微量元素对甘草中甘草酸形成与积累影响的研究
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摘要
甘草,具有调和诸药的功效,为中国传统大宗中药材。然而,对野生甘草过度的挖掘和开发,导致了全世界范围内其野生种质资源近乎灭绝。人工种植甘草是解决当前甘草资源短缺的有效途径。但是,人工种植甘草的甘草酸含量普遍低于野生甘草。甘草酸是甘草入药的主要化学成分。因此,稳定和提高人工种植甘草质量的研究成为近年的一个研究热点。本研究的目的是定量分析B、Mn、Zn和Mo等4种微量元素对甘草根中主要的次生代谢产物、初生代谢产物、中间合成前体物质、微量元素水平及存在形态、甘草酸生物合成关键酶基因表达的影响。乌拉尔甘草(Glycyrrhizia uralensis Fisch.)(以下简称甘草)为研究材料,上述4种微量元素,分别设置低、中和高浓度处理。本研究可为人工定向调控甘草品质,开发人工栽培甘草专用微量元素肥料提供研究依据。取得的主要研究结果如下：
1.B、Mn、Zn和Mo等4种微量元素中,对甘草酸形成与积累具有明显促进作用的是Zn、Mo和Mn等3种元素。但不同施肥方式(土壤条施和叶面喷施)间作用结果略有差异。其中叶面喷施可以显著提高甘草根中甘草酸含量的是Mn和Zn元素三种浓度处理(0.05%,0.1%和0.15%),中、高浓度Mo元素处理(0.1%和0.15%)及高浓度B元素处理；根部土壤条施对甘草根中甘草酸积累有显著促进作用的元素及相应处理浓度分别是中、高浓度的Mo元素(75g／hm2和100g／hm2)和Mn元素(6kg／hm2和9kg／hm2),低、中及高浓度的Zn元素(3.4kg／hm2、4.0kg／hm2和4.5kg／hm2)。
2.施用微量元素后甘草根中甘草酸含量以当年10月和5月含量显著高于其他月份,说明人工栽培甘草适宜的采收期应是春末夏初和秋末这两个时期。
3.能够促进甘草酸生物合成关键酶——鲨烯合成酶(SQS)基因表达的是高和极高浓度的Zn元素(0.15%和0.3%)和中、高和极高浓度的Mo元素(0.1%,0.15%和0.3%)；Zn元素所有处理浓度均可明显提高β-香树脂醇合成酶(β-AS)基因的表达,随着Mo元素处理浓度的提高,β-AS基因的表达量增加。
4.在甘草根中首次检测到甘草酸合成的中间前体物质——角鲨烯,但含量较低。建立了相应的高效液相及气相色谱质谱检测方法。4种微量元素处理对角鲨烯含量具有极显著提高作用的是高浓度的Zn元素(0.15%)和高浓度的Mo元素(0.15%)处理,此外,中、低浓度的Zn元素(0.1%和0.05%)和中浓度的Mo元素(0.1%)对甘草根中角鲨烯的积累也具有显著的促进作用。各处理下6月份角鲨烯含量极显著高于其他月份。甘草根中角鲨烯含量与甘草酸含量间存在显著正相关关系。
5.微量元素处理后对甘草根中的物质组分也产生了明显影响,其中4种微量元素高浓度(0.15%)处理显著提高了甘草根中粗纤维的含量,中、高浓度(0.1%和0.15%)B、Mn、Zn元素则显著提高了甘草根中粗蛋白的含量,粗脂肪含量在Mn元素的三种浓度(0.05%、0.1%和0.15%)处理下及中、高浓度(0.1%和0.15%)Mo元素、中、低浓度(0.1%和0.05%)Zn元素和高浓度(0.15%)B元素处理下均显著高于对照处理。
6.对甘草酸与各物质组分及其他次生代谢产物进行逐步回归分析、通经分析表明,总黄酮含量与甘草酸间直接通径系数最大为1.0467,其次为多糖,直接通径系数为0.5593,再次为角鲨烯,直接通径系数为0.4294,其后依次为粗蛋白(直接通径系数为0.4037),粗脂肪(直接通径系数为0.1745)和甘草苷(直接通径系数为-0.8097)。回归方程为Y=-1.17+0.097x1+0.072x3+1.99x4-0.47x5+0.49x6+0.019x7(其为中x1粗蛋白,x3为粗脂肪,x4为角鲨烯,x5为甘草苷,x6为总黄酮,x7为多糖)
7.根施微量元素处理后,甘草根中微量元素水平产生了明显变化,存在Zn元素和Mn元素的富集。
8.甘草根中微量元素初级形态分析表明,各元素的总提取率在1.71-60.06%之间。Zn元素的提取率最高,为60.06%,其次为Mn,Cu和Ca,同时,Zn元素的浸留比为1.682,也是所有被测元素中最大值,可以认为Zn元素是甘草中最特征的元素。次级形态分析表明甘草根中Cu、Mn、Ca、Mg 4种元素主要以游离态存在,Cu、Mn、Ca、Mg、Zn稳定态含量均大于50%,且Cu、Mn、Ca、Mg、Zn元素大部分以无机态形式存在。各元素以多种形态并存
基于上述实验结果,初步推断微量元素Zn和Mo通过提高甘草酸生物合成关键酶基因的表达最终促进了甘草酸的形成和积累。Mn元素处理对SQS和β-AS基因表达量影响不大,推测其可能是通过在甘草体内富集,影响甘草光合作用、促进同化物质运输等方式促进了甘草酸的积累。
Glycyrrhiza uralensis Fisch. is among the most important and widely used medicinal plants in traditional Chinese medicine. Wild G. uralensis germplasm resources are to be extinguished due to much over excavation and exploitation. Fortunately, artificial cultivation of G uralensis in a large scale of growing areas has been an effective alternative strategy to produce and supply the medicinal plant. However, the glycyrrhizic acid content in cultivated G. uralensis is lower than that in the wild plants. Glycyrrhizic acid is the major chemical to generate medical functions. It is believed that glycyrrhizic acid is mainly responsible for the pharmacological activities of G. uralensis. Therefore, increase of glycyrrhizic acid content in the roots of cultivated plants of G. uralensis would be economically important. The objectives of this investigation were to characterize the effects of applying four micro-elements, B, Mn, Zn, and Mo on the secondary metabolites, primary metabolites, gene of squalene synthase (SQS) andβ-amyrin synthase (β-AS) expression in cultivated G. uralensis. Artificially planted G uralensis was treated with different concentrations of B, Mn, Zn, Mo. The study should provide research-based data for improving cultivation of G. uralensis by increasing its medicinal quality. Major results are described as follows:
