Journal of Plant Physiology 224–225 (2018) 30–48

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Journal of Plant Physiology

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The compensation effects of physiology and yield in cotton after drought T

stress
Jing Niu1, Siping Zhang1, Shaodong Liu, Huijuan Ma, Jing Chen, Qian Shen, Changwei Ge,
⁎ ⁎
Xiaomeng Zhang, Chaoyou Pang , Xinhua Zhao
Institute of Cotton Research of CAAS/State Key Laboratory of Cotton Biology, Anyang, Henan 455000, China

A R T I C LE I N FO A B S T R A C T

Keywords: The objective of this study was to investigate the root growth compensatory effects and cotton yield under
Cotton drought stress. The results indicate that the root dry weight, boll weight, and cotton yield increased in both the
Drought drought-resistant cultivar (CCRI-45) and the drought-sensitive cultivar (CCRI-60). Compensation effects were
Compensation exhibited under the three-day drought stress treatment at a soil relative water content (SRWC) of 60% and 45%
Physiology
during the seedling stage, and flowering and boll-forming stage over two years. The yield of the drought-re-
Yield
sistant cultivar (CCRI-45) was higher than the control, however, following the six-day 45% SRWC drought
treatments, the yield of the drought-sensitive cultivar (CCRI-60) was lower than the control. The soluble sugar
content, proline content, superoxide dismutase (SOD) activity, and peroxidase (POD) activity of the roots in-
creased under drought stress and then decreased after re-watering, although the values remained higher than
those of the controls for a short period. These physiological measures may represent stress reactions and thus
may not indicate factors that result in compensation effects. However, catalase (CAT) activity and gibberellic
acid (GA) content of the roots decreased under drought stress. After re-watering, the CAT activity and the GA
content increased and were significantly higher than those of the controls under the six-day 60% SRWC and 45%
SRWC drought treatments. The abscisic acid (ABA) content of the roots increased under drought stress. After re-
watering, the ABA content decreased to a lower level under the three and six-day 60% SRWC and 45% SRWC
drought treatments than in the controls. According to an analysis of various indicators, the interaction between
ABA and GA signals may play an important role in root growth compensatory effects. In summary, the results
demonstrate that moderate drought stress is beneficial to root growth and yield. This conclusion is of great
significance to improving our understanding of the maximum utilization of limited water resources.

1. Introduction high-yield agriculture has become a priority.

With global climate change, the causes and impacts of frequently
occurring extreme meteorological events have drawn increasing
worldwide attention (Trenberth et al., 2015; Powell and Reinhard,
2016). Drought, which is caused by dramatically reduced precipitation,
is an extreme meteorological event that greatly affects agriculture
production and social economic development, and the frequency,
duration, and spatial extent of droughts have exhibited increasing
trends (Wilhite et al., 2014). Cotton is the world’s most important
textile fiber crop, and it provides the majority of the fiber used in the
textile industry. However, cotton yield is hindered by water and salt
stress when the plant is grown in arid and semi-arid areas. Therefore, to
exploit limited water resources fully, maximizing high-efficiency and
Since the early 1970s, scholars have been exploring the compen-
sation effects of crops under stress. Acevedo et al. (1971) found that
leaves undergo a fast growing period that fully compensates for losses
during drought stress, and the biomass of plants that were exposed to
short-term and moderate drought stress was higher than that of the
controls. Since then, multiple studies on different crops have been
performed, and the results have shown that crops generally exhibit a
compensation effect after moderate drought stress (Kramer, 1989;
Adalsteinsson, 1994). Martin and Jamieson (1996) drew the conclusion
that the total dry weight decreased under drought stress before flow-
ering. However, after re-watering, the grain yield increased because the
single grain weight and harvest index were enhanced. The results of
that study showed that the transshipment of the carbon library to grain

Abbreviations: SRWC, soil relative water content; SOD, superoxide dismutase; POD, peroxidase; CAT, catalase; GA, gibberellic acid; ABA, abscisic acid
⁎
Corresponding authors.
E-mail addresses: niujing890321@163.com, 1259637820@qq.com (J. Niu), zhsp5337@163.com (S. Zhang), chypang@163.com (C. Pang), zhaoxinhua1968@126.com (X. Zhao).
1
These authors contributed equally to this study.

