小偃54和8602及其杂交后代小偃41和小偃81旗叶光合特性及光破坏防御能力比较
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摘要
小麦旗叶的光合速率及光合有效时间是影响小麦产量的重要因素之一。大田生产中,外在的环境因素很难控制,因此,通过栽培和育种等手段改善叶片自身的光合特性和光破坏防御能力,是提高其光合速率,延长其光合有效时间的有效途径之一。
本研究通过测定叶绿体色素含量、叶片气体交换参数、叶绿素荧光参数、820nm吸收参数、抗氧化酶活性、D1蛋白及PsaA蛋白的含量,比较了小偃54(XY54)和8602及其杂交后代小偃41(XY41)和小偃81(XY81)花后旗叶光合特性及光破坏防御能力的差异。主要结果如下:
(1)在开花期,4个品种(系)的叶绿素含量不同,基本表现为杂交后代高于双亲,且以XY81为最高;8602品系的光合速率(Pn)和羧化效率(CE)最高,XY41品系的气孔导度(Gs)最低,而XY54品种的CE最低。4个小麦品种(系)的胞间CO2浓度(Ci)、K点可变荧光(F_K)占F_J–F_O荧光振幅的比率(Wk)、光系统II(PSII)最大量子效率(Fv/Fm)、电子传递到Q_A下游的概率(Ψo)、电子从系统间电子传递体传递给光系统I受体侧电子受体的概率(δRo)、820nm相对可变透射光(ΔI/Io)及PSII的实际光化学效率(ΦPSII)无显著差异。小麦开花后,XY54的叶绿素含量、Pn、Gs、CE、F_v/F_m、δRo及ΔI/Io的下降幅度和Wk的上升幅度显著大于其他3个品系,但Ci、Ψo和ΦPSII与其他3个品系没有显著差异。8602和两个杂交后代的上述参数的变化趋势在整个发育期内基本相同。当叶绿素含量相同时,XY54的Pn相对其他3个品系较高。在灌浆后期,4个小麦品种的光合速率及光合光反应的日变化基本相同。以上结果表明,与XY54相比,XY41和XY81的光合速率下降幅度较小,可能与他们的光合暗反应能力不同有关,而不是由于三者的叶绿素含量、气孔导度及光反应的能力不同所造成,杂交后代XY81和XY41可能遗传了8602品系较高的光合暗反应特性。
(2)在开花期,两个杂交后代XY81和XY41的类胡萝卜素含量高于8602,但与XY54基本相同;4个品种(系)的ΦPSII/ΦCO_2、非光化学猝灭(NPQ)、D1蛋白和PsaA蛋白的含量、过氧化氢酶(CAT)和抗坏血酸过氧化物酶(APX)的活性都没有显著差异;XY54超氧化物歧化酶(SOD)的活性较其他3个品系低。小麦开花后,XY81和XY41类胡萝卜素含量的下降速率较XY54慢,但与8602类似;4个品种(系)的NPQ逐渐增加,增加幅度基本相同;XY54的SOD活性的绝对值在花后的整个衰老过程中低于其他3个品系,且上升幅度显著高于其他3个品系;APX活性的下降幅度却比其他3个品系大;4个品种(系)CAT的活性的变化基本相同。在开花后期,与其他3个品系相比,XY54的ΦPSII/ΦCO_2增加的幅度更快,D1蛋白和PsaA蛋白的降解幅度加快。以上结果表明,两个杂交后代XY81和XY41可能遗传了其亲本XY54较高的类胡萝卜素含量,但是其类胡萝卜素的降解却与8602的某些性状相关。在开花后期,XY54依赖光抑制的热耗散的比例可能更高。小偃54的抗氧化酶活性相对其他3个品系较低,对超氧阴离子及过氧化氢的清除能力比较弱,从而加重了对XY54的光破坏。
Photosyntheis and its active period are important factors determing wheat production. In wheat production, it is hard to control the external conditions that affect photosynthesis. However, it is feasible to enhance photosynthesis and extend its active period by perfecting the photosynthetic characteristics and photoprotection capacity of wheat leaves through cultivating and breeding.
