光系统Ⅱ蛋白磷酸化在植物光破坏防御中的作用
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摘要
光合作用是地球上最重要的化学反应 是唯一可以把太阳光能转变成生物
体所需化学能的过程 是地球上多种多样生命活动的能量基础 尽管太阳光能
是植物光合作用的原初推动力 但过多的光能也会造成光合效率降低 即发生
光抑制 在其它胁迫条件同时存在时 强光对植物光合作用的抑制作用更加明
显 甚至导致光合机构的破坏 光系统 II PSII 复合体是光破坏的首发部位
在长期的进化过程中 植物演化形成了一系列防御光破坏的机制 保护光合机
构免受光破坏 我们的研究表明 PSII 蛋白的磷酸化在光破坏防御中具有重要
的保护作用
一 大豆叶片 D1 蛋白磷酸化/去磷酸化本身不影响 PSII 电子传递
活性
为了探讨 PSII 功能和 D1 蛋白磷酸化之间的关系 在利用蛋白激酶抑制剂
FSBA 处理并改变大豆叶片中 PSII 核心组分 D1 蛋白的磷酸化水平之后 观测
了叶片叶绿素荧光参数 PSII 电子传递活性和 D1 蛋白含量的变化 同时 利
用 SDS-聚丙烯酰胺凝胶电泳和蛋白免疫印迹的方法分析磷酸化的 D1* 和非
磷酸化的 D1 D1 蛋白 得到如下结果 1 在暗适应的大豆叶片中存在高
水平磷酸化的 D1 蛋白 2 小时的 FSBA 1mmol/L 处理可以使磷酸化的 D1
蛋白全部去磷酸化 2 磷酸化 D1 蛋白的去磷酸化没有造成 D1 蛋白的净损失
也没有引起大豆叶片叶绿素荧光参数的明显变化 3 PSII 的电子传递活性
(H2O → 1,4-BQ) 没有因为磷酸化 D1 蛋白的完全去磷酸化而发生明显的变化
根据以上结果得出结论 大豆叶片 PS II 复合体核心组分 D1 蛋白的磷酸化/去
磷酸化本身并不影响 PSII 反应中心的功能
III
光系统 II 蛋白磷酸化在植物光破坏防御中的作用
二 大豆和南瓜叶片 PSII 可逆失活的不同机制
通过对饱和光响应的比较研究 探讨大豆和南瓜叶片光破坏防御机制的差
异 一方面 大豆和南瓜叶片在饱和光下产生一些相同的变化
1 经过 3 小
时饱和光处理 它们的初始叶绿素荧光参数 Fo 都明显升高 而 PSII 光化学效
率 Fv/Fm 都明显降低 但经过随后的 3 小时暗恢复后 Fo 和 Fv/Fm 都可以恢
复到初始暗对照的水平 2 在饱和光处理和随后的暗恢复过程中都没有发生
D1 蛋白的净损失 另一方面 经过饱和光处理 3 小时后 大豆和南瓜叶片还发
生一些不同的变化 1 大豆叶片的低温荧光参数 F685 和 F685/F735 都明显降
低 但南瓜叶片的这两个参数却没有明显的变化2 大豆叶片的部分 LHCII
从 PSII 核心复合体上脱离 但南瓜叶片却没有发生 LHCII 的脱离 3 在饱和
光下测定的南瓜类囊体 PSII 电子传递活性明显降低 而大豆类囊体 PSII 电子
传递活性没有明显变化 此外 在经过充分暗适应的大豆叶片检测到部分 D1
蛋白处于磷酸化状态 而在充分暗适应的南瓜叶片却没有检测到磷酸化的 D1
蛋白 以上结果显示 大豆和南瓜叶片对光破坏具有相同的防御策略----PSII
可逆失活 但却涉及不同的机制 大豆叶片的 PSII 失活涉及 LHCII 从 PSII 核
心复合体脱离 而南瓜叶片的 PSII 失活不涉及 LHCII 的脱离
三 LHCII 脱离保护 PSII 反应中心免受光破坏
通过在饱和光下蛋白激酶抑制剂 FSBA 处理 研究了 LHCII 脱离在 PSII
光破坏防御中的作用 获得如下结果 1 饱和光照明可以引起部分 LHCII 从
PSII 核心复合体上脱离 但 FSBA 处理可以抑制 LHCII 脱离 2 大豆叶片经
过 3 小时饱和光处理后 初始荧光 Fo 明显升高 而 PSII 光化学效率 Fv/Fm 显
著降低 但经过随后 3 小时暗恢复后 Fo 和 Fv/Fm 基本可以恢复到初始暗对照
的水平 而 FSBA 处理增大它们的变化程度 并且放置暗中 3 小时后不能恢复
3 在没有 FSBA 存在的情况下 饱和光照明对 PSII 电子传递活性和 D1 蛋白
含量没有明显的影响 但是在 FSBA 存在时 3 小时饱和光照明引起 PSII 电子
传递活性和 D1 蛋白含量的明显降低 以类囊体为实验材料进行的体外实验得
出类似的结果 4 饱和光照明可以诱导大豆叶片 PSII 核心蛋白 D1 D2 和
IV
光系统 II 蛋白磷酸化在植物光破坏防御中的作用
CP43磷酸化水平的升高 但引起LHCII蛋白磷酸化水平的降低 饱和光下 FSBA
可以抑制 D1 D2 和 CP43 的蛋白磷酸化 这些结果表明 在饱和光下 LHCII
脱离可以保护 PSII 反应中心免受光破坏 而且 PSII 核心蛋白的磷酸化可能在
LHCII 脱离过程中发挥重要的作用
Photosynthesis, which is the only way to convert the sunlight energy to the
chemical energy to maintain all living organisms, is the most important chemical
reaction on earth. Even though sunlight is the ultimate energy source for the
photosynthesis, it can also harm plants. The efficiency of photosynthesis can be
significantly reduced in plants when exposed to high light. This phenomenon of
decline in photosynthetic efficiency is referred as photoinhibition. Particularly under
environmental stress, such as low temperature and water deficiency, too much light
can cause the photodamage to the photosynthetic apparatus and the primary target
for the damage is photosystem II. However, plants have evolved an array of
protective mechanisms against photodamage. Our experimental results indicate that
PSII protein phosphorylation plays an important role in protecting PSII from
photodamage.
1. D1 protein phosphorylation/dephosphorylation alone has no
