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Abstract Light triggers an enhancement of global translation during photomorphogenesis in Arabidopsis, but little is known about the underlying mechanisms. The phosphorylation of the α-subunit of eukaryotic initiation factor 2 (eIF2α) at a conserved serine residue in the N-terminus has been shown as an important mechanism for the regulation of protein synthesis in mammalian and yeast cells. However, whether the phosphorylation of this residue in plant eIF2α plays a role in regulation of translation remains elusive. Here, we show that the quadruple mutant of SUPPRESSOR OF PHYA-105 family members (SPA1-SPA4) display repressed translation efficiency after light illumination. Moreover, SPA1 directly phosphorylates the eIF2α C-terminus under light conditions. The C-term-phosphorylated eIF2α promotes translation efficiency and photomorphogenesis, whereas the C-term-unphosphorylated eIF2α results in a decreased translation efficiency. We also demonstrate that the phosphorylated eIF2α enhances ternary complex assembly by promoting its affinity to eIF2β and eIF2γ. This study reveals a unique mechanism by which light promotes translation via SPA1-mediated phosphorylation of the C-terminus of eIF2α in plants. 

Abstract Light serves as a pivotal environmental cue regulating various aspects of plant growth and development, including seed germination, seedling de-etiolation, and shade avoidance. Within this regulatory framework, the basic helix–loop–helix transcription factors known as phytochrome-interacting factors (PIFs) play an essential role in orchestrating responses to light stimuli. Phytochromes, acting as red/far-red light receptors, initiate a cascade of events leading to the degradation of PIFs (except PIF7), thereby triggering transcriptional reprogramming to facilitate photomorphogenesis. Recent research has unveiled multiple post-translational modifications that regulate the abundance and/or activity of PIFs, including phosphorylation, dephosphorylation, ubiquitination, deubiquitination, and SUMOylation. Moreover, intriguing findings indicate that PIFs can influence chromatin modifications. These include modulation of histone 3 lysine 9 acetylation (H3K9ac), as well as occupancy of histone variants such as H2A.Z (associated with gene repression) and H3.3 (associated with gene activation), thereby intricately regulating downstream gene expression in response to environmental cues. This review summarizes recent advances in understanding the role of PIFs in regulating various signaling pathways, with a major focus on photomorphogenesis. 

Abstract In addition to providing the radiant energy that drives photosynthesis, sunlight carries signals that enable plants to grow, develop and adapt optimally to the prevailing environment. Here we trace the path of research that has led to our current understanding of the cellular and molecular mechanisms underlying the plant's capacity to perceive and transduce these signals into appropriate growth and developmental responses. Because a fully comprehensive review was not possible, we have restricted our coverage to the phytochrome and cryptochrome classes of photosensory receptors, while recognizing that the phototropin and UV classes also contribute importantly to the full scope of light-signal monitoring by the plant. 

Abstract The phytochrome (phy) family of sensory photoreceptors modulates developmental programs in response to ambient light. Phys also control gene expression in part by directly interacting with the bHLH class of transcription factors, PHYTOCHROME-INTERACTING FACTORS (PIFs), and inducing their rapid phosphorylation and degradation. Several kinases have been shown to phosphorylate PIFs and promote their degradation. However, the phosphatases that dephosphorylate PIFs are less understood. In this study, we describe 4 regulatory subunits of the Arabidopsis (Arabidopsis thaliana) protein PHOSPHATASE 2A (PP2A) family (B′α, B′β, B″α, and B″β) that interact with PIF3 in yeast 2-hybrid, in vitro and in vivo assays. The pp2ab″αβ and b″αβ/b′αβ mutants display short hypocotyls, while the overexpression of the B subunits induces longer hypocotyls compared with the wild type (WT) under red light. The light-induced degradation of PIF3 is faster in the b″αβ/b′αβ quadruple mutant compared with that in the WT. Consistently, immunoprecipitated PP2A A and B subunits directly dephosphorylate PIF3-MYC in vitro. An RNA-sequencing analysis shows that B″α and B″β alter global gene expression in response to red light. PIFs (PIF1, PIF3, PIF4, and PIF5) are epistatic to these B subunits in regulating hypocotyl elongation under red light. Collectively, these data show an essential function of PP2A in dephosphorylating PIF3 to modulate photomorphogenesis in Arabidopsis. 

