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Phototropins, namely, phototropin 1 (phot1) and phototropin 2 (phot2), mediate chloroplast movement to maximize photosynthetic efficiency and prevent photodamage in plants. Phot1 primarily functions in chloroplast accumulation process, whereas phot2 mediates both chloroplast avoidance and accumulation responses. The avoidance response of phot2-mediated chloroplasts under high-intensity blue light (HBL) limited the understanding of the function of phot1 in the chloroplast accumulation process at the HBL condition. In this study, we showed that the phot2 mutant exhibits a chloroplast accumulation response under HBL, which is defective when the root phototropism 2 (RPT2) gene is mutated in the phot2 background, mimicking the phenotype of the phot1 phot2 double mutant. A further analysis revealed that the expression of RPT2 was induced by HBL and the overexpression of RPT2 could partially enhance the chloroplast accumulation response under HBL. These results confirmed that RPT2 also participates in regulating the phot1-mediated chloroplast accumulation response under HBL. In contrast, RPT2 functions redundantly with neural retina leucine zipper (NRL) protein for chloroplast movement 1 (NCH1) under low-light irradiation. In addition, no chloroplast accumulation response was detected in the phot2 jac1 double mutant under HBL, which has been previously observed in phot2 rpt2 and phot1 phot2 double mutants. Taken together, our results indicated that phot1 mediates the HBL-induced chloroplast accumulation response in an RPT2-dependent manner and is also regulated by j-domain protein required for chloroplast accumulation response 1 (JAC1).

Stomata in the epidermis of plants play essential roles in the regulation of photosynthesis and transpiration. Stomata open in response to blue light (BL) by phosphorylation-dependent activation of the plasma membrane (PM) H+-ATPase in guard cells. Under water stress, the plant hormone abscisic acid (ABA) promotes stomatal closure via the ABA-signaling pathway to reduce water loss. We established a chemical screening method to identify compounds that affect stomatal movements in Commelina benghalensis. We performed chemical screening using a protease inhibitor (PI) library of 130 inhibitors to identify inhibitors of stomatal movement. We discovered 17 PIs that inhibited light-induced stomatal opening by more than 50%. Further analysis of the top three inhibitors (PI1, PI2, and PI3; inhibitors of ubiquitin-specific protease 1, membrane type-1 matrix metalloproteinase, and matrix metalloproteinase-2, respectively) revealed that these inhibitors suppressed BL-induced phosphorylation of the PM H+-ATPase but had no effect on the activity of phototropins or ABA-dependent responses. The results suggest that these PIs suppress BL-induced stomatal opening at least in part by inhibiting PM H+-ATPase activity but not the ABA-signaling pathway. The targets of PI1, PI2, and PI3 were predicted by bioinformatics analyses, which provided insight into factors involved in BL-induced stomatal opening.

Photomorphogenesis refers to photoreceptor-mediated morphological changes in plant development that are triggered by light. Multiple photoreceptors and transcription factors (TFs) are involved in the molecular regulation of photomorphogenesis. Likewise, light can also modulate the outcome of plant–virus interactions since both photosynthesis and many viral infection events occur in the chloroplast. Despite the apparent association between photosynthesis and virus infection, little is known about whether there are also interplays between photomorphogenesis and plant virus resistance. Recent research suggests that plant–virus interactions are potentially regulated by several photoreceptors and photomorphogenesis regulators, including phytochromes A and B (PHYA and PHYB), cryptochromes 2 (CRY2), phototropin 2 (PHOT2), the photomorphogenesis repressor constitutive photomorphogenesis 1 (COP1), the NAM, ATAF, and CUC (NAC)-family TF ATAF2, the Aux/IAA protein phytochrome-associated protein 1 (PAP1), the homeodomain-leucine zipper (HD-Zip) TF HAT1, and the core circadian clock component circadian clock associated 1 (CCA1). Particularly, the plant growth promoting brassinosteroid (BR) hormones play critical roles in integrating the regulatory pathways of plant photomorphogenesis and viral defense. Here, we summarize the current understanding of molecular mechanisms linking plant photomorphogenesis and defense against viruses, which represents an emerging interdisciplinary research topic in both molecular plant biology and virology.

