CHAPTER THREE

The Physiology of Potassium

in Crop Production
Derrick M. Oosterhuis*,1, Dimitra A. Loka*,
Eduardo M. Kawakami* and William T. Pettigrew†
*University of Arkansas, Department of Crop, Soil and Environmental Sciences, Fayetteville, AR, USA
†ARS-USDA, Stoneville, MS, USA
1Corresponding author: e-mail address: [email protected]

Contents
1.  Introduction204
2.  Physiology of Potassium Nutrition 204
2.1  Enzyme and Organic Compound Synthesis Regulation 205
2.2  Water Relations 206
2.3  Leaf Movements 208
2.4  Meristematic Growth and Plant Growth Regulation 209
2.5  Stomatal Regulation 211
2.6 Photosynthesis 212
2.7 Respiration 214
2.8  Assimilate Transport 214
2.9  Nitrate Transport–Potassium Interactions 215
2.10  Potassium Channels 217
3.  Stress Mitigation 218
3.1  Drought Stress 218
3.2  Cold Stress 219
3.3  Salt Stress 220
3.4  Biotic Stress 221
3.5  Potassium and Stress Signaling 222
4.  Summary223
References223

Abstract
Potassium (K) plays a major role in the basic functions of plant growth and develop-
ment. In addition, K is also involved in numerous physiological functions related to
plant health and tolerance to biotic and abiotic stress. However, deficiencies occur
widely resulting in poor growth, lost yield, and reduced fiber quality. This review
describes the physiological functions of K and the role in stress relief and also provides
some agronomic aspects of K requirements, diagnosis of soil and plant potassium
status, and amelioration. The physiological processes described include enzymes and

Advances in Agronomy, Volume 126 © 2014 Elsevier Inc.

ISSN 0065-2113, http://dx.doi.org/10.1016/B978-0-12-800132-5.00003-1 All rights reserved. 203
204 Derrick M. Oosterhuis et al.

organic compound synthesis regulation, water relations and stomates, photosynthesis,

transport, cell signaling, and plant response to drought stress, cold stress, salt stress, as
well as biotic stresses.

1.  INTRODUCTION
Potassium (K) is the mineral element, next to nitrogen, required in
the largest amount by plants. For optimal growth and productivity, modern
crop production requires a large amount of K, particularly during reproduc-
tive development. The K requirement for optimal plant growth is 2–5% of
the plant dry weight (Marschner, 1995), however, the amount of K utilized
by the crop varies depending on the crop species, and the quantity of K+
in the soil available to the plants. Additionally, K uptake is influenced by
the environmental conditions during the growing season and the manage-
ment practices used (Mullins et al., 1997). Proper plant nutrition for optimal
crop productivity requires that nutrient deficiencies be avoided. However,
potassium deficiencies have been reported to occur all around the world
for a variety of reasons such as soil types and management practices (Ren-
gel and Damon, 2008), or removal of crop residues for use in the biofuel
industry (Romhel and Kirkby, 2010). In addition, even though farmers in
the USA and elsewhere are using substantially more commercial fertilizer
than 40 years ago the ratio of nitrogen: potassium input has been signifi-
cantly decreased from 100:63 in 1960 to 100:27 in 2000 (Maene, 2001).
This has prompted a renewed focus on K management with some emphasis
on understanding K fertilizer requirements and use by the plants. An effi-
cient fertilizer regime requires an accurate knowledge of the nutrient status
of the soil, as well as a reliable tissue analysis during the season to fine-tune
the fertility status and avoid any unforeseen deficiencies. Fundamental to
this is an understanding of the role of the nutrient in plant metabolism and
yield formation. This review provides an overview of the physiology of K
nutrition in crop growth, and provides an overview of K fertility require-
ments and deficiency symptoms in field crops.

2.  PHYSIOLOGY OF POTASSIUM NUTRITION

Potassium (K) is the most abundant cation in the plant cells, with
high mobility within short-distance transport (i.e., between individual cells
and between neighbor tissues), as well as within long distance transloca-
tion through the xylem and phloem (Marschner, 1995). Since K is not
The Physiology of Potassium in Crop Production 205

metabolized into organic compounds in the plant, K is predominately pres-

ent in the cationic form (Wyn Jones et al., 1979). Potassium can be stored,
both in the cell cytoplasm and vacuole, and the distribution among these
two locations is the major factor for determining the K function in the
plant (Marschner, 1995). These characteristics convey K as a major nutrient
responsible for controlling many physiological and biochemical processes
in the plant, such as enzyme activation, cell osmotic potential regulation,
soluble and insoluble molecular anions neutralization, and cell pH stabiliza-
tion (Marschner, 1995). This section of this review will summarize the role
of K in a few key aspects of plant physiology.

2.1  Enzyme and Organic Compound Synthesis Regulation

In the plant, K is responsible for activation and/or stimulation of a number of
enzymes (Suelter, 1970). The activation of enzymes is resulted from a change
in the conformation of the inactive enzyme structure caused by a ligation of
K to a specific site in the protein (Marschner, 1995). Potassium availability has
an influence in the activity of more than 60 enzymes (Wyn Jones and Pollard,
1983) involved in a variety of metabolic process as mainly related to protein
and carbohydrates synthesis (Marschner, 1995; Mengel et al., 2001).
The optimum concentration of K for maximum enzyme activation is
about 50 mM (Nitsos and Evans, 1969). However cytoplasmic concentra-
tion of K is known to be between 100 and 200 mM (Leigh and Wyn Jones,
1984) in order to optimize protein synthesis in the cell (Wyn Jones et al.,
1979). For this reason, cytoplasmic K concentration is maintained constant
by the pool of K in the vacuole (Leigh and Wyn Jones, 1984; Walker et al.,
1996), and changes in the level of cytoplasmic K will be observed only when
vacuolar K concentration has been depleted to a minimal vital concentra-
tion (Walker et al., 1996). The importance of K in protein synthesis is clear
when a decrease in N incorporation into protein is exhibited by K deficient
plants (Marschner, 1995; Mengel et al., 2001). Amino acid polymerization
is one of the main steps where K has been shown to regulate protein turn-
over in plants (Conway, 1964). In the review of Mengel et al. (2001), a K
effect on ribulose bisphosphate carboxylase activity, nitrate reduction pro-
cess, and ribosome polypeptide syntheses have been described. Furthermore
Marschner (1995) reported a central role of K in the translation process, and
activity and synthesis of nitrate reductase.
In carbohydrates synthesis, K has been reported to affect a number of
enzymes including, glucose starch synthase (Hawker et al., 1974, 1979), glu-
cose pyrophosphorylase (Hawker et al., 1979), β-amylase (Berg et al., 2009),
206 Derrick M. Oosterhuis et al.

sucrose synthase (Berg et al., 2009), invertase (Ward, 1960), amylase (Li et al.,
1997), phosphofructosekinase (Lauchli and Pfluger, 1978), pyruvate kinase
(Memon et al., 1985; Matoh et al., 1988). Among all enzymes affected by K
availability, pyruvate kinase is probably the most important. Pyruvate kinase
has a central role in plant metabolism because it regulates the conversion of
phosphoenol pyruvate to pyruvate (Kayne, 1973). Armengaud et al. (2009),
studying the effect of K nutrition in Arabidopsis roots, observed that despite
the fact that a number of enzymes involved in the glycolysis and N assimila-
tion processes were regulated by K, the primary effect of low K availability
on metabolic disorders were directly related to inhibition of pyruvate kinase
activity.The central role of K on carbohydrates synthesis has been described
to be the main reasons for the presence of high concentration of reduc-
ing sugars, and low starch content in K deficient plants (Marschner, 1995;
Amtmann et al., 2008). In cotton, K fertilization increased leaf protein con-
tent (Akhtar et al., 2009), and decreased leaf starch (Bednarz and Ooster-
huis, 1999; Akhtar et al., 2009) and sucrose (Zhao et al., 2001). Similarly,
Pettigrew (1999) reported that K application decreased leaf glucose, root
starch, root glucose, and root fructose content. This effect on carbohydrates
was possibly due to an increase in metabolites utilization and translocation
(Bednarz and Oosterhuis, 1999). The effect of K on phloem translocation
will be discussed in more details later in this review.

2.2  Water Relations

Potassium plays an integral role in plant–water relations (Hsiao and Läuchli,
1986) and is involved in numerous physiological functions within the plant
where water is involved including stomatal opening and closing, assimilate
translocation, enzyme activation, and leaf heliotropic movements.Water poten-
tial, the thermodynamic energy status of water, is well recognized as an indi-
cator of plant–water status (Begg and Turner, 1976). Water potential consists
of various components, the main ones being osmotic potential and pressure
potential.The magnitude of these two components regulates water potential.
While organic compounds synthesis is regulated by cytoplasmic K con-
centration, water potential is mainly affected by K concentration in the
cell vacuole (Hsiao and Lauchi, 1986). Potassium salts (e.g., KNO3, KCl,
K-malate) are the common forms of K in the vacuoles, which have a major
role in regulating cell osmotic potential in order to maintain adequate tur-
gor pressure for cell functioning (MacRobbie, 1977). The K content in the
vacuole has a sole purpose of “biophysical” regulation, and no biochemical
functions have been described (Leigh and Wyn Jones, 1984). Potassium is the
The Physiology of Potassium in Crop Production 207

most important vacuolar solute that regulates cell osmoregulation process

(Mengel and Arneke, 1982; Beringer et al., 1986; Morgan, 1992). However,
in contrast with the cytoplasmic pool, the vacuolar K can be replaced by
other solutes, i.e., Na (Bednarz and Oosterhuis, 1999), Mg, and amino acids
for maintenance of vital water potential values, when K availability is low
(Leigh and Wyn Jones, 1984). Vacuolar K content can be variable, ranging
from 9 to 174 mM depending on plant species and growing medium (Hsiao
and Lauchi, 1986). Leigh and Wyn Jones (1984) reported that the critical
minimum concentration of K in the vacuole is about 10–20 mM, and if
vacuolar K concentration falls below these values an effect in the cytoplas-
mic K pool can occur resulting in disruption of vital metabolic processes
(i.e., enzyme activation and protein synthesis). In cotton, K fertilization was
reported to increase turgor pressure and decrease water and osmotic poten-
tial values (Pervez et al., 2004). Pettigrew (1999) observed that application
of K to cotton plants resulted in an increase of 17% in turgor pressure, and
did not affect water and osmotic potential of leaf tissues.
Another important aspect of K in regulating plant–water relation attri-
butes is the possible role of K in cell osmotic adjustment. Osmotic adjust-
ment is the process of increasing solute concentration in the cell vacuole in
order to maintain lower values of water potential during periods of salin-
ity and/or water deficit stress (Hsiao et al., 1976; Morgan, 1984; Taiz and
Zeigher, 2010). In water-stressed cotton, although osmotic adjustment is
known to occur both in leaf and root tissue, a greater effect was observed
in roots compared to leaves (Oosterhuis and Wullshleger, 1987). Potassium
ions have been reported to accumulate under water and/or salinity stress in
different crops, such as sorghum (Sorghum bicolor) (Weimberg et al., 1982),
sunflower (Helianthus annuus, L.) (Jones et al., 1980), beans (Vicia faba L.)
(Mengel and Arneke, 1982), and annual clovers (Trifolium sp.) (Iannucci
et al., 2002). Morgan (1992) observed that 78% of the osmotic adjustment
in water-stressed wheat (Triricum aestivum L.) plants was attributed to K, and
only 22% to organic solutes. In water-stressed chickpeas (Cicer arietinum), K
importance in regulation of osmotic adjustment processes decreased with
plant age (Moinuddin and Imas, 2007). In contrast, K does not appear to
play a major role in osmotic adjustment of cotton plants. In an early study
of Cutler and Rains (1978), K concentration did not change with water
potential of drought stressed cotton; in this case organic solutes (i.e., soluble
sugars and malate) were the main regulator of the osmoregulation process.
Similarly, Stark (1991) observed that although K accumulated in leaves of
salt-stressed cotton plants, K by itself was not a precondition for osmotic
208 Derrick M. Oosterhuis et al.

adjustment regulation. Results of no effect of K on osmotic adjustment have

also been reported in millet (Pennisetum glaucum) (Ashraf et al., 2002), maize
(Zea mays L.) (Sharp et al., 1990), sorghum (Turner et al., 1978), and panic
grass (Panicum scribnerianum) (Ford and Wilson, 1981). However, K has been
reported to stimulate osmotically active solute, such as malate (Beringer,
1978 cited by Moinuddin and Imas, 2007) and proline (­Weimberg et al.,
1982), thus it is likely that K can also indirectly affect osmotic adjustment
of plants.

