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Unique
properties of plant phosphoinositide metabolism: A characteristic
of plant phosphoinositide metabolism is that PtdIns(4)P levels are
typically 20 times higher than those of PtdIns(4,5)P2,
suggesting that PtdIns(4)P may be the key regulator of cellular
metabolism and a signaling molecule in its own right, which however
has yet to be characterized.
Major
differences between plants, yeasts and animals are evident in the
plasma membrane-associated phospholipase C (PLC), which hydrolyzes
PtdIns(4,5)P2 to produce Ins(1,4,5)P3. In
animal cells, there are three major families of PLCs: PLCb, which
is regulated by G-proteins, PLCg, which is regulated by phosphotyrosine
receptors, and PLCd, which is regulated by Ca2+ and possibly
other mechanisms. Plants and yeast contain only the d-family of
PLCs. In addition, unlike the animal and yeast enzymes, none of
the plant d family of PLCs contain a PH domain. The PLC-PH domain
is thought to facilitate enzyme substrate interaction as well as
feedback regulation. These differences in PLCs and the fact that
the plant PtdIns(4,5)P2-PLC is primarily membrane associated
led to the hypothesis that in plant membranes PtdIns(4,5)P2
hydrolysis would occur continuously, unless the lipid was bound
and protected by proteins such as profilin.
Metabolization
of the intermediate Ins(1,4,5)P3 in plants also differs
from animals. While in animal cells Ins(1,4,5)P3 is dephosphorylated
by a 5'phosphatase (ptase), in plants it can alternatively be dephosphorylated
by a 5'ptase or a 1'ptase.
Although
many aspects of the phosphoinositide pathway have been evolutionarily
conserved between plants, yeast and animals, plasma membrane signaling
appears to have evolved to the specification of the organism. For
example, plants lack the class I PtdIns 3-kinase that is involved
in signal transduction via phosphotyrosine-coupled plasma membrane
receptors in animal cells. Class I PtdIns 3- kinases phosphorylate
PtdIns 4P, PtdIns 5P, and PtdIns(4,5)P2 and produce PtdIns(3,4)P2,
PtdIns(3,5)P2 and PtdIns(3,4,5)P3. The plant
PtdIns 3-kinase only phosphorylates PtdIns and produces PtdIns(3)P.
PtdIns 3-kinase appears to be essential for plant growth and has
been reported to be involved in vacuolar trafficking and transcription
initiation; however, there is no evidence that it is directly involved
in plasma membrane signaling.
Cellular
functions of Phosphoinositides
The
inositol headgroup
Cellular
localization of phosphoinositides
Phosphoinositides
as central mediators of plant growth responses
Unique
properties of plant phosphoinositide metabolism
Inositol
1,4,5-trisphosphate regulates intracellular Ca2+ levels
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