Development 105, 147-153 (1989)
Printed in Great Britain © The Company of Biologists Limited 1989
147
Presence of basic fibroblast growth factor in the early Xenopus embryo
J. M. W. SLACK and H. V. ISAACS
Imperial Cancer Research Fund, Developmental Biology Unit, Department of Zoology, University of Oxford, South Parks Road, Oxford,
OX13PS
Summary
Mesoderm-inducing activity can be extracted from
Xenopus embryos, eggs or whole ovary. It binds to
heparin and can be neutralized by heparin or anti-bFGF
but not by anti-TGF/?. Two molecular forms can be
identified by Western blotting and have molecular
weights of about 19 and 14K. The content in embryos is
about 7 units g~' (approximately 7 ng ml~*) which would
be sufficient for it to be acting as an endogenous inducer
of ventral mesoderm. Attempts to detect TGF/5-like
inducing factors in embryos were not successful.
Key words: Xenopus oocyte, egg, blastula, heparin-binding
growth factors, fibroblast growth factor, transforming
growth factor beta, mesoderm-including factors.
Introduction
Much interest has been aroused recently by the possi-
bility of identifying the morphogens responsible for
mesoderm induction in the early amphibian embryo.
This is the process by which the mesoderm is induced
from the animal hemisphere of the blastula by signals
from the vegetal region (Nieuwkoop, 1969; Dale et al.
1985; Gurdon et al. 1985; Jones & Woodland, 1987).
Recently we reported that a small group of heparin-
binding growth factors (HBGFs) are active as meso-
derm-inducing agents and the relatively high degree of
specificity suggested to us that at least the ventral
mesoderm induction within the embryo was mediated
by an HBGF (Slack et al. 1987). However, it has now
been found that another type of growth factor, TGF/J-2,
is active (Rosa et al. 1988) and a mesoderm-inducing
factor recently purified from a Xenopus cell line is also
thought to belong to the TGF/3 family (Smith, 1987;
Smith et al. 1988). So which, if any, of these factors is
really involved in mesoderm induction in the early
embryo? A clue was provided by Kimelman &
Kirschner (1987) who detected an mRNA of the bFGF
type in Xenopus embryos. We now show that a meso-
derm-inducing factor can be isolated from Xenopus
blastulae. It is similar to bFGF by biochemical and
immunological criteria and has a similar mesoderm-
inducing activity in vitro. The quantity present is small
but sufficient for it to be responsible for induction in
vivo. This is consistent with, although does not yet
finally prove, the involvement of bFGF as a morphogen
in mesoderm induction.
Materials and methods
Extraction of HBGF from Xenopus material
Eggs and embryos were dejellied with 2% cysteine hydro-
chloride pH7-9 and stored at -70°C before use, as were
whole ovaries from adult females. The final protocol adopted
was as follows: 25-100 g of ovary, unfertilized eggs or em-
bryos were homogenized in 2-3 vols of 0-15M-ammonium
sulphate containing lmM-PMSF and 100 nin-pepstatin A.
Homogenization was by Ultra turrax blender for 3x1 min at
maximum speed, the temperature being kept below 10°C with
ice. All subsequent steps up to the Amicon concentration
were carried out at cold room temperature. The pH was
adjusted to 4-5 with acetic acid and the homogenate was
stirred for 1 h on ice. It was then centrifuged at 150000g for
one hour and lipid was removed by straining through nylon
mesh. The supernatant was neutralized with ammonia and
fractionated by adding, successively, 0-24 g ml"
1
(40%) and
0'2gml"' (70%) solid ammonium sulphate followed by
centrifugation at 12000g for 10min. The 40-70% cut was
dialysed overnight against 0-6M-NaCl, 20mM-Tris pH7-0 and
then applied to a 2 ml column of heparin Sepharose (Sigma or
Pharmacia) equilibrated in the same buffer. The column was
washed in the same buffer and eluted with 2M-NaCl, 20 HIM-
Tris-HCl pH7-0. Ovalbumin (Sigma, mol. wt std) was added
as carrier and the product was concentrated to 1 ml with an
Amicon ultrafiltration cell and YM10 membrane. This ma-
terial is referred to below as the 'heparin-bound fraction'.
Earlier preparations differed in that the initial homogenate
was centrifuged less vigorously (25 OOOg, 30 mins), a 40-80 %
ammonium sulphate precipitate was taken, and also that a
CM-Sephadex column at pH6 was employed before the
heparin column.
Western blots
The samples were separated using 15 % SDS