The Physiological,Biochemical and Molecular Roles of Brassinosteroids and Salicylic Acid in Plant Processes and Salt Tolerance

Plant hormones regulate plant growth and development by affecting an array of cellular, physiological, and developmental processes, including, but not limited to, cell division and elongation, stomatal regulation, photosynthesis, transpiration, ion uptake and transport, initiation of leaf, flower and fruit development, and senescence. Environmental factors such as salinity, drought, and extreme temperatures may cause a reduction in plant growth and productivity by altering the endogenous levels of plant hormones, sensitivity to plant hormones, and/or signaling pathways. Molecular and physiological studies have determined that plant hormones and abiotic stresses have interactive effects on a number of basic biochemical and physiological processes, leading to reduced plant growth and development. Various strategies have been considered or employed to maximize plant growth and productivity under environmental stresses such as salt-stress. A fundamental approach is to develop salt-tolerant plants through genetic means. Breeding for salt tolerance, however, is a long-term endeavor with its own complexities and inherent difficulties. The success of this approach depends, among others, on the availability of genetic sources of tolerance and reliable screening techniques, identification and successful transfer of genetic components of tolerance to desired genetic backgrounds, and development of elite breeding lines and cultivars with salt tolerance and other desirable agricultural characteristics. Such extensive processes have delayed development of successful salt-tolerant cultivars in most crop species. An alternative and technically simpler approach is to induce salt tolerance through exogenous application of certain plant growth–regulating compounds. This approach has gained significant interest during the past decade, when a wealth of new knowledge has become available on the beneficial roles of the six classes of plant hormones (auxins, gibberellins, cytokinins, abscisic acid, ethylene, and brassinosteroids) as well as several other plant growth–regulating substances (jasmonates, salicylates, polyamines, triacontanol, ascorbic acid, and tocopherols) on plant stress tolerance. Among these, brassinosteroids (BRs) and salicylic acid (SA) have been studied most extensively. Both BRs and SA are ubiquitous in the plant kingdom, affecting plant growth and development in many different ways, and are known to improve plant stress tolerance. In this article, we review and discuss the current knowledge and possible applications of BRs and SA that could be used to mitigate the harmful effects of salt-stress in plants. We also discuss the roles of exogenous applications of BRs and SA in the regulation of various biochemical and physiological processes leading to improved salt tolerance in plants.     

Regulation in Plant Stress Tolerance by a Potential Plant Growth Regulator, 5-Aminolevulinic Acid

Exogenous application of different plant growth regulators is a well-recognized strategy to alleviate stress-induced adverse effects on different crop plants by regulating a variety of physiobiochemical processes such as photosynthesis, chlorophyll biosynthesis, nutrient uptake, antioxidant metabolism, and protein synthesis, which are directly or indirectly involved in the mechanism of stress tolerance. Of various environmental factors, salinity, drought, and extreme temperature (low or high) considerably diminish plant growth and yield by modulating endogenous levels as well as signaling pathways of plant hormones. Of various plant hormones/regulators, a potential plant growth regulator, 5-aminolevulinic acid (ALA), is known to be effective in counteracting the injurious effects of various abiotic stresses in plants. Until now the mechanisms behind ALA regulation of growth under stress have not been fully elucidated. It is also not yet clear how far growth and yield in different crops can be promoted by exogenous application of ALA and whether this ALA-induced growth and yield promotion is cost-effective. Thus, in this review we discuss at length the effects of ALA in regulating growth and development in plants under a variety of abiotic stress conditions, including salinity, drought, and temperature stress. Furthermore, advances in the functional and regulatory interactions of this plant growth regulator with plant stress tolerance, as well as the effective mode of exogenous application of ALA in inducing stress tolerance in plants are also comprehensively discussed in this review. In the future, overaccumulation of ALA in plants through manipulation of gene(s) could enhance plant stress tolerance. Thus, genetic manipulation of plants with the goal of attaining increased synthesis/accumulation of ALA and hence improved stress tolerance under stress conditions is an important area for research.     

