Acceleration Of Flowering Research Articles

Functional analysis of PHYB polymorphisms in Arabidopsis thaliana collected in Patagonia.

Arabidopsis thaliana shows a wide range of natural genetic variation in light responses. Shade avoidance syndrome is a strategy of major adaptive significance that includes seed germination, elongation of vegetative structures, leaf hyponasty, and acceleration of flowering. Previously, we found that the southernmost Arabidopsis accession, collected in the south of Patagonia (Pat), is hyposensitive to light and displays a reduced response to shade light. This work aimed to explore the genetic basis of the shade avoidance response (SAR) for hypocotyl growth by QTL mapping in a recently developed 162 RIL population between Col-0 and Pat. We mapped four QTL for seedling hypocotyl growth: WL1 and WL2 QTL in white light, SHADE1 QTL in shade light, and SAR1 QTL for the SAR. PHYB is the strongest candidate gene for SAR1 QTL. Here we studied the function of two polymorphic indels in the promoter region, a GGGR deletion, and three non-synonymous polymorphisms on the PHYB coding region compared with the Col-0 reference genome. To decipher the contribution and relevance of each PHYB-Pat polymorphism, we constructed transgenic lines with single or double polymorphisms by using Col-0 as a reference genome. We found that single polymorphisms in the coding region of PHYB have discrete functions in seed germination, seedling development, and shade avoidance response. These results suggest distinct functions for each PHYB polymorphism to the adjustment of plant development to variable light conditions.

EARLY FLOWERING 3 and Photoperiod Sensing in Brachypodium distachyon.

The proper timing of flowering, which is key to maximize reproductive success and yield, relies in many plant species on the coordination between environmental cues and endogenous developmental programs. The perception of changes in day length is one of the most reliable cues of seasonal change, and this involves the interplay between the sensing of light signals and the circadian clock. Here, we describe a Brachypodium distachyon mutant allele of the evening complex protein EARLY FLOWERING 3 (ELF3). We show that the elf3 mutant flowers more rapidly than wild type plants in short days as well as under longer photoperiods but, in very long (20 h) days, flowering is equally rapid in elf3 and wild type. Furthermore, flowering in the elf3 mutant is still sensitive to vernalization, but not to ambient temperature changes. Molecular analyses revealed that the expression of a short-day marker gene is suppressed in elf3 grown in short days, and the expression patterns of clock genes and flowering time regulators are altered. We also explored the mechanisms of photoperiodic perception in temperate grasses by exposing B. distachyon plants grown under a 12 h photoperiod to a daily night break consisting of a mixture of red and far-red light. We showed that 2 h breaks are sufficient to accelerate flowering in B. distachyon under non-inductive photoperiods and that this acceleration of flowering is mediated by red light. Finally, we discuss advances and perspectives for research on the perception of photoperiod in temperate grasses.

The two clock proteins CCA1 and LHY activate VIN3 transcription during vernalization through the vernalization-responsive cis-element.

Vernalization, a long-term cold-mediated acquisition of flowering competence, is critically regulated by VERNALIZATION INSENSITIVE 3 (VIN3), a gene induced by vernalization in Arabidopsis. Although the function of VIN3 has been extensively studied, how VIN3 expression itself is upregulated by long-term cold is not well understood. In this study, we identified a vernalization-responsive cis-element in the VIN3 promoter, VREVIN3, composed of a G-box and an evening element (EE). Mutations in either the G-box or the EE prevented VIN3 expression from being fully induced upon vernalization, leading to defects in the vernalization response. We determined that the core clock proteins CIRCADIAN CLOCK-ASSOCIATED 1 (CCA1) and LATE-ELONGATED HYPOCOTYL (LHY) associate with the EE of VREVIN3, both in vitro and in vivo. In a cca1 lhy double mutant background harboring a functional FRIGIDA allele, long-term cold-mediated VIN3 induction and acceleration of flowering were impaired, especially under mild cold conditions such as at 12°C. During prolonged cold exposure, oscillations of CCA1/LHY transcripts were altered, while CCA1 abundance increased at dusk, coinciding with the diurnal peak of VIN3 transcripts. We propose that modulation of the clock proteins CCA1 and LHY participates in the systems involved in sensing long-term cold for the activation of VIN3 transcription.

