Growth and physiological responses of sugarcane to drought stress at an early growth stage

ISSN 2588-1299  
VJAS 2019; 2(4): 451-460  
Vietnam Journal  
of Agricultural  
Sciences  
Growth and Physiological Responses of  
Sugarcane to Drought Stress at an Early  
Growth Stage  
Bui The Khuynh, Vu Ngoc Thang, Vu Dinh Chinh &  
Pham Thi Thom  
Faculty of Agronomy, Vietnam National University of Agriculture, Hanoi 131000, Vietnam  
Abstract  
A pot experiment was conducted in a net house to evaluate the effects  
of drought stress (a 20-day water withholding treatment from 100-  
120 days after planting) on the growth and physiology of five  
sugarcane cultivars. The results showed that water stress at an early  
stage significantly affected sugarcane growth and physiology.  
Water stress resulted in reductions in plant height, stalk diameter,  
and leaf number of sugarcane, in addition to reductions in the  
photosynthetic pigment content, Fv/Fm, and SPAD (Soil Plant  
Analysis Development) readings after the 20-day withholding  
water period (120 DAP), and in stem, root, and leaf fresh weights,  
and leaf area at 150 DAP. Besides, drought stress led to increases  
in stomata density and decreases in stomata length. Variation was  
also found among the cultivars in response to water stress.  
Significant genotypic differences in stem fresh weight and leaf  
area under water stress among the cultivars were observed. The  
highest value of stem fresh weight under stressed conditions was  
recorded in ROC22 (50.6g), followed by QĐ159 (46.5g), ROC16  
(46.2g), ROC10 (46.1g), and VL06 (44.4g). However, the highest  
DTI was recorded in ROC16, followed by VL06, ROC10, QĐ93-  
159, and ROC22, respectively.  
Keywords  
Sugarcane, drought stress, growth, physiological response  
Introduction  
Received: June 9, 2019  
Drought is considered a major abiotic stress that limits crop  
production, resulting in severe reductions of their growth rate and  
development (Begcy et al., 2012). Sugarcane (Saccharum spp.) is an  
important crop used in the production of approximately 60% of the  
global sugar supply as well as in the production of ethanol and bio-  
energy (Amalraj et al., 2010). As a C4 plant with a high  
photosynthetic capacity, sugarcane is highly dependent on water  
Accepted: November 2, 2019  
Correspondence to  
ORCID  
Khuynh T. Bui  
752X  
451  
Growth and Physiological Responses of Sugarcane to Drought Stress at an Early Growth Stage  
availability. Sugarcane is mostly cultivated in  
tropical and sub-tropical rain-fed regions on both  
sides of the equator (from 35o North to 35o South)  
(Silva et al., 2013). In many of these regions,  
rainfall does not provide the amount of water  
required for the growth and photosynthetic  
activities of the crop, and thus water shortages  
remain a major limiting factor for achieving high  
yields of sugarcane. Even in humid tropical areas,  
inequality in rainfall distribution leads to  
reductions in growth or yield loss (Silva et al.,  
2013). A yield loss up to 60% due to drought has  
been recorded in sugarcane (Robertson et al.,  
1999). In Vietnam, the largest growing region of  
sugarcane is located along the central coast where  
drought is a recurring problem. Apart from  
University of Agriculture. The mature stalks (8  
months old) of five cultivars were selected, cut  
into short pieces of which each had one active  
bud, and used for propagating in germination  
trays containing moist sand. The uniformly  
germinated seedlings (buds 5-7cm in height)  
were then transplanted into plastic containers  
with the dimensions of 25cm in diameter and  
35cm in height, and filled with 15kg of dry soil.  
A two factorial experiment was laid out with  
five replications following  
a
completely  
randomized design (CRD). The two water  
regimes (field capacity as the control and drought  
stress followed by recovery) were designated  
factor A, while the five sugarcane cultivars were  
designated factor B. Water was supplied to all the  
treatments at the field capacity level from the  
time of seed cane planting to 100 days after  
planting (100 DAP). For the drought stress  
treatment, water was withheld for 20 days from  
100 to 120 DAP. Re-watering then was done in  
the drought stress treatment to bring the soil back  
to the level of field capacity and maintained in  
that condition from 120 to 150 DAP. Soil  
moisture was regularly measured at a depth of 20  
cm using a soil tester Takemura DM15 (Japan)  
every five days during the drought stress period.  
Mean values of the recorded soil moistures in the  
stressed treatment ranged from 78-85%, 75-82%,  
62-68%, and 46-54% on the 5th, 10th, 15th, and  
20th days of withholding water, respectively.  
irrigation  
practices,  
drought  
tolerance  
improvement is required to alleviate drought and  
water deficit problems, and to reduce yield losses.  
