三萜人参皂苷对人参、西洋参等植物生长发育的效应研究
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摘要
人参Panax ginseng C.A.Mey.和西洋参Panax quinquefolium L是我国名贵的中药材,二者均为五加科多年生宿根性草本植物。人参、西洋参忌地性都极强,重茬栽参会发生烧须、须根脱落、植株多病及存苗率低等现象,严重影响了经济效益。人参和西洋参连作障碍的形成主要是由于土壤退化、病原菌增多及化感自毒作用。三萜皂苷是人参、西洋参主要药效成分之一,积累于参根的韧皮部和周皮中以及须根中,现已分离鉴定出的单体皂苷超过40种。三萜皂苷是人参、西洋参根系分泌物中的特有物质,是个多组分的代谢产物,具有重要的生理生态功能。人参皂苷作为一种化学防御物质,对人参和西洋参的种子萌发、愈伤组织的形成以及某些土传病原菌繁殖都具有调控作用。但关于其对人参等植物生长发育的影响以及单体皂苷的化感活性尚未见报道。本实验采用HPLC检测了不同地点、不同采样时间2-5年生参地土壤中人参皂苷的含量；采用人参幼苗为受体,探讨低中高(25,50和100mg·L-1)三种浓度人参总皂苷及其分离出的3种单体Re、Rg1和Rb1对人参早期生长的化感调控作用；以水为溶剂从4年生的西洋参中提取人参皂苷,测定了其不同浓度处理后西洋参幼苗的生理变化；考察了人参皂苷对白菜、黄瓜、小麦及绿豆4种常见作物主根及不定根发育的影响；研究了在自然土壤状态下,不同浓度(0.125,0.25和0.5g·kg-1)的人参皂苷对人参幼苗生长发育的化感效应。主要研究结论如下：
1.不同采集时间、采集地点和生长年限不同,参地土壤中总皂苷含量明显不同。随着生长年限的延长,土壤中人参皂苷含量不断增加,林地和农田栽培的多年生人参土壤中,6月采集的样品中皂苷含量远大于8月份；而西洋参根际土壤中8月采集的样品中皂苷含量远大于6月份。参地土壤中检测到的皂苷单体有Rb1、Rb2、Rb3、Re、Rd和Rg1含量呈不规律变化。不同生长年限的林地人参干土样中总皂苷的含量介于0.031-0.332mg·g-1(平均为0.0795mg·g-1),林地西洋参干土样中总皂苷的含量约在0.025-0.154mg·g-1(平均为0.0732mg·g-1),而平地人参干土样中总皂苷的含量介于0.002-0.076mg·g-1(平均为0.0213mg·g-1)。
2.高浓度的TGS和单体Rb1处理对人参的幼苗和幼根的生长发育表现出明显的抑制作用,而单体Rg1和Re却对人参幼苗的各项生长指标总体表现为微弱的促进作用。人参皂苷抑制了人参幼苗的根系活力,抑制效应依次为：Rb1>TGS>Rg1>Re,其中TGS和Rb1处理组人参根系活力的变化趋势相近,均表现为低浓度根系活力升高而高浓度下降；低浓度Rg1处理组根系活力较低,但随着浓度的增加根系活力急剧上升而后开始下降；而Re处理组的根系活力则随着浓度的升高而缓慢增高。低浓度皂苷处理后,4种化合物对叶绿素的合成均有促进作用；尽管幼苗叶绿素合成的趋势随着处理浓度的升高而降低,但各处理的叶绿素含量仍然都高于对照。透射电镜观察到人参幼根细胞在人参皂苷的诱导培养下,发生了质壁分离和细胞器降解,高浓度TGS处理的根尖细胞质壁分离严重,液泡膜分解,其它细胞器的结构也变得模糊。100mg·L-1Rb1处理后,部分液泡分解,细胞核扭曲变形,核膜模糊不清。人参根尖在高浓度的人参皂苷处理后因细胞功能受损而不能完成正常的生理活动。
3.以水为溶剂从西洋参根中提取人参皂苷对西洋参幼根伸长表现为典型的“低促高抑”浓度效应；对幼根的鲜质量及干质量均无明显影响；中高浓度均对根系活力有明显抑制效应。随着皂苷浓度和处理时间的延长,超氧阴离子自由基(O2-)的产生速率不断加快。对西洋参叶片中可溶性蛋白和叶绿素的合成有轻微的促进作用,可溶性蛋白含量在高浓度处理达48h时降到最低,叶绿素含量随着处理时间的延长而呈逐渐下降趋势,但仍高于对照。叶片丙二醛(MDA)的含量和相对电导率均随着皂苷浓度的增加和处理时间的延长而缓慢升高。高浓度人参皂苷处理48h后,人参根尖细胞核膜膨胀、核仁变形,液泡膜分解,线粒体结构不完整。不同浓度人参皂苷对西洋参早期生长发育具有化感效应,且对地下部分的影响大于地上部分。
4.人参皂苷处理液对4种常见作物的主根及不定根的发育影响明显不同。随着处理质量浓度的升高,小麦、白菜、黄瓜主根根系活力及根长、根鲜质量均呈下降趋势。而各浓度人参皂苷处理对黄瓜和绿豆下胚轴不定根的数量、根长、根鲜质量及根系活力的影响均未达到显著水平,但二者抗氧化酶的活性都微有升高。人参皂苷对4种栽培作物的主根发育均有一定抑制作用,但对小麦和黄瓜主根生长的抑制作用更强,而对黄瓜和绿豆不定根发育的影响却不明显。
5.人参皂苷与新林地土混合盆栽发现,人参幼苗地上部分的苗鲜质量无明显变化,但幼根的伸长受到严重抑制,根系活力显著下降,高浓度处理根长比对照降低7.0%,根系活力降低了69.3%,根鲜质量降低了45.5%。蛋白质合成没有受到明显,叶绿素的合成却有所增加,虽然在高浓度处理有下降的趋势但仍然高于对照组。高浓度处理使幼根的颜色变为黄褐色；内源激素IAA含量随浓度变化呈先升高后降低趋势,各处理GA3含量均明显降低,而ABA含量在低浓度处理变化不明显,但在高浓度处理后含量迅速增加到对照的2.39倍。
因此得出结论：人参三萜皂苷及其三种单体都明显干扰了人参、西洋参的生长及发育,而且这种干扰作用还与三萜皂苷的种类及浓度密切相关。对其它常见几种农作物早期根系的形成有一定的影响,但不如对人参和西洋参的化感效应显著。人参皂苷可能在自身及其它植物的生长发育过程中作为一种化学手段参与了群体形成与稳定的调控。
