氮硫配施对小麦产量和品质的调控及其生理学基础
摘要
试验于2005-2007年在河南农业大学科教示范园区进行,以2个不同穗型的国审中筋高产小麦品种为材料,研究了不同氮水平下施硫对小麦光合特性、氮、硫同化代谢、积累运转关键酶、蔗糖合成酶活性、激素含量、氮素和硫素的积累与运转、籽粒灌浆特性及产量、主要品质指标的影响。主要试验结果如下:
1.硫对小麦光合速率和源库糖合成的影响与对照(S0)相比,豫农949在N240和N330水平下,施硫提高了旗叶Chl含量、可溶性蛋白含量和旗叶净光合速率,提高花后旗叶可溶性糖和蔗糖的含量和灌浆前期籽粒中可溶性糖和蔗糖的含量,加速了灌浆后期籽粒中糖的降解;兰考矮早8在N240和N330水平下提高了旗叶可溶性蛋白含量、花后旗叶可溶性糖和蔗糖的含量和灌浆前期籽粒中可溶性糖和蔗糖的含量,加速了灌浆后期籽粒中糖的降解,旗叶Chl含量和净光合速率N330S60处理较N330S0处理有所降低。说明施用硫肥能够增强光合“源”器官碳同化物的供应能力,促进了籽粒对叶源端同化物的转化、利用能力,为淀粉合成提供了充足的底物供应。
2.硫对小麦氮、硫同化酶活性的影响硝酸还原酶、谷氨酰胺合成酶和谷氨酸丙酮酸转氨酶是氮素同化的三个关键酶。OASS裂解酶,是硫代谢的关键酶。在N240水平下施硫可以增加旗叶中三个酶的活性,提高氮素的同化能力。在2种氮水平下,2个小麦品种旗叶三种酶活性变化趋势基本一致,即NR于花后0~5d,GS于花后5d,GPT于花后10d,分别达到最大值,然后下降;2个小麦品种籽粒GS在花后5~10d,GPT在花后10~20d,分别保持较高的活性值,然后呈迅速下降趋势。与对照相比,在N240水平下S60处理显著提高了旗叶GS和GPT活性值;在N330水平下,S60处理在大部分时期提高了旗叶GS和GPT的活性值。籽粒GS和GPT活性值与旗叶变化趋势一致。与对照相比,增施硫肥均降低了两品种旗叶和籽粒中乙酰丝氨酸水解酶(OASS)的活性。
3.硫对小麦碳、氮、硫积累运转的影响与对照相比,在中氮(N240)和高氮(N330)水平下,施硫可以促进小麦干物质的积累和向籽粒的转运,N240S60比N330S60更利于促进花前营养器官贮藏物质向结实器官的运转,两品种相比较,从开花到成熟期豫农949单株干物质积累量明显高于兰考矮早8。与对照相比,施硫提高了开花期叶、茎鞘氮和硫的积累量以及成熟期籽粒氮、硫的含量,减小了成熟期氮素在植株体内的累积量,2品种相比较,豫农949较兰考矮早8的施硫处理更能有效地提高氮素的吸收、利用率。与对照相比,施硫提高了花前植株体积累氮素的输出能力,同时提高了输出氮素对籽粒氮的贡献率,提高了硫素在植株体内的移动率,利于硫素及时供应到营养器官,减小了植株体和籽粒的N/S比。
4.硫对小麦内源激素含量变化的影响与对照相比,施硫提高了灌浆前期旗叶细胞分裂素(ZR,IAA)的含量,在灌浆后期减小了旗叶IAA的含量,提高了花后0~15d和30d时赤霉素(GA3)的含量,减小了灌浆末期旗叶脱落酸(ABA)的含量。施用硫肥,能够在灌浆之前促进和保持叶片的生理功能,而在灌浆后期则延缓了功能叶片的衰老,说明硫素能够通过改善内源激素的含量,来调控小麦叶片的衰老进程。与对照相比,ZR含量花后5~10d时籽粒S60>S0,花后10d时籽粒IAA含量S60>S0。施硫能够提高灌浆高峰到来之前籽粒的ZR和IAA含量,说明施用硫肥促进了前期细胞分裂,增加了库容,提高了贮藏能力,促进了有机物质向籽粒的转运。
5.硫对小麦产量及其构成因素的影响与对照相比,两个小麦品种在2种氮水平下,施硫在提高产量构成因素的同时增加小麦籽粒产量。N240水平下施硫,豫农949和兰考矮早8两品种籽粒产量分别增加12.74%和6.15%, N330水平下施硫,籽粒产量分别增加16.41%和10.63%,2品种均为N330水平下施硫增产幅度高,但2品种N330S60和N240S60两处理相比,差异均不显著。与对照相比,S60处理提高了豫农949的穗数、穗粒数、千粒重,差异均达显著水平;兰考矮早8穗数、穗粒数均为S60>S0,穗数差异达显著水平,穗粒数差异不显著。千粒重N330S60>N330S0,差异达显著水平,N240S60 6.硫对小麦籽粒主要品质性状的影响与对照相比,两个小麦品种在2种氮水平下,施硫提高了贮藏蛋白和结构蛋白的含量,减小了淀粉的直/支比。与对照相比,2品种在2种氮水平下施硫,提高了小麦面粉的峰值粘度、低谷粘度和最终粘度,增加了面粉的形成时间和粉质指数,提高幅度分别为5.75-45.86%和2.61-23.45%,同时提高了面粉稳定时间,但差异未达显著水平。与对照相比,施硫提高了面团的拉伸面积和延伸度,施硫对豫农949拉伸阻力的影响,差异达显著水平。说明施用硫肥能够改善小麦的营养品质,并改善了面团的流变学特性,改进了小麦的加工品质。与对照相比,施硫减小了2小麦品种面粉的PPO活性,提高了面粉的亮度(L*值),2品种相比较而言,豫农949的面粉和面片L*均高于兰考矮早8,而a*、b*则均低于兰考矮早8。说明施用硫肥,能够在改善小麦加工品质的同时提高感官的舒适度。
Field experiment were carried out on the farm of Henan agricultural university during 2005-2007 using two winter wheat cultivars with different spike i.e. Yunong949(with multiply spike)and Lankao Aizao 8 (with large spike).The objective of this study was to clarify the effects of different nitrogen combined sulphur application on key enzyme activity involved in nitrogen and sulphur assimilation SS and SPS in flag leaves and grains and photosynthetic characteristics in flag leaves accumulation and transportation of dry matter nitrogen and sulphur grain yield and main quality traits of two winter wheat cultivars with different spikes so as to provide some experimental foundations for reasonable sulfur application of winter wheat. The results were summarized as follows:
