The Crop Journal 9 (2021) 249–256

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The Crop Journal

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

Synergistic and antagonistic interactions between potassium and

magnesium in higher plants
Kailiu Xie a,b, Ismail Cakmak c, Shiyu Wang a, Fusuo Zhang d,e, Shiwei Guo a,d,⇑
a
Jiangsu Provincial Key Lab of Solid Organic Waste Utilization, Jiangsu Collaborative Innovation Center of Solid Organic Wastes, Educational Ministry Engineering Center of
Resource-saving Fertilizers, Nanjing Agricultural University, Nanjing 210095, Jiangsu, China
b
School of Land Resources and Environment, Jiangxi Agricultural University, Nanchang 330045, Jiangxi, China
c
Faculty of Engineering and Natural Sciences, Sabanci University, 34956 Istanbul, Turkey
d
International Magnesium Institute, College of Resources and Environment, Fujian Agriculture and Forestry University, Fuzhou 350002, Fujian, China
e
College of Resources and Environmental Sciences, China Agricultural University, Beijing 100193, China

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

Article history: Magnesium (Mg) affects various critical physiological and biochemical processes in higher plants, and its
Received 11 March 2020 deficiency impedes plant growth and development. Although potassium (K)-induced Mg deficiency in
Revised 27 September 2020 agricultural production is widespread, the specific relationship of K with Mg and especially its compet-
Accepted 3 November 2020
itive nature is poorly understood. This review summarizes current knowledge on the interactions
Available online 25 November 2020
between K and Mg with respect to their root uptake, root-to-shoot translocation and distribution in
plants. Their synergistic effects on certain physiological functions are also described. The antagonistic
Keywords:
effect of K on Mg is stronger than that of Mg on K in root absorption and transport within plants, indi-
Antagonism
Functional replacement
cating that the balanced use of K and Mg fertilizers is necessary for sustaining high plant-available Mg
Magnesium and alleviating K-induced Mg deficiency, especially in plant species with high K demand or in high-
Potassium available-K soil. The relationship between Mg and K in plant tissues may be antagonistic or synergistic
Synergy depending on plant species, cell type, leaf age, source- and sink organs. There are synergistic effects of
K and Mg on photosynthesis, carbohydrate transport and allocation, nitrogen metabolism, and turgor reg-
ulation. Definition of optimal K/Mg ratios for soils and plant tissues is desirable for maintaining proper
nutritional status in plants, leading to a physiological state supporting crop production. Future research
should concentrate on identifying the physiological and molecular mechanisms underlying the interac-
tions between K and Mg in a given physiological function.
Ó 2020 Crop Science Society of China and Institute of Crop Science, CAAS. Production and hosting by
Elsevier B.V. on behalf of KeAi Communications Co., Ltd. This is an open access article under the CC BY-NC-
ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).

Contents

1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ........... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 250

2. Interaction between Mg and K during root uptake . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ........... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 251
3. Interaction between Mg and K during translocation and distribution in shoots . . . . . . . . . . ........... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 252
4. Functional synergism between Mg and K in photosynthesis, phloem loading, and nitrogen metabolism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 253
5. Effects of balanced fertilization of K and Mg on crop yield and quality . . . . . . . . . . . . . . . . . ........... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 254
6. Conclusions. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ........... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 254
CRediT authorship contribution statement . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ........... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 254
Declaration of competing interest . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ........... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 254
Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ........... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 254
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ........... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 254

⇑ Corresponding author.
E-mail address: sguo@njau.edu.cn (S. Guo).

https://doi.org/10.1016/j.cj.2020.10.005
2214-5141/Ó 2020 Crop Science Society of China and Institute of Crop Science, CAAS. Production and hosting by Elsevier B.V. on behalf of KeAi Communications Co., Ltd.
This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).
K. Xie, I. Cakmak, S. Wang et al. The Crop Journal 9 (2021) 249–256

