生长调节剂对油菜氮素再利用的影响及其与氮素利用效率的关系研究
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摘要
作为一种重要的有着传统种植优势的油料作物,油菜已经越来越受到大家的关注。随着高产优质新品种的不断涌现,养分利用效率低已成为限制油菜生产进一步发展的新瓶颈,尤其是油菜对氮素的需求量大,而利用率较低,直接制约着油菜产业的发展。因此,如何提高油菜氮素利用率已经成为整个植物营养学研究领域的热点。在植物的整个生长发育期间,不论是生长代谢还是形态建成方面,植物激素都起着重要的调节作用。但是,植物激素对油菜生长后期氮素再利用、进而对油菜氮效率的调节作用方面的研究目前报道较少。
本试验采用两个油菜品种(分别用A和B表示),在常氮和氮胁迫两个供氮水平下,研究角果发育期涂抹不同生长调节剂(生长素IAA、细胞分裂素6-BA及脱落酸ABA)溶液于角果,对油菜生长后期氮素再利用和氮效率的影响及其影响机理,为揭示油菜氮效率的生理机制,探索提高油菜氮效率的可行性途径提供科学依据。
1.油菜产量、产量构成因素及氮素生理效率的测定结果表明,三种生长调节剂均能不同程度地提高籽粒产量、氮素生理效率、单株角果数和每角果粒数,但对于千粒重没有产生什么影响。不同生长调节剂对以上各指标的影响不同,ABA主要对每角果粒数增效作用大,IAA和6-BA则主要对籽粒产量和单株角果数贡献较大。各生长调节剂对同一指标的影响因氮水平高低和品种不同而有所差异,以籽粒产量为基础的氮素生理效率为例,A品种在常氮条件下以ABA效应好,氮胁迫条件下以6-BA的效应最好；而B品种则均以IAA的效果最好。另外,常氮水平下的生物产量氮素生理效率及籽粒产量氮素生理效率低于氮胁迫水平。比较两个供氮水平的籽粒产量、产量构成因素及氮素生理效率,籽粒产量和单株角果数,常氮条件高于氮胁迫条件；氮素生理效率则常氮条件低于氮胁迫条件；千粒重和每角果粒数在两个氮水平之间没有差异。
2.油菜生长后期氮素转运比例和转运量的测定结果表明,三种生长调节剂处理后,氮素向籽粒的转运比例和转运量增加,损失量减少。但不同生长调节剂的影响程度因供氮水平和品种不同而有所不同。从氮素转运再分配的增加效果来看,常氮条件下IAA对A品种的贡献大,ABA对B品种贡献大,这种优势同时也表现在较低的氮素损失量方面；氮胁迫条件下,A、B两品种均是以IAA的贡献大。比较两个供氮水平的氮素转运比例及转运量,无论是向籽粒还是向角果的转运,两个品种均表现为常氮条件下高于氮胁迫条件。比较两个品种的氮素转运比例及转运量,无论是向籽粒还是向角果的转运,均表现为A品种的转运比例及转运量略高于B品种,但差异不显著。
3.油菜生长后期氮素转运相关酶活性的测定结果表明,三种生长调节剂都可不同程度地提高氮素转运相关酶活性。其中,对蛋白水解酶提高效应最好的是ABA；而IAA对谷氨酰胺合成酶活性和谷氨酸合成酶活性有相对好的提高效果；6-BA对酶的活性提高幅度不如前两种生长调节剂大。氮水平和品种对三种酶的活性的影响来看,总体上表现为常氮条件低于氮胁迫条件、品种A高于品种B的趋势。
4.韧皮部汁液成分的测定结果表明,三种生长调节剂都可不同程度地增加韧皮部汁液中的L-谷氨酰胺含量、可溶性糖含量和游离氨基酸含量。IAA和ABA对L-谷氨酰胺含量的提高作用相对较大,6-BA对可溶性糖含量的提高作用相对大,三种生长调节剂对游离氨基酸含量的影响,因品种而异。氮水平和品种对韧皮部汁液成分的影响来看,L-谷氨酰胺含量为常氮条件高于氮胁迫条件、可溶性糖含量和游离氨基酸含量在两氮素水平间几乎没有差异；L-谷氨酰胺含量和可溶性糖含量表现为A品种略低于B品种,游离氨基酸含量在两个品种之间差异性不大。
5.对油菜籽粒蛋白质含量和油分含量的测定结果表明,生长调节剂能增加油菜籽粒蛋白质含量及蛋白质产量,但对油分含量的影响较小,但因为生长调节剂增加了籽粒产量,因而能显著提高油分产量。从各生长调节剂对提高油分产量方面的贡献来看,A品种在常氮水平下以6-BA的贡献较大,氮胁迫条件下以ABA的贡献较大；而B品种在两种供氮水平下均以IAA的贡献较大。从氮水平和品种对籽粒蛋白质含量和油分含量的影响来看,蛋白质含量、蛋白质产量和油分产量表现为常氮条件高于氮胁迫条件,油分含量则反之；蛋白质含量、蛋白质产量和油分产量A品种均高于B品种,油分含量则在两个品种之间没有差异。
Oilseed Rape, as a significant oil crop with planting advantage, has aroused widespread concern increasingly. With the new oilseed rape variety springing up possessing high yield and high quality, low nutrient utilization efficiency has become a new bottleneck, which constrain the further development of oilseed rape production. Especially because of high nitrogen (N) demand and low utilization rate, the development of rape career are restricted. As a result, how to raise the N fertilizer efficiency has become a central issue in plant nutrition research area. In the whole plant growth and development period, phytohormones play a critical regulatory role in the process of physiological metabolism and morphogenesis. However, effect of phytohormones on N reutilization and N efficiency regulation were few reported in recently.