Abstract Glucosinolates (GSLs) are defensive secondary metabolites produced by Brassicaceae species in response to abiotic and biotic stresses. The biosynthesis of GSL compounds and the expression of GSL-related genes are highly modulated by endogenous signals (i.e. circadian clocks) and environmental cues, such as temperature, light, and pathogens. However, the detailed mechanism by which light signaling influences GSL metabolism remains poorly understood. In this study, we found that a light-signaling factor, ELONGATED HYPOCOTYL 5 (HY5), was involved in the regulation of GSL content under light conditions in Arabidopsis (Arabidopsis thaliana). In hy5-215 mutants, the transcript levels of GSL pathway genes were substantially upregulated compared with those in wild-type (WT) plants. The content of GSL compounds was also substantially increased in hy5-215 mutants, whereas 35S::HY5-GFP/hy5-215 transgenic lines exhibited comparable levels of GSL-related transcripts and GSL content to those in WT plants. HY5 physically interacts with HISTONE DEACETYLASE9 and binds to the proximal promoter region of MYB29 and IMD1 to suppress aliphatic GSL biosynthetic processes. These results demonstrate that HY5 suppresses GSL accumulation during the daytime, thus properly modulating GSL content daily in Arabidopsis plants. 

SUMMARY As sessile organisms, plants encounter dynamic and challenging environments daily, including abiotic/biotic stresses. The regulation of carbon and nitrogen allocations for the synthesis of plant proteins, carbohydrates, and lipids is fundamental for plant growth and adaption to its surroundings. Light, one of the essential environmental signals, exerts a substantial impact on plant metabolism and resource partitioning (i.e., starch). However, it is not fully understood how light signaling affects carbohydrate production and allocation in plant growth and development. An orphan gene unique toArabidopsis thaliana, namedQUA‐QUINE STARCH(QQS) is involved in the metabolic processes for partitioning of carbon and nitrogen among proteins and carbohydrates, thus influencing leaf, seed composition, and plant defense in Arabidopsis. In this study, we show that PHYTOCHROME‐INTERACTING bHLH TRANSCRIPTION FACTORS (PIFs), including PIF4, are required to suppressQQSduring the period at dawn, thus preventing overconsumption of starch reserves.QQSexpression is significantly de‐repressed inpif4andpifQ, while repressed by overexpression ofPIF4, suggesting that PIF4 and its close homologs (PIF1, PIF3, and PIF5) act as negative regulators ofQQSexpression. In addition, we show that the evening complex, including ELF3 is required for active expression ofQQS, thus playing a positive role in starch catabolism during night‐time. Furthermore,QQSis epigenetically suppressed by DNA methylation machinery, whereas histone H3 K4 methyltransferases (e.g., ATX1, ATX2, and ATXR7) and H3 acetyltransferases (e.g., HAC1 and HAC5) are involved in the expression ofQQS. This study demonstrates that PIF light signaling factors help plants utilize optimal amounts of starch during the night and prevent overconsumption of starch before its biosynthesis during the upcoming day. 

Abstract Plants respond to high ambient temperature by implementing a suite of morphological changes collectively termed thermomorphogenesis. Here we show that the above and below ground tissue-response to high ambient temperature are mediated by distinct transcription factors. While the central hub transcription factor, PHYTOCHROME INTERCTING FACTOR 4 (PIF4) regulates the above ground tissue response, the below ground root elongation is primarily regulated by ELONGATED HYPOCOTYL 5 (HY5). Plants respond to high temperature by largely expressing distinct sets of genes in a tissue-specific manner. HY5 promotes root thermomorphogenesis via directly controlling the expression of many genes including the auxin and BR pathway genes. Strikingly, the above and below ground thermomorphogenesis is impaired inspaQ. Because SPA1 directly phosphorylates PIF4 and HY5, SPAs might control the stability of PIF4 and HY5 to regulate thermomorphogenesis in both tissues. These data collectively suggest that plants employ distinct combination of SPA-PIF4-HY5 module to regulate tissue-specific thermomorphogenesis. 

