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Topic: PHOTOPERIODISM

Course Title: PLANT PHYSIOLOGY AND METABOLISM

Paper: BOT-G-CC-4-4-TH

Unit: 8

Semester: IV

Name of the Teacher: Mrs. Rinku Halder Sahu

Name of the Department: Botany (Morning)


Perception of the Photoperiodic Stimulus and Presence of a Floral 
Hormone: 

I​t is now well established that the​ photoperiodic stimulus is perceived


by the leaves.​ As a result, a floral hormone is produced in the leaves
which is then translocated to the apical tip, subsequently causing the
initiation of floral primordia.

That the photoperiodic stimulus is perceived by the leaves can be shown


by simple ex​periments on cocklebur (​Xanthium pennsylvanicum​), a short
day plant.

● Cocklebur plant will flower if it has previously been kept under


short-day conditions (Fig. 18.2 A).
● If the plant is defoliated and then kept under short day condition, it will
not flower (Fig. 18.2 B).
● Flowering will also occur even if all the leaves of the plant except one
leaf have been removed (Fig. 18.2 C).
● If a cocklebur plant, whether intact or defoliated, is kept under long
day conditions it will not flower (Fig. 18.2 D, E).
● But, if even one of its leaves is exposed to short day conditions and
the rest are under long day photoperiods, flowering will occur (Fig.
18.2 F).
The photoperiodic stimulus can be transmitted from one branch of
the plant to another branch.​ For example,

● If in a two branched cocklebur plant one branch is exposed to short


day and other to long day photoperiod, flowering occurs on both the
branches (Fig. 18.3 A).
● Flowering also occurs if one branch is kept under long day conditions
and another branch from which all the leaves except one have been
removed is exposed to short day condition (Fig. 18.3 B).
● However, if one branch is exposed to long photoperiod and the other
has been defoliated under short day condition, flow​ering will not occur
in any of the branches (Fig. 18.3 C).
Gibberellins and the Flowering Response:  
It is often proposed​ ​that Pfr stimulates gibberellin production and this GA in
turn stimulates the production of floral hormone called florigen.
Importance of GA is due to the following effects of that hormone:
1. External GA can mimic the effect of red light in some cases.
2. GA prompts flowering in some LDPs under short day conditions,
mainly rosette plants which bolt before flowering.
3. GA also inhibits flowering in some SDPs.
4. Anti gibberellin nullifies the above effect of GA.
5. Natural GA from induced LDP also causes flowering in non induced
plants.
Although GA can induce flowering but they can not be considered to be the
primary floral hormone mainly because of the following reasons:
1. In LDPs, vegetative growth and differentiation of floral primordia
occur simultaneously but GA on the other hand first stimulates
vegetative growth (bolting) and then only floral primordia are
formed.
2. GAs have been unable to induce flowering in all short day
plants under a non inductive cycle.
Florigen concept: 

​A mobile molecule that is synthesized in leaves in response to the


favourable photoperiod and it migrates through the vascular system to the
apical meristem to promote floral initiation.

The name florigen has been proposed by Chailakhyan (1936) for the
unknown chemical stimulus acting as floral hormone. The significant
evidence of the existence of that substance comes from interspecific
grafting experiments.

Grafting experiment of florigen existence: 

● Chailakhyan removed the leaves from the upper half of


Chrysanthemum​ (SDP) but left the leaves on lower half untouched.
● He then exposed the lower half to a short day- the plant did not
flower.
● He concluded that day length does not have an effect directly on the
buds but causes the leaves to manufacture a hormone that moves
from the leaves to the buds and induce flowering.
● He concluded that floral stimulus is translocated from the leaf of one
photo induced plant to another non photo induced plant.
● Florigen is believed to be synthesized in the photo induced leaf and
from there moved to the shoot apices for floral evocation , through
phloem.
Grafting experiments in cocklebur plants ​(​Xanthium strumarium)​ have even
proved that the floral hormone can be trans located from one plant to
another. For example,

● if one branched cocklebur plant (Fig. 18.4 A) which has been


exposed to short day conditions is grafted to another cocklebur plant
kept under long day conditions, flowering occurs on both the plants
(Fig. 18.4 B).
● Obviously the flo​ral hormone has been transmitted to the receptor
plant through graft union.
● But if a cocklebur plant is grafted to another similar plant both of
which have been kept under long day condi​tions, flowering will not
occur on either of the two plants (Fig. 18.4 C).
It has also been indicated that the floral hormone may be identical in
short-day and long- day plants. For example, grafting experiments between
certain long-day plants (eg: ​Silene armeria)​ and short-day plants( e.g:
Xanthium strumarium)​ have shown that flowering occurs on both the plants
even if one of them has been kept under non-inductive photoperiods.

But Hamner and Bonner (1938) tested about 246 different kinds of extracts
from SDPs, LDPs, and DNPs but none had any flower evoking effect in non
-inductive plants. So, it is believed that floral stimulus ( florigen) may have
several components which interact with one another in such a way that
some biochemical and cellular changes initiate floral primordia.

Chailakhyan later modified his florigen concept and proposed that

● Florigen is two hormones rather than one, a ​gibberellin​ and a


hypothetical hormone, he called ​anthesin​.
● He suggested that LDPs can produce anthesin under any day length
but GA only under long days,
● SDPs can produce gibberellins under any day length but anthesin
only under short day,
● DNPs can produce both anthesin and GA in any day length.
● Thus a plant can flower only when both GA and anthesin are present.
According to a scheme proposed by Brian (1958), a gibberellin like
hormone is produced in the leaves during the photoperiod somewhat
as follows:

The precursor may be slightly stimulatory or inactive or antagonistic to the


gibberellin-like hormone. Red irradiations promote the conversion of the
precursor to the gibberellin-like hor​mone. In the dark there is a slow
reconversion of the gibberellin-like hormone to the precursor.

This reconversion is accelerated by far-red irradiations. It is further


presumed that high concentration of the gibberellin-like hormone leads to
the synthesis of florigen in long-day plants. In short-day plants the
synthesis of florigen takes place when the level of gibberellin-like hormone
is low. But, flowering eventually follows once the florigen synthesis has
taken place in both the cases.

The whole scheme is diagrammatically shown below:


Nature of the Floral Hormone: 
Recent researches are indicative of ‘florigen’ to be a macromolecule unlike
other plant growth hormones which are rather small molecules. This
macromolecule may possibly be a RNA or protein molecule which is trans
located from the leaf to the apical tips (or meristems) via phloem in
photo-induced plants (Corbesier and Coupland, 2005).

I​n ​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. However, in 2007 another 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".
Here FT acts like a molecular “alarm-clock”, evoking a complex genetic
scenario, which culminates in flower formation.

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