5-氨基乙酰丙酸调控干旱胁迫下油菜幼苗生长的生理机制研究
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摘要
以甘蓝型油菜品种浙双758(Brassica napus cv. Zheshuang758)为试验材料,利用生理学、分子生物学等方法研究了5-氨基乙酰丙酸(ALA)对干旱胁迫下油菜幼苗活性氧清除系统、光合作用、水分关系、矿质元素的吸收及转运以及叶绿素合成的影响,探讨了其调控干旱胁迫下油菜生长的机理。所得主要结果如下：
1.研究了根部施用ALA对聚乙二醇(PEG)诱导的水分缺失胁迫下油菜子叶期幼苗生长及活性氧清除系统的影响。PEG诱导的水分缺失胁迫抑制幼苗生长的同时增加了子叶中丙二醛(MDA)和活性氧自由基(ROS, O2和H202)的积累,提高了抗氧化酶的活性以及谷胱甘肽和抗坏血酸还原型和氧化型的比值。对根部施用低浓度的ALA (0.1-1mg L-1)提高了地上部鲜重和叶绿素含量,而10mgL-1ALA则对幼苗的生长表现出抑制作用。在水分缺失胁迫下,低浓度的ALA促进了叶绿素的积累,进一步提高了谷胱甘肽和抗坏血酸还原型和氧化型的比值。同时,与PEG单独处理相比,1mg L-1ALA显著地降低了受胁迫子叶中MDA的含量和ROS的积累,进一步提升了抗氧化酶的活性,上调了POD、CAT和GR的表达水平。以上结果表明,低浓度的ALA在水分缺失胁迫下能够通过提高ASA-GSH循环,提升抗氧化酶活性以及上调部分抗氧化酶基因的表达以保证植物能够有效地清除ROS,从而缓解子叶期幼苗受到的水分缺失胁迫。
2.研究了叶面喷施ALA对基质造成的干旱胁迫下油菜四叶期幼苗抗氧化酶活性及基因表达的影响。干旱胁迫增加了叶片中MDA和H2O2的积累,增强了CAT、 POD、 APX和GR的活性,上调了APX和GR的转录水平。在复水处理后,MDA和H202的含量以及APX和GR的表达量均恢复到正常水平。在干旱胁迫下,与对照(清水处理)相比,叶面喷施ALA有效地缓解了MDA和H2O2的积累,增加了抗氧化酶的活性,并上调了CAT和POD的表达。以上结果表明,叶片喷施ALA能通过增强CAT、 POD、 APX和GR的活性以及上调CAT和POD的表达来促进受胁迫叶片对ROS的清除。
3.研究了外源ALA对干旱胁迫下油菜幼苗光合作用的调控机理。干旱胁迫抑制了幼苗地上部和根部的生长,减少了叶片中叶绿素、淀粉以及可溶性蛋白的含量,降低了叶片净光合速率(Pn)、光系统Ⅱ(PSⅡ)的最大和实际光合量子产量,下调了卡尔文循环相关基因的表达,但却诱导了还原糖和蔗糖的积累。复水处理4d后,大多数指标均能恢复到正常水平。叶面喷施ALA能够部分地缓解干旱胁迫对油菜生长的抑制。在干旱胁迫下,与清水处理相比,ALA促进了叶绿素的积累,提高了叶片的Pn和气孔导度(gs),提升了光化学焠灭系数(qP)和非光化学焠灭系数(NPQ),上调了卡尔文循环相关基因的表达,同时在维持淀粉和可溶性蛋白含量正常水平的情况下进一步促进了还原糖和蔗糖的积累。这些结果表明,叶面喷施ALA对受干旱胁迫植株的光合作用的增强可能与气孔导度的增加、叶绿素含量的升高、植物光保护能力的提升以及卡尔文循环相关基因表达的上调有关。
4.研究了外源ALA对干旱胁迫下油菜水分关系、有机渗透物质的积累以及无机离子吸收的影响。干旱胁迫降低了油菜幼苗的叶片相对含水量和水势,诱导了叶片和根中可溶性糖、游离氨基酸和脯氨酸的积累,抑制了根系对矿质元素的吸收。复水处理4d后,叶片中脯氨酸和可溶性糖含量恢复到正常水平,但根系对矿质离子的吸收仍没有完全恢复。在干旱胁迫下,ALA降低了叶片中脯氨酸含量,促进了叶片中Na+的外排,提高了K+/Na+比值以及叶片相对含水量和水势,从而缓解干旱胁迫对油菜幼苗造成的渗透胁迫。但是在干旱胁迫下,与清水处理相比,ALA对根系中矿质元素吸收的影响并不显著。
5.研究了外源ALA在PEG诱导的水分缺失胁迫下对黄化的油菜子叶转绿过程中ALA积累、叶绿素合成以及相关基因表达的影响。低浓度的ALA促进了内源ALA的积累和叶绿素的合成,但其对HEMA的转录水平没有影响。水分缺失胁迫对叶绿素合成的抑制与其下调了HEMA和ChlH的表达并上调了FC的表达有关。而在水分缺失胁迫下,外源ALA处理对叶绿素合成过程中相关基因的表达并没有明显的调控作用。这些结果表明外源ALA对叶绿素生物合成的调节作用并不是发生在转录水平上。
Oilseed rape (Brassica napus L. cv. Zheshuang758) seedlings was used to investigate the effects of ALA on reactive oxygen species (ROS) scavenging system, photosynthesis, water relations, ion uptake, and chlorophyll synthesis under drought stress through physiological and molecular biological experiments. The results are as follows:
