氮素对茶树叶片品质成分影响机理研究
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摘要
茶树是我国一种重要的经济作物,其叶片中含有丰富的代谢产物,容易受到环境和栽培措施的影响。氮(N)对茶树生长及品质成分有重要的影响。茶树生长需要适宜的施N量,从而在保证最大经济效益的同时,避免过量施N对环境造成的危害和对茶叶品质的降低。同时,茶树对不同形态的N具有不同的吸收特性,而对于其原初机制及调控机理缺少研究报道。本文主要研究结果如下：
1.茶树叶片品质成分浸提方法比较及应用
本文比较了不同浸提剂(甲醇、乙醇和高氯酸)和不同研磨方法(球磨机和研钵研磨),优化出一种针对茶树鲜叶浸提的快速前处理方法(70%乙醇,球磨机研磨5分钟)。该方法可以较好的浸提茶树鲜叶中的茶多酚和氨基酸。同时比较了不同前处理叶片对茶叶品质成分浸提效果的影响,并应用于茶树不同叶位品质成分差异比较。
2.基于代谢谱分析氮素水平和光照对茶树叶片品质成分影响
设立不同施N水平与光/暗处理的交互试验,探讨基于1H-NMR的茶树叶片代谢组学是否可以有效的区分不同处理的样本。1H-NMR分析结果表明,光／暗处理对茶树代谢组影响较大,两个处理样本可以显著分离,其中,差异较大的组分为茶氨酸(Thea)、葡萄糖(Gluc)和蔗糖(Suc)。不同施N水平下茶树新梢二叶代谢组没有显著差异。而定量分析结果表明,施N水平和光／暗处理对儿茶素组分均有显著影响。
3.氮素水平对茶树叶片氮素营养及SPAD值影响
测定叶片N含量是诊断茶树N素状况的一项重要方法。本文通过盆栽结合大田试验,对施N水平影响茶树产量及茶树叶片N素水平进行了研究,同时,研究了叶绿素仪(SPAD)在茶树N素营养快速诊断上的适用性。结果表明,茶树新梢产量,全N含量和游离氨基酸总量与施N水平呈线性加平台的关系。SPAD值、叶绿素含量及全N含量三者之间呈显著线性正相关,SPAD值随着N素水平增加同样表现为线性加平台的趋势。结果表明SPAD可以作为快速诊断方法用于指导茶树施N,尤其是缺N茶树的推荐施肥。本文还探讨了测定位点、叶位、天气状况、田间原位或离体测定方式、表面清洗以及样本量等因素对SPAD测试精度的影响。
4.茶树对不同形态氮源吸收代谢差异机制研究
对于主要的两种无机N源铵态氮(NH4+-N)和硝态氮(N03--N),茶树偏好NH4+-N。本文利用15N标记营养液培养的茶苗,通过测定其吸收15N量计算吸收动力学参数,来比较茶树对不同N源吸收系统的差异。同时,运用气相色谱-质谱联用仪(GC-MS)分析15N进入主要氨基酸组分的丰度,从而了解不同形态N源同化途径的差异机制。结果表明,’5NH4+-N培养的茶苗吸收的15N量显著高于‘5N03--N培养的茶苗。其吸收动力学曲线表明茶树吸收不同N源均存在高、低两个亲和吸收系统。计算得到的动力学参数最大吸收速度(Vmax)值15NH4+-N为54.5nmol/g DW/min,高于15NO3--N39.3(nmol/g DW/min),两种N源的表观米氏常数(Km)值分别为0.06mmol/L和0.16mmol/L。15NH4+-N培养的茶苗同化15N的量远远高于15N03--N培养的茶苗,尤其是谷氨酰胺(Gln)的量。Gln、谷氨酸(Glu)、Thea、丝氨酸(Ser)和天冬氨酸(Asp)是茶树N同化的主要游离氨基酸组分。15NH4+-N培养的茶苗合成Thea比15N03--N培养的茶苗快。茶树直接吸收的NH4+和N03同化生成的NH4+均通过谷氨酰胺合成酶-谷氨酸合酶(GC-GOGAT)途径同化为氨基酸。综上所述,相较于同化转运途径,茶树偏铵特性主要原因在于吸收系统的差异。Thea合成与茶树偏铵特性相关性不大。不同N素水平和不同N源的预培养抑制了茶树NH4+-N和N03--N的吸收,对于N同化过程也有影响。通过在吸收液中加入Thea和抑制剂进行分析,结果表明,茶树吸收N03--N不仅受茶树体内Thea的影响,还受N03-离子本身的影响,而茶树根系对NH4+的吸收主要受其体内NH4+或N03-两种离子本身的影响,不受Thea的影响。
Tea [Camellia sinensis (L.) O. Kuntze] is an economically important perennial crop species that contains abundant secondary metabolites. Nitrogen (N) is a key element in the life-histories of tea plants. The biosynthesis and accmulation of the quality-related metabolites in young shoots of tea plants are largely affected by N supply. The objectives of this study were to investigate the functional relationship between N nutrition and accumulation of primary and secondary metabolites in tea plants. The results are as follows:
1. Evaluation of methods for extraction of quality-related constituents in tea leaves
A simple and quick method was developed and optimised for the analysis of polyphenols and free amino acids in tea fresh shoots (leaves) in this study. The use of70%Ethanol and grinded by ball mill for5min has been proved to be the most suitable method. In addition, we compared the effect of different dried methods of tea shoots (leaves) on the quality-related constituents and evaluate it to compare the metabolites in tea plant leaves of different positions.