1. Introduction
Magnesium ion (Mg2+) is one of the most abundant cations in
living plant cells, which usually require 1.5–3.5 g kg 1 for optimal
plant growth [1]. Magnesium is vital for several plant physiological
and biochemical processes, including photosynthesis, carbohy-
drate partitioning, and protein synthesis, that affect plant growth
and yield formation [2–5]. In addition to being the centrally bound
ion of the chlorophyll molecule, Mg is an activator of more than
300 enzymes (Fig. 1) [1,6]. The early response of plants to Mg defi-
ciency is an impaired partitioning of photoassimilates between
source and sink organs, resulting in accumulation of carbohydrates
in leaves (source organs) and inhibition of growth of sink organs
such as roots, nodules, and seeds [4,5,7,8]. Symptoms of Mg defi-
ciency appear as leaf interveinal chlorosis, with the development
of chlorotic and necrotic lesions in later stages, particularly under
high light intensity [9,10].
Magnesium deficiency can be simply a consequence of very low
amounts of plant-available Mg in soils, resulting, for example, from
the loss of soil Mg via leaching, especially in acidic and sandy soils.
Because of its large hydrated ionic radius and its correspondingly
weak adsorption to soil colloids, Mg2+ is highly prone to leaching
[11,12]. Leaching losses of Mg2+ below the root zone can be up to
90 kg Mg ha 1, as shown in a sugarcane-cultivated soil in Brazil
[13]. An absolute Mg deficiency might also be associated with
long-term unbalanced crop fertilization neglecting input of magne-
sium fertilizer, and intensive production systems leading to severe
Mg depletion in soil caused by Mg removal by crops [11,12,14].
Removal of Mg with the harvested part of plants may reach up to
100 kg MgO ha 1 [15].
Remarkably, because of the complex competition between Mg2+
and other soil cations, high soluble concentrations of Mg2+ in the
soil solution do not necessarily mean that it is plant-available.
Acidic soils are usually highly saturated with cations such as H+,
Al3+, and Mn2+, which induce Mg deficiency in plants by interfering
with root Mg2+ uptake [11,16,17]. Addition of high Mg2+ alleviates
the inhibition of plant growth and development by many toxic
heavy metals, such as copper and cadmium [18,19]. An antagonis-
tic relationship between iron and Mg has also been described [19].
Application of liming materials to acidic soils is common practice
to correct the pH; however, in the case of calcitic limestone, this
agronomic practice may alter the availability of K+ and Mg2+ to
plants owing to several chemical processes occurring after liming,
such as increased soil adsorption and reduction in root uptake
owing to competition from Ca2+ [20,21]. Continuous excessive
application of K+ and NH+4 fertilizers in cropping systems can
increase the risk of Mg deficiency, resulting from antagonistic
interference with plant Mg2+ uptake [11,22]. As the best-known
nutrient interaction in plant mineral nutrition, the antagonistic
interaction between K and Mg has been widely studied and is con-
sidered the main cause of Mg deficiency. Accordingly, this review
focuses on this interaction.
The locations commonly associated with K-induced Mg defi-
ciency are usually found in tropical and subtropical regions with
high rainfall and soil acidity as well as high soil concentrations of
cationic nutrients such as K and Ca [11,12,23]. K-induced Mg defi-
ciency often occurs in crops with high demand for K for the forma-
tion of yield and quality, such as banana, potato, and sugarcane,
especially at their critical developmental stages with rapid carbo-
hydrate accumulation [5]. For example, up to 1.6 t of K application
per ha was required for banana [24], and imbalanced nutrition
with Mg in banana due to excessive application of K fertilizers
may impair the nutritional composition and value of banana fruits.
In South Africa, use of high amounts of K fertilizers induces Mg
deficiency by reducing root Mg2+ uptake [25]. In some instances,
growers and crop advisors fail to recognize K-induced Mg-
deficiency symptoms in leaves, especially in the case of greenhouse
crops in North China, where high amounts of K fertilizers are com-
monly applied [26]. Although calcareous soil is usually considered
to be rich in available Mg and K in North China, K-induced Mg defi-
ciency frequently occurs because excessive K fertilizer increases
soil-available K to over 600 mg kg 1, higher than its optimum soil
concentration (240–300 mg kg 1) [26,27]. Insufficient or unbal-
anced fertilization with K and Mg is widespread in agricultural pro-
duction, resulting in imbalances in their distribution in plants and
thereby limiting plant growth as well as the nutritional quality of
edible parts of plants [28].
Elucidating K–Mg interactions in plants may help to clarify their
physiological consequences to plant growth and yield improve-
ment, and provide a theoretical basis for balancing fertilization

