The Arabidopsis TALE homeobox gene ATH1 controls floral competency through positive regulation of FLC

Floral induction is controlled by a plethora of genes acting in different pathways that either repress or promote floral transition at the shoot apical meristem (SAM). During vegetative development high levels of floral repressors maintain the Arabidopsis SAM as incompetent to respond to promoting factors. Among these repressors, FLOWERING LOCUS C (FLC) is the most prominent. The processes underlying downregulation of FLC in response to environmental and developmental signals have been elucidated in considerable detail. However, the basal induction of FLC and its upregulation by FRIGIDA (FRI) are still poorly understood. Here we report the functional characterization of the ARABIDOPSIS THALIANA HOMEOBOX 1 (ATH1) gene. A function of ATH1 in floral repression is suggested by a gradual downregulation of ATH1 in the SAM prior to floral transition. Further evidence for such a function of ATH1 is provided by the vernalization-sensitive late flowering of plants that constitutively express ATH1. Analysis of lines that differ in FRI and/or FLC allele strength show that this late flowering is caused by upregulation of FLC as a result of synergism between ATH1 overexpression and FRI. Lack of ATH1, however, results in attenuated FLC levels independently of FRI, suggesting that ATH1 acts as a general activator of FLC expression. This is further corroborated by a reduction of FLC-mediated late flowering in fca-1 and fve-1 autonomous pathway backgrounds when combined with ath1. Since other floral repressors of the FLC clade are not significantly affected by ATH1, we conclude that ATH1 controls floral competency as a specific activator of FLC expression.     

Differential regulation of trichome formation on the adaxial and abaxial leaf surfaces by gibberellins and photoperiod in Arabidopsis thaliana (L.) Heynh.

In wild-type (WT) Columbia and Landsberg erecta ecotypes of Arabidopsis thaliana (L.) Heynh., trichomes are present on the adaxial surfaces of all rosette leaves but are absent from the abaxial surfaces of the first-formed leaves. We have determined that both long-day (LD) photoperiod and gibberellin (GA) stimulate trichome formation. WT plants grown in LD conditions produce the first abaxial trichome on earlier leaves than plants grown in short-day (SD) conditions. Photoperiod sensitivity of abaxial trichome formation on WT plants develops gradually over time, reaching the maximum sensitivity about 24 d after germination. Application of gibberellic acid to WT plants growing in SD conditions accelerates the onset of abaxial trichomes. Conversely, application of 20 to 80 mg L-1 paclobutrazol, a GA biosynthesis inhibitor, to wild-type plants suppresses trichome initiation on the abaxial epidermis. The GA-deficient mutants ga1-5 and ga4-1 and the GA-insensitive mutant gai-1 exhibit delayed onset of abaxial trichomes when grown in LD conditions. The null mutant ga1-3 produces completely glabrous leaves when grown in SD conditions. Application of gibberellic acid to glabrous ga1-3 plants consistently induces earlier formation of trichomes on the adaxial epidermis than on the abaxial epidermis, demonstrating a difference between the adaxial and abaxial surfaces in their response to GA with regard to trichome formation.     

Reversion of flowering in Glycine Max (Fabaceae)

Photoperiodic changes, if occurring before a commitment to flowering is established, can alter the morphological pattern of plant development. In this study, Glycine max (L.) Merrill cv. Ransom plants were initially grown under an inductive short-day (SD) photoperiod to promote flower evocation and then transferred to a long-day (LD) photoperiod to delay flower development by reestablishing vegetative growth (SD-LD plants). Some plants were transferred back to SD after 4-LD exposures to repromote flowering (SD-LD-SD plants). Alterations in organ initiation patterns, from floral to vegetative and back to floral, are characteristic of a reversion phenomenon. Morphological features that occurred at the shoot apical meristem in SD, LD, SD-LD, and SD-LD-SD plants were observed using scanning electron microscopy (SEM). Reverted plants initiated floral bracts and resumed initiation of trifoliolate leaves in the two-fifths floral phyllotaxy prior to terminal inflorescence development. When these plants matured, leaf-bract intermediates were positioned on the main stem instead of trifoliolate leaves. Plants transferred back to a SD photoperiod flowered earlier than those left in LD conditions. Results indicated that in plants transferred between SDs and LDs, photoperiod can influence organ initiation in florally evoked, but not committed, G. max plants.     

