Non-inductive Photoperiods Research Articles

Supplemental Low-Irradiance Mono/Polychromatic LED Lighting Significantly Enhances Floral Biology of the Long-Day F1 Hybrid Strawberry ‘Soraya’ (Fragaria x ananassa Duch.)

Floral and vegetative responses of the strawberry (Fragaria x ananassa Duch.) to specific light wavelengths are not well documented. LED lights make it feasible for precise exposure to specific wavelengths during a 24 h cycle to alter growth responses regulated by phytochromes and cryptochromes and thereby potentially enhance fruit productivity in both a controlled environment and field systems or to enhance stolon production for controlled environment propagation. This research developed a systematic method to assess the effects of supplemental, low-irradiance LED lighting on strawberry flowering and vegetative biology. Growth of the long-day F1 seed-propagated cultivar ‘Soraya’ was evaluated during and following 6 or 12 weeks of exposure to supplemental red (660 nm), far-red (730 nm), blue (454 nm), or incandescent lighting at various times during the dark period of a 24 h cycle under a 10 h non-inductive photoperiod at non-inductive temperatures (>27/18 °C, day/night). Treatment effects were monitored via flower mapping and phenology during treatment, field and greenhouse production after treatment, and floral scores derived by ranking treatment effects within the evaluation method and then combining them into a single, simple score. The most promising treatment for enhancing the floral nature of plug plants was exposure to far-red + red light as a 5 h night interruption. This treatment increased inflorescence production in the greenhouse by 285% and resulted in multi-branched, floral plants with the potential for enhancing yield in either greenhouse or field production. Greenhouse runner production increased by 483% following exposure to incandescent lighting at the beginning of the dark period; thus, this treatment or one using a spectral distribution similar to incandescent may be suitable for enhancing vegetative propagation in controlled environments.

GmFT3a fine-tunes flowering time and improves adaptation of soybean to higher latitudes.

Onset of flowering of plants is precisely controlled by extensive environmental factors and internal molecular networks, in which FLOWERING LOCUS T (FT) is a key flowering integrator. In soybean, a typical short-day plant, 11 FT homologues are found in its genome, of which several homologues are functionally diversified in flowering pathways and the others including GmFT3a are yet unknown. In the current study, we characterized GmFT3a, which is located on the same chromosome as the flowering promoters GmFT2a and GmFT5a. Overexpression of GmFT3a significantly promoted flowering of Arabidopsis under the inductive long-day (LD) photoperiod. GmFT3a over-expressed soybean also flowered earlier than the control under LD, but they were not significantly different under inductive short-day (SD) conditions, indicating that GmFT3a acts as a flowering promoter in the non-inductive photoperiod in soybean. Compared with other GmFT homologues, GmFT3a exhibited a slighter effect in flowering promotion than GmFT2a, GmFT5a and GmFT2b under LD conditions. GmFT3a promoted flowering by regulating the expression of downstream flowering-related genes and also affected the expression of other GmFTs. According to the re-sequencing data, the regional distributions of two major haplotypes in 176 soybean varieties were analyzed. The varieties with GmFT3a-Hap2 haplotype matured relatively early, and relative higher expression of GmFT3a was detected in early maturing varieties, implying that Hap2 variation may contribute to the adaptation of soybean to higher latitude regions by increasing expression level of genes in metabolism and signaling pathways. The early flowering germplasm generated by overexpression of GmFT3a has potential to be planted at higher latitudes where non-inductive long day is dominant in the growing season, and GmFT3a can be used to fine-tune soybean flowering and maturity time and improve the geographical adaptation.

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.

