REVIEW ARTICLE
Mod Rheumatol (2005) 15:383–390 © Japan College of Rheumatology and Springer-Verlag Tokyo 2005
DOI 10.1007/s10165-005-0430-x
Gerhard E. Grossmayer · Luis E. Munoz · Udo S. Gaipl
Sandra Franz · Ahmed Sheriff · Reinhard E. Voll
Joachim R. Kalden · Martin Herrmann
Removal of dying cells and systemic lupus erythematosus
Received: July 29, 2005 / Accepted: August 23, 2005
Abstract Systemic lupus erythematosus (SLE) is a very
heterogeneous systemic autoimmune disease, in which
autoantibody synthesis against nuclear constituents is the
main immunological characteristic. These autoantibodies
underwent affinity maturation and isotype switching. Addi-
tionally, T-cell tolerance against nuclear autoantigens
should be affected in these autoimmune patients. Nuclear
material derived from apoptotic and/or necrotic cells may
serve as an important source of autoantigens. However,
dead and dying cells as well as cellular debris are rapidly
removed from tissues by phagocytes without eliciting in-
flammation or immune responses under healthy conditions.
During apoptosis nuclear components are strongly modified
through enzymatic reactions. If these cells are not timely
cleared, those autoantigens may be released, taken up, and
presented by dendritic cells in tissues or presented by
follicular dendritic cells in lymph nodes to T and B cells,
respectively. This could be a mechanism for breaking the
peripheral self-tolerance. In this article we focus on the
deficient clearance of apoptotic cells in SLE patients and its
importance in development of this autoimmune disease.
Key words Apoptosis · Autoimmunity · Clearance · Necro-
sis · Systemic lupus erythematosus (SLE)
G.E. Grossmayer · L.E. Munoz · U.S. Gaipl · S. Franz · A. Sheriff ·
J.R. Kalden · M. Herrmann (*)
Institute for Clinical Immunology, Department of Medicine 3,
Friedrich-Alexander University of Erlangen-Nuremberg,
Glückstrasse 4a, 91054 Erlangen, Germany
Tel. +49-9131-85-36345; Fax +49-9131-85-35776
e-mail: martin.herrmann@med3.imed.uni-erlangen.de
R.E. Voll
IZKF Research Group N2, Nikolaus-Fiebiger Center of Molecular
Medicine, Erlangen, Germany
G.E.G. and L.E.M. contributed equally to this work.
Introduction
The immune mechanisms of the pathogenesis of systemic
lupus erythematosus (SLE) are still not fully understood.
This disease shows a wide array of clinical manifestations
resulting from inflammatory reactions in multiple organs.
Immunological abnormalities observed in the autoimmune
disease SLE are characterized by autoantibody synthesis
against nuclear constituents, immune complex deposits in
tissues, and complement activation. Antinuclear autoanti-
bodies are not exclusively generated in SLE, but also in
other autoimmune diseases like scleroderma, rheumatoid
arthritis, and Sjögren’s syndrome. Several genetic and envi-
ronmental factors as well as infections have been implicated
as important elements of the disease (reviewed in
Herrmann et al.
1
and Kuenkele et al.
2
). How intracellular
proteins become targets of immune responses is important
in understanding the etiology of autoimmune diseases.
Anti-double-stranded DNA antibodies (anti-dsDNA)
are one of the most important criteria for the diagnosis of
the disease. They show high-affinity binding activity in con-
trast to low-affinity naturally occurring anti-dsDNA anti-
bodies from normal healthy donors (NHD), which did not
go through the isotype switch from IgM to IgG or IgA.
Affinity maturation of immunoglobulins for the antigen and
isotype switching are processes dependent on T-cell help.
This takes place in the germinal centers (GCs) of lymphoid
tissues. The role of T cells in the development of chronic
autoimmunity in SLE is supported by the fact that patients
with SLE respond to T-cell specific immunosuppressive
drugs like cyclosporin A.
3
Although human DNA is known to be poorly immuno-
genic and does not bear T-cell epitopes, DNA–histone com-
plexes and nucleosomes are released in SLE patients in high
amounts into the circulation.
4,5
The nucleosomes are targets
of anti-dsDNA autoantibodies and of anti-histone autoanti-
bodies, as shown by enzyme-linked immunosorbent assays
with autoimmune sera and monoclonal antibodies.
6,7
Those
autoantibodies are predominantly of the IgG isotype and
molecular analyses revealed a high degree of somatic muta-