The essential nature of Zn and widespread Zn deficiency in plants under field conditions underlie the great interest of researchers in the regulation of plant Zn homeostasis. Here, the current ...knowledge of plant Zn homeostasis regulation, mainly in A. thaliana, is reviewed. The plant Zn homeostasis machinery is regulated largely at the transcriptional level. Local regulation in response to changes in cellular Zn status is based on the transcription factors bZIP19 and bZIP23, which sense changes in free Zn2+ concentrations in the cell. However, there are likely other unidentified ways to sense cellular free Zn2+ concentrations in addition to the well-known bZIP19 and bZIP23 factors. In recent years, the existence of a shoot-derived systemic Zn deficiency signal, which is involved in the upregulation of Zn transport from roots to shoots, was demonstrated. Additionally, rates of mRNA degradation of Zn homeostasis genes are likely regulated by changes in cellular Zn status. In addition to the regulation of Zn transport, other mechanisms for the regulation of plant Zn homeostasis exist. “Zn sparing” mechanisms could be involved in the decrease in plant Zn requirements under Zn deficiency. Additionally, autophagy is probably regulated by local Zn status and involved in Zn reutilization at the cellular level. Current issues related to studying Zn homeostasis regulation are discussed.
•Free-Zn-sensing transcription factors are the main regulators of the cellular Zn response.•Systemic signalling of shoot Zn status regulates gene expression in roots.•Zn status influences both mRNA synthesis and mRNA degradation of Zn homeostasis genes.•Autophagy and Zn sparing likely participate in plant Zn homeostasis.
Tree growth is highly sensitive to water deficit. At the same time, growth processes substantially influence tree performance under water stress by changing the root-absorbing surface, ...leaf-transpiring surface, amount of conducting xylem, etc. Drought-induced growth suppression is often higher in conifers than in broadleaf species. This review is devoted to the relations between the growth of coniferous plants and their performance under increasing climate aridization in the temperate and boreal zones of the Northern Hemisphere. For adult trees, available evidence suggests that increasing the frequency and severity of water deficit would be more detrimental to those plants that have higher growth in favorable conditions but decrease growth more prominently under water shortage, compared to trees whose growth is less sensitive to moisture availability. Not only the overall sensitivity of growth processes to water supply but also the asymmetry in response to lower-than-average and higher-than-average moisture conditions can be important for the performance of coniferous trees under upcoming adverse climate change. To fully understand the tree response under future climate change, the responses to both drier and wetter years need to be analyzed separately. In coniferous seedlings, more active growth is usually linked with better drought survival, although physiological reasons for such a link can be different. Growth stability under exacerbating summer water deficit in coniferous plants can be maintained by more active spring growth and/or by a bimodal growth pattern; each strategy has specific advantages and drawbacks. The optimal choice of growth strategy would be critical for future reforestation programs.
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•Growth processes are highly sensitive to water stress in conifers.•Lower growth sensitivity to moisture is linked with lower drought-induced mortality.•Resource allocation can lead to an asymmetric growth response to a changed water supply.•More active growth is linked with better seedling performance under drought stress.•Stable growth can be maintained by active spring growth or a bimodal growth pattern.
Efficient post-drought recovery of growth and assimilation enables a plant to return to its undisturbed state and functioning. Unlike annual plants, trees suffer not only from the current drought, ...but also from cumulative impacts of consecutive water stresses which cause adverse legacy effects on survival and performance. This review provides an integrated assessment of ecological, physiological and molecular evidence on the recovery of growth and photosynthesis in trees, with a view to informing the breeding of trees with a better ability to recover from water stress. Suppression of recovery processes can result not only from stress damage but also from a controlled downshift of recovery as part of tree acclimation to water-limited conditions. In the latter case, recovery processes could potentially be activated by turning off the controlling mechanisms, but several obstacles make this unlikely. Tree phenology, and specifically photoperiodic constraints, can limit post-drought recovery of growth and photosynthesis, and targeting these constraints may represent a promising way to breed trees with an enhanced ability to recover post-drought. The mechanisms of photoperiod-dependent regulation of shoot, secondary and root growth and of assimilation processes are reviewed. Finally, the limitations and trade-offs of altering the photoperiodic regulation of growth and assimilation processes are discussed.