In this study, gas exchange, chlorophyll fluorescence, 820nm transmission, pigments, antioxidant enzymes, D1 protein and PsaA protein were assayed to compare photosynthetic characteristics and capacity of photoprotection of flag leaves of Xiaoyan 54 (XY54) and 8602, and their hybrids, Xiaoyan 41 (XY41) and Xiaoyan 81 (XY81) after anthesis. In this way, we expect to clear the differences in photosynthetic characteristics and photoprotection capacity between the parients and their hybrids, and then provide some valuable information for wheat breeding. The main results are as follows:
(1) At the period of anthesis, there were some differences among these four cultivars (lines) in the content of chlorophylls. The pigment contents were higher in the hybrids, especially XY81, than their parents. Photosynthetic rate (Pn) and carboxylation efficiency (CE) were the highest in 8602 whereas stomatal conductance (Gs) was lower in XY41 and CE was the lowest in XY54. These four cultivars (lines) had similar intercellular CO2 concentration (Ci), ratio of variable F_K to the amplitude F_J–F_O (Wk), the maximum quantum yield of PSII (F_v/F_m), efficiency that an electron moves further than QA (Ψo), efficiency with which an electron from the intersystem electron carriers moves to reduce end electron acceptors at the PSI acceptor side (δRo), relative variable transmission at 820nm (ΔI/Io), and actually photochemical efficiency of PSII (ΦPSII). After anthesis, decreases in the pigment contents, Pn, CE, Gs, Fv/Fm,δRo andΔI/Io, and increases in Wk were faster in XY54 in contrast with other three cultivars (lines). No difference was detected in Ci,Ψo, orΦPSII among these four cultivars (lines). The change of above parameters were similar in 8602 and the hybrids during the whole period after anthesis. With similar chlorophyll contents, Pn was higher in XY54 in contrast with other three cultivars (lines). At late filling period, the four cultivars (lines) exhibited identically diurnal change in Pn and photosynthetic light reactions. The above results indicate that, compared with XY 54, the relatively slower decrease in photosynthetic rate in XY41 and XY81 was not correlated with their chlorophyll contents, stomatal conductance and photosynthetic light reaction capacities, but with their different dark reaction capacities. XY41 and XY81 may inherit the high dark reaction capacity from 8602, one of their parents.
(2) At the period of anthesis, the content of carotenoids in XY81 and XY41 was similar as that in XY54, but higher than that in 8602. No differences were found among the four cultivars (lines) in terms of ratio ofΦPSII/ΦCO_2, non-photochemical quenching (NPQ), the content of D1 protein and PsaA protein, and the activity of catalase (CAT) or ascorbate peroxidase (APX). However, the activity of superoxide dismutase (SOD) was higher in XY54 compared with other three cultivars (lines). After anthesis, the content of carotenoids in XY81 and XY41 decreased slower than that in XY54, but identically with that in 8602. NPQ increased similarly in the four cultivars (generations). The activity of SOD increased whereas that of APX decreased faster in XY54 in contrast with those three generations. CAT showed a similar activity in the four cultivars (lines) after anthesis. At the late stage after anthesis, compared with those three cultivars (lines), XY54 had a faster increase in ratio ofΦPSII/ΦCO_2 and more pronounced degradation of D1 protein and PsaA protein. The above results suggest that the two hybrids might inherbit the higher carotenoids contents of XY54, but their degradation of carotenoids was correlated with the genetic characteristics from 8602. At the late stage after anthesis, heat dissipation depedent on photoinhibition was more predominant in XY54. XY54 might enhance its Mehler reaction to consume more absorbed light energy. During Mehler reaction, more produced superoxide anion was converted to hydrogen peroxide by SOD in XY54. Nevertheless, such accumulated hydrogen peroxide was not scavenged timely by APX. As a result, the photodamage became more severe in XY54.
本研究通过测定叶绿体色素含量、叶片气体交换参数、叶绿素荧光参数、820nm吸收参数、抗氧化酶活性、D1蛋白及PsaA蛋白的含量,比较了小偃54(XY54)和8602及其杂交后代小偃41(XY41)和小偃81(XY81)花后旗叶光合特性及光破坏防御能力的差异。主要结果如下:
(1)在开花期,4个品种(系)的叶绿素含量不同,基本表现为杂交后代高于双亲,且以XY81为最高;8602品系的光合速率(Pn)和羧化效率(CE)最高,XY41品系的气孔导度(Gs)最低,而XY54品种的CE最低。4个小麦品种(系)的胞间CO2浓度(Ci)、K点可变荧光(F_K)占F_J–F_O荧光振幅的比率(Wk)、光系统II(PSII)最大量子效率(Fv/Fm)、电子传递到Q_A下游的概率(Ψo)、电子从系统间电子传递体传递给光系统I受体侧电子受体的概率(δRo)、820nm相对可变透射光(ΔI/Io)及PSII的实际光化学效率(ΦPSII)无显著差异。小麦开花后,XY54的叶绿素含量、Pn、Gs、CE、F_v/F_m、δRo及ΔI/Io的下降幅度和Wk的上升幅度显著大于其他3个品系,但Ci、Ψo和ΦPSII与其他3个品系没有显著差异。8602和两个杂交后代的上述参数的变化趋势在整个发育期内基本相同。当叶绿素含量相同时,XY54的Pn相对其他3个品系较高。在灌浆后期,4个小麦品种的光合速率及光合光反应的日变化基本相同。以上结果表明,与XY54相比,XY41和XY81的光合速率下降幅度较小,可能与他们的光合暗反应能力不同有关,而不是由于三者的叶绿素含量、气孔导度及光反应的能力不同所造成,杂交后代XY81和XY41可能遗传了8602品系较高的光合暗反应特性。
(2)在开花期,两个杂交后代XY81和XY41的类胡萝卜素含量高于8602,但与XY54基本相同;4个品种(系)的ΦPSII/ΦCO_2、非光化学猝灭(NPQ)、D1蛋白和PsaA蛋白的含量、过氧化氢酶(CAT)和抗坏血酸过氧化物酶(APX)的活性都没有显著差异;XY54超氧化物歧化酶(SOD)的活性较其他3个品系低。小麦开花后,XY81和XY41类胡萝卜素含量的下降速率较XY54慢,但与8602类似;4个品种(系)的NPQ逐渐增加,增加幅度基本相同;XY54的SOD活性的绝对值在花后的整个衰老过程中低于其他3个品系,且上升幅度显著高于其他3个品系;APX活性的下降幅度却比其他3个品系大;4个品种(系)CAT的活性的变化基本相同。在开花后期,与其他3个品系相比,XY54的ΦPSII/ΦCO_2增加的幅度更快,D1蛋白和PsaA蛋白的降解幅度加快。以上结果表明,两个杂交后代XY81和XY41可能遗传了其亲本XY54较高的类胡萝卜素含量,但是其类胡萝卜素的降解却与8602的某些性状相关。在开花后期,XY54依赖光抑制的热耗散的比例可能更高。小偃54的抗氧化酶活性相对其他3个品系较低,对超氧阴离子及过氧化氢的清除能力比较弱,从而加重了对XY54的光破坏。
Photosyntheis and its active period are important factors determing wheat production. In wheat production, it is hard to control the external conditions that affect photosynthesis. However, it is feasible to enhance photosynthesis and extend its active period by perfecting the photosynthetic characteristics and photoprotection capacity of wheat leaves through cultivating and breeding.
