不同玉米骨干自交系氮素利用的差异蛋白质组学初步研究
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摘要
玉米(Zea mays L)是世界上最重要的食粮之一,平均单产居首位。就目前全球情况而言,增加氮肥的施用量是提高玉米产量的一条重要措施。然而氮肥施用只能在一定程度上增加其产量,施用过量不但给生产者带来更多的经济负担,而且带来严重的环境问题。我国西南地区玉米主要种植于较贫瘠的土壤中。因此研究玉米的骨干自交系,尤其是对氮素反应敏感度不同的自交系,氮素利用差异的机理,有利于搞清楚玉米耐低氮品种在低氮胁迫下启动应急的机制,从而为选育高肥料利用效率品种奠定基础。
本研究选取在育种中广泛应用玉米骨干自交系耐瘠薄的黄早四和不耐瘠薄的Mo17为材料。研究二者在无土栽培条件下以硝酸钙为唯一氮源且施氮浓度梯度为0mmol/L、2mmol/L、4mmol/L、8mmol/L下株型、干物质积累差异,并以双向电泳技术分别研究在上述栽培条件下黄早四、Mo17叶片蛋白质组表达的差异,得到以下结论:
1.黄早四在前期的营养生长中株高叶面积不如Mo17,干物质积累量也比Mo17少,但是在后期成熟时的干物质积累总量和Mo17相当,生殖生长的干物质积累比Mo17多,并且每个生长时期黄早四在不同施氮水平间的差异表现均没有Mo17明显,黄早四在4mmol/L时能够获得与8mmol/L相近的生物学产量,据此推断黄早四比Mo17更具有耐瘠薄的特点。
2.黄早四、Mo17在苗期和拔节期的各个施氮水平叶片表达的蛋白质组进行双向电泳分析,检测到各个时期Mo17表达的蛋白点均少于黄早四。这可能与黄早四比Mo17更耐瘠薄有关。分别分析相同自交系同一时期不同施氮水平的双向电泳图谱,对其中28个差异表达的蛋白点进行肽质量指纹图谱分析,结果表明28个蛋白中包括了大量的未知蛋白,其中27个蛋白点与玉米以外的其他植物种属同源。
3.黄早四和Mo17拔节期均检验到1,5-二磷酸核酮糖羧化酶/加氧酶大亚基且这一蛋白的表达量在黄早四和Mo17中均随着施氮浓度的增加呈上升的趋势。在相同施氮水平下黄早四1,5-二磷酸核酮糖羧化酶/加氧酶大亚基表达量高于Mo17,这可能是黄早四比Mo17更耐低氮胁迫的原因之一。
4.在黄早四苗期和Mo17的拔节期均检测到了半胱氨酸蛋白酶抑制蛋白,证明半胱氨酸蛋白酶抑制蛋白作为一种逆境诱导蛋白,对玉米氮素胁迫起作用。在黄早四苗期中半胱氨酸蛋白酶抑制蛋白随着施氮浓度的增加表达量呈献上升的趋势,而Mo17拔节期的情况正好相反。这可能是黄早四对施氮水平较Mo17不敏感的原因。
5.在Mo17拔节期检测到抗病基因编码的一类NBS-LRR蛋白,这种蛋白首次在玉米的氮素胁迫中被检测到。推测该蛋白对玉米对氮素的反应起作用。但这一蛋白质在氮代谢过程中的具体功能以及如何参与整个过程,还需要进一步研究。
Maize (Zea mays L) is one of the most important foods and the average yield is highest in the world. All over the world, increase the nitrogen fertilizer application rate is an important measure to increase maize production right now. However, application of nitrogen fertilizer only increase yield to a certain extent, excessive use not only bring more economic burden to producers and caused serious environmental problems. Study the Elite Maize Inbred Lines with different sensitivity of nitrogen response especially and the different mechanism of nitrogen use are conducive to clear the emergency activate mechanism in low nitrogen stress of bear low nitrogen corn variety, lay the foundation for breeding the variety with high use efficiency of fertilizer.
