Journal of Applied Botany and Food Quality 87, 147 - 156 (2014), DOI:10.5073/JABFQ.2014.087.022

1Department of Botany and Microbiology, King Saud University, Riyadh, Saudi Arabia
2University College of Agriculture, University of Sargodha, Sargodha-Pakistan

3Department of Botany, University of Agriculture, Faisalabad-Pakistan

Water deficit-induced regulation of growth, gas exchange, chlorophyll fluorescence, 
inorganic nutrient accumulation and antioxidative defense mechanism 

in mungbean [Vigna radiata (L.)Wilczek]
A. Afzal 3, I. Gulzar 3, M. Shahbaz 3, M. Ashraf 1, 2*

(Received April 16, 2014)

* Corresponding author

Summary
The study was conducted to appraise the influence of water defi-
cit conditions on growth, yield, gas exchange characteristics, and 
antioxidative defense system in two mungbean [Vigna radiata (L.) 
Wilczek] lines, 97001 and 97012. The plants of both lines were 
grown in equal weight plastic pots for 30 days under normal natural 
conditions, after which time two drought regimes [control (well-
watering and 60 % field capacity)] were applied. Data for various 
attributes were recorded after 30 days of drought application while 
at maturity, yield attributes were recorded. Water deficit conditions 
caused a considerable reduction in growth attributes, net CO2 as-
similation rate, stomatal conductance, electron transport ratio, total 
phenolics, leaf Ca2+ and yield attributes. Imposition of water deficit 
conditions significantly increased leaf tocopherol contents and acti-
vity of catalase in both mungbean lines. Both lines showed a con-
siderable variation in growth attributes, the line 97001 being better in 
performance compared with 97012 under water deficit conditions. 

Introduction 
Of various abiotic stresses, drought stress holds an important posi-
tion on the globe (FARAHANI et al., 2009; KUSVURAN et al., 2011). 
The main cause of this stress is the high rate of evapo-transpiration, 
particularly in arid and semi-arid regions having low precipitation 
rate (JALEEL et al., 2007; NAKAYAMA et al., 2007; SHAHBAZ et al., 
2011a, b). Drought is believed to influence plants from metabolic 
compartments to an individual organism (CHOLUJ et al., 2004). It 
can also alter the morpho-physiological and biochemical attributes 
of plants (ALI et al., 2008; RAHBARIAN et al., 2011; SHAHBAZ et al., 
2011a). Although drought stress has a marked inhibitory effect on 
plants at every growth and development phase, its imposition at 
early growth stages causes considerable reduction in both cell ex-
pansion and its elongation (SHAO et al., 2008; ASHRAF et al., 2013).  
Drought stress mainly hampers the rate of carbon assimilation which 
results in decreased growth and yield of most crops (JABEEN et al., 
2008; MAFAKHERI et al., 2010) as well as it significantly reduces 
chlorophyll fluorescence attributes (PIPER et al., 2007). 
Under drought stress, a majority of plants accumulate active osmo-
lytes in their cells, the process referred to as osmotic adjustment 
(AFKARI et al., 2009). These osmolytes support the plant’s defense 
mechanism to nullify the adverse effects of drought (SHAO et al., 
2005) as these osmolytes play a pivotal role in the maintenance of 
water absorption and cell turgor potential under stress conditions 
(CHAVES et al., 2009; MAHMOOD et al., 2009). Osmotic adjustment 
is believed to occur in all plant parts like stem, leaf, root and fruit 
(SAGLAM et al., 2010).
Under drought stress, a variety of reactive oxygen species (ROS) 
are produced, which result in oxidative damage of cell membranes 
(MAHESWARI and DUBEY, 2009). In order to scavenge these activated 
oxygen species, plants produce a number of enzymatic antioxidants 

