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Type III CaM did not induce NAD kinase at any Ca 2+ level (Karita et al. 2004 ). Ca 2+
spike frequency optimizes gene expression (Li et al. 1998 ). Calcium oscillations
increase the effi ciency and specifi city of gene expression (Dolmetsch et al. 1998 ).
A combination of changes in all Ca 2+ parameters produced by a particular signal
determines Ca 2+ signature (Luan et al. 2002 ).
4.12
Ca 2+ Sensors in Ca 2+ Signal Transduction
The calcium signature is perceived by different Ca 2+ -binding proteins (Kudla et al.
2010 ). These intracellular Ca 2+ -binding proteins are also known as Ca 2+ sensors.
The changes in [Ca 2+ ] cyt concentrations are monitored by the Ca 2+ sensors and the
Ca 2+ signals are subsequently decoded and propagated downstream to activate plant
defense responses. Ca 2+ signaling pathways are composed of molecular relays; the
fi rst runner after Ca 2+ is Ca 2+ “sensor”, which monitors temporal and spatial changes
in Ca 2+ concentrations. Several Ca 2+ sensors have been identifi ed in plants (Fig. 4.4 ).
Calmodulin
(CaM)
Calmodulin-like
proteins
Calcineurin B-
like (CBL)
proteins
Ca 2+ -dependent
and calmodulin-
independent
protein kinases
(CDPKs)
Calcium
sensors
Ca 2+ -CaM-
dependent kinases
(CCaMK)
NADPH oxidase
Copines
Phospholipase D
Fig. 4.4
Calcium-binding proteins as calcium sensors
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