P. ginseng and P. quinquefolium, all belonging to Araliaceae family, are widely cultivated in China, and used in traditional herbal medicine. However, they cannot be planted consecutively on the same land otherwise their yield and quality will be greatly reduced. The obstacle of re-plantation has been considered to be caused by the soil sickness, pathogenic fungal accumulation, allellopathy and autotoxicity. Ginsenoside is the major pharmaceutically-active component in ginseng and American ginseng, which can be found in their root exudates and old ginseng soils. However, the allelopathic activity of ginsenoside and its monomers and their influences on the recipient plant are less studied. Our studies aim:(i) to detect ginsenoside content in the ginseng of different growth-year and in different collection time and plot via HPLC method;(ii) to understand the allelopathic effects of the total ginsenoside (TGS) and its monomer fractions Rb1, Re and Rg1on ginseng seedling growth and development at different concentration(25mg·L-1,50mg·L-1and100mg·L-1);(iii) to examine the reaction of American ginsengs treated with ginsenoside of different concentration, extracted from4-year American ginseng with water;(iv) to study the influence of ginsenoside on the roots and adventitious roots of four cultivars-cabbage, cucumber, wheat, and mung bean;(v) to test the allelopathic effects of ginsenoside on ginseng seedling growth when they were planted in natural soils mixed with0.125g·kg-1,0.25g·kg-1and0.5g·kg-1TGS. The key conclusions are showed as below:
1. The TGS content of different growth-year ginsengs and ginsengs in different collection time and plot is distinctively different. The TGS contents extracted from the dried soils increased constantly as growth year prolonged. In both understory and farmland ginseng soils, the TGS content in June were higher than that in August. On the contrary, the TGS content in American ginseng soils in August were higher than that in June, and TGS content increased with growth year increasing. Rb1, Rb2, Rb3, Re, Rd and Rg1were detected in the ginseng soils of3-5year-old. The TGS content in the dried ginseng soils was approximately from0.031to0.332mg·g-1(average:0.0795mg·g-1); the TGS content in American ginseng soils was from0.025to0.154mg·g-1(average:0.0732mg·g-1), whereas the TGS contents in farmland soil were0.002-0.076mg·g-1(average:0.0213mg·g-1).
