光周期途径核心因子CO的开花调控机制

CONSTANS（CO）基因是生物钟和开花时间基因之间监测日照长度的重要元件,在光周期途径中发挥核心功能。CO可以整合光信号和生物钟信号,诱导开花途径整合子FLOWERINGLOCUST（F即和SUPPRESSOROF OVEREXPRESSION OF CONSTANS 1（SOC1）的表达,进而促进植株开花。本文综述CO基因的开花调控机制,并结合CO基因的研究现状展望了其未来的研究方向。     

AGL24 acts in concert with SOC1 and FUL during Arabidopsis floral transition

Arabidopsis plants flower in response to long days (LDs). Exposure of leaves to inductive day lengths activates expression of FLOWERING LOCUS T (FT) protein which moves to the shoot apical meristem (SAM) to induce developmental reprogramming. SUPPRESSOR OF OVEREXPRESSION OF CONSTANS 1 (SOC1) and FRUITFULL (FUL) are induced by FT at the apex. We previously screened the SAM for mRNAs of genes required to promote the floral transition in response to photoperiod, and conducted detailed expression and functional analyses on several putative candidates. Here, we show that expression of AGAMOUS-LIKE 24 (AGL24) is detected at the SAM under SD conditions and increases upon exposure to LDs. Mutations in AGL24 further delay flowering of a soc1 ful double mutant, suggesting that flowering is controlled by AGL24 partly independently of SOC1 and FUL.     

Flowering time regulation produces much fruit

Many of the molecular details regarding the promotion of flowering in response to prolonged exposure to cold temperatures (vernalization) and daylength have recently been elucidated in Arabidopsis. The daylength and vernalization pathway converge in the regulation of floral promoters referred to as floral integrators. In the meristem, vernalization promotes flowering through the epigenetic repression of the floral repressor FLOWERING LOCUS C. This allows for the induction of floral integrators by CONSTANS under inductive long days. In the vasculature of leaves, CONSTANS protein is produced only in long days where it acts to promote the expression of FLOWERING LOCUS T (FT). FT protein is then translocated to the meristem where it acts to promote floral induction. Thus a detailed molecular framework for the regulation of flowering time has now been established in Arabidopsis.     

Integration of photoperiod and cold temperature signals into flowering genetic pathways in Arabidopsis

Appropriate timing of flowering is critical for propagation and reproductive success in plants. Therefore, flowering time is coordinately regulated by endogenous developmental programs and external signals, such as changes in photoperiod and temperature. Flowering is delayed by a transient shift to cold temperatures that frequently occurs during early spring in the temperate zones. It is known that the delayed flowering by short-term cold stress is mediated primarily by the floral repressor FLOWERING LOCUS C (FLC). However, how the FLC-mediated cold signals are integrated into flowering genetic pathways is not fully understood. We have recently reported that the INDUCER OF CBF EXPRESSION 1 (ICE1), which is a master regulator of cold responses, FLC, and the floral integrator SUPPRESSOR OF OVEREXPRESSION OF CONSTANS 1 (SOC1) constitute an elaborated feedforward-feedback loop that integrates photoperiod and cold temperature signals to regulate seasonal flowering in Arabidopsis. Cold temperatures promote the binding of ICE1 to FLC promoter to induce its expression, resulting in delayed flowering. However, under floral inductive conditions, SOC1 induces flowering by blocking the ICE1 activity. We propose that the ICE1-FLC-SOC1 signaling network fine-tunes the timing of photoperiodic flowering during changing seasons.     

Potent induction of Arabidopsis thaliana flowering by elevated growth temperature

The transition to flowering is an important event in the plant life cycle and is modulated by several environmental factors including photoperiod, light quality, vernalization, and growth temperature, as well as biotic and abiotic stresses. In contrast to light and vernalization, little is known about the pathways that mediate the responses to other environmental variables. A mild increase in growth temperature, from 23 °C to 27 °C, is equally efficient in inducing flowering of Arabidopsis plants grown in 8-h short days as is transfer to 16-h long days. There is extensive natural variation in this response, and we identify strains with contrasting thermal reaction norms. Exploiting this natural variation, we show that FLOWERING LOCUS C potently suppresses thermal induction, and that the closely related floral repressor FLOWERING LOCUS M is a major-effect quantitative trait locus modulating thermosensitivity. Thermal induction does not require the photoperiod effector CONSTANS, acts upstream of the floral integrator FLOWERING LOCUS T, and depends on the hormone gibberellin. Analysis of mutants defective in salicylic acid biosynthesis suggests that thermal induction is independent of previously identified stress-signaling pathways. Microarray analyses confirm that the genomic responses to floral induction by photoperiod and temperature differ. Furthermore, we report that gene products that participate in RNA splicing are specifically affected by thermal induction. Above a critical threshold, even small changes in temperature can act as cues for the induction of flowering. This response has a genetic basis that is distinct from the known genetic pathways of floral transition, and appears to correlate with changes in RNA processing.     

