Salt-Induced Changes in Cytosolic pH and Photosynthesis in Tobacco and Potato Leaves.

Salinity is one of the most common factors limiting the productivity of crops. The damaging effect of salt stress on many vital plant processes is mediated, on the one hand, by the osmotic stress caused by large concentrations of Na+ and Cl- outside the root and, on the other hand, by the toxic effect of these ions loaded in the cell. In our work, the influence of salinity on the changes in photosynthesis, transpiration, water content and cytosolic pH in the leaves of two important crops of the Solanaceae family-tobacco and potato-was investigated. Salinity caused a decrease in photosynthesis activity, which manifested as a decrease in the quantum yield of photosystem II and an increase in non-photochemical quenching. Along with photosynthesis limitation, there was a slight reduction in the relative water content in the leaves and a decrease in transpiration, determined by the crop water stress index. Furthermore, a decrease in cytosolic pH was detected in tobacco and potato plants transformed by the gene of pH-sensitive protein Pt-GFP. The potential mechanisms of the salinity influence on the activity of photosynthesis were analyzed with the comparison of the parameters' dynamics, as well as the salt content in the leaves.

A Theoretical Analysis of Relations between Pressure Changes along Xylem Vessels and Propagation of Variation Potential in Higher Plants.

Variation potential (VP) is an important long-distance electrical signal in higher plants that is induced by local damages, influences numerous physiological processes, and participates in plant adaptation to stressors. The transmission of increased hydraulic pressure through xylem vessels is the probable mechanism of VP propagation in plants; however, the rates of the pressure transmission and VP propagation can strongly vary. We analyzed this problem on the basis of a simple mathematical model of the pressure distribution along a xylem vessel, which was approximated by a tube with a pressure gradient. It is assumed that the VP is initiated if the integral over pressure is more than a threshold one, taking into account that the pressure is transiently increased in the initial point of the tube and is kept constant in the terminal point. It was shown that this simple model can well describe the parameters of VP propagation in higher plants, including the increase in time before VP initiation and the decrease in the rate of VP propagation with an increase in the distance from the zone of damage. Considering three types of the pressure dynamics, our model predicts that the velocity of VP propagation can be stimulated by an increase in the length of a plant shoot and also depends on pressure dynamics in the damaged zone. Our results theoretically support the hypothesis about the impact of pressure variations in xylem vessels on VP propagation.

Influence of electrical signals on pea leaf reflectance in the 400-800-nm range.

Local action of stressors induces generation and propagation of electrical signals (ESs), which influence numerous physiological processes (including photosynthesis, expression of genes, production of phytohormones, etc.) in undamaged parts of plants; i.e. they induce a systemic stress response. Development of methods of remote sensing of this response (in particular, optical methods) is an important practical task for agricultural and ecological monitoring. However, this problem is not sufficiently researched. Earlier, we reported that ESs influence the photochemical reflectance index, which can be calculated on the basis of reflected light at 531 and 570 nm, and these changes are connected with photosynthetic changes. The aim of the current work is investigation of the influence of ESs on reflectance at broad spectral bands (400-500 nm, 500-600 nm, 600-700 nm and 700-800 nm). We showed that burning-induced ESs caused transient increase of intensity of reflected light at the all investigated spectral bands of visible light: reflectance at 600-700 nm had the maximal magnitude of changes and reflectance at 700-800 nm had the minimal magnitude of changes. Dynamics of the reflectance changes were distinguished from dynamics of photosynthetic changes, induced by ESs; i.e. ESs-induced changes in reflectance seem to be weakly connected with the photosynthetic response. Thus, our results show that changes in reflectance at broad spectral bands can also be used for remote sensing of the ESs-induced systemic stress response in plants.

Parameters of electrical signals and photosynthetic responses induced by them in pea seedlings depend on the nature of stimulus.

Local damage induces generation and propagation of variation potentials (VPs) that affect physiological processes in plants. The aims of the work presented here were to investigate parameters of VP induced by burning, heating and mechanical injury in pea seedlings, and to undertake a theoretical analysis of the mechanisms underlying the differences in VP parameters and a study of the photosynthetic responses caused by VPs induced by the damaging factors. The velocity of propagation of burn-induced VP decreased with distance from the damaged area whereas the velocities of heating- and injury-induced VPs were constant. The amplitudes of burn- and heating-induced VPs did not depend on distance whereas the amplitude of VP induced by mechanical injury decreased. VP propagation has been simulated on the basis of wound substance spread. The simulation revealed two possible ways of wound substance propagation: turbulent diffusion from the damaged area and secondary active production in intact cells. The photosynthetic response (decrease in the quantum yield of PSII and raising the level of non-photochemical fluorescence quenching (NPQ)) developed in case of VP entering the intact leaf under heating and burn but was not registered after mechanical injury. An increase in NPQ level was biphasic under burn in comparison with a single-phase one under heating, and the NPQ amplitude was slightly higher under burn. We suggest that differences in photosynthetic responses may be determined by the parameters of VPs induced by stimuli of different nature.

Mathematical Models of Electrical Activity in Plants.

