FLS – 601 CROP REGULATION IN ORNAMENTAL CROPS

Lecture No. 2 : Photoperiodism- History - Photoperiodic

stimulus - Chailakhyan’s hypothesis -Classification of plants
based on the photoperiodism - Concept of critical day length.
 PHOTOPERIODISM
• Photoperiodism is the phenomenon of physiological changes that occur in plants
in response to relative length of day and night (i.e. photoperiod).

• The response of the plants to the photoperiod, expressed in the form of flowering
is also called as photoperiodism.
 HISTORY OF PHOTOPERIODISM
1852 – Henfrey - Suggested that the daylength at different latitudes in summer might be
responsible for the natural distribution of plants
1910 – Julien Tournois - Worked on hops (Humulus sp.) and Cannabis [short-day plants (SDP)] and
concluded that shorter daylength can accelerate their flowering
1918 – George Klebs - Determined that extended daylength can accelerate flowering of cobweb
sedum [Sempervivum funkii, a long-day plant (LDP)] during winter
1920 – Garner & Allard - Formally proposed the photoperiod flowering phenomenon. They observed
Maryland Mammoth (Tobacco cv.) growing under natural summer conditions
and moved plants to a ventilated darkroom during the afternoon to induce
flowering. They suggested that shortening daylength can induce flowering in
tobacco.
1938 – Hamner & Bonner - Exposed plants to various day lengths with a constant night length or to
various night lengths with a constant day length and found that the length of
the day was not very important for flowering in Xanthium but that it was
essential that the dark period exceeded 8½ hours.
 SIGNIFICANCE OF PHOTOPERIODISM
The knowledge of photoperiodism and its application has great economical potential as
follows

• The yield of tubers, corms, bulbs and rhizomes can be increased substantially by
increasing or decreasing the duration of day or night.
• Annuals may be grown twice or thrice in a year.
• Perennials might flower throughout the year.
• Hybridization experiments have got a fillip because different varieties growing in
different areas with different durations and flowering at different times are made to grow
and flower side by side by artificially controlling their photoperiods.
• Understanding the concept of photoperiodism has helped to choose photo insensitive
varieties and cultivars. These cultivars are best suited in intensive agriculture.
 PHOTOPERIODIC INDUCTION

• The relative length of day and night is known as photoperiod. An appropriate

photoperiod in 24 hours cycle constitutes one inductive cycle.
• Plants require one or more inductive cycles for flowering.
• The number of inductive cycles required for flowering differs from species to species.
• If a plant receives sufficient inductive cycle in an unfavourable photoperiods, it still
flowers. This means flowering also occurs if a plant receives inductive cycles after
intervals of unfavourable photoperiods, i. e., discontinuous inductive cycles. It is called
photoperiodic induction.
• An increase in the number of inductive cycles can cause early flowering.

• For example, Xanthium (SDP) needs one inductive cycle and 64 days for flowering.
But it can flower in 13 days if provided with 4-8 inductive cycles.

• Continuous inductive cycles help more in early flowering than discontinuous

inductive cycles.

• Some plants require more than one inductive cycle. For example, Biloxi soybean
(SDP) requires two inductive cycles, Salvia occidentalis (LDP) requires 17 inductive
cycles, Plantago lanceolata (LDP) requires 25 inductive cycles.
 PERCEPTION OF PHOTOPERIODIC STIMULUS
Knott (1934) first observed that photoperiodic induction is perceived by young expanded
leaves.
Later Knott’s work was confirmed by Chailakhyan (1936).He divided chrysanthemum
plants into four groups and varied the regime of light and dark cycles in all the four
groups by using light proof cases.
The four groups of plants are as follows:
Group A- Entire plant continuously received long day treatment
Group B- Lower leaf portion received short day treatment while upper defoliated
portion received long day treatment
Group C- Lower leafy portion received long day treatment while upper defoliated
portion received short day treatment
Group D- Entire plant continuously received short day treatment
• He observed that flowering occurred in those plants where the leaves received short
day treatment (Group B&D) but it failed in those plants where the leaves received long
day treatment (Group A&C).
• On the basis of above observation he concluded that short day stimulus is perceived by
the leaves.

Floral stimulus
• The flowering stimulus produced in the photo induced leaves is translocated to the
shoot apices for flower evocation.
• It has been observed that floral stimulus is similar in long and short day plants by
grafting experiments.
• If a leaf from a photo induced plant is removed and
then grafted on a non induced plant, then this plant
flowers.
• Apparently a chemical substance is produced
during photo inductive cycle, which is transmitted
during grafting to non induced plants and evokes
flowering.
• Chailakhyan (1936) named this flower inducing
chemical substance as Florigen.
• The flowering stimulus travels through phloem but
independent to the transport of photosynthates.
 THEORIES OF FLOWERING
1. Bunning’s hypothesis
2. Chailakhyan’s hypothesis

Bunning’s hypothesis:

• Bunning (1958) assumes the presence of endogenous rhythms (Oscillator which

consist of two half cycles.
• The first half cycle occurs in day and is called photophilous phase. During this,
anabolic process predominates including flowering in plants.
• The other half cycle is dark, sensitive and is called skotophilous phase. In this,
catabolic process (dehydration of starch) predominates.
• SD plants have a critical day length of 9 hours. This period falls within the
photophilous phase.
• Light during scotophil phase will inhibit photo process initiated during photophase.
• The L.D. plants have a critical day length of 15 hours and some light falls in the skoto
philous phase. Under these conditions in L.D. plants will flower.
• In S.D. plants oscillator is present close to skoto philous phase, while in L.D. plants it
is close to photo philous phase.

