全 文 ：基金项目：黑龙江农垦总局课题 (No. HNK125B-02-03).
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The Growth and Physiololgical Characteristics of Uniconazole (S3307)
on Coix Seedlings Under Drought Stress
HUANG Yu-Lan XIANG Jun-Liang YIN Kui-De*
Heilongjiang Bayi Agricultural University, Daqing 163319, China
*Corresponding author, yinkuide@163.com
Abstract Coix (Coix lachryma- jobi) is a kind of medicine and food crop, while drought is the most main
factor limiting its growth. To explore the applicative effects of uniconazile (S3307) on coix seedlings under
drought stress, the growth parameters, anti- oxidant enzyme activities, chlorophyll content and chloroplast
ultrastructure in Yiliao No. 5 under drought stress were investigated through nutrient solution cultivation in
greenhouse. The results showed that the stem diameter and ground biomass were significantly improved with
9 mg/L S3307 under drought stress (P＜0.05). Compared with the S0 (drought group), stem diameter were
increased for 16.19%, and the above-ground and under-ground biomass increased for 19.56% and 22.75%,
respectively. The activities of peroxidase (POD), catalase (CAT) and superoxide dismutase (SOD) were
significantly enhanced (P＜0.05), while POD and CAT activities of roots were higher than those of leafs.
Total chlorophyll content in the S3 (9 mg/L S3307 group) was the highest in all the treatments, which was
increased by 7.35% compared to S0 (drought treatment group) ( P＜0.05). Meanwhile, S3307 addition could
improve chloroplast and root cell ultrastructure and lighten the damage to grana ultrastructure under drought
stress, which was consistent with increasing chlorophyll content. All the analysis showed that 9 mg/L S3307
concentration was effective on alleviating the injury of coix seedlings under drought stress. This study can
clarify physiological mechanism of coix drought resistance and provide a theoretical basis for S3307 in coix
production and practice of drought resistance.
Keywords Uniconazile (S3307), Drought stress, Coix seedlings, Antioxidant enzyme, Ultrastructure
烯效唑(S3307)对干旱胁迫下薏苡幼苗生长及生理特性的影响
黄玉兰 向君亮 殷奎德*
黑龙江八一农垦大学，大庆163319
*通讯作者，yinkuide@163.com
摘 要 薏苡(Coix lachryma-jobi)是一种药食兼用的作物，干旱是限制薏苡种植和生长最为主要的因
素。为探讨烯效唑(uniconazole, S3307)在薏苡苗期抗旱方面的应用效果，本研究采用营养液水培法，以
薏辽5号为材料，研究外源S3307对干旱胁迫下薏苡生长指标、抗氧化酶活性、叶绿体含量及超微结构的
影响。结果表明，9 mg/L S3307显著提高了干旱胁迫下薏苡幼苗的茎粗和生物量(P＜0.05)。茎粗比干旱
组(S0)提高了16.19%，地上、地下生物量比S0分别提高了19.56%和22.75%。另外，9 mg/L S3307组中，过
氧化物酶 (peroxidase, POD)，过氧化氢酶(catalase, CAT) 和超氧化物歧化酶 (superoxide dismutase, SOD)
活性显著增加(P＜0.05)，其中根系的抗氧化酶活性要显著高于叶片(P＜0.05)。在所有处理组中，9 mg/L
Online system: http://www.jabiotech.org
农 业 生 物 技 术 学 报
Journal of Agricultural Biotechnology
2017, 25(2): 231~239
DOI: 10.3969/j.issn.1674-7968.2017.02.007
网络出版时间：2017-01-03 16:53:42
网络出版地址：http://www.cnki.net/kcms/detail/11.3342.S.20170103.1653.006.html
农业生物技术学报
Journal of Agricultural Biotechnology
Drought is one of the most severe agriculture
problems limiting growth and production of crops
(Pastori, Foyer, 2002), which leads to moisture deple-
tion in soil and water potential deficient in plant tis-
sues. When plants are subjected to drought stress,
various reactive oxygen species (ROS) are generat-
ed, such as superoxide, H 2O2 and hydroxyl radicals,
which cause oxidative damage to plants (Sminoff,
1993). Water is an essential reagent and solvent for
chemical reactions and metabolites translocation
(Crusciol et al., 2008). Therefore, drought has a cru-
cial impact on current agriculture production. Coix
(Coix lachryma- jobi) is commonly known as Jobs
tears, and is a member of the grass family in the tribe
Maydeae. Recently, the high protein nutrient of Jobs
tears is more and more concerned by people. Many
scholars has carried out the studies of pharmacology,
cell genetics, activities on coix, while there is less re-
port on physiology and biochemistry during cultiva-
tion process. Meanwhile, coix is a crop of being keen
on water, which habits and characteristics are similar
to rice (Oryza sativa). Therefore, drought is more sen-
sitive than other cereal crops for coix.
