Expression Patterns of OsPIL11,a Phytochrome-Interacting Factor in Rice,and Preliminary Analysis of Its Roles in Light Signal Transduction

The expression patterns of OsPILll, one of six putative phytochrome-interacting factors, were analyzed in different organs of transgenic tobacco （Nicotiana tabacum）. The expression of OsPIL 11 was organ-specific and was regulated by leaf development, abscisic acid （ABA）, jasmonic acid （JA） and salicylic acid （SA）. To further explore the role of OsPIL 11 in plant light signal transduction, a plant expression vector of OsPILll was constructed and introduced into tobacco. When grown under continuous red light, OsPILll-overexpressed transgenic tobacco exhibited shorter hypocotyls and larger cotyledons and leaves compared to wild-type seedlings. When grown under continuous far-red light, however, transgenic and wild-type seedlings showed similar phenotypes. These results indicate that OsPILll is involved in red light induced de-etiolation, but not in far-red light induced de-etiolation in transgenic tobacco, which lays the foundation for dissecting the function of OsPIL11 in phytochrome-mediated light signal transduction in rice.

Regulated Degradation of HFR1 Desensitizes Light Signaling in Arabidopsis

Arabidopsis seedlings undergo photomorphogenesis in the light and etiolation in the dark. HFR1, a bHLH transcription factor, is required for both phytochrome A (phyA)-mediated far-red and cryptochrome 1 (cry1)-mediated blue light signaling. We report that HFR1 is a short-lived protein in darkness and is degraded through a 26S proteasome-dependent pathway. Light, irrespective of its quality, enhances HFR1 protein accumulation via promoting its stabilization. We demonstrate that HFR1 physically interacts with COP1 and that COP1 exhibits ubiquitin ligase activity toward HFR1 in vitro. In addition, we show that COP1 is required for degradation of HFR1 in vivo. Furthermore, plants overexpressing a C-terminal 161 amino acid fragment of HFR1 (CT161) display enhanced photomorphogenesis, suggesting an autonomous function of CT161 in promoting light signaling. This truncated HFR1 gene product is more stable than the full-length HFR1 protein in darkness, indicating that the COP1-interacting N-terminal portion of HFR1 is essential for COP1-mediated destabilization of HFR1. These results suggest that light enhances HFR1 protein accumulation by abrogating COP1-mediated degradation of HFR1, which is necessary and sufficient for promoting light signaling. Additionally, our results substantiate the E3 ligase activity of COP1 and its critical role in desensitizing light signaling.

Modulation of starch synthesis in Arabidopsis via phytochrome B-mediated light signal transduction

Starch is a major storage carbohydrate in plants and is critical in crop yield and quality.Starch synthesis is intricately regulated by internal metabolic processes and external environmental cues;however,the precise molecular mechanisms governing this process remain largely unknown.In this study,we revealed that high red to far-red(high R:FR)light significantly induces the synthesis of leaf starch and the expression of synthesis-related genes,whereas low R:FR light suppress these processes.Arabidopsis phytochrome B(phyB),the primary R and FR photoreceptor,was identified as a critical positive regulator in this process.Downstream of phyB,basic leucine zipper transcription factor ELONGATED HYPOCOTYL5(HY5)was found to enhance starch synthesis,whereas the basic helix-loop-helix transcription factors PHYTOCHROME INTERACTING FACTORs(PIF3,PIF4,and PIF5)inhibit starch synthesis in Arabidopsis leaves.Notably,HY5 and PIFs directly compete for binding to a shared G-box cis-element in the promoter region of genes encoding starch synthases GBSS,SS3,and SS4,which leads to antagonistic regulation of their expression and,consequently,starch synthesis.Our findings highlight the vital role of phyB in enhancing starch synthesis by stabilizing HY5 and facilitating PIFs degradation under high R:FR light conditions.Conversely,under low R:FR light,PIFs predominantly inhibit starch synthesis.This study provides insight into the physiological and molecular functions of phyB and its downstream transcription factors HY5 and PIFs in starch synthesis regulation,shedding light on the regulatory mechanism by which plants synchronize dynamic light signals with metabolic cues to module starch synthesis.

