银杏叶次生代谢产物的环境诱导机制及其调控
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摘要
本文研究了干旱胁迫、光强处理和光质处理条件下2年生银杏苗生长规律和根、茎、叶中黄酮和萜内酯含量的年动态变化,同步测定了叶中次生代谢关键酶PAL活性,采用RT-PCR方法分析黄酮和萜内酯代谢过程中14个关键酶基因表达情况,从终产物、蛋白质、核酸3个水平探讨银杏应对环境变化的过程和分子机制,主要结论如下:
1、干旱胁迫抑制银杏苗生长。干旱胁迫对叶中黄酮和萜内酯代谢影响较大,对茎和根内含量影响较小。银杏叶黄酮和萜内酯含量年动态变化呈双峰曲线,PAL酶活性与银杏叶黄酮含量变化趋势相似。中度干旱条件下叶中总黄酮醇苷含量最大,黄酮代谢途径上游PAL、C4H、CHS基因表达与黄酮含量升高的趋势相对应,下游F3H、FLS、ANS、ANR基因表达与黄酮含量同步变化。轻度干旱有利于叶和根中萜内酯积累,萜内酯代谢MVA途径中HMGR、MVD基因协同调控白果内酯BB代谢,MEP途径中DXS、DXR、MECT、MECPs基因调控总二萜内酯的合成,下游GGPPs基因表达与总萜内酯含量的变化趋势一致。
2、轻度遮阴有利于银杏生长。银杏叶黄酮和萜内酯含量年动态变化呈单峰曲线,不同光强条件下银杏叶PAL活性与黄酮含量变化并不对应。轻度遮阴条件下,叶中总黄酮醇苷含量最大,其代谢途径下游基因表达与黄酮代谢关系紧密。重度遮阴有利于叶中萜内酯积累,其MVA代谢途径中基因表达与白果内酯BB含量变化趋势完全相符,而MEP代谢途径中仅有MECT和MECPs基因表达与二萜类物质含量变化趋势基本一致,下游GGPPs基因表达与总萜内酯代谢同步变化。
3、红光处理促进银杏苗地上部分生长,而蓝光有利于地下部分生长。银杏叶黄酮和萜内酯含量年动态变化呈双峰曲线,PAL活性与黄酮含量变化并不对应。红光有利于总黄酮醇苷含量的积累,黄酮代谢下游关键酶基因表达与含量变化趋势基本相同。蓝光有利于萜内酯积累,萜内酯MVA代谢途径中关键酶基因表达与白果内酯BB含量变化趋势不同步,MEP代谢途径中关键酶基因和GGPPs基因表达与总萜内酯含量变化相一致。
This dissertation presented the study on the growth and main medicinal compositions of Ginkgo seedlings of two years under drought stress, light intensity and light quality, and the annual dynamic variations of the total flavonoids, terpene lactones in leaves, roots and stems from April to November. Meanwhile, the study also determined PAL activity of the key enzyme in secondary metabolism in leaves and analyzed the expression of 14 key enzyme genes in the metabolistic process of flavonoids, terpene lactones by RT-PCR. Then, it discessed the process of adaptation to the enviornment and its molecular mechanism from the perspectives of products, protein and nucleic acid of Gingko biloba. The main results were showed as following:
1. Drought stress inhibited the growth of Ginkgo seedlings. It exerted greater influence on the metabolism of flavonoids, terpene lactones in leaves than their contents in stems and roots. Contents of flavonoids and terpene lactones in leaves were correlated significantly with drought stress. The content variation of flavonoids and terpene lactones in leaves displayed a double peaks curve and PAL activity and flavonoids content shared a similar changing pattern. Flavonoids reached its highest level in moderate drought. In the early stage of flavonoids metabolism, expressions of PAL, C4H and CHS genes corresponded with the ascending content of flavonoids, and those of F3H、FLS、ANS、ANR genes in later stage of flavonoid metabolism were entirely the same as changes of flavonoid content. Light drought helped in the accumulation of terpene lactones in leaves and roots. In the metabolism of terpene lactones through MVA , genes HMGR and MVD jointly regulated the metabolism of bilobalide, while genes DXS, DXR, MECT and MECPs in the metabolism of terpene lactones through MEP controled synthesis of diterpene lactones, and the experession of gene GGPPs corresponded with the content of total terpene .
2. Light shade facilitated the growth of Ginkgo biloba. The variation of flavonoids and terpene lactones in the leaves showed a single-peak curve, and PAL activity and flavonoids content in the leaves didn’t correspond to each other under different light intensity. In light shade treatment, flavonoids reached its peak in content, in whose later stage of metabolism, genetic expression and flavonoids mebabolism remained close relationship; Severe shade faciliated the accumulation of terpene lactones in leaves, in the metabolistic channel of which the genetic expression of MVA was entirely corresponding with the variation of bilobalide content, while in the metabolistic channel of MEP the expression of genes MECT and MECPs basically agreed with the variation of diterpene lactones content, and the experession of gene GGPPs in later matabolistic stage varied with the metabolism of total terpene lactones.
3. Red light helped in the growth of aboveground parts, while blue light the growth of underground parts. The variation of flavonoids content and terpene lactones displayed a double-peak curve, and PAL activity didn’t correspond with the variation of flavonoids content in the leaves. Red light was in favor of the flavonoids accumulation and the genetic expression of the key enzyme of later stage of flavonoids metabolism was basically in accordance with the variation of flavonoid contents. Blue light was favorable to the accumulation of terpene lactones in leaves and roots. The genetic expression of the key enzyme in the metabolism of MVA in terpene lactones was not in accordance with the variation of bilobalide contents but that of the key genes in MEP metabolism and GGPPs conformed with the content variation of terpene lactones in total.