Successful selection and breeding for cultivars  
with drought tolerance can only be achieved when  
the mechanisms underlying drought tolerance in  
sugarcane are fully understood. Better  
understanding of the physiological mechanisms  
employed by the plant to better cope with drought  
and water deficits will enhance the success rate in  
selecting and breeding sugarcane for drought  
tolerance (Silva et al., 2013). Thus, the objectives  
of this study were to evaluate the effects of a water  
deficit on the growth of five sugarcane cultivars  
and analyze the genotypic differences in their  
levels of drought tolerance.  
Measurements  
Materials and Methods  
Plant height, stalk diameter, and leaf number  
were measured at 100, 120 (end of the drought  
stress treatment), and 150 DAP (30 days after re-  
watering). The number of fully expanded leaves  
was counted, and the center point of the stalk was  
measured for stalk diameter using a vernier caliper.  
Plant culture and water deficit treatment  
Five sugarcane cultivars (Table 1) were  
used in this study. A pot experiment was  
conducted in greenhouse conditions at the  
Faculty of Agronomy, Vietnam National  
Table 1. Origin of the five sugarcane cultivars used in the study  
Cultivar  
Origin  
QĐ93-159 (Yuetang 93-159)  
ROC10  
GSIRI, China  
TSRI, Taiwan  
TSRI, Taiwan  
TSRI, Taiwan  
TSRI, Taiwan  
ROC16  
ROC22  
Vien Lam 6 (VL6)  
Note: NTSRI: Taiwan Sugar Research Institute; GSIRI: Guangzhou Sugarcane Industry Research Institute.  
Vietnam Journal of Agricultural Sciences  
452  
Bui The Khuynh et al. (2019)  
Stomatal density and size were measured  
according to the methods of Camargo & Marenco  
(2011). The youngest fully expanded leaves (5  
leaves and 4 samples per leaf) were selected, and  
the midrib vein was removed. Stomatal density  
and stomatal size were determined from nail  
polish imprints taken from both the adaxial and  
abaxial leaf surfaces. Stomata were counted by  
calculating the view field using a stage  
micrometer placed under the microscope (Nikon  
Ys100) with a 10x objective lens. Stomatal size  
was measured from a sample of 20 stomata per  
leaf at 100x magnification.  
(version 7.2) and Sigmaplot 12.5. Treatment  
means were compared using the least significant  
difference (LSD) test.  
Results and discussion  
Plant height, stalk diameter, and leaf number  
Drought stress can reduce leaf size, decrease  
stem growth and root expansion, increase hair  
density on leaves and stems, and alter plant and  
water relations (Farooq et al., 2012). After 20  
days of withholding water, significant reductions  
in plant height, stalk diameter, and leaf number  
were recorded in the five cultivars (Table 2). This  
is consistent with the findings of Silva et al.  
(2008) and Smith et al. (2005) who reported that  
cane elongation and stalk height were negatively  
and strongly affected by drought. At 150 DAP  
(30 days after re-watering), the plant height and  
stalk diameter of all the cultivars had not fully  
recovered as large reductions in stalk diameter  
and plant height were still observed in the  
stressed treatments among the five cultivars.  
Significant differences among the five sugarcane  
cultivars were recorded in plant height, stalk  
diameter, and leaf number in the stressed period  
(120 DAP) and after the recovery period (150  
DAP). At 120 DAP, the highest plant height was  
observed in ROC16 (140.5cm), followed by  
ROC22, QĐ93-159, ROC10, and VL06 with  
values of 126.0, 112.8, 98.7, and 79.5cm,  
respectively. The reduction of stem height may  
have been caused by the low soil water potential  
which resulted in smaller leaf sizes and a smaller  
number of leaves per plant (Reddy et al., 2003).  
After the recovery period (150 DAP), stalk  
diameters ranged from 10.9mm (in VL06) to  
16.1mm (in ROC10). Jangpromma et al. (2012)  
reported that drought stress did not lead to  
significant variations in stalk diameter among  
sugarcane cultivars after a 10-day water  
withholding period. These differences compared  
with the current study may indicate differences  
among the cultivars, experimental conditions,  
and duration of the stress treatment. A drought  
occurring for more than 10 days was seen to result  
in severe reductions in root development and  
yield (Jangpromma et al., 2012).  
Chlorophyll a fluorescence and photosynthetic  
pigments measurements  
The emission of chlorophyll a fluorescence  
was evaluated at 12:00 pm, using a modulated  
fluorometer (Opti-Sciences, OS30p+, USA), as  
described by Medeiros et al. (2013). The initial  
fluorescence (F0), the maximum fluorescence  
(Fm), and the maximum quantum efficiency of  
photosystem II (Fv /Fm) were then recorded after  
30 minutes of dark adaptation using leaf-clips on  
the first fully expanded leaf.  
For the quantification of chlorophyll a,  
chlorophyll b, and the carotenoids, 0.1g of fresh,  
fully expanded leaves was collected, ground, and  
soaked in 10mL of 90% acetone for 48h.  
