不同氮素水平下烤烟苗期生理特性及差异蛋白组学研究
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摘要
烟草对肥料相当敏感,施肥是烟草栽培中的重要环节之一,不同的烤烟品种对氮素营养需求表现不同。本研究以烤烟品种K326、翠碧1号(CB-1)和红花大金元(以下简称红大)为试验材料,采用水培方法,研究了3个不同氮水平(低氮: 1.0 mmol/L;中氮:4.0 mmol/L;高氮:8.0 mmol/L)处理对不同烤烟品种苗期生理特性的影响,同时利用双向电泳技术分析K326苗期叶片蛋白质差异表达的影响。其结果如下:
1、对3个氮素水平处理的不同烤烟品种苗期光合特性的研究表明:随氮素水平的提高,烤烟叶片中叶绿素含量增加,烟株叶片的光合速率(Pn)加快。氮素浓度从1.0mmol/L到4.0 mmol/L时,有利于提高叶片中碳代谢相关酶的活性和增加光合作用相关产物的累积;氮素浓度进一步增加到8 mmol/L时,碳代谢相关酶的活性和光合作用相关产物的累积量有所下降。品种间比较得出,K326和CB-1的Pn对高氮水平比较敏感;3个烤烟品种的碳代谢相关酶活性均是中氮水平处理下最高,但品种间蔗糖合成酶(SS)、蔗糖磷酸合成酶(SPS)活性差异不显著;而K326的酸性转化酶(AI)活性最大,与CB-1和红大比,差异达显著水平;CB-1在低氮和中氮水平光合产物累积量最高,其次是红大,K326最低,高氮时红大最大,这表明品种间碳代谢能力存在差异。
2、增加氮素水平提高了叶片中全氮、全钾、镁、硝态氮、游离氨基酸的含量以及谷氨酰胺合成酶(GS)的活性,但对叶片中全磷和烟碱含量影响不明显。硝酸还原酶(NR)活性在中氮水平时最高,氮素过多过少均会影响NR活性。品种间比较,K326叶片中全氮、全磷和全钾的含量比CB-1和红大相比均较高,,但全氮、全磷含量差异不显著,且在高氮条件叶片中GS活性和NR活性较高;CB-1在高氮水平下,叶片中全氮、全磷、全钾、镁的含量及NR的活性最低。
3、提高氮素水平,增加了烤烟苗期叶片中过氧化氢酶(CAT)的活性,降低了丙二醛(MDA)和过氧化氢(H_2O_2)的含量,减轻了烟株膜脂氧化程度。低氮和高氮水平处理均可增加叶片超氧化物歧化酶(SOD)活性。延缓了叶片的衰老,利于烤烟苗期的生长。品种间比较,SOD活性:K326>CB-1>红大;MDA在K326中最低,其余两品种无显著差异,CAT与H_2O_2在红大中分别最低与最高,其余两品种无显著差异,说明品种间在响应氮素胁迫机制上存在的差异。
4、3个氮素水平处理下,K326苗期叶片蛋白差异表达的分析发现,有18种蛋白质对氮素响应发生差异表达,这些蛋白与光合作用和碳同化、逆境胁迫、物质和能量代谢以及功能蛋白相关。光合作用的蛋白随着氮素水平的提高出现上调表达,可见氮素可以提高光合速率;逆境胁迫相关蛋白在高氮和低氮下出现上调表达,说明缺氮和氮过剩都会诱导植株逆境蛋白的表达;与能量代谢相关的蛋白在高氮水平下上调表达,说明高氮处理烤烟苗期氮代谢旺盛。
总而言之,烤烟对氮素的响应是由外界信号诱导的。在烤烟栽培上可以通过调节烟株的氮素营养供给调控烤烟的碳氮代谢,有利于提高烤烟的适产和优质。品种之间比较,K326具有较高的磷含量,较低的蔗糖含量和可溶性糖含量,在高氮水平下,根系硝酸盐含量较高,这可能与其具有较强的耐高氮能力和“浓香型”烟叶风格的形成有关;CB-1在不同氮素水平下叶绿素含量最高,且在低氮和中氮水平下,蔗糖明显高于K326和红大,这可能有利于“清香型”烟叶风格的形成;红大在不同氮素浓度水平下,CAT活性较低,H_2O_2含量较高,且在高氮水平下Pn,全钾含量也较低,这可能与红大抗病性比较弱有关。
Tobacco is highly sensitive to fertilizer. Fertilization is one of the important link of tobacco cultivation. Different flue-cured tobacco cultivar show different nitrogen nutritional requirements. Three flue-cured tobacco varieties, K326、CB-1、Red Flower Big Jinyuan (hereinafter referred to as Red Big), were used in this study. The effects of three nitrogen level [1.0 mmol/L (low), 4 mmol/L (normal) and 8.0 mmol/L (high) NH_4NO_3] on physiological characteristics of seedlings were investigated. Also, the differentially expressed proteins of tobacco K326 leaves under different nitrogen level were studied using two-dimensional electrophoresis to study. The major results are as follow:
1. Chlorophyll (Chl) content and photosynthetic rate(Pn) in flue-cured tobacco leaves increased with increasing nitrogen supply. As nitrogen supply increased from 0.5 mmol/L to 2.0 mmol/L, the activities of carbon metabolism enzymes and the accumulation of photosynthetic product in leaves increased, then decreased under 8.0 mmol/L. By comparision, we found that photosynthesis was more sensitive to high nitrogen in K326 and CB-1 than in Big Red. The activities of carbon metabolism enzymes are highest in normal nitrogen level. No significant difference was found in sucrose synthase (SS)、sucrose phosphate synthase (SPS) among different varieties. The activity of acid invertase (AI) was significantly higher in K326 leaves than in CB-1 and Red Big leaves. This is advantage for the formation of“luzhou-flovor”smoke. On the low and normal nitrogen levels, CB-1 had the highest accumulation of photosynthetic product, followed by Red Big and K326. On the high nitrogen level, Red Big ad the highest accumulation of photosynthetic product. Based on these results, it can be concluded that the ability of the carbon metabolism varied with tobacco varieties.
