渗透胁迫下小麦幼苗体内多胺形态、定位与功能
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摘要
干旱是困扰世界粮食生产的主要问题。研究植物耐旱机理以及探索提高植物抗旱能力途径一直是各国科研工作者关注的热点之一。小麦是我国和全世界主要的粮食作物。多胺是生物体内广泛存在的一类低分子量脂肪族含氮碱，是调控植物生长和发育的重要生理活性物质。然而，有关多胺与植物抗旱性的研究较为少见，特别是有关水分胁迫下游离态腐胺(fPut)累积的意义存在着较大分歧；有关结合态多胺与水分胁迫关系的研究就更为罕见；目前，尚缺乏水分胁迫下有关共价和非共价结合态多胺的细胞定位以及多胺参与水分胁迫信号转导功能报告。因此，笔者选择生长于不同生态区抗旱性不同的两个小麦品种豫麦18(抗旱性强)和扬麦9号(抗旱性弱)为试材，用聚乙二醇(PEG-6000)模拟自然干旱对幼苗根际进行渗透胁迫，研究多胺的种类和形态的变化、细胞中定位及功能。研究结果如下：
渗透胁迫7d，小麦幼苗叶片三种游离态多胺：游离态腐胺(fPut)、游离态亚精胺(fSpd)、游离态精胺(fSpm)和高氯酸不溶性共价结合态(PISCC)-Put均上升，但是豫麦18号的fSpd、fSpm和PISCC-Put的上升幅度明显大于扬麦9号，而扬麦9号的fPut的上升幅度明显大于豫麦18号，所以导致了在渗透胁迫条件下，豫麦18号的fSpd+fSpm／fPut的比值明显大于扬麦9号。渗透胁迫也引起了幼苗叶片中的PSCC-PAs(PScC-Put、PSCC-Spd和PSCC-Spm)含量的上升，但是在两个小麦品种之间的上升幅度没有差异。这些结果表明，渗透胁迫条件下，小麦幼苗叶片的fSpd、fSpm及PISCC-Put含量的上升有利于增强小麦幼苗的抗胁迫能力，而幼苗叶片的PSCC-PAs与幼苗的抗渗能力影响不大。外源多胺和抑制剂处理实验以及以胚芽鞘为材料的实验验证了这一结论。
渗透胁迫下多胺的动态变化研究表明，渗透胁迫6h，两个品种叶片的fPut含量都急剧升高，豫麦18的升高幅度大于扬麦9号，但在胁迫6h后fPut含量急剧下降，并稳定在200nmol／g Fw左右的水平，然而扬麦9号的fPut水平在胁迫6h后仅稍有下降，随后稳定在300 nmol／g Fw左右的水平。两个品种的fSpd和fSpm水平在胁迫6h内与对照无大差别，然而在胁迫6h后，豫麦18的fSpd和fSpm急剧升高，12h后，分别稳定在150-200 nmol／gFW和80-100nmol／g Fw水平，而扬麦9号的fSpd和fSpm含量在胁迫48h内变化不明显。检测PISCC-PAs发现，随着胁迫时间的延长，两个品种叶片的PISCC-Put逐渐升高，但是豫麦18在胁迫12h后的升高幅度明显大于扬麦9号。这些结果表明，小麦幼苗对渗透胁迫的适应不仅表现为胁
刘怀攀南京农业大学博士学位论文:渗透胁迫下小麦幼苗体内多胺形态、定位与功能
迫初期叶片内游离态腐胺(fPut)的大量累积，而且更重要的是这些fPut能否及时
转化为游离态亚精胺(fspd)和游离态精胺(fSPm)以及高氛酸不溶性的共价结合态
腐胺(PISCC一Put)o
豫麦18根系质膜上的NCC一Spd、NCC一Spm和PISCC一Put含量以及H’一ATPase活性
在渗透胁迫下明显上升，而扬麦9号的变化不明显。豫麦18的P工sCC一Spd含量在渗
透胁迫下的上升幅度明显大于扬麦9号。暗示了渗透胁迫下，小麦幼苗根质膜上的
NCC一Spd、NCC一Spm、PISCC一Put和PISCC一Spd通过提高质膜H’一^TPase的活性而增强
幼苗的杭渗透胁迫能力。
扬麦9号根系液泡膜H+一ATPase和H‘一PPase活性在渗透胁迫下明显下降，而豫麦
18下降不明显;豫麦18根系液泡膜上NCC一Spd和PISCC一Put含量在渗透胁迫下的上
升幅度明显大于扬麦9号。暗示了渗透胁迫下，液泡膜上NCC一Spd以及PIScC一Put有
才}!于维持膜上H‘一ATPase和H‘一PPase活性的稳定，从而有助于提高小麦幼苗的抗渗透