Fig. 1. The major physiological functions of K and Mg in plant cells. (1) Free Mg2+ and K+ together regulate cation–anion balance and serve as osmotically active ions in
turgor regulation of cells. (2) Both K and Mg play critical roles in photosynthesis, and functional synergy or functional substitution between K and Mg in photosynthesis has
been found in plants. (3) Both K and Mg are activators or cofactors of key enzymes controlling plant physiological and biochemical processes. (4) Mg plays a vital regulatory
role in cellular energy balance, interacting with the pyrophosphate moiety of nucleotide tri- and diphosphates. The energy-rich compounds Mg-ADP and Mg-ATP, which is in
equilibrium with the free Mg2+ pool and is required for K+ absorption across root cell membranes, represent the primary complexed Mg pools in the cytosol. (5) and (6) As
cations, Mg2+ and K+ play important roles in maintaining cellular pH balance, and K+, a major osmolyte in vacuoles, is essential for stomatal opening.

250
K. Xie, I. Cakmak, S. Wang et al.
with K and Mg. The competition between Mg2+ and K+ is generally
considered as unidirectional and governed only by K+, an increase
in K+ concentration reduces Mg2+ uptake, and this inhibition occurs
mainly in plant roots [29]. Evidence is now mounting that their
interaction occurs not only during root uptake but probably also
during root-to-shoot translocation, distribution, and utilization
[12,30,31].
Although K–Mg interaction in plants has been extensively stud-
ied, the synergistic and antagonistic relationships between K and
Mg and its agronomic and physiologic relevance in crop nutrition
have not been well characterized and reviewed. The objectives of
this review are (i) to summarize and analyze the current knowl-
edge of K-Mg interactions with respect to the root absorption,
translocation, and distribution of these cations in plants; and (ii)
to highlight important future research areas including the physio-
logical and molecular mechanism underlying K–Mg interaction,
and functional synergism between Mg and K.
2. Interaction between Mg and K during root uptake
Root absorption of Mg2+ may involve two mechanisms, which
are either highly specific and insensitive to K+ or highly sensitive
to K+ [12,32]. In previous studies [33,34], Mg2+ uptake at the root
surface was inhibited at K+ concentrations greater than 20 lmol
L 1. The generally accepted competition mechanism of Mg2+ with
K+ is a lack of specificity of the individual uptake systems for the
two cations [1,35]. The inhibitory effect of K+ on Mg2+ uptake
may result from competition for metabolically produced binding
compounds or active sites present on the plasma membrane [36].
In Arabidopsis, the Shaker K+ channel AKT1 and KUP/HAK/KT
transporter HAK5, which are expressed primarily in roots and
function in K+ uptake from the external environment, mediate
almost all K+ absorption in roots [37]. Under most conditions, the
majority of K+ is taken up by AKT1, not HAK5, because AKT1 func-
tions in both the low-affinity and high-affinity ranges. The high-
Fig. 2. Possible competitive effects and mechanisms of Mg2+ with K+ in plants.
(1) Mg2+ uptake is facilitated by two systems: a non-selective ion channel (NSCC)
and an H+/Mg2+ exchanger (MHX). NSCC is also capable of K+ transport in leaves and
roots cells [42]. (2) and (3) The two members of class II transporters (OsHKT2;4 and
TaHKT2;1) in rice and wheat were originally described to transport K+, and evidence
[38,39] suggests that they also transport Mg2+ in plants. (4) and (5) A high-affinity
K+ transporter gene (OsHAK1) was up-regulated under Mg starvation in rice roots,
and expression of a putative Mg transporter gene (OsMGT) was up-regulated under
K starvation [46], suggesting that an interaction between Mg and K regulates the
expression of genes controlling their uptake. (AKT, a K+ selective channel).
251
The Crop Journal 9 (2021) 249–256
affinity plant K+ transporter (HKT) family is divided into two sub-
groups (HKT1 and HKT2). Most HKT2 members function as Na+/
K+ transporters with a role in maintaining Na+/K+ homeostasis in
plants, but HKT2;4 seems to be an exception, as it shows perme-
ability to a wide range of cations. The two members of class II
transporters responsible for K transport, OsHKT2;4 in rice and
TaHKT2;1 in wheat, have been suggested to mediate Mg2+ trans-
port in plants (Fig. 2) [38,39], and it can be limited and depend
on competing K+ concentrations in plants [38]. In plants, Mg2+
uptake by roots is mediated mainly by MGT family members,
and OsMGT1 in rice and AtMGT6 in Arabidopsis have been identi-
fied as plasma membrane-localized transporters mediating root
Mg uptake [40,41]. Mg2+ uptake is also facilitated by a non-