Extended photoperiods after flowering increase the rate of dry matter production and nitrogen assimilation in mid maturing soybean cultivars

• Extended photoperiods after flowering prolonged reproductive growth. • Vegetative and reproductive biomass increased under extended photoperiods. • Crop growth rate increased, and duration of the growth period was lengthened. • Nitrogen assimilation increased with no change in tissue nitrogen concentration. In soybeans, exposure to non-inductive photoperiods ( i.e ., long or extended days) after flowering triggers a number of responses, including slower development of reproductive structures, enhanced vegetative growth and increased production of flowers. The larger number of flowers under extended photoperiods results in an increase of the number of fruits, which eventually may translate into greater grain yield. The main goal of this work was to assess whether C and N assimilation rates increase in soybeans subjected to long days after flowering. Soybean cultivars NS4619 and NS5019 were planted in the field during the normal planting season in 2016−17 and 2018−19. At flowering, two treatments were imposed: natural (Nat) or extended photoperiods (Ext) in which daylength was prolonged by 4 h with low intensity illumination from light-emitting diode lamps delivering less than 10 μmol m −2 s −1 of photosynthetically active radiation at the top of the canopy. Extended photoperiods prolonged reproductive growth, thereby delaying crop maturity. Growth of the main stem and branches was promoted by Ext, with plants showing greater number of nodes and main stem + branches dry weight at maturity. The number of pods and seeds increased by 69–87 % and 31–76 %, respectively, in Ext, and pod and seed dry weight increased significantly in 2016−17. Averaged over the whole reproductive period, crop growth rate increased under Ext in 2016−17 and 2018−2019. Ext promoted post-flowering vegetative growth more than growth of seeds, therefore harvest index was lower under Ext. The growth increase under Ext was larger if the biomass of abscised leaf blades and petioles was taken into account. Consistently with increased biomass under Ext, nitrogen accumulation by the crop was higher under extended photoperiods, with no dilution effect of N due to increased dry mass. About half of the N in plant tissues derived from biological nitrogen fixation, irrespective of photoperiodic condition, which implied that both, nitrogen fixation and uptake of soil nitrogen increased under long photoperiods. In brief, both the amounts of C and N assimilated by the crop were higher under Ext. Understanding the molecular, biochemical and physiological basis of post-flowering photoperiodic effects might offer clues to increase the yield potential of soybeans.

Systematic analyses of the MIR172 family members of Arabidopsis define their distinct roles in regulation of APETALA2 during floral transition.

MicroRNAs (miRNAs) play important roles in regulating flowering and reproduction of angiosperms. Mature miRNAs are encoded by multiple MIRNA genes that can differ in their spatiotemporal activities and their contributions to gene regulatory networks, but the functions of individual MIRNA genes are poorly defined. We functionally analyzed the activity of all 5 Arabidopsis thaliana MIR172 genes, which encode miR172 and promote the floral transition by inhibiting the accumulation of APETALA2 (AP2) and APETALA2-LIKE (AP2-LIKE) transcription factors (TFs). Through genome editing and detailed confocal microscopy, we show that the activity of miR172 at the shoot apex is encoded by 3 MIR172 genes, is critical for floral transition of the shoot meristem under noninductive photoperiods, and reduces accumulation of AP2 and TARGET OF EAT2 (TOE2), an AP2-LIKE TF, at the shoot meristem. Utilizing the genetic resources generated here, we show that the promotion of flowering by miR172 is enhanced by the MADS-domain TF FRUITFULL, which may facilitate long-term silencing of AP2-LIKE transcription, and that their activities are partially coordinated by the TF SQUAMOSA PROMOTER-BINDING-LIKE PROTEIN 15. Thus, we present a genetic framework for the depletion of AP2 and AP2-LIKE TFs at the shoot apex during floral transition and demonstrate that this plays a central role in floral induction.

