Potassium: a vital nutrient mediating stress tolerance in plants

Nutrients have been known to affect stress conditions, in fact, nutrient deprivations are stress conditions for plants itself. Likewise, three important nutrients Nitrogen (N), Phosphorus (P) and Potassium (K) mediates major stress responses in plants. Here, involvement of K has been discussed briefly in plant stress response along with its impact on plant development. K has been regarded as immensely important nutrient in agriculture, hence, its deficiency triggers various signaling cascades, finally enabling plants to activate stress adaptation responses. So far, K⁺ has been reported to play pivotal role in various abiotic stresses such as drought, cold, water stresses etc. However, the exact mechanism and interplay of these different abiotic stress regulation by K⁺ is not completely explored and demand further functional investigations. The in-depth understanding of components involved in K⁺ sensing, transport, and homeostasis will enable plant biologist to engineer crop varieties tolerant to abiotic stresses and nutrient deficient soil in near future.

     

What is the role of putrescine accumulated under potassium deficiency?

Biomarker metabolites are of increasing interest in crops since they open avenues for precision agriculture, whereby nutritional needs and stresses can be monitored optimally. Putrescine has the potential to be a useful biomarker to reveal potassium (K⁺) deficiency. In fact, although this diamine has also been observed to increase during other stresses such as drought, cold or heavy metals, respective changes are comparably low. Due to its multifaceted biochemical properties, several roles for putrescine under K⁺ deficiency have been suggested, such as cation balance, antioxidant, reactive oxygen species mediated signalling, osmolyte or pH regulator. However, the specific association of putrescine build-up with low K⁺ availability in plants remains poorly understood, and possible regulatory roles must be consistent with putrescine concentration found in plant tissues. We hypothesize that the massive increase of putrescine upon K⁺ starvation plays an adaptive role. A distinction of putrescine function from that of other polyamines (spermine, spermidine) may be based either on its specificity or (which is probably more relevant under K⁺ deficiency) on a very high attainable concentration of putrescine, which far exceeds those for spermidine and spermine. putrescine and its catabolites appear to possess a strong potential in controlling cellular K⁺ and Ca²⁺, and mitochondria and chloroplasts bioenergetics under K⁺ stress.     

Improvement of Cerium on Photosynthesis of Maize Seedlings Under a Combination of Potassium Deficiency and Salt Stress

Added Ce³⁺ can partly substitute for Ca²⁺ or Mg²⁺ and improve photosynthesis under the deficiency of these elements, but very few studies focused on photosynthetic improvement in maize seedlings caused by K⁺ deficiency, salt stress, especially a combination of K⁺ deficiency and salt stress. In the present study, the effects of Ce³⁺ on the photosynthesis of maize seedlings under the three different stresses were investigated. The results showed that added Ce³⁺ under various stresses increased the ratios of free water/bound water and of K⁺/Na⁺, the pigment contents, the values of Fv/Fm, Y(II), ETR(II), Y(NPQ), Qp, qL, NPQ, and qN of photosystem II (PSII), the values of Y(I) and ETR(I) of photosystem I (PSI) and the expression levels of LhcII cab1 and rbcL, and decreased the values of Y(NO) and Y(NA). This implied that added Ce³⁺ depressed ion toxicity, photodamage of PSII, and acceptor side constraints of PSI, and enhanced adjustable energy dissipation, the responses of photochemistry, and carbon assimilation caused by K⁺ deficiency, salt stress, and the combination of K⁺ deficiency and salt stress. However, Ce³⁺ mitigation of photosynthetic inhibition in maize seedlings caused by the combined stresses was greater than that of salt stress, and Ce³⁺ mitigation under salt stress was greater than that under K⁺ deficiency. In addition, the results also showed that Ce³⁺ cannot improve photosynthesis and growth of maize seedlings under K⁺ deficiency by substituting for K⁺.     

Transport,signaling, and homeostasis of potassium and sodium in plants

Potassium （K＋） is an essential macronutrient in plants and a lack of K＋ significantly reduces the potential for plant growth and development. By contrast, sodium （Na＋）, while beneficial to some extent, at high concentrations it disturbs and inhibits various physiological processes and plant growth. Due to their chemical similarities, some functions of K＋ can be undertaken by Na＋ but K＋ homeostasis is severely affected by salt stress, on the other hand. Recent advances have highlighted the fascinating regulatory mechanisms of K＋ and Na＋ transport and signaling in plants. This review summarizes three major topics： （i） the transport mechanisms of K＋ and Na＋ from the soil to the shoot and to the cellular - compartments; （ii） the mechanisms through which plants sense and respond to K＋ and Na＋ availability; and （iii） the components involved in maintenance of K＋/Na＋ homeostasis in plants under salt stress.     

