甜菜苗期对干旱适应的生理生化及分子机制研究
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摘要
甜菜是世界上重要的糖料作物,在目前甜菜生产中,干旱已成为制约其生产和产业发展的重要因素,降低甜菜对水分缺乏的敏感性和提高其抗(耐)旱性已成为甜菜生产亟待解决的难题。因此,深入研究甜菜抗旱机制,对加速甜菜抗旱育种,扩大甜菜种植范围,提高其生产潜力,促进干旱、半干旱地区的经济发展意义重大。本论文对国内外30份甜菜种质资源的抗旱性进行了综合评价,依据评价结果选出抗旱性强和抗旱性弱的两个品种为试验材料,对其抗旱生理生化及分子机制进行系统研究,取得如下主要研究结果:
1、通过反复干旱存活率鉴定与多指标主成分和隶属函数分析相结合,对30份甜菜种质资源抗旱性进行综合评价,并通过田间试验对评价结果进行了验证,从中鉴定并筛选出抗旱性较强的材料6份,抗旱性较弱的材料4份,20份为中间型,并为甜菜种质资源苗期抗旱性大规模筛选提供了新技术。
2、干旱胁迫下,通过诱导保护酶系基因的表达,在保护酶系SOD、POD及抗氧化性物质Car、GSH的共同协调作用下,使抗旱材料HI0466细胞结构得到很好的保护,表现为活性氧可维持在较低水平,膜脂过氧化程度小,使细胞膜透性小,抗失水能力强,表现出较强的抗旱性。
3、水分胁迫下,脯氨酸、甜菜碱、可溶性糖、可溶性蛋白在渗透调节中的作用显著,其中脯氨酸、甜菜碱在P5CS基因、BADH基因的实时、过量表达下,成为两种甜菜的主要渗透调节物质,其含量的增加及增幅的增强可提高甜菜抗旱性。
4、干旱胁迫会影响甜菜的光合作用并产生光抑制。干旱胁迫下,净光合速率、PS II原初光能转化效率降低,PS II潜在活性中心受损。相比之下,抗旱性强的甜菜净光合速率下降的幅度小,复水后自身修复能力强,具有较为完善的非光化学淬灭机制,可通过对过剩光能的消耗来避免或减轻光合机构受损,使抵抗干旱的能力在一定程度得以提高。同时发现轻度水分胁迫时气孔限制是影响光合作用的主要因素,而胁迫加重时非气孔限制占主导。
5、水分胁迫下甜菜ABA含量有不同程度的升高,而IAA含量在根系表现为先升高后降低。通过ABA的显著增加可增强对干旱胁迫的调节,而根系IAA含量在胁迫初期的提高可使其根系生长潜能增强。在水分胁迫下两种激素相互协调,趋于促进根系生长和加速气孔关闭,抗旱甜菜在胁迫初期趋向于促进根系生长的能力较强,而在重度胁迫下通过减缓生长速率、促进气孔关闭来适应干旱胁迫的优势明显。
6、在水分胁迫下,分子量约为54KD的蛋白与甜菜体内水分的变化关系较为密切,抗旱性强的甜菜在胁迫后期该蛋白大量出现,推测其可能参与了抗旱过程。质谱分析表明其主要成分为核酮糖-1,5-二磷酸羧化酶,在水分胁迫过程中可对光合作用进行调控来增强对干旱的适应性。
Sugarbeet(Beta vulgaris L.) is an important sugar crop in the world, but its yield and quality are attacked severely by drought stress. Reducing it’s sensitivity towards water deficiency and increasing the drought resistance will be important field of sugar beet breeding research. So, making known the mechanism of drought resistance during seedling stage, speeding up the proceedings of drought resistance breeding research are great significantly. The paper got comprehensive evaluation on the drought resistance of 30 sugarbeet genetic resources from home and abroad, two varieties (represented respectively the high and low drought resistance)are selected for study materials according to the above mentioned evaluation results. The physiological, biochemical and molecular mechanisms of the adaptation to drought stress are analyzed widely. The major results studied in the paper are abstracted as follows:
1、The drought resistance of 30 sugarbeet genetic resources are evaluated comprehensively by the viability test combining principal component analysis and factor analysis, verification is conducted by field experiments, which provided new technology for drought–resistance screening during seedling stage for sugar beet.
2、By the combined action of the protective enzyme systems (SOD and POD) and antioxidants (Car and GSH), the HI0466 genotype showed higher drought resistance due to the better cell structure protection with significant lower active oxygen and less membrane lipid peroxidation production etc..
3、The action of proline, betaine, soluble sugar and soluble protein in osmotic adjustment were extremely remarkable, the first two factors of which(proline and betaine) became the main osmotic adjustment substances with the P5CS and BADH genes’real–time expression.
4、Drought stress may interfere with sugar beet photosynthesis, causing Photoinhibition. The net photosynthesis rate and primary light energy translation of PSⅡwere decreased under drought stress but the variety with higher drought resistance was less affected, which can repair itself after flooding and had higher non-photochemical quenching. The results also showed that stomata was the major influencing factor on photosynthesis under slight drought stress, while the opposite for serious one.
5、The ABA content increased both in the two beet genotype and IAA content increased first, and then decreased under drought stress, this trend contributed to the drought stress regulation. The two hormones work cooperatively to enhance root growth and stomatal closure.
