甜菜氮素同化与蔗糖代谢机理研究及其人工调控
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摘要
本研究以当前甜菜主栽品种标准偏高糖型的甜研7号为试验材料,于2006年至2007年在东北农业大学校内和实验实习基地进行。试验采用四因素五水平二次正交旋转组合设计,四个试验处理因素分别为NO_3~--N、NH_4~+-N、MgO、P_2O_5。系统地研究了甜菜氮素同化和蔗糖代谢关键酶活性与主要产物的变化动态,硝态氮、铵态氮、镁肥和磷肥对氮素同化和蔗糖代谢关键酶的调控途径,分析了氮素同化和蔗糖代谢的相关关系,氮素同化和蔗糖代谢与产质量的相关关系,阐明了甜菜氮素同化和蔗糖代谢的机理,并进一步在氮素同化和蔗糖代谢的交叉点上明确了氮素同化和蔗糖代谢的互作机理,揭示了氮、磷和镁肥对甜菜氮素同化和蔗糖代谢及产质量的调控效应。本研究填补了甜菜生理的基础性资料,可为种质资源的鉴定、评价和改造利用及制定合理栽培措施提供理论依据。研究结果表明:
1.甜菜氮素同化关键酶活性的变化规律:从时间上,叶片中enNRA、GSA、GOGATA和GDHA峰值均在块根增长期之前;块根中,GOGATA在前期活性较高,而GSA和GDHA在后期影响较大。在空间上,除苗期略低于叶片外,其它各时期块根的GSA均远远高于叶片;叶片GOGATA除了在叶丛形成期和叶丛形成末期略低于块根外,其它各时期均远远高于块根;而叶片GDHA均高于块根。且GSA、GOGATA和GDHA在块根和叶片中的变化动态均不同。氮素同化关键酶间呈不同程度相关。在叶丛形成期之前主要是内源和外源NRA间、NRA与GSA间及块根和叶片GSA间显著正相关;在叶丛形成期及之后主要是GSA与GOGATA间显著负相关。氮素同化关键酶活性与块根产量和品质的关系:在块根增长期之前与块根的产量和产糖量关系密切;从块根增长期开始,与块根含糖率相关均极显著。且叶丛形成期之前叶片的关键酶活性与产量相关密切;叶丛形成期之后直到块根增长末期块根的GSA对产量和品质的影响最大;到糖分积累期NRA成为影响甜菜块根含糖率最主要因素。
甜菜氮素同化主要产物含量的变化动态:块根可溶性蛋白含量在块根增长期之前基本呈缓慢下降趋势,之后大幅增加;叶片可溶性蛋白含量在整个生育期呈逐渐增加趋势。块根全氮量从苗期到块根增长末期一直下降,之后略有上升;叶片全氮量从苗期到叶丛形成末期一直下降,之后直到块根增长期上升,随后呈下降趋势。氮素同化主要产物间呈现不同程度的相关关系。氮素同化产物与块根的产量和品质在整个生育时期基本均呈不同程度相关。且在块根增长期全氮量和可溶性蛋白含量与块根产量和品质的相关系数同时达到较高水平,可以作为块根增长期预测产量、含糖率和产糖量的参考指标。
同时甜菜氮素同化关键酶与主要同化产物在不同时期呈现不同程度相关。
2.甜菜蔗糖代谢关键酶活性的变化规律:块根中SPSA、SS分解方向活性、NIA和AIA的变化动态相似,叶片中SPSA与SS合成方向活性变化动态相似,其它各关键酶活性的变化动态均不同。从空间来看,块根的SPSA、SS合成方向活性(除苗期)、NIA在整个生育时期均高于叶片;块根的SS分解活性在中后期低于叶片,其它时期均高于叶片;块根的AIA在苗期和后期高于叶片,在中期低于叶片。SS合成活性与SPSA相比,在块根中,除苗期和叶丛形成初期略低于SPSA,其它时期均表现为高于SPSA;在叶片中,二者的活性大小接近。NIA与AIA相比,块根和叶片NIA的峰值均低于AIA。SS分解活性与NIA和A1A相比,块根中SS分解活性除叶丛形成初期大于NIA和AIA之外,其它时期均小于NIA和AIA;叶片中三种酶均处于较低水平,差异不大。不同时期蔗糖代谢关键酶间相关关系:苗期块根中糖代谢分解方向酶间的同步性很强;其它各时期SPSA和SSA(包括合成和分解方向)与其它糖代谢关键酶呈不同程度相关。
甜菜蔗糖代谢各主要产物的变化动态:块根和叶片可溶性总糖含量均呈单峰曲线变化,峰值分别在块根增长期和叶丛形成末期。块根的果糖含量在整个生育时期呈波动变化,分别在叶丛形成期和块根增长期各有一个波谷。叶片的果糖含量呈“V”型变化,在叶丛形成初期降至最低水平。块根的蔗糖含量从苗期到块根增长期逐渐增加,之后略有下降,但直到糖分积累期仍保持较高水平。叶片的蔗糖含量从苗期到叶丛形成末期呈缓慢增加趋势,之后略微下降,在块根增长期又缓慢上升。且整个生育期块根的可溶性总糖、果糖和蔗糖含量均大于叶片。蔗糖代谢主要产物间在不同时期呈现不同程度的相关关系。且叶丛形成初期果糖是块根糖代谢的最主要产物,叶丛形成期蔗糖是块根糖代谢的最主要产物。而整个生育时期果糖在叶片可溶性总糖中所占比例一直处于领先地位,同时从块根增长期直到糖分积累期叶片中蔗糖含量也迅速增加。并且块根中果糖和蔗糖含量在整个生育期间均为同步变化,叶片中果糖和蔗糖含量从块根增长期到糖分积累期也是同步变化的。
甜菜蔗糖代谢关键酶与主要产物的关系:从部位来看,蔗糖代谢关键酶活性在前期和叶丛形成期之后与叶片糖代谢主要产物、在叶丛形成期与块根糖代谢主要产物相关显著。从时期来看,苗期糖代谢分解方向的酶类对产物的影响均较大;在叶丛形成初期分解和合成方向的酶类均具有一定影响;在叶丛形成期合成方向酶类的影响略占优势;叶丛形成末期分解方向酶类的影响优势变大;在块根增长期和块根增长末期合成方向酶类作用又大于分解;糖分积累期,蔗糖代谢分解和合成方向的酶类对蔗糖代谢产物均具有一定影响。
甜菜蔗糖代谢关键酶活性与产量和品质在整个生育期基本都显著相关。蔗糖代谢主要产物与产量和品质均呈不同程度相关(除苗期和块根增长末期)。
3.甜菜氮素同化关键酶与蔗糖代谢关键酶的关系:NRA与块根SS分解方向活性、SPSA在生育前期关系较密切,与NIA和AIA在后期表现很强的同步性。块根中GSA与块根的糖代谢各关键酶活性间从叶丛形成末期到块根增长末期均呈不同程度的正相关,而叶片GSA与糖代谢关键酶活性在生育初期和末期相关更为密切,而在中期主要与叶片的糖代谢关键酶活性呈不同程度相关。叶片GOGATA与SPSA在苗期极显著正相关;叶丛形成初期与糖代谢各酶活性相关均不显著;叶丛形成期到块根增长末期与糖代谢合成方向酶类显著相关;糖分积累期与糖代谢合成与分解方向酶类均呈不同程度相关。
甜菜氮素同化关键酶与蔗糖代谢主要产物的关系:叶片NRA主要与果糖和蔗糖含量关系密切。块根GSA与蔗糖代谢主要产物含量在整个生育期相关均不显著,而叶片GSA在生育初期和后期对叶片的果糖和可溶性总糖含量影响较大,在中期与块根果糖和可溶性总糖含量关系密切。叶片GOGATA在叶丛形成初期与块根蔗糖代谢主要产物均呈同步变化趋势,在叶丛形成末期变化不再同步,到块根增长末期只与块根果糖含量同步变化。
甜菜氮素同化主要产物与蔗糖代谢关键酶的关系:块根可溶性蛋白含量在整个生育期与块根糖代谢各酶活性基本都呈极显著负相关。叶片可溶性蛋白含量与叶片糖代谢各酶活性均呈显著负相关;与块根糖代谢各酶活性基本均呈显著正相关。块根、叶片全氮量与糖代谢各酶的相关关系相似,SSA(分解和合成方向)和NIA对块根和叶片全氮量的影响均较大。
氮素同化与蔗糖代谢主要产物间的关系:块根可溶性蛋白含量与蔗糖代谢主要产物含量间在苗期和生育末期相关较弱,在中期相关较密切。叶片可溶性蛋白与叶片蔗糖、可溶性总糖含量在叶丛形成初期和叶丛形成期显著正相关,其它时期相关均不显著。块根全氮量与蔗糖代谢主要产物含量从苗期到叶丛形成期关系均较密切,而从叶丛形成末期到块根增长期仅与叶片蔗糖代谢主要产物含量相关紧密。并且块根全氮量与蔗糖代谢主要产物含量间均为负相关关系。叶片全氮量与蔗糖代谢主要产物含量间在苗期和糖分积累期关系均较弱;而从叶丛形成初期到块根增长末期关系密切。
4.甜菜氮糖代谢的相关生理指标的变化动态:块根与叶片C/N从苗期到块根增长期均呈增加趋势,之后略有下降。且整个生育期块根C/N均高于叶片。叶绿素含量从苗期到叶丛形成初期呈下降趋势,之后迅速上升,到叶丛形成期达到最高,随后下降,直至糖分积累期降至最低水平。块根干物质积累量在整个生育期呈增加趋势;叶片干物质积累量从苗期到块根增长期呈增加趋势,之后下降,块根增长末期之后又略有上升。冠根比在整个生育时期一直呈下降趋势。在叶丛形成末期冠根比接近1。因此,可用叶丛形成末期冠根比来反映甜菜植株的生长状况及生长中心转移情况,适时进行人工调控。
甜菜氮糖代谢的相关生理指标与产量和品质的关系:叶丛形成初期和叶丛形成末期与产量和品质关系最为密切,并以叶绿素含量、块根干重和冠根比的影响较大。且在整个生育期相关生理指标对含糖率的影响均极显著,对产量的影响均未达到极显著水平,对产糖量的影响程度在各时期不同。
5.硝态氮、铵态氮、镁肥和磷肥四种处理因素在不同时期对甜菜氮素同化和蔗糖代谢关键酶活性调控效应不同,而且两处理因素之间存在明显的交互作用,在交互作用中对氮素同化和蔗糖代谢关键酶活性起主导作用的因子不同,同时在不同生育时期四种处理因素对甜菜氮素同化和蔗糖代谢关键酶活性的贡献率不同。硝态氮和铵态氮在整个生育期均具有不同程度的贡献率;镁肥和磷肥在后期的贡献率较高。研究还得到四种处理因素影响显著时期甜菜氮素同化和蔗糖代谢关键酶活性最高时四个处理因子组合。
6.不同处理因素对甜菜产量、含糖率和产糖量的影响不同。对产量和产糖量的贡献率顺序均为:镁肥＞硝态氮＞铵态氮＞磷肥,对含糖率的贡献率顺序为:镁肥＞铵态氮＞硝态氮＞磷肥。研究还得到甜菜块根产量、含糖率和产糖量最高时硝态氮、铵态氮、镁肥和磷肥四个处理因子组合。