Phenological features of development for the apricot cultivars in connection with the changing climate

The aim of the research was to study the influence of weather and climatic environmental factors on the formation of the apricot productivity of various origins. The objects of research were 45 apricot cultivars planted on the collection area in the Nikitsky Botanical Gardens. Phenological observations, yield accepting and determination of plant productivity of different apricot cultivars were done in accordance with generally accepted methods. When analyzing the data of phenological observations during the past 13 years due to climate change, acceleration of flowering and ripening have not been identified. The excess of monthly average precipitation with low monthly average air temperatures leads to a delay of the beginning of flowering of apricot plants by 1-3 weeks. The excess of monthly average air temperature during the period of formation (April-May) and fruit ripening (June-July), as well as a significant decrease in the amount of precipitation relative to their long-term average, accelerates the fruit ripening period. For the studied cultivars, it ranged from 1 to 3 weeks, in comparison with long-term data. Apricot cultivars and forms that better adapted to new growing conditions and have several valuable characteristics were selected: Large Early, Bucuria, Magister, Mandule Kajszi, Sanatate, Stokk, Ketch-Pshar (early or late ripening is combined with high yield; later flowering is combined with early or late fruit ripening). They are of interest for production testing, amateur gardeners and breeders.

No evidence of flowering synchronization upon floral volatiles for a short lived annual plant species: revisiting an appealing hypothesis

BackgroundSelf-incompatible plants require simultaneous flowering mates for crosspollination and reproduction. Though the presence of flowering conspecifics and pollination agents are important for reproductive success, so far no cues that signal the flowering state of potential mates have been identified. Here, we empirically tested the hypothesis that plant floral volatiles induce flowering synchrony among self-incompatible conspecifics by acceleration of flowering and flower opening rate of non-flowering conspecifics. We exposed Brassica rapa Maarssen, a self-incompatible, in rather dense patches growing annual, to (1) flowering or non-flowering conspecifics or to (2) floral volatiles of conspecifics by isolating plants in separate containers with a directional airflow. In the latter, odors emitted by non-flowering conspecifics were used as control.ResultsDate of first bud, duration of first flower bud, date of first flower, maximum number of open flowers and flower opening rate were not affected by the presence of conspecific flowering neighbors nor by floral volatiles directly.ConclusionsThis study presents a compelling approach to empirically test the role of flower synchronization by floral volatiles and challenges the premises that are underlying this hypothesis. We argue that the life history of the plant as well as its interaction with pollinators and insect herbivores, as well as the distance over which volatiles may serve as synchronization cue, set constraints on the fitness benefits of synchronized flowering which needs to be taken into account when testing the role of floral volatiles in synchronized flowering.

Light spectral and thermal properties govern biomass allocation in tomato through morphological and physiological changes

