全 文 ：人参皂苷对 4 种栽培作物早期根系发育的化感效应*
张秋菊1,2, 张爱华1, 杨摇 鹤1, 张连学1**
(1 吉林农业大学中药材学院, 吉林 长春摇 130118; 2 通化师范学院生物系, 吉林 通化摇 134001)
摘要: 人参、 西洋参能通过根系分泌三萜皂苷等化感物质, 严重影响后茬人参的生长, 但对人参以外的植
物是否具有化感效应尚不清楚。 本实验研究了不同质量浓度的人参皂苷对小麦、 白菜、 黄瓜及绿豆 4 种常
见栽培作物早期根系发育的影响, 结果发现人参皂苷处理液 (25、 50 和 100 mg·L-1) 对 4 种作物主根及
不定根的发育影响不尽相同。 随着处理质量浓度的升高, 小麦、 白菜、 黄瓜根系活力分别比同组 CK明显
降低, 根长、 根鲜重也呈降低趋势。 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
重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。 总之, 人参皂苷对 4 种栽培
作物的主根发育均有抑制作用, 尤其对小麦、 黄瓜主根生长的抑制作用较强; 但对黄瓜和绿豆不定根发育
的影响不明显。
关键词: 人参皂苷; 根系; 发育; 下胚轴; 不定根; 化感效应
中图分类号: Q 945摇 摇 摇 摇 摇 摇 摇 文献标识码: A摇 摇 摇 摇 摇 摇 摇 文章编号: 2095-0845(2012)04-391-06
Allelopathic Effects of Ginsenosides from Panax ginseng
on the Root Development of Four Cultivar Plants
in Their Early Growth Stages
ZHANG Qiu鄄Ju1,2, ZHANG Ai鄄Hua1, YANG He1, ZHANG Lian鄄Xue1**
(1 College of Chinese Medicinal Materials, Jilin Agricultural University, Changchun 130118, China;
2 Department of Biology, Tonghua Normal University, Tonghua 134001, China)
Abstract: Some allelochemicals, including ginsenosides secreted from the roots of ginseng and American ginseng,
have been considered to play a regulatory role in population regeneration. This study was conducted to investigate the
allelopathic effects of ginsenosides on the root development of several cultivar plants at the early growth stage. The
different inhibitions in wheat, cabbage, and cucumber roots resulting from ginsenosides treatment at different con鄄
centrations were observed. The root vigor of the three receptor plants constantly decreased compared with the control
when the treatment concentration increased; the root length and root fresh weight exhibited similar trends. However,
ginsenoside treatment showed negligible effect on the hypocotyl adventitious roots of cucumber and mung bean. The
root number, root length, root fresh weight, and root vigor showed no significant changes; however, the antioxidant
enzyme activities were slightly enhanced compared with that of the control. These results indicate that the ginsen鄄
osides inhibited the root growth of all indicator plants, and these inhibitory effects were stronger on wheat and cu鄄
cumber than on cabbage. However, no evidence indicates that ginsenosides inhibited the growth of hypocotyl cuttings
of cucumber and mung bean.
Key words: Ginsenosides; Root; Development; Hypocotyls; Adventitious roots; Allelopathic effect
植 物 分 类 与 资 源 学 报摇 2012, 34 (4): 391 ~ 396
Plant Diversity and Resources摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 DOI: 10. 3724 / SP. J. 1143. 2012. 11172
*
**
Foundation items: The National Natural Science Foundation of China (31070316) and National Science & Technology Pillar Program during
the Eleventh Five鄄year Plan Period (2007BAI38B01)
Author for correspondence; E鄄mail: zlx863@ 163. com
Received date: 2012-04-05, Accepted date: 2012-05-16
作者简介: 张秋菊 (1968-) 女, 博士, 副教授, 主要从事植物生理生态学研究。 E鄄mail: zhangqiuju5515@ 163. com
摇 Panax ginseng C. A. Mey. is an important tra鄄
ditional herb that belongs to the Araliaceae family
and is widely cultivated in the Changbai mountain
area of Jilin Province (Liu and Xiao, 1992). Gin鄄
seng cannot be cultivated consecutively on the same
plot of land within 30 years, or its yield and quality
would be greatly reduced (Jin et al., 2006).