1. The interaction of5-aminolevulinic acid (ALA:0,0.1,1,10and100mg L-1) and water-deficit stress induced by polyethylene glycol (PEG6000:0,-0.3MPa) on growth and ROS scavenging system in oilseed rape (B. napus L.) seedlings was investigated. Water-deficit stress reduced chlorophyll content, and non-enzymatic antioxidants levels, while it enhanced the malondialdehyde (MDA) content, ROS production, enzymatic antioxidants activities, oxidized glutathione ratio and ascorbate ratio. Application of ALA with lower dosages (0.1-1mg L-1) by root soaking significantly increased shoot fresh weight and chlorophyll content in seedlings, whereas moderately higher dosage of ALA (10mg L-1) hampered the growth. The study also indicated that0.1and1mg L-1ALA improved chlorophyll content and reduced/oxidized glutathione ratio and ascorbate ratio as compared to the seedlings under water stress. Compared to those of PEG treatment alone,1mg L-1ALA decreased the MDA and ROS content, further enhanced the activities of antioxidant enzymes, and up-regulated the expression of POD, CAT and GR under water stress conditions. These results indicated that0.1to1mg L-1ALA could alleviate the negative effects of water-deficit stress through maintaining a relative high ratio of GSH/GSSG and ASA/DHA, enhancing the activities of the specific antioxidant enzymes and inducing the expression of the specific antioxidant enzyme genes.
2. The effects of ALA on the accumulation of reactive oxygen species (ROS), and activities and gene expression patterns of antioxidant enzymes in four-leaf stage seedlings under dehydration and rehydration conditions were studied. Seedlings were imposed to drought stress (40%of water holding capacity) and subsequently were sprayed with ALA (30mg/L), and then were rehydrated for four days. Dehydration increased MDA and H2O2contents, enhanced activities of CAT, POD, APX and GR, and up-regulated the expression of APX and GR. After rehydration, MDA and H2O2contents, and the expression of APX and GR were restored to normal level. Exogenous ALA application alleviated the accumulation of MDA and H2O2, increased the activities of CAT, POD, APX and GR, and enhanced the transcript level of CAT and POD in drought treated plants, compared to those of plants sprayed with water. Our results demonstrated that ALA could effectively activate antioxidant enzyme system through improving the activities of CAT, POD, APX and GR, and the expression of CAT and POD under drought stress.
