The adaption of plants to stressful environments depends on long-distance responses in plant organs, which themselves are remote from sites of perception of external stimuli. Jasmonic acid (JA) and its derivatives are known to be involved in plants’ adaptation to salinity. However, to our knowledge, the transport of JAs from roots to shoots has not been studied in relation to the responses of shoots to root salt treatment. We detected a salt-induced increase in the content of JAs in the roots, xylem sap, and leaves of pea plants related to changes in transpiration. Similarities between the localization of JA and lipid transfer proteins (LTPs) around vascular tissues were detected with immunohistochemistry, while immunoblotting revealed the presence of LTPs in the xylem sap of pea plants and its increase with salinity. Furthermore, we compared the effects of exogenous MeJA and salt treatment on the accumulation of JAs in leaves and their impact on transpiration. Our results indicate that salt-induced changes in JA concentrations in roots and xylem sap are the source of accumulation of these hormones in leaves leading to associated changes in transpiration. Furthermore, they suggest the possible involvement of LTPs in the loading/unloading of JAs into/from the xylem and its xylem transport.

Adaptation to changes in the environment depends, in part, on signaling between plant organs to integrate adaptive response at the level of the whole organism. Changes in the delivery of hormones from one organ to another through the vascular system strongly suggest that hormone transport is involved in the transmission of signals over long distances. However, there is evidence that, alternatively, systemic responses may be brought about by other kinds of signals (e.g., hydraulic or electrical) capable of inducing changes in hormone metabolism in distant organs. Long-distance transport of hormones is therefore a matter of debate. This review summarizes arguments for and against the involvement of the long-distance transport of cytokinins in signaling mineral nutrient availability from roots to the shoot. It also assesses the evidence for the role of abscisic acid (ABA) and jasmonates in long-distance signaling of water deficiency and the possibility that Lipid-Binding and Transfer Proteins (LBTPs) facilitate the long-distance transport of hormones. It is assumed that proteins of this type raise the solubility of hydrophobic substances such as ABA and jasmonates in hydrophilic spaces, thereby enabling their movement in solution throughout the plant. This review collates evidence that LBTPs bind to cytokinins, ABA, and jasmonates and that cytokinins, ABA, and LBTPs are present in xylem and phloem sap and co-localize at sites of loading into vascular tissues and at sites of unloading from the phloem. The available evidence indicates a functional interaction between LBTPs and these hormones.

Little information is available on how rhizosphere bacteria affect abscisic acid (ABA) levels in plants and whether these bacterial effects are associated with improved plant water status. In this study, we tested the hypothesis that the stimulation of plant growth may be associated with the ability of ABA to increase the hydraulic conductivity of roots through the up-regulation of aquaporin. To do this, we studied the effect of bacteria capable of producing ABA on a barley mutant deficient in this hormone. Measurements of hydraulic conductivity of the ABA-deficient barley mutant Az34 showed that its tissues exhibited a reduced ability to conduct water, which correlated with lower ABA content in plants. The inoculation of Bacillus subtilis IB-22 stimulated the growth of both the mutant and its parent variety. Also, under the influence of bacteria, the ABA content in plants increased, and the increase was more significant in the mutant. This effect was accompanied by an increase in hydraulic conductivity in the roots of the ABA-deficient mutant, and immunolocalization using antibodies against PIP2;1 and PIP2;2 aquaporins revealed an increase in their abundance. Thus, the results obtained support the hypothesis about the importance of a sufficiently high ABA content in plants to maintain the abundance of aquaporins, hydraulic conductivity and the growth of barley plants.