It has been well documented that plant growth
regulators (PGRs) play important roles in crop pro-
duction and resistance to environmental stress (Part
et al., 2008). As a member of the triazole PGRs fami-
ly, uniconazile (S3307) is increasingly applied in
crops as a plant growth retardant (Wu et al., 2013).
Some studies showed that S3307 exerts beneficial ef-
fects on alleviating various abiotic stresses, such as
drought stress, salt injury, extreme temperature dam-
age, waterlogging, sulphur dioxide and herbicide
stress (Zhang et al., 2007; Duan et al., 2008; Chaum
et al., 2009; Zhang et al., 2012). For example, S3307
treatment had signifcantly improved morphological
characteristics, protective enzyme system activity
and increased relvtive chlorophyll content for Pinelli-
aternata. (Thunb.) under drought stress (Cheng,
2010). S3307 soaking treatment can increase soluble
protein content in root and stem dry weight of soy-
bean (Glycine max) under drought stress (Zhang,
2016). Consequently, S3307 is applying more and
more frequently in agriculture.
To show the effectiveness of S3307 in mitigat-
ing the adverse effects of drought stress and investi-
gate possible physiological mechanisms of drought
stress in coix seedlings, growth, antioxidases activi-
ties, chlorophyll content and chloroplast ultrastruc-
ture were studied. All the results will help us to evalu-
ate the possibility of S3307 application for improv-
ing coix drought resistance.
1 Material and Methods
1.1 Plant materials and uniconazole treatments
Yiliao No. 5 (Coix lachryma- jobi) was obtained
from Coix Research Center of Heilongjiang Bayi Ag-
ricultural University, Heilongjiang Province, China.
The experiment was conducted in plant culture room
in Agricultural Department of Heilongjiang Bayi Ag-
ricultural University. Coxi seeds were sterilized with
10% NaClO solution for 30 min, rinsed thoroughly 3
times with distilled water and allowed to germinate
in the dark. After 96 h, germinated seedlings were re-
transferred to Hoaglands nutrient solution, which
seedlings were grown in a greenhouse at 25~30 ℃
with 13 h day/11 h night photoperiod, with 60% ~
70% humidity. The nutrient solution was renewed ev-
ery other day, and pH was 5.8. When the seedlings
grew up to two leaves, seedlings were used for the
experiments. A wide range of concentrations of uni-
S3307组中叶绿体含量最高，比干旱组增加7.35% (P＜0.05)。同时，S3307能够减轻干旱胁迫下对叶片和
根系细胞超微结构的影响；9 mg/L S3307能够有效减轻干旱胁迫对薏苡幼苗的伤害。研究结果为明确
S3307缓解薏苡抗旱能力的生理机制提供了基础资料，同时为S3307在薏苡抗性生产中的应用提供理论
依据。
关键词 烯效唑(S3307)，干旱胁迫，薏苡幼苗，抗氧化酶，超微结构
中图分类号 S513 文献标识码 A
232
conazile (S3307) was examined in previous tests.
Furthermore, the adequate range of S3307 solution
was 0~15 mg/L (Mei et al., 2014). Therefore, coix
seedlings were pretreated in 0, 3, 6, 9, 12 and 15 mg/
L S3307 solution concentration for 72 h at 25 ℃ ,
named with S0, S1, S2, S3, S4 and S5, respectively.
