Plants have developed complex systems of perception and signaling to adapt to changing environmental conditions. Electrical signaling is one of the most promising candidates for the regulatory mechanisms of the systemic functional response under the local action of various stimuli. Long-distance electrical signals of plants, such as action potential (AP), variation potential (VP), and systemic potential (SP), show specificities to types of inducing stimuli. The systemic response induced by a long-distance electrical signal, representing a change in the activity of a complex of molecular-physiological processes, includes a nonspecific component and a stimulus-specific component. This review discusses possible mechanisms for transmitting information about the nature of the stimulus and the formation of a specific systemic response with the participation of electrical signals induced by various abiotic factors.

Plants are constantly exposed to environmental stresses. Local stimuli sensed by one part of a plant are translated into long-distance signals that can influence the activities in distant tissues. Changes in levels of phytohormones in distant parts of the plant occur in response to various local stimuli. The regulation of hormone levels can be mediated by long-distance electrical signals, which are also induced by local stimulation. We consider the crosstalk between electrical signals and phytohormones and identify interaction points, as well as provide insights into the integration nodes that involve changes in pH, Ca²⁺ and ROS levels. This review also provides an overview of our current knowledge of how electrical signals and hormones work together to induce a systemic response.

Abscisic acid (ABA) is an important hormone in plants that participates in their acclimation to the action of stressors. Treatment by exogenous ABA and its synthetic analogs are a potential way of controlling the tolerance of agricultural plants; however, the mechanisms of influence of the ABA treatment on photosynthetic processes require further investigations. The aim of our work was to investigate the participation of inactivation of the plasma membrane H⁺-ATP-ase on the influence of ABA treatment on photosynthetic processes and their regulation by electrical signals in peas. The ABA treatment of seedlings was performed by spraying them with aqueous solutions (10⁻⁵ M). The combination of a Dual-PAM-100 PAM fluorometer and GFS-3000 infrared gas analyzer was used for photosynthetic measurements; the patch clamp system on the basis of a SliceScope Pro 2000 microscope was used for measurements of electrical activity. It was shown that the ABA treatment stimulated the cyclic electron flow around photosystem I and decreased the photosynthetic CO₂ assimilation, the amplitude of burning-induced electrical signals (variation potentials), and the magnitude of photosynthetic responses relating to these signals; in contrast, treatment with exogenous ABA increased the heat tolerance of photosynthesis. An investigation of the influence of ABA treatment on the metabolic component of the resting potential showed that this treatment decreased the activity of the H⁺-ATP-ase in the plasma membrane. Inhibitor analysis using sodium orthovanadate demonstrated that this decrease may be a mechanism of the ABA treatment-induced changes in photosynthetic processes, their heat tolerance, and regulation by electrical signals.

A local leaf burning causes variation potential (VP) propagation, a decrease in photosynthesis activity, and changes in the content of phytohormones in unstimulated leaves in pea plants. The VP-induced photosynthesis response develops in two phases: fast inactivation and long-term inactivation. Along with a decrease in photosynthetic activity, there is a transpiration suppression in unstimulated pea leaves, which corresponds to the long-term phase of photosynthesis response. Phytohormone level analysis showed an increase in the concentration of jasmonic acid (JA) preceding a transpiration suppression and a long-term phase of the photosynthesis response. Analysis of the spatial and temporal dynamics of electrical signals, phytohormone levels, photosynthesis, and transpiration activity showed the most pronounced changes in the more distant leaf from the area of local stimulation. The established features are related to the architecture of the vascular bundles in the pea stem.

Cytosolic pH (pH_cyt) regulates a wide range of cellular processes in plants. Changes in pH_cyt occurring under the effect of different stressors can participate in signal transmission. The dynamics of pH_cyt under the action of external factors, including significant factors for open ground crops such as temperature, remains poorly understood, which is largely due to the difficulty of intracellular pH registration using standard methods. In this work, model plants of potato (one of the essential crops) expressing a fluorescent ratiometric pH sensor Pt-GFP were created. The calibration obtained in vivo allowed for the determination of the pH_cyt values of the cells of the leaves, which is 7.03 ± 0.03 pH. Cooling of the whole leaf caused depolarization and rapid acidification of the cytosol, the amplitude of which depended on the cooling strength, amounting to about 0.2 pH units when cooled by 15 °C. When the temperature rises to 35–40 °C, the cytosol was alkalized by 0.2 pH units. Heating above the threshold temperature caused the acidification of cytosol and generation of variation potential. The observed rapid changes in pH_cyt can be associated with changes in the activity of H⁺-ATPases, which was confirmed by inhibitory analysis.

Plants are able to perceive diverse environmental factors and form an appropriate systemic functional response. Systemic responses are induced by stimulus-specific long-distance signals that carry information about the stimulus. Variation potential is proposed as a candidate for the role of such a signal. Here, we focus on the mechanisms that determine the specificity of the variation potential under the action of different local stimuli. Local stimuli such as heating, burning and wounding cause variation potential, the parameters of which differ depending on the type of stimulus. It was found that the stimulus-specific features of the hydraulic signal monitored by changes in leaf thickness and variation potential, such as a greater amplitude upon heating and burning and a significant amplitude decrement upon burning and wounding, were similar. The main features of these signals are the greater amplitude upon heating and burning, and a significant amplitude decrement upon burning and wounding. Together with the temporal correspondence of signal propagation, this evidence indicates a role for the hydraulic signal in the induction of stimulus-specific variation potential. Experiments using mechanosensitive channel inhibitors have demonstrated that the hydraulic signal contributes more to the induction of the variation potential in the case of rapidly growing stimuli, such as burning and wounding, than in the case of gradual heating. For thermal stimuli (gradual heating and burning), a greater contribution, compared to wounding, of the chemical signal related to reactive oxygen species to the induction of the variation potential was demonstrated. Thus, the specificity of the parameters of the variation potential is determined by the different contributions of hydraulic and chemical signals.