effect on the electron transport activity of photosystem II in soybean
leaves
In order to explore the relationship between the phosphorylation of D1 protein
and the function of PSII, the changes in chlorophyll fluorescence parameters, PSII
electron transport activity and D1 protein amount were observed after
dephosphorylation of phosphorylated D1 proteins caused by FSBA
(5’-p-fluorosulfonylbenzoyl adenosine, an inhibitor of protein kinase) treatment.
VI
光系统 II 蛋白磷酸化在植物光破坏防御中的作用
Also, phosphorylated and non-phosphorylated D1 proteins (D1* and D1,
respectively) were analyzed by sodium dodecyl sulfate-polyacrylamide gel
electrophoresis (SDS-PAGE) and Western Blotting. The following results were
obtained. (1) A high level of phosphorylated D1 protein (D1*) was observed in
dark-adapted soybean leaves and D1* could be completely dephosphorylated by
FSBA (1 mmol/L) treatment for 2 h. (2) Dephosphorylation of D1* resulted in
neither substantial net loss of D1 proteins nor the significant changes in value of
chlorophyll fluorescence parameters. (3) The electron transport activity of PS II
(H2O → 1,4-BQ) was not changed significantly after D1* was completely
dephosphorylated. Based on these results, it is concluded that D1 protein
phosphorylation/dephosphorylation alone has no significant effect on the function of
PS II reaction center in soybean leaves.
2. Different mechanisms for Photosystem II reversible inactivation
in pumpkin and soybean leaves at saturating light
A comparative investigation on the response to saturating light was made
between pumpkin and soybean leaves. Some similar responses were observed in the
both species. (1) After saturating illumination for 3 h, the original fluorescence Fo
increased while the PSII photochemical efficiency Fv/Fm declined significantly, but
these parameters could largely recover to the levels of dark-adapted leaves after 3 h
of subsequent dark recovery. (2) No net loss of the D1 proteins occurred after the
saturating illumination. However, soybean and pumpkin leaves also had some
different responses to the saturating illumination. (1) Low temperature fluorescence
parameters, F685 and F685/F735 decreased significantly in soybean but not in
pumpkin leaves. (2) Part of LHCII dissociated from PSII complexes in soybean
leaves but not in pumpkin leaves, as shown by the results of sucrose density gradient
centrifugation and the SDS-PAGE. (3) The light-saturated PSII electron transport
activity declined significantly in pumpkin thylakoids but not in soybean thylakoids.
In addition, a high level of phosphorylated D1 proteins was found in dark-adapted
soybean leaves but not in dark-adapted pumpkin leaves. These results may imply
that at excessive light soybean and pumpkin have the same protective strategy
against photodamage, reversible inactivation of PSII, but two different mechanisms,
namely the reversible inactivation is related to the dissociation of LHCII in soybean
but not in pumpkin leaves.
