Journal of Photochemistry & Photobiology, B: Biology 160 (2016) 86–95

Contents lists available at ScienceDirect

Journal of Photochemistry & Photobiology, B: Biology

journal homepage: www.elsevier.com/locate/jphotobiol

Chlorophyll fluorescence induction kinetics and yield responses in rainfed

crops with variable potassium nutrition in K deficient semi-arid alfisols
Ch. Srinivasarao, Arun K. Shanker ⁎, Sumanta Kundu, Sharanbhoopal Reddy
Central Research Institute for Dryland Agriculture, Santoshnagar, Saidabad P.O, Hyderabad 500 059, India

a r t i c l e i n f o a b s t r a c t

Article history: Optimum potassium (K) nutrition in semi-arid regions may help crop plants to overcome constraints in their
Received 3 October 2015 growth and development such as moisture stress, leading to higher productivity of rainfed crops, thus judicious
Accepted 31 March 2016 K management is essential. A study was conducted to evaluate the importance of K nutrition on physiological
Available online 13 April 2016
processes like photosynthesis through chlorophyll a fluorescence and chlorophyll fluorescence induction kinetics
(OJIP) of rainfed crops viz., maize (Zea mays L.), pearl millet (Pennisetum glaucum), groundnut (Arachis hypogaea),
Keywords:
Plant nutrition
sunflower (Helianthus annuus), castor (Ricinus communis L.) and cotton (Gossypium hirsutum) under water stress
Chlorophyll fluorescence conditions by studying their growth attributes, water relations, yield, K uptake and use efficiency under varied K
OJIP levels. Highest chlorophyll content was observed under K60 in maize and pearl millet. Narrow and wide Chl a:b
Water relations ratio was observed in castor and groundnut respectively. The fluorescence yield decreased in the crops as K dos-
Nutrient Use efficiency age increased, evidenced by increasing of all points (O, J, I and P) of the OJIP curves. The fluorescence transient
curve for K60 was lower than K0 and K40 for all the crops. Potassium levels altered the fluorescence induction
and impaired photosynthetic systems in all the crops studied. There was no distinct trend observed in leaf
water potential of crops under study. Uptake of K was high in sunflower with increased rate of K application.
Quantitatively, K uptake by castor crop was lesser compared to all other crops. Our results indicate that the
yield reduction under low K was due to the low capacity of the crops to translocate K from non-
photosynthetic organs such as stems and petioles to upper leaves and harvested organs and this in turn influ-
enced the capacity of the crops to produce a high economic yield per unit of K taken up thus reducing utilization
efficiency of K.
© 2016 Elsevier B.V. All rights reserved.

1. Introduction
Potassium (K) is the third most essential macronutrient needed by
plants to complete their life cycle successfully, after nitrogen and phos-
phorus. Essentiality of K in plant life cycle is evidenced by its role in ac-
tivating about 80 enzymes which are involved in various physiological
processes such as energy metabolism, starch synthesis, nitrate reduc-
tion, photosynthesis and sugar degradation. As a component of plant cy-
toplasmic solution, K reduces loss of water from leaf surfaces by
regulating stomatal closure and increases water uptake efficiency of
roots from soil mainly due to the role of K in regulating cellular osmotic
potentials [1]. Thus, K has a positive impact on improved drought toler-
ance of plants growing under water shortage conditions [2] in rainfed
regions. Agriculture in rainfed regions suffers from constraints like
poor soil fertility particularly K, low soil moisture availability/retention,
and reduced nutrient use efficiency which directly affect the perfor-
mance of crop plants resulting in reduced productivity. The general
⁎ Corresponding author at: ICAR-Central Research Institute for Dryland Agriculture,
Santoshnagar, Hyderabad 500 059, India.
E-mail address: arunshank@gmail.com (A.K. Shanker).
practice in these regions is to only use NP rich fertilizers excluding K fer-
tilization in major rainfed crops such as maize (Zea mays L.), pearl millet
(Pennisetum glaucum), cotton (Gossypium hirsutum), groundnut
(Arachis hypogaea), sunflower (Helianthus annuus) and castor (Ricinus
communis L.) which are also known to be K exhaustive crops. Thus, con-
tinuous cropping in absence of optimum K nutrition resulted in deple-
tion of soil K reserve such as non-exchangeable K fraction [3,4]. Many
studies across Indian soils and elsewhere in the world are becoming K
deficit in different soil types [5]; this resulted in K deficiency in soils
and sub-optimal K nutrition is an important productivity constraint par-
ticularly in rainfed dryland regions, highly vulnerable weather condi-
tions mainly elevated temperature and untimely rainfall.
Semi-arid alfisol regions in general experience droughts in quick
succession mainly due to monsoon failures which have a serious impact
on crop yields. Deficit rainfall due to delayed onset or breaks in south
west monsoon is of major concern in rainfed agriculture that reduced
productivity of major crops [6]. Effective K management is one of the
key approaches that will help the plant to overcome these situations
resulting in normal yields rather than critical loss of crop.
During the photosynthetic process, light is absorbed by the antenna
molecules within the photosynthetic membrane and the absorbed

http://dx.doi.org/10.1016/j.jphotobiol.2016.03.052
1011-1344/© 2016 Elsevier B.V. All rights reserved.
 C. Srinivasarao et al. / Journal of Photochemistry & Photobiology, B: Biology 160 (2016) 86–95
energy is transferred as excitation energy. There are two ways that de-
termine the fate of this energy one is that the energy is trapped at a re-
action center and used in photochemistry, and the other is that it is
dissipated mainly in the form of heat and fluorescence. This fluores-
cence emission offers interesting opportunities in studying the plants'
response to environmental conditions. The properties of this emitted
fluorescence are basically determined by the pigments that absorb
them, the transfer of energy by the excitation process, and the nature
and orientation of the fluorescing pigments. Chlorophyll (Chl) fluores-
cence is a non-invasive method to monitor the alterations of photosyn-
thetic processes [7] by measuring the emitted radiation from the crop. It
is a useful indicator as it gives important information about the state of
health of a photosynthetic apparatus and it is a valuable tool for plant
studies from leaf to ecosystem levels [8,9]. Chl fluorescence can be char-
acterized by different phases: Chlorophyll Fluorescence Induction Ki-
netics (OJIP) is fluorescence transient corresponding to the redox
states of photosystem PSII and PSI, and to the efficiencies of electron
transfer through the intersystem chain to the end electron acceptors
at the PSI acceptor side [10,11]. It is known that K impacts photosynthe-
sis of the crop canopy via solar radiation interception, and we thus hy-
pothesize that increasing doses of K will have an impact on the
growth and development and yield of crops, there is a dearth of litera-
ture on the relationship between of Chl fluorescence and K levels in
rainfed crops. Hence, current study was planned to evaluate the impor-
tance of K nutrition on crops and with the objective to study the effects
of applied K on Chl a fluorescence in addition to exploring the possibil-
ities to surmount the effects of early, mid-season and terminal droughts
that become common occurrences in rainfed regions affecting perfor-
mance of rainfed crops like maize, pearl millet, cotton, groundnut, sun-
flower and castor.
2. Material and Methods
2.1. Details of Experiment Site
A series of field experiments were carried out on different rainfed
crops at Gunegal Research Farm of ICAR-Central Research Institute for
Dryland Agriculture (CRIDA), Hyderabad (India) during summer season
of 2012 and 2013. Location coordinates are 17° 05′ 01.19″ N 78° 40′
06.32″ E and the average annual rainfall is 690.0 mm, mean annual tem-
perature was 25.7 °C. Mean monthly rainfall and temperature during
experiment period are presented in Fig. 1. Soils are sandy loam in tex-
ture, slightly acidic in reaction (pH 5.1), EC was in normal range
(0.08–0.1 dS m−1), low in organic carbon (3.4 g kg−1), available N
(141–153 kg ha−1), medium in available P (16–19 kg ha−1) and avail-
able K (154–188 kg ha−1). Among micronutrients Zn was deficient
(0.48 mg kg−1).
2.2. Crops and Treatment Details
Popular varieties of major rainfed crops viz., maize (DHM-117),
pearl millet (Shanti), cotton (MRC-7347 BG-1, MRC-7351 BG-1 and
Fig. 1. Mean monthly rainfall and temperature during experiment period.
87
NCS-207 Mallika Bt-2) groundnut (JL-24), sunflower (KBSH-1) and cas-
tor (DCS-9) of the region were selected for the experiment. There were
three treatments comprising of 0, 40 and 60 kg ha−1 K respectively de-
noted as K0, K40 and K60. Three K levels respectively depict nil, medium
and maximum K recommendations for a crop. The levels of K were final-
ized with reference to package of practices suggested for different crops
of the region by State Agricultural University. Details of the spacing,
sowing and harvesting of each crop is given in Table 1 and monthly
mean rainfall and temperature during experiment period is given in
Fig. 1. Except differential dose of K application all other nutrients were
supplied uniformly as per recommendation considering soil test values.
2.3. Carotenoids and Chlorophyll Content of Leaf Tissue
Carotenoids and chlorophyll content in plant leaf were measured
(mgg−1 of fresh weight) following the procedure outlined by Wellburn
[12] and equations developed by Arnon [13] were used to calculate Chla,
Chlb and Total Chl are as follows,