Cuscuta australis (dodder) parasite eavesdrops on the host plants’ FT signals to flower

Many plants use environmental cues, including seasonal changes of day length (photoperiod), to control their flowering time. Under inductive conditions, FLOWERING LOCUS T (FT) protein is synthesized in leaves, and FT protein is a mobile signal, which is able to travel to the shoot apex to induce flowering. Dodders (Cuscuta, Convolvulaceae) are root- and leafless plants that parasitize a large number of autotrophic plant species with varying flowering time. Remarkably, some dodder species, e.g., Cuscuta australis, are able to synchronize their flowering with the flowering of their hosts. Detailed sequence inspection and expression analysis indicated that the FT gene in dodder C. australis very likely does not function in activating flowering. Using soybean host plants cultivated under inductive and noninductive photoperiod conditions and soybean and tobacco host plants, in which FT was overexpressed and knocked out, respectively, we show that FT-induced flowering of the host is likely required for both host and parasite flowering. Biochemical analysis revealed that host-synthesized FT signals are able to move into dodder stems, where they physically interact with a dodder FD transcription factor to activate dodder flowering. This study demonstrates that FTs can function as an important interplant flowering signal in host-dodder interactions. The unique means of flowering regulation of dodder illustrates how regressive evolution, commonly found in parasites, may facilitate the physiological synchronization of parasite and host, here allowing the C. australis parasite to time reproduction exactly with that of their hosts, likely optimizing parasite fitness.

Effect of phyB and phyC loss-of-function mutations on the wheat transcriptome under short and long day photoperiods

BackgroundPhotoperiod signals provide important cues by which plants regulate their growth and development in response to predictable seasonal changes. Phytochromes, a family of red and far-red light receptors, play critical roles in regulating flowering time in response to changing photoperiods. A previous study showed that loss-of-function mutations in either PHYB or PHYC result in large delays in heading time and in the differential regulation of a large number of genes in wheat plants grown in an inductive long day (LD) photoperiod.ResultsWe found that under non-inductive short-day (SD) photoperiods, phyB-null and phyC-null mutants were taller, had a reduced number of tillers, longer and wider leaves, and headed later than wild-type (WT) plants. The delay in heading between WT and phy mutants was greater in LD than in SD, confirming the importance of PHYB and PHYC in accelerating heading date in LDs. Both mutants flowered earlier in SD than LD, the inverse response to that of WT plants. In both SD and LD photoperiods, PHYB regulated more genes than PHYC. We identified subsets of differentially expressed and alternatively spliced genes that were specifically regulated by PHYB and PHYC in either SD or LD photoperiods, and a smaller set of genes that were regulated in both photoperiods. We found that photoperiod had a contrasting effect on transcript levels of the flowering promoting genes VRN-A1 and PPD-B1 in phyB and phyC mutants compared to the WT.ConclusionsOur study confirms the major role of both PHYB and PHYC in flowering promotion in LD conditions. Transcriptome characterization revealed an unexpected reversion of the wheat LD plants into SD plants in the phyB-null and phyC-null mutants and identified flowering genes showing significant interactions between phytochromes and photoperiod that may be involved in this phenomenon. Our RNA-seq data provides insight into light signaling pathways in inductive and non-inductive photoperiods and a set of candidate genes to dissect the underlying developmental regulatory networks in wheat.

Constitutive expression of CsGI alters critical night length for flowering by changing the photo-sensitive phase of anti-florigen induction in chrysanthemum

Chrysanthemum is a typical short day (SD) flowering plant that requires a longer night period than a critical minimum duration to successfully flower. We identified FLOWERING LOCUS T-LIKE 3 (FTL3) and ANTI-FLORIGENIC FT/TFL1 FAMILY PROTEIN (AFT) as a florigen and antiflorigen, respectively, in a wild diploid chrysanthemum (Chrysanthemum seticuspe). Expression of the genes that produce these proteins, CsFTL3 and CsAFT, is induced in the leaves under SD or a noninductive photoperiod, respectively, and the balance between them determines the progression of floral transition and anthesis. However, how CsFTL3 and CsAFT are regulated to define the critical night length for flowering in chrysanthemum is unclear. In this study, we focused on the circadian clock-related gene GIGANTEA (GI) of C. seticuspe (CsGI) and generated transgenic C. seticuspe plants overexpressing CsGI (CsGI-OX). Under a strongly inductive SD (8 L/16D) photoperiod, floral transition occurred at almost the same time in both wild-type and CsGI-OX plants. However, under a moderately inductive (12 L/12D) photoperiod, the floral transition in CsGI-OX plants was strongly suppressed, suggesting that the critical night length for flowering was lengthened for CsGI-OX plants. Under the 12 L/12D photoperiod, CsAFT was upregulated in CsGI-OX plants. Giving a night break (NB) 10 h after dusk was the most effective time to inhibit flowering in wild-type plants, while the most effective time for NB was extended to dawn (12 and 14 h after dusk) in CsGI-OX plants. In wild-type plants, a red-light pulse delivered 8 or 10 h after dusk induced maximal CsAFT expression, but the length of the time period over which CsAFT could be induced by red light was extended until subjective dawn in CsGI-OX plants. Therefore, CsGI-OX plants required a longer dark period to maintain lower levels of CsAFT, and their critical night length for flowering was thus lengthened. These results suggested that CsGI has an important role in the control of photoperiodic flowering through shaping the gate for CsAFT induction by light in chrysanthemum.