https://doi.org/10.1016/j.jplph.2018.03.001
Received 14 November 2017; Received in revised form 2 March 2018; Accepted 2 March 2018
Available online 09 March 2018
0176-1617/ © 2018 The Authors. Published by Elsevier GmbH. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/BY-NC-ND/4.0/).
J. Niu et al.
increased significantly under a moderate water deficit during the filling
stage, thereby increasing production (Foulkes et al., 2007). Yang et al.
(2001) and Hsiao et al. (2007) also found similar results. Additionally,
cotton is often considered a drought-resistant crop and exhibits infinite
growth. The root, leaf, bud, and flower can be grown repeatedly with
re-watering to compensate for injuries under drought stress. Studies
have shown that the cotton yield and fiber strength of certain cotton
cultivars were improved when the irrigation volume was moderately
decreased (Dağdelen et al., 2009; Papastylianou and Argyrokastritis,
2014).
The root is an integral plant organ that is involved in the acquisition
of nutrients and water; the synthesis of plant hormones, organic acids,
and amino acids; and the anchorage of plants (Chloupek et al., 2010;
Středa et al., 2012). Root morphology and physiology are closely as-
sociated with the growth and development of aboveground plant ma-
terial. The root is the initial part affected by soil drought, and its
morphological and physiological characteristics are closely correlated
with plant drought resistance (Gajri et al., 1989). Therefore, studying
the compensation effect of cotton roots is of great significance to cotton
production and water-saving irrigation. The aims of this study were to
(i) clarify the compensation effects of cotton root dry weight, yield,
quality, and root physiological indices and (ii) identify the mechanisms
underlying the root compensation effects.
2. Materials and methods
2.1. Treatment, experimental design, and management
Field experiments were conducted at the experimental farm of the
Institute of Cotton Research, Chinese Academy of Agricultural Sciences,
Anyang, Henan Province, China (36°06′N, 114°21′E) during the cotton
growing seasons of 2014 and 2015. The chemical properties of the field
soil were measured for two years (attached Table A1), and the me-
teorological data are shown in the attached Table A2.
The cultivars CCRI-45 and CCRI-60 were planted in plastic pots
(33 cm × 706.5 cm2) filled with 27 kg of sandy loam soil, with five
plants per pot in the seedling stage (for a total of 216 pots) and one
plant per pot in the flowering and boll-forming period (for a total of 270
pots). The experiment was designed for the seedling stage, and flow-
ering and boll-forming stage. Three water treatments were imposed on
the plants: 75% ( ± 5%) of field capacity (control, CK), 60% ( ± 5%) of
field capacity (mild drought stress, D1), and 45% ( ± 5%) of field ca-
pacity (moderate drought stress, D2). The maximum amount of water
the soil can retain by capillary action is known as the field capacity
(FC). Changes in the soil relative water content (SRWC) during the
experiments are shown in Fig. A1. Drought stress lasted for three days,
six days, and nine days for each treatment, and then all plants under-
went re-watering.
In both years, the experiment was arranged in a completely ran-
domized design. Cotton was grown in a plastic greenhouse to prevent
exposure to natural rainfall, and the cultivation management methods
were consistent (except the water treatments). The SRWC was de-
termined by weighing the soil samples, which were collected every two
days. The soil fresh weight (FW) was recorded, the samples were dried
to a constant weight, and then the final dry soil weight (DW) was re-
corded. The water mass content of the soil (in percentage terms) was
equal to (FW − DW)/DW × 100, and the SRWC was equal to SRWC
(%) = SWC/FC × 100. The SRWC was maintained within the pre-
scribed levels during the experiment.
2.2. Determination of the root biomass, agronomic performance, yield, and
fiber quality of cotton
Traditional methods were used to obtain all the roots, which were
rinsed with water after blistering for two hours. The roots were weighed
after drying to a constant weight at 80 °C. The plant height, the number
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Journal of Plant Physiology 224–225 (2018) 30–48
of fruiting branch nodes, and the bolls per plant were counted from the
tagged plants, and the mean values were measured and expressed as the
number per plant. The weights of fully opened and matured bolls
picked from ten plants were recorded, and the mean boll weight was
calculated to estimate the yield of cotton. The fiber length, uniformity
index, micronaire, elongation rate, and fiber strength were measured by
the Ministry of Agriculture on Cotton Quality Inspection Center.
2.3. Determining the proline content, soluble sugar content, soluble protein
content, and antioxidant enzyme activity
The proline content was determined according to the Bates method
(1973). Dry roots (0.15 g) were homogenized in 5 mL of 3% sulfosa-
licylic acid at 100 °C for 20 min in a water bath and then filtrated. The
filtrate (2 mL) was mixed with acid-ninhydrin (2 mL) and glacial acetic
acid (2 mL) in a test tube. The reaction mixture was incubated in a
water bath at 100 °C for 30 min, cooled to room temperature, and then
extracted with toluene (5 mL). The toluene, which contains chromo-
phores, was aspirated, and the absorbance was then measured at
520 nm. The soluble sugar content of the roots was determined using
the anthrone-sulfuric acid method (Yemm and Willis, 1954). The dry
roots (0.1 g) were homogenized in 5 mL of 80% ethanol at 80 °C for
30 min in a water bath, and then the supernatant was collected. The
above steps were repeated twice. The filtrate (2 mL) was mixed with
anthrone-sulfuric acid (2 mL) in ice water, and the reaction mixture was
then incubated in a water bath for 10 min at 100 °C and then cooled to
room temperature. Afterwards, the absorbance was measured at
620 nm.
Fresh root tissues (0.3 g) were ground with 5 mL of 50 mM phos-
phate buffer (pH 7.0), which was prepared by mixing NaH2 PO4·H2O
and Na2HPO4·7H2O and then centrifuging at 12000g for 20 min at 4 °C.
Then, the supernatant was collected to measure the antioxidant enzyme
activity and the soluble protein content. The superoxide dismutase
(SOD) activity was determined using reagent kits (obtained from the
Nanjing Jiancheng Bioengineering Institute, Nanjing, Jiangsu Province,
China, WST-1method). The peroxidase (POD) and catalase (CAT) ac-
tivities were measured using the guaiacol method and the ultraviolet
absorption method (Li et al., 2000a). The soluble protein content was
determined from the same enzyme extract using Brilliant Blue G-250
dye (Bradford, 1976).
2.4. Determining the ABA and GA content
Approximately 0.5 g of fresh sample was collected, and the en-
dogenous ABA and GA were extracted according to a previously re-
ported method (Yang et al., 2001) using an enzyme-linked im-
munosorbent assay (ELISA). The ELISA kits were manufactured by the
Phytohormones Research Institute, China Agricultural University.
2.5. Data analysis
The results were analyzed via analyses of variance (ANOVAs) using
Statistical Analysis System 8.1. The means were analyzed using the
least significant difference (LSD) method at P = 0.05 (LSD 0.05).
3. Results
3.1. Effect of drought stress on the cotton yield and fiber quality
The yield of cotton increased under the three-day and six-day
drought treatments (Tables 1, A3 and A4). In both years, the yields of
two cultivars were higher than those of the controls under the three and
six-day 60% SRWC and 45% SRWC drought treatments. During the
seedling period, the highest yields of CCRI-45 (the drought-tolerant
cultivar) were observed under the six-day 60% SRWC drought treat-
ments in 2014 and 2015, exhibiting yields 27.27% and 22.34% higher
J. Niu et al. Journal of Plant Physiology 224–225 (2018) 30–48

Table 1
Changes in the cotton yield and yield composition under drought stress after re-watering during the seedling stage in 2015.

Varieties

CCRI-45 CCRI-60

Treatment Boll number per plant Boll weight (g) Lint (%) Lint cotton yield (kg Hm−2) Boll number per plant Boll weight (g) Lint (%) Lint cotton yield (kg Hm−2)

CK 19.67a 5.59bc 0.39a 2122.19b 20.00a 5.36bc 0.39a 2042.01ab

D1L3 20.33a 5.54bc 0.41a 2261.34ab 21.00a 5.43bc 0.39a 2215.98ab
D2L3 21.33a 5.87ab 0.40a 2486.77ab 21.33a 5.59ab 0.39a 2334.53a
D1L6 21.00a 6.43a 0.39a 2596.32a 20.00a 6.09a 0.40a 2420.99a
D2L6 20.33a 5.37bc 0.42a 2269.53ab 16.33b 5.57ab 0.39a 1744.72bc
D1L9 16.00b 5.37bc 0.38a 1634.99c 12.33c 5.20bc 0.38a 1195.04c
D2L9 12.00c 5.16c 0.38a 1188.51c 13.00c 4.85c 0.38a 1184.29c

Note: CK: SRWC (75 ± 5)%; D1: SRWC (60 ± 5)%; D2: SRWC (45 ± 5)%; L3: drought stress for three days; L6: drought stress for six days; and L9: drought stress for nine days. In the
columns, different letters indicate significant differences at the 0.05 probability level.

Table 2
Changes in the final agronomic traits of cotton under drought stress after re-watering during the seedling stage in 2015.