Chla mg g‐1 ¼ 0:0127 A663–0:00269 A645


Chlb mg g‐1 ¼ 0:0029 A663–0:00468 A645


Total Chl mg g‐1 ¼ 0:0202 A663 þ 0:00802 A645
2.4. Chlorophyll Fluorescence Measurements
The OJIP fluorescence transient was measured using a portable fluo-
rometer (FluorPen FP 100; Photon Systems Instruments; Drasov, Czech
Republic). Fully developed youngest leaves were selected for the mea-
surements. The leaves were dark-adapted for 30 min before starting
the measurements using leaf clips provided by the manufacturer. Mea-
surements were done three times on the adaxial leaf surface. For each
measurement, two places per leaf were selected in one plant. Data
were analyzed from 10 measurements (five plants with two places
per leaf, and one leaf per plant) for each treatment and control. The
method followed was according to Thwe and Kasemsap [8]. The OJIP
fluorescence parameters were calculated based on the formulas
shown in Table 2. Minimum fluorescence (F0) was measured at 50 μs
when all PSII reaction centers are open and it is defined as the O step,
followed by the J step (at 2 ms), the I step (at 60 ms) and at maximum
fluorescence (FM) when all PSII reaction centers are closed, known as
the P step [10]. The OJIP test represents a translation of the original
data to biophysical parameters that quantify the energy flow through
PS II [14]. From each OJIP fluorescence induction, specific energy fluxes
per reaction center were analyzed and compared.
2.5. Leaf Water Potential
Measurement of leaf water potential is a better indicator for plant
physiological response under water shortage conditions. For all crops
leaf water potential was measured timely within field using Psypro™
Wescor water potential system.
2.6. Soil Sampling and Analysis
Soil samples from three depths i.e., 0–0.2, 0.2–0.4 and 0.4–0.6 m
were collected with the help of tube augar before and after every crop.
Soil samples were air dried, gently ground and passed through a 2-
mm sieve. Further these processed samples were analyzed for basic
physico-chemical properties viz., pH and EC, respectively indicating
acidity–alkalinity and salt concentration of the soil. A known quantity
of soil (20 g) was vigorously shaken after adding distilled water (1:2.5
soil:water ratio) [15]. After a time period pH of soil was determined
88 C. Srinivasarao et al. / Journal of Photochemistry & Photobiology, B: Biology 160 (2016) 86–95

Table 1
Spacing, sowing and harvesting details of each crop.

Crop Spacing (cm) Sowing date Harvesting date

2012 2013 2012 2013

Maize 60 × 25 10th June 28th June 17th October 8th November

Pearl millet 45 × 15 11th June 18th July 28th September 14th October
Cotton 60 × 45 12th June 28th June 22nd October 4th November
2nd November 13th November
13th November 2nd December
(1st, 2nd and 3rd pickings, respectively) (1st, 2nd and 3rd pickings, respectively)
Groundnut 30 × 10 11th June 18th July 14th October 6th November
Sunflower 60 × 30 10th June 18th July 9th October 15th November
Castor 90 × 60 14th June 28th June 2nd November 13th November

using VSI-01 ATC Deluxe pH meter. Whereas, EC was determined very
next day using VSI-01 ATC Deluxe conductivity meter once after clear
supernatant was observed. Soil organic carbon (SOC) content (g kg−1)
was determined by wet digestion with acid-dichromate [16]. Major nu-
trient availability viz., N was determined by alkaline permanganate
method using Kjeldahl distillation unit [17], P was determined by
extracting soils with Olsen's extractant (0.5 M NaHCO3, pH 8.5) and
subsequently knowing the concentration at 660 nm using spectropho-
tometer (T-60 UV Visible Spectrophotometer, PG Instruments) [18];
and exchangeable K was extracted using neutral normal ammonium ac-
etate (pH 7.0) extractant and concentration was determined using
Systronics Flame Photometer 128 [19]. Whereas, micronutrients con-
centration (Fe, Zn, Cu and Mn) was determined using Atomic Absorp-
tion Spectrophotometer (AAS- Perkin Elmer AAS 800) instrument
after extracting with DTPA (0.005 M DTPA. 0.01 M CaCl2, 0.1 M TEA;
pH 7.3) extractant (1:2 soil:extractant) [20]. Available B was deter-
mined through hot water extraction [21].
and stover, lint samples were also analyzed to know their nutrient com-
position following digestion with diacid mixture [22].
2.8. Determination of Potassium in Plant Samples
Uptake of potassium by the crops under study was measured by rep-
resentative plant samples collection from respective treatment plots.
Plants collected were washed gently with water to remove dust parti-
cles on surface of leaves and other parts. Once after washing samples
were oven dried (at 60 ± 5 °C) at lab until no difference in moisture
content was observed. Then after samples were ground into fine pow-
der using grinder for further analysis. To determine nutrient content
in plant samples, were digested in diacid mixture (1:10 sample: diacid
ratio) till colorless supernatant was obtained. Supernatant obtained
was made up into a known volume. Then aliquot made was used for K
analysis using Flame Photometer (Systronics Flame Photometer 128).
The K uptake by various crops was calculated using the following
formula,