In this dissertation,Tianyan 7 of NZ sugar beet was selected as experimental sample,and experiments were made in the campus and practice base of NEAU from 2006 to 2007. Experimental factors were as follows:NO_3~--N,NH_4~+-N,MgO and P_2O_5.In experiments the quadratic rotation-orthogonal combination design was adopted.We comprehensively and systematically studied on dynamic changes and regulation approaches of key enzymes and main products of nitrogen assimilation and sucrose metabolism of sugar beet,the mechanism of nitrogen assimilation and sucrose metabolism,further analyzed interaction mechanism between nitrogen assimilation and sucrose metabolism,and announced the regulation effect of nitrogen, phosphorus and magnesium fertilizer on nitrogen assimilation,sucrose metabolism,yield and quality.This dissertation will fill in the infrastructure data of sugar beet physiology,provide theoretical basis for identifying,evaluating,reforming and using germplasm resources,and making reasonable cultivation measurement.Results of this experiment are as following:
1.The variation laws of the activities of key enzymes of nitrogen assimilation were different at each stage and location of sugar beet.In leaves the peak values of enNRA,GSA,GOGATA and GDHA were all before the tuber growth stage.In roots GOGATA was higher at the prophase and GSA and GDHA were bigger in the late period.In location,GSA of roots at other stages was much higher than that of leaves besides at seedling stage;GOGATA of leaves was very higher than that of roots besides in the foliage formation period and the end period of the foliage formation;and GDHA of leaves was bigger than that of roots.And the dynamic changes of GSA,GOGATA and GDHA in roots and leaves were all different.The relationships were different among the key enzymes of nitrogen assimilation.Before the foliage formation period there were significantly positive correlations between enNRA and exNRA,NRA and GSA and GSA of roots and leaves. After the foliage formation period there were significantly negative correlations between GSA and GOGATA.At different stages the relationships were different between the activities of key enzymes of nitrogen assimilation and yield and quality.Before the root growth stage there were closely relationships between them and yield and sugar yield,and from the root growth stage extremely correlations were observed with sugar contents.And before the foliage formation period the key enzymes activities of nitrogen assimilation in leaves were significantly related with yield, from the foliage formation period to the end stage of tuber growth there were significant correlations between GSA of roots and yield and quality,and at the sugar accumulation stage NRA became the main factor.In the end stage of tuber growth the effect of GSA of roots on yield and quality were highest.
The dynamic changes of the main products contents of nitrogen assimilation of sugar beet were different.The contents of soluble protein of roots slowly decreased before the root growth stage,quickly increased after the root growth stage,and at the sugar accumulation stage got the highest level.The contents of soluble protein of leaves increased with growth and development stage,decreased from seedling stage to the end of August,after the end stage of tuber growth increased.The total contents of nitrogen of leaves decreased from seedling stage to the end period of the foliage formation,from the end period of the foliage formation to the root growth stage increased,and till the sugar accumulation stage decreased.At different stages the correlations among the main products of nitrogen assimilation were different.At different stages the relationships were different between the contents of main products of nitrogen assimilation and yield and quality.From seedling stage to the sugar accumulation stage the main products of nitrogen assimilation were related with yield and quality.And at the root growth stage the correlation coefficients between the contents of total nitrogen and soluble protein and yield and quality attained to the higher level at the same time,so the contents of total nitrogen and soluble protein can be used as reference indexes to predict yield and quality of sugar beet in the root growth period.
And at different growth and development stages there were different correlations between the key enzymes and main products of nitrogen assimilation of sugar beet.
