Photoperiodism

It is the physiological reaction of organisms to the length of day or night. It

occurs in plants and animals. When it comes to plants in particular,
Photoperiodism can be defined as the developmental responses (such as
physiology of flowering, production of seeds and development of fruits) in
plants to the relative lengths of the light and dark periods.
Many flowering plants (angiosperms) use a photoreceptor protein, such as
phytochrome or cryptochrome to sense seasonal changes in night length,
or photoperiod, which they take as signals to flower. In a further subdivision,
obligate photoperiodic plants absolutely require a long or short enough
night before flowering (so they basically need darkness regardless of
the time to flower), whereas facultative photoperiodic plants are more
likely to flower under the appropriate light conditions (that is, they require
day-time to flower regardless of the period), but will eventually flower
regardless of night length.
In 1920, W. W. Garner and H. A. Allard published their discoveries on
photoperiodism and based on their theories, they felt it was the length of
daylight was critical, but then it was later discovered that it the length of
night-time was the controlling factor. Photoperiodic flowering plants are
classified as long-day plants and short-day plants, even though night is
the critical factor, because of the initial misunderstanding about daylight
being the controlling factor. Each plant has a different length critical
photoperiod, or critical night length.
Modern biologists believe that it is the coincidence of the active forms of
phytochrome or cryptochrome, created by light during the daytime, with the
rhythms of the circadian clock (!) that allows plants to measure the length of
the night. Other than flowering, photoperiodism in plants includes the growth
of stems or roots during certain seasons, or the loss of leaves. Using artificial
lighting to induce long days and nights, short-day plants will bloom but longday plants will note

Long-day
plants

A long-day plant flowers only when the day length exceeds the critical
photoperiod. These plants typically flower in the northern hemisphere during
late spring or early summer as days are getting longer. In the northern
hemisphere, the longest day of the year is 21st of June (solstice). After that,
days grow shorter (i.e. nights grow longer) until 21st of December (solstice).
This situation is reversed in the southern hemisphere (i.e. longest day is 21
December and shortest day is 21 June). In some parts of the world, however,
"winter" or "summer" might refer to rainy versus dry seasons, respectively,
rather than the coolest or warmest time of year.
Some long-day obligate plants are:

Carnation (Dianthus)

Henbane (Hyoscyamus)

Oat (Avena)

Ryegrass (Lolium)

Clover (Trifolium)

Bellflower (Campanula carpatica)

Barley (Hordeum vulgare)

Lettuce (Lactuca sativa)

Wheat (Triticum aestivum)

Turnip (Brassica rapa)

Short-day
plants
Short-day plants flower when the day lengths are less than their critical
photoperiod. They cannot flower under long days or if a pulse of artificial
light is shone on the plant for several minutes during the middle of the night;
they require a consolidated period of darkness before floral development can

begin. Natural nighttime light, such as moonlight or lightning, is not sufficient

brightness or duration to interrupt flowering
In general, short-day (i.e. long-night) plants flower as days grow shorter (and
nights grow longer) after 21 June in the northern hemisphere, which is during
summer or fall. The length of the dark period required to induce flowering
differs among species and varieties of a species.
Photoperiod affects flowering when the shoot is induced to produce
floral buds instead of leaves and lateral buds. Note that some
species must pass through a "juvenile" period during which they
cannot be induced to flowercommon cocklebur is an example of a
plant species with a remarkably short period of juvenility and plants
can be induced to flower when quite small.
Some short-day obligate plants are:

Chrysanthemum

Coffee

Poinsettia

Strawberry

Tobacco,

Common duckweed, (Lemna minor)

Cocklebur (Xanthium)

Maize tropical cultivars only

var. Maryland Mammoth

Some short-day facultative plants are:

Hemp (Cannabis)

Cotton (Gossypium)

Rice

Sugar cane

Day-neutral
plants
Day-neutral plants, such as cucumbers, roses and tomatoes, do not
initiate flowering based on photoperiodism at all! They flower regardless of
the night length. They may initiate flowering after attaining a certain overall
developmental stage or age, or in response to alternative environmental
stimuli, such as vernalization (a period of low temperature), rather than in
response to photoperiod

Vernalization
of the spring)

(from Latin: vernus,

It is the acquisition of a plant's ability to flower or germinate in the spring

after exposure to the prolonged cold of winter. After vernalization, plants
would have acquired the ability to flower, but they may require additional
seasonal cues or weeks of growth before they will actually flower.
Many plants grown in temperate climates require vernalization and must
experience a period of low winter temperature to initiate or accelerate the
flowering process. This ensures that reproductive development and seed
production occurs in spring and summer, rather than in autumn. The needed
cold is often expressed in chill hours. Typical vernalization temperatures are
between 5 and 10degree Celsius (40 and 50 degree Fahrenheit).

For many perennial plants, such as fruit-tree species, a period of cold is

needed to break dormancy, prior to flowering. Many monocarpic annuals and
biennials, including some ecotypes of Arabidopsis thaliana and winter
cereals such as wheat, must go through a prolonged period of cold before
flowering occurs.