selective ion channel (NSCC), which is also permeable to K+
(Fig. 2) [42]. Guo [43] reported that AtCNGC10 (Arabidopsis thaliana
cyclic nucleotide-gated channel 10), located in the plasma mem-
brane, is involved in Mg2+ uptake and long-distance transport. In
contrast to MGTs, AtCNGC10 is not specific for Mg2+ transport,
given that it also regulates K+ transport (Fig. 3) [43–45]. These
molecular mechanisms provide direct evidence for K–Mg antago-
nism in the process of root absorption. A high-affinity K transporter
gene (OsHAK1) was prominently up-regulated in Mg-starved rice
root, and expression of a putative Mg transporter gene (OsMGT)
was markedly up-regulated under K starvation [46], suggesting
that K-Mg interaction might regulate expression of the genes,
which thus control K+ or Mg2+ uptake (Fig. 2). In response to defi-
ciency of a given nutrient, plants may indirectly increase the
uptake of other nutrients by overexpressing or up-regulating non-
specific transporters with the effect of improving the uptake of the
deficient nutrient [47]. The study of K or Mg transporter genes
under corresponding deficiency conditions contributes to under-
standing K–Mg interaction at the molecular level. The above-
mentioned mechanism, such as competition for active sites or
transporters, could partly explain the synergism or antagonism
between K+ and Mg2+ in root uptake or transport in plants. The
molecular basis of K–Mg interaction during root uptake awaits fur-
ther study.
Indirect results or effects in K–Mg interaction during root
uptake are also reported from many studies. Mg2+ uptake was
stimulated at low K+ concentration and inhibited at high K+ con-
centration [29]. In rice, Mg2+ concentration in shoot and root was
significantly reduced with increasing K+ concentrations in growth
medium [48], and a similar negative effect of high K supply on
Mg2+ uptake was found in safflower [49]. Reduction in root Mg2+
uptake by high K+ concentration may involve blocking of nonspeci-
fic Mg2+ transporters by K+ [12]. In contrast, the effects of increas-
ing Mg supply on K+ uptake in plants are inconsistent, and usually,
no or a slight inhibitory effect of high Mg2+ on K+ uptake has been
reported [11,12]. At low K levels in the growth medium, K+ concen-
tration in rice leaves showed no change with increasing Mg2+ sup-
ply; however, an increase in Mg2+ concentration in the medium
significantly reduced K+ uptake at high K levels [30,48]. At ade-
quate K levels, K+ concentration in plants under Mg deficiency
was increased or not affected. For example, K+ uptake was signifi-
cantly increased under insufficient Mg supply in sugar beet, sun-
flower, onion, banana, and Citrus sinensis seedlings [4,50–53],
whereas in perennial ryegrass and safflower, Mg supply had no
effect on K absorption [49,54]. Plant K+ concentrations were
decreased with supply or resupply of Mg in coffee and maize
[55,56], whereas increased levels of Mg showed a minimal effect
on leaf K levels in sugarcane [25]. These conflicting observations
suggest that there is a moderate synergistic, an antagonistic, or
even no effect of Mg2+ on K+ uptake, and the inconsistency among
research groups likely depends on the ratio of K to Mg used in the
growth medium and plant species used in the experiments. Mg2+
exerts little effect on K+ absorption, in part because very specific
K. Xie, I. Cakmak, S. Wang et al. The Crop Journal 9 (2021) 249–256

Fig. 3. Interactions between Mg and K in plants. (A) Uptake, translocation, and distribution. K+ has a strongly antagonistic effect on Mg2+ uptake, but there is no consistently
verifiable influence of Mg2+ on K+ uptake, owing to the nonspecificity of Mg2+ uptake systems. In general, K+ inhibits Mg2+ translocation from roots to shoots, but the
mechanism remains unclear. Guo [43] reported that AtCNGC10, localized in the plasma membrane, is involved in Mg2+ uptake and long-distance transport, and it also
regulates K+ transport in Arabidopsis. However, because not all transporters responsible for root-to-shoot translocation of Mg2+ have been identified, the effects of Mg2+ on K+
translocation are not well characterized. The antagonistic effect on ionic distribution was most significant in older leaves. (B) Synergism or substitution in function. K and Mg
nutrition greatly affects photosynthetic electron transport, formation of photo-assimilates, phloem sucrose loading, and nitrogen metabolism. (SKOR, the K+ loading
transporter in xylem; CNGC10, cyclic nucleotide-gated channel 10, involved in long-distance Mg2+ transport; AA, amino acid; NRT2, nitrate transporter 2; NR, nitrate
reductase; NiR, nitrite reductase; GS, glutamine synthetase; Glu, glutamic acid).