Absorbance readings at the wavelengths of 663,  
647, and 470nm were measured in  
a
spectrophotometer and used for the calculation of  
the chlorophyll a, chlorophyll b, and carotenoid  
contents (Arnold, 1949), and expressed as mg g-  
1
fresh weight (FW). Root fresh weight, stem  
fresh weight, leaf fresh weight, and leaf area  
were measured at 150 DAP. SPAD chlorophyll  
readings were measured at 100, 110, and 120  
DAP using portable chlorophyll meters (SPAD-  
502, Minolta, Japan). Drought tolerant index  
(DTI) was determined following the methods of  
Hoang et al. (2018) where DTI equaled the dry  
stem weight under stressed conditions divided by  
the dry weight under well-watered conditions.  
Data analysis  
Analysis of variance (ANOVA) tests on the  
measured traits was performed using CropStat  
453  
Growth and Physiological Responses of Sugarcane to Drought Stress at an Early Growth Stage  
Table 2. Plant height, stalk diameter, and leaf number of different sugarcane genotypes before (100 days after planting, DAP) and  
after (120 DAP) a 20 day-drought, and 30 days after re-watering (150 DAP)  
Plant height (cm plant-1)  
Stalk diameter (mm)  
Leaf number (leaves plant-1)  
Treatment  
100 DAP 120 DAP 150 DAP 100 DAP 120 DAP 150 DAP 100 DAP 120 DAP 150 DAP  
ROC10 FC  
82.3  
84.0  
100.0  
96.6  
103.7  
90.3  
89.5  
85.6  
53.6  
61.5  
14.6  
83.2  
98.7  
97.0  
87.6  
105.2  
92.2  
152.0  
146.0  
200.0  
160.0  
175.3  
148.4  
158.3  
148.5  
141.5  
83.1  
8.7  
8.3  
8.6  
8.9  
8.2  
7.7  
8.5  
8.3  
6.0  
6.2  
2.0  
8.4  
8.7  
7.9  
8.4  
13.1  
9.4  
17.1  
15.1  
14.2  
12.4  
17.9  
13.8  
17.3  
13.0  
11.6  
10.3  
2.2  
7.0  
7.0  
5.3  
5.6  
6.6  
6.0  
5.3  
5.3  
6.3  
6.3  
0.8  
7.0  
5.5  
6.3  
5.3  
9.7  
6.7  
7.0  
5.7  
9.0  
6.3  
7.0  
6.3  
8.6  
5.6  
1.4  
8.2  
6.3  
7.7  
6.7  
10.7  
7.0  
8.0  
4.7  
8.7  
6.3  
9.3  
7.7  
6.0  
4.3  
1.8  
8.8  
6.3  
7.5  
8.5  
Stressed  
FC  
ROC16  
ROC22  
161.7  
119.3  
139.0  
112.3  
117.5  
108.0  
92.4  
12.2  
8.7  
Stressed  
FC  
12.2  
8.2  
Stressed  
FC  
QĐ93-  
12.4  
8.7  
159  
Stressed  
FC  
VL06  
8.1  
Stressed  
68.9  
7.1  
LSD0.05C*W  
12.0  
16.9  
1.6  
Mean  
ROC10  
ROC16  
ROC22  
98.7  
149.4  
180.1  
161.8  
153.4  
11.2  
10.4  
10.2  
10.6  
16.1  
13.3  
15.9  
15.2  
140.5  
126.0  
112.8  
QĐ93-  
159  
VL06  
57.5  
10.3  
86.0  
83.6  
6.5  
79.5  
8.5  
112.3  
11.9  
165.5  
137.4  
7.5  
6.6  
1.4  
7.6  
1.2  
10.9  
1.5  
6.3  
0.5  
6.1  
6.0  
0.3  
7.9  
7.2  
1.0  
5.1  
1.3  
LSD0.05C  
Mean  
FC  
123.3  
99.6  
5.4  
8.0  
11.6  
8.4  
15.6  
12.9  
1.0  
8.3  
8.5  
Stressed  
8.0  
6.1  
6.0  
LSD0.05W  
CV%  
0.9  
0.7  
0.6  
0.8  
10.1  
6.3  
6.5  
14.8  
9.6  
8.9  
11.1  
14.7  
Note: C: cultivar; W: water treatment; FC: field capacity (control); Stressed: drought stress treatment.  
Stomatal density and size  
length in the sugarcane cultivars under stressed  
conditions compared to the control. Similar results  
were observed by Meng et al. (1999) and Xu &  
Zhou (2008) in which stomatal density was  
negatively correlated with stomatal length under  
different drought stress conditions. An increase  
in stomatal density and a decrease in stomatal  
size are regarded as important adaptations of  
plants to drought and water deficits (Xu & Zhou,  
2008). Although stomatal density was found to  
increase under drought stress, the number of  
stomata per leaf decreased as the leaf area was  
reduced (Xu & Zhou, 2008). However, Yang et  
al. (2007) showed that stomatal density varies  
according to drought intensification. An increase  
in stomatal density was recorded under light and  
moderate drought, but a decrease was observed  
in severe drought.  
