甘草品种选育基础及HMGR基因克隆研究
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摘要
甘草中甘草酸等药用活性成分含量是影响其药材质量的主要因素。因此,进行高甘草酸等高药用活性成分含量品种的选育,建立良种繁育体系成为获得优质栽培甘草的重要途径。本论文以甘草优良品种选育需要开展的基础研究工作为主线,以内蒙古杭锦旗栽培甘草为主要试验材料,通过形态学观察,石蜡切片制作和显微镜观察,主要药用活性成分含量测定,以及基因克隆等方法,对甘草的有性生殖特性、种质资源筛选和甘草酸代谢途径中关键酶HMGR基因的克隆和表达特征三部分内容进行了研究。主要研究结果如下。
1.观察和分析了甘草花部形态特征、单花发育过程、单花开花动态和花序花数、结荚花序果实数及单荚种子数的数量特征。在甘草群体中,发现了花旗瓣呈紫色花萼深紫色和花旗瓣呈白色花萼颜色接近全绿的两种花色的变异,本文将这两种变异定义为深紫花和白花变异类型。对甘草的开花习性研究发现,一日内,以10:00-16:00开花最多,占开花总量的38.9%;下午20:00至第二天早晨6:00次之,占开花总量的35.3%。经过多种方法比较研究发现甘草为白花授粉中闭花受精的植物。
2.筛选了适合甘草花粉生活力和柱头活性检测的方法,利用田间授粉法研究了甘草的花粉生活力和柱头活性。结果表明,一天当中在12:00时花粉的生活力最高;对柱头活性的检测结果表明,在去雄后第5天柱头已经没有活性。对甘草的小孢子发生及雄配子体发育过程观察发现,可以通过花蕾的外部形态来推测其雄蕊的发育程度;观察到了甘草的大孢子发生及雌配子体发育过程;在对甘草去雄后不同时间授粉材料的荧光显微镜的观察中发现授粉后6h花粉管进入胚珠受精。
3.对甘草的形态结构变异和药用活性成分含量的关系进行了研究,提出可以用一些形态指标进行选种。
(1)对甘草的外部形态与甘草苷和甘草酸含量的相关性分析表明,四年生移栽甘草中顶叶宽和小叶宽与甘草酸和甘草苷含量具有显著的相关性或相关趋势;对茎皮颜色和茎附属物(刺毛多少)赋值以后与甘草酸和甘草苷含量的相关分析表明,在一定程度上,茎皮颜色越深,茎皮刺毛数越多,甘草酸含量越低。对一年生直播甘草的分析表明,主根长度与甘草苷含量达到显著正相关水平,根重与甘草酸和甘草苷含量均呈一定程度的正相关;甘草苷含量与甘草酸含量呈极显著正相关关系,说明甘草酸和甘草苷的积累速度具有协同性。
(2)通过对深紫花变异类型、白花变异类型和普通类型的甘草酸、甘草苷和异甘草素含量的分析和比较,发现白花类型的药材质量较好。
(3)不同种质甘草叶片的外部形态和内部结构存在差异。考虑黄酮苷和黄酮苷元等次生代谢物质可能跟叶片栅栏组织中的分泌结构相关,对不同种质甘草的这些结构进行了计数,从利用地上部作为黄酮生产原料的资源开发角度考虑,可以利用甘草叶片内部结构中的分泌组织数量推测不同甘草种质的黄酮含量。
4.通过对栽培在内蒙古杭锦旗的四个种源材料的主要药效成分含量进行比较,发现46号种源的种质较为优良。通过对两个种源的萌发期和幼苗期生理特性和一年生根系的药效成分含量进行比较,说明宁夏高沙窝种源为比较优良的种质。
5.根据同源基因的保守区设计引物,结合基因组行走技术,得到甘草HMGR基因,其长度为2662bp,本研究将其命名为GuHMGR1。GuHMGR1基因在DNA水平上由3个外显子和2个内含子组成;GuHMGR1的cDNA全长为2330bp,其中包括1518bp的开放阅读框、311bp的5‘非翻译区和501bp的3‘非翻译区。该基因编码区共翻译成505个氨基酸。保守结构域分析发现,GuHMGR1在结构上归属于HMGR-CoA还原酶Ⅰ类家族。
6.对GuHMGR1在甘草不同生长时期器官中的表达特性结果表明,GuHMGR1在被检测不同时期的根、叶中都有表达,但不同样本中的表达水平不同。表现为根部的表达水平高于叶部,主根和老叶中的表达水平相应高于须根和幼叶。对筛选出的白花和深紫花类型的两个甘草酸含量不同植株的GuHMGR1基因特征分析结果表明,两者在GuHMGR1基因组部分DNA序列上存在碱基差异。
The medicinal composition content,such as glycyrrhizin,in liquorice is the limited factor of medicinal material quality.So,to obtain superior quality cultivation liquorice,it is necessary to perform the high medicinal composition content breeding including glycyrrhizin and establish the good variety propagation system.Therefore,a study was performed using the base of good variety breeding of liquorice as the main line,the Hangjinqi liquorice from inner Mongolia as the major research materials.The study was conducted by morphology observation,paraffin section making and microscopic observation,the content determination of major medicinal composition and gene cloning to study the three parts of contents,including of sexual reproduction properties,germplasm resources evaluation and the gene clone and expression characteristics of the HMGR,which is the key enzyme in the metabolic pathway of liquorice.The main results were as follows:
1.Observing and analyzing the floral characteristics,the development process of a single floral,flowering dynamics of a single floral and the quantitative characteristics of inflorescence and pod seeds.The variation of flower color in liquorice colony was found,this text named them as modena type and white type.The flowering rate was 38.9%from 10:00 to 16:00,and it was 35.3%from 20:00 to 6:00 the next day.The study determined finally the breeding system of liquorice,which was cleistogamy by various ways,simultaneity,the stigma could accept other pollens such as the pollen which was given by artificial pollination.