After 72 h of treatment, drought stress was imposed
on the seedlings in the presence of 20% polyethylene
glycol (PEG) 6000 solution (-0.8 MPa) by immers-
ing the roots in the solution and three replications
were used. According to growth parameters, concen-
tration of S3307 was selected as the most appropriate
concentration for further experiment. Samples of the
coix leaves and roots were harvested at the same
time points, which were 0, 24, 48, 72 and 96 h under
PEG stress for antioxidant enzyme activities. Mean-
while chlorophyll content and leaf chloroplast ultra-
structure were tested after 72 h of the PEG stress.
1.2 Growth parameters measurement
After 72 h of the PEG stress, thirty seedlings
were utilized for measuring plant and root height,
while stem diameter being measured with vernier cal-
iper in minimum value 0.02 mm. The packaged
roots, stems and leaves of those ten seedlings by
draft paper were oven- dried at 105 ℃ for 30 min,
and then dried at 80 ℃ to constant weight. The dry
weights were measured by electronic balance. The re-
sult of above- ground and under- ground biomasses
were in dry weight (mg).
1.3 Enzyme activity assays
Peroxidase (POD), catalase (CAT) and superox-
ide dismutase (SOD) were extracted as Maia et al.
(2010). Enzymes were extracted from 0.5 g leaves or
roots with a prechilled mortar being added 0.1 mol/L
phosphate buffer (pH 7.8) and were centrifuged at 15
000 ×g for 15 min. The supernatants were used to as-
say enzyme activity, which operations for the en-
zyme extractions preparation were performed at 4 ℃.
SOD activity was measured according to Li
(2000). The tubes were placed under fluorescent
lamps (4 000 lx) for 30 min. One unit of SOD was
defined as the amount of extract that gave half-maxi-
mal inhibition of nitroblue tetrazolium reduction in
the experimental conditions. POD activity was deter-
mined according to Cakmak et al.,(1993). The differ-
ent absorbance was recorded with guaiacol oxidation
at 470 nm. CAT activity was assayed according to
Gong et al. (2005). The reaction was initiated by add-
ing enzyme extract and the activity was calculated by
monitoring the initial rate of H2O2 disappearance at
240 nm.
1.4 Chlorophyll content
Chlorophyll content was estimated according to
Strain and Svec (1996). The fresh leaves taken from
different treatments were extracted with 1∶1 alcohol
and acetone mixture. The OD was measured by using
a visible spectrophotometer at 663 and 645 nm wave-
lengths. Chlorophyll content were calculated by the
following equation:
Ca (mg/mL)=11.64(OD663)−2.16(OD645)
Cb (mg/mL)= 0.97(OD645)−3.94(OD663)
Ct=Ca+Cb=8.02OD663+20.21OD645
Ct: Total chlorophyll content; Ca: Chlorophyll a
content; Cb: Chlorophyll b content.
1.5 Leaf and root ultrastructure
The leaves and roots at the same position on 3
parts of each treatment were taken and repaired for ≤
(1 mm×5 mm). Then the sample segments were im-
mediately put into 2.5% glutaraldehyde for 24 h at
4 ℃, with 1% glutaraldehyde 4 h to be fixed. The sec-
tions were dehydrated in ethanol gradually, and fur-
ther treated for being transparent, paraffining, embed-
ding, production, dyeing, sealing and being observed
by the electron microscope (Wang, Li, 2007).
1.6 Statistical analysis
All the datas were the mean values of 3 repli-
cates at least, which results were as X ±SD. The datas
were subjected to statistical analysis by Microsoft Ex-
cel and SPSS 20.0 for testing the significance of dif-
ferences.
2 Results and Analysis
烯效唑(S3307)对干旱胁迫下薏苡幼苗生长及生理特性的影响
The Growth and Physiololgical Characteristics of Uniconazole (S3307) on Coix Seedlings Under Drought Stress 233
农业生物技术学报
Journal of Agricultural Biotechnology
2.1 Effects of S3307 on growth parameters of
coix seedlings under PEG stress
Exogenous S3307 may be involved in the regu-
lation of coix seedling response to PEG stress. The
plant height, root length, stem diameter and dry
weight of above-ground and blow-ground biomass of
stressed plants largely reduced, whereas S3307 addi-
tion groups improved some growth parameters under
PEG stress. Compared with S0, the plant stem diame-
ter in S3 was increased by 16.19% (P＜0.05), and
the above- ground and blow- ground biomasses in-
creased by 19.56% and 22.75% (P＜0.05), respec-
tively. There is no significant difference between S3
and CK in stem diameter, above- ground and blow-
ground biomass (P>0.05). (Table 1).
