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Gibberellins
Gibberellins
• Gibberellins are usually associated with plant height. If you apply them to a dwarf plant, the
plant will grow very high. It can overcome genetic dwarfism. While a normal plant treated
with gibberellins shows no response. It can also induce flowering in plants.
• Discovery: In Japan, rice is an economically important plant. Towards the end of the 19th
century, a disease affected the rice plants and resulted in loss of almost 40% of the crop. The
disease was called bakanae or foolish seedling. The plant grows exclusively high that they
cannot support themselves. They become sterile (no seeds), and chlorosis takes place
(yellowing-plants/pale → no chlorophyll produced) and eventually die. They started doing
research and they managed to find a relation between this disease and a fungus called
Gibberella fujikuroi. Later, the scientists in the West became interested in studying this
disease, and they noted that we don’t need the living fungus to infect. They managed to
extract organic chemical from the fungus, and this chemical, when applied to a normal plant,
was enough to cause the same symptoms. They called these chemicals gibberellins.
• Gibberellins has a carboxyl group → Gibberellic acid (GA). There are more than 100 types of
GA, numbered in the order of their discovery. They were discovered in almost all plants but
never in bacteria. These GA have a complicated structure that might remind you of
cholesterol. 4 rings fused together (either 19 or 20 Carbon atoms). They are lipids derived
from isoprene (5 Carbons).
• The synthesis of GA is very similar to cholesterol synthesis, and this same pathway is shared
with carotenoids, ABA, steroids, plastoquinone, and cytokinins (they all start with Acetyl-CoA
precursor and an IPP intermediate):
Acetyl-CoA →→→ Mevalonate →→→ IPP (isopentylpyrophosphate)
• Mevalonate-dependent Pathway:
• Mevalonate-independent Pathway:
- Starts in the plastids.
- It involves G3P and pyruvate.
- No mevalonate intermediate
• The synthesis of GA usually occurs in the embryos of the seeds, in the end leaves, in the roots,
and in intercalary meristems of upper internodes.
• Once GAs are synthesized, they are transported. The transport of GA is not polar. It occurs in
sieve tubes of the phloem. They control physiological activities when they reach target cells
and then they will be inactivated.
• Inactivation of GA:
- Adding OH at a key position → structure modification (example: adding a hydroxyl group
to GA4 converts it to GA34 which is much less active)
- Formation of conjugates → stored form of GAs by attaching them to a molecule. These
molecules are usually small metabolizes (glucose) to form an ester bond or an ether bond.
Whatever is the bond, gibberellin glucoside will be formed and this is a stored form.
• Synthetic compounds:
- There are many synthetic compounds that act as anti-gibberellins or growth retardants.
They inhibit key enzymes in the synthesis process, preventing the synthesis of
gibberellins.
- Example: AMO1618, Phosphon D, and Cycocel (CCC) prevent the conversion of GGPP to
form CPP (the first cyclic step) by inhibiting the enzyme CPS (copalyl pyrophosphate
synthase) → the production of the GA will be blocked.
Some of them even act on more enzymes (phosphon D can act on the next step that
converts CPP to kaurene → it inhibits Kaurene synthase enzyme).
The chemistry of these compounds have been identified, but these are commercial
names.
4. Flowering:
- For the GA to have effect on flowering, two conditions must be met: Light photoperiod
and convenient temperature (cold?) (though some plants show no response to both).
These two factors are needed for gene expression and transcription, which will lead to
the formation of GA and other hormones.
Based on light:
- Long day plants (LDP): the plant flowers when the light period exceeds a certain critical
length. (ex: 16 continuous hours of light). They require long days.
- Short day plants (SDP): The plant will flower if the length of the day is shorter than a
critical length.
- Neutral day plants (NDP): ex: tomato; it flowers when it reaches a certain number of
internodes (no light needed).
Actually, the most important factor in photoperiod is dark rather than the light.
- In the plant, there is a protein pigment called phytochrome. This phytochrome is present
in 2 interconvertible forms:
Pr (absorbs light in the red region 650-680 nm).
Pfr (absorbs light in the far red region 710-740 nm).
When Pr absorbs light in the red region, it will be converted to Pfr →cascade → production
of GA → the plant will flower.
In dark, Pfr is reverted back to Pr → dark reversion → dark periods limit the amount of Pfr
present.
(Pr has a tetra-pyrrole structure (open pyrrole rings/ heme group opened))
- Some plants require cold treatment to flower (vernalization) like carrots and apples.
Ex: Carrots are biennial plants that require 2 growing seasons: vegetative growth and
flowering growth.
- Plants that require long days or vernalization can be induced to flower by adding GA, since
their flowering necessitates the buildup of GA.
- GA are also known to determine sex of flower (whether flower is male or female). Recall
that the flower is made up of 4 floral parts: sepals-petals-stamen-pistil.
Most flowers are hermaphrodite (perfect flowers), having both male and female organs.
Some species, on the other hand, are imperfect (unisexual). Such flowers could be carried
on the same plant (monoecious-cucumber), while others are dioecious (separate plants-
spinach). So GA are known to determine sex. It is known to induce stamenate flowers and
inhibit pistilate flowers from forming.
Note that sex determination can be carried out by other hormones also (ethylene).
6. Parthenocarpy:
It is the production of seedless fruits. Some plants can produce parthenocarpy fruits by
application of GA (ex: pome and stone fruits/ peaches and other fruits that have a stone
seed)
Apples are not responsive to auxins (only GA). Other fruits might be responsive to both
or none.
When two hormones are involved, one hormone usually induces an enzyme that leads to
the production of another hormone.
- Starch is a large molecule needed for germination. It should be broken into glucose and
mobilized. We need amylase (hydrolytic enzyme) to do so. Amylase (and other hydrolytic
enzymes) are produced in the aleurone layer de novo (new enzymes) and will have to
move to the endosperm and hydrolyze the starch so that glucose can move to the
embryo.
- GA are needed for the expression of the genes needed for the synthesis of amylase. GA
is usually present in the embryo and will move to the aleurone layer.
- How can we prove this?
If we remove the embryo. The aleurone layer can no longer produce amylase.
If we add GA → Amylase (and hydrolytic enzymes are expressed).
- Hydrolytic enzymes can be also synthesized and secreted in the part of scutellum facing
the endosperm (it is rich in both ER and GA, which concentrates secretory substances).