The experiment was conducted by daubing solutions of different growth regulators such as auxin(IAA), cytokinin(6-BA) and abscisic acid(ABA) to rape pod with two rapeseed varieties(A and B) under normal and deficient N conditions. The effect of N reutilization and efficiency during later stages of rape and corresponding mechanisms were studied, aiming at revealing physiological mechanisms of N utilization efficiency in rape, and providing a scientific basis for improving N utilization efficiency in rape.
The results of the yield, yield components and nitrogen physiological efficiency indicated that the growth regulators significantly increased the rapeseed yield, N physiological efficiency, single pod number and grain number per pod, but had little effect on weight of thousand seeds. Different growth regulators had different influences on above indexes. ABA mainly impacted on grain number per pod, while IAA and6-BA on the grain yield and plant pod number. The effect of each growth regulator on single index differed in rape variety difference and different N treatments. IAA exhibited optimized effect on nitrogen physiological efficiency in rape variety A under normal N condition, while6-BA had a good effect under N deficient condition. As to rape variety B, IAA exhibited best impact under normal and deficient N conditions. In addition, nitrogen physiological efficiency of biomass and grain yield in normal N condition was lower than that in N deficient condition. Comparing with the grain yield, yield components and N physiological efficiency of two N levels, we found that grain yield and plant pod number in normal N condition were higher than those in N deficiency, nitrogen physiological efficiency in normal N condition was lower than that in deficient N condition, and there were little difference in weight of1000-grain weight and grain number per pod at two N levels.
The results of nitrogen transport ratio and transport volume showed that three kinds of growth regulators could increase the nitrogen transport ratio and transport volume to grain, and reduce the losses. But the effect of different growth regulators on them differed in applied N levels and different varieties. IAA could pose greater contribution to nitrogen translocation and redistribution in variety A, and ABA in variety B. This advantage, meanwhile, also exhibited in the aspect of low N loss. IAA had a greater contribution to two rape varieties under N stress. On the normal N application condition, the amount of nitrogen redistribution and its proportion toward seeds and siliques was larger than the results of N stress condition. And the breed A are higher than breed B in two N levels, but the difference was not significant.
The activity of proteoltic enzyme (PE) and the glutamine synthetase (GS) in blades and the activity of glutamate z-oxoghitarate aminotransferase (GOGAT) in seeds were determined during the later growing period. The results suggested that three kinds of plant growth regulators, to some extent, could improve the activity of enzyme related to N transport. ABA could improve the activity of PE, and IAA is effective on the active of GS and GOGAT. Compared with IAA and ABA, the percentage of increasing activity by6-BA was lower. In general, the enzyme activity in normal N condition is lower than that under N stress condition, and the enzyme activity in breed A is higher than that in breed B.
The contents of L-glutamine, free amino acid and soluble sugar in phloem juice were measured. The results showed that phytohormones could increase the contents of L-glutamine, soluble sugar and free amino acid to varying degrees. IAA and ABA are more beneficial in L-glutamine content, while6-BA is useful to soluble sugar. To the free amino acid content, the influence is different from varieties. Different levels of N and varieties had different effects on the content of phloem juice. The content of L-glutamine showed that the normal N application condition was better for its improvement than that under N stress condition, and the content in breed A was lower than that in breed B. The content of soluble sugar in breed A is slightly lower than that in breed B, but few differences between the two N levels. The free amino acid content in two N levels and two varieties are barely different.
