全 文 ：外来入侵植物蒺藜草的化感效
应研究
马婉捷，缪绅裕 *，陶文琴，许镇健，王厚麟，陈
健辉，黄丽宜 （广州大学生命科学学院，广东
广州 510006）
摘 要 [目的] 探讨外来入侵植物蒺藜草的
潜在生态危害。 [方法] 利用种子萌发法研究
了不同浓度蒺藜草的水提液对 3 种茄科作物
种子萌发和幼苗生长的影响。 [结果] 培养 5
d， 不同水提液处理组的番茄萌发速度指数与
对照组相比无显著差异，但茄子的萌发速度指
数随处理浓度的上升而显著下降；所有辣椒种
子未见萌发。 培养 7 d，番茄根长随处理浓度
上升而受到抑制，茎长和综合化感效应则表现
为促进作用；所有处理组的茄子萌发率显著低
于对照组，较高浓度处理组的根长、茎长和综
合化感效应表现为受抑制。 辣椒萌发率受明
显抑制，最高浓度组完全未萌发。 培养 14 d，
茄子的所有指标均受较明显抑制。 最高处理
浓度组辣椒各指标受明显抑制。 [结论] 3 种作
物对蒺藜草化感物质的敏感性依次为茄子>
辣椒>番茄，表明蒺藜草对某些作物生长产生
不同程度的抑制作用，在农业生产上应注意加
强防治。
关键词 蒺藜草；化感效应；外来入侵植物；茄
科；作物
基金项目 广州市教育科学“十二五”规划项目
（12A008）； 广州市教育局科普项目 （12A102，
13B022-02）； 广 州 市 科 技 计 划 资 助 项 目
（2012KP024）。
作者简介 马婉捷（1992-），女，广东海丰人，
本科， 专业： 生物科学。 * 通讯作者，E-mail：
miaoshy@gzhu.edu.cn。
收稿日期 2014-02-24
修回日期 2014-06-11
Study on the Allelopathic Effects of Alien Invasive
Species Cenchrus echinatus on Seed Germina-
tion and Seedling Growth of Solanaceae Crops
Wanjie MA, Shenyu MIAO*, Wenqin TAO, Zhenjian XU, Houlin WANG, Jianhui CHEN, Liyi HUANG
School of Life Sciences, Guangzhou University, Guangzhou 510006, China
Supported by the Twelfth Five-Year Guideline Project of Education Sciences of
Guangzhou City (12A008); Popular Science Project of Bureau of Education of
Guangzhou Municipality (12A102, 13B022-02); Science and Technology Planning
Project of Guangdong Province (2012KP024).
*Corresponding author. E-mail: miaoshy@gzhu.edu.cn
Received: February 24, 2014 Accepted: June 11, 2014A
Agricultural Science & Technology, 2014, 15(6): 885-889
Copyright訫 2014, Information Institute of HAAS. All rights reserved Agricultural Basic Science and Technology
T here are approximately 25Cenchrus species in the world,which are distributed in tropical
and temperate areas and mainly origi-
nate in temperate arid regions of Africa
and the America. According to Flora of
China, two Cenchrus species were
found in China. To be specific, C. echi-
natus is mainly distributed in Guang-
dong, Hainan, Fujian, Taiwan, Hong
Kong, Guangxi and southern Yunnan;
C. calyculatus is mainly distributed
in sandy soil of seaside areas in
Fushun City, Liaoning Province [1]. In
recent years, several new Cenchrus
species were found in China, such
as C. longispinus [2], C. pauciflorus [3],
C. ciliaris and C. indius [4]. In January
2010, C. echinatus was included in the
list of the second batch of alien inva-
sive species by the Ministry of Envi-
ronmental Protection of the People’s
Republic of China[5].
