Protoplasma
DOI 10.1007/s00709-016-1037-0
ORIGINAL ARTICLE
Potassium up-regulates antioxidant metabolism and alleviates
growth inhibition under water and osmotic stress in wheat
(Triticum aestivum L)
Mohammad Abass Ahanger 1 & R. M. Agarwal 1
Received: 12 July 2016 / Accepted: 19 October 2016
# Springer-Verlag Wien 2016
Abstract Pot experiments were conducted to find out the ef-
fectivity of K on Triticum aestivum L cultivars. Polyethylene
glycol 6000 (PEG 6000) was used as an osmoticum to induce
osmotic stress under sand culture setting up the water potential
of external solution at −3 and −5 bars. In pots, plants were
raised under restricted and normal irrigation and K was applied
in varying doses (0, 20, 40, 60 kg ha−1) and estimation of
different physiological and biochemical parameters was done
at two developmental stages, i.e., preflowering and flowering.
Supplementation of K resulted in obvious increase in growth
and activity of antioxidant enzymes in both normal and stressed
plants. Added potassium increased total phenols and tannins
thereby strengthening the components of both the enzymatic
as well as non-enzymatic antioxidant system. Under both
normal and stressed conditions, K-fed plants experienced sig-
nificant increase in the synthesis of osmolytes like free proline,
amino acids, and sugars which assumes special significance in
growth under water stress conditions. Wheat plants accumulat-
ing greater K were able to counteract the water stress-induced
changes by maintaining lower Na/K ratio.
Keywords Antioxidants . Polyphenols . Osmolytes .
Potassium . Water stress . Triticum aestivum L
Handling Editor: Néstor Carrillo
Electronic supplementary material The online version of this article
(doi:10.1007/s00709-016-1037-0) contains supplementary material,
which is available to authorized users.
* Mohammad Abass Ahanger
ahangerma@gmail.com
1
School of Studies in Botany, Jiwaji University, Gwalior 474011,
India
Introduction
Water stress is one of the important environmental constraints
in arid climates and dry-land agricultural ecosystems which
influence plant survival. Effect of water stress on growth and
other physiological aspects and the mechanisms underlying
the tolerance strategies involving potassium has been studied
(Chaves and Davies 2010; Benlloch-Gonzalez et al. 2015).
Reduced availability of water, whether chronic or sporadic,
leads to reduction in growth and hence losses in yield (Jatav
et al. 2014). A set of morphological, physiological, and
biochemical changes are initiated in plants in response to wa-
ter deficit so as to withstand the existing conditions. Every
response pattern has a correlation with the modification of
yield and the physiological traits (Jatav et al. 2012, 2014).
In view of the importance of K in major physiological pro-
cesses, sufficiently large amounts of potassium must be
absorbed from the soil and subsequently transported to various
organs. For example, changes in the ionic concentration, espe-
cially cations of xylem sap can bring change in the volume of
pectin pit membranes resulting in modulations in the hydraulic
conductivity (Zwieniecki et al. 2001). In most areas of the
globe, available soil K is either low or is considerably fluctuat-
ing. In order to meet the demand-driven needs, plants have
evolved strategies e.g., enhancement in the capacity for high-
affinity K+ uptake from the soil, redistribution of absorbed K+
between the cytosolic and vacuolar pools for ensuring cytosolic
homeostasis, and alterations in the developmental pattern of
root system. These mechanisms contribute to efficient K uptake
for preserving the vital functions and hence ensuring better
growth when K availability in soil becomes a limiting factor
(Cherel et al. 2014). K availability is also influenced by the
complex soil dynamics as well as the root-soil interactions.
Low K status triggers expression of high affinity K transporters
and associated signaling cascades. Reactive oxygen species

(ROS) molecules and phytohormones like auxin, ethylene, and
jasmonic acid may have important role in sensing the K
deficiency (Ashley et al. 2006).
Enhanced hydraulic conductivity due to K application can
benefit the plant in maintaining the turgor, stomatal move-
ments, and gas exchange rate as has been observed in laurel
plants (Oddo et al. 2012). K deficiency obstructs plant metab-
olism and decreases chlorophyll content and the photosynthet-
ic efficiency (Tiwari et al. 1998) by affecting the activity of
ribulose 1,5-bisphosphate carboxylase/oxygenase and the as-
sociated photosynthetic attributes (Weng et al. 2007).
Improved K uptake has been recognized to contribute in mit-
igating the oxidative stress by modulating ROS metabolism
(Soledad et al. 2015) and improving K/Na ratio (Jatav et al.
2014; Ahanger et al. 2015).
Exposure of plants to various environmental stresses ac-
crue production of ROS-triggering oxidation of proteins, chlo-
rophylls, lipids, nucleic acids, carbohydrates, impairment of
enzyme activity, etc. (Mittler 2002; Ahmad et al. 2010;
Demidchik 2015), inducing anomalies in several important
cellular biochemical pathways operating in cellular organelles
like chloroplast and mitochondria (Miyake 2010). ROS acti-
vate calcium (Ca2+) and potassium (K+) permeable cation
channels at the plasma membrane, mediating Ca2+-based sig-
naling events, K+ ion leakage, and programmed cell death
(Pottosin et al. 2012; Shabala and Pottosin 2014). Both enzy-
matic and non-enzymatic components of antioxidant defense
system are closely coordinated to each other for neutralizing
the ROS (Ahmad et al. 2010). Plants up-regulate the activity
of antioxidant enzymes, superoxide dismutase (SOD),
catalase (CAT), ascorbate peroxidase (APX), and glutathione
reductase (GR) and the production of non-enzymatic antioxi-
dants, i.e., reduced glutathione (GSH), ascorbic acid (AsA),
and phenols (Lee et al. 2015).
Plants tend to accumulate low molecular weight
compounds such as free amino acids, free proline, non-
structural carbohydrates, and quaternary ammonium
compounds including betaines to cope up with stressful con-
ditions. Accumulation of these compounds leads to reduction
in the osmotic potential by helping in restoration and mainte-
nance of the potential gradient between the plant cell and
external soil solution. Puniran-Hartley et al. (2014) have
observed a strong correlation between enhanced accumulation
of compatible osmolytes and oxidative damage tolerance in
salt-stressed wheat and barley seedlings. In addition to the
compatible organic solutes, ions like K also contribute signif-
icantly to the maintenance of tissue water content and
deficiency of K increases stress susceptibility (Jatav et al.
2014). Nio et al. (2011) have demonstrated that K+ contributes
considerably towards osmoticum in water-stressed wheat
cultivars with low osmotic adjustment potential. Plants grow-
ing in arid and semi-arid regions of globe exhibit symptoms of
dehydration that have been accepted to be associated with low
M.A. Ahanger, R.M. Agarwal
potassium status of the agricultural lands. K improves growth
and yield performance of many crops (Tiwari et al. 1998;
Umar 2006; Sharma et al. 2006; Jatav et al. 2012, 2014).
In fact, K is known for its role in water relations and pho-
tosynthesis, nevertheless, reports regarding K protecting
plants under water stress are scanty, particularly related with
the antioxidant metabolism. K being a dominant cation can
restore plant growth under stress by altering antioxidant me-
tabolism, nitrogen assimilation, and accumulating the
osmotica. Following experiments were conducted on two
cultivars of wheat to test this hypothesis.
Material and methods
Plant material and growth conditions
Two wheat (Triticum aestivum L) cultivars (viz LOK-1 and
MP-1203) were procured from Rajmata Vijayraje Scindhia
Krishi Vishwa Vidhyalaya (RVSKVV), Gwalior for the pres-
ent study. Seeds were surface sterilized with 0.01 % HgCl2
followed by repeated washings with distilled water, and ex-
periments were conducted under following setup:
Pot experiments using soil substrate
Bottom-perforated pots were filled with garden soil (sandy
clay loam) having slightly alkaline pH and 0.1366 dS m−1
conductivity with native concentration of K as 13 mg g−1 soil.
Recommended doses of nitrogen (in the form of urea) and
phosphorous (single super phosphate) fertilizers (120 and
60 kg ha−1, respectively) and varying doses of K (in the form
of K2O), i.e., 0, 20, 40, and 60 kg ha −1 were applied.
Replicates receiving only nitrogen and phosphorous but no
potassium (0 kg ha−1) served as control. Water stress was
imposed by withholding irrigation at vegetative stage and dur-
ing rainfall transparent polythene was used as rain-out shelter
to impose water stress. Flag leaves were analyzed at two
developmental stages, i.e., preflowering and flowering.
Pot experiments using sand substrate
Screening experiments were conducted (Table 1) to ascertain
the effective concentrations, and based on that, 1 and 10 mM
K2O with and without polyethylene glycol B6000^ were sub-
sequently used. Surface-sterilized seeds were sown in 18 cm
diameter pots filled with acid-washed sand. After germination,
ten seedlings were maintained in each pot and were fed with
200 mL modified full-strength Hoagland’s solution with 0, 1,
and 10 mM K on alternate days. Ten-day-old seedlings were
exposed to water stress for another 15 days using PEG 6000 at
−3 and −5 bars water potential (Michel et al. 1983). PEG 6000
was prepared in freshly prepared Hoagland nutrient solution,
Potassium up-regulates antioxidant metabolism