It is well known that light is a crucial environmental factor that has a fundamental role in plant growth and development from seed germination to fruiting. For this process, plants contain versatile and multifaceted photoreceptor systems to sense variations in the light spectrum and to acclimate to a range of ambient conditions. Five main groups of photoreceptors have been found in higher plants, cryptochromes, phototropins, UVR8, zeitlupes, and phytochromes, but the last one red/far red wavelengths photoreceptor is the most characterized. Among the many responses modulated by phytochromes, these molecules play an important role in biotic and abiotic stress responses, which is one of the most active research topics in plant biology, especially their effect on agronomic traits. However, regarding the light spectrum, it is not surprising to consider that other photoreceptors are also part of the stress response modulated by light. In fact, it has become increasingly evident that cryptochromes, which mainly absorb in the blue light region, also act as key regulators of a range of plant stress responses, such as drought, salinity, heat, and high radiation. However, this information is rarely evidenced in photomorphogenetic studies. Therefore, the scope of the present review is to compile and discuss the evidence on the abiotic stress responses in plants that are modulated by cryptochromes.

Phototropins (phot1 and phot2) are blue-light receptors that control cotyledon flattening and positioning under strong light; however, their functional redundancy restricts our understanding of the specific roles of phot2. To identify the factors responsible for phot2-dependent cotyledon flattening and growth, we screened for light-insensitive mutants among mutagenized phot1 mutants in Arabidopsis thaliana. The double mutant phot1 lea1 (leaf expansion associated 1), which is defective in cotyledon flattening and positioning but not the phototropic response was selected. This mutant phenotype could be alleviated by constitutively expressing MORE AXILLARY GROWTH 2 (MAX2), indicating that LEA1 was allelic to MAX2. The max2 mutants (max2-2 and max2-3) are defective in cotyledon flattening, which is similar to that of the phot1 phot2 mutants. Moreover, the amounts of MAX2 transcripts are inhibited in leaves of phot1 mutant. However, the additional disruption of PHOT1 gene in max2-2 or max2-3 did not affect their phenotype, including MAX2-mediated inhibition of hypocotyl elongation. By contrast, phototropins-mediated hypocotyl phototropism was not regulated by MAX2. Together, these results suggest that cotyledon flattening was mediated by both phototropins and MAX2 signaling, but the relationship between two pathways need further study.

Rising temperatures during growing seasons coupled with altered precipitation rates presents a challenging task of improving crop productivity for overcoming such altered weather patterns and cater to a growing population. Light is a critical environmental factor that exerts a powerful influence on plant growth and development ranging from seed germination to flowering and fruiting. Higher plants utilize a suite of complex photoreceptor proteins to perceive surrounding red/far-red (phytochromes), blue/UV-A (cryptochromes, phototropins, ZTL/FKF1/LKP2), and UV-B light (UVR8). While genomic studies have also shown that light induces extensive reprogramming of gene expression patterns in plants, molecular genetic studies have shown that manipulation of one or more photoreceptors can result in modification of agronomically beneficial traits. Such information can assist researchers to engineer photoreceptors via genome editing technologies to alter expression or even sensitivity thresholds of native photoreceptors for targeting aspects of plant growth that can confer superior agronomic value to the engineered crops. Here we summarize the agronomically important plant growth processes influenced by photoreceptors in crop species, alongwith the functional interactions between different photoreceptors and phytohormones in regulating these responses. We also discuss the potential utility of synthetic biology approaches in photobiology for improving agronomically beneficial traits of crop plants by engineering designer photoreceptors.

Phytochromes are the principle photoreceptors in light-regulated plant development, primarily acting via translocation of the light-activated photoreceptor into the nucleus and subsequent gene regulation. However, several independent lines of evidence indicate unambiguously that an additional cytoplasmic signaling mechanism must exist. Directional responses in filament tip cells of the moss Physcomitrella patens are steered by phy4 which has been shown to interact physically with the blue light receptor phototropin at the plasma membrane. This complex might perceive and transduce vectorial information leading to cytoskeleton reorganization and finally a directional growth response. We developed yeast two-hybrid procedures using photochemically functional, full-length phy4 as bait in Physcomitrella cDNA library screens and growth assays under different light conditions, revealing Pfr-dependent interactions possibly associated with phytochrome cytoplasmic signaling. Candidate proteins were then expressed in planta with fluorescent protein tags to determine their intracellular localization in darkness and red light. Of 14 candidates, 12 were confirmed to interact with phy4 in planta using bimolecular fluorescence complementation. We also used database information to study their expression patterns relative to those of phy4. We discuss the likely functional characteristics of these holophytochrome-interacting proteins (HIP’s) and their possible roles in signaling.

