Plants respond to the changing seasons to initiate developmental programmes precisely at particular times of year. Flowering is the best characterized of these seasonal responses, and in temperate climates it often occurs in spring. Genetic approaches in Arabidopsis thaliana have shown how the underlying responses to changes in day length (photoperiod) or winter temperature (vernalization) are conferred and how these converge to create a robust seasonal response. Recent advances in plant genome analysis have demonstrated the diversity in these regulatory systems in many plant species, including several crops and perennials, such as poplar trees. Here, we report progress in defining the diverse genetic mechanisms that enable plants to recognize winter, spring and autumn to initiate flower development.

The genetic basis of flowering responses to seasonal cues

Transcriptional Regulatory Network of Plant Heat Stress Response

http://europepmc.org/articles/PMC3796012?pdf=render

Flowering time regulation: photoperiod- and temperature-sensing in leaves.

Many plants use information about changing day length (photoperiod) to align their flowering time with seasonal changes to increase reproductive success. A mechanism for photoperiodic time measurement is present in leaves, and the day-length-specific induction of the FLOWERING LOCUS T (FT) gene, which encodes florigen, is a major final output of the pathway. Here, we summarize the current understanding of the molecular mechanisms by which photoperiodic information is perceived in order to trigger FT expression in Arabidopsis as well as in the primary cereals wheat, barley, and rice. In these plants, the differences in photoperiod are measured by interactions between circadian-clock-regulated components, such as CONSTANS (CO), and light signaling. The interactions happen under certain day-length conditions, as previously predicted by the external coincidence model. In these plants, the coincidence mechanisms are governed by multilayered regulation with numerous conserved as well as unique regulatory compon...

Photoperiodic flowering: time measurement mechanisms in leaves.

Phytochrome-interacting factors (PIFs) are members of the Arabidopsis thaliana basic helix-loop-helix family of transcriptional regulators that interact specifically with the active Pfr conformer of phytochrome (phy) photoreceptors. PIFs are central regulators of photomorphogenic development that act to promote stem growth, and this activity is reversed upon interaction with phy in response to light. Recently, significant progress has been made in defining the transcriptional networks directly regulated by PIFs, as well as the convergence of other signaling pathways on the PIFs to modulate growth. Here, we summarize and highlight these findings in the context of PIFs acting as integrators of light and other signals. We discuss progress in our understanding of the transcriptional and posttranslational regulation of PIFs that illustrates the integration of light with hormonal pathways and the circadian clock, and we review seedling hypocotyl growth as a paradigm of PIFs acting at the interface of these signals. Based on these advances, PIFs are emerging as required factors for growth, acting as central components of a regulatory node that integrates multiple internal and external signals to optimize plant development.

/pdf/pifs-systems-integrators-in-plant-development-1uesk8que0.pdf

PIFs: systems integrators in plant development

N(6)-Methyladenosine RNA Modification Regulates Shoot Stem Cell Fate in Arabidopsis.

The capacity to respond to day length, photoperiodism, is crucial for flowering plants to adapt to seasonal change. The photoperiodic control of flowering in plants is mediated by a long-distance mobile floral stimulus called florigen that moves from leaves to the shoot apex. Although the proteins encoded by FLOWERING LOCUS T (FT) in Arabidopsis and its orthologs in other plants are identified as the long-sought florigen, whether their transport is a simple diffusion process or under regulation remains elusive. Here we show that an endoplasmic reticulum (ER) membrane protein, FT-INTERACTING PROTEIN 1 (FTIP1), is an essential regulator required for FT protein transport in Arabidopsis. Loss of function of FTIP1 exhibits late flowering under long days, which is partly due to the compromised FT movement to the shoot apex. FTIP1 and FT share similar mRNA expression patterns and subcellular localization, and they interact specifically in phloem companion cells. FTIP1 is required for FT export from companion cells to sieve elements, thus affecting FT transport through the phloem to the SAM. Our results provide a mechanistic understanding of florigen transport, demonstrating that FT moves in a regulated manner and that FTIP1 mediates FT transport to induce flowering.

/pdf/ftip1-is-an-essential-regulator-required-for-florigen-1srn28j463.pdf

FTIP1 is an essential regulator required for florigen transport.