Melatonin is among one of the promising agents able to protect agricultural plants from the adverse action of different stressors, including salinity. We aimed to investigate the effects of melatonin ...priming (0.1, 1.0 and 10 µM) on salt-stressed potato plants (125 mM NaCl), by studying the growth parameters, photochemical activity of photosystem II, water status, ion content and antioxidant system activity. Melatonin as a pleiotropic signaling molecule was found to decrease the negative effect of salt stress on stolon formation, tissue water content and ion status without a significant effect on the expression of Na
/H
-antiporter genes localized on the vacuolar (
to
) and plasma membrane (
). Melatonin effectively decreases the accumulation of lipid peroxidation products in potato leaves in the whole range of concentrations studied. A melatonin-induced dose-dependent increase in Fv/Fm together with a decrease in uncontrolled non-photochemical dissipation Y(NO) also indicates decreased oxidative damage. The observed protective ability of melatonin was unlikely due to its influence on antioxidant enzymes, since neither SOD nor peroxidase were activated by melatonin. Melatonin exerted positive effects on the accumulation of water-soluble low-molecular-weight antioxidants, proline and flavonoids, which could aid in decreasing oxidative stress. The most consistent positive effect was observed on the accumulation of carotenoids, which are well-known lipophilic antioxidants playing an important role in the protection of photosynthesis from oxidative damage. Finally, it is possible that melatonin accumulated during pretreatment could exert direct antioxidative effects due to the ROS scavenging activity of melatonin molecules.
Heavy metals and aluminum are among the most significant abiotic factors that reduce the productivity and quality of crops in acidic and contaminated soils. The protective effects of brassinosteroids ...containing lactone are relatively well-studied under heavy metal stress, but the effects of brassinosteroids containing ketone are almost unstudied. Moreover, there are almost no data in the literature on the protective role of these hormones under polymetallic stress. The aim of our study was to compare the stress-protective effects of lactone-containing (homobrassinolide) and ketone-containing (homocastasterone) brassinosteroids on the barley plant's resistance to polymetallic stress. Barley plants were grown under hydroponic conditions; brassinosteroids, increased concentrations of heavy metals (Mn, Ni, Cu, Zn, Cd, and Pb), and Al were added to the nutrient medium. It was found that homocastasterone was more effective than homobrassinolide in mitigating the negative effects of stress on plant growth. Both brassinosteroids had no significant effect on the antioxidant system of plants. Both homobrassinolide and homocastron equally reduced the accumulation of toxic metals (except for Cd) in plant biomass. Both hormones improved Mg nutrition of plants treated with metal stress, but the positive effect on the content of photosynthetic pigments was observed only for homocastasterone and not for homobrassinolide. In conclusion, the protective effect of homocastasterone was more prominent compared to homobrassinolide, but the biological mechanisms of this difference remain to be elucidated.
We investigated the specific features of copper and zinc excess action on the roots of canola (Brassica napus L.) plants. Copper rapidly accumulated in canola root cells and reached saturation during ...several hours of treatment, whereas the root zinc content increased relatively slowly. Excessive copper and zinc entry inside the cell resulted in significant cell damage, as evidenced by alterations in plasmalemma permeability and decreases in cellular enzymatic activity. Zinc excess specifically damaged root hair cells, which correlated with a pronounced elevation of their labile zinc level. In vitro, we showed that reduced glutathione (GSH) readily reacted with copper ions to form complexes with blocked sulfhydryl groups. In contrast, zinc ions were ineffective as glutathione blockers, and glutathione molecules did not lose their specific chemical activity in the presence of Zn2+ ions. The effect of copper and zinc excess on the glutathione pool in canola root cells was analysed by a combination of biochemical determination of total and oxidized glutathione contents and fluorescent staining of free reduced glutathione with monochlorobimane dye. Excess copper led to dose-dependent diminution of free reduced glutathione contents in the root cells, which could not be explained by the loss of total cellular glutathione or its oxidation. In contrast, we observed little effect of much higher intracellular zinc concentrations on the free reduced glutathione content. We concluded that GSH plays an important role in copper excess, but not zinc excess chelation, in canola root cells.
•Copper and zinc differed in the pattern of their accumulation in root cells.•Intracellular availability of copper and zinc sharply increased under their excess.•Cell damage correlated with the level of metal intracellular availability.•Glutathione participated in the copper chelation, but not in zinc chelation in vivo.
Brassinosteroids are promising agents for alleviating the negative effects of salinity on plants, but the mechanism of their protective action is far from being understood. We investigated the effect ...of pretreatment with 24-epibrassinolide (24-EBL) on the photosynthetic and physiological parameters of potato plants under progressive salinity stress caused by root application of 100 mM NaCl. Salinity clearly inhibited primary photosynthetic processes in potato plants by reducing the contents of photosynthetic pigments, photosynthetic electron transport and photosystem II (PSII) maximal and effective quantum yields. These negative effects of salinity on primary photosynthetic processes were mainly due to toxic ionic effects on the plant’s ability to oxidize the plastoquinone pool. Pretreatment with 24-EBL alleviated this stress effect and allowed the maintenance of plastoquinone pool oxidation and the efficiency of photosystem II photochemistry to be at the same levels as those in unstressed plants; however, the pretreatment did not affect the photosynthetic pigment content. 24-EBL pretreatment clearly alleviated the decrease in leaf osmotic potential under salinity stress. The stress-induced increases in lipid peroxidation and proline contents were not changed under brassinosteroid pretreatment. However, 24-EBL pretreatment increased the peroxidase activity and improved the K
+
/Na
+
ratio in potato leaves, which were likely responsible for the protective 24-EBL action under salt stress.