Plants have evolved with complex sensory systems to recognize signals from multiple environmental conditions. A light signal is one of the most important environmental factors that regulates not only photomorphogenesis but also the developmental strategy of plants throughout their life cycle. The molecular mechanisms of the light signaling modules and the interactions between light and other environmental signals have been studied extensively. However, to enhance plant growth, particularly in crop production, we need to gain a deeper understanding of how light regulates plant development within gene regulatory networks (GRNs). Understanding GRNs is important to identify not only the novel genes and transcription factors in light signaling pathways but also the factors that connect light signaling and other environmental signals. Weighted gene co-expression network analysis (WGCNA) has been used to study GRN. We applied WGCNA to 58 RNA-seq samples of wild-type Arabidopsis grown under different light treatments and built the gene co-expression networks. We identified 14 different modules that are significantly associated with different light treatments. Among them, the honeydew1 and ivory display significant association with the dark-grown seedlings. Many hub genes identified from these modules are significantly enriched in light responses, including responses to red, far-red, blue light, light stimulus, auxin responses, and photosynthesis. Although we found many known transcription factors in these modules, we also identified several unknown genes and transcription factors that are significantly associated with the honeydew1 module and highly differentially expressed between dark and light conditions. To examine whether the hub genes in the honeydew1 module play a role in light signaling, we isolated mutants in selected hub genes and measured hypocotyl lengths under dark, red, and far-red light conditions. These assays showed that four hub genes are involved in regulating light signaling pathways. This study provides a new approach to identifying novel genes in GRNs underlying light responses in Arabidopsis. 

Plant phytochromes are well-studied photoreceptors that sense red and far-red light, regulating photomorpho- genic development. Molecular signaling mechanisms of phytochrome A (phyA) and phyB largely overlap, especially in regulation of PHYTOCHROME-INTERACTING FACTORs (PIFs) and E3 ligase complexes composed of CONSTITUTIVELY PHOTOMORPHOGENIC 1 (COP1) and SUPPRESSORs OF phyA-105 (SPAs). However, the differences in their molecular signaling mechanisms remain unclear. Constitutively active mutants of phyB (YVB) and NLS-fused phyA (YVA:NLS) mediate light-independent seedling development, leading to constitutive photomorphogenic (cop) phenotypes in their transgenic Arabidopsis plants. Interestingly, YVB interacted with PIF3 independently of light, but YVA showed little interaction. In this study, we investigated distinct signaling mechanisms underlying the similar cop phenotypes given by YVB and YVA:NLS. Our findings indicated that YVA efficiently inactivate the COP1/SPA complex, leading to accumulation of ELONGATED HYPOCOTYL 5 (HY5) and subsequent expression of its target genes HY5 and HYH. YVB induced light-independent PIF3 and PIF1 degra- dation, in addition to HY5 accumulation. Moreover, co-expression of PIF3 in the YVB plant significantly attenuated the cop phenotypes, but minimal effects were observed in the YVA:NLS plant. In particular, PIF3 negatively regulated the interaction between YVB and COP1, which decreased HY5 accumulation in the YVB plant co-expressing PIF3. Furthermore, when transferred from light to dark, PIF3 was highly accumulated in phyB-5, whereas HY5 is degraded faster in phyA-201 compared to that in Ler. Collectively, our results suggest HY5 accumulation as the molecular bases for the cop phenotypes and also indicate that phyB is more important for regulating PIF3, whereas phyA effectively inactivates the COP1/SPA complex relative to PIF3 degradation.