2.The variation laws of the key enzymes activities of sucrose metabolism were different at each stage.In roots,the dynamic changes of SPSA,NIA,AIA and the decomposition activities of SS were similar,and in leaves the ones of SPSA and SS were similar.The dynamic changes of activities of other key enzymes were different.In sampling location,SPSA,the synthesis activities of SS(besides seedling stage) and NIA of roots were higher than that of leaves at whole growth and development stage;at the middle and late stage the decomposition activities of SS of roots were smaller than that of leaves,at other stages were higher than leaves;AIA of roots were higher than leaves at seedling and late stage,at the middle stage were lower leaves.With SPSA,in roots the synthesis activities of SS at each stage were higher than SPSA except for were lower than SPSA;in leaves the activities of them were very close.With AIA,the maximums of NIA of roots and leaves were all lower than AIA.With NIA and AIA,the decomposition activities of SS of roots were all lower than NIA and AIA besides in the beginning period of the foliage formation, and the activities of 3 enzymes in leaves were all low,and the diversity were small.At different stages the correlations of the key enzymes of sugar metabolism were different.At seedling stage there were significantly positive correlations among the decomposition enzymes of sugar metabolism of roots;in the beginning period of the foliage formation there were significantly positive correlations between SPSA and the synthesis activities of SS in leaves and the decomposition enzymes of leaves,and in roots there were significantly positive correlations between the synthesis and decomposition activities of SS;in the foliage formation period there were correlations among each enzymes besides the synthesis activities of SS;there were significantly positive correlations among each enzymes of roots and leaves in the end period of the foliage formation and among each enzymes of roots at root growth stage;there were correlations between SPSA and the synthesis activities of SS in roots and each enzymes.In the end stage of tuber growth there were significant correlations between SPSA of leaves,the synthesis activities of SS of roots and each enzymes of sucrose metabolism at sugar accumulation stage.
The dynamic changes of the main products contents of sucrose metabolism of sugar beet were different.The total contents of soluble sugar in roots and leaves formed a single peak curve,and their peak values were in roots growth period and the end period of the foliage formation respectively.The fructose contentss of roots changed like wave motion,its troughs were in the foliage formation and roots growth period respectively.The change of fructose contents in leaves was like "V",and its minimum was in the beginning period of the foliage formation.The contents of sucrose in roots gradually increased from seedling to roots growth stage,and in the end stage of tuber growth slightly decreased,but kept higher level till sugar accumulation stage.The contents of sucrose in leaves slowly increased from seedling to the end period of the foliage formation;in the end period of the foliage formation got the maximum;later slightly decreased;from the sugar accumulation stage slowly increased again.At whole growth and development stage the contents of soluble sugar,fructose and sucrose of roots were all higher than those of leaves.At different stages the correlations among the main products of sucrose metabolism were different.And above-mentioned analysis showed:in the foliage formation initial period fructose was the most important product of sucrose metabolism;in the foliage formation period sucrose was uppermost product.At whole stage the contents of fructose in soluble sugar of leaves was higher at all time, and at the same time the contents of sucrose in leaves from the root growth to sugar accumulation stage quickly increased.And at whole stage the contents of fructose and sucrose in roots all changed all together;the contents of fructose and sucrose in leaves all changed together from the root growth to sugar accumulation stage.