AcFT promotes kiwifruit invitro flowering when overexpressed and Arabidopsis flowering when expressed in the vasculature under its own promoter.

Kiwifruit (Actinidia chinensis) has three FLOWERING LOCUS T (FT) genes, AcFT, AcFT1, and AcFT2, with differential expression and potentially divergent roles. AcFT was previously shown to be expressed in source leaves and induced in dormant buds by winter chilling. Here, we show that AcFT promotes flowering in A. chinensis, despite a short sequence insertion not present in other FT‐like genes. A 3.5‐kb AcFT promoter region contained all the regulatory elements required to mediate vascular expression in transgenic Arabidopsis thaliana (Arabidopsis). The promoter activation was initially confined to the veins in the distal end of the leaf, before extending to the veins in the base of the leaf, and was detected in inductive and noninductive photoperiods. The 3‐kb and 2.7‐kb promoter regions of AcFT1 and AcFT2, respectively, demonstrated different activation patterns in Arabidopsis, corresponding to differential expression in kiwifruit. Expression of AcFT cDNA from the AcFT promoter was capable to induce early flowering in transgenic Arabidopsis in noninductive photoperiods. Further, expression of AcFT cDNA fused to the green fluorescent protein was detected in the vasculature and was also capable to advance flowering in noninductive photoperiods. Taken together, these studies implicate AcFT in regulation of kiwifruit flowering time and as a candidate for kiwifruit florigen.

Growth and Development of Two Perennial Grasses in Ambient Light Conditions during their Natural Dormant Period

Phenological and morphological behaviors under variable temporal and spatial environmental conditions provide essential information for breeding native perennial warm‐season grasses for bioenergy and ecosystem goods and services. Among various environmental factors, natural photoperiodic signals can affect both vegetative and reproductive growth of perennial species. In this study, field‐collected plants of switchgrass (Panicum virgatum L.) and prairie cordgrass (Spartina pectinata Link) were subjected to ambient photoperiods of either increasing or decreasing daylength during the dormant period of their normal growth cycles in a temperature‐controlled greenhouse. Upland switchgrass reacted to a short‐day treatment (decreasing or increasing daylength <12 h) with a less synchronized dormancy‐breaking pattern than prairie cordgrass. Prairie cordgrass showed a strict daylength overriding dormancy‐breaking pattern. However, phenological and morphological traits observed during the growing season through reproductive development and senescence indicated that the capacity to reach sexual maturity for both species was not greatly affected by subjection to noninductive photoperiod immediately after breaking dormancy. Overall, our results indicated that switchgrass coped with unfamiliar natural photoperiod conditions better than prairie cordgrass in terms of vegetative and reproductive growth.