Expression analysis of phytochromes A, B and floral integrator genes during the entry and exit of grapevine-buds from endodormancy

A common molecular regulatory pathway that involves PHYA, PHYB and floral integrator genes CONSTANS (CO), FLOWERING LOCUS T (FT) and SUPRESSOR OF OVEREXPRESSION OF CO1 (SOC1) has been suggested to participate in the regulation of photoperiod dependent processes such as flowering and dormancy. In grapevines (Vitis vinifera L.), decreasing photoperiod and low temperatures trigger the transition of buds into endodormancy (ED), a process that is accompanied by drastic changes in gene expression of VvPHYA and B in leaves. To analyse the relationship of VvPHYA, VvPHYB, and grape homologues of Arabidopsis floral integrator genes VvCO, VvFT, VvMADS8, with ED, a comparative expression analysis of these genes was performed in grapevine-leaves and buds before, during and after the transition of buds into ED. The expression of all the above genes in the bud-tissue, and the fact that photoperiod regulates differently the expression of VvPHYA and B in buds than in leaves, suggests that the bud might be an autonomous or semi-autonomous organ in perceiving and transducing the photoperiod signal. On the other hand, the coordinated down-regulation of VvFT in leaves and buds during the transition of buds into ED, and its subsequent up-regulation following the application of dormancy-breaking compounds, hydrogen cyanamide (HC) and sodium azide, suggests that VvFT could play a key role in stimulating bud-growth by repressing their entry into ED.     

The quest for florigen: a review of recent progress

The photoperiodic induction of flowering is a systemic process requiring translocation of a floral stimulus from the leaves to the shoot apical meristem. In response to this stimulus, the apical meristem stops producing leaves to initiate floral development; this switch in morphogenesis involves a change in the identity of the primordia initiated and in phyllotaxis. The physiological study of the floral transition has led to the identification of several putative floral signals such as sucrose, cytokinins, gibberellins, and reduced N-compounds that are translocated in the phloem sap from leaves to the shoot apical meristem. On the other hand, the genetic approach developed more recently in Arabidopsis thaliana allowed the discovery of many genes that control flowering time. These genes function in 'cascades' within four promotive pathways, the 'photoperiodic', 'autonomous', 'vernalization', and 'gibberellin' pathways, which all converge on the 'integrator' genes SUPPRESSOR OF OVEREXPRESSION OF CO 1 (SOC1) and FLOWERING LOCUS T (FT). Recently, several studies have highlighted a role for a product of FT as a component of the floral stimulus or 'florigen'. These recent advances and the proposed mode of action of FT are discussed here.     

The E3 Ubiquitin Ligase HOS1 Regulates Arabidopsis Flowering by Mediating CONSTANS Degradation Under Cold Stress

The timing of flowering is coordinated by a web of gene regulatory networks that integrates developmental and environmental cues in plants. Light and temperature are two major environmental determinants that regulate flowering time. Although prolonged treatment with low nonfreezing temperatures accelerates flowering by stable repression of FLOWERING LOCUS C (FLC), repeated brief cold treatments delay flowering. Here, we report that intermittent cold treatments trigger the degradation of CONSTANS (CO), a central activator of photoperiodic flowering; daily treatments caused suppression of the floral integrator FLOWERING LOCUS T (FT) and delayed flowering. Cold-induced CO degradation is mediated via a ubiquitin/proteasome pathway that involves the E3 ubiquitin ligase HIGH EXPRESSION OF OSMOTICALLY RESPONSIVE GENE 1 (HOS1). HOS1-mediated CO degradation occurs independently of the well established cold response pathways. It is also independent of the light signaling repressor CONSTITUTIVE PHOTOMORPHOGENIC 1 (COP1) E3 ligase and light wavelengths. CO has been shown to play a key role in photoperiodic flowering. Here, we demonstrated that CO served as a molecular hub, integrating photoperiodic and cold stress signals into the flowering genetic pathways. We propose that the HOS1-CO module contributes to the fine-tuning of photoperiodic flowering under short term temperature fluctuations, which often occur during local weather disturbances.     