Properties of Shaker-type Potassium Channels in Higher Plants

Potassium (K⁺), the most abundant cation in biological organisms, plays a crucial role in the survival and development of plant cells, modulation of basic mechanisms such as enzyme activity, electrical membrane potentials, plant turgor and cellular homeostasis. Due to the absence of a Na⁺/K⁺ exchanger, which widely exists in animal cells, K⁺ channels and some type of K⁺ transporters function as K⁺ uptake systems in plants. Plant voltage-dependent K⁺ channels, which display striking topological and functional similarities with the voltage-dependent six-transmembrane segment animal Shaker-type K⁺ channels, have been found to play an important role in the plasma membrane of a variety of tissues and organs in higher plants. Outward-rectifying, inward-rectifying and weakly-rectifying K⁺ channels have been identified and play a crucial role in K⁺ homeostasis in plant cells. To adapt to the environmental conditions, plants must take advantage of the large variety of Shaker-type K⁺ channels naturally present in the plant kingdom. This review summarizes the extensive data on the structure, function, membrane topogenesis, heteromerization, expression, localization, physiological roles and modulation of Shaker-type K⁺ channels from various plant species. The accumulated results also help in understanding the similarities and differences in the properties of Shaker-type K⁺ channels in plants in comparison to those of Shaker channels in animals and bacteria.     

Expression of animal CED-9 anti-apoptotic gene in tobacco modifies plasma membrane ion fluxes in response to salinity and oxidative stress

Apoptosis, one form of programmed cell death (PCD), plays an important role in mediating plant adaptive responses to the environment. Recent studies suggest that expression of animal anti-apoptotic genes in transgenic plants may significantly improve a plant’s ability to tolerate a variety of biotic and abiotic stresses. The underlying cellular mechanisms of this process remain unexplored. In this study, we investigated specific ion flux “signatures” in Nicotiana benthamiana plants transiently expressing CED-9 anti-apoptotic gene and undergoing salt- and oxidative stresses. Using a range of electrophysiological techniques, we show that expression of CED-9 increased plant salt and oxidative stress tolerance by altering K⁺ and H⁺ flux patterns across the plasma membrane. Our data shows that PVX/CED-9 plants are capable of preventing stress-induced K⁺ efflux from mesophyll cells, so maintaining intracellular K⁺ homeostasis. We attribute these effects to the ability of CED-9 to control at least two types of K⁺-permeable channels; outward-rectifying depolarization-activating K⁺ channels (KOR) and non-selective cation channels (NSCC). A possible scenario linking CED-9 expression and ionic relations in plant cell is suggested. To the best of our knowledge, this study is the first to link “ion flux signatures” and mechanisms involved in regulation of PCD in plants.     

Genetic Control and Mechanisms of Salt and Hyperosmotic Stress Resistance in Cyanobacteria

Exposure to high concentrations of environmental NaCl exerts two stress effects on living cells, increasing the osmotic pressure and the concentration of inorganic ions. Salt stress dramatically suppresses the photosynthetic activity in cells of phototrophic organisms, such as cyanobacteria. During salt adaptation, cyanobacterial cells accumulate osmoprotectors, export excessive Na⁺ with the help of Na⁺/H⁺ antiporters, and actively absorb K⁺ with the help of K⁺-transporting systems. These physiological processes are accompanied by induction or suppression of several genes involved in salt adaptation. The review considers the main mechanisms responsible for the resistance of cyanobacterial cells to salt and hyperosmotic stresses. Special emphasis is placed on recent achievements in studying the genetic control of salt resistance and regulation of gene expression during adaptation of cyanobacteria to salt and hyperosmotic stresses.     

The combined effect of salinity and heat reveals a specific physiological,biochemical and molecular response in tomato plants

Many studies have described the response mechanisms of plants to salinity and heat applied individually; however, under field conditions some abiotic stresses often occur simultaneously. Recent studies revealed that the response of plants to a combination of two different stresses is specific and cannot be deduced from the stresses applied individually. Here, we report on the response of tomato plants to a combination of heat and salt stress. Interestingly, and in contrast to the expected negative effect of the stress combination on plant growth, our results show that the combination of heat and salinity provides a significant level of protection to tomato plants from the effects of salinity. We observed a specific response of plants to the stress combination that included accumulation of glycine betaine and trehalose. The accumulation of these compounds under the stress combination was linked to the maintenance of a high K⁺ concentration and thus a lower Na⁺/K⁺ ratio, with a better performance of the cell water status and photosynthesis as compared with salinity alone. Our findings unravel new and unexpected aspects of the response of plants to stress combination and provide a proposed list of enzymatic targets for improving crop tolerance to the abiotic field environment.     

