的马蔺光合日变化、光适应特性、光合机构的控制和马蔺对盐碱土和干旱的光合特点,结果如下:
1、马蔺在7～8月份生长速率较快,其平均株高为50～90cm,分蘖枝为5～8个,生物量为47.2g/株。8月份以后生物量逐渐减小。马蔺的生长过程呈S型曲线变化。生物量干重增重过程大致分为缓慢、快速和停止生长3个生长阶段。马蔺叶片中Na~+、K~+和脯氨酸的含量随着盐碱度的变化而改变,盐碱度较大,即pH高的月份,马蔺叶片中Na~+、K~+和脯氨酸的含量也同样增加,叶片细胞的膜透性增强。渗透调节物质的增加,降低了叶片水势,维持了细胞膨压,保持了马蔺在盐碱土的正常生长。
2、与对照相比处理材料叶片中ABA含量显著增加,外源ABA的加入降低了盐碱胁迫引起的MDA和H_2O_2量的增加,延缓了盐碱胁迫引起的CAT活性的衰减,提高了GR活性。盐碱长期胁迫明显增加了马蔺叶片ABA含量,但是不同植物根系中ABA含量存在着差异。在8天内,马蔺和大豆根系的ABA含量与土壤中ABA含量相差不大,根系ABA流向土壤,使土壤ABA水平在短期内提高,与土壤ABA之间建立了平衡关系。玉米根系ABA明显高于土壤ABA水平,与土壤之间没有交换关系,这主要与玉米根系内皮层的凯氏带有关。
3、马蔺叶片Pn和Tr均呈现双峰曲线,具有光合午休现象。这种现象出现的原因是由于较高的太阳辐射引起叶片温度的升高,叶片的羧化效率(CE)和表观量子产额(AQY)的下降造成的。
4、在马蔺叶片的成熟衰退过程中,光合能力、羧化效率、表观量子效率都逐渐下降,但是PSⅡ最大光化学效率在整个生育期基本维持恒定。同时叶片的羧化效率(CE)和RuBP的再生能力(A_(350))二者下降的幅度高于表观量子产额(AQY)和电子传递能力(J_(max))。也就是说,在马蔺叶片光合衰退过程中,光饱和时的光合速率及光饱和点的下降幅度显著大于CO_2饱和时的光合速率及CO_2饱和点的下降幅度,A_(350)/J_(max)逐渐下降,说明光合暗反应过程的衰退幅度大于光反应过程的衰退幅度。
5、马蔺叶片原初光化学效率(F_v/F_m),实际光化学效率(ΦPSⅡ)的日变化与光强有关,中午PSⅡ功能下调,有明显的光抑制。在光合有效辐射(PAR)逐渐增强时,F_v/F_m和ΦPSⅡ降低,非光化学猝灭(NPQ)增加,激发能由PSⅠ向PSⅡ分配,说明引起马蔺光抑制的原因除了NPQ之外,分配给PSⅡ能量增加后进一步促进PSⅡ可逆失活是非常重要的因素。PAR逐渐减小时,光抑制逐渐减缓,光化学效率恢复,这主要是NPQ减小,尤其是激发能分配不平衡性系数(β/α-1)下降,PSⅡ能量负荷减轻,失活的PSⅡ恢复活性的结果。
6、盐碱(含盐量0.36%、pH8.90)处理对马蔺叶片PSⅡ光化学活性(F_v/F_m)影响不大,但降低了电子传递速率(ETR)。盐碱伴随着干旱胁迫明显降低了PSⅡ最大光化学效率(F_v/F_m)、光化学猝灭(qP)和PSⅡ反应中心的光能捕获效率(F_v′/F_m′),分配给PSⅠ的激发能减少了33.3%,分配给PSⅡ激发能的份额增加了18.1%,激发能分配失衡。进一步揭示了盐碱和干旱胁迫下,非光化学耗散(NPQ)的增加和状态转换(由状态2向状态1转换)是马蔺减弱逆境伤害的重要适应机制。
7、盐胁迫降低了马蔺光合作用和气孔导度,对F_v/F_m没有影响。在盐胁迫的基础上碱胁迫却降低了F_v/F_m。外源Ca~(2+)和ABA可以明显缓解盐胁迫下的光抑制,外源Ca~(2+)对碱胁迫作用不大,ABA发挥重要的作用。因此,pH值对马蔺光合作用的影响比NaCl更为严重。
For several years, Iris lactea Pall, var chinensis(Fisch .) Koidz is bred a tolerant herbage to salt and alkali soil as virescence. In this paper, Ecophysiology of Iris lactea Pall, var chinensis(Fisch.) Koidz were studied to investigate growth, osmotic regulation, photosynthetic diurnal changes, photosynthetic apparatus characteristics grown in salt and alkali soil. The results were followings:1. A study was conducted to determine growth characteristics, capacity of osmotic adjustment of Iris Lactea Pall, var Chinensis Koidz grown in salt-alkali soil of Songnen plain in China. Iris Lactea Pall, var Chinensis Koidz grew more rapid on July and August, plant height is 50~90cm, and has 5-8 branches. The biomass per plant is 47.2g/plant during growth and development period. The process of Iris growth is divided three stages: slow growth stage, rapid growth stage and stop growth stage. Substances of osmotic adjustment were analyzed to determine the capacity of osmotic adjustment and adaptation in Iris leaves. Accumulation of Na~+ and K~+, proline concentration and improvement of cell membrane penetration were conducted to adjusted very rapidly to salt and alkali content during each month. With the accumulation osmotic substances, water potentials and osmotic potentials in Iris leaves decreased, cell turgid pressure increased or maintained to preserve cell, leaf and then whole plant growth under salt and drought condition.2. The results showed that the ABA contents