Florigen is a systemic signal for the transition to flowering in plants. .. Chailakhyan enunciated the concept of a specific flowering hormone and called it . Molecular nature of florigen. (A) Concept of florigen. Florigen is generated in leaf vasculature, transported to the shoot apical meristem and promote flowering. Riv Biol. Jan-Apr;97(1) “Florigen “: an intriguing concept of plant biology. Pennazio S(1). Author information: (1)Istituto di Virologia vegetale del.
|Published (Last):||3 September 2018|
|PDF File Size:||5.20 Mb|
|ePub File Size:||19.85 Mb|
|Price:||Free* [*Free Regsitration Required]|
Florigen is a mobile flowering signal in plants that has a strong impact on plant reproduction and is considered one of the important targets for crop improvement. Recent advances in molecular genetics, cell biology and structural biology in plants revealed the presence of intercellular receptors for florigen, a transcriptional complex essential for florigen to function, and also shed light on the molecular basis of pleiotropic function of florigen beyond flowering.
Furthermore, cutting-edge technologies, such as live cell imaging and next generation sequencing revealed the precise distribution of florigen and transcriptional targets of the florigen activation complex FAC during early stages of floral transition.
These understandings will help future crop improvement through the regulation of flowering and other plant developmental processes. Florigen is a systemic signal that initiates flowering in plants Chailakhyan It is synthesized in leaves and transported to the shoot apical meristem SAM where it promotes floral transition Fig. The molecular function of florigen has long been an important question in the field of flowering studies Tamaki et al.
The identification of florigen facilitated the understanding of its molecular mechanism for the perception and function of FT florigen in detail Tsuji and Taoka A combination of recent advances in techniques such as next generation sequencing and live cell imaging identified its precise distribution and overall function in the shoot apical meristem Tamaki et al.
Molecular nature of florigen. A Concept of florigen. Florigen is generated in leaf vasculature, transported to the shoot apical meristem and cooncept flowering. Square at the top of the stem indicate the enlargement of the shoot apex. Floriven the enlarged picture, green and yellow ovals indicate leaf or flower primordia, respectively. B Cristal structure of Hd3a florigen. Regions colored by blue, yellow and green indicate the regions essential for receptor binding, regions essential for transcriptional activation and other essential residues for florigen function, respectively.
Regulation of flowering time is an important target for plant breeding because the control of flowering to a favorable time provides successful grain production in a given cropping area Jung and Muller Flowering at unfavorable seasons causes loss of yield due to insufficient growth of photosynthetic organs or poor fertility due to heat or cold stress during reproduction.
Thus, understanding the mechanisms of flowering, and especially of the function of florigen, can contribute to novel breeding techniques in crops to produce cultivars that can start their reproductive stage at optimal seasons Tsuji et al. Recent advances in plant genomics help in identifying flprigen regulators of flowering control in many crop species, and also contribute to the understanding of the diversity for future improvements of floral transition.
Studies on the control of flowering include many important areas of plant sciences, such as photoperiodic flowering pathway and day length perception, molecular phonology in field-grown plants, and natural concepy of flowering time Hori et al. Understanding the function of florigen can contribute to many of these research fields because florigen is highly conserved across flowering plants Tsuji et al.
Here I summarize our current understanding of the molecular function of florigen, focusing mainly on rice Oryza sativa florigen Heading date 3a Hd3a. Recent advances in molecular genetics in plants revealed florigenn molecular nature of florigen as a doncept protein named FT, which satisfies the major prerequisites for florigen as a systemic floral signal Corbesier et al. These prerequisites were determined from the long history of classical physiological studies, and they are as follows: Here I will briefly introduce the molecular nature of florigen florigenn describing how FT proteins are recognized.
Photoperiodic flowering has long been considered as a systemic event and the efforts to characterize the genetic factors regulating this process succeeded through molecular genetic analysis of model plants Andres and Coupland Mutants for flowering time regulation in Arabidopsis and quantitative trait loci QTL floriigen of heading date in rice provided rich information for understanding the mechanisms of photoperiodic flowering.
