全 文 ：干旱胁迫对 5个楸树无性系渗
透调节物质的影响
董 蕾 1，李吉跃 1*，王军辉 2，谢 堃 1，苏 艳 1
（1.华南农业大学林学院，广东广州 510642；
2.中国林业科学研究院林业研究所，国家林业
局林木培育重点实验室，北京 100091）
摘 要 [目的]了解干旱胁迫对楸树（Catalpa
bungei） 无性系叶片渗透调节物质的影响。 [方
法]采用盆栽塑料薄膜封盆法，通过土壤含水量
与叶水势变化对干旱胁迫程度进行划分，并研
究 5个楸树无性系在不同干旱程度下叶片渗透
调节物质 （游离脯氨酸、 可溶性糖及可溶性蛋
白）含量的变化。 [结果]通过土壤含水量与叶水
势的变化，将干旱胁迫过程分为正常（CK）、轻
度（LD）、中度（MD）和重度（SD）干旱，其区间为
正常干旱 （土壤含水量 97.49%， 叶水势-0.54
MPa）、轻度干旱（土壤含水量 59.96%，叶水势-
1.28MPa）、中度干旱（土壤含水量 34.19%，叶水
势-2.32 MPa）、重度干旱（土壤含水量 14.52%，
叶水势-2.99 MPa）；5 个楸树无性系土壤含水
量与叶水势拟合关系均以指数拟合最佳，平均
R2达到 0.989 3 （P<0.001）；5 个楸树无性系叶
片游离脯氨酸含量随干旱胁迫程度的加剧显
著上升（P<0.01），其中无性系 015-1 和无性系
7080在 SD 下分别达到正常状态的 34.39 倍和
33.41 倍，达极显著水平（P<0.001）；无性系 1-3
则在 LD 时快速上升，迅速达（855.46±227.52）
μg/g Fw； 叶片可溶性蛋白含量表现出不同趋
势，无性系 7080 在各阶段均最低，正常状态仅
为（1.644±0.137） mg/g Fw，而无性系 1-3在 CK
时可溶性蛋白含量较高，在 LD时迅速下降，体
现出不同的响应模式。 [结论]5个楸树无性系叶
片渗透调节物质对干旱胁迫均有一定的响应，
以游离脯氨酸为最主要渗调物质，可溶性蛋白
为辅，未见可溶性糖有显著贡献；其中无性系
7080 在各渗透调节物质含量变化上表现出较
高的能力，体现出较好的抗旱性，而无性系 1-3
则对于干旱胁迫响应最迅速。
关键词 干旱胁迫，楸树无性系，渗透调节物质
基金项目 “十二五”农村领域国家科技计划课
题（2012BAD21B03；2012BAD21B0304）。
作者简介 董蕾 (1984-)，女，天津人，助理研
究员， 博士， 主要从事楸树水分生理研究，E-
mail：icefox1984@aliyun.com。 * 通讯作者，李吉
跃（1959-），男，四川金堂人 ，教授，博士生导
师。主要从事森林培育及栽培生理生态方面研
究，E-mail：ljyymy@vip.sina.com。
收稿日期 2013-08-02
修回日期 2013-09-02
W ith global climate changes,drought has become a criti-cal factor limiting the sus-
tainable development of agriculture
and forestry worldwide [1-2]. In China,
drought is the major constraint affect-
ing plant growth and development and
population regeneration in Northwest
China [3], which has significant impact
on physiological and ecological pro-
cesses of plants. Drought stress leads
to water metabolic imbalance and im-
paired physiological processes of
plants, and plants can adapt to water
stress through a series of physiological
and biochemical changes. Specifically,
osmotic regulation is an important re-
sponse mechanism. Under stress
conditions, plants increase intracellular
solute concentration and reduce os-
Effects of Drought Stress on Osmotic Regulation
Substances of Five Catalpa bungei Clones
Lei DONG1, Jiyue LI1*, Junhui WANG2, Kun XIE1, Yan SU1
1.College of Forestry, South China Agricultural University, Guangzhou 510642, China;
2. Institute of Forestry, Chinese Academy of Forestry, Beijing 100091, China
Supported by National Scientific and Technological Project in Rural Areas in the Twelfth
Five Year Plan (2012BAD21B03, 2012BAD21B0304).