K+ transporters belonging to the high-affinity transport systems in
root cells are not competitively blocked (antagonized) by Mg2+
[38]. The effects of Mg2+ on root K+ uptake might be explained by
effects on the active absorption of K+ across root cell membranes,
given that this process depends on ATPase activity, which is dis-
rupted under Mg deficiency [2,55]. In brief, the antagonistic effect
of Mg on K absorption is not as strong as the inhibitory effect of K
on root Mg uptake, and the magnitude of K and Mg antagonism
seems to be greatly affected by the K/Mg ratio in the growth
medium.
3. Interaction between Mg and K during translocation and
distribution in shoots
Nutrients are transferred to shoots via xylem loading after
uptake from the soil by roots. Synergistic and antagonistic effects
between K and Mg also occur during their transport from root to
shoot and distribution within plants (Fig. 3). Mg2+ transport from
roots into shoots markedly decreased with increasing K supply in
wheat plants [57,58]. Similar depressive effects of high K+ on
Mg2+ transport were also found in rice [48], tomato [59], and Pinus
radiate [60]. In tomato, the K/Mg ratio in the shoots increased sig-
nificantly under high K+ and was higher than that in roots, indicat-
ing that high K+ inhibits Mg2+ root–shoot translocation [59]. A
reduction in the translocation of Mg2+ may be an indirect conse-
quence of a reduction of its uptake at the root. In contrast, K+
translocation was not affected by Mg2+ concentration in wheat for-
age [58], and there was even an increase in K+ translocation with
increasing Mg2+ [48]. Ohno and Grunes [58] reported that both in
soil and hydroponics experiments, Mg2+ concentration in shoot
252
was depressed with increasing K supply because of the inhibitory
effect of K+ on Mg2+ translocation; however, Mg supply did not
affect the total uptake or concentration of K+ in plants. It is clear
that K exerts a strong antagonistic effect on Mg transport, while
Mg exerts either a synergistic or no effect on K transport into
shoots. Karley and White [61] showed that K+ and Mg2+ behave dif-
ferently during xylem and phloem transport, although both are
highly mobile cations. Compared with K+, Mg2+ is more easily
absorbed by parenchymal cells owing to its higher valence. When
the K/Mg ratio becomes imbalanced because of high K+ concentra-
tions, the transport rate of K+ may be much higher than that of
Mg2+ [62]. The major contributor to K+ translocation to the shoot
is the SKOR channel, which is involved in xylem K+ loading [63].
The transcript levels of SKOR in tomato and Arabidopsis were
downregulated under N, P and S deprivation, partly contributing
to the reduction in K+ translocation to shoot [64]; however, to date,
there is little evidence that K+ transport is affected by Mg2+. There
is an overall lack of knowledge of the effects of K+ on Mg2+ trans-
port because the Mg2+ transport proteins in plants are yet to be
identified [39,65]. One candidate, the cation channel AtCNGC10
involved in xylem-mediated long-distance Mg2+ transport, might
allow K+ to inhibit Mg2+ translocation [43,45]. The physiological
and molecular mechanisms mediating the transport of Mg2+ and
K+, including the transporters involved in root-to-shoot transloca-
tion of Mg, should be studied to further elucidate the relationship
of K+ and Mg2+ during root-to-shoot translocation.
Another link between K+ and Mg2+ is their distribution in plant
tissues after root-to-shoot translocation. The distribution of K+
among shoot organs was affected by low Mg supply in sugar beet
[4]. In most cases, when the K/Mg ratio supplied to plants is unbal-
anced, the relationship between K+ and Mg2+ is often found to be
K. Xie, I. Cakmak, S. Wang et al.
antagonistic in source organs but synergistic in sinks. For example,
in potato, high K application was effective in reducing Mg concen-
trations in leaves, but not in roots and tubers [66]. In pear, an
antagonistic effect between K and Mg occurred in leaves, but syn-
ergetic effects of K on Mg in fruits were observed [67]. The positive
relationship in sink organs was attributed partly to the synergistic
and positive effects of K and Mg on the export of photoassimilates
into sink organs such as fruits [68,69]. After root-to-shoot translo-
cation, both K+ and Mg2+ are preferentially delivered to developing
tissues (the sink); moreover, a K/Mg imbalance in tissues can trig-