Integration of flowering signals in winter-annual Arabidopsis

Photoperiod is the primary environmental factor affecting flowering time in rapid-cycling accessions of Arabidopsis (Arabidopsis thaliana). Winter-annual Arabidopsis, in contrast, have both a photoperiod and a vernalization requirement for rapid flowering. In winter annuals, high levels of the floral inhibitor FLC (FLOWERING LOCUS C) suppress flowering prior to vernalization. FLC acts to delay flowering, in part, by suppressing expression of the floral promoter SOC1 (SUPPRESSOR OF OVEREXPRESSION OF CONSTANS1). Vernalization leads to a permanent epigenetic suppression of FLC. To investigate how winter-annual accessions integrate signals from the photoperiod and vernalization pathways, we have examined activation-tagged alleles of FT and the FT homolog, TSF (TWIN SISTER OF FT), in a winter-annual background. Activation of FT or TSF strongly suppresses the FLC-mediated late-flowering phenotype of winter annuals; however, FT and TSF overexpression does not affect FLC mRNA levels. Rather, FT and TSF bypass the block to flowering created by FLC by activating SOC1 expression. We have also found that FLC acts as a dosage-dependent inhibitor of FT expression. Thus, the integration of flowering signals from the photoperiod and vernalization pathways occurs, at least in part, through the regulation of FT, TSF, and SOC1.     

Overexpression of COL9, a CONSTANS-LIKE gene, delays flowering by reducing expression of CO and FT in Arabidopsis thaliana

CONSTANS (CO) is an important floral regulator in the photoperiod pathway, integrating the circadian clock and light signal into a control for flowering time. It is known that CO promotes flowering in Arabidopsis under long-day conditions. CONSTANS-LIKE 9 (COL9) is a member of the CONSTANS-LIKE gene family, encoding a nuclear protein. The expression of COL9 is regulated by the circadian clock in the photoperiod pathway and is detected in various organs. Unexpectedly, overexpression of COL9 in transgenic Arabidopsis resulted in delayed flowering, while co-suppression lines and a transferred DNA (T-DNA) knockout line showed earlier flowering under long-day conditions. Overexpression of COL9 did not enhance the late-flowering phenotype in a co mutant background. Double overexpressors produced by overexpression of CO in COL9 transgenic lines showed an early flowering phenotype similar to single CO overexpressors. The pattern of oscillation of a number of circadian-associated genes remained unchanged in the COL9 transgenic lines. Compared with wild-type plants, the abundance of CO and FLOWERING LOCUS T (FT) mRNA was reduced in the COL9 overexpression lines. Our results indicate that COL9 is involved in regulation of flowering time by repressing the expression of CO, concomitantly reducing the expression of FT and delaying floral transition.     

FLC调控植物成花的分子机制研究新进展

FLC是植物成花关键抑制因子, 主要通过结合到其下游2个关键的成花促进基因(FT和SOC1)启动子上而抑制二者的表达。此外, 还可以与其它调控因子结合调控开花。然而, 关于FLC在成花调控中的具体分子机制仍需深入研究。该文主要结合8条成花调控遗传途径, 梳理近年来与FLC相关的新进展, 并展望了未来的研究方向。     

From the model to the crop: genes controlling tuber formation in potato

Photoperiod regulates many different developmental processes, including floral induction in several species and tuber formation in potato. Research in Arabidopsis led to the identification of FLOWERING LOCUS T (FT) as a main component of the florigen or mobile flowering promoting signal produced in the leaves. A similar mobile signal or tuberigen has been reported to induce tuber formation in potato, recent evidence obtained in our laboratory indicates that a potato homolog of FT encodes this signal. Flowering regulators, like CONSTANS and miR172, also play a role in tuberization, although it remains unclear whether these regulators function in identical pathways. Here, we highlight differential regulation of these genes in flowering and tuberization control and discuss on their possible tuberization-related function.