2.3  Leaf Movements

Certain plants exhibit reversible leaf movements in response to environmental
conditions (Satter and Galston, 1973). Leaf movements (paraheliotropic) in
leguminous crops have been well documented (Kawashima, 1969; Oosterhuis
et al., 1985; Berg and Heuchlin, 1990), and have also been reported in other
crops such as cotton (diaheliotropic) (Miller, 1975). The organ of movement
of the leaf is the pulvinus (or the pulvinule of a leaflet) situated at the point
where the petiole joins the leaflet lamina (Satter and Galston, 1981). Differen-
tial changes in osmotic potential in different parts of the pulvinus have been
used to explain the movements (Carlson, 1973; Gorton, 1987). Leaf move-
ments in Albizzia and Samanea are apparently controlled by differential turgor
changes in the pulvinal motor cells (Satter and Galston, 1981) which, in turn,
appear to be a consequence of K+ flux into and out of the pulvinus (Sat-
ter and Galston, 1973; Schrempf et al., 1976). Oosterhuis and Walker (1984)
reported that the bending and straightening of soybean (Glycine max) leaflets
under conditions of water stress were due to differential changes in osmotic
potential (Ψs) and turgor (Ψp) in the ventral and dorsal sides of the pulvinule
associated with changes in K concentration.The greatest change in Ψs and Ψp
was shown to coincide with the maximum rate of change in leaflet angle with
the onset of water stress.They suggested that under conditions of water stress
K+ flux may have the role of regulating movements of the leaflet by inducing
changes in turgor in opposing sides of the pulvinule. Ion channels are under-
stood to be the conduits for the movement of K+ (Koller, 2000), and water
channels (aquaporins) serve as the water conduits through the pulvinule cell
membranes (Moshelion et al., 2002; Uehlein and Kaldenhoff, 2008).
The importance of K in the regulation of cell osmotic potential and
subsequently turgor pressure, results in an indirect effect of K in other phys-
iological processes, such as cell growth, stomata movement, and photosyn-
thesis, which will be discussed in more details throughout this review.
The Physiology of Potassium in Crop Production 209

2.4  Meristematic Growth and Plant Growth Regulation

Potassium is essential for cell growth, a vital process for adequate plant func-
tioning and development (Marschner, 1995; Hepler et al., 2001). The most
acceptable concept for plant cell elongation is known as the acid growth.
This theory involves three different processes: acid-induced cell wall loosen-
ing, osmoregulation, cell wall synthesis, and deposition (Rayle and C ­ leland,
1992). The cell growth is initiated with an acidification of the cell apo-
plast that triggers cell wall loosening and activation of hydrolyzing enzymes
(Hager et al., 1971). This acid stimulus is regulated by the ATPase in the
plasmalemma that pumps H+ from the cytoplasm to the apoplast. The role
of K in this process is to stimulate and control the ATPase cycle in the cell
(Mengel et al., 2001). Potassium has been reported to stimulate ATPase in
chloroplast inner envelope vesicles (Shingles and McCarty, 1994) and to
regulate the activity of ATPase by controlling the dephosphorylation of
subunit in the ATPase structure (Buch-Pedersen et al., 2006). The impor-
tance of osmoregulation in cell growth is to regulate wall stress relaxation
in the cell (Taiz and Zeiger, 2010) and the role of K on cell osmoregulation
was described above. The major constraint for cell growth is the rigid cell
wall, and without wall stress relaxation, the cell growth would only result
in thicker cell walls.The stress relaxation is responsible for a mechanical cell
expansion, and it is regulated by high turgor pressure that is caused by water
intake, due to a decrease in osmotic potential (Taiz and Zeiger, 2010). The
last steps for the cell growth process include cell wall synthesis and deposi-
tion. Cell walls are composed by cellulose, hemicellulose, lignin, pectin, and
protein (Taiz and Zeiger, 2010). As previously mentioned, K is important
for carbohydrates and protein synthesis, thus K can indirectly affect cell
growth by regulating the production of major components utilized in cell
wall metabolism. Furthermore, K can also affect the availability of cell wall
constituents by regulating phloem transport, this effect will be discussed
later in the review. In addition to being one of the cell wall components,
proteins can also be responsible for regulating the acid-induced cell wall
loosening process (Taiz and Zeiger, 2010).
Another important role of K in cell growth is the regulation of plant
growth regulators (Mengel et al., 2001). Auxin (IAA) activity is the primary
controller of the acid growth process (Rayle and Cleland, 1992). However,
IAA-induced cell elongation is dependent of protein synthesis (Cocucci
and Rosa, 1980), since K regulates cell protein turnover, the occurrence
of a combined effect between K and IAA can exist. Claussen et al. (1997),
studying the relationship between K channels and IAA, observed that the
210 Derrick M. Oosterhuis et al.

IAA-induced cell growth in maize did not occur in the absence of K+

ions. Furthermore, an effect of IAA on K channel activity has also been
reported (Thiel and Weise, 1999). A synergistic effect between gibberel-
lin (GA) and K has also been reported (Marschner, 1995). La Guardia and
Benlloch (1980) observed that GA regulates plant remobilization of K and
that stem elongation was favored when both GA and K were present. In
grape fruits application of GA has been reported to increase the concentra-
tion of K in the fruit (Zhenming et al., 2008). In wheat, the presence of K
has been reported to be indispensable for the GA induced stem elongation
process (Chen et al., 2001). Similar results were observed by Nishizawa et al.
(2002), in which they reported that K must be taken up from the soil, in
order for wheat stem elongation to occur. Potassium has also been reported
to affect the role of cytokinin in plants (Mengel et al., 2001). Green and
Muir (1978) observed that the stimulation of cucumber cotyledons growth
caused by cytokinin was enhanced by the presence of K. In contrast to
K causing an increase in the growth response of cytokinin, K application
has been reported to decrease ethylene evolution in cucumber seedlings
(Green, 1983). Similarly, drought stressed sunflower grown in adequate lev-
els of K exhibited a decrease in ethylene synthesis in comparison to plants
under low K conditions (Benlloch-Gonzalez et al., 2010). Ethylene produc-
tion appears to be important in the tolerance and regulation of K deficiency
stress response in plants (Jung et al., 2009). Green and Muir (1979) studying
the relationship among K, cytokinin, and abscisic acid (ABA) on cotyledon
expansion, observed that both the cytokinin growth stimulation and ABA
growth inhibition was higher with the presence of K. They concluded that
the ABA growth inhibition was mediated by an interference in the uptake
and accumulation of K in the plant. Furthermore, ABA has been reported
to regulate the transport and accumulation of K in roots of higher plants
(Roberts and Snowman, 1999); and also to decrease K uptake by inhibiting
the K+/H+ ion exchange system (Watanabe and Takahashi, 1997). Other
compounds, such as jasmonic acid (Ashley et al., 2006) and polyamines
(Adams et al., 1990; Sarjala and Kaunisto, 1993) have also been reported to
be affected by K availability. In addition, polyamines are also involved in the
regulation of K channels (Kusano et al., 2008).
In cotton, the fiber development is probably the most evident process in
which K affects cell growth. As previously mentioned, water potential regu-
lation is an important event in cell growth. Dhindsa et al. (1975) reported
that K and malate accounted for more than 50% of the fiber osmotic poten-
tial during fiber growth.They also observed that the level of K peaked when
The Physiology of Potassium in Crop Production 211

fiber growth was the highest and that fiber growth was adversely affected
by the absence of K. Similarly, Ruan et al. (2001) reported that high expres-
sion of K+ transporters occurs during cotton fiber development. The K
availability is known to have an influence in cotton growth, with effects on
plant leaf area index (LAI), number of main stem nodes, plant height, and
plant dry matter (Pettigrew and Merdith, 1997). Reddy and Zhao (2005)
reported that K fertilization increased plant growth and biomass partition-
ing to fruits. Growth of cotton roots is also affected by K, Zhang et al.
(2009) observed that lack of K decreased root growth, as a result of low IAA
and high ethylene activity. Additional K effects on cotton growth have been
described in different studies (Bednarz and Oosterhuis, 1999; ­Pettigrew,
1999, 2003; Zhao et al., 2001).

2.5  Stomatal Regulation

Stomatal functioning is important for providing CO2 for photosynthesis
while keeping water losses to a minimal level, thus the regulation of
stomata opening and closing is crucial for efficient plant productiv-
ity. The mechanism of stomates movement is known to be regulated
by turgor pressure, which is controlled mostly by K ion concentration
(Marschner, 1995). The concentration of K on stomates guard cells,
depending on plant species, can increase from 100 mM, when closed, to
800 mM, when opened (Taiz and Zeiger, 2010). A detailed explanation
of the stomates activity has been described by Taiz and Zeiger (2010).
In summary, light radiation into the plant cell activates three important
metabolic processes involved in stomates opening: proton pump ATPase,
solute uptake, and organic solute synthesis. The H+ pumping ATPase
creates an electrochemical potential that favor the uptake of K and the
companion anions Cl− and/or malate. The increase of these solutes and
sucrose in the guard cell vacuole cause a decrease in the osmotic poten-
tial. Then, an uptake of water occurs, increasing turgor pressure, which
results in stomate opening. In contrast, the closure of stomates is mainly
regulated by ABA activity. In this process, the ABA stimulates Ca uptake
into the cell, which blocks the K+ channels and favors the extrusion of
anions (Cl−) into the cell apoplast. The increase in the intercellular Ca
inhibits the proton pump ATPase, causing a depolarization in the cell
membrane, resulting in extrusion of vacuolar and cytoplasmic K+ to the
cell apoplast. Thus, stomates close due to a decrease in turgor pressure,
caused by high osmotic potential due to low intercellular solute con-
centration.
212 Derrick M. Oosterhuis et al.

Recently, an unexpected role of K in plant stomatal regulation was

described by Benlloch-Gonzalez et al. (2008). They observed that K depri-
vation increased stomatal conductivity of water-stressed plants. Report of
increased water uptake and decreased water use efficiency under K defi-
ciency also support this evidence (Founier et al., 2005). However, since
K is important in the stomatal opening process, the lack of K is expected
to decrease, not increase, plant stomatal conductivity. An interaction effect
between ABA and ethylene has been proposed to explain this behavior
(Benlloch-González et al., 2010). As previously mentioned, ABA regulates
stomatal closure and ethylene synthesis increases under K deficiency. Inhi-
bition of stomatal closure caused by ethylene interference with the activity
of ABA has been reported (Tanaka et al., 2005). Benlloch-Gonzalez et al.
(2010) could not confirm this effect, but they observed that increased sto-
matal conductance in K deficient water-stressed plants disappeared with
application of an ethylene synthesis inhibitor.They concluded that the high
stomatal conductance under deprivation of K, could be a mechanism to
increase xylem sap movement, in order to avoid K deficiency in the plant.