RACK1A promotes hypocotyl elongation by scaffolding light signaling components in Arabidopsis

Plants deploy versatile scaffold proteins to intricately modulate complex cell signaling.Among these,RACK1A(Receptors for Activated C Kinase 1A)stands out as a multifaceted scaffold protein functioning as a central integrative hub for diverse signaling pathways.However,the precise mechanisms by which RACK1A orchestrates signal transduction to optimize seedling development remain largely unclear.Here,we demonstrate that RACK1A facilitates hypocotyl elongation by functioning as a flexible platform that connects multiple key components of light signaling pathways.RACK1A interacts with PHYTOCHROME INTERACTING FACTOR(PIF)3,enhances PIF3 binding to the promoter of BBX11 and down-regulates its transcription.Furthermore,RACK1A associates with ELONGATED HYPOCOTYL 5(HY5)to repress HY5 biochemical activity toward target genes,ultimately contributing to hypocotyl elongation.In darkness,RACK1A is targeted by CONSTITUTIVELY PHOTOMORPHOGENIC(COP)1 upon phosphorylation and subjected to COP1-mediated degradation via the 26 S proteasome system.Our findings provide new insights into how plants utilize scaffold proteins to regulate hypocotyl elongation,ensuring proper skoto-and photo-morphogenic development.

Light regulation of horticultural crop nutrient uptake and utilization

Plants demonstrate dynamic changes in molecular structures under fluctuating light conditions.Accumulating evidence suggests that light plays a vital role in plant growth and morphogenesis.In particular,light has a role in the absorption and utilization of nutrients in plants.Despite significant progress in understanding the mechanism of nutrient acquisition and assimilation,how light affects and regulates ion uptake remains a question.Studies in model plants,Arabidopsis thaliana,suggest that light affects the nutrient utilization in roots through a complex regulatory network;nonetheless,the molecular mechanisms underlying the various effects of light on these processes in crop plants remain fragmentary.In this review,we discuss the light effects(light quality,light intensity,and photoperiod)on nutrient uptake and utilization in horticultural crops for optimizing crop productivity and increasing fertilizer use efficiency.

The DELLA-ABI4-HY5 module integrates light and gibberellin signals to regulate hypocotyl elongation

Plant growth is coordinately controlled by various environmental and hormonal signals,of which light and gibberellin(GA)signals are two critical factors with opposite effects on hypocotyl elongation.Although interactions between the light and GA signaling pathways have been studied extensively,the detailed regulatory mechanism of their direct crosstalk in hypocotyl elongation remains to be fully clarified.Previously,we reported that ABA INSENSITIVE 4(ABI4)controls hypocotyl elongation through its regulation of cellelongation-related genes,but whether it is also involved in GA signaling to promote hypocotyl elongation is unknown.In this study,we showthat promotion of hypocotyl elongation by GA is dependent on ABI4 activation.DELLAs interact directly with ABI4 and inhibit its DNA-binding activity.In turn,ABI4 combined with ELONGATED HYPOCOTYL 5(HY5),a key positive factor in light signaling,feedback regulates the expression of the GA2ox GA catabolism genes and thus modulates GA levels.Taken together,our results suggest that the DELLA-ABI4-HY5 module may serve as a molecular link that integrates GA and light signals to control hypocotyl elongation.

Temporal control of the Aux/IAA genes BnIAA32 and BnIAA34 mediates Brassica napus dual shade responses

Precise responses to changes in light quality are crucial for plant growth and development.For example,hypocotyls of shade-avoiding plants typically elongate under shade conditions.Although this typical shade-avoidance response(TSR)has been studied in Arabidopsis(Arabidopsis thaliana),the molecular mechanisms underlying shade tolerance are poorly understood.Here we report that B.napus(Brassica napus)seedlings exhibit dual shade responses.In addition to the TSR,B.napus seedlings also display an atypical shade response(ASR),with shorter hypocotyls upon perception of early-shade cues.Genome-wide selective sweep analysis indicated that ASR is associated with light and auxin signaling.Moreover,genetic studies demonstrated that phytochrome A(BnphyA)promotes ASR,whereas BnphyB inhibits it.During ASR,YUCCA8 expression is activated by early-shade cues,leading to increased auxin biosynthesis.This inhibits hypocotyl elongation,as young B.napus seedlings are highly sensitive to auxin.Notably,two non-canonical AUXIN/INDOLE-3-ACETIC ACID(Aux/IAA)repressor genes,BnIAA32 and BnIAA34,are expressed during this early stage.BnIAA32 and BnIAA34 inhibit hypocotyl elongation under shade conditions,and mutations in BnIAA32 and BnIAA34 suppress ASR.Collectively,our study demonstrates that the temporal expression of BnIAA32 and BnIAA34 determines the behavior of B.napus seedlings following shade-induced auxin biosynthesis.