including: superoxide dismutase (SOD), glutathione reductase (GR), 
catalase (CAT) and peroxidase (POD), as well as non-enzymatic 
antioxidants such as ascorbic acid (AsA), glutathione, α-tocopherol, 
flavonoids and carotenoids (SGHERRI et al., 2000). It is reported 
in different studies that the oxidative cellular damage in drought 
stressed plants is related to the ability of their antioxidant systems 
(LIU et al., 2009; BASU et al., 2010).
Mungbean [Vigna radiata (L.) Wilzeck] is cultivated in many 
regions of the world because of its considerable nutritional value 
particularly for people encountering malnutrition (ALLAHMORADI 
et al., 2011). Despite this, being a leguminous plant it possesses con-
siderable N fixing ability (NADEEM et al., 2004). Although mung-
bean crop requires less management practices (SADEGHIPOUR, 2009), 
sufficient availability of water is attributed for better crop producti-
vity and vice versa (KRAMER and BOYER, 1997). Being a favorite 
crop of arid and semi-arid regions it must have to face drought con-
ditions of varying intensity and time period (LADRERA et al., 2007). 
Mungbean production is adversely affected by the increase in drought 
prone area world-over (POSTEL, 2000). The use of drought resistant 
genotypes is one of the viable means of attaining better yield under 
water deficit conditions (ENNAJEH et al., 2010). In order to develop 
drought resistant varieties/lines it is necessary to have the complete 
knowledge of plant behavior to drought stress (JALEEL et al., 2009). 
The variability of morpho-physiological attributes under water de-
ficit conditions helps to detect resistant genotypes/lines for better 
productivity under stress environment (NAM et al., 2001).
In view of considerable inhibitory effects of drought stress on mung-
bean crop the premier objective of the present investigation was to 
examine how far this stress regulates some key physio-biochemical 
attributes involved in growth and development of mungbean plant.

Materials and methods
To assess the effect of water deficit conditions on mungbean, a pot 
experiment was conducted in Old Botanical Garden, Department of 
Botany, University of Agriculture, Faisalabad-Pakistan. Two lines of 
mungbean (lines 97001 and 97012) were obtained from the Ayub 
Agricultural Research Institute, Faisalabad. Fifteen seeds of each 
line of mungbean were sown in each of the plastic pots (24.5 cm 
diameter and 28 cm deep) filled with equal weight dry soil. Two 
drought stress treatments [Control (well-watered i.e. normal water-
ing as per requirement of the crop) and 60 % field capacity (FC)] 
were applied on 30-day old plants. The experiment was laid out in a 
completely randomized design with four replications of each experi-
mental unit. The weight of each plastic pots with filled soil and the 
water contents present at the time of sowing were already known, 
therefore moisture contents present in soil of pots equal to Control 
and 60 % FC were calculated for drought treatments. When water 
contents of each pot were at field capacity, then 15 seed of each line 
of pulse crop were sown. After 15 days of germination thinning of 
plants were done and 5 plants per pot were maintained. Plants were 
irrigated normally according to their requirements till 30 days before 



148 A. Afzal, I. Gulzar, M. Shahbaz, M. Ashraf

treatment start. After one month of normal growth of plants, drought 
stress levels were maintained. After 30 days of drought treatment, 
two plants were harvested from each of replicate, washed with dis-
tilled water and recorded data for shoot and root fresh weights and 
shoot and root lengths. The samples were oven-dried at 65 °C up 
to their constant weight and then dry weights recorded. In addition, 
data for following attributes were also recorded:

Gas Exchange Characteristics
A portable infra-red gas analyzer (IRGA) (ACD LCA-4 Analytical 
Development, Hoddesdon, UK) was used to determine the net pho-
tosynthetic rate (A), transpiration rate (E), stomatal conductance (g  s), 
water use efficiency (A/E), and internal CO2 concentration (Ci) on 
fully expanded leaves. Following adjustments/values of the instru-
ment were recorded/maintained during its operation: 403.3 mmol 
m-2 s-1 for molar flow of air, 99.9 kPa atmospheric pressure, 6.0 
to 8.9 mbar water vapor pressure, 1711 μmol m-2 s-1 PAR, 28.4 to 
27.9 °C leaf temperature and 352 μmol mol-1 ambient CO2 concen-
tration.

