镧对UV-B辐射胁迫下大豆幼苗内源激素与类黄酮代谢影响
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摘要
臭氧层衰减诱发UV-B辐射抵达地球表面增加,对陆生植物的影响引起世人广泛关注。探讨稀土对UV-B辐射胁迫伤害植物的影响与机理,是稀土环境植物学研究的拓展和延伸,其结果可为减轻UV-B辐射伤害植物的防护对策提供参考。本文采用水培方法,以油料作物大豆(Glycine max)为实验材料,以生理学、生物化学及物理化学方法为研究手段;以La(III)对UV-B辐射胁迫下大豆幼苗生长、内源激素与芳香族氨基酸含量、类黄酮代谢关键酶的影响与机理为内容;从环境生态角度,生理生化及酶分子水平,揭示La(III)对UV-B辐射胁迫下植物内源激素和类黄酮代谢的影响与机理;为科学评价稀土对UV-B辐射胁迫伤害植物的影响;丰富UV-B辐射生态学理论,提供参考,主要研究
结果如下:
(1) UV-B辐射胁迫下,大豆幼苗根系活力下降,抑制了营养元素吸收,破坏叶片形态,降低叶片光合作用,最终导致整个植物干物质累积量降低,此效应随UV-B辐射胁迫剂量增加而加剧。La(III)(20 mg/L)能提高大豆根系活力、叶绿素含量、矿质元素含量,进而促进光合作用,提高叶片叶面积及大豆幼苗鲜(干)重,促进大豆幼苗的生长。La(III)(20 mg/L)的介入使逆境中的大豆幼苗也能尽量维持对光能的吸收和利用,从而满足植物自身恢复所需的能量和物质,缓解UV-B辐射对叶片损伤。高剂量La(III)(60 mg/L)降低大豆根系活力、叶绿素含量、矿质元素含量,进而抑制光合作用,降低叶片叶面积及大豆幼苗鲜(干)重,抑制大豆幼苗的生长。La(III)(60 mg/L)+UV-B辐射胁迫处理对大豆幼苗起到了协同抑制作用。La(III)(60 mg/L)的介入加剧了植物自身恢复所需能量和物质的减少,使逆境中的大豆幼苗表现出受到复合胁迫伤害的作用。适量La(III)可缓解UV-B辐射胁迫对大豆生长的抑制作用,而高剂量La(III)可加剧UV-B辐射胁迫对大豆生长的抑制作用。
(2) UV-B辐射胁迫下,大豆幼苗IAA、GA和CTK含量下降,ABA、ETH含量和IAAO活性增加,器官生长发育减缓、形态结构改变、生物量下降,此效应随UV-B辐射胁迫剂量增加而加剧。La(III)(20 mg/L)能提高UV-B辐射胁迫下大豆幼苗IAA、GA和CTK含量、降低ABA含量、IAAO活性和乙烯含量,减轻UV-B辐射胁迫对植物生长的抑制,缓解植株生长与类黄酮合成的矛盾,减缓UV-B辐射对植株伤害。La(III)(20 mg/L)对植物的影响,可能与La(III)(20 mg/L)增加类黄酮含量,减轻UV-B辐射胁迫伤害,稳定细胞膜结构,维持植物水势,降低植物对胁迫的应激反应强度有关。La(III)(60 mg/L)能降低UV-B辐射胁迫下大豆幼苗IAA、GA和CTK含量、促进ABA含量、IAAO活性升高,加剧UV-B辐射胁迫对植物生长的抑制,进而加剧UV-B辐射胁迫对植株伤害。复合胁迫使细胞增殖与生长、器官生长发育减缓、形态结构改变,生物量下降。大豆幼苗内源乙烯释放量在La(III)(20 mg/L)和UV-B辐射胁迫共同作用下降低,源于1-氨基环丙烷-1-羧酸(ACC)含量和氨基环丙烷羧酸合酶活性的减少。La(III)(20 mg/L)通过加速大豆幼苗中自由基的清除,提高细胞膜保护系统活性和保持细胞膜透性来抑制乙烯的生成。La(III)(60 mg/L)表现出相反的结果。
(3) UV-B辐射胁迫下,大豆幼苗Trp含量下降,Phe、Tyr含量,DAHPS活性升高。其结果导致植物自身IAA含量下降,抑制植物生长。Phe、Tyr含量及DAHPS活性升高,使植物向着类黄酮代谢途径的方向生长,进而竞争性抑制了有机化合物向着植物生长的方向运行;与此同时,植物次生代谢作用被增强(类黄酮含量提高),植物把更多的资源分配于防御机制,从而导致生长减缓,生物量下降。La(III)(20 mg/L)通过提高Trp含量,有利于植物IAA含量的提高,同时通过提高大豆幼苗DAHPS活性及Phe、Tyr含量来促进类黄酮合成。类黄酮含量增加减少了进入植株体内的UV-B辐射净通量,减轻了UV-B辐射胁迫对大豆幼苗的伤害。La(III)(60 mg/L)通过抑制大豆幼苗Trp、Phe、Tyr含量及DAHPS活性进而抑制类黄酮合成。类黄酮含量减少从而加剧了进入植株体内的UV-B辐射净通量,增加了UV-B辐射胁迫对大豆幼苗的伤害。La(III)(60 mg/L)+UV-B辐射胁迫处理对大豆幼苗表现出复合胁迫伤害作用。
