B-BOX proteins:Multi-layered roles of molecular cogs in light-mediated growth and development in plants.

B-box containing proteins (BBXs) are a class of zinc-ligating transcription factors or regulators that play essential roles in various physiological and developmental processes in plants. They not only directly associate with target genes to regulate their transcription, but also interact with other transcription factors to mediate target genes' expression, thus forming a complex transcriptional network ensuring plants' adaptation to dynamically changing light environments. This review summarizes and highlights the molecular and biochemical properties of BBXs, as well as recent advances with a focus on their critical regulatory functions in photomorphogenesis (de-etiolation), shade avoidance, photoperiodic-mediated flowering, and secondary metabolite biosynthesis and accumulation in plants.

Using Magnetic Fields to Enhance the Seed Germination, Growth, and Yield of Plants.

Geomagnetic field (GMF) protects living organisms on the Earth from the radiation coming from space along with other environmental factors during evolution, and it has affected the growth and development of plants. Many researchers have always been interested in investigating these effects in different aspects. In this chapter, we focus on the methods of using different types of magnetic fields (MFs) to investigate the dimensions of their biological effects on plants. The aim is to increase seed germination, growth characters, and yield of plants using the following methods: (1) Using MFs lower than GMF to study effects of GMF on the growth and yield of plants. (2) Using reversed magnetic fields (RMFs) lower than GMF to study its effects on the growth and development of plants during evolution. (3) Using static magnetic fields (SMFs) higher than GMF and reversed SMFs to study effects of the south (S) and north (N) magnetic pole on plants. (4) Using electromagnetic fields (EMFs) to increase and accelerate seed germination, growth, and yield of plants, and establish the status of plants against other environmental stresses. (5) Using magnetized water (MW) to improve plant seed germination, growth, and yield. (6) Using high gradient magnetic field (HGMF) to study magneto-tropism in plants. In this chapter, we recommend application of various types of MFs to study their biological effects on plants to improve crop production.

Environmentally adaptive reshaping of plant photomorphogenesis by karrikin and strigolactone signaling.

Coordinated morphogenic adaptation of growing plants is critical for their survival and propagation under fluctuating environments. Plant morphogenic responses to light and warm temperatures, termed photomorphogenesis and thermomorphogenesis, respectively, have been extensively studied in recent decades. During photomorphogenesis, plants actively reshape their growth and developmental patterns to cope with changes in light regimes. Accordingly, photomorphogenesis is closely associated with diverse growth hormonal cues. Notably, accumulating evidence indicates that light-directed morphogenesis is profoundly affected by two recently identified phytochemicals, karrikins (KARs) and strigolactones (SLs). KARs and SLs are structurally related butenolides acting as signaling molecules during a variety of developmental steps, including seed germination. Their receptors and signaling mediators have been identified, and associated working mechanisms have been explored using gene-deficient mutants in various plant species. Of particular interest is that the KAR and SL signaling pathways play important roles in environmental responses, among which their linkages with photomorphogenesis are most comprehensively studied during seedling establishment. In this review, we focus on how the phytochemical and light signals converge on the optimization of morphogenic fitness. We also discuss molecular mechanisms underlying the signaling crosstalks with an aim of developing potential ways to improve crop productivity under climate changes.

Morphogenetic and physiological effects of LED spectra on the apical buds of Ficus carica var. Black Jack.

The use of artificial light sources such as light-emitting diodes (LEDs) has become a prerequisite in tissue culture studies to obtain morphogenetic enhancements on in vitro plants. This technology is essential for developmental enhancements in the growing plant cultures due to its light quality and intensity greatly influencing the in vitro growing explants at a cellular level. The current study investigates the effects of different light-emitting diode (LED) spectra on the growth of apical buds of Ficus carica var. Black Jack. Ficus carica, commonly known as figs is rich in vitamins, minerals, and phytochemicals capable of treating microbial infections and gastric, inflammatory, and cardiac disorders. Apical buds of Ficus carica var. Black Jack, presented morphogenetic changes when grown under six different LED spectra. The highest multiple shoots (1.80 per growing explant) and healthy growing cultures were observed under the blue + red LED spectrum. Wound-induced callus formation was observed on apical buds grown under green LED spectrum and discolouration of the growing shoots were observed on the cultures grown under far-red LED spectrum. Multiple shoots obtained from the blue + red LED treatment were rooted using 8 µM indole-3-acetic acid (IAA), and the rooted plantlets were successfully acclimatised. Compared with the other monochromatic LEDs, blue + red proved to be significantly better for producing excellent plant morphogeny. It is apparent that blue and red LED is the most suitable spectra for the healthy development of plants. The findings have confirmed that the combination of blue + red LED can potentially be used for enhancing growth yields of medicinally and commercially important plants.