Varieties

CCRI-45 CCRI-60

Treatment Height (cm) Fruiting branch nodes Fruiting positions Shedding rate Height (cm) Fruiting branch nodes Fruiting positions Shedding rate
number number (%) number number (%)

CK 96.33a 14.33ab 55.00a 64.21c 113ab 13.67a 58.33a 65.69bc

D1L3 99.00a 15.67a 54.00a 62.30cd 115.33ab 14.00a 57.67ab 63.62bc
D2L3 102.67a 14.33ab 56.33a 62.12cd 117.67a 13.67a 58.67a 63.47bc
D1L6 104.33a 14.67ab 54.00a 61.11d 118.33a 13.33a 53.00c 62.13c
D2L6 98.67a 12.00bc 56.00a 63.68cd 110.00b 12.00ab 53.67bc 69.55ab
D1L9 82.33b 12.00bc 51.00a 68.62c 113ab 13.67a 58.33a 65.69bc
D2L9 78.67b 9.67c 48.67a 74.80a 115.33ab 14.00a 57.67ab 63.62bc

Note: CK: SRWC (75 ± 5)%; D1: SRWC (60 ± 5)%; D2: SRWC (45 ± 5)%; L3: drought stress for three days; L6: drought stress for six days; and L9: drought stress for nine days. In the
columns, different letters indicate significant differences at the 0.05 probability level.

Table 3
Changes in the cotton fiber quality under drought stress after re-watering during the seedling stage in 2015.

Varieties

CCRI-45 CCRI-60

Treatment Fiber length Uniformity index Micronaire Elongation rate Fiber strength Fiber length Uniformity index Micronaire Elongation rate Fiber strength
(mm) (%) (%) (cN/tex) (mm) (%) (%) (cN/tex)

CK 29.47a 83.73a 5.30a 6.70a 28.80c 28.03a 83.23a 5.67a 6.60a 27.63ab
D1L3 29.73a 85.90a 5.17a 6.73a 28.93bc 28.43a 84.20a 5.30a 6.60a 28.87a
D2L3 29.53a 84.90a 5.27a 6.67a 30.30ab 28.57a 84.37a 5.43a 6.60a 28.80a
D1L6 29.90a 86.20a 5.43a 6.70a 30.70a 28.97a 84.73a 5.63a 6.63a 29.30a
D2L6 29.40a 85.20a 5.40a 6.67a 31.70a 28.33a 84.33a 5.73a 6.60a 27.57ab
D1L9 29.93a 83.20a 5.60a 6.70a 27.60cd 27.83a 82.87a 5.50a 6.60a 25.97bc
D2L9 29.33a 82.07a 5.57a 6.67a 26.77d 27.57a 82.37a 5.57a 6.57a 24.90c

Note: CK: SRWC (75 ± 5)%; D1: SRWC (60 ± 5)%; D2: SRWC (45 ± 5)%; L3: drought stress for three days; L6: drought stress for six days; and L9: drought stress for nine days. In the
columns, different letters indicate significant differences at the 0.05 probability level.

than those of the control, respectively (Tables 1 and A3). The yields of
CCRI-60 (the drought-sensitive cultivar) were highest under the three-
day 45% SRWC and six-day 60% SRWC treatments in 2014 and 2015,
exhibiting yields 18.92% and 18.56% higher than those of the control,
respectively (Tables 1 and A3). Compared with CCRI-60, the yield and
the increment of CCRI-45 were higher under the six-day 60% SRWC
treatments. The increase in production was primarily related to the
increase in the boll weight, which was measured by analyzing the yield
component indices (Tables 2, A5 and A6). In both the drought-tolerant
and the drought-sensitive cultivars, the nine-day 60% SRWC and 45%
SRWC treatments resulted in significantly lower yields compared to
those of the controls. The elongation rate of CCRI-45 increased under
the three and six-day 60% SRWC and 45% SRWC treatments, but de-
creased under the nine-day 60% SRWC and 45% SRWC treatments. The
elongation rate of CCRI-60 increased under the three-day 60% SRWC
and 45% SRWC treatments and the six-day 60% SRWC treatments, but
it decreased under the six-day 60% SRWC treatments and the nine-day
60% SRWC and 45% SRWC treatments (Tables 3, A7 and A8).
3.2. Effects of drought stress on cotton roots
The increase in yield may have been caused primarily by the com-
pensation effects of the root. Therefore, we further analyzed the
changes in the root dry weights under drought stress and re-watering.
As shown in Figs. 1, A2 and A3, the root dry weight of the two varieties
increased under the three and six-day 60% SRWC and 45% SRWC
treatments, although the root dry weights of CCRI-60 were lower than
those of the controls under the six-day 45% SRWC droughts in 2014.

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J. Niu et al. Journal of Plant Physiology 224–225 (2018) 30–48

Fig. 1. Changes in the cotton root dry weight under drought stress and re-watering during the seedling stage in 2015.
Note: L3: drought stress for three days; L3-5: fifth day after re-watering following three-day drought stress; L3-10: tenth day after re-watering following three-day drought stress; L6:
drought stress for six days; L6-5: fifth day after re-watering following six-day drought stress; L6-10: tenth day after re-watering following six-day drought stress; L9: drought stress for nine
days; L9-5: fifth day after re-watering following nine-day drought stress; and L9-10: tenth day after re-watering following nine-day drought stress. In the columns, different letters indicate
significant differences at the 0.05 probability level.