Table 1 Growth, photosynthetic pigments, and nitrate reductase activity in wheat (Triticum aestivum L cultivar LOK-1) seedlings grown with different
potassium treatments (7 DAS)

Characteristics T. aestivum L cultivar LOK-1

Control 1 mM K 10 mM K′ 50 mM K′ 100 mM K

Shoot length (cm) 7.36±0.28c 8.40±0.06b 11.50±0.26a 7.60±0.06c 4.80±0.20d

Root length (cm) 12.58±0.75b 12.29±0.39b 13.51±0.62a 09.63±0.24c 06.73±0.51d
Shoot fresh weight (g) 0.392 ±0.007c 0.416 ±0.015b 0.463 ±0.028a 0.391 ±0.021c 0.278 ±0.024d
Root fresh weight (g) 0.372 ±0.007c 0.460 ±0.05a 0.425 ±0.04ab 0.288 ±0.02d 0.174 ±0.01e
Shoot dry weight (g) 0.0467 ±0.002c 0.0504 ±0.002b 0.0535 ±0.002a 0.0450 ±0.003c 0.0340 ±0.002d
Root dry weight (g) 0.0271 ±0.001b 0.0274 ±0.001b 0.0295 ±0.001a 0.0252 ±0.002c 0.0178 ±0.004d
Chlorophyll a (mg g−1 FW) 0.5748±0.008c 0.6248±0.007b 0.6731±0.015a 0.5802±0.024c 0.3968±0.029d
Chlorophyll b (mg g−1 FW) 0.3790±0.025c 0.4189±0.015b 0.4741±0.032a 0.4105±0.044b 0.2395±0.006d
Total chlorophylls (mg g−1 FW) 0.9367±0.027d 1.0175±0.026b 1.1491±0.021a 0.9967±0.044c 0.6207±0.044e
Carotenoids (mg g−1 FW) 0.2642±0.014c 0.2923±0.011b 0.3497±0.020a 0.2733±0.008c 0.1890±0.009e
Shoot nitrate reductase activity 0.6219±0.033cd 0.9453±0.177b 1.0131±0.033a 0.6344±0.061c 0.3736±0.016e
(μM NO2− released h−1 g−1 FW)
Root nitrate reductase activity 0.5416±0.025c 0.7272±0.051b 0.8325±0.073a 0.5097±0.041d 0.3410±0.006e
(μM NO2− released h−1 g−1 FW)

Data presented is mean of four replicates (±SE). Data followed by same letter are not significantly different at p < 0.05

and plants were grown in different K- and PEG 6000-induced
water stress treatments independently and in conjugation viz;
1 mM K2O, 10 mM K2O, −3 bars, −3 bars + 1 mM K2O, −3
bars + 10 mM K2O, −5 bars, −5 bars + 1 mM K2O, and −5
bars + 10 mM K2O maintaining a control having no K and
PEG 6000 and different parameters as outlined below were
worked out at 25 DAS.
Relative water content
Relative water content (RWC) of leaf was measured in fully
expanded leaves following Weatherley’s (1950) method, and
calculation was done using the formula given below:
FW−DW
RWC ð%Þ ¼  100
TW−DW
Determination of photosynthetic pigments
Fresh leaves were macerated in 80 % acetone (v/v), and optical
density of the supernatant was recorded at 645, 663, and 480
−nm (Arnon 1949).
Determination of lipid peroxidation and protease activity
Lipid peroxidation was determined by measuring the
malondialdehyde (MDA) formation. Absorbance of the
supernatant was read at 532 and 600 nm, and MDA content
was calculated using an extinction coefficient (ε) of
155 mM−1 cm−1 (Heath and Packer 1968). For estimation of
protease (EC 3.4.21.40) activity, method of Green and
Neurath (1954) was adopted and absorbance was taken at
660 nm and activity was expressed as micrograms of tyrosine
released per milligram of protein.
Assay of antioxidant enzymes
Fresh plant tissue was homogenized in chilled extraction mix-
ture (100 mM, pH 7.8) containing PVP (0.1 %) and EDTA
(0.5 mM) using chilled mortar and pestle. Homogenate was
centrifuged at 12,000×g for 15 min at 4 °C, and supernatant
was used for enzyme assay. For extraction of APX, extraction
buffer was supplemented with 2 mM ascorbate.
SOD (EC 1.15.1.1) activity was estimated by monitoring
the ability of enzyme extract to inhibit the photochemical
reduction of NBT (Dhindsa et al. 1981). Absorbance was
recorded at 560 nm, and the amount of protein causing 50 %
photoreduction was considered 1 unit of enzyme activity and
expressed as enzyme unit (EU) per milligram of protein.
Activity of CAT (EC 1.11.1.6) was measured by monitor-
ing the disappearance of H2O2 at 240 nm for 2 min. Activity
was calculated using extinction coefficient (ε) of
0.036 mM−1 cm−1 and was expressed as EU per milligram
of protein (Aebi 1984).
APX (EC 1.11.1.1) was spectrophotometrically assayed in
accordance with Nakano and Asada (1981). H2O2-dependent
oxidation of ascorbate was followed by change in the absor-
bance at 290 nm. APX activity was calculated using extinction
coefficient (ε) of 2.8 mM−1 cm−1 and expressed as EU per
milligram of protein.
GR (EC 1.6.4.2) was assayed by monitoring the glutathi-
one dependent oxidation of nicotinamide adenine dinucleotide