Light Induced Regulation Pathway of Anthocyanin Biosynthesis in Plants.

Anthocyanins are natural pigments with antioxidant effects that exist in various fruits and vegetables. The accumulation of anthocyanins is induced by environmental signals and regulated by transcription factors in plants. Numerous evidence has indicated that among the environmental factors, light is one of the most signal regulatory factors involved in the anthocyanin biosynthesis pathway. However, the signal transduction of light and molecular regulation of anthocyanin synthesis remains to be explored. Here, we focus on the research progress of signal transduction factors for positive and negative regulation in light-dependent and light-independent anthocyanin biosynthesis. In particular, we will discuss light-induced regulatory pathways and related specific regulators of anthocyanin biosynthesis in plants. In addition, an integrated regulatory network of anthocyanin biosynthesis controlled by transcription factors is discussed based on the significant progress.

Functional comparison of the WD-repeat domains of SPA1 and COP1 in suppression of photomorphogenesis.

The Arabidopsis COP1/SPA complex acts as a cullin4-based E3 ubiquitin ligase to suppress photomorphogenesis in darkness. It is a tetrameric complex of two COP1 and two SPA proteins. Both COP1 and SPA are essential for the activity of this complex, and they both contain a C-terminal WD-repeat domain responsible for substrate recruitment and binding of DDB1. Here, we used a WD domain swap-approach to address the cooperativity of COP1 and SPA proteins. We found that expression of a chimeric COP1 carrying the WD-repeat domain of SPA1 mostly complemented the cop1-4-mutant phenotype in darkness, indicating that the WD repeat of SPA1 can replace the WD repeat of COP1. In the light, SPA1-WD partially substituted for COP1-WD. In contrast, expression of a chimeric SPA1 protein carrying the WD repeat of COP1 did not rescue the spa-mutant phenotype. Together, our findings demonstrate that a SPA1-type WD repeat is essential for COP1/SPA activity, while a COP1-type WD is in part dispensible. Moreover, a complex with four SPA1-WDs is more active than a complex with only two SPA1-WDs. A homology model of SPA1-WD based on the crystal structure of COP1-WD uncovered two insertions and several amino acid substitutions at the predicted substrate-binding pocket of SPA1-WD.

Sound Waves Promote Arabidopsis thaliana Root Growth by Regulating Root Phytohormone Content.

Sound waves affect plants at the biochemical, physical, and genetic levels. However, the mechanisms by which plants respond to sound waves are largely unknown. Therefore, the aim of this study was to examine the effect of sound waves on Arabidopsis thaliana growth. The results of the study showed that Arabidopsis seeds exposed to sound waves (100 and 100 + 9k Hz) for 15 h per day for 3 day had significantly longer root growth than that in the control group. The root length and cell number in the root apical meristem were significantly affected by sound waves. Furthermore, genes involved in cell division were upregulated in seedlings exposed to sound waves. Root development was affected by the concentration and activity of some phytohormones, including cytokinin and auxin. Analysis of the expression levels of genes regulating cytokinin and auxin biosynthesis and signaling showed that cytokinin and ethylene signaling genes were downregulated, while auxin signaling and biosynthesis genes were upregulated in Arabidopsis exposed to sound waves. Additionally, the cytokinin and auxin concentrations of the roots of Arabidopsis plants increased and decreased, respectively, after exposure to sound waves. Our findings suggest that sound waves are potential agricultural tools for improving crop growth performance.

Light regulates stomatal development by modulating paracrine signaling from inner tissues.

