JOURNAL OF INTERFERON & CY'FOKINE RESEARCH 23:319-327 (2003)
© Mary Ann Liebert, Inc.
Differential Regulation of Cytokine Gene Expression by Avian
Heterophils During Receptor-Mediated Phagocytosis of
Opsonized and Nonopsonized Salmonella enteritidis
MICHAEL H. KOGUT,' LISA ROTHWELL. 2 and PETE KAISER2
ABSTRACT
Internalization of pathogens by phagocytic cells triggers the innate immune response, which in turn regulates
the acquired response. Phagocytes express a variety of receptors that are involved in recognition of patho-
gens, including (1) pattern recognition receptors (PRR), which recognize conserved motifs, (2) complement
receptors (CR), which recognize complement-opsonized pathogens, and (3) Fe receptors (FcR), which recog-
nize antibody-opsonized pathogens. Recognition of microbes is accompanied by the induction of multiple cell
processes, including the production of proinflammatory and anti-inflammatory cytokines and chemokines.
The objective of the present experiments was to use probes to known avian proinflammatory and anti-in-
flammatory cytokines and 'I'aqMan technology to ascertain levels of cytokine gene expression in avian het-
er()phils following receptor-mediated phagocytosis of either nonopsonized Salmonella enferitidis (SE), serum-
opsonized SE, or IgG-opsonized SE. Expression of interleukin-6 (IL-6) and IL-8, considered in mammals as
a proinflammatory chemokine, were upregulated following exposure to the nonopsonized or the opsonized
SE. However, mRNA expression for IL-18 and interferon-'y (IFN-y) was downregulated, and the expression
of mRNA for the anti-inflammatory cytokine transforming growth factor-134 ('FGF- 1 64). was upregulated. In-
terestingly, IL-
1
1 3
mRNA expression was significantly upregulated in heterophils that phagocytized either the
nonopsonized SE via PRRs or IgG-opsonized SE via FcRs, whereas serum-opsonized SE phagocytized by CRs
induced a downregulation of IL-113 mRNA. These results suggest that signaling interactions initiated by re-
ceptor recognition of the microbe surface differentially regulate the induction of inflammatory cytokines in
avian heterophils.
It
-
IJ.
INTRODUCTION
M
1CROI3IAL INVASION OF THE Hosr is followed by a series
of events designed to control (recognition. phagocytosis,
microhicidal activities) and eventually resolve (inflammation
and acquired immunity) the infection.° The immediate re-
.sponse to invasive pathogens, clearance via the inflammatory
response, and activation of the appropriate acquired responses
are all coordinated and orchestrated by the innate host de-
fenses. )
Recognition of potential pathogenic microbes by the innate
immune system is the function of a restricted class of cellular
receptors known as the nonclonal pattern recognition receptors
(PRRs), such as the toll-like receptors (TLRs). The innate sys-
tem uses these germ-line encoded receptors to recognize evo-
lutionarily conserved molecular motifs (pathogen-associated
'USDA-ARS, SPARC. College Station, TX 77845,
Institute for Animal Health. Compton. Berkshire RG20 7N. U.K.
molecular patterns {PAMP5]) of infectious microbes. 23 The
phagocytic process is triggered by two distinct cell surface re-
ceptors, one involving the recognition of the carhoxy-terminus
of immunoglohulin (Ig) molecules by the Fe receptor (FcR) and
the other where complement receptors (CR) recognize com-
plement fragments
.(1.5
Likewise, some PRRs, such as mannose
receptors and scavenger receptors, have been shown to be in-
volved in the phagocytosis process as well. 5 The end prod-
uct of microbial recognition and phagocytosis by cells of the
innate response is the activation of intracellular signaling path-
ways that initiate cellular processes, such as activation of mi-
crobicidal killing mechanisms, the production of proinfiamma-
tory and anti-inflammatory cytokines. and the production of
costimulatory molecules required for antigen presentation to the
acquired immune system.9' )
As the first cells to migrate to a site of infection, polymor-
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