(4) UV-B辐射胁迫下,PAL活性、C4H活性、4CL活性、CHS活性及类黄酮含量升高,但在恢复期急剧下降。La(III)(20 mg/L)的介入有效提高PAL活性、C4H活性、4CL活性、CHS活性及类黄酮含量为植物抵御UV-B辐射胁迫提供屏障。La(III)(60 mg/L)的介入抑制了PAL活性、C4H活性、4CL活性、CHS活性及类黄酮含量,不利于类黄酮的合成,对植物抵御UV-B辐射胁迫起负作用。La(III)(20 mg/L)和UV-B辐射均能促进类黄酮代谢关键酶活性及类黄酮含量升高,但其机理不一致,前者是主动的促进,而后者是植物面对胁迫时的一种应激反应,是被动的提高。La(III)(20 mg/L)对类黄酮代谢关键酶活性及类黄酮含量调控效果与胁迫强度有关,即对低剂量UV-B辐射胁迫的缓解效果优于高剂量。
(5)通过分子动力学模拟计算,发现CHS蛋白表面存在大量负电荷集中区。根据化学原则,La(III)易与CHS蛋白相互作用,导致CHS蛋白结构发生改变。UV-B辐射导致Tyr和Trp残基的荧光同时被淬灭,CHS肽链上的酰氨基吸收峰及芳香氨基酸(Tyr和Trp)残基带的紫外吸收峰值下降,从而改变了分子的空间结构,降低CHS活性。低剂量La(III)与CHS肽链中的酰氨基团上的O或N原子及CHS肽链上的芳香氨基酸(Tyr和Trp)残基发生作用后,能导致CHS发生微结构的变化,稳定CHS酶结构,从而对活性中心产生微扰作用,进而影响CHS活性,促进类黄酮的合成,降低UV-B辐射诱导自由基的产生。高剂量La(III)直接伤害CHS酶结构,影响活性中心,导致CHS酶活性降低。
Atmospheric pollutants such as chlorofluorocarbons have caused severe depletion of stratospheric ozone, particularly in the Antarctic, where the Springtime“Ozone Hole”has greatly increased the amounts of damaging ultraviolet-B radiation reaching the Earth’s surface. This study used the method of water culture, and oil-bearing crops of soybean (Glycine max), as the test material, to explore the effect of La(III) and UV-B radiation on endogenous hormones and flavonoids metabolism and mechanism within the seedlings. Research methods used involve physiology, biochemistry, molecular spectroscopy technology and simulation technology. This study investigated the effects of La(III) on the content of endogenous hormones and aromatic amino acids, mechanism of key enzymes of flavonoid metabolism in soybean seedlings under UV-B radiation stress. It revealed the effects and mechanism of La(III) on endogenous hormones and metabolism of flavonoids in plants under UV-B radiation stress from an environmental and ecological perspective, physiological, biochemical and enzyme conformation levels.