phosphate (NADPH) at 340 nm (Foyer and Halliwell 1976).
GR activity was calculated using extinction coefficient of
6.2 mM−1 cm−1 and expressed as EU per milligram of protein.
Estimation of ascorbic acid and reduced glutathione
Ascorbic acid content was determined spectrophotometrically
following Omayl et al. (1979). GSH was determined by the
method of Moron et al. (1979). Computations were done
using standard curves of AsA and GSH for estimation of
AsA and GSH, respectively.
Determination of phenylalanine ammonia lyase activity
and contents of phenols and tannins
Activity of phenylalanine ammonia lyase (PAL) was estimated
following Zucker (1965). Optical density was recorded at
290 nm, and the activity was expressed as nanomolars of
trans-cinnamic acid formed per minute per milligram of protein.
Total phenols were extracted in 80 % ethanol Malick and
Singh’s (1980) method, and concentration of phenols was
expressed in milligrams per gram weight, equivalent to
catechol. Tannins were estimated in accordance with the
method of Swain and Hills (1959) using Folin-Denis reagent
in alkaline medium and reading absorbance at 700 nm. Tannin
content was expressed as milligrams per gram weight tannic
acid equivalents.
Total antioxidant activity
The free radical scavenging activity of methanolic plant
extract was measured using the method described by
Shimada et al. (1992) using 1,1-diphenyl-2-picrylhydrazyl
(DPPH). Absorbance was recorded at 517 nm, and the activity
was expressed as the percentage of DPPH scavenging.
Estimation of free proline, amino acids and total sugars
Free proline estimation was done following Bates et al.
(1973). Free amino acids were determined following the
method outlined in Sadasivam and Manickam (2004). Total
free sugars were estimated using anthrone method (Fong et al.
1953; Jain and Guruprasad 1989).
Estimation of potassium and sodium
K and Na were estimated flame photometrically after
digesting dry plant material in tri-acid mixture (H2SO4 +
HNO3 + HClO4 in 9:3:1 ratio) and reading on a digital flame
photometer employing K and Na filters separately and using
standard curves of K and Na for computation.
M.A. Ahanger, R.M. Agarwal
Statistical analysis
Data presented is the mean of four replicates with
standard error (±SE) calculated. Data was statistically
analyzed for ANOVA using SPSS software version 17
and significant difference was calculated at 0.05 % level
of significance. Data followed by same letter are not sig-
nificantly different by LSD test at p < 0.05.
Results
Effect of K supplementation on photosynthetic pigments
Photosynthetic pigments were reduced in plants subjected
to stress conditions (Table 2). K supplementation as-
suaged the deleterious effects of the stress to considerable
extent. Under normal irrigation, K enhanced the chloro-
phyll pigments which reached maximum in plants
supplied with higher potassium dose (60 kg ha−1), and
this trend was observed in the plants maintained under
restricted irrigation as well and at both the developmental
stages (Table 2).
K-fed wheat cultivars exhibit reduced oxidative damage
Lipid peroxidation measured in terms of MDA content in-
creased with the intensity of stress under both experimental
setup (Figs. 1 and 2). In sand cultures, seedlings raised with
solutions of lower water potential (−5 bars) exhibited in-
creased lipid peroxidation (Fig. 1). Seedlings raised with K
alone showed significant reduction in the lipid peroxidation,
and K also mitigated the deleterious impact of both osmotic
and water stress to some extent (Figs. 1 and 2). In pot exper-
iments, positive effect of K application was apparent by in-
creasing K supply from 20 to 60 kg ha−1, and this positive
influence of added K was maintained even under restricted
irrigation regime (Fig. 2). Application of K at 60 kg ha−1 re-
duced lipid peroxidation maximally both under normal as well
as water deficit conditions (Fig. 2). K-deficient plants, under
normal and deficit irrigation conditions, exhibited significant-
ly increased lipid peroxidation as compared with the K-fed
counterparts (Fig. 2).
In sand culture setup, PEG 6000-induced osmotic
stress up-regulated the activity of protease (Fig. 3).
Observed values for protease activity in plants receiving
1 and 10 mM K showed a reduction of 9.6 and 37.7 %,
respectively, as compared with control plants (Fig. 3).
Ten-millimolar K proved to be effective with PEG 6000
by reducing the protease activity by 20.6 and 7.4 % at −3
and −5 bars, respectively (Fig. 3).
Table 2 Chlorophyll and carotenoid (mg g−1 FW) content in wheat (Triticum aestivum L cultivar LOK-1 and MP-1203) grown under normal (N) and restricted (S) irrigation and different potassium doses
at different developmental stages

Cultivar K levels (kg ha−1) Irrigation regime Photosynthetic pigments

Preflowering Flowering
Potassium up-regulates antioxidant metabolism

Chlorophyll a Chlorophyll b Total chlorophyll Carotenoids Chlorophyll a Chlorophyll b Total chlorophyll Carotenoids

LOK-1 0 N 0.672±0.024d 0.385±0.030f 0.990±0.016d 0.344±0.003i 0.948±0.04d 0.601±0.034b 1.569±0.06d 0.620±0.038e