6、The changes of water in sugar beet were closely related to 54KD protein under drought stress. The 54KD protein may participate to drought resistance process presenting in large quantity during the later period of stress. Mass Spectrometric Analysis showed that its main component was ribulose-1,5-bisphosphate carboxylase(Rubisco), which can regulate photosynthesis to enhance the drought resistance adaption.
1、通过反复干旱存活率鉴定与多指标主成分和隶属函数分析相结合,对30份甜菜种质资源抗旱性进行综合评价,并通过田间试验对评价结果进行了验证,从中鉴定并筛选出抗旱性较强的材料6份,抗旱性较弱的材料4份,20份为中间型,并为甜菜种质资源苗期抗旱性大规模筛选提供了新技术。
2、干旱胁迫下,通过诱导保护酶系基因的表达,在保护酶系SOD、POD及抗氧化性物质Car、GSH的共同协调作用下,使抗旱材料HI0466细胞结构得到很好的保护,表现为活性氧可维持在较低水平,膜脂过氧化程度小,使细胞膜透性小,抗失水能力强,表现出较强的抗旱性。
3、水分胁迫下,脯氨酸、甜菜碱、可溶性糖、可溶性蛋白在渗透调节中的作用显著,其中脯氨酸、甜菜碱在P5CS基因、BADH基因的实时、过量表达下,成为两种甜菜的主要渗透调节物质,其含量的增加及增幅的增强可提高甜菜抗旱性。
4、干旱胁迫会影响甜菜的光合作用并产生光抑制。干旱胁迫下,净光合速率、PS II原初光能转化效率降低,PS II潜在活性中心受损。相比之下,抗旱性强的甜菜净光合速率下降的幅度小,复水后自身修复能力强,具有较为完善的非光化学淬灭机制,可通过对过剩光能的消耗来避免或减轻光合机构受损,使抵抗干旱的能力在一定程度得以提高。同时发现轻度水分胁迫时气孔限制是影响光合作用的主要因素,而胁迫加重时非气孔限制占主导。
5、水分胁迫下甜菜ABA含量有不同程度的升高,而IAA含量在根系表现为先升高后降低。通过ABA的显著增加可增强对干旱胁迫的调节,而根系IAA含量在胁迫初期的提高可使其根系生长潜能增强。在水分胁迫下两种激素相互协调,趋于促进根系生长和加速气孔关闭,抗旱甜菜在胁迫初期趋向于促进根系生长的能力较强,而在重度胁迫下通过减缓生长速率、促进气孔关闭来适应干旱胁迫的优势明显。
6、在水分胁迫下,分子量约为54KD的蛋白与甜菜体内水分的变化关系较为密切,抗旱性强的甜菜在胁迫后期该蛋白大量出现,推测其可能参与了抗旱过程。质谱分析表明其主要成分为核酮糖-1,5-二磷酸羧化酶,在水分胁迫过程中可对光合作用进行调控来增强对干旱的适应性。
Sugarbeet(Beta vulgaris L.) is an important sugar crop in the world, but its yield and quality are attacked severely by drought stress. Reducing it’s sensitivity towards water deficiency and increasing the drought resistance will be important field of sugar beet breeding research. So, making known the mechanism of drought resistance during seedling stage, speeding up the proceedings of drought resistance breeding research are great significantly. The paper got comprehensive evaluation on the drought resistance of 30 sugarbeet genetic resources from home and abroad, two varieties (represented respectively the high and low drought resistance)are selected for study materials according to the above mentioned evaluation results. The physiological, biochemical and molecular mechanisms of the adaptation to drought stress are analyzed widely. The major results studied in the paper are abstracted as follows:
1、The drought resistance of 30 sugarbeet genetic resources are evaluated comprehensively by the viability test combining principal component analysis and factor analysis, verification is conducted by field experiments, which provided new technology for drought–resistance screening during seedling stage for sugar beet.
2、By the combined action of the protective enzyme systems (SOD and POD) and antioxidants (Car and GSH), the HI0466 genotype showed higher drought resistance due to the better cell structure protection with significant lower active oxygen and less membrane lipid peroxidation production etc..
3、The action of proline, betaine, soluble sugar and soluble protein in osmotic adjustment were extremely remarkable, the first two factors of which(proline and betaine) became the main osmotic adjustment substances with the P5CS and BADH genes’real–time expression.
4、Drought stress may interfere with sugar beet photosynthesis, causing Photoinhibition. The net photosynthesis rate and primary light energy translation of PSⅡwere decreased under drought stress but the variety with higher drought resistance was less affected, which can repair itself after flooding and had higher non-photochemical quenching. The results also showed that stomata was the major influencing factor on photosynthesis under slight drought stress, while the opposite for serious one.
5、The ABA content increased both in the two beet genotype and IAA content increased first, and then decreased under drought stress, this trend contributed to the drought stress regulation. The two hormones work cooperatively to enhance root growth and stomatal closure.
6、The changes of water in sugar beet were closely related to 54KD protein under drought stress. The 54KD protein may participate to drought resistance process presenting in large quantity during the later period of stress. Mass Spectrometric Analysis showed that its main component was ribulose-1,5-bisphosphate carboxylase(Rubisco), which can regulate photosynthesis to enhance the drought resistance adaption.
引文
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