The root dry weights of the two varieties decreased significantly under
the nine-day 60% SRWC and 45% SRWC treatments during both years.
After re-watering, the roots grew rapidly and the root dry weight in-
creased. On the tenth day after re-watering, the root dry weights of both
varieties were highest under the three-day 45% SRWC and six-day 60%
SRWC treatments during the seedling stage in 2014 and 2015. Com-
pared with the root dry weight of the controls, that of CCRI-45 in-
creased by 31.21% and 19.77% (Figs. 1 and A2), and the root dry
weight of CCRI-60 increased by 9.36% and 20.13% in 2014 and 2015,
respectively (Figs. 1 and A2). Although the root dry weights increased
under the nine-day 60% SRWC and 45% SRWC treatments, the dry
weights were still lower than those of the controls (Figs. 1, A2 and A3).
3.3. Changes in the root physiology under drought stress
The physiological characteristics of the roots are closely correlated
with the root growth and drought resistance of the plant. As the degree
and duration of drought stress increased, the soluble sugar content
(Figs. 3, A6 and A7), proline content (Figs. 3, A6 and A8), and SOD
(Figs. 5, A11 and A12) and POD (Figs. 5, A11 and A13) activities of
both varieties of cotton increased. After re-watering, these values de-
creased, although they were still higher than those of the controls
during the two stages. During the seedling period, the soluble protein
content of the cotton root decreased under the three and nine-day 60%
SRWC and 45% SRWC treatments, but they increased under the six-day
60% SRWC and 45% SRWC treatments. After re-watering, the soluble
protein contents returned to normal levels or higher levels than those of
the controls (Figs. 4, A9 and A10).
The CAT activity showed an opposite trend relative to that of the
SOD and POD activities. After re-watering, the CAT activity of the
cotton root increased, thereby demonstrating the compensation effect.
The CAT activity of cotton roots was higher than that of the controls
under the three and six-day 60% SRWC and 45% SRWC treatments.
However, under the nine-day drought stress, the value remained lower
than that of the controls (Figs. 6, A14 and A15). Compared with the
CAT activity of the controls, that of CCRI-45 plants was highest under
the six-day 60% SRWC drought treatments in 2014 and 2015, in-
creasing by 14.65% and 45.63%, respectively. The CAT activity of
CCRI-60 was highest under the three day and six-day 60% SRWC
treatments in 2014 and 2015, increasing by 13.12% and 26.48%, re-
spectively (Figs. 6 and A14). Additionally, the CAT activity increments
of CCRI-45 were higher than those of CCRI-60 under the six-day 60%
SRWC drought treatments.
As the degree and duration of drought stress increased, the ABA
content of the roots increased. After re-watering, the ABA content de-
creased (Figs. 7, A16 and A17). On the tenth day after re-watering, the
ABA content returned to normal levels or lower levels than in the
controls. Compared with the ABA content of the controls, those of CCRI-
45 decreased by 13.31% and 14.48% under the six-day 60% SRWC
treatments in 2014 and 2015, respectively, and the ABA contents of
CCRI-60 decreased by 19.90% and 15.63% under the three-day 45%
SRWC and six-day 60% SRWC treatments (Figs. 7 and A16), respec-
tively, during the seedling period in 2014 and 2015.
However, the GA content of the roots increased under the three-day
60% SRWC and 45% SRWC treatments and then decreased under the
six and nine-day 60% SRWC and 45% SRWC treatments. After re-wa-
tering, the GA content increased (Figs. 7, A16 and A18). On the tenth
day after re-watering, the GA content of the roots was significantly

Fig. 2. Relationship between the increased percentages of yield and root dry weight under drought stress after re-watering during the seeding stage in 2015.
Note: * and ** indicate significant differences at 0.05 and 0.01 probability levels, respectively (n = 6, R0.05 = 0.811, and R0.01 = 0.917).

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J. Niu et al. Journal of Plant Physiology 224–225 (2018) 30–48

Fig. 3. Changes in the soluble sugar and proline content in cotton roots under drought stress and re-watering during the seedling stage in 2015.
Note: L3: drought stress for three days; L3-5: fifth day after re-watering following three-day drought stress; L3-10: tenth day after re-watering following three-day drought stress; L6:
drought stress for six days; L6-5: fifth day after re-watering following six-day drought stress; L6-10: tenth day after re-watering following six-day drought stress; L9: drought stress for nine
days; L9-5: fifth day after re-watering following nine-day drought stress; and L9-10: tenth day after re-watering following nine-day drought stress. Changes in the soluble sugar content
2015: A and B; changes in the proline content in 2015: C and D. In the columns, different letters indicate significant differences at the 0.05 probability level.

Fig. 4. Changes in the soluble protein content in cotton roots under drought stress and re-watering during the seedling stage in 2015.
Note: L3: drought stress for three days; L3-5: fifth day after re-watering following three-day drought stress; L3-10: tenth day after re-watering following three-day drought stress; L6:
drought stress for six days; L6-5: fifth day after re-watering following six-day drought stress; L6-10: tenth day after re-watering following six-day drought stress; L9: drought stress for nine
days; L9-5: fifth day after re-watering following nine-day drought stress; and L9-10: tenth day after re-watering following nine-day drought stress. In the columns, different letters indicate
significant differences at the 0.05 probability level.

higher than that of the controls under the three and six-day 60% SRWC
and 45% SRWC treatments, but it was still significantly lower than that
of the controls under the nine-day 60% SRWC and 45% SRWC treat-
ments. The GA content of CCRI-45 was the highest under the six-day
60% SRWC drought stress in 2014 and 2015 and, compared with the
controls, increasing by 49.68% and 38.95%, respectively. The GA
content of CCRI-60 was the highest under the three-day 45% SRWC and
the six-day 60% SRWC treatments in 2014 and 2015, increasing by
45.50% and 32.37% (Figs. 7 and A16), respectively, during the seedling
period in 2014 and 2015. The GA increments of CCRI-45 were higher
than those of CCRI-60 under the six-day 60% SRWC drought.
4. Discussion
In recent years, the frequency and degree of drought have been on
the rise. Cotton often encounters long-term drought stress, which ser-
iously affects the cotton yield. However, there is a positive regulation
mechanism that can compensate the effect of drought stress after re-
watering to some extent. In this paper, we systematically studied the
response of each physiological index of two cultivars under drought
stress, as well as the intensity of the compensation effect and the re-
sponse of yield and quality after re-watering at the seedling stage, and
flowering and boll-forming stage. The study showed that soluble sugar,
proline, SOD, POD, and ABA showed an increasing trend under the
drought stress, while GA and CAT decreased. After re-watering, the
recovery of all the physiological indices and the root dry weight and
yield was comparable with the control.
However, the resilience of cotton is different under different degrees
of drought stress, showing an overcompensation effect, equal com-
pensation effect, and partial compensation effect. The study found that
the changes in CAT, GA, and ABA were closely related to cotton yield
after re-watering. The results provide some theoretical basis for
drought-tolerant cultivation and water-saving irrigation, and explain
the drought-resistant mechanism of cotton and the compensation effect.
4.1. Moderate drought stress can promote cotton growth
In this study, the data showed that the boll number and yield were
higher than those of the controls under the three and six-day 60%
SRWC and 45% SRWC treatments during the seedling stage, and

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Fig. 5. Changes in the SOD and POD activity in cotton root under drought stress and re-watering during the seedling stage in 2015.
Note: L3: drought stress for three days; L3-5: fifth day after re-watering following three-day drought stress; L3-10: tenth day after re-watering following three-day drought stress; L6:
drought stress for six days; L6-5: fifth day after re-watering following six-day drought stress; L6-10: tenth day after re-watering following six-day drought stress; L9: drought stress for nine
days; L9-5: fifth day after re-watering following nine-day drought stress; and L9-10: tenth day after re-watering following nine-day drought stress. Changes in the SOD activity in 2015: A
and B; changes in the POD activity in 2015: C and D. In the columns, different letters indicate significant differences at the 0.05 probability level.