Water relation attributes
A fully expanded second leaf was excised at dawn and its mid rib 
was used in Scholander type pressure chamber (Arimad-2-Japan) to 
obtain water potential. The same leaf that used for water potential 
stored in freezer at -20 oC to use it for osmotic potential. After one 
week the frozen leaf was thawed and the sap was extracted by press-
ing it with glass rod. The extracted sap was used to determine the 
osmotic potential by using the osmometer (Wescor 5520). Turgor 
potential was calculated as the difference between water potential 
and osmotic potential. 

Chlorophyll fluorescence 
Before measurements of Chlorophyll fluorescence, the leaf samples 
were kept in dark for 30 min by using light-exclusion clips to the 
surface of leaves. Chlorophyll fluorescence was determined using 
an OS5p Modulator Fluorometer (ADC BioScientific Ltd, Great 
AmwellHerts, UK) according to STRASSER et al. (1995). 

Determination of mineral ions
The dried ground leaf or root material (100 mg) was digested with 
2 ml H2SO4 following the method of WOLF (1982). The volume of 
the extract was brought up to 50 ml with distilled water, filtered and 
used determining mineral elements. Leaf and root potassium (K+) 
and calcium (Ca2+) contents were determined using a flame pho-
tometer (Jenway, PFP-7, UK). Nitrogen was determined following 
the KJELDAHL method as described by BREMNER et al. (1965) while 
phosphorus was determined spectrophotometrically following meth-
od of JACKSON (1962).

Extraction of antioxidant enzymes
Antioxidant enzymes were extracted from fresh leaf samples (0.5 g 
each sample) in 10 ml of phosphate buffer (50 mM with pH 7.8) at 
4 ºC. The homogenate was then centrifuged at 12000 × g at 4 ºC for 
20 min and it was centrifuged again at 15000 × g for 10 min.  The 
supernatant was used for determining the activities of antioxidant 
enzymes. 

Superoxide dismutase (SOD)
The protocol described by GIANNOPOLITIS and RIES (1977) was 
used for the determination of SOD activity. It was determined as the 

enzyme ability to inhibit photochemical reduction of nitrobluetetra-
zolium (NBT). The 3 ml reaction mixture consisted of 50 mM phos-
phate buffer of 7.8 pH, distilled water, methionine 13 mM, 50 μM 
NBT, 50 μl enzyme extract and 1.3 μM riboflavin. The reaction solu-
tions were then kept under light (15 W fluorescent lamps) for 15 min 
at 78 μmol m-2 s-1. The absorbance of the reaction mixture was read 
at 560 nm with a UV-visible spectrophotometer (U2020 IRMECO). 
One unit activity of SOD was defined as the amount of enzyme re-
quired to cause 50 % inhibition of the rate of NBT photoreduction as 
compared to the sample that lacked the plant enzyme extract.

Activities of catalase (CAT) and peroxidase (POD)
The method described by CHANCE and MAEHLY (1955) was used to 
appraise the activities of CAT and POD on protein amount basis. The 
reaction solution for CAT contained phosphate buffer and H2O2 of 50 
and 5.9 mM, respectively. Addition of 0.1 ml enzyme extract to the 
reaction mixture initiated the reaction. After every 20 s the changes 
in the absorbance of the reaction mixture were observed at 240 nm. 
The reaction mixture for POD consisted of phosphate buffer, guai-
acol, and H2O2 with molar values as 50, 20 and 40 mM, respectively, 
and 0.1 ml of the enzyme extract. At 470 nm, the absorbance was 
taken after every 20 s. The enzyme activity was assessed on pro-
tein basis, while one unit of CAT considered equivalent to 0.01 units 
per min change in absorbance and one unit of POD defined as the 
0.01 units per min change in absorbance.

Determination of non-enzymatic antioxidants
Total phenolics
JULKENEN-TITTO (1985) proposed a method which was used to 
determine total phenolics. Leaf fresh material (50 mg) was homo-
genized in 80 % acetone. The homogenized material was centrifuged 
at 10,000 × g for 10 min, removed the pellet and the supernatant was 
used for the determination of phenolics. Then 100 μl of the super-
natant were mixed with 1 ml of Folin-Ciocalteau,s reagent. In ad-
dition, 2.0 ml distilled water and 5 ml of 20 % Na2CO3 were also 
added. The mixture was vortexed and absorbance read at 750 nm 
using a UV-Visible spectrophotometer (IRMECO U2020).