C. echinatus is an annual herb
native to tropical America, which is
mainly distributed in 33° S-33° N trop-
ical and subtropical regions[6]. In China,
C. echinatus was collected for the first
Abstract [Objective] This study aimed to investigate the potential ecological dam-
ages of alien invasive species Cenchrus echinatus. [Method] By using seed germi-
nation method, the effects of different concentrations of C. echinatus aqueous ex-
tracts on seed germination and seedling growth of three Solanaceae crops were in-
vestigated. [Result] After incubated for 5 d, no significant differences were observed
in germination speed indices of Lycopersicum esculentum seeds between aqueous
extract groups and control groups; germination speed indices of Solanum melongena
seeds were significantly reduced with the increasing concentration of Cenchrus echi-
natus aqueous extracts; however, Capsicum frutescens seeds in various groups
were not germinated. After incubated for 7 d, root lengths of Lycopersicum esculen-
tum seedlings were inhibited with the increasing concentration of Cenchrus echinatus
aqueous extracts, while stem lengths and comprehensive allelopathic effects were
promoted; germination rates of Solanum melongena seeds in various groups were
all significantly lower than control groups; root length, stem length and comprehen-
sive allelopathic effects in higher concentration groups were inhibited; germination
rates of Capsicum frutescens seeds were significantly inhibited, and those in the
highest concentration group were not generated. After incubated for 14 d, all indica-
tors of Solanum melongena were significantly inhibited; various indicators of Cap-
sicum frutescens in the highest concentration group were significantly inhibited.
[Conclusion] The sensitivity of three Solanaceae crops to Cenchrus echinatus aque-
ous extracts presented an overall order of Solanum melongena > Capsicum
frutescens > Lycopersicum esculentum, suggesting that Cenchrus echinatus can to
some extent inhibit the growth of several crops, which should be strictly prevented
and controlled in agricultural production.
Key words Cenchrus echinatus; Allelopathic effect; Alien invasive plant; Solanaceae;
Crops
DOI:10.16175/j.cnki.1009-4229.2014.06.018
Agricultural Science & Technology 2014
time in Lanyu Island, Taiwan Province
in 1934. C. echinatus blooms through-
out the year in humid areas, which
can be propagated by seeds (com-
monly germinate in Acanthospermum)
and can be attached to clothing, ani-
mal furs or goods to spread. As a
weed species seriously harming the
production of a variety of fruits and
crops such as peanuts and sweet
potatoes, Cenchrus echinatus may
puncture the skins of humans and ani-
mals or hurt the eyes, mouths and
tongues of animals when mixed into
feed or forage, which can rapidly ex-
pand the occupy the gaps between
bare or newly reclaimed lands[6].
At present, very little information
is available on C. echinatus in China.
Tu et al.[7] elucidated the structure of a
new saponin from Tribulus terrestris by
using high-field gradient NMR tech-
niques. Zhang et al.[8] conducted char-
acteristic analysis, microscopic iden -
tification and TLC identification of
T. terrestris. Hu et al. [9] found T. ter-
restris in Putuo Mountain of Zhejiang
Province. Silva et al. [10] evaluated the
biological activities of seed extracts
and stilbene resveratrol-derived com-
pounds in Tribulus seeds and roots.
Dan et al. [11] investigated the effect of
the growth stage of Cenchrus echina-
tus on weed suppression imposed by
Atrazine. Fidelis et al. [12] surveyed the
emergences of C. echinatus and Cos-
mos sulphureus with different sowing
depths in two seasons. Nascimento
et al.[13] analyzed the allelopathic activ-
ity of C. echinatus. extracts on weeds
and crops. The allelopathic effects of
methanol extracts from stems and
roots of Cenchrus planted in the
greenhouse and laboratory on seed
germination, root length and stem
length of Panicum maximum, Ama-
ranthus hypochondriacus, Physalis
ixocarpa, Trifolium alexandrinum and
Lolium perenne were analyzed; results
indicated that the inhibitory or promo-
tion effects of methanol extracts from
stems and roots of Cenchrus on seed
germination and seedling growth de-
pended on the concentration of ex-
tracts and species of receptor plants.
Overall, the inhibitory effect of stem
extracts was lower than that of root
extracts[13]. In this study, the allelopathic
effects of aqueous extracts of C. echi-
natus on seed germination and
seedling growth of three Solanaceae
crops were investigated, which provid-
ed theoretical basis for the prevention
and control of alien invasive species
C. echinatus.
Materials and Methods
Materials
C. echinatus shoots were collect-
ed as experimental materials from
Guizhuwei, Shangying Town, Lufeng
City, Guangdong Province; seeds of
three Solanaceae crops were pur-
chased from Guangdong Academy of
Agricultural Sciences as receptor ma-
terials, including tomato (Lycoper-
sicum esculentum) (Jinguozhuang-
yuan), pepper (Capsicum frutescens)
(F1 generation of Xinla No.8) and egg-
plant (Solanum melongena) (Chang-
feng·Hongfei, F1 generation of Zi-
hongchangqie).