Electrical activity plays an important role in plant life; in particular, electrical responses can participate in the reception of the action of stressors (local electrical responses and oscillations) and signal transduction into unstimulated parts of the plant (action potential, variation potential and system potential). Understanding the mechanisms of electrical responses and subsequent changes in physiological processes and the prediction of plant responses to stressors requires the elaboration of mathematical models of electrical activity in plant organisms. Our review describes approaches to the simulation of plant electrogenesis and summarizes current models of electrical activity in these organisms. It is shown that there are numerous models of the generation of electrical responses, which are based on various descriptions (from modifications of the classical Hodgkin-Huxley model to detailed models, which consider ion transporters, regulatory processes, buffers, etc.). A moderate number of works simulate the propagation of electrical signals using equivalent electrical circuits, systems of excitable elements with local electrical coupling and descriptions of chemical signal propagation. The transmission of signals from a plasma membrane to intracellular compartments (endoplasmic reticulum, vacuole) during the generation of electrical responses is much less modelled. Finally, only a few works simulate plant physiological changes that are connected with electrical responses or investigate the inverse problem: reconstruction of the type and parameters of stimuli through the analysis of electrical responses. In the conclusion of the review, we discuss future perspectives on the simulation of electrical activity in plants.

Variation potential in higher plants: Mechanisms of generation and propagation.

Long-distance intercellular electrical signals, including variation potential (VP) in higher plants, are a potential mechanism of coordinate functional responses in different plant cells under action of stressors. VP, which is caused by damaging factors (e.g., heating, crushing), is transient depolarization with an irregular shape. It can include a long-term depolarization and fast impulse depolarization ('AP-like' spikes). Mechanisms of VP generation and propagation are still under investigation. It is probable that VP is a local electrical response induced by propagation of hydraulic wave and (or) chemical agent. Both hypotheses are based on numerous experimental results but they predict VP velocities which are not in a good accordance with speed of variation potential propagation. Thus combination of hydraulic and chemical signals is the probable mechanism of VP propagation. VP generation is traditionally connected with transient H(+)-ATPase inactivation, but AP-like spikes are also connected with passive ions fluxes. Ca(2+) influx is a probable mechanism which triggers H(+)-ATPase inactivation and ions channels activation at VP.

Ionic nature of burn-induced variation potential in wheat leaves.

Variation potential (VP) in higher plants cells is a transitory depolarization of the plasma membrane occurring in response to external damage. The effects of VP on different physiological processes are actively studied, but little is known about their ionic nature, which limits the interpretation of VP-induced functional changes. It is thought that VP generation is based on transient inactivation of plasma membrane proton pumps and is not connected to passive ionic fluxes. To study burn-induced VP in wheat seedlings, we measured membrane electric potential and cell input resistance. Cell input resistance decreased during VP generation, indicating that ionic channels were activated. In addition, VP amplitude decreased when the extracellular calcium concentration was lowered. When anion channels were blocked by ethacrynic acid addition, the VP had poor depolarization speed and amplitude. A decrease in the chlorine gradient by extracellular chlorine concentration shift leads to lowering of the VP amplitude and depolarization speed. This result indicates the role of chlorine efflux in depolarization phase formation. The VP repolarization is connected to potassium ion efflux, that is confirmed by repolarization suppression under addition of the potassium channel blocker tetraethylammonium (TEA) and an increase in the extracellular potassium concentration. We also showed that the addition of a proton pump inhibitor leads to membrane potential depolarization and inhibition of VP generation. These results suggest that the VP may be formed not only by transient suppression of proton pumps but also by passive ionic fluxes through the membrane.

Simulation of variation potential in higher plant cells.

Variation potential (VP), a propagating electrical signal unique to plants, induces a number of changes in many physiological processes. However, the mechanisms of its generation and propagation are still under discussion and require experimental and theoretical analysis, including VP simulations. The mathematical model for VP formation in plants has been worked out and is based on our previous description of electrophysiological processes in higher plant cells, including plasma membrane ion transport systems (K(+), Cl(-) and Ca(2+) channels, H(+) and Ca(2+)-ATPase, 2H(+)/Cl(-) symporter and H(+)/K(+) antiporter) and their regulation, ion concentration changes in cells and extracellular spaces and buffers in cytoplasm and apoplast. In addition, the VP model takes into account wound substance diffusion, which is described by a one-dimensional diffusion equation, and ligand-gated Ca(2+) channels, which are activated by this substance. The VP model simulates the experimental dependence of amplitude, velocity and shape of VP on the distance from the wounding site and describes the influence of metabolic inhibitors, divalent cation chelators and anion channel blockers on the generation of this electrical reaction, as shown in experiments. Thus, our model favorably simulates VP in plants and theoretically supports the role of wound substance diffusion and Ca(2+) influx in VP development.

The mechanism of propagation of variation potentials in wheat leaves.

Here we examined the mechanism of propagation of variation potential (VP) induced by burning in wheat leaves. Participation of hydraulic and chemical mechanisms in VP transmission was analyzed by optical coherent tomography and a radioactive tracer method, respectively. The speed of the hydraulic signal considerably exceeded the VP velocity. Investigation of a chemical substance spreading from the zone of local wounding was based on experimental data for radioactive marker transmission derived with a one-dimensional diffusion equation. The speed of the marker transmission was in accordance with VP velocity. The elimination of the potential transmission of a chemical signal by a timed severing of the leaf between the burn site and the recorded site blocked VP propagation. We suggest that a VP is formed by the transmission of a wound substance, the velocity of which is likely increased by hydraulic wave propagation.