2.7. Plant Sample Analysis K uptake ðkg=haÞ ¼ %K content in plant  yield obtained=100

Representative plant samples were collected from every treatment

and processed for analysis. Grain and stover samples of each crop 2.9. K Use Efficiency
were analyzed individually, with regard to cotton along with grain
Efficiency of applied K for different crops was calculated through ef-
Table 2 ficiency parameters such as agronomic efficiency (AE, kg crop yield in-
Parameters of the steps of fluorescence induction. crease per kg nutrient applied); apparent recovery efficiency (RE, kg
Parameter Formula explanation. nutrient taken up per kg nutrient applied); and physiological efficiency
abbreviation. (PE, kg yield increase per kg nutrient taken up) [23].
F0F50μs fluorescence intensity at 50μs
Fj Fluorescence intensity at J-step (at 2 ms) 2.10. Statistical Analysis
Fi Fluorescence intensity at i-step (at 60 ms)
FM Maximal fluorescence intensity Statistical analysis of data was performed using the Windows based
FV FV = FM − F0 (maximal variable fluorescence)
Vj Vj = (Fj − F0)/(FM − F0)
SPSS program (Version 11., SPSS, Chicago, IL, USA 2001). The SPSS pro-
Vi Vi = (Fi − F0)/(FM − F0) cedure was used to analyze variance and to determine statistical signif-
M0 or (dV/dt)0 M0 = TR0/RC − ET0/RC = 4 (F300 − F0)/(FM − F0) icance of treatment effects. Simple correlation coefficients and
Area Area between fluorescence curve and FM (background regression equations were developed to obtain relationship between K
subtracted)
content and leaf water potential, fluor pen readings.
Fix area Area below the fluorescence curve between F40μs and F1s
(background subtracted)
φ_P0 φ _P0 = 1 − (F0/FM) (or FV/FM) 3. Results
ψ_0 ψ_0 = 1 − Vj
φ_E0 φ _E0 = (1 − (F0/FM)) ∗ ψ_0 3.1. Pigments
φ_D0 φ_D0 = 1 − φ_P0 − (F0/FM)
φ_Pav φ_Pav = φ _P0 (SM/tFM) tFM = time to reach FM (in ms)
rRC2 rRC2 = ChlRC/Chltotal Effect of K application on the chlorophyll pigments viz., chlorophyll
ψ_ET2o = 1 − Fj/FM a, chlorophyll b and their ratio, total chlorophyll and carotenoids are
ABS/RC ABS/RC = M0 ∗ (1/Vj) ∗ (1/φ_P0) given in Tables 4 and 5. In general, chlorophyll a was significantly higher
TR0/RC TR0/RC = M0 ∗ (1/Vj)
in K60 treatments as compared to K40 in maize, pearl millet, sunflower,
ET0/RC ET0/RC = M0 ∗ (1/Vj) ∗ ψ _0
DI0/RC DI0/RC = (ABS/RC) − (TR0/RC) cotton and castor excluding groundnut. Highest values (6.37) of chloro-
PIAbs PIAbs = [rRC2/(1 − rRC2)] ∗ [φ_P0/(1 − φ_P0)] ∗ phyll a were seen in maize at 60 kg ha−1 treatment. Chlorophyll a was
[ψ_ET2o/(1 − ψ_ET2o)] generally higher in the K60 as compared to the other treatments in all
PItotal PItotal = PIAbs ∗ {(1 − Vi)/(1 − Vj)}/[1 − {(1 − Vi)/(1 − Vj)}] the crops except sunflower and castor wherein there was a slight de-
Source: Strasser et al. (2000); Stirbet and Govindjee (2011). crease in the values at higher levels of K application. Chlorophyll b
 C. Srinivasarao et al. / Journal of Photochemistry & Photobiology, B: Biology 160 (2016) 86–95
showed a similar trend as seen in chlorophyll a, K60 treatment showed
significantly higher values when compared with the K40 treatment in
maize, pearl millet, groundnut and sunflower. In the case of cotton
and castor chlorophyll b content was on par between K60 and K40, sig-
nificant increase in comparison to control. In maize and pearl millet
total chlorophyll was significantly different when compared among all
the treatments. In sunflower and castor 60 kg ha −1 was significantly
higher than the other two treatments that were on par. There was no
significant difference seen in groundnut crop in all the K levels. Chloro-
phyll a:b ratio trend was observed similar to that of total chlorophyll in
maize, pearl millet, cotton and sunflower. Values of control and K40
treatments were on par whereas K60 values were differed significantly.
But, in groundnut all the levels of K exhibited significant different values
of chlorophyll a:b ratio having decreasing trend with increasing K doses
i.e., 10.84 (K0), 7.79 (K40) and 3.34 (K60).
Effect of different K doses on carotenoids level (mg g−1 of fresh
weight) in leaves of major rainfed crops are presented in Table 5. Appli-
cation of K at 40 kg ha−1 to maize showed comparatively higher carot-
enoids than K0 and K60 but, Pearl millet showed no variation in leaf
carotenoids levels having on par values under different K additions. Oil-
seed crops were observed with slightly reduced carotenoids levels with
every increment of K. Carotenoid levels in groundnut and castor leaves
were 3.00 and 4.98; 2.06 and 4.91; and 1.90 and 4.35 under K0, K40 and
K60 treatments. Cotton crop also showed higher carotenoids levels
under no K additions (4.79) followed by K60 (4.19) and K40 (3.88).
3.2. Leaf Water Status
Leaf water potential (MPa) of major rainfed crops grown influenced
by different K levels are given in Table 6. Leaf water potential was signif-
icantly different in cereals and millets, oilseeds and fiber crop. Relatively
oilseeds showed high (negative) water potential values ranging from
− 2.80 (castor, K0) to − 5.41 (sunflower, K60). Significant difference
in leaf water potential values of cotton crop under varied K levels was
observed viz., −0.05 (K0), −0.17 (K40) and − 0.76 (K60) while con-
trasting values were there for maize i.e., − 1.86 (K0), − 0.03 (K40)
and − 0.02 (K60). On par leaf water potential values were in K0
Fig. 2. OJIP chlorophyll a fluorescence transients of castor (a) and cotton (b) at different levels of potassium.
89
(−5.04) and K40 (−4.69) treatments under groundnut crop and K40
(−4.86) and K60 (−5.09) treatments of castor crop.
3.3. OJIP Fluorescence Transient Curves
OJIP chlorophyll a fluorescence transients of six crops viz., maize,
pearl millet, groundnut, sunflower, castor and cotton grown under dif-
ferent K are depicted in Figs. 2–4. The crops under both control (K0)
and other K treatments showed a typical chlorophyll a polyphasic fluo-
rescence rise OJIP. However, the fluorescence yield decreased in the
crops as K dosage increased, this is evidenced by increasing of all points
(O, J, I and P) of the OJIP curves (Figs. 2–4). The fluorescence transient
curve for K60 was lower than K0 and K40 for all the crops. In castor
(Fig. 2a) K60 level clearly showed a marked difference in the transient
curve with the two phases I and P flattening as against the curves ob-
served in K0 and K40. In the case of cotton (Fig. 2b) the curves for K0
and K40 are the same for the I and P phases whereas the there is a re-
duction in chlorophyll fluorescence arbitrary units for the higher K
level i.e., K60. OJIP chlorophyll a fluorescence transients of groundnut
is given in Fig. 3a and it was observed that K60 level exhibited higher
values at chlorophyll fluorescence arbitrary units during the J step and
started to decrease when it approached 10 ms going into the I phase
of the curve, I and P phases were plateaus, this pattern was not observed
in any of the crops studied. In sunflower (Fig. 3b) all the three levels of K
viz., 0, 40 and 60 kg K ha−1 exhibited a clear demarcation with the chlo-
rophyll a fluorescence transient curve, for K60 being the lowest. In Pearl
millet (Fig. 4a) the I step of the curve between 10 and 100 ms was
cluttered together for all the treatments, the curves separated towards
the end of the P step with K60 being the lowest. In maize (Fig. 4b) the
chlorophyll a fluorescence transient curves were separated clearly
with the K60 the lowest below the K0 and K40.
3.4. Maximum Quantum Yield of Primary Photochemistry (Fv/Fm) and its
Efficiency (ψ0)
Maximum quantum yield of primary photochemistry (Fv/Fm) and its
efficiency (ψ0) of the six crops studied under different K levels is depicted
90 C. Srinivasarao et al. / Journal of Photochemistry & Photobiology, B: Biology 160 (2016) 86–95