1. Zn, Mo and Mn had significant effects on quality of G. uralensis although the effects of micro-elements were slight different in foliar application and applying micro-element fertilizers into the soil. Glycyrrhizic acid contents with micro-element spraying treatment were increased at applying Mn and Zn of all three concentrations(0.05%,0.1% and 0.15%), Mo of medium to high concentrations(0.1% and 0.15%).
2. Early summer (May) or late fall (October) were the best harvesting months due to higher glycyrrhizic acid contents in roots of G. uralensis.
3. The expression of SQS andβ-AS gene were significantly higher with applying Zn of three concentrations(0.05%,0.1% and 0.15%) and Mo of medium to high concentrations(0.1% and 0.15%) than other treatments.
4. Squalene was detected in roots of G. uralensis, but its content was relatively low. Mo with medium to high concentrations(0.1% and 0.15%) and Zn with three concentrations(0.05%,0.1% and 0.15%) elevated the contents of squalene. Squalene content in June was the highest in a year. Significantly positive correlations were found between the contents of glycyrrhizic acid and squalene.
5. The chemical composition of the ingredients in the roots of G. uralensis was changed with four micro-elements treatments. Four micro-elements with highest concentration(0.15%) had remarkable influence on crude fiber content in roots of G. uralensis. B, Mn and Zn of medium to high concentrations(0.10% and 0.15%) increaseed the content of crude protein. Mn with three concentrations(0.05%,0.1% and 0.15%) increased crude fat, and so did Mo of medium to high concentrations(0.10% and 0.15%), Zn of low to medium concentrations(0.05% and 0.1%) and high concentration of B(0.15%).
6. The relationship between glycyrrhizic acid and other metabolites were analyzed via path correlation analysis. The sequence of direct path coefficient were total flavonoids, polysaccharides, squalene, crude protein, crude fat and liquiritin. The regression equation was: Y=-1.17+0.097x1+0.072x3+1.99x4-0.47x5+0.49x6+0.019x7, in which x1 as crude protein, x3 as crude fat, x4 as squalene, x5 as liquiritin, x6 as total flavonoids, x7 as polysaccharides.
7. Relative enrichment in Zn and Mn was found in roots of G. uralensis after B, Mn, Zn, Mo treatments.
8. The analyses of Fe, Zn, Cu, Mn, Ca, Mg in G. uralensis showed that the total extraction rate of microelement ranged from 1.71-60.06%, among them the highest was Zn, second was Mn, meanwhile the immerse-residue ratio of Zn was also the highest among the microelements, therefore, Zn could be identified as the characteristic element in G. uralensis. The analyses also indicated that the contents of Cu、Mn、Ca、Mg, and Zn in dissociative and stable forms were more than 50%, the majority of Cu、Mn、Ca、Mg and Zn were in inorganic forms. The second forms of Cu、Fe、Mn、Ca、Mg、Zn in the roots of artificially planted G. uralensis were presented in dissociative and non-dissociative, labile and stable, organic and inorganic forms at the same time.