Fig. 6. Changes in the CAT activity in cotton roots under drought stress and re-watering during the seedling stage in 2015.
Note: L3: drought stress for three days; L3-5: fifth day after re-watering following three-day drought stress; L3-10: tenth day after re-watering following three-day drought stress; L6:
drought stress for six days; L6-5: fifth day after re-watering following six-day drought stress; L6-10: tenth day after re-watering following six-day drought stress; L9: drought stress for nine
days; L9-5: fifth day after re-watering following nine-day drought stress; and L9-10: tenth day after re-watering following nine-day drought stress. In the columns, different letters indicate
significant differences at the 0.05 probability level.

flowering and boll-forming stage, and the drought-tolerant cultivar
(CCRI-45) showed a greater compensation effect than the drought-
sensitive cultivar (CCRI-60). These findings have also been gained in
previous studies: the cotton yields of some cotton cultivars improved
when the drip irrigation volume was moderately reduced during the
growth stage (Wanjura et al., 2002; Ertek and Kanber, 2003; Kang et al.,
2012), and the fiber quality decreased under an irrigation deficit
(Dağdelen et al., 2009; Ünlü et al., 2011). However, possibly due to the
long-term drought, the boll number and yield decreased under the nine-
day 60% and 45% SRWC treatments, which is similar to the findings of
a previous study (Wang et al., 2013). The above findings proved that a
certain range of drought stress can promote the yield of cotton. They
also showed that the intensity of the compensation effect varied in
cultivars. However, the cotton yield will suffer serious irreversible da-
mage when this drought range is exceeded.
Roots, as the main organs for water and nutrient uptake, play an
important role in the formation of plant drought resistance
(Ramamoorthy et al., 2017). Thus, we hypothesized that the increase in
yield may be caused largely by the compensation effects in the root.
Studies have found that the cotton root growth and root biomass in-
creased under moderate drought stress during the seedling stage (Li
et al., 2000b) and the budding stage (Yan et al., 2009); we also obtained
similar results. Studies have not been able to explain the changes oc-
curring in cotton root after re-watering. Therefore, we decided to focus
on these changes in our study. After re-watering, the root system had a
stronger water absorbing capacity, the roots grew rapidly, and the root
dry weights increased, especially under the six-day 60% SRWC drought
stress treatment. In addition, the increments of CCRI-45 (the drought-
resistant cultivar) were higher than those of CCRI-60 (the drought-
sensitive cultivar). In rice (Chu et al., 2014) and corn (Liang et al.,
2008; Qi et al., 2010) the root growth also accelerated, especially in the
lateral roots and root hairs, and the root biomass was significantly in-
creased after undergoing moderate drought. However, because the
cotton suffered irreversible damage after a drought of nine days, the
root dry weights were significantly lower after re-watering. These re-
sults indicate that moderate drought stress is beneficial to root growth,
and a strong root system not only improves the drought resistance but
also promotes the recovery of cotton growth.
An analysis of the yield and root dry weight showed that the yield
and root dry weight are closely correlated after re-watering (Figs. 2, A4
and A5), suggesting that the root dry weight might be used as an in-
dicator of the drought resistance of a crop.

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J. Niu et al. Journal of Plant Physiology 224–225 (2018) 30–48

Fig. 7. Changes in the ABA and GA content in cotton root under drought stress and re-watering during the seedling stage in 2015.
Note: L3: drought stress for three days; L3-5: fifth day after re-watering following three-day drought stress; L3-10: tenth day after re-watering following three-day drought stress; L6:
drought stress for six days; L6-5: fifth day after re-watering following six-day drought stress; L6-10: tenth day after re-watering following six-day drought stress; L9: drought stress for nine
days; L9-5: fifth day after re-watering following nine-day drought stress; and L9-10: tenth day after re-watering following nine-day drought stress. Changes in the ABA content in 2015 (A
and B); changes in the GA content in 2015 (C and D). In the columns, different letters indicate significant differences at the 0.05 probability level.

4.2. Close correlation between cotton growth compensation effects and ABA
and GA in roots after re-watering
When subjected to drought stress, the root system first senses and
changes its physiology to resist drought stress. The morphological
characteristics of the roots change in correspondence with the changes
in physiology. Our finding showed that soluble sugar, proline, SOD,
POD, and ABA exhibited increasing trends, while GA and CAT de-
creased under the drought stress, which is similar to the findings of
many previous studies (Jiang and Huang, 2001; Türkan et al., 2005;
Deeba et al., 2012; Sekmen et al., 2014; Shan et al., 2015; Devnarain
et al., 2016; Wang et al., 2016b). After re-watering, all the physiological
indices recovered. However, the intensity of the compensation effects
varied. Through our analysis, we speculate there are three physiological
indicators (ABA, GA, and CAT) that may be closely related to com-
pensation.
It is well known that endogenous plant hormones are important
signaling substances that regulate plant responses under stress. Plants
regulate the metabolism to adapt to or withstand stress by changing
various endogenous hormones. Previous studies have proven that the
increased ABA content under drought stress leads to adjustments in the
pore size, the protection of the structure and function of cell membrane,
the inhibition of plant growth, and a reduction in the metabolic rate,
thereby improving the drought resistance of plants (Zhang et al., 2006;
Kondo et al., 2012; Kowitcharoen et al., 2015; Wang et al., 2016a; Lv
et al., 2017). In this study, we focused on measuring the change in ABA
and its relationship to root compensation after re-watering. After re-
watering, the ABA content decreased and was even lower than that of
the controls under the three and six-day 60% SRWC and 45% SRWC
treatments, but was still higher than that of the controls under the nine-
day 60% SRWC and 45% SRWC treatments. In addition, the root dry
weights and yields were higher than those of the controls under the
three and six-day 60% SRWC and 45% SRWC treatments, but lower
than those of the controls under the nine-day 60% SRWC and 45%
SRWC droughts. These results suggest that ABA improvd the drought
resistance of cotton and was beneficial to the compensation effect. The
ABA contents of the cotton roots did not decrease on the sixth day after
re-watering during the flowering and boll-forming stages in Xinjiang,
which was likely caused by climate differences (Luo et al., 2013).
GA, as a hormone to promote plant growth, can promote cell
elongation and growth. According to their physiological functions, GA
levels are reduced under drought stress, improving drought resilience of
plants by reducing plant growth rate and reducing plant energy con-
sumption, and increased after re-watering. The increments of GA were
also higher under the moderate drought stress treatments than under
the severe drought stress (Luo et al., 2013; Yan et al., 2012). Wang et al.
(2012) and Albacete et al. (2014) also demonstrated that GA could
improve the adversity resistance abilities of plants. We found that, after
re-watering, the GA content of the roots increased and was higher than
the normal levels under the three and six-day 60% SRWC and 45%
SRWC treatments, which resulted in a large compensation effect.
However, the GA content was still significantly lower than the normal
levels under the nine-day 60% SRWC and 45% SRWC drought treat-
ments.
After re-watering, the ABA content was reduced, relieving the in-
hibition of cotton roots growth, and GA content increased, promoting
the root growth and making up for the damage caused by the drought.
We speculated that the growth of the roots and the final cotton yield
were improved under the mutual regulation of ABA and GA, providing
the compensation effect. Simultaneously, the osmotic regulation and
antioxidant enzymes were able to maintain the plant cell pressure and
eliminate ROS to protect the cells and provide the material basis and
energy for root growth (Fig. 8). CAT is an important antioxidant en-
zyme in the plant system that eliminates the excess H2O2 produced by
the action of the SOD, slowing plant peroxidation (Ozkur et al., 2009).
However, the CAT activity increased when SOD activity decreased after
re-watering following drought stress. Studies have found that the CAT
activity of the roots decreased under drought stress (Wu et al., 2016;
Wang et al., 2016b). After re-watering, the CAT activity increased and
was higher than that of the controls under the three and six-day 60%
SRWC and 45% SRWC treatments. In addition, the yields of cotton were
higher than those of the controls under the same treatments. However,
after re-watering, the CAT activity did not return to normal levels under
the nine day 60% SRWC and the 45% SRWC drought treatments, thus