Developmental outcomes are shaped by the interplay between intrinsic and external factors. The production of stomata-essential pores for gas exchange in plants-is extremely plastic and offers an excellent system to study this interplay at the cell lineage level. For plants, light is a key external cue, and it promotes stomatal development and the accumulation of the master stomatal regulator SPEECHLESS (SPCH). However, how light signals are relayed to influence SPCH remains unknown. Here, we show that the light-regulated transcription factor ELONGATED HYPOCOTYL 5 (HY5), a critical regulator for photomorphogenic growth, is present in inner mesophyll cells and directly binds and activates STOMAGEN. STOMAGEN, the mesophyll-derived secreted peptide, in turn stabilizes SPCH in the epidermis, leading to enhanced stomatal production. Our work identifies a molecular link between light signaling and stomatal development that spans two tissue layers and highlights how an environmental signaling factor may coordinate growth across tissue types.

Lighting Direction Affects Leaf Morphology, Stomatal Characteristics, and Physiology of Head Lettuce (Lactuca sativa L.).

Plants are exposed to numerous biotic and abiotic stresses, and light is one of the most important factors that influences the plant morphology. This study was carried out to examine how the lighting direction affected the plant morphology by investigating the growth parameters, epidermal cell elongation, stomatal properties, and physiological changes. Seedlings of two head lettuce (Lactuca sativa L.) cultivars, Caesar Green and Polla, were subjected to a 12 h photoperiod with a 300 µmol·m-2·s-1 photosynthetic photon flux density (PPFD) provided by light emitting diodes (LEDs) from three directions: the top, side, and bottom, relative to the plants. Compared with the top and side lighting, the bottom lighting increased the leaf angle and canopy by stimulating the epidermal cell elongation in leaf midrib, reduced the leaf number and root biomass, and induced large stomata with a low density, which is associated with reduced stomatal conductance and carbohydrate contents. However, the proline content and quantum yield exhibited no significant differences with the different lighting directions in both cultivars, which implies that the plants were under normal physiological conditions. In a conclusion, the lighting direction had a profound effect on the morphological characteristics of lettuce, where the plants adapted to the changing lighting environments.

Light-induced degradation of SPA2 via its N-terminal kinase domain is required for photomorphogenesis.

Arabidopsis (Arabidopsis thaliana) CONSTITUTIVELY PHOTOMORPHOGENIC1 (COP1) and members of the SUPPRESSOR OF PHYTOCHROMEA-105 (SPA) protein family form an E3 ubiquitin ligase that suppresses light signaling in darkness by polyubiquitinating positive regulators of the light response. COP1/SPA is inactivated by light to allow photomorphogenesis to proceed. Mechanisms of inactivation include light-induced degradation of SPA1 and, in particular, SPA2, corresponding to a particularly efficient inactivation of COP1/SPA2 by light. Here, we show that SPA3 and SPA4 proteins are stable in the light, indicating that light-induced destabilization is specific to SPA1 and SPA2, possibly related to the predominant function of SPA1 and SPA2 in dark-grown etiolating seedlings. SPA2 degradation involves cullin and the COP10-DEETIOLATED-DAMAGED-DNA BINDING PROTEIN (DDB1) CDD complex, besides COP1. Consistent with this finding, light-induced SPA2 degradation required the DDB1-interacting Trp-Asp (WD)-repeat domain of SPA2. Deletion of the N-terminus of SPA2 containing the kinase domain led to strong stabilization of SPA2 in darkness and fully abolished light-induced degradation of SPA2. This prevented seedling de-etiolation even in very strong far-red and blue light and reduced de-etiolation in red light, indicating destabilization of SPA2 through its N-terminal domain is essential for light response. SPA2 is exclusively destabilized by phytochrome A in far-red and blue light. However, deletion of the N-terminal domain of SPA2 did not abolish SPA2-phytochrome A interaction in yeast nor in vivo. Our domain mapping suggests there are two SPA2-phytochrome A interacting domains, the N-terminal domain and the WD-repeat domain. Conferring a light-induced SPA2-phyA interaction only via the WD-repeat domain may thus not lead to COP1/SPA2 inactivation.