2.6 Photosynthesis
Potassium impacts photosynthesis of the crop canopy via two mechanisms:
(1) solar radiation interception, and (2) photosynthesis per unit leaf area.
Collectively, these two phenomenon regulate the pool of photo assimilates
available for plant growth.
One of the more obvious consequences of plant growth under K defi-
cient conditions is a reduction in the plant stature (Cassman et al., 1989;
Ebelhar and Varsa, 2000; Heckman and Kamprath, 1992; Pettigrew and
Meredith, 1997). Accompanying this reduction in plant stature is an overall
reduction in LAI (Jordan-Meille and Pellerin, 2004; Kimbrough et al., 1971;
Pettigrew and Meredith, 1997) for the crop canopy. Reductions in both the
overall number of leaves produced and in the size of individual leaves lead to
this reduced overall LAI seen in K deficient conditions. Smaller size of the
individual leaves was related to a reduced leaf area expansion as observed
with soybean leaves (Huber, 1985) and maize (Jordan-Meille and Pellerin,
2004). This lower leaf area expansion under K deficient conditions is most
likely related to the role potassium plays in lowering the osmotic potential
and thereby raising the turgor pressure to drive cell expansion (Dhindsa
et al., 1975; Mengel and Arneke, 1982).
Not only is the sunlight intercepting leaf surface area diminished when
plants are grown under K deficient conditions, but the rate of photosynthesis
The Physiology of Potassium in Crop Production 213

per unit of that leaf surface area is also reduced (Bednarz et al., 1998; Huber,
1985; Longstreth and Nobel, 1980; Pier and Berkowitz, 1987; Wolf et al.,
1976). Potassium impacts photosynthesis through influences on both sto-
matal and nonstomatal aspects of photosynthesis. The role that K plays in
regulating stomatal aperture is well established and has previously been dis-
cussed in detail earlier in this chapter and therefore will not be dwelt within
this section.This stomatal aperture regulation controls the flow of CO2 into
and the flow of H2O vapor out of the intercellular spaces, thus affecting the
level of CO2 available at the reaction site for photosynthesis.
Nonstomatal factors can also be impacted by the potassium level to
regulate the photosynthetic rate. Much of the nonstomatal potassium effects
are tied into the role that potassium plays in photophosphorylation, rather
than the effect that it has on the enzymes involved in carbon assimila-
tion (Huber, 1985). Peoples and Koch (1979), however, reported reduced
Rubisco activities caused by potassium deficiencies. Huber (1985) then
countered and speculated that this response reported by Peoples and Koch
(1979) was more due to reduced enzyme synthesis rather than reduce activ-
ity from the individual enzymes. This reduced photophosphorylation seen
under K+ deficient conditions is related to an inner chloroplast membrane
ATPase that maintains a high stromal pH needed for the efficient conver-
sion from light energy to chemical energy by pumping protons out of the
stroma into the cytosol while allowing K+ flux into the stroma (Berkowitz
and Peters, 1993). An adequate potassium supply is critical for maintain-
ing optimal activity of this ATPase (Shingles and McCarty, 1994). In addi-
tion, the reduced translocation of carbon assimilates out of the chloroplast
(Ashley and Goodson, 1972; Mengel and Haeder, 1977; Mengel and Viro,
1974) could lead to feedback inhibition in the nonstomatal component of
the photosynthetic process (Pettigrew, 2008; Cakmak, 2005). There is also
evidence that plants not receiving adequate potassium levels can have an
increased production of reactive oxygen species (ROS) in the photosyn-
thetic tissue that can lead to photooxidative damage under higher light
intensities (Cakmak, 2005). This decreased efficiency in processing excited
electrons created by sunlight capture for K+ deficient plants is not surprising
considering the overall reduced photosynthesis and photoassimilate trans-
port exhibited by those plants.
Although both stomatal and nonstomatal photosynthetic factors can
be impacted by the level of available potassium, the timing and extent of
any potassium deficiency development can dictate which of these factors
plays the predominant role in regulating the photosynthetic production.
214 Derrick M. Oosterhuis et al.

For example, Bednarz et al. (1998) reported that during the early onset of a
developing potassium deficiency, stomatal conductance was principle com-
ponent regulating photosynthesis. However, as the potassium deficiency
became more pronounced and extreme, nonstomatal or biochemical factors
emerge as the overriding factors for the decreased photosynthesis.
The result of this combined potassium deficiency induced reduction in
overall LAI, solar radiation interception, and photosynthetic rate per unit
leaf area is the generation of a smaller pool of photosynthetic assimilates
available for growth. Ultimately, a smaller pool of photosynthetic assimilates
will reduce the yield levels that can be attained and compromise the quality
of the lint that is produced (Pettigrew, 2008).

2.7 Respiration
Similarly to photosynthesis, dark respiration has also been reported to be
affected by K levels since its function depends on the sum of nonstructural
carbohydrates and not on the previous day total assimilation (Cunningham
and Syvertsen, 1977). Under conditions of K deficiency dark respiration
rates were initially increased until the deficiency became severe, after which
dark respiration was suppressed (Okamoto, 1969). It was hypothesized that
respiration rates were increased due to enhanced mitochondrial activity, and
that was supported by Yeo et al. (1977) who in experiments on maize under
limited K supply observed significantly higher numbers of mitochondria
per cell in roots, stems, and leaves.

2.8  Assimilate Transport

Transpiration may influence translocation of carbon and nitrogen com-
pounds from production sites to sinks. Potassium has been reported to
control transpiration rates through its effect on stomatal function (Blatt,
1988) and consequently the root–shoot transport of mineral salts, nitrate,
and amino acids (Ben-Zioni et al., 1971; Marschner, 1995; Schobert et al.,
1998). The membrane potential of the sieve tube/companion cell com-
plex is controlled by K concentrations (Wright and Fischer, 1981). Since
apoplasmic phloem loading of sucrose in source leaves mediated by a
proton–sucrose cotransport requires a steep transmembrane pH gradient,
high potassium concentrations are needed for efficient phloem loading
of sucrose. Deeken et al. (2002) in experiments with Arabidopsis reported
that loss of the AKT2/3 potassium channel resulted in decreased phloem
loading of sucrose. In addition, K influences the rate of phloem loading
not only by promoting the efflux of assimilates into the apoplast prior to
The Physiology of Potassium in Crop Production 215

phloem loading (Mengel and Haeder, 1977; Doman and Geiger, 1979), but
also by regulating activation and function of invertase in the sink organs
(Oparka, 1990). Furthermore, K not only is essential for maintenance of
osmotic and pH gradients between the phloem and the parenchyma cells
within the sieve tubes that are required for phloem loading and transport
of assimilates (Marschner, 1995), but also provides the energy needed for
the transmembrane phloem re-loading processes (Gajdanowicz et al., 2011).
Carbohydrate translocation, therefore, is largely dependent on plant K levels
with many researchers reporting that lower than optimum K levels result in
accumulation of carbohydrates in several plant species (Haeder et al., 1973;
Mengel and Viro, 1974; Geiger and Conti, 1983; Cakmak et al., 1994a,b;
Amtmann et al., 2008; Amtmann and Armengaud, 2009), including cot-
ton (Bednarz and Oosterhuis, 1999; Pettigrew, 1999; Zhao et al., 2001). In
addition to the increased carbohydrate concentrations in the leaves of K
deficient cotton plants, Zhao et al. (2001) noticed that stem sucrose con-
centrations of K deficient plants were significantly lower compared to the
control, suggesting either an inhibition of sucrose entry in the transport
pool or a compromise in the phloem-loading mechanism. In support of
those observations, Ashley and Goodson (1972) observed that insufficient
K severely reduced the translocation 14C-labeled photosynthate. Transloca-
tion rates are also dependent on transpiration rates. However, Bednarz et al.
(1998) reported that K starvation increased transpiration rates while the
opposite was observed by Zhao et al. (2001) and Pervez et al. (2004). More
research needs to be focused on the effect of K supply on photosynthate
translocation and phloem loading in cotton. Furthermore, studies need to
be extended to include nitrogen compound translocation since similar inhi-
bitions and accumulations have been reported for amino acids and other
nitrogen compounds to occur under K deficiency in tobacco (Nicotiana
tabacum L.) (Koch and Mengel, 1974), as well as in rice (Oryza sativa L.),
soybean, and sunflower (Yamada et al., 2002).

2.9  Nitrate Transport–Potassium Interactions

Nitrogen exists in several ionic and nonionic forms in soil, however, the
two monovalent ionic forms, anion NO3− and cation NH4+ are the forms
mostly taken up by plants (Marschner, 1995). Research has revealed that
a positive relationship exists between N and K (Cai and Qin, 2006) and
increased K rates are required at higher N rates (Mondal, 1982). However,
nitrogen form, rate and timing of application have been reported to affect
K behavior in the soil as well as its uptake by the plants. NH4+ has similar
216 Derrick M. Oosterhuis et al.

charges and hydrated diameters with K+ (Wang and Wu, 2010) and it
has been reported that NH4+ inhibits the high-affinity transport system
which is functional primarily at low external K+ concentrations (<1 mM)
while the low-affinity transport system, which operates primarily at high
external K+ levels remains relatively unaffected (Nieves-Cordones et al.,
2007; Szczerba et al., 2009). On the other hand, NO3− is a univalent
anion that serves as a counterion to K+ (Lu and Li, 2005, Maathuis, 2009)
and it has been shown that NO3− uptake is more efficient with K+ as
counterion compared to Ca2+, Mg2+, or Na+ and similarly K+ uptake
has been reported to increase with nitrate fertilization (Pettersson and
­Jensen, 1983). K+ is also involved in the NO3+ transport in the xylem to
the shoot since after retranslocation of K-malate to the roots and sub-
sequent decarboxylation of the organic acids, HCO3− is exchanged for
NO3− (Ben-Zioni et al., 1971; Marschner et al., 1996). Lu and Li (2005)
observed that using NH4+-N as the only source of N caused a decrease
in K uptake compared to NO3−N, however, NH4+ resulted in more K
being translocated to the leaves than NO3−N in terms of the quantities
of xylem-transported K. K+ depletion of the nutrient solution has been
reported to increase NH4+ uptake while it suppressed NO3− absorption,
translocation, and assimilation and resulting in reduction in leaf nitrate
reductase activity (Ali et al., 1991). K+ deficiency was also reported to
result in downregulation of root nitrate transporters (Armengaud et al.,
2004), which was quickly reversed after resupply of K+. Apart from nitrate
reductase, asparaginase, another important enzyme in N metabolism was
shown to be controlled by K supply (Sodek et al., 1980) while pyruvate
kinase, a central integrator of C and N metabolism was suggested to be
the main target of K+ deficiency (Amtmann et al., 2008). Additionally,
K+ deficiency has been observed to result in amino acids, amide, and
polyamine accumulation in tissues (Evans and Sorger, 1966) indicating
that K+ plays a pivotal role in the transport of N compounds to the site of
protein synthesis and their further stabilization while a number of studies
has shown that K+ application enhances protein and amino acids content
in grain crops (Yang et al., 2004). Furthermore Koch and Mengel (1974)
reported that adequate supply of K was needed for optimum translo-
cation of amino acids and nitrate as well as incorporation of nitrogen
into proteins in tobacco. Reduced K+ uptake and accumulation has been
shown to occur under NH4+ toxicity conditions since NH4+ significantly
suppresses the high-affinity pathway system and especially KUP trans-
porters (Martinez-Cordero et al., 2004).
The Physiology of Potassium in Crop Production 217

2.10  Potassium Channels

Potassium is taken up against its concentration gradient through K+ trans-
porters and channels located in the plasma membrane of root cells (Ashley
et al., 2006) while two independent transport mechanisms are employed:
(1) a selective, high-affinity pathway that allows active K+ uptake at low
concentrations (<1 mM) and involves plasma membrane proton pumping
ATPase complexes, and (2) a passive, low-affinity, channel-mediated path-
way that mainly functions at high external concentrations (>1 mM) (Epstein
et al., 1963; Britto and Kronzucker, 2008). A highly mobile nutrient, not
only within short-distance transport, such as between individual cells and
neighboring tissues, but also within long distance transport, such as through
xylem and phloem, K is transported throughout the plant with the help of
three channel families, Shaker, tandem-pore (TPK), and inwardly rectifying
(Kir), and three transporter families, KUP/HAK/KT (K+/H+ symporters),
HKT (K+/H+ or K+/Na+ transporters), and CPA (cation/H+ antiport-
ers) (Cherel, 2004; Amtmann and Armengaud, 2009; Wang and Wu, 2010).
Shaker type channels are involved in K uptake, long distance K transport,
and K release, and they mainly occur in the plasma membrane. They have a
high selectivity for K and channel gating is activated in response to changes
in membrane potential with depolarization of membrane resulting in out-
ward rectifying K channels (K moving out of the cytosol) and hyperpolar-
ization of membrane resulting in inward rectifying K channels (K moving
into the cytosol). Additionally, Shaker type channels are the main K conduits
during cellular movement exemplified by stomatal function. Furthermore,
a main component of low-affinity K uptake pathway in the roots has been
identified as an inward rectifying K channel of the Shaker family (AKT1
for Arabidopsis) (Hirsch et al., 1998) and is mainly expressed in the plasma
membrane of root cortical and epidermal cells (Maathuis and Sanders,
1996). Two pore K (TPK) channels in contrast to Shaker channels are less
if any affected by changes in the membrane potential (Gobert et al., 2007).
They are involved in K homeostasis and they have been reported to regulate
membrane potential; however, they are not characterized in great detail.
KUP/HAK/KT (K+/H+ symporters) are capable of both low and high
affinity of K+ (Ahn et al., 2004; Rodriguez-Navarro and Rubio, 2006).
Their expression has been shown to increase under K starvation conditions
(Armengaud et al., 2004; Gierth et al., 2005). Their roles include high-
affinity K+ uptake at the root:soil boundary, intracellular distribution, tur-
gor driven growth (Rigas et al., 2001; Elumalai et al., 2002), and general K
homeostasis (Rodriguez-Navarro and Rubio, 2006).
218 Derrick M. Oosterhuis et al.