2. Motabolic profiling reveals the effects of different nitrogen application rate and light/dark treatment on quality of fresh tea leaves
The different cultivation methods affect tea quality by altering the basic metabolite profiles. In this study, the metabolome changes were investigated in fresh tea shoots by HPLC and1H-NMR coupled with a multivariate data set. The effect of light/dark treatment was observed through the metabolomics method but there were no difference among N application rates. However, the HPLC results showed both variation.
3. Response of foliar nitrogen and SPAD readings of tea plants to nitrogen fertilization
Maximization of benefit while offsetting deteriorating effects on quality and environmental problems of N fertilization requires reliable N recommendation in tea plantations. Determination of N concentrations of tissues is an important diagnostic means for N status of plants. Pot and field experiments were carried out to investigate the response of yield and total N of mature leaves of clone Longjing43to N and to evaluate the feasibility using SPAD-502as a diagnostic tool. The yield, total N and total free amino acids showed similar patterns with an initial linear increase followed by a platform to increasing N fertilization. A significant positive linear relationship was observed between the SPAD reading and chlorophyll contents and total N. Furthermore the SPAD reading responded to N fertilization same as for the yield, increasing with increasing N fertilization until a platform observed. The SPAD reading was likely an indicator of N status and a potentially useful and quick diagnostic tool for N fertilization recommendation especially in the range of N deficiency. Those factors which may influence the precision of SPAD measurement, such as the point of test within a leaf, selection of leaves from twig (position), weather condition, in situ or in vitro analysis, surface cleaning and the number of leaves for a composite sample were also investigated.
4. Characterization and regulation of ammonium and nitrate uptake and metabolism in roots of tea plants
It has been pointed out that tea prefers ammonium (NH4+-N) over nitrate (NO3--N) as an inorganic N source.15N studies were conducted using hydroponically grown tea plants to clarify the characteristics of uptake and assimilation of NH4+and NO3-by tea roots. The total15N was detected and kinetic parameters were calculated after feeding15NH4+or15NO3-to tea plants. The process of N assimilation was studied by monitoring the dynamic15N abundance in the free amino acids of tea plant roots by Gas Chromatography-Mass spectrometry (GC-MS). Tea plants supplied with15NH4+absorbed significantly more15N than those supplied with15NO3-. The kinetics of15NH4+and15NO3-influx in tea plants followed a classic biphasic pattern, demonstrating the action of a high affinity transport system (HATS) and a low affinity transport system (LATS). The Maximum rate of uptake (Vmax) value for NH4+uptake was54.5nmol/g DW/min, which was higher than that observed for NO3-(39.3nmol/g DW/min). Apparent michaelis constant (Km) estimates were approximately0.06mmol/L for NH4+and0.16mmol/L for NO3-, indicating a higher rate of NH4+absorption in tea plant roots. Tea plants fed with15NH4+accumulated larger amounts of assimilated N, especially glutamine (Gln), compared with those fed with15NO3-. Gln, glutamic acid (Glu), theanine (Thea), serine (Ser) and aspartic acid (Asp) were the main free amino acids that were labeled with15N under both conditions. The rate of N assimilation into Thea in the roots of NO3--supplied tea plants was quicker than in NH4+-supplied tea plants. NO3-uptake in roots, rather than reduction or transport within the plant, seems to be the main factor limiting the growth of tea plants supplied with NO3-as the sole N source. The NH4+absorbed by tea plants directly, as well as that produced by NO3-reduction, were assimilated through the glutamine synthetase-glutamine oxoglutarate aminotransferase (GS-GOGAT) pathway in tea plant roots. The15N labeling experiments show that there is no direct relationship between the Thea synthesis and the preference of tea plants for NH4+.