Fig. 3. OJIP chlorophyll a fluorescence transients of groundnut (a) and sunflower (b) at different levels of potassium.

in Fig. 5a and b, respectively. In all the crops except castor and cotton the
K60 showed significantly higher Fv/Fm ratio as compared to the K0, K40
levels. In the case of castor and cotton the K40, K60 treatments were on
par with each other whereas both were significantly different from K0.
The maximum quantum yield of primary photochemistry was differed
significantly from each other with varied doses of K application in case
of groundnut, maize, Pearl millet and sunflower. In castor and cotton,
highest values of Fv/Fm ratio were observed in K60 as compared to
other crops although they were not varied significantly from K40. Compa-
rable trends were observed for efficiency of primary photochemistry
(ψ0). The marked difference in the pattern was observed in cotton that re-
ceived K60 and varied significantly from K40 which was contrasting to Fv/
Fm in cotton. Similar to Fv/Fm ratio, the highest values for the efficiency of
primary photochemistry was seen in cotton crop.

Fig. 4. OJIP chlorophyll a fluorescence transients of pearl millet (a) and maize (b) at different levels of potassium.
 C. Srinivasarao et al. / Journal of Photochemistry & Photobiology, B: Biology 160 (2016) 86–95
Fig. 5. Maximum quantum yield of primary photochemistry (FV/FM) (a) and efficiency (Psi0, ψ0) (b) of crops at different levels of potassium.
3.5. Photochemical quenching (qP) and nonphotochemical quenching
(npq)
Photochemical quenching (qP) and nonphotochemical quenching
(npq) are shown in Fig. 6a and b respectively. Cotton and castor exhib-
ited highest levels of photochemical quenching under all three K doses,
with levels K40 and K60 exhibiting significantly increased values in
comparison with control treatment (K0). Alternatively groundnut,
maize, Pearl millet and sunflower exhibited significant increase in
values under each level of K applied. Divergent results observed for
nonphotochemical quenching (npq). Significant differences exhibiting
decreasing trend with increasing dosage of applied K. In the case of cot-
ton and castor K40 and K60 did not exhibit significant differences,
whereas this was not the case in other crops.
3.6. Crop Yield
Effect of K application at different rates on yield (grain, straw and
total) of major rainfed crops are given in Table 7. Highest grain yield
(t ha−1) was obtained in maize under the K60 (4.46) followed by K40
(4.27) which was also significantly higher than control yields (3.13). A
similar trend was seen in straw yield of maize; whereas harvest index
(HI) has not differed significantly among treatments. Pearl millet con-
siderably yielded better with extra additions of K i.e., 0.55 (K40) and
0.68 (K60) than control (0.29) that received no K. Straw yield and har-
vest index recorded significant values under K60 and K40 over control.
Unlike in maize, HI of Pearl millet showed significant difference among
treatments such as 6.40 (K0), 10.71 (K40) and 12.56 (K60). A slight
Fig. 6. Photochemical quenching (qP) (a) and Non photochemical quenching (b) of crops at different levels of potassium.
91
effect of K application on grain and stover yield of oilseeds was observed
as compared to control. Groundnut and castor crops resulted in signifi-
cant grain yields under K40 (0.54 and 1.11) and K60 (0.49 and 1.12)
than control (0.40 and 0.73), respectively. Harvest index was signifi-
cantly different from each other i.e., 22.94 (K0), 27.35 (K40) and 16.27
(K60). There was no grain yield difference in sunflower crop between
control and K40 treatments having 1.61 kg ha−1, but was significantly
higher in K60 (1.81 kg ha−1), but relatively no significant difference in
straw yield (≈ 5 kg ha−1) between the levels of K applied. Grain and
straw yield in castor was significantly higher in K60 than K40 and K0.
This was not the case in HI wherein the values were on par with each
other in all the treatments. Grain and lint yield followed the same pat-
tern in cotton i.e., respectively 0.68 and 2.16 with K40 and 0.71 and
2.60 with K40 recorded significantly higher yield than control (0.53
and 2.05). Straw yield in cotton exhibited no significant difference (av-
erage yield of 3.5 kg ha−1) among the treatments. There was a good HI
for cotton with K60 (38.76), significantly higher values when compared
to K40 (33.42) and control (34.35).
3.7. Potassium Uptake
Potassium uptake ability of various rainfed crops at different K levels
were presented in Table 3. Sunflower crop shown higher K uptake with
a subsequent increase in K dose having 48.89, 59.91 and 100.15 kg ha−1
with no, 40 and 60 kg ha−1 of K application. There is good amount of K
uptake was observed in sunflower with every 20 kg increment. Castor
crop comparatively was with low K uptake efficiency ranging from
8.75 (K0) to 16.87 kg ha−1 (K60). Among cereals and millets, Pearl
92 C. Srinivasarao et al. / Journal of Photochemistry & Photobiology, B: Biology 160 (2016) 86–95

Table 3 Table 5
Potassium uptake (kg ha−1) by major rainfed crops at different potassium levels. Carotenoids level (mg g−1 of fresh weight) in leaves at different potassium levels in major
rainfed crops.
Crop K uptake (kg ha−1)
Crop Potassium level
K0 K40 K60
K0 K40 K60
Cereals and millets
C B A
Maize 13.45 16.48 32.32 Cereals and millets
Pearl millet 40.97C 50.74B 59.11A Maize 3.52C 6.03A 5.12B
Pearl millet 4.89A 4.75A 5.01A
Oilseeds
Groundnut 13.45C 16.48B 32.32A Oilseeds
Sunflower 48.89C 59.91B 100.15A Groundnut 3.00A 2.06B 1.90B
Castor 8.75C 15.28 B 16.87A Sunflower 3.44A 3.47A 2.89B
Castor 4.98A 4.91A 4.35B
Fiber crops
Cotton 36.85C 45.06B 64.77A Fiber crops
Cotton 4.79A 3.88B 4.19A
Capital letters represent level of significance at P = 0.05 among various treatments and
different letters indicate significant difference. Capital letters represent level of significance at P = 0.05 among various treatments and
different letters indicate significant difference.