Haloxylon ammodendron (Amaranthaceae) fruit development delay caused by post-flowering non-inductive photoperiod

Haloxylon ammodendron (C. A. Mey.) is one of the economically and ecologically important desert trees used for sand fixation. The ovary of H. ammodendron is found not to swell after flowering in spring until at the end of August or early September in western China. However, what happens for ovary at anatomic level in that period and which crucial ecological factor regulates the phenomenon of H. ammodendron have not been fully understood. To characterize the phenomenon and explore the crucial environmental regulating factors, we carried out the morphological and anatomic observations at the different development stages of the fruits and three single-factor experiments (low air temperature, sufficient soil moisture, and short day length). Our results showed that under the natural conditions, the ovary of H. ammodendron after flowering developed slowly and the morphological changes of fruits were not significant for the period from May to August and after late August or early September; and then the ovary developed rapidly and matured in October. Cell division in embryo was observed to start approximately 25 days after flowering (DAF) and just developed to globular embryo stage at mid-August. Photoperiod was identified as the pivotal environmental factor regulating the fruit development of H. ammodendron. Moreover, the threshold value of day length for the fruit development was 14.0 h. A long day (>14.0 h) treatment began from 5 DAF could delay fruit development of H. ammodendron while a short day (<14.0 h) treatment could accelerate it. Moreover, a further longer day treatment (>15.0 h) could also delay fruit development even when they had developed for a long time (110 DAF). The present study indicated that H. ammodendron adopted a reproductive strategy of delayed fruit development and this strategy helps it survive and obtain offspring in harsh desert habitats.

Photoperiod-insensitive floral transition in chrysanthemum induced by constitutive expression of chimeric repressor CsLHY-SRDX

A wide variety of physiological processes including flowering are controlled by the circadian clock in plants. In Arabidopsis, LATE ELONGATED HYPOCOTYL (LHY) and CIRCADIAN CLOCK ASSOCIATED 1 (CCA1) constitute the central oscillator, and their gain of function and loss of function disrupt the circadian clock and affect flowering time through FLOWERING LOCUS T (FT), a gene encoding a florigen. Chrysanthemum is a typical short-day (SD) plant and responds to shortening of day length by the transition from the vegetative to reproductive phase. We identified FLOWERING LOCUS T-LIKE 3 (FTL3) and ANTI-FLORIGENIC FT/TFL1 FAMILY PROTEIN (AFT) as a florigen and antiflorigen, respectively, in a wild diploid chrysanthemum (Chrysanthemum seticuspe f. boreale). CsFTL3 and CsAFT are induced under SD or a noninductive photoperiod, respectively, and their balance determines the floral transition and anthesis. Meanwhile, the time-keeping mechanism that regulates the photoperiodic flowering in chrysanthemum is poorly understood. Here, we focused on a LHY/CCA1-like gene called CsLHY in chrysanthemum. We fused CsLHY to a gene encoding short transcriptional repressor domain (SRDX) and constitutively expressed it in chrysanthemum. Although the transcription of clock-related genes was conditionally affected, circadian rhythm was not completely disrupted in CsLHY-SRDX transgenic plants. These plants formed almost the same number of leaves before floral transition under SD and long-day conditions. Thus, CsLHY-SRDX chrysanthemum showed photoperiod-insensitive floral transition, but further development of the capitulum was arrested, and anthesis was not observed. Simultaneously with the flowering phenotype, CsFTL3 and CsAFT were downregulated in CsLHY-SRDX transgenic plants. These results suggest that CsLHY-SRDX affects CsFTL3 and CsAFT expression and causes photoperiod-insensitive floral transition without a severe defect in the circadian clock.

Within variety flowering time variation leads to yield variation in Sri Lankan traditional rice “Sudu wee”

