Crosstalk of the Brassinosteroid Signalosome with Phytohormonal and Stress Signaling Components Maintains a Balance between the Processes of Growth and Stress Tolerance

Brassinosteroids (BRs) are a class of phytohormones, which regulate various processes during plant life cycle. Intensive studies conducted with genetic, physiological and molecular approaches allowed identification of various components participating in the BR signaling—from the ligand perception, through cytoplasmic signal transduction, up to the BR-dependent gene expression, which is regulated by transcription factors and chromatin modifying enzymes. The identification of new components of the BR signaling is an ongoing process, however an emerging view of the BR signalosome indicates that this process is interconnected at various stages with other metabolic pathways. The signaling crosstalk is mediated by the BR signaling proteins, which function as components of the transmembrane BR receptor, by a cytoplasmic kinase playing a role of the major negative regulator of the BR signaling, and by the transcription factors, which regulate the BR-dependent gene expression and form a complicated regulatory system. This molecular network of interdependencies allows a balance in homeostasis of various phytohormones to be maintained. Moreover, the components of the BR signalosome interact with factors regulating plant reactions to environmental cues and stress conditions. This intricate network of interactions enables a rapid adaptation of plant metabolism to constantly changing environmental conditions

Exploring the Brassinosteroid Signaling in Monocots Reveals Novel Components of the Pathway and Implications for Plant Breeding

Brassinosteroids (BRs) are a class of steroidal phytohormones which are key regulators of diverse processes during whole life cycle of plants. Studies conducted in the dicot model species Arabidopsis thaliana have allowed identification and characterization of various components of the BR signaling. It is currently known that the BR signaling is interconnected at various stages with other phytohormonal and stress signaling pathways. It enables a rapid and e cient adaptation of plant metabolism to constantly changing environmental conditions. However, our knowledge about mechanism of the BR signaling in the monocot species is rather limited. Thus, identification of new components of the BR signaling in monocots, including cereals, is an ongoing process and has already led to identification of some monocot-specific components of the BR signaling. It is of great importance as disturbances in the BR signaling influence architecture of mutant plants, and as a consequence, the reaction to environmental conditions. Currently, the modulation of the BR signaling is considered as a target to enhance yield and stress tolerance in cereals, which is of particular importance in the face of global climate change

The role of strigolactones in nutrient-stress responses in plants

Strigolactones (SLs) are a new group of plant hormones, which have been intensively investigated during the last few years. The wide spectrum of SLs actions, including the regulation of shoot/root architecture, and the stimulation of the interactions between roots and fungi or bacteria, as well as the stimulation of germination of parasitic plants, indicates that this group of hormones may play an important role in the mechanisms that control soil exploration, and the root-mediated uptake of nutrients. Current studies have shown that SLs might be factors that have an influence on the plant response to a deficiency of macronutrients. Experimental data from the last four years have confirmed that the biosynthesis and exudation of SLs are increased under phosphorus and nitrogen deficiency. All these data suggest that SLs may regulate the complex response to nutrient stress, which include not only the modification of the plant developmental process, but also the cooperation with other organisms in order to minimize the effects of threats. In this paper the results of studies that indicate that SLs play an important role in the response to nutrient stress are reviewed and the consequences of the higher biosynthesis and exudation of SLs in response to phosphorus and nitrogen deficiency are discussed

Comprehensive Overview of the Brassinosteroid Biosynthesis Pathways: Substrates, Products, Inhibitors, and Connections

Brassinosteroids (BRs) as a class of steroid plant hormones participate in the regulation of numerous developmental processes, including root and shoot growth, vascular differentiation, fertility, flowering, and seed germination, as well as in responding to environmental stresses. During four decades of research, the BR biosynthetic pathways have been well studied with forward- and reverse genetics approaches. The free BRs contain 27, 28, and 29 carbons within their skeletal structure: (1): 5a-cholestane or 26- nor-24a-methyl-5a-cholestane for C27-BRs; (2) 24a-methyl-5a-cholestane, 24b-methyl- 5a-cholestane or 24-methylene-5a-cholestane for C28-BRs; (3) 24a-ethyl-5acholestane, 24(Z)-ethylidene-5a-cholestane, 25-methyl-5a-campestane or 24- methylene-25-methyl-5a-cholestane for C29-BRs, as well as different kinds and orientations of oxygenated functions in A- and B-ring. These alkyl substituents are also common structural features of sterols. BRs are derived from sterols carrying the same side chain. The C27-BRs without substituent at C-24 are biosynthesized from cholesterol. The C28-BRs carrying either an a-methyl, b-methyl, or methylene group are derived from campesterol, 24-epicampesterol or 24-methylenecholesterol, respectively. The C29-BRs with an a-ethyl group are produced from sitosterol. Furthermore, the C29 BRs carrying methylene at C-24 and an additional methyl group at C-25 are derived from 24- methylene-25-methylcholesterol. Generally, BRs are biosynthesized via cycloartenol and cycloartanol dependent pathways. Till now, more than 17 compounds were characterized as inhibitors of the BR biosynthesis. For nine of the inhibitors (e.g., brassinazole and YCZ-18) a specific target reaction within the BR biosynthetic pathway has been identified. Therefore, the review highlights comprehensively recent advances in our understanding of the BR biosynthesis, sterol precursors, and dependencies between the C27-C28 and C28-C29 pathways