In the history of agriculture, farmers observed a traditional distinction

between "winter cereals," whose seeds require chilling, and "spring cereals,
whose seeds can be sown in spring and flower soon thereafter. The word
"vernalization" is a translation of "Jarovization," a word coined by Trofim
Lysenko to describe a chilling process he used to make the seeds of winter
cereals behave like spring cereals ("Jarovoe" in Russian). Scientists had also
discussed how some plants needed cold temperatures to flower, as early as
the 18th century, with the German plant physiologist Gustav Gassner often
mentioned for his 1918 paper.
Lysenko's 1928 paper on vernalization and plant physiology drew wide
attention due to its practical consequences for Russian agriculture. Severe
cold and lack of winter snow had destroyed many early winter wheat
seedlings. By treating wheat seeds with moisture as well as cold, Lysenko
induced them to bear a crop when planted in spring.. Later however, Lysenko
inaccurately asserted that the vernalized state could be inherited - i.e., that
the offspring of a vernalized plant would behave as if they themselves had
also been vernalized and would not require vernalization in order to flower
quickly

Arabidopsis thaliana, also known as "thale cress," is a much-studied model

species. In 2000, the entire genome of its five chromosomes was completely
sequenced. Some variants, called "winter annuals", require vernalization to
flower; others ("summer annuals") do not. The genes that underlie this
difference in plant physiology have been intensively studied.
The reproductive phase change of A. thalliana occurs by a sequence of two
related events: first, the bolting transition (flower stalk elongates), then the
floral transition (first flower appears). Bolting is a robust predictor of flower
formation, and hence a good indicator for vernalization research.

In Arabidopsis winter annuals, the apical meristem is the part of the plant
that needs to be chilled. Vernalization of the meristem appears to confer
competence to respond to floral inductive signals on the meristem. A
vernalized meristem retains competence for as long as 300 days in the
absence of an inductive signal.
Before vernalization, flowering is repressed by the action of a gene called
Flowering Locus Controller (FLC). Vernalization activates a gene called
Frigida (FRI), which progressively turns off FLC expression over a period of six
weeks.

Devernalization
It is possible to devernalize a plant by exposure to high temperatures
subsequent to vernalization. For example, commercial onion growers store
seeds at low temperatures, but devernalize them before planting, because
they want the plant's energy to go into enlarging its bulb (underground
stem), not making flowers.

Phytochrome
Phytochrome is a photoreceptor, a pigment that plants use to detect light.
It is sensitive to light in the red and far-red region of the visible spectrum.
Many flowering plants use it to regulate the time of flowering based on the
length of day and night (photoperiodism) and to set circadian rhythms.
It also regulates other responses including the germination of
seeds(photoblasty), elongation of seedlings, the size, shape and number of
leaves, the synthesis of chlorophyll, and the straightening of the epicotyl or
hypocotyl hook of dicot seedlings. It is found in the leaves of most plants.
Biochemically, phytochrome is a protein with a bilin chromophore.
Phytochrome has been found in most plants including all higher plants; very
similar molecules have been found in several bacteria. A fragment of a
bacterial phytochrome now has a solved three-dimensional protein structure.
Other plant photoreceptors include cryptochromes and
phototropins, which are sensitive to light in the blue and ultra-violet
regions of the spectrum
Phytochromes are characterised by a red/far-red photochromicity.
Photochromic pigments change their "colour" (spectral absorbance
properties) upon light absorption. In the case of phytochrome the ground
state is Pr, the r indicating that it absorbs red light particularly strongly. The
absorbance maximum is a sharp peak 650670 nm, so concentrated
phytochrome solutions look turquoise-blue to the human eye. But once a red
photon has been absorbed, the pigment undergoes a rapid conformational
change to form the Pfr state. Here fr indicates that now not red but far-red
(also called "near infra-red"; 705740 nm) is preferentially absorbed.

This shift in absorbance is apparent to the human eye as a slightly more

greenish colour. When Pfr absorbs far-red light it is converted back to Pr.
Hence, red light makes Pfr, far-red light makes Pr. In plants at least Pfr is the
physiologically active or "signaling" state.

Chemistry of
Phytochrome
Chemically, phytochrome consists of a chromophore, a single bilin molecule
consisting of an open chain of four pyrrole rings, bonded to the protein
moiety. It is the chromophore that absorbs light, and as a result changes the
conformation of bilin and subsequently that of the attached protein,
changing it from one state or isoform to the other.
The phytochrome chromophore is usually phytochromobilin, and is closely
related to phycocyanobilin (the chromophore of the phycobiliproteins
used by cyanobacteria and red algae to capture light for
photosynthesis) and to the bile pigment bilirubin (whose structure is also
affected by light exposure, a fact exploited in the phototherapy of jaundiced
newborns). The term "bili" in all these names refers to bile. Bilins are derived
from the closed tetrapyrrole ring of heame by an oxidative reaction catalysed
by heame oxygenase to yield their characteristic open chain. Chlorophyll too
is derived from heame. In contrast to bilins, heame and chlorophyll carry a
metal atom in the center of the ring, iron or magnesium, respectively.
The Pfr state passes on a signal to other biological systems in the cell, such
as the mechanisms responsible for gene expression. Although this
mechanism is almost certainly a biochemical process, it is still the subject of
much debate. It is known that although Phytochromes are synthesized in the
cytosol and the Pr form is localized there, the Pfr form, when generated by
light illumination, is translocated to the cell nucleus. This implies a role of
phytochrome in controlling gene expression, and many genes are known to
be regulated by phytochrome, but the exact mechanism has still to be fully
discovered. It has been proposed that phytochrome, in the Pfr form, may act
as a kinase, and it has been demonstrated that phytochrome in the Pfr form
can interact directly with transcription factors.