Phytochrome Signaling： Time to Tighten up the Loose Ends

Phytochromes are red and far-red light photoreceptors that play fundamental roles in controlling many aspects of plant growth and development in response to light. The past two decades have witnessed the mechanistic elucidation of the action mode of phytochromes, including their regulation by external and endogenous factors and how they exert their function as transcriptional regulators. More importantly, recent advances have substantially deepened our understanding on the integration of the phytochromemediated signal into other cellular and developmental processes, such as elongation of hypocotyls, shoot branching, circadian clock, and flowering time, which ofteninvolves complex intercellular and interorgan signaling. Based on these advances, this review illustrates a blueprint of our current understanding of phytochrome signaling and its crosstalk with other signaling pathways, and also points out still open questions that need to be addressed in the future.

BZS1, a B-box Protein, Promotes Photomorphogenesis Downstream of Both Brassinosteroid and Light Signaling Pathways

Photomorphogenesis is controlled by multiple signaling pathways, including the light and brassinosteroid （BR） pathways. BR signaling activates the BZR1 transcription factor, which is required for suppressing photomorphogen- esis in the dark, We identified a suppressor of the BR hypersensitive mutant bzrl-lD and named it bzrl-lD suppressorl- Dominant （bzsl-D）. The bzsl-D mutation was caused by overexpression of a B-box zinc finger protein BZS1, which is transcriptionally repressed by BZR1. Overexpression of BZS1 causes de-etiolation in the dark, short hypocotyls in the light, reduced sensitivity to BR treatment, and repression of many BR-activated genes. Knockdown of BZS1 by co-suppression partly suppressed the short hypocotyl phenotypes of BR-deficient or insensitive mutants. These results support that BZSl is a negative regulator of BR response. BZS1 overexpressors are hypersensitive to different wavelengths of light and loss of function of BZS1 reduces plant sensitivity to light and partly suppresses the constitutive photomorphogenesis 1 （cop1） mutant in the dark, suggesting a positive role in light response. BZS1 protein accumulates at an increased level after light treatment of dark-grown BZSl-OXplants and in the cop1 mutants, and BZS1 interacts with COP1 in vitro, suggesting that light regulates BZS1 through COPl-mediated ubiquitination and proteasomal degradation. These results demonstrate that BZS1 mediates the crosstalk between BR and light pathways.

Phytochrome Signaling in Green Arabidopsis Seedlings： Impact Assessment of a Mutually Negative phyB-PIF Feedback Loop