Leaf stomata play a central role in controlling  
the exchange of CO2 and water vapor (Xu &  
Zhou, 2008), and thus, are a crucial part of the  
stress response of plants to water deficits. Under  
drought, reductions in the photosynthetic rate can  
be caused by both stomatal and non-stomatal  
activities, depending on the drought intensity and  
species (Chaves et al., 2003). Our results revealed  
that the stomatal density was higher on the abaxial  
surface than on the adaxial surface under both the  
stressed and control (FC) conditions (Figure 1).  
Drought stress led to significant increases in the  
stomatal density on the abaxial surface of the  
cultivars ROC10, ROC22, and QĐ93-159, and on  
the adaxial surface of the cultivars ROC10,  
ROC22, and VL06. In addition, the results  
showed a decrease in the mean values of stomatal  
Vietnam Journal of Agricultural Sciences  
454  
Bui The Khuynh et al. (2019)  
(B)  
(A)  
FC  
FC  
Stressed  
Stressed  
100  
80  
60  
40  
20  
0
300  
250  
200  
150  
100  
50  
**  
*
ns  
*
*
*
ns  
**  
ns  
ns  
0
ROC10 ROC16 ROC22 QĐ 159 VL 06  
ROC10 ROC16 ROC22 QĐ 159 VL 06  
Genotype  
Genotype  
(D)  
(C)  
ns  
FC  
Stressed  
FC  
Stressed  
**  
*
*
25  
**  
*
40  
30  
*
*
*
*
20  
15  
m  
m  
20  
10  
0
10  
5
0
ROC10 ROC16 ROC22 QĐ 159 VL 06  
ROC10 ROC16 ROC22 QĐ 159 VL 06  
Genotype  
Genotype  
Note: ns = not significant; * = significant at p = 0.05; ** = significant at p = 0.01.  
Figure 1. Mean values of stomatal density on the abaxial (A) and adaxial (B) surfaces,  
and mean values of stomatal length on the abaxial (C) and adaxial (D) surfaces among cultivars  
Photosynthetic pigments  
chlorophyll a content, drought stress also  
reduced the chlorophyll b content by 61%  
compared to the control (FC), leading to changes  
in the chlorophyll a/b ratio. However, no  
significant differences in carotenoid content  
were observed regardless of the cultivar or water  
treatment. Similar results have also been observed  
by Ashraf & Harris (2013) and Mafakheri et al.  
(2010), who reported that reductions in the  
chlorophyll content from drought stress are due to  
Droughts have negative impacts on  
photosynthetic pigments. Depending on the  
drought intensity and duration, drought can cause  
damage to pigments or even deterioration of  
thylakoid membranes (Ashraf & Harris, 2013).  
In this study, the results showed a decrease in  
both chlorophyll a and b content as consequences  
of drought stress in all the cultivars (Table 3).  
While a decrease of 36% was found in the  
455  
Growth and Physiological Responses of Sugarcane to Drought Stress at an Early Growth Stage  
Table 3. Effects of drought stress at an early stage on photosynthetic pigments at 120 DAP  
Treatment  
FC  
Chla (mg g-1)  
0.946  
0.729  
0.955  
0.651  
0.890  
0.763  
0.936  
0.704  
0.954  
0.579  
0.163  
0.838  
0.803  
0.827  
0.820  
0.767  
0.011  
0.936  
0.685  
0.072  
0.1  
Chlb (mg g-1)  
0.338  
0.258  
0.343  
0.213  
0.351  
0.235  
0.339  
0.209  
0.343  
0.220  
0.043  
0.298  
0.278  
0.293  
0.274  
0.282  
0.0285  
0.343  
0.227  
0.018  
8.3  
Carotenoid (mg g-1)  
0.208  
0.203  
0.207  
0.203  
0.206  
0.208  
0.207  
0.198  
0.202  
0.197  
0.009  
0.206  
0.205  
0.207  
0.203  
0.202  
0.006  
0.206  
0.202  
0.041  
2.6  
Chla/Chlb  
Total chl  
ROC10  
ROC16  
ROC22  
QĐ93-159  
VL06  
2.799  
1.284  
Stressed  
2.826  
0.987  
FC  
2.784  
1.298  
Stressed  
FC  
3.056  
0.864  
2.536  
1.241  
Stressed  
FC  
3.247  
0.998  
2.761  
1.275  
Stressed  
FC  
3.368  
0.913  
2.781)  
1.297  
Stressed  
2.632  
0.799  
LSD0.05C*W  
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
Mean  
ROC10  
ROC16  
ROC22  
QĐ93-159  
VL06  
LSD0.05C  
Mean  
FC  
Stressed  
LSD0.05W  
CV%  
Note: C: cultivar; W: water treatment; FC: field capacity (control); Stressed: drought stress treatment.  