Sri Lankan traditional rice is an important breeding resource for development of rice for ecological adaptation during the era of climate change. There are around 2000 accessions belonging to more than 500 varieties comprised of one to several accessions in each with wider morphological differences, flowering times and yield variations. Genetic identity of Sri Lankan traditional rice accessions belonging to each rice variety is not established at molecular level. Variety Sudu wee comprises of around 30 accessions at PGRC. Sudu wee could be useful in re-introduction programmes of traditional rice due to relatively shorter crop duration. The objective of the present study was to evaluate the diversity of 29 Sudu wee accessions from PGRC, based on days to flowering, (DF), 12 morphological and 2 physiological characters during non-inductive photoperiod season. DF varied from 62 to 106 days while accession 4193 did not flower during the experimental period of nearly 7 months. Leaf temperature at heading (LeT) and Leaf temperature difference at heading (LeTD), vegetative morphological characters and yield components varied among Sudu wee accessions. Seventy nine percent of total observed variation among accessions was explained through Principal Component Analysis. In the dendogram of accessions, 10 clusters were formed at the rescale distance of 5. Seed morphology indicated the genetic similarity among accessions while significantly different DF could be an indication of flowering time gene variation among similar genotypes. Increased DF reduced the grain yield increasing the vegetative growth among accessions of Sudu wee . Our results would be useful in future breeding for manipulating flowering time and yield within similar genetic backgrounds.

Morphological Variation of Vegetative Phase among Ma Wee (Oryza sativa L.) Accessions during Non-Inductive Growing Season for Flowering

Ma wee (Oryza sativa L.) had been a popular traditional rice in the past due to desirable agronomic traits for adverse environments. Ma wee accessions (Accns) are known to be photoperiod sensitive for flowering initiation which flower during Maha (short day) season. Understanding morphological variation of Ma wee is useful in future analysis for genetic diversity. Thirty nine Accns of Ma wee varieties used for the present study with the objective of determining variation of morphological responses when Accns remained in vegetative phase under non-inductive photoperiod for flower initiation at 210 days after sowing. Thirty nine Accns from plant genetic resources center, Sri Lanka were grown in Kamburupitiya with four replicates in a completely randomised design at the end of Maha season in 2013/2014. Quantitative characters of plant height (PH), total tiller number (TTN), total leaf number (TLN), culm number (CN) and ligules length (LL) and qualitative characters of ligule color (LC), ligule shape (LS), internode color (IC), culm strength (CS) were recorded. None of the 39 Accns flowered by 210 days and PH, TTN, TLN, CN, and LL varied from 87.5±1.61 cm, 2±0.37, 8±0.84, 2±0.43, 1.8±0.46 cm to 199±1.36 cm, 22±2.09, 136±2.20, 24±2.03 and 4.18±0.25 cm respectively. The lowest PH, TTN, TLN, and CN were recorded from Accn 6253. Except for Accn 6253, rests 38 Accns were over 1 m in PH. Qualitative characters varied among clusters without any uniformity for a given cluster. CS varied among Accns from weak, intermediate to strong. LC and IC varied from green to white and green to yellow respectively. Acute to acuminate and two cleft LS were also observed. Principal Component Analysis followed by Cluster Analysis was performed using data from quantitative characters. First two of PCs explained over 80% of total morphological variations. The first PC represented the PH, TTN, TLN and CN whereas PC2 represented the LL. Eight clusters were obtained at rescaled distance five in the dendogram. Variation among Accns within clusters for qualitative characters may be an indication of genetic diversity. Keywords: Ma wee, Morphological variation, Vegetative phase

Florigen-Encoding Genes of Day-Neutral and Photoperiod-Sensitive Maize Are Regulated by Different Chromatin Modifications at the Floral Transition.

The activity of the maize (Zea mays) florigen gene ZEA CENTRORADIALIS8 (ZCN8) is associated with the floral transition in both day-neutral temperate maize and short-day (SD)-requiring tropical maize. We analyzed transcription and chromatin modifications at the ZCN8 locus and its nearly identical paralog ZCN7 during the floral transition. This analysis was performed with day-neutral maize (Zea mays ssp. mays), where flowering is promoted almost exclusively via the autonomous pathway through the activity of the regulatory gene indeterminate1 (id1), and tropical teosinte (Zea mays ssp. parviglumis) under floral inductive and noninductive photoperiods. Comparison of ZCN7/ZCN8 histone modification profiles in immature leaves of nonflowering id1 mutants and teosinte grown under floral inhibitory photoperiods reveals that both id1 floral inductive activity and SD-mediated induction result in histone modification patterns that are compatible with the formation of transcriptionally competent chromatin environments. Specific histone modifications are maintained during leaf development and may represent a chromatin signature that favors the production of processed ZCN7/ZCN8 messenger RNA in florigen-producing mature leaf. However, whereas id1 function promotes histone H3 hyperacetylation, SD induction is associated with increased histone H3 dimethylation and trimethylation at lysine-4. In addition, id1 and SD differently affect the production of ZCN7/ZCN8 antisense transcript. These observations suggest that distinct mechanisms distinguish florigen regulation in response to autonomous and photoperiod pathways. Finally, the identical expression and histone modification profiles of ZCN7 and ZCN8 in response to floral induction suggest that ZCN7 may represent a second maize florigen.