in leaves after salt and alkali treats increased markedly. The addition of exogenous ABA decreased the increment of MDA and H2O2 contents and retarded the attenuation of CAT activity, which were induced by salt and alkali treats, and enhanced the GR activity. The content of leaf ABA increased under long-term salt and alkali stress, but root ABA differed among plants. However, in corn roots which possess a well-developed exodermis, alkaline and saline conditions in the rhizosphere did not reduce the endogenous ABA concentration, because the leaching of ABA from corn roots into the rhizosphere was lower than that from Iris and soybean roots. ABA efflux from corn and Iris and soybeab roots into the soil solution was observed only during the first days of the experiments and thereafter became substantially decreased.3. The data acquired in this study showed that diurnal changes of Pn and Tr were two-peak-type-curve, and was marked midday depression of Pn, which could be fully accounted for Gs changes. Carboxylic efficiency(CE) photosynthetic rate at saturation for intercellular CO2 concentration(A_350) apparent quantum yield(AQY) and photosynthetic rate at light saturation(Pmax) decreased in the midday. The result demonstrated that midday depression could be attributed to leaf temperature rising caused by the increasing of strong solar radiation (PAR).4. The results showed that Pn decreased with leaf ripping, and CO_2 carboxylic efficiency (CE), apparent quantum yield(AQY) declined at same time, but maximum photochemical efficiency(Fv/Fm) had little changed during leaf senescence of Iris lactea Pall, var chinensis(Fisch.) Koidz. During senescence of Iris lactea Pall, var chinensis(Fisch.) Koidz, CE and A_350 (net photosynthetic rate at light saturation) declined more rapid than AQY and J_max (net photosynthetic rate at CO_2 saturation (net photosynthetic rate at light saturation). This result showed that dark reaction or carbon cycle of photosynthesis declined more rapid than light reaction.5. The data acquired in this study showed that the diurnal changes of the maximum PS II photochemical efficiency (F_v/F_m) and actual photochemical efficiency of PS II (OPS II) were related to light intensity, the functional photosystem II reaction centers were decreased to cause significant photoinhibition. With the increasing of photosynthetic actual irradiation (PAR) to simulate
PAR diurnal change, Fv/Fm and OPS II decreased, nonphotochemical quenching (NPQ) increased, and the fraction of excited energy allocated to PS II increased to enhance PS II burden carrying excited energy. These results indicated that photoinhibition in Iris leaves are caused primarily by increasing of excited energy allocated to PS II to enhance reversible inactivation of PS II centers except NPQ increasing. With the decreasing of PAR, Fv/Fm and OPS II increased, NPQ and unbalance index of excited energy allocated to PS II and PS I (p/a-1) declined respectively. This indicated that