The potassium transporter OsHAK21 functions in the maintenance of ion homeostasis and tolerance to salt stress in rice

The intracellular potassium (K⁺) homeostasis, which is crucial for plant survival in saline environments, is modulated by K⁺ channels and transporters. Some members of the high‐affinity K⁺ transporter (HAK) family are believed to function in the regulation of plant salt tolerance, but the physiological mechanisms remain unclear. Here, we report a significant inducement of OsHAK21 expression by high‐salinity treatment and provide genetic evidence of the involvement of OsHAK21 in rice salt tolerance. Disruption of OsHAK21 rendered plants sensitive to salt stress. Compared with the wild type, oshak21 accumulated less K⁺ and considerably more Na⁺ in both shoots and roots, and had a significantly lower K⁺ net uptake rate but higher Na⁺ uptake rate. Our analyses of subcellular localizations and expression patterns showed that OsHAK21 was localized in the plasma membrane and expressed in xylem parenchyma and individual endodermal cells (putative passage cells). Further functional characterizations of OsHAK21 in K⁺ uptake‐deficient yeast and Arabidopsis revealed that OsHAK21 possesses K⁺ transporter activity. These results demonstrate that OsHAK21 may mediate K⁺ absorption by the plasma membrane and play crucial roles in the maintenance of the Na⁺/K⁺ homeostasis in rice under salt stress.     

Herbivore exposure alters ion fluxes and improves salt tolerance in a desert shrub

Plants have evolved complex mechanisms that allow them to withstand multiple environmental stresses, including biotic and abiotic stresses. Here, we investigated the interaction between herbivore exposure and salt stress of Ammopiptanthus nanus, a desert shrub. We found that jasmonic acid (JA) was involved in plant responses to both herbivore attack and salt stress, leading to an increased NaCl stress tolerance for herbivore-pretreated plants and increase in K⁺/Na⁺ ratio in roots. Further evidence revealed the mechanism by which herbivore improved plant NaCl tolerance. Herbivore pretreatment reduced K⁺ efflux and increased Na⁺ efflux in plants subjected to long-term, short-term, or transient NaCl stress. Moreover, herbivore pretreatment promoted H⁺ efflux by increasing plasma membrane H⁺-adenosine triphosphate (ATP)ase activity. This H⁺ efflux creates a transmembrane proton motive force that drives the Na⁺/H⁺ antiporter to expel excess Na⁺ into the external medium. In addition, high cytosolic Ca²⁺ was observed in the roots of herbivore-treated plants exposed to NaCl, and this effect may be regulated by H⁺-ATPase. Taken together, herbivore exposure enhance s A. nanus tolerance to salt stress by activating the JA-signalling pathway, increasing plasma membrane H ⁺ - ATPase activity, promoting cytosolic Ca²⁺ accumulation, and then restricting K⁺ leakage and reducing Na⁺ accumulation in the cytosol.     

Potassium Substitution by Sodium in Plants

Soil salinity is an ever-increasing constraint to crop productivity worldwide especially in countries with irrigated agriculture. In contrast to all the soil reclamation strategies to decrease salt concentrations in root zone, the use of sodium (Na⁺) in plant nutrition may be an interesting tactic. The roles of potassium (K⁺) and Na⁺ in plant nutrition suggest that K⁺ is the only monovalent cation which is essential for most higher plants and is involved in three important functions, i.e., enzyme activation, charge balance and osmoregulation. Plants need a small amount but high concentration of K⁺ for specific functions in the cytoplasm and a major portion (～90%) of it is localized in vacuoles, where it acts as an osmoticum. Maintenance of osmotic potential in vacuoles, a nonspecific function of K⁺, can be achieved by other cations such as Na⁺. For decades an ample amount of work has been done on the substitution of K⁺ by Na⁺ in plant nutrition. In this regard, Na⁺ has the potential to replace K⁺ for some of its functions. In some plants, supplementation of Na⁺ in reduced amounts can eliminate K⁺ deficiency symptoms under limited K⁺ supply. Thus, the question of K⁺ substitution by Na⁺ in plant physiology is not only of academic interest but has considerable practical implications in relation to fertilizer management and plant growth in salt-affected environments. In this review, we discuss the possibilities of K⁺ substitution by Na⁺ under specific soil and environmental conditions.     

β-aminobutyric acid-mediated tobacco tolerance to potassium deficiency

??-Aminobutyric acid (BABA) is known as a nonprotein amino acid that can enhance the capacity of plants to resist various biotic and abiotic stresses. However, the role of BABA in protecting tobacco (Nicotiana tabacum L.) against potassium deficiency is not well understood. In this study, we demonstrated that the appropriate concentration of BABA promoted tobacco growth and enhanced plant tolerance to potassium deficiency. BABA protected tobacco from the reduction in the primary root length and chlorophyll content and increased K⁺ uptake via the regulation of expression of potassium up-taking-related NtHAK, NtLKS, NKT1, and NtKC genes. In addition, BABA affected the balancing between hydrogen peroxide level and peroxidase activity under low-potassium stress. Together, our results suggest that BABA plays a role in enhancing potassium deficiency stress tolerance by increasing K⁺ uptake, at least in part, via a ROS-dependent mechanism.     

