Effect of pro-inflammatory stimuli on mucin expression
and inhibition by secretory leucoprotease inhibitor
Siobhan Griffin, Tomas P. Carroll,
Catherine M. Greene, Shane J. O’Neill,
Clifford C. Taggart* and Noel G. McElvaney
Respiratory Research Division, Department of Medicine,
Royal College of Surgeons in Ireland, Education and
Research Centre, Beaumont Hospital, Dublin 9, Ireland.
Summary
Stimuli-induced expression of certain mucin genes
has been demonstrated to occur as a result of ligand-
dependent activation of the epidermal growth factor
receptor (EGFR). In particular, MUC5AC expression
can be induced by cigarette-smoke, neutrophil
elastase and lipopolysaccharide (LPS) following acti-
vation of tumour necrosis factor a-converting
enzyme. We now show that a large of number of
stimuli relevant to the cystic fibrosis lung – neutrophil
elastase, LPS, Pam
3Cys-Ser-(Lys)4 Hydrochloride (a
lipopeptide analogue), CpG DNA (which mimics bac-
terial DNA) and cystic fibrosis bronchoalveolar lavage
fluid – can activate MUC1 and 2 expression as well as
MUC5AC expression in lung epithelial cells via an
EGFR-dependent mechanism. In addition, we demon-
strate that the immunomodulatory anti-protease,
secretory leucoprotease inhibitor, can inhibit stimuli-
induced MUC1, 2 and 5AC expression via a mecha-
nism that is primarily dependent on the inhibition of
transforming growth factor type alpha release. There-
fore, mucin gene expression, induced by cystic fibro-
sis respiratory stimuli, can be inhibited by secretory
leucoprotease inhibitor indicating its potential impor-
tance as an anti-mucin agent in cystic fibrosis and
other chronic lung diseases characterized by mucus
hypersecretion.
Introduction
Hypersecretion of mucus is a common pathophysiological
feature of cystic fibrosis (CF) and other inflammatory lung
diseases. Mucus obstruction of the airways in CF, asthma
and chronic bronchitis contributes significantly to mortality
and morbidity in these conditions. Mucus, which is com-
posed mostly of water and ions with a small percentage of
protein (5%), plays an important role in host defence
where it binds bacteria and removes it via the mucociliary
ladder to the upper airways and oesophagus where it is
either expectorated or ingested. To date nine MUC genes
have been described in the lung – MUC1, 2, 4, 5AC, 5B,
7, 8, 13 – and very recently, MUC19 (Voynow 2002; Chen
et al., 2004). MUC2, 5AC and 5B are secreted, gel-
forming mucins and MUC1, 4 and 13 are membrane asso-
ciated (Voynow 2002). Although MUC7 is secreted it does
not possess the cysteine-rich D domain (a feature of
MUC2, 5AC and 5B) and therefore does not oligomerize
or contribute to the gel phase of airway secretion (Bobek
et al., 1993). MUC5AC is the most prominent mucin of
normal airway secretions and its expression is increased
in nasal epithelium of individuals with CF and allergic
rhinitis (Voynow et al., 1998). In addition, MUC2 expres-
sion is elevated in CF nasal epithelium compared with
healthy controls (Li et al., 1997). MUC1 has recently been
shown to be an adhesion site for Pseudomonas aerugi-
nosa and binding of Pseudomonas to MUC1 results in
activation of an intracellular mitogen-activated protein
kinase pathway (Lillehoj et al., 2001; 2004).
In the context of airway inflammation, mucin gene
expression can be activated by a number of stimuli. Inter-
leukin (IL)-9 can regulate mucin expression via the human
calcium-activated chloride channel, hCLCA1 (Hauber
et al., 2003). Oxidative stress and activated neutrophils
can also increase MUC5AC expression via the epidermal
growth factor receptor (EGFR). Neutrophil elastase (NE)
has been shown to regulate MUC5AC expression both
through an oxidant mechanism and via a tumour necrosis
factor a-converting enzyme (TACE-EGFR pathway;
Fischer and Voynow, 2002; Kohri et al., 2002): other
stimuli have also been shown to regulate MUC5AC
expression via the TACE-EGFR pathway including ciga-
rette smoke, lipopolysaccharide (LPS) and supernatant
from Gram-negative bacteria (Dohrman et al., 1998; Shao
et al., 2003; 2004). Gram-positive bacteria-derived lipote-
ichoic acid (LTA) can activate MUC2 expression through
the platelet-activating factor receptor and a disintegrin
and metalloproteinase (ADAM)-10 resulting in the libera-
tion of heparin-binding epidermal growth factor (HB-EGF)
and activation of the EGFR (Lemjabbar and Basbaum,
2002). Although a number of these stimuli – LPS and LTA
– are ligands for the Toll-like receptor family of proteins,
Received 26 June, 2006; revised 23 August, 2006; accepted 28
August, 2006. *For correspondence. E-mail ctaggart@rcsi.ie; Tel.
(+353) 1 8093712; Fax (+353) 1 8093808.
Cellular Microbiology (2007) 9(3), 670–679 doi:10.1111/j.1462-5822.2006.00819.x
First published online 4 October 2006
© 2006 The Authors
Journal compilation © 2006 Blackwell Publishing Ltd