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B p65 Mediates the Assembly and Activation of the TNF-Responsive Element of the Murine Monocyte Chemoattractant-1 Gene1
Department of Microbiology and Immunology, Emory University School of Medicine, Emory University, Atlanta, GA 30322
TNF-
transcriptionally regulates murine monocyte chemoattractant
protein-1 (MCP-1) expression. Three approaches were used to determine
the mechanism by which TNF regulates MCP-1. Mutation analysis showed
that two distal 
B sites, a novel dimethylsulfate-hypersensitive
sequence, and a promoter proximal SP-1 site were required for TNF
induction. Although the B sites and the hypersensitive sequence
function as a NF-B-mediated enhancer, regulating induction by TNF,
stereospecific alignment of the B sites was not critical.
Trans-activation studies conducted by cotransfection of
p50 and/or p65 expression vectors with MCP-1 constructions showed that
TNF regulates MCP-1 through NF-B. Examination of MCP-1 induction in
NF-B-disrupted embryonic fibroblasts showed that p65 was necessary
for both the induction and the TNF-induced protein occupancy of the
enhancer in vivo. The action of the antioxidant inhibitor of NF-B
activation, pyrrolidine dithiocarbamate, in wild-type and NF-B
mutant cells was examined. The results suggested that TNF activates
NF-B through both pyrrolidine dithiocarbamate-sensitive and
-insensitive mechanisms. This study illustrates the crucial role for
NF-B p65 in the induction of the MCP-1 gene by TNF and in the
assembly of a NF-B dependent enhancer in vivo.
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