2. Nitrogen supply increased the contents of total nitrogen, K~+, Mg~(2+), amino acids, NO_3~-N and the activities of glutamine synthetase (GS) in leaves, but did not significantly affect the contents of total phosphorus and nicotine. Nitrate reductase (NR) activity was the highest in normal nitrogen leaves, both low and high nitrogen decreased its activity. Total contents of N, P and K+ were higher in K326 leaves than in Red Big and CB-1 ones.In high nitrogen level,GS activity and NR activity are high. compare with K326 and Red Big ,CB-1 has lowest N ,P K~+ and Mg~(2+) the activity of NR is low also.
3. Nitrogen supply increased the activity of catalase (CAT), decreased the contents of malondialdehyde (MDA) and H_2O_2 in leaves, thus decreasing the membrane-lipid peroxidation. Both low and high nitrogen increased leaf activity of superoxide dismutase (SOD) This might help to delay aging and increase the quality of tobacco. The activity of SOD was the highest in K326 leaves, followed in CB-1 and Red Big ones. Leaf content of MDA was lower in K326 leaves than in Red Big and CB-1 ones.. The activity of CAT was lower in Red Big leaves than in K326 and CB-1 ones, while the content of H_2O_2 was higher in the former. Based on these results, it can be concluded that the mechanism in response to nitrogen is different among three varieties.
4. In tobacco K326 leaves, 18 proteins were differentially expressed under different nitrogen levels. These proteins are related to photosynthesis and carbon assimilation, stress, material and energy metabolism and function proteins. The expression of photosynthesis related proteins were up-regulated with increasing nitrogen supply, meaning that nitrogen play a significant role in promoting photosynthesis. The expression of stress-related proteins was up-regulated in both low nitrogen and high nitrogen. The expression of energy related proteins was up-regulated under high nitrogen, meaning that nitrogen metabolism in seedlings is active
In conclusion, the response of flue-cured to nitrogen was induced by external signal. In tobacco cultivation, it could change nitrogen nutrient supply to regulate the carbon and nitrogen metabolisn in order to improve yields and quality of tobacco leaves. It could be concluded that normal nitrogen is conducive to growth and the production of high quality tobacco. K326 has the highest phosphorus content, lowest sucrose and soluble sugar contents, in high notrogen level,it has highest NO_3~-N content, this may concern about strong resistance to high notrogen and the formation of“Luzhou-flavor”tobacco style ; CB-1 The highest Chl content in leaves, on low and normal nitrogen levels,has higher sucrose content than K326 and Red Big, this may concern about to format the“Qing scent”tobacco style. CAT activity and H_2O_2 content is lowest in Red Big under each nitrogen levels, Pn and k~+ content is lowest in high nitrogen level, this may relate to weaker relevant of Red Big.