millet showed high K uptake followed by maize; among oilseeds uptake
trend was higher in sunflower followed by groundnut and castor. After
sunflower, cotton crop extracted more of applied K from soil i.e., 36.85
under K0; 45.06 under K40 and 64.77 kg ha−1 under K60 levels. In
maize and groundnut crops significant increase i.e., almost double quan-
tity of K uptake was observed with 60 kg ha−1 of K application (32.32)
as compared to 40 kg ha−1 (16.48). In general, all crops have shown an
increased trend in K uptake with increasing K dose. Among crops K up-
take trend was in the order of sunflower N cotton ≈ Pearl
milletN maize ≈ groundnut N castor.
3.8. Apparent Recovery Efficiency (RE), Physiological Efficiency (PE) and
Agronomic Efficiency (AE)
Efficiency of applied K and performance of crop under varied K levels
expressed using apparent recovery efficiency (RE), physiological effi-
ciency (PE) and agronomic efficiency (AE) are depicted in Fig. 7. Castor
crop exhibited the highest PE of above 45 among all the crops studied,
with K40 and K60 additions followed by groundnut at K40. In sunflower
crop the PE was lowest having less than 5 under both K40 and K60.
Groundnut crop at K40 showed high PE as compared to other crops. Re-
garding AE, maize crop was observed with significant performance
under both levels of applied K than any other crop. Even AE was also
lowest in sunflower in both the levels of K applied. In castor although
PE was high this did not reflect in the AE. Apparent recovery efficiency
(RE) was significantly very high in maize as compared to other crops.
Sunflower crop observed with increased RE with K augmentation
i.e., K40 and K60. While the other crops not differed much and showed
on par RE values with K40 and K60.
4. Discussion
4.1. Effect of K Nutrient Management on Chlorophyll Pigments and Caroten-
oids Concentration
Application of K influenced chlorophyll content, chlorophyll a:b ratio
and subsequently total chlorophyll content and it was observed that
high dose of K improved levels of chlorophyll. However, there was a dif-
ferential pattern observed in each crop showing increased rate of K ap-
plication increased chlorophyll-a concentration though no effect on
total chlorophyll concentration. Application of K at 112 kg ha−1 in cot-
ton increased chlorophyll a but did not increase total chlorophyll con-
centration [24]. Deficiency of K reduced chlorophyll a and total leaf
chlorophyll concentration in cotton [25]. Similarly, differential response
was observed for carotenoid concentration of various crops to applied K.
Influence of major factors such as genetic potential of crop, potentiality
of crops to synthesize pigment, plant K nutrition on the chlorophyll and
carotenoid biosynthesis pathways might have resulted in differential
chlorophyll and carotenoids concentration.
4.2. Effect of K Fertilization on Leaf Water Potential
Leaf water potential provides an integrated measurement of soil
water potential as it is sensed by the plants. A close relationship be-
tween K nutritional status and plant drought tolerance has been dem-
onstrated. Past studies already confirmed K application improves plant
resistance to extreme drought stress as it affects water transport in en-
tire to plant maintain cell pressure, regulation of stomata opening and
closing [26,27]. Application of K had a differential influence on leaf

Table 4
Chlorophyll pigment (mg g−1 of fresh weight) levels in leaves of major rainfed crops.

Crop Potassium level (kg ha−1)

K0 K40 K60

Chl a Chl b Total a/b ratio Chl a Chl b Total a/b RATIO Chl a Chl b Total a/b ratio

Cereals and millets

Maize 2.83C 1.37B 4.20C 2.07B 3.76B 1.78A 5.54B 2.12B 6.37A 1.32B 7.69A 4.82A
Pearl millet 2.95C 1.57C 4.52C 1.87B 3.99B 1.91B 5.91B 2.09B 5.29A 2.11A 7.40A 2.51A

Oilseeds
Groundnut 2.69A 0.25C 2.93A 10.84A 2.51B 0.32B 2.84A 7.79B 2.31B 0.69A 3.00A 3.34C
Sunflower 4.42A 0.98A 5.40A 4.53A 4.37A 0.99A 5.35A 4.43A 3.58C 0.88B 4.47B 4.07B
Castor 3.44B 2.05A 5.49A 1.68B 3.94A 1.86A 5.80A 2.13A 3.04C 1.89A 4.93B 1.61B

Fiber crops
Cotton 3.91B 1.48A 5.40A 2.64B 3.19C 1.12B 4.32B 2.84B 4.40A 1.14B 5.54A 3.86A

Capital letters represent level of significance at P = 0.05 among various treatments and different letters indicate significant difference.
 C. Srinivasarao et al. / Journal of Photochemistry & Photobiology, B: Biology 160 (2016) 86–95 93

Table 6
Leaf water potential of major rainfed crops different potassium levels.

Crop Potassium level

K0 K40 K60

Cereals and millets

Maize −1.86A −0.03B −0.02B
Pearl millet −0.14B −0.17A −0.01C

Oilseeds
Groundnut −5.04A −4.69A −3.83B
Sunflower −3.18C −4.81B −5.41A
Castor −2.80B −4.86A −5.09A

Fiber crops
Cotton −0.05C −0.17B
Capital letters represent level of significance at P = 0.05 among various treatments and
different letters indicate significant difference.
water potential of crops. External K source may induce drought toler-
ance by enhancing internal K concentration under water shortage con-
ditions. Reduced leaf water potential under limited K supply reduced
aquaporin activity, consequently suppressing root hydraulic conduc-
tance and water supply to the growing stem for diameter expansion
and leaf for transpiration [26,28]. This could also explain the reduction
in yield that was observed in the plants that were treated with low
dose of K. Results obtained further reveal the close relationship between
K-nutrition and plant water status in the crops studied. Red Soils of
southern plateau with low K containing mica are deficient in K and
rainfed crops in absence of external K application often suffer from op-
timum K requirements and in turn are affected negatively during mois-
ture stress conditions.
4.3. Chlorophyll Fluorescence Kinetics
Mineral nutrition strongly affects the photosynthetic processes in
crops and particularly under different K levels, as evidenced in current
study that K levels not only affected the pigment concentrations but
also the excitation energy transfer processes in the leaves. There are
not many studies conducted on effects of K nutrition on crops OJIP
fluorescence transient curves, maximum quantum yield of primary
photochemistry (Fv/Fm) and its efficiency (ψ0) and photochemical
quenching (qP) and nonphotochemical quenching (npq), Our results
indicate that different levels of K in different crops altered the response
of chlorophyll a fluorescence in comparison with the control. The OJIP
fluorescence transient strongly reduced under low K levels mainly at
the 40 kg ha −1 level. Other parameters such as energy fluxes per reac-
tion center and the performance index were also affected by K nutrition.
In vivo and rapid detection of chlorophyll a fluorescence was used to
detect K level induced effects on light utilization efficiency of the
Fig. 7. Apparent recovery efficiency (RE), physiological efficiency (PE) and agronomic
−0.76A
efficiency (AE) of applied K of major rainfed crops different potassium levels.
photosynthetic machinery. Higher K application in all rainfed crops in-
creased F0 and decreased Fm resulting in a decline in the values of Fv/
Fm, Fv/F0 and Fm/F0 ratios. The F0 represents the estimation of the rel-
ative size of the antenna pigments of the PS II complex especially under
nutrient limiting conditions [29]. This shows damage to the PS II reac-
tion center that in turn results in decreased level of absorption of light
and subsequently causes an increase in unused emitted light. Our re-
sults obtained in relation to the increased F0 level could be due to the
impact of lower K on PS II reaction center in tandem with the reduction
in transfer efficiency of the energy absorbed in antenna chlorophyll a to
reaction center of PS II.
The OJIP fluorescence curve markedly decreased under low K levels
mainly at the P level. Similarly, energy fluxes per reaction center were
also affected by K levels. Fluorescence reduction was more prominent
at the P level which significantly reduced under lower K additions in
all the crops. A lower FM value in control treatment corresponds to a re-
duction of the maximum quantum yield of primary photochemistry
(Fv/Fm), since F0 does not change significantly. The decrease of Fm is
associated with the increase of the non-photochemical de-excitation
constant and the decrease of F0 indicates an enhancement of the overall
de-excitation constants [30]. Conversely, the increase of F0 is considered
as an irreversible damage to the PSII and indicates that heat dissipation
occurs in an uncontrolled manner, this was seen in the npq pattern ob-
served in the crops as a response to K nutrition in the present study,
which could have resulted in production of excess of excitation within
the leaves.
The Fm was sensitive to levels of K and F0 was found to be non-
sensitive. Low Fm indicates an increase of npq when the crops receive
or uptake low K in respective treatments. The decreasing trend of Fm
and low Fv/Fm was also observed in two soybean cultivars highlighting
the effect of low K stress on photosynthetic apparatus [31]. Additionally,
the decreased efficiency (ψ0) in low K treated crops suggests that the
possibility of a trapped excitation that moves an electron into the