S 0.517±0.010h 0.317±0.002i 0.825±0.015ef 0.313±0.004k 0.720±0.03f 0.434±0.034d 1.166±0.10f 0.476±0.005g
20 N 0.680±0.032c 0.408±0.008e 1.101±0.036b 0.367±0.003h 1.100±0.06b 0.635±0.016b 1.810±0.09b 0.729±0.037c
S 0.523±0.015h 0.362±0.010g 0.859±0.027e 0.336±0.008j 0.812±0.04e 0.490±0.007cd 1.256±0.05g 0.554±0.037f
40 N 0.748±0.008b 0.494±0.021b 1.271±0.138a 0.391±0.007f 1.321±0.04a 0.708±0.007a 2.000±0.06a 0.795±0.003b
S 0.666±0.004d 0.418±0.008e 1.195±0.02ab 0.349±0.004i 0.958±0.03d 0.539±0.025c 1.506±0.04e 0.595±0.004e
60 N 0.855±0.013a 0.560±0.017a 1.271±0.049a 0.437±0.006de 1.394±0.03a 0.735±0.005a 2.164±0.04a 0.862±0.004a
S 0.691±0.020c 0.450±0.007c 1.002±0.004c 0.378±0.027g 1.004±0.004c 0.605±0.008b 1.625±0.11c 0.643±0.021d
MP-1203 0 N 0.529±0.046h 0.334±0.004h 0.838±0.03ef 0.518±0.013d 0.828±0.025e 0.399±0.009de 1.082±0.053h 0.518±0.013g
S 0.463±0.010i 0.300±0.012j 0.668±0.01g 0.421±0.03e 0.564±0.038g 0.338±0.010e 0.909±0.049i 0.421±0.037h
20 N 0.561±0.042g 0.369±0.012g 0.933±0.03d 0.585±0.017c 0.857±0.026e 0.484±0.011c 1.338±0.042e 0.585±0.017ef
S 0.478±0.041i 0.329±0.003h 0.698±0.008g 0.459±0.01d 0.707±0.006f 0.378±0.006e 1.041±0.061h 0.459±0.016g
40 N 0.653±0.048e 0.427±0.016d 1.037±0.04c 0.668±0.012b 1.001±0.003c 0.504±0.005c 1.586±0.039d 0.668±0.012d
S 0.524±0.016h 0.371±0.012f 0.888±0.068e 0.537±0.02cd 0.792±0.035f 0.419±0.006d 1.089±0.009h 0.537±0.023f
60 N 0.643±0.052e 0.470±0.006c 1.116±0.01b 0.728±0.009a 1.034±0.060c 0.591±0.020bc 1.682±0.049c 0.728±0.009c
S 0.585±0.009f 0.424±0.008d 0.984±0.01d 0.599±0.008c 0.902±0.004d 0.469±0.004d 1.331±0.044e 0.599±0.008e

Data presented is mean of four replicates (±SE). Data followed by same letter are not significantly different at p < 0.05

Fig. 1 Lipid peroxidation (μM MDA g−1 FW) in leaves of wheat
(Triticum aestivum L cultivar LOK-1) plants grown under PEG 6000-
induced osmotic stress and potassium treatments (25 DAS). Data
followed by same letter are not significantly different at p < 0.05
Influence of K on enzymatic and non-enzymatic
components of antioxidants
T. aestivum L subjected to water/osmotic stress showed up-
regulation of the enzymatic antioxidant defense system, and
Fig. 2 Lipid peroxidation (μM MDA g−1 FW) in flag leaf of wheat
(Triticum aestivum L cultivars LOK-1 and MP-1203) grown under
normal (N) and restricted (S) irrigation and different potassium doses at
preflowering (a) and flowering (b) stages. Data followed by same letter
are not significantly different at p < 0.05
M.A. Ahanger, R.M. Agarwal
Fig. 3 Protease (μg tyrosine released mg−1 protein) activity in leaves of
wheat (Triticum aestivum L cultivar LOK-1) seedlings grown under PEG
6000-induced osmotic stress and potassium treatments (25 DAS). Data
followed by same letter are not significantly different at p < 0.05
cultivar LOK-1 maintained higher activity of antioxidant en-
zymes (Figs. 4 and 5). Water-stressed plants exhibited higher
activity of antioxidant enzymes (SOD, CAT, APX, GR) as
compared with the normally irrigated counterparts, and added
potassium enhanced the activity further both under normal as
well as stressed conditions at both the developmental stages;
however, cultivar difference was apparent (Fig. 5a–h). In
water-stressed plants, SOD, CAT, APX, and GR increased
by 21.8, 5.02, 13.1, and 15.02 % in LOK-1 and 22.2, 6.32,
12.5, and 17.3 % in MP-1203; however, supplementation of
60 kg ha−1 K further enhanced the activity of SOD, CAT,
APX, and GR by 51.06, 45.5, 79.9, and 60.7 % in LOK-1
and 59.84, 54.01, 45.1, and 58.78 % in MP-1203 (Fig. 5a–h).
Similar trend was observed in sand cultures (Fig. 4).
Water/osmotic stress enhanced AsA and GSH content at
both the developmental stages. Increase was greater in K treat-
ments both under normal and restricted irrigation conditions
(Figs. 6a, b and 7c, d).
K supplementation enhances metabolite fraction and PAL
activity
Water/osmotic-stressed seedlings exhibited increased content
of phenols (Fig. 8a). An obvious increase in PAL activity was
observed in seedlings treated with K (Fig. 8b). K (1 and
10 mM) caused an increase of 25.4 and 56.1 % in PAL activ-
ity, and the increase was also observed when K was applied to
the seedlings exposed to water/osmotic stress using PEG 6000
(Fig. 8b). Total phenols and tannins increased in plants ex-
posed to water stress and K treatments under normal as well
as restricted irrigation regime (Fig. 9a–d). At flowering stage,
an increase of 32.0 and 47.8 % in total phenols and 30.9 and
24.6 % in tannins was observed in LOK-1 and MP-1203 cul-
tivars, respectively, when K was applied to water-stressed
plants at the rate of 60 kg ha−1 (Fig. 9a–d).
Potassium up-regulates antioxidant metabolism

Fig. 4 Superoxide dismutase,

catalase, ascorbate peroxidase and
glutathione reductase (EU mg−1
protein) activity in leaves (a–d) of
wheat (Triticum aestivum L
cultivar LOK-1) seedlings grown
under PEG 6000-induced osmotic
stress and potassium treatments
(25 DAS). Data followed by same
letter are not significantly
different at p < 0.05

