Pedosphere 33(1): 105–115, 2023

doi: 10.1016/j.pedsph.2022.06.025
ISSN 1002-0160/CN 32-1315/P
© 2023 Soil Science Society of China
Published by Elsevier B.V. and Science Press

Potassium sources, microorganisms and plant nutrition: Challenges and future

research directions

Abdoulaye SOUMARE1,2,3,∗ , Djibril SARR1 and Abdala G. DIÉDHIOU2,3,4

1 UFR ST, Departement of Agroforestry, Assane Seck University of Ziguinchor (UASZ), Ziguinchor BP 523 (Senegal)
2 LaboratoireCommun de Microbiologie (LCM), IRD/ISRA/Université Cheikh Anta Diop (UCAD), Centre de Recherche de Bel Air, Dakar BP 1386 (Senegal)
3 Centre d’Excellence Africain en Agriculture pour la Sécurité Alimentaire et Nutritionnelle (CEA-AGRISAN), UCAD, Dakar BP 5005 (Senegal)
4 Département de Biologie Végétale, Faculté des Sciences et Techniques, UCAD, Dakar BP 5005 (Senegal)

(Received January 5, 2022; revised March 12, 2022; accepted April 19, 2022)

ABSTRACT
Until recently, potassium (K) has not received considerable attention because of the general belief that soils contain ample amounts of this element. In
addition, low rates of K fertilizer application in agriculture have led to rapid depletion of K in the rhizosphere soil in many underdeveloped countries. This
results in various negative impacts, including preventing optimum utilization of applied nitrogen and phosphorus fertilizers. To compensate for these losses,
massive use of K fertilizers in agriculture has been suggested. Potassium fertilizers are manufactured from rock minerals, particularly sylvite (KCl) and
carnallite (KCl·MgCl2 ·6H2 O). Unfortunately, to date, there is no cost-effective technology available for converting rock minerals into potassic fertilizers.
Potassium-solubilizing microorganisms (KSMs) can release K from soil/minerals into plant-available forms, which could be a sustainable option. The
possibility of using KSMs as efficient biofertilizers to improve crop production has been increasingly highlighted by researchers. In this review, the existing
forms of K in soils and their availability and dynamic equilibrium are discussed. In addition, different K fertilizers and their advantages and disadvantages for
crops are described. Furthermore, the microorganisms usually reported as K solubilizers, the research progress on KSMs, and future insights on the use of
these KSMs in agriculture are reviewed. Screening and analyses of the published literature show that organic acid production is the common mechanism of K
solubilization by bacteria and fungi. This review may serve as a proposal for the future research avenues identified here.
Key Words: biofertilizer, crop production, organic acid, K solubilization, K-solubilizing microorganisms, rock minerals

Citation: Soumare A, Sarr D, Diédhiou A G. 2023. Potassium sources, microorganisms and plant nutrition: Challenges and future research directions.
Pedosphere. 33(1): 105–115.

INTRODUCTION their efficiency, increase costs, and damage the environment.

Potassium (K) is one of the 16 elements essential for the
growth and development of animals, humans, and plants. For
plants, K is the third most important nutrient after nitrogen
(N) and phosphorus (P) and is the most abundant nutrient
after N in photosynthetic tissues of land plants. For some
plant species, such as cotton and banana, the demand for K
is higher than that for N and P (Mora et al., 2012). These
latter are often added to soils through fertilization, whereas
the K requirement of crops is often neglected, leading to K
deficiencies in soils. Usually, soils contain large amounts
of K, but unfortunately, only 1%–2% of the total soil K
is immediately available, and the rest is bound to other
minerals. This available amount is not sufficient to meet the
crop requirements. Therefore, K deficiency is one of the
major constraints in crop production (Meena et al., 2015). To
meet the world’s demand of crop production, the application
of K fertilizers is recommended for diverse crops. Howe-
ver, the mass application of these fertilizers can decrease
∗ Corresponding author. E-mail: ablaysoumare@yahoo.fr, asoumare@univ-zig.sn.
