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.[24] 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. [25] 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. [24] 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. [26][27] 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.[28] 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". [29] The initial article[30] that described
FT mRNA as flowering stimuli was retracted by the authors themselves. [31]

Triggers of gene transcription[edit]

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.[32]
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 protein in animals and plants,
controls gene transcriptions.[33]

Flowering mechanism[edit]
A biological mechanism is proposed based on the information we have above. Light is the
flowering signal of Arabidopsis thaliana. Light activates photo-receptors[32] 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. This causes an
influx of calcium ions in the plant. These ions bind to calmodulin and the Ca2+/CaM signaling
system triggers[33] 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. [32]

References[edit]
1. ^ Taiz, Lincoln (2018). "17: Flowering and Fruit Development".  Fundamentals of plant
physiology. Zeiger, Eduardo,, Møller, I. M. (Ian Max), 1950-, Murphy, Angus S. New York,
NY. ISBN 978-1-60535-790-4.  OCLC 1035316853.
2. ^ Taiz, Lincoln (2018). "17: Flowering and Fruit Development".  Fundamentals of plant
physiology. Zeiger, Eduardo,, Møller, I. M. (Ian Max), 1950-, Murphy, Angus S. New York, NY.
p. 474.  ISBN  978-1-60535-790-4. OCLC  1035316853.
3. ^ Taiz, Lincoln (2018). "17: Flowering and Fruit Development".  Fundamentals of plant
physiology. Zeiger, Eduardo,, Møller, I. M. (Ian Max), 1950-, Murphy, Angus S. New York, NY.
p. 477.  ISBN  978-1-60535-790-4. OCLC  1035316853.
4. ^ Jump up to:a b Taiz, Lincoln (2018). "17: Flowering and Fruit Development".  Fundamentals of
plant physiology. Zeiger, Eduardo,, Møller, I. M. (Ian Max), 1950-, Murphy, Angus S. New York,
NY. p.  478. ISBN 978-1-60535-790-4.  OCLC 1035316853.
5. ^ Taiz, Lincoln (2018). "17: Flowering and Fruit Development".  Fundamentals of plant
physiology. Zeiger, Eduardo,, Møller, I. M. (Ian Max), 1950-, Murphy, Angus S. New York, NY.
p. 478.  ISBN  978-1-60535-790-4. OCLC  1035316853.
6. ^ Jump up to:a b c d e Taiz, Lincoln (2018). "17: Flowering and Fruit
Development".  Fundamentals of plant physiology. Zeiger, Eduardo,, Møller, I. M. (Ian Max),
1950-, Murphy, Angus S. New York, NY. p.  478. ISBN 978-1-60535-790-4.  OCLC 1035316853.
7. ^ Jump up to:a b Taiz, Lincoln (2018). "17: Flowering and Fruit Development".  Fundamentals of
plant physiology. Zeiger, Eduardo,, Møller, I. M. (Ian Max), 1950-, Murphy, Angus S. New York,
NY. p.  479. ISBN 978-1-60535-790-4.  OCLC 1035316853.
8. ^ Jump up to:a b Taiz, Lincoln (2018). "17: Flowering and Fruit Development".  Fundamentals of
plant physiology. Zeiger, Eduardo,, Møller, I. M. (Ian Max), 1950-, Murphy, Angus S. New York,
NY. p.  482. ISBN 978-1-60535-790-4.  OCLC 1035316853.
9. ^ Jump up to:a b c Turck F, Fornara F, Coupland G (2008). "Regulation and identity of florigen:
FLOWERING LOCUS T moves center stage"  (PDF). Annual Review of Plant Biology. 59: 573–
94.  doi:10.1146/annurev.arplant.59.032607.092755. hdl:11858/00-001M-0000-0012-374F-8. PMI
D 18444908.
10. ^ Mizoguchi T, Wright L, Fujiwara S, Cremer F, Lee K, Onouchi H, et  al. (August
2005). "Distinct roles of GIGANTEA in promoting flowering and regulating circadian rhythms in
Arabidopsis". The Plant Cell.  17  (8): 2255–
70.  doi:10.1105/tpc.105.033464. PMC  1182487.  PMID  16006578.
11. ^ Jump up to:a b Yanovsky MJ, Kay SA (September 2002). "Molecular basis of seasonal time
measurement in Arabidopsis".  Nature.  419  (6904): 308–
12.  Bibcode:2002Natur.419..308Y. doi:10.1038/nature00996. PMID 12239570.
12. ^ Valverde F, Mouradov A, Soppe W, Ravenscroft D, Samach A, Coupland G (February
2004). "Photoreceptor regulation of CONSTANS protein in photoperiodic
flowering". Science. 303 (5660): 1003–
6. Bibcode:2004Sci...303.1003V. doi:10.1126/science.1091761.  hdl:11858/00-001M-0000-0012-
3C25-D.  PMID  14963328.
13. ^ An H, Roussot C, Suárez-López P, Corbesier L, Vincent C, Piñeiro M, et al. (August