The barley EST DNA Replication and Repair Database (bEST-DRRD) as a tool for the identification of the genes involved in DNA replication and repair

Background: The high level of conservation of genes that regulate DNA replication and repair indicates that they may serve as a source of information on the origin and evolution of the species and makes them a reliable system for the identification of cross-species homologs. Studies that had been conducted to date shed light on the processes of DNA replication and repair in bacteria, yeast and mammals. However, there is still much to be learned about the process of DNA damage repair in plants.Description: These studies, which were conducted mainly using bioinformatics tools, enabled the list of genes that participate in various pathways of DNA repair in Arabidopsis thaliana (L.) Heynh to be outlined; however, information regarding these mechanisms in crop plants is still very limited. A similar, functional approach is particularly difficult for a species whose complete genomic sequences are still unavailable. One of the solutions is to apply ESTs (Expressed Sequence Tags) as the basis for gene identification. For the construction of the barley EST DNA Replication and Repair Database (bEST-DRRD), presented here, the Arabidopsis nucleotide and protein sequences involved in DNA replication and repair were used to browse for and retrieve the deposited sequences, derived from four barley (Hordeum vulgare L.) sequence databases, including the " Barley Genome version 0.05" database (encompassing ca. 90% of barley coding sequences) and from two databases covering the complete genomes of two monocot models: Oryza sativa L. and Brachypodium distachyon L. in order to identify homologous genes. Sequences of the categorised Arabidopsis queries are used for browsing the repositories, which are located on the ViroBLAST platform. The bEST-DRRD is currently used in our project during the identification and validation of the barley genes involved in DNA repair.Conclusions: The presented database provides information about the Arabidopsis genes involved in DNA replication and repair, their expression patterns and models of protein interactions. It was designed and established to provide an open-access tool for the identification of monocot homologs of known Arabidopsis genes that are responsible for DNA-related processes. The barley genes identified in the project are currently being analysed to validate their function

The Impact of Mutations in the HvCPD and HvBRI1 Genes on the Physicochemical Properties of the Membranes from Barley Acclimated to Low/High Temperatures

1) Background: The study characterized barley mutants with brassinosteroid (BR) biosynthesis and signaling disturbances in terms of the physicochemical/structural properties of membranes to enrich the knowledge about the role of brassinosteroids for lipid metabolism and membrane functioning. (2) Methods: The Langmuir method was used to investigate the properties of the physicochemical membranes. Langmuir monolayers were formed from the lipid fractions isolated from the plants growing at 20 C and then acclimated at 5 C or 27 C. The fatty acid composition of the lipids was estimated using gas chromatography. (3) Results: The BR-biosynthesis and BR-signaling mutants of barley were characterized by a temperature-dependent altered molar percentage of fatty acids (from 14:0 to 20:1) in their galactolipid and phospholipid fractions in comparison to wild-type (WT). For example, the mutants had a lower molar percentage of 18:3 in the phospholipid (PL) fraction. The same regularity was observed at 5 C. It resulted in altered physicochemical parameters of the membranes (Alim, coll, Cs1). (4) Conclusions: BR may be involved in regulating fatty acid biosynthesis or their transport/incorporation into the cell membranes. Mutants had altered physicochemical parameters of their membranes, compared to the WT, which suggests that BR may have a multidirectional impact on the membrane-dependent physiological processes

HSP Transcript and Protein Accumulation in Brassinosteroid Barley Mutants Acclimated to Low and High Temperatures

In temperature stress, the main role of heat-shock proteins (HSP) is to act as molecular chaperones for other cellular proteins. However, knowledge about the hormonal regulation of the production of the HSP is quite limited. Specifically, little is known about the role of the plant steroid hormones—brassinosteroids (BR)—in regulating the HSP expression. The aim of our study was to answer the question of how a BR deficit or disturbances in its signaling affect the accumulation of the HSP90, HSP70, HSP18, and HSP17 transcripts and protein in barley growing at 20C (control) and during the acclimation of plants at 5 C and 27 C. In barley, the temperature of plant growth modified the expression of HSPs. Furthermore, the BR-deficient mutants (mutations in the HvDWARF or HvCPD genes) and BR-signaling mutants (mutation in the HvBRI1 gene) were characterized by altered levels of the transcripts and proteins of the HSP group compared to the wild type. The BR-signaling mutant was characterized by a decreased level of the HSP transcripts and heat-shock proteins. In the BR-deficient mutants, there were temperature-dependent cases when the decreased accumulation of the HSP70 and HSP90 transcripts was connected to an increased accumulation of these HSP. The significance of changes in the accumulation of HSPs during acclimation at 27 C and 5 C is discussed in the context of the altered tolerance to more extreme temperatures of the studied mutants (i.e., heat stress and frost, respectively)