1. Under the condition of nitrogen fertilization of 240kg/hm2 and 330kg/hm2Comparing with no sulphur application treatment sulphur could increase the Chl content soluble protein content and the flag of the net photosynthetic rate(Pn) increased spending after the flag leaf soluble sugar and the sugar content and early grain filling in soluble sugar and sugar content accelerated the grain filling in the latter part of the sugar Degradation. Improve the flag of the soluble protein content after spending flag leaf soluble sugar and the sugar content and early grain filling in soluble sugar and sugar content accelerated the grain filling in the latter part of the degradation of sugar of LankaoAizao8 Flag leaf Chl content and net photosynthetic rate N330S60 deal with than dealing with lower N330S0. Sulfur fertilizer application that can enhance photosynthesis "source" organ carbon assimilation of supply capacity promote the grain of assimilation-leaf source of transformation capacity utilization for the synthesis of starch to provide an adequate supply of substrate.
2. Sulphur fertilization could improve the activity of nitrate reductase (NR) Glutamine Synthetase (GS) and glutamic-pyruvic transaminaser (GPT) which was the key enzymes of nitrogen assimilation and so improved the Nitrogen assimilation ability. OASS lyase sulfur metabolism is the key enzyme. Under the condition of nitrogen fertilization of 240kg/hm2 sulphur could increase the nitrogen assimilation capacity of three enzyme in flag leaves. Under two kinds of nitrogen level the variation tendencies of three enzymes activity were uniform basically namely NR GS and GPT reach maximum 0~5d after anthesis 5d after anthesis 10d after anthesis respectively then began to drop. The two cultivar enzyme activity in grains namely GS and GPT reach maximum 5-10d after anthesis 10-20d after anthesis respectively. Compared with control S60 treatment significantly increased the flag of the GS and GPT activity value in N240 level S60 handle most of the time increased GS and GPT activity in flag leaf in N330 level. GS and GPT variation tendencies were uniform basically in grains. Compared with control sulphur can decrease the OASS activity of two wheat cultivars in flag leaf and grains.