Leaf ascorbic acid contents 
The amount of ascorbic acid in the mungbean leaves was determined 
following MUKHERJEE and CHOUDHRI (1983). Fresh leaves (0.25 g) 
were ground in 10 ml of 6 % TCA. The mixture was centrifuged for 
10 min at 4 ºC at 1000 × g. An aliquot of 2 ml of 2 % dinitrophenyl 
hydrazine solution was added to 4 ml of supernatant. One drop of 
thiourea (10 % thiourea prepared in 70 % ethanol) was added to the 
mixture, and boiled the mixture for 20 min in a water bath.  The mix-
ture was placed in ice to reduce the temperature to about 25 ºC, then 
added 5 ml of 80 % sulphuric acid (v/v) at 0 ºC and the absorbance 
read at 530 nm. The ascorbic acid content was quantified against 
a standard curve which was prepared by known concentrations of 
ascorbic acid.

Estimation of leaf tocopherol content 
The method of BAKER et al. (1980) was used for the determination 
of leaf alpha-tocopherol concentration. A mixture of 20 ml of pe-
troleum ether and ethanol (2:1:6, v/v) was used to grind the fresh 
leaves (1.0 g each sample). The mixture was centrifuged at 10,000 × 
g for 20 min. An aliquot of 200 μl of 2 % 2, 2- dipyridyl (prepared 
in ethanol) was added to 1 ml of supernatant, mixed the mixture and 
placed it in the dark for 5 min. The absorbance was read at 520 nm 
using a spectrophotometer.



 Influence of water deficit conditions on mungbean 149

Yield attributes
Data for yield attributes like number of pods per plant, number of 
seed per plant, seed yield per plant and 100-seed weight were col-
lected at the maturity of the mungbean crop.

Statistical analysis
A two-way analysis of variance (ANOVA) of data for all attributes 
was calculated using the COSTAT computer program. 

Results
Imposition of water deficit conditions (normal watering and 60 % 
field capacity) significantly  reduced shoot and root fresh and dry 
weights of two mungbean lines (97001and 97012) (Tab. 1; Fig. 1). 
Both lines showed a significant variation in root dry weight as line 
97001 performed better than 97012, while in shoot fresh and dry 
weights and root fresh weight both lines showed a uniform behavior 
under both well-watered and drought stress conditions. 

Imposition of water deficit conditions markedly (P ≤ 0.001) re-
duced shoot length of the two mungbean lines i.e. 97001 and 97012 
(Tab. 1; Fig. 1). Of both lines, 97001 showed better performance in 
shoot length as compared to 97012. In root length, the influence of 
drought stress was not prominent for both mungbean lines and the 
lines did not show any variation in this attribute under water deficit 
conditions (Tab. 1; Fig. 1).
Leaf water potential increased while osmotic potential decreased 
prominently due to imposition of drought stress regimes (Tab. 1; 
Fig. 1). Imposition of water deficit conditions caused an increase in 
leaf turgor pressure (Fig. 1). However, both lines showed a uniform 
behavior with respect to water relation attributes under well-watered 
and drought stress conditions. 
Various drought stress regimes proved to be non-significant for 
photochemical quenching (qP), co-efficient of non-photochemical 
quenching (qN), non-photochemical quenching (NPQ) and efficiency 
of photosystem-II (Fv/Fm) in the two mungbean lines (97001 and 
97012). In addition, the lines did not show any difference with re-
spect to all these chlorophyll fluorescence attributes under various 

Tab. 1:  Mean squares from analyses of variance of data for morphological, physiological, biochemical and yield attributes of mungbean [Vigna radiata (L.) 
Wilczek] when 30 day old plants were subjected to drought stress

 SOV df Shoot f. wt. Shoot d. wt. Root f. wt. Root d. wt. Shoot length Root length Water Osmotic 
         potential potential