Effect of K on total antioxidant activity
K caused significant increase in total antioxidant activity an-
alyzed as the DPPH radical scavenging potential. Osmotic
stress (−3 and −5 bars) resulted in enhancement in DPPH
radical scavenging activity, and K supplementation increased
it further (Fig. 10).
Effect of K on osmotic constituents
At flowering stage, maximum percent increase in free proline,
free amino acids, and free sugars under normal irrigation was
25.3, 45.6, and 25.2 % in LOK-1 and 25.7, 29.2, and 42.7 %
in MP-1203, respe,tively, when K was applied at the rate of
60 kg ha−1. However, when K (60 kg ha−1) was applied to
water-stressed plants, free proline, free amino acids, and free
sugars increased by 32.5, 50.9, and 31.7 % in LOK-1 and
40.7, 33.6, and 43.9 % in MP-1203, respectively (Fig. 11a–f).
Effect of water stress on K and Na
Water stress caused considerable decline in the uptake of K
and Na (Tables S1 and S2). K supplementation resulted in
significant reduction in Na content (Table S2). Both the culti-
vars accumulated higher content of K in flag leaf as compared
with stem and root (Table S1). Application of K resulted in
substantial reduction of Na in flag leaf than in root; however,
maximum effectivity was observed in plants supplemented
with 60 kg ha−1 K. Under normal irrigation, at flowering
stage, Na in flag leaf was reduced by 22.8 % in LOK-1 and
16.4 % in MP-1203, respectively (Table S2), therefore causing
significant reduction in Na/K ratio (Table 3).
Effect on leaf area, RWC and yield attributes
Added K (60 kg ha−1) exerted a positive influence on growth
under normal irrigation regime, and the effect was also
apparent under restricted irrigation (Table 4). K-treated plants
maintained higher water contents and leaf area. Enhancing K
application caused a significant increase in spike length, num-
ber of spikelets, and grain yield in both the cultivars (Table 4).
Discussion
Potassium has a critical role in maintaining the cellular func-
tioning of plants, although it does not form part of any meta-
bolic product, but its deficiency causes obvious effects on
growth and physiological attributes of plants (Tiwari et al.
1998; Sharma and Agarwal 2002; Sharma et al. 2006; Umar
2006; Jatav et al. 2014; Ahanger et al. 2015).
Photosynthetic pigments registered a decrease in plants
subjected to stressed conditions. However, supplementation
of K not only enhanced the synthesis of pigments but also
assuaged the deleterious effects of the stress to considerable
extent. K supplementation enhanced the cholorphyll pigments
which reached maximum in plants supplied with higher K
dose (60 kg ha−1) under normal irrigation, and such a trend
was found even in the plants maintained under restricted irri-
gation at both the developmental stages indicating the role of
K in improving the photosynthetic efficiency of wheat
 M.A. Ahanger, R.M. Agarwal

Fig. 5 Superoxide dismutase,

catalase, ascorbate peroxidase and
glutathione reductase
(EUp < 0.05mg−1 protein) activity
in flag leaf of wheat (Triticum
aestivum L cultivars LOK-1 and
MP-1203) grown under normal
(N) and restricted (S) irrigation
and different potassium doses at
preflowering (a–d) and flowering
(e–h) stages. Data followed by
same letter are not significantly
different at p < 0.05

cultivars. Role of K in maintaining the chlorophyll biosynthet-
ic rate can be an important step in K-induced tolerance in
plants. Reduction in chlorophyll synthesis in water-stressed
seedlings of rice has been attributed to decreased synthesis
of intermediates like glutamate-1-semialdehyde (GSA) and
5-aminolevulinic acid which can alter the gene expression of
certain enzymes involved in chlorophyll biosynthesis (Dalal
and Tripathy 2012). Reduced cholorphyll content in stressed
plants/seedlings has direct influence on the synthesis of
photoassimilates and growth. K supplementation mitigated
the adverse effects of water and osmotic stress considerably
by enhancing cholorphyll content. Stresses alter the function-
ing of pigment protein complex and cause hindrance in the de
novo synthesis of proteins and associated pigment molecules
(Murata et al. 2007).
Lipid peroxidation measured in terms of MDA content
formation increased with the intensity of stress. K treatments
significantly reduced the formation of MDA, mitigating the
deleterious impact of both water/osmotic stress to some
extent. Peroxidation of lipids is one of the key effects of
stress-induced oxidative damage. Increased lipid peroxida-
tion, a direct effect of ROS on polyunsaturated components
of membranes, affects membrane stability resulting in leakage
and hence cell damage (Ahmad et al. 2010; Gill and Tuteja
2010). ROS causes base modification in DNA, and ROS-
induced lipid peroxidation triggers formation of exocyclic
Potassium up-regulates antioxidant metabolism

Fig. 6 Ascorbic acid (mg g−1 FW) and reduced glutathione (nM g−1 FW)
content in leaves (a, b) of wheat (Triticum aestivum L cultivar LOK-1)
seedlings grown under PEG 6000-induced osmotic stress and potassium
treatments (25 DAS). Data followed by same letter are not significantly
different at p < 0.05
Fig. 8 Total phenol (mg g−1 FW) content and phenyl alanine ammonia
lyase (nM trans-cinnamic acid min-1 mg-1 protein) activity in leaves (a-b)
of wheat (Triticum aestivum L cultivar LOK-1) seedlings grown under
PEG 6000-induced osmotic stress and potassium treatments (25
DAS). Data followed by same letter are not significantly different at
p < 0.05

Fig. 7 Ascorbic acid (mg g−1

FW) and reduced glutathione
(nM g−1 FW) content in flag leaf
of wheat (Triticum aestivum L
cultivars LOK-1 and MP-1203)
grown under normal (N) and
restricted (S) irrigation and
different potassium doses at
preflowering (a, b) and flowering
(c, d) stages. Data followed by
same letter are not significantly
different at p < 0.05