*Corresponding author. E-mail: ljyymy@vip.sina.com
Received: August 2, 2013 Accepted: September 2, 2013A
Agricultural Science & Technology, 2013, 14(9): 1335-1343
Copyright訫 2013, Information Institute of HAAS. All rights reserved Resources and Environment
Abstract [Objective] This study aimed to investigate the effects of drought stress on
osmotic regulation substances of Catalpa bungei clones. [Method] Potting experi-
mental was carried out with plastic film sealing method, to differentiate the degrees
of drought stress based on changes in soil water content and leaf water potential
and investigate variations in contents of osmotic regulation substances (free proline,
soluble sugar and soluble protein) in leaves of five Catalpa bungei clones under dif-
ferent degrees of drought stresses. [Result] According to changes in soil water con-
tent and leaf water potential, the whole process of drought stress was divided into
normal level (CK, with soil water content of 97.49% and leaf water potential of -0.54
MPa), light drought (LD with soil water content of 59.96% and leaf water potential of
-1.28 MPa), mediate drought (MD with soil water content of 34.19% and leaf water
potential of -2.32 MPa) and severe drought (SD with soil water content of 14.52%
and leaf water potential of -2.99 MPa). The soil water content and leaf water po-
tential of five Catalpa bungei clones reached the highest correlation in exponential
fitting, with an average R2 of 0.989 3(P<0.001). The free proline content in leaves of
five Catalpa bungei clones increased rapidly with the increasing degree of drought
stress (P<0.001). To be specific, free proline contents of Catalpa bungei clones
015 -1 and 7080 were 34.39 and 33.41 times of the normal level under severe
drought conditions, which reached an extremely significant level (P<0.001); the free
proline content of Catalpa bungei clone 1-3 rapidly increased to (855.46±227.52) μg/g
Fw under light drought conditions. The soluble protein content in leaves showed dif-
ferent variation trends. To be specific, the soluble protein content of Catalpa bungei
clone 7080 was the lowest at various drought stages and reached (1.644±0.137)
mg/g Fw under normal conditions; the soluble protein content of Catalpa bungei
clone 1-3 was relatively high under normal conditions and was reduced rapidly un-
der light drought conditions, showing different response patterns. [Conclusion] Osmot-
ic regulation substances in leaves of five Catalpa bungei clones all have certain re-
sponse to drought stress. Free proline is the most important osmotic regulation sub-
stance, followed by soluble protein, while soluble sugar makes no significant contri-
bution. According to changes in content of osmotic regulation substances, Catalpa
bungei clone 7080 has relatively high capacity and good drought resistance, while
Catalpa bungei clone 1-3 has the rapidest response to drought stress.
Key words Drought stress; Catalpa bungei clones; Osmotic regulation substances
DOI:10.16175/j.cnki.1009-4229.2013.09.030
Agricultural Science & Technology 2013
Table 1 Growth statuses of five Catalpa bungei clones
Clones Seedling height∥cm Ground diameter∥mm Whole plant leaf area∥m2
004-1 135.92± 21.63 17.02± 0.97 1.990 7±0.032
015-1 118.47± 3.18 14.83± 0.82 1.780 0±0.051
7080 141.00± 3.56 15.10± 1.16 1.489 2±0.071
1-3 144.00± 4.55 15.44± 0.86 1.120 2±0.036
1-4 133.32± 7.41 15.74± 0.81 1.260 9±0.048
motic potential through metabolic ac-
tivities to decline the water potential
and continue to absorb water, thus
maintaining turgor pressure and nor-
mal physiological metabolic activities[4].
Numerous studies have confirmed that
the content of osmotic regulation sub-
stances of plants under water stress is
positively correlated with the drought
resistance[5-7]. To be specific, K+, Cl-,
Ca2+ and Mg2+ are the main inorganic
substances; soluble sugar, soluble
protein and free proline are the main
organic substances. In addition, a
large number of studies have com-
pared the drought resistance of differ-
ent plants based on the content of os-
motic regulation substances. For in-
stance, Xu et al.[8] compared the chang-
es in osmotic regulation substances of
Robinia pseudoacacia, Acer truncatum
Bunge and Platycladus orientalis un-
der water stress and found that the os-
motic regulation mechanisms of differ-
ent tree species are different. Reyizha
et al. [9] investigated osmotic regulation
substances of different varieties of
black currant and reported that Shi-
jiguang has the highest drought resis-
tance. Liu et al.[10] analyzed the drought
resistance of Rosmarinus officinalis
based on the physiological response
of the release of volatile organic com-
pounds and osmotic regulation under
drought stress.