Abscisic acid (ABA) is one of the main participants in the regulation of plant responses to water deficiency. Knowledge of the ABA signal transduction pathways in gymnosperms is rather limited, ...especially in comparison with those in angiosperms. Seedlings of Scots pine and Norway spruce are known for their contrasting behaviour strategies under water deficit. To characterize the possible role of ABA in these differences, ABA dynamics were investigated under conditions of water deficit in seedlings of these two species. The content of ABA and its catabolites was followed in the roots and needles of seedlings of Pinus sylvestris and Picea abies under conditions of polyethylene glycol (PEG)-induced water deficiency (−0.15 and −0.5 MPa) for 10 days. The expression of the main genes for ABA-biosynthetic enzymes was also analysed. ABA showed more pronounced stress-dependent dynamics in pine roots than in spruce roots, whereas in needles, the response was greater for spruce than pine. The ABA increase during drought was mainly due to de novo synthesis and the shift in the balance between ABA synthesis and catabolism towards synthesis. The ABA-glucosyl ester did not serve as a reserve for the release of free ABA under water deficiency. The expression levels of the main ABA biosynthetic genes showed a weak or no correlation with changes in ABA content under water stress, i.e., the ABA content in the seedlings of both species was not directly linked to the transcript levels of the main ABA biosynthetic genes. Less-pronounced stress-induced changes in ABA in pine needles than in spruce needles may be related to pine seedlings having a less conservative strategy of growth and maintenance of water balance under water deficit.
•Scots pine and Norway spruce differed in water deficit-induced ABA dynamics.•Changes in ABA content were mostly due to de novo biosynthesis.•ABA-glucosyl ester did not serve as a major source of active ABA under water stress.•ABA content was not directly linked to the transcript levels of ABA biosynthetic genes.
The tight connection between the deterioration of xylem function and plant mortality under drought is well recognized. However, a lack of mechanistic understanding of how substantial conductivity ...loss influences plant performance under drought and during post‐drought recovery hinders our ability to model tree responses to drought stress. We artificially induced a loss of 50% of xylem conducting area in Scots pine and Norway spruce saplings by stem notching and investigated plant performance under drought and during post‐drought recovery. Plant mortality, xylem hydraulic conductivity, leaf water status and stomatal conductance were measured. We observed no preferential mortality of top plant parts (above the notches) compared to basal plant parts (below the notches), and no consistent trend in hydraulic conductivity loss was observed between top and basal parts of dying plants. Stem hydraulic conductivity, water status of the needles and stomatal conductance changed similarly between the top and basal parts during drought and post‐drought recovery, which indicated the substantial hydraulic overcapacity of the stems. The recovery of stomatal conductance demonstrated prominent hysteresis due to non‐hydraulic stomatal limitations. The results obtained are highly important for modelling the influence of plant hydraulic impairment on plant performance under drought and during post‐drought recovery.
Forest trees are subjected to multiple stressors during their long lifetime and therefore require effective and finely regulated stress-protective systems. Stressors can induce protective systems ...either directly or with the involvement of stress memory mechanisms. Stress memory has only begun to be uncovered in model plants and is unexplored in coniferous species. Therefore, we studied the possible role of stress memory in the regulation of the accumulation of stress-protective compounds (heat shock proteins, dehydrins, proline) in the needles of naturally grown Scots pine and Norway spruce trees subjected to the subsequent action of long-term (multiyear) and short-term (seasonal) water shortages. Although the water deficit was relatively mild, it significantly influenced the pattern of expression of stress memory-related heat shock factor (HSF) and SWI/SNF genes, indicating the formation of stress memory in both species. In spruce, dehydrin accumulation was increased by water shortage in a manner compatible with Type II stress memory. The accumulation of HSP40 in spruce needles was positively influenced by long-term water shortage, but this increase was unlikely to be of biological importance due to the concomitant decrease in HSP70, HSP90 and HSP101 accumulation. Finally, proline accumulation was negatively influenced by short-term water deficit in spruce. In pine, no one protective compound accumulated in response to water stress. Taken together, the results indicate that the accumulation of stress-protective compounds was generally independent of stress memory effects both in pine and in spruce.
•Mild water deficit influenced the pattern of expression of stress memory genes.•Dehydrin accumulation was consistent to type II stress memory in Norway spruce.•For Scots pine, no memory effects on accumulation of protective compounds were found.