We investigated the effects of light spectral and thermal properties on biomass allocation among vegetative and reproductive structures in tomato (Solanum lycopersicum L. ‘Merlice’) plants during growth and development and to determine its underlying mechanisms. Plants were grown in a greenhouse under a DLI of 3.4 mol m2 day−1 with/without supplemental lighting and a 16-h photoperiod, provided at equal photosynthetic photon flux density (PPFD of 230 μmol m‒2 s‒1 either from overhead high-pressure sodium (HPS) lamps or intracanopy light-emitting diode (LED) arrays: red light alone (R100; 100% of photosynthetically active radiation (%PAR)), red plus low far-red (R100+FR21), red plus high far-red (R100+FR41), and red plus blue (R61+B39). Supplemental lighting substantially increased total biomass and fruit yield, but to a greater extent by LED lighting regardless of spectrum. However, spectral contribution to leaf morphological traits and biomass allocation pattern varied greatly among the light treatments. HPS-supplemented plants had higher leaf temperature, photosynthetic rate (Pn), and transpiration rate (E) in the upper foliar canopy (161 to 240 cm from the base of the shoot) and contained higher water content in all tissues, particularly in roots by 2-fold, compared to LED-supplemented plants. They also had thinner, smaller leaves and allocated a higher fraction of total biomass to vegetative tissues, demonstrating their allocation strategies for effective transpiration and heat dissipation. Meanwhile, LED-supplemented plants allocated biomass preferentially to reproductive structures at the expense of vegetative tissues. Inclusion of higher FR or B with R light decreased leaf biomass fraction relative to total biomass but increased leaf dry mass and leaf thickness. Particularly, supplemental FR light significantly lowered the water content of leaves and fruits compared to B light. Long-term exposure of tomato plants to low R:FR was associated with reductions in leaf area, chlorophyll content, and vegetative shoot biomass fraction, increases in leaf thickness and fruit biomass, and acceleration of flowering and fruiting. Light-induced biomass allocation changed between vegetative and reproductive structures during plant growth and development. FR light had the most pronounced effect in this regard by significantly shifting dry-mass accumulation to fruits, via more efficient photosynthetic mechanisms and conserved water use. We conclude that light spectral and thermal properties affect biomass allocation among plant parts during tomato growth and development, and such responses involve morphological and physiological changes in tomato plants, ultimately affecting crop performance and yield.

Epigenetic regulation of agronomical traits in Brassicaceae.

Epigenetic regulation, covalent modification of DNA and changes in histone proteins are closely linked to plant development and stress response through flexibly altering the chromatin structure to regulate gene expression. In this review, we will illustrate the importance of epigenetic influences by discussing three agriculturally important traits of Brassicaceae. (1) Vernalization, an acceleration of flowering by prolonged cold exposure regulated through epigenetic silencing of a central floral repressor, FLOWERING LOCUS C. This is associated with cold-dependent repressive histone mark accumulation, which confers competency of consequence vegetative-to-reproductive phase transition. (2) Hybrid vigor, in which an F1 hybrid shows superior performance to the parental lines. Combination of distinct epigenomes with different DNA methylation states between parental lines is important for increase in growth rate in a hybrid progeny. This is independent of siRNA-directed DNA methylation but dependent on the chromatin remodeler DDM1. (3) Self-incompatibility, a reproductive mating system to prevent self-fertilization. This is controlled by the S-locus consisting of SP11 and SRK which are responsible for self/non-self recognition. Because self-incompatibility in Brassicaceae is sporophytically controlled, there are dominance relationships between S haplotypes in the stigma and pollen. The dominance relationships in the pollen rely on de novo DNA methylation at the promoter region of a recessive allele, which is triggered by siRNA production from a flanking region of a dominant allele.

Stress-induced flowering: the third category of flowering response.

The switch from vegetative growth to reproductive growth, i.e. flowering, is the critical event in a plant's life. Flowering is regulated either autonomously or by environmental factors; photoperiodic flowering, which is regulated by the duration of the day and night periods, and vernalization, which is regulated by low temperature, have been well studied. Additionally, it has become clear that stress also regulates flowering. Diverse stress factors can induce or accelerate flowering, or inhibit or delay it, in a wide range of plant species. This article focuses on the positive regulation of flowering via stress, i.e. the induction or acceleration of flowering in response to stress that is known as stress-induced flowering - a new category of flowering response. This review aims to clarify the concept of stress-induced flowering and to summarize the full range of characteristics of stress-induced flowering from a predominately physiological perspective.