Therefore, in order to prevent K fertilizer shortage, there
is a need to develop novel technologies to accelerate the K
release from soil reserves and to investigate the bio-activation
of rock K deposits for direct application. Rhizosphere mi-
crobes, especially K-solubilizing microorganisms (KSMs)
such as arbuscular mycorrhizal fungi, bacteria, and other
fungi, could be an efficient and sustainable alternative to
meet the K demand for crop improvement (Boubekri et al.,
2021). In fact, these microorganisms can effectively release
K from soil K reserves, thereby decreasing the need for
additional K in some agricultural soils. Agricultural soil par-
ticulates contain K-bearing minerals, such as illite, biotite,
orthoclase, mica, and feldspar (Teotia et al., 2017). Certain
species of microorganisms belonging to fungi (Aspergillus
spp. and Aspergillus terreus), actinomycetes, and other bac-
teria (Pseudomonas spp., Bacillus mucilaginosus, Bacillus
edaphicus, Bacillus circulans, Acidithiobacillus ferrooxi-
dans, and Paenibacillus spp.) have long been recognized to
release K from inert K-bearing minerals (Lian et al., 2008).
106
Thus, using these soil microorganisms as biofertilizers has
been investigated in several studies. Current public concerns
about the side effects of agrochemicals and the increasing
interest in low-input agriculture have emphasized the need to
promote the application of KSM biofertilizers in agriculture.
Although KSMs are present in diverse soils, their number
and K solubilization capacity depend on soil type and cli-
matic conditions. Therefore, screening for efficient KSMs in
vitro has become a hotspot for research over several years
(Rajawat et al., 2016). Numerous studies have suggested
that the application of KSMs as biofertilizers will reduce
the use of agrochemicals and might be more beneficial and
economical for sustainable crop production. This review
presents the importance of K for plants and the status of
commercial K fertilizers in terms of their production, use,
and repartition. Furthermore, it emphasizes the search for an
alternative and effective indigenous source of K for plants to
maintain the K status in soils and sustain crop production.
The main mechanism of K solubilization was investigated
through a comprehensive analysis and interpretation of the
existing literature. For this purpose, data from multiple stu-
dies were analyzed to allow a more precise and reliable
comprehension of the current knowledge on KSMs and their
underlying mechanisms.
K IN SOIL
Soil minerals from silt, clay, and sand are natural reser-
voirs of K. Based on their availability to crops, K is often
subdivided into four fractions: solution K, exchangeable K,
non-exchangeable or fixed K, and mineral K (K in primary
mineral structure). These forms are in dynamic equilibrium
(Fig. 1), which is controlled by the exchange properties,
mineral makeup, and weathering rate of the soil. The relative
abundances of the different forms of K are in the following
order of mineral K (> 90%–98% of total K) > fixed K (1%–
10% of total K) > available K (1%–2% of total K). Their
availability to plants is in the following order of solution K
> exchangeable K > fixed K > mineral K (Kirkman et al.,
1994). The most available forms exist in soil solution or
are adsorbed onto soil colloidal surfaces as exchangeable
Fig. 1 Forms of soil K and their dynamic equilibrium.
A. SOUMARE et al.
K. The exchangeable K is electrostatically retained on the
outer surface of clay minerals and organic substances, while
non-exchangeable K exists predominantly in silicate forms; it
is held between adjacent tetrahedral layers of minerals and is
not readily available. The most common K-bearing minerals
in soils are orthoclase, feldspar, biotite, illite, aluminosi-
licate, mica, and muscovite (Etesami et al., 2017). Among
these rock minerals, feldspar and mica represent 90%–98%
of the total (Zhang and Kong, 2014), and K release from
these two minerals is too slow to be of much agronomic
use. Mineral K availability to plants depends on the degree
of weathering, while K cation availability depends on the
relative amounts of other cations, including calcium (Ca)
and magnesium (Mg). This implies that K availability does
not depend only on the K content, because K, Ca, and Mg
are strongly antagonistic to each other. For instance, some
studies have shown that excess Mg suppresses or inhibits
the uptake of K (Fageria, 1974) and Ca (Rains and Epstein,
1967) through competitive processes.
Besides the K present in soil minerals, potash deposits
exist worldwide. The global geological potash deposits are
estimated at 210 billion tonnes, accounting for 2.1%–2.3%
of the earth’s crust. The deposits are mainly distributed
in Canada, Brazil, Germany, Belarus, Jordan, Israel, New
Mexico, and China. Almost 90% of these reserves are concen-
trated in Canada, Russia, and Belarus (Fig. 2). World produ-
ction reached approximately 45.6 million tonnes in 2021,
with the highest consumption in Asia and South America
(Valdez et al., 2020). Potash deposits are generally in the form
of sylvinite, i.e., a combination of sylvite (KCl) with halite
(NaCl), arcanite (K2 SO4 ), or carnallite (KCl·MgCl2 ·6H2 O).
Other rock K deposits, such as nepheline syenites, phono-
lites, and trachytes, are occasionally found but less studied.