胁迫能力。
豫麦18根系线粒体膜H’一ATPase活性和膜上筑基(一SH)的含量在渗透胁迫下的
下降幅度明显小于扬麦9号，而膜上NCC一SPd和Pl SCC一Put含量在渗透胁迫下的上升
幅度明显大于扬麦9号.暗示了渗透胁迫下，小麦幼苗根线粒体膜上的NCC一SPd和
PISCC一Put可能通过维持膜上筑基含量和H’一ATPase活性的稳定来提高小麦幼苗的抗
渗透胁迫能力.
与其它细胞器不同，除了NCC一Put外，小麦幼苗类囊体膜上结合态多胺含量在渗
透胁迫下都有所下降，且豫麦18的NCC一Spd、PISCC一Put和PISCC一Spd含量的下降幅
度明显小于扬麦9号，而扬麦9号的NCC一SPm在渗透胁迫下明显下降，豫麦18的
NCC一Spm变化不明显。暗示了渗透胁迫下，小麦幼苗类囊体膜上NcC一SPd、PISCC一Spd
和PlsCC一Put含量的下降，可能是叶绿体光合作用对水分胁迫敏感的原因之一。
NCC一SPd、P工SCC一Spd和PISCC一Put水平在胁迫条件下的稳定有利于增强小麦幼苗适应
渗透胁迫的能力。
豫麦18根系核蛋白体上NCC一Spm、PISCC一Put和PISCC一Spd含量在渗透胁迫一的
上升幅度明显大于扬麦9号，像麦18的NCC一Spd含量在渗透胁迫下的上升明显，而
扬麦9号的NCC一SPd含量变化不明显.暗示了渗透胁迫下，小麦幼苗根核蛋白体上
NCC一Spm、NCc一Spd、pIScC一put和pISCC一Spd水平的上升可能通过维持核蛋白体构象
和功能的稳定以及调节基因的表达而增强幼苗适应渗透胁迫的能力。
渗透胁迫初期小麦幼苗根fPut的积累受ABA
Drought is a problem worldwide that threatens food supplies. To investigate the mechanism of plant response to drought stress and to improve the drought tolerance has been one of major considerations for a long time. Wheat (Triticum aestivum L.) is one of the important food crops. Polyamines (PAs) are biologically ubiquitous aliphatic amines with low molecular weight that are implicated in many aspects of growth and development. However, the studies on the relationship between polyamines and tolerance of plant to water stress remain disagreement, especially on the significance of great accumulation of putrescine (Put) under water stress, and at the same time the studies on relationship between conjugated polyamines and water stress are still lacking. The subcellular localization of conjugated polyamines in plants under osmotic stress and involvement of polyamine in water stress-induced signaling pathway have not yet been documented. Here, two wheat cultivars with different drought stress-tolerance (Yumai No.