Table 7
Yield of major rainfed crops (grain, straw and total) (t ha−1) at different potassium levels.

Crop Potassium level

K0 K40 K60

Grain Straw HI Total Grain Straw HI Total Grain Straw HI Total

Cereals and millets

Maize 3.13B 10.25B 23.39B 13.38C 4.27A 11.68A 26.78A 15.95B 4.46A 12.40A 26.45A 16.86A
Pearl millet 0.29C 4.19B 6.40C 4.48B 0.55B 4.61A 10.71B 5.17A 0.68A 4.77A 12.56A 5.45A

Oilseeds
Groundnut 0.40B 1.35B 22.94B 1.75B 0.54A 1.42B 27.35A 1.96B 0.49A 2.54A 16.27C 3.04A
Sunflower 1.61B 4.75A 25.30A 6.36B 1.61B 5.00A 24.38A 6.61BA 1.81A 5.25A 25.60A 7.06A
Castor 0.73B 0.86B 45.94A 1.59B 1.11A 1.48A 42.84A 2.58A 1.12A 1.42A 44.07A 2.54A

Fiber crops
Cotton 0.53 (2.05)B 3.39A 34.35B 5.98B 0.68(2.16) A 3.49A 33.42B 6.45B 0.71 (2.60)A 3.61A 38.76A 6.71A