Inhibition of root elongation is an important growth response to salinity, which is thought to be regulated by the accumulation of jasmonates and auxins in roots. Nevertheless, the mechanisms of the interaction of these hormones in the regulation of the growth response to salinity are still not clear enough. Their better understanding depends on the study of the distribution of jasmonates and auxins between root cells. This was achieved with the help of immunolocalization of auxin (indoleacetic acid) and jasmonates on the root sections of pea plants. Salinity inhibited root elongation and decreased the size of the meristem zone and the length of cells in the elongation zone. Immunofluorescence based on the use of appropriate, specific antibodies that recognize auxins and jasmonates revealed an increased abundance of both hormones in the meristem zone. The obtained data suggests the participation of either auxins or jasmonates in the inhibition of cell division, which leads to a decrease in the size of the meristem zone. The level of only auxin and not jasmonate increased in the elongation zone. However, since some literature evidence argues against inhibition of root cell division by auxins, while jasmonates have been shown to inhibit this process, we came to the conclusion that elevated jasmonate is a more likely candidate for inhibiting root meristem activity under salinity conditions. Data suggests that auxins, not jasmonates, reduce cell size in the elongation zone of salt-stressed plants, a suggestion supported by the known ability of auxins to inhibit root cell elongation.

On March 3, 2024, sad news spread among Russian plant physiologists: an outstanding scientist and wonderful person, Farida Minnikhanovna Shakirova, died. Friends and colleagues exchanged letters in an attempt to honor her memory. Members of Editorial board of the journal “Ecobiotech”, created by colleagues F.M. Shakirova, decided to write about her scientific activities and her amazing qualities: kindness, dedication to science, efficiency, talent as a researcher, willingness to share her experience with colleagues. We would like young colleagues who did not have the opportunity to know her personally to learn from her example.

Increased auxin levels in root primordia are important in controlling root branching, while their interaction with abscisic acid (ABA) likely regulates lateral root development in water-deficient plants. The role of ABA accumulation in regulating root branching was investigated using immunolocalization to detect auxin (indoleacetic acid, IAA) and ABA (abscisic acid) in root primordia of the ABA-deficient barley mutant Az34 and its parental genotype (cv. Steptoe) barley plants. Osmotic stress strongly inhibited lateral root branching in Steptoe plants, but hardly affected Az34. Root primordial cells of Steptoe plants had increased immunostaining for ABA but diminished staining for IAA. ABA did not accumulate in root primordia of the Az34, and IAA levels and distribution were unaltered. Treating Az34 plants with exogenous ABA decreased root IAA concentration, while increasing root primordial ABA accumulation and decreasing root primordial IAA concentration. Although ABA treatment of Az34 plants increased the root primordial number, it decreased the number of visible emerged lateral roots. These effects were qualitatively similar to that of osmotic stress on the number of lateral root primordia and emerged lateral roots in Steptoe. Thus ABA accumulation (and its crosstalk with auxin) in root primordia seems important in regulating lateral root branching in response to water stress.

The phytoremediation of soil contaminated with petroleum depends on the association of plants with rhizosphere bacteria capable of promoting plant growth and destroying petroleum hydrocarbonates. In the present work, we studied the effects of "Lenoil" biological product containing bacteria Pseudomonas turukhanskensis IB 1.1, capable of destroying petroleum hydrocarbons on Secale cereale L plants, which previously proved to be resistant to the weak oil pollution of gray forest soil and to the composition of microorganisms in their rhizosphere. The composition of microorganisms in the rhizosphere of rye roots was studied, morphometric parameters of shoots and roots of rye plants were estimated, and hormone concentration was immunoassayed under conditions of 4% petroleum pollution of the soil. Addition of petroleum to the soil increased the content of oligonitrophilic (by 24%) and hydrocarbon-oxidizing (by 33%) microorganisms; however, the content of cellulolytic (by 12.5 times) microorganisms in the rhizosphere decreased. The use of Lenoil led to a further increase in the number of cellulolytic (by 5.6 times) and hydrocarbon-oxidizing (by 3.8 times) microorganisms and a decrease in the number of oligonitrophilic (by 22.7%) microorganisms in the rhizosphere. Under petroleum pollution, the content of auxins (IAA), zeatin riboside, zeatin nucleotide, and zeatin decreased, while the content of abscisic acid (ABA) increased in the shoots of rye plants. Lenoil treatment led to an eight-fold increase in the IAA content in the roots and a decrease in the ABA content in the aerial part and in the roots. It was shown for the first time that the treatment of petroleum-contaminated soil with "Lenoil" increased root mass due to the development of lateral roots, concurrent with high root IAA content. Petroleum pollution increased the deposition of lignin and suberin in the roots, which strengthened the apoplastic barrier and, thus, reduced the infiltration of toxic components. The deposition of suberin and lignin decreased under "Lenoil" treatment, indicating a decrease in the concentration of toxic petroleum components in the soil degraded by the bacteria. Thus, the biological preparation reduced the growth-inhibiting effect of petroleum on rye plants by increasing the content of growth-stimulating phytohormones (IAA and cytokinins) and reducing the content of ABA, justifying the importance of further study of relevant hormones for the improvement of phytoremediation.