18 cv., drought-tolerant; Yangmai No.9 cv., drought-sensitive) were used as experimental materials to study the changes in PA kinds and forms, cellular localization and function of polyamines in seedlings subjected to PEG-6000 osmotic stress. The results are as follow:
PEG osmotic stress led to increases of three free polyamine level (fPut, fSpd and fSpm) in the leaves of two wheat cultivar seedlings. FSpd, fSpm and PISCC-Put (perchloric acid insoluble covalently conjugated Put) levels in leaves of Yumai No. 18 cv. (drought-tolerant) increased more markedly than those in Yangmai No.9 cv. (drought-sensitive) after polyethylene glycol (PEG) 6,000 treatment for 7 d, while fPut level has a more significant increase in Yangmai No.9. Thus Yumai No. 18 cv. gave a higher ratio of (fSpd + fSpm) / fPut than Yangmai No.9 cv. in leaves in response to osmotic stress. PEG osmotic stress also induced increases of perchloric acid soluble covalently conjugated polyamine ( PSCC-PA: PSCC-Put, PSCC-Spd and PSCC-Spm) levels in leaves, but such a increase has no aparently difference between the two cv. wheat seedlings. The results suggested that fSpd, fSpm and PISCC-Put in leaves enhance the adaptation of the seedlings to osmotic stres.
This hypothesis has also been demonstrated by the results of experiments with exogenous polyamines and inhibitors and was consistent with the results of experiments on wheat coleoptiles.
FPut level increased greatly in leaves of two cv. wheat seedlings with PEG treatment for 6 h, but fPut level in Yumai No. 18 cv. decreased sharply to a low level (200 nmol/g FW) after 6 h, while in Yangmai No.9 cv. a high level of fPut (300 nmol/g FW) was remained after 6 h. PEG osmotic stress affected slightly the fSpd and fSpm levels in leaves of two cv. wheat seedlings during PEG treatment for 6 h. However, after 6 h, the levels of fSpd and fSpm in leaves of Yumai No. 18 cv. increased greatly, while the increase in Yangmai No.9 cv. was slight. We also found that Yumai No. 18 cv. have greater increases of PISCC-Put than Yangmai No.9 cv. in leaves after PEG treatment for 12 h. These results suggested that tolerance of wheat seedlings to osmotic stress-induced injury is attributed not only to the great increase of fPut in leaves at prophase, but also to the conversion of fPut to fSpd, fSpm and PISCC-Put in time. This hypothesis has also been demonstrated by the results of experiments with exogenous polyamines and inhibitors.
Under osmotic stress, significant increases of NCC-Spd, NCC-Spm and PISCC-Put levels and activity of H+-ATPase were observed in root plasma membrane (PM) of drought-tolerant Yumai No. 18 cv. wheat seedling, while no obvious change was found in drought-sensitive Yangmai No.9 cv. Furthermore, Yumai No. 18 gave more obvious increase of PISCC-Spd than Yangmai No. 9. The results suggested that the tolerance of the wheat seedlings to osmotic stress was associated with the activity of H+-ATPase and the contents of NCC-Spd and NCC-Spm and CC-Put and CC-Spd in PM of the seedling roots.