Huangzaosi and Mo17 the Elite Maize Inbred Lines was studied in this paper, they were widely used in breeding, Huangzaosi was seen as bear low nitrogen variety and Mo17 was not bear low nitrogen. Study the difference of the plant type and dry matter accumulation between these two varieties under taking the Ca(NO_3)_2 for the nitrogen source and setting a gradient of 0mmol /L、2mmol / L、4mmol / L、8mmol / L soil-free cultivation condition. After that using 2D electrophoresis technology analyze the differences of the leaf proteome expression. The follows are the results:
1. In the earlier vegetative growth period Huangzaosi's leaf area is smaller, length is shorter and the material accumulation quantity is also less than Mo17. But in later period of mature time Huangzaosi's dry material accumulation quantity are almost the same to Mo17. This means Huangzaosi accumulation more dry material than Mo17 in mature time. And Huangzaosi's differences exhibitions are less than Mo17 between gradients. Huangzaosi have almost the same biological productions between gradients 4mmol/L and 8mmol / L. So mine infers according to the above is Huangzoasi is not as sensitive as Mo17 to the nitrogen.
2. Analysis Huangzaosi and Mo17 leaf expressing proteome in each nitrogrn levels during seeding stage and shooting stage with 2D-electrophoretic. The result shows that Huangzaosi expressing more protein spots than Mo17, maybe this is the reason why Mo17 is as sensitive as Huangzaosi to the nitrogen. Analysis the same inbreeding line 2D picture, using peptide mass fingerprinting analysis 28 spots the result shows that most of the protein spots are unknown protein and come from other kind of plants.
3. Ribulose-1,5-bisphosphate carboxylase/ oxygenase large subunit has been found in both Huangzaosi and Mo17 in the seeding stage. And the ribulose-1,5-bisphosphate carboxylase / oxygenase large subunit expression level increased with the nitrogen gradient in both Huangzaosi and Mo17. In the same nitrogen gradient ribulose-l,5-bisphosphate carboxylase / oxygenase large subunit expression level in Huangzaosi is higher than Mol7. This maybe the reason Huangzoasi is not as sensitive as Mo17 between the nitrogen levels.
4. Cysteine protease inhibitor is found in both seeding stage of Huangzoasi and jointing stage of Mo17. This proved the cysteine protease inhibitor is as a kind of adversity stress protein also has effect in maize stress resistance. In the seeding stage of Huangzoasi the cysteine protease inhibitor expression level is increased with the nitrogen gradient, but in the jointing stage of Mo17 the cysteine protease inhibitor expression level is decreased with the nitrogen gradient. This maybe another reason for Huangzoasi is not as sensitive as Mo17 between the nitrogen levels.
5. A class of NBS-LRR protein which is known coding by disease-resistant gene is found in Mo17 jointing stage, this is the first time this protein found in adversity stress maize. Extrapolation this protein has an effect to corn's resistance. But how this protein participates in the entire process of nitrogen metabolism, and the protein's function are not known. So further study about this protein are need.
本研究选取在育种中广泛应用玉米骨干自交系耐瘠薄的黄早四和不耐瘠薄的Mo17为材料。研究二者在无土栽培条件下以硝酸钙为唯一氮源且施氮浓度梯度为0mmol/L、2mmol/L、4mmol/L、8mmol/L下株型、干物质积累差异,并以双向电泳技术分别研究在上述栽培条件下黄早四、Mo17叶片蛋白质组表达的差异,得到以下结论:
1.黄早四在前期的营养生长中株高叶面积不如Mo17,干物质积累量也比Mo17少,但是在后期成熟时的干物质积累总量和Mo17相当,生殖生长的干物质积累比Mo17多,并且每个生长时期黄早四在不同施氮水平间的差异表现均没有Mo17明显,黄早四在4mmol/L时能够获得与8mmol/L相近的生物学产量,据此推断黄早四比Mo17更具有耐瘠薄的特点。
2.黄早四、Mo17在苗期和拔节期的各个施氮水平叶片表达的蛋白质组进行双向电泳分析,检测到各个时期Mo17表达的蛋白点均少于黄早四。这可能与黄早四比Mo17更耐瘠薄有关。分别分析相同自交系同一时期不同施氮水平的双向电泳图谱,对其中28个差异表达的蛋白点进行肽质量指纹图谱分析,结果表明28个蛋白中包括了大量的未知蛋白,其中27个蛋白点与玉米以外的其他植物种属同源。
3.黄早四和Mo17拔节期均检验到1,5-二磷酸核酮糖羧化酶/加氧酶大亚基且这一蛋白的表达量在黄早四和Mo17中均随着施氮浓度的增加呈上升的趋势。在相同施氮水平下黄早四1,5-二磷酸核酮糖羧化酶/加氧酶大亚基表达量高于Mo17,这可能是黄早四比Mo17更耐低氮胁迫的原因之一。
4.在黄早四苗期和Mo17的拔节期均检测到了半胱氨酸蛋白酶抑制蛋白,证明半胱氨酸蛋白酶抑制蛋白作为一种逆境诱导蛋白,对玉米氮素胁迫起作用。在黄早四苗期中半胱氨酸蛋白酶抑制蛋白随着施氮浓度的增加表达量呈献上升的趋势,而Mo17拔节期的情况正好相反。这可能是黄早四对施氮水平较Mo17不敏感的原因。
5.在Mo17拔节期检测到抗病基因编码的一类NBS-LRR蛋白,这种蛋白首次在玉米的氮素胁迫中被检测到。推测该蛋白对玉米对氮素的反应起作用。但这一蛋白质在氮代谢过程中的具体功能以及如何参与整个过程,还需要进一步研究。