The main research results of this dissertation were shown as follows:
(1) UV-B radiation stress effected grouth in soybean seedlings.The lack of green areas on leaf surfaces and rate of leaf folding were increased, leaf area, chlorophyll content, mineral elements, root activity, root length, fresh(dry) weight were decreased in soybean seedlings under UV-B radiation stress. This decline is related with UV-B radiation dosage. La(III)(20 mg/L) could increase soybean root activity, chlorophyll content, mineral elements, in order to promote photosynthesis; increased leaf area and fresh (dry) weight of leaves of soybean seedling promotes growth. seedlings was able to maintain absorption and utilization of light energy to meet the energy demand and materials needed for plant self-repair, to alleviate UV-B radiation on leaf damage after La(III)(20 mg/L) intervention. High doses of La (III)(60 mg/L)could reduce soybean root activity, chlorophyll content, mineral elements, thereby inhibiting photosynthesis; and lower leaf area and fresh (dry) weight of leaves in soybean seedling, thus inhibiting the growth of soybean seedlings. La(III)(20 mg/L) could alleviate the UV-B radiation stress on plant growth inhibition, La(III)(20 mg/L) could increase inhibition of UV-B radiation stress on plant growth.
(2) UV-B radiation stress causes a decrease in IAA, GA and CTK content and increased ABA content, IAAO activity and ETH content, plant growth, organ growth and development of mitigation, change in morphology, decrease in biomass, and thus harming the plants. La(III)(20 mg/L) in plants exposed to UV-B radiation stress can increase IAA, GA and CTK content and reduce ABA content, IAAO activity and ETH content, reduced inhibition caused by UV-B radiation on plant growth, and to plant growth and ease the contradiction of flavonoid synthesis, reduce UV-B radiation on plant injury. The effect of La(III)(20 mg/L) on plants may be related to La(III)(20 mg/L) causing an increase in flavonoids, reducing damage caused by UV-B radiation, stability of membrane structure, maintenance of plant water potential, and a lower intensity of stress response in plants. La(III)(60 mg/L) in soybean seedlings exposed to UV-B radiation stress, showed an decrease in IAA, GA and CTK content, increase in ABA content and IAAO activity, thus increasing plant injury caused by UV-B radiation. Combined stress leads to decreased cell proliferation and growth, organ growth and development of mitigation, morphology change, and a decrease in biomass. Endogenous ETH productionwas reduced in soybean seedlings under the action of La(III)(20 mg/L)+UV-B radiation stress, showed a decrease in 1-amino-cyclopropane-1-carboxylic acid (ACC) content and that of amino-cyclopropane carboxyl acid synthase activity, which reduces the amount of ethylene converted from 1-aminocyclopropane-1-carboxylic acid (ACC). La(III)(20 mg/L) accelerates the removal of free radicals to increase cell membrane protection system activity and maintain cell membrane permeability to inhibit ethylene production. However, La(III)(60 mg/L) shows opposite results.
(3) Trp content in soybean seedlings decreased, while Phe, Tyr, and DAHPS activity increased in soybean seadlinngs under UV-B stress. As a result, IAA content in plants was decreased and plant growth was inhibited. La(III)(20 mg/L) increases Trp, Phe, Tyr content and DAHPS activity, is beneficial for increasing plant IAA content, while by increasing the activity of soybean seedlings DAHPS and Phe, Tyr content to promote the synthesis of flavonoids. Increase of flavonoids would reduce the amount of UV-B radiation absorbed by the plant, thus reducing damage caused by UV-B radiation on soybean seedling. La(III)(60 mg/L) of soybean seedlings inhibited flavonoids synthesis by inhibiting Trp, Phe, Tyr content and DAHPS activity. This decrease in flavonoids thus increased the amount of UV-B radiation absorbed in the plant and damage on soybean seedling was increased.
(4) The PAL activity, C4H activity, 4CL activity, CHS activity and flavonoid content were increased in soybean seedlings under UV-B radiation stress, but declined sharply in the recovery period. La(III)(20 mg/L) effectively enhanced PAL activity, C4H activity, 4CL activity, CHS activity and flavonoid content to provide barriers against UV-B radiation stress in plants. La(III)(60 mg/L) inhibited PAL activity, C4H activity, 4CL activity, CHS activity and flavonoid content, and played a negative role under UV-B radiation stress of the plants. La(III)(20 mg/L) and UV-B radiation could promote the activities of key enzymes of flavonoid metabolism, and increased flavonoid content, but the mechanism was inconsistent, the former is a stress response by the plants, which has a passive increase.