At different growth and development stages there were different correlations between the key enzymes and main products of sucrose metabolism of sugar beet.To sampling locations there were significant correlations between the key enzymes activities of sucrose metabolism and the main products of sucrose metabolism of leaves at prophase and after the foliage formation period;in the foliage formation period there were significant correlations between the key enzymes activities of sucrose metabolism and the main products of sucrose metabolism of roots.With stages at seedling stage the decomposition enzymes of sucrose metabolism had significant effect on the products;in the beginning period of the foliage formation the contents of main products were affected significantly by the decomposition and synthesis enzymes;in the foliage formation period the effect of the synthesis enzymes was slightly in the ascendant;in the end period of the foliage formation effect of the decomposition enzymes on main products became more important;at root growth stage and in the end stage of tuber growth effect of the synthesis enzymes on main products was higher than the decomposition enzymes.
At all stages there were almost significant correlations between the key enzymes activities of sucrose metabolism and yield,sugar contents and sugar yield.At all stages there were significant correlations between the main products of sucrose metabolism and yield,sugar contents and sugar yield besides at seedling stage and in.the end stage of tuber growth.At the same time the main products were all related extremely with sugar contents,and at root growth stage were all significantly related with yield,sugar contents and sugar yield.
3.At different growth and development stages there were different correlations between the key enzymes of nitrogen assimilation and of sucrose metabolism of sugar beet.The relationships between NRA and the decomposition activities of SS and SPSA of roots were close at prophase, and at late stage there were significantly positive correlations between NIA and AIA.From the end period of the foliage formation to the end stage of tuber growth there were significantly positive correlations between GSA of roots and the key enzymes of sugar metabolism of roots at different levels.GSA of leaves were closely related with the key enzymes of sugar metabolism at initial and telophase stage,and were related with the key enzymes of sugar metabolism of leaves in the middle stage.At different stages GOGATA of leaves were related with the key enzymes of sugar metabolism at different levels.
At different stages the relationships between the key enzymes of nitrogen assimilation and the main products of sucrose metabolism were different.NRA of leaves was related significantly with the fructose contents of roots.The correlations of GSA of roots and the main products of sucrose metabolism were not remarkable.There were significant correlations between GSA of leaves and the contents of fructose and soluble sugar of leaves at the initial and late stage,and in the middle period the relationships between the contents of fructose and soluble sugar were close.GOGATA of leaves were related significantly positively with the main products of sucrose metabolism of roots in the beginning period of the foliage formation,were related extremely negatively with the main products in the end period of the foliage formation,and were related extremely positively with the fructose contents of roots in the end stage of tuber growth.
At different stages the relationships of the main products of nitrogen assimilation and the key enzymes of sucrose metabolism were different.At whole growth and development stages there were extremely significant negative correlations between the contents of soluble protein of roots and the enzymes activities of sucrose metabolism of roots.The contents of soluble protein in leaves were related significantly negatively with the enzymes activities of sucrose metabolism of leaves,were related significantly positively with the enzymes activities of sucrose metabolism of roots except for at the foliage formation period were related extremely significantly negatively with the synthesis activities of SS of roots.The correlations between the total contents of nitrogen in roots or leaves and the key enzymes of sucrose metabolism were similar.Effect of SSA and NIA on the total contents of Nitrogen in roots and leaves were all significant.
At different stages the relationships of the main products of nitrogen assimilation and the ones of sucrose metabolism were different.In the middle stage the contents of soluble protein in roots were related significantly with the main products contents of sucrose metabolism,in the beginning period of the foliage formation and the foliage formation period there were significantly positive correlations between the contents of soluble protein in leaves and the contents of sucrose and soluble sugar in leaves.The total contents of N in roots were related significantly with the main products contents of sucrose metabolism from seedling to foliage formation period,were related remarkable only with the main products contents of sucrose metabolism from the end period of the foliage formation to the turb growth period.The correlations between the total contents of N in leaves and the main products contents of sucrose metabolism were close from the beginning period of the foliage formation to the end stage of tuber growth.
4.The dynamic changes of related physiological indexes of nitrogen and sucrose metabolism in sugar beet were all different.The dynamic changes of C/N in roots and leaves were similar. From seedling stage to root growth stage C/N all increased,decreased slightly after the end stage of tuber growth.At all stages C/N of roots were higher than that of leaves.The contents of chlorophyll decreased from seedling stage to the beginning period of the foliage formation,later quickly rose,in the foliage formation period got the peak value,from 9~(th) July descended again,till sugar accumulation stage attained to the lowest level.The dry weights of roots increased with the growth and development stage;the dry weights of leaves enhanced from seedling stage to root growth stage,later decreased,after the end stage of tuber growth little increased.The ratio of crown and root decreased with the growth and development stage,was near 1 in the end period of the foliage formation.So it can be used as reference indexes to reflect the status of growth and the center growth of sugar beet.
At different stages the relationships between related physiological indexes of nitrogen and sucrose metabolism and yield,sugar contents and sugar yield were different in sugar beet.In the beginning period of the foliage formation and the end period of the foliage formation related physiological indexes were significantly related with yield and quality,and effects of chlorophyll contents,dry weight of roots and ratio of crown and root on yield and quality were larger.And at whole stages related physiological indexes were related extremely significantly with the sugar contents,were related significantly with yield,were related with sugar yield at different levels.
5.At different stages regulation effects of NO_3~--N,NH_4~+-N,MgO and P_2O_5 on the key enzymes activities of nitrogen assimilation and sucrose metabolism were different.There were obvious interactions between two factors,and in interaction the main factor were different.At the same time the contributor of four treatments on the key enzymes activities of nitrogen assimilation and sucrose metabolism were different.And we obtain the different treatment combinations at the highest activities of key enzymes of nitrogen assimilation and sucrose metabolism of sugar beet when the effects of four factors were outstanding.