Brassinosteroid-Mediated Stress Responses

Brassinosteroids (BRs) are a group of naturally occurring plant steroidal compounds with wide-ranging biological activity that offer the unique possibility of increasing crop yields through both changing plant metabolism and protecting plants from environmental stresses. In recent years, genetic and biochemical studies have established an essential role for BRs in plant development, and on this basis BRs have been given the stature of a phytohormone. A remarkable feature of BRs is their potential to increase resistance in plants to a wide spectrum of stresses, such as low and high temperatures, drought, high salt, and pathogen attack. Despite this, only a few studies aimed at understanding the mechanism by which BRs promote stress resistance have been undertaken. Studies of the BR signaling pathway and BR gene-regulating properties indicate that there is cross-talk between BRs and other hormones, including those with established roles in plant defense responses such as abscisic acid, jasmonic acid, and ethylene. Recent studies aimed at understanding how BRs modulate stress responses suggest that complex molecular changes underlie BR-induced stress tolerance in plants. Analyses of these changes should generate exciting results in the future and clarify whether the ability of BRs to increase plant resistance to a range of stresses lies in the complex interactions of BRs with other hormones. Future studies should also elucidate if BRI1, an essential component of the BR receptor, directly participates in stress response signaling through interactions with ligands and proteins involved in plant defense responses.     

植物激素在苔藓生长发育与逆境响应过程中的作用机制研究进展

植物激素是由植物自身代谢产生的一类从产生部位移动到作用部位发挥调控功能的微量小分子有机物质,在植物生长发育、响应环境胁迫过程中起到关键作用.苔藓植物作为早期登陆的非维管植物,处于陆生植物进化早期的阶段,具有许多不同于维管植物的形态和生理特征.大部分苔藓中普遍存在8种主要的植物激素及其衍生物(包括ABA、JA、ET、SA...     

Physiological and molecular mechanisms of brassinosteroid-induced tolerance to high and low temperature in plants

Brassinosteroids (BRs) are plant hormones that were isolated for the first time in the 1970s. This group currently includes more than 70 compounds that differ in their structure and physiological activity. BRs are present in plants in a free form or in the form of conjugates. BRs are known as plant growth regulators, but they also play a role in the plant response to environmental stresses. In the case of plants that are exposed to low/high temperature, exogenous BRs can counteract growth inhibition and reduce biomass losses as well as increase plant survival. BRs show a multidirectional activity in regulating the metabolism of plants exposed to extreme temperatures. The following BRs actions can be distinguished: changes in membrane physicochemical properties, regulation of the expression of selected genes (including stress-responsive genes), as well as indirect effects on metabolism through other hormones or signalling molecules (such as hydrogen peroxide). This review summarizes the current knowledge about the effects of BRs on the physiological and biochemical processes that occur in plants during exposure to low or high temperatures.     

植物对盐胁迫响应的信号转导途径

植物通过调控复杂的信号网络来应对盐胁迫。近年来,随着植物基因工程技术的发展,对植物在盐胁迫下信号转导系统的研究取得了一定进展。本文以拟南芥为代表,对盐胁迫下参与调控植物耐盐生理响应的两大类主要信号转导途径——Ca2＋依赖型信号转导通路和丝裂原活化蛋白激酶（MAPK）级联反应途径的研究进展进行综述,主要介绍参与各信号转导通路的组件及诱发的耐盐生理响应等方面,并对该研究领域存在的问题及今后可能的研究方向进行展望。     

Brassinosteroid-mediated stress tolerance in Arabidopsis shows interactions with abscisic acid,ethylene and salicylic acid pathways

Background  

Brassinosteroids (BRs) play crucial roles in plant development and also promote tolerance to a range of abiotic stresses. Although much has been learned about their roles in plant development, the mechanisms by which BRs control plant stress responses and regulate stress-responsive gene expression are not fully known. Since BR interacts with other plant hormones, it is likely that the stress tolerance conferring ability of BR lies in part in its interactions with other stress hormones.     