Fig. 9 Total phenol (mg g−1 DW)
and tannin (mg g−1 DW) content
in flag leaf of wheat (Triticum
aestivum L cultivars LOK-1 and
MP-1203) grown under normal
(N) and restricted (S) irrigation
and different potassium doses at
preflowering (a-b) and flowering
(c-d) stages. Data followed by
same letter are not significantly
different at p < 0.05
DNA adducts, leading to induction of endogenous lesions (del
Maestro et al. 1981; Tuteja et al. 2001). Products of lipid
peroxidation include strong nucleophiles that attack DNA
and result in the formation of mutagenic adducts (Marnett
1999). Reduced lipid peroxidation due to K application is
indicative of reduced ROS accumulation in potassium-
treated plants, resulting in enhanced membrane stability for
the optimal functioning of cell. In Vicia faba L, Siddiqui
et al. (2012) have demonstrated reduced oxidative stress in
K-treated plants. Supplementing K salts improve growth by
providing stability to membranes through up-regulating the
antioxidant defense system (enzymatic as well as non-
enzymatic) and maintenance of tissue water content
Fig. 10 Total antioxidant activity in leaves of wheat (Triticum aestivum L
cultivar LOK-1) seedlings grown under PEG 6000-induced osmotic
stress and potassium treatments (25 DAS). Data followed by same letter
are not significantly different at p < 0.05
M.A. Ahanger, R.M. Agarwal
(Ahanger et al. 2015). The present study indicates that K can
be instrumental in preventing the oxidative stress-induced
damage to membrane lipids through its obvious role in main-
taining the activity of the antioxidant enzymes. Seedlings ex-
posed to PEG 6000-induced osmotic stress exhibited higher
activity of protease whereas, K alone as well as applied to the
seedlings/plants experiencing PEG 6000-induced stress
showed reduced activity of protease. Stress-induced proteoly-
sis may also be associated with the oxidative damage caused
by the excessive production of ROS. Proteases mediate the
removal of misfolded and mistargeted proteins (Kidric et al.
2014). Pandey et al. (2004) and Hameed et al. (2011) have
also reported increased protease activity in water-stressed rice
and wheat, respectively. K supplementation down-regulated
the activity of protease, suggesting the involvement of
potassium in enhancing protein stability through their proper
folding and subsequent targeting indicating role of K in
modulating protease activity under salt and osmotic stress.
T. aestivum L subjected to water and osmotic stress showed
up-regulation of the enzymatic antioxidant defense system
which was further enhanced by K supplementation. LOK-1
was more responsive to K, maintaining higher activity of
antioxidant enzymes and the contents of GSH and AsA.
Water-stressed plants exhibited higher activity of antioxidant
enzymes, SOD, CAT, APX, GR, and GPX and contents of
GSH and AsA. Potassium supplementation caused enhance-
ment in the ascorbic acid content with maximal increase
noticed in water-stressed plants treated with higher K dose
(60 kg ha−1). Plants (both at preflowering and flowering stages)
raised under normal as well as restricted irrigation exhibited
increased GSH accumulation with the application of K. GSH
and AsA are the ubiquitous components of the ascorbate-
Potassium up-regulates antioxidant metabolism

Fig. 11 Free proline, free

aminoacids and total free sugars
(mg g−1 DW) in flag leaf of wheat
(Triticum aestivum L cultivars
LOK-1 and MP-1203) grown
under normal (N) and restricted
(S) irrigation and different
potassium doses at preflowering
(a–c) and flowering (d, e) stages.
Data followed by same letter are
not significantly different at
p < 0.05

Table 3 Na/K ratio of wheat

(Triticum aestivum L cultivars Cultivar K levels Irrigation Na/K ratio
LOK-1 and MP-1203) grown (kg ha−1) regime
under normal (N) and restricted Preflowering Flowering
(S) irrigation and different
potassium doses at preflowering Flag Stem Root Flag Stem Root
and flowering stage (Data based leaf leaf
on tables S1 and S2)
LOK-1 0 N 0.131 0.099 0.3471 0.1021 0.0920 0.263
S 0.154 0.103 0.396 0.0958 0.1167 0.278
20 N 0.103 0.085 0.305 0.0726 0.0765 0.206
S 0.100 0.101 0.332 0.0602 0.0845 0.229
40 N 0.072 0.051 0.268 0.0564 0.0596 0.178
S 0.083 0.063 0.281 0.0417 0.0594 0.192
60 N 0.046 0.035 0.226 0.0428 0.0446 0.134
S 0.059 0.055 0.250 0.0301 0.0436 0.143
MP-1203 0 N 0.136 0.102 0.495 0.1146 0.1083 0.332
S 0.145 0.105 0.518 0.1048 0.1433 0.343
20 N 0.093 0.072 0.352 0.0861 0.0987 0.218
S 0.105 0.077 0.416 0.0766 0.1342 0.247
40 N 0.066 0.058 0.292 0.0659 0.0713 0.192
S 0.059 0.060 0.332 0.0586 0.0946 0.176
60 N 0.047 0.047 0.255 0.0538 0.0608 0.168
S 0.053 0.046 0.281 0.0393 0.0760 0.155
 M.A. Ahanger, R.M. Agarwal

Table 4 Height, flag leaf area, RWC, spike length, number of spikelets, 100 grain weight, and biomass of wheat (Triticum aestivum L cultivars LOK-1
and MP-1203) grown under normal (N) and restricted (S) irrigation and different potassium doses at flowering stage

Cultivar K levels Irrigation Height (cm) Leaf area RWC (%) Spike length Number of 100 grain Biomass
(kg ha−1) regime (cm2) (cm) spikelets per weight (g/pot)
Preflowering Flowering spike

LOK-1 0 N 43.1±0.2d 61.0±2.2g 5.73±0.02f 77.50±1.5d 5.30±0.06h 8.66±0.3h 4.20±0.24e 19.0±0.2h

S 35.6±0.3g 51.0±2.1h 4.80±0.03j 71.00±1.3g 4.10±0.13j 5.66±0.5k 4.03±0.08f 13.3±1.7j
20 N 46.9±1.9b 71.4±0.9d 6.18±0.02h 77.47±1.1d 6.70±0.46ef 10.33±0.4g 5.07±0.36c 29.3±0.4f
S 38.2±0.4f 63.8±0.8g 5.70±0.06fg 74.49±3.8e 5.16±0.08i 6.66±0.4j 4.42±0.40d 15.0±1.3i
40 N 50.6±1.0a 78.5±0.5c 6.73±0.04cd 81.76±3.7b 8.20±0.06d 11.33±0.4f 5.41±0.45b 34.3±1.7e
S 43.6±0.2d 66.7±1.7f 5.80±0.06f 73.01±0.3ef 5.66±0.28g 7.33±0.4i 4.19±0.10ef 18.0±0.2h
60 N 50.7±0.3a 80.4±1.3b 6.66±0.04d 86.59±3.5a 8.50±0.40c 11.66±0.4f 5.73±0.52a 37.0±0.8d
S 45.0±0.7c 67.0±1.1f 5.76±0.02f 76.73±3.6de 5.23±0.04h 7.66±0.3i 4.49±0.34d 23.3±3.7g
MP-1203 0 N 40.8±1.3e 76.8±0.8c 6.65±0.06d 73.11±3.0e 7.9±0.06e 14.00±0.5c 3.71±0.29h 39.0±1.3c
S 36.0±1.1g 62.1±0.6g 5.43±0.04i 70.84±5.4h 5.83±0.42g 8.33±0.4h 3.12±0.06k 23.3±3.1g
20 N 44.0±1.3c 81.2±0.5b 6.80±0.03c 74.20±0.9e 8.73±0.44c 17.00±0.6b 3.90±0.12g 39.3±7.5c
S 37.7±0.2f 67.5±0.8f 6.08±0.02e 70.18±1.7h 6.70±0.20ef 10.66±0.3g 3.34±0.08i 29.6±3.5f
40 N 47.3±0.2b 88.8±2.2a 7.29±0.02b 78.69±3.1c 9.33±0.08a 18.33±0.4a 3.96±0.17g 47.6±1.1b
S 37.3±0.5f 72.4±0.8d 6.13±0.04h 71.61±2.7g 7.26±0.17de 13.66±0.3d 3.37±0.35i 28.0±3.3f
60 N 49.3±0.6a 86.4±0.8a 7.43±0.01a 81.14±0.7b 9.23±0.11ab 18.66±0.8a 4.14±0.06ef 51.3±3.5a
S 41.5±0.9e 69.9±0.5e 6.63±0.01d 74.53±2.1e 6.40±0.40f 12.66±0.4e 3.33±0.18ij 33.0±2.6e