the recovery of PS II photochemical efficiency is caused primarily by renewing PS II activation owing to unloading the energy burden of PS II.6. Salt and alkali treatment (the content of salt 0.36%,pH 8.90) alone had no effect on the maximal photochemistry of PS II (Fv/Fm), but decreased apparent photosynthetic electron transport rate(ETR).However, the salt and alkali treatment accompanied with drought decreased Fv/Fm and the efficiency of excitation captured open PS II(Fv'/Fm'), and state transition (from II to I) caused 33.3% decrease in allocation of excited energy to PS I and 18.1% increase to PS II. At the same time, the salt and alkali treatment accompanied with drought resulted in serious imbalance of excited energy distributions between two photosystems, enhanced the pressure on PS II .It is suggested that Nonphotochemical quenching (NPQ) could play a key role in adaptation mechanism of plant to stress.7. NaCl treatment alone had no effect on the maximal photochemistry of PSII. However, the NaCl treatment modified pH stress on PSII photochemistry in Iris leaves, which was manifested by a lesser pH-induced decrease in photochemical quenching (qP), efficiency of excitation energy capture by open PSII reaction centers (Fv7Fm'), and quantum yield of PSII electron transport (OPSII). Additions of calcium and ABA could alleviate photoinhibition in Iris leaves treated by NaCl. But Additions of calcium has no effects on alleviating under alkaline stress, and ABA does.
1、马蔺在7～8月份生长速率较快,其平均株高为50～90cm,分蘖枝为5～8个,生物量为47.2g/株。8月份以后生物量逐渐减小。马蔺的生长过程呈S型曲线变化。生物量干重增重过程大致分为缓慢、快速和停止生长3个生长阶段。马蔺叶片中Na~+、K~+和脯氨酸的含量随着盐碱度的变化而改变,盐碱度较大,即pH高的月份,马蔺叶片中Na~+、K~+和脯氨酸的含量也同样增加,叶片细胞的膜透性增强。渗透调节物质的增加,降低了叶片水势,维持了细胞膨压,保持了马蔺在盐碱土的正常生长。
2、与对照相比处理材料叶片中ABA含量显著增加,外源ABA的加入降低了盐碱胁迫引起的MDA和H_2O_2量的增加,延缓了盐碱胁迫引起的CAT活性的衰减,提高了GR活性。盐碱长期胁迫明显增加了马蔺叶片ABA含量,但是不同植物根系中ABA含量存在着差异。在8天内,马蔺和大豆根系的ABA含量与土壤中ABA含量相差不大,根系ABA流向土壤,使土壤ABA水平在短期内提高,与土壤ABA之间建立了平衡关系。玉米根系ABA明显高于土壤ABA水平,与土壤之间没有交换关系,这主要与玉米根系内皮层的凯氏带有关。
3、马蔺叶片Pn和Tr均呈现双峰曲线,具有光合午休现象。这种现象出现的原因是由于较高的太阳辐射引起叶片温度的升高,叶片的羧化效率(CE)和表观量子产额(AQY)的下降造成的。
4、在马蔺叶片的成熟衰退过程中,光合能力、羧化效率、表观量子效率都逐渐下降,但是PSⅡ最大光化学效率在整个生育期基本维持恒定。同时叶片的羧化效率(CE)和RuBP的再生能力(A_(350))二者下降的幅度高于表观量子产额(AQY)和电子传递能力(J_(max))。也就是说,在马蔺叶片光合衰退过程中,光饱和时的光合速率及光饱和点的下降幅度显著大于CO_2饱和时的光合速率及CO_2饱和点的下降幅度,A_(350)/J_(max)逐渐下降,说明光合暗反应过程的衰退幅度大于光反应过程的衰退幅度。
5、马蔺叶片原初光化学效率(F_v/F_m),实际光化学效率(ΦPSⅡ)的日变化与光强有关,中午PSⅡ功能下调,有明显的光抑制。在光合有效辐射(PAR)逐渐增强时,F_v/F_m和ΦPSⅡ降低,非光化学猝灭(NPQ)增加,激发能由PSⅠ向PSⅡ分配,说明引起马蔺光抑制的原因除了NPQ之外,分配给PSⅡ能量增加后进一步促进PSⅡ可逆失活是非常重要的因素。PAR逐渐减小时,光抑制逐渐减缓,光化学效率恢复,这主要是NPQ减小,尤其是激发能分配不平衡性系数(β/α-1)下降,PSⅡ能量负荷减轻,失活的PSⅡ恢复活性的结果。
6、盐碱(含盐量0.36%、pH8.90)处理对马蔺叶片PSⅡ光化学活性(F_v/F_m)影响不大,但降低了电子传递速率(ETR)。盐碱伴随着干旱胁迫明显降低了PSⅡ最大光化学效率(F_v/F_m)、光化学猝灭(qP)和PSⅡ反应中心的光能捕获效率(F_v′/F_m′),分配给PSⅠ的激发能减少了33.3%,分配给PSⅡ激发能的份额增加了18.1%,激发能分配失衡。进一步揭示了盐碱和干旱胁迫下,非光化学耗散(NPQ)的增加和状态转换(由状态2向状态1转换)是马蔺减弱逆境伤害的重要适应机制。
7、盐胁迫降低了马蔺光合作用和气孔导度,对F_v/F_m没有影响。在盐胁迫的基础上碱胁迫却降低了F_v/F_m。外源Ca~(2+)和ABA可以明显缓解盐胁迫下的光抑制,外源Ca~(2+)对碱胁迫作用不大,ABA发挥重要的作用。因此,pH值对马蔺光合作用的影响比NaCl更为严重。
