A Calmodulin Binding Protein from Arabidopsis
Is Induced by Ethylene and Contains
a DNA-Binding Motif
A. S. N. Reddy,
1
Vaka S. Reddy, and Maxim Golovkin
Department of Biology and Program in Cell and Molecular Biology, Colorado State University,
Fort Collins, Colorado 80523
Received October 23, 2000
Calmodulin (CaM), a key calcium sensor in all eu-
karyotes, regulates diverse cellular processes by inter-
acting with other proteins. To isolate CaM binding
proteins involved in ethylene signal transduction, we
screened an expression library prepared from
ethylene-treated Arabidopsis seedlings with
35
S-
labeled CaM. A cDNA clone, EICBP (Ethylene-Induced
CaM Binding Protein), encoding a protein that inter-
acts with activated CaM was isolated in this screening.
The CaM binding domain in EICBP was mapped to the
C-terminus of the protein. These results indicate that
calcium, through CaM, could regulate the activity of
EICBP. The EICBP is expressed in different tissues
and its expression in seedlings is induced by ethylene.
The EICBP contains, in addition to a CaM binding
domain, several features that are typical of transcrip-
tion factors. These include a DNA-binding domain at
the N terminus, an acidic region at the C terminus, and
nuclear localization signals. In database searches a
partial cDNA (CG-1) encoding a DNA-binding motif
from parsley and an ethylene up-regulated partial
cDNA from tomato (ER66) showed significant similar-
ity to EICBP. In addition, five hypothetical proteins in
the Arabidopsis genome also showed a very high se-
quence similarity with EICBP, indicating that there
are several EICBP-related proteins in Arabidopsis.
The structural features of EICBP are conserved in all
EICBP-related proteins in Arabidopsis, suggesting
that they may constitute a new family of DNA binding
proteins and are likely to be involved in modulating
gene expression in the presence of ethylene. © 2000
Academic Press
Key Words: calmodulin binding protein; ethylene;
transcription factor; ankyrin repeat; DNA-binding do-
main; Arabidopsis.
Calcium (Ca
2+
) is an important intracellular messen-
ger in plants. In recent years, a large number of hor-
monal signals, and biotic and abiotic stresses have
been shown to elevate cytoplasmic Ca
2+
([Ca
2+
]
cyt
) lev-
els transiently (1–5). Four major families of proteins
that sense changes in [Ca
2+
]
cyt
have been identified in
plants (5, 6). These include (i) calmodulin (CaM) and
its isoforms, (ii) CaM-like proteins, (iii) Ca
2+
-
dependent and CaM-independent protein kinases (CD-
PKs) that are found only in plants and some protozo-
ans, and (iv) Ca
2+
-binding proteins without EF-hand
motifs (5– 8). Of these Ca
2+
sensors, CaM is highly
conserved among all eukaryotes and contains four Ca
2+
binding EF-hand motifs (1, 6, 7). Activated CaM (Ca
2+
/
CaM complex) interacts with a variety of unrelated
proteins (CaM binding proteins, CBPs) and regulates
their activity/function (5–7). To understand Ca
2+
me-
diated signaling pathways in plants it is essential to
isolate and identify CaM binding proteins. Screening of
expression libraries with labeled CaM has resulted in
isolation of a limited number of CBPs in plants (5,
9 –11). Interestingly, several plant CBPs have no ho-
mologs in nonplant systems (5).
Ethylene, a two-carbon gaseous plant hormone, reg-
ulates diverse physiological processes during plant
growth and development (12). Seed germination, seed-
ling growth, fruit ripening, senescence, plant defense
responses, abscission and sex determination are some
of the processes that are regulated by ethylene (12, 13).
Using genetic and molecular approaches significant
advances have been made in elucidating the ethylene
signal transduction pathway (13–16). Isolation of mu-
tants that are insensitive to ethylene or show ethylene
response in the absence of ethylene and cloning of
corresponding genes from the model plant Arabidopsis
have resulted in identification of many components of
the ethylene signal transduction pathway (13–19). In
addition to characterization of ETR/ERS and CTR
gene products in the ethylene signal transduction
1
To whom correspondence and reprint requests should be ad-
dressed. Fax: 970-491-0649. E-mail: reddy@lamar.colostate.edu.
Biochemical and Biophysical Research Communications 279, 762–769 (2000)
doi:10.1006/bbrc.2000.4032, available online at http://www.idealibrary.com on
762 0006-291X/00 $35.00
Copyright © 2000 by Academic Press
All rights of reproduction in any form reserved.