1、对3个氮素水平处理的不同烤烟品种苗期光合特性的研究表明:随氮素水平的提高,烤烟叶片中叶绿素含量增加,烟株叶片的光合速率(Pn)加快。氮素浓度从1.0mmol/L到4.0 mmol/L时,有利于提高叶片中碳代谢相关酶的活性和增加光合作用相关产物的累积;氮素浓度进一步增加到8 mmol/L时,碳代谢相关酶的活性和光合作用相关产物的累积量有所下降。品种间比较得出,K326和CB-1的Pn对高氮水平比较敏感;3个烤烟品种的碳代谢相关酶活性均是中氮水平处理下最高,但品种间蔗糖合成酶(SS)、蔗糖磷酸合成酶(SPS)活性差异不显著;而K326的酸性转化酶(AI)活性最大,与CB-1和红大比,差异达显著水平;CB-1在低氮和中氮水平光合产物累积量最高,其次是红大,K326最低,高氮时红大最大,这表明品种间碳代谢能力存在差异。
2、增加氮素水平提高了叶片中全氮、全钾、镁、硝态氮、游离氨基酸的含量以及谷氨酰胺合成酶(GS)的活性,但对叶片中全磷和烟碱含量影响不明显。硝酸还原酶(NR)活性在中氮水平时最高,氮素过多过少均会影响NR活性。品种间比较,K326叶片中全氮、全磷和全钾的含量比CB-1和红大相比均较高,,但全氮、全磷含量差异不显著,且在高氮条件叶片中GS活性和NR活性较高;CB-1在高氮水平下,叶片中全氮、全磷、全钾、镁的含量及NR的活性最低。
3、提高氮素水平,增加了烤烟苗期叶片中过氧化氢酶(CAT)的活性,降低了丙二醛(MDA)和过氧化氢(H_2O_2)的含量,减轻了烟株膜脂氧化程度。低氮和高氮水平处理均可增加叶片超氧化物歧化酶(SOD)活性。延缓了叶片的衰老,利于烤烟苗期的生长。品种间比较,SOD活性:K326>CB-1>红大;MDA在K326中最低,其余两品种无显著差异,CAT与H_2O_2在红大中分别最低与最高,其余两品种无显著差异,说明品种间在响应氮素胁迫机制上存在的差异。
4、3个氮素水平处理下,K326苗期叶片蛋白差异表达的分析发现,有18种蛋白质对氮素响应发生差异表达,这些蛋白与光合作用和碳同化、逆境胁迫、物质和能量代谢以及功能蛋白相关。光合作用的蛋白随着氮素水平的提高出现上调表达,可见氮素可以提高光合速率;逆境胁迫相关蛋白在高氮和低氮下出现上调表达,说明缺氮和氮过剩都会诱导植株逆境蛋白的表达;与能量代谢相关的蛋白在高氮水平下上调表达,说明高氮处理烤烟苗期氮代谢旺盛。
总而言之,烤烟对氮素的响应是由外界信号诱导的。在烤烟栽培上可以通过调节烟株的氮素营养供给调控烤烟的碳氮代谢,有利于提高烤烟的适产和优质。品种之间比较,K326具有较高的磷含量,较低的蔗糖含量和可溶性糖含量,在高氮水平下,根系硝酸盐含量较高,这可能与其具有较强的耐高氮能力和“浓香型”烟叶风格的形成有关;CB-1在不同氮素水平下叶绿素含量最高,且在低氮和中氮水平下,蔗糖明显高于K326和红大,这可能有利于“清香型”烟叶风格的形成;红大在不同氮素浓度水平下,CAT活性较低,H_2O_2含量较高,且在高氮水平下Pn,全钾含量也较低,这可能与红大抗病性比较弱有关。
Tobacco is highly sensitive to fertilizer. Fertilization is one of the important link of tobacco cultivation. Different flue-cured tobacco cultivar show different nitrogen nutritional requirements. Three flue-cured tobacco varieties, K326、CB-1、Red Flower Big Jinyuan (hereinafter referred to as Red Big), were used in this study. The effects of three nitrogen level [1.0 mmol/L (low), 4 mmol/L (normal) and 8.0 mmol/L (high) NH_4NO_3] on physiological characteristics of seedlings were investigated. Also, the differentially expressed proteins of tobacco K326 leaves under different nitrogen level were studied using two-dimensional electrophoresis to study. The major results are as follow:
1. Chlorophyll (Chl) content and photosynthetic rate(Pn) in flue-cured tobacco leaves increased with increasing nitrogen supply. As nitrogen supply increased from 0.5 mmol/L to 2.0 mmol/L, the activities of carbon metabolism enzymes and the accumulation of photosynthetic product in leaves increased, then decreased under 8.0 mmol/L. By comparision, we found that photosynthesis was more sensitive to high nitrogen in K326 and CB-1 than in Big Red. The activities of carbon metabolism enzymes are highest in normal nitrogen level. No significant difference was found in sucrose synthase (SS)、sucrose phosphate synthase (SPS) among different varieties. The activity of acid invertase (AI) was significantly higher in K326 leaves than in CB-1 and Red Big leaves. This is advantage for the formation of“luzhou-flovor”smoke. On the low and normal nitrogen levels, CB-1 had the highest accumulation of photosynthetic product, followed by Red Big and K326. On the high nitrogen level, Red Big ad the highest accumulation of photosynthetic product. Based on these results, it can be concluded that the ability of the carbon metabolism varied with tobacco varieties.