2.The study filtered the detection method of the pollen viability and the stigma activity.Finally,the experiment of hand-pollination in the field was adopted.The pollen viability was the highest at 12:00 in daytime,the stigma had no activity in the fifth day after artificial emasculation.The results about microsporogenesis and male gametophyte development showed that it would estimate the development degree of stamen according to the external morphology of the flower buds.The development degree of pistil before fertilization were observed in the study.The fluorescence microscope observation results about the pollinated material in the different time after emasculation showed that the pollen tube entered into the ovule to fertilize 6h after pollination.
3.The results of correlation analysis between agronomic characteristics and the content of glycyrrhizin and liquiritin content showed that the relationship of parietal and the leaflets width and the glycyrrhizin and liquiritin content was significant or had the trend in the four years transplanted liquorice,the analysis result between stem bark color and stem appendants(chaeta numbers) with the glycyrrhizin content showed that the stem bark color was deeper and the chaeta number was much more, the glycyrrhizin content was higher to some extent.The analysis result about the one year planting liquorice showed that the correlation of tap root length with liquiritin content was positive significantly,the correlation of root weight and the glycyrrhizin and liquiritin content was positive to some extent.The correlation of glycyrrhizin and liquiritin was positive significantly,which indicated that the accumulation rate of two medicinal composition content had a certain coordination.
4.Analyzing and comparing the glycyrrhizin,liquiritin and isoliquiritigenin content among the modena,white flower type and commen flower color type,the white type was the selected better type.The Gaoshawo(from Gansu province) provenience showed a better growth status in plain region by comparing with the Hangjinqi(from inner Mongolia) provenience of the seedling physiological characterization and the medicinal composition content of the annual root.Comparing the major medicinal composition content among four proveniences in Hangjinqi(from inner Mongolia),the provenience 46 was the better breeding material.
5.The anatomical structure of leaves of liquorice was observed to reveal the drought-resistant mechanism of Glycyrrhiza L.A selected way of high medicinal composition content was put forward which was the quantities of secretory tissues in the internal structure of the leaves.Considering the aerial parts as the flavonoids production materials,the selected order was G.glabra L.> G.inflate Batat.> G. uralensis Fisch.among the Glycyrrhiza L.,while,the selected order among the G. uralensis Fisch.was provenience 41> provenience 14> provenience 46.
5.A full length DNA encoding HMGR(designated as GuHMGR1) was isolated from liquorice(G.uralensis Fisch.) based on degenerated PCR and genome walking approach.The comparison between the GuHMGR1 genomic sequence with the cDNA of Pisum sativum showed that the genomic DNA contained two introns.The lengths of three exons were 1206 bp,181 bp and 943 bp,respectively.The lengths of the two introns were 87 bp and 245 bp,respectively.The cDNA of GuHMGR1 was 2330 bp in length and contained a 1518 bp ORF,311bp 5' untranslated region and 501bp 3' untranslated region,encoding a protein of 505 amino acids.The analyzed result of conserved domain of GuHMGR1 found that the GuHMGR1 was ascribed to the I family of reductase super family in structure.