体所需化学能的过程 是地球上多种多样生命活动的能量基础 尽管太阳光能
是植物光合作用的原初推动力 但过多的光能也会造成光合效率降低 即发生
光抑制 在其它胁迫条件同时存在时 强光对植物光合作用的抑制作用更加明
显 甚至导致光合机构的破坏 光系统 II PSII 复合体是光破坏的首发部位
在长期的进化过程中 植物演化形成了一系列防御光破坏的机制 保护光合机
构免受光破坏 我们的研究表明 PSII 蛋白的磷酸化在光破坏防御中具有重要
的保护作用
一 大豆叶片 D1 蛋白磷酸化/去磷酸化本身不影响 PSII 电子传递
活性
为了探讨 PSII 功能和 D1 蛋白磷酸化之间的关系 在利用蛋白激酶抑制剂
FSBA 处理并改变大豆叶片中 PSII 核心组分 D1 蛋白的磷酸化水平之后 观测
了叶片叶绿素荧光参数 PSII 电子传递活性和 D1 蛋白含量的变化 同时 利
用 SDS-聚丙烯酰胺凝胶电泳和蛋白免疫印迹的方法分析磷酸化的 D1* 和非
磷酸化的 D1 D1 蛋白 得到如下结果 1 在暗适应的大豆叶片中存在高
水平磷酸化的 D1 蛋白 2 小时的 FSBA 1mmol/L 处理可以使磷酸化的 D1
蛋白全部去磷酸化 2 磷酸化 D1 蛋白的去磷酸化没有造成 D1 蛋白的净损失
也没有引起大豆叶片叶绿素荧光参数的明显变化 3 PSII 的电子传递活性
(H2O → 1,4-BQ) 没有因为磷酸化 D1 蛋白的完全去磷酸化而发生明显的变化
根据以上结果得出结论 大豆叶片 PS II 复合体核心组分 D1 蛋白的磷酸化/去
磷酸化本身并不影响 PSII 反应中心的功能
III
光系统 II 蛋白磷酸化在植物光破坏防御中的作用
二 大豆和南瓜叶片 PSII 可逆失活的不同机制
通过对饱和光响应的比较研究 探讨大豆和南瓜叶片光破坏防御机制的差
异 一方面 大豆和南瓜叶片在饱和光下产生一些相同的变化
1 经过 3 小
时饱和光处理 它们的初始叶绿素荧光参数 Fo 都明显升高 而 PSII 光化学效
率 Fv/Fm 都明显降低 但经过随后的 3 小时暗恢复后 Fo 和 Fv/Fm 都可以恢
复到初始暗对照的水平 2 在饱和光处理和随后的暗恢复过程中都没有发生
D1 蛋白的净损失 另一方面 经过饱和光处理 3 小时后 大豆和南瓜叶片还发
生一些不同的变化 1 大豆叶片的低温荧光参数 F685 和 F685/F735 都明显降
低 但南瓜叶片的这两个参数却没有明显的变化2 大豆叶片的部分 LHCII
从 PSII 核心复合体上脱离 但南瓜叶片却没有发生 LHCII 的脱离 3 在饱和
光下测定的南瓜类囊体 PSII 电子传递活性明显降低 而大豆类囊体 PSII 电子
传递活性没有明显变化 此外 在经过充分暗适应的大豆叶片检测到部分 D1
蛋白处于磷酸化状态 而在充分暗适应的南瓜叶片却没有检测到磷酸化的 D1
蛋白 以上结果显示 大豆和南瓜叶片对光破坏具有相同的防御策略----PSII
可逆失活 但却涉及不同的机制 大豆叶片的 PSII 失活涉及 LHCII 从 PSII 核
心复合体脱离 而南瓜叶片的 PSII 失活不涉及 LHCII 的脱离
三 LHCII 脱离保护 PSII 反应中心免受光破坏
通过在饱和光下蛋白激酶抑制剂 FSBA 处理 研究了 LHCII 脱离在 PSII
光破坏防御中的作用 获得如下结果 1 饱和光照明可以引起部分 LHCII 从
PSII 核心复合体上脱离 但 FSBA 处理可以抑制 LHCII 脱离 2 大豆叶片经
过 3 小时饱和光处理后 初始荧光 Fo 明显升高 而 PSII 光化学效率 Fv/Fm 显
著降低 但经过随后 3 小时暗恢复后 Fo 和 Fv/Fm 基本可以恢复到初始暗对照
的水平 而 FSBA 处理增大它们的变化程度 并且放置暗中 3 小时后不能恢复
3 在没有 FSBA 存在的情况下 饱和光照明对 PSII 电子传递活性和 D1 蛋白
含量没有明显的影响 但是在 FSBA 存在时 3 小时饱和光照明引起 PSII 电子
传递活性和 D1 蛋白含量的明显降低 以类囊体为实验材料进行的体外实验得
出类似的结果 4 饱和光照明可以诱导大豆叶片 PSII 核心蛋白 D1 D2 和
IV
光系统 II 蛋白磷酸化在植物光破坏防御中的作用
CP43磷酸化水平的升高 但引起LHCII蛋白磷酸化水平的降低 饱和光下 FSBA
可以抑制 D1 D2 和 CP43 的蛋白磷酸化 这些结果表明 在饱和光下 LHCII
脱离可以保护 PSII 反应中心免受光破坏 而且 PSII 核心蛋白的磷酸化可能在
LHCII 脱离过程中发挥重要的作用
Photosynthesis, which is the only way to convert the sunlight energy to the
chemical energy to maintain all living organisms, is the most important chemical
reaction on earth. Even though sunlight is the ultimate energy source for the
photosynthesis, it can also harm plants. The efficiency of photosynthesis can be
significantly reduced in plants when exposed to high light. This phenomenon of
decline in photosynthetic efficiency is referred as photoinhibition. Particularly under
environmental stress, such as low temperature and water deficiency, too much light
can cause the photodamage to the photosynthetic apparatus and the primary target
for the damage is photosystem II. However, plants have evolved an array of
protective mechanisms against photodamage. Our experimental results indicate that
PSII protein phosphorylation plays an important role in protecting PSII from
photodamage.
1. D1 protein phosphorylation/dephosphorylation alone has no
effect on the electron transport activity of photosystem II in soybean