Depending on their habitat conditions, plants can greatly change the growth rate of their roots. However, the mechanisms of such responses remain insufficiently clear. The influence of a low level of illumination on the content of endogenous auxins, their localization in leaves and transport from shoots to roots were studied and related to the lateral root branching of barley plants. Following two days' reduction in illumination, a 10-fold reduction in the emergence of lateral roots was found. Auxin (IAA, indole-3-acetic acid) content decreased by 84% in roots and by 30% in shoots, and immunolocalization revealed lowered IAA levels in phloem cells of leaf sections. The reduced content of IAA found in the plants under low light suggests an inhibition of production of this hormone under these conditions. At the same time, two-fold downregulation of the LAX3 gene expression, facilitating IAA influx into the cells, was detected in the roots, as well as a decline in auxin diffusion from shoots through the phloem by about 60%. It was suggested that the reduced emergence of lateral roots in barley under a low level of illumination was due to a disturbance of auxin transport through the phloem and down-regulation of the genes responsible for auxin transport in plant roots. The results confirm the importance of the long distance transport of auxins for the control of the growth of roots under conditions of low light. Further study of the mechanisms that control the transport of auxins from shoots to roots in other plant species is required.

Background Plant growth promoting rhizobacteria (PGPR) often increase leaf area. Since water loss by transpiration is proportional to leaf area, adaptive responses will be required if further growth is not to be compromised by plant water deficits. However, studies of water relations of plants growing in soil enriched with PGPR have yielded conflicting results and an analysis of the causes of this contradiction are needed if PGPR application is to be utilized successfully in practical farming. Scope We discuss the effects of PGPR on stomatal conductance, transpiration, leaf relative water content, leaf water potential, root growth and hydraulic conductance. We assess their importance for maintaining water balance and sustaining and promoting plant growth following bacterial inoculation. We also assess the involvement of PGPR on hormone concentrations in planta (primarily abscisic acid, ABA) in regulating these processes. Conclusions PGPR exert variable effects on plant water relations. For example, both increased and decreased stomatal conductance and relative water content have been reported. Effectiveness of these alternative responses is discussed in relation to soil water availability. Plant responses to PGPR can be classified in terms of: (1) stimulating root growth thereby increasing capacity for water uptake and compensating for increased transpirational load (2) promoting stomatal closure and reduced transpiration, (3) increased activity of water channels (AQPs) in roots resulting in greater hydraulic conductivity; (4) osmotic adjustment that helps maintain leaf turgor under decreased leaf hydration.;

Plants frequently experience hypoxia due to flooding caused by intensive rainfall or irrigation, when they are partially or completely submerged under a layer of water. In the latter case, some resistant plants implement a hypoxia avoidance strategy by accelerating shoot elongation, which allows lifting their leaves above the water surface. This strategy is achieved due to increased water uptake by shoot cells through water channels (aquaporins, AQPs). It remains a puzzle how an increased flow of water through aquaporins into the cells of submerged shoots can be achieved, while it is well known that hypoxia inhibits the activity of aquaporins. In this review, we summarize the literature data on the mechanisms that are likely to compensate for the decline in aquaporin activity under hypoxic conditions, providing increased water entry into cells and accelerated shoot elongation. These mechanisms include changes in the expression of genes encoding aquaporins, as well as processes that occur at the post-transcriptional level. We also discuss the involvement of hormones, whose concentration changes in submerged plants, in the control of aquaporin activity.