2. The four compounds all affected the root vigor (RV) and root length (RL) of ginseng seedling in different ways. The TGS and the Rb1treatments both showed a dose-dependent effect on RV; The RV enhanced drastically as the TGS and Rb1concentration increased, and then when their concentration moved to some certain level which is relatively high the RV began to decline. When the Rgl concentration was low, the RV was inhibited. And as the Rgl concentration went up, the RV increased as well. But just like the TGS and the Rbl, the RV then turned down quickly. As Re concentration increased, the RV enhanced slowly but constantly. For those ginsengs that were treated with ginsenoside at lower concentration (25mg·L-1), the chlorophyll synthesis was promoted. The promotion effect of the four compounds was in order:TGS>Rg1>Re>Rb1, and even if the promoting effect grew weaker as concentration increased, the chlorophyll content was still higher than that of the control. Electron microscope showed that the plasma membranes were detached from the cell wall, the structures of mitochondria and vacuolus were also dissolved under higher TGS and Rbl concentration. Hence, these abnormal physiological parameters in plant growth process indicated the responsibility of ginsenoside concentration.
3. The ginsenoside extracted from American ginseng with the deionzed water showed a dose-dependent effect on the root elongation. However there was no significant difference between the biomass of seedlings and roots before and after. The root vigor was inhibited significantly as Ginsenoside concentration increased. The production of O2-in seedlings sped up markedly as ginsenoside concentration increased and exposure time prolonged. The synthesis of soluble protein and chlorophyll in the seedlings was slightly stimulated by the ginsenoside. The promoting effect declined as the concentration increased and the treatment time prolonged, but the content is still higher than that of the control. Otherwise, the MDA content and the relative electric conductivity in leaves and roots increased slightly compared with the control as the concentration increased and the application time prolonged. The treated ultra-structure of root-tip cells showed that the nuclear membrane and nucleolus deformed and the vacuole membrane dissolved with the treatment time prolonging. The ginsenoside affected the underground growth of American ginseng more than aboveground growth. Taken together, these findings indicated that ginsenoside affected the early growth of American ginseng.
4. The alleopathic effect of TGS on the root growth of four cultivar species was conspicuously different.The RV, RL and RFW (root fresh weight) of wheat, cabbage and cucumber declined constantly as ginsenoside concentration ascended. However, the ginsenoside treatment had almost no influence on the hypocotyl adventitious roots of cucumber and mung bean. The RL, RFW, RV and Root number of mung bean adventitious roots had no significant response to ginsenoside, although the antioxidant enzymes activity rose a little bit over the control. Thereby, ginsenoside inhibited the root growth of the four indicator plants, and had stronger inhibitory effects on wheat and cucumber than on cabbage. However, there was no evidence to indicate that ginsenoside affected the hypocotyl cuttings growth in cucumber and mung bean.