Enhanced flowering of the F1 long-day strawberry cultivars ‘Tarpan’ and ‘Gasana’ with nitrogen and daylength management

Interest in sustainable self-sufficiency, particularly when considering produce has increased. Strawberries, especially particularly the seed propagated, long-day, F1 hybrids such as ‘Tarpan’ and ‘Gasana’, fit self-sufficient, sustainable production models well. They provide aesthetic as well as culinary benefits producing colorful, attractive blossoms and flavorful fruit. To determine if ‘Tarpan’ and ‘Gasana’ flowering would respond to photoperiod and nitrogen, seedlings were fertilized with 100 or 800 ppm nitrogen for 4 weeks in September beginning one week after exposure to either short days, the natural photoperiod, or long days, the natural photoperiod supplemented with 24 hours of incandescent radiation. Plants were then greenhouse forced under both photoperiods and floral phenology evaluated. Photoperiod and N fertility during floral initiation in September affected subsequent flowering of both cultivars. The response was rapid with significant differences observed 4 weeks after the commencement of treatment. Both cultivars responded with increased rate (enhanced precocity) and intensity (enhanced inflorescence/flower number) of flowering with elevated N. In ‘Gasana’ elevated N accelerated flowering by 2–3 weeks. There was a slight acceleration of flowering in lower N plants with long-day forcing, however, elevated N was much more effective in accelerating flowering than long-day forcing. In ‘Tarpan’, long-day forcing and elevated N were equally effective in accelerating flowering. Inflorescence production per plant or crown and the number of flowers per plant were enhanced with elevated N in both cultivars. Long-day forcing stimulated the number of inflorescences produced per crown in both cultivars. The number of flowers per plant in ‘Tarpan’ was also enhanced by long-day initiation or forcing. Elevated N, short-day initiation or long-day forcing enhanced the number of flowers per inflorescence in ‘Gasana’. Elevated N under long-day initiation or under short-day initiation when followed by long-day forcing enhanced the number of flowers per inflorescence in ‘Tarpan’.

Changes in the germination process and growth of pea in effect of laser seed irradiation

Abstract The aim of this study was to determine the effect of pre-sowing helium-neon (He-Ne) laser irradiation of pea seeds on changes in seed biochemical processes, germination rate, seedling emergence, growth rate, and yield. The first experimental factor was exposure to laser radiation: D0 - no irradiation, D3 - three exposures, D5 - five exposures, and the harvest dates were the second factor. Pre-sowing treatment of pea seeds with He-Ne laser light increased the concentrations of amylolytic enzymes and the content of indole-3-acetic acid (IAA) in pea seeds and seedlings. The exposure of seeds to He-Ne laser light improved the germination rate and uniformity and modified growth stages, which caused acceleration of flowering and ripening of pea plants. Laser light stimulation improved the morphological characteristics of plants by increasing plant height and leaf surface area. Irradiation improved the yield of vegetative and reproductive organs of pea, although the effects varied at the different growth stages. The increase in the seed yield resulted from a higher number of pods and seeds per plant, whereas no significant changes were observed in the number of seeds per pod. Both radiation doses exerted similarly stimulating effects on pea growth, development, and yield.

Genetic and epigenetic control of plant heat responses.

Plants have evolved sophisticated genetic and epigenetic regulatory systems to respond quickly to unfavorable environmental conditions such as heat, cold, drought, and pathogen infections. In particular, heat greatly affects plant growth and development, immunity and circadian rhythm, and poses a serious threat to the global food supply. According to temperatures exposing, heat can be usually classified as warm ambient temperature (about 22–27°C), high temperature (27–30°C) and extremely high temperature (37–42°C, also known as heat stress) for the model plant Arabidopsis thaliana. The genetic mechanisms of plant responses to heat have been well studied, mainly focusing on elevated ambient temperature-mediated morphological acclimation and acceleration of flowering, modulation of circadian clock and plant immunity by high temperatures, and thermotolerance to heat stress. Recently, great progress has been achieved on epigenetic regulation of heat responses, including DNA methylation, histone modifications, histone variants, ATP-dependent chromatin remodeling, histone chaperones, small RNAs, long non-coding RNAs and other undefined epigenetic mechanisms. These epigenetic modifications regulate the expression of heat-responsive genes and function to prevent heat-related damages. This review focuses on recent progresses regarding the genetic and epigenetic control of heat responses in plants, and pays more attention to the role of the major epigenetic mechanisms in plant heat responses. Further research perspectives are also discussed.