植物荫蔽胁迫的激素信号响应

植物的生长发育与光信号密切相关, 外界光强、光质的变化会改变植物的生长发育状态。在自然或人工生态系统中, 植株个体的光环境往往会被其周围植物所影响, 导致荫蔽胁迫, 其主要表现为光合有效辐射以及红光与远红光比值(R:FR)降低。荫蔽胁迫对植物生长发育的多个时期均有影响, 如抑制种子萌发、促进幼苗下胚轴伸长及促进植物花期提前等, 这对农业生产不利, 会导致作物产量以及品质的降低。植物激素是调控植物生长发育的关键内源因子。大量研究表明, 生长素(IAA)、赤霉素(GA)及油菜素甾醇(BR)等植物激素均参与介导植物的荫蔽胁迫响应。当植物处于荫蔽胁迫时, 光信号的改变会影响植物激素的合成及信号转导。不同植物激素对荫蔽胁迫的响应各不相同, 但其信号通路之间却存在互作关系, 从而形成复杂的网络状调控路径。该文总结了几种主要植物激素(生长素、赤霉素、油菜素甾醇及乙烯)响应荫蔽胁迫的机理, 重点论述了荫蔽胁迫对植物激素合成及信号通路的影响, 以及植物激素调控荫蔽胁迫下植物生长的分子机理, 并对未来潜在的研究热点进行了分析。     

Brassinosteroid signal transduction--choices of signals and receptors

Small signaling molecules that mediate cell-cell communication are essential for developmental regulation in multicellular organisms. Among them are the steroids and peptide hormones that regulate growth in both plants and animals. In plants, brassinosteroids (BRs) are perceived by the cell surface receptor kinase BRI1, which is distinct from the animal steroid receptors. Identification of components of the BR signaling pathway has revealed similarities to other animal and plant signal transduction pathways. Recent studies demonstrated that tomato BRI1 (tBRI1) perceives both BR and the peptide hormone systemin, raising new questions about the molecular mechanism and evolution of receptor-ligand specificity.     

Induction of Abiotic Stress Tolerance by Salicylic Acid Signaling

The role of salicylic acid (SA) as a key molecule in the signal transduction pathway of biotic stress responses has already been well described. Recent studies indicate that it also participates in the signaling of abiotic stresses. The application of exogenous SA could provide protection against several types of stresses such as high or low temperature, heavy metals, and so on. Although SA may also cause oxidative stress to plants, partially through the accumulation of hydrogen peroxide, the results published so far show that the preliminary treatment of plants with low concentrations of SA might have an acclimation-like effect, causing enhanced tolerance toward most kinds of abiotic stresses due primarily to enhanced antioxidative capacity. The effect of exogenous SA depends on numerous factors such as the species and developmental stage of the plant, the mode of application, and the concentration of SA and its endogenous level in the given plant. Recent results show that not only does exogenous SA application moderate stress effects, but abiotic stress factors may also alter the endogenous SA levels in the plant cells. This review compares the roles of SA during different abiotic stresses.     

生长素与油菜素甾醇相互作用机制的研究进展

植物生长发育是一个复杂、精细的调控过程,涉及细胞、组织和器官间多层次的信息交流,激素在其间发挥了重要调控作用．生长素和油菜素甾醇（BR）都能促进植物伸长,随着对其作用机制研究的深入,人们发现它们协同调控很多生理过程,对二者作用机制和信号转导的相互作用研究也成为激素研究领域的热点之一．对生长素和BR转导途径的揭示及直接下游基因的大规模发掘为二者通过相互作用调控不同生理过程的机制研究提供了重要线索．生长素和BR的相互作用涉及到下游基因转录的调控、信号组分互作以及合成代谢和极性运输等多层次的调控．     

Identification and Characterization of Brassinosteroid Biosynthesis and Signaling Pathway Genes in Solanum tuberosum

Russian Journal of Plant Physiology - Brassinosteroids (BRs) are plant hormones which promote plant growth and development. Their biosynthetic pathway and signal transduction pathway have been well...     