The reversibly red （R）/far-red （FR）-Iight-responsive phytochrome （phy） photosensory system initiates both the deetiolation process in dark-germinated seedlings upon first exposure to light, and the shade-avoidance process in fully deetiolated seedlings upon exposure to vegetational shade. The intracellular signaling pathway from the light-activated photoreceptor conformer （Pfr） to the transcriptional network that drives these responses involves direct, physical inter- action of Pfr with a small subfamily of bHLH transcription factors, termed Phy-lnteracting Factors （PIFs）, which induces rapid PIF proteolytic degradation. In addition, there is evidence of further complexity in light-grown seedlings, whereby phyB-PIF interaction reciprocally induces phyB degradation, in a mutually-negative, feedback-loop configuration. Here, to assess the relative contributions of these antagonistic activities to the net phenotypic readout in light-grown seedlings, we have examined the magnitude of the light- and simulated-shade-induced responses of a pentuple phyBpiflpif3pif4pif5 （phyBpifq） mutant and various multiple pif-mutant combinations. The data （1） reaffirm that phyB is the predominant, if not exclusive, photoreceptor imposing the inhibition of hypocotyl elongation in deetiolating seedlings in response to pro- longed continuous R irradiation and （2） show that the PIF quartet （PIF1, PIF3, PIF4, and PIF5） retain and exert a dual capacity to modulate hypocotyl elongation under these conditions, by concomitantly promoting cell elongation through intrinsic transcriptional-regulatory activity, and reducing phyB-inhibitory capacity through feedback-loop-induced phyB degrada- tion. In shade-exposed seedlings, immunoblot analysis shows that the shade-imposed reduction in Pfr levels induces increases in the abundance of PIF3, and mutant analysis indicates that PIF3 acts, in conjunction with PIF4 and PIF5, to promote the known shade-induced acceleration of hypocotyl elongation. Conversely, although the quadruple pifq mutant displays clearly reduced hypocotyl elongation compared to wild-type in response to prolonged shade, immunoblot analysis detects no elevation in phyB levels in the mutant seedlings compared to the wild-type during the majority of the shade-induced growth period, and phyB levels are not robustly correlated with the growth phenotype across the pif-mutant combinations compared. These results suggest that PIF feedback modulation of phyB abundance does not play a dominant role in modulating the magnitude of the PIF-promoted, shade-responsive phenotype under these conditions. In seedlings grown under diurnal light-dark cycles, the data show that FR-pulse-induced removal of Pfr at the beginning of the dark period （End-of-Day-FR （EOD-FR） treatment） results in longer hypocotyls relative to no EOD-FR treatment and that this effect is attenuated in the pif-mutant combinations tested. This result similarly indicates that the PIF quartet members are capable of intrinsically promoting hypocotyl cell elongation in light-grown plants, independently of the effects of PIF feedback modulation of photoactivated-phyB abundance.

Coordination of Plastid and Light Signaling Pathways upon Development of Arabidopsis Leaves under Various Photoperiods

Plants synchronize their cellular and physiological functions according to the photoperiod （the length of the light period） in the cycle of 24 h. Photoperiod adjusts several traits in the plant life cycle, including flowering and senes- cence in annuals and seasonal growth cessation in perennials. Photoperiodic development is controlled by the coordinated action of photoreceptors and the circadian clock. During the past 10 years, remarkable progress has been made in under- standing the molecular mechanism of the circadian clock, especially with regard to the transition of Arabidopsis from the vegetative growth to the reproductive phase. Besides flowering photoperiod also modifies plant photosynthetic struc- tures and traits. Light signals controlling biogenesis of chloroplasts and development of leaf photosynthetic structures are perceived both by photoreceptors and in chloroplasts. In this review, we provide evidence suggesting that the photope- riodic development of Arabidopsis leaves mimics the acclimation of plant to various light intensities. Furthermore, the chloroplast-to-nucleus retrograde signals that adjust acclimation to light intensity are proposed to contribute also to the signaling pathways that control photoperiodic acclimation of leaves.

Long-distance blue light signalling regulates phosphate deficiency-induced primary root growth inhibition

Although roots are mainly embedded in the soil, recent studies revealed that light regulates mineral nutrient uptake by roots. However, it remains unclear whether the change in root system architecture in response to different rhizosphere nutrient statuses involves light signaling. Here, we report that blue light regulates primary root growth inhibition under phosphate-deficient conditions through the cryptochromes and their downstream signaling factors. We showed that the inhibition of root elongation by low phosphate requires blue light signal perception at the shoot and transduction to the root. In this process, SPA1 and COP1 play a negative role while HY5 plays a positive role. Further experiments revealed that HY5 is able to migrate from the shoot to root and that the shoot-derived HY5 autoactivates root HY5 and regulates primary root growth by directly activating the expression of LPR1, a suppressor of root growth under phosphate starvation. Taken together, our study reveals a regulatory mechanism by which blue light signaling regulates phosphate deficiency-induced primary root growth inhibition, providing new insights into the crosstalk between light and nutrient signaling.