6.Effects of different treatment factors on yield,sugar contents and sugar yield were different. Contributors on yield and sugar yield were all in order:MgO＞NO_3~--N＞NH_4~+-N＞P_2O_5,and contributor on sugar contents were in order:MgO＞NH_4~+-N＞NO_3~--N＞P_2O_5.And we obtain the different treatment combinations at the highest yield,sugar contents and sugar yield of sugar beet.
1.甜菜氮素同化关键酶活性的变化规律:从时间上,叶片中enNRA、GSA、GOGATA和GDHA峰值均在块根增长期之前;块根中,GOGATA在前期活性较高,而GSA和GDHA在后期影响较大。在空间上,除苗期略低于叶片外,其它各时期块根的GSA均远远高于叶片;叶片GOGATA除了在叶丛形成期和叶丛形成末期略低于块根外,其它各时期均远远高于块根;而叶片GDHA均高于块根。且GSA、GOGATA和GDHA在块根和叶片中的变化动态均不同。氮素同化关键酶间呈不同程度相关。在叶丛形成期之前主要是内源和外源NRA间、NRA与GSA间及块根和叶片GSA间显著正相关;在叶丛形成期及之后主要是GSA与GOGATA间显著负相关。氮素同化关键酶活性与块根产量和品质的关系:在块根增长期之前与块根的产量和产糖量关系密切;从块根增长期开始,与块根含糖率相关均极显著。且叶丛形成期之前叶片的关键酶活性与产量相关密切;叶丛形成期之后直到块根增长末期块根的GSA对产量和品质的影响最大;到糖分积累期NRA成为影响甜菜块根含糖率最主要因素。
甜菜氮素同化主要产物含量的变化动态:块根可溶性蛋白含量在块根增长期之前基本呈缓慢下降趋势,之后大幅增加;叶片可溶性蛋白含量在整个生育期呈逐渐增加趋势。块根全氮量从苗期到块根增长末期一直下降,之后略有上升;叶片全氮量从苗期到叶丛形成末期一直下降,之后直到块根增长期上升,随后呈下降趋势。氮素同化主要产物间呈现不同程度的相关关系。氮素同化产物与块根的产量和品质在整个生育时期基本均呈不同程度相关。且在块根增长期全氮量和可溶性蛋白含量与块根产量和品质的相关系数同时达到较高水平,可以作为块根增长期预测产量、含糖率和产糖量的参考指标。
同时甜菜氮素同化关键酶与主要同化产物在不同时期呈现不同程度相关。
2.甜菜蔗糖代谢关键酶活性的变化规律:块根中SPSA、SS分解方向活性、NIA和AIA的变化动态相似,叶片中SPSA与SS合成方向活性变化动态相似,其它各关键酶活性的变化动态均不同。从空间来看,块根的SPSA、SS合成方向活性(除苗期)、NIA在整个生育时期均高于叶片;块根的SS分解活性在中后期低于叶片,其它时期均高于叶片;块根的AIA在苗期和后期高于叶片,在中期低于叶片。SS合成活性与SPSA相比,在块根中,除苗期和叶丛形成初期略低于SPSA,其它时期均表现为高于SPSA;在叶片中,二者的活性大小接近。NIA与AIA相比,块根和叶片NIA的峰值均低于AIA。SS分解活性与NIA和A1A相比,块根中SS分解活性除叶丛形成初期大于NIA和AIA之外,其它时期均小于NIA和AIA;叶片中三种酶均处于较低水平,差异不大。不同时期蔗糖代谢关键酶间相关关系:苗期块根中糖代谢分解方向酶间的同步性很强;其它各时期SPSA和SSA(包括合成和分解方向)与其它糖代谢关键酶呈不同程度相关。
甜菜蔗糖代谢各主要产物的变化动态:块根和叶片可溶性总糖含量均呈单峰曲线变化,峰值分别在块根增长期和叶丛形成末期。块根的果糖含量在整个生育时期呈波动变化,分别在叶丛形成期和块根增长期各有一个波谷。叶片的果糖含量呈“V”型变化,在叶丛形成初期降至最低水平。块根的蔗糖含量从苗期到块根增长期逐渐增加,之后略有下降,但直到糖分积累期仍保持较高水平。叶片的蔗糖含量从苗期到叶丛形成末期呈缓慢增加趋势,之后略微下降,在块根增长期又缓慢上升。且整个生育期块根的可溶性总糖、果糖和蔗糖含量均大于叶片。蔗糖代谢主要产物间在不同时期呈现不同程度的相关关系。且叶丛形成初期果糖是块根糖代谢的最主要产物,叶丛形成期蔗糖是块根糖代谢的最主要产物。而整个生育时期果糖在叶片可溶性总糖中所占比例一直处于领先地位,同时从块根增长期直到糖分积累期叶片中蔗糖含量也迅速增加。并且块根中果糖和蔗糖含量在整个生育期间均为同步变化,叶片中果糖和蔗糖含量从块根增长期到糖分积累期也是同步变化的。
甜菜蔗糖代谢关键酶与主要产物的关系:从部位来看,蔗糖代谢关键酶活性在前期和叶丛形成期之后与叶片糖代谢主要产物、在叶丛形成期与块根糖代谢主要产物相关显著。从时期来看,苗期糖代谢分解方向的酶类对产物的影响均较大;在叶丛形成初期分解和合成方向的酶类均具有一定影响;在叶丛形成期合成方向酶类的影响略占优势;叶丛形成末期分解方向酶类的影响优势变大;在块根增长期和块根增长末期合成方向酶类作用又大于分解;糖分积累期,蔗糖代谢分解和合成方向的酶类对蔗糖代谢产物均具有一定影响。
甜菜蔗糖代谢关键酶活性与产量和品质在整个生育期基本都显著相关。蔗糖代谢主要产物与产量和品质均呈不同程度相关(除苗期和块根增长末期)。
3.甜菜氮素同化关键酶与蔗糖代谢关键酶的关系:NRA与块根SS分解方向活性、SPSA在生育前期关系较密切,与NIA和AIA在后期表现很强的同步性。块根中GSA与块根的糖代谢各关键酶活性间从叶丛形成末期到块根增长末期均呈不同程度的正相关,而叶片GSA与糖代谢关键酶活性在生育初期和末期相关更为密切,而在中期主要与叶片的糖代谢关键酶活性呈不同程度相关。叶片GOGATA与SPSA在苗期极显著正相关;叶丛形成初期与糖代谢各酶活性相关均不显著;叶丛形成期到块根增长末期与糖代谢合成方向酶类显著相关;糖分积累期与糖代谢合成与分解方向酶类均呈不同程度相关。
甜菜氮素同化关键酶与蔗糖代谢主要产物的关系:叶片NRA主要与果糖和蔗糖含量关系密切。块根GSA与蔗糖代谢主要产物含量在整个生育期相关均不显著,而叶片GSA在生育初期和后期对叶片的果糖和可溶性总糖含量影响较大,在中期与块根果糖和可溶性总糖含量关系密切。叶片GOGATA在叶丛形成初期与块根蔗糖代谢主要产物均呈同步变化趋势,在叶丛形成末期变化不再同步,到块根增长末期只与块根果糖含量同步变化。
甜菜氮素同化主要产物与蔗糖代谢关键酶的关系:块根可溶性蛋白含量在整个生育期与块根糖代谢各酶活性基本都呈极显著负相关。叶片可溶性蛋白含量与叶片糖代谢各酶活性均呈显著负相关;与块根糖代谢各酶活性基本均呈显著正相关。块根、叶片全氮量与糖代谢各酶的相关关系相似,SSA(分解和合成方向)和NIA对块根和叶片全氮量的影响均较大。
氮素同化与蔗糖代谢主要产物间的关系:块根可溶性蛋白含量与蔗糖代谢主要产物含量间在苗期和生育末期相关较弱,在中期相关较密切。叶片可溶性蛋白与叶片蔗糖、可溶性总糖含量在叶丛形成初期和叶丛形成期显著正相关,其它时期相关均不显著。块根全氮量与蔗糖代谢主要产物含量从苗期到叶丛形成期关系均较密切,而从叶丛形成末期到块根增长期仅与叶片蔗糖代谢主要产物含量相关紧密。并且块根全氮量与蔗糖代谢主要产物含量间均为负相关关系。叶片全氮量与蔗糖代谢主要产物含量间在苗期和糖分积累期关系均较弱;而从叶丛形成初期到块根增长末期关系密切。
4.甜菜氮糖代谢的相关生理指标的变化动态:块根与叶片C/N从苗期到块根增长期均呈增加趋势,之后略有下降。且整个生育期块根C/N均高于叶片。叶绿素含量从苗期到叶丛形成初期呈下降趋势,之后迅速上升,到叶丛形成期达到最高,随后下降,直至糖分积累期降至最低水平。块根干物质积累量在整个生育期呈增加趋势;叶片干物质积累量从苗期到块根增长期呈增加趋势,之后下降,块根增长末期之后又略有上升。冠根比在整个生育时期一直呈下降趋势。在叶丛形成末期冠根比接近1。因此,可用叶丛形成末期冠根比来反映甜菜植株的生长状况及生长中心转移情况,适时进行人工调控。