Catalpa bungei is a precious high-
quality timber species and famous or-
namental tree species cultivated tradi-
tionally in warm temperate and sub-
tropical regions of China[11-12], which is
loved by the masses due to its excel-
lent material quality and beautiful tree
form[13-14]. In recent years, many schol-
ars have investigated main biologi-
cal characteristics and the situation
of Catalpa bungei germplasm re-
sources [15-16]. Studies of Catalpa bungei
clones are mostly focused on differ-
ences in response to salt stress be-
tween various clones[17], growth status
after fertilization[18] and relationship be-
tween chlorophyll fluorescence and
growth [19], etc . In addition, studies of
water physiology of Catalpa bungei
are mostly focused on comparisons of
photosynthetic changes between dif-
ferent varieties [20]. However, few re-
searches have been reported on
changes in osmotic regulation sub-
stances between different Catalpa
bungei clones under drought stress [21].
In this study, different Catalpa bungei
clones were used for potting experi-
ments under drought conditions, to in-
vestigate the osmotic regulation
mechanism of Catalpa bungei under
drought stress and compare the
drought resistance of different Catalpa
bungei clones according to changes in
water metabolism and osmotic regula-
tion substances of five Catalpa bungei
clones with different drought resis-
tance screened by Chen et al.[21] under
different water gradients, which provid-
ed reference for the selection of af-
forestation tree species in Northwest
China.
Materials and Methods
Status of experimental plot
The experimental plot is located at
105° 48′ E, 34° 29′ N, in the green-
house of Xiaolongshan Forestry Re-
search Institute in Tianshui City of
Gansu Province, with an altitude of
about 1 450 m. The forestry centre be-
longs to temperate semi-humid mon-
soon climate with an average annual
rainfall of about 600-800 mm, average
annual evaporation of about 1 290
mm, average temperature of about 11
℃ , extreme maximum temperature of
39 ℃ , extreme minimum temperature
of -19.2 ℃ and frost-free period of
about 180 d[22].
Experimental materials
Catalpa bungei belongs to Catal-
pa of Bignoniaceae, with height of 20-
30 m, diameter at breast height of 1-2
m, small branch opening angles,
dense crowns, grey-brown barks, shal-
low longitudinal cracks, purple-brown
and smooth twigs, opposite or ternate-
whorled leaves, corymbose raceme,
bisexual flowers, sharp tops, flower-
ing period from April to May, fruit
ripening period from August to
September, elongated capsules[15].
Experimental materials were col-
lected from the nursery of Xiaolong-
shan Forestry Research Institute in
Tianshui City of Gansu Province. Fifty
two-year-old plantlets of five Catalpa
bungei clones (004-1, 1-3, 1-4, 7080
and 015-1) with relatively consistent
growth were selected, and the growth
statuses were shown in Table 1. In
early March 2012, Catalpa bungei
seedlings were transplanted into 35
cm×35cm×30cm (bottomdiameter×
upper diameter × height) pots, one
seedling per plot. In order to prevent
the loss of water and fertilizer, plastic
trays were adopted for each pot, and
the pots were sheathed with double-
deck white plastic bags. The matrix
was collected from Xiaolongshan for-
est seed base of Gansu Province, with
a volume ratio of peat ∶forest soil ∶fowl
dung = 2∶7∶1. The physical and chemi-
cal nature of matrix was: pH =6.20,
containing 74.44 g/kg organic matter,
3.03 g/kg total nitrogen, 1.47 g/kg total
phosphorus, 20.29 g/kg total potassi-
um, 325.36 g/kg alkali-hydrolyzable
nitrogen, 131.88 g/kg available phos-
phorus and 372.94 g/kg rapidly avail-
able phosphorus. The bulk density
was 0.96 g/cm3; field moisture capacity
was (46.59±4.05)％; total porosity was
63.64% . Each plot was filled with
about 20 kg of matrix. The fertilization
experiment was carried out from
seedling recovering period to late May.
Experimental methods
At the beginning (day 0), early pe-
riod (day 3, day 6), medium period
(day 8, day 10) and late period (day
13, day 18) of the experiment, the light
intensity, temperature and humidity of
the greenhouse were measured using
hand-held weather station (Kestrel ∶
14500, U.S.) and Li∶cor 6400 (Licor, U.
S.) every 2 h since 8:00 in the morn-
ing. During the experiment process,
the average levels of environmental
factors were shown in Fig.1.
Normal watering management
was conducted for experimental
1336
Agricultural Science & Technology2013
seedlings. The physiological and bio-
chemical indicators of Catalpa bungei
seedlings were measured in typical
sunny day. Pots were sealed with plas-
tic film (sheathed from the bottom to
the top and sealed at the bottom of the
stem), to prevent water loss. At differ-
ent periods of drought, the following
indicators were measured (each indi-
cator was measured three times; to be
specific, soil water content and leaf
water potential were measured on day
0, day 2, day 4, day 6, day 10, day 13
and day 18; osmotic regulation sub-
stances were sampled destructively on
day 0, day 4, day 7, day 11 and day
18):
1) Soil water content:
Three pots of Catalpa bungei
seedlings of each clone with consis-
tent growth were selected and marked.