Potash deposits were formed over geological time from the
evaporation of seawater salts. K exists in seawater as KCl at
a concentration of 380 mg L−1 K+ . Different precipitation
methods have been attempted to recover K from seawater.
Fig. 2 The top 10 countries in terms of potash reserve in the world.
K, MICROORGANISMS AND PLANT NUTRITION
However, these processes face practical problems and require
a large amount of energy. Potash deposits are currently the
commercial source of K fertilizers, even if their exploitation
is difficult, expensive, and unsustainable. Fortunately, some
natural biological processes are used by plants and microbes
for the bioactivation of K in soil or rock deposits.
RHIZOSPHERES AS ISLANDS OF K FERTILITY
Root exudates and K availability
Plant roots exudate different compounds, including car-
bohydrates, organic acids, amino acids, phenolic compounds,
enzymes, gaseous molecules (such as CO2 and H2 ), and
inorganic ions such as HCO− − +
3 , OH , and H (Dakora and
Phillips, 2002), which are directly or indirectly involved in
K solubilization. For instance, Yang et al. (2019) showed
that, under K-deficiency treatments, tobacco plants exudate a
large amount of organic acids and activate a greater amount
of K-bearing minerals. The organic acids from root exudates
or plant and animal residue decomposition promote silicate
breakdown and lead to the release of K from silicate minerals
with K in their crystal structures (Teotia et al., 2017). Some
Poaceae species release phytosiderophores, which are natu-
ral K- and Fe(III)-solubilizing compounds, from plant roots
and contribute to K availability (Carvalhais et al., 2011).
Recently, Balogh-Brunstad et al. (2008) hypothesized a new
mechanism of K mineral dissolution through the formation
of biofilms on the surfaces of minerals closely associated
with certain bacterial strains. Similar to the P solubilization
process, K solubilization is also accompanied by a decrease
in pH, which ultimately leads to the release of K ions from
mineral K by protonation and acidification. Indeed, H+ in
the soil or soil solution and CO2 from root respiration are
directly related to the release of K from minerals. Waks-
man and Starkey (1931) showed that CO2 from microbial
decomposition of organic matter or root respiration leads to
orthoclase degradation and K release. Plant species differ
in their ability to acidify soil rhizosphere to access non-
exchangeable K (White et al., 2021). For instance, legumes
reduce the rhizosphere pH more effectively than cereals;
through N2 fixation, legumes absorb more cations than an-
ions and release protons (Zhou et al., 2009). Furthermore,
the loss of symbiotically fixed N through NO− 3 -N leaching
may contribute to legume rhizosphere acidification (Haynes,
1983) and, therefore, to K availability and uptake. In addition,
root exudates stimulate microbial communities and provide
the driving force for the development of active microbial
populations in the rhizosphere, thereby being involved in K
solubilization.
K solubilization by rhizosphere microorganisms
The rhizosphere is a narrow zone of soil, in which
107
beneficial microorganisms interact with rhizodeposits and
promote plant nutrition and health. Many rhizobacteria and
rhizofungi can release K from insoluble silicate minerals
through solubilization mechanisms, usually by releasing
chelating organic and inorganic acids (Rogers and Bennett,
2004). Microbes generally excrete organic acids, such as
citric, tartaric, and oxalic acids. For instance, Sheng and He
(2006) reported that the solubilization of illite and feldspar
by rhizospheric microorganisms is associated with produ-
cing 2-ketogluconic, propionic, malic, tartaric, citric, oxalic,
gluconic, succinic, and fumaric acids. Among these compo-
unds, gluconic, oxalic, 2-ketogluconic, and succinic acids
are the most efficient acids involved in the solubilization
of insoluble K (Meena et al., 2016). Several Bacillus and
Pseudomonas strains have been reported to release K from
rock minerals (such as mica, illite, feldspar, and orthoclase)
through the production of oxalic, citric, fumaric, and tartaric
acids (Girgis et al., 2008; Archana et al., 2012). According
to Malinovskaya et al. (1990), feldspar solubilization by B.
mucilaginosus and B. edaphicus is associated with citric,
tartaric, and oxalic acid production. Tartaric acid is the most
frequent agent for mineral K solubilization (Zarjani et al.,
2013). In addition to organic acid production (Uroz et al.,
2009), there are other mechanisms by which microorganisms
solubilize inorganic K, such as production of siderophores
(Vassileva et al., 1997) and exopolysaccharides (EPSs) (An-
janadevi et al., 2016), secretion of phenolic compounds and
humic substances, and complexation between various metal
ions, such as Ca, aluminum, and iron (Fe). For instance,
polysaccharides are involved in solubilization by binding
to metals (Dominguez-Nuez et al., 2016). Groudev (1987)
stated that silicate mineral dissolution is enhanced by EPSs,
extrapolysaccharides, and mucilaginous compounds. Simi-
larly, Anjanadevi et al. (2016) showed that K solubilization
by Bacillus sp. was correlated with the production of EPSs in
the culture medium. Currently, more than 500 siderophores
are produced by microbes, and these compounds could play
a vital role in the solubilization of elements, such as silicon,
Fe, P, and K, in a liquid medium containing muscovite and
biotite (Hutchens et al., 2003; Sharma et al., 2013).