2004). "CONSTANS acts in the phloem to regulate a systemic signal that induces photoperiodic
flowering of Arabidopsis".  Development. 131 (15): 3615–
26.  doi:10.1242/dev.01231.  PMID  15229176.
14. ^ Corbesier L, Vincent C, Jang S, Fornara F, Fan Q, Searle I, et al. (May 2007). "FT
protein movement contributes to long-distance signaling in floral induction of
Arabidopsis". Science. 316 (5827): 1030–
3. Bibcode:2007Sci...316.1030C.  doi:10.1126/science.1141752. hdl:11858/00-001M-0000-0012-
3874-C.  PMID  17446353.
15. ^ Mathieu J, Warthmann N, Küttner F, Schmid M (June 2007). "Export of FT protein from
phloem companion cells is sufficient for floral induction in Arabidopsis".  Current Biology. 17 (12):
1055–60. doi:10.1016/j.cub.2007.05.009. PMID 17540570.
16. ^ Jaeger KE, Wigge PA (June 2007). "FT protein acts as a long-range signal in
Arabidopsis". Current Biology.  17  (12): 1050–4.  doi:10.1016/j.cub.2007.05.008.  PMID  17540569.
17. ^ Jump up to:a b c d e f Tsuji H (September 2017).  "Molecular function of florigen".  Breeding
Science. 67 (4): 327–332. doi:10.1270/jsbbs.17026.  PMC 5654465. PMID 29085241.
18. ^ Taiz L (2018).  Fundamentals of Plant Physiology. 198 Madison Avenue, New York, NY
10016: Oxford University Press. p.  488. ISBN 9781605357904.
19. ^ Jump up to:a b Nakamura Y, Andrés F, Kanehara K, Liu YC, Dörmann P, Coupland G (April
2014). "Arabidopsis florigen FT binds to diurnally oscillating phospholipids that accelerate
flowering". Nature Communications. 5  (1):
3553.  doi:10.1038/ncomms4553. PMC  3988816.  PMID  24698997.
20. ^ Taoka K, Ohki I, Tsuji H, Furuita K, Hayashi K, Yanase T, et al. (July 2011). "14-3-3
proteins act as intracellular receptors for rice Hd3a florigen".  Nature.  476  (7360): 332–
5. doi:10.1038/nature10272. PMID 21804566.
21. ^ Wang JW, Czech B, Weigel D (August 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.
22. ^ Jump up to:a b <Eshed Y, Lippman ZB (November 2019).  "Revolutions in agriculture chart a
coursresee for targeted breeding of old and new crops".  Science.  366  (6466):
eaax0025.  doi:10.1126/science.aax0025. PMID 31488704.
23. ^ Pnueli L, Carmel-Goren L, Hareven D, Gutfinger T, Alvarez J, Ganal M, et  al. (June
1998). "The SELF-PRUNING gene of tomato regulates vegetative to reproductive switching of
sympodial meristems and is the ortholog of CEN and TFL1".  Development. 125 (11): 1979–
89.  PMID  9570763.
24. ^ Jump up to:a b Chaïlakhyan MK (1985). "Hormonal regulation of reproductive development in
higher plants".  Biologia Plantarium. 27 (4–5): 292–302.  doi:10.1007/BF02879865.
25. ^ Chaïlakhyan MK (1975). "Substances of plant flowering".  Biologia Plantarium. 17: 1–
11.  doi:10.1007/BF02921064.
26. ^ Zeevaart JA (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.
27. ^ Bernier G, Havelange A, Houssa C, Petitjean A, Lejeune P (October
1993). "Physiological Signals That Induce Flowering". The Plant Cell.  5 (10): 1147–
1155.  doi:10.1105/tpc.5.10.1147.  PMC 160348. PMID 12271018.
28. ^ Notaguchi M, Abe M, Kimura T, Daimon Y, Kobayashi T, Yamaguchi A, et al.
(November 2008).  "Long-distance, graft-transmissible action of Arabidopsis FLOWERING
LOCUS T protein to promote flowering".  Plant & Cell Physiology.  49  (11): 1645–
58.  doi:10.1093/pcp/pcn154. PMID 18849573.
29. ^ Böhlenius H, Eriksson S, Parcy F, Nilsson O (April
2007). "Retraction".  Science.  316(5823):
367.  doi:10.1126/science.316.5823.367b.  PMID  17446370.
30. ^ Huang T, Böhlenius H, Eriksson S, Parcy F, Nilsson O (September 2005). "The mRNA
of the Arabidopsis gene FT moves from leaf to shoot apex and induces
flowering". Science. 309 (5741): 1694–
6. Bibcode:2005Sci...309.1694H.  doi:10.1126/science.1117768. PMID 16099949.  S2CID 92623
31.
31. ^ Böhlenius H, Eriksson S, Parcy F, Nilsson O (April
2007). "Retraction".  Science.  316(5823):
367.  doi:10.1126/science.316.5823.367b.  PMID  17446370.
32. ^ Jump up to:a b c Mizoguchi T, Wright L, Fujiwara S, Cremer F, Lee K, Onouchi H, et al.
(August 2005). "Distinct roles of GIGANTEA in promoting flowering and regulating circadian
rhythms in Arabidopsis". The Plant Cell.  17  (8): 2255–
70.  doi:10.1105/tpc.105.033464. PMC  1182487.  PMID  16006578.
33. ^ Jump up to:a b Gagliano M, Renton M, Depczynski M, Mancuso S (May 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. PMID 24390479.