When external [K+] becomes increasingly low, K+ uptake needs to be

energized and this occurs through H+ coupled systems that have been shown
to operate in root plasma membranes.The coupling stoichiometry is 1:1 and
K+:H+ symports can drive 106 fold K accumulation (Maathuis and Sanders,
1996). KUP/HAK/KT:K+ uptake permeases, CHXs:cation-H+ exchang-
ers are encoding K+/H+ symporters (Britto and Kronzucker, 2008; Zhao
et al., 2008) and are found to catalyze K+ influx to cells at low apoplastic
[K+] (Zhao et al., 2008). Similarly to KUP/HAK/KT family of transporters,
HKT have been reported to be expressed in roots and they are involved in
efflux and influx of ions however, they are relatively permeable to K+.
K+ transporter and channel families have been extensively investigated
in Arabidopsis with less information existing on crops, such as barley (Hor-
deum vulgare L.), wheat, beans, rice, maize, tomato (Lycopersicon esculentum L.)
(Ashley et al., 2006; Szczerba et al., 2009).

3.  STRESS MITIGATION

3.1  Drought Stress
As mentioned above, K levels significantly affect the efficiency of plant photo-
synthetic machinery with lower than optimum levels resulting in significant
decreases in CO2 fixation due to its involvement in ribulose bisphosphate
carboxylase/oxygenase activation and stomatal function through turgor
regulation. Water deficit stress significantly reduces photosynthetic rates due
to stomatal limitations, decreases in leaf stomatal conductance that result in
decreased CO2 fixation rates (Lawlor and Cornic, 2002), as well as due to
nonstomatal or metabolic impairment where activities of photosynthetic
enzymes, such as ribulose bisphosphate carboxylase/oxygenase and ribulose
bisphosphate are disturbed (Gimenez et al., 1992; Medrano et al., 1997;
Tezara et al., 1999). Efficiency and function of adenosine triphosphate syn-
thase has also been reported to be reduced under conditions of water stress
(Younis et al., 1979; Tezara et al., 1999) resulting in disturbances of stro-
mal pH values (Berkowitz and Gibbs, 1983) and concomitant reductions
in energy production. Most importantly, however, due to the decreased
efficiency of the photosynthetic mechanism to utilize the incoming light
energy, production of ROS is increased (Lawlor and Cornic, 2002).
Water-stressed chloroplasts have been observed to suffer increased leak-
age of K resulting in further suppression of photosynthesis (Sen Gupta and
Berkowitz, 1987) while high K levels have been associated with maintenance
of optimum pH values in the chloroplast’s stroma and optimal function of
The Physiology of Potassium in Crop Production 219

photosynthetic mechanism (Pier and Berkowitz, 1987). Research with spin-

ach (Spinacia oleracea L.) and wheat indicated that water-stressed plants con-
taining higher than optimum quantities of K were able to maintain efficient
photosynthetic activity (Berkowitz and Whalen, 1985; Pier and Berkowitz,
1987) due to the alleviating effects of K on chloroplast stromal pH balance
and dehydration. Further investigation on wheat also reported that detri-
mental effects of water stress on photosynthetic activity were minimized
when K supply was sufficient (Sen Gupta et al., 1989) and similar results
were also obtained from mung beans (Vigna radiata L.) and cowpea (Vigna
unguiculata L.) (Sangakkara et al., 2000) leading researchers to suggest that
modification of K plant concentrations can maintain CO2 assimilation rates
by regulating stomatal function and balancing cell water relations (Cakmak
and Engels, 1999; Mengel et al., 2001). Research with cotton has indicated
the importance of K supply on the photosynthetic functions. However,
limited attention has been given to the combination of K and water stress.
Bar-Tsur and Rudich (1987) observed that K-deprived cotton plants were
able to survive successive water stress, nevertheless with significant growth
inhibition.

3.2  Cold Stress

Lower than optimum or freezing temperatures result in photooxida-
tive damage to plant chloroplasts and reduce their CO2 fixation capacity
since their membrane structure is significantly damaged (Marschner, 1995;
Thomashow, 1999). Increased ROS production has also been related to
cold or freezing temperatures (Prasad et al., 1994; Huner et al., 1998; Foyer
et al., 2002; Devi et al., 2012) due to the impaired photosynthetic electron
transport chain. In addition, leaf stomatal conductance rates were signifi-
cantly decreased (Ort, 2002) and similar reductions have also been observed
for ribulose bisphosphate carboxylase activity, while membrane fluidity was
greatly affected (Makino et al., 1994; Allen and Ort, 2001). Beringer and
Trolldenier (1978) indicated that a high cell K concentration has the ability
to enhance cold tolerance by decreasing the cell’s osmotic potential. Fur-
ther research revealed a positive correlation between K availability and cold
stress tolerance, with lower than optimum K concentrations escalating the
negative effects of cold stress (Kafkafi, 1990; Yermiyahu and Kafkafi, 1990).
In contrast, increased K levels enhanced plant defenses against cold stress
by acting as an osmolyte, lowering the freezing point of sap and prevent-
ing cell dehydration (Kafkafi, 1990; Kant and Kafkafi, 2002). In addition,
significant yield losses and extensive leaf damage due to cold temperatures
220 Derrick M. Oosterhuis et al.

were reported to occur under low K fertilization, while the effects were
alleviated once K supply was increased in a number of vegetable crops such
as potato (Solanum tuberosum L.) and tomato (Hakerlerler et al., 1997). In
experiments with maize, Farooq et al. (2008) observed that seed treatment
with KCl was able to enhance frost tolerance by increasing production of
antioxidant enzymes, such as catalase, ascorbate peroxidase, and superoxide
dismutase and similar results were reported by Devi et al. (2012) in ginseng
(Panax ginseng L.).

3.3  Salt Stress

Increased salt levels inhibit plant growth by inducing both osmotic and ionic
stresses (Shabala and Cuin, 2007) while also damaging the photosynthetic
machinery (Sudhir and Murthy, 2004; Chaves et al., 2009). High Na levels
in the soil solution significantly reduce K uptake by the plant, while in the
cytoplasm water is driven out of the cell vacuole resulting in decreased cell
turgor (Yeo et al., 1991; Zhu et al., 1997). High concentrations of Na cat-
ions compete in the soil with K cations substantially reducing their uptake
by the plants (Zhu, 2003). As a consequence, higher concentrations of Na
in the cytoplasm compete with K for uptake sites at the plasma membrane,
including both low-affinity and high-affinity transporters (Shabala et al.,
2006) and more Na crosses the plasma membrane resulting in a significant
membrane depolarization (Shabala et al., 2005). Under such circumstances
passive K uptake is impossible and an increased K leakage from the cell is
observed (Shabala and Cuin, 2007). Furthermore, in order for the cell to
retain metabolic functions through osmotic adjustment, significant quanti-
ties of ATP are used for de novo synthesis of compatible osmolytes synthe-
sis, making active K acquisition even more problematic.
Regarding CO2 fixation, photosynthetic capacity and quantum yield
of oxygen evolution have been reported to significantly decrease under
conditions of high Na concentrations (Sudhir and Murthy, 2004; Chaves
et al., 2009) resulting in detrimental increases of ROS production and fur-
ther chlorophyll and membrane damage (Kaya et al., 2001; Cakmak, 2005).
Higher K levels as well as increased capacity of plants to accumulate K have
been associated with increased salt resistance initially in Arabidopsis (Liu
and Zhu, 1997; Zhu et al., 1998) and later in wheat (Rascio et al., 2001;
Santa-Maria and Epstein, 2001), cucumber (Cucumis sativus L.), and pepper
(Piper nigrum L.) (Kaya et al., 2001) due to potassium’s ability to enhance
the plants’ antioxidative mechanism. In cotton, a relatively salt tolerant crop
with a threshold level of 7.7 dS m−1 (Maas, 1986), research has been mainly
The Physiology of Potassium in Crop Production 221

focused on the effects of partial replacement of K+ with Na+ (Cooper

et al., 1967; Joham and Amin, 1964) and their effect on cotton yield. Later
research indicated that substituting K supply with appropriate amounts of
Na could be beneficial for cotton yield (Chen, 1992; Zhang et al., 2006)
while ­Bednarz and Oosterhuis (1999) observed that a similar treatment
could delay K deficiency development.

3.4  Biotic Stress

Disease resistance is genetically controlled but it is mediated through physi-
ological and biochemical processes and is interrelated with the nutritional
status of the plant or pathogen. It is generally observed that highly resistant
plants are less affected by alterations in nutrition than plants tolerant to
disease, and highly susceptible plants may remain susceptible with nutri-
ent conditions that greatly increase the resistance of intermediate or tol-
erant plants (Huber and Arny, 1985). Potassium plays a significant role in
plant protection against biotic stresses since high concentrations of K have
been reported to alleviate detrimental effects of disease and pest infesta-
tions (Perrenoud, 1990; Prabhu et al., 2007). This has been attributed to the
regulation by K of primary metabolic plant functions. In more detail, high
levels of K in the plant promote the synthesis of high molecular weight
compounds, such as proteins, starch and cellulose while simultaneously sup-
pressing the formation of soluble sugars, organic acids, and amides, com-
pounds indispensable for feeding pathogens and insects (Marschner, 1995;
Amtmann et al., 2008). In addition, the accumulation of inhibitory amino
acids, phytoalexins, phenols and auxins is dependent on the level of K
(Perrenoud, 1990) while K deficiency results in inorganic N accumulation,
due to poor translocation, and phenols (with fungicidal properties) break
down (Kiraly, 1976). Exuded arginine that inhibits sporangial germination
of Phytophthora infestans on potato increases as the level of K increases (Alten
and Orth, 1941). Additionally, canopy discoloration due to K deficiency has
been observed to be more prone to attack by parasites, while cracks and
lesions on leaf surfaces that develop under low K supply provide additional
access (Krauss, 2001). In tomato, Kirali (1976) observed that higher K supply
successfully suppressed disease incidence, and similar results were reported
in soybean (Mondal et al., 2001) and wheat (Sweeney et al., 2000). K appli-
cation on cotton has been reported to decrease root rot (Phymatotrichum
omnivorum) (Tsai and Bird, 1975), wilt (Verticilium alboatrum) (Hafez et al.,
1975), as well as wilt and root rot caused by Fusarium oxysporum sp. (Prabhu
et al., 2007). Ramasami and Shanmugam (1976) reported that K application
222 Derrick M. Oosterhuis et al.

decreased seedling blight (Rhizoctonia solani) in cotton, however, the oppo-

site results were observed by Blair and Curl (1974). Miller (1969) reported
that increased K application rates decreased leaf blight (Cercospora gossypina–
Alternaria solani complex) while KNO3 significantly reduced the severity of
Alternaria leaf blight of cotton at the middle canopy level (Bhuiyan et al.,
2007). Furthermore, reductions from nematode (Meloidogyne incognita, Roty-
lenchulus reniformis) damage due to high K supply were also reported by
Oteifa and Elgindi (1976). Cotton cultivars exuding more carbohydrate
and K and less Mg and Ca have been reported to have less seedling
disease and establish higher plant populations than plants exuding less
­carbohydrate and K (Tsai and Bird, 1975).

