Protoplasma (2003) 221: 117–127
DOI 10.1007/s00709-002-0059-y
Summary. Evidence is rapidly accumulating that low-activity
NAD(P)H oxidases homologous to that in phagocytic cells gener-
ate reactive oxygen species as signaling intermediates. In this review
we discuss evidence that signaling NAD(P)H oxidases in part
influence normal and malignant cell division by activating the
redox-regulated transcription factor nuclear factor kB. The roles of
growth-regulatory NAD(P)H oxidases in human airway smooth
muscle and malignant melanoma are used as examples.
Keywords: NADPH oxidase; Nuclear factor kB; NAD(P)H oxidase
subunit p22
phox
; NAD(P)H oxidase subunit gp91
phox
; Superoxide
anion; Melanoma.
Introduction
Reactive oxygen species (ROS) were once viewed
only as vehicles of chemical cellular destruction but
are now also appreciated to be elegant, carefully con-
trolled cellular switches for signaling cascades impor-
tant in cell growth, cell death, and inflammation. One
potentially important source for signaling ROS is the
NOX family of NAD(P)H oxidases similar but distinct
from the membrane NADPH oxidase used by phago-
cytes for host defense.These NOX oxidases have been
cloned for homologs (NOX1, NOX3, NOX4, and
NOX5, with NOX2 corresponding to gp91
phox
) and
characterized by Lambeth and co-workers (Cheng
et al. 2001). We will concentrate only on the impact of
NOX oxidases in cellular signaling.
Most investigative efforts to link NOX oxidases to
cell signaling events have occurred in the area of vas-
cular diseases (Griendling et al. 2000), where the sig-
naling NAD(P)H oxidase has been rapidly and
carefully explored in endothelium (Gorlach et al.
2000) and vascular smooth muscle (Lassegue et al.
2001). However, important functions for these
NAD(P)H oxidases are rapidly emerging in other
areas. A constitutively active NAD(P)H oxidase has
been recently shown responsible for increased ROS
production and the hyperproliferative state of skin
fibroblasts cultured from patients with systemic scle-
rosis (Sambo et al. 2001). Thus, NOX oxidases are
likely to be important in mediating events in a number
of diseases.
Components of NAD(P)H oxidase signaling
While a signaling role for these NAD(P)H oxidases is
now generally accepted, it is still unclear which signal-
ing cascades are responsible for mediating NAD(P)H
oxidase effects. Signaling is best worked out in vascu-
lar smooth muscle, where the NAD(P)H oxidase is
thought to mediate many of the effects of angiotensin
II on vascular remodeling (Griendling et al. 2000,
Irani 2001). In cultured rat vascular smooth muscle
cells, expression of adenovirus-linked antisense oligo-
nucleotides to NOX1 significantly decreases activation
of p38 mitogen-activated protein kinase (p38 MAP
kinase) and the phosphatidylinositol 3 kinase(PI 3-
kinase)-activated Akt/protein kinase B (Akt) stimu-
lated by angiotensin II or platelet-derived growth
PROTOPLASMA
Printed in Austria
Redox signaling of NF-kB by membrane NAD(P)H oxidases in normal
and malignant cells
S. S. Brar
1
, T. P. Kennedy
1,
*, M. Quinn
2
, and J. R. Hoidal
3
1
Department of Internal Medicine, Carolinas Medical Center, Charlotte, North Carolina
2
Department of Veterinary Molecular Biology, Montana State University, Bozeman, Montana
3
Department of Medicine, University of Utah, Salt Lake City, Utah
Received May 4, 2002; accepted July 26, 2002; published online May 21, 2003
© Springer-Verlag 2003
* Correspondence and reprints: Department of Internal Medicine,
Carolinas Medical Center, P.O. Box 32861, Charlotte, NC 28232,
U.S.A.