Soil water content at the depth of 5-6
cm was determined using high-preci-
sion EM50 soil moisture meter
(Decagon, US), to obtain the volumet-
ric water content, thereby calculating
the mass water content according to
the soil bulk density in accordance with
the following equation:
Soil mass water content (% ) =
Soil volumetric water content (%) / Soil
bulk density ×100%
2) Leaf water potential:
Six pots of Catalpa bungei
seedlings of each clone with consis-
tent growth were selected and marked.
Leaf water potential was determined
using ARIMAD 3000 plant pressure
chamber detector (MRC, Israel) before
dawn.
3) Free proline content:
Free proline content was mea-
sured using acid ninhydrin method[23].
4) Soluble protein content:
Soluble protein content was mea-
sured using Kaumas blue method[23].
5) Soluble sugar content:
Soluble sugar content was mea-
sured using anthrone-sulfuric acid
method[23].
Results and Analysis
Differentiation of drought degrees
Under water stress conditions,
water potential is one of the most sen-
sitive indicators reflecting water stress
in plants[24], which can be used to de-
scribe the stress level and compare
the drought resistance of plants. In
soil-plant-atmospheric circulation sys-
tems (SPAC), soil water content and
leaf water potential can be adopted as
a continuum to reflect the water loss or
water condition of plants [25]. Numerous
studies have indicated that soil water
content and leaf water potential are
very closely correlated; leaf water po-
tential is reduced with the decrease of
soil water content[26]. Analysis of vari-
ance showed that leaf water potentials
of different Catalpa bungei clones var-
ied significantly during the process of
drought stress. Under normal water
conditions, leaf water potentials of five
Catalpa bungei clones had no signifi-
cant difference; with the increase of
drought stress, leaf water potentials of
five Catalpa bungei clones were all
significantly reduced, but differences
between various clones declined;
when Catalpa bungei seedlings died
under drought stress, leaf water po-
tentials of various Catalpa bungei
clones varied significantly. Throughout
the process of drought stress, leaf
water potential of Catalpa bungei
clone 7 080 declined insignificantly un-
der light drought, decreased signifi-
cantly under mediate drought, and
reached the minimum level (-3.18 ±
0.32 MPa) when seedlings died under
drought stress, which was significantly
lower than that of other four Catalpa
bungei clones, indicating that Catalpa
bungei clone 7 080 can rapidly reduce
the leaf water potential under drought
stress to regulate the seedling water
potential and soil water potential gradi-
ent, thereby reducing the ability of wa-
ter loss, suggesting relatively high
drought tolerance.
During the process of water
stress, soil water content and leaf wa-
ter potential of five Catalpa bungei
clones were both significantly reduced
with the increase of drought stress
(Table 2). Under normal water condi-
tions, the average soil water content of
five Catalpa bungei clones was
44.54%±0.52%, with no significant dif-
ference between various Catalpa
bungei clones; with the extension of
water stress duration, soil water con-
tent was significantly reduced; to be
specific, soil water content of Catalpa
bungei clone 004-1 declined rapidly
and greatly, indicating that transpira-
tion water consumption of Catalpa
bungei clone 004-1 is higher than that
of other four clones, which may be due
to that Catalpa bungei clone 004-1 has
the maximum leaf area (Table 1).