1、干旱胁迫抑制银杏苗生长。干旱胁迫对叶中黄酮和萜内酯代谢影响较大,对茎和根内含量影响较小。银杏叶黄酮和萜内酯含量年动态变化呈双峰曲线,PAL酶活性与银杏叶黄酮含量变化趋势相似。中度干旱条件下叶中总黄酮醇苷含量最大,黄酮代谢途径上游PAL、C4H、CHS基因表达与黄酮含量升高的趋势相对应,下游F3H、FLS、ANS、ANR基因表达与黄酮含量同步变化。轻度干旱有利于叶和根中萜内酯积累,萜内酯代谢MVA途径中HMGR、MVD基因协同调控白果内酯BB代谢,MEP途径中DXS、DXR、MECT、MECPs基因调控总二萜内酯的合成,下游GGPPs基因表达与总萜内酯含量的变化趋势一致。
2、轻度遮阴有利于银杏生长。银杏叶黄酮和萜内酯含量年动态变化呈单峰曲线,不同光强条件下银杏叶PAL活性与黄酮含量变化并不对应。轻度遮阴条件下,叶中总黄酮醇苷含量最大,其代谢途径下游基因表达与黄酮代谢关系紧密。重度遮阴有利于叶中萜内酯积累,其MVA代谢途径中基因表达与白果内酯BB含量变化趋势完全相符,而MEP代谢途径中仅有MECT和MECPs基因表达与二萜类物质含量变化趋势基本一致,下游GGPPs基因表达与总萜内酯代谢同步变化。
3、红光处理促进银杏苗地上部分生长,而蓝光有利于地下部分生长。银杏叶黄酮和萜内酯含量年动态变化呈双峰曲线,PAL活性与黄酮含量变化并不对应。红光有利于总黄酮醇苷含量的积累,黄酮代谢下游关键酶基因表达与含量变化趋势基本相同。蓝光有利于萜内酯积累,萜内酯MVA代谢途径中关键酶基因表达与白果内酯BB含量变化趋势不同步,MEP代谢途径中关键酶基因和GGPPs基因表达与总萜内酯含量变化相一致。
This dissertation presented the study on the growth and main medicinal compositions of Ginkgo seedlings of two years under drought stress, light intensity and light quality, and the annual dynamic variations of the total flavonoids, terpene lactones in leaves, roots and stems from April to November. Meanwhile, the study also determined PAL activity of the key enzyme in secondary metabolism in leaves and analyzed the expression of 14 key enzyme genes in the metabolistic process of flavonoids, terpene lactones by RT-PCR. Then, it discessed the process of adaptation to the enviornment and its molecular mechanism from the perspectives of products, protein and nucleic acid of Gingko biloba. The main results were showed as following:
1. Drought stress inhibited the growth of Ginkgo seedlings. It exerted greater influence on the metabolism of flavonoids, terpene lactones in leaves than their contents in stems and roots. Contents of flavonoids and terpene lactones in leaves were correlated significantly with drought stress. The content variation of flavonoids and terpene lactones in leaves displayed a double peaks curve and PAL activity and flavonoids content shared a similar changing pattern. Flavonoids reached its highest level in moderate drought. In the early stage of flavonoids metabolism, expressions of PAL, C4H and CHS genes corresponded with the ascending content of flavonoids, and those of F3H、FLS、ANS、ANR genes in later stage of flavonoid metabolism were entirely the same as changes of flavonoid content. Light drought helped in the accumulation of terpene lactones in leaves and roots. In the metabolism of terpene lactones through MVA , genes HMGR and MVD jointly regulated the metabolism of bilobalide, while genes DXS, DXR, MECT and MECPs in the metabolism of terpene lactones through MEP controled synthesis of diterpene lactones, and the experession of gene GGPPs corresponded with the content of total terpene .
2. Light shade facilitated the growth of Ginkgo biloba. The variation of flavonoids and terpene lactones in the leaves showed a single-peak curve, and PAL activity and flavonoids content in the leaves didn’t correspond to each other under different light intensity. In light shade treatment, flavonoids reached its peak in content, in whose later stage of metabolism, genetic expression and flavonoids mebabolism remained close relationship; Severe shade faciliated the accumulation of terpene lactones in leaves, in the metabolistic channel of which the genetic expression of MVA was entirely corresponding with the variation of bilobalide content, while in the metabolistic channel of MEP the expression of genes MECT and MECPs basically agreed with the variation of diterpene lactones content, and the experession of gene GGPPs in later matabolistic stage varied with the metabolism of total terpene lactones.
3. Red light helped in the growth of aboveground parts, while blue light the growth of underground parts. The variation of flavonoids content and terpene lactones displayed a double-peak curve, and PAL activity didn’t correspond with the variation of flavonoids content in the leaves. Red light was in favor of the flavonoids accumulation and the genetic expression of the key enzyme of later stage of flavonoids metabolism was basically in accordance with the variation of flavonoid contents. Blue light was favorable to the accumulation of terpene lactones in leaves and roots. The genetic expression of the key enzyme in the metabolism of MVA in terpene lactones was not in accordance with the variation of bilobalide contents but that of the key genes in MEP metabolism and GGPPs conformed with the content variation of terpene lactones in total.
引文
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