Role of strigolactones : Signalling and crosstalk with other phytohormones

Plant hormones play important roles in controlling how plants grow and develop. While metabolism provides the energy needed for plant survival, hormones regulate the pace of plant growth. Strigolactones (SLs) were recently defined as new phytohormones that regulate plant metabolism and, in turn, plant growth and development. This group of phytohormones is derived from carotenoids and has been implicated in a wide range of physiological functions including regulation of plant architecture (inhibition of bud outgrowth and shoot branching), photomorphogenesis, seed germination, nodulation, and physiological reactions to abiotic factors. SLs also induce hyphal branching in germinating spores of arbuscular mycorrhizal fungi (AMF), a process that is important for initiating the connection between host plant roots and AMF. This review outlines the physiological roles of SLs and discusses the significance of interactions between SLs and other phytohormones to plant metabolic responses

Disturbances in the Biosynthesis or Signalling of Brassinosteroids That Are Caused by Mutations in the HvDWARF, HvCPD and HvBRI1 Genes Increase the Tolerance of Barley to the Deacclimation Process

Tolerance to deacclimation is an important physiological feature in plants in the face of global warming, which is resulting in incidents of increases in winter temperatures. The aim of the work was to explore how disturbances in the signalling and synthesis of brassinosteroids (BR) influence the deacclimation tolerance of barley. One group of mutants and their reference cultivars (Bowman and Delisa) was cold-acclimated, deacclimated and then tested for frost tolerance at − 12 °C. After cold acclimation, the second group of plants was additionally exposed to frost (− 6 °C) and then, deacclimated and tested for frost tolerance at − 12 °C. The deacclimated brassinosteroid mutants were characterised by an increased tolerance to frost, and consequently, had a higher tolerance to deacclimation than their wild-type cultivars. The mechanism of this phenomenon may be partly explained by analysing the hormonal homeostasis in the crowns. For all of the tested plants, a characteristic feature of the response to the deacclimation phase was an increase in the growth-promoting hormones and abscisic acid compared to the cold acclimation phase. The increase was greater in the BR-deficient (BW084) and BR-insensitive (BW312) mutants compared to the Bowman reference cultivar. Mutant 522DK was characterised by a lower accumulation of total cytokinins and gibberellins as well as an enhanced auxin deactivation compared to the Delisa. In the second group, when the plants were exposed to a temperature of − 6 °C before deacclimation, the hormonal homeostasis was further altered in both the mutants and reference cultivars, but all of the mutants had a higher frost tolerance than the wild types

Mutations in the HvDWARF, HvCPD and HvBRI1 Genes-Involved in Brassinosteroid Biosynthesis/Signalling : Altered Photosynthetic Efficiency, Hormonal Homeostasis and Tolerance to High/Low Temperatures in Barley

Brassinosteroids (BR) are steroid phytohormones that are involved in the growth and stress response in plants, but the precise mechanisms of their action are still being discovered. In our study we have used BR-deficient barley mutants 522DK and BW084 (which carry missense mutations in the HvDWARF and HvCPD genes, respectively). We have also used a BR-signalling mutant that harbors missense substitutions in the HvBRI1 gene. Our aim was (1) to find out if the content of phytohormones in the mutants grown at 20 °C is different than in the wild types and whether/how the content of phytohormones changes after plant acclimation at temperatures of 5 °C and 27 °C?, (2) to characterise the effectiveness of the light reactions of photosynthesis of the barley mutants in comparison to wild types at various temperatures, and (3) to verify the impact of mutations on the tolerance of barley to high and low temperatures. Hormonal characteristics of the BR mutants of barley show the complexity of the interactions between BR and other plant hormones that are additionally modified by temperature and possibly by other factors. The results suggest the participation of BR in auxin catabolism. Further, BR appears to play a role in maintaining the ABA–ABAGlc balance. As for the gibberellin content in plants at a temperature of 20 °C, more in-depth studies will be required to explain the contradictory effects regarding the accumulation of GA3, GA4 and GA5, which appears to be dependent on the type of mutation and connected to the BR level. A fast-kinetic chlorophyll a fluorescence analysis has revealed that the mutants had lower values of energy absorption than the wild types, but the values of the energy transferred via the electron-transport chain was maintained at the wild-type level. We presumed that BR are involved in regulating plant acclimation to extreme (low/high) temperatures, thus the BR-deficient and BR-signalling mutants should be less tolerant to low/high temperatures when compared to the wild types. Unexpectedly, all of the mutants showed a higher tolerance to high temperatures than the wild types. The BW084 and BW312 mutants were less tolerant to frost than the wild type, but 522DK had a similar frost tolerance as the reference wild-type cultivar