渗透胁迫7d，小麦幼苗叶片三种游离态多胺：游离态腐胺(fPut)、游离态亚精胺(fSpd)、游离态精胺(fSpm)和高氯酸不溶性共价结合态(PISCC)-Put均上升，但是豫麦18号的fSpd、fSpm和PISCC-Put的上升幅度明显大于扬麦9号，而扬麦9号的fPut的上升幅度明显大于豫麦18号，所以导致了在渗透胁迫条件下，豫麦18号的fSpd+fSpm／fPut的比值明显大于扬麦9号。渗透胁迫也引起了幼苗叶片中的PSCC-PAs(PScC-Put、PSCC-Spd和PSCC-Spm)含量的上升，但是在两个小麦品种之间的上升幅度没有差异。这些结果表明，渗透胁迫条件下，小麦幼苗叶片的fSpd、fSpm及PISCC-Put含量的上升有利于增强小麦幼苗的抗胁迫能力，而幼苗叶片的PSCC-PAs与幼苗的抗渗能力影响不大。外源多胺和抑制剂处理实验以及以胚芽鞘为材料的实验验证了这一结论。
渗透胁迫下多胺的动态变化研究表明，渗透胁迫6h，两个品种叶片的fPut含量都急剧升高，豫麦18的升高幅度大于扬麦9号，但在胁迫6h后fPut含量急剧下降，并稳定在200nmol／g Fw左右的水平，然而扬麦9号的fPut水平在胁迫6h后仅稍有下降，随后稳定在300 nmol／g Fw左右的水平。两个品种的fSpd和fSpm水平在胁迫6h内与对照无大差别，然而在胁迫6h后，豫麦18的fSpd和fSpm急剧升高，12h后，分别稳定在150-200 nmol／gFW和80-100nmol／g Fw水平，而扬麦9号的fSpd和fSpm含量在胁迫48h内变化不明显。检测PISCC-PAs发现，随着胁迫时间的延长，两个品种叶片的PISCC-Put逐渐升高，但是豫麦18在胁迫12h后的升高幅度明显大于扬麦9号。这些结果表明，小麦幼苗对渗透胁迫的适应不仅表现为胁
刘怀攀南京农业大学博士学位论文:渗透胁迫下小麦幼苗体内多胺形态、定位与功能
迫初期叶片内游离态腐胺(fPut)的大量累积，而且更重要的是这些fPut能否及时
转化为游离态亚精胺(fspd)和游离态精胺(fSPm)以及高氛酸不溶性的共价结合态
腐胺(PISCC一Put)o
豫麦18根系质膜上的NCC一Spd、NCC一Spm和PISCC一Put含量以及H’一ATPase活性
在渗透胁迫下明显上升，而扬麦9号的变化不明显。豫麦18的P工sCC一Spd含量在渗
透胁迫下的上升幅度明显大于扬麦9号。暗示了渗透胁迫下，小麦幼苗根质膜上的
NCC一Spd、NCC一Spm、PISCC一Put和PISCC一Spd通过提高质膜H’一^TPase的活性而增强
幼苗的杭渗透胁迫能力。
扬麦9号根系液泡膜H+一ATPase和H‘一PPase活性在渗透胁迫下明显下降，而豫麦
18下降不明显;豫麦18根系液泡膜上NCC一Spd和PISCC一Put含量在渗透胁迫下的上
升幅度明显大于扬麦9号。暗示了渗透胁迫下，液泡膜上NCC一Spd以及PIScC一Put有
才}!于维持膜上H‘一ATPase和H‘一PPase活性的稳定，从而有助于提高小麦幼苗的抗渗透
胁迫能力。
豫麦18根系线粒体膜H’一ATPase活性和膜上筑基(一SH)的含量在渗透胁迫下的
下降幅度明显小于扬麦9号，而膜上NCC一SPd和Pl SCC一Put含量在渗透胁迫下的上升
幅度明显大于扬麦9号.暗示了渗透胁迫下，小麦幼苗根线粒体膜上的NCC一SPd和
PISCC一Put可能通过维持膜上筑基含量和H’一ATPase活性的稳定来提高小麦幼苗的抗
渗透胁迫能力.
与其它细胞器不同，除了NCC一Put外，小麦幼苗类囊体膜上结合态多胺含量在渗
透胁迫下都有所下降，且豫麦18的NCC一Spd、PISCC一Put和PISCC一Spd含量的下降幅
度明显小于扬麦9号，而扬麦9号的NCC一SPm在渗透胁迫下明显下降，豫麦18的
NCC一Spm变化不明显。暗示了渗透胁迫下，小麦幼苗类囊体膜上NcC一SPd、PISCC一Spd
和PlsCC一Put含量的下降，可能是叶绿体光合作用对水分胁迫敏感的原因之一。
NCC一SPd、P工SCC一Spd和PISCC一Put水平在胁迫条件下的稳定有利于增强小麦幼苗适应
渗透胁迫的能力。
豫麦18根系核蛋白体上NCC一Spm、PISCC一Put和PISCC一Spd含量在渗透胁迫一的
上升幅度明显大于扬麦9号，像麦18的NCC一Spd含量在渗透胁迫下的上升明显，而
扬麦9号的NCC一SPd含量变化不明显.暗示了渗透胁迫下，小麦幼苗根核蛋白体上
NCC一Spm、NCc一Spd、pIScC一put和pISCC一Spd水平的上升可能通过维持核蛋白体构象
和功能的稳定以及调节基因的表达而增强幼苗适应渗透胁迫的能力。
渗透胁迫初期小麦幼苗根fPut的积累受ABA
Drought is a problem worldwide that threatens food supplies. To investigate the mechanism of plant response to drought stress and to improve the drought tolerance has been one of major considerations for a long time. Wheat (Triticum aestivum L.) is one of the important food crops. Polyamines (PAs) are biologically ubiquitous aliphatic amines with low molecular weight that are implicated in many aspects of growth and development. However, the studies on the relationship between polyamines and tolerance of plant to water stress remain disagreement, especially on the significance of great accumulation of putrescine (Put) under water stress, and at the same time the studies on relationship between conjugated polyamines and water stress are still lacking. The subcellular localization of conjugated polyamines in plants under osmotic stress and involvement of polyamine in water stress-induced signaling pathway have not yet been documented. Here, two wheat cultivars with different drought stress-tolerance (Yumai No.
