Shade Avoidance 3 Mediates Crosstalk Between Shade and Nitrogen in Arabidopsis Leaf Development

After nitrogen treatments, plant leaves become narrower and thicker, and the chlorophyll content increases. However, the molecular mechanisms behind these regulations remain unknown. Here, we found that the changes in leaf width and thickness were largely compromised in the shade avoidance 3 (sav3) mutant. The SAV3 gene encodes an amino-transferase in the auxin biosynthesis pathway. Thus, the crosstalk between shade and nitrogen in Arabidopsis leaf development was investigated. Both hypocotyl elongation and leaf expansion promoted by the shade treatment were reduced by the high-N treatment; high-N-induced leaf narrowing and thickening were reduced by the shade treatment; and all of these developmental changes were largely compromised in the sav3 mutant. Shade treatment promoted SAV3 expression, while high-N treatment repressed SAV3 expression, which then increased or decreased auxin accumulation in cotyledons/leaves, respectively. SAV3 also regulates chlorophyll accumulation and nitrogen assimilation and thus may function as a master switch responsive to multiple environmental stimuli.

Basic Protein Modules Combining Abscisic Acid and Light Signaling in Arabidopsis

It is generally accepted that plants use the complex signaling system regulated by light and abscisic acid (ABA) signaling components to optimize growth and development in different situations. The role of ABA–light interactions is evident in the coupling of stress defense reactions with seed germination and root development, maintaining of stem cell identity and stem cell specification, stem elongation and leaf development, flowering and fruit formation, senescence, and shade avoidance. All these processes are regulated jointly by the ABA–light signaling system. Although a lot of work has been devoted to ABA–light signal interactions, there is still no systematic description of central signaling components and protein modules, which jointly regulate plant development. New data have emerged to promote understanding of how ABA and light signals are integrated at the molecular level, representing an extensively growing area of research. This work is intended to fill existing gaps by using literature data combined with bioinformatics analysis.

Understanding the Shade Tolerance Responses Through Hints From Phytochrome A-Mediated Negative Feedback Regulation in Shade Avoiding Plants

Based on how plants respond to shade, we typically classify them into two groups: shade avoiding and shade tolerance plants. Under vegetative shade, the shade avoiding species induce a series of shade avoidance responses (SARs) to outgrow their competitors, while the shade tolerance species induce shade tolerance responses (STRs) to increase their survival rates under dense canopy. The molecular mechanism underlying the SARs has been extensively studied using the shade avoiding model plant Arabidopsis thaliana, while little is known about STRs. In Aarabidopsis, there is a PHYA-mediated negative feedback regulation that suppresses exaggerated SARs. Recent studies revealed that in shade tolerance Cardamine hirsuta plants, a hyperactive PHYA was responsible for suppressing shade-induced elongation growth. We propose that similar signaling components may be used by shade avoiding and shade tolerance plants, and different phenotypic outputs may result from differential regulation or altered dynamic properties of these signaling components. In this review, we summarized the role of PHYA and its downstream components in shade responses, which may provide insights into understanding how both types of plants respond to shade.

Shade suppresses wound-induced leaf repositioning through a mechanism involving PHYTOCHROME KINASE SUBSTRATE (PKS) genes

Shaded plants challenged with herbivores or pathogens prioritize growth over defense. However, most experiments have focused on the effect of shading light cues on defense responses. To investigate the potential interaction between shade-avoidance and wounding-induced Jasmonate (JA)-mediated signaling on leaf growth and movement, we used repetitive mechanical wounding of leaf blades to mimic herbivore attacks. Phenotyping experiments with combined treatments on Arabidopsis thaliana rosettes revealed that shade strongly inhibits the wound effect on leaf elevation. By contrast, petiole length is reduced by wounding both in the sun and in the shade. Thus, the relationship between the shade and wounding/JA pathways varies depending on the physiological response, implying that leaf growth and movement can be uncoupled. Using RNA-sequencing, we identified genes with expression patterns matching the hyponastic response (opposite regulation by both stimuli, interaction between treatments with shade dominating the wound signal). Among them were genes from the PKS (Phytochrome Kinase Substrate) family, which was previously studied for its role in phototropism and leaf positioning. Interestingly, we observed reduced shade suppression of the wounding effect in pks2pks4 double mutants while a PKS4 overexpressing line showed constitutively elevated leaves and was less sensitive to wounding. Our results indicate a trait-specific interrelationship between shade and wounding cues on Arabidopsis leaf growth and positioning. Moreover, we identify PKS genes as integrators of external cues in the control of leaf hyponasty further emphasizing the role of these genes in aerial organ positioning.