Data presented is mean of four replicates (±SE). Data followed by same letter are not significantly different at p < 0.05

glutathione pathway. Plants exposed to stressful environmental
conditions are reported to maintain higher activity of antioxi-
dant enzymes for quick removal of toxic ROS therefore,
protecting cell from the oxidative damage. SOD catalyzes the
conversion of O2− into H2O2 and O2whereas, H2O2 is further
acted upon by either CAT or APX through ascorbate-
peroxidase pathway. CAT eliminates H2O2 from the cell. Like
CAT, APX also scavenges H2O2 through its involvement in an
intriguing ROS scavenging pathway, ascorbate-peroxidase
cycle which also uses GR, MDHAR, DHAR, and non-
enzymatic components, AsA and glutathione (Mittler 2002;
Ahmad et al. 2010). Overproduction of ROS cause oxidation
of polyunsaturated fatty acids resulting in loss of membrane
function; however, seedlings/plants raised with potassium
supplementation exhibited lower lipid peroxidation suggesting
improved membrane stability. K reduces production of ROS by
down-regulating the activity of ROS, generating NADH oxi-
dases (Cakmak 2005). GSH helps in maintaining the levels of
ascorbate by acting as reductant in dehydroascorbate reductase
(DHAR)-mediated reduction of dehydroascorbate (DHA) to
ascorbate, and continuous supply of GSH to this reaction is
maintained by the NADPH-dependent reduction of GSSG by
GR (Gill and Tuteja 2010). K supplementation resulted in the
optimal activity of APX and GR and enhanced synthesis of
AsA and GSH. Both GSH and AsA have important role in
several important developmental processes including stress tol-
erance. By participating in ROS removal, AsA protects photo-
synthetic rate and electron transport chain (Miyaji et al. 2015).
Increased GR activity ensures optimal NADH availability for
accepting electrons of photosynthetic ETC, minimizing the
chances of O2− generation. Enhancement in the activity of an-
tioxidant enzymes due to potassium supplementation as noticed
in the present study may contribute to protection of the photo-
synthetic apparatus by lowering production of toxic ROS. Not
many reports are available focusing on the role of K in the
modulation of antioxidant defense system. Our results, i.e.,
increased AsA in water/osmotic-stressed seedlings/plants are
corroborative with the results of Manivannan et al. (2007),
Alam et al. (2014), and Shafiq et al. (2015). Mineral nutrition-
induced growth improvement under stressful conditions is
mainly achieved through the maintenance of GSH and AsA
levels (Fatma et al. 2014; Iqbal et al. 2015). Khare et al.
(2015) have also demonstrated increased GSH and AsA
content in salt-stressed rice, resulting in over-expression of dif-
ferent isozymes of APX and GR. Positive effects of K as no-
ticed in the present study on the antioxidant components could
contribute to stress tolerance through maintenance of the redox
balance and protection of membranes and proteins mitigating
the stress-triggered anomalies in important cellular pathways.
T. aestivum cultivars subjected to water stress showed
increase in accumulation of phenols and phenol derivatives
including tannins. Under normal irrigation, accumulation of
phenols increased with increasing K application, and water
stress imposition further increased their synthesis. MP-1203
maintained lower contents of metabolites as compared with
LOK-1, indicating cultivar differences. Accumulation of
Potassium up-regulates antioxidant metabolism
phenols in water/osmotic-stressed seedlings/plants probably
imparted greater DPPH radical scavenging potential.
Enhanced accumulation of metabolites in K-treated and
osmotic-stressed seedlings was also accompanied with im-
proved activity of PAL. K has been reported to increase the
synthesis of phenolics including tannins and phytic acid;
however, reports on the regulation of their synthesis under
stress conditions as affected by K availability are a few
(Tomar and Agarwal 2013). Secondary plant metabolites
(phenylpropanoids) are believed to have an active role in
metabolic processes of plants and are important for the
plant-environment interactions. Secondary metabolites in-
cluding phenols and flavonoids possess beneficial regulatory
roles in protecting plants from several biotic and abiotic
threats by their active involvement in free radical scavenging
and strengthening the cell wall structures from the stress-
induced alterations. Secondary metabolites protect plants from
bacterial, fungal, and viral infections (Kennedy and
Wightman 2011). K-induced increase in synthesis of phenols,
tannins, and phytic acid contributing to the maintenance of
growth has been indicated. Accumulation of phenols, tannins,
and flavonoids in response to stress has been reported by other
workers as well (Tomar and Agarwal 2013; Kaur and Zhawar
2015). Polyphenols act as pro-oxidants and can terminate the
free radical chain reactions (Kagan and Tyurina 2006). K-
supplemented plants in the present study exhibited reduced
lipid peroxidation, indicating role of potassium in preventing
stress-induced oxidative damage. Hypericum brasiliense
subjected to water stress exhibited greater accumulation of phe-
nols like rutin, kaempherol, quercetin, 1,5-dihydroxyxanthone,
and isouliginosin B (de Abreu and Mazzafera 2005). In
Hypoxis hemerocallidea, cadmium and aluminum stresses
reportedly induced accumulation of phenols and flavonoids ac-
companied by higher DPPH radical-scavenging activity (Okem
et al. 2015). Phenol derivatives share a crosstalk with important
signaling molecules like nitric oxide for optimizing the nitrogen
assimilatory pathways (Klein et al. 2015). Metabolite accumu-
lation can discharge antioxidant function when enzymatic
antioxidants and low molecular weight antioxidants get
depleted due to severe oxidative stress (Brunetti et al. 2015).
K application enhanced the synthesis of compatible
osmolytes in plants which increased with increasing doses.
Substantial increase in accumulation of osmolytes was noticed
in plants grown under restricted irrigation regimes and accu-
mulation of osmolytes increased as the plants advanced to
flowering stage; however, free amino acids registered a de-
cline with the advancement from preflowering to flowering
stages. Accumulation of compatible osmolytes during
stressful environmental conditions and with the addition of
K indicates protective role of K for plants through osmotic
adjustment. Accumulation of sugars resulting from the enzy-
matic degradation mediated by amylases may contribute to the
maintenance of tissue turgor potential under stressed condition
(Pandey et al. 2004). Accumulation of osmotic constituents
like free proline, sugars, and amino acids contribute to the
maintenance of cell volume through turgor maintenance,
protecting important cellular structures including proteins
(Buchanan et al. 2000). Accumulation of low molecular
weight compatible solutes in the tissues tend to bring the tissue
water potential much below the external soil solution so that
water absorption gets less affected (Chen and Jiang 2010).
Osmotic stress induced by PEG increased accumulation of
free sugars in Zea mays L (Velazquez-Marquez et al. 2015).
Proline-synthesizing pathway is up-regulated as compared
with catabolizing, which is down-regulated under stressed
conditions. Reddy et al. (2015) found significant decrease in
lipid peroxidation, increase in photosynthetic attributes, and
subsequent alleviation of salt stress-induced changes due to
increased proline accumulation in Sorghum bicolor as a result
of over-expressing proline biosynthesis enzymes. Apart from
their obvious role in osmoregulation, sugars, amino acids, and
proline can exert protective functions by acting as antioxidants
(Keunen et al. 2013; Kishor and Sreenivasulu 2014).
Enhanced proline accumulation protects the free radical scav-
enging potential of antioxidant enzymes (Ozden et al. 2009).
Increased proline accumulation due to K supplementation
could probably help plants revive growth quickly when stress
is relieved. Accumulated proline reservoirs are depleted for
providing energy supply for fast growth recovery after stress
release and can also mediate regulated growth transition from
vegetative to reproductive phase up to the seed initiation
(Kishor and Sreenivasulu 2014). Proline protects proteins,
membranes, and DNA by inhibiting the formation of
conjugated dienes and malonaldehyde, the products of lipid
peroxidation (Matysik et al. 2002). Sugars maintain the car-
boxylase activity of Rubisco with concomitant reduction in its
oxygenase activity under salt stress (Sivakumar et al. 2002).
When entering into the pentose phosphate pathway, sugars
help in developing reducing power for GSH production hence,
contributing to H2O2 elimination. It has been suggested that
sugars, especially water-soluble oligo and polysaccharides,
e.g., fructans, might be effective candidates to capture ROS
during stresses and in co-ordination with phenolic compounds
can form an integrated redox system for quenching of ROS
thereby helping in oxidative stress mitigation (Bolouri-
Moghaddam et al. 2010). Increased contents of sugars and
phenols during stresses and K treatments found in the present
experiments might have contributed towards the production of
GSH, resulting in enhanced potential for counteracting the
stress-induced ROS damage.
Irrigated plants showed efficient K uptake. Plants raised
under restricted water irrigation showed a decline in K and
Na content (Tables S1 and S2). Roots were found to maintain
higher concentrations of sodium as compared with leaf and
stem indicating selective absorption and uptake. K is the dom-
inant cation in plant tissues and stress imposition induces K

