In this paper, we discuss biological potassium acquisition and utilization processes over an evolutionary timescale, with emphasis on modern vascular plants. The quintessential osmotic and electrical functions of the K+ ion are shown to be intimately tied to K+-transport systems and membrane energization. Several prominent themes in plant K+-transport physiology are explored in greater detail, including: (1) channel mediated K+ acquisition by roots at low external [K+]; (2) K+ loading of root xylem elements by active transport; (3) variations on the theme of K+ efflux from root cells to the extracellular environment; (4) the veracity and utility of the “affinity” concept in relation to transport systems. We close with a discussion of the importance of plant-potassium relations to our human world, and current trends in potassium nutrition from farm to table.

Compartmental analysis by tracer efflux (CATE) is fundamental to examinations of membrane transport, allowing study of solute movement among subcellular compartments with high temporal, spatial, and chemical resolution. CATE can provide a wealth of information about fluxes and pool sizes in complex systems, but is a mathematically intensive procedure, and there is a need for software designed to fully, easily, and dynamically analyse results from CATE experiments. Here we present vaCATE (Visualized Automation of Compartmental Analysis by Tracer Efflux), a software package that meets these criteria. A robust suite of test cases using CATE datasets from experiments with intact rice (Oryza sativa L.) root systems reveals the high fidelity of vaCATE and the ease with which parameters can be extracted, using a three-compartment model and a curve-stripping procedure to distinguish them on the basis of variable exchange rates. vaCATE was developed using Python 2.7 and can be used in most situations where compartmental analysis is required. Funding Statement: This work was supported by the Natural Sciences and Engineering Research Council of Canada (NSERC) and the Ontario Graduate Scholarship Fund (OGS).

Provision of silicon (Si) to roots of rice (Oryza sativa L.) can alleviate salt stress by blocking apoplastic, transpirational bypass flow of Na+ from root to shoot. However, little is known about how Si affects Na+ fluxes across cell membranes. Here, we measured radiotracer fluxes of 24Na+, plasma membrane depolarization, tissue ion accumulation, and transpirational bypass flow, to examine the influence of Si on Na+ transport patterns in hydroponically grown, salt-sensitive (cv. IR29) and salt-tolerant (cv. Pokkali) rice. Si increased growth and lowered [Na+] in shoots of both cultivars, with minor effects in roots; neither root nor shoot [K+] were affected. In IR29, Si lowered shoot [Na+] via a large reduction in bypass flow, while in Pokkali, where bypass flow was small and not affected by Si, this was achieved mainly via a growth dilution of shoot Na+. Si had no effect on unidirectional 24Na+ fluxes (influx and efflux), or on Na+-stimulated plasma-membrane depolarization, in either IR29 or Pokkali. We conclude that, while Si can reduce Na+ translocation via bypass flow in some (but not all) rice cultivars, it does not affect unidirectional Na+ transport or Na+ cycling in roots, either across root cell membranes or within the bulk root apoplast.

Vascular plants are major intermediaries in the global water cycle, and are highly adapted to both facilitate and resist water fluxes, such as during root uptake, translocation in the xylem, and transpiration by leaves. Here, we summarize the contributions to a Special Issue on water in the Journal of Plant Physiology, which cluster around the theme of control and facilitation of water movement in plants. We conclude with an editorial view of the need for plant physiologists to consider larger cultural issues surrounding water use, especially in terms of the increasing agricultural demand for water to produce animal feed, with its associated trophic nutritive losses and environmental damage.

A systematic analysis of NaCl-dependent, plasma-membrane depolarization (∆∆Ψ) in rice roots calls into question the current leading model of rapid membrane cycling of Na+ under salt stress.