The phototropin blue light (BL) photoreceptors, phot1 and phot2, mediate a number of organellar, cellular, and tissue/organ-level responses in plants including: phototropism, chloroplast accumulation and avoidance, stomatal opening, leaf expansion, and petiole/leaf orientation (Liscum et al., 2014). Not surprisingly, given the contribution of these responses to the optimization of photosynthetic efficiency and plant growth, several studies have described an intimate relationship between phot function and plant fitness (Galen et al., 2004, 2007a,b; Takemiya et al., 2005; Goh et al., 2009; Crepy and Casal, 2015). So how do phots induce such a wide variety of critical physiological responses? Though this remains a largely unanswered question, one apparent common early response has received recurrent attention; namely, BL-induced intracellular movement of phots (Sakamoto and Briggs, 2002; Knieb et al., 2004; Kong et al., 2006, 2007, 2013a,b; Aihara et al., 2008; Han et al., 2008; Kong and Nagatani, 2008; Sullivan et al., 2008, 2010, 2016b; Wan et al., 2008; Zienkiewicz et al., 2008; Kaiserli et al., 2009; Ding et al., 2011; Aggarwal et al., 2014; Kami et al., 2014; Preuten et al., 2015; Ishishita et al., 2016). Key observations made during these studies will be discussed here, keeping always in mind an important question: Does, or will, the study of phot movement improve our fundamental understanding of phot function, or are these events merely a red herring?

International audience; Guard cells are specialized cells located at the leaf surface delimiting pores whichcontrol gas exchanges between the plant and the atmosphere. To optimize the CO$_2$uptake necessary for photosynthesis while minimizing water loss, guard cells integrateenvironmental signals to adjust stomatal aperture. The size of the stomatal pore isregulated by movements of the guard cells driven by variations in their volume andturgor. As guard cells perceive and transduce a wide array of environmental cues, theyprovide an ideal system to elucidate early events of plant signaling. Reversible proteinphosphorylation events are known to play a crucial role in the regulation of stomatalmovements. However, in some cases, phosphorylation alone is not sufficient to achievecomplete protein regulation, but is necessary to mediate the binding of interactorsthat modulate protein function. Among the phosphopeptide-binding proteins, the 14-3-3 proteins are the best characterized in plants. The 14-3-3s are found as multipleisoforms in eukaryotes and have been shown to be involved in the regulation of stomatalmovements. In this review, we describe the current knowledge about 14-3-3 roles inthe regulation of their binding partners in guard cells: receptors, ion pumps, channels,protein kinases, and some of their substrates. Regulation of these targets by 14-3-3 proteins is discussed and related to their function in guard cells during stomatalmovements in response to abiotic or biotic stresses.

During the course of green plant evolution, numerous light responses have arisen that optimize their growth under fluctuating light conditions. The blue light receptor phototropin mediates several photomovement responses at the tissue, cellular and organelle levels. Chloroplast photorelocation movement is one such photomovement response, and is found not only in most green plants, but also in some red algae and photosynthetic stramenopiles. In general, chloroplasts move towards weak light to maximally capture photosynthetically active radiation (the chloroplast accumulation response), and they move away from strong light to avoid photodamage (the avoidance response). In land plants, chloroplast movement is dependent on specialized actin filaments, chloroplast-actin filaments (cp-actin filaments). Through molecular genetic analysis using Arabidopsis thaliana, many molecular factors that regulate chloroplast photorelocation were identified. In this Perspective, we discuss the evolutionary history of the molecular mechanism for chloroplast photorelocation movement in green plants in view of cp-actin filaments.

Plant phototropism, the ability to bend toward or away from light, is predominantly controlled by blue-light photoreceptors, the phototropins. Although phototropins have been well-characterized in Arabidopsis thaliana, their evolutionary history is largely unknown. In this study, we complete an in-depth survey of phototropin homologs across land plants and algae using newly available transcriptomic and genomic data. We show that phototropins originated in an ancestor of Viridiplantae (land plants + green algae). Phototropins repeatedly underwent independent duplications in most major land-plant lineages (mosses, lycophytes, ferns, and seed plants), but remained single-copy genes in liverworts and hornworts—an evolutionary pattern shared with another family of photoreceptors, the phytochromes. Following each major duplication event, the phototropins differentiated in parallel, resulting in two specialized, yet partially overlapping, functional forms that primarily mediate either low- or high-light responses. Our detailed phylogeny enables us to not only uncover new phototropin lineages, but also link our understanding of phototropin function in Arabidopsis with what is known in Adiantum and Physcomitrella (the major model organisms outside of flowering plants). We propose that the convergent functional divergences of phototropin paralogs likely contributed to the success of plants through time in adapting to habitats with diverse and heterogeneous light conditions.