3. The effects of exogenous foliar application of ALA on plant photosynthesis were investigated through analyzing changes in gas exchange parameters, chlorophyll fluorescence parameters, and expression levels of some Calvin cycle related genes of oilseed rape under drought stress and recovery conditions. Drought suppressed the growth of shoots and roots, decreased chlorophyll, starch and soluble protein contents, declined net photosynthetic rate (Pn) as well as maximum and actual quantum yield of photosystem II (PSII), and down-regulated the transcript levels of Calvin cycle related genes, while it stimulated the accumulation the hexose and sucrose. After rehydration for4d, the growth of drought-treated seedlings was restored to normal level for the most physiological parameters. ALA partially alleviated the growth inhibition of shoots and roots. Foliar application of ALA enhanced Pn and gs, increased the photochemical quenching (qP) and non-photochemical quenching (NPQ), induced the expression levels of Calvin cycle related genes, further improved the accumulation of hexose and sucrose accompanied with a maintenance of starch and soluble protein contents in leaves, compared to those of sprayed with water, under drought stress. These results indicate ALA may enhance the photosynthesis through increasing gs and chlorophyll content, improving photoprotection capacity and up-regulating the expression of Calvin cycle related genes.
4. The effects of ALA on water relations, osmolytes accumulation, and ion uptake under drought stress were investigated in oilseed rape seedlings. Drought stress decreased leaf relative water content (RWC) and water potential (WP), accumulating soluble sugars, free amino acid and proline in leaf and root, and inhibited the ion uptake by root. After rehydration for4d, the proline and soluble sugars contents were restored to normal level, while the ion uptake was not recovered except for Mg and Fe. Foliar application of ALA may help leaf maintain relatively higher leaf RWC and WP, decrease proline content and improve K+/Na+ratio by reducing Na+content in leaves to alleviate the osmotic stress induced by drought. However, compared to those of sprayed with water, ALA application has no significant effect on root ion uptake under drought stress.
5. The effects of exogenous ALA on chlorophyll and endogenous ALA biosynthesis in six-day-old etiolated oilseed rape cotyledons during the de-etiolated stage were investigated. ALA treatment with low dosage enhanced the accumulation of endogenous ALA and chlorophyll, while the expression level of HEMA was not affected. The decrease of chlorophyll biosynthesis induced by water-defict stress was assumed to be related to the down-regulation of HEMA and ChlH and up-regulation of FC. However, exogenous ALA has no effects on the expression of chlorophyll synthesis related genes under PEG treatment. These results suggested that exogenous ALA might not be involved in the regulation of transcriptional level in chlorophyll biosynthesis.
