Rangkuman hal 484-490

GIBBERELLIN SIGNAL TRANSDUCTION:

CEREAL ALEURONE LAYER
The biochemical and molecular mechanisms, which are probably common to all gibberellin
responses,

have been studied most extensively in relation to the gibberellin-

stimulated synthesis and secretion of -amylase in
cereal aleurone layers (Jacobsen et al. 1995).

Gibberellin from the Embryo Induces -Amylase

Production by Aleurone Layers
Cereal grains (caryopses; singular caryopsis) can be divided
into three parts: the diploid embryo, the triploid
endosperm, and the fused testa–pericarp (seed coat–fruit
wall). The embryo part consists of the plant embryo proper,
along with its specialized absorptive organ, the scutellum
(plural scutella), which functions in absorbing the solubilized
food reserves from the endosperm and transmitting
them to the growing embryo. The endosperm is composed
of two tissues: the centrally located starchy endosperm and
the aleurone layer

1. GA3-stimulated -amylase production was shown to

be blocked by inhibitors of transcription and translation.
2. Heavy-isotope- and radioactive-isotope-labeling
studies demonstrated that the stimulation of -amylase
activity by gibberellin involved de novo synthesis
of the enzyme from amino acids, rather than activation
of preexisting enzyme.
Definitive molecular evidence now shows that gibberellin
acts primarily by inducing the expression of the protoplasts
and the production of the blue color was shown to
be stimulated by gibberellin (Jacobsen et al. 1995).
The partial deletion of known sequences of bases from
-amylase promoters from several cereals indicates that the
sequences conferring gibberellin responsiveness, termed
gibberellin response elements, are located 200 to 300 base pairs
upstream of the transcription start site

Proteins
on the Plasma Membrane
Acell surface localization of the gibberellin receptor is suggested
from the fact that gibberellin that has been bound to
microbeads that are unable to cross the plasma membrane is
still active in inducing -amylase production in aleurone
protoplasts (Hooley et al. 1991). In addition, microinjection
of GA3 into aleurone protoplasts had no effect, but when the
protoplasts were immersed in GA3 solution, they produced
-amylase (Gilroy and Jones 1994). These results suggest that
gibberellin acts on the outer face of the plasma membrane.

Gibberellins are a family of compounds defined by their

structure. They now number over 125, some of which are
found only in the fungus Gibberella fujikuroi. Gibberellins
induce dramatic internode elongation in certain types of
plants, such as dwarf and rosette species and grasses

Gibberellins are identified and quantified by gas chromatography

combined with mass spectrometry, following
separation by high-performance liquid chromatography.
Bioassays may be used to give an initial idea of the gibberellins
present in a sample. Only certain GAs, notably
GA1 and GA4, are responsible for the effects in plants; the
others are precursors or metabolites.
Gibberellins are terpenoid compounds, made up of isoprene
units. The first compound in the isoprenoid pathway
committed to gibberellin biosynthesis is ent-kaurene. The
biosynthesis up to ent-kaurene occurs in plastids. ent-Kaurene
is converted to GA12—the precursor of all the other
gibberellins—on the plastid envelope and then on the
endoplasmic reticulum via cytochrome P450 monooxygenases.
Commonly a hydroxylation at C-13 also takes place
to give GA53.