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Finkler et al. ( 2007 ) found that the Ca 2+ /CaM-responsive CAMTAs bind to the
ABA-responsive ABRE cis elements in Arabidopsis , suggesting a link between
Ca 2+ -responsive transcription factors and ABA-responsive cis -elements. CAMTA3
directly interacts with the promoter of the EDS1 gene, a regulator of salicylic acid
levels, and represses its expression (Du et al. 2009 ). Ca 2+ /calmodulin binding to
CAMTA3 is required for the suppression of plant defense, indicating a direct role of
Ca 2+ /calmodulin in regulating the function of CAMTA3 (Du et al. 2009 ).
4.14.11
Calmodulin-Binding OsCBT, NtER1, and AtSR1
Transcription Factors
Some transcription factors of the basic helix-loop-helix family were shown to bind
calmodulin (Onions et al. 2000 ). A CaM-binding transcription factor, OsCBT
( O ryza s ativa CaM-binding transcription factor), has been isolated from rice (Choi
et al. 2005 ). It contains a CG-1 homology DNA binding domain, three ankyrin
repeats, a putative transcriptional activation domain, and fi ve putative CaM-binding
motifs (Choi et al. 2005 ). OsCBT has two different types of functional CaM-binding
domains, an IQ motif, and a Ca 2+ -dependent motif (Choi et al. 2005 ).
Some ethylene-responsive genes-encoded proteins in tobacco, tomato, rapeseed,
Arabidopsis have structural similarity to the rice CaM-binding transcription factor,
OsCBT (Choi et al. 2005 ). The tobacco early ethylene-responsive gene NtER1 encodes
a CaM-binding protein (Yang and Poovaiah 2000 ). One NtER1 homolog ( AtSR1 ) and
fi ve related genes ( AtSR2-6 ) have been identifi ed in Arabidopsis and they are desig-
nated as AtSR (for A rabidopsis t haliana S ignal R esponsive ) genes (Yang and Poovaiah
2002a ). These six genes exhibit rapid and differential response to signal elicitors such
as ethylene, jasmonate (JA), salicylic acid, ABA, and H 2 O 2 . Ca 2+ /CaM binds to a
23-mer peptide in all AtSRs that corresponds to the CaM-binding region of NtER1.
Each AtSR has a conserved structural feature with a DNA-binding domain in the N
terminus and a CaM-binding domain in the C terminus. AtSR1 targets the nucleus and
specifi cally recognizes a novel 6-bp CGCG box (A/C/G) CGCG (G/T/C). The multi-
ple CGCG cis -elements are found in promoters of various genes involved in ET- and
ABA signaling systems. The results suggest that AtSR may interact with ET and ABA
signaling systems (Yang and Poovaiah 2002a ).
4.14.12
Calmodulin-Binding WRKY Transcription Factors
WRKY proteins constitute a large class of transcription factors. More than 70
WRKY genes have been reported in Arabidopsis (Eulgem et al. 2000 ; Robatzek and
Somssich 2001 ; Dong et al. 2003 ; Bhattarai et al. 2010 ). About 109 WRKY genes
have been identifi ed in rice (Qu and Zhu 2006 ; Liu et al. 2005 , 2007 ; Qiu et al. 2007 ).
The WRKY transcription factor family is defi ned by a domain of 60 amino acids,
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