SPAD readings and maximum photochemical  
efficiency (Fv/Fm)  
damage of the photosynthetic pigments and  
deterioration of the thylakoid membranes. The  
reduction of the chlorophyll content is mainly  
due to the destruction of pigments caused by  
oxidative damage when the plant is exposed to  
severe drought stress. Generally, drought stress  
leads to a greater reduction of the chlorophyll b  
content than that of chlorophyll a and thus,  
results in an increased ratio of chlorophyll a/b  
(Ashraf & Harris, 2013). Under adverse  
conditions such as drought, plants employ  
different physiological and metabolic strategies  
for survival. For instance, plants can synthesize  
antioxidants such as ascorbate, glutathione, and  
flavonoids (Medeiros et al., 2013), together  
with increasing the activity of antioxidant  
enzymes including peroxidase, superoxide  
dismutase, and catalases.  
The maximum photochemical efficiency  
(Fv/Fm) is positively correlated with the  
photosynthesis rate of a plant and thus, is used as  
an important parameter for quantifying a plant’s  
response to drought stress (Silva et al., 2013).  
Maintaining a similar Fv/Fm ratio between plants  
under drought stress and plants under well-  
irrigated conditions indicates a high efficiency of  
carbon assimilation and hence, the plant is better  
able to adapt to the stressed conditions (Silva et  
al., 2007). No significant differences in the  
Fv/Fm ratios were found among all the cultivars  
at 100 and 110 DAP. All the cultivars had a  
Fv/Fm ratio of 0.74 to 0.78. However, at 120  
DAP (20 days of withholding water), significant  
differences in the Fv/Fm ratios were observed  
Vietnam Journal of Agricultural Sciences  
456  
Bui The Khuynh et al. (2019)  
Table 4. SPAD meter readings and Fv/Fm ratios of different sugarcane cultivars before (100 days after planting, DAP), after 10  
days (110 DAP), and after 20 days of drought (120 DAP)  
Fv/Fm  
SPAD readings  
Treatment  
FC  
100 DAP  
110 DAP  
120 DAP  
100 DAP  
110 DAP  
120 DAP  
150 DAP  
ROC10  
ROC16  
ROC22  
0.765  
0.767  
0.791  
0.781  
0.788  
0.786  
0.780  
0.775  
0.764  
0.783  
0.211  
0.776  
0.764  
0.786  
0.741  
0.784  
0.704  
0.771  
0.756  
0.780  
0.701  
0.161  
0.742  
0.633  
0.773  
0.679  
0.738  
0.581  
0.755  
0.550  
0.752  
0.610  
0.427  
53.6  
51.0  
52.2  
52.7  
50.7  
53.5  
48.3  
53.6  
47.7  
52.2  
4.5  
43.3  
44.2  
46.0  
45.2  
45.7  
46.1  
46.8  
46.1  
44.2  
45.3  
1.9  
45.5  
36.9  
44.2  
40.1  
45.7  
40.9  
43.7  
34.0  
43.9  
36.4  
2.9  
39.0  
44.4  
42.8  
44.3  
42.6  
44.9  
41.7  
43.1  
39.1  
42.6  
2.1  
Stressed  
FC  
Stressed  
FC  
Stressed  
FC  
QĐ93-  
159  
Stressed  
FC  
VL06  
Stressed  
LSD 0.05C*W  
Mean  
ROC10  
ROC16  
ROC22  
QĐ93-159  
VL06  
0.766  
0.786  
0.787  
0.778  
0.773  
0.149  
0.770  
0.763  
0.744  
0.763  
0.762  
0.114  
0.687  
0.726  
0.659  
0.653  
0.681  
0.303  
52.3  
52.4  
52.1  
51.0  
49.9  
3.2  
43.8  
45.6  
45.9  
46.4  
44.8  
1.4  
41.2  
42.2  
43.3  
38.9  
40.2  
2.1  
41.7  
43.5  
43.8  
42.4  
40.9  
1.4  
LSD0.05C  
Mean  
FC  
0.778  
0.778  
0.145  
0.779  
0.742  
0.132  
0.752  
0.610  
0.191  
50.5  
52.6  
2.2  
45.2  
45.3  
0.9  
44.6  
37.7  
1.3  
41.2  
43.9  
2.0  
Stressed  
LSD 0.05W  
CV%  
1.6  
1.2  
3.7  
5.1  
2.5  
4.2  
2.7  
Note: C: cultivar; W: water treatment; FC: field capacity (control); Stressed: drought treatment.  
among cultivars. The highest Fv/Fm ratio value  
was found in ROC16 (0.726) while the rest of  
the cultivars had no significant differences.  