According to the percentage of
soil mass water content accounting for
field moisture capacity and changes in
leaf water potential of Catalpa bungei
clones at different periods of drought,
the whole drought stress period can be
divided into four stages (Table 2). Day
0 was set as the control (CK), Catalpa
bungei seedlings were not under
stress, soil water content accounted
for about 96.89%±1.22% of field mois-
ture capacity, and leaf water potential
ranged from -0.51 to -0.56 MPa; light
drought stage (LD) lasted from the be-
ginning of drought stress to day 4, soil
water content accounted for about
70% of field moisture capacity, and
leaf water potential ranged from -1.48
to -20.20 MPa; mediate drought stage
(MD) lasted from day 5 to day 10, soil
water content and leaf water potential
were both reduced; severe drought
stage (SD) lasted from day 11 to the
death of plants (day 18), leaf water po-
tential reached -3.09±0.14 MPa, while
soil water content accounted for only
about 15% of field moisture capacity,
with high degree of drought stress, and
Catalpa bungei seedlings could not
Fig.1 Daily change of environmental factors
1337
Agricultural Science & Technology 2013
Table 2 Soil water content and leaf water potential of five Catalpa bungei clones under
different drought gradients
Drought stress
duration∥d Clones
Mass water
soil
content∥%
Water soil
content
rate∥%
Leaf water
potential∥
MPa
Drought gradient
0 004-1 45.56 99.71 -0.56 Normal (CK)
015-1 44.83 98.12 -0.52
7080 43.01 94.13 -0.55
1-3 44.31 96.98 -0.56
1-4 45.01 98.51 -0.51
2 004-1 30.33 66.38 -0.93 Light drought (LD)
015-1 35.59 77.89 -0.90
7080 33.58 73.50 -0.66
1-3 32.54 71.22 -0.72
1-4 32.19 70.45 -0.91
4 004-1 20.20 44.21 -2.02
015-1 20.76 45.43 -1.76
7080 25.98 56.87 -1.48
1-3 21.40 46.83 -1.75
1-4 21.37 46.78 -1.63
6 004-1 9.21 32.15 -2.16 Mediate drought (MD)
015-1 9.56 33.93 -2.29
7080 11.54 35.26 -1.94
1-3 9.57 32.96 -2.55
1-4 9.52 31.84 -2.42
10 004-1 8.73 29.11 -2.63
015-1 8.24 28.04 -2.39
7080 8.08 27.69 -2.36
1-3 7.21 25.79 -2.29
1-4 7.10 25.53 -2.20
13 004-1 6.18 13.53 -3.07 Severe drought (SD)
015-1 6.91 15.12 -2.76
7080 7.33 16.05 -2.71
1-3 7.28 15.94 -3.04
1-4 7.28 15.94 -2.89
18 004-1 5.92 12.96 -3.06
015-1 6.47 14.15 -3.00
7080 6.29 13.77 -3.18
1-3 6.51 14.25 -3.09
1-4 6.18 13.53 -3.14
survive but died.
With soil water content as the in-
dependent variable x, leaf water po-
tential as the dependent variable y,
logarithmic, exponential and power
functions were fit. Results showed that
three fitting methods could all achieve
good results for Catalpa bungei
clones, with correlation coefficient of
above 0.9 (P<0.1); comparisons be-
tween three fitting methods indicated
that five Catalpa bungei clones
showed higher correlation in exponen-
tial fitting (P<0.05), followed by loga-
rithmic fitting, while five Catalpa bungei
clones showed the lowest correlation
in power fitting (Table 3). However,
comparative analysis of all the experi-
mental data of five Catalpa bungei
clones showed that exponential rela-
tionship led to the best fitting result,
with correlation coefficient of 0.989 3
(P < 0.001) (Fig.2).
By using exponential equations of
each clone in Table 3, leaf water po-
tential was calculated with the average
soil mass water content under differ-
ent degrees of drought stresses as
the independent variable, which was
shown in Table 4. According to the
calculation results of fitting equations,
Catalpa bungei clone 7080 constantly
maintained the minimum level of leaf
water potential. Compared with the
actual operation, fitting equations can
better reflect the ability of Catalpa
bungei clone 7080 to regulate leaf
water potential, with better explana-
tion. However, adjusted values of leaf
water potential of other four Catalpa
bungei clones under various soil
moisture conditions had no significant
difference.
Changes in osmotic regulation
substances with the degree of
drought stress
Changes in free proline content
Under arid environments, the rapid in-
crease of free proline content in leaves
is resulted from the metabolic regula-
tion of plant leaves under drought
stress[27]. As shown in Fig.3, free pro-
line content in leaves of five Catalpa
bungei clones increased rapidly, with
differences between various clones.