The inhibitory effects exerted by the pretreatment of tea plants by different level of N or two N source upon N uptake and assimilation by the roots of tea plants have been showed in this study. The regulation of NO3-uptake by Thea and NO3-were showed in our study, and the uptake of NH4+was affected mainly by the NH4+and NO3-.
1.茶树叶片品质成分浸提方法比较及应用
本文比较了不同浸提剂(甲醇、乙醇和高氯酸)和不同研磨方法(球磨机和研钵研磨),优化出一种针对茶树鲜叶浸提的快速前处理方法(70%乙醇,球磨机研磨5分钟)。该方法可以较好的浸提茶树鲜叶中的茶多酚和氨基酸。同时比较了不同前处理叶片对茶叶品质成分浸提效果的影响,并应用于茶树不同叶位品质成分差异比较。
2.基于代谢谱分析氮素水平和光照对茶树叶片品质成分影响
设立不同施N水平与光/暗处理的交互试验,探讨基于1H-NMR的茶树叶片代谢组学是否可以有效的区分不同处理的样本。1H-NMR分析结果表明,光／暗处理对茶树代谢组影响较大,两个处理样本可以显著分离,其中,差异较大的组分为茶氨酸(Thea)、葡萄糖(Gluc)和蔗糖(Suc)。不同施N水平下茶树新梢二叶代谢组没有显著差异。而定量分析结果表明,施N水平和光／暗处理对儿茶素组分均有显著影响。
3.氮素水平对茶树叶片氮素营养及SPAD值影响
测定叶片N含量是诊断茶树N素状况的一项重要方法。本文通过盆栽结合大田试验,对施N水平影响茶树产量及茶树叶片N素水平进行了研究,同时,研究了叶绿素仪(SPAD)在茶树N素营养快速诊断上的适用性。结果表明,茶树新梢产量,全N含量和游离氨基酸总量与施N水平呈线性加平台的关系。SPAD值、叶绿素含量及全N含量三者之间呈显著线性正相关,SPAD值随着N素水平增加同样表现为线性加平台的趋势。结果表明SPAD可以作为快速诊断方法用于指导茶树施N,尤其是缺N茶树的推荐施肥。本文还探讨了测定位点、叶位、天气状况、田间原位或离体测定方式、表面清洗以及样本量等因素对SPAD测试精度的影响。
4.茶树对不同形态氮源吸收代谢差异机制研究
对于主要的两种无机N源铵态氮(NH4+-N)和硝态氮(N03--N),茶树偏好NH4+-N。本文利用15N标记营养液培养的茶苗,通过测定其吸收15N量计算吸收动力学参数,来比较茶树对不同N源吸收系统的差异。同时,运用气相色谱-质谱联用仪(GC-MS)分析15N进入主要氨基酸组分的丰度,从而了解不同形态N源同化途径的差异机制。结果表明,’5NH4+-N培养的茶苗吸收的15N量显著高于‘5N03--N培养的茶苗。其吸收动力学曲线表明茶树吸收不同N源均存在高、低两个亲和吸收系统。计算得到的动力学参数最大吸收速度(Vmax)值15NH4+-N为54.5nmol/g DW/min,高于15NO3--N39.3(nmol/g DW/min),两种N源的表观米氏常数(Km)值分别为0.06mmol/L和0.16mmol/L。15NH4+-N培养的茶苗同化15N的量远远高于15N03--N培养的茶苗,尤其是谷氨酰胺(Gln)的量。Gln、谷氨酸(Glu)、Thea、丝氨酸(Ser)和天冬氨酸(Asp)是茶树N同化的主要游离氨基酸组分。15NH4+-N培养的茶苗合成Thea比15N03--N培养的茶苗快。茶树直接吸收的NH4+和N03同化生成的NH4+均通过谷氨酰胺合成酶-谷氨酸合酶(GC-GOGAT)途径同化为氨基酸。综上所述,相较于同化转运途径,茶树偏铵特性主要原因在于吸收系统的差异。Thea合成与茶树偏铵特性相关性不大。不同N素水平和不同N源的预培养抑制了茶树NH4+-N和N03--N的吸收,对于N同化过程也有影响。通过在吸收液中加入Thea和抑制剂进行分析,结果表明,茶树吸收N03--N不仅受茶树体内Thea的影响,还受N03-离子本身的影响,而茶树根系对NH4+的吸收主要受其体内NH4+或N03-两种离子本身的影响,不受Thea的影响。
Tea [Camellia sinensis (L.) O. Kuntze] is an economically important perennial crop species that contains abundant secondary metabolites. Nitrogen (N) is a key element in the life-histories of tea plants. The biosynthesis and accmulation of the quality-related metabolites in young shoots of tea plants are largely affected by N supply. The objectives of this study were to investigate the functional relationship between N nutrition and accumulation of primary and secondary metabolites in tea plants. The results are as follows:
1. Evaluation of methods for extraction of quality-related constituents in tea leaves
A simple and quick method was developed and optimised for the analysis of polyphenols and free amino acids in tea fresh shoots (leaves) in this study. The use of70%Ethanol and grinded by ball mill for5min has been proved to be the most suitable method. In addition, we compared the effect of different dried methods of tea shoots (leaves) on the quality-related constituents and evaluate it to compare the metabolites in tea plant leaves of different positions.