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J. Niu et al. Journal of Plant Physiology 224–225 (2018) 30–48

Fig. 8. Response of cotton roots and yield to drought stress and re-watering.

leading to significantly lower values than those of the controls. This
finding shows that a high CAT activity may improve the root growth
and final yield. Nevertheless, the above conclusion still requires further
study to improve our understanding of the cotton compensation me-
chanism.
5. Conclusions
(CCRI-45) was stronger than that of the drought-sensitive cultivar
(CCRI-60), showing that the compensation ability varies in cultivars. In
addition, the compensation conditions were different for different
varieties and climates; therefore, the compensation effect mechanisms
of cotton require further investigation. This conclusion has important
significance for maximizing limited water resources and promoting
cotton cultivation in arid regions.

This study found that cotton root growth could be stimulated by Funding
moderate drought stress during the seedling stage, and flowering and
boll-forming stage (Appendix A), which results in the promotion of root This work was supported by Special Fund for Agro-scientific
growth that in turn prevents yield reductions or even leads to increased Research in the Public Interest (201503117).
yields. However, the yields were significantly reduced by long-duration
droughts. An analysis of the physiological indices, root dry weight, and Acknowledgments
yield showed that ABA, GA, and CAT play an important role in the
compensation effect, and the physiological metabolism and growth of We gratefully acknowledge the help of the teachers and workers
the roots were promoted by interactions among these factors. from the experimental station at the Institute of Cotton Research of the
Additionally, the compensation ability of the drought-resistant cultivar Chinese Academy of Agricultural Sciences.

Appendix A

Table A1
Chemical properties of the soil used for the experiments in 2014 and 2015.

Year Organicmatter (g/kg) Total N Available P (mg/ Available K (mg/

(g/kg) kg) kg)

2014 7.9 1.5 18.4 143.5

2015 8.7 1.6 18.9 187.0

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 Table A2
Air temperature, total duration of sunshine and rainfall, and mean relative humidity at the experimental site during May to October in 2014 and 2015 in the Henan Anyang.
J. Niu et al.

Year

2014 2015

Month MDT (°C) MDTmax (°C) MDTmin (°C) Accumulated Sunshine Mean relative Total MDT (°C) MDTmax (°C) MDTmin (°C) Accumulated Sunshine Mean relative Total
temperature (°C) hours (h) humidity (%) rainfall temperature (°C) hours (h) humidity (%) rainfall
(mm) (mm)

May 22.78 30.03 15.33 691.31 273.34 61.15 31.80 20.71 27.17 14.50 617.54 239.90 27.51 59.70
June 25.99 32.53 20.23 779.65 217.15 58.66 37.20 26.06 32.62 19.66 781.69 193.55 17.45 19.90
July 26.87 33.34 21.77 832.90 223.40 14.24 129.40 27.02 33.65 21.50 837.76 221.16 12.94 72.40
August 25.10 31.45 20.21 778.21 215.30 4.55 46.10 25.86 32.24 20.87 801.53 193.81 25.04 29.00
September 20.61 26.50 16.05 590.09 121.98 15.94 156.10 21.13 27.61 15.71 633.87 176.91 4.98 17.20
October 16.74 23.43 11.25 436.87 158.79 26.57 4.30 15.68 22.91 8.94 333.11 176.18 0.37 18.20
Mean 23.02 29.55 17.47 684.84 201.66 30.19 67.48 22.74 29.37 16.86 667.58 200.25 14.72 36.07

MDT, MDTmin, and MDTmax stand for mean daily temperature, mean daily minimum temperature and mean daily maximum temperature, respectively. Accumulated temperature were calculated from maximum and minimum temperatures as
∑[(max. temp. + min. temp.)/2–15 °C].

38
Table A3
Changes in the cotton yield and yield composition under drought stress after re-watering during the seedling stage in 2014.

Varieties

CCRI-45 CCRI-60

Treatment Boll number per plant Boll weight (g) Lint (%) Lint cotton yield (kg Hm−2) Boll number per plant Boll weight (g) Lint (%) Lint cotton yield (kg Hm−2)

CK 17.67b 4.43e 40.06b 1550.17c 19.00ab 4.07d 39.36cd 1507.61b

D1L3 18.33ab 4.68d 40.22b 1708.58bc 19.67a 4.03d 40.08b 1570.64b
D2L3 18.33ab 4.77c 40.35b 1744.71b 18.67ab 4.76a 40.79a 1792.73a
D1L6 20.00a 4.82b 41.36a 1972.91a 17.00b 4.58b 39.84bc 1536.98b
D2L6 19.33ab 5.07a 39.86b 1932.98a 14.00c 4.25c 39.41cd 1161.27c
D1L9 14.00c 4.60d 38.34c 1221.83d 10.33d 4.26c 39.11de 852.56d
D2L9 9.67d 4.62d 37.88c 836.55e 10.33d 4.12cd 38.76e 816.75d
Journal of Plant Physiology 224–225 (2018) 30–48
J. Niu et al. Journal of Plant Physiology 224–225 (2018) 30–48

Table A4
Changes in the cotton yield and yield composition under drought stress after re-watering during the flowering and boll-forming stage in 2014 and 2015.