Nuclear factor Y (NF-Y) is a conserved heterotrimeric transcription factor complex that binds to the CCAAT motifs within the promoter region of many genes. In plants, a large number of genes code for variants of each NF-YA, B or C subunit that can assemble in a combinatorial fashion. Here, we report the discovery of an Arabidopsis NF-Y complex that exerts epigenetic control over flowering time by integrating environmental and developmental signals. We show that NF-Y interacts with CONSTANS in the photoperiod pathway and DELLAs in the gibberellin pathway, to directly regulate the transcription of SOC1, a major floral pathway integrator. This NF-Y complex binds to a unique cis-element within the SOC1 promoter to modulate trimethylated H3K27 levels, partly through a H3K27 demethylase REF6. Our findings establish NF-Y complexes as critical mediators of epigenetic marks that regulate the response to environmental or intrinsic signals in plants.

/pdf/nuclear-factor-y-mediated-h3k27me3-demethylation-of-the-soc1-12y49i3uj5.pdf

Nuclear factor Y-mediated H3K27me3 demethylation of the SOC1 locus orchestrates flowering responses of Arabidopsis

Floral transition in Arabidopsis is tightly controlled by complex genetic regulatory networks in response to endogenous and environmental flowering signals. SUPPRESSOR OF OVEREXPRESSION OF CONSTANS 1 (SOC1) and SHORT VEGETATIVE PHASE (SVP), two key MADS-domain transcription factors, perceive these signals and function as antagonistic flowering regulators. To understand how these factors mediate floral transition, we mapped in vivo binding sites of SOC1 and SVP using chromatin immunoprecipitation followed by hybridization to whole-genome tiling arrays (ChIP-chip). Genes that encoded proteins with transcription regulator activity and transcription factor activity were the most enriched groups of genes of those bound by SOC1 and SVP, which indicates their central roles in flowering regulatory networks. In combination with gene expression microarray studies, we further identified the genes whose expression was controlled directly by SOC1 or SVP. Among the common direct targets identified, APETALA2 (AP2)-like genes that repress FT and SOC1 expression were down-regulated by SOC1, but up-regulated by SVP, revealing a complex feedback regulation among the key genes that determine the integration of flowering signals. SOC1 regulatory regions were also accessed by SOC1 itself and SVP, suggesting that self-activation and repression by SVP contribute to the control of SOC1 expression. In addition, ChIP-chip analysis demonstrated that miR156e and miR172a, which are involved in the regulation of AP2-like genes, were direct targets of SOC1 and SVP, respectively. Taken together, these findings revealed that feedback regulatory loops mediated by SOC1 and SVP are essential components of the gene regulatory networks that underpin the integration of flowering signals during floral transition.

https://onlinelibrary.wiley.com/doi/pdf/10.1111/j.1365-313X.2012.04919.x

Genome‐wide identification of SOC1 and SVP targets during the floral transition in Arabidopsis

Flowering plants perceive photoperiodic signals in leaves to generate mobile stimuli required for the induction of flower formation at shoot apices Although FLOWERING LOCUS T (FT) has been identified as part of the mobile floral stimuli in Arabidopsis thaliana, the mechanisms underlying long-distance movement of FT from leaves to shoot apices remain largely unclear Here we show that a heavy-metal-associated (HMA) domain-containing protein, SODIUM POTASSIUM ROOT DEFECTIVE 1 (NaKR1), is activated by CONSTANS (CO) under long-day conditions and regulates long-distance movement of FT in Arabidopsis Loss of function of NaKR1 compromises FT transport to shoot apices through sieve elements, causing late flowering under long-day conditions NaKR1 and FT share similar expression patterns and subcellular localization, and interact with each other in vivo Grafting experiments demonstrate that NaKR1 promotes flowering through mediating FT translocation from leaves to shoot apices Thus, photoperiodic control of floral induction requires NaKR1-mediated long-distance delivery of florigenic signals

Lu Liu

Papers

N(6)-Methyladenosine RNA Modification Regulates Shoot Stem Cell Fate in Arabidopsis.

FTIP1 is an essential regulator required for florigen transport.

Nuclear factor Y-mediated H3K27me3 demethylation of the SOC1 locus orchestrates flowering responses of Arabidopsis

Genome‐wide identification of SOC1 and SVP targets during the floral transition in Arabidopsis

NaKR1 regulates long-distance movement of FLOWERING LOCUS T in Arabidopsis.