甜菜氮糖代谢的相关生理指标与产量和品质的关系:叶丛形成初期和叶丛形成末期与产量和品质关系最为密切,并以叶绿素含量、块根干重和冠根比的影响较大。且在整个生育期相关生理指标对含糖率的影响均极显著,对产量的影响均未达到极显著水平,对产糖量的影响程度在各时期不同。
5.硝态氮、铵态氮、镁肥和磷肥四种处理因素在不同时期对甜菜氮素同化和蔗糖代谢关键酶活性调控效应不同,而且两处理因素之间存在明显的交互作用,在交互作用中对氮素同化和蔗糖代谢关键酶活性起主导作用的因子不同,同时在不同生育时期四种处理因素对甜菜氮素同化和蔗糖代谢关键酶活性的贡献率不同。硝态氮和铵态氮在整个生育期均具有不同程度的贡献率;镁肥和磷肥在后期的贡献率较高。研究还得到四种处理因素影响显著时期甜菜氮素同化和蔗糖代谢关键酶活性最高时四个处理因子组合。
6.不同处理因素对甜菜产量、含糖率和产糖量的影响不同。对产量和产糖量的贡献率顺序均为:镁肥＞硝态氮＞铵态氮＞磷肥,对含糖率的贡献率顺序为:镁肥＞铵态氮＞硝态氮＞磷肥。研究还得到甜菜块根产量、含糖率和产糖量最高时硝态氮、铵态氮、镁肥和磷肥四个处理因子组合。
In this dissertation,Tianyan 7 of NZ sugar beet was selected as experimental sample,and experiments were made in the campus and practice base of NEAU from 2006 to 2007. Experimental factors were as follows:NO_3~--N,NH_4~+-N,MgO and P_2O_5.In experiments the quadratic rotation-orthogonal combination design was adopted.We comprehensively and systematically studied on dynamic changes and regulation approaches of key enzymes and main products of nitrogen assimilation and sucrose metabolism of sugar beet,the mechanism of nitrogen assimilation and sucrose metabolism,further analyzed interaction mechanism between nitrogen assimilation and sucrose metabolism,and announced the regulation effect of nitrogen, phosphorus and magnesium fertilizer on nitrogen assimilation,sucrose metabolism,yield and quality.This dissertation will fill in the infrastructure data of sugar beet physiology,provide theoretical basis for identifying,evaluating,reforming and using germplasm resources,and making reasonable cultivation measurement.Results of this experiment are as following:
1.The variation laws of the activities of key enzymes of nitrogen assimilation were different at each stage and location of sugar beet.In leaves the peak values of enNRA,GSA,GOGATA and GDHA were all before the tuber growth stage.In roots GOGATA was higher at the prophase and GSA and GDHA were bigger in the late period.In location,GSA of roots at other stages was much higher than that of leaves besides at seedling stage;GOGATA of leaves was very higher than that of roots besides in the foliage formation period and the end period of the foliage formation;and GDHA of leaves was bigger than that of roots.And the dynamic changes of GSA,GOGATA and GDHA in roots and leaves were all different.The relationships were different among the key enzymes of nitrogen assimilation.Before the foliage formation period there were significantly positive correlations between enNRA and exNRA,NRA and GSA and GSA of roots and leaves. After the foliage formation period there were significantly negative correlations between GSA and GOGATA.At different stages the relationships were different between the activities of key enzymes of nitrogen assimilation and yield and quality.Before the root growth stage there were closely relationships between them and yield and sugar yield,and from the root growth stage extremely correlations were observed with sugar contents.And before the foliage formation period the key enzymes activities of nitrogen assimilation in leaves were significantly related with yield, from the foliage formation period to the end stage of tuber growth there were significant correlations between GSA of roots and yield and quality,and at the sugar accumulation stage NRA became the main factor.In the end stage of tuber growth the effect of GSA of roots on yield and quality were highest.