Plants can diffuse organic compounds into the
environment, which then accumulate in the rhizo鄄
sphere zone and affect the growth and development of
neighboring plants both positively and negatively, a
phenomenon called allelopathy ( Inderjit and Duke,
2003). The allelochemicals are mainly plant second鄄
ary metabolites that can be released via volatilization,
root exudation, leaching, and plant residue decompo鄄
sition in soil. Researchers believe that the irregular
cultivation of ginseng is caused by soil deterioration,
pathogenic fungal accumulation, and allelochemicals
in soil (Chen et al., 2006; Zhang et al., 2008; Lei
et al., 2010a). Hence, a rational rotation is one of
the workable solutions to the limitations in ginseng re鄄
planting. However, some allelochemicals found in
ginseng soil not only inhibits ginseng growth, but also
disrupts the growth of other plants, such as wheat,
cabbage, maize, burdock, and rhubarb, among oth鄄
ers (Liu and Wang, 2010; Lei et al., 2010b).
Ginsenosides are major pharmaceutically active
compounds found in the different tissues of ginseng,
especially in the roots (Zhang et al., 2005). Yang
et al. (1982) first detected ginsenosides in the
root exudates of ginseng. Ginsenosides collected from
American ginseng is present at levels approximating
0. 06% of the soil dry weight (Nicol et al., 2003).
Reports have confirmed that ginsenosides in ginseng
and American ginseng can affect the growth of soil鄄
borne pathogens, as well as seed germination, an鄄
tioxidant enzyme activities, and root vigor through
the action of active allelochemicals (Yousef and Ber鄄
nards, 2006; Zhang et al., 2009; Zhang et al.,
2011; Lei et al., 2011).
However, the effect of ginsenosides in old gin鄄
seng soil on other cultivar plants remains to be deter鄄
mined. Furthermore, no studies on the effects of al鄄
lelochemicals on adventitious root development have
been reported to date. The present study aims to (i)
demonstrate the allelopathic effects of ginsenosides
on the root growth and development of wheat, cab鄄
bage, and cucumber, and (ii) determine the allelo鄄
pathic activity of ginsenosides on the adventitious
roots of cucumber and mung bean.
Materials and methods
Plant materials
The dried roots of four鄄year鄄old ginseng were
obtained from Fusong town, Jilin Province, China.
Wheat (Triticum aestivum L. ), cabbage (Brassica
chinensis L. ), cucumber (Cucumis sativus L. ), and
mung bean (Vigna radiate R. Wilozak) were chosen
as indicator plants in the allelopathy bioassay and hy鄄
pocotyl adventitious root development experiments.
All tested seeds were purchased at the seed鄄market.
Ginsenosides extraction and identification
Total ginsenoside ( TGS) was prepared from
ginseng roots according to Zhang et al. ( 2011 ).
Distilled water was added to the root powder ( v 颐 w
=10 颐 1), and the mixture was soaked overnight,
extracted three times in an ultrasonicator for 30 mi鄄
nutes, and vacuum鄄filtered. The combined filtrate
was loaded onto D101 extraction columns. Eluents
with 80% ethanol were evaporated to dryness and
purified using anhydrous ethanol, and the TGS con鄄
tent was determined via high鄄performance liquid
chromatography (HPLC > 80% ). The TGS was dis鄄
solved and diluted in distilled water to three concen鄄
trations namely, 25 mg·L-1, 50 mg·L-1, and 100
mg·L-1 and the solutions were stored at 4 益 in the
dark for use in the subsequent experiments.