The stomatal closure of salt-stressed plants reduces transpiration bringing about the maintenance of plant tissue hydration. The aim of this work was to test for any involvement of aquaporins (AQPs) in stomatal closure under salinity. The changes in the level of aquaporins in the cells were detected with the help of an immunohistochemical technique using antibodies against HvPIP2;2. In parallel, leaf sections were stained for abscisic acid (ABA). The effects of salinity were compared to those of exogenously applied ABA on leaf HvPIP2;2 levels and the stomatal and leaf hydraulic conductance of barley plants. Salinity reduced the abundance of HvPIP2;2 in the cells of the mestome sheath due to it being the more likely hydraulic barrier due to the deposition of lignin, accompanied by a decline in the hydraulic conductivity, transpiration, and ABA accumulation. The effects of exogenous ABA differed from those of salinity. This hormone decreased transpiration but increased the shoot hydraulic conductivity and PIP2;2 abundance. The difference in the action of the exogenous hormone and salinity may be related to the difference in the ABA distribution between leaf cells, with the hormone accumulating mainly in the mesophyll of salt-stressed plants and in the cells of the bundle sheaths of ABA-treated plants. The obtained results suggest the following succession of events: salinity decreases water flow into the shoots due to the decreased abundance of PIP2;2 and hydraulic conductance, while the decline in leaf hydration leads to the production of ABA in the leaves and stomatal closure.

The influence of illumination on the level of endogenous auxins content and localization and their transport from shoots to roots in regulating lateral root branching of barley plants was investigated. The four times reducing of illumination was found to suppress the lateral roots formation in two days after beginning of exposure. The decrease in roots and shoots IAA content and in the intensity of phloem cells staining for auxins on leaf sections were observed already in a day after the light was reduced. A decline in auxin content in the roots was appeared to a greater extent compared to the shoots. At the same time, suppression of LAX3 gene responsible for the cell auxin influx was found in the roots along with a decrease in the rate of auxin flow through the phloem from shoots. It was assumed the inhibition of the lateral roots growth in barley under reduced level of irradiance in the photosynthetically active region was due to impaired phloem transport of auxin and a decrease in the level of expression of the auxin transporter in plant roots.

Reports on importance of aquaporins for plant salt-resistance are rather contradictory. We have studied the consequences of expression of barley (Hordeum vulgare L.) aquaporin gene, HvPIP2;1, in transgenic tobacco plants on their water relations and capacity to grow under salinity. Detection of HvPIP2;1 protein and mRNA by immunolocalization and in situ hybridization, respectively, confirmed that expression of this gene occurred in most of leaf cells. In the absence of salt stress, improvements in water status resulting from HvPIP2;1 expression were manifested in increased relative water content (RWC) best noticeable in young leaves, where expression of their own tobacco gene NtPIP2;1 was relatively low. Whole small leaves and leaf discs overexpressing HvPIP2;1, exposed to moderate level of salinity (100 and 300 mM), were characterized by greater accumulation of fresh mass, but their fresh mass decreased faster when concentration of NaCl increased to 400 and 500 mM. The ability to increase fresh mass was manifested only when the gradient between the leaf and the solution of NaCl was maintained due to accumulation of osmotics. Thus the effect of expression of an alien aquaporin gene on leaf growth under salinity depends on capacity of plants for osmotic adjustment.