5. The result of the experiment done with the soils from the new understory land indicated that there was no significant change in the biomass of the ginseng seedlings treated with ginsenoside, whereas the root elongation was reduced significantly by7.0%under higher concentration, the RV was declined by69.3%, and the fresh weight of roots slid by45.5%. The soluble protein content had little changes, yet the chlorophyll content had been in an uptrend. Even though the chlorophyll content decreased under higher concentration, it was still higher than that of the control. The color of ginseng roots changed to brown when TGS reached the highest. The IAA content first increased and then decreased as concentration ascended; the GA3content in all the treatments declined; the ABA content had no significant difference at lower concentration, but it increased sharply to2.39times than of the control when treated with high ginsenoside concentratrion.
Hence, we concluded that triterpenoid ginsenoside and their three monomers all disturbed the growth and development of ginseng and American ginseng, and this disturbance was affected by the types and the concentrations of ginsenosides. The ginsenoside do had allelopathic effect on other plants, yet it was lower than that on ginsengs and American ginsengs. It was found that triterpenoid ginsenoside may participate in the regulation of the formation and stability of plant population.
1.不同采集时间、采集地点和生长年限不同,参地土壤中总皂苷含量明显不同。随着生长年限的延长,土壤中人参皂苷含量不断增加,林地和农田栽培的多年生人参土壤中,6月采集的样品中皂苷含量远大于8月份；而西洋参根际土壤中8月采集的样品中皂苷含量远大于6月份。参地土壤中检测到的皂苷单体有Rb1、Rb2、Rb3、Re、Rd和Rg1含量呈不规律变化。不同生长年限的林地人参干土样中总皂苷的含量介于0.031-0.332mg·g-1(平均为0.0795mg·g-1),林地西洋参干土样中总皂苷的含量约在0.025-0.154mg·g-1(平均为0.0732mg·g-1),而平地人参干土样中总皂苷的含量介于0.002-0.076mg·g-1(平均为0.0213mg·g-1)。
2.高浓度的TGS和单体Rb1处理对人参的幼苗和幼根的生长发育表现出明显的抑制作用,而单体Rg1和Re却对人参幼苗的各项生长指标总体表现为微弱的促进作用。人参皂苷抑制了人参幼苗的根系活力,抑制效应依次为：Rb1>TGS>Rg1>Re,其中TGS和Rb1处理组人参根系活力的变化趋势相近,均表现为低浓度根系活力升高而高浓度下降；低浓度Rg1处理组根系活力较低,但随着浓度的增加根系活力急剧上升而后开始下降；而Re处理组的根系活力则随着浓度的升高而缓慢增高。低浓度皂苷处理后,4种化合物对叶绿素的合成均有促进作用；尽管幼苗叶绿素合成的趋势随着处理浓度的升高而降低,但各处理的叶绿素含量仍然都高于对照。透射电镜观察到人参幼根细胞在人参皂苷的诱导培养下,发生了质壁分离和细胞器降解,高浓度TGS处理的根尖细胞质壁分离严重,液泡膜分解,其它细胞器的结构也变得模糊。100mg·L-1Rb1处理后,部分液泡分解,细胞核扭曲变形,核膜模糊不清。人参根尖在高浓度的人参皂苷处理后因细胞功能受损而不能完成正常的生理活动。
3.以水为溶剂从西洋参根中提取人参皂苷对西洋参幼根伸长表现为典型的“低促高抑”浓度效应；对幼根的鲜质量及干质量均无明显影响；中高浓度均对根系活力有明显抑制效应。随着皂苷浓度和处理时间的延长,超氧阴离子自由基(O2-)的产生速率不断加快。对西洋参叶片中可溶性蛋白和叶绿素的合成有轻微的促进作用,可溶性蛋白含量在高浓度处理达48h时降到最低,叶绿素含量随着处理时间的延长而呈逐渐下降趋势,但仍高于对照。叶片丙二醛(MDA)的含量和相对电导率均随着皂苷浓度的增加和处理时间的延长而缓慢升高。高浓度人参皂苷处理48h后,人参根尖细胞核膜膨胀、核仁变形,液泡膜分解,线粒体结构不完整。不同浓度人参皂苷对西洋参早期生长发育具有化感效应,且对地下部分的影响大于地上部分。
4.人参皂苷处理液对4种常见作物的主根及不定根的发育影响明显不同。随着处理质量浓度的升高,小麦、白菜、黄瓜主根根系活力及根长、根鲜质量均呈下降趋势。而各浓度人参皂苷处理对黄瓜和绿豆下胚轴不定根的数量、根长、根鲜质量及根系活力的影响均未达到显著水平,但二者抗氧化酶的活性都微有升高。人参皂苷对4种栽培作物的主根发育均有一定抑制作用,但对小麦和黄瓜主根生长的抑制作用更强,而对黄瓜和绿豆不定根发育的影响却不明显。