6.The result of GuHMGR1 expression pattern in different tissues showed that GuHMGR1 was expressed in different growth period of roots and leaves,but the expression level was different in the different transcripts,the expression level in the roots was higher than in the leaves,the expression level in the taproots and old leaves was higher than in the lateral roots and new leaves.The results of GuHMGR1 characteristics in the liquorice with different glycyrrhizin content,one belonged to white flower type,and the other belonged to modena flower type,showed that the two type had base differences in the part DNA sequence.
1.观察和分析了甘草花部形态特征、单花发育过程、单花开花动态和花序花数、结荚花序果实数及单荚种子数的数量特征。在甘草群体中,发现了花旗瓣呈紫色花萼深紫色和花旗瓣呈白色花萼颜色接近全绿的两种花色的变异,本文将这两种变异定义为深紫花和白花变异类型。对甘草的开花习性研究发现,一日内,以10:00-16:00开花最多,占开花总量的38.9%;下午20:00至第二天早晨6:00次之,占开花总量的35.3%。经过多种方法比较研究发现甘草为白花授粉中闭花受精的植物。
2.筛选了适合甘草花粉生活力和柱头活性检测的方法,利用田间授粉法研究了甘草的花粉生活力和柱头活性。结果表明,一天当中在12:00时花粉的生活力最高;对柱头活性的检测结果表明,在去雄后第5天柱头已经没有活性。对甘草的小孢子发生及雄配子体发育过程观察发现,可以通过花蕾的外部形态来推测其雄蕊的发育程度;观察到了甘草的大孢子发生及雌配子体发育过程;在对甘草去雄后不同时间授粉材料的荧光显微镜的观察中发现授粉后6h花粉管进入胚珠受精。
3.对甘草的形态结构变异和药用活性成分含量的关系进行了研究,提出可以用一些形态指标进行选种。
(1)对甘草的外部形态与甘草苷和甘草酸含量的相关性分析表明,四年生移栽甘草中顶叶宽和小叶宽与甘草酸和甘草苷含量具有显著的相关性或相关趋势;对茎皮颜色和茎附属物(刺毛多少)赋值以后与甘草酸和甘草苷含量的相关分析表明,在一定程度上,茎皮颜色越深,茎皮刺毛数越多,甘草酸含量越低。对一年生直播甘草的分析表明,主根长度与甘草苷含量达到显著正相关水平,根重与甘草酸和甘草苷含量均呈一定程度的正相关;甘草苷含量与甘草酸含量呈极显著正相关关系,说明甘草酸和甘草苷的积累速度具有协同性。
(2)通过对深紫花变异类型、白花变异类型和普通类型的甘草酸、甘草苷和异甘草素含量的分析和比较,发现白花类型的药材质量较好。
(3)不同种质甘草叶片的外部形态和内部结构存在差异。考虑黄酮苷和黄酮苷元等次生代谢物质可能跟叶片栅栏组织中的分泌结构相关,对不同种质甘草的这些结构进行了计数,从利用地上部作为黄酮生产原料的资源开发角度考虑,可以利用甘草叶片内部结构中的分泌组织数量推测不同甘草种质的黄酮含量。
4.通过对栽培在内蒙古杭锦旗的四个种源材料的主要药效成分含量进行比较,发现46号种源的种质较为优良。通过对两个种源的萌发期和幼苗期生理特性和一年生根系的药效成分含量进行比较,说明宁夏高沙窝种源为比较优良的种质。
5.根据同源基因的保守区设计引物,结合基因组行走技术,得到甘草HMGR基因,其长度为2662bp,本研究将其命名为GuHMGR1。GuHMGR1基因在DNA水平上由3个外显子和2个内含子组成;GuHMGR1的cDNA全长为2330bp,其中包括1518bp的开放阅读框、311bp的5‘非翻译区和501bp的3‘非翻译区。该基因编码区共翻译成505个氨基酸。保守结构域分析发现,GuHMGR1在结构上归属于HMGR-CoA还原酶Ⅰ类家族。
6.对GuHMGR1在甘草不同生长时期器官中的表达特性结果表明,GuHMGR1在被检测不同时期的根、叶中都有表达,但不同样本中的表达水平不同。表现为根部的表达水平高于叶部,主根和老叶中的表达水平相应高于须根和幼叶。对筛选出的白花和深紫花类型的两个甘草酸含量不同植株的GuHMGR1基因特征分析结果表明,两者在GuHMGR1基因组部分DNA序列上存在碱基差异。
The medicinal composition content,such as glycyrrhizin,in liquorice is the limited factor of medicinal material quality.So,to obtain superior quality cultivation liquorice,it is necessary to perform the high medicinal composition content breeding including glycyrrhizin and establish the good variety propagation system.Therefore,a study was performed using the base of good variety breeding of liquorice as the main line,the Hangjinqi liquorice from inner Mongolia as the major research materials.The study was conducted by morphology observation,paraffin section making and microscopic observation,the content determination of major medicinal composition and gene cloning to study the three parts of contents,including of sexual reproduction properties,germplasm resources evaluation and the gene clone and expression characteristics of the HMGR,which is the key enzyme in the metabolic pathway of liquorice.The main results were as follows:
1.Observing and analyzing the floral characteristics,the development process of a single floral,flowering dynamics of a single floral and the quantitative characteristics of inflorescence and pod seeds.The variation of flower color in liquorice colony was found,this text named them as modena type and white type.The flowering rate was 38.9%from 10:00 to 16:00,and it was 35.3%from 20:00 to 6:00 the next day.The study determined finally the breeding system of liquorice,which was cleistogamy by various ways,simultaneity,the stigma could accept other pollens such as the pollen which was given by artificial pollination.