 Drought (D) 1 117.0** 2.772* 0.25* 0.024** 907.5*** 5.881ns 0.035* 0.453*

 Lines (L) 1 7.659ns 0.336ns 0.086ns 0.009* 247.3* 1.051ns 0.002ns 0.016ns

 D × L 1 14.35ns 0.235ns 0.242* 0.004ns 83.26ns 1.626ns 0.0002ns 0.022ns

 Error 12 11.68 0.466 0.045 0.002 28.83 4.027 0.004 0.050

 SOV df Turgor A E gs A/E Ci Ci/Ca Fv/Fm
   potential

 Drought (D) 1 0.695** 29.00*** 0.360* 0.013* 7.777ns 17835.6** 0.144ns 0.006ns

 Lines (L) 1 0.038ns 1.357ns 0.123ns 0.002ns 1.127ns 715.6ns 0.006ns 0.003ns

 D × L 1 0.019ns 0.601ns 0.240ns 0.003ns 13.20ns 1410.0ns 0.012ns 0.149ns

 Error 12 0.053 0.366 0.065 0.002 4.965 1559.1 0.013 0.039

 SOV df ETR NPQ qN qP Total Leaf Leaf ascorbic  CAT  
       phenolics tocopherols acid

 Drought (D) 1 69.72* 0.000ns 0.001ns 0.234ns 3.597** 0.039* 0.0005ns 11.22*

 Lines (L) 1 0.562ns 0.006ns 0.000ns 0.034ns 0.163ns 0.002ns 0.0001ns 25.49**

 D × L 1 1.440ns 0.000ns 0.000ns 0.730* 0.007ns 0.024ns 0.00002ns 8.390ns

 Error 12 8.642 0.017 0.006 0.091 0.295 0.005 0.0002 2.151

 SOV df POD SOD Total soluble MDA Leaf K+ Root K+ Leaf Ca2+ Root Ca2+
     proteins

 Drought (D) 1 0.899ns 0.291ns 4.155* 0.987ns 7.562ns 2.641ns 15.02** 1.891*

 Lines (L) 1 13.28ns 0.238ns 0.049ns 3.822ns 9.00ns 9.766ns 5.641ns 0.141ns

 D × L 1 429.4** 0.025ns 0.131ns 4.077ns 0.25ns 0.016ns 5.641ns 0.141ns

 Error 12 29.22 0.125 0.929 3.43 4.302 3.620 1.411 0.2243

 SOV df Leaf P Root P Leaf N Root N Number of Number of 100-seed Seed yield   
       pods/plant seeds/plant weight per plant

 Drought (D) 1 2.806* 0.226ns 0.111ns 0.005ns 37.82** 5681.4** 39.74ns 12.73**

 Lines (L) 1 0.64ns 0.331ns 0.162ns 0.011ns 0.01ns 606.4ns 34.47ns 0.022ns

 D × L 1 0.81ns 0.226ns 0.111ns 0.099ns 6.76ns 1113.9ns 133.7ns 0.228ns

 Error 12 0.546 0.374 0.183 0.046 3.262 413.6 69.31 0.972

*, **, *** = significant at 0.05, 0.01, and 0.001 (probability) levels, respectively
ns  =  non-significant



150 A. Afzal, I. Gulzar, M. Shahbaz, M. Ashraf

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Fig. 1:  Growth, water relations and chlorophyll fluorescence attributes of mungbean [Vigna radiata (L.) Wilczek] when 30 day old plants were subjected to 
drought stress (Mean ± S.E.; n = 4).