Analysis of the molecular basis of flowering time variation in Arabidopsis accessions

Allelic variation at the FRI (FRIGIDA) and FLC (FLOWERING LOCUS C) loci are major determinants of flowering time in Arabidopsis accessions. Dominant alleles of FRI confer a vernalization requirement causing plants to overwinter vegetatively. Many early flowering accessions carry loss-of-function fri alleles containing one of two deletions. However, some accessions categorized as early flowering types do not carry these deletion alleles. We have analyzed the molecular basis of earliness in five of these accessions: Cvi, Shakhdara, Wil-2, Kondara, and Kz-9. The Cvi FRI allele carries a number of nucleotide differences, one of which causes an in-frame stop codon in the first exon. The other four accessions contain nucleotide differences that only result in amino acid substitutions. Preliminary genetic analysis was consistent with Cvi carrying a nonfunctional FRI allele; Wil-2 carrying either a defective FRI or a dominant suppressor of FRI function; and Shakhdara, Kondara, and Kz-9 carrying a functional FRI allele with earliness being caused by allelic variation at other loci including FLC. Allelic variation at FLC was also investigated in a range of accessions. A novel nonautonomous Mutator-like transposon was found in the weak FLC allele in Landsberg erecta, positioned in the first intron, a region required for normal FLC regulation. This transposon was not present in FLC alleles of most other accessions including Shakhdara, Kondara, or Kz-9. Thus, variation in Arabidopsis flowering time has arisen through the generation of nonfunctional or weak FRI and FLC alleles.     

Carbon Dioxide and Flowering in Pharbitis nil Choisy

The effects of photoperiod on floral and vegetative development of Pharbitis nil were modified by atmospheric CO₂ concentrations maintained during plant growth. Short day (SD) photoperiods caused rapid flowering in Pharbitis plants growing in 0.03 or 0.1% CO₂, while plants in long day (LD) conditions remained vegetative. At 1 or 5% CO₂, however, flower buds were developed under both the SD and LD photoperiods. Flowering was earliest in the plants exposed to SD at low CO₂ concentrations which formed floral buds at stem node 3 or 4. At high CO₂ concentrations, floral buds did not form until stem node 6 or 7. Both high CO₂ concentrations and LD photoperiods tended to enhance stem elongation and leaf formation.     

EARLY IN SHORT DAYS 1 (ESD1) encodes ACTIN-RELATED PROTEIN 6 (AtARP6), a putative component of chromatin remodelling complexes that positively regulates FLC accumulation in Arabidopsis

We have characterized Arabidopsis esd1 mutations, which cause early flowering independently of photoperiod, moderate increase of hypocotyl length, shortened inflorescence internodes, and altered leaf and flower development. Phenotypic analyses of double mutants with mutations at different loci of the flowering inductive pathways suggest that esd1 abolishes the FLC-mediated late flowering phenotype of plants carrying active alleles of FRI and of mutants of the autonomous pathway. We found that ESD1 is required for the expression of the FLC repressor to levels that inhibit flowering. However, the effect of esd1 in a flc-3 null genetic background and the downregulation of other members of the FLC-like/MAF gene family in esd1 mutants suggest that flowering inhibition mediated by ESD1 occurs through both FLC-and FLC-like gene-dependent pathways. The ESD1 locus was identified through a map-based cloning approach. ESD1 encodes ARP6, a homolog of the actin-related protein family that shares moderate sequence homology with conventional actins. Using chromatin immunoprecipitation (ChIP) experiments, we have determined that ARP6 is required for both histone acetylation and methylation of the FLC chromatin in Arabidopsis.     