For several years, Iris lactea Pall, var chinensis(Fisch .) Koidz is bred a tolerant herbage to salt and alkali soil as virescence. In this paper, Ecophysiology of Iris lactea Pall, var chinensis(Fisch.) Koidz were studied to investigate growth, osmotic regulation, photosynthetic diurnal changes, photosynthetic apparatus characteristics grown in salt and alkali soil. The results were followings:1. A study was conducted to determine growth characteristics, capacity of osmotic adjustment of Iris Lactea Pall, var Chinensis Koidz grown in salt-alkali soil of Songnen plain in China. Iris Lactea Pall, var Chinensis Koidz grew more rapid on July and August, plant height is 50~90cm, and has 5-8 branches. The biomass per plant is 47.2g/plant during growth and development period. The process of Iris growth is divided three stages: slow growth stage, rapid growth stage and stop growth stage. Substances of osmotic adjustment were analyzed to determine the capacity of osmotic adjustment and adaptation in Iris leaves. Accumulation of Na~+ and K~+, proline concentration and improvement of cell membrane penetration were conducted to adjusted very rapidly to salt and alkali content during each month. With the accumulation osmotic substances, water potentials and osmotic potentials in Iris leaves decreased, cell turgid pressure increased or maintained to preserve cell, leaf and then whole plant growth under salt and drought condition.2. The results showed that the ABA contents in leaves after salt and alkali treats increased markedly. The addition of exogenous ABA decreased the increment of MDA and H2O2 contents and retarded the attenuation of CAT activity, which were induced by salt and alkali treats, and enhanced the GR activity. The content of leaf ABA increased under long-term salt and alkali stress, but root ABA differed among plants. However, in corn roots which possess a well-developed exodermis, alkaline and saline conditions in the rhizosphere did not reduce the endogenous ABA concentration, because the leaching of ABA from corn roots into the rhizosphere was lower than that from Iris and soybean roots. ABA efflux from corn and Iris and soybeab roots into the soil solution was observed only during the first days of the experiments and thereafter became substantially decreased.3. The data acquired in this study showed that diurnal changes of Pn and Tr were two-peak-type-curve, and was marked midday depression of Pn, which could be fully accounted for Gs changes. Carboxylic efficiency(CE) photosynthetic rate at saturation for intercellular CO2 concentration(A_350) apparent quantum yield(AQY) and photosynthetic rate at light saturation(Pmax) decreased in the midday. The result demonstrated that midday depression could be attributed to leaf temperature rising caused by the increasing of strong solar radiation (PAR).4. The results showed that Pn decreased with leaf ripping, and CO_2 carboxylic efficiency (CE), apparent quantum yield(AQY) declined at same time, but maximum photochemical