Preparation of aqueous extracts
Dried stems and leaves were cut
into 1 cm segments; 100 g of plant
segments were weighed, soaked into
1 000 ml of distilled water, placed for
24 h and filtrated to obtain water ex-
tract liquid (0.10 g/ml), which was
sealed and stored in a refrigerator at
4 ℃ before use. The water extract liq-
uid was distilled with diluted water into
0.02, 0.04, 0.06, 0.08 and 0.10 g/ml
aqueous extracts; distilled water was
used as a control (CK, 0.00 g/ml); petri
dishes and other experimental appa-
ratus were disinfected with dilute hy-
drochloric acid.
Seed germination and early seed-
ling growth
By using seed germination
method, 30 seeds of the same crop
species were placed on each of the
petri dishes coated with double layers
of filter paper, and added with different
concentrations of aqueous plant ex-
tracts. Thus, six different concentration
treatments were conducted including
control group, and each treatment was
repeated three times. Seeds were in-
cubated in an incubator at 25 ℃ under
light intensity≥ 3 500 lx for 12 h/d.
The number of generated seeds
in petri dishes was recorded regularly
every day. After germination, root
length and stem length of seedlings
were measured using a digital caliper.
Generated seeds were identified as
those with 1-2 mm radicles. Specifi-
cally, Lycopersicum esculentum seeds
were rapidly germinated and the ger-
mination experiment lasted 7 d; how-
ever, the germination experiments of
Solanum melongena and Capsicum
frutescens lasted 14 d. Finally, seed
germination rate, seedling root length
and stem length, allelopathic effect in-
dex, and comprehensive allelopathic
effect index of each crop species were
calculated.
Data processing and statistics
Germination rate Germination rate
(%) = (Number of seeds germinated
within a specified time / Total number
of seeds) × 100[14-15]
Germination speed index Germi-
nation speed index was calculated
based on the seed germination status
during the first five days:
Germination speed index=2(5X1+
4X2+3X3+2X4+X5)
To be specific, X1 indicates the to-
tal number of germinated seeds; X2,
X3, X4 and X5 indicate total number of
seeds germinated on the 2nd, 3rd, 4th
and 5th d, respectively[14].
Allelopathic effect index Allelo-
pathic effect index = 1-C/T (T≥C) or
RI = T/C-1(T
ue in control groups; T indicates the
value in experimental groups. Allelo-
pathic effect index >0 suggests pro-
motion effect; allelopathic effect in-
dex <0 suggests inhibitory effect. The
absolute value is related to the intensi-
ty of allelopathic effect[14-15].
Comprehensive allelopathic effect
index Comprehensive allelopathic
effect index = Allelopathic effect index
of (germination rate + root length +
stem length)/3.
To be specific, comprehensive al-
lelopathic effect index >0 suggests
promotion effect; comprehensive al-
lelopathic effect index <0 suggests in-
hibitory effect. The absolute value is
related to the intensity of allelopathic
effect[14-15].
Statistical analysis All the experi-
mental data were analyzed using Ex-
cel and SPSS statistical software, to
investigate the significance of differ-
ences with control groups.
Results and Analysis
Germination speed index and ger-
mination rate of crop seeds
Germination speed indices of the
886
Agricultural Science & Technology2014
seeds of three Solanaceae crops incu-
bated in different concentrations of
Cenchrus echinatus aqueous extracts
were calculated. As could be seen in
Table 1, no significant differences
were observed among germination
speed indices of Lycopersicum escu-
lentum seeds in various treatments;
germination speed indices of Solanum
melongena seeds treated with differ-
ent concentrations of Cenchrus echi-
natus aqueous extracts were signifi-
cantly lower compared with control
groups; none of Capsicum frutescens
seeds in various treatments were
generated. To be specific, germina-
tion speed indices of the seeds of
different crop species exhibited an
overall order of Lycopersicum escu-
lentum > Solanum melongena > Cap-
sicum frutescens.