In summary, the glycyrrhizic acid biosynthesis and accumulation were increased by enhancing gene expression of SQS andβ-AS which were two key enzymes in synthesis of glycyrrhizic acid under Zn and Mo treatments, while Mn application enriched its content in the roots and promote the assimilate transportation, consequently enhanced glycyrrhizic acid formation and accumulation.
1.B、Mn、Zn和Mo等4种微量元素中,对甘草酸形成与积累具有明显促进作用的是Zn、Mo和Mn等3种元素。但不同施肥方式(土壤条施和叶面喷施)间作用结果略有差异。其中叶面喷施可以显著提高甘草根中甘草酸含量的是Mn和Zn元素三种浓度处理(0.05%,0.1%和0.15%),中、高浓度Mo元素处理(0.1%和0.15%)及高浓度B元素处理；根部土壤条施对甘草根中甘草酸积累有显著促进作用的元素及相应处理浓度分别是中、高浓度的Mo元素(75g／hm2和100g／hm2)和Mn元素(6kg／hm2和9kg／hm2),低、中及高浓度的Zn元素(3.4kg／hm2、4.0kg／hm2和4.5kg／hm2)。
2.施用微量元素后甘草根中甘草酸含量以当年10月和5月含量显著高于其他月份,说明人工栽培甘草适宜的采收期应是春末夏初和秋末这两个时期。
3.能够促进甘草酸生物合成关键酶——鲨烯合成酶(SQS)基因表达的是高和极高浓度的Zn元素(0.15%和0.3%)和中、高和极高浓度的Mo元素(0.1%,0.15%和0.3%)；Zn元素所有处理浓度均可明显提高β-香树脂醇合成酶(β-AS)基因的表达,随着Mo元素处理浓度的提高,β-AS基因的表达量增加。
4.在甘草根中首次检测到甘草酸合成的中间前体物质——角鲨烯,但含量较低。建立了相应的高效液相及气相色谱质谱检测方法。4种微量元素处理对角鲨烯含量具有极显著提高作用的是高浓度的Zn元素(0.15%)和高浓度的Mo元素(0.15%)处理,此外,中、低浓度的Zn元素(0.1%和0.05%)和中浓度的Mo元素(0.1%)对甘草根中角鲨烯的积累也具有显著的促进作用。各处理下6月份角鲨烯含量极显著高于其他月份。甘草根中角鲨烯含量与甘草酸含量间存在显著正相关关系。
5.微量元素处理后对甘草根中的物质组分也产生了明显影响,其中4种微量元素高浓度(0.15%)处理显著提高了甘草根中粗纤维的含量,中、高浓度(0.1%和0.15%)B、Mn、Zn元素则显著提高了甘草根中粗蛋白的含量,粗脂肪含量在Mn元素的三种浓度(0.05%、0.1%和0.15%)处理下及中、高浓度(0.1%和0.15%)Mo元素、中、低浓度(0.1%和0.05%)Zn元素和高浓度(0.15%)B元素处理下均显著高于对照处理。
6.对甘草酸与各物质组分及其他次生代谢产物进行逐步回归分析、通经分析表明,总黄酮含量与甘草酸间直接通径系数最大为1.0467,其次为多糖,直接通径系数为0.5593,再次为角鲨烯,直接通径系数为0.4294,其后依次为粗蛋白(直接通径系数为0.4037),粗脂肪(直接通径系数为0.1745)和甘草苷(直接通径系数为-0.8097)。回归方程为Y=-1.17+0.097x1+0.072x3+1.99x4-0.47x5+0.49x6+0.019x7(其为中x1粗蛋白,x3为粗脂肪,x4为角鲨烯,x5为甘草苷,x6为总黄酮,x7为多糖)
7.根施微量元素处理后,甘草根中微量元素水平产生了明显变化,存在Zn元素和Mn元素的富集。
8.甘草根中微量元素初级形态分析表明,各元素的总提取率在1.71-60.06%之间。Zn元素的提取率最高,为60.06%,其次为Mn,Cu和Ca,同时,Zn元素的浸留比为1.682,也是所有被测元素中最大值,可以认为Zn元素是甘草中最特征的元素。次级形态分析表明甘草根中Cu、Mn、Ca、Mg 4种元素主要以游离态存在,Cu、Mn、Ca、Mg、Zn稳定态含量均大于50%,且Cu、Mn、Ca、Mg、Zn元素大部分以无机态形式存在。各元素以多种形态并存
基于上述实验结果,初步推断微量元素Zn和Mo通过提高甘草酸生物合成关键酶基因的表达最终促进了甘草酸的形成和积累。Mn元素处理对SQS和β-AS基因表达量影响不大,推测其可能是通过在甘草体内富集,影响甘草光合作用、促进同化物质运输等方式促进了甘草酸的积累。
Glycyrrhiza uralensis Fisch. is among the most important and widely used medicinal plants in traditional Chinese medicine. Wild G. uralensis germplasm resources are to be extinguished due to much over excavation and exploitation. Fortunately, artificial cultivation of G uralensis in a large scale of growing areas has been an effective alternative strategy to produce and supply the medicinal plant. However, the glycyrrhizic acid content in cultivated G. uralensis is lower than that in the wild plants. Glycyrrhizic acid is the major chemical to generate medical functions. It is believed that glycyrrhizic acid is mainly responsible for the pharmacological activities of G. uralensis. Therefore, increase of glycyrrhizic acid content in the roots of cultivated plants of G. uralensis would be economically important. The objectives of this investigation were to characterize the effects of applying four micro-elements, B, Mn, Zn, and Mo on the secondary metabolites, primary metabolites, gene of squalene synthase (SQS) andβ-amyrin synthase (β-AS) expression in cultivated G. uralensis. Artificially planted G uralensis was treated with different concentrations of B, Mn, Zn, Mo. The study should provide research-based data for improving cultivation of G. uralensis by increasing its medicinal quality. Major results are described as follows:
1. Zn, Mo and Mn had significant effects on quality of G. uralensis although the effects of micro-elements were slight different in foliar application and applying micro-element fertilizers into the soil. Glycyrrhizic acid contents with micro-element spraying treatment were increased at applying Mn and Zn of all three concentrations(0.05%,0.1% and 0.15%), Mo of medium to high concentrations(0.1% and 0.15%).
2. Early summer (May) or late fall (October) were the best harvesting months due to higher glycyrrhizic acid contents in roots of G. uralensis.
3. The expression of SQS andβ-AS gene were significantly higher with applying Zn of three concentrations(0.05%,0.1% and 0.15%) and Mo of medium to high concentrations(0.1% and 0.15%) than other treatments.
4. Squalene was detected in roots of G. uralensis, but its content was relatively low. Mo with medium to high concentrations(0.1% and 0.15%) and Zn with three concentrations(0.05%,0.1% and 0.15%) elevated the contents of squalene. Squalene content in June was the highest in a year. Significantly positive correlations were found between the contents of glycyrrhizic acid and squalene.
5. The chemical composition of the ingredients in the roots of G. uralensis was changed with four micro-elements treatments. Four micro-elements with highest concentration(0.15%) had remarkable influence on crude fiber content in roots of G. uralensis. B, Mn and Zn of medium to high concentrations(0.10% and 0.15%) increaseed the content of crude protein. Mn with three concentrations(0.05%,0.1% and 0.15%) increased crude fat, and so did Mo of medium to high concentrations(0.10% and 0.15%), Zn of low to medium concentrations(0.05% and 0.1%) and high concentration of B(0.15%).
6. The relationship between glycyrrhizic acid and other metabolites were analyzed via path correlation analysis. The sequence of direct path coefficient were total flavonoids, polysaccharides, squalene, crude protein, crude fat and liquiritin. The regression equation was: Y=-1.17+0.097x1+0.072x3+1.99x4-0.47x5+0.49x6+0.019x7, in which x1 as crude protein, x3 as crude fat, x4 as squalene, x5 as liquiritin, x6 as total flavonoids, x7 as polysaccharides.
7. Relative enrichment in Zn and Mn was found in roots of G. uralensis after B, Mn, Zn, Mo treatments.
8. The analyses of Fe, Zn, Cu, Mn, Ca, Mg in G. uralensis showed that the total extraction rate of microelement ranged from 1.71-60.06%, among them the highest was Zn, second was Mn, meanwhile the immerse-residue ratio of Zn was also the highest among the microelements, therefore, Zn could be identified as the characteristic element in G. uralensis. The analyses also indicated that the contents of Cu、Mn、Ca、Mg, and Zn in dissociative and stable forms were more than 50%, the majority of Cu、Mn、Ca、Mg and Zn were in inorganic forms. The second forms of Cu、Fe、Mn、Ca、Mg、Zn in the roots of artificially planted G. uralensis were presented in dissociative and non-dissociative, labile and stable, organic and inorganic forms at the same time.
In summary, the glycyrrhizic acid biosynthesis and accumulation were increased by enhancing gene expression of SQS andβ-AS which were two key enzymes in synthesis of glycyrrhizic acid under Zn and Mo treatments, while Mn application enriched its content in the roots and promote the assimilate transportation, consequently enhanced glycyrrhizic acid formation and accumulation.
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