5.人参皂苷与新林地土混合盆栽发现,人参幼苗地上部分的苗鲜质量无明显变化,但幼根的伸长受到严重抑制,根系活力显著下降,高浓度处理根长比对照降低7.0%,根系活力降低了69.3%,根鲜质量降低了45.5%。蛋白质合成没有受到明显,叶绿素的合成却有所增加,虽然在高浓度处理有下降的趋势但仍然高于对照组。高浓度处理使幼根的颜色变为黄褐色；内源激素IAA含量随浓度变化呈先升高后降低趋势,各处理GA3含量均明显降低,而ABA含量在低浓度处理变化不明显,但在高浓度处理后含量迅速增加到对照的2.39倍。
因此得出结论：人参三萜皂苷及其三种单体都明显干扰了人参、西洋参的生长及发育,而且这种干扰作用还与三萜皂苷的种类及浓度密切相关。对其它常见几种农作物早期根系的形成有一定的影响,但不如对人参和西洋参的化感效应显著。人参皂苷可能在自身及其它植物的生长发育过程中作为一种化学手段参与了群体形成与稳定的调控。
P. ginseng and P. quinquefolium, all belonging to Araliaceae family, are widely cultivated in China, and used in traditional herbal medicine. However, they cannot be planted consecutively on the same land otherwise their yield and quality will be greatly reduced. The obstacle of re-plantation has been considered to be caused by the soil sickness, pathogenic fungal accumulation, allellopathy and autotoxicity. Ginsenoside is the major pharmaceutically-active component in ginseng and American ginseng, which can be found in their root exudates and old ginseng soils. However, the allelopathic activity of ginsenoside and its monomers and their influences on the recipient plant are less studied. Our studies aim:(i) to detect ginsenoside content in the ginseng of different growth-year and in different collection time and plot via HPLC method;(ii) to understand the allelopathic effects of the total ginsenoside (TGS) and its monomer fractions Rb1, Re and Rg1on ginseng seedling growth and development at different concentration(25mg·L-1,50mg·L-1and100mg·L-1);(iii) to examine the reaction of American ginsengs treated with ginsenoside of different concentration, extracted from4-year American ginseng with water;(iv) to study the influence of ginsenoside on the roots and adventitious roots of four cultivars-cabbage, cucumber, wheat, and mung bean;(v) to test the allelopathic effects of ginsenoside on ginseng seedling growth when they were planted in natural soils mixed with0.125g·kg-1,0.25g·kg-1and0.5g·kg-1TGS. The key conclusions are showed as below:
1. The TGS content of different growth-year ginsengs and ginsengs in different collection time and plot is distinctively different. The TGS contents extracted from the dried soils increased constantly as growth year prolonged. In both understory and farmland ginseng soils, the TGS content in June were higher than that in August. On the contrary, the TGS content in American ginseng soils in August were higher than that in June, and TGS content increased with growth year increasing. Rb1, Rb2, Rb3, Re, Rd and Rg1were detected in the ginseng soils of3-5year-old. The TGS content in the dried ginseng soils was approximately from0.031to0.332mg·g-1(average:0.0795mg·g-1); the TGS content in American ginseng soils was from0.025to0.154mg·g-1(average:0.0732mg·g-1), whereas the TGS contents in farmland soil were0.002-0.076mg·g-1(average:0.0213mg·g-1).