Similar to germination indices of
seeds incubated for 5 d, no significant
differences were observed among ger-
mination indices of Lycopersicum es-
culentum seeds incubated for 7 d in
various treatments; germination in-
dices of Solanum melongena seeds
treated with different concentrations of
Cenchrus echinatus aqueous extracts
were significantly lower compared with
control groups and the germination
rates were gradually reduced with the
increasing concentration of aqueous
extracts; germination rates of Cap-
sicum frutescens seeds were also re-
duced with the increasing concentra-
tion of aqueous extracts; no significant
differences were observed between
low-concentration (0.02 g/ml) aqueous
extract treatment groups and control
groups, but high-concentration (0.04
g/ml) aqueous extract treatment
groups varied significantly or extreme-
ly significantly compared with control
groups.
After incubated for 14 d, germina-
tion rates of Solanum melongena
seeds treated with high concentration
(0.08 g/ml) and higher concentration
(0.10 g/ml) of Cenchrus echinatus
aqueous extracts were significantly
lower compared with control groups,
which indicated that Solanum melon-
gena seeds were gradually adapted
to allelopathic substances in Cenchrus
echinatus. No significant differences
were observed in germination rates
of Capsicum frutescens seeds be-
tween low-concentration aqueous ex-
tract treatment groups and control
groups, while germination rates of
C. frutescens seeds in other groups
varied significantly or extremely sig-
nificantly compared with control
groups.
Root length and stem length of
young seedlings
After incubated for 7 d, root
lengths of Lycopersicum esculentum
seedlings were slightly reduced with
the increase of aqueous extract con-
centration, which exhibited no signifi-
cant differences among various
groups; stem lengths of L. esculen-
tum seedlings increased slightly with
the increase of aqueous extract con-
centration, which exhibited no signifi-
cant differences among various
groups. Root lengths of Solanum mel-
ongena seedlings treated with low
concentration and lower concentration
of Cenchrus echinatus aqueous ex-
tracts increased slightly compared with
controls; root lengths of Solanum mel-
ongena seedlings treated with moder-
ate or higher concentration of
Cenchrus echinatus aqueous extracts
were reduced slightly with the increase
of aqueous extract concentration; root
lengths of Solanum melongena
seedlings treated with the highest
concentration of Cenchrus echinatus
aqueous extracts were significantly re-
duced compared with control groups.
Stem length of Solanum melongena
seedlings exhibited similar variation
trends to root lengths, but no signifi-
cant differences were observed am-
ong various groups. Capsicum
frutescens seeds treated with the
highest concentration of Cenchrus
echinatus aqueous extracts were not
generated, resulting in the extremely
significantly lower root lengths com-
pared with control groups; root lengths
of Capsicum frutescens seeds in other
aqueous extract treatments were
slightly improved compared with con-
trol groups but the increments were
gradually reduced with the increasing
concentration of aqueous extract.
Stem lengths in various groups all re-
mained unchanged.
After incubated for 14 d, root
lengths and stem lengths of Solanum
melongena seedlings were reduced
with the increasing concentration of
Cenchrus echinatus aqueous extracts.
Compared with control groups, root
lengths and stem lengths of Capsicum
frutescens seedlings increased first
and the declined with the increasing
concentration of Cenchrus echinatus
aqueous extracts; exhibiting no signifi-
Table 1 Germination of crop seeds incubated for 5, 7 and 14 d
Crop Treatmentg/ml
5 d 7 d 14 d
Germination speed
index
Germination
rate∥%
Germination
rate∥%
Lycopersicum 0(CK) 398±8.05 92.22±6.94 -
esculentum 0.02 413±21.00 96.67±3.33 -
0.04 385±14.77 92.22±5.09 -
0.06 414±30.24 92.22±6.94
0.08 423±12.87 93.33±3.33
0.10 407±13.85 93.33±6.67 -
Solanum 0(CK) 230±5.14 64.44±5.09 86.68±7.02
melongena 0.02 110±19.00** 37.78±1.92** 80.00±6.67
0.04 62±7.58** 33.33±5.77** 80.00±6.67
0.06 54±3.46** 31.11±3.85** 77.78±3.85
0.08 40±10.19** 30.00±3.33** 68.89±1.92*
0.10 21±3.00** 18.89±1.92** 66.67±8.28*
Capsicum 0(CK) 0 16.67±1.77 85.56±1.92
frutescens 0.02 0 16.67±1.33 80.00±11.55
0.04 0 13.33±0.82 74.44±6.62*
0.06 0 8.89±1.09** 71.11±6.18*
0.08 0 6.67±0.77** 65.56±8.44**
0.10 0 0** 5.56±0.92**
* represents a significant difference (P<0.05); ** represents an extremely significant
difference (P<0.01); - represents indicator undetermined.