HI—harvest index; capital letters represent level of significance at P = 0.05 among various treatments and different letters indicate significant difference, values in parenthesis are lint yield
of cotton.
94 C. Srinivasarao et al. / Journal of Photochemistry & Photobiology, B: Biology 160 (2016) 86–95
electron transport chain beyond QA was reduced and thus these treat-
ments exhibited low maximum quantum yield of primary photochem-
istry (Fv/Fm) and efficiency (ψ0).
Potassium levels altered the fluorescence induction and impaired
photosynthetic systems in all the crops studied. Further, we also see
that the OJIP test is a good indicator to detect fluorescence induction
and photosynthetic activity of the PSII RC complex as affected by K nu-
trition. Disorganization of the chloroplast ultrastructure and inhibition
of electron transport processes due to lower levels of K and impairing
the transport of the electrons in the whole of electron transport chain
from PS II to PS I is a possible explanation for the decrease in photosyn-
thetic rate in K limited crops and this was reflected in the yield of the
crops as seen in this study. In addition, the decline in Fv/Fm ratio under
nutrient limiting conditions can be an indicator of photoinhibition or
some other kind of injury to PS II components. Since the Fv/Fo ratio de-
scribes the activity of PS II while Fm/Fo signifies electron transport rate
through PS II, in the present study, it is possible that K influenced the
activities of PS II and thus decreased these ratios. Decreased level of qP
also suggested the structural and functional photo damages of PS II
under low K.
4.4. Yield Trend of Rainfed Crops as Influenced by K Fertilization
Yield of the crops in terms of grain and straw yield and lint in the
case of cotton and also the harvest index was affected by K levels in
crops studied. Various studies confirmed the response of varied level
of K in different crops [32–36]. Reduction in the yield in the major
crops under low K could have been due to the limited effect of K on pho-
tosynthesis and water status. Efficient use of K during the various stages
need not necessarily translate into K efficiency for economic yield, there
is a possibility that source sink relationship of the crops could have been
altered and more of the assimilates could have been partitioned to the
leaves. It is also known that the pattern of phloem loading is affected
by K, wherein sucrose export to the root is reduced in K-deficient plants,
and adequate K supply is known to play a crucial role in phloem trans-
location of assimilates, this could explain the lower drymatter accumu-
lation in the reproductive structures under lower K levels. Increase of
harvest index was observed in two C4 cereal crops like maize and
Pearl millet compared to oilseed crops (groundnut, sunflower and cas-
tor) and fiber crop (cotton). Again within the C4 cereal crops, increase
of harvest index was higher at K0 to K40 levels of fertilization compared
to K40 to K60 level of fertilization. The difference in harvest index
influenced the efficiency with which applied K could be used to trans-
late into higher grain yield than the capacity to utilize K to produce bio-
mass in the crops. A high harvest index is fundamental to efficient
utilization of all resources taken up by the plant and is therefore of sig-
nificant interest to breeders [37]. The yield reduction observed also
could be due to the low capacity of the crops to translocate K from
non-photosynthetic organs such as stems and petioles to upper leaves
and harvested organs and this in turn could have an influence on the ca-
pacity of the crops to produce a high economic yield per unit of K taken
up.
4.5. Potassium Uptake and Use Efficiency
Uptake of K was seen to be directly correlated to the dosage applied
in all the crops, the higher the dosage there was higher uptake, differen-
tial uptake and utilization efficiency of K in terms of recovery efficiency
was clearly recorded in the study. The K-recovery process is thus seen
here in our experiment as a complex process which comprises of a mix-
ture of uptake and utilization efficiency mechanisms. Compared to
maize, requirement of K is high for Pearl millet as it needs higher K
(nearly double the amount) to produce unit quantity of produce [38].
But, maize crop showed significant increase in K uptake with each incre-
ment of K addition. This may be attributed to effective K uptake that was
influenced by the grand vegetative growth of maize. A similar trend of
early plant K concentration, early K uptake and early leaf K concentra-
tion was observed in maize with increased K fertilization viz., 0, 34.0,
68.1, 136.1, and 204.1 kg K2O/ha− 1 broadcasted before tillage [39].
Among various crops under study sunflower had an increased K uptake
with every additions of K. Sunflower is a highly K responding crop and
uptakes a maximum quantity of applied K hence making it K exhaustive
crop. There are evidences that sunflower crop on average uptakes near-
ly 126 kg of K to produce kg of crop yield. Fiber crop cotton had the
highest K uptake that may be influenced by the stage of crop as K uptake
increases immediately once bolls are set which are major sink for K. Up-
take of K often exceeds nitrogen in crops such as cotton, maize, sugar-
cane and sunflower ranging from 408 to 625 kg K2O/ha under
intensive cropping system every season. During entire crop growth pe-
riod translocation and redistribution of K within and among vegetative
and reproductive parts of cotton plant occurs [40].
Physiological and morphological traits such as leaf characters, plant
canopy, growth and development, root hairs, root exudates, rooting
density and distribution, rooting depth, plant need for K, [41] and geno-
typic characters of crop plants might have influenced the K uptake [42].
The rooting pattern of the crops under study might have influenced K
uptake and recovery. Crops efficiency in K uptake is influenced by effi-
cient root interception due to larger surface area of contact between
roots and soil subsequently enabling increased uptake at the root–soil
interface to maintain a larger diffusive gradient towards roots [37].
Trend of K uptake by various crops is sunflower N Pearl millet N
cotton N maize N groundnut N castor as evidenced by different studies
[43] across the country. A similar tendency of K uptake was observed
in different rainfed crops under study. A difference in K uptake was in-
fluenced by rate of K applied in maize and soybean crop and resulted
in accumulation of K in grain and straw respectively 60 and 9% for soy-
bean and 57 and 7% for maize [44].
5. Conclusion
Physiological growth and development of major rainfed crops under
varied rates of K application differed significantly influencing nutrient
uptake and ultimate yield. Potassium application considerably in-
creased total chlorophyll (Chl a + Chl b) and carotenoid content in
maize and Pearl millet whereas, chlorophyll content of other crops did
not differ much even with K addition. Depiction of soil–plant water re-
lations through leaf water potential measurements differed among
crops. Application of K upto 60 kg ha− 1 induced monocots to grow
well than oilseeds and cotton under rainfed conditions. Sunflower, a
known K exhaustive crop comparatively extracted maximum amount
of applied K that resulted in high RE but low PE and AE. Use efficiency
of applied K was better i.e., high PE in castor and groundnut. In our re-
sults, we observed that the fluorescence yield decreased in the crops
as K dosage increased, evidenced by increasing of all points (O, J, I
and P) of the OJIP curves. Chlorophyll fluorescence parameters
viz., maximum quantum yield of primary photochemistry (Fv/Fm)
and its efficiency (ψ0) and photochemical quenching (qP) and
nonphotochemical quenching (npq) where affected by K applica-
tion rates. The fluorescence transient curve for K60 was lower
than K0 and K40 for all the crops. This study highlights the impor-
tance of K nutrition in rainfed crops in one or the other way that di-
rectly gives acceptable yields. Alfisols in south India being K
deficient heavy uptake of K over a period of time excluding K appli-
cation in nutrient management may lead these soils severely deficit
in K that ultimately affects physiology of crop plants resulting in re-
duced yields.
Acknowledgments
The authors are thankful to ICAR for providing laboratory and field
facilities for the study.
 C. Srinivasarao et al. / Journal of Photochemistry & Photobiology, B: Biology 160 (2016) 86–95
References
[1] N.C. Brady, R.R. Weil, The Nature and Properties of Soils, Pearson/Prentice Hall, New
Jersey, USA, 2010.
[2] A.P. Singh, Potassium in Soil–Plant System: a Wider Perspective, 39th Dr. R.V.
Tamhane Memorial Lecture, 77th Annual Convention of Indian Society of Soil Sci-