Some fungal strains such as Aspergillus, Penicillium, and
Fusarium can solubilize rock K and potassium aluminum
silicate through the mechanisms similar to those of bacteria
(Lopes et al., 2019). Fungi also produce extracellular poly-
meric substances that adsorb and accumulate cations and
decrease the saturation of these elements. Fungi can enhance
physical weathering by transferring loose material and rock
fragments and releasing organic anions and acids at their
hyphal tips. Furthermore, several studies (Van Schll et al.,
2006; Pinzari et al., 2022) have reported that ectomycorrhizal
fungi can actively weather silicates such as muscovite, K-
vermiculite, phlogopite, and plagioclase to mine nutrients.
108
According to Van Schll et al. (2006), the ectomycorrhizal
hyphae of Paxillus involutus exude low-molecular-weight
organic anions, including citrate and oxalate, accelerating
mineral weathering. These two organic acids are the most
common exudates of ectomycorrhizal fungi. Several other
fungal strategies based on defined metabolic activities and
gene expression have been reported by Pinzari et al. (2022).
Bibliometric analyses of the literature on K solubilization
mechanisms by microorganisms are shown in Fig. 3. Data
were obtained from the Google Scholar and Elsevier Scopus
databases, which are among the most important sources of ci-
tation data. We identified 50 publications that presented KSM
solubilization mechanism(s) and the compounds involved.
The microbial genera found in the literature were Bacillus
(34%) and Pseudomonas (20%) for bacteria and Penicillium
(47%), Aspergillus (20%), and Trichoderma (20%) for fungi
(Fig. S1, see Supplementary Material for Fig. S1). Their
predominance may be related to their catabolic versatility.
Fig. 3 Hierarchical clustering of K-solubilizing microorganisms (KSMs) based on the organic compounds produced to solubilize K. The most secreted acids
are positioned following each cluster. Results are generated from bibliometric analyses of the literature on K solubilization mechanisms using the Google
Scholar and Elsevier Scopus databases.
A. SOUMARE et al.
The hierarchical clustering of KSMs, based on the organic
compounds produced to solubilize K, was performed using
the package FactoMineR in R software (v.3.6.1) and three
clusters were identified. Cluster 3 included the bacteria most
frequently reported as producers of oxalic, lactic, gluconic,
acetic, and citric organic acids for K solubilization (Fig. 3).
These are the most common organic compounds. The bacte-
ria in Cluster 1 mainly produce fumaric, propionic, and citric
organic acids for K solubilization. However, the bacteria
in Cluster 2 present the most diverse solubilization mecha-
nisms; in addition to organic acids, they also produce EPSs,
siderophores, and other organic ligands. The predominant
acids for bacteria are the same as those for fungi, i.e., citric,
gluconic, oxalic, and succinic acids (Fig. S2, see Supplemen-
tary Material for Fig. S2). Although this analytical approach
provides an overview of the K solubilization mechanisms and
the involved compounds, it also has limitations and pitfalls.
Indeed, the predominance of organic acids in the literature
K, MICROORGANISMS AND PLANT NUTRITION
can be explained by the fact that previous studies targeted
these compounds before to identify them later. Therefore,
other potential mechanisms were ignored, and research on
new mechanisms remains scarce. Thus, research into develo-
ping an efficient protocol using insoluble K for the isolation
and characterization of KSMs is crucial.
