Development 108, 173-183 (1990)
Printed in Great Britain © T h e Company of Biologists Limited 1990
173
The biological effects of XTC-MIF: quantitative comparison with Xenopus
bFGF
J. B. A. GREEN, G. HOWES, K. SYMES*, J. COOKE and J. C. SMITH
Laboratory of Embryogenesis, National Institute for Medical Research, The Ridgeway, Mill Hill, London NW71AA, UK
* Present address: Department of Cell and Molecular Biology, 385 LSA, University of California, Berkeley, CA 94720, USA
Summary
Mesoderm in Xenopus and other amphibian embryos is
induced by signals from the vegetal hemisphere acting
on equatorial or animal hemisphere cells. These signals
are diffusible and two classes of candidate signal mol-
ecule have been identified: the fibroblast growth factor
(FGF) and transforming growth factor /? (TGF-/?) types.
In this paper, we compare the effects of cloned Xenopus
basic FGF (A1>FGF) and electophoretically homo-
geneous XTC-MIF (a TGF-0-like factor obtained from a
Xenopus cell line) on animal pole explants. We find that
they have a similar minimum active concentration
(0.1-0.2ngml"') but that, nonetheless, XTC-MIF is at
least 40 times more active in inducing muscle. In
general, we find that the two factors cause inductions of
significantly different characters in terms of tissue type,
morphology, gene expression and timing.
At low concentrations (0.1-1.0 ng ml"
1
) both factors
induce the differentiation of 'mesenchyme' and 'meso-
thelium' as well as blood-like cells. These latter cells do
not, however, react with an antibody to Xenopus globin.
This raised the possibility that the identification of red
blood cells in other studies on mesoderm induction might
have been mistaken, but combinations of animal pole
regions with ventral vegetal pole regions confirmed that
genuine erythrocytes are formed. The identity of the
blood-like cells formed in response to the inducing
factors remains unknown. At higher concentrations
XTC-MIF induces neural tissue, notochord, pronephros
and substantial and often segmented muscle. By con-
trast, -YbFGF only induces significant amounts of muscle
above 24ngml~
1
and even then this is much less than
that induced by XTC-MIF.
For both factors an exposure of less than 30min is
effective. Competence of animal pole cells to respond to
A1)FGF is completely lost by the beginning of gastru-
lation (stage 10) while competence to XTC-MIF is
detectable until somewhat later (stage 11). Since animal
pole tissue is known to be able to respond to the natural
inducer at least until stage 10, and perhaps until stage
10.5, this suggests that bFGF cannot be the sole inducer
of mesoderm in vivo.
Taken together, these results are consistent with XTC-
MIF being a dorsoanterior inducer and AT)FGF a
ventroposterior inducer, suggesting that body pattern is
established by the interaction of two types of inducing
signal. This model is discussed in view of the qualitative
and quantitative differences between the factors.
Key words: mesoderm induction, mesoderm-inducing
factors, bFGF, XTC-MIF, amphibian embryo, Xenopus
laevis.
Introduction
The mesoderm of amphibian embryos arises through an
inductive interaction during which a signal from the
vegetal hemisphere of the embryo acts on overlying
animal pole cells (reviewed by Dawid et al. 1989;
Gurdon et al. 1989; Smith, 1989). Recently, two meso-
derm-inducing factors (MIFs) have been derived from
homologous sources. XTC-MIF is produced by the
Xenopus XTC cell line and may be related to TGF-/J
(Smith, 1987; Rosa et al. 1988; Smith et al. 1988). Basic
fibroblast growth factor (bFGF) has been cloned from a
Xenopus oocyte cDNA library and the protein is known
to be present in the egg and early embryo (Slack et al.
1987; Kimelman and Kirschner, 1987; Kimelman et al.
1988; Slack and Isaacs, 1989). Both XTC-MIF and
bFGF induce mesodermal cell types from Xenopus
blastula ectoderm but until now most work has been
done only with partially purified XTC-MIF and with
bFGF from bovine brain. The limited information that
is available suggests that XTC-MIF induces dorsal,
intermediate and ventral mesodermal cell types while
bFGF induces only intermediate and ventral tissue (see
Smith, 1987; Godsave et al. 1988; Smith et al. 1988).
In this paper, we undertake a direct quantitative
comparison of the effects of the two Xenopus-dtrived
MIFs in their pure forms. While they display a similar
potency for mesoderm induction, the two factors are