Monoseeding Increases Peanut (Arachis hypogaea L.) Yield by Regulating Shade-Avoidance Responses and Population Density

We aimed to elucidate the possible yield-increasing mechanisms through regulation of shade-avoidance responses at both physiological and molecular levels under monoseeding. Our results revealed that monoseeding decreased the main stem height but increased the main stem diameter and the number of branches and nodes compared to the traditional double- and triple-seeding patterns. The chlorophyll contents were higher under monoseeding than that under double- and triple-seeding. Further analysis showed that this, in turn, increased the net photosynthetic rate and reallocated higher levels of assimilates to organs. Monoseeding induced the expression patterns of Phytochrome B (Phy B) gene but decreased the expression levels of Phytochrome A (Phy A) gene. Furthermore, the bHLH transcription factors (PIF 1 and PIF 4) that interact with the phytochromes were also decreased under monoseeding. The changes in the expression levels of these genes may regulate the shade-avoidance responses under monoseeding. In addition, monoseeding increased pod yield at the same population density through increasing the number of pods per plant and 100-pod weight than double- and triple-seeding patterns. Thus, we inferred that monoseeding is involved in the regulation of shade-avoidance responsive genes and reallocating assimilates at the same population density, which in turn increased the pod yield.

Ubiquitin-specific proteases UBP12 and UBP13 promote shade avoidance response by enhancing PIF7 stability

Changes in light quality caused by the presence of neighbor proximity regulate many growth and development processes of plants. PHYTOCHROME INTERACTING FACTOR 7 (PIF7), whose subcellular localization, DNA-binding properties, and protein abundance are regulated in a photoreversible manner, plays a central role in linking shade light perception and growth responses. How PIF7 activity is regulated during shade avoidance responses has been well studied, and many factors involved in this process have been identified. However, the detailed molecular mechanism by which shade light regulates the PIF7 protein level is still largely unknown. Here, we show that the PIF7 protein level regulation is important for shade-induced growth. Two ubiquitin-specific proteases, UBP12 and UBP13, were identified as positive regulators in shade avoidance responses by increasing the PIF7 protein level. The ubp12-2w/13–3 double mutant displayed significantly impaired sensitivity to shade-induced cell elongation and reproduction acceleration. Our genetic and biochemical analysis showed that UBP12 and UBP13 act downstream of phyB and directly interact with PIF7 to maintain PIF7 stability and abundance through deubiquitination.

Comparative phenotypic and transcriptomic analyses unravel conserved and distinct mechanisms underlying shade avoidance syndrome in Brassicaceae vegetables

Background Plants grown under shade are exposed to low red/far-red ratio, thereby triggering an array of altered phenotypes called shade avoidance syndrome (SAS). Shade negatively influences plant growth, leading to a reduction in agricultural productivity. Understanding of SAS is crucial for sustainable agricultural practices, especially for high-density indoor farming. Brassicaceae vegetables are widely consumed around the world and are commonly cultivated in indoor farms. However, our understanding of SAS in Brassicaceae vegetables and their genome-wide transcriptional regulatory networks are still largely unexplored. Results Shade induced common signs of SAS, including hypocotyl elongation and reduced carotenoids/anthocyanins biosynthesis, in two different Brassicaceae species: Brassica rapa (Choy Sum and Pak Choy) and Brassica oleracea (Kai Lan). Phenotype-assisted transcriptome analysis identified a set of genes induced by shade in these species, many of which were related to auxin biosynthesis and signaling [e.g. YUCCA8 (YUC8), YUC9, and INDOLE-3-ACETIC ACID INDUCIBLE (IAAs)] and other phytohormones signaling pathways including brassinosteroids and ethylene. The genes functioning in plant defense (e.g. MYB29 and JASMONATE-ZIM-DOMAIN PROTEIN 9) as well as in biosynthesis of anthocyanins and glucosinolates were repressed upon shade. Besides, each species also exhibited distinct SAS phenotypes. Shade strongly reduced primary roots and elongated petioles of B. oleracea, Kai Lan. However, these SAS phenotypes were not clearly recognized in B. rapa, Choy Sum and Pak Choy. Some auxin signaling genes (e.g. AUXIN RESPONSE FACTOR 19, IAA10, and IAA20) were specifically induced in B. oleracea, while homologs in B. rapa were not up-regulated under shade. Contrastingly, shade-exposed B. rapa vegetables triggered the ethylene signaling pathway earlier than B. oleracea, Kai Lan. Interestingly, shade induced the transcript levels of LONG HYPOCOTYL IN FAR-RED 1 (HFR1) homolog in only Pak Choy as B. rapa. As HFR1 is a key negative regulator of SAS in Arabidopsis, our finding suggests that Pak Choy HFR1 homolog may also function in conferring higher shade tolerance in this variety. Conclusions Our study shows that two Brassicaceae species not only share a conserved SAS mechanism but also exhibit distinct responses to shade, which will provide comprehensive information to develop new shade-tolerant cultivars that are suitable for high-density indoor farms.