ISOLATION AND CHARACTERIZATION OF KSMS
FROM RHIZOSPHERE SOILS
The Aleksandrov selective agar medium is generally used
for the isolation and quantitative assessment of KSMs. The
medium contains 5.0 g L−1 glucose, 0.5 g L−1 magnesium
sulfate, 0.005 g L−1 ferric chloride, 0.1 g L−1 calcium car-
bonate, 2 g L−1 calcium phosphate, 2 g L−1 K-bearing
minerals, and 3% agar (Aleksandrov et al., 1967). Inspired
by the identification of P solubilizers using the Pikovskaya
agar plate method, the plate assay has been developed for
KSM isolation. The formation of a visible halo/zone on
agar plate is considered the main criterion for KSM isola-
tion. After incubation, colonies exhibiting clear zones of
K solubilization are selected. This plate assay method was
recently optimized by amending Aleksandrov medium with
acid-base indicator dyes, such as phenol red, bromothymol
blue, and bromocresol purple (Rajawat et al., 2016). These
indicators enhance the efficiency of screening and shorten
the detection time of KSMs from 4–5 to 2–3 d. However, the
plate assay method fails when the halo is inconspicuous or
absent, because many isolates do not show any clear zone on
agar plates (Nautiyal, 1999). Hence, the direct measurement
of K solubilization in broth assay becomes necessary. There-
fore, screened isolates were grown in Aleksandrov broth
to quantitatively measure the released K using the flame
photometric method (Boubekri et al., 2021). In addition,
molecular markers, such as 16S ribosomal RNA and internal
transcribed spacers, were used to assess the genetic diversity
of K-solubilizing microorganisms (Wang et al., 2015).
MAJOR FUNCTIONS OF K IN CROP PRODUCTION
AND QUALITY
Potash fertilizer ensures optimal plant growth, and the
K+ requirement for plant growth changes with the develop-
mental stage, crop species, and quantity of K+ available in
soil. K is involved in many physiological processes that are
vital to plant nutrient and water uptake. The optimal cyto-
plasmic concentration for enzyme activity is approximately
100–200 mmol L−1 . Furthermore, K plays an important role
in many fundamental physiological and metabolic processes,
such as controlling ion homeostasis, maintaining turgor (Hu
et al., 2017), reducing transpiration, preventing energy loss,
activating enzymes, and promoting disease resistance. K is
required for CO2 assimilation during photosynthesis and N
109
assimilation (Hu et al., 2015). K also increases amino acid
transport, especially in developing seeds (Pettigrew, 2008).
In addition to improving the utilization of N, K increases the
vitamin C content, thereby influencing protein formation in
plants (Srinivasarao et al., 2007). These different functions
explain why K is often described as a crucial element for
crop production.
K improves N use efficiency through improved N uptake
and storage of carbohydrates in roots, whereas K deficiency
inhibits N absorption and lowers NO− 3 content in the leaves
(Cakmak, 2010; Hu et al., 2017). According to Xu et al.
(2020), K+ and NO− 3 are often positively correlated. Howe-
ver, an antagonistic relationship was observed between K+
and NH+ 4 . Furthermore, K is also necessary for plants to
obtain a significant response to applied P (Haeder et al.,
1973). K intensifies the translocation of P and increases
photosynthesis and carbohydrate production (Ahmed et al.,
2020) as well as the productivity and quality of agricultural
produce (Dhillon et al., 2019). Because K is involved in many
plant activities, its deficiency increases plant susceptibility
to disease (Rawat et al., 2016; Mikhailova et al., 2019),
whereas high K concentrations enhance the transcriptional
response to salinity and drought stresses (Anschtz et al.,
2014). Many deficiency symptoms are associated with K de-
ficiency. Under K deficiency, discoloration gradually appears
on the margins of younger leaves, as illustrated in Fig. 4, in
some crops and tree plants. Moreover, K deficiency leads
to low yield and deterioration of product quality, because
plants become susceptible to drought, overwatering, extreme
temperatures, and diseases caused by nematodes and pests
(Wang et al., 2013; Bahrami-Rad and Hajiboland, 2017). The
current context is marked by increased aridity worldwide;
consequently, K is becoming more critical for its role in
plant stress response. Hence, the availability and uptake of K
are essential for decreasing disease incidence and improving
drought, salinity, and cold resistance in various types of plant
communities (Wang et al., 2013).
ADVANTAGES AND DISADVANTAGES OF K FER-
TILIZER APPLICATION
K in silicate structures can hardly be used by plants when
added as fertilizer. Therefore, manufacturing K fertilizers
is required for crop growth in soils depleted of available
K. Currently, K fertilizers are obtained from sedimentary
deposit rocks. The most frequently manufactured and used
forms are KNO3 , K2 SO4 , K2 CO3 , and KCl. Among these
products, KCl (more than 95% of K fertilizer) is the most
widely used form for agronomic crops. However, its appli-
cation is subject to some precautions, because KCl as a
fertilizer can increase soil salinity, damaging the plants and
other organisms present in the soil. K2 SO4 and KNO3 are
110
Fig. 4 K deficiency symptoms in some crop and tree plants. Symptoms are often seen as necrosis of the leaf tips or margins and orangish discoloration from
the tip, progressing along the margins towards the base of the leaves.