CsTFL1, a constitutive local repressor of flowering, modulates floral initiation by antagonising florigen complex activity in chrysanthemum

Chrysanthemums require repeated cycles of short-day (SD) photoperiod for successful anthesis, but their vegetative state is strictly maintained under long-day (LD) or night-break (NB) conditions. We have previously demonstrated that photoperiodic flowering of a wild diploid chrysanthemum (Chrysanthemum seticuspe f. boreale) is controlled by a pair of systemic floral regulators, florigen (CsFTL3) and anti-florigen (CsAFT), produced in the leaves. Here, we report the functional characterisation of a local floral regulator, CsTFL1, a chrysanthemum orthologue of TERMINAL FLOWER 1 gene in Arabidopsis. Constitutive expression of CsTFL1 in C. seticuspe (CsTFL1-ox) resulted in extremely late flowering under SD and prevented up-regulation of floral meristem identity genes in shoot tips and leaves. Bimolecular fluorescence complementation assay showed that both CsTFL1 and CsFTL3 interacted with CsFDL1, a bZIP transcription factor FD homologue, in the nucleus. The transient gene expression assay indicated that CsTFL1 suppresses flowering by directly antagonising the flower inductive activity of the CsFTL3-CsFDL1 complex. Our results suggest that strict maintenance of vegetative state under non-inductive photoperiod is achieved by the coordinated action of both the systemic floral inhibitor and local floral inhibitor CsTFL1, which is constitutively expressed in shoot tips.

Nitric oxide is involved in salicylic acid-induced flowering of Lemna aequinoctialis Welw.

Salicylic acid (SA) is a well-known inducer of flowering in Lemna under both non-inductive and inductive photoperiod conditions. However, the underlying mechanism is not well understood. Herein, we report for the first time that nitric oxide (NO) is partially involved in SA-induced flowering in L. aequinoctialis (Syn. L. paucicostata Hegelm.). Our results demonstrated that SA-induced flowering is significantly reduced by exogenous application of NO scavengers; 2-(4-carboxyphenyl)-4,4,5,5-tetramethylimidazoline-1-oxyl-3-oxide and methylene blue, nitric oxide synthase inhibitors; N-ω-nitro-l-arginine and N-ω-nitro-l-arginine-methyl ester hydrochloride, and nitrate reductase inhibitor; sodium tungstate in two strains of Lemna viz. 6746 and LP6. Altogether our present findings shed a light on the new role of NO in SA-induced flowering and open interesting directions that need further investigation.

Functional characterization of duplicated Suppressor of Overexpression of Constans 1-like genes in petunia.

Flowering time is strictly controlled by a combination of internal and external signals that match seed set with favorable environmental conditions. In the model plant species Arabidopsis thaliana (Brassicaceae), many of the genes underlying development and evolution of flowering have been discovered. However, much remains unknown about how conserved the flowering gene networks are in plants with different growth habits, gene duplication histories, and distributions. Here we functionally characterize three homologs of the flowering gene SUPPRESSOR OF OVEREXPRESSION OF CONSTANS 1 (SOC1) in the short-lived perennial Petunia hybrida (petunia, Solanaceae). Similar to A. thaliana soc1 mutants, co-silencing of duplicated petunia SOC1-like genes results in late flowering. This phenotype is most severe when all three SOC1-like genes are silenced. Furthermore, expression levels of the SOC1-like genes UNSHAVEN (UNS) and FLORAL BINDING PROTEIN 21 (FBP21), but not FBP28, are positively correlated with developmental age. In contrast to A. thaliana, petunia SOC1-like gene expression did not increase with longer photoperiods, and FBP28 transcripts were actually more abundant under short days. Despite evidence of functional redundancy, differential spatio-temporal expression data suggest that SOC1-like genes might fine-tune petunia flowering in response to photoperiod and developmental stage. This likely resulted from modification of SOC1-like gene regulatory elements following recent duplication, and is a possible mechanism to ensure flowering under both inductive and non-inductive photoperiods.