Year

2014 2015

Varieties Treatment Boll number per Boll weight Lint (%) Lint cotton yield Boll number per Boll weight Lint (%) Lint cotton yield
plant (g) (kg Hm−2) plant (g) (kg Hm−2)

CCRI-45 CK 16.67b 5.10a 40.11cd 1688.28b 19.67a 5.80ab 39.11a 2213.53b

D1L3 17.33ab 5.11a 40.86c 1791.57b 19.67a 5.81ab 39.69a 2243.41a
D2L3 17.67ab 4.83b 40.77c 1720.79b 20.33a 6.03ab 39.44a 2410.49a
D1L6 17.67ab 5.17a 41.71b 1886.12ab 20.00a 6.21ab 40.09a 2456.14a
D2L6 19.33a 5.11a 42.76a 2093.03a 19.33a 6.33a 41.28a 2501.99a
D1L9 13.67c 5.12a 39.59d 1374.29c 14.67b 5.62bc 38.69a 1581.94c
D2L9 10.33d 5.11a 37.94e 990.33d 11.00c 5.12c 39.42a 1100.15c
CCRI-60 CK 17.33b 5.00a 39.17cd 1681.02c 19.00a 5.68ab 38.53a 2064.82b
D1L3 17.67b 4.97a 39.68c 1722.27c 20.00a 5.70ab 39.29a 2224.94a
D2L3 18.67ab 4.99a 40.97ab 1886.61b 20.67a 5.83ab 39.76a 2363.65a
D1L6 19.67a 4.31a 41.79a 2071.58a 20.00a 6.20a 39.48a 2427.26a
D2L6 17.33b 4.78a 40.68b 1666.50c 15.33b 6.12a 39.70a 1854.88a
D1L9 14.67c 4.39a 38.61de 1223.48d 11.33c 5.42ab 39.47a 1207.66c
D2L9 9.33d 4.32a 38.05e 753.06e 11.33c 5.05c 39.97a 1131.14c

Note: CK: SRWC (75 ± 5)%; D1: SRWC (60 ± 5)%; D2: SRWC (45 ± 5)%; L3: drought stress for three days; L6: drought stress for six days; and L9: drought stress for nine days. In the
columns, different letters indicate significant differences at the 0.05 probability level.

Table A5
Changes in the final agronomic traits of cotton under drought stress after re-watering during the seedling stage in 2014.

Varieties

CCRI-45 CCRI-60

Treatment Height (cm) Fruiting branch nodes Fruiting positions Shedding rate Height (cm) Fruiting branch nodes Fruiting positions Shedding rate
number number (%) number number (%)

CK 90.33a 12.67a 46.33b 61.88c 109.00ab 12.33a 50.00a 61.78c

D1L3 91.00a 13.67a 47.67b 61.45c 110.00ab 13.33a 51.67a 61.79c
D2L3 95.33a 13.33a 46.67b 60.69c 113.00a 12.67a 50.67a 63.07c
D1L6 96.33a 13.67a 53.00a 62.14c 110.00ab 12.67a 53.33a 68.13ab
D2L6 93.00a 11.33a 52.33a 62.97bc 105.00ab 12.00ab 47.33a 70.41ab
D1L9 77.33b 11.67a 45.00c 68.70ab 95.33c 11.33ab 35.67b 69.98ab
D2L9 73.67b 10.33a 39.00c 75.04a 90.00c 10.00b 39.33b 73.38a

Table A6
Changes in the final agronomic traits of cotton under drought stress after re-watering during the flowering and boll-forming stage in 2014 and 2015.

Year

2014 2015

Varieties Treatment Height (cm) Fruiting branch Fruiting positions Shedding rate Height (cm) Fruiting branch Fruiting positions Shedding rate
nodes number number (%) nodes number number (%)

CCRI-45 CK 84.67a 13.67a 49.33ab 66.09bc 93.67a 14.33ab 60.00a 67.20bc

D1L3 85.33a 13.67a 49.67ab 64.94cd 95.67a 15.67a 59.00a 66.69bc
D2L3 86.33a 13.33a 51.33a 65.51cd 96.33a 15.67a 60.33a 66.25c
D1L6 87.00a 13.33a 51.67a 65.79bc 94.00a 13.33b 56.67ab 64.74c
D2L6 85.67a 13.33a 48.33ab 59.92d 92.00a 13.33b 53.00bc 63.59c
D1L9 72.00b 10.33b 47.33ab 71.05b 80.00b 10.33c 50.67c 71.06a
D2L9 70.67b 9.67b 46.33b 77.68a 76.00b 9.33c 43.67d 74.77a
CCRI-60 CK 110.33ab 12.33a 49.33a 64.85bc 120.00a 13.33ab 56.00a 66.11b
D1L3 110.67ab 12.67a 48.67ab 63.66bc 122.00a 14.00a 58.33a 65.60b
D2L3 111.67ab 12.33a 50.33a 62.84bc 122.67a 12.33ab 58.00a 64.05b
D1L6 112.33a 11.67ab 51.00a 61.37c 121.67a 12.00ab 55.00ab 63.56b
D2L6 111.67ab 11.33ab 46.67ab 62.69c 108.33b 11.67ab 47.33c 67.60ab
D1L9 101.33c 11.00b 45.67ab 67.83b 107.33b 11.00c 42.33c 73.47a
D2L9 100.67c 9.00b 43.67b 78.51a 103.67b 8.33c 43.33c 73.69a

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J. Niu et al. Journal of Plant Physiology 224–225 (2018) 30–48

Table A7
Changes in the cotton fiber quality under drought stress after re-watering during the seedling stage in 2014.

Varieties

CCRI-45 CCRI-60

Treatment Fiber length Uniformity Micronaire Elongation rate Fiber strength Fiber length Uniformity Micronaire Elongation rate Fiber strength
(mm) index (%) (%) (cN/tex) (mm) index (%) (%) (cN/tex)

CK 29.84ab 84.20a 3.55a 5.70b 32.99a 28.22c 83.13a 3.59bc 5.47a 30.22ab
D1L3 29.68ab 84.90ab 3.85a 5.77b 33.9a 28.34bc 83.27a 3.75bc 5.83a 30.83a
D2L3 30.16a 84.80ab 3.94a 6.00ab 33.23a 29.17a 83.57a 4.32a 5.87a 30.61a
D1L6 29.93ab 85.37a 4.01a 6.00ab 33.47a 28.73b 83.48a 4.04ab 5.73a 30.67a
D2L6 29.86ab 85.20a 3.78a 6.13ab 32.60a 28.39bc 83.13a 3.96ab 5.07a 29.50ab
D1L9 29.69ab 84.43ab 3.83a 6.53a 31.05b 28.28c 82.49b 3.48c 5.20a 28.46b
D2L9 29.56b 84.03b 3.83a 5.73b 30.60b 28.19c 82.12b 3.65c 4.93a 28.42b

Table A8
Changes in the cotton fiber quality under drought stress after re-watering during the flowering and boll-forming stage in 2014 and 2015.