Drought stress only led to a significant  
reduction in the Fv/Fm ratio in the stressed  
treatment compared with the control at 120  
DAP (20 days of water withholding). A Fv/Fm  
ratio value of less than 0.75 indicates the  
beginning of stress and, therefore, a reduction  
in the photosynthetic capacity of the plant  
(Maxwell & Johnson, 2010). Low values of the  
Fv/Fm ratio have also been reported by Graca  
et al. (2010) and Silva et al. (2007) in  
sugarcane under severe drought.  
sugarcane, corn, and papaya (Jangproma et al.,  
2010). Our results showed that drought  
significantly reduced the SCMRs of plants under  
stressed conditions at the end of the drought  
stress treatment (120 DAP), which was  
consistent with the findings of Silva et al. (2007).  
A drought imposed 90 DAP led to reductions in  
the SCMRs, and more severe reductions were  
recorded in susceptible genotypes. SCMRs,  
therefore, can be used for the identification of  
drought-tolerant genotypes (Silva et al., 2007).  
Significant differences in the SCMRs were  
observed among the five cultivars at the end of  
the drought stress treatment. The highest mean  
SCMR value was recorded in ROC22 (43.8),  
followed by ROC16 (42.2), and was lowest in  
QĐ93-159 (38.6). Under stressed conditions, the  
A SPAD chlorophyll meter reading (SCMR)  
is considered a rapid assessment of the  
chlorophyll content in many crops including  
457  
Growth and Physiological Responses of Sugarcane to Drought Stress at an Early Growth Stage  
SCMRs were maintained at an average of 40 in  
ROC22 and ROC16, which were significantly  
higher compared to those in ROC10, VL06, and  
QĐ93-159. This suggests a higher capacity of the  
ROC22 and ROC16 plants to conserve their  
photosynthetic pigment content during drought  
stress. According to Silva et al. (2013), a SCMR  
below 40 in sugarcane indicates the start of  
chlorophyll deficiency, which affects the  
photosynthetic activities of the plants. Thus, the  
low SCMRs (below 40) recorded in ROC10,  
VL06, and QĐ93-159 indicate drought stress  
sensitivities. No significant difference in terms of  
the SCMRs was found between the control and  
stressed treatments at 30 days after re-watering  
(150 DAP).  
Plant fresh weight, leaf area (LA), and  
drought tolerance index (DTI) at recovery  
(150 DAP)  
Sugarcane plants exposed to prolonged  
drought have been reported to experience a  
reduction in growth (Jaiphong et al., 2016). Our  
results showed that root, leaf, and stem fresh  
weights, and leaf area were significantly reduced  
in plants under the drought treatment relative to  
the control (Table 5). Fresh root weight was  
reduced by 51.5% compared to the drought stress  
treatment at 150 DAP. This matches with the  
findings in early studies by Medeiros et al.  
(2013), Jaiphong et al. (2016), and Silva et al.  
(2013). Significant differences in root fresh  
weight, leaf fresh weight, stem fresh weight, and  
Table 5. Effects of drought stress at an early stage on the plant fresh weight, leaf area, and drought tolerance index (DTI) of five  
cultivars at 150 DAP  
Root fresh weight  
(g plant-1)  
Leaf fresh weight  
(g plant-1)  
Stem fresh weight  
(g plant-1)  
Leaf area  
(cm2)  
Drought tolerance  
index (DTI)  
Treatment  
ROC10  
FC  
57.2  
30.7  
50  
158.6  
76.5  
79.3  
46.1  
71.2  
46.2  
96.7  
50.6  
80.5  
46.5  
64.7  
44.4  
4.4  
18.2  
9.5  
-
0.59  
-
Stressed  
FC  
ROC16  
ROC22  
QĐ93-159  
VL06  
188.8  
78.8  
22.5  
10.5  
23  
Stressed  
FC  
31.3  
93.1  
41.3  
58.1  
21.2  
46.5  
26.7  
4.1  
0.69  
-
176.5  
115.6  
164.6  
82.5  
Stressed  
FC  
17.1  
14.9  
8.8  
0.55  
-
Stressed  
FC  
0.57  
-
91.3  
17.2  
5.7  
Stressed  
54.9  
0.68  
LSD 0.05C*W  
10.5  
1.9  
Mean  
ROC10  
ROC16  
ROC22  
QĐ93-159  
43.9  
40.6  
67.2  
39.6  
117.5  
133.8  
146.1  
123.5  
62.7  
58.7  
73.6  
63.5  
13.9  
16.5  
20.1  
11.8  
VL06  
36.6  
73.1  
54.5  
11.4  
LSD0.05C  
2.8  
7.4  
3.1  
1.3  
Mean  
FC  
60.9  
30.2  
1.8  
155.9  
81.6  
4.7  
78.5  
46.8  
2
19.2  
10.3  
0.9  
Stressed  
LSD 0.05W  
CV%  
5.2  
5.2  
4.2  
7.8  
Note: C: cultivar; W: water treatment; FC: field capacity (control); Stressed: drought treatment.  