3. Compared with control sulfur can promote the accumulation of dry matter and grain to the transit of two cultivars under two nitrogen levels (240 and 330 kg N hm-2). N240S60 than N330S60 spend more conducive to the promotion of organ storage before the nutritional substances to the functioning of solid organs. Comparing two cultivar Yunong 949 dry matter accumulation was significantly higher than that as early as LankaoAizao 8 in plant from Blossom to maturity. Compared with control sulfur increased flowering period leaves stem and sheath nitrogen sulfur accumulation and nitrogen and sulfur content of grains and reducing the nitrogen content in plants at maturity. Comparing two cultivar Yunong949 have more effective than LankaoAizao 8 enhancing the absorption and utilization rate of nitrogen.Compared with control sulfur enhance the accumulation of nitrogen output capacity while increasing the output of grain nitrogen nitrogen contribution Increaseing the sulfur mobile rate of plant conducive to the timely supply of sulfur to nutrition organs reducing the the N / S ratio of plant and seed.
4. Compared with control sulfur could increase the cytokinin of flag leaf pre-filling decrease the IAA content and improved GA3 content 0~15d after anthesis 30d after anthesis respectively decreased the ABA content of flag leaf in the end of filling. Sulfur fertilizer application can promote and maintain physiological function in pre-filling and delayed the function of the aging leaves.The results indicate that sulfur able to improve the content of the hormone endogenous then control the aging process of leaves. Compared control S60 treatment is greater than S0 of ZR content in grains 5~10d after anthesis but S60 treatment is greater than S0 of IAA content 10d after anthesis. Sulfur can improve the ZR and IAA content before the peak of grain filling. The results showed that sulphur can promote the cell division and increase of storage capacity and promote the organic material to the transshipment of grain.
5. Comparing with no sulphur application treatment under different nitrogen sulphur treatments could increase the grain yield and the yield components of two wheat cultivars. Yunong949 amd LankaoAizao8 varieties yield grains improve 12.74% and 6.15%respectively in N240 level yield grains improve 16.41% and 10.63% respectively in N330 level. Two varieties are high production rate in N330 level. However two varieties N330S60 and N240S60 two treatment compared to no significant difference. Comparing with control S60 improve the handling of ears Kernels per spike grain weight of Yunong949 the difference reached significant levels. Ears and Kernels per spike were S60 greater than S0 of LankaoAizao8 and the Kernels per spike reached significant difference. 1000-grain weight N330S60> N330S0 significant differences in the level N240S60 6. Comparing with no sulphur application treatment under different nitrogen sulphur treatments could increase the storage protein and structure protein content Reduce the rate of amylase/amylopectin Increase the time and the formation of flour silty index the development time and Farinograph quality improve 5.75-45.86% and 2.61-23.45% respectively at the same time improve the stability time of flour but the difference was not up significant level. Comparing control sulphur could increase the area and extensibility of flour the impact of paste resistance reach significant level. The results showed that sulfur fertilizer application could improve the nutritional quality and improve the Rheological properties of dough and improve processing quality in wheat. Comparing control sulphur application could reduce the PPO activity improve the brightness of the flour. 2 varieties of comparison Yunong949 has higher lightless than LankaoAizao8 but a * b * are lower than it. As a result sulfur fertilizer application could improve the processing of wheat at the same time improve the quality of sensory comfort.