2. The four compounds all affected the root vigor (RV) and root length (RL) of ginseng seedling in different ways. The TGS and the Rb1treatments both showed a dose-dependent effect on RV; The RV enhanced drastically as the TGS and Rb1concentration increased, and then when their concentration moved to some certain level which is relatively high the RV began to decline. When the Rgl concentration was low, the RV was inhibited. And as the Rgl concentration went up, the RV increased as well. But just like the TGS and the Rbl, the RV then turned down quickly. As Re concentration increased, the RV enhanced slowly but constantly. For those ginsengs that were treated with ginsenoside at lower concentration (25mg·L-1), the chlorophyll synthesis was promoted. The promotion effect of the four compounds was in order:TGS>Rg1>Re>Rb1, and even if the promoting effect grew weaker as concentration increased, the chlorophyll content was still higher than that of the control. Electron microscope showed that the plasma membranes were detached from the cell wall, the structures of mitochondria and vacuolus were also dissolved under higher TGS and Rbl concentration. Hence, these abnormal physiological parameters in plant growth process indicated the responsibility of ginsenoside concentration.
3. The ginsenoside extracted from American ginseng with the deionzed water showed a dose-dependent effect on the root elongation. However there was no significant difference between the biomass of seedlings and roots before and after. The root vigor was inhibited significantly as Ginsenoside concentration increased. The production of O2-in seedlings sped up markedly as ginsenoside concentration increased and exposure time prolonged. The synthesis of soluble protein and chlorophyll in the seedlings was slightly stimulated by the ginsenoside. The promoting effect declined as the concentration increased and the treatment time prolonged, but the content is still higher than that of the control. Otherwise, the MDA content and the relative electric conductivity in leaves and roots increased slightly compared with the control as the concentration increased and the application time prolonged. The treated ultra-structure of root-tip cells showed that the nuclear membrane and nucleolus deformed and the vacuole membrane dissolved with the treatment time prolonging. The ginsenoside affected the underground growth of American ginseng more than aboveground growth. Taken together, these findings indicated that ginsenoside affected the early growth of American ginseng.
4. The alleopathic effect of TGS on the root growth of four cultivar species was conspicuously different.The RV, RL and RFW (root fresh weight) of wheat, cabbage and cucumber declined constantly as ginsenoside concentration ascended. However, the ginsenoside treatment had almost no influence on the hypocotyl adventitious roots of cucumber and mung bean. The RL, RFW, RV and Root number of mung bean adventitious roots had no significant response to ginsenoside, although the antioxidant enzymes activity rose a little bit over the control. Thereby, ginsenoside inhibited the root growth of the four indicator plants, and had stronger inhibitory effects on wheat and cucumber than on cabbage. However, there was no evidence to indicate that ginsenoside affected the hypocotyl cuttings growth in cucumber and mung bean.
5. The result of the experiment done with the soils from the new understory land indicated that there was no significant change in the biomass of the ginseng seedlings treated with ginsenoside, whereas the root elongation was reduced significantly by7.0%under higher concentration, the RV was declined by69.3%, and the fresh weight of roots slid by45.5%. The soluble protein content had little changes, yet the chlorophyll content had been in an uptrend. Even though the chlorophyll content decreased under higher concentration, it was still higher than that of the control. The color of ginseng roots changed to brown when TGS reached the highest. The IAA content first increased and then decreased as concentration ascended; the GA3content in all the treatments declined; the ABA content had no significant difference at lower concentration, but it increased sharply to2.39times than of the control when treated with high ginsenoside concentratrion.
Hence, we concluded that triterpenoid ginsenoside and their three monomers all disturbed the growth and development of ginseng and American ginseng, and this disturbance was affected by the types and the concentrations of ginsenosides. The ginsenoside do had allelopathic effect on other plants, yet it was lower than that on ginsengs and American ginsengs. It was found that triterpenoid ginsenoside may participate in the regulation of the formation and stability of plant population.
引文
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