recommended for crops sensitive to Cl− . The K concentra-
tion in commercial fertilizers is reported on either an oxide
basis (CK2 O ) or an elemental basis (CK ), which are related
using the following formula: CK2 O = 1.2CK . A summary of
the advantages and disadvantages of common K fertilizers
is shown in Table I. Currently, these soluble fertilizers are
shipped over long distances at high costs, making them less
accessible to low-income small-scale farmers. Improving K
use efficiency (KUE) and developing natural K fertilizers
with little or no processing will reduce the costs of K fertili-
zers. In this respect, the use of multi-nutrient silicate rock
fertilizers as a low-cost option was tested (Harley and Gilkes,
2000). However, the research on the effectiveness of silicate
rock fertilizers in agricultural practices is conflicting (Van
A. SOUMARE et al.
Straaten, 2006). Currently, concerted efforts are being made
to understand the combined effects of K rock material and
inoculation with KSMs on nutrient availability in soils and
the growth of different crops (Meena et al., 2016; Soumare
et al., 2020a).
IMPROVING K USE EFFICIENCY
Annually, over 30 million tonnes of K fertilizers are
applied to agricultural fields worldwide (Bahadur et al.,
2016). The demand increased at a rate of 2.5% between
2014 and 2019 and was distributed as follows: 56% in Asia,
27% in America, 11% in Europe, and 6% in Africa (Basak
et al., 2017). However, the excessive application of K from
chemical fertilizers leads to environmental concerns without
K, MICROORGANISMS AND PLANT NUTRITION 111

TABLE I
Advantages and disadvantages of common K fertilizers used for crop improvement
K fertilizer
Potassium nitrate (KNO3 , 46%)
Potassium chloride (KCl, 60%–62%)
Potassium sulfate (K2 SO4 , 54%)
Potassium phosphate (KH2 PO4 , 34%)
Advantages
Good water solubility, containing nitrate, preventing
soil acidification
Highly soluble, low price, effective, also containing
35% Cl
Good water solubility, increasing sweetness of fruit
Highly soluble product, source of both P and K,
foliar application
Disadvantage(s)
Expensive
Salt accumulation in the soil, unsuitable for
salt-sensitive crops, cannot be applied to acidic soil
Expensive, increasing soil acidity under long-term use
Leading to leaf damage with repeated applications of
foliar K solutions in high concentrations

improving yield, whereas K deficiency results in reduced
crop yields. In fact, K+ from fertilizers is easily leached
beyond the root zone, and the losses can be equivalent to
90–300 kg K+ ha−1 . Therefore, it is important to improve
KUE to achieve sustainable agriculture and meet the global
food demand.
Usually, the following strategies are proposed to increase
KUE by plants: i) to increase root volume by improving the
root architecture and root hair development. A large root
volume increases root surface area that may increase the
absorption of K and other minerals (Shin, 2014). Significant
advances in depicting root traits, root architecture, and their
controlling molecular mechanisms have been made using
phenotyping and molecular and genetic tools (Badri and
Vivanco, 2009). Once fully deciphered, the genetic basis of
regulating root exudation will provide important knowledge
to enhance root exudation and, therefore, KUE; ii) to increase
K uptake by breeding new varieties with high KUE. This
can be achieved by identifying the beneficial traits in large
germ-plasm collections. Molecular marker-assisted selection
using quantitative trait locus (QTL) mapping technologies
will help to achieve this goal (White et al., 2021); iii) to
increase K mobility. Some microorganisms, such as Bacillus
spp., and some plant species, through their root exudates,
have been reported to increase K mobility.
In addition to these strategies, polymer-coated fertilizer
technology, using compounds such as KCl, has been develo-
ped to increase KUE. Coated fertilizers are in the form of
pellets or beads. Alkyd-type resins and polyurethane are
the two main resins used in such coatings. This technology
improves KUE by gradually releasing coated K nutrient and
minimizing K leaching (Wang et al., 2015). Many studies
showed that the application of controlled-release K improves
KUE and, therefore, crop yields (Singh et al., 2010, Boubekri
et al., 2021). Foliar fertilization is another management stra-
tegy for K, which may help to maximize crop yield and
quality through better K absorption. Currently, microorga-
nisms such as KSMs show great potential in improving K
availability and thus KUE in an environmentally friendly
and cost-effective manner.
APPLICATION OF KSMS TO CROP PRODUCTION
Several studies have indicated that using KSMs as
bio-fertilizers can reduce agrochemical consumption while
improving crop production. In this regard, Meena et al.