2. Motabolic profiling reveals the effects of different nitrogen application rate and light/dark treatment on quality of fresh tea leaves
The different cultivation methods affect tea quality by altering the basic metabolite profiles. In this study, the metabolome changes were investigated in fresh tea shoots by HPLC and1H-NMR coupled with a multivariate data set. The effect of light/dark treatment was observed through the metabolomics method but there were no difference among N application rates. However, the HPLC results showed both variation.
3. Response of foliar nitrogen and SPAD readings of tea plants to nitrogen fertilization
Maximization of benefit while offsetting deteriorating effects on quality and environmental problems of N fertilization requires reliable N recommendation in tea plantations. Determination of N concentrations of tissues is an important diagnostic means for N status of plants. Pot and field experiments were carried out to investigate the response of yield and total N of mature leaves of clone Longjing43to N and to evaluate the feasibility using SPAD-502as a diagnostic tool. The yield, total N and total free amino acids showed similar patterns with an initial linear increase followed by a platform to increasing N fertilization. A significant positive linear relationship was observed between the SPAD reading and chlorophyll contents and total N. Furthermore the SPAD reading responded to N fertilization same as for the yield, increasing with increasing N fertilization until a platform observed. The SPAD reading was likely an indicator of N status and a potentially useful and quick diagnostic tool for N fertilization recommendation especially in the range of N deficiency. Those factors which may influence the precision of SPAD measurement, such as the point of test within a leaf, selection of leaves from twig (position), weather condition, in situ or in vitro analysis, surface cleaning and the number of leaves for a composite sample were also investigated.
4. Characterization and regulation of ammonium and nitrate uptake and metabolism in roots of tea plants
It has been pointed out that tea prefers ammonium (NH4+-N) over nitrate (NO3--N) as an inorganic N source.15N studies were conducted using hydroponically grown tea plants to clarify the characteristics of uptake and assimilation of NH4+and NO3-by tea roots. The total15N was detected and kinetic parameters were calculated after feeding15NH4+or15NO3-to tea plants. The process of N assimilation was studied by monitoring the dynamic15N abundance in the free amino acids of tea plant roots by Gas Chromatography-Mass spectrometry (GC-MS). Tea plants supplied with15NH4+absorbed significantly more15N than those supplied with15NO3-. The kinetics of15NH4+and15NO3-influx in tea plants followed a classic biphasic pattern, demonstrating the action of a high affinity transport system (HATS) and a low affinity transport system (LATS). The Maximum rate of uptake (Vmax) value for NH4+uptake was54.5nmol/g DW/min, which was higher than that observed for NO3-(39.3nmol/g DW/min). Apparent michaelis constant (Km) estimates were approximately0.06mmol/L for NH4+and0.16mmol/L for NO3-, indicating a higher rate of NH4+absorption in tea plant roots. Tea plants fed with15NH4+accumulated larger amounts of assimilated N, especially glutamine (Gln), compared with those fed with15NO3-. Gln, glutamic acid (Glu), theanine (Thea), serine (Ser) and aspartic acid (Asp) were the main free amino acids that were labeled with15N under both conditions. The rate of N assimilation into Thea in the roots of NO3--supplied tea plants was quicker than in NH4+-supplied tea plants. NO3-uptake in roots, rather than reduction or transport within the plant, seems to be the main factor limiting the growth of tea plants supplied with NO3-as the sole N source. The NH4+absorbed by tea plants directly, as well as that produced by NO3-reduction, were assimilated through the glutamine synthetase-glutamine oxoglutarate aminotransferase (GS-GOGAT) pathway in tea plant roots. The15N labeling experiments show that there is no direct relationship between the Thea synthesis and the preference of tea plants for NH4+.
The inhibitory effects exerted by the pretreatment of tea plants by different level of N or two N source upon N uptake and assimilation by the roots of tea plants have been showed in this study. The regulation of NO3-uptake by Thea and NO3-were showed in our study, and the uptake of NH4+was affected mainly by the NH4+and NO3-.
引文
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