FT in Arabidopsis and Hd3a in rice are genes identified through these studies, and they are orthologs that encode proteins similar to the florogen binding protein PEBP in animals Kardailsky et al. PEPB is a small globular protein with the size of 20 to 25 kD, and shares small pocket on the surface which can interact with the small anions in animal homologs. Both genes are expressed from the phloem of leaves when flowering-promotive day lengths are given—long days for Arabidopsis and short days for rice.
In rice, the precise sites for Hd3a promoter activity, Hd3a mRNA accumulation and Hd3a protein distribution were examined using transgenic plants, and results suggested that Hd3a protein is the mobile floral signal in rice Tamaki et al. Hd3a promoter is active in the phloem of the leaf blade under short day conditions, and Hd3a mRNA is detected only from the leaf blades. On the other hand, Hd3a protein, which was visualized by a fusion protein with green fluorescent protein GFP expressed from Hd3a promoter, accumulates in the SAM.
These observations provided evidence to show that translated Hd3a protein moves from leaf phloem tissue to the SAM, where it initiates floral transition. The results obtained in studies using Arabidopsis, which included the grafting of FT -expressing plants, support this conclusion Notaguchi et al.
However, the precise mechanism for the complex formation has been poorly understood. Molecular genetics and biochemical analyses from rice indicated that the interaction between FT and FD was mediated by another class of FT interacting protein, namedand cellular imaging study indicated a dynamic process of complex formation Fig.
Formation of florigen activation complex FAC. A Scheme for FAC formation in a shoot apex cell of rice. After florigen Hd3a reaches the cell at the shoot apex, Hd3a interacts with proteins in the cytoplasm, then translocated to the nucleus to form FAC with FD.
The resultant FAC activates downstream gene expression. The black bar and orange rectangle in the nucleus indicate a promoter and a downstream gene for FAC, and arrow indicate transcriptional activation.
“Florigen “: an intriguing concept of plant biology.
Two molecules of florigen Hd3a magentaa dimer of florigen receptor blueand C-terminus region of OsFD1 green sticks and red balls essential for direct binding are solved as structure of Florigej.
Several studies have shown a similar complex formation in the rice homologs of Arabidopsis FT and FD, Hd3a and OsFD1, but interestingly, direct interactions in vitro using purified proteins have not been observed. This suggests the presence of additional components that can mediate the interaction of Hd3a and OsFD1 in the cell. This missing link was identified from the assay using another class of Hd3a interacting protein,because interacts directly with Hd3a and OsFD1 through different interfaces on the surface ofand addition of results in the formation of Hd3aOsFD1 complex in vitro.
They form dimers to build their W-shaped structure.
Mutation analysis revealed that the interaction between Hd3a and OsFD1 depends on the interaction of with both the proteins and the function of Hd3a depends completely on its ability to form a complex with OsFD1 through Since the complex formation is essential for Hd3a florigen function, this is named as florigen activation complex FAC Fig.
Cellular imaging of how FAC is formed in the cell revealed that proteins act as intracellular receptors for rice Hd3a florigen. Hd3a forms a subcomplex with in the cytoplasm, where there is an accumulation of The presence of OsFD1 results in the translocation of Hd3a subcomplex from the cytoplasm into the nucleus where OsFD1 localizes.
Based on this dynamic mechanism of complex formation, proteins are considered as the receptor of Hd3a florigen Fig. Another question on florigen function is how florigen changes its distribution and coordinates downstream gene expression.
Early studies of rice florigen imaging failed to obtain detailed localization during floral transition and subsequent inflorescence development Tamaki et al.
Recent advances in the imaging of living plant cells and organs enabled the observation of Hd3a accumulation precisely at early stages of the inflorescence development. Hd3a-GFP is less abundant in the newly developed branch primordia in the inflorescences primary branch meristembut it accumulates at the late stages of branch primordia. This may contribute to restrict precocious initiation of floral meristem in the inflorescences.
In fact, overexpression of Hd3a-GFP from phloem reduced the number of inflorescence branches because of precocious conversion of branch meristem to the floral meristem Tamaki et al. In addition, recent analysis of the genetic basis of heterosis identified florigen function in the inflorescence architecture and the yield of crops Krieger et al.
In tomato, optimal level of expression for genes encoding FAC components contributes fruit yield through the regulation of the timing of floral meristem development Jiang et al.