efficiency(Fv/Fm) had little changed during leaf senescence of Iris lactea Pall, var chinensis(Fisch.) Koidz. During senescence of Iris lactea Pall, var chinensis(Fisch.) Koidz, CE and A_350 (net photosynthetic rate at light saturation) declined more rapid than AQY and J_max (net photosynthetic rate at CO_2 saturation (net photosynthetic rate at light saturation). This result showed that dark reaction or carbon cycle of photosynthesis declined more rapid than light reaction.5. The data acquired in this study showed that the diurnal changes of the maximum PS II photochemical efficiency (F_v/F_m) and actual photochemical efficiency of PS II (OPS II) were related to light intensity, the functional photosystem II reaction centers were decreased to cause significant photoinhibition. With the increasing of photosynthetic actual irradiation (PAR) to simulate
PAR diurnal change, Fv/Fm and OPS II decreased, nonphotochemical quenching (NPQ) increased, and the fraction of excited energy allocated to PS II increased to enhance PS II burden carrying excited energy. These results indicated that photoinhibition in Iris leaves are caused primarily by increasing of excited energy allocated to PS II to enhance reversible inactivation of PS II centers except NPQ increasing. With the decreasing of PAR, Fv/Fm and OPS II increased, NPQ and unbalance index of excited energy allocated to PS II and PS I (p/a-1) declined respectively. This indicated that the recovery of PS II photochemical efficiency is caused primarily by renewing PS II activation owing to unloading the energy burden of PS II.6. Salt and alkali treatment (the content of salt 0.36%,pH 8.90) alone had no effect on the maximal photochemistry of PS II (Fv/Fm), but decreased apparent photosynthetic electron transport rate(ETR).However, the salt and alkali treatment accompanied with drought decreased Fv/Fm and the efficiency of excitation captured open PS II(Fv'/Fm'), and state transition (from II to I) caused 33.3% decrease in allocation of excited energy to PS I and 18.1% increase to PS II. At the same time, the salt and alkali treatment accompanied with drought resulted in serious imbalance of excited energy distributions between two photosystems, enhanced the pressure on PS II .It is suggested that Nonphotochemical quenching (NPQ) could play a key role in adaptation mechanism of plant to stress.7. NaCl treatment alone had no effect on the maximal photochemistry of PSII. However, the NaCl treatment modified pH stress on PSII photochemistry in Iris leaves, which was manifested by a lesser pH-induced decrease in photochemical quenching (qP), efficiency of excitation energy capture by open PSII reaction centers (Fv7Fm'), and quantum yield of PSII electron transport (OPSII). Additions of calcium and ABA could alleviate photoinhibition in Iris leaves treated by NaCl. But Additions of calcium has no effects on alleviating under alkaline stress, and ABA does.
引文
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