Florigen (or flowering hormone) is the hypothesized hormone-like molecule responsible for controlling and/or triggering flowering in plants. Florigen is produced in the leaves, and acts in the shoot apical meristem of buds and growing tips. It is known to be graft-transmissible, and even functions between species. However, despite having been sought since the 1930s, the exact nature of florigen is still a mystery.
Central to the hunt for florigen is an understanding of how plants use seasonal changes in day length to mediate flowering—a mechanism known as photoperiodism. Plants which exhibit photoperiodism may be either 'short day' or 'long day' plants, which in order to flower require short days or long days respectively, although plants in fact distinguish day length from night length.
The current model suggests the involvement of multiple different factors. Research into florigen is predominately centred on the model organism and long day plant, Arabidopsis thaliana. Whilst much of the florigen pathways appear to be well conserved in other studied species, variations do exist. The mechanism may be broken down into three stages: photoperiod-regulated initiation, signal translocation via the phloem, and induction of flowering at the shoot apical meristem.
In Arabidopsis thaliana, the signal is initiated by the production of messenger RNA (mRNA) coding a transcription factor called CONSTANS (CO). CO mRNA is produced approximately 12 hours after dawn, a cycle regulated by the plant's biological clock. This mRNA is then translated into CO protein. However CO protein is stable only in light, so levels stay low throughout short days and are only able to peak at dusk during long days when there is still a little light. CO protein promotes transcription of another gene called Flowering Locus T (FT). By this mechanism, CO protein may only reach levels capable of promoting FT transcription when exposed to long days. Hence, the transmission of florigen—and thus, the induction of flowering—relies on a comparison between the plant's perception of day/night and its own internal biological clock.
At the shoot apical meristem, the FT protein interacts with a transcription factor (FD protein) to activate floral identity genes, thus inducing flowering. Specifically, arrival of FT at the shoot apical meristem and formation of the FT/FD heterodimer is followed by the increased expression of at least one direct target gene, APETALA 1 (AP1), along with other targets, such as SOC1 and several SPL genes, which are targeted by a microRNA.
This section needs expansion. You can help by adding to it. (April 2009)
Florigen was first described by Soviet Armenian plant physiologist Mikhail Chailakhyan, who in 1937 demonstrated that floral induction can be transmitted through a graft from an induced plant to one that has not been induced to flower. Anton Lang showed that several long-day plants and biennials could be made to flower by treatment with gibberellin, when grown under a non-flower-inducing (or non-inducing) photoperiod. This led to the suggestion that florigen may be made up of two classes of flowering hormones: Gibberellins and Anthesins. It was later postulated that during non-inducing photoperiods, long-day plants produce anthesin, but no gibberellin while short-day plants produce gibberellin but no anthesin. However, these findings did not account for the fact that short-day plants grown under non-inducing conditions (thus producing gibberellin) will not cause flowering of grafted long-day plants that are also under noninductive conditions (thus producing anthesin).
As a result of the problems with isolating florigen, and of the inconsistent results acquired, it has been suggested that florigen does not exist as an individual substance; rather, florigen's effect could be the result of a particular ratio of other hormones. However, more recent findings indicate that florigen does exist and is produced, or at least activated, in the leaves of the plant and that this signal is then transported via the phloem to the growing tip at the shoot apical meristem where the signal acts by inducing flowering. In Arabidopsis thaliana, some researchers have identified this signal as mRNA coded by the FLOWERING LOCUS T (FT) gene, others as the resulting FT protein. First report of FT mRNA being the signal transducer that moves from leaf to shoot apex came from the publication in Science Magazine. However, in 2007 other group of scientists made a breakthrough saying that it is not the mRNA, but the FT Protein that is transmitted from leaves to shoot possibly acting as "Florigen". The initial article that described FT mRNA as flowering stimuli was retracted by the authors themselves.