BBX11 promotes red light-mediated photomorphogenic development by modulating phyB-PIF4 signaling

phytochrome B(phyB)acts as the red light photoreceptor and negatively regulates the growth-promoting factor PHYTOCHROME INTERACTING 4(PIF4)through a direct physical interaction,which in turn changes the expression of a large number of genes.phyB-PIF4 module regulates a variety of biological and developmental processes in plants.In this study,we demonstrate that B-BOX PROTEIN 11(BBX11)physically interacts with both phyB and PIF4.BBX11 negatively regulates PIF4 accumulation as well as its biochemical activity,consequently leading to the repression of PIF4-controlled genes’expression and promotion of photomorphogenesis in the prolonged red light.This study reveals a regulatory mechanism that mediates red light signal transduction and sheds a light on phyB-PIF4 module in promoting red light-dependent photomorphognenesis.

Coordinated Shoot and Root Responses to Light Signaling in Arabidopsis

Light is one of the most important environmental signals and regulates many biological processes in plants.Studies on light-regulated development have mainly focused on aspects of shoot growth,such as deetiolation,cotyledon opening,inhibition of hypocotyl elongation,flowering,and anthocyanin accumulation.However,recent studies have demonstrated that light is also involved in regulating root growth and development in Arabidopsis.In this review,we summarize the progress in understanding how shoots and roots coordinate their responses to light through different light-signaling components and pathways,including the COP1(CONSTITUTIVELY PHOTOMORPHOGENIC 1),HY5(ELONGATED HYPOCOTYL 5),and MYB73/MYB77(MYB DOMAIN PROTEIN 73/77)pathways.

The involvement of the N-terminal PHR domain of Arabidopsis cryptochromes in mediating light signaling

Light is a key environmental cue that fundamentally regulates all aspects of plant growth and development,which is mediated by the multiple photoreceptors including the blue light photoreceptors cryptochromes(CRYs).In Arabidopsis,there are two well-characterized homologous CRYs,CRY1 and CRY2.Whereas CRYs are flavoproteins,they lack photolyase activity and are characterized by an Nterminal photolyase-homologous region(PHR)domain and a C-terminal extension domain.It has been established that the C-terminal extension domain of CRYs is involved in mediating light signaling through direct interactions with the master negative regulator of photomorphogenesis,COP1.Recent studies have revealed that the N-terminal PHR domain of CRYs is also involved in mediating light signaling.In this review,we mainly summarize and discuss the recent advances in CRYs signaling mediated by the N-terminal PHR domain,which involves the N-terminal PHR domain-mediated dimerization/oligomerization of CRYs and physical interactions with the pivotal transcription regulators in light and phytohormone signaling.

PIFs‑and COP1‑HY5‑mediated temperature signaling in higher plants

Plants have to cope with the surrounding changing environmental stimuli to optimize their physiological and devel-opmental response throughout their entire life cycle.Light and temperature are two critical environmental cues that fluctuate greatly during day-night cycles and seasonal changes.These two external signals coordinately control the plant growth and development.Distinct spectrum of light signals are perceived by a group of wavelength-specific photoreceptors in plants.PIFs and COP1-HY5 are two predominant signaling hubs that control the expression of a large number of light-responsive genes and subsequent light-mediated development in plants.In parallel,plants also transmit low or warm temperature signals to these two regulatory modules that precisely modulate the responsive-ness of low or warm temperatures.The core component of circadian clock ELF3 integrates signals from light and warm temperatures to regulate physiological and developmental processes in plants.In this review,we summarize and discuss recent advances and progresses on PIFs-,COP1-HY5-and ELF3-mediated light,low or warm temperature signaling,and highlight emerging insights regarding the interactions between light and low or warm temperature signal transduction pathways in the control of plant growth.

New proposal of modulation of amplitude-squeezed state of light by intensity variation of a low-frequency coherent message signal

Squeezed state of light explores a new era in noiseless communication and data processing recently breaking the quantum limit of noise. We propose a new mechanism of modulating an amplitude-squeezed signal with the instantaneous intensity variation of a coherent signal. The modulating signal is a coherent light where the amplitude-squeezed light takes the role of a carrier signal.