1.研究了根部施用ALA对聚乙二醇(PEG)诱导的水分缺失胁迫下油菜子叶期幼苗生长及活性氧清除系统的影响。PEG诱导的水分缺失胁迫抑制幼苗生长的同时增加了子叶中丙二醛(MDA)和活性氧自由基(ROS, O2和H202)的积累,提高了抗氧化酶的活性以及谷胱甘肽和抗坏血酸还原型和氧化型的比值。对根部施用低浓度的ALA (0.1-1mg L-1)提高了地上部鲜重和叶绿素含量,而10mgL-1ALA则对幼苗的生长表现出抑制作用。在水分缺失胁迫下,低浓度的ALA促进了叶绿素的积累,进一步提高了谷胱甘肽和抗坏血酸还原型和氧化型的比值。同时,与PEG单独处理相比,1mg L-1ALA显著地降低了受胁迫子叶中MDA的含量和ROS的积累,进一步提升了抗氧化酶的活性,上调了POD、CAT和GR的表达水平。以上结果表明,低浓度的ALA在水分缺失胁迫下能够通过提高ASA-GSH循环,提升抗氧化酶活性以及上调部分抗氧化酶基因的表达以保证植物能够有效地清除ROS,从而缓解子叶期幼苗受到的水分缺失胁迫。
2.研究了叶面喷施ALA对基质造成的干旱胁迫下油菜四叶期幼苗抗氧化酶活性及基因表达的影响。干旱胁迫增加了叶片中MDA和H2O2的积累,增强了CAT、 POD、 APX和GR的活性,上调了APX和GR的转录水平。在复水处理后,MDA和H202的含量以及APX和GR的表达量均恢复到正常水平。在干旱胁迫下,与对照(清水处理)相比,叶面喷施ALA有效地缓解了MDA和H2O2的积累,增加了抗氧化酶的活性,并上调了CAT和POD的表达。以上结果表明,叶片喷施ALA能通过增强CAT、 POD、 APX和GR的活性以及上调CAT和POD的表达来促进受胁迫叶片对ROS的清除。
3.研究了外源ALA对干旱胁迫下油菜幼苗光合作用的调控机理。干旱胁迫抑制了幼苗地上部和根部的生长,减少了叶片中叶绿素、淀粉以及可溶性蛋白的含量,降低了叶片净光合速率(Pn)、光系统Ⅱ(PSⅡ)的最大和实际光合量子产量,下调了卡尔文循环相关基因的表达,但却诱导了还原糖和蔗糖的积累。复水处理4d后,大多数指标均能恢复到正常水平。叶面喷施ALA能够部分地缓解干旱胁迫对油菜生长的抑制。在干旱胁迫下,与清水处理相比,ALA促进了叶绿素的积累,提高了叶片的Pn和气孔导度(gs),提升了光化学焠灭系数(qP)和非光化学焠灭系数(NPQ),上调了卡尔文循环相关基因的表达,同时在维持淀粉和可溶性蛋白含量正常水平的情况下进一步促进了还原糖和蔗糖的积累。这些结果表明,叶面喷施ALA对受干旱胁迫植株的光合作用的增强可能与气孔导度的增加、叶绿素含量的升高、植物光保护能力的提升以及卡尔文循环相关基因表达的上调有关。
4.研究了外源ALA对干旱胁迫下油菜水分关系、有机渗透物质的积累以及无机离子吸收的影响。干旱胁迫降低了油菜幼苗的叶片相对含水量和水势,诱导了叶片和根中可溶性糖、游离氨基酸和脯氨酸的积累,抑制了根系对矿质元素的吸收。复水处理4d后,叶片中脯氨酸和可溶性糖含量恢复到正常水平,但根系对矿质离子的吸收仍没有完全恢复。在干旱胁迫下,ALA降低了叶片中脯氨酸含量,促进了叶片中Na+的外排,提高了K+/Na+比值以及叶片相对含水量和水势,从而缓解干旱胁迫对油菜幼苗造成的渗透胁迫。但是在干旱胁迫下,与清水处理相比,ALA对根系中矿质元素吸收的影响并不显著。
5.研究了外源ALA在PEG诱导的水分缺失胁迫下对黄化的油菜子叶转绿过程中ALA积累、叶绿素合成以及相关基因表达的影响。低浓度的ALA促进了内源ALA的积累和叶绿素的合成,但其对HEMA的转录水平没有影响。水分缺失胁迫对叶绿素合成的抑制与其下调了HEMA和ChlH的表达并上调了FC的表达有关。而在水分缺失胁迫下,外源ALA处理对叶绿素合成过程中相关基因的表达并没有明显的调控作用。这些结果表明外源ALA对叶绿素生物合成的调节作用并不是发生在转录水平上。
Oilseed rape (Brassica napus L. cv. Zheshuang758) seedlings was used to investigate the effects of ALA on reactive oxygen species (ROS) scavenging system, photosynthesis, water relations, ion uptake, and chlorophyll synthesis under drought stress through physiological and molecular biological experiments. The results are as follows:
1. The interaction of5-aminolevulinic acid (ALA:0,0.1,1,10and100mg L-1) and water-deficit stress induced by polyethylene glycol (PEG6000:0,-0.3MPa) on growth and ROS scavenging system in oilseed rape (B. napus L.) seedlings was investigated. Water-deficit stress reduced chlorophyll content, and non-enzymatic antioxidants levels, while it enhanced the malondialdehyde (MDA) content, ROS production, enzymatic antioxidants activities, oxidized glutathione ratio and ascorbate ratio. Application of ALA with lower dosages (0.1-1mg L-1) by root soaking significantly increased shoot fresh weight and chlorophyll content in seedlings, whereas moderately higher dosage of ALA (10mg L-1) hampered the growth. The study also indicated that0.1and1mg L-1ALA improved chlorophyll content and reduced/oxidized glutathione ratio and ascorbate ratio as compared to the seedlings under water stress. Compared to those of PEG treatment alone,1mg L-1ALA decreased the MDA and ROS content, further enhanced the activities of antioxidant enzymes, and up-regulated the expression of POD, CAT and GR under water stress conditions. These results indicated that0.1to1mg L-1ALA could alleviate the negative effects of water-deficit stress through maintaining a relative high ratio of GSH/GSSG and ASA/DHA, enhancing the activities of the specific antioxidant enzymes and inducing the expression of the specific antioxidant enzyme genes.