Determination of the Role of GI, CO, and FT Genes, and Ca2+/CaM triggers Gene TranscriptionEdit
There are three genes involved in clock-controlled flowering pathway, GIGANTEA (GI), CONSTANS (CO), and FLOWERING LOCUS T (FT). Constant overexpression of GI from the Cauliflower mosaic virus 35S promoter causes early flowering under short day so an increase in GI mRNA expression induces flowering. Also, GI increases the expression of FT and CO mRNA, and FT and CO mutants showed later flowering time than GI mutant. In other words, functional FT and CO genes are required for flowering under short day. In addition, these flowering genes accumulate during light phase and decline during dark phase, which are measured by green fluorescent protein. Thus, their expressions oscillate during the 24-hour light-dark-cycle. In conclusion, the accumulation of GI mRNA alone or GI, FT, and CO mRNA promote flowering in Arabidopsis thaliana and these genes expressed in the temporal sequence GI-CO-FT.
Action potential triggers calcium flux into neurons in animal or root apex cells in plants. The intracellular calcium signals are responsible for regulation of many biological functions in organisms. For instance, Ca2+ binding to calmodulin, a Ca2+-binding proteins in aninals and plants, controls gene transcriptions.
A Possible Mechanism for FloweringEdit
A biological mechanism is proposed based on the information we have above. Light is the flowering signal of Arabidopsis thaliana. Light activates photo-receptors and triggers signal cascades in plant cells of apical or lateral meristems. Action potential is spread via the phloem to the root and more voltage-gated calcium channels are opened along the stem. There is increase in calcium ions influx in plant cells. These ions bind to calmodulin and the Ca2+/CaM signaling system triggers the expression of GI mRNA or FT and CO mRNA. The accumulation of GI mRNA or GI-CO-FT mRNA during the day causing the plant to flower.
- Garner W.W.; Allard H.A. (1920). "Effect of the relative length of day and night and other factors of the environment on growth and reproduction in plants". Journal of Agricultural Research. 18 (7): 553–606. Bibcode:1920MWRv...48..415G. doi:10.1175/1520-0493(1920)48<415b:EOTRLO>2.0.CO;2.
- Turck, F.; Fornara, F.; Coupland, G. (2008). "Regulation and Identity of Florigen: FLOWERING LOCUS T Moves Centre Stage" (PDF). Annual Review of Plant Biology. 59: 573–594. doi:10.1146/annurev.arplant.59.032607.092755. PMID 18444908.
- Mizoguchi, T.; Wright, L.; Fujiwara, S.; Cremer, F.; Lee, K.; et al. (2005). "Distinct roles of GIGANTEA in promoting flowering and regulating circadian rhythms in Arabidopsis". Plant Cell. 17 (8): 2255–2270. doi:10.1105/tpc.105.033464. PMC 1182487. PMID 16006578.
- Yanovsky, M.J.; Kay, S.A. (2002). "Molecular basis of seasonal time measurement in Arabidopsis". Nature. 419 (6904): 308–312. Bibcode:2002Natur.419..308Y. doi:10.1038/nature00996. PMID 12239570.
- Valverde, F.; Mouradov, A.; Soppe, W.; Ravenscroft, D.; Samach, A.; Coupland, G. (2004). "Photoreceptor regulation of CONSTANS protein in photoperiodic flowering". Science. 303 (5660): 1003–1006. Bibcode:2004Sci...303.1003V. doi:10.1126/science.1091761. PMID 14963328.
- An, Hailong; Roussot, Clotilde; Suárez-López, Paula; Corbesier, Laurent; Vincent, Coral; Piñeiro, Manuel; Hepworth, Shelley; Mouradov, Aidyn; Justin, Samuel (2004-08-01). "CONSTANS acts in the phloem to regulate a systemic signal that induces photoperiodic flowering of Arabidopsis". Development. 131 (15): 3615–3626. doi:10.1242/dev.01231. ISSN 0950-1991. PMID 15229176.
- Corbesier, L.; Vincent, C.; Jang, S.; Fornara, F.; Fan, Q.; et al. (2007). "FT protein movement contributes to long distance signalling in floral induction of Arabidopsis". Science. 316 (5827): 1030–1033. Bibcode:2007Sci...316.1030C. doi:10.1126/science.1141752. PMID 17446353.
- Mathieu J, Warthmann N, Küttner F, Schmid M (2007). "Export of FT protein from phloem companion cells is sufficient for floral induction in Arabidopsis". Curr Biol. 17 (12): 1055–60. doi:10.1016/j.cub.2007.05.009. PMID 17540570.