In rice, extensive genome-wide association studies for heterosis identified Hd3a as the candidate gene that contributes to the expression of heterosis Huang et al. It is interesting to investigate the molecular mechanism that limits florigen Hd3a distribution or its transport in the SAM to optimize inflorescence development. Molecules that may control florigen transport are suggested from the recent molecular genetic analysis of the mutants in Arabidopsis and rice Liu et al. Thus, identification of these downstream genes is important for understanding florigen function.
“Florigen “: an intriguing concept of plant biology.
Interestingly, recent RNA-seq experiments identified that the expression of transposable elements TEs genes cobcept regulated by florigen Hd3a in rice Tamaki et al. Hd3a induces silencing of a subset of TEs at the early stages of floral transition.
The silenced TEs are distributed in similar proportion to all the TEs in the rice genome which are concentrated around the centromeres International Rice Genome Sequencing Projectindicating that the mechanisms by which Hd3a silences Flofigen may be a generic one that can affect it in a genome-wide manner. The possible mechanisms of this regulation include changes of expressions or activities in DNA methylation enzymes and RNA-dependent DNA methylation pathways because these pathways silences TEs and changes of their activities affect TE expressions in genome-wide manner reviewed in Matzke and Mosher The silencing of TEs upon flowering can contribute to the defense of the genome against the activity of TEs and it is possible that the changes in TE expression can act as controlling elements for the genes that play a role during floral transition in rice Tamaki et al.
Molecular basis of pleiotropic vlorigen of florigen. FAC containing FD as a transcription factor subunit promotes flowering right.
On the other hand, FAC with transcription factors TFs other than FD, such as FD-like proteins, promotes processes other than flowering such as storage organ formation and leaf development. Tuber formation in potato is induced by short days, and classical grafting experiments have suggested the involvement of a mobile tuberization signal Navarro et al.
Florigen – Wikipedia
This signal is generated in leaves upon exposure to short days and transported to the underground stem or stolons where it initiates tuberization Navarro et concpet.
This mechanism is quite similar to that of florigen and therefore, the tuber forming signal or tuberigen was considered as a molecule related to florigen.
This suggests that the exchange of transcription factors in FAC can change the function of FT fkorigen on the developmental context. Same conclusions were obtained from molecular genetic analysis of growth cessation in poplar and leaf development and lateral branching in rice Tsuji et al. From these evidences, it is clear that florign functions of florigen at the molecular level beyond flowering are cocnept of FD function and the exchange of transcription factor components of FAC at the specific developmental context Tsuji et al.
The molecular nature of the systemic floral signal, florigen, is a protein product encoded by the FT gene, which is highly conserved across flowering plants. FT is expressed in leaves and transported to the SAM. In SAM cells, FT interacts with its receptor in the cytoplasm and is translocated into the nucleus to interact with the transcription factor Comcept.
FT distribution in the SAM flprigen its function during inflorescence development, and its contribution to the yield of some crops, through the regulation of balance between inflorescence branching and floral meristem formation.
Besides flowering, florigen shows multiple functions such as formation of storage organs and contributes to plant architecture. The exchange of transcription factors in the FAC is the molecular basis of this pleiotropic function. Understanding the molecular function of florigen by cutting-edge technologies florrigen help future crop improvement programs through the regulation of flowering and other plant developmental processes.
Shimamoto passed away on 28th SeptemberNara Institute of Science and Technology for encouraging the florigen studies. National Center for Biotechnology InformationU. Journal List Breed Sci v. Published online Jul Author information Article notes Copyright and License information Disclaimer.
Received Mar 16; Accepted Apr This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.
Tlorigen Florigen is a mobile flowering signal in plants that has a strong impact on plant reproduction and is considered one of the important targets for crop improvement. Introduction Florigen is a systemic signal that initiates flowering in plants Chailakhyan Open in a separate window. Molecular nature of florigen Recent advances in molecular genetics in plants revealed the molecular nature of florigen as a globular protein named FT, which satisfies the major prerequisites for florigen as a systemic floral signal Corbesier et al.
Florigen receptor and activation complex The next important question is how FT forigen exerts floral transition in the SAM.