Effect of the hope FT-B1 allele on wheat heading time and yield components.

Precise regulation of flowering time is critical for plant reproductive success and, in cereals, to maximize grain yields. Seasonal cues including temperature and day length are integrated to regulate the timing of flowering. In temperate cereals, extended periods of cold (vernalization) release the repression of FLOWERING LOCUS T1 (FT1), which is upregulated in the leaves in response to inductive long-day photoperiods. FT1 is a homolog of rice HD3a, which encodes a protein transported from leaves to the shoot apical meristem to induce flowering. A rare FT-B1 allele from the wheat variety "Hope" has been previously shown to be associated with an early flowering phenotype under long-day photoperiods. Here, we demonstrate that the Hope FT-B1 allele accelerates flowering even under short days, and that it is epistatic to the VERNALIZATION 1 (VRN1) gene. On average, the introgression of Hope FT-B1 into 6 genetic backgrounds resulted in 2.6 days acceleration of flowering (P<0.0001) and 4.1% increase in spike weight (P=0.0093), although in one variety, it was associated with a decrease in spike weight. These results suggest that the Hope FT-B1 allele could be useful in wheat breeding programs to subtly accelerate floral development and increase adaptation to changing environments.

Multiple FLC haplotypes defined by independent cis-regulatory variation underpin life history diversity in Arabidopsis thaliana

Relating molecular variation to phenotypic diversity is a central goal in evolutionary biology. In Arabidopsis thaliana, FLOWERING LOCUS C (FLC) is a major determinant of variation in vernalization--the acceleration of flowering by prolonged cold. Here, through analysis of 1307 A. thaliana accessions, we identify five predominant FLC haplotypes defined by noncoding sequence variation. Genetic and transgenic experiments show that they are functionally distinct, varying in FLC expression level and rate of epigenetic silencing. Allelic heterogeneity at this single locus accounts for a large proportion of natural variation in vernalization that contributes to adaptation of A. thaliana.

Recovery of dicer-like 1-late flowering phenotype by miR172 expressed by the noncanonical DCL4-dependent biogenesis pathway.

MicroRNAs (miRNAs) act as down-regulators of gene expression, and play a dominant role in eukaryote development. In Arabidopsis thaliana, DICER-LIKE 1 (DCL1) is the main processor in miRNA biogenesis, and dcl1 mutants show various developmental defects at the early stage of embryogenesis or at gamete formation. However, miRNAs responsible for the respective developmental stages of the dcl1 defects have not been identified. Here, we developed a DCL1-independent miRNA expression system using the unique DCL4-dependent miRNA, miR839. By replacing the mature sequence in the miR839 precursor sequence with that of miR172, one of the most widely conserved miRNAs in angiosperms, we succeeded in expressing miR172 from a chimeric miR839 precursor in dcl1-7 plants and observed the repression of miR172 target gene expression. In parallel, the DCL4-dependent miR172 expression rescued the late flowering phenotype of dcl1-7 by acceleration of flowering. We established the DCL1-independent miRNA expression system, and revealed that the reduction of miR172 expression is responsible for the dcl1-7 late flowering phenotype.

Elevated Levels of MYB30 in the Phloem Accelerate Flowering in Arabidopsis through the Regulation of FLOWERING LOCUS T

In Arabidopsis thaliana, the R2R3 MYB-like transcription factor MYB30 is a positive regulator of the pathogen-induced hypersensitive response and of brassinosteroid and abscisic acid signaling. Here, we show that MYB30 expressed under the control of the strong phloem-specific SUC2 promoter accelerates flowering both in long and short days. Early flowering is mediated by elevated expression of FLOWERING LOCUS T (FT), which can be observed in the absence and presence of CONSTANS (CO), the main activator of FT. CO-independent activation by high MYB30 expression results in FT levels that remain below those observed in the wild-type plants, which show an additive CO-dependent activation. In contrast, TWIN SISTER OF FT (TSF) is repressed in plants expressing high levels of MYB30 in the phloem. In transient assays, MYB30 and CO additively increase the activity of a reporter construct driven by a 1 kb FT promoter. Acceleration of flowering by MYB30 does not require the presence of salicylic acid and is independent of FLC. Taken together, increased levels of MYB30, which was reported to be induced in response to the perception of pathogens, can accelerate flowering and MYB30 may thus be a candidate to mediate cross-talk between gene networks involved in biotic stress perception and flowering time.