Magnetic Field Treatments Improves Sunflower Yield by Inducing Physiological and Biochemical Modulations in Seeds.

Magnetic seed enhancement has been practicing as a promising tool to improve germination and seedling growth of low vigor seeds stored under suboptimal conditions, but there is still ambiguity regarding the prospects for magnetism in oilseeds. Present study elucidates the potential of magnetic seed stimulation to improve sunflower germination, growth and yield. Germination and emergence tests were performed to optimize the strength of the magnetic field to sunflower seed enhancement. The seeds were directly exposed to magnetic field strengths of 50, 100 and 150 millitesla (mT) for 5, 10 and 15 min (min) and then standard germination tests were performed. Secondly, the emergence potential of untreated seeds was compared with seed exposed to hydropriming, priming with 3% moringa leaf extract (MLE), priming with magnetically treated water (MTW) for 10 min and priming with 3% MLE solution prepared in magnetically treated water (MTW + MLE). Germination, emergence, seedling growth and seed biochemical properties were used to select the best treatment for field evaluation. The results of the study revealed that magnetic seed treatment with 100 mT for 10 min and seed priming with 3% MLE solution in magnetically treated water (MTW + MLE) significantly improved emergence, crop growth rate and sunflower yield.

Light signalling shapes plant-plant interactions in dense canopies.

Plants growing at high densities interact via a multitude of pathways. Here, we provide an overview of mechanisms and functional consequences of plant architectural responses initiated by light cues that occur in dense vegetation. We will review the current state of knowledge about shade avoidance, as well as its possible applications. On an individual level, plants perceive neighbour-associated changes in light quality and quantity mainly with phytochromes for red and far-red light and cryptochromes and phototropins for blue light. Downstream of these photoreceptors, elaborate signalling and integration takes place with the PHYTOCHROME INTERACTING FACTORS, several hormones and other regulators. This signalling leads to the shade avoidance responses, consisting of hyponasty, stem and petiole elongation, apical dominance and life cycle adjustments. Architectural changes of the individual plant have consequences for the plant community, affecting canopy structure, species composition and population fitness. In this context, we highlight the ecological, evolutionary and agricultural importance of shade avoidance.

Light from below matters: Quantifying the consequences of responses to far-red light reflected upwards for plant performance in heterogeneous canopies.

In vegetation stands, plants receive red to far-red ratio (R:FR) signals of varying strength from all directions. However, plant responses to variations in R:FR reflected from below have been largely ignored despite their potential consequences for plant performance. Using a heterogeneous rose canopy, which consists of bent shoots down in the canopy and vertically growing upright shoots, we quantified upward far-red reflection by bent shoots and its consequences for upright shoot architecture. With a three-dimensional plant model, we assessed consequences of responses to R:FR from below for plant photosynthesis. Bent shoots reflected substantially more far-red than red light, causing reduced R:FR in light reflected upwards. Leaf inclination angles increased in upright shoots which received low R:FR reflected from below. The increased leaf angle led to an increase in simulated plant photosynthesis only when this low R:FR was reflected off their own bent shoots and not when it reflected off neighbour bent shoots. We conclude that plant response to R:FR from below is an under-explored phenomenon which may have contrasting consequences for plant performance depending on the type of vegetation or crop system. The responses are beneficial for performance only when R:FR is reflected by lower foliage of the same plants.

Light- and hormone-mediated development in non-flowering plants: An overview.

MAIN CONCLUSION: Light, hormones and their interaction regulate different aspects of development in non-flowering plants. They might have played a role in the evolution of different plant groups by conferring specific adaptive evolutionary changes. Plants are sessile organisms. Unlike animals, they lack the opportunity to abandon their habitat in unfavorable conditions. They respond to different environmental cues and adapt accordingly to control their growth and developmental pattern. While phytohormones are known to be internal regulators of plant development, light is a major environmental signal that shapes plant processes. It is plausible that light-hormone crosstalk might have played an important role in plant evolution. But how the crosstalk between light and phytohormone signaling pathways might have shaped the plant evolution is unclear. One of the possible reasons is that flowering plants have been studied extensively in context of plant development, which cannot serve the purpose of evolutionary comparisons. In order to elucidate the role of light, hormone and their crosstalk in the evolutionary adaptation in plant kingdom, one needs to understand various light- and hormone-mediated processes in diverse non-flowering plants. This review is an attempt to outline major light- and phytohormone-mediated responses in non-flowering plant groups such as algae, bryophytes, pteridophytes and gymnosperms.