Discovery of
Phytochrome
The phytochrome pigment was discovered by Sterling Hendricks and Harry
Borthwick at the USDA-ARS Beltsville Agricultural Research Center in
Maryland during a period from the late 1940s to the early 1960s. Using a
spectrograph built from borrowed and war-surplus parts, they discovered
that red light was very effective for promoting germination or triggering
flowering responses. The red light responses were reversible by far-red light,
indicating the presence of a photoreversible pigment.
The phytochrome pigment was identified using a spectrophotometer
in 1959 by biophysicist Warren Butler and biochemist Harold
Siegelman. Butler was also responsible for the name, phytochrome.
In 1996 a gene in the newly sequenced genome of the cyanobacterium
Synechocystis was noticed to have a weak similarity to those of plant
phytochromes, the first evidence of phytochromes outside the plant
kingdom. Jon Hughes in Berlin and Clark Lagarias at UC Davis subsequently
showed that this gene indeed encoded a bona fide phytochrome (named
Cph1) in the sense that it is a red/far-red reversible chromoprotein.
Presumably plant phytochromes are derived from an ancestral
cyanobacterial phytochrome, perhaps by gene migration from the chloroplast
to the nucleus. Subsequently phytochromes have been found in other
prokaryotes including Deinococcus radiodurans and Agrobacterium
tumefaciens. In Deinococcus phytochrome regulates the production of
light-protective pigments, however in 1983 the laboratories of Peter Quail
and Clark Lagarias reported the chemical purification of the intact
phytochrome molecule, and in 1985 the first phytochrome gene sequence
was published by Howard Hershey and Peter Quail. By 1989, molecular
genetics and work with monoclonal antibodies that more than one type of
phytochrome existed; for example, the pea plant was shown to have at least
two phytochrome types (then called type I (found predominantly in darkgrown seedlings) and type II (predominant in green plants)).

It is now known by genome sequencing that Arabidopsis has five

phytochrome genes (PHYA - E) but that rice has only three (PHYA - C). While
this probably represents the condition in several di- and monocotyledonous
plants, many plants are polyploid. Hence maize, for example, has six

Phytochromes - phyA1, phyA2, phyB1, phyB2, phyC1 and phyC2. While all
these Phytochromes have significantly different protein components, they all
use phytochromobilin as their light-absorbing chromophore. Phytochrome A
or phyA is rapidly degraded in the Pfr form - much more so than the other
members of the family. In the late 1980s, the Vierstra lab showed that phyA
is degraded by the ubiquitin system, the first natural target of the system to
be identified in eukaryotes.
in Synechocystis and Agrobacterium the biological function of these
pigments is still unknown.
In 2005, the Vierstra and Forest labs at the University of Wisconsin published
a three-dimensional structure of the photosensory domain of Deinococcus
phytochrome. This breakthrough paper revealed that the protein chain forms
a knot - a highly unusual structure for a protein.

Genetic
Engineering and
Phytochrome
Around 1989 several laboratories were successful in producing transgenic
plants which produced elevated amounts of different phytochromes
(overexpression). In all cases the resulting plants had conspicuously short
stems and dark green leaves. Harry Smith and co-workers at Leicester
University in England showed that by increasing the expression level of
phytochrome A (which responds to far-red light), shade avoidance responses
can be altered.[5] As a result, plants can expend less energy on growing as
tall as possible and have more resources for growing seeds and expanding
their root systems. This could have many practical benefits: for example,
grass blades that would grow more slowly than regular grass would not
require mowing as frequently, or crop plants might transfer more energy to
the grain instead of growing taller.

Significance of
Phytochrome
Protein

They mainly help in the development of the plant-physiology of

flowering, development of fruits, and production of seeds.
They are sensitive to prolonged duration of day and darkness. Thus,
they undergo a conformational change in the 4th Carbon of the pyrrole
ring structure to trigger the action of the protein.
They ensure that the developmental changes in plants take place at
the right time.
They ensure that the dormancy in plants is broken at the right time.

***
THE NATIONAL DEGREE COLLEGE (AUTONOMOUS)
Basavangudi, Bangalore-560 098

A brief report on-

Physiology
of flowering

By,
Arun S
09NCBS1002
VI semester B.Sc. section

Geometric representation of a pyrrole molecule