The dynamic changes of the main products contents of nitrogen assimilation of sugar beet were different.The contents of soluble protein of roots slowly decreased before the root growth stage,quickly increased after the root growth stage,and at the sugar accumulation stage got the highest level.The contents of soluble protein of leaves increased with growth and development stage,decreased from seedling stage to the end of August,after the end stage of tuber growth increased.The total contents of nitrogen of leaves decreased from seedling stage to the end period of the foliage formation,from the end period of the foliage formation to the root growth stage increased,and till the sugar accumulation stage decreased.At different stages the correlations among the main products of nitrogen assimilation were different.At different stages the relationships were different between the contents of main products of nitrogen assimilation and yield and quality.From seedling stage to the sugar accumulation stage the main products of nitrogen assimilation were related with yield and quality.And at the root growth stage the correlation coefficients between the contents of total nitrogen and soluble protein and yield and quality attained to the higher level at the same time,so the contents of total nitrogen and soluble protein can be used as reference indexes to predict yield and quality of sugar beet in the root growth period.
And at different growth and development stages there were different correlations between the key enzymes and main products of nitrogen assimilation of sugar beet.
2.The variation laws of the key enzymes activities of sucrose metabolism were different at each stage.In roots,the dynamic changes of SPSA,NIA,AIA and the decomposition activities of SS were similar,and in leaves the ones of SPSA and SS were similar.The dynamic changes of activities of other key enzymes were different.In sampling location,SPSA,the synthesis activities of SS(besides seedling stage) and NIA of roots were higher than that of leaves at whole growth and development stage;at the middle and late stage the decomposition activities of SS of roots were smaller than that of leaves,at other stages were higher than leaves;AIA of roots were higher than leaves at seedling and late stage,at the middle stage were lower leaves.With SPSA,in roots the synthesis activities of SS at each stage were higher than SPSA except for were lower than SPSA;in leaves the activities of them were very close.With AIA,the maximums of NIA of roots and leaves were all lower than AIA.With NIA and AIA,the decomposition activities of SS of roots were all lower than NIA and AIA besides in the beginning period of the foliage formation, and the activities of 3 enzymes in leaves were all low,and the diversity were small.At different stages the correlations of the key enzymes of sugar metabolism were different.At seedling stage there were significantly positive correlations among the decomposition enzymes of sugar metabolism of roots;in the beginning period of the foliage formation there were significantly positive correlations between SPSA and the synthesis activities of SS in leaves and the decomposition enzymes of leaves,and in roots there were significantly positive correlations between the synthesis and decomposition activities of SS;in the foliage formation period there were correlations among each enzymes besides the synthesis activities of SS;there were significantly positive correlations among each enzymes of roots and leaves in the end period of the foliage formation and among each enzymes of roots at root growth stage;there were correlations between SPSA and the synthesis activities of SS in roots and each enzymes.In the end stage of tuber growth there were significant correlations between SPSA of leaves,the synthesis activities of SS of roots and each enzymes of sucrose metabolism at sugar accumulation stage.
The dynamic changes of the main products contents of sucrose metabolism of sugar beet were different.The total contents of soluble sugar in roots and leaves formed a single peak curve,and their peak values were in roots growth period and the end period of the foliage formation respectively.The fructose contentss of roots changed like wave motion,its troughs were in the foliage formation and roots growth period respectively.The change of fructose contents in leaves was like "V",and its minimum was in the beginning period of the foliage formation.The contents of sucrose in roots gradually increased from seedling to roots growth stage,and in the end stage of tuber growth slightly decreased,but kept higher level till sugar accumulation stage.The contents of sucrose in leaves slowly increased from seedling to the end period of the foliage formation;in the end period of the foliage formation got the maximum;later slightly decreased;from the sugar accumulation stage slowly increased again.At whole growth and development stage the contents of soluble sugar,fructose and sucrose of roots were all higher than those of leaves.At different stages the correlations among the main products of sucrose metabolism were different.And above-mentioned analysis showed:in the foliage formation initial period fructose was the most important product of sucrose metabolism;in the foliage formation period sucrose was uppermost product.At whole stage the contents of fructose in soluble sugar of leaves was higher at all time, and at the same time the contents of sucrose in leaves from the root growth to sugar accumulation stage quickly increased.And at whole stage the contents of fructose and sucrose in roots all changed all together;the contents of fructose and sucrose in leaves all changed together from the root growth to sugar accumulation stage.
