Unit IV: Phytohormones

Lecture 6
(Gulzar A. Bhat)

Gibberellins signalling

Gibberellins induces GA-induced gene expression by deactivating

repressors (DELLA proteins)

 Bioactive gibberellins promote plant growth and development by

promoting the degradation of the DELLA proteins, a family of nuclear
growth repressors. DELLA proteins are characterized by the presence of
a DELLA motif (aspartate-glutamate-leucine-leucine-alanine). For
example gibberellins act by deactivating repressors (DELLA proteins),
such as SPY (spindly), GAI (gibberellin insensitive), and RGA
(repressor of ga1) en route to both an increase in cell elongation and the
production of α-amylase.

 In a Gibberellin (GA)-deficient cell (Figure A), the transmembrane GA

receptor is inactive in the absence of GA signal. In this situation, SPY is
an active O-GlcNAc transferase that catalyzes the addition of a signal
GlcNAc residue (from UDP-GlcNAc) via an O linkage to specific serine
and/or threonine residues of target proteins, RGA and GAI. Active RGA
and GAI function as repressors of transcription, and they indirectly or
directly inhibit the expression of GA-induced genes.

 In the presence of gibberellins (Figure B), the GA receptor is activated

by binding of bioactive gibberellin. The gibberellin signal inhibits RGA
and GAI repressors both directly and by deactivating SPY. In the absence
of repression by RGA and GAI, GA-induced genes are transcribed.
 (A)

(B)

Figure A & B: Effect of Gibberellins on GA induced gene repressors (SPY, GAI and
RGA)
 De-repression in presence of soluble gibberellin receptor (GID1)
(Figure C): In absence of gibberellins, the DELLA proteins restrain
growth by sequestering transcription factors into inactive protein
complexes. The binding of gibberellins to its soluble receptor GID1
protein permits the interaction between GID1 and DELLA proteins. The
formation of GA-GID1-DELLA complex enhances the interaction
between DELLA and SCF complex, resulting in polyubiquitination of
DELLA and its targeting for degradation via 26S proteasome pathway

Figure C: Degradation of DELLA proteins in presence gibberellins

Physiological role of Gibberellins

1. Gibberellin signaling in Dormancy and Seed Germination

 Seed contains embryo that is arrested to develop into plant with

appropriate environmental conditions to continue their life cycles.
 Breaking of seed dormancy to germination is controlled by some physical
factors (light, temperature and moisture) and by the endogenous growth
regulating hormones.
 Gibberellin stimulates the seed germination whereas; ABA is involved in
the establishment and maintenance of dormancy.
 Gibberellin exerts its influence in two manners, fist by increasing the
growth potential of embryo and second by inducing the expression of
hydrolytic enzymes. During seed germination embryonic GA is released
that triggers the weakness of seed cover by stimulating gene expression
involved in cell expansion and modification.
 Gibberellins represent a natural regulator of the processes involved in
seed germination to stimulate the production of hydrolytic enzyme i.e., α-
amylase, in the aleurone layer of germinating cereal grains.

Cereal grains can be divided into 3 parts i.e., embryo, endosperm and seed
coat. The endosperm is composed of the aleurone layer and centrally located
starchy endosperm. The starchy endosperm, typically non-living at maturity,
consists of thin walled cells with starch grains surrounded by aleurone layer,
having thick cell wall with protein bodies.

 As consequences, the stored food reserves of the starchy endosperms are

broken down into soluble sugars, amino acids, and other products that are
transported to the growing embryo.
Thus gibberellins stimulate mobilization of nutrient reserves during
germination of cereal grains.
 Figure: Structure seed and response to gibberellins during germination

 Gibberellin biosynthesis occurs in the embryo synthesis in response to

water uptake by seeds. These gibberellins diffuse into aleurone layer and
stimulate the synthesis of α-amylase and other hydrolytic enzymes.

 The expression of α-amylase gene in barley aleurone layers is

upregulated by gibberellins, mediated through GA-MYB transcription
factors. This process is mediated by the transcription of a specific
transcription factor, GA-MYB, which binds to the upstream region of
the α-amylase gene, thus switching it on. The gibberellin receptor is
located on the surface of aleurone cells. G-proteins and cyclic GMP have
been implicated as members of the signal transduction chain on the way
to GA-MYB.
Figure: Mechanism of induction of α-amylase synthesis in barley aleurone layers by gibberellins
2. Stimulate Stem elongation and growth
 Gibberellins induce the stem elongation in dwarf and rosette plant like
auxins but it is not involved in acidification of cell wall. It is believed that
gibberellins activate the enzyme xyloglucan endotransglycosylase
(XET) that facilitates the penetration of expansins into the cell wall. So
cell wall elongation of auxin and gibberellin is additive.
 Gibberellins also stimulate stem growth by promoting cell division.
During cell cycle gibberellins result in transition of cell from G 1 to S
phase leading to an increase in mitotic activity. These hormones also
induce the expression of cyclin dependent kinases (CDKs) which are
involved in cell cycle regulation.