ger the redistribution of these two mobile elements among leaves
(the source), requiring phloem-targeted Mg transporters that also
remain to be identified. In the case of typical source organs such
as mature or old leaves, strong antagonism was more obvious
(Fig. 3) [55,70], suggesting that the regulation of K and Mg remobi-
lization from source organs might depend on the K/Mg ratio in tis-
sues. For example, Diem and Godbold [70] reported that the K/Mg
ratio in elongating (sink) leaves of Populus trichocarpa was
increased from 3.75 at 0 lmol L 1 K to 8.31 at 4000 lmol L 1 K
supply; however, in older (source) leaves, the same ratio was
increased more sharply, from 0.25 under low K to 13.35 under high
K. Similarly, in Arabidopsis, short-term Mg deficiency reduced the
contents of K in mature but not in expanding (young) leaves, a
finding attributed to the stimulated nutrient retranslocation [71].
Most minerals are sorted into vacuoles for subcellular compart-
mentation after transport into cells [72]. Both intracellular K and
Mg are important for maintaining osmotic balance, enzyme activa-
tion, and cellular pH control (Fig. 1). Transport of Mg2+ or K+ into
vacuoles has been proposed to act in maintenance of cation–anion
balance and osmotic potential, as well as to store excessive Mg2+ or
K+ [6], where they could substitute for each other (Fig. 1). For
example, under serpentine-rich growth conditions (high Mg:Ca
ratios but N, P and K deficiency), Mg2+ as the dominant cation is
responsible for turgor regulation [73,74]. Diem and Godbold [70]
reported that an increase of Mg2+ concentrations in vacuoles com-
pensated the reduction of K+ at low K levels, but that K+ deficit in
the cytoplasm was not balanced by Mg2+. This observation sug-
gests that K+ in vacuoles responsible for maintaining ionic and
osmotic balance can be substituted by Mg2+, but that K+ in cyto-
plasm responsible for physiological and biochemical reactions
may not be. Conn and Gilliham [75] compared ion distribution in
monocot and dicot plants, showing that Mg2+ concentration in
mesophyll was greater than in epidermis in monocots, whereas
Mg2+ in dicots was accumulated mainly in epidermis. However,
in Arabidopsis (a dicot) leaves, Mg2+ concentration was highest in
vacuoles of the mesophyll rather than of epidermal or bundle
sheath cells [73], and after Mg supply to leaves was increased,
Mg2+ concentration increased primarily in the vacuoles of palisade
and spongy mesophyll cells [76]. These results suggest that the
antagonistic or synergistic relationship between Mg and K in dif-
ferent tissues of plants depends on plant species, sink- and source
organ, leaf age, cell type, and the roles of these elements in plants.
4. Functional synergism between Mg and K in photosynthesis,
phloem loading, and nitrogen metabolism
Focus is also given to the effects of K and Mg on formation,
transport, and distribution of carbohydrates and nitrogen (N)
metabolism. The productivity of plants is greatly affected by their
capacity i) to fix atmospheric carbon into organic carbon by photo-
synthesis, ii) to translocate the assimilated carbon from source into
sink organs such as roots, flowers, and seeds, and iii) to use the
assimilated carbon in sink tissues for growth and development
[6]. It is well known [68,69,77–79] that the K and Mg nutritional
253
The Crop Journal 9 (2021) 249–256
status of plants strongly affects these physiological processes in
plants.
A deficiency of K or Mg resulted in a substantial decline in pho-
tosynthesis [69,80]. Given the differing forms and roles of the two
elements in plants, their effects on photosynthetic rate may vary in
different ways. As a dominant osmolyte, K+ accumulating in guard
cells regulates stomatal morphology and function [81], thus affect-
ing stomatal conductance. Given that K+ plays vital roles in main-
taining chloroplast osmoregulation and pH stability, its deficiency
may reduce the activity and content of the Rubisco enzyme and
lead to chlorophyll degeneration [79,82]. In contrast, most Mg2+
is bound in or incorporated into cellular compartments [76], with
the highest concentrations in chloroplasts [61]. Mg2+ is probably
best known for its central position in the chlorophyll molecule,
and facilitates a well-structured organization of grana and stroma
lamellae. Accordingly, Mg deficiency exerts adverse effects on the