2.The study filtered the detection method of the pollen viability and the stigma activity.Finally,the experiment of hand-pollination in the field was adopted.The pollen viability was the highest at 12:00 in daytime,the stigma had no activity in the fifth day after artificial emasculation.The results about microsporogenesis and male gametophyte development showed that it would estimate the development degree of stamen according to the external morphology of the flower buds.The development degree of pistil before fertilization were observed in the study.The fluorescence microscope observation results about the pollinated material in the different time after emasculation showed that the pollen tube entered into the ovule to fertilize 6h after pollination.
3.The results of correlation analysis between agronomic characteristics and the content of glycyrrhizin and liquiritin content showed that the relationship of parietal and the leaflets width and the glycyrrhizin and liquiritin content was significant or had the trend in the four years transplanted liquorice,the analysis result between stem bark color and stem appendants(chaeta numbers) with the glycyrrhizin content showed that the stem bark color was deeper and the chaeta number was much more, the glycyrrhizin content was higher to some extent.The analysis result about the one year planting liquorice showed that the correlation of tap root length with liquiritin content was positive significantly,the correlation of root weight and the glycyrrhizin and liquiritin content was positive to some extent.The correlation of glycyrrhizin and liquiritin was positive significantly,which indicated that the accumulation rate of two medicinal composition content had a certain coordination.
4.Analyzing and comparing the glycyrrhizin,liquiritin and isoliquiritigenin content among the modena,white flower type and commen flower color type,the white type was the selected better type.The Gaoshawo(from Gansu province) provenience showed a better growth status in plain region by comparing with the Hangjinqi(from inner Mongolia) provenience of the seedling physiological characterization and the medicinal composition content of the annual root.Comparing the major medicinal composition content among four proveniences in Hangjinqi(from inner Mongolia),the provenience 46 was the better breeding material.
5.The anatomical structure of leaves of liquorice was observed to reveal the drought-resistant mechanism of Glycyrrhiza L.A selected way of high medicinal composition content was put forward which was the quantities of secretory tissues in the internal structure of the leaves.Considering the aerial parts as the flavonoids production materials,the selected order was G.glabra L.> G.inflate Batat.> G. uralensis Fisch.among the Glycyrrhiza L.,while,the selected order among the G. uralensis Fisch.was provenience 41> provenience 14> provenience 46.
5.A full length DNA encoding HMGR(designated as GuHMGR1) was isolated from liquorice(G.uralensis Fisch.) based on degenerated PCR and genome walking approach.The comparison between the GuHMGR1 genomic sequence with the cDNA of Pisum sativum showed that the genomic DNA contained two introns.The lengths of three exons were 1206 bp,181 bp and 943 bp,respectively.The lengths of the two introns were 87 bp and 245 bp,respectively.The cDNA of GuHMGR1 was 2330 bp in length and contained a 1518 bp ORF,311bp 5' untranslated region and 501bp 3' untranslated region,encoding a protein of 505 amino acids.The analyzed result of conserved domain of GuHMGR1 found that the GuHMGR1 was ascribed to the I family of reductase super family in structure.
6.The result of GuHMGR1 expression pattern in different tissues showed that GuHMGR1 was expressed in different growth period of roots and leaves,but the expression level was different in the different transcripts,the expression level in the roots was higher than in the leaves,the expression level in the taproots and old leaves was higher than in the lateral roots and new leaves.The results of GuHMGR1 characteristics in the liquorice with different glycyrrhizin content,one belonged to white flower type,and the other belonged to modena flower type,showed that the two type had base differences in the part DNA sequence.
引文
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