887
Agricultural Science & Technology 2014
cant differences except in the highest
concentration group.
Allelopathic effect index and com-
prehensive allelopathic effect index
After incubated for 7 d and 14 d
respectively, allelopathic effect indices
and comprehensive allelopathic effect
indices of various indicators of different
crops were calculated. As could be
seen from Table 3, in various treat-
ments of Lycopersicum esculentum,
Cenchrus echinatus aqueous extracts
possessed negative effects on root
lengths, positive effects on stem
lengths and positive comprehensive
allelopathic effects, which indicated
that Cenchrus echinatus aqueous
extracts exhibited certain influences on
seed germination and seedling growth
of Lycopersicum esculentum.
Cenchrus echinatus aqueous extracts
possessed negative effects on germi-
nation rates of Solanum melongena
seeds; however, after incubated for 7
d, the effects on root lengths and stem
lengths and the comprehensive allelo-
pathic effects were positive in low con-
centration groups and negative in high
concentration groups; after incubated
for 14 d, the effects on root lengths
and stem lengths and the comprehen-
sive allelopathic effects were all nega-
tive in various groups, suggesting that
Cenchrus echinatus aqueous extracts
had significant adverse effects on
seed germination and seedling growth
of Solanum melongena. The effects
on germination rates of Capsicum
frutescens seeds were basically nega-
tive; the effects on root lengths and
stem lengths and the comprehensive
allelopathic effects were all negative in
the highest concentration group; com-
prehensive allelopathic effects in high-
er concentration groups were negative
after incubated for 7 d and positive af-
ter incubated for 14 d, which indicated
that Capsicum frutescens seedlings
were gradually adapted to the al-
lelopathic effects of Cenchrus echi-
natus aqueous extracts .
Table 2 Root length and stem length of crop seeds incubated for 7 and 14 d
Crop Treatmentg/ml
7 d 14 d
Root
length∥mm
Stem
length∥mm
Root
length∥mm
Stem
length∥mm
Lycopersicum 0(CK) 48.82±11.79 20.69±5.11 - -
esculentum 0.02 47.45±9.85 25.29±7.17 - -
0.04 45.60±8.13 25.66±5.97 - -
0.06 41.55±8.07 26.97±8.22 - -
0.08 39.51±7.00 29.31±8.43 - -
0.10 38.61±7.20 30.86±7.51 - -
Solanum 0(CK) 22.59±4.43 8.73±2.89 29.84±6.21 23.22±5.18
melongena 0.02 24.75±3.78. 13.04±3.45 23.92±5.54 18.59±6.77
0.04 23.91±3.71 13.67±2.30 22.39±5.38 18.63±5.08
0.06 20.05±4.83 10.37±3.04 22.24±5.91 17.40±4.85
0.08 18.94±4.09 7.61±2.32 20.65±4.56 15.76±4.90
0.10 12.79±4.54* 6.42±2.12 18.52±6.32 15.23±5.02
Capsicum 0(CK) 2.36±0.93 0 17.10±5.80 14.82±4.30
frutescens 0.02 3.08±0.67 0 19.95±5.63 16.09±5.03
0.04 2.92±0.92 0 23.05±8.54 16.62±5.71
0.06 2.85±0.81 0 24.58±7.23 18.47±4.89
0.08 2.61±0.68 0 23.78±6.42 16.04±4.31
0.10 0** 0 16.21±5.02 8.16±2.56*
* represents a significant difference (P<0.05); ** represents an extremely significant
difference (P<0.01); - represents indicator undetermined.