Bioassay
Twenty grains of wheat and cucumber seeds (or
50 grains for cabbage) were surface鄄sterilized and
individually sown in 20 cm伊20 cm伊10 cm culture
box containing clean glass sand. Approximately 10
mL each of the previously prepared solutions (25,
50, and 100 mg·L-1) of TGS was added to contain鄄
293摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 植 物 分 类 与 资 源 学 报摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 第 34 卷
ers following a previously described procedure (Nic鄄
ol et al., 2003; Zhang et al., 2011). The controls
received only deionized water. All boxes were sealed
with plastic films and incubated in the dark at 25 益
followed by a 12 h / 12 h light / dark cycle until germi鄄
nation. The glass sands were kept moist with the
same volume of half鄄strength Hoagland爷 s solution
(pH 5. 6) throughout the entire experiment period.
The root length (RL) and root fresh weight (RFW)
were measured after six days incubation. The root
length / seedling height value (R / S) was calculated.
Root vigor ( RV) was measured according to the
method prescribed by Gao (2006).
Hypocotyls adventitious root incubation
Presoaked cucumber and mung bean seeds were
placed on two layers of moistened filter paper on a
10 cm鄄diameter Petri鄄dish and incubated for 48 h in
the dark at 25 益 . The uniform radicals were collect鄄
ed, and their root tips were cut off. The hypocotyls
were cut into 1. 5 cm segments and placed in a con鄄
tainer filled with glass sand at 20 cuttings per box.
The treated groups were irrigated with or without 10
mL of 25 mg·L-1 to 100 mg·L-1 TGS. All treat鄄
ments and controls were kept in 1 / 2 half鄄strength
Hoagland爷 s solution to maintain the appropriate
moisture. After six days of incubation in the dark at
25 益, the roots were harvested, and their growth pa鄄
rameters were measured. Antioxidant enzyme activi鄄
ties were measured after treated 48 h of treatment.
Superoxide dismutase (SOD), peroxidase (POD),
and catalas ( CAT) activities were determined fol鄄
lowing the method prescribed by Gao (2006).
Statistical analysis
All treatments were arranged in a completely
randomized design, with four replications each. The
response index ( RI) was calculated following the
method of Williamson (1998) using the equation, RI
=1鄄C / T (T逸C) or RI = T / C鄄1 (Tcontrol, and T means treatment value. The data were
analyzed via one鄄way ANOVA using the SPSS 17. 0
statistical software (SPSS Inc., USA), followed by
multiple comparisons via the least significant differ鄄
ence (LSD) tests at the 0. 05 or 0. 01 level.
Results and discussion
Allelopathic activities of TGS on wheat, cabbage,
and cucumber roots
The allelopathic potential of TGS from ginseng
on wheat, cabbage, and cucumber root growth and
development during an early growth stage was deter鄄
mined. The inhibitory rate increased with the in鄄
crease in treatment concentration ( Table 1). The
changes in the RL and RFW of the three receptor
plants showed similar trends with low to high TGS
concentrations. However, significant differences in
the RV values of wheat, cabbage, and cucumber
were found. The strongest inhibition of RV was ob鄄
served in cucumber after treatment with 100 mg·L-1
TGS, whereas the minimum RV was detected in cab鄄
bage at a lower concentration. These results suggest
that TGS exhibited similar allelopathic effects on di鄄
cotyledonous and monocotyledonous plant species.
Moreover, TGS may play the same role during the
early growth of cultivar plants other than ginseng and
American ginseng. However, the cause of the differ鄄
ent allelopathy on cabbage RV cannot be determined.
