Florigen

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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
disputed.

Contents

 1Mechanism
o 1.1Initiation
o 1.2Translocation
o 1.3Flowering
o 1.4Antiflorigen
 2Research history
 3Triggers of gene transcription
 4Flowering mechanism
 5References
 6External links

Mechanism[edit]
Essentially, to understand Florigen, you must first understand how flowering works.
For a plant to begin flowering, it must make its changes to the shoot apical
meristem (SAM).[1] However, there are factors the plant must first consider before it
begins this process such as the environment but even more specifically, light. It is
through "the evolution of both internal and external control systems that enables
plants to precisely regulate flowering so that it occurs at the optimal time for
reproductive success."[2] The way the plant determines this optimal time is through
day-night periods through the use of Photoperiodism. Although it was originally
thought that the accumulation of photosynthetic products controlled the flowering of
plants, two men by the names of Wightman Garner and Henry Allard proved it was
not.[3] They instead found that it was a matter of day length rather than the
accumulation of the products within the plants that affected their flowering abilities.
Flowering plants fall into two main photoperiodic response categories:

1. "Short-day plants (SDPs) flower only in short days (qualitative SDPs), or their

flowering is accelerated by short days (quantitative SDPs)"[4]
2. "Long-day plants (LDPs) flower only in long days (qualitative LDPs), or their
flowering is accelerated by long days (quantitative LDPs)"[4]
These types of flowering plants are differentiated by the whether the day has
exceeded some duration - usually calculated by 24-hour cycles - known as
the critical day length.[5] It is also important to note that there is no absolute value
for the minimum day length as it varies greatly amid species. Until the correct
amount of day length is reached, the plants ensure no flowering results. They do so
through adaptations like preventing immature plants from responding to inadequate
day lengths.[6] Plants also have the ability to prevent the response of the
photoperiodic stimulus until a certain temperature is reached. [6] Species like winter
wheat that rely on just that.[6] The wheat require a cold period before being able to
respond to the photoperiod.[6] This is known as vernalization or overwintering.[6]
This ebb-and-flow of flowering in plants is essentially controlled by an internal clock
known as the endogenous oscillator.[7] It is thought that these internal pacemakers
"are regulated by the interaction of four sets of genes expressed in the dawn,
morning, afternoon, and evening hours [and that] light may augment the amplitude of
the oscillations by activating the morning and evening genes." [7] The rhythms
between these different genes are generated internally in the plants, starts with the
leaves, but requires an environmental stimulus such as light. The light essentially
stimulates the transmission of a floral stimulus (florigen) to the shoot apex when the
correct amount of day-length is perceived.[8] This process is known as photoperiodic
induction and is a photoperiod-regulated process that is also dependent on the
endogenous oscillator.[8]
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. [9] 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.
Initiation[edit]
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
circadian rhythms, and is then translated into CO protein.[10][11] 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 some light. [11][12] CO protein
promotes transcription of another gene called Flowering Locus T (FT). [13] 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. [9]
Translocation[edit]
The FT protein resulting from the short period of CO transcription factor activity is
then transported via the phloem to the shoot apical meristem.[14][15][16]
Flowering[edit]
Florigen is a systemically mobile signal that is synthesized in leaves and the
transported via the phloem to the shoot apical meristem (SAM) where it initiates
flowering.[17][18] In Arabidopsis, the FLOWERING LOCUS T (FT) genes encode for the
flowering hormone and in rice the hormone is encoded by Hd3a genes thereby
making these genes orthologs.[17] It was found though the use of transgenic plants
that the Hd3a promoter in rice is located in the phloem of the leaf along with
the Hd3a mRNA. However, the Hd3a protein is found in neither of these places but
instead accumulates in the SAM which shows that Hd3a protein is first translated in
leaves and then transported to the SAM via the phloem where floral transition is
initiated; the same results occurred when looked at Arabidopsis.[17] These results
conclude that FT/Hd3a is the florigen signal that induces floral transition in plants.
Upon this conclusion, it became important to understand the process by which the
FT protein causes floral transition once it reaches the SAM. The first clue came with
looking at models from Arabidposis which suggested that a bZIP domain containing
transcription factor, FD, is somehow interacting with FT to form a transcriptional
complex that activates floral genes.[17] Studies using rice found that there is an
interaction between Hd3a and OsFD1, homologs of FT and FD respectively, that is
mediated by the 14-3-3 protein GF14c.[17][19] The 14-3-3 protein acts as intracellular
florigen receptor that interacts directly with Hd3a and OsFD1 to form a tri-protein
complex called the florigen activation complex (FAC) because it is essential for
florigen function.[17] The FAC works to activate genes needed to initiate flowering at
the SAM; flowering genes in Arabidopsis include AP1, SOC1 and several SPL
genes, which are targeted by a microRNA and in rice the flowering gene
is OsMADS15 (a homolog of AP1).[19][20][21]
Antiflorigen[edit]
Florigen is regulated by the action of an antiflorigen. [22] Antiflorigens are hormones
that are encoded by the same genes for florigen that work to counteract its function.
[22]
 The antiflorigen in Arabidopsis is TERMINAL FLOWER1 (TFL1)[9] and in tomato it
is SELF PRUNING (SP)