HY5-HDA9 orchestrates the transcription of HsfA2 to modulate salt stress response in Arabidopsis

Integration of light signaling and diverse abiotic stress responses contribute to plant survival in a changing environment.Some reports have indicated that light signals contribute a plant’s ability to deal with heat,cold,and stress.However,the molecular link between light signaling and the saltresponse pathways remains unclear.We demonstrate here that increasing light intensity elevates the salt stress tolerance of plants.Depletion of HY5,a key component of light signaling,causes Arabidopsis thaliana to become salinity sensitive.Interestingly,the small heat shock protein(sHsp)family genes are upregulated in hy5-215 mutant plants,and HsfA 2 is commonly involved in the regulation of these sH sps.We found that HY5directly binds to the G-box motifs in the HsfA2promoter,with the cooperation of HISTONE DEACETYLASE 9(HDA9),to repress its expression.Furthermore,the accumulation of HDA9 and the interaction between HY5 and HDA9 are significantly enhanced by salt stress.On the contrary,high temperature triggers HY5 and HDA9 degradation,which leads to dissociation of HY5-HDA9from the HsfA2 promoter,thereby reducing salt tolerance.Under salt and heat stress conditions,fine tuning of protein accumulation and an interaction between HY5 and HDA9 regulate HsfA2 expression.This implies that HY5,HDA9,and HsfA2play important roles in the integration of light signaling with salt stress and heat shock response.

Role of Arabidopsis RAP2.4 in Regulating Light-and Ethylene-Mediated Developmental Processes and Drought Stress Tolerance

Light and the plant hormone ethylene regulate many aspects of plant growth and development in an overlapping and interdependent fashion. Little is known regarding how their signal transduction pathways cross-talk to regulate plant development in a coordinated manner. Here, we report functional characterization of an AP2/DREB-type transcription factor, Arabidopsis RAP2.4, in mediating light and ethylene signaling. Expression of the RAP2.4 gene is down-regulated by light but up-regulated by salt and drought stresses. RAP2.4 protein is constitutively targeted to the nucleus and it can bind to both the ethylene-responsive GCC-box and the dehydration-responsive element （DRE）. We show that RAP2.4 protein possesses an intrinsic transcriptional activation activity in yeast cells and that it can activate a reporter gene driven by the DRE cis-element in Arabidopsis protoplasts. Overexpression of RAP2.4 or mutation in RAP2.4 cause altered expression of representative light-, ethylene-, and drought-responsive genes. Although no salient phenotype was observed with a rap2.4 loss-of-function mutant, constitutive overexpression of RAP2.4 results in defects in multiple developmental processes regulated by light and ethylene, including hypocotyl elongation and gravitropism, apical hook formation and cotyledon expansion, flowering time, root elongation, root hair formation, and drought tolerance. Based on these observations, we propose that RAP2.4 acts at or downstream of a converging point of light and ethylene signaling pathways to coordinately regulate multiple developmental processes and stress responses.

MYC2： The Master in Action

Jasmonates （JAs） are plant hormones with essential roles in plant defense and development. The basic- helix-loop-helix （bHLH） transcription factor （TF） MYC2 has recently emerged as a master regulator of most aspects of the jasmonate （JA） signaling pathway in Arabidopsis. MYC2 coordinates JA-mediated defense responses by antagonistically regulating two different branches of the JA signaling pathway that determine resistance to pests and pathogens, respectively. MYC2 is required for induced systemic resistance （ISR） triggered by beneficial soil microbes while MYC2 function is targeted by pathogens during effector-mediated suppression of innate immunity in roots. Another notable function of MYC2 is the regulation of crosstalk between the signaling pathways of JA and those of other phytohormones such as abscisic acid （ABA）, salicylic acid （SA）, gibberellins （GAs）, and auxin （IAA）. MYC2 also regulates interactions between JA signaling and light, phytochrome signaling, and the circadian clock, MYC2 is involved in JA-regulated plant development, lateral and adventitious root formation, flowering time, and shade avoidance syndrome. Related bHLH TFs MYC3 and MYC4 also regulate both overlapping and distinct MYC2-regulated functions in Arabidopsis while MYC2 orthologs act as ＇master switches＇ that regulate JA-mediated biosynthesis of secondary metabolites. Here, we briefly review recent studies that revealed mechanistic new insights into the mode of action of this versatile TF.