Maize (Zea mays L) is one of the most important foods and the average yield is highest in the world. All over the world, increase the nitrogen fertilizer application rate is an important measure to increase maize production right now. However, application of nitrogen fertilizer only increase yield to a certain extent, excessive use not only bring more economic burden to producers and caused serious environmental problems. Study the Elite Maize Inbred Lines with different sensitivity of nitrogen response especially and the different mechanism of nitrogen use are conducive to clear the emergency activate mechanism in low nitrogen stress of bear low nitrogen corn variety, lay the foundation for breeding the variety with high use efficiency of fertilizer.
Huangzaosi and Mo17 the Elite Maize Inbred Lines was studied in this paper, they were widely used in breeding, Huangzaosi was seen as bear low nitrogen variety and Mo17 was not bear low nitrogen. Study the difference of the plant type and dry matter accumulation between these two varieties under taking the Ca(NO_3)_2 for the nitrogen source and setting a gradient of 0mmol /L、2mmol / L、4mmol / L、8mmol / L soil-free cultivation condition. After that using 2D electrophoresis technology analyze the differences of the leaf proteome expression. The follows are the results:
1. In the earlier vegetative growth period Huangzaosi's leaf area is smaller, length is shorter and the material accumulation quantity is also less than Mo17. But in later period of mature time Huangzaosi's dry material accumulation quantity are almost the same to Mo17. This means Huangzaosi accumulation more dry material than Mo17 in mature time. And Huangzaosi's differences exhibitions are less than Mo17 between gradients. Huangzaosi have almost the same biological productions between gradients 4mmol/L and 8mmol / L. So mine infers according to the above is Huangzoasi is not as sensitive as Mo17 to the nitrogen.
2. Analysis Huangzaosi and Mo17 leaf expressing proteome in each nitrogrn levels during seeding stage and shooting stage with 2D-electrophoretic. The result shows that Huangzaosi expressing more protein spots than Mo17, maybe this is the reason why Mo17 is as sensitive as Huangzaosi to the nitrogen. Analysis the same inbreeding line 2D picture, using peptide mass fingerprinting analysis 28 spots the result shows that most of the protein spots are unknown protein and come from other kind of plants.
3. Ribulose-1,5-bisphosphate carboxylase/ oxygenase large subunit has been found in both Huangzaosi and Mo17 in the seeding stage. And the ribulose-1,5-bisphosphate carboxylase / oxygenase large subunit expression level increased with the nitrogen gradient in both Huangzaosi and Mo17. In the same nitrogen gradient ribulose-l,5-bisphosphate carboxylase / oxygenase large subunit expression level in Huangzaosi is higher than Mol7. This maybe the reason Huangzoasi is not as sensitive as Mo17 between the nitrogen levels.
4. Cysteine protease inhibitor is found in both seeding stage of Huangzoasi and jointing stage of Mo17. This proved the cysteine protease inhibitor is as a kind of adversity stress protein also has effect in maize stress resistance. In the seeding stage of Huangzoasi the cysteine protease inhibitor expression level is increased with the nitrogen gradient, but in the jointing stage of Mo17 the cysteine protease inhibitor expression level is decreased with the nitrogen gradient. This maybe another reason for Huangzoasi is not as sensitive as Mo17 between the nitrogen levels.
5. A class of NBS-LRR protein which is known coding by disease-resistant gene is found in Mo17 jointing stage, this is the first time this protein found in adversity stress maize. Extrapolation this protein has an effect to corn's resistance. But how this protein participates in the entire process of nitrogen metabolism, and the protein's function are not known. So further study about this protein are need.
引文
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