Human adult T cell leukaemia-derived factor
(ADF) was first described as an IL-2 receptor α chain
(IL-2R α, Tac) upregulating factor produced by cell
lines from patients with adult T-cell leukemia.
1
The
purification and cloning of human ADF showed that it
represents a mammalian homologue of bacterial thiore-
doxin (TRX). ADF/human TRX has an active thiol-
group (-Cys-Gly-Pro-Cys-) and acts as an endogenous
thiol-related reducing protein.
2,3
TRX functions as
hydrogen donor mediating the conversion of ribo-
nucleotides to deoxyribonucleotides, the degradation
of insulin, and the stabilization of glucocorticoid re-
ceptors. Furthermore, thioredoxin is involved in an
electron-transfer system common to a variety of cells
and scavanges free radicals.
4
Despite its intracellular function as disulfide reduc-
ing enzyme and its lack of a signal sequence the protein
has been found to exert several effects extracellularily.
5
It has been shown that TRX/ADF is actively secreted
by a variety of normal and transformed human cells,
including fibroblasts, airway epithelial cells, and acti-
vated B and T cells. The secretory mechanism for ADF
shares several features with but is not identical to the
alternative pathway described for IL-1β.
6
For human ADF it has been demonstrated that sev-
eral cytokine-like functions are related to this protein.
For example, an autocrine growth factor produced
by the EBV-transformed B-cell line 3B6
7
named
3B6-IL-1, and MP6-BCGF, a B-cell growth factor de-
rived from the T-cell hybridoma MP6,
8
were shown to
be identical to ADF. A factor enhancing cytotoxicity of
eosinophils (eosinophil cytotoxicity-enhancing factor,
ECEF), which is produced by activated U937 cells, is
closely related or identical to ADF.
9
In line with this,
purified recombinant ADF enhanced cytotoxicity and
migration of human eosinophils induced by C5a.
10
In
addition, recombinant ADF has been described to
inhibit the cytotoxicity mediated by TNF and anti-Fas
antibodies on U937 cells.
11
In contrast to the large array of studies analysing
effects of human ADF, very little is known about the
expression and regulating acivities of the murine homo-
logue. We therefore expressed this molecule in E. coli,
established a purification protocol and analysed the effect
of murine ADF on the proliferation of murine T cells.
6 CYTOKINE, Vol. 8, No. 1 (January), 1996: pp 6–13
EXPRESSION AND CO-CYTOKINE FUNCTION
OF MURINE THIOREDOXIN/ADULT T CELL
LEUKAEMIA-DERIVED FACTOR (ADF)
Horst Blum, Martin Röllinghoff, André Gessner
Human ADF (adult T cell leukaemia-derived factor), an isoform of thioredoxin, promotes
proliferation of certain human lymphoid cell lines and is involved in many thiol-dependent
reducing reactions. To study functional aspects of the murine homologue, we established
inducible overexpression of murine ADF in E. coli and a purification method which led to
an apparently homogeneous 14 kDa protein. This recombinant ADF was tested in prolifer-
ation assays with murine Th2 cells (D 10.G4.1) and CTLL-2 cells. In synergy with IL-2, IL-4,
IL-7 and IL-9 A D F displayed co-cytokine activity. These proliferative effects were neutral-
ized by an affinity-purified polyclonal rabbit anti-A D F antiserum. The effects of A D F were
critically dependent on the presence of 2-mercaptoethanol. Bacterial thioredoxin had simi-
lar effects on the proliferation of murine T cells. Thus, the thiol-related reducing capacity of
these proteins is essential for their growth promoting activity. A s investigated at the levels of
mRNA and protein in several murine cell clones and lines as well as in mouse tissues A D F is
expressed ubiquitously. Finally it could be demonstrated by competitive PCR that in contrast
to cytokine mRNAs (e.g. IL-4 and IL-13) the expression of ADF mRNA in murine Th2
clones and spleen cells is not influenced by stimulation of these cells through the T cell recep-
tor complex. Murine ADF therefore represents a protein constitutively expressed in a wide
variety of cells with the capacity to enhance the proliferative effect of several cytokines on
murine T cells.
© 1996 A cademic Press Limited
From the Institut für Klinische Mikrobiologie und Immunologie der
Universität Erlangen-Nürnberg, Wasserturmstr. 3, D-91054
Erlangen, Germany
Correspondence to: André Gessner
Received 3 April 1995; accepted for publication 18 August 1995
© 1996 Academic Press Limited
1043-4666/96/010006+8 $12.00/0
KEY WORDS: Thioredoxin/Adult T cell leukemia-derived
factor/co-cytokine function/IL-2/IL-4/IL-7/IL-9