Under normal water conditions, the
free proline level is relatively low of
(207.38 ± 111.5) μg/g Fw averagely,
with no significant difference between
five Catalpa bungei clones. Under light
drought stress conditions, the free
proline content in leaves of five Catal-
pa bungei clones increased significant-
ly; to be specific, the free proline con-
tent of Catalpa bungei clone 1-3 was
significantly higher than that of other
four clones, which reached (855.46±
227.52) μg/g Fw; free proline contents
of Catalpa bungei clones 7080 and 1-
4 were the lowest with no significant
Fig.2 Exponential fitting equation between
leaf water potential and soil water
content of five Catalpa bungei clones
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Agricultural Science & Technology2013
Table 4 Variance analysis of leaf water potential of Catalpa bungei clones under drought stress
Leaf water potential
under drought stress
(MPa)
Clones
Fitting equation
CK (soil water
content of 44.54%)
LD (soil water
content of 27.39%)
MD (soil water
content of 8.88%)
SD (soil water
content of 6.64%)
004-1 y = 3.818 3e-0.042x -0.588 -1.209 -2.630 -2.889
015-1 y =3.683 5e-0.042x -0.567 -1.166 -2.537 -2.787
7080 y = 3.736e-0.0345x -0.804 -1.452 -2.750 -2.971
1-3 y =3.892e-0.045x -0.524 -1.135 -2.610 -2.887
1-4 y = 3.714 4e-0.043x -0.598 -1.208 -2.581 -2.829
Table 3 Correlation between leaf water potential and soil water content of different Catalpa
bungei clones
Clones Fitting equation Correlation coefficient (R 2) Fitting pattern
004-1 y=-1.186ln(x) + 5.147 0.940 Logarithmic
y=3.818 3e-0.042x 0.962 Exponential
y=13.013x-0.765 0.900 Power
015-1 y=-1.143ln(x) + 4.976 8 0.973 Logarithmic
y=3.683 5e-0.042x 0.982 Exponential
y=13.182x-0.777 0.917 Power
7080 y=-1.228ln(x) + 5.154 0.948 Logarithmic
y=3.736e-0.0345x 0.947 Exponential
y=14.668x-0.833 0.917 Power
1-3 y=-1.247ln(x) + 5.28 0.923 Logarithmic
y=3.892e-0.045x 0.955 Exponential
y=15.129x-0.833 0.908 Power
1-4 y=-1.171ln(x) + 5.03 0.928 Logarithmic
y=3.714 4e-0.043x 0.973 Exponential
y=13.196x-0.787 0.910 Power
difference. As the drought intensifies,
the free proline content in leaves in-
creased significantly. Under mediate
drought stress conditions, the free pro-
line level of Catalpa bungei clone 015-
1 was significantly higher than that of
other four clones , which reached
(3 948.17 ± 221.02) μg/g Fw; under
light drought stress, Catalpa bungei
clone 1-3 had the highest free proline
content and the lowest increasing rate
that was enhanced by only 1.85 times
than before. Under severe drought
stress to death, the average free pro-
line content in leaves of various
Catalpa bungei clones reached
(6427.89 ± 301.19) μg/g Fw. To be
specific, free proline contents of Catal-
pa bungei clones 7080 and 015-1 had
no significant difference, which
reached (6 835.14 ±148.00) μg/g Fw
and (7 028.84 ±311.73) μg/g Fw, re-
spectively; free proline contents of
Catalpa bungei clones 1-4 and 004-1
were the lowest, with no significant dif-
ference.
Changes in soluble sugar content
As an osmotic regulation substance of
plants, soluble sugar has been con-
sidered an important physiological in-
dicator of the response of plants to ad-
versity stress. Increasing soluble sugar
content can decline the osmotic po-
tential of plants, which is conducive to
maintaining normal moisture require-
ments of plants under stress. Accord-
ing to changes in soluble sugar con-
tent of five Catalpa bungei clones un-
der different degrees of drought
stresses (Fig.4), the variation of solu-
ble sugar content of five Catalpa
bungei clones during the whole pro-
cess of drought could be divided into
three trends: ① continuously increas-
ing: soluble sugar contents of Catalpa
bungei clones 7080, 1 -4 and 015 -1
constantly increased with the increas-
ing degree of drought stress; specifi-
cally, the soluble sugar content in
leaves of Catalpa bungei clone 7080
under normal water conditions was sig-
nificantly lower than that of other four
Catalpa bungei clones (P<0.05), which
increased rapidly with the increasing
degree of drought stress. ② maintain-
ing stable after reducing: the soluble
sugar content in leaves of Catalpa
bungei clone 1-3 was higher than that
of four Catalpa bungei clones (with no
significant difference) under normal
water conditions and was significantly
lower than that of four Catalpa bungei
clones under drought conditions. ③
Severely declining: the soluble sugar
content in leaves of Catalpa bungei
clone 004-1 increased with the in-
creasing degree of drought stress but
significantly declined under severe
drought conditions.
Changes in soluble protein content
Plants under stress will produce stress
proteins through different metabolic
processes and physiological changes,
to eliminate, repair or recover the
damages caused by environmental
stresses [28 ] . Most of the soluble pro -
teins in plants are enzymes involving
Fig.3 Changed in free proline content of five Catalpa bungei clones
1339
Agricultural Science & Technology 2013
in various metabolic processes.
Therefore, determining changes in
soluble protein content under drought
stress is conducive to understanding
plant resistance and total metabolism.