(5) The method of physical chemistry, molecular dynamics simulation was used in the in vitro experiments. It showed that changes in molecular structure of CHS under UV-B radiation stress is UV-B radiation reduced the molecular basis of CHS activity; it was found that a large number of negative charge clusters on CHS protein surface, indicating that La(III) may be a relationship directly with the CHS proteins. La(III) acted with acylamino group O, or N atom on CHS peptide chain directly, which lead to change CHS micro structure, increase/decrease CHS activity, promote/inhibit the synthesis of flavonoids.
结果如下:
(1) UV-B辐射胁迫下,大豆幼苗根系活力下降,抑制了营养元素吸收,破坏叶片形态,降低叶片光合作用,最终导致整个植物干物质累积量降低,此效应随UV-B辐射胁迫剂量增加而加剧。La(III)(20 mg/L)能提高大豆根系活力、叶绿素含量、矿质元素含量,进而促进光合作用,提高叶片叶面积及大豆幼苗鲜(干)重,促进大豆幼苗的生长。La(III)(20 mg/L)的介入使逆境中的大豆幼苗也能尽量维持对光能的吸收和利用,从而满足植物自身恢复所需的能量和物质,缓解UV-B辐射对叶片损伤。高剂量La(III)(60 mg/L)降低大豆根系活力、叶绿素含量、矿质元素含量,进而抑制光合作用,降低叶片叶面积及大豆幼苗鲜(干)重,抑制大豆幼苗的生长。La(III)(60 mg/L)+UV-B辐射胁迫处理对大豆幼苗起到了协同抑制作用。La(III)(60 mg/L)的介入加剧了植物自身恢复所需能量和物质的减少,使逆境中的大豆幼苗表现出受到复合胁迫伤害的作用。适量La(III)可缓解UV-B辐射胁迫对大豆生长的抑制作用,而高剂量La(III)可加剧UV-B辐射胁迫对大豆生长的抑制作用。
(2) UV-B辐射胁迫下,大豆幼苗IAA、GA和CTK含量下降,ABA、ETH含量和IAAO活性增加,器官生长发育减缓、形态结构改变、生物量下降,此效应随UV-B辐射胁迫剂量增加而加剧。La(III)(20 mg/L)能提高UV-B辐射胁迫下大豆幼苗IAA、GA和CTK含量、降低ABA含量、IAAO活性和乙烯含量,减轻UV-B辐射胁迫对植物生长的抑制,缓解植株生长与类黄酮合成的矛盾,减缓UV-B辐射对植株伤害。La(III)(20 mg/L)对植物的影响,可能与La(III)(20 mg/L)增加类黄酮含量,减轻UV-B辐射胁迫伤害,稳定细胞膜结构,维持植物水势,降低植物对胁迫的应激反应强度有关。La(III)(60 mg/L)能降低UV-B辐射胁迫下大豆幼苗IAA、GA和CTK含量、促进ABA含量、IAAO活性升高,加剧UV-B辐射胁迫对植物生长的抑制,进而加剧UV-B辐射胁迫对植株伤害。复合胁迫使细胞增殖与生长、器官生长发育减缓、形态结构改变,生物量下降。大豆幼苗内源乙烯释放量在La(III)(20 mg/L)和UV-B辐射胁迫共同作用下降低,源于1-氨基环丙烷-1-羧酸(ACC)含量和氨基环丙烷羧酸合酶活性的减少。La(III)(20 mg/L)通过加速大豆幼苗中自由基的清除,提高细胞膜保护系统活性和保持细胞膜透性来抑制乙烯的生成。La(III)(60 mg/L)表现出相反的结果。
(3) UV-B辐射胁迫下,大豆幼苗Trp含量下降,Phe、Tyr含量,DAHPS活性升高。其结果导致植物自身IAA含量下降,抑制植物生长。Phe、Tyr含量及DAHPS活性升高,使植物向着类黄酮代谢途径的方向生长,进而竞争性抑制了有机化合物向着植物生长的方向运行;与此同时,植物次生代谢作用被增强(类黄酮含量提高),植物把更多的资源分配于防御机制,从而导致生长减缓,生物量下降。La(III)(20 mg/L)通过提高Trp含量,有利于植物IAA含量的提高,同时通过提高大豆幼苗DAHPS活性及Phe、Tyr含量来促进类黄酮合成。类黄酮含量增加减少了进入植株体内的UV-B辐射净通量,减轻了UV-B辐射胁迫对大豆幼苗的伤害。La(III)(60 mg/L)通过抑制大豆幼苗Trp、Phe、Tyr含量及DAHPS活性进而抑制类黄酮合成。类黄酮含量减少从而加剧了进入植株体内的UV-B辐射净通量,增加了UV-B辐射胁迫对大豆幼苗的伤害。La(III)(60 mg/L)+UV-B辐射胁迫处理对大豆幼苗表现出复合胁迫伤害作用。
(4) UV-B辐射胁迫下,PAL活性、C4H活性、4CL活性、CHS活性及类黄酮含量升高,但在恢复期急剧下降。La(III)(20 mg/L)的介入有效提高PAL活性、C4H活性、4CL活性、CHS活性及类黄酮含量为植物抵御UV-B辐射胁迫提供屏障。La(III)(60 mg/L)的介入抑制了PAL活性、C4H活性、4CL活性、CHS活性及类黄酮含量,不利于类黄酮的合成,对植物抵御UV-B辐射胁迫起负作用。La(III)(20 mg/L)和UV-B辐射均能促进类黄酮代谢关键酶活性及类黄酮含量升高,但其机理不一致,前者是主动的促进,而后者是植物面对胁迫时的一种应激反应,是被动的提高。La(III)(20 mg/L)对类黄酮代谢关键酶活性及类黄酮含量调控效果与胁迫强度有关,即对低剂量UV-B辐射胁迫的缓解效果优于高剂量。