Acceleration of flowering in Arabidopsis thaliana by Cape Verde Islands alleles of FLOWERING H is dependent on the floral promoter FD

Flowering time in the model plant Arabidopsis thaliana is regulated by both external environmental signals and internal developmental pathways. Natural variation at the FLOWERING H (FLH) locus has previously been described, with alleles present in the Cape Verde Islands accession causing early flowering, particularly after vernalization. The mechanism of FLH-induced early flowering is not understood. Here, the integration of FLH activity into the known flowering time pathways is described using molecular and genetic approaches. The identification of molecular markers that co-segregated with the FLH locus allowed the generation of multiple combinations of FLH alleles with mutations in flowering time genes in different flowering pathways. Combining an early flowering FLH allele with mutations in vernalization pathway genes that regulate FLC expression revealed that FLH appears to act in parallel to FLC. Surprisingly, the early flowering allele of FLH requires the floral integrator FD, but not FT, to accelerate flowering. This suggests a model in which some alleles of FLH are able to affect the FD-dependent activity of the floral activator complex.

The receptor-like kinase ERECTA contributes to the shade-avoidance syndrome in a background-dependent manner.

Plants growing at high densities perceive a decrease in the red to far-red (R/FR) ratio of incoming light. These changes in light quality trigger a suite of responses collectively known as the shade-avoidance syndrome (SAS) including hypocotyl and stem elongation, inhibition of branching and acceleration of flowering. Quantitative trait loci (QTLs) were mapped for hypocotyl length to end-of-day far-red (EOD), a simulated shade-avoidance response, in recombinant inbred line (RIL) populations of Arabidopsis thaliana seedlings, derived from Landsberg erecta (Ler) and three accessions (Columbia, Col; Nossen, No-0; and Cape Verde Islands, Cvi-0). Five loci were identified as being responsible for the EOD response, with a positive contribution of Ler alleles on the phenotype independently of the RIL population. Quantitative complementation analysis and transgenic lines showed that PHYB is the candidate gene for EODRATIO5 in the Ler × Cvi-0 RIL population, but not for two co-localized QTLs, EODRATIO1 and EODRATIO2 mapped in the Ler × No-0 and Ler × Col RIL populations, respectively. The ERECTA gene was also implicated in the SAS in a background-dependent manner. For hypocotyl length EOD response, a positive contribution of erecta alleles was found in Col and Van-0, but not in Ler, Cvi-0, Hir-1 or Ws. Furthermore, pleiotropic effects of ERECTA in the EOD response were also detected for petiole and lamina elongation, hyponastic growth, and flowering time. The results show that the analysis of multiple mapping populations leads to a better understanding of the SAS genetic architecture. Moreover, the background- and trait-dependent contribution of ERECTA in the SAS suggest that its function in shaded natural environments may be relevant for some populations in different phases of plant development. It is proposed that ERECTA is involved in canalization processes buffering the genetic variation of the SAS against environmental light fluctuations.