(2016) reported that silicate-dissolving KSMs could free K
from insoluble minerals and increase K availability from
84.8% in uninoculated soil to 127.9% in inoculated soil.
Similarly, Singh et al. (2010) used waste mica as the sole
source of K and showed that three KSMs (B. mucilaginosus,
Azotobacter chroococcum, and Rhizobium spp.) significantly
improved K assimilation in both maize and wheat. Bacte-
rial inoculation also increased N and P contents in the
aboveground plant components (Subhashini, 2015). Etesami
et al. (2017) also reported that tomato plants inoculated
with silicate-dissolving bacterial strain B. mucilaginosus
increased their biomass by 125% and nutrient (K and P)
uptake by more than 150% compared to uninoculated plants.
Similar results were observed in sorghum (Badr et al., 2006),
Sudan grass (Basak and Biswas, 2012), and potato (Abdel-
Salam and Shams, 2012). Compared to bacteria, there are
very few studies on fungi; nevertheless, some strains of
filamentous fungi and yeasts have been reported to act as K
solubilizers. For example, Maity et al. (2019) showed that
the K-solubilizing fungus Penicillium pinophilum increased
pomegranate fruit yield by 116.9% when used in conjunction
with K feldspar at 200 g plant−1 . Torulaspora globosa (Sac-
charomycetaceae) releases K from alkaline ultramafic rock
powder (Rosa-Magri et al., 2012). Strain B. mucilaginosus
in the presence of mica as the sole source of K increased
the dry biomass and K uptake in wheat and maize com-
pared with uninoculated controls (Singh et al., 2010). Glowa
et al. (2003) showed the ability of ectomycorrhizal fungi
Piloderma spp. to release K from biotite, microline, and
chlorite, thereby improving K+ nutrition in plants. Apart
from the K-solubilizing activity, KSMs have other beneficial
effects, including the production of phytohormones such as
indolyl-3-acetic acid, cytokinins, and giberellins (Soumare
et al., 2021b), N fixation (Soumare et al., 2020b), P solu-
bilization and mineralization (Soumare et al., 2020a), 1-
aminocyclopropane-1-carboxylic acid production (Boubekri
et al., 2021), and biological control of pathogens (Gopalakri-
shnan et al., 2020). In addition to their role in solubilization,
112 A. SOUMARE et al.

organic acids produced by KSMs can be helpful in mitigating
the toxic effects of heavy metals (Archana et al., 2012).
Through an indirect mechanism, some of these microorga-
nisms decompose organic materials that can produce strong
acids, such as H2 SO4 and HNO3 , which solubilize K-bearing
minerals. Despite these important results, the large-scale
application of KSMs in crop production is greatly hindered
by their variability in performance under field conditions.
Table II provides some examples of the agronomical ef-
fectiveness of the direct application of K-bearing rocks with
KSMs to crops in greenhouse conditions.
At the field scale, the application of KSMs as bioferti-
lizers in agriculture remains low. Only a few results have
been reported under field conditions. For example, in a field
experiment B. mucilaginosus showed a high capability to
mobilize K from K-containing minerals and improve nutrient
uptake and growth of cucumber and pepper on Inceptisol
in Korea (Han and Lee, 2005). Another field experiment by
Subramanian et al. (2006) showed a significant increase in
rice grain yield (5 218 vs. 4 419 kg ha−1 for the control) due
to inoculation with silicate-solubilizing bacteria.
CHALLENGES FOR COMMERCIAL K BIOFERTILI-
ZERS AT THE INDUSTRIAL LEVEL
To date, few biofertilizers based on KSMs are available
on the market, and most of them are based on Frateuria spe-
cies. For instance, Symbion-K, Green K, ABTEC Bio-Potash,
UPTAKE, and K Soil B® are based on Frateuria aurantia and
ADVEN is based on Frateuria sp., a K-mobilizing bacterium
(Kore et al., 2020). Other products are based on Bacillus
spp., such as POTAK, a liquid formulation of K-mobilizing
Bacillus licheniformis. Overall, there are few biofertilizers
based on K-solubilizing and/or mobilizing microbes com-
pared to N and P fertilizers, which account for 79% and 14%,
respectively, of the global biofertilizer market (Soumare
et al., 2020b). This shows that KSMs remain less studied
and exploited, perhaps because of their slow influence on
crop yield and few conclusive results in the field owing to a
lack of knowledge and awareness about biological K fertili-
zers among farmers. Consequently, further investigation is
required to improve the performance and use of KSMs as
efficient microbial inoculants. Genetic approaches may help
in screening plant genotypes for their capability to release
organic acids, i.e., to use K mineral reserves in the soil.