At different growth and development stages there were different correlations between the key enzymes and main products of sucrose metabolism of sugar beet.To sampling locations there were significant correlations between the key enzymes activities of sucrose metabolism and the main products of sucrose metabolism of leaves at prophase and after the foliage formation period;in the foliage formation period there were significant correlations between the key enzymes activities of sucrose metabolism and the main products of sucrose metabolism of roots.With stages at seedling stage the decomposition enzymes of sucrose metabolism had significant effect on the products;in the beginning period of the foliage formation the contents of main products were affected significantly by the decomposition and synthesis enzymes;in the foliage formation period the effect of the synthesis enzymes was slightly in the ascendant;in the end period of the foliage formation effect of the decomposition enzymes on main products became more important;at root growth stage and in the end stage of tuber growth effect of the synthesis enzymes on main products was higher than the decomposition enzymes.
At all stages there were almost significant correlations between the key enzymes activities of sucrose metabolism and yield,sugar contents and sugar yield.At all stages there were significant correlations between the main products of sucrose metabolism and yield,sugar contents and sugar yield besides at seedling stage and in.the end stage of tuber growth.At the same time the main products were all related extremely with sugar contents,and at root growth stage were all significantly related with yield,sugar contents and sugar yield.
3.At different growth and development stages there were different correlations between the key enzymes of nitrogen assimilation and of sucrose metabolism of sugar beet.The relationships between NRA and the decomposition activities of SS and SPSA of roots were close at prophase, and at late stage there were significantly positive correlations between NIA and AIA.From the end period of the foliage formation to the end stage of tuber growth there were significantly positive correlations between GSA of roots and the key enzymes of sugar metabolism of roots at different levels.GSA of leaves were closely related with the key enzymes of sugar metabolism at initial and telophase stage,and were related with the key enzymes of sugar metabolism of leaves in the middle stage.At different stages GOGATA of leaves were related with the key enzymes of sugar metabolism at different levels.
At different stages the relationships between the key enzymes of nitrogen assimilation and the main products of sucrose metabolism were different.NRA of leaves was related significantly with the fructose contents of roots.The correlations of GSA of roots and the main products of sucrose metabolism were not remarkable.There were significant correlations between GSA of leaves and the contents of fructose and soluble sugar of leaves at the initial and late stage,and in the middle period the relationships between the contents of fructose and soluble sugar were close.GOGATA of leaves were related significantly positively with the main products of sucrose metabolism of roots in the beginning period of the foliage formation,were related extremely negatively with the main products in the end period of the foliage formation,and were related extremely positively with the fructose contents of roots in the end stage of tuber growth.
At different stages the relationships of the main products of nitrogen assimilation and the key enzymes of sucrose metabolism were different.At whole growth and development stages there were extremely significant negative correlations between the contents of soluble protein of roots and the enzymes activities of sucrose metabolism of roots.The contents of soluble protein in leaves were related significantly negatively with the enzymes activities of sucrose metabolism of leaves,were related significantly positively with the enzymes activities of sucrose metabolism of roots except for at the foliage formation period were related extremely significantly negatively with the synthesis activities of SS of roots.The correlations between the total contents of nitrogen in roots or leaves and the key enzymes of sucrose metabolism were similar.Effect of SSA and NIA on the total contents of Nitrogen in roots and leaves were all significant.
At different stages the relationships of the main products of nitrogen assimilation and the ones of sucrose metabolism were different.In the middle stage the contents of soluble protein in roots were related significantly with the main products contents of sucrose metabolism,in the beginning period of the foliage formation and the foliage formation period there were significantly positive correlations between the contents of soluble protein in leaves and the contents of sucrose and soluble sugar in leaves.The total contents of N in roots were related significantly with the main products contents of sucrose metabolism from seedling to foliage formation period,were related remarkable only with the main products contents of sucrose metabolism from the end period of the foliage formation to the turb growth period.The correlations between the total contents of N in leaves and the main products contents of sucrose metabolism were close from the beginning period of the foliage formation to the end stage of tuber growth.
4.The dynamic changes of related physiological indexes of nitrogen and sucrose metabolism in sugar beet were all different.The dynamic changes of C/N in roots and leaves were similar. From seedling stage to root growth stage C/N all increased,decreased slightly after the end stage of tuber growth.At all stages C/N of roots were higher than that of leaves.The contents of chlorophyll decreased from seedling stage to the beginning period of the foliage formation,later quickly rose,in the foliage formation period got the peak value,from 9~(th) July descended again,till sugar accumulation stage attained to the lowest level.The dry weights of roots increased with the growth and development stage;the dry weights of leaves enhanced from seedling stage to root growth stage,later decreased,after the end stage of tuber growth little increased.The ratio of crown and root decreased with the growth and development stage,was near 1 in the end period of the foliage formation.So it can be used as reference indexes to reflect the status of growth and the center growth of sugar beet.
At different stages the relationships between related physiological indexes of nitrogen and sucrose metabolism and yield,sugar contents and sugar yield were different in sugar beet.In the beginning period of the foliage formation and the end period of the foliage formation related physiological indexes were significantly related with yield and quality,and effects of chlorophyll contents,dry weight of roots and ratio of crown and root on yield and quality were larger.And at whole stages related physiological indexes were related extremely significantly with the sugar contents,were related significantly with yield,were related with sugar yield at different levels.
5.At different stages regulation effects of NO_3~--N,NH_4~+-N,MgO and P_2O_5 on the key enzymes activities of nitrogen assimilation and sucrose metabolism were different.There were obvious interactions between two factors,and in interaction the main factor were different.At the same time the contributor of four treatments on the key enzymes activities of nitrogen assimilation and sucrose metabolism were different.And we obtain the different treatment combinations at the highest activities of key enzymes of nitrogen assimilation and sucrose metabolism of sugar beet when the effects of four factors were outstanding.
6.Effects of different treatment factors on yield,sugar contents and sugar yield were different. Contributors on yield and sugar yield were all in order:MgO＞NO_3~--N＞NH_4~+-N＞P_2O_5,and contributor on sugar contents were in order:MgO＞NH_4~+-N＞NO_3~--N＞P_2O_5.And we obtain the different treatment combinations at the highest yield,sugar contents and sugar yield of sugar beet.
引文
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