1.硫对小麦光合速率和源库糖合成的影响与对照(S0)相比,豫农949在N240和N330水平下,施硫提高了旗叶Chl含量、可溶性蛋白含量和旗叶净光合速率,提高花后旗叶可溶性糖和蔗糖的含量和灌浆前期籽粒中可溶性糖和蔗糖的含量,加速了灌浆后期籽粒中糖的降解;兰考矮早8在N240和N330水平下提高了旗叶可溶性蛋白含量、花后旗叶可溶性糖和蔗糖的含量和灌浆前期籽粒中可溶性糖和蔗糖的含量,加速了灌浆后期籽粒中糖的降解,旗叶Chl含量和净光合速率N330S60处理较N330S0处理有所降低。说明施用硫肥能够增强光合“源”器官碳同化物的供应能力,促进了籽粒对叶源端同化物的转化、利用能力,为淀粉合成提供了充足的底物供应。
2.硫对小麦氮、硫同化酶活性的影响硝酸还原酶、谷氨酰胺合成酶和谷氨酸丙酮酸转氨酶是氮素同化的三个关键酶。OASS裂解酶,是硫代谢的关键酶。在N240水平下施硫可以增加旗叶中三个酶的活性,提高氮素的同化能力。在2种氮水平下,2个小麦品种旗叶三种酶活性变化趋势基本一致,即NR于花后0~5d,GS于花后5d,GPT于花后10d,分别达到最大值,然后下降;2个小麦品种籽粒GS在花后5~10d,GPT在花后10~20d,分别保持较高的活性值,然后呈迅速下降趋势。与对照相比,在N240水平下S60处理显著提高了旗叶GS和GPT活性值;在N330水平下,S60处理在大部分时期提高了旗叶GS和GPT的活性值。籽粒GS和GPT活性值与旗叶变化趋势一致。与对照相比,增施硫肥均降低了两品种旗叶和籽粒中乙酰丝氨酸水解酶(OASS)的活性。
3.硫对小麦碳、氮、硫积累运转的影响与对照相比,在中氮(N240)和高氮(N330)水平下,施硫可以促进小麦干物质的积累和向籽粒的转运,N240S60比N330S60更利于促进花前营养器官贮藏物质向结实器官的运转,两品种相比较,从开花到成熟期豫农949单株干物质积累量明显高于兰考矮早8。与对照相比,施硫提高了开花期叶、茎鞘氮和硫的积累量以及成熟期籽粒氮、硫的含量,减小了成熟期氮素在植株体内的累积量,2品种相比较,豫农949较兰考矮早8的施硫处理更能有效地提高氮素的吸收、利用率。与对照相比,施硫提高了花前植株体积累氮素的输出能力,同时提高了输出氮素对籽粒氮的贡献率,提高了硫素在植株体内的移动率,利于硫素及时供应到营养器官,减小了植株体和籽粒的N/S比。
4.硫对小麦内源激素含量变化的影响与对照相比,施硫提高了灌浆前期旗叶细胞分裂素(ZR,IAA)的含量,在灌浆后期减小了旗叶IAA的含量,提高了花后0~15d和30d时赤霉素(GA3)的含量,减小了灌浆末期旗叶脱落酸(ABA)的含量。施用硫肥,能够在灌浆之前促进和保持叶片的生理功能,而在灌浆后期则延缓了功能叶片的衰老,说明硫素能够通过改善内源激素的含量,来调控小麦叶片的衰老进程。与对照相比,ZR含量花后5~10d时籽粒S60>S0,花后10d时籽粒IAA含量S60>S0。施硫能够提高灌浆高峰到来之前籽粒的ZR和IAA含量,说明施用硫肥促进了前期细胞分裂,增加了库容,提高了贮藏能力,促进了有机物质向籽粒的转运。
5.硫对小麦产量及其构成因素的影响与对照相比,两个小麦品种在2种氮水平下,施硫在提高产量构成因素的同时增加小麦籽粒产量。N240水平下施硫,豫农949和兰考矮早8两品种籽粒产量分别增加12.74%和6.15%, N330水平下施硫,籽粒产量分别增加16.41%和10.63%,2品种均为N330水平下施硫增产幅度高,但2品种N330S60和N240S60两处理相比,差异均不显著。与对照相比,S60处理提高了豫农949的穗数、穗粒数、千粒重,差异均达显著水平;兰考矮早8穗数、穗粒数均为S60>S0,穗数差异达显著水平,穗粒数差异不显著。千粒重N330S60>N330S0,差异达显著水平,N240S60
Field experiment were carried out on the farm of Henan agricultural university during 2005-2007 using two winter wheat cultivars with different spike i.e. Yunong949(with multiply spike)and Lankao Aizao 8 (with large spike).The objective of this study was to clarify the effects of different nitrogen combined sulphur application on key enzyme activity involved in nitrogen and sulphur assimilation SS and SPS in flag leaves and grains and photosynthetic characteristics in flag leaves accumulation and transportation of dry matter nitrogen and sulphur grain yield and main quality traits of two winter wheat cultivars with different spikes so as to provide some experimental foundations for reasonable sulfur application of winter wheat. The results were summarized as follows:
1. Under the condition of nitrogen fertilization of 240kg/hm2 and 330kg/hm2Comparing with no sulphur application treatment sulphur could increase the Chl content soluble protein content and the flag of the net photosynthetic rate(Pn) increased spending after the flag leaf soluble sugar and the sugar content and early grain filling in soluble sugar and sugar content accelerated the grain filling in the latter part of the sugar Degradation. Improve the flag of the soluble protein content after spending flag leaf soluble sugar and the sugar content and early grain filling in soluble sugar and sugar content accelerated the grain filling in the latter part of the degradation of sugar of LankaoAizao8 Flag leaf Chl content and net photosynthetic rate N330S60 deal with than dealing with lower N330S0. Sulfur fertilizer application that can enhance photosynthesis "source" organ carbon assimilation of supply capacity promote the grain of assimilation-leaf source of transformation capacity utilization for the synthesis of starch to provide an adequate supply of substrate.