structure and function of chloroplasts, the activity of Rubisco, the
capacity of light reactions in the stroma, and the partitioning of
carbon among sink organs [31,68,69], and thereby on carbon
assimilation. Although K and Mg differ in their physiological roles
associated with photosynthesis, their interaction may have syner-
gistic effects on regulation of leaf photosynthetic capacity [48,66],
which needs a balanced supply of K and Mg [59]. The few studies
that have investigated the effects on photosynthesis of the interac-
tions between K and Mg have not identified their mechanisms.
Both K and Mg play critical roles in phloem loading and trans-
port of sugars into sink organs, and their deficiency led to the accu-
mulation of sucrose in source leaves [2,5,83–85]. K promotes
loading of sucrose into phloem channels and activates H+-ATPase,
which resides in the plasma membrane of sieve tube cells and con-
tributes to the maintenance of sucrose loading into phloem chan-
nels [69,86]. As with K, there are two potential reasons for the
impaired phloem loading of sucrose under Mg deficiency: (i) a
reduction of Mg-ATP availability, which leads to impairments in
the H+-ATPase activity and proton gradient formation [3,68,87],
and (ii) a direct effect on sucrose symporter activity (BvSUT1, a
sucrose/H+ symporter expressed in companion cells in sugar beets)
[83]. Koch [66] suggested that although both low K and Mg supply
increased transcript levels of H+/sucrose symporters, the increase
was less pronounced during Mg deficiency, which probably owing
to sugar accumulation in different cell compartments. These obser-
vations indicate that adequate Mg and K supply is required to guar-
antee sufficient re-translocation of assimilates into harvest
products, such as grain, fruits, and tubers.
Magnesium and K are also required for protein biosynthesis and
affect root absorption, utilization, and metabolism of N in plants
[88–90]. Nitrate uptake in roots can be affected by Mg2+ and K+
[50,91,92] via regulation of NRT2 (nitrate) transporters in Ara-
bidopsis [92] and soybean [93]. K is crucial to the activity of key
enzymes required in protein synthesis (such as nitrate reductase)
[89,92], ribosome synthesis, and mRNA turnover [94,95]. Approxi-
mately 75% of leaf Mg appears to be associated either directly or
indirectly with protein synthesis, owing to its roles in ribosomal
structure and function and N metabolism [33]. Mg deficiency
reduced the activities of nitrate reductase and glutathione syn-
thase (Fig. 3) [48,96], resulting in a decrease in protein concentra-
tion and an increase in amounts of amino acids and amines in tea
and soybean mature leaves [72,90]. However, there are few reports
of functional substitution between Mg and K in N metabolism.
Ding et al. [48] reported that Mg could partially replace K in nitrate
reduction and ammonium assimilation (Fig. 3).
These results support the hypothesis that certain non-specific
functions in the plant, at least in part, can be performed by one
cation in the absence of another. The function of K and Mg in plants
is characterized by crosstalk and redundancy. In summary, there
are synergistic effects of Mg and K on physiological processes such
K. Xie, I. Cakmak, S. Wang et al.
as photosynthesis, carbohydrate allocation, and nitrogen metabo-
lism (Fig. 3), and Mg can partially replace K in some functions.
The commonality of these functions appears to be responsible for
physiological synergism between K and Mg in plants.
5. Effects of balanced fertilization of K and Mg on crop yield and
quality
Because of the antagonistic interactions of K+ and Mg2+ in the
process of their absorption, translocation, and distribution, plants
need to maintain a homeostatic balance between K+ and Mg2+
response to changing nutrient status in the soil for optimal growth
and development. In soils, a K/Mg imbalance (such as may be
caused by unbalanced fertilization) limits the growth and survival
of many plant species. Ertiftik [97] found that certain combinations
of K and Mg (80 kg K2O ha 1 and 40 kg MgO ha 1) maximized the
yield components of maize. Gerendás and Führs [28] concluded
that quality parameters decisive for horticultural crops, such as
total soluble solid concentrations and acidity, often more closely
correlated with K/Mg ratios than with Mg2+ concentrations alone.
The fertilizer ratio of K/Mg required for best yield may be different