Control of Tiller Growth of Rice by OsSPL14 and Strigolactones, Which Work in Two Independent Pathways

The architecture of rice is greatly influenced by the growth of tillers, i.e. vegetative shoot branches. OsSPL14, a member of the SQUAMOSA PROMOTER BINDING PROTEIN-LIKE (SPL) genes, and strigolactones (SLs) are known to suppress outgrowth of tiller buds. OsSPL14 also regulates panicle development. We show that OsSPL14 mRNA accumulates in leaf primordia during the vegetative phase and in the primordia of bracts, or modified leaves, in the panicles, but not in the meristems. OsSPL14 is a target of miR156, and accumulation of OsSPL14 transcripts is negatively regulated by miR156. The enhancement of the expression level of OsSPL14 by the introduction of the mOsSPL14 gene, in which the miR156 cleavage site is mutated, resulted in an increase in the plastochron, an acceleration of flowering and a decrease in tiller number in the wild type and in dwarf10-2, an SL-deficient mutant. Our analysis suggests that OsSPL14 and SLs function in parallel pathways to suppress tiller growth. SLs exuded from roots trigger germination of root parasitic plants that can cause severe damage to crop productivity. SL-deficient mutants, however, exhibit an excess branching phenotype which is usually undesirable for productivity. Our results indicate that OsSPL14 can be used to manipulate the branching patterns of SL-deficient mutants. We also confirmed that this strategy is applicable to Arabidopsis. A greater understanding of the OsSPL14 and SL pathways and their interactions may help in the production of root parasite-resistant crops.

Double loss-of-function mutation in EARLY FLOWERING 3 and CRYPTOCHROME 2 genes delays flowering under continuous light but accelerates it under long days and short days: an important role for Arabidopsis CRY2 to accelerate flowering time in continuous light

The photoperiodic response is one of the adaptation mechanisms to seasonal changes of lengths of day and night. The circadian clock plays pivotal roles in this process. In Arabidopsis, LHY, CCA1, ELF3, and other clock proteins play major roles in maintaining circadian rhythms. lhy;cca1 double mutants with severe defects in circadian rhythms showed accelerated flowering under short days (SDs), but delayed flowering under continuous light (LL). The protein level of the floral repressor SVP increased in lhy;cca1 mutants under LL, and the late-flowering phenotype of lhy;cca1 mutants was partially suppressed by svp, flc, or elf3. ELF3 interacted with both CCA1 and SVP, and elf3 suppressed the SVP accumulation in lhy;cca1 under LL. These results suggest that the unique mechanism of the inversion of the flowering response of lhy;cca1 under LL may involve both the ELF3-SVP/FLC-dependent and -independent pathways. In this work, elf3-1 seeds were mutagenized with heavy-ion beams and used to identify mutation(s) that delayed flowering under LL but not long days (LDs) or SDs even without ELF3. In this screening, seven candidate lines named suppressor of elf3 1 (self1), sel3, sel5, sel7, sel14, sel15, and sel20 were identified. Genetic analysis indicated that sel20 was a new deletion allele of a mutation in the blue light receptor, CRY2. A late-flowering phenotype and decrease of FT expression in the elf3;sel20 double mutant was obvious under LL but not under SDs or LDs. These results indicated that the late-flowering phenotype in the double mutant elf3;sel20 as well as in lhy;cca1 was affected by the presence of darkness. The results suggest that CRY2 may play more essential roles in the acceleration of flowering under LL than LDs or SDs.

Function of B-BOX under shade

Plants are capable of perceiving changes in the light environment and finely adjust their growth and development. Reductions of red to far-red ratio (R:FR) generated by an increase of the plant canopy above the plant are sensed by the phytochrome system triggering the shade-avoidance syndrome (SAS) that includes elongation of vegetative structures, reduction of branching and acceleration of flowering. Albeit the SAS is a strategy of major adaptative significance in plant communities, involving massive changes in gene expression, our knowledge of the SAS signaling network is still fragmented. By a selection and characterization of a T-DNA mutant with a long hypocotyl under shade, we identified BBX21, a protein with two B-box domains involved in the SAS. BBX21 belongs to a small eight member family of B-box containing proteins with both opposite and additive functions in the SAS signaling. BBX21 down-regulates the gene expression of auxin, brassinosteroid and ethylene signaling pathway components under shade. Furthermore BBX21 is a transcription factor that interacts genetically with COP1. We propose a model in which a dynamic balance of positive and negative B-box transcriptional regulators acts as a gas-and-brake mechanism into the COP1 signaling to regulate the expression of SAS.