The determination of rapeseed oil and protein indicated that growth regulators could increase the grain protein content and protein content of rapeseed, but little effect in the oil content. However, because growth regulators increased the grain yield, the oil yield got significantly improved. The growing contribution of regulators in oil production improvement are varied. For breed A, on the normal N application condition,6-BA is better. However, ABA was more useful under N stress condition. For breed B, IAA is more effective in two N levels. The influences of N levels and rape varieties on grain protein and oil content were analysed. The protein content, protein yield and oil yield showed that normal N condition is better for its improvement than N stress, but oil contein is the opposite results. Breed A is higher than breed B in protein content, protein yield and oil yield, but the oil content is no difference between two varieties.
本试验采用两个油菜品种(分别用A和B表示),在常氮和氮胁迫两个供氮水平下,研究角果发育期涂抹不同生长调节剂(生长素IAA、细胞分裂素6-BA及脱落酸ABA)溶液于角果,对油菜生长后期氮素再利用和氮效率的影响及其影响机理,为揭示油菜氮效率的生理机制,探索提高油菜氮效率的可行性途径提供科学依据。
1.油菜产量、产量构成因素及氮素生理效率的测定结果表明,三种生长调节剂均能不同程度地提高籽粒产量、氮素生理效率、单株角果数和每角果粒数,但对于千粒重没有产生什么影响。不同生长调节剂对以上各指标的影响不同,ABA主要对每角果粒数增效作用大,IAA和6-BA则主要对籽粒产量和单株角果数贡献较大。各生长调节剂对同一指标的影响因氮水平高低和品种不同而有所差异,以籽粒产量为基础的氮素生理效率为例,A品种在常氮条件下以ABA效应好,氮胁迫条件下以6-BA的效应最好；而B品种则均以IAA的效果最好。另外,常氮水平下的生物产量氮素生理效率及籽粒产量氮素生理效率低于氮胁迫水平。比较两个供氮水平的籽粒产量、产量构成因素及氮素生理效率,籽粒产量和单株角果数,常氮条件高于氮胁迫条件；氮素生理效率则常氮条件低于氮胁迫条件；千粒重和每角果粒数在两个氮水平之间没有差异。
2.油菜生长后期氮素转运比例和转运量的测定结果表明,三种生长调节剂处理后,氮素向籽粒的转运比例和转运量增加,损失量减少。但不同生长调节剂的影响程度因供氮水平和品种不同而有所不同。从氮素转运再分配的增加效果来看,常氮条件下IAA对A品种的贡献大,ABA对B品种贡献大,这种优势同时也表现在较低的氮素损失量方面；氮胁迫条件下,A、B两品种均是以IAA的贡献大。比较两个供氮水平的氮素转运比例及转运量,无论是向籽粒还是向角果的转运,两个品种均表现为常氮条件下高于氮胁迫条件。比较两个品种的氮素转运比例及转运量,无论是向籽粒还是向角果的转运,均表现为A品种的转运比例及转运量略高于B品种,但差异不显著。
3.油菜生长后期氮素转运相关酶活性的测定结果表明,三种生长调节剂都可不同程度地提高氮素转运相关酶活性。其中,对蛋白水解酶提高效应最好的是ABA；而IAA对谷氨酰胺合成酶活性和谷氨酸合成酶活性有相对好的提高效果；6-BA对酶的活性提高幅度不如前两种生长调节剂大。氮水平和品种对三种酶的活性的影响来看,总体上表现为常氮条件低于氮胁迫条件、品种A高于品种B的趋势。
4.韧皮部汁液成分的测定结果表明,三种生长调节剂都可不同程度地增加韧皮部汁液中的L-谷氨酰胺含量、可溶性糖含量和游离氨基酸含量。IAA和ABA对L-谷氨酰胺含量的提高作用相对较大,6-BA对可溶性糖含量的提高作用相对大,三种生长调节剂对游离氨基酸含量的影响,因品种而异。氮水平和品种对韧皮部汁液成分的影响来看,L-谷氨酰胺含量为常氮条件高于氮胁迫条件、可溶性糖含量和游离氨基酸含量在两氮素水平间几乎没有差异；L-谷氨酰胺含量和可溶性糖含量表现为A品种略低于B品种,游离氨基酸含量在两个品种之间差异性不大。
5.对油菜籽粒蛋白质含量和油分含量的测定结果表明,生长调节剂能增加油菜籽粒蛋白质含量及蛋白质产量,但对油分含量的影响较小,但因为生长调节剂增加了籽粒产量,因而能显著提高油分产量。从各生长调节剂对提高油分产量方面的贡献来看,A品种在常氮水平下以6-BA的贡献较大,氮胁迫条件下以ABA的贡献较大；而B品种在两种供氮水平下均以IAA的贡献较大。从氮水平和品种对籽粒蛋白质含量和油分含量的影响来看,蛋白质含量、蛋白质产量和油分产量表现为常氮条件高于氮胁迫条件,油分含量则反之；蛋白质含量、蛋白质产量和油分产量A品种均高于B品种,油分含量则在两个品种之间没有差异。
Oilseed Rape, as a significant oil crop with planting advantage, has aroused widespread concern increasingly. With the new oilseed rape variety springing up possessing high yield and high quality, low nutrient utilization efficiency has become a new bottleneck, which constrain the further development of oilseed rape production. Especially because of high nitrogen (N) demand and low utilization rate, the development of rape career are restricted. As a result, how to raise the N fertilizer efficiency has become a central issue in plant nutrition research area. In the whole plant growth and development period, phytohormones play a critical regulatory role in the process of physiological metabolism and morphogenesis. However, effect of phytohormones on N reutilization and N efficiency regulation were few reported in recently.