Coix seedling growth situation of different treat-
ment groups were seen in Figure 1. The coix seed-
lings were growing well in CK (no wilting phenome-
non); There was an obvious wilting phenomenon in
S0, while the wilting phenomenon had been alleviat-
ed in S3 (Figure 1).
2.2 Effects of S3307 on antioxidant enzymes acti⁃
vities in coix seedlings under PEG stress
SOD, POD and CAT are the major antioxidant
enzymes associated with scavenging the reactive oxy-
gen species (ROS). POD, CAT and SOD activities
were increased under PEG stress. After S3307 addi-
tion, the CAT, SOD and POD activities in coix leafs
and roots exhibited the same trend as shown in Fig-
ure 2. The SOD, POD and CAT activity in leafs in-
creased by 10.46% , 18.16% and 80.56% , and those
in roots increased by 5.54% , 3.96% and 50.96% in
S3 compared with S0, respectively (P＜0.05). With
the increasing of the S3307 concentration, the antiox-
idant enzymes activities showed the trend of first in-
creasing then decreasing. Final results showed that
antioxidant enzymatic activities of coix seedling
could remarkably improve under 9 mg/L S3307.
Effects of different treatment time on the SOD,
POD and CAT activities of coix seedlings under
drought stress were shown in Figure 3. There is the
same tendency of first rising, then decline for SOD,
POD and CAT activity with stress time. The SOD
and CAT activity reached maximum at 24 h, while
POD at 48 h. The activity of SOD, POD and CAT in
leafs at 96 h increased by 16.93% , 9.26% and
49.45% , respectively, and increased by 9.3% ,
80.20% and 56.76% in roots at 96 h, respectively,
which showed SOD, POD and CAT activities of coix
seedlings presented increasing trend at 96 h.
2.3 Effect of S3307 on chlorophyll content in
coix leafs under PEG stress
It is considered that chlorophyll is the extremely
Table 1 Effects of exogenous S3307 on the growth of coix seedlings under drought stress
Treatment
CK
S0
S1
S2
S3
S4
S5
Plant height/cm
27.29±0.55a
24.37±0.69b
24.25±0.52b
24.09±0.52b
23.43±1.23b
22.05±1.45b
22.35±0.69b
Stem diameter/mm
0.430±0.014ab
0.389±0.013b
0.407±0.004b
0.420±0.009ab
0.452±0.016a
0.425±0.008a
0.434±0.012ab
Root length/cm
23.56±1.28a
22.54±1.24a
22.39±1.10a
21.95±1.21a
21.87±1.16a
21.78±1.12a
21.45±1.46a
Above-ground biomass
(10 plants) /mg
1.3340±0.0590a
1.1182±0.0655b
1.2035±0.0227ab
1.2557±0.0540a
1.3369±0.0208a
1.3027±0.0342a
1.3233±0.0129a
Blow-ground biomass
(10 plants) /mg
1.2489±0.1184a
1.0159±0.0679b
1.0814±0.0453ab
1.1021±0.0577ab
1.2470±0.0726a
1.1769±0.0271a
1.1832±0.0413ab
CK: Control group; S0: Drought group; S1, S2, S3, S4, S5: PEG+3, 6, 9, 12, 15 mg/L uniconazile (S3307), respectively. Different
letters indicate significant difference (P＜ 0.05). The same below
CK S0 S3
Figure 1 Effects of different treatment group on coix
seedling growth situation under drought stress
234
important pigment for photosynthesis in green
plants, and photosynthesis can improve and grana
and grana lamella numbers in plant can increase un-
der higher content of chlorophyll (Anderson et al.,
1973). The content of chlorophyll b in S3 was in-
creased by 11.37% compared with S0 (P＜0.05).
And total chlorophyll content in S3 was the highest
in all treatments, which was increased by 7.35% com-
pared with S0 (P＜0.05) (Figure 4). These results
showed that photosynthesis of coix seedlings could
improve under S3307 by promoting chlorophyll syn-
thesis.