2. The effects of ALA on the accumulation of reactive oxygen species (ROS), and activities and gene expression patterns of antioxidant enzymes in four-leaf stage seedlings under dehydration and rehydration conditions were studied. Seedlings were imposed to drought stress (40%of water holding capacity) and subsequently were sprayed with ALA (30mg/L), and then were rehydrated for four days. Dehydration increased MDA and H2O2contents, enhanced activities of CAT, POD, APX and GR, and up-regulated the expression of APX and GR. After rehydration, MDA and H2O2contents, and the expression of APX and GR were restored to normal level. Exogenous ALA application alleviated the accumulation of MDA and H2O2, increased the activities of CAT, POD, APX and GR, and enhanced the transcript level of CAT and POD in drought treated plants, compared to those of plants sprayed with water. Our results demonstrated that ALA could effectively activate antioxidant enzyme system through improving the activities of CAT, POD, APX and GR, and the expression of CAT and POD under drought stress.
3. The effects of exogenous foliar application of ALA on plant photosynthesis were investigated through analyzing changes in gas exchange parameters, chlorophyll fluorescence parameters, and expression levels of some Calvin cycle related genes of oilseed rape under drought stress and recovery conditions. Drought suppressed the growth of shoots and roots, decreased chlorophyll, starch and soluble protein contents, declined net photosynthetic rate (Pn) as well as maximum and actual quantum yield of photosystem II (PSII), and down-regulated the transcript levels of Calvin cycle related genes, while it stimulated the accumulation the hexose and sucrose. After rehydration for4d, the growth of drought-treated seedlings was restored to normal level for the most physiological parameters. ALA partially alleviated the growth inhibition of shoots and roots. Foliar application of ALA enhanced Pn and gs, increased the photochemical quenching (qP) and non-photochemical quenching (NPQ), induced the expression levels of Calvin cycle related genes, further improved the accumulation of hexose and sucrose accompanied with a maintenance of starch and soluble protein contents in leaves, compared to those of sprayed with water, under drought stress. These results indicate ALA may enhance the photosynthesis through increasing gs and chlorophyll content, improving photoprotection capacity and up-regulating the expression of Calvin cycle related genes.
4. The effects of ALA on water relations, osmolytes accumulation, and ion uptake under drought stress were investigated in oilseed rape seedlings. Drought stress decreased leaf relative water content (RWC) and water potential (WP), accumulating soluble sugars, free amino acid and proline in leaf and root, and inhibited the ion uptake by root. After rehydration for4d, the proline and soluble sugars contents were restored to normal level, while the ion uptake was not recovered except for Mg and Fe. Foliar application of ALA may help leaf maintain relatively higher leaf RWC and WP, decrease proline content and improve K+/Na+ratio by reducing Na+content in leaves to alleviate the osmotic stress induced by drought. However, compared to those of sprayed with water, ALA application has no significant effect on root ion uptake under drought stress.
5. The effects of exogenous ALA on chlorophyll and endogenous ALA biosynthesis in six-day-old etiolated oilseed rape cotyledons during the de-etiolated stage were investigated. ALA treatment with low dosage enhanced the accumulation of endogenous ALA and chlorophyll, while the expression level of HEMA was not affected. The decrease of chlorophyll biosynthesis induced by water-defict stress was assumed to be related to the down-regulation of HEMA and ChlH and up-regulation of FC. However, exogenous ALA has no effects on the expression of chlorophyll synthesis related genes under PEG treatment. These results suggested that exogenous ALA might not be involved in the regulation of transcriptional level in chlorophyll biosynthesis.
引文
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