drought stress regimes (Tab. 1; Fig. 1 and 2). Water stress caused a 
slight decrease (P ≤ 0.05) in electron transport ratio (ETR) of both 
mungbean lines. The behavior of the lines was uniform under both 
well-watered and water deficit conditions. 
Water deficit conditions caused a significant decrease in net CO2 
assimilation rate, transpiration rate, stomatal conductance and sub-
stomatal CO2 concentration of the two mungbean lines (Tab. 1; 
Fig. 2).  However, water use efficiency and Ci/Ca ratio were not al-
tered by water deficit conditions (Tab. 1; Fig. 2). Both lines showed 
a uniform behavior with respect to all the above-mentioned gas ex-
change characteristics. 
Drought stress caused a slight increase (P ≤ 0.05) in leaf tocophe-
rol contents in the two mungbean lines while it did not affect leaf 
ascorbic acid (Tab. 1; Fig. 2). Total phenolic concentration decreased 
significantly (P ≤ 0.01) due to drought stress. Both mungbean lines 
showed a uniform behavior in all the earlier mentioned attributes 
under well watered and water deficit conditions. Imposition of water 

deficit conditions showed a non-significant effect on leaf MDA con-
tent and the variation between the two mungbean lines with respect 
to this attribute was also non-significant (Tab. 1; Fig. 3).
Activities of superoxide dismutase and peroxidase, and amount of 
total soluble proteins did not change due to water deficit conditions 
(Tab. 1; Fig. 3). Both mungbean lines did not show any variation 
under both well watered and water deficit conditions. The activity 
of catalase markedly (P ≤ 0.05) increased under water deficit condi-
tions. The lines also varied prominently as line 97001 showed more 
catalase activity than that of mungbean line 97012. 
Water deficit conditions did not alter leaf K+, N and root K+, P and 
N contents in two mungbean lines, while it caused a significant 
decrease in leaf Ca2+ and increase in leaf P and root Ca2+ in both 
mungbean lines under well watered and drought stress conditions 
(Tab. 1; Fig. 3). Both mungbean lines showed uniform response to 
water deficit conditions due to root and leaf mineral elements and 
they did not show prominent difference in various ion contents. 



 Influence of water deficit conditions on mungbean 151
 
 

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Fig. 2:  Gas exchange characteristics and antioxidants of mungbean [Vigna radiata (L.) Wilczek]  when 30 day old plants were subjected to drought stress (Mean 
± S.E.; n = 4).

Number of pods per plant, number of seeds per plant and seed yield 
per plant of two mungbean lines decreased significantly (P ≤ 0.01) 
due to imposition of drought stress (Tab. 1; Fig. 4). Water deficit 
conditions did not alter 100-seed weight prominently. Both mung-
bean lines did not show prominent difference with respect to yield 
attributes under various drought stress regimes.

Discussion
Crop growth and yield is adversely affected by drought stresss 
(ASHRAF, 2010), which is ascribed to drought-induced impairment 
in a number of metabolic processes involved in regulation of growth 
and development (CHAVES et al., 2009). Drought-induced growth re-
duction has been reported extensively in many crops like mungbean 
(ZARE et al., 2012), chicory (ASGHARI et al., 2009), barley (FATEH 
et al., 2012), wheat (SHAHBAZ et al., 2011b) etc. In the present study, 

change in growth was appraised in terms of change in plant height, 
which decreased markedly under drought stress conditions. These 
results are in agreement with some previous studies on mungbean 
(UDDIN et al., 2013), chickpea (SHAMSI, 2010), and Ocimum basili-
cum (ALISHAH et al., 2006). Such reduction in plant height under 
water deficit conditions was ascribed to reduced cell division, expan-
sion and elongation (HUSSAIN et al., 2008). 
The effect of drought stress on root growth is the matter of contro-
versy in view of a number of earlier published reports. In the present 
study, the root length showed a significant decrease under drought 
stress which does not conform to some earlier published reports on 
mungbean (RANAWAKE et al., 2011) as well as on Catharanthus ro-
seus (JALEEL et al., 2008). However, our results are in accordance 
with the findings of TERZI and KADIOGLU (2006) for Ctenanthe 
setosa in which a significant decrease in root length was observed. 
Water deficit conditions alter water relation parameters in most 



152 A. Afzal, I. Gulzar, M. Shahbaz, M. Ashraf

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Fig. 3:  Activities of antioxidants and mineral nutrients of mungbean [Vigna radiata (L.) Wilczek]  when 30 day old plants were subjected to drought stress 
(Mean ± S.E.; n = 4).