Vietnam Journal of Agricultural Sciences  
458  
Bui The Khuynh et al. (2019)  
leaf area were found among sugarcane cultivars.  
The mean values of stem fresh weight ranged  
from 54.5g (in VL06) to 73.6g (in ROC22).  
Significant differences in stem fresh weight and  
leaf area were also found among the cultivars  
under drought stress. The highest value of stem  
fresh weight under stressed conditions was  
recorded in ROC22 (50.6g), followed by QĐ93-  
159 (46.5g), ROC16 (46.2g), ROC10 (46.1g),  
and VL06 (44.4g). Leaf area also varied among  
the sugarcane cultivars and ranged from 11.4cm2  
in VL06 to 20.1cm2 in ROC22. A reduction in  
plant growth under water deficit conditions can  
be seen as the consequence of the decrease in cell  
elongation caused by the interruption of water  
flow from the xylem to surrounding elongation  
cells, the increase in cell sap concentration, and  
the dehydration of the protoplasm (Nonami,  
and VL06 (44.4g). However, the highest DTI  
was recorded in ROC16, followed by VL06,  
ROC10, QĐ93-159, and ROC22, respectively.  
Conflicts of Interest  
The authors declare no conflicts of interest.  
References  
Amalraj R. S., Selvaraj N., Veluswamy G. K., Rananujan  
R. P., Muthurajan R. & Palaniyandi M. (2010).  
Sugarcane proteomics: Establishment of a protein  
extraction method for 2-DE in stalk tissues and  
initiation of sugarcane proteome reference map.  
Electrophoresis. 31: 1959-1974.  
Arnon D. I. (1949). Copper enzymes in isolated  
chloroplasts, polyphenoxidase in Beta vulgaris. Plant  
Physiology. 24(1): 1-15.  
Ashraf M. & Harris P. J. C. (2013). Photosynthesis under  
stressful environments: An overview. Photosynthetica.  
51(2): 163-190.  
1998;  
Larcher,  
2003).  
Low  
biomass  
accumulation of sugarcane, when exposed to  
drought stress, can be explained by reductions in  
light interception, plant extension rate, and  
photosynthetic capacity (Koonjah et al., 2006).  
The drought tolerance index (DTI) has been used  
as an important parameter in evaluating the  
tolerance ability of crops. Among the sugarcane  
cultivars, the highest DTI was recorded in  
ROC16, followed by VL06, ROC10, QĐ93-159,  
and ROC22, respectively.  
Begcy K., Mariano E. D., Gentile A., Lembke C. G.,  
Zingaretti S. M., Souza G. M. & Menossi M. (2012).  
A novel stress-induced sugarcane gene confers  
tolerance to drought, salt and oxidative stress in  
transgenic tobacco plants. PLoS One. 7:e44697.  
Camargo M. A. B. & Marenco R. A. (2011). Density, size  
and distribution of stomata in 35 rainforest tree species  
in Central Amazonia. Acta Amazonia. 41(2): 205-212.  
Chaves M. M., Maroco J. P. & Pereira J. S. (2003).  
Understanding plant responses to drought-from genes  
to the whole plant. Functional Plant Biology. 30: 239-  
264.  
Conclusions  
Farooq M., Hussain M., Wahid A. & Siddique K. H. M.  
(2012). Drought stress in plants: an overview. In:  
Aroca R. (Eds.) Plant responses to drought stress.  
Springer, Berlin/Heidelberg. 1-33.  
Drought stress at an early growth stage (100-  
120 DAP) significantly affected the growth and  
physiological characteristics of five sugarcane  
cultivars. A 20-day water withholding treatment  
resulted in reductions in plant height, stalk  
diameter, and leaf number of sugarcane, in  
addition to reductions in photosynthetic pigment  
content, Fv/Fm, and SPAD readings at 120 DAP,  
and in stem, root, and leaf fresh weights, and leaf  
area at 150 DAP (30 days after re-watering).  
Furthermore, drought stress led to reductions in  
stomatal density and increases in stomatal length.  
Variation was also found among the cultivars in  
response to drought stress. The highest value of  
stem fresh weight under stressed conditions was  
recorded in ROC22 (50.6g), followed by QĐ93-  
159 (46.5g), ROC16 (46.2g), ROC10 (46.1g),  
Graça J. P., Rodrigues F. A., Farias J. R. B., Oliveira M. C.  
N., Hoffmann-Campo C. B. & Zingaretti S. M. (2010).  
Physiological parameters in sugarcane cultivars  
submitted to water deficit. Brazilian Journal of Plant  
Physiology. 22: 189-197.  
Hoang D. T., Hiroo T. & Yoshinobu K. (2018). Nitrogen  
use efficiency and drought tolerant ability of various  
sugarcane varieties under drought stress at early  
growth stage. Plant Production Science. 22: 250-261.  