2. Nitrogen supply increased the contents of total nitrogen, K~+, Mg~(2+), amino acids, NO_3~-N and the activities of glutamine synthetase (GS) in leaves, but did not significantly affect the contents of total phosphorus and nicotine. Nitrate reductase (NR) activity was the highest in normal nitrogen leaves, both low and high nitrogen decreased its activity. Total contents of N, P and K+ were higher in K326 leaves than in Red Big and CB-1 ones.In high nitrogen level,GS activity and NR activity are high. compare with K326 and Red Big ,CB-1 has lowest N ,P K~+ and Mg~(2+) the activity of NR is low also.
3. Nitrogen supply increased the activity of catalase (CAT), decreased the contents of malondialdehyde (MDA) and H_2O_2 in leaves, thus decreasing the membrane-lipid peroxidation. Both low and high nitrogen increased leaf activity of superoxide dismutase (SOD) This might help to delay aging and increase the quality of tobacco. The activity of SOD was the highest in K326 leaves, followed in CB-1 and Red Big ones. Leaf content of MDA was lower in K326 leaves than in Red Big and CB-1 ones.. The activity of CAT was lower in Red Big leaves than in K326 and CB-1 ones, while the content of H_2O_2 was higher in the former. Based on these results, it can be concluded that the mechanism in response to nitrogen is different among three varieties.
4. In tobacco K326 leaves, 18 proteins were differentially expressed under different nitrogen levels. These proteins are related to photosynthesis and carbon assimilation, stress, material and energy metabolism and function proteins. The expression of photosynthesis related proteins were up-regulated with increasing nitrogen supply, meaning that nitrogen play a significant role in promoting photosynthesis. The expression of stress-related proteins was up-regulated in both low nitrogen and high nitrogen. The expression of energy related proteins was up-regulated under high nitrogen, meaning that nitrogen metabolism in seedlings is active
In conclusion, the response of flue-cured to nitrogen was induced by external signal. In tobacco cultivation, it could change nitrogen nutrient supply to regulate the carbon and nitrogen metabolisn in order to improve yields and quality of tobacco leaves. It could be concluded that normal nitrogen is conducive to growth and the production of high quality tobacco. K326 has the highest phosphorus content, lowest sucrose and soluble sugar contents, in high notrogen level,it has highest NO_3~-N content, this may concern about strong resistance to high notrogen and the formation of“Luzhou-flavor”tobacco style ; CB-1 The highest Chl content in leaves, on low and normal nitrogen levels,has higher sucrose content than K326 and Red Big, this may concern about to format the“Qing scent”tobacco style. CAT activity and H_2O_2 content is lowest in Red Big under each nitrogen levels, Pn and k~+ content is lowest in high nitrogen level, this may relate to weaker relevant of Red Big.
引文
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