(5)通过分子动力学模拟计算,发现CHS蛋白表面存在大量负电荷集中区。根据化学原则,La(III)易与CHS蛋白相互作用,导致CHS蛋白结构发生改变。UV-B辐射导致Tyr和Trp残基的荧光同时被淬灭,CHS肽链上的酰氨基吸收峰及芳香氨基酸(Tyr和Trp)残基带的紫外吸收峰值下降,从而改变了分子的空间结构,降低CHS活性。低剂量La(III)与CHS肽链中的酰氨基团上的O或N原子及CHS肽链上的芳香氨基酸(Tyr和Trp)残基发生作用后,能导致CHS发生微结构的变化,稳定CHS酶结构,从而对活性中心产生微扰作用,进而影响CHS活性,促进类黄酮的合成,降低UV-B辐射诱导自由基的产生。高剂量La(III)直接伤害CHS酶结构,影响活性中心,导致CHS酶活性降低。
Atmospheric pollutants such as chlorofluorocarbons have caused severe depletion of stratospheric ozone, particularly in the Antarctic, where the Springtime“Ozone Hole”has greatly increased the amounts of damaging ultraviolet-B radiation reaching the Earth’s surface. This study used the method of water culture, and oil-bearing crops of soybean (Glycine max), as the test material, to explore the effect of La(III) and UV-B radiation on endogenous hormones and flavonoids metabolism and mechanism within the seedlings. Research methods used involve physiology, biochemistry, molecular spectroscopy technology and simulation technology. This study investigated the effects of La(III) on the content of endogenous hormones and aromatic amino acids, mechanism of key enzymes of flavonoid metabolism in soybean seedlings under UV-B radiation stress. It revealed the effects and mechanism of La(III) on endogenous hormones and metabolism of flavonoids in plants under UV-B radiation stress from an environmental and ecological perspective, physiological, biochemical and enzyme conformation levels.
The main research results of this dissertation were shown as follows:
(1) UV-B radiation stress effected grouth in soybean seedlings.The lack of green areas on leaf surfaces and rate of leaf folding were increased, leaf area, chlorophyll content, mineral elements, root activity, root length, fresh(dry) weight were decreased in soybean seedlings under UV-B radiation stress. This decline is related with UV-B radiation dosage. La(III)(20 mg/L) could increase soybean root activity, chlorophyll content, mineral elements, in order to promote photosynthesis; increased leaf area and fresh (dry) weight of leaves of soybean seedling promotes growth. seedlings was able to maintain absorption and utilization of light energy to meet the energy demand and materials needed for plant self-repair, to alleviate UV-B radiation on leaf damage after La(III)(20 mg/L) intervention. High doses of La (III)(60 mg/L)could reduce soybean root activity, chlorophyll content, mineral elements, thereby inhibiting photosynthesis; and lower leaf area and fresh (dry) weight of leaves in soybean seedling, thus inhibiting the growth of soybean seedlings. La(III)(20 mg/L) could alleviate the UV-B radiation stress on plant growth inhibition, La(III)(20 mg/L) could increase inhibition of UV-B radiation stress on plant growth.