Domestication reduces phenotypic plasticity in chaya (Cnidoscolus aconitifolius (Mill.) I.M. Johnst)

Background: Natural selection optimizes phenotypic plasticity in plants found in environmentally variable habitats. However, it is unclear how artificial selection has affected the phenotypic plasticity of crops. Reduced plasticity in crop yield is often considered a desirable feature in cultivated plants; however, limited phenotypic plasticity in this and other traits may also affect the ability of crops to cope with environmental variation. Study species: Wild and domesticated chaya (Cnidoscolus aconitifolius (Mill.) I.M. Johnst). Question: How domestication has affected the phenotypic plasticity of vegetative traits in response to the light environment? Methods: Leaf area, leaf perimeter, leaf specific area, leaf production, trichome density, stem elongation, growth in stem diameter and slenderness were measured in clones of wild and domesticated plants. These clones were allocated to two contrasting light treatments: fully exposed to sun vs. placement beneath a shade cloth. The phenotypic traits and reaction norms were compared between the genotypes of wild and domesticated plants. Results: Lower plasticity in leaf production and slenderness was observed in the domesticated compared to the wild plants. Leaf production and slenderness are associated with the shade avoidance syndrome, which was evident in wild plants but not manifested in domesticated plants. Reduced plasticity in leaf production also suggests yield stability. Conclusion: Artificial selection reduces phenotypic plasticity in the yield of chaya and in its response to variation in light availability.

Effects of partial replacement of red by green light in the growth spectrum on photomorphogenesis and photosynthesis in tomato plants

The artificial light used in growth chambers is usually devoid of green (G) light, which is considered to be less photosynthetically efficient than blue (B) or red (R) light. To verify the role of G light supplementation in the spectrum, we modified the RB spectrum by progressively replacing R light with an equal amount of G light. The tomato plants were cultivated under 100 µmol m–2 s–1 of five different combinations of R (35–75%) and G light (0–40%) in the presence of a fixed proportion of B light (25%) provided by light-emitting diodes (LEDs). Substituting G light for R altered the plant’s morphology and partitioning of biomass. We observed a decrease in the dry biomass of leaves, which was associated with increased biomass accumulation and the length of the roots. Moreover, plants previously grown under the RGB spectrum more efficiently utilized the B light that was applied to assess the effective quantum yield of photosystem II, as well as the G light when estimated with CO2 fixation using RB + G light-response curves. At the same time, the inclusion of G light in the growth spectrum reduced stomatal conductance (gs), transpiration (E) and altered stomatal traits, thus improving water-use efficiency. Besides this, the increasing contribution of G light in place of R light in the growth spectrum resulted in the progressive accumulation of phytochrome interacting factor 5, along with a lowered level of chalcone synthase and anthocyanins. However, the plants grown at 40% G light exhibited a decreased net photosynthetic rate (Pn), and consequently, a reduced dry biomass accumulation, accompanied by morphological and molecular traits related to shade-avoidance syndrome.

DcPAR1 is Required for Development and Carotenoid Synthesis in the Dark-Grown Carrot Taproot

Light stimulates carotenoid synthesis in plants during photomorphogenesis through the expression of PHYTOENE SYNTHASE (PSY), a key gene in carotenoid biosynthesis. The orange Daucus carota (carrot) synthesizes and accumulates high amounts of carotenoids in the taproot that grows underground. Contrary to other organs, light impairs carrot taproot development and represses the expression of carotenogenic genes such as DcPSY1 and DcPSY2 reducing carotenoid accumulation. By means of an RNA-seq, in previous analysis we observed that carrot PHYTOCHROME RAPIDLY REGULATED 1 (DcPAR1) is more expressed in the underground grown taproot respect to those grown in light. PAR1 is a transcriptional cofactor with a negative role in the shade avoidance syndrome regulation in Arabidopsis thaliana through the dimerization with PHYTOCHROME INTERACTING FACTORs (PIFs), allowing a moderate synthesis of carotenoids. Here we show that overexpressing AtPAR1 in carrot produces an increment of carotenoids in taproots grown underground as well as higher DcPSY1 expression. The high identity of AtPAR1 and DcPAR1 let us to suggest a functional role of DcPAR1 that was verified through the in vivo binding to AtPIF7 and the overexpression in Arabidopsis, where it increments AtPSY expression and carotenoid accumulation together with a photomorphogenic phenotype. Finally, DcPAR1 antisense carrot lines presented a dramatic decrease in carotenoids levels and in the relative expression of key carotenogenic genes as well as impairment in taproot development. These results let us to propose that DcPAR1 is a key factor for secondary root development, plastid differentiation and carotenoid synthesis in carrot taproot grown underground.One-sentence summaryDcPAR1 is a key factor for secondary root development, plastid differentiation and carotenoid synthesis in carrot taproot grown underground.