Using flowering times and leaf numbers to model the phases of photoperiod sensitivity in Antirrhinum majus L

A model has been developed that can be used to determine the phases of sensitivity to photoperiod for seedlings subjected to reciprocal transfers at regular intervals between long (LD) and short day (SD) conditions. The novel feature of this approach is that it enables the simultaneous analysis of the time to flower and number of leaves below the inflorescence. A range of antirrhinum cultivars were grown, all of which were shown to be quantitative long-day plants. Seedlings were effectively insensitive to photoperiod when very young (juvenile). However, after the end of the juvenile phase, SD delayed flowering and increased the number of leaves below the inflorescence. Plants transferred from LD to SD showed a sudden hastening of flowering and a decrease in leaf number once sufficient LD had been received for flower commitment. Photoperiod had little effect on the rate of flower development. The analysis clearly identified major cultivar differences in the length of the juvenile phase and the photoperiod-sensitive inductive phase in both LD and SD.     

Rates of Growth and Cell Division in the Shoot Apex of Silene During the Transition to Flowering

The growth rates of the shoot apex during and after floral inductionwere measured in Silene, a long-day plant. Plants were inducedto flower with 4 or more long days (LD) but not with 3 longdays or with short days (SD). The rate of increase of cell numberin the apical dome, above the youngest leaf pair, was exponentialand in plants given 3 LD remained the same as in plants in SD.In plants induced to flower with 7 LD, until the end of theinductive period the rate of increase of cell number in theapical dome remained the same as in plants in SD. Only whenthe apex began to enlarge as the first stage in the formationof the flower did the growth rate of the apical dome increase.The rates of increase of cell numbers in the apex correspondedto mean cell generation times of 20 to 33 h for plants in SD,for plants given 3 LD, and during the 7 days of induction forplants given 7 LD, and 6 to 8 h for induced plants when flowerformation was beginning. The distribution of cell division in the apex was examined bytreating plants with colchicine and noting in sections the positionsof the resulting metaphases. In vegetative apices and also inapices undergoing transition to flowering the whole of the apicaldome appeared to consist of cells dividing at a similar rate. The rate of leaf initiation during induction was the same asin vegetative, non-induced plants.     

拟南芥春化作用相关基因FLC的表达调控及天然早花突变体的研究进展

植物开花是从营养生长到生殖状态的重要发育转变,是多种内在因子和环境因素共同作用的结果。在拟南芥开花调控网络中,开花抑制基因FLC处于枢纽地位。FLC的表达受许多来自环境和生长发育的信号调控,主要包括：PAF1复合体、SWR1复合体成员,FRI依赖途径、自主途径和春化作用途径基因。本文主要综述了影响FLC表达的春化相关基因及天然早花突变体的研究进展,并根据最新的研究成果提出该研究领域的研究方向和重点。     

The Control of Apical Bud Growth and Senescence by Auxin and Gibberellin in Genetic Lines of Peas

Pea (Pisum sativum L.) lines G2 (dwarf) and NGB1769 (tall) (Sn Hr) produce flowers and fruit under long (LD) or short (SD) days, but senesce only under LD. Endogenous gibberellin (GA) levels were inversely correlated with photoperiod (over 9-18 h) and senescence: GA20 was 3-fold and GA1 was 10- to 11-fold higher in flowering SD G2 shoots, and the vegetative tissues within the SD apical bud contained 4-fold higher levels of GA20, as compared with the LD tissues. Prefloral G2 plants under both photoperiods had GA1 and GA20 levels similar to the flowering plants under LD. Levels of indole-3-acetic acid (IAA) were similar in G2 shoots in LD or SD; SD apical bud vegetative tissues had a slightly higher IAA content. Young floral buds from LD plants had twice as much IAA as under SD. In NGB1769 shoots GA1 decreased after flower initiation only under LD, which correlated with the decreased growth potential. We suggest that the higher GA1 content of G2 and NGB1769 plants under SD conditions is responsible for the extended vegetative growth and continued meristematic activity in the shoot apex. This and the increased IAA level of LD floral buds may play a role in the regulation of nutrient partitioning, since more photosynthate partitions of reproductive tissue under LD conditions, and the rate of reproductive development in LD peas is faster than under SD.     