(2) UV-B radiation stress causes a decrease in IAA, GA and CTK content and increased ABA content, IAAO activity and ETH content, plant growth, organ growth and development of mitigation, change in morphology, decrease in biomass, and thus harming the plants. La(III)(20 mg/L) in plants exposed to UV-B radiation stress can increase IAA, GA and CTK content and reduce ABA content, IAAO activity and ETH content, reduced inhibition caused by UV-B radiation on plant growth, and to plant growth and ease the contradiction of flavonoid synthesis, reduce UV-B radiation on plant injury. The effect of La(III)(20 mg/L) on plants may be related to La(III)(20 mg/L) causing an increase in flavonoids, reducing damage caused by UV-B radiation, stability of membrane structure, maintenance of plant water potential, and a lower intensity of stress response in plants. La(III)(60 mg/L) in soybean seedlings exposed to UV-B radiation stress, showed an decrease in IAA, GA and CTK content, increase in ABA content and IAAO activity, thus increasing plant injury caused by UV-B radiation. Combined stress leads to decreased cell proliferation and growth, organ growth and development of mitigation, morphology change, and a decrease in biomass. Endogenous ETH productionwas reduced in soybean seedlings under the action of La(III)(20 mg/L)+UV-B radiation stress, showed a decrease in 1-amino-cyclopropane-1-carboxylic acid (ACC) content and that of amino-cyclopropane carboxyl acid synthase activity, which reduces the amount of ethylene converted from 1-aminocyclopropane-1-carboxylic acid (ACC). La(III)(20 mg/L) accelerates the removal of free radicals to increase cell membrane protection system activity and maintain cell membrane permeability to inhibit ethylene production. However, La(III)(60 mg/L) shows opposite results.
(3) Trp content in soybean seedlings decreased, while Phe, Tyr, and DAHPS activity increased in soybean seadlinngs under UV-B stress. As a result, IAA content in plants was decreased and plant growth was inhibited. La(III)(20 mg/L) increases Trp, Phe, Tyr content and DAHPS activity, is beneficial for increasing plant IAA content, while by increasing the activity of soybean seedlings DAHPS and Phe, Tyr content to promote the synthesis of flavonoids. Increase of flavonoids would reduce the amount of UV-B radiation absorbed by the plant, thus reducing damage caused by UV-B radiation on soybean seedling. La(III)(60 mg/L) of soybean seedlings inhibited flavonoids synthesis by inhibiting Trp, Phe, Tyr content and DAHPS activity. This decrease in flavonoids thus increased the amount of UV-B radiation absorbed in the plant and damage on soybean seedling was increased.
(4) The PAL activity, C4H activity, 4CL activity, CHS activity and flavonoid content were increased in soybean seedlings under UV-B radiation stress, but declined sharply in the recovery period. La(III)(20 mg/L) effectively enhanced PAL activity, C4H activity, 4CL activity, CHS activity and flavonoid content to provide barriers against UV-B radiation stress in plants. La(III)(60 mg/L) inhibited PAL activity, C4H activity, 4CL activity, CHS activity and flavonoid content, and played a negative role under UV-B radiation stress of the plants. La(III)(20 mg/L) and UV-B radiation could promote the activities of key enzymes of flavonoid metabolism, and increased flavonoid content, but the mechanism was inconsistent, the former is a stress response by the plants, which has a passive increase.
(5) The method of physical chemistry, molecular dynamics simulation was used in the in vitro experiments. It showed that changes in molecular structure of CHS under UV-B radiation stress is UV-B radiation reduced the molecular basis of CHS activity; it was found that a large number of negative charge clusters on CHS protein surface, indicating that La(III) may be a relationship directly with the CHS proteins. La(III) acted with acylamino group O, or N atom on CHS peptide chain directly, which lead to change CHS micro structure, increase/decrease CHS activity, promote/inhibit the synthesis of flavonoids.
引文
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