Year

2014 2015

Varieties Treatment Fiber Uniformity Micronaire Elongation Fiber Fiber Uniformity Micronaire Elongation Fiber
length index (%) rate (%) strength (cN/ length index (%) rate (%) strength (cN/
(mm) tex) (mm) tex)

CCRI-45 CK 30.37b 84.20b 3.81a 5.20b 32.26a 28.33bc 83.27bc 5.47a 6.67a 28.80c
D1L3 30.50ab 84.67ab 3.91a 5.27b 32.26a 28.77bc 84.77ab 5.47a 6.67a 28.93bc
D2L3 30.84ab 84.83ab 4.03a 5.73ab 32.41a 29.63ab 85.13ab 5.03b 6.63a 30.30ab
D1L6 31.27a 85.00ab 4.12a 5.40ab 32.57a 29.77ab 86.23a 5.70a 6.70a 30.70a
D2L6 30.38b 85.73a 4.17a 6.53a 32.31a 30.23a 86.23a 5.37ab 6.63a 31.70a
D1L9 30.11bc 84.18b 4.16a 5.20b 32.31a 27.23cd 82.57cd 5.43a 6.63a 27.60cd
D2L9 29.59c 82.83c 3.86a 5.33b 32.24a 26.70d 81.00d 5.37ab 6.63a 26.77d
CCRI-60 CK 28.34ab 81.67bc 4.00a 4.77ab 29.37a 28.00ab 82.17bc 5.70ab 6.60a 27.63ab
D1L3 28.52ab 82.00abc 4.08a 5.03ab 29.52a 28.50ab 83.60ab 5.50ab 6.60a 28.87a
D2L3 28.61ab 82.63ab 4.13a 4.90b 29.73a 28.53ab 84.53a 5.60ab 6.57a 28.80a
D1L6 28.92a 83.00a 4.23a 5.33a 29.56a 29.17a 84.73a 5.33b 6.60a 29.30a
D2L6 29.06a 82.53abc 4.16a 5.13ab 29.55a 29.47a 83.27ab 5.67ab 6.60a 27.57ab
D1L9 28.06ab 81.13bc 3.89a 5.17ab 29.23a 27.03bc 80.73bc 5.90a 6.57a 25.97bc
D2L9 27.47b 80.70c 3.78a 5.13ab 29.22a 26.03c 80.30c 5.67ab 6.53a 24.90c

Fig. A1. The relative water content of soil in 2014 and 2015.
Note: CK: regular irrigation; D1: soil relative water content of 60% ± 5%; D2: soil relative water content of 45% ± 5%. L3: drought for three days; L6: drought for six days; and L9:
drought for nine days. The relative water content of the soil during the seedling stage in 2014 and 2015 (A and C), the relative water content of the soil during the flowering and boll-
forming stage in 2014 and 2015 (B and D).

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J. Niu et al. Journal of Plant Physiology 224–225 (2018) 30–48

Fig. A2. Changes in the cotton root dry weight under drought stress and re-watering during the seedling stage in 2014.
Note: L3: drought stress for three days; L3-5: fifth day after re-watering following three-day drought stress; L3-10: tenth day after re-watering following three-day drought stress; L6:
drought stress for six days; L6-5: fifth day after re-watering following six-day drought stress; L6-10: tenth day after re-watering following six-day drought stress; L9: drought stress for nine
days; L9-5: fifth day after re-watering following nine-day drought stress; and L9-10: tenth day after re-watering following nine-day drought stress. The changes in root dry weights of the
drought-resistant cotton variety CCRI-45 in 2014 (A), the changes in root dry weights of the drought-sensitive cotton variety CCRI-60 in 2014 (B). In the columns, different letters indicate
significant differences at the 0.05 probability level. The same applies below.

Fig. A3. Changes in the cotton root dry weight under drought stress and re-watering during the flowering and boll-forming stage in 2014 and 2015.
Note: The changes in root dry weights of the drought-resistant cotton variety CCRI-45 in 2014 and 2015: A and B, the changes in root dry weights of the drought-sensitive cotton variety
CCRI-60 in 2014 and 2015: C and D. In the columns, different letters indicate significant differences at the 0.05 probability level. The same applies below.

Fig. A4. Relationship between the increased percentages of yield and root dry weight under drought stress after re-watering during the seedling stage in 2014.
Note: * and ** indicate significant differences at the 0.05 and 0.01 probability levels, respectively (n = 6, R0.05 = 0.811, and R0.01 = 0.917).

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J. Niu et al. Journal of Plant Physiology 224–225 (2018) 30–48

Fig. A5. Relationship between the increased percentages of yield and root dry weight under drought stress after re-watering during the flowering and boll-forming stage in 2014 and
2015.

Fig. A6. Changes in the soluble sugar and proline content in cotton roots under drought stress and re-watering during the seedling stage in 2014.
Note: Changes in the soluble sugar content in 2015: A and B; changes in the proline content in 2015: C and D. In the columns, different letters indicate significant differences at the 0.05
probability level.

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J. Niu et al. Journal of Plant Physiology 224–225 (2018) 30–48

Fig. A7. Changes in the soluble sugar content in cotton roots under drought stress and re-watering during the flowering and boll-forming stage in 2014 and 2015.

Fig. A8. Changes in the proline content in cotton roots under drought stress and re-watering during the flowering and boll-forming stage in 2014 and 2015.

Fig. A9. Changes in the soluble protein content in cotton roots under drought stress and re-watering during the seedling stage in 2014.

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Fig. A10. Changes in the soluble protein content in cotton roots under drought stress and re-watering during the flowering and boll-forming stage in 2014 and 2015.

Fig. A11. Changes in the SOD and POD activity in cotton root under drought stress and re-watering during the seedling stage in 2014.

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J. Niu et al. Journal of Plant Physiology 224–225 (2018) 30–48

Fig. A12. Changes in the SOD activity in cotton root under drought stress and re-watering during the flowering and boll-forming stage in 2014 and 2015.

Fig. A13. Changes in the POD activity in cotton roots under drought stress and re-watering during the flowering and boll-forming stage in 2014 and 2015.

Fig. A14. Changes in the CAT activity in cotton roots under drought stress and re-watering during the seedling stage in 2014.

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Fig. A15. Changes in the CAT activity in cotton roots under drought stress and re-watering during the flowering and boll-forming stage in 2014 and 2015.

Fig. A16. Changes in the ABA and GA content in cotton root under drought stress and re-watering during the seedling stage in 2014.

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J. Niu et al.
Fig. A17. Changes in the ABA content in cotton root under drought stress and re-watering during the flowering and boll-forming stage in 2014 and 2015.
Fig. A18. Changes in the GA content in cotton roots under drought stress and re-watering during the flowering and boll-forming stage in 2014 and 2015.
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