3. Bolting and flowering

 In many herbaceous plants, the early period of growth shows rossette

habit with short stem and small leaves. In rosette plants internode gaps
between nodes of stem do not expand so all leaves remain clustered
together due to inactive interclary meristem.
 Under short days, the rosette habit is retained while under long days
bolting occurs i.e. the stem elongates rapidly and is converted into polar
axis bearing flower primordia. This bolting can also be induced in such
plants by the application of gibberellins that stimulates interclary
meristem even under non-inductive short days.
 Figure: Effect of gibberellin treatment on rosette plants

4. Commercial applications of gibberellins

 The major uses of gibberellins, applied as a spray or dip, are to manage

fruit crops, to malt barley, and to increase sugar yield in sugarcane.

i. A major use of gibberellins is to increase the stalk length of seedless

grapes. Because of the shortness of the individual fruit stalks,
bunches of seedless grapes are too compact and the growth of the
berries is restricted. Gibberellin stimulates the stalks to grow longer,
thereby allowing the grapes to grow larger by alleviating compaction,
and it promotes elongation of the fruit.

Figure: Gibberellins induces the growth of seedless fruits

 ii. The principal commercial use of gibberellins is in the production of
table grapes, such as the „„Thompson Seedless.‟‟
iii. Sugarcane (Saccharum officinarum) is one of relatively few plants
that store their carbohydrate as sugar (sucrose) instead of starch.
Sugarcane is a giant perennial grass that can grow from 4 to 6 m tall.
The sucrose is stored in the central vacuoles of the internode
parenchyma cells. Spraying the crop with gibberellin can increase
the yield of raw cane. This increase is a result of the stimulation of
internode elongation during the winter season.
iv. Malting is the first step in the brewing process. During malting,
barley seeds are allowed to germinate at temperatures that maximize
the production of hydrolytic enzymes by the aleurone layer.
Gibberellin is sometimes used to speed up the malting process. The
germinated seeds are then dried and pulverized to produce “malt,”
consisting mainly of a mixture of amylolytic (starch-degrading)
enzymes and partly digested starch.

5. Dormancy of buds

 In temperate regions the buds formed in autumn remain dormant until

next spring due to severe cold. This dormancy of buds can be broken by
gibberellin treatments. In potato also, there is a dormant period after
harvest, but the application of gibberellin sprouts them vigorously.
Assignment 3:
Q.1.Gibberellins were first discovered in Japan when rice plants were suffering from bakane
(the foolish seedling disease) caused by a fungus Gibberella fujikuroi. Which property of
gibberellin caused foolish seedling disease in rice?

Q.2.While eating watermelons, you wish it was seedless. As a plant biochemist can you
suggest any method by which this can be achieved?

Q.3.A farmer growing a particular variety of grape plant in vineyard, observes the following:
A. Fruit size normally remained small
B. Natural seed abortion
C. Development of fungal infection as the pedicles are small in size due to which
moisture is retained in the bunches of grapes.
Expert suggested spraying gibberellic acid during the fruit development. The treatment
would help in getting rid of
a) A, B and C
b) Only A and B
c) Only A and C
d) Only B and C
Q.4. In Signal transduction pathway of which phytohormones genes SPY, DELLA, RGA are
associated
a) Auxin
b) Cytokinin
c) Abscisic acid
d) Gibberellic acid
Q.5. The dwarfing gene which was responsible for green revolution is involved in signal
transduction of over expression of
a) Gibberellic acid
b) Cytokinin
c) Abscisic acid
d) Ethylene
Q.6. How would gibberellic acid be beneficial to plants?
a) By stimulating stem growth
b) By promoting seed storage reserve accumulation
c) By promoting fruit ripening
d) By delaying leaf senescence
Q. 7. The plant hormone gibberellic acid is generally not associated with
a) Stem elongation
b) Nutrient mobilization during seed germination
c) Parthenogenesis
d) Malting
Q.8. To increase sugar production in sugarcanes, they are sprayed with
a) IAA
b) Cytokinin
c) Gibberellic acid
d) Ethylene

Further readings:

1. Ramwant, G and Chakrabarty, SK (2013) Gibberellic acid in plant: Still a mystery

unresolved. Plant Signaling & Behaviour, 8:9, e25504.
2. Sponsel, VM. and Hedden, P. (2004) Gibberellin, biosynthesis and inactivation. In
Plant Hormones: Biosynthesis, Signal Transduction, Action! (Davies, P. J., ed.), pp.
63–94, Springer, Dordrecht.
3. Taiz, L and Zeiger, E (2010) Plant physiology. 5 th Edition, Sinauer Associates Inc.,
Sunderland.