Comparative physiological analysis in the tolerance to salinity and drought individual and combination in two cotton genotypes with contrasting salt tolerance

Soil salinity and drought are the two most common and frequently co‐occurring abiotic stresses limiting cotton growth and productivity. However, physiological mechanisms of tolerance to such condition remain elusive. Greenhouse pot experiments were performed to study genotypic differences in response to single drought (4% soil moisture; D) and salinity (200 mM NaCl; S) stress and combined stresses (D + S) using two cotton genotypes Zhongmian 23 (salt‐tolerant) and Zhongmian 41 (salt‐sensitive). Our results showed that drought and salinity stresses, alone or in combination, caused significant reduction in plant growth, chlorophyll content and photosynthesis in the two cotton genotypes, with the largest impact visible under combined stress. Interestingly, Zhongmian 23 was more tolerant than Zhongmian 41 under the three stresses and displayed higher plant dry weight, photosynthesis and antioxidant enzymes activities such as superoxide dismutase (SOD), peroxidase (POD) catalase (CAT) and ascorbate peroxidase (APX) activities compared to control, while those parameters were significantly decreased in salt‐stresses Zhongmian 41 compared to control. Moreover, Na⁺/K⁺‐ATPase activity was more enhanced in Zhongmian 23 than in Zhongmian 41 under salinity stress. However, under single drought stress and D + S stress no significant differences were observed between the two genotypes. No significant differences were detected in Ca²⁺/Mg²⁺‐ATPase activity in Zhongmian 41, while in Zhongmian 23 it was increased under salinity stress. Furthermore, Zhongmian 23 accumulated more soluble sugar, glycine‐betaine and K⁺, but less Na⁺ under the three stresses compared with Zhongmian 41. Obvious changes in leaf and root tips cell ultrastructure was observed in the two cotton genotypes. However, Zhongmian 23 was less affected than Zhongmian 41 especially under salinity stress. These results give a novel insight into the mechanisms of single and combined effects of drought and salinity stresses on cotton genotypes.     

The molecular mechanism of plasma membrane H+-ATPases in plant responses to abiotic stress

Plasma membrane H⁺-ATPases (PM H⁺-ATPases) are critical proton pumps that export protons from the cytoplasm to the apoplast. The resulting proton gradient and difference in electrical potential energize various secondary active transport events. PM H⁺-ATPases play essential roles in plant growth, development, and stress responses. In this review, we focus on recent studies of the mechanism of PM H⁺-ATPases in response to abiotic stresses in plants, such as salt and high pH, temperature, drought, light, macronutrient deficiency, acidic soil and aluminum stress, as well as heavy metal toxicity. Moreover, we discuss remaining outstanding questions about how PM H⁺-ATPases contribute to abiotic stress responses.     

Physiological and molecular aspects of salt stress in plants

The study of salt stress mechanisms in plants has become an important issue for the modern agricultural development, climate change, and global food crisis. The plant response to high salt concentrations is complex and comprehensive; it includes many different processes, which should be correctly coordinated. The effect of excessive salt concentrations on plants results in osmotic stress and creates an ionic inbalance due to the accumulation of toxic ions, such as Cl^? and, especially, Na⁺. Salt stress also has negative impact on mineral homeostasis, in particular Ca²⁺ and K⁺. The progress in transcryptomics, genomics, and molecular biology revealed a new gene families that participate in the formation of salt stress response in plants. This review describes the fundamental principles and mechanisms of plant salt tolerance, maintenance of ion homeostasis. In this paper the detailed analysis of the maine transport membrane systems responsible for the transport of ions and their role in plant salt stress were conducted. The perspectives and directions for the further biotechnological and genetic improvement of salt tolerance in plants are underlied.     

Changes in cellular distribution regulate SKD1 ATPase activity in response to a sudden increase in environmental salinity in halophyte ice plant

Halophyte Mesembryanthemum crystallinum L. (ice plant) rapidly responds to sudden increases in salinity in its environment by activating specific salt-tolerant mechanisms. One major strategy is to regulate a series of ion transporters and proton pumps to maintain cellular Na⁺/K⁺ homeostasis. Plant SKD1 (suppressor of K⁺ transport growth defect 1) proteins accumulate in cells actively engaged in the secretory processes, and play a critical role in intracellular protein trafficking. Ice plant SKD1 redistributes from the cytosol to the plasma membrane hours after salt stressed. In combination with present knowledge of this protein, we suggest that stress facilitates SKD1 movement to the plasma membrane where ADP/ATP exchange occurs, and functions in the regulation of membrane components such as ion transporters to avoid ion toxicity.