Inoculation with plant growth-promoting rhizobacteria can increase plant salt resistance. We aimed to reveal bacterial effects on the formation of apoplastic barriers and hormone concentration in relation to maintaining ion homeostasis and growth of salt-stressed plants. The rhizosphere of a durum wheat variety was inoculated with cytokinin-producing Bacillus subtilis and auxin-producing Pseudomonas mandelii strains. Plant growth, deposition of lignin and suberin and concentrations of sodium, potassium, phosphorus and hormones were studied in the plants exposed to salinity. Accumulation of sodium inhibited plant growth accompanied by a decline in potassium in roots and phosphorus in shoots of the salt-stressed plants. Inoculation with both bacterial strains resulted in faster appearance of Casparian bands in root endodermis and an increased growth of salt-stressed plants. B. subtilis prevented the decline in both potassium and phosphorus concentrations and increased concentration of cytokinins in salt-stressed plants. P. mandelii decreased the level of sodium accumulation and increased the concentration of auxin. Growth promotion was greater in plants inoculated with B. subtilis. Increased ion homeostasis may be related to the capacity of bacteria to accelerate the formation of Casparian bands preventing uncontrolled diffusion of solutes through the apoplast. We discuss the relative impacts of the decline in Na accumulation and maintenance of K and P content for growth improvement of salt-stressed plants and their possible relation to the changes in hormone concentration in plants.

Hypoxia as a stress condition occurs due to a decrease in the availability of oxygen in the environment as a result of natural waterlogging of soils or suboptimal watering. In laboratory conditions, when growing plants in hydroponics, hypoxia was modeled by stopping the aeration of the nutrient solution, which led to a rapid decrease of oxygen concentration in it. Under these conditions was observed, the formation of aerenchyma, through which oxygen can penetrate the roots from the shoot. In addition, the luminescence of berberine complexes with lignin and suberin in the cell membranes of the outer layers of the cortex increased, which indicates the beginning of the formation of apoplastic barriers in this area, which in turn can reduce the diffusion of oxygen into the solution on the way of its transport to the apical part of the root along aerenchyma. In the morphological features the most striking was the decrease in the length of the root under hypoxic conditions, which reduses the path of oxygen transport from the shoot to the root tips. However, the decrease in the rate of shoot growth and photosynthesis under the influence of hypoxia was insignificant, which confirms the adaptive significance of the aerenchyma and apoplast barriers formation that provide oxygen supply to the roots.

Lipid transfer proteins (LTPs) participate in many important physiological processes in plants, including adaptation to stressors, e.g., salinity. Here we address the mechanism of this protective action of LTPs by studying the interaction between LTPs and abscisic acid (ABA, a “stress” hormone) and their mutual participation in suberin deposition in root endodermis of salt-stressed pea plants. Using immunohistochemistry we show for the first time NaCl induced accumulation of LTPs and ABA in the cell walls of phloem paralleled by suberin deposition in the endoderm region of pea roots. Unlike LTPs which were found localized around phloem cells, ABA was also present within phloem cells. In addition, ABA treatment resulted in both LTP and ABA accumulation in phloem cells and promoted root suberization. These results suggested the importance of NaCl-induced accumulation of ABA in increasing the abundance of LTPs and of suberin. Using molecular modeling and fluorescence spectroscopy we confirmed the ability of different plant LTPs, including pea Ps-LTP1, to bind ABA. We therefore hypothesize an involvement of plant LTPs in ABA transport (unloading from phloem) as part of the salinity adaptation mechanism.

Local effects on plant roots or shoots are accompanied by a change of plant hormones concentration providing signal transduction in plants and their adaptation to changing environmental conditions. It is known that plants respond to drought stress by increasing the concentration of abscisic acid (ABA) in xylem sap which leads to decrease in stomatal conductance to prevent plant water loss from transpirational pathways. Earlier, we found a rapid leaf ABA accumulation in barley and wheat plants under influence of the shortterm salinity. However, the mechanism of the salt stress induced rapid accumulation of abscisic acid in plant leaves remained unclear. The aim of this study was to investigate whether the salt-induced rapid accumulation of abscisic acid in the leaves was the result of its inflow from the roots (as a root signal). In our experiments the short-term salinity did not increase but decreased the concentration of abscisic acid in xylem sap of wheat plants. Thus, detected accumulation of ABA in the leaves did not result from an increase in its inflow from the roots. Apparently leaf ABA accumulation already detected in 15 min after the onset of salinity stress could be a local reaction due to its metabolism in the shoot itself. The decrease in the leaf water potential could induce the change of abscisic acid metabolism pathways in the shoot that led to the accumulation of this hormone. A small short-term accumulation of ABA in the roots did not lead to an increase in their hydraulic conductivity.