3.5  Potassium and Stress Signaling

Apart from its major role in plant protection from a variety of abiotic
and biotic stresses, K has been suggested to be significantly involved in
plant stress responses (Ashley et al., 2006; Wang and Wu, 2010). Depolar-
ization of the membrane is observed in response to other fungal elici-
tors (Rossard et al., 2006) as well as to herbivory (Maffei et al., 2007).
Changes in the external K concentration have a great impact on the
membrane potential because the conductance of the plasma membrane
is larger for K than any other ion. A high external K concentration
depolarizes the membrane while a low external concentration hyperpo-
larizes the membrane (Amtmann et al., 2008). Potassium deficiency not
only induced K transporter upregulation (Gierth et al., 2005; Lee et al.,
2007), but additionally controlled stress-signaling mechanisms such as
ROS synthesis (Schachtman and Shin, 2007) as well as hormone syn-
thesis (Ashley et al., 2006). ROS production, and especially production
of hydrogen peroxide (H2O2), increased under conditions of limited K
supply (Shin and Schachtman, 2004) with H2O2 further activating K
uptake mechanisms in order to increase K plant status. In addition, genes
regulating jasmonic acid and auxin synthesis were upregulated at low
plant K concentrations (Armengaud et al., 2004) while ethylene pro-
duction rates were increased two-fold in K-starved Arabidopsis plants
compared to control plants (Shin and Schachtman, 2004). In cotton,
research indicated that K deficiency resulted in increased abscisic acid
concentrations in the roots and the xylem sap while cytokinins (zeatin
riboside and isopentenyl adenosine) levels were decreased in the xylem
sap and leaves (Wang et al., 2012) indicating that K affects the hormonal
balance in cotton.
The Physiology of Potassium in Crop Production 223

4.  SUMMARY
This review has described the fundamental role K plays in plant
growth and crop development. Its involvement in several physiological
functions, such as water relations, enzyme activation, stomatal regulation
and photosynthesis, assimilate and nitrate transport was summarized and
potassium deficiency was described and related to them. In addition, potas-
sium’s major role in plant health and tolerance to abiotic and biotic stress as
well as stress signaling was underlined. However, research has mainly focused
on Arabidopsis with little information existing on crop species. Considering
potassium’s major role in yield development and quality research should
focus on expanding its scope in major crop species hence, enable farmers to
achieve optimal utilization of K fertilization and additionally provide targets
for future genetic improvement efforts.

REFERENCES
Adams, D.O., Franke, K.E., Christensen, L.P., 1990. Elevated putrescine levels in grapevine
leaves that display symptoms of potassium deficiency. Am. J. Enol.Vitic. 41, 121–125.
Ahn, S.J., Shin, R., Schachtman, D.P., 2004. Expression of KT/KUP genes in Arabidopsis and
the role of root hairs in K+ uptake. Plant Physiol. 134, 1135–1145.
Akhtar, M.E., Khan, M.Z., Ahmad, S., Ashraf, M., Sardar, A., 2009. Effect of potassium on
micromorphological and chemical composition of three cotton (Gossypium hirsutum
L.) genotypes. Afr. J. Biotech. 8, 3511–3518.
Ali, A.A., Ikeda, M.,Yamada,Y., 1991. Effects of supply of K, Ca, and Mg on the absorption
and assimilation of ammonium- and nitrate- N in tomato plants. Soil. Sci. Plant Nutr.
37, 283–289.
Allen, D.J., Ort, D.R., 2001. Impact of chilling temperatures on photosynthesis in warm
climate plants. Trends Plant Sci. 6, 36–42.
Alten, F., Orth, H., 1941. Untersuchungen über den aminosäurengehalt und die anfälligkeit
der kartoffel gegen die kraut- und kollenfäule (Phytophthora infestans). Phytopathol. Z.
13, 243–247.
Amtmann, A., Troufflard, S., Armengaud, P., 2008. The effect of potassium nutrition on pest
and disease resistance in plants. Physiol. Plant 133, 682–691.
Amtmann, A., Armengaud, P., 2009. Effects of N, P, K, and S on metabolism: new knowledge
gained from multilevel analysis. Curr. Opin. Plant Biol. 12, 275–283.
Armengaud, P., Breitling, R.M., Amtmann, A., 2004. The potassium-dependent transcrip-
tome of Arabidopsis reveals a prominent role of jasmonic acid in nutrient signaling. Plant
Phys. 136, 2556–2576.
Armengaud, P., Sulpice, R., Miller, A.J., Stitt, M., Amtmann, A., Gibon, Y., 2009. Multilevel
analysis of primary metabolism provides new insights into the role of potassium nutrition
for glycolysis and nitrogen assimilation in Arabidopsis roots. Plant Phys. 150, 772–785.
Ashley, D.A., Goodson, R.D., 1972. Effect on time and plant potassium status on 14C-labled
photosynthate movement in cotton. Crop Sci. 12, 686–690.
Ashley, M.K., Grant, M., Grabov, A., 2006. Plant responses to potassium deficiencies: a role
for potassium transport proteins. J. Exp. Bot. 57, 425–436.
224 Derrick M. Oosterhuis et al.

Ashraf, M., Ashfaq, A., Ashraf, M.Y., 2002. Effects of increased supply of potassium on growth
and nutrient content in pearl millet under water stress. Biol. Plant 45, 141–144.
Bar-Tsur, A., Rudich, J., 1987. Osmotic adjustment of cotton to moderate potassium chlo-
ride stress and subsequent water stress during early stage of development. Agron. J. 79,
166–171.
Bednarz, C.W., Oosterhuis, D.M., Evans, R.D., 1998. Leaf photosynthesis and carbon isotope dis-
crimination of cotton in response to potassium deficiency. Environ. Exp. Bot. 39, 131–139.
Bednarz, C.W., Oosterhuis, D.M., 1999. Physiological changes associated with potassium
deficiency in cotton. J. Plant Nutr. 22, 303–313.
Begg, J.E., Turner, N.C., 1976. Crop water deficits. Adv. Agron. 28, 161–217.
Benlloch-Gonzalez, M., Arquero, O., Fournier, J.M., Barranco, D., Benlloch, M., 2008. K
starvation inhibits water-stress-induced stomatal closure. J. Plant Physiol. 165, 623–630.
Benlloch-González, M., Romera, J., Cristescu, S., Harren, F., Fournier, J.M., Benlloch, M.,
2010. K starvation inhibits water-stress-induced stomatal closure via ethylene synthesis
in sunflower plants. J. Exp. Bot. 61, 11–39.
Ben-Zioni, A., Vaadia, Y., Lips, S.H., 1971. Nitrate uptake by roots as regulated by nitrate
reduction products of the shoot. Physiol. Plant 24, 288–290.
Berg,W.K., Cunningham, S.M., Brouder, S.M., Joern, B.C., Johnson, K.D.,Volenec, J.J., 2009.
Influence of phosphorus and potassium on alfalfa yield, taproot C and N pools, and
transcript levels of key genes after defoliation. Crop Sci. 49, 974–982.
Berg,V.S., Heuchelin, S., 1990. Leaf orientation of soybean seedlings: l. effect of water poten-
tial and photosynthetic photon flux density on paraheliotropism. Crop Sci. 30, 631–638.
Beringer, H., 1978. Functions of potassium in plant metabolism with particular reference
to yield. In: Potassium in soils and crops. Intl. Potash Institute, New Delhi, India, pp.
185–202.
Beringer, H., Trolldenier, G., September 4–8, 1978. The influence of K nutrition on the
response of plants to environmental stress. In: “Potassium Research-Review and Trends”
Proceedings of 11th Congress of the International Potash Institute, Bern, Switzerland,
pp. 189–222.
Beringer, H., Koch, K., Lindhauer, M.G., 1986. Sucrose accumulation and osmotic potentials
in sugar beet at increasing levels of potassium nutrition. J. Sci. Food Agric. 37, 211–218.
Berkowitz, G.A., Gibbs, M., 1983. Reduced osmotic potential inhibition of photosynthesis.
Site-specific effects of osmotically induced stromal acidification. Plant Phys. 71, 905–911.
Berkowitz, G.A., Whallen, C., 1985. Leaf K+ interaction with water stress inhibition of non-
stomatal controlled photosynthesis. Plant Phys. 79, 189–193.
Berkowitz, G.A., Peters, J.S., 1993. Chloroplast inner envelope ATPase acts as a primary H+
pump. Plant Physiol. 102, 261–267.
Bhuiyan, S.A., Boyd, M.C., Dougall, A.J., Martin, C., Hearnden, M., 2007. Effects of foliar
application of potassium nitrate on suppression of Alternaria leaf blight of cotton (Gos-
sypium hirsutum) in northern Australia. Aust. Plant Pathol. 36, 462–465.
Blair,W.C., Curl, E.A., 1974. Influence of fertilization regimes on the rhizosphere microflora
and Rhizoctonia disease of cotton seedlings. Proc. Am. Phytopath. Soc. 1, 28–29.
Blatt, M.R., 1988. Potassium-dependent, bipolar of K+ channels in guard cells. J. Membr.
Biol. 102, 235–246.
Britto, D.T., Kronzucker, H.J., 2008. Cellular mechanisms of potassium transport in plants.
Physiol. Plant 133, 637–650.
Buch-Pedersen, M.J., Rudashevskaya, E.L., Berner, T.S., Venema, K., Palmgren, M.G., 2006.
Potassium as an intrinsic uncoupler of the plasma membrane H-ATPase. J. Biol. Chem.
281, 38285–38292.
Cai, Z.C., Qin, S.W., 2006. Dynamics of crop yields and soil organic carbon in a long-
term fertilization experiment in the Huang-Huai-Hai Plain of China. Geoderma 136,
708–715.
The Physiology of Potassium in Crop Production 225