The experiment was conducted by daubing solutions of different growth regulators such as auxin(IAA), cytokinin(6-BA) and abscisic acid(ABA) to rape pod with two rapeseed varieties(A and B) under normal and deficient N conditions. The effect of N reutilization and efficiency during later stages of rape and corresponding mechanisms were studied, aiming at revealing physiological mechanisms of N utilization efficiency in rape, and providing a scientific basis for improving N utilization efficiency in rape.
The results of the yield, yield components and nitrogen physiological efficiency indicated that the growth regulators significantly increased the rapeseed yield, N physiological efficiency, single pod number and grain number per pod, but had little effect on weight of thousand seeds. Different growth regulators had different influences on above indexes. ABA mainly impacted on grain number per pod, while IAA and6-BA on the grain yield and plant pod number. The effect of each growth regulator on single index differed in rape variety difference and different N treatments. IAA exhibited optimized effect on nitrogen physiological efficiency in rape variety A under normal N condition, while6-BA had a good effect under N deficient condition. As to rape variety B, IAA exhibited best impact under normal and deficient N conditions. In addition, nitrogen physiological efficiency of biomass and grain yield in normal N condition was lower than that in N deficient condition. Comparing with the grain yield, yield components and N physiological efficiency of two N levels, we found that grain yield and plant pod number in normal N condition were higher than those in N deficiency, nitrogen physiological efficiency in normal N condition was lower than that in deficient N condition, and there were little difference in weight of1000-grain weight and grain number per pod at two N levels.
The results of nitrogen transport ratio and transport volume showed that three kinds of growth regulators could increase the nitrogen transport ratio and transport volume to grain, and reduce the losses. But the effect of different growth regulators on them differed in applied N levels and different varieties. IAA could pose greater contribution to nitrogen translocation and redistribution in variety A, and ABA in variety B. This advantage, meanwhile, also exhibited in the aspect of low N loss. IAA had a greater contribution to two rape varieties under N stress. On the normal N application condition, the amount of nitrogen redistribution and its proportion toward seeds and siliques was larger than the results of N stress condition. And the breed A are higher than breed B in two N levels, but the difference was not significant.
The activity of proteoltic enzyme (PE) and the glutamine synthetase (GS) in blades and the activity of glutamate z-oxoghitarate aminotransferase (GOGAT) in seeds were determined during the later growing period. The results suggested that three kinds of plant growth regulators, to some extent, could improve the activity of enzyme related to N transport. ABA could improve the activity of PE, and IAA is effective on the active of GS and GOGAT. Compared with IAA and ABA, the percentage of increasing activity by6-BA was lower. In general, the enzyme activity in normal N condition is lower than that under N stress condition, and the enzyme activity in breed A is higher than that in breed B.
The contents of L-glutamine, free amino acid and soluble sugar in phloem juice were measured. The results showed that phytohormones could increase the contents of L-glutamine, soluble sugar and free amino acid to varying degrees. IAA and ABA are more beneficial in L-glutamine content, while6-BA is useful to soluble sugar. To the free amino acid content, the influence is different from varieties. Different levels of N and varieties had different effects on the content of phloem juice. The content of L-glutamine showed that the normal N application condition was better for its improvement than that under N stress condition, and the content in breed A was lower than that in breed B. The content of soluble sugar in breed A is slightly lower than that in breed B, but few differences between the two N levels. The free amino acid content in two N levels and two varieties are barely different.
The determination of rapeseed oil and protein indicated that growth regulators could increase the grain protein content and protein content of rapeseed, but little effect in the oil content. However, because growth regulators increased the grain yield, the oil yield got significantly improved. The growing contribution of regulators in oil production improvement are varied. For breed A, on the normal N application condition,6-BA is better. However, ABA was more useful under N stress condition. For breed B, IAA is more effective in two N levels. The influences of N levels and rape varieties on grain protein and oil content were analysed. The protein content, protein yield and oil yield showed that normal N condition is better for its improvement than N stress, but oil contein is the opposite results. Breed A is higher than breed B in protein content, protein yield and oil yield, but the oil content is no difference between two varieties.
引文
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