Cysteine-rich receptor-like kinases (CRKs) are transmembrane proteins characterized by the presence of two domains of unknown function 26 (DUF26) in their ectodomain. The CRKs form one of the largest groups of receptor-like protein kinases in plants, but their biological functions have so far remained largely uncharacterized. We conducted a large-scale phenotyping approach of a nearly complete crk T-DNA insertion line collection showing that CRKs control important aspects of plant development and stress adaptation in response to biotic and abiotic stimuli in a non-redundant fashion. In particular, the analysis of reactive oxygen species (ROS)-related stress responses, such as regulation of the stomatal aperture, suggests that CRKs participate in ROS/redox signalling and sensing. CRKs play general and fine-tuning roles in the regulation of stomatal closure induced by microbial and abiotic cues. Despite their great number and high similarity, large-scale phenotyping identified specific functions in diverse processes for many CRKs and indicated that CRK2 and CRK5 play predominant roles in growth regulation and stress adaptation, respectively. As a whole, the CRKs contribute to specificity in ROS signalling. Individual CRKs control distinct responses in an antagonistic fashion suggesting future potential for using CRKs in genetic approaches to improve plant performance and stress tolerance. Receptor-like kinases (RLKs) are important regulators in signal transduction in plants. However, the large number of RLKs and their high sequence similarity has hampered the analysis of RLKs. One of the largest subgroups of RLKs, the cysteine-rich receptor-like kinases (CRKs), has been suggested to be involved in mediating the effects of reactive oxygen species (ROS). While ROS are recognized as important signalling elements with a large variety of roles in plants, their ligands and achievement of signalling specificity remain unknown. Using insertion mutants we analysed the roles of CRKs in plant development and stress responses and show that CRKs have important roles as mediators of signalling specificity during regulation of stomatal aperture. Our study shows that, despite their large number and high sequence conservation, individual CRKs have intriguingly distinct functions in different aspects of plant life. This makes the CRKs promising candidates for future studies of their biochemical function.

This review confronts the classical view of plant immune defence and light acclimation with recently published data. Earlier findings have linked plant immune defences to nucleotide‐binding site leucine‐rich repeat (NBS‐LRR)‐dependent recognition of pathogen effectors and to the role of plasma membrane‐localized NADPH‐dependent oxidoreductase (AtRbohD), reactive oxygen species (ROS) and salicylic acid (SA). However, recent results suggest that plant immune defence also depends on the absorption of excessive light energy and photorespiration. Rapid changes in light intensity and quality often cause the absorption of energy, which is in excess of that required for photosynthesis. Such excessive light energy is considered to be a factor triggering photoinhibition and disturbance in ROS/hormonal homeostasis, which leads to cell death in foliar tissues. We highlight here the tight crosstalk between ROS‐ and SA‐dependent pathways leading to light acclimation, and defence responses leading to pathogen resistance. We also show that LESION SIMULATING DISEASE 1 (LSD1) regulates and integrates these processes. Moreover, we discuss the role of plastid–nucleus signal transduction, photorespiration, photoelectrochemical signalling and ‘light memory’ in the regulation of acclimation and immune defence responses. All of these results suggest that plants have evolved a genetic system that simultaneously regulates systemic acquired resistance (SAR), cell death and systemic acquired acclimation (SAA).

In plants, receptor-like protein kinases play essential roles in signal transduction by recognizing extracellular stimuli and activating the downstream signalling pathways. Cysteine-rich receptor-like kinases (CRKs) constitute a large subfamily of receptor-like protein kinases, with 44 members in Arabidopsis thaliana. They are distinguished by the novel C-X8-C-X2-C motif (DUF26) in the extracellular domains. One of them, CRK5, is an important component of the biochemical machinery involved in the regulation of essential physiological processes. Functional characterization of crk5 mutant plants showed their clear phenotype, manifested by impaired stomatal conductance and accelerated senescence. This phenotype correlated with accumulation of reactive oxygen species, higher foliar levels of ethylene and salicylic acid, and increased transcript abundance for genes associated with signalling pathways corresponding to these hormones. Moreover, the crk5 plants displayed enhanced cell death and oxidative damage in response to ultraviolet radiation. Complementation of CRK5 mutation managed to recover the wild-type phenotype, indicating an essential role of this gene in the regulation of growth, development, and acclimatory responses.

In the natural environment, plants are simultaneously exposed to multivariable abiotic and biotic stresses. Typical abiotic stresses are changes in temperature, light intensity and quality, water stress (drought, flood), microelements availability, salinity, air pollutants, and others. Biotic stresses are caused by other organisms, such as pathogenic bacteria and viruses or parasites. This review presents the current state-of-the-art knowledge on programmed cell death in the cross-tolerance phenomena and its conditional molecular and physiological regulators, which simultaneously regulate plant acclimation, defense, and developmental responses. It highlights the role of the absorbed energy in excess and its dissipation as heat in the induction of the chloroplast retrograde phytohormonal, electrical, and reactive oxygen species signaling. It also discusses how systemic- and network-acquired acclimation and acquired systemic resistance are mutually regulated and demonstrates the role of non-photochemical quenching and the dissipation of absorbed energy in excess as heat in the cross-tolerance phenomenon. Finally, new evidence that plants evolved one molecular system to regulate cell death, acclimation, and cross-tolerance are presented and discussed.

In Arabidopsis thaliana, cysteine-rich receptor-like kinases (CRKs) constitute a large group of membrane-localized proteins which perceive external stimuli and transduce the signal into the cell. Previous reports based on their loss-of-function phenotypes and expression profile support their role in many developmental and stress-responsive pathways. Our study revealed that one member of this family, CRK5, acts as a negative regulator of leaf aging. Enrichment of the CRK5 promoter region in W-box cis-elements demonstrated that WRKY transcription factors control it. We observed significantly enhanced WRKY53 expression in crk5 and reversion of its early-senescence phenotype in the crk5 wrky53 line, suggesting a negative feedback loop between these proteins antagonistically regulating chlorophyll a and b contents. Yeast-two hybrid assay showed further that CRK5 interacts with several proteins involved in response to water deprivation or calcium signaling, while gas exchange analysis revealed a positive effect of CRK5 on water use efficiency. Consistent with that, the crk5 plants showed disturbed foliar temperature, stomatal conductance, transpiration, and increased susceptibility to osmotic stress. These traits were fully or partially reverted to wild-type phenotype in crk5 wrky53 double mutant. Obtained results suggest that WRKY53 and CRK5 are antagonistic regulators of chlorophyll synthesis/degradation, senescence, and stomatal conductance.