plants. For example, imposition of water deficit stress reduced the 
water potential and osmotic potential of plants of melon (KUS-
VURAN, 2012), wild jujube (MARAGHNI et al., 2011), and common 
bean (GULER et al., 2011; ZALVET et al., 2005). But in the present 
study, leaf water potential and osmotic potential did not change sig-
nificantly by imposition of drought stress while leaf turgor potential 
increased significantly in mungbean plants due to drought stress.
Drought stress caused a marked reduction in net CO2 assimilation 
rate (A) which is in agreement with many previous studies on differ-
ent crops, e.g. wheat (KAMRAN et al., 2009; SHAHBAZ et al., 2011b), 
sunflower (VANAJA et al., 2011), kidney bean (MIYASHITA et al., 
2005), grapevine (ZOSFI et al., 2008), and chickpea (RAHBARIAN 
et al., 2011). Stress-induced reduction in photosynthetic capacity is 
believed to be due to a number of factors including reduced leaf 
area, damage to photosynthetic machinery, initiation of leaf senes-
cence before maturity, lipid peroxidation in chloroplast and changes 

in pigment and protein composition (ANJUM et al., 2011). Reduction 
in A might also be due to reduction in Rubisco carboxylation ac-
tivity and regeneration of RuBP (LAWLOR and CORNIC, 2002). Fur-
thermore, low leaf water contents also leads to impaired metabolic 
system leading to less production of photo-assimilates (LAWLOR and 
CORNIC, 2002) under water deficit conditions. In the present study, 
drought stress slightly reduced the rate of transpiration (E) and sto-
matal conductance (gs), but did not affect water use efficiency (A/E) 
in mungbean plants. These results with respect to E and gs are in 
agreement with many previous studies on different crops e.g. barley 
(FATEH et al., 2012), forest plants (KEENAN et al., 2010), chickpea 
(MAFAKHERI et al., 2010), and rice (HALDER and BURRAGE, 2004) 
while contradictory with respect to A/E. Drought stress also caused 
a significant reduction in internal CO2 concentration (Ci) and Ci/Ca 
ratio in mungbean plants as has been earlier observed in soybean 
(ANJUM et al., 2011), forest plants (KEENAN et al., 2010) and wheat 



 Influence of water deficit conditions on mungbean 153

(BOGALE et al., 2011), but in contrast, these results do not agree with 
those for chickpea (MAFAKHERI et al., 2010) and rice (HALDER and 
BURRAGE, 2004). Furthermore, no significant relations were found 
between water deficit conditions and chlorophyll fluorescence at-
tributes. These results are analogous to what has been reported in 
chickpea (KHAMSSI et al., 2010) and olive (PETRIDIS et al., 2012). 
This might be due to low close relation of leaf water potential with 
chlorophyll fluorescence transients (JEFFERIES, 1992) and leaf water 
potential might have not affected these attributes. Decrease in A may 
also cause by the photodamage of PSII, but in the present study, 
drought stress did not affect the efficiency of PSII showing that light 
did not cause damage to PSII in mungbean under drought stress con-
ditions (BAKER and HORTON, 1987). Decrease in A is also related to 
stomatal conductance as both are reduced under drought stress con-
ditions, but these stomatal effects did not affect the efficiency of PSII 
(Fv/Fm) (FLEXAS and MEDRANO, 2002). Some coastal plants also 
maintained their Fv/Fm value even under drought stress conditions 
(DEMATTOS et al., 1997). In our study, drought stress did not cause 
any severe damage to PSII, and increased photorespiration might 
also be a factor causing protection to PSII from drought-induced ad-
verse effects (FLEXAS and MEDRANO, 2002).
The activity of enzymatic anti-oxidants, most specifically of super-
oxide dismutase (SOD), did not alter due to drought stress in 
mungbean plants which is analogous to the findings of TERZI and 
KADIOGLU (2006) in Ctenanthe setosa and VARGA et al. (2012) in 
wheat. Peroxidase (POD) and catalase (CAT) in mungbean plants 
also remained unchanged under water deficit conditions which were 
in accordance with the findings of GAMBLE and BURKE (1984) who 
reported a non-significant change in the activity of CAT in wheat 
under water deficit conditions. One of the possible reasons is that 
CAT has low affinity for ROS particularly for H2O2 as compared 
to ascorbate peroxidase (MITTLER, 2002). Furthermore, MDA and 
ascorbic acid (AsA) contents in mungbean plants also did not change 
under drought regimes. In our study, damage to membrane in the 
form of lipid peroxidation as shown by MDA contents is not se-