ence, Ludhiana, India 2012, pp. 1–16.
[3] Ch. Srinivasarao, Sumanta Kundu, B.K. Ramachandrappa, Sharanbhoopal Reddy,
Rattan Lal, B. Venkateswarlu, K.L. Sahrawat, R. Prakash Naik, Potassium release char-
acteristics, potassium balance, and fingermillet (Eleusine coracana G.) yield sustain-
ability in a 27-year long experiment on an Alfisol in the semi- arid tropical India,
Plant Soil 374 (2014) 315–330.
[4] Ch. Srinivasarao, Sharan Bhoopal Reddy, Sumanta Kundu, Potassium nutrition and
management in Indian agriculture, Indian J. Fertil. 10 (5) (2014) 58–80.
[5] Bijay-Singh, Yadvinder-Singh, P. Imas, X. Jian-Chang, Potassium nutrition of the rice-
wheat cropping system, Adv. Agron. 81 (2004) 203–259.
[6] Y.G. Prasad, M. Maheswari, S. Dixit, Ch. Srinivasarao, A.K. Sikka, B. Venkateswarlu, N.
Sudhakar, S. Prabhu Kumar, A.K. Singh, A.K. Gogoi, A.K. Singh, Y.V. Singh, A. Mishra,
Smart Practices and Technologies for Climate Resilient Agriculture, ICAR-Central Re-
search Institute for Dryland Agriculture, Hyderabad, India, 2014 1–76.
[7] G.H. Krause, E. Weis, Chlorophyll fluorescence and photosynthesis, the basics, Annu.
Rev. Plant Physiol. Plant Mol. Biol. 42 (1991) 313–349.
[8] A.A. Thwe, P. Kasemsap, Quantification of OJIP fluorescence transient in tomato
plants under acute ozone stress, Kasetsart J. (Nat. Sci.) 48 (2014) 665–675.
[9] W.W. Adams III, B. Demming-Adams, in: G.C. Papageorgiou, Govindjee (Eds.), Chlo-
rophyll Fluorescence as a Tool to Monitor Plant Response to Environment, Springer,
Dordrecht, The Netherlands 2004, pp. 583–604.
[10] R.J. Strasser, A. Srivastava, M. Tsimilli- Michael, in: M. Yunus, U. Pathre, P. Mohanty
(Eds.), The Fluorescence Transient as a Tool to Characterize and Screen Photosyn-
thetic Samples, Taylor and Francis, New York, NY, USA 2000, pp. 445–483.
[11] R.J. Strasser, M. Tsimilli-Michael, A. Srivastava, in: G.C. Papageorgiou, Govindjee
(Eds.), Analysis of the Chlorophyll a Fluorescence Transient, Springer, Dordrecht,
The Netherlands 2004, pp. 321–362.
[12] A.R. Wellburn, The spectral determination of chlorophylls a and b, as well as total
carotenoids, using various solvents with spectrophotometers of different resolution,
J. Plant Physiol. 144 (1994) 307–313.
[13] D.I. Arnon, Copper enzymes in isolated chloroplasts, polyphenoxidase in beta
vulgaris, Plant Physiol. 24 (1949) 1–15.
[14] L.B. Thach, A. Shapcott, S. Schmidt, C. Critchley, The OJIP fast fluorescence rise char-
acterizes Graptophyllum species and their stress responses, Photosynth. Res. 94
(2007) 423–436.
[15] W.H. Herdershort, H. Lalande, M. Duquette, in: M.R. Carter (Ed.), Soil Reaction and
Exchangeable Acidity, Lewis Publishers, Boca Raton, FL. 1993, pp. 141–145.
[16] D.W. Nelson, L.E. Sommers, in: D.L. Sparks, A.L. Page, P.A. Helmke, R.H. Loeppert, P.N.
Soltanpour, M.A. Tabatabai, C.T. Johnston, M.E. Sumner (Eds.), Total Carbon, Organic
Carbon, and Organic Matter, Part 3, Chemical Methods, SSSA Book Series No. 5, SSSA
and ASA, Madison, WI 1996, pp. 961–1010.
[17] B.V. Subbiah, G.L. Asija, A rapid procedure for the determination of available nitro-
gen in soils, Curr. Sci. 25 (1956) 259–260.
[18] S. Kuo, in: D.L. Sparks, A.L. Page, P.A. Helmke, R.H. Loeppert, P.N. Soltanpour, M.A.
Tabatabai, C.T. Johnston, M.E. Sumner (Eds.), Phosphorus, Part 3, Chemical Methods,
Soil Science Society of America Book Series No. 5, SSSA and ASA, Madison, WI 1996,
pp. 869–919.
[19] P. Helmke, D.L. Sparks, in: D.L. Sparks, A.L. Page, P.A. Helmke, R.H. Loeppert, P.N.
Soltanpour, M.A. Tabatabai, C.T. Johnston, M.E. Sumner (Eds.), Lithium, Sodium, Po-
tassium, Rubidium, and Cesium, Part 3, Chemical Methods, Soil Science Society of
America Book Series No. 5, SSSA and ASA, Madison, WI 1996, pp. 551–574.
[20] W.L. Lindsay, W.A. Norvell, Development of a DTPA soil test for zinc, iron, manga-
nese, and copper, Soil Sci. Soc. Am. J. 42 (1978) 421–428.
[21] R. Keren, in: D.L. Sparks, A.L. Page, P.A. Helmke, R.H. Loeppert, P.N. Soltanpour, M.A.
Tabatabai, C.T. Johnston, M.E. Sumner (Eds.), Boron, Part 3, Chemical Methods, Soil
Science Society of America Book Series No. 5, SSSA and ASA, Madison, WI 1996,
pp. 603–626.
[22] K.L. Sahrawat, G. Ravi Kumar, J.K. Rao, Evaluation of triacid and dry ashing proce-
dures for determining potassium, calcium, magnesium, iron, zinc, manganese and
copper in plant materials, Commun. Soil Sci. Plant Anal. 33 (2002) 95–102.
95
[23] A.R. Mosier, J.K. Syers, J.R. Freney, Agriculture and the Nitrogen Cycle, Assessing the
Impacts of Fertilizer Use on Food Production and the Environment, Scope-65, Island
Press, London, 2004.
[24] J.L. Fridgen, J.J. Varco, Dependency of cotton leaf nitrogen, chlorophyll, and reflec-
tance on nitrogen and potassium availability, Agron. J. 96 (1) (2004) 63–69.
[25] D.M. Oosterhuis, C.W. Bednarz, in: T. Ando, K. Fujita, T. Mae, H. Matsumoto, S. Mori, J.
Sekiya (Eds.), Physiological Changes during the Development of Potassium Deficien-
cy in Cotton, Kluwer Academic Publishers, Japan 1997, pp. 347–351.
[26] Y. Wang, K. Noguchi, N. Ono, S. Inoue, I. Terashima, T. Kinoshita, Over expression of
plasma membrane H+-ATPase in guard cells promotes light-induced stomatal
opening and enhances plant growth, Proc. Natl. Acad. Sci. U. S. A. 111 (2014)
533–538.
[27] G.R. Kudoyarova, I.C. Dodd, D.S. Veselov, S.A. Rothwell, S.Y. Veselov, Common and
specific responses to availability of mineral nutrients and water, J. Exp. Bot. 17
(2015) 45–52.
[28] S. Kanai, R.E. Moghaieb, H.A. El-Shemy, R. Panigrahi, P.K. Mohapatra, J. Ito, K. Fujita,
Potassium deficiency affects water status and photosynthetic rate of the vegetative
sink in green house tomato prior to its effects on source activity, Plant Sci. 180 (2)
(2011) 368–374.
[29] Z.A. Huang, D.A. Jiang, Y. Yang, J.W. Sun, S.H. Jin, Effects of nitrogen deficiency on gas
exchange, chlorophyll fluorescence, and antioxidant enzymes in leaves of rice
plants, Photosynthetica 42 (2004) 357–364.
[30] J. Harbinson, A.E. Prinzenberg, W. Kruijer, M.G. Aarts, High throughput screening
with chlorophyll fluorescence imaging and its use in crop improvement, Curr.
Opin. Biotechnol. 23 (2) (2012) 221–226.
[31] X.T. Li, P. Cao, X.G. Wang, M.J. Cao, H.Q. Yu, Comparison of gas exchange and chloro-
phyll fluorescence of low-potassium-tolerant and-sensitive soybean [Glycine max
(L.)Merr.] cultivars under low-potassium condition, Photosynthetica 49 (4) (2011)
633–636.
[32] M.W. Clover, A.P. Mallarino, Corn and soybean tissue potassium content responses
to potassium fertilization and relationships with grain yield, Soil Sci. Soc. Am. J. 77
(2013) 630–642.
[33] S. Qiu, J. Xie, S. Zhao, X. Xu, Y. Hou, X. Wang, J. Jin, Long-term effects of potassium
fertilization on yield, efficiency, and soil fertility status in a rain-fed maize system
in northeast China, Field Crop Res. 163 (2014) 1–9.
[34] S. Zhao, P. He, S. Qiu, L. Jia, M. Liu, J. Jin, A.M. Johnston, Long-term effects of potassi-
um fertilization and straw return on soil potassium levels and crop yields in north-
central China, Field Crop Res. 169 (2014) 116–122.
[35] N. Sui, Z. Zhou, C. Yu, R. Liu, C. Yang, F. Zhang, Y. Meng, Yield and potassium use ef-
ficiency of cotton with wheat straw incorporation and potassium fertilization on
soils with various conditions in the wheat–cotton rotation system, Field Crop Res.
172 (2015) 132–144.
[36] Q. Lu, D. Jiaa, Y. Zhang, X. Dai, M. He, Split application of potassium improves yield
and end-use quality of winter wheat, Agron. J. 106 (2014) 1411–1419.
[37] Z. Rengel, P.M. Damon, Crops and genotypes differ in efficiency of potassium uptake
and use, Physiol. Plant. 133 (4) (2008) 624–636.
[38] K.G. Pillai, L. Suseela Devi, M.V. Sholapurkar, M.K. Jagannadh, Effect of continuous
cropping and manuring in selected cropping systems, Fertil. News 32 (6) (1987)
31–37.
[39] M.W. Clover, A.P. Mallarino, P.A. Barbagelata, Corn and Soybean Grain Yield and
Concentration of Potassium in Plant Tissues and Soil as Affected by Potassium Fertil-
ization, Proceedings of the Thirty-Seventh North Central Extension-Industry Soil
Fertility Conference, Vol. 23 2007, pp. 119–125.
[40] F. Yang, M. Du, X. Tian, A. Egrinya Eneji, L. Duan, Z. Li, Plant growth regulation en-
hanced potassium uptake and use efficiency in cotton, Field Crop Res. 163 (2014)
109–118.
[41] W.T. Pettigrew, Potassium influences on yield and quality production for maize,
wheat, soybean and cotton, Physiol. Plant. 133 (2008) 670–681.
[42] C. Zörb, M. Senbayram, E. Peiter, Potassium in agriculture — status and perspectives,
J. Plant Physiol. 171 (2013) 656–669.
[43] Fertilizer Statistics, Fifty eighth ed., Fertilizer Association of India, New Delhi, India,
2012–13.
[44] R.R. Oltmans, A.P. Mallarino, Potassium uptake by corn and soybean, recycling to
soil, and impact on soil test potassium, Soil Sci. Soc. Am. J. 79 (1) (2015) 314–327.