2.4 Leaf and root ultrastructure in coix see⁃
dlings under PEG stress
Chloroplasts are the sites of plant photosynthe-
sis and chlorophyll existence. The shape of chloro-
plast was ellipse and granum lamella structure was
clearly visible, which had evident billayer structure
and there was no plasmolysis (Figure 5 A~C).
Drought stress decreased the water level in the cells
and caused serious plasmolysis, and the chloroplast
shape was deformed and misty, and lamellae became
invisible, meanwhile there were some plastoglobules
under drought stress (Figure 5 D~F). Mesophyll chlo-
roplasts were still close to the plasma membrane and
regularly arranged under S3307, which had no too
much difference between S3 and S0, which were
some substantial starch grains (Figure 5 G~I).
The effect of exogenous S3307 on root ultra-
structure in coix seedlings under drought stress were
showed in Figure 6. The root cell structure and cyto-
membrane were intact. There was no plasmolysis
phenomenon and mitochondria form plumpness, and
structure was clear. The nucleus had a double mem-
brane and clear nucleoli (Figure 6 A~C). There was
an evident alteration for PEG groups in root cell,
which presented obvious plasmolysis, and the mito-
chondria structure was vague. The nucleoli could not
be observed in nucleus and part karyotheca was deg-
radated (Figure 6 D~F). The mitochondria in root
cell returned to normal and plasmolysis phenomenon
had been relieved in S3; The nucleus membrane
structure was integrity and nucleolar was clear, and
there was an even distribution in nucleus chromatin
(Figure 6 G~I).
3 Discussion
Drought is the major adverse environmental
threat, which not only affects plant growth and devel-
opment but also has evolved series of physiological
and biochemical strategies (Wang, Li, 2007). S3307
can alleviate various abiotic and biotic stresses as re-
viewed, including dwarfing and thicker roots and
shoots, above-ground and blow-ground biomass and
so on (Mei et al., 2014; Sean et al., 2014), and Simi-
lar effects were also achieved in this experiment. The
stem diameter, above- ground and blow- ground bio-
Figure 2 Effects of different concentration uniconazile on
the SOD (A), POD (B) and CAT (C) activities of coix seed⁃
lings under drought stress
SO
D
ac
tiv
ity
/(U
·g-
1 FW
) 300
250
200
150
100
50
0 CK S0 S1 S2 S3 S4 S5
Group
Leaf
Root
c
c
c
ab b b a
a a a a a ab
A
CK S0 S1 S2 S3 S4 S5
Group
Leaf
Root
PO
D
ac
tiv
ity
/(U
·g-
1 FW
) 300
250
200
150
100
50
0
c
b
a
b
a
b
a
b
a
b
a
b
a
b
B
CK S0 S1 S2 S3 S4 S5
Group
Leaf
Root
C
AT
ac
tiv
ity
/(U
·g-
1 FW
) 14
12
10
8
6
4
2
0
f e f
e
d
d
c
a
b
a
b b
c
C
烯效唑(S3307)对干旱胁迫下薏苡幼苗生长及生理特性的影响
The Growth and Physiololgical Characteristics of Uniconazole (S3307) on Coix Seedlings Under Drought Stress 235
农业生物技术学报
Journal of Agricultural Biotechnology
mass in S3 group were greater than drought group in
the experiment. Therefore, it is reasonable to hypoth-
esize that uniconazole could alleviate drought stress
in coix seedlings. SOD, CAT and POD are the major
antioxidant enzymes in plants (Allen, 1995; Zhu et
al., 2004), which can efficiently clear the ROS pro-
duction in plants, such as O2-, H2O2, and OH1- and so
on. In the present experiment, drought stress en-
hanced the activities of POD, SOD and CAT, which
could induce the accumulation of O 2- and H2O2,
while the antioxidant enzymes activities will appear
remarkable decline at 72 h. Compared with the
drought stress, application of S3307 further in-
creased the activities of antioxidant enzymes, which
protected the plant from oxidative damage that was
caused by drought stress, and the SOD, POD and
CAT activity was still enhanced at 96 h. It was simi-
lar to early work in wheat (Triticum aestivum) (Duan
et al., 2008), marine microalgae (Mei et al., 2014)
and soybean (Giycine max)(Zhang et al., 2007) under
drought stress.
Chlorophyll is extremely significant for the ab-
sorption, conversion and utilization of light energy.