EARLY FLOWERING3 Regulates Flowering in Spring Barley by Mediating Gibberellin Production and FLOWERING LOCUS T Expression

EARLY FLOWERING3 (ELF3) is a circadian clock gene that contributes to photoperiod-dependent flowering in plants, with loss-of-function mutants in barley (Hordeum vulgare), legumes, and Arabidopsis thaliana flowering early under noninductive short-day (SD) photoperiods. The barley elf3 mutant displays increased expression of FLOWERING LOCUS T1 (FT1); however, it remains unclear whether this is the only factor responsible for the early flowering phenotype. We show that the early flowering and vegetative growth phenotypes of the barley elf3 mutant are strongly dependent on gibberellin (GA) biosynthesis. Expression of the central GA biosynthesis gene, GA20oxidase2, and production of the bioactive GA, GA1, were significantly increased in elf3 leaves under SDs, relative to the wild type. Inhibition of GA biosynthesis suppressed the early flowering of elf3 under SDs independently of FT1 and was associated with altered expression of floral identity genes at the developing apex. GA is also required for normal flowering of spring barley under inductive photoperiods, with chemical and genetic attenuation of the GA biosynthesis and signaling pathways suppressing inflorescence development under long-day conditions. These findings illustrate that GA is an important floral promoting signal in barley and that ELF3 suppresses flowering under noninductive photoperiods by blocking GA production and FT1 expression.

Identification of high long-day leaf number cultivars and prevention of premature budding by cold pre-treatment for fine control of flowering in summer-to-autumn-flowering chrysanthemum (Chrysanthemum morifolium Ramat.)

SummaryWhen maintained continuously under long-day (non-inductive) photoperiodic conditions, chrysanthemum (Chrysanthemum morifolium Ramat.), a short-day plant, eventually initiates flower buds. This floral initiation is termed ‘premature budding’ in commercial chrysanthemum production. Premature budding, which reduces cut-flower quality, is often observed in summer-to-autumn-flowering cultivars. The timing of flower initiation under non-inductive photoperiods depends on the number of leaves that a plant has formed below the terminal flower bud. This leaf number indicates whether a cultivar will form flower buds prematurely during a non-inductive photoperiod. Two summer-flowering, ten summer-to-autumn-flowering, and six autumn-flowering chrysanthemum cultivars were grown under controlled conditions to measure the number of leaves produced under a non-inductive photoperiod with a night-break. The average number of leaves per plant ranged from 13.8 for ‘Tasogare’, to 69.1 for ‘Furamu’. These data can be used to identify cultivars that will exhibit minimal flower-bud formation under night-break conditions. The numbers of leaves were greater in plants subjected to a 12-week cold treatment prior to planting, suggesting that premature budding could be inhibited by a prolonged cold pre-treatment. We also examined whether the cold treatment of stock plants was an effective means to prevent premature budding. The incidence of premature budding was reduced in rooted cuttings obtained from cold-treated stock plants of the summer-to-autumn-flowering cultivars ‘Nagano Queen’, ‘Ariesu’, ‘Sei-un’, ‘Oruka’, and ‘Floral Yuka’. Without a cold treatment, cuttings from these cultivars showed premature budding. This result suggests that a prolonged cold pre-treatment of stock plants could prevent both premature budding and reduced cut-flower quality in summer-to-autumn-flowering chrysanthemum cultivars.