The trans-membrane carrier AtENT3 is known to transport externally supplied cytokinin ribosides and thus promote uptake by cells. However, its role in distributing either exogenous or endogenous cytokinins within the intact plant has not hitherto been reported. To test this, we used ent3-1 mutant Arabidopsis seedlings in which the gene is not expressed due to a T-DNA insertion, and examined the effect on the concentration and distribution of either endogenous cytokinins or exogenous trans-zeatin riboside applied to the roots. In the mutant, accumulation of endogenous cytokinins in the roots was reduced and capacity to deliver externally supplied trans-zeatin riboside to the shoots was increased suggesting involvement of equilibrative nucleoside (ENT) transporter in the control of cytokinin distribution in the plants. Roots of ent3-1 were longer in the mutant in association with their lower cytokinin concentration. We concluded that the ENT3 transporter participates in partitioning endogenous cytokinins between the apoplast and the symplast by facilitating their uptake by root cells thereby limiting cytokinin export to the shoots through the xylem. Dilution of the mineral nutrient solution lowered endogenous cytokinin concentration in the roots of both wild type (WT) and ent3-1 plants accompanied by promotion of root elongation. Nevertheless, cytokinin content was lower, while roots were longer in the ent3-1 mutant than in the WT under either normal or deficient mineral nutrition suggesting a significant role of ENT3 transporter in the control of cytokinin level in the roots and the rate of their elongation.

An ABA-deficient barley mutant (Az34) and its parental cultivar (Steptoe) were compared. Plants of salt-stressed Az34 (100 mmol m−3 NaCl for 10 days) grown in sand were 40% smaller than those of “Steptoe”, exhibited a lower leaf relative water content and lower ABA concentrations. Rhizosphere inoculation with IB22 increased plant growth of both genotypes. IB22 inoculation raised ABA in roots of salt-stressed plants by supplying ABA exogenously and by up-regulating ABA synthesis gene HvNCED2 and down-regulating ABA catabolic gene HvCYP707A1. Inoculation partially compensated for the inherent ABA deficiency of the mutant. Transcript abundance of HvNCED2 and related HvNCED1 in the absence of inoculation was 10 times higher in roots than in shoots of both mutant and parent, indicating that ABA was mainly synthesized in roots. Under salt stress, accumulation of ABA in the roots of bacteria-treated plants was accompanied by a decline in shoot ABA suggesting bacterial inhibition of ABA transport from roots to shoots. ABA accumulation in the roots of bacteria-treated Az34 was accompanied by increased leaf hydration, the probable outcome of increased root hydraulic conductance. Thereby, we tested the hypothesis that the ability of rhizobacterium (Bacillus subtilis IB22) to modify responses of plants to salt stress depends on abscisic acid (ABA) accumulating in roots.

The role of reactive oxygen species (ROS) in ABA-induced increase in hydraulic conductivity was hypothesized to be dependent on an increase in aquaporin water channel (AQP) abundance. Single ABA application or its combination with ROS manipulators (ROS scavenger ascorbic acid and NADPH oxidase inhibitor diphenyleneiodonium chloride (DPI)) were studied on detached roots of barley plants. We measured the osmotically driven flow rate of xylem sap and calculated root hydraulic conductivity. In parallel, immunolocalization of ABA and HvPIP2;2 AQPs was performed with corresponding specific antibodies. ABA treatment increased the flow rate of xylem, root hydraulic conductivity and immunostaining for ABA and HvPIP2;2, while the addition of antioxidants prevented the effects of this hormone. The obtained results confirmed the involvement of ROS in ABA effect on hydraulic conductivity, in particular, the importance of H2O2 production by ABA-treated plants for the effect of this hormone on AQP abundance.