Cakmak, I., Hengeler, C., Marschner, H., 1994a. Partitioning of shoot and root dry matter
and carbohydrates in bean plants suffering from Phosphorus, potassium, and magnesium
deficiency. J. Exp. Bot. 45, 1245–1251.
Cakmak, I., Hengeler, C., Marschner, H., 1994b. Changes in phloem export of sucrose in
leaves in response to phosphorus, potassium and magnesium deficiency. J. Exp. Bot. 45,
1251–1257.
Cakmak, I., Engels, C., 1999. Role of mineral nutrients in photosynthesis and yield forma-
tion. In: Rengel, Z. (Ed.), Mineral Nutrition of Crops: Mechanisms and Implications.
The Haworth Press, New York, pp. 141–168.
Cakmak, I., 2005. The role of potassium in alleviating detrimental effects of abiotic stresses
in plants. J. Plant Nutr. Soil Sci. 168, 521–530.
Carlson, J.G., 1973. Morphology. In: Caldwell, B.E. (Ed.), Soybeans: Improvement, Produc-
tion and Uses, pp. 17–95. Am. Soc. Agron., Madison, WI.
Cassman, K.G., Kerby, T.A., Roberts, B.A., Bryant, D.C., Brouder, S.M., 1989. Differential
response to two cotton cultivars to fertilizer and soil potassium. Agron. J. 81, 870–876.
Chaves, M.M., Flexas, J., Pinheiro, C., 2009. Photosynthesis under drought and salt stress:
regulation mechanisms from whole plant to cell. Ann. Bot. 103, 551–560.
Chen, G.A., 1992. Na effect on the growth of cotton and K and Na absorption. Soil 24, 201–204.
Chen, L., Nishizawa, T., Higashitani, A., Suge, H., Wakui,Y., Takeda, K., Takahashi, H., 2001.
A variety of wheat tolerant to deep‐seeding conditions: elongation of the first internode
depends on the response to gibberellin and potassium. Plant Cell Environ. 24, 469–476.
Cherel, I., 2004. Regulation of K+ channel activities in plants: from physiological to molecu-
lar aspects. J. Exp. Bot. 55, 337–351.
Claussen, M., Lüthe, H., Blatt, M., Böttger, M., 1997. Auxin-induced growth and its linkage
to potassium channels. Planta 201, 227–234.
Cocucci, M.C., Rosa, S.D., 1980. Effects of canavanine on IAA‐and fusicoccin‐stimulated
cell enlargement, proton extrusion and potassium uptake in maize coleoptiles. Physiol.
Plant 48, 239–242.
Conway,T.W., 1964. On the role of ammonium or potassium ion in amino acid polymeriza-
tion. Proc. Natl. Acad. Sci. 51, 1216–1220.
Cooper, R.B., Blaser, R.E., Brown, R.E., 1967. Potassium nutrition effects on net photosyn-
thesis and morphology of Alfalfa. Soil. Sci. Soc. Am. J. 31, 231–235.
Cunningham, G.L., Syvertsen, J.P., 1977. The effect of nonstructural carbohydrate levels on
dark CO2, release in creosotebush. Photosynthetica 11, 291–295.
Cutler, J.M., Rains, D.W., 1978. Effects of water stress and hardening on the internal water
relations and osmotic constituents of cotton leaves. Physiol. Plant 42, 261–268.
Deeken, R., Geiger, D., Fromm, J., Kokoreva, O., Ache, P., Langenfeld-Heyser, R., Sauer, N.,
May, T., Hedrich, R., 2002. Loss of the AKT2/3 potassium channel affects sugar loading
into the phloem of Arabidopsis. Planta 216, 334–344.
Devi, B.S.R., Kim, J.Y., Selvi, S.K., Lee, S., Yang, D.C., 2012. Influence of potassium nitrate
on antioxidant level and secondary metabolite genes under cold stress in Panax ginseng.
Rus. J. Plant Physiol. 59, 318–325.
Dhindsa, R.S., Beasley, C.A., Ting, I.P., 1975. Osmoregulation in cotton fiber: accumulation
of potassium and malate during growth. Plant Physiol. 56, 394–398.
Doman, D.C., Geiger, D.R., 1979. Effect of exogenously supplied foliar potassium on phloem
loading in Beta vulgaris L. Plant Physiol. 64, 528–533.
Ebelhar, S.A.,Varsa, E.C., 2000.Tillage and potassium placement effects on potassium utiliza-
tion by corn and soybean. Commun. Soil. Sci. Plant Anal. 31, 11–14.
Elumalai, R.P., Nagpal, P., Reed, J.W., 2002. A mutation in Arabidopsis KT2/KUP2 potassium
transporter gene affects shoot cell expansion. Plant Cell 14, 119–131.
Epstein, E., Rains, D., Elzam, O., 1963. Resolution of dual mechanisms of potassium absorp-
tion by barley roots. Proc. Natl. Acad. Sci. 49, 684–692.
226 Derrick M. Oosterhuis et al.

Evans, H.J., Sorger, G.J., 1966. Role of mineral elements with emphasis on the univalent
cations. Ann. Rev. Plant Phys. 17, 47–76.
Farooq, M., Aziz, T., Chemma, Z.A., Hussian, M., Khaliq, A., 2008. Activation of antioxidant
system by KCl improves the chilling tolerance in hybrid maize. J. Agron. Crop Sci. 194,
438–448.
Ford, C.W., Wilson, J.R., 1981. Changes in levels of solutes during osmotic adjustment to
water stress in leaves of four tropical pasture species. Aust. J. Plant Physiol. 8, 77–91.
Fournier, J.M., Roldan, A.M., Sanchez, C., Alexandre, G., Benlloch, M., 2005. K starvation
increases water uptake in whole sunflower plants. Plant Sci. 168, 823–829.
Foyer, C.H., Vanacker, H., Gomez, L.D., Harbinson, J., 2002. Regulation of photosynthesis
and antioxidant metabolism in maize leaves at optimal and chilling temperatures: review.
Plant Physiol. Biochem. 40, 659–668.
Gajdanowicz, P., Michard, E., Sandmann, M., Rocha, M., Dreyer, I., 2011. Potassium gradi-
ents serve as a mobile energy source in plant vascular tissues. Proc. Natl. Acad. Sci. 108,
864–869.
Geiger, D.R., Conti,T.R., 1983. Relation of increased potassium nutrition to photosynthesis
and translocation of carbon. Plant Physiol. 71, 141–144.
Gierth, M., Maser, P., Schroeder, J.I., 2005. The potassium transporter AtHAK5 functions in
K-deprivation induced high affinity K uptake and AKT1 K channel contribution to K
uptake kinetics in Arabidopsis roots. Plant Physiol. 137, 1105–1114.
Gimenez, C., Mitchell, V.J., Lawlor, D.W., 1992. Regulation of photosynthetic rate of two
sunflower hybrids under water stress. Plant Physiol. 98, 516–524.
Gobert, A., Isayenkov, S., Voelker, C., Czempinski, K., Maathuis, F.J.M., 2007. The two-
pore channel TPK1 gene encodes the vacuolar K+ conductance and plays a role in K+
homeostasis. Proc. Natl. Acad. Sci. 104, 10726–10731.
Gorton, H.L., 1987.Water relations in pulvini from Samanea saman. Plant Physiol. 83, 945–950.
Green, J.F., Muir, R.M., 1979. An analysis of the role of potassium in the growth effects of
cytokinin, light and abscisic acid on cotyledon expansion. Physiol. Plant 46, 19–24.
Green, J.F., Muir, R.M., 1978. The effect of potassium on cotyledon expansion induced by
cytokinins. Physiol. Plant 43, 213–218.
Green, J., 1983. The effect of potassium and calcium on cotyledon expansion and ethylene
evolution induced by cytokinins. Physiol. Plant 57, 57–61.
Haeder, H.E., Mengel, K., Forster, H., 1973. The effect of potassium on translocation of
photosynthates and yield patterns of potato plants. J. Sci. Food Agric. 24, 1479–1487.
Hafez, A.A.R., Stout, P.R., DeVay, J.E., 1975. Potassium uptake by cotton in relation to Ver-
ticillium wilt. Agron. J. 67, 359–361.
Hager, A., Menzel, H., Krauss, A., 1971. Versuche und hypothese zur primarwirkung des
auxins beim streckungswachstum. Planta 100, 47–75.
Hakerlerler, H., Oktay, M., Eryuce, N.,Yagmur, B., 1997. Effect of potassium sources on the
chilling tolerance of some vegetable seedlings grown in hotbeds. In: Johnston, A.E. (Ed.),
Food Security in the WANA Region, the Essential Need for Balanced Fertilization, pp.
317–327. Basel, Switzerland.
Hawker, J.S., Marschner, H., Downton, W.J.S., 1974. Effects of sodium and potassium on
starch synthesis in leaves. Funct. Plant Biol. 1, 491–501.
Hawker, J.S., Marschner, H., Krauss, A., 1979. Starch synthesis in developing potato tubers.
Physiol. Plant 46, 25–30.
Heckman, J.R., Kamprath, E.J., 1992. Potassium accumulation and corn yield related to
potassium fertilizer rate and placement. Soil. Sci. 56, 141–148.
Hepler, P.K., Vidali, L., Cheung, A.Y., 2001. Polarized cell growth in higher plants. Annu.
Rev. Cell. Dev. Biol. 17, 159–187.
Hirsch, R.E., Lewis, B.D., Spalding, E.P., Sussman, M.R., 1998. A role for the AKT1 potas-
sium channel in plant nutrition. Science 280, 918–921.
The Physiology of Potassium in Crop Production 227

Hsiao, T.C., Läuchli, A., 1986. A role for potassium in plant-water-relations. In: Tinker, B.,
Läuchli, A. (Eds.), Advances in Plant Nutrition, pp. 281–311(Praeger Scientific, New
York).
Hsiao, T.C., Acevedo, E., Fereres, E., Henderson, D., 1976. Water stress, growth, and osmotic
adjustment. Phil. Trans. R. Soc. Lon. B. Biol. Sci. 273, 479–500.
Huber, S.C., 1985. Role of potassium in photosynthesis and respiration. In: Munson, R.D.
(Ed.), Potassium in Agriculture. ASA, CSSA, SSSA, Madison, WI, pp. 369–391.
Huber, D.M., Arny, D.C., 1985. Interaction of potassium with plant disease. In: Munson, R.D.
(Ed.), Potassium in Agriculture. ASA, CSSA, SSSA, Madison, WI, pp. 467–488.
Huner, N.P.A., Oquist, G., Sarhan, F., 1998. Energy balance and acclimation to light and cold.
Trends Plant Sci. 3, 224–230.
Iannucci, A., Russo, M., Arena, L., Di Fonzo, N., Martiniello, P., 2002.Water deficit effects on
osmotic adjustment and solute accumulation in leaves of annual clovers. Eur. J. Agron.
16, 111–122.
Joham, H.E., Amin, J.V., 1964. Role of sodium in the potassium nutrition of cotton. Soil.
Sci. 99, 220–226.
Jones, M.M., Osmond, C.B., Turner, N.C., 1980. Accumulation of solutes in leaves of sor-
ghum and sunflower in response to water deficits. Aust. J. Plant Physiol. 7, 193–205.
Jordan-Meille, I., Pellerin, S., 2004. Leaf area establishment of a maize (Zea mays L.) field crop
under potassium deficiency. Plant Soil. 265, 75–92.
Jung, J.Y., Shin, R., Schachtman, D.P., 2009. Ethylene mediates response and tolerance to
potassium deprivation in Arabidopsis. Plant Cell 21, 607–621.
Kafkafi, U., 1990.The functions of plant K in overcoming environmental stress situations. In:
22nd Colloquium. International Potash Institute, Bern, Switzerland, pp. 81–93.
Kant, S., Kafkafi, U., 2002. Potassium and abiotic stresses in plants. In: Pasricha, N.S., Bansal, S.K.
(Eds.), Plant Roots:The Hidden Half. Marcel Dekker Incorporated, New York, pp. 435–449.
Kawashima, R., 1969. Studies on the leaf orientation-adjusting movement in soybean plants.
I.The leaf orientation-adjusting movement and light intensity on leaf surface. Proc. Crop
Sci. Soc. Jpn. 38, 718–729 (in Japanese with English summary).
Kaya, C., Kimak, H., Higgs, D., 2001. Enhancement of growth and normal growth param-
eters by foliar application of potassium and phosphorus in tomato cultivars grown at
high (NaCl) salinity. J. Plant Nutr. 24, 357–367.
Kayne, F.J., 1973. In: Boyer, P.D. (Ed.), The Enzymes, Vol. 8. Academic Press, New York, pp.
353–382.
Kimbrough, E.L., Blaser, R.E., Wolf, D.D., 1971. Potassium effects on regrowth of alfalfa
(Medicago sativa L.). Agron. J. 63, 836–839.
Kirali, Z., 1976. Plant disease resistance as influenced by biochemical effects of nutrients in
fertilizers. In: “Fertilizer Use and Plant Health. Proceedings of 12th College of Interna-
tional Potash Institute, pp. 33–46. Izmir, Turkey.
Koch, K., Mengel, K., 1974.The influence of the level of potassium supply to young tobacco
plants (Nicotiana tabacum L.) on short-term uptake and utilisation of nitrate nitrogen
(15N). Sci. Food Agric. 25, 165–471.
Koller, D., 2000. Plants in search of sunlight. Adv. Bot. Res. 33, 36–131.
Krauss, A., 2001. Potassium and biotic stress. In: Workshop on Potassium in Argentina’s Agri-
cultural Systems. Intl. Potash Institute, Buenos Aires, Argentina, pp. 281–293.
Kusano, T., Berberich, T., Tateda, C., Takahashi, Y., 2008. Polyamines: essential factors for
growth and survival. Planta 228, 367–381.
La Guardia, M.D., Benlloch, M., 1980. Effects of potassium and gibberellic acid on stem
growth of whole sunflower plants. Physiol. Plant 49, 443–448.
Läuchli, A., Pflüger, R., 1978. Potassium transport through plant cell membranes and meta-
bolic role of potassium in plants. In: Potasium Research Review and Trends. Intl. Potash
Institute, Bern, Switzerland, pp. 111–164.
228 Derrick M. Oosterhuis et al.