As shown in Fig.5, soluble protein con-
tents in leaves of Catalpa bungei
clones 1-3 and 004-1 was reduced un-
der light drought conditions and in-
creased with the increasing degrees of
drought stresses, while soluble protein
contents in leaves of Catalpa bungei
clones 7080, 015-1 and 1-4 increased
constantly with the increasing degrees
of drought stresses from (1.937 ±
0.121) mg/g Fw under normal condi-
tions to (4.270±0.351) mg/g Fw (aver-
age of three clones). Overall, the solu-
ble protein content in leaves of Catalpa
bungei clone 7080 was significantly
lower than that of other clones at dif-
ferent drought stages; the soluble pro-
tein content in leaves of Catalpa
bungei clone 1-3 was significantly
higher than that of other clones under
normal conditions, which could rapidly
reduced to a low level that was con-
sistent with other clones under drought
stress; under severe drought condi-
tions, soluble protein contents of
Catalpa bungei clones 1-4, 015-1 and
004-1 were relatively high, with an av-
erage of (4.711±0.294) mg/g Fw (with
no difference between various clones),
which was significantly higher than
that of Catalpa bungei clones 1-3 and
7080.
Conclusion and Discussion
Correlation between soil water con-
tent and leaf water potential of five
Catalpa bungei clones
Predawn leaf water potential rep-
resents the water recovery status of
plants, which can be used to degree of
water stress in plants[29-30]. Compared
with water potentials in other tissues,
leaf water potential can not only reflect
the constraint degrees of various
physiological activities of plants in
growing seasons under different envi-
ronmental water conditions, but also
directly show the water status in vari-
ous tissues of plants[31]. In addition, a
large number of studies have been
carried out on the relationship between
soil water content and leaf water po-
tential. Guo et al.[32] adopted piecewise
fitting method and found that the effect
of changes in soil water content on leaf
water potential of seedlings in the
morning had a threshold, the rela-
tionship between soil water content
and leaf water potential could be de-
scribed by a linear equation when soil
water content was greater than the
threshold and described by a expo-
nential equation when soil water con-
tent was lower than the threshold. In
this study, leaf water potential and soil
water content of five Catalpa bungei
clones during the process of drought
stress showed an exponential rela-
tionship, and the average level also
reached the highest correlation in ex-
ponential fitting (R2 = 0.989 3, P <
0.001), indicating that the result of ex-
ponential fitting in Catalpa bungei is
the best, which is consistent with the
research of Chen et al. [21] but different
from other tree species. He et al. [33 -34]
reported that Chinese white poplar
reaches the best result with logarith-
mic fitting, while Chinese scholartree
reaches the best result with binomial
relationship. According to these com-
parison results, different fitting rela-
tionships between leaf water potential
and soil water content of plants may
be unique characteristics of different
tree species and reflect their drought
tolerance, which has great significance
for the identification of intraspecific and
interspecific drought resistance and
water conservation.
By using separate fitting method,
the exponential fitting equation of leaf
water potential and soil water content
of five Catalpa bungei clones was ob-
tained (Table 3). According to the re-
sults of correction calculation based on
the average level of soil water content
under different degrees of drought
stresses (Table 4), it is clearly found
that the leaf water potential of Catalpa
bungei clone 7080 is significantly lower
than that of other four clones under dif-
ferent levels of soil water content. The
level of leaf water potential can repre-
sent the ability of plants to absorb wa-
ter from the soil or neighboring cells,
and plants under drought stress can
effectively prevent water loss into the
soil by reducing leaf water potential,
which is also reported by Li et al.[35] and
Li et al. [36]. Among the five Catalpa
bungei clones in this study, 7080
shows higher leaf water potential and
drought regulation ability than other
four clones.
Comparison between osmotic reg-
ulation substances and drought
tolerance of five Catalpa bungei
clones
Numerous studies have showed
that damages of plants under drought
stress are closely related to the
Fig.4 Changed in soluble sugar content of five Catalpa bungei clones
Fig.5 Changed in soluble protein content of five Catalpa bungei clones
1340
Agricultural Science & Technology2013
changes in osmotic regulation sub-
stances. Greatly accumulating osmotic
regulation substances can increase
water retention capacity of plants and
stabilize membrane system, thereby
resisting the drought stress injury[37-38].
Therefore, the determination of dy-
namic changes in contents of soluble
sugar, soluble protein and free proline
under drought stress is conducive to
effectively investigating differences in
drought resistance ability and drought
response mechanism between various
clones.
Free proline contents of five
Catalpa bungei clones increase rapidly
with the increasing degree of drought
stress, indicating certain drought re-
sponse in different approaches. A
large number of studies have shown
that free proline contents of plants un-
der drought stress vary significantly.