Chilakhyan’s hypothesis:
• This hypothesis assumes that flowering hormone – florigen is a complex of two types
of substances – gibberellin and anthesins. Gibberellin is essential for growth of the
plant stems and anthesins are required for flower formation.
• According to him, flowering in all annual seed
plants requires two phases: (i) Floral stem
formation phase (ii) Flower formation phase.
• First phase involves increased carbohydrate
metabolism and respiration with increased
content of GA in leaves.
• Second phase requires intensive nitrogen
metabolism, higher content of anthesins in
leaves and nucleic acid metabolites in stem
buds.
• Long day conditions favour the first phase while
short day conditions favour second phase. In
long day plants gibberellins are critical, while
anthesins are critical in short day plants.
However, anthesin is hypothetical; it has not
been isolated as yet.
 FLORIGEN
Florigen: 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.
Florigen is a hypothetical flowering hormone.

Translocation of florigen:
a) Translocation with same plant:
This hormone is formed in the leaves. Then this hormone multiplies itself. It is the
unique property of this hormone. Then it is translocated through the phloem to the
apical and lateral buds. It causes flowering in them.
b) Translocation between different plants of same species:
Florigen can move from one plant to the other by graft union.
Cont..
It can be proved by a simple experiment. A number of xanthium seedlings grown under
long days. The adjacent branches of a number of plants are grafted together in a series.
Only the terminal branches are exposed to short days.
It causes flowering of all the plants. Therefore, florigen moves from one plant to the other
through the graft union. It can also move both in the upward and downward directions.
c) Translocation between different plants of different species:
Two different taxonomically related plants can also be grafted together.
Tobacco is a short-day plant. Hyoscyamus is a long day plant. Both belong to the family
Solanaceae.
Now branch of tobacco is grafted to Hyoscyamus. They are kept under short day
conditions.
Cont..
Tobacco starts flowering. Interestingly
Hyoscyamus plants also start flowering.
Thus, florigen formed in tobacco is
translocated to Hyoscyamus. It causes
flowering in it.
Similarly, when they were placed in long
day condition again both produces flower.
It indicates florigen is universal flower
forming substance. It found in all the plants
like long day, short day, day neutral plants
etc.
Furthermore, florigen of monocots can
cause flowering in dicot and vice-versa.
Chemical nature of florigen:
 A number of attempts have been made to extract florigen hormone.
 Hamner and others (1964) extracted a substance from the inflorescence of Xanthium.
 But this substance was not effective in stimulating flowering of even Xanthium plants.
Some plant physiologist suggested that some of these hormones might be acting as
florigen. Experiments were performed on these hormones.

• Application of auxin causes flowering only in one of the short-day plant pineapple.
• Gibberellin causes flowering only in rosette long day plant like Hyoscyamus and
vernalizable plant like carrot. GA causes flowering in only one short day plant.
• Similarly, abscisic acid and ethylene also cause flowering in few species.
• Chailakhyan suggested that two factors are involved in the flowering stimulus.
• First factor is gibberellins and the second is anthesins.
• Both gibberellin and anthesins collectively acts a florigen but, cannot proved
experimentally.
• Therefore, no one hormone is taken as florigen. It is possible that the flowering
stimulus is transmitted by mixture of hormones.
CLASSIFICATION OF PLANTS BASED ON PHOTOPERIODISM
Depending upon the duration of photoperiod, plants are classified into three
categories.
• Short day plants
• Long day plants
• Day neutral plants
Short day plants

• These plants require a relatively short day light period (usually 8-10 hours) and a
continuous dark period of about 14-16 hours for subsequent flowering. These plants
are also known as long-night plants
E.g. chrysanthemum, Gerbera, African marigold, Cosmos, Celosia, zinnia.
• In short day plants, the dark period is critical and must be continuous. If this dark
period is interrupted with a brief exposure of red light (660-665 nm), the short day
plant will not flower.
• However, the inhibitory effect of red light can be overcome by a subsequent
exposure with far-red light (730-735 mm wavelength)
• Prolongation of the continuous dark period initiates early flowering.
Long day plants

• These plants require longer day light period

(usually 14-16 hours) in a 24 hours cycle for
subsequent flowering. These plants are also
called as short night plants.

E.g. Aster, Carnation, Hibiscus,

gladiolus, Tuberose, crossandra,
snapdragon, salvia, petunia.

• A brief exposure of red light in the dark

period or the prolongation of light period
stimulates flowering in long day plants.
Day-neutral plants
Day-neutral plants do not flower in response to daylight changes. They flower when
they reach a particular stage of maturity or because of some other cue like
temperature or water, etc. This is the most common kind of flowering pattern.
E.g. Rose, French marigold, Geranium, begonia.
Intermediate categories

Long short day plants

These are short day plants but must be exposed to long days during early
periods of growth for subsequent flowering. E.g. Jasmine.

Short long day plants

These are long day plants but must be exposed to short day during early
periods of growth for subsequent flowering. E.g. certain varieties of wheat and rye.
 CRITICAL DAY LENGTH
“Critical day length” refers to the specific duration of day-light that triggers certain
physiological responses in plants, particularly in relation to flowering and other
developmental processes.

Different plant species have evolved to respond to specific day length conditions, which is
often referred to as photoperiodism.

There are three main categories based on a plant's response to day length:

SDP initiate flowering when day length is shorter than CDL. Example include
chrysanthemums and poinsettia
LDP initiate flowering when day length is longer than CDL. Example include larkspur
DNP are not significantly affected by day length regarding their flowering. They tend to
flower based on other factors like age or environmental conditions. Examples include rose,
zinnia and sunflower.