2. Second Messenger : Calcium ions
ā¢ Calcium (Ca2+) is an important cellular second messenger for diverse
developmental processes and environmental responses.
ā¢ cytosolic Ca2+ is the only second messenger to be involved in diverse pathways and
respond to numerous stimuli.
ā¢ Various environmental and developmental processes induces transient
increases in cytosolic Ca2+ {the cellular calcium levels, termed as calcium
signatures},
ā¢ developmental processes including root growth, stomatal movement, pollen growth,
ā¢ environmental cues including light, abiotic stress responses, and plant-microbe
interactions.
ā¢ The concentration of Ca2+ is delicately balanced by the presence of
āCa2+ storesā like vacuoles, endoplasmic reticulum, mitochondria and cell
wall.
3. Why is Ca2+ are excluded from the cytosol?
ā¢ One reason is that Ca2+ binds water much less tightly and precipitates
phosphate.
ā¢ Hence, cells have evolved ways to isolate this dangerous divalent,
ā¢ to simply reduce its cytosolic levels
ā¢ to use its binding energy for signal transduction.
ā¢ Unlike complex molecules, Ca2+ cannot be chemically altered. Thus, to
exert control over Ca2+, cells must chelate, compartmentalize, or extrude it.
ā¢ Hundreds of cellular proteins have been adapted to bind Ca2+
ā¢ in some cases simply to buffer or lower Ca2+ levels, and
ā¢ in others to trigger cellular processes.
ā¢ The local nature of Ca2+ signaling is intimately tied to this large range of
affinities.
4. Mechanisms of intracellular calcium ion
concentration changes
ā¢ The mechanisms of intracellular concentration changes include
ā¢ calcium-induced calcium release and
ā¢ calcium dependent inactivation of calcium release.
ā¢ The removal of Ca2+ from the cytosol against its electrochemical
gradient to either the apoplast or to intracellular organelles requires
energized āactiveā transport.
ā¢ Ca2+-ATPases and
ā¢ H+/Ca2+ antiporters
are the key proteins catalyzing this movement.
5. Calcium signatures are generated by the coordinated action of various
Ca2+ influx and efflux systems including channels, pumps, and exchangers
located at different cellular membranes.
ā¢ The cytoplasmic Ca2+ level is low in
resting cells.
ā¢ Low cytoplasmic [Ca2+] is maintained by
extrusion via
ā¢ plasma membrane Ca2+ ATPase (PMCA) and
ā¢ smooth endoplasmic reticular Ca2+ ATPase
(SERCA) transporters.
ā¢ The Na/Ca exchanger (NCX), a major
secondary regulator of [Ca2+], is
electrogenic,
ā¢ exchanging three Na ions for one Ca2+.
ā¢ Intracellular Ca2+ hyperpolarizes many
cells by activating K+ channels, and in
some cells, Clā channels.
ā¢ This decreases CaV channel activity but
increases the driving force across active
Ca2+-permeant channels.
6. The Ca2+ signaling network: An Overview
ā¢ In excitatory Ca2+ signaling, plasma membrane
ion channels are triggered to open by changes
in voltage, or extra- or intracellular ligand
binding.
ā¢ Initial increases in [Ca2+] trigger more release,
primarily from ER via Ca2+-sensitive ryanodine
receptors (RyR).
ā¢ G protein-coupled receptor (GPCR) or receptor
tyrosine kinase-mediated activation of PLC
cleaves PIP2 into inositol (1,4,5) trisphosphate
(IP3) and diacylglycerol (DAG).
ā¢ IP3 is a ligand for the intracellular IP3R channel
spanning the membrane of the ER.
ā¢ GPCRs catalyze the exchange of guanosine
diphosphate (GDP) for GTP on GĪ± subunits,
releasing active GĪ± and GĪ²Ī³ subunits that in
turn activate PLCĪ².
ā¢ RTKs dimerize upon ligand binding, auto-
phosphorylate, and interact with other signaling
proteins to activate PLCĪ³.