Jaiphong T., Tominaga J., Watanabe K., Nakabaru M.,  
Takaragawa H., Suwa R., Ueno M. & Kawamitsu Y  
(2016). Effects of duration and combination of drought  
and flood conditions on leaf photosynthesis, growth  
and sugar content in sugarcane, Plant Production  
Science. 19: 427-437.  
Jangpromma N., Thammasirirak S., Jailsil P. & Songsri P.  
(2012). Effects of drought and recovery from drought  
459  
Growth and Physiological Responses of Sugarcane to Drought Stress at an Early Growth Stage  
stress on above ground and root growth, and water use  
efficiency in sugarcane (Saccharum officinarum L.).  
Australian Journal of Crop Science. 6: 1298-1304.  
Research. 111: 373-382.  
Robertson M. J., Inman-Bamber N. G., Muchow R. C. &  
Wood A. W. (1999). Physiology and productivity of  
sugarcane with early and mid-season water deficit.  
Field Crop Research. 64: 211-227.  
Jangpromma N., Songsri P. & Jaisil P. (2010). Rapid  
assessment of chlorophyll content in sugarcane using a  
SPAD chlorophyll meter across different water stress  
conditions. Asian Journal of Plant Science. 9: 368-374.  
Reddy T. Y., Reddy V. R. & Anbumozhi V. (2003).  
Physiological response of groundnut (Arachis hypogea  
L.) to drought stress and its amelioration: A critical  
review. Plant Growth Regulation. 41: 75-88.  
Koonjah S. S., Walker S., Singels A., Van Antwerpen R.  
& Nayamuth A. R. (2006). A quantitative study of  
water  
stress  
effect  
on  
sugarcane  
Silva M. A., Jifon J. L., Santos C. M., Jadoski C. J. & Silva  
J. A. G. (2013). Photosyntheitc capacity and water use  
efficiency in sugarcane genotypes subject to water  
deficit during early growth phase. Brazilian Archives  
of Biology and Technology. 56: 735-748.  
photosynthesis. Proceedings of the South African  
Sugarcane Technologists’ Association. 80: 148-158.  
Larcher W. (2003). Physiological plant ecology:  
Ecophysiology and stress physiology of functional  
groups (4th ed). Springer-Verlag. 401-416.  
Silva M. A., Jifon J. L., Silva J. A. G. & Sharma V. (2007).  
Use of physiological parameters as fast tools to screen  
for drought tolerance in sugarcane. Brazilian Journal  
of Plant Physiology. 19: 735-748.  
Mafakheri A., Siosemardeh A. & Bahramnejad B. (2010).  
Effect of drought stress on yield, proline and  
chlorophyll contents in three chickpea cultivars.  
Australian Journal of Crop Science. 4: 580-585.  
Silva M. A., Silva J. A. G., Enciso J., Sharma V. & Jifon J.  
(2008). Yield components as indicators of drought  
tolerance of sugarcane. Scientia Agricola. 65: 620-627.  
Maxwell K. & Johnson G. N. (2000). Chlorophyll  
florescence: a practical guide. Journal of Experimental  
Botany. 51: 659-668.  
Smith D. M., Inman-Bamber N. G. & Thorburn P. J. (2005).  
Growth and function of the sugarcane root system.  
Field Crops Research. 92: 169-183.  
Meng L., Li L., Chen W., Xu Z. & Liu L. (1999). Effect of  
water stress on stomatal density, length, width and net  
photosynthetic rate in rice leaves. Journal of Shenyang  
Agricultural University. 30: 477-480.  
Xu Z. & Zhou Z. (2008). Responses of leaf stomatal density  
to water status and its relationship with photosynthesis  
in a grass. Journal of Experimental Botany. 59: 3317-  
3325.  
Medeiros D. B., Silva E. C., Nogueira R. J. M. C., Teixeira  
M. M., Buckeridge M. S. (2013). Physiological  
limitations in two sugarcane varieties under water  
suppression and after recovering. Theoretical and  
Experimental Plant Physiology. 25: 213-222.  
Yang L., Han M., Zhou G. & Li J. (2007). The changes of  
water-use efficiency and stoma density of Leymus  
chinensis along Northeast China Transect. Acta  
Ecologica Sinica. 27: 16-24.  
Nonami H. (1998). Plant water relations and control of cell  
elongation at low water potentials. Journal of Plant  
Vietnam Journal of Agricultural Sciences  
460  
pdf 10 trang yennguyen 11100 Free
Bạn đang xem tài liệu "Growth and physiological responses of sugarcane to drought stress at an early growth stage", để tải tài liệu gốc về máy hãy click vào nút Download ở trên

File đính kèm:

  • pdfgrowth_and_physiological_responses_of_sugarcane_to_drought_s.pdf