Reyizha et al. [9] compared and ana-
lyzed various black currant clones and
found that under little and moderate
drought stresses, the free proline con-
tent of black currant reaches 1.9 and
4.7 times of that under normal condi-
tions, respectively. Xu et al. [8] com-
pared and analyzed Robinia pseu-
doacacia, Acer truncatum Bunge and
Platycladus orientalis and found that
the free proline content of Robinia
pseudoacacia under severe drought
stress can reach 10 times of normal
level, while free proline contents of Ac-
er truncatum Bunge and Platycladus
orientalis are reduced. Compared to
other species, Catalpa bungei clones
have higher free proline regulation
ability. To be specific, proline contents
in leaves of Catalpa bungei clones
015-1 and 7080 died of drought are
34.39 and 34.39 times higher than the
normal level (with no significant differ-
ence), showing the most significant
changes among five Catalpa bungei
clones, which suggests that proline
regulation is the major osmotic regula-
tion pattern of Catalpa bungei clones
015-1 and 7080 under drought stress.
However, the proline content of Catal-
pabungei clone 1-3 under light drought
increases the rapidest and greatest, in-
dicating that Catalpa bungei clone 1-3
has the rapidest response to drought
stress.
Under drought stress conditions,
the osmotic regulation of soluble sugar
and free proline in plant leaves is a
mutual compensation strategy [ 39 - 40 ] .
Among five Catalpa bungei clones in
this study, the soluble sugar content in
leaves of Catalpa bungei clone 7080
under normal conditions is significantly
lower than that of other clones, which
rapidly increases to an insignificant
level under drought stress, suggesting
that Catalpa bungei clone 7080 has
rapid response to drought and great
osmotic regulation ability. However,
soluble sugar contents of Catalpa
bungei clones 1-3, 1-4 and 015-1
have not reached significant level,
which is consistent with the significant-
ly increasing free proline contents of
these three clones, indicating that their
osmotic regulation abilities may not be
realized by soluble sugar but free pro-
line. Many studies have reported the
effects of soluble sugar in drought
stress. For instance, Wang et al. [41] in-
vestigated osmotic regulation sub-
stances of four tree species and found
that the soluble sugar content of Os-
tryopsis davidiana has the most signifi-
cant response to drought stress, while
soluble sugar contents of Liaodong
oak and vetchleaf sophora show no
significant change under drought
stress. Similarly, Xu et al. [8] also re-
ported that the increase level of solu-
ble sugar content of Robinia pseudoa-
cacia under drought stress is not high,
which is different from the response
mechanisms of Acer truncatum Bunge
and Platycladus orientalis. These re-
sults are similar to the present study,
suggesting that different species have
different response mechanisms of os-
motic regulation under drought stress.
The accumulation of reactive oxy-
gen species in plants caused by water
stress will lead to lipid peroxidation and
destroyed enzymes, nucleic acids and
other molecules, thereby inducing the
production of stress resistant proteins
to maintain a low osmotic potential of
plant cells and avoid damages caused
by water stress. Therefore, the level of
soluble protein content can be used as
a comparative indicator of plant
drought resistance[42-43]. Tong et al.[3] in-
vestigated Populus diversifolia under
drought stress and found that the sol-
uble protein content increases first and
then declined with the increasing de-
gree of drought stress. Wang et al. [41]
found that the soluble protein content
of Acer stenolobum Rehd.var. mega-
lophyllum declined first and then in-
creased with the increasing degree of
drought stress. Reyizha et al.[9] investi-
gated changes in the soluble protein
content of black currant under drought
stress and found that the soluble pro-
tein content of black currant was re-
duced constantly. These studies show
that changes in soluble protein con-
tents in leaves between different
species or different clones are differ-
ent, which might be related with the
different response patterns of various
species. In this study, soluble protein
contents of five Catalpa bungei clones
show an overall increasing trend, sug-
gesting that under drought stress,
Catalpa bungei produces stress resis-
tant proteins to avoid water stress in-
jury; however, the soluble protein con-
tent of Catalpa bungei clone 7080 is
significantly lower than that of other
four clones during the process of
drought stress, suggesting that the os-
motic regulation function of Catalpa
bungei clone 7080 is not controlled by
soluble protein; the soluble protein
content of Catalpa bungei clone 1-3 is
significantly higher than that of other
clones under normal conditions and is
rapidly reduced under light drought
stress; based on the free proline con-
tent, it can be found that Catalpa
bungei clone 1-3 is rapidly responsive
under light drought stress, and its os-
motic regulation is mainly controlled by
changes in the soluble protein content.
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