Recently, Arif et al. (2020) showed that plants can be bred
or genetically modified to release hormones or exudates that
attract and maintain beneficial microbiomes. However, plant-
microbe molecular dialogue and specific signaling molecules
from plants and microorganisms are poorly understood.
More proteomics and metabolomics studies are required
to maximize beneficial plant-microbiome interactions (Arif
et al., 2020; Batista and Singh, 2021). In addition, enginee-
ring of genes responsible for K solubilization into efficient
microbes with good colonization potential and developing
plants that more efficiently use K may be worthwhile goals to
manage and utilize soil microbial communities for agronomic
purposes. Screening functionally expressed genes involved
in the synthesis and release of acids via metagenomic ap-
proaches is another area of research worth exploring (Sindhu
et al., 2016). It is important to understand root-associated
microbiome functioning, their usefulness, and potential
applications under natural conditions (Arif et al., 2020). Fur-
thermore, the solubilization of K and P are similar in several
ways. Hence, it is necessary to develop new K and P fertilizers
with slow solubility using P- and K-solubilizing microorga-
nisms. Therefore, the direct application of P- and K-bearing
rock materials may be agronomically and economically use-
ful (Supanjani et al., 2006; Boubekri et al., 2021). In fact,
this simplicity-based approach does not involve any chemical
processes and is more ecologically sound, because it results
in better adaptation to environmental changes.

TABLE II
Examples of agronomical effectiveness of direct application of K-bearing minerals with K-solubilizing microorganisms (KSMs) to crops in greenhouse
conditions
K-bearing mineral tested KSM(s) used Crop boosted Experiment scale Reference(s)
Mica Streptomyces Wheat Pots Mikhailouskaya and Tcherhysh, 2005;
Pettgrew, 2008; Boubekri et al., 2021
Illite powder Bacillus megaterium Eggplant Pots Han and Lee, 2005
B. megaterium var. Cucumber, pepper Pots Han et al., 2006
phosphaticum Soybean, cotton Pots Pettigrew, 2008
Bacillus edaphicus Rape, cotton Pots Sheng and He, 2006
Waste mica Streptomyces Sorghum Field, pots Gopalakrishnan et al., 2013
B. megaterium var. Maize Pots Singh et al., 2010;
phosphaticum Abdel-Salam and Shams, 2012
K-feldspar Bacillus cereus Potato Pots Ahmed et al., 2020
Enterobacter hormaechei, Okra Pots Prajapati et al., 2013
fungal Aspergillus terreus
E. hormaechei KSB-8 Cucumber Pots Prajapati and Modi, 2016
Bacillus pasteurii Peanut, sesame Pots Youssef et al., 2010
K, MICROORGANISMS AND PLANT NUTRITION
MULTIDISCIPLINARY RESEARCH CONTRIBUTIONS
TO KSM BIOFORMULATION
Providing new formulations with the potential to ful-
fill their expectations requires exploring multidisciplinary
research involving biotechnology, nanotechnology, agro-
biotechnology, chemical engineering, and material sciences.
At the crossroads of these diverse disciplines, several tech-
niques have been developed, such as bioencapsulation, seed
coating, and cell- or spore-lyophilized beads with cryoprotec-
tants, such as mannitol, microcrystalline cellulose, granular
inoculants, and dried polymeric carriers. These contempo-
rary techniques increase microbial cell survival, delivery, and
performance. For instance, bioformulations supplemented
with EPSs help to protect strains from extreme pH, osmotic
shock, desiccation, toxic substances, predators, and radiation
(Chaudhary et al., 2020). Recently, simpler formulations
of potent KSMs using K-bearing rock powder as a carrier
material have been suggested by several authors (Anjanadevi
et al., 2016; Soumare et al., 2021a).
CONCLUSIONS
This review shows that the potential application of KSMs
in agriculture is still under exploration, and studies are
currently confined mostly to the laboratory scale. Most
publications have shown that the main mechanisms of K
solubilization are organic acid production and rhizosphere
acidification. In addition, the literature analysis shows that
the number of published articles related to K solubilization
mechanisms is scattered and lower compared to that of
K solubilization microbes. The groups of microorganisms
involved in bio-solubilization and those currently used in bio-
formulation are identified and categorized and their potential
for improving crop production is discussed.
ACKNOWLEDGEMENT
We would like to thank Younes En-Nahli and Dr. Sitor
Ndour for their support with the statistical analyses.
SUPPLEMENTARY MATERIAL
Supplementary material for this article can be found in
the online version.
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