The role of miR156 in developmental transitions in Nicotiana tabacum

Plants undergo a series of developmental transitions during their life cycle. After seed germination, plants pass through two distinct phases: the vegetative phase in which leaves are produced and the reproductive phase in which flowering occurs. Based on the reproductive competence and morphological changes, the vegetative phase can be further divided into juvenile and adult phases. Here, we demonstrate that the difference between juvenile and adult phase of Nicotiana tabacum is characterized by the changes in leaf size, leaf shape as well as the number of leaf epidermal hairs(trichomes). We further show that miR156, an age-regulated microR NA, regulates juvenile-to-adult phase transition in N. tabacum. Overexpression of miR156 results in delayed juvenile-to-adult transition and flowering. Together, our results support an evolutionarily conserved role of miR156 in plant developmental transitions.     

Morphological and anatomical evaluation of adult and juvenile leaves of olive plants

The olive tree (Olea europaea L.), like many other woody plants, has a long juvenile period in which the plant is not able to produce flowers. Knowledge of the moment when the plant is capable of flowering is important for breeding programs and also for determining the physiological basis for sexual reproductive behavior, but currently the only indicator of that moment is the actual flowering. In many species, the juvenile-to-adult phase shift includes changes in leaf structure known as heteroblasty, that is, varied form of successive leaves on the same plant. Some differences have been observed between juvenile and adult olive leaves, particularly in size and form, but to our knowledge, no complete systematic study has been carried out. In this research, we measured size, morphology and anatomy for juvenile and adult leaves of olive plants grown from seeds. Differences were found in most of the parameters studied, including leaf size, form, mesophyll thickness, layers of palisade parenchyma and quantity of peltate trichomes, which were generally significant but overlapping between the two leaf types. The most consistent and striking difference was the presence of an organized layer of subepidermal cells only in the abaxial mesophyll of adult leaves. This characteristic could be a simple and effective criterion of phase change in the olive tree.     

FRIGIDA-independent variation in flowering time of natural Arabidopsis thaliana accessions

FRIGIDA (FRI) and FLOWERING LOCUS C (FLC) are two genes that, unless plants are vernalized, greatly delay flowering time in Arabidopsis thaliana. Natural loss-of-function mutations in FRI cause the early flowering growth habits of many A. thaliana accessions. To quantify the variation among wild accessions due to FRI, and to identify additional genetic loci in wild accessions that influence flowering time, we surveyed the flowering times of 145 accessions in long-day photoperiods, with and without a 30-day vernalization treatment, and genotyped them for two common natural lesions in FRI. FRI is disrupted in at least 84 of the accessions, accounting for only approximately 40% of the flowering-time variation in long days. During efforts to dissect the causes for variation that are independent of known dysfunctional FRI alleles, we found new loss-of-function alleles in FLC, as well as late-flowering alleles that do not map to FRI or FLC. An FLC nonsense mutation was found in the early flowering Van-0 accession, which has otherwise functional FRI. In contrast, Lz-0 flowers late because of high levels of FLC expression, even though it has a deletion in FRI. Finally, eXtreme array mapping identified genomic regions linked to the vernalization-independent, late-flowering habit of Bur-0, which has an alternatively spliced FLC allele that behaves as a null allele.     

Photoperiodic studies on growth and development of Impatiens balsamina L.

Summary In the short-day plant Impatiens balsamina it was found that, while floral buds are initiated with 3 short-day (SD) cycles, at least 8 such cycles are required for flowering. The numbers of floral buds and open flowers bear a linear relationship with the number of SD cycles. The induced floral buds revert to vegetative growth unless the plants receive the minimum number of SD cycles needed for flowering, this reversion occurring in a basipetal direction. The rate of extension growth of the stem increases with increasing numbers of SD cycles. The high rate is maintained longer in plants receiving 32 or more SD cycles, but the subsequent fall is also steeper in these plants than in plants receiving less inductive cycles. Senescence also occurs in these plants and appears to be related to the magnitude of reproductive development and the high rate of extension growth.