efflux through roots leading to reduced tissue expansion and
production of photoassimilates thereby hampering tissue water
balance. It is the reduced tissue water content which is respon-
sible for the altered transpiration rates and the reduced stem
expansion and hydraulic conductance (Kanai et al. 2011).
Plants prefer K over Na, and fine balance between these two
is maintained by the efficient working of transporters like
NHX, SOS, and HKT which prevent the over-accumulation
of Na+ ion, hence maintaining K/Na ratio (Shabala et al.
2016). Anschutz et al. (2014) while linking post-translational
regulation of K+ channels with PCD suggest that K in addition
to being an essential nutrient is also involved in a variety of
plant responses to environmental stresses. Different abiotic and
biotic stresses reportedly (Shabala and Pottosin 2014) disturb
K+ homeostasis, reflecting on K channels and transport
proteins and consequently on absorption and usage. Shabala
et al. (2016) have analyzed regulation of membrane trans-
porters under stress conditions, highlighting its importance in
breeding strategies for improving abiotic stress tolerance. K
treatments reduced the oxidative stress by probably maintain-
ing the sufficient availability of K+ for quick displacement of
Na+, averting the toxic effects of salinity reflected in reduced
lipid peroxidation. Reduced Na uptake concomitant with
increased K uptake in water-stressed plants evident in our
results is corroborative with the other reports (Akram 2013;
Jatav et al. 2014; Vijayalakshmi et al. 2014).
Wheat exposed to water stress exhibited considerable re-
duction in yield and yield associated attributes like length of
spike, number of spikelets, leaf area, grain weight, and bio-
mass yield (Table 4). Cultivar differences were obvious with
LOK-1 showing relatively better stress withstanding potential
than MP-1203. A significant reduction in the yield of wheat
cultivars due to water stress has been reported by others as
well (Tiwari et al. 1998; Hameed et al. 2011; Jatav et al. 2014).
Flowering, a critical developmental phase is vulnerable to
environmental stresses and exposure to salt and water stress
hastened flowering. Alterations in the flowering time brought
about by stresses are an evolutionary adaptation exhibited by
stress-challenged plants to maximize the chances of reproduc-
tion (Kazan and Lyons 2016).
Conclusions
It can be concluded that K supplementation improved growth
of wheat cultivars by up-regulating antioxidant metabolism
for quick removal of reactive free radicals. Furthermore,
wheat plants accumulated highest amounts of K at higher
doses which resulted in improved growth performance under
normal as well as restricted irrigation regime. Enhanced
accumulation of K in such plants depicts the demand driven
requirement of K for the maintenance of osmolarity for better
extraction of water. Future studies could be beneficial in
M.A. Ahanger, R.M. Agarwal
providing better understanding about the underlying mecha-
nisms for K-induced stress tolerance.
Acknowledgments Financial assistance from Jiwaji University,
Gwalior (F/DEV/2013-14/33) to the first author in the form of a JU
Research Fellowship is gratefully acknowledged. Help received from
Prof. Nafees A. Khan, Department of Botany, Aligarh Muslim
University is thankfully acknowledged. Thanks are also due to Dr.
Mohd Asgher for sharing SPSS and Sigma plot software.
Compliance with ethical standards
Conflict of interest Authors have no conflict of interest
Authors’ contribution Present manuscript forms part of a PhD work
of MA Ahanger which was conceived and designed by Prof. RM
Agarwal. MA Ahanger performed experiments and wrote the first draft
of the manuscript, and RM Agarwal cross checked the results and
manuscript.
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