Table 3 Comprehensive allelopathic effects of seeds or young seedlings incubated for 7 and 14 d
Crop Treat-ment∥g/ml
7 d 14 d
Germination
rate
Root
length
Stem
length
Comprehensive
allelopathic
effect
Germination
rate
Root
length
Stem
length
Comprehensive
allelopathic
effect
Lycopersicum 0.02 0.048 -0.028 0.182 0.067 - - - -
esculentum 0.04 0.000 -0.066 0.194 0.043 - - - -
0.06 0.000 -0.149 0.233 0.028 - - - -
0.08 0.012 -0.191 0.294 0.038 - - - -
0.10 0.012 -0.209 0.330 0.044 - - - -
Solanum 0.02 -0.414 0.087 0.331 0.001 -0.077 -0.198 -0.199 -0.158
melongena 0.04 -0.483 0.055 0.361 -0.022 -0.077 -0.250 -0.198 -0.175
0.06 -0.517 -0.112 0.158 -0.157 -0.103 -0.255 -0.251 -0.203
0.08 -0.534 -0.162 -0.128 -0.275 -0.205 -0.308 -0.321 -0.278
0.10 -0.707 -0.434 -0.265 -0.469 -0.231 -0.379 -0.344 -0.318
Capsicum 0.02 0.000 0.234 - 0.117 -0.065 0.143 0.079 0.052
frutescens 0.04 -0.200 0.192 - -0.004 -0.130 0.258 0.108 0.079
0.06 -0.467 0.172 - -0.148 -0.169 0.304 0.198 0.111
0.08 -0.600 0.096 - -0.252 -0.234 0.281 0.076 0.041
0.10 -1.000 -1.000 - -1.000 -0.935 -0.052 -0.449 -0.479
* represents a significant difference (P<0.05); ** represents an extremely significant difference (P<0.01); - represents indicator
undetermined.
888
Agricultural Science & Technology2014
Responsible editor: Xiaohui FAN Responsible proofreader: Xiaoyan WU
Conclusion and Discussion
Different responses of seed germi-
nation and seedling growth of dif-
ferent crop species to C. echinatus
aqueous extracts
Overall, the sensitivity of three
crop species to C. echinatus aqueous
extracts presented an order of Sola-
num melongena >Capsicum frutesc-
ens >Lycopersicum esculentum. Ac-
cording to previous study [16], the in-
hibitory effects of aqueous extracts
of Leucaena leucocephala leaves on
different crop species presented an
overall order of Brassica juncea >
B. parachinensis > B. pekinensis.
Different adaptabilities of various
organs of the same crop species to
allelopathic substances in Cen-
chrus echinatus aqueous extracts
For instance, C. echinatus aque-
ous extracts exhibited inhibitory effects
on root lengths and promotion effects
on stem lengths of Lycopersicum es-
culentum seedlings, which also inhibit-
ed the germination rates of Capsicum.
frutescens seeds, but promoted root
lengths and stem lengths of C. frute-
scens seedlings in low concentration
groups. Results of the present study
were consistent to the research of
Wedelia trilobata that confirmed W. tr-
ilobata aqueous extracts exhibited in-
hibitory effects on seed germination
and root length of Paspalum notatum
and Cynodon dactylon, but had no
effects on stem length [17]. In addition,
aqueous extracts of Leucaena leuco-
cephala leaves could also inhibit the
root length of receptor plants[16].
Different adaptabilities of the same
tissues of the same crop species to
allelopathic substances in Cen-
chrus echinatus aqueous extracts
at different incubation stages
The inhibitory effects on germina-
tion rates of Solanum melongena
seeds incubated for 7 d were higher
compared with those incubated for 14
d; however, the inhibitory effects on
root length, stem length and compre-
hensive allelopathic effect indices basi-
cally increased with the increase of in-
cubation time. On the contrary, the in-
hibitory effects on germination rates of
Capsicum frutescens seeds were re-
duced with the increase of incubation
time.
Allelopathic effects of alien inva-
sive species are rather complex.
Therefore, allelopathic effects on dif-
ferent crops should be analyzed sepa-
rately to investigate the effect rules. In
this study, significant inhibitory effects
of Cenchrus echinatus aqueous ex-
tracts on various tissues of Solanum
melongena, Capsicum frutescens and
Lycopersicum esculentum were ob-
served, suggesting that Cenchrus
echinatus may exhibit allelopathic ef-
fects on other crops, which should be
strictly prevented and controlled in
crop-growing areas to reduce the eco-
logical risk.
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