from that for quality in the same crop. Zengin [98] reported that
maximum potato tuber yields were achieved with a K/Mg fertilizer
ratio of 3 (120 kg K2O ha 1 with 40 kg MgO ha 1) applied to acidic
soils deficient in K and Mg, whereas a K/Mg ratio of 1.6 led to the
highest starch concentration in tubers, reflecting good potato qual-
ity [99]. These findings emphasize the potential of a balanced
between K and Mg fertilization to increase crop yield and quality.
Suchartgul [100] reported that when the K/Mg ratios in the soil
and leaves were 2.0 and 3.0, respectively, the growth of rubber
was best. When the K/Mg ratio in the shoot reached 22, dry matter
of rice attained its highest value [48]. Maximum values were gen-
erally obtained with combined K + Mg applications, which adjusted
defective balances between soil exchangeable K and Mg cations.
The positive effects of balanced K and Mg nutrition are most likely
represented by the maintenance of phloem loading and the trans-
portation of sugars from source to sink organs.
The saturation values of K and Mg were respectively 5% and 10%
[101], which would be sufficient for plants to absorb K and Mg, and
thus the ideal ratio of these two cations in soil would be K:
Mg = 0.5. However, the balances between the two elements found
in different soils or species discussed above differed from those
recommended values. Generally, studies of the recommended val-
ues of K/Mg ratio in soil or plants have produced various results,
owing probably to the diverse experimental conditions: soil and
crop types, form and rate of K and Mg treatments, soil properties,
age and position of leaves, and growth stage. To alleviate K-
induced Mg-deficiency risk in plant species with high K demand
or in soils with high available K, more attention should be paid
to K–Mg interaction. We recommend that overapplication of K fer-
tilizer should be avoided in these situations. This topic represents
an interesting scientific question for future study.
6. Conclusions
Root uptake and shoot transport of Mg2+ in plants are inhibited
by increased K supply, which can exert detrimental effects on pro-
ductivity and nutritional quality of harvested products. However,
there are inconsistent effects of Mg on absorption and transloca-
tion of K+, which probably depend on the plant species studied
and application rates and ratios of K and Mg in growth medium.
There are synergistic effects of Mg and K on photosynthesis, carbo-
hydrate transport and allocation, N metabolism, and osmotic bal-
ance of cells. To avoid antagonistic interaction between K and Mg
and its adverse consequences on yield and nutritional quality, par-
254
The Crop Journal 9 (2021) 249–256
ticular attention should be paid to balanced nutrition with K and
Mg. Ranges for optimal K/Mg ratios should be defined for both soils
and plant tissues. However, determination of such K/Mg ratios is
challenging, because they are influenced by multiple soil chemical
and physical conditions as well as plant factors. The physiological
and molecular mechanisms proposed in this review do not fully
explain the nature of K-Mg interaction, particularly in the context
of translocation from roots to shoots. Future research should focus
on clarification of the physiological and molecular mechanisms
underlying the antagonistic and synergistic interactions between
K and Mg, during their transport within plants and in their physi-
ological functions, with the ultimate goal of increasing nutrient-
use efficiency in crops.
CRediT authorship contribution statement
Kailiu Xie: Visualization, Writing - original draft. Ismail Cak-
mak: Writing—review & editing. Shiyu Wang: Visualization.
Fusuo Zhang: Conceptualization. Shiwei Guo: Conceptualization,
Funding acquisition, Project Administration, Writing—review &
editing.
Declaration of competing interest
The authors declare that they have no known competing finan-
cial interests or personal relationships that could have appeared
to influence the work reported in this paper.
Acknowledgments
This work was supported by the National Key Research and
Development Program of China (2016YFD0200901 and
2016YFD0200305), the Fundamental Research Funds for the Cen-
tral Universities (KJQN201514 and KYZ201625). We specially
appreciate Prof. Chunjian Li of China Agricultural University for
helpful comments that helped us to greatly improve the
manuscript.
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