INTRODUCTION
In normal adults, myelopoiesis occurs in the bone marrow,
where cells of the medullar stroma, locally produced or
extramedullar cytokines, and extracellular matrix, form the
haematopoietic bone marrow microenvironment. The inter-
action of both stromal and haematopoietic cells with
cytokines and the associated extracellular matrix molecules
controls the progressive commitment and differentiation of
myeloid stem cells into mature blood cells. These controls
involve both the maintenance of blood cell homeostasis and
the capacity to increase production of one or several cell
lines in a coordinated and strictly controlled manner. The
controls of myeloid cell production and differentiation are
thus necessarily complex and tightly regulated.
In contrast to this tight regulation of the bone marrow
haematopoietic environment stands the fact that partial or
full haematopoiesis can be observed in extramedullar loci,
like spleen or liver, in many pathological situations in adult
life. This is frequent in diseases where the bone marrow
microenvironment is either modified, as in myelofibrosis,
or occupied by new cell populations, as in neoplasias
involving bones. This phenomenon can be understood as a
passive dislodging of haematopoiesis into sites that have
had the haematopoietic function in earlier embryonic devel-
opment: it thus represents a case of myeloid metaplasia. On
the other hand, in chronic tissue inflammatory reactions, a
local myeloid proliferation may be observed. In schistoso-
miasis, both in humans and in experimental models, pro-
liferation of mono-macrophagic, neutrophil, eosinophil and
megakaryocytic cell lineages, alone or in combination, have
been described, in association with diffuse or granuloma-
tous inflammatory reactions to parasites (Borojevic and
Carvalho, 1981; Borojevic et al., 1981, 1983, 1989a,b;
Clark et al., 1988; El-Cheikh et al., 1991). Since in schis-
tosomiasis the general bone marrow function was not ham-
pered, this phenomenon has to be understood as an active
displacement of myelopoiesis outside the normal myelopoi-
477
Journal of Cell Science 104, 477-484 (1993)
Printed in Great Britain © The Company of Biologists Limited 1993
In chronic murine schistosomiasis, extramedullar
myelopoiesis was observed, with proliferation of myeloid
cells in liver parenchyma and in periovular granulomas.
We have studied the question of whether cells obtained
from granulomatous connective tissue may act as
myelopoietic stroma, supporting long-term myeloid pro-
liferation. Primary cell lines (GR) were obtained in vitro
from periovular granulomas, induced in mouse livers by
Schistosoma mansoni infection. These cells were char-
acterized as myofibroblasts, and represent liver con-
nective tissue cells involved in fibro-granulomatous
reactions. They were able to sustain survival and pro-
liferation of the multipotent myeloid cell lines FDC-P1
and DA-1 (dependent on interleukin-3 and/or granulo-
cyte-macrophage colony stimulating factor, GM-CSF)
without the addition of exogenous growth factors. This
stimulation was dependent upon myeloid cell attach-
ment to the GR cell layer; GR cell-conditioned medium
had no activity. Primary murine skin fibroblasts could
not sustain myelopoiesis. The endogenous growth-factor
was identified as GM-CSF by neutralization assays with
monoclonal antibodies. The stimulation of myelopoiesis
occurred also when GR cells had been fixed with glu-
tardialdehyde. The observed stimulatory activity was
dependent upon heparan sulphate proteoglycans
(HSPGs) associated with GR cell membranes. It could
be dislodged from the cell layer with heparin or a high
salt buffer. Our results indicate a molecular interaction
between endogenous growth-factor and HSPGs; this
interaction may be responsible for the stabilization and
presentation of growth factors in myelopoietic stromas,
mediating extramedullar proliferation of myeloid cells
in periovular granulomas.
Key words: proteoglycan, myelopoiesis, granuloma,
schistosomiasis
SUMMARY
Cell membrane-associated proteoglycans mediate extramedullar myeloid
proliferation in granulomatous inflammatory reactions to schistosome
eggs
Márcio Alvarez-Silva, Luiz-Cláudio F. da Silva and Radovan Borojevic
*
Departamento de Bioquímica, Instituto de Química, Universidade Federal do Rio de Janeiro, Caixa Postal 68021, 21944 Rio
de Janeiro, Brazil
*
Author for correspondence