Light spectra affect the in vitro shoot development of Cedrela fissilis Vell. (Meliaceae) by changing the protein profile and polyamine contents.

The light spectrum quality is an important signal for plant growth and development. We evaluated the effects of different light spectra on the in vitro shoot development of Cedrela fissilis and its proteomic and polyamine (PA) profiles. Cotyledonary and apical nodal segments were grown under different light emitting diodes (LED) and fluorescent lamps. Shoots from cotyledonary nodal segments cultured with 6-benzyladenine (BA) that were grown under WmBdR LED showed increased length and higher fresh and dry matter compared to shoots grown under fluorescent lamps. A nonredundant protein databank generated by transcriptome sequencing and the de novo assembly of C. fissilis improved, and almost doubled, the protein identification compared to a Citrus sinensis databank. A total of 616 proteins were identified, with 23 up- and 103 down-accumulated in the shoots under WmBdR LEDs compared to fluorescent lamps. Most differentially accumulated proteins in shoots grown under the WmBdR LED lamp treatment compared to the fluorescent lamp treatment are involved in responding to metabolic processes, stress, biosynthetic and cellular protein modifications, and light stimulus processes. Among the proteins, the up-accumulation of argininosuccinate synthase was associated with an increase in the free putrescine content and, consequently, with higher shoot elongation under WmBdR LED. The down-accumulation of calreticulin, heat shock proteins, plastid-lipid-associated protein, ubiquitin-conjugating enzymes, and ultraviolet-B receptor UVR8 isoform X1 could be related to the longer shoot length noted under LED treatment. This study provides important data related to the effects of the light spectrum quality on in vitro morphogenesis through the modulation of specific proteins and free putrescine biosynthesis in C. fissilis, an endangered wood species from the Brazilian Atlantic Forest of economic and ecological relevance. The nonredundant protein databank of C. fissilis is available via ProteomeXchange under identifier PXD018020.

Allosteric deactivation of PIFs and EIN3 by microproteins in light control of plant development.

Buried seedlings undergo dramatic developmental transitions when they emerge from soil into sunlight. As central transcription factors suppressing light responses, PHYTOCHROME-INTERACTING FACTORs (PIFs) and ETHYLENE-INSENSITIVE 3 (EIN3) actively function in darkness and must be promptly repressed upon light to initiate deetiolation. Microproteins are evolutionarily conserved small single-domain proteins that act as posttranslational regulators in eukaryotes. Although hundreds to thousands of microproteins are predicted to exist in plants, their target molecules, biological roles, and mechanisms of action remain largely unknown. Here, we show that two microproteins, miP1a and miP1b (miP1a/b), are robustly stimulated in the dark-to-light transition. miP1a/b are primarily expressed in cotyledons and hypocotyl, exhibiting tissue-specific patterns similar to those of PIFs and EIN3 We demonstrate that PIFs and EIN3 assemble functional oligomers by self-interaction, while miP1a/b directly interact with and disrupt the oligomerization of PIFs and EIN3 by forming nonfunctional protein complexes. As a result, the DNA binding capacity and transcriptional activity of PIFs and EIN3 are predominantly suppressed. These biochemical findings are further supported by genetic evidence. miP1a/b positively regulate photomorphogenic development, and constitutively expressing miP1a/b rescues the delayed apical hook unfolding and cotyledon development of plants overexpressing PIFs and EIN3 Our study reveals that microproteins provide a temporal and negative control of the master transcription factors' oligomerization to achieve timely developmental transitions upon environmental changes.

Effect of Shading on Development, Yield and Quality of Bastard Balm Herb (Melittis melissophyllum L.).