In this study, gas exchange, chlorophyll fluorescence, 820nm transmission, pigments, antioxidant enzymes, D1 protein and PsaA protein were assayed to compare photosynthetic characteristics and capacity of photoprotection of flag leaves of Xiaoyan 54 (XY54) and 8602, and their hybrids, Xiaoyan 41 (XY41) and Xiaoyan 81 (XY81) after anthesis. In this way, we expect to clear the differences in photosynthetic characteristics and photoprotection capacity between the parients and their hybrids, and then provide some valuable information for wheat breeding. The main results are as follows:
(1) At the period of anthesis, there were some differences among these four cultivars (lines) in the content of chlorophylls. The pigment contents were higher in the hybrids, especially XY81, than their parents. Photosynthetic rate (Pn) and carboxylation efficiency (CE) were the highest in 8602 whereas stomatal conductance (Gs) was lower in XY41 and CE was the lowest in XY54. These four cultivars (lines) had similar intercellular CO2 concentration (Ci), ratio of variable F_K to the amplitude F_J–F_O (Wk), the maximum quantum yield of PSII (F_v/F_m), efficiency that an electron moves further than QA (Ψo), efficiency with which an electron from the intersystem electron carriers moves to reduce end electron acceptors at the PSI acceptor side (δRo), relative variable transmission at 820nm (ΔI/Io), and actually photochemical efficiency of PSII (ΦPSII). After anthesis, decreases in the pigment contents, Pn, CE, Gs, Fv/Fm,δRo andΔI/Io, and increases in Wk were faster in XY54 in contrast with other three cultivars (lines). No difference was detected in Ci,Ψo, orΦPSII among these four cultivars (lines). The change of above parameters were similar in 8602 and the hybrids during the whole period after anthesis. With similar chlorophyll contents, Pn was higher in XY54 in contrast with other three cultivars (lines). At late filling period, the four cultivars (lines) exhibited identically diurnal change in Pn and photosynthetic light reactions. The above results indicate that, compared with XY 54, the relatively slower decrease in photosynthetic rate in XY41 and XY81 was not correlated with their chlorophyll contents, stomatal conductance and photosynthetic light reaction capacities, but with their different dark reaction capacities. XY41 and XY81 may inherit the high dark reaction capacity from 8602, one of their parents.
(2) At the period of anthesis, the content of carotenoids in XY81 and XY41 was similar as that in XY54, but higher than that in 8602. No differences were found among the four cultivars (lines) in terms of ratio ofΦPSII/ΦCO_2, non-photochemical quenching (NPQ), the content of D1 protein and PsaA protein, and the activity of catalase (CAT) or ascorbate peroxidase (APX). However, the activity of superoxide dismutase (SOD) was higher in XY54 compared with other three cultivars (lines). After anthesis, the content of carotenoids in XY81 and XY41 decreased slower than that in XY54, but identically with that in 8602. NPQ increased similarly in the four cultivars (generations). The activity of SOD increased whereas that of APX decreased faster in XY54 in contrast with those three generations. CAT showed a similar activity in the four cultivars (lines) after anthesis. At the late stage after anthesis, compared with those three cultivars (lines), XY54 had a faster increase in ratio ofΦPSII/ΦCO_2 and more pronounced degradation of D1 protein and PsaA protein. The above results suggest that the two hybrids might inherbit the higher carotenoids contents of XY54, but their degradation of carotenoids was correlated with the genetic characteristics from 8602. At the late stage after anthesis, heat dissipation depedent on photoinhibition was more predominant in XY54. XY54 might enhance its Mehler reaction to consume more absorbed light energy. During Mehler reaction, more produced superoxide anion was converted to hydrogen peroxide by SOD in XY54. Nevertheless, such accumulated hydrogen peroxide was not scavenged timely by APX. As a result, the photodamage became more severe in XY54.
引文
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