Lawlor, M.M., Cornic, G., 2002. Photosynthetic carbon assimilation and associated metabo-
lism in relation to water deficits in higher plants. Plant Cell Environ. 25, 275–294.
Lee, S.C., Lan, W.Z., Kim, B.G., Li, L., Cheong,Y.H., Pandey, G.K., Buchanan, B.B., Luan, S.,
2007. A protein phosphorylation/dephosphorylation network regulates a plant potas-
sium channel. Proc. Natl. Acad. Sci. 104, 15959–15964.
Leigh, R.A.,Wyn Jones, R.G., 1984. A hypothesis relating critical potassium concentrations for
growth to the distribution and functions of this ion in the plant cell. New. Phytol. 97, 1–13.
Li, R.,Volenec, J.J., Joern, B.C., Cunningham, S.M., 1997. Potassium and nitrogen effects on
carbohydrate and protein metabolism in alfalfa roots. J. Plant Nutr. 20, 511–529.
Liu, J., Zhu, J.K., 1997. Proline accumulation and salt stress-induced gene expression in a
salt-hypersensitive mutation of Arabidopsis. Plant Physiol. 114, 591–596.
Longstreth, D.J., Nobel, P.S., 1980. Nutrient influences on leaf photosynthesis: effects of nitro-
gen, phosphorus, and potassium for Gossypium hirsutum L. Plant Physiol. 65, 541–543.
Lu, Y.X., Li, C.J., 2005. Transpiration, potassium uptake and flow in tobacco as affected by
nitrogen forms and nutrient levels. Ann. Bot. 95, 991–998.
Maas, E.V., 1986. Salt tolerance of plants. Appl. Agric. Res. 1, 12–26.
Maathuis, F.J.M., Sanders, D., 1996. Mechanisms of potassium absorption by higher plant
roots. Physiol. Plant 96, 158–168.
Maathuis, F.J.M., 2009. Physiological functions of mineral macronutrients. Curr. Opin. Plant
Biol. 12, 250–258.
MacRobbie, E.A.C., 1977. Functions of ion transport in plant cells and tissues. Int. Rev.
Biochem. 13, 211–247.
Maene, L.M., 2001. Global Potassium Fertilizer Situation, Current Use and Perspectives. 3–5
December. International Potash Institute, New Delhi, India, pp. 1–10.
Maffei, M.E., Mithofer, A., Boland, W., 2007. Insects feeding on plants: rapid signals and
responses preceding the induction of phytochemical release. Phytochem 68, 2946–2959.
Makino, A., Nakano, H., Mae, T., 1994. Effects of growth temperature on the responses of
ribulose-1,5-bisphosphate carboxylase, electron transport components, and sucrose syn-
thesis enzymes to leaf nitrogen in rice and their relationships to photosynthesis. Plant
Physiol. 105, 1231–1238.
Marschner, H., 1995. Functions and mineral nutrients. In: Mineral Nutrition of Higher
Plants. Academic press, London, UK, pp. 213–255.
Marshner, H., Kirkby, E.A., Cakmak, I., 1996. Effect of mineral nutritional status on shoot-root
partitioning of photoassimilates and cycling of mineral nutrients. J. Exp. Bot. 47, 1255–1263.
Martinez-Cordero, M.A., Vicente, M., Francisco, R., 2004. Cloning and functional char-
acterization of the high-affinity K+ transporter HAK1 of pepper. Plant Mol. Biol. 56,
413–421.
Matoh, T., Yasuoka, S., Ishikawa, T., Takahashi, E., 1988. Potassium requirement of pyruvate
kinase extracted from leaves of halophytes. Physiol. Plant 74, 675–678.
Medrano, H., Parry, M.A., Socias, X., Lawlor, D.W., 1997. Long term water stress inactivates
rubisco in subterranean clover. Ann. Appl. Biol. 131, 491–501.
Memon, A.R., Siddiqi, M.Y., Class, A.D.M., 1985. Efficiency of K utilization by barley vari-
eties: activation of pyruvate kinase. J. Exp. Bot. 36, 79–90.
Mengel, K.,Viro, M., 1974. Effects on potassium supply on the transport of photosynthates to
the fruits of tomatoes (Lycopersicon esculentum). Plant Physiol. 30, 295–300.
Mengel, K., Haeder, H.E., 1977. Effect of potassium supply on the rate phloem sap exuda-
tion and the composition of phloem sap of Ricinus communis. Plant Physiol. 59, 282–284.
Mengel, K., Arneke, W.W., 1982. Effect of potassium on the water potential, the pressure
potential, the osmotic potential and cell elongation in leaves of Phaseolus vulgaris. Physiol.
Plant 54, 402–408.
Mengel, K., Kirkby, E.A., Kosegarten, H., Appel,T., 2001. Principles of Plant Nutrition. Klu-
wer Academic Publishers, Dordrecht, the Netherlands, pp. 481–511.
The Physiology of Potassium in Crop Production 229

Miller, J.W., 1969. The effect of soil moisture and plant nutrition on the Cercospora-Alternaria
leaf blight complex of cotton in Missouri. Phytopathology 59, 767–769.
Miller, C.S., 1975. Short interval leaf movements of cotton. Plant Physiol. 55, 562–566.
Moinuddin, Imas, P., 2007. Evaluation of potassium compared to other osmolytes in relation
to osmotic adjustment and drought tolerance of chickpea under water deficit environ-
ments. J. Plant Nutr. 30, 517–535.
Mondal, S.S., 1982. Potassium nutrition at high levels of N fertilization on rice. In: Potash
Review. Intl. Potash Institute, Bern, Switzerland, pp. 1–4.
Mondal, S.S., Pramanik, C.K., Das, J., 2001. Effect of nitrogen and potassium on oil yield,
nutrient uptake and soil fertility in soybean (Glycine max) – sesame (Sesamum indicum)
intercropping system. Indian J. Agric. Sci. 71, 44–46.
Morgan, J.M., 1984. Osmoregulation and water stress in higher plants. Ann. Rev. Plant Phys.
35, 299–319.
Morgan, J.M., 1992. Osmotic components and properties associated with genotypic differ-
ences in osmoregulation in wheat. Funct. Plant Biol. 19, 67–76.
Moshelion, M., Becker, D., Biela, A., Uehlein, N., Hedrich, R., Otto, B., Levi, H., Moran,
N., Kaldenhoff, R., 2002. Plasma membrane aquaporins in the motor cells of Samanea
saman: diurnal and circadian regulation. Plant Cell 14, 727–739.
Mullins, G.L., Burmester, C.H., Reeves, D.W., 1997. Cotton response to in-row subsoiling
and potassium fertilizer placement in Alabama. Soil. Till. Res. 40, 145–154.
Nieves-Cordones, M., Martínez-Cordero, M.A., Martínez, V., Rubio, F., 2007. An NH4+-
sensitive component dominates high-affinity K+ uptake in tomato plants. Plant Sci. 172,
273–280.
Nishizawa, T., Chen, L., Higashitani, A., Takahashi, H., Takeda, K., Suge, H., 2002. Responses
of the first internodes of hong mang mai wheat to ethylene, gibberellins and potassium.
Plant Prod. Sci. 5, 93–100.
Nitsos, R.E., Evans, H.J., 1969. Effects of univalent cations on the activity of particulate
starch synthetase. Plant Physiol. 44, 1260–1266.
Okamoto, S., 1969. The respiration in leaf discs from young taro plants under a moderate
potassium deficiency. Soil. Sci. Plant Nutr. 15, 274–279.
Oosterhuis, D.M., Walker, S., Eastham, J., 1985. Soybean leaflet movements as an indicator of
crop water stress. Crop Sci. 25, 1101–1106.
Oosterhuis, D.M.,Walker, S., 1984. Changes in the soybean leaflet pulvinule during inversion
with water stress. Plant Phys. Suppl. 75, 176.
Oosterhuis, D.M., Wullschleger, S.D., 1987. Osmotic adjustment in cotton (Gossypium hirsu-
tum L.) leaves and roots in response to water stress. Plant Physiol. 84, 1154–1157.
Oparka, K.J., 1990. What is phloem unloading? Plant Physiol. 94, 393–396.
Ort, D.R., 2002. Chilling-induced limitations on photosynthesis in warm climate plants:
contrasting mechanisms. Environ. Control Biol. 40, 7–18.
Oteifa, B.A., Elgindi, A.Y., 1976. Potassium nutrition of cotton (Gossypium barbadense, L) in
relation to nematode infection by Meloidogyne incognita and Rotylenchulus reniformis. In:
“Fertilizer Use and Plant Health”. Proceedings of 12th College of International Potash
Institute, pp. 301–306. Izmir, Turkey.
Peoples, T.R., Koch, D.W., 1979. Role of potassium in carbon dioxide assimilation in Medi-
cago sativa L. Plant Physiol. 63, 878–881.
Perrenoud, S., 1990. Potassium and plant health. In: IPI-Research Topics No. 3. Intl. Potash
Institute, Basel, Switzerland, p. 365.
Pervez, H., Ashraf, M., Makhdum, M.I., 2004. Influence of potassium nutrition on gas
exchange characteristics and water relations in cotton (Gossypium hirsutum L.). Photo-
synthetica 42, 251–255.
Pettersson, S., Jensen, P., 1983. Variations among species and varieties in uptake and utiliza-
tion of potassium. Plant Soil. 72, 231–237.
230 Derrick M. Oosterhuis et al.

Pettigrew, W.T., 2003. Relationship between insufficient potassium and crop maturity in
cotton. Agron. J. 95, 1323–1329.
Pettigrew,W.T., 2008. Potassium influences on yield and quality production for maize, wheat,
soybean and cotton. Physiol. Plant 133, 670–681.
Pettigrew, W.T., 1999. Potassium deficiency increases specific leaf weights and leaf glucose
levels in field-grown cotton. Agron. J. 91, 962–968.
Pettigrew, W.T., Meredith, W.R., 1997. Dry matter production, nutrient uptake, and growth
of cotton as affected by potassium fertilization. J. Plant Nutr. 20, 531–548.
Pier, P.A., Berkowitz, G.A., 1987. Modulation of water stress effects on photosynthesis by
altered leaf K+. Plant Physiol. 85, 655–661.
Prabhu, A.S., Fageria, N.K., Huber, D.M., Rodriguez, F.A., 2007. Potassium and plant disease.
In: Datnoff, L.E., Elmer,W.H., Huber, D.M. (Eds.), Mineral Nutrition and Plant Disease.
The American Phytopathological Soc. Press, Saint Paul, MN, pp. 57–78.
Prasad, T.K., Anderson, M.D., Martin, B.A., Stewart, C.R., 1994. Evidence of chilling-
induced oxidative stress in maize seedlings and a regulatory role for hydrogen peroxide.
Plant Cell 6, 65–74.
Ramasami, R., Shanmugam, N., 1976. Effect of nutrients on the incidence of Rhizoctonia
seedling disease of cotton. Indian Phytopathol. 29, 465–466.
Rascio, A., Russo, M., Mancuzzo, L., Platani, C., Nicastro, G., Di Fonzo, N., 2001. Enhanced
osmotolerance of a wheat mutant selected for potassium accumulation. Plant Sci. 160,
441–448.
Rayle, D.L., Cleland, R.E., 1992. The acid growth theory of auxin-induced cell elongation
is alive and well. Plant Physiol. 99, 1271–1274.
Reddy, K.R., Zhao, D., 2005. Interactive effects of elevated CO2 and potassium deficiency
on photosynthesis, growth, and biomass partitioning of cotton. Field Crops Res. 94,
201–213.
Rengel, Z., Damon, P.M., 2008. Crops and genotypes differ in efficiency of uptake and use.
Physiol. Plant 133, 624–636.
Rigas, S., Debrosses, G., Haralampidis, K., Vicente-Agullo, F., Feldmann, K.A., Grabov, A.,
Dolan, L., Hatzopoulos, P., 2001.TRH1 encodes a potassium transporter required for tip
growth in Arabidopsis root hairs. Plant Cell 13, 139–151.
Roberts, S.K., Snowman, B.N., 1999. The effects of ABA on channel-mediated K+ transport
across higher plant roots. J. Exp. Bot. 51, 1585–1594.
Rodriguez-Navarro, A., Rubio, F., 2006. High-affinity potassium and sodium transport sys-
tems in plants. J. Exp. Bot. 57, 1149–1160.
Romheld,V., Kirkby, E.A., 2010. Research in potassium in agriculture: needs and prospects.
Plant Soil. 335, 155–180.
Rossard, S., Luini, E., Perault, J.M., Bonmont, J., Roblin, G., 2006. Early changes in mem-
brane permeability, production of oxidative burst and modification of PAL activity
induced by ergosterol in cotyledons of Mimosa pudica. J. Exp. Bot. 57, 1245–1252.
Ruan, Y.L., Llewellyn, D.J., Furbank, R.T., 2001. The control of single-celled cotton fiber
elongation by developmentally reversible gating of plasmodesmata and coordinated
expression of sucrose and K transporters and expansin. Plant Cell 13, 47–60.
Sangakkara, U.R., Frehner, M., Nosberger, J., 2000. Effect of soil moisture and potassium fer-
tilizer on shoot water potential, photosynthesis and partitioning of carbon in mungbean
and cowpea. J. Agron. Crop Sci. 185, 201–207.
Santa-Maria, G.E., Epstein, E., 2001. Potassium/sodium selectivity in wheat and the amphi-
ploid cross wheat × Lophopyrum elongatum. Plant Sci. 160, 523–534.
Sarjala, T., Kaunisto, S., 1993. Needle polyamine concentrations and potassium nutrition in
scots pine. Tree Phys. 13, 87–96.
Satter, R.L., Galston, A.W., 1973. Leaf movements: Rosetta stone of plant behavior. Biosci-
ence 23, 407–416.
The Physiology of Potassium in Crop Production 231