The aim of the study was to assess the effects of Melittis melissophyllum shading on its development and accumulation of phenolics. Their content (verbascoside, apiin, luteolin-7-O-glucoside, coumarin, 3,4-dihydroxycoumarin, o-coumaric acid 2-O-glucoside as well as o-coumaric, p-coumaric, chlorogenic, caffeic, ferulic and cichoric acid) was determined in the herb using HPLC-DAD. The results showed that the content of abovementioned flavonoids and phenolic acids was highest in plants grown under full sunlight. On the other hand, a higher content of coumarin was observed in shaded plants, especially after the seed-setting stage. A similar tendency was noted for the amount of chlorophyll a and b. The content of hydrogen peroxide and malondialdehyde, the activity of polyphenol oxidase and catalase and the antioxidant capacity of plant extracts (measured using DPPH, ABTS and FRAP assays) were found to be the highest in the plants grown in full sunlight. However, the plants grown in moderate (30%) shade were found to thrive best.

Measurement Method for Height-Independent Vegetation Indices Based on an Active Light Source.

A coefficient CW, which was defined as the ratio of NIR (near infrared) to the red reflected spectral response of the spectrometer, with a standard whiteboard as the measuring object, was introduced to establish a method for calculating height-independent vegetation indices (VIs). Two criteria for designing the spectrometer based on an active light source were proposed to keep CW constant. A designed spectrometer, which was equipped with an active light source, adopting 730 and 810 nm as the central wavelength of detection wavebands, was used to test the Normalized Difference Vegetation Index (NDVI) and Ratio Vegetation Index (RVI) in wheat fields with two nitrogen application rate levels (NARLs). Twenty test points were selected in each kind of field. Five measuring heights (65, 75, 85, 95, and 105 cm) were set for each test point. The mean and standard deviation of the coefficient of variation (CV) for NDVI in each test point were 3.85% and 1.39% respectively, the corresponding results for RVI were 2.93% and 1.09%. ANOVA showed the measured VIs possessed a significant ability to discriminate the NARLs and had no obvious correlation with the measurement heights. The experimental results verified the feasibility and validity of the method for measuring height-independent VIs.

PCH1 and PCHL Directly Interact with PIF1, Promote Its Degradation, and Inhibit Its Transcriptional Function during Photomorphogenesis.

PHOTOPERIODIC CONTROL OF HYPOCOTYL 1 (PCH1) and PCH1-LIKE (PCHL) were shown to directly bind to phytochrome B (phyB) and suppress phyB thermal reversion, resulting in plants with dramatically enhanced light sensitivity. Here, we show that PCH1 and PCHL also positively regulate various light responses, including seed germination, hypocotyl gravitropism, and chlorophyll biosynthesis, by physically interacting with PHYTOCHROME INTERACTING FACTOR 1 (PIF1) and CONSTITUTIVE PHOTOMORPHOGENIC 1 (COP1). PCH1 and PCHL interact with PIF1 both in the dark and light, and regulate PIF1 abundance. Moreover, PCH1 and PCHL facilitate the physical interaction between phyB and PIF1 in vivo to promote the light-induced degradation of PIF1. PCH1 and PCHL also inhibit the DNA-binding ability of PIF1 to negatively regulate the expressions of PIF1 target genes. In addition, PCH1 and PCHL interact with COP1 and undergo degradation through the 26S proteasome pathway in the dark. Consistently, pch1 suppresses cop1 phenotype in darkness. Collectively, our study reveals a novel mechanism by which PCH1 and PCHL regulate diverse light responses not only by stabilizing phyB Pfr form but also by directly interacting with PIF1 and COP1, providing a molecular understanding of the control of hypocotyl growth by these proteins.

Regulation of Photomorphogenic Development by Plant Phytochromes.

Photomorphogenesis and skotomorphogenesis are two key events that control plant development, from seed germination to flowering and senescence. A group of wavelength-specific photoreceptors, E3 ubiquitin ligases, and various transcription factors work together to regulate these two critical processes. Phytochromes are the main photoreceptors in plants for perceiving red/far-red light and transducing the light signals to downstream factors that regulate the gene expression network for photomorphogenic development. In this review, we highlight key developmental stages in the life cycle of plants and how phytochromes and other components in the phytochrome signaling pathway play roles in plant growth and development.