Many studies found that environment stress had sig-
nificant effect on chlorophyll content, among which
drought stress could decrease the biosynthesis of
chlorophyll and increase the formation of ROS,
which accelerated the breakdown of chlorophyll
(Yordanov et al., 2000; Agastian et al., 2000). In this
study, S3307 could increase chlorophyll content,
which might be related with S3307 enhancing endog-
enous abscisic acid (ABA) content. While, exoge-
nous ABA performs a positive function in Plasma
source ion implantation (PSII) under water stress
conditions (Kai et al., 2014). Chloroplasts are ex-
Figure 3 Effects of different treatment time on the POD, SOD and CAT activities of coix leaf and root under drought
stress
Figure 4 Effects of exogenous uniconazile on chlorophyll
concentration of coix seedlings under drought stress
236
tremely sites of plant photosynthesis and chlorophyll
existence. Some stressful conditions will lead to the
disruption of chloroplast structure, furthermore dam-
age the photosynthesis processes in plant (Yu et al.,
2005). Similar changes took place in this study. Un-
der PEG stress, serous plasmolysis and the chloro-
plast shape was deformed, misty and lamellae be-
came invisible, meanwhile there were some cosmio-
philic particles under drought stress. S3307 could
lighten the damage to grana ultrastructure, improve
the integrity of chloroplasts membrane structure, and
the reason might be S3307 could enhance antioxi-
dant enzymes, decrease H2O2 and membrane lipid
peroxide to a certain extent (Kai et al., 2014). Re-
searchers had reported that most ABA was high cor-
related with biosynthetic pathway genes and photo-
synthesis (Kilian et al., 2007; Pinheiro, Chaves,
2011). Exogenous ABA treatment was reported to in-
duce the up-regulation of most chloroplast and allevi-
ate the drought damage (Kai et al., 2014), which
might be the mechanism of S3307 improving photo-
synthesis and chloroplast ultrastructure. Meanwhile,
S3307 could also relieve the damage of root cell ul-
trastructure under PEG stress, and the reason might
be that S3307 enhanced antioxidant enzymes and de-
creased membrane lipid peroxide. Especially, S3307
had a positive effect on the stability of memebrane
system and could relieve the injury extent for meme-
brane system, mainly reducing MDA content, which
has been proved in other researches (Kilian et al.,
2007; Pinheiro, Chaves, 2011).
4 Conclusion
To explore the effects of S3307 on coix seed-
lings under drought stress, some growth parameters
and physiological indexes were investigated under
drought stress. The results showed that the stem di-
ameter and ground biomass, the activities of POD,
CAT, SOD and total chlorophyll content were signifi-
cantly improved in 9 mg/L S3307 group. Meanwhile,
S3307 could also lighten the damage to grana ultra-
structure under drought stress. All the analysis
showed that 9 mg/L S3307 was effective on alleviat-
ing the injury of coix seedlings under drought stress.
5 μm 1 μm 200 nm
5 μm 1 μm 100 nm
5 μm 5 μm 5 μm
A B C
D E F
G H I
Figure 5 Effect of exogenous S3307 on leaf ultrastructure in coix seedlings under drought stress
A~C：CK；D~F：S0；G~I：S3; S: Starch grains; PM: Plasmalemma; Pg: Plastoglobules.
烯效唑(S3307)对干旱胁迫下薏苡幼苗生长及生理特性的影响
The Growth and Physiololgical Characteristics of Uniconazole (S3307) on Coix Seedlings Under Drought Stress 237
农业生物技术学报
Journal of Agricultural Biotechnology
This study can clarify physiological mechanism of
S3307 relieving drought stress and provide a theoreti-
cal basis for S3307 in coix production and practice
of drought resistance.
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Figure 6 Effect of exogenous S3307 on root ultrastructure in coix seedlings under drought stress
A~C：CK；D~ F：S0；G~I：S3; Mi: Mitochondria; PM: Plasmalemma; N: Nucleu
238
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(责任编辑 靳晓霞)
烯效唑(S3307)对干旱胁迫下薏苡幼苗生长及生理特性的影响
The Growth and Physiololgical Characteristics of Uniconazole (S3307) on Coix Seedlings Under Drought Stress 239