Table 1摇 Response index (RI) of TGS on root development in wheat, cabbage and cucumber
TGS
concentration
/ mg·L-1
RL
wheat cabbage cucumber
RFW
wheat cabbage cucumber
RV
wheat cabbage cucumber
25 -0. 031* -0. 144摇 -0. 252* -0. 124 -0. 152 -0. 355** -0. 238 +0. 034 -0. 262*
50 -0. 027* -0. 428* -0. 188摇 -0. 085 -0. 186 -0. 307* 摇 -0. 480** -0. 019 -0. 038摇
100 -0. 105** -0. 509* -0. 282* -0. 238* -0. 225* -0. 426** -0. 268 -0. 080 -0. 908**
RI +indicates growth promotion; RI -indicates growth inhibition. * and ** indicate significant differences between the treatments at the 0. 05 and
0. 01 levels respectively. RL: root length, RFW: root fresh weight, RV: root vigor
3934 期摇 摇 摇 摇 ZHANG Qiu鄄Ju et al. : Allelopathic Effects of Ginsenosides from Panax ginseng on the Root …摇 摇 摇 摇 摇
摇 The R / S is typically used to describe the corre鄄
lation between aboveground and underground seed鄄
ling growth. In this study, the R / S values ( seed鄄
lings data not shown) of wheat and cucumber all de鄄
clined compared with that of the control, except that
of cabbage. In particular, the R / S value of cucum鄄
ber showed the most significant decline at 24. 2% ,
14. 7% , and 25. 3% comparing with the control.
Therefore, the roots may be more sensitive to
TGS than the seedlings. This finding is consistent
with a previous report on allelopathic mechanism
(Weir et al., 2004; Zhang et al., 2011).
Allelopathic activities of TGS on cucumber and
mung bean adventitious root
The changes in the root number, RL, RFW,
and RV of adventitious roots of cucumber and mung
bean hypocotyl cuttings after TGS treatment were de鄄
termined respectively ( Table 2). TGS showed no
significant effect on the number of adventitious roots
of the two plants. The number of adventitious roots
of mung bean slightly increased with increased TGS
concentration, whereas that of cucumber decreased
only at higher concentrations. Higher TGS concen鄄
trations slightly inhibited the adventitious root devel鄄
opment in cucumber, whereas lower concentrations
promoted the process. The maximum RV index was
observed after 100 mg·L-1 TGS treatment. This
dose鄄dependent response, which is a common allelo鄄
chemical phenomenon, has been documented in va鄄
rious plants (Zhang et al., 2008; Duke, 2010; Bi
et al., 2010). By contrast, both the RL and RFW
were stimulated after TGS application at different
concentrations, whereas the RV of mung bean de鄄
creased. Nevertheless, no evidence to indicates that
the effect of TGS on the adventitious roots of hypo鄄
cotyls is greater more than that on the roots.
Allelopathic activities of TGS on the antioxidant
enzyme of cucumber and mung bean adventi鄄
tious roots
In response to environmental stress, plants can
produce reactive oxygen species ( ROS ), which
cause oxidative damage to the cell membrane. How鄄
ever, ROS scavengers such as SOD, POD, and CAT
protect against the damaging effects of ROS (Qian et
al., 2009; Duke, 2010). SOD is primarily respon鄄
sible for ROS scavenging, whereas POD and CAT
are primarily responsible for the decomposition of
peroxidant. Exposure to allelochemicals, significant鄄
ly stimulates the SOD and POD activities of some
plant species. (Weir et al., 2004; Huang et al.,
2010). Table 3 clearly indicates the different chan鄄
ges in the antioxidant enzyme activities of the adven鄄
titious roots of two plants in response to different
TGS concentrations. The SOD activities in the cu鄄
cumber adventitious root were drastically enhanced,
but the POD activities of the adventitious root of cu鄄
cumber increased compared with that of the control.
The maximum POD appeared after 100 mg·L-1 TGS
treatment. Meanwhile, the CAT activities also
slightly increased compared with the SOD and POD
activities, but no significant difference was found
between the values. However, all enzyme activities
in the mung bean adventitious root showed no signifi鄄
cant change at the three concentrations, although
they showed increasing trends.
Table 2摇 RI of TGS on the adventitious root development in cucumber and mung bean
TGS
concentration
/ mg·L-1
RL
cucumber mug bean
RFW
cucumber mug bean
RV
cucumber mug bean
25 +0. 070 +0. 054 +0. 051 +0. 022 +0. 085 -0. 147
50 -0. 075 +0. 102 -0. 003 +0. 077 -0. 019 -0. 206
100 -0. 038 +0. 028 +0. 013 +0. 008 -0. 148* -0. 294
*P<0. 05; ** P<0. 01. For other notes, see Table 1.
493摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 植 物 分 类 与 资 源 学 报摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 第 34 卷
Table 3摇 RI of TGS on the antioxidant enzyme activities of the cucumber and mung bean adventitious roots
TGS
concentration
/ mg·L-1
SOD / U·g-1 FW·min-1
cucumber mung bean
POD / U·g-1 FW·min-1
cucumber mung bean
CAT / U·g-1 FW·min-1
cucumber mung bean
0 (CK) 39. 11依0. 25 35. 13依0. 08 115. 61依3. 87 122. 43依2. 05 30. 16依0. 56 28. 32依0. 21
25 46. 42依0. 39 31. 87依0. 45 123. 82依0. 65 127. 45依0. 94 34. 23依0. 18 26. 28依0. 55
50 57. 73依0. 48* 39. 08依0. 27 138. 77依1. 41 129. 72依1. 56 36. 87依0. 47 35. 13依0. 34
100 59. 32依0. 16* 49. 54依0. 95 152. 11依5. 01* 138. 17依3. 75 41. 13依2. 08 40. 76依0. 58
*P<0. 05; ** P<0. 01. For other notes, see Table 1
摇 Underground plants are highly susceptible to
soil allelochemicals during the different seedling sta鄄
ges, possibly due to their high metabolic speed (In鄄
derjit and Duke, 2003). In a similar manner, the
allelopathic effect of TGS on root and adventitious
root development can be detected in the very early
growth stage of several cultivar species in the present
study. These findings suggest that TGS accumulated
in old ginseng soil not only disrupted the reseeding
of ginseng, but also affected the population estab鄄
lishment of other crops. In fact, a few distantly re鄄
lated rotational crops (rice, asarum, maize, and al鄄
falfa, among others) were collected and in an old
ginseng plot (Yang et al., 2008). The results of the
current study would be beneficial to future studies on
the rotation system of ginseng. In addition, the dif鄄
ferent mechanism of action of TGS as allelochemicals
on the main root and adventitious root growth should
be the focus of future studies.
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中国科技核心期刊、 中国农业核心期刊、
全国中文核心期刊、 全国优秀农业期刊
《植物遗传资源学报》 征订启事
《植物遗传资源学报》 是中国农业科学院作物科学研究所和中国农学会主办的学术期刊, 为中国
科技论文统计源期刊、 中国科学引文数据库来源期刊 (核心期刊)、 中国核心期刊 (遴选) 数据库收
录期刊、 中国学术期刊综合评价数据库统计源期刊, 又被 《中国生物学文摘》 和中国生物学文献数
据库、 中文科技期刊数据库收录。 据中国期刊引证研究报告统计, 2011 年度 《植物遗传资源学报》
影响因子 1. 396。 影响因子在自然科学与工程技术类学科排序第 9 名。
报道内容为大田、 园艺作物, 观赏、 药用植物, 林用植物、 草类植物及其一切经济植物的有关植
物遗传资源基础理论研究、 应用研究方面的研究成果、 创新性学术论文和高水平综述或评论。 诸如,
种质资源的考察、 收集、 保存、 评价、 利用、 创新, 信息学、 管理学等; 起源、 演化、 分类等系统
学; 基因发掘、 鉴定、 克隆、 基因文库建立、 遗传多样性研究。
双月刊, 大 16 开本, 128 页。 定价 20 元, 全年 120 元。 各地邮局发行。
邮发代号: 82-643。 国内刊号 CN11-4996 / S, 国际统一刊号 ISSN1672-1810。
本刊编辑部常年办理订阅手续, 如需邮挂每期另加 3 元。
地址: 北京市中关村南大街 12 号摇 中国农业科学院 《植物遗传资源学报》 编辑部
邮编: 100081摇 电话: 010-82105794摇 010-82105796 (兼传真)
网址: www. zwyczy. cn; E鄄mail: zwyczyxb2003@ 163. com摇 zwyczyxb2003@ sina. com
693摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 植 物 分 类 与 资 源 学 报摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 第 34 卷