Cytokinins: metabolism and function in plant adaptation to environmental stresses

In plants, the cytokinin (CK) phytohormones regulate numerous biological processes, including responses to environmental stresses, via a complex network of CK signaling. By an unknown mechanism, stress signals are perceived and transmitted through the His-Asp phosphorelay, an important component of the CK signal transduction pathway, triggering CK-responsive genes. Because of the intensive crosstalk between CKs and abscisic acid (ABA), modulation of CK levels and their signal transduction affects both ABA-dependent and ABA-independent pathways, enabling plant adaptation to adverse conditions. This review presents our current understanding of the functions of CKs and CK signaling in the regulation of plant adaptation to stress. Biotechnological strategies based on the modulation of CK levels have been examined with the aim of stabilizing agriculture yields.     

Regulation of Arabidopsis thaliana Ku genes at different developmental stages under heat stress

Ku, a heterodimeric protein consisting of 70- and 80-kDa subunits, is involved in many cellular processes, such as DNA replication, cell cycle regulation and heat shock response. Moreover, the expression of Arabidopsis thaliana Ku genes (AtKu) is modulated by certain plant hormones through several signal transduction pathways. This study investigated how AtKu are regulated by heat stress. AtKu expression in 3-week-old young seedlings was down-regulated by heat stress in a time-dependent manner, as examined using real-time quantitative PCR, GUS reporter systems, and western blotting analysis. Additionally, the heat-induced repression of AtKu was mediated through the abscisic acid (ABA) biosynthetic pathway, as shown by the reversal of AtKu suppression in the ABA biosynthesis mutant, aba3, and by an increase in the ABA level as analyzed by reverse-phase high performance liquid chromatography. Heat stress-induced regulation of AtKu repression also involved ethylene signaling, DNA repair pathways, and fatty acid synthesis. Furthermore, AtKu expression was repressed in stems, rosette leaves, and cauline leaves in 4-5-week-old plants under heat stress, whereas it remained unchanged in roots and primary inflorescence, indicating that heat differentially modulated AtKu expression in distinct tissues of Arabidopsis.     

Phospholipase D is a negative regulator of proline biosynthesis in Arabidopsis thaliana

Accumulation of proline has been observed in a large number of plant species in response to drought and salt stresses, suggesting a key role of this amino acid in plant stress adaptation. Upstream components of the proline biosynthesis signal transduction pathways are still poorly defined. We provide experimental evidence that phospholipase D (PLD) is involved in the regulation of proline metabolism in Arabidopsis thaliana. The application of primary butyl alcohols, which divert part of PLD-derived phosphatidic acid by transphosphatidylation, stimulated proline biosynthesis even without hyperosmotic constraints. Moreover, application of primary butyl alcohols enhanced the proline responsiveness of seedlings to mild hyperosmotic stress. These data indicate that some PLDs are negative regulators of proline biosynthesis and that plants present a higher proline responsiveness to hyperosmotic stress when this regulator is abolished. We clearly demonstrate that PLD signaling for proline biosynthesis is similar to RD29A gene expression and different from the abscisic acid-dependent RAB18 gene expression. Our data reveal that PLDs play positive and negative roles in hyperosmotic stress signal transduction in plants, contributing to a precise regulation of ion homeostasis and plant salt tolerance.     

Role of salicylic acid on physiological and biochemical mechanism of salinity stress tolerance in plants

Salinity stress is one of the major abiotic stresses affecting plant growth and productivity globally. In order to improve the yields of plants growing under salt stress bear remarkable importance to supply sustainable agriculture. Acclimation of plants to salinized condition depends upon activation of cascade of molecular network involved in stress sensing/perception, signal transduction, and the expression of specific stress-related genes and metabolites. Isolation of salt overly sensitive (SOS) genes by sos mutants shed us light on the relationship between ion homeostasis and salinity tolerance. Regulation of antioxidative system to maintain a balance between the overproduction of reactive oxygen species and their scavenging to keep them at signaling level for reinstating metabolic activity has been elucidated. However, osmotic adaptation and metabolic homeostasis under abiotic stress environment is required. Recently, role of phytohormones like Abscisic acid, Jasmonic acid, and Salicylic acid in the regulation of metabolic network under osmotic stress condition has emerged through crosstalk between chemical signaling pathways. Thus, abiotic stress signaling and metabolic balance is an important area with respect to increase crop yield under suboptimal conditions. This review focuses on recent developments on improvement in salinity tolerance aiming to contribute sustainable plant yield under saline conditions in the face of climate change.