- Jaeger, Katja E.; Wigge, Philip A. (2007). "FT Protein Acts as a Long-Range Signal in Arabidopsis". Current Biology. 17 (12): 1050–1054. doi:10.1016/j.cub.2007.05.008.
- Wigge PA, Kim MC, Jaeger KE, Busch W, Schmid M, Lohmann JU, Weigel D (2005). "Integration of spatial and temporal information during floral induction in Arabidopsis". Science. 309 (5737): 1056–1059. Bibcode:2005Sci...309.1056W. doi:10.1126/science.1114358. PMID 16099980.
- Abe, M.; Kobayashi, Y.; Yamamoto, S.; Daimon, Y.; Yamaguchi, A.; et al. (2005). "FD, a bZIP protein mediating signals from the floral pathway integrator FT at the shoot apex". Science. 309 (5737): 1052–1056. Bibcode:2005Sci...309.1052A. doi:10.1126/science.1115983. PMID 16099979.
- Wang JW, Czech B, Weigel D (2009). "miR156-regulated SPL transcription factors define an endogenous flowering pathway in Arabidopsis thaliana". Cell. 138 (4): 738–49. doi:10.1016/j.cell.2009.06.014. PMID 19703399.
- Chaïlakhyan, M.K. (1985). "Hormonal regulation of reproductive development in higher plants". Biologia Plantarium. 27 (4–5): 292–302. doi:10.1007/BF02879865.
- Chaïlakhyan, M.K. (1975). "Substances of plant flowering". Biologia Plantarium. 17: 1–11. doi:10.1007/BF02921064.
- Zeevaart, J.A.D. (1976). "Physiology of flower formation". Annual Review of Plant Physiology and Plant Molecular Biology. 27: 321–348. doi:10.1146/annurev.pp.27.060176.001541.
- Bernier, G.; Havelange, A.; Houssa, C.; Petitjean, A. & Lejeune, P. (1993). "Physiological signals that induce flowering". Plant Cell. 5 (10): 1147–1155. doi:10.1105/tpc.5.10.1147. PMC 160348. PMID 12271018.
- Notaguchi, Michitaka; Mitsutomo Abe; Takahiro Kimura; Yasufumi Daimon; Toshinori Kobayashi; Ayako Yamaguchi; Yuki Tomita; Koji Dohi; Masashi Mori; Takashi Araki (2008). "Long-Distance, Graft-Transmissible Action of Arabidopsis FLOWERING LOCUS T Protein to Promote Flowering". Plant Cell Physiol. 49 (11): 1645–1658. doi:10.1093/pcp/pcn154. PMID 18849573. Retrieved 29 January 2011.
- Bohlenius, H.; Eriksson, S.; Parcy, F.; Nilsson, O. (2007). "Retraction". Science. 316 (5823): 367b. doi:10.1126/science.316.5823.367b.
- Huang, T.; Böhlenius, Henrik; Eriksson, Sven; Parcy, François; Nilsson, Ove (2005). "The mRNA of the Arabidopsis Gene FT Moves from Leaf to Shoot Apex and Induces Flowering". Science. 309 (5741): 1694–1696. Bibcode:2005Sci...309.1694H. doi:10.1126/science.1117768.
- Mizoguchi, Tsuyoshi (August 2005). "Distinct Roles of GIGANTEA in Promoting Flowering and Regulating Circadian Rhythms in Arabidopsis". The Plant Cell. 17 (8): 2255–2270. doi:10.1105/tpc.105.033464. PMC 1182487.
- Gagliano, Monica (January 5, 2014). "Experience teaches plants to learn faster and forget slower in environments where it matters". Oecologia. 175 (1): 63–72. Bibcode:2014Oecol.175...63G. doi:10.1007/s00442-013-2873-7.
- Zeevaart, J.A.D. (August 2006). "Florigen Coming of Age after 70 Years". The Plant Cell. 18 (8): 1783–1789. doi:10.1105/tpc.106.043513. PMC 1533981. PMID 16905662.
- Zeevaart, J.A.D. (September 2007). "FT Protein, not mRNA, is the Phloem-Mobile Signal for Flowering". Plant Physiology Online.
- Blooming Marvelous: Long sought-after flowering signal on Science’s list of Breakthroughs of 2005