Satter, R.L., Galston, A.W., 1981. Mechanisms of control of leaf movements. Ann. Rev. Plant
Phys. 32, 83–110.
Schobert, C., Baker, L., Komor, E., Hayashi, H., Chino, M., Lucas,W.J., 1998. Identification of
immunologically related proteins in sieve-tube exudate collected from monocotyledon-
ous and dicotyledonous plants. Planta 206, 245–252.
Schrempf, M., Satter, R.L., Galston, A.W., 1976. Potassium-linked chloride fluxes during
rhythmic leaf movement of Albizia Julibrissin. Plant Physiol. 58, 190–192.
Sen Gupta, A., Berkowitz, G.A., 1987. Osmotic adjustment, symplast volume, and nonsto-
matally mediated water stress inhibition of photosynthesis in wheat. Plant Physiol. 85,
1040–1047.
Sen Gupta, A., Berkowitz, G.A., Pier, P.A., 1989. Maintenance of photosynthesis at low leaf
water potential in wheat. Role of potassium status and irrigation history. Plant Physiol.
89, 1358–1365.
Shabala, L., Cuin, T.A., Newman, I.A., Shabala, S., 2005. Salinity induced ion flux patterns
from the excised roots of Arabidopsis SOS mutants. Planta 222, 1041–1050.
Shabala, S., Demidichic, V., Shabala, L., Cuin, T.A., Smith, S.J., Miller, A.J., Davies, J.M.,
Newman, I.A., 2006. Extracellular Ca2+ ameliorates NaCl induced loss from Arabidopsis
root and leaf cells by controlling plasma membrane K permeable channels. Plant Physiol.
141, 1653–1665.
Shabala, S., Cuin, T.A., 2007. Potassium transport and plant salt tolerance. Physiol. Plant 133,
651–669.
Schachtman, D., Shin, R., 2007. Nutrient sensing and signaling: NPKS. Annu. Rev. Plant
Biol. 58, 47–69.
Sharp, R.E., Hsiao, T.E., Silk, W.K., 1990. Growth of the maize primary root at low water
potentials. II. Role of growth and deposition of hexose and potassium in osmotic adjust-
ment. Plant Physiol. 93, 1337–1346.
Shin, R., Schachtman, D.P., 2004. Hydrogen peroxide mediates plant root cell response to
nutrient deprivation. Proc. Natl. Acad. Sci. 101, 8827–8832.
Shingles, R., McCarty, R.E., 1994. Direct measurement of ATP-dependent proton concen-
tration changes and characterization of a K-stimulated ATPase in pea chloroplast inner
envelope vesicles. Plant Physiol. 106, 731–737.
Sodek, L., Lea, P.J., Miflin, D.J., 1980. Distribution and properties of a potassium-dependent
asparaginase isolated from developing seeds of Pisum sativum and other plants. Plant
Physiol. 65, 22–26.
Stark, C., 1991. Osmotic adjustment and growth of Salt‐stressed cotton as improved by a
bioregulator. J. Agron. Crop Sci. 167, 326–334.
Sudhir, P., Murthy, S.D.S., 2004. Effects of salt stress on basic processes of photosynthesis.
Photosynthetica 42, 481–486.
Suelter, C.H., 1970. Enzymes activated by monovalent cations. Science 168, 789–795.
Sweeney, D.W., Granade, G.V., Eversmeyer, M.G., Whitney, D.A., 2000. Phosphorus, potas-
sium, chloride, and fungicide effects on wheat yield and leaf rust severity. J. Plant Nutr.
23, 1267–1281.
Szczerba, M.W., Britto, D.T., Kronzucker, H.J., 2009. K+ transport in plants: physiology and
molecular biology. J. Plant Physiol. 166, 447–466.
Taiz, L., Zeiger, E., 2010. In: Plant Physiology. Sinauer Associates Inc., Sunderland, MA.
Tanaka, Y., Sano, T., Tamaoki, M., Nakajima, N., Kondo, N., Hasezawa, S., 2005. Ethyl-
ene inhibits abscisic acid-induced stomatal closure in Arabidopsis. Plant Physiol. 138,
2337–2343.
Tezara, W., Mitchell,V.J., Driscoll, S.D., Lawlor, D.W., 1999. Water stress inhibits plant photo-
synthesis by decreasing coupling factor and ATP. Nature 401, 914–917.
Thiel, G.,Weise, R., 1999. Auxin augments conductance of K inward rectifier in maize cole-
optile protoplasts. Planta 208, 38–45.
232 Derrick M. Oosterhuis et al.

Thomashow, M.F., 1999. Plant cold acclimation: freezing tolerance genes and regulatory
mechanisms. Annu. Rev. Physiol. Plant Mol. Biol. 50, 571–599.
Tsai, H.Y., Bird, L.S., 1975. Microbiology of host pathogen interactions for resistance to
seedling disease and multiadversity resistance in cotton. In: Proceedings of the Beltwide
Cotton Production Research Conference, pp. 39–45. Memphis, TN.
Turner, N.C., Begg, J.E.,Tonnet, M.L., 1978. Osmotic adjustment of sorghum and sunflower
crops in response to water deficits and its influence on the water potential at which
stomata close. Funct. Plant Biol. 5, 597–608.
Uehlein, N., Kaldenhoff, R., 2008. Aquaporins and plant leaf movements. Ann. Bot. 101, 1–4.
Walker, D.J., Leigh, R.A., Miller, A.J., 1996. Potassium homeostasis in vacuolate plant cells.
Proc. Natl. Acad. Sci. 93, 10510–10514.
Wang, Y., Wu, W.H., 2010. Plant sensing and signaling in response to K+ deficiency. Mol.
Plant 3, 280–287.
Wang,Y., Du, M., Eneji, E., Wang, B., Tian, X., 2012. Mechanism of phytohormone involve-
ment in feedback regulation of cotton leaf senescence induced by potassium deficiency.
J. Exp. Bot. 63, 5887–5901.
Ward, G.M., 1960. Potassium in plant metabolism. III. Some carbohydrates changes in the wheat
seedling associated with varying rates of potassium supply. Can. J. Plant Sci. 40, 729–735.
Watanabe, H., Takahashi, K., 1997. Effects of abscisic acid, fusicoccin, and potassium on
growth and morphogenesis of leaves and internodes in dark-grown rice seedlings. Plant
Growth Regul. 109, 109–114.
Weimberg, R., Lerner, H.R., Poljakoff‐Mayber, A., 1982. A relationship between potassium
and proline accumulation in salt‐stressed Sorghum bicolor. Physiol. Plant 55, 5–10.
Wolf, D.D., Kimbrough, E.L., Blaser, R.E., 1976. Photosynthetic efficiency in alfalfa with
increasing potassium nutrition. Crop Sci. 16, 292–294.
Wright, J.P., Fisher, D.B., 1981. Measurement of the sieve tube membrane potential. Plant
Physiol. 67, 845–848.
Wyn Jones, R.G., Brady, C.J., Speirs, J., 1979. Ionic and osmotic relations in plant cells. In:
Laidman, D.L.,Wyn Jones, R.G. (Eds.), Recent Advances in the Biochemistry of Cereals.
Academic Press, London, UK, pp. 63–103.
Wyn Jones, R.J., Pollard, A., 1983. Proteins, enzymes and inorganic ions. In: Lauchli, A.,
Pirson, A. (Eds.), Encyclopedia of Plant Physiology. Springer, Berlin, pp. 528–562.
Yamada, S., Osaki, M., Shimano,T.,Yamada, M., Ito, M., Permana, A.T., 2002. Effect of potas-
sium nutrition on current photosynthesized carbon distribution to carbon and nitrogen
compounds among rice, soybean, and sunflower. J. Plant Nutr. 25, 1957–1973.
Yang, X.E., Liu, J.X., Li, H., Luo, A.C., 2004. Potassium internal use efficiency, relative to
growth vigor, potassium distribution and carbohydrate allocation in rice genotypes.
J. Plant Nutr. 27, 837–852.
Yeo, A.R., Kramer, D., Lauchli, A., Gullasch, J., 1977. Ion distribution in salt-stressed mature
Zea mays roots in relation to ultra-structure and retention of sodium. J. Exp. Bot. 28, 17–29.
Yeo, A.R., Lee, K.S., Izard, P., Boursier, P.J., Flowers, T.J., 1991. Short-term and long-term
effects of salinity on leaf growth in rice (Oryza sativa L.). J. Exp. Bot. 42, 881–889.
Yermiyahu, U., Kafkafi, U., 1990. Yield increase and stem brittle decrease in response to
increasing concentrations of potassium and NO3−/NH4+ in White Carnation CV. Stand.
Hassadeh 90, 742–746.
Younis, H.M., Boyer, J.S., Govindjee, 1979. Conformation and activity of chloroplast factor
exposed to low chemical potential of water in cells. Biochim. Biophys. Acta 548, 328–340.
Zhang,Y., Li, Q., Zhou, X., Zhai, C., Li, R., 2006. Effects of partial replacement of potassium
by sodium on cotton seedlings development and yield. J. Plant Nutr. 29, 1845–1854.
Zhang, Z.,Yang, F., Tian, X., 2009. Coronatine-induced lateral formation in cotton (Gossy-
pium hirsutum) seedlings under potassium-sufficient and –deficient conditions in rela-
tion to auxin. J. Plant Nutr. Soil Sci. 172, 435–444.
The Physiology of Potassium in Crop Production 233

Zhao, D., Oosterhuis, D.M., Bednarz, C.W., 2001. Influence of potassium deficiency on
photosynthesis, chlorophyll content, and chloroplast ultrastructure of cotton plants. Pho-
tosynthetica 39, 103–109.
Zhao, J., Cheng, N.H., Motes, C.M., Blancaflor, E.B., Moore, M., Gonzales, N., Padmanaban,
S., Sze, H., Ward, J.M., Hirschi, K.D., 2008. AtCHX13 is a plasma membrane K+ trans-
porter. Plant Physiol. 148, 796–807.
Zhenming, N., Xuefeng, X., Yi, W., Tianzhong, L., Jin, K., Zhenhai, H., 2008. Effects of
leaf-applied potassium, gibberellin and source-sink ratio on potassium absorption and
distribution in grape fruits. Sci. Hort. 115, 164–167.
Zhu, J., Hasegawa, P.M., Bressan, R.A., 1997. Molecular aspects of osmotic stress in plants.
Crit. Rev. Plant Sci. 16, 253–277.
Zhu, J.K., Liu, J., Xiong, L., 1998. Genetic analysis of salt tolerance in Arabidopsis thaliana:
evidence of a critical role for potassium nutrition. Plant Cell 10, 1181–1192.
Zhu, J.K., 2003. Regulation of ion homeostasis under salt stress. Curr. Opin. Plant Biol. 6,
441–445.