leaves
In order to explore the relationship between the phosphorylation of D1 protein
and the function of PSII, the changes in chlorophyll fluorescence parameters, PSII
electron transport activity and D1 protein amount were observed after
dephosphorylation of phosphorylated D1 proteins caused by FSBA
(5’-p-fluorosulfonylbenzoyl adenosine, an inhibitor of protein kinase) treatment.
VI
光系统 II 蛋白磷酸化在植物光破坏防御中的作用
Also, phosphorylated and non-phosphorylated D1 proteins (D1* and D1,
respectively) were analyzed by sodium dodecyl sulfate-polyacrylamide gel
electrophoresis (SDS-PAGE) and Western Blotting. The following results were
obtained. (1) A high level of phosphorylated D1 protein (D1*) was observed in
dark-adapted soybean leaves and D1* could be completely dephosphorylated by
FSBA (1 mmol/L) treatment for 2 h. (2) Dephosphorylation of D1* resulted in
neither substantial net loss of D1 proteins nor the significant changes in value of
chlorophyll fluorescence parameters. (3) The electron transport activity of PS II
(H2O → 1,4-BQ) was not changed significantly after D1* was completely
dephosphorylated. Based on these results, it is concluded that D1 protein
phosphorylation/dephosphorylation alone has no significant effect on the function of
PS II reaction center in soybean leaves.
2. Different mechanisms for Photosystem II reversible inactivation
in pumpkin and soybean leaves at saturating light
A comparative investigation on the response to saturating light was made
between pumpkin and soybean leaves. Some similar responses were observed in the
both species. (1) After saturating illumination for 3 h, the original fluorescence Fo
increased while the PSII photochemical efficiency Fv/Fm declined significantly, but
these parameters could largely recover to the levels of dark-adapted leaves after 3 h
of subsequent dark recovery. (2) No net loss of the D1 proteins occurred after the
saturating illumination. However, soybean and pumpkin leaves also had some
different responses to the saturating illumination. (1) Low temperature fluorescence
parameters, F685 and F685/F735 decreased significantly in soybean but not in
pumpkin leaves. (2) Part of LHCII dissociated from PSII complexes in soybean
leaves but not in pumpkin leaves, as shown by the results of sucrose density gradient
centrifugation and the SDS-PAGE. (3) The light-saturated PSII electron transport
activity declined significantly in pumpkin thylakoids but not in soybean thylakoids.
In addition, a high level of phosphorylated D1 proteins was found in dark-adapted
soybean leaves but not in dark-adapted pumpkin leaves. These results may imply
that at excessive light soybean and pumpkin have the same protective strategy
against photodamage, reversible inactivation of PSII, but two different mechanisms,
namely the reversible inactivation is related to the dissociation of LHCII in soybean
but not in pumpkin leaves.
引文
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