vere (non-significant due to drought stress) and this might be a main 
reason for the non-significant effect of drought stress on the acti-
vities of antioxidant enzymes. In addition, activities of antioxidant 
enzymes depend on a number of factors like plant age, drought stress 
duration, tolerance level of a particular species (DECARVALHO, 2008).  
Drawing correlation between induction of antioxidants and degree of 
drought tolerance among species of a same genus or even cultivars 
of a same species has been found not so easy (LOGGINI et al., 1999). 
These results are not in agreement with those reported for Withania 
somnifera (JALEEL et al., 2009) and tomato fruit (GHORBANLI et al., 
2012) under drought stress conditions but tocopherol was increased 
by the application of water stress, similar to that reported in canna 
cultivars (ZHANG et al., 2013). Increase in tocopherol under water 
deficit conditions might be due to active expression of genes which 
are responsible for the synthesis of tocopherols (ZHU, 2002; ZONG 
et al., 2009). TERZI and KADIOGLU (2006) observed an increase in 
MDA content during early phase of drought while a decrease in later 
phase in Ctenanthe setosa. Leaf phenolics in the present study de-
creased by the imposition of stress which is contrary to that found in 
maize (HURA et al., 2008) and olive (PETRIDIS et al., 2012), while in 
contrast, an increase in phenolic contents was reported in horsegram 
(BHARDWAJ and YADAV, 2012). 
Water deficit conditions are believed to generally reduce nutrient 
uptake in plants (BALIGAR et al., 2001) and plants with small or no 
reduction in nutrient uptake are considered drought resistant. How-
ever, great variation has been observed in nutrient uptake in even 
genotypes of a same species under drought stress conditions (SHAH-
BAZ et al., 2011a, b). In the present study, drought stress had a non-
significant effect on leaf and root N and K+ except P which decreased 
in mungbean plant roots under drought stress. Similar results have 
been observed in wheat (SHAHBAZ et al., 2011b), while in contrast, 
higher K+ accumulation has been reported in drought stressed com-
mon bean (ZADEHBAGHERI et al., 2012) and okra (KUSVURAN et al., 
2011) plants. A non-significant influence of drought stress was also 
observed on shoot and root K+ for Cenchrus ciliarus and Cyanodon 

&& & & & & & & & &
& & & & & & & & &
& & & & & & & & &
& & & & & & & & &
& & & & & & & & &
& & & & & & & & &
& & & & & & & & &
& & & & & & & & &
& & & & & & & & &
& & & & & & & & &
& & & & & & & & &
& & &
& & &
& & &
& & &
& & &
& & &
& & &
& & &
& & &
& & &
& & &

Fig. 4:  Leaf and root N and yield attributes of mungbean [Vigna radiata (L.) Wilczek]  when 30 day old plants were subjected to drought stress (Mean ± S.E.; 
n = 4).



154 A. Afzal, I. Gulzar, M. Shahbaz, M. Ashraf

dactylon (AKRAM et al., 2008). 
In conclusion, water deficit conditions significantly reduced all 
growth attributes, net CO2 assimilation rate, internal CO2 concentra-
tion, Ci /Ca ratio, root P, total soluble proteins, phenolics and seed 
yield, but they markedly increased turgor potential, and tocopherol 
contents. 

Acknowledgements
The last author gratefully acknowledges the funding from the Pa-
kistan Academy of Sciences (PAS)(Grant No. 5-9/PAS/7536). The 
results presented in this paper are a part of M. Sc. studies of Miss 
Aneela Afzal and Miss Iqra Gulzar.

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Address of the corresponding author:
E-mail: ashrafbot@yahoo.com