18 cv., drought-tolerant; Yangmai No.9 cv., drought-sensitive) were used as experimental materials to study the changes in PA kinds and forms, cellular localization and function of polyamines in seedlings subjected to PEG-6000 osmotic stress. The results are as follow:
PEG osmotic stress led to increases of three free polyamine level (fPut, fSpd and fSpm) in the leaves of two wheat cultivar seedlings. FSpd, fSpm and PISCC-Put (perchloric acid insoluble covalently conjugated Put) levels in leaves of Yumai No. 18 cv. (drought-tolerant) increased more markedly than those in Yangmai No.9 cv. (drought-sensitive) after polyethylene glycol (PEG) 6,000 treatment for 7 d, while fPut level has a more significant increase in Yangmai No.9. Thus Yumai No. 18 cv. gave a higher ratio of (fSpd + fSpm) / fPut than Yangmai No.9 cv. in leaves in response to osmotic stress. PEG osmotic stress also induced increases of perchloric acid soluble covalently conjugated polyamine ( PSCC-PA: PSCC-Put, PSCC-Spd and PSCC-Spm) levels in leaves, but such a increase has no aparently difference between the two cv. wheat seedlings. The results suggested that fSpd, fSpm and PISCC-Put in leaves enhance the adaptation of the seedlings to osmotic stres.
This hypothesis has also been demonstrated by the results of experiments with exogenous polyamines and inhibitors and was consistent with the results of experiments on wheat coleoptiles.
FPut level increased greatly in leaves of two cv. wheat seedlings with PEG treatment for 6 h, but fPut level in Yumai No. 18 cv. decreased sharply to a low level (200 nmol/g FW) after 6 h, while in Yangmai No.9 cv. a high level of fPut (300 nmol/g FW) was remained after 6 h. PEG osmotic stress affected slightly the fSpd and fSpm levels in leaves of two cv. wheat seedlings during PEG treatment for 6 h. However, after 6 h, the levels of fSpd and fSpm in leaves of Yumai No. 18 cv. increased greatly, while the increase in Yangmai No.9 cv. was slight. We also found that Yumai No. 18 cv. have greater increases of PISCC-Put than Yangmai No.9 cv. in leaves after PEG treatment for 12 h. These results suggested that tolerance of wheat seedlings to osmotic stress-induced injury is attributed not only to the great increase of fPut in leaves at prophase, but also to the conversion of fPut to fSpd, fSpm and PISCC-Put in time. This hypothesis has also been demonstrated by the results of experiments with exogenous polyamines and inhibitors.
Under osmotic stress, significant increases of NCC-Spd, NCC-Spm and PISCC-Put levels and activity of H+-ATPase were observed in root plasma membrane (PM) of drought-tolerant Yumai No. 18 cv. wheat seedling, while no obvious change was found in drought-sensitive Yangmai No.9 cv. Furthermore, Yumai No. 18 gave more obvious increase of PISCC-Spd than Yangmai No. 9. The results suggested that the tolerance of the wheat seedlings to osmotic stress was associated with the activity of H+-ATPase and the contents of NCC-Spd and NCC-Spm and CC-Put and CC-Spd in PM of the seedling roots.
引文
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