2. Sulphur fertilization could improve the activity of nitrate reductase (NR) Glutamine Synthetase (GS) and glutamic-pyruvic transaminaser (GPT) which was the key enzymes of nitrogen assimilation and so improved the Nitrogen assimilation ability. OASS lyase sulfur metabolism is the key enzyme. Under the condition of nitrogen fertilization of 240kg/hm2 sulphur could increase the nitrogen assimilation capacity of three enzyme in flag leaves. Under two kinds of nitrogen level the variation tendencies of three enzymes activity were uniform basically namely NR GS and GPT reach maximum 0~5d after anthesis 5d after anthesis 10d after anthesis respectively then began to drop. The two cultivar enzyme activity in grains namely GS and GPT reach maximum 5-10d after anthesis 10-20d after anthesis respectively. Compared with control S60 treatment significantly increased the flag of the GS and GPT activity value in N240 level S60 handle most of the time increased GS and GPT activity in flag leaf in N330 level. GS and GPT variation tendencies were uniform basically in grains. Compared with control sulphur can decrease the OASS activity of two wheat cultivars in flag leaf and grains.
3. Compared with control sulfur can promote the accumulation of dry matter and grain to the transit of two cultivars under two nitrogen levels (240 and 330 kg N hm-2). N240S60 than N330S60 spend more conducive to the promotion of organ storage before the nutritional substances to the functioning of solid organs. Comparing two cultivar Yunong 949 dry matter accumulation was significantly higher than that as early as LankaoAizao 8 in plant from Blossom to maturity. Compared with control sulfur increased flowering period leaves stem and sheath nitrogen sulfur accumulation and nitrogen and sulfur content of grains and reducing the nitrogen content in plants at maturity. Comparing two cultivar Yunong949 have more effective than LankaoAizao 8 enhancing the absorption and utilization rate of nitrogen.Compared with control sulfur enhance the accumulation of nitrogen output capacity while increasing the output of grain nitrogen nitrogen contribution Increaseing the sulfur mobile rate of plant conducive to the timely supply of sulfur to nutrition organs reducing the the N / S ratio of plant and seed.
4. Compared with control sulfur could increase the cytokinin of flag leaf pre-filling decrease the IAA content and improved GA3 content 0~15d after anthesis 30d after anthesis respectively decreased the ABA content of flag leaf in the end of filling. Sulfur fertilizer application can promote and maintain physiological function in pre-filling and delayed the function of the aging leaves.The results indicate that sulfur able to improve the content of the hormone endogenous then control the aging process of leaves. Compared control S60 treatment is greater than S0 of ZR content in grains 5~10d after anthesis but S60 treatment is greater than S0 of IAA content 10d after anthesis. Sulfur can improve the ZR and IAA content before the peak of grain filling. The results showed that sulphur can promote the cell division and increase of storage capacity and promote the organic material to the transshipment of grain.
5. Comparing with no sulphur application treatment under different nitrogen sulphur treatments could increase the grain yield and the yield components of two wheat cultivars. Yunong949 amd LankaoAizao8 varieties yield grains improve 12.74% and 6.15%respectively in N240 level yield grains improve 16.41% and 10.63% respectively in N330 level. Two varieties are high production rate in N330 level. However two varieties N330S60 and N240S60 two treatment compared to no significant difference. Comparing with control S60 improve the handling of ears Kernels per spike grain weight of Yunong949 the difference reached significant levels. Ears and Kernels per spike were S60 greater than S0 of LankaoAizao8 and the Kernels per spike reached significant difference. 1000-grain weight N330S60> N330S0 significant differences in the level N240S60
引文
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