The nitrogen (N)-use efficiency of agricultural plants is notoriously poor. Globally, about 50% of the N fertilizer applied to cropping systems is not absorbed by plants, but lost to the environment as ammonia (NH3), nitrate (NO3-), and nitrous oxide (N2O, a greenhouse gas with 300 times the heat-trapping capacity of carbon dioxide), raising agricultural production costs and contributing to pollution and climate change. These losses are driven by volatilization of NH-3 and by a matrix of nitrification and denitrification reactions catalysed by soil microorganisms (chiefly bacteria and archaea). Here, we discuss mitigation of the harmful and wasteful process of agricultural N loss via biological nitrification inhibitors (BNIs) exuded by plant roots. We examine key recent discoveries in the emerging field of BNI research, focusing on BNI compounds and their specificity and transport, and discuss prospects for their role in improving agriculture while reducing its environmental impact.

Nitrogen (N) and potassium (K) are the two most abundantly acquired mineral elements by plants, and their acquisition pathways interact in complex ways. Here, we review pivotal interactions with respect to root acquisition, storage, translocation and metabolism, between the K+ ion and the two major N sources, ammonium (NH4+ ) and nitrate (NO3- ). The intersections between N and K physiology are explored at a number of organizational levels, from molecular-genetic processes, to compartmentation, to whole plant physiology, and discussed in the context of both N-K cooperation and antagonism. Nutritional regulation and optimization of plant growth, yield, metabolism and water-use efficiency are also discussed.

Although the global nitrogen (N) cycle is largely driven by soil microbes, plant root exudates can profoundly modify soil microbial communities and influence their N transformations. A detailed understanding is now beginning to emerge regarding the control that root exudates exert over two major soil N processes - nitrification and N2 fixation. We discuss recent breakthroughs in this area, including the identification of root exudates as nitrification inhibitors and as signaling compounds facilitating N-acquisition symbioses. We indicate gaps in current knowledge, including questions of how root exudates affect newly discovered microbial players and N-cycle components. A better understanding of these processes is urgent given the widespread inefficiencies in agricultural N use and their links to N pollution and climate change.

Although deemed a "non-essential" mineral nutrient, silicon (Si) is clearly beneficial to plant growth and development, particularly under stress conditions, including salinity and drought. Here, we review recent research on the physiological, biochemical, and molecular mechanisms underlying Si-induced alleviation of osmotic and ionic stresses associated with salinity and drought. We distinguish between changes observed in the apoplast (i.e., suberization, lignification, and silicification of the extracellular matrix; transpirational bypass flow of solutes and water), and those of the symplast (i.e., transmembrane transport of solutes and water; gene expression; oxidative stress; metabolism), and discuss these features in the context of Si biogeochemistry and bioavailability in agricultural soils, evaluating the prospect of using Si fertilization to increase crop yield and stress tolerance under salinity and drought conditions.

Rapid sodium cycling across the plasma membrane of root cells is widely thought to be associated with Na(+) toxicity in plants. However, the efflux component of this cycling is not well understood. Efflux of Na(+) from root cells is believed to be mediated by Salt Overly-Sensitive-1, although expression of this Na(+)/H(+) antiporter has been localized to the vascular tissue and root meristem. Here, we used a chambered cuvette system in which the distal root of intact salinized barley and Arabidopsis thaliana plants (wild-type and sos1) were isolated from the bulk of the root by a silicone-acrylic barrier, so that we could compare patterns of (24)Na(+) efflux in these two regions of root. In barley, steady-state release of (24)Na(+) was about four times higher from the distal root than from the bulk roots. In the distal root, (24)Na(+) release was pronouncedly decreased by elevated pH (9.2), while the bulk-root release was not significantly affected. In A. thaliana, tracer efflux was about three times higher from the wild-type distal root than from the wild-type bulk root and also three to four times higher than both distal- and bulk-root fluxes of Atsos1 mutants. Elevated pH also greatly reduced the efflux from wild-type roots. These findings support a significant role of SOS1-mediated Na(+) efflux in the distal root, but not in the bulk root.

Carbon dioxide (CO2) concentrations in the earth’s atmosphere are projected to rise from current levels near 400ppm to over 700ppm by the end of the 21st century. Projections over this time frame must take into account the increases in total net primary production (NPP) expected from terrestrial plants, which result from elevated CO2 (eCO2) and have the potential to mitigate the impact of anthropogenic CO2 emissions. However, a growing body of evidence indicates that limitations in soil nutrients, particularly nitrogen (N), the soil nutrient most limiting to plant growth, may greatly constrain future carbon fixation. Here, we review recent studies about the relationships between soil N supply, plant N nutrition, and carbon fixation in higher plants under eCO2, highlighting key discoveries made in the field, particularly from free-air CO2 enrichment (FACE) technology, and relate these findings to physiological and ecological mechanisms.

Potassium (K(+)) acquisition in roots is generally described by a two-mechanism model, consisting of a saturable, high-affinity transport system (HATS) operating via H(+)/K(+) symport at low (<1mM) external [K(+)] ([K(+)]ext), and a linear, low-affinity system (LATS) operating via ion channels at high (>1mM) [K(+)]ext. Radiotracer measurements in the LATS range indicate that the linear rise in influx continues well beyond nutritionally relevant concentrations (>10mM), suggesting K(+) transport may be pushed to extraordinary, and seemingly limitless, capacity. Here, we assess this rise, asking whether LATS measurements faithfully report transmembrane fluxes. Using (42)K(+)-isotope and electrophysiological methods in barley, we show that this flux is part of a K(+)-transport cycle through the apoplast, and masks a genuine plasma-membrane influx that displays Michaelis-Menten kinetics. Rapid apoplastic cycling of K(+) is corroborated by an absence of transmembrane (42)K(+) efflux above 1mM, and by the efflux kinetics of PTS, an apoplastic tracer. A linear apoplastic influx, masking a saturating transmembrane influx, was also found in Arabidopsis mutants lacking the K(+) transporters AtHAK5 and AtAKT1. Our work significantly revises the model of K(+) transport by demonstrating a surprisingly modest upper limit for plasma-membrane influx, and offers insight into sodium transport under salt stress.

The efflux of sodium (Na+) ions across the plasma membrane of plant root cells into the external medium is surprisingly poorly understood. Nevertheless, Na+ efflux is widely regarded as a major mechanism by which plants restrain the rise of Na+ concentrations in the cytosolic compartments of root cells and, thus, achieve a degree of tolerance to saline environments. In this review, several key ideas and bodies of evidence concerning root Na+ efflux are summarized with a critical eye. Findings from decades past are brought to bear on current thinking, and pivotal studies are discussed, both “purely physiological”, and also with regard to the SOS1 protein, the only major Na+ efflux transporter that has, to date, been genetically characterized. We find that the current model of rapid transmembrane sodium cycling (RTSC), across the plasma membrane of root cells, is not adequately supported by evidence from the majority of efflux studies. An alternative hypothesis cannot be ruled out, that most Na+ tracer efflux from the root in the salinity range does not proceed across the plasma membrane, but through the apoplast. Support for this idea comes from studies showing that Na+ efflux, when measured with tracers, is rarely affected by the presence of inhibitors or the ionic composition in saline rooting media. We conclude that the actual efflux of Na+ across the plasma membrane of root cells may be much more modest than what is often reported in studies using tracers, and may predominantly occur in the root tips, where SOS1 expression has been localized.

Unidirectional influx and efflux of nutrients and toxicants, and their resultant net fluxes, are central to the nutrition and toxicology of plants. Radioisotope tracing is a major technique used to measure such fluxes, both within plants, and between plants and their environments. Flux data obtained with radiotracer protocols can help elucidate the capacity, mechanism, regulation, and energetics of transport systems for specific mineral nutrients or toxicants, and can provide insight into compartmentation and turnover rates of subcellular mineral and metabolite pools. Here, we describe two major radioisotope protocols used in plant biology: direct influx ( DI) and compartmental analysis by tracer efflux ( CATE). We focus on flux measurement of potassium ( K+) as a nutrient, and ammonia/ ammonium ( NH3/ NH4 +) as a toxicant, in intact seedlings of the model species barley ( Hordeum vulgare L.). These protocols can be readily adapted to other experimental systems ( e. g., different species, excised plant material, and other nutrients/ toxicants). Advantages and limitations of these protocols are discussed.

K(+) channels are among the best-characterized classes of membrane protein in plants. Nevertheless, in-planta demonstrations of traits emerging from molecular characterizations have often been insufficient or lacking altogether. Such linkages are, however, critical to our basic understanding of plant nutrition and to addressing 'real-world' issues that are faced in environmental and agricultural settings. Here, we cover some of the recent advances in K(+) acquisition with particular focus on voltage-gated K(+) channel functioning and regulation in roots, and highlight where linkages to in-planta behavior have been successfully made and, conversely, where such linkages are yet to be made.

Britto, D. T., Balkos, K. D., Becker, A., Coskun, D., Huynh, W. Q. and Kronzucker, H. J. 2014. Potassium and nitrogen poising: Physiological changes and biomass gains in rice and barley. Can. J. Plant Sci. 94: 1085–1089. Soil nitrogen, potassium, and water are three of the most important factors influencing, often interdependently, the growth of plants. Maximizing plant growth is not simply a matter of maximizing the availability of these and other nutrients; indeed, excess supply can be deleterious to plant performance. Rather, optimal performance may come about by adjusting the supply of each of the disparate factors required for plant growth, not only individually, but in relation to one another. In our work investigating the nutritional maximization of plant growth, we have found that altering the ratios of N and K provided to seedlings of cereal grasses can result in very substantial increases in vegetative biomass accrual, e.g., >220% of low-K+controls, in short-term studies with rice, the world's most important cereal grain, and even greater gains in grain yield, in the longer term. Hence, the findings in our laboratory are of direct relevance to the aim of NSERC's Green Crop Network, which was to contribute to the amelioration of climate change by improvement of carbon capture and sequestration in crop plants. In addition, these findings may help to increase the world's food supply, the security of which is sometimes at odds with proposed means to thwart climate change. Our work in this area has also led to a potential breakthrough of a more fundamental sort in plant nutritional biology, which may in itself have important practical implications: evidence that aquaporin-type transport proteins conduct rapid NH3fluxes into roots at toxic levels of external ammonia/ammonium.

Standard procedures for the tracing of ion fluxes into roots of plants are described here, with emphasis on cations, especially potassium (K(+)). We focus in particular on the measurement of unidirectional influx by use of radiotracers and provide a brief introduction to compartmental analysis by tracer efflux (CATE).

Fluxes of NH3, but not NH4+, account for futile cycling across both plasmalemma and tonoplast in roots of barley, resulting in a thermodynamic NH3 equilibrium between cytosol, vacuole, and external solution, and aquaporins are likely mediators of these fluxes. Futile transmembrane NH3/NH4+ cycling in plant root cells, characterized by extremely rapid fluxes and high efflux to influx ratios, has been successfully linked to NH3/NH4+ toxicity. Surprisingly, the fundamental question of which species of the conjugate pair (NH3 or NH4+) participates in such fluxes is unresolved. Using flux analyses with the short-lived radioisotope 13N and electrophysiological, respiratory, and histochemical measurements, we show that futile cycling in roots of barley (Hordeum vulgare) seedlings is predominately of the gaseous NH3 species, rather than the NH4+ ion. Influx of 13NH3/13NH4+, which exceeded 200 µmol g–1 h–1, was not commensurate with membrane depolarization or increases in root respiration, suggesting electroneutral NH3 transport. Influx followed Michaelis-Menten kinetics for NH3 (but not NH4+), as a function of external concentration (Km = 152 µm, Vmax = 205 µmol g–1 h–1). Efflux of 13NH3/13NH4+ responded with a nearly identical Km. Pharmacological characterization of influx and efflux suggests mediation by aquaporins. Our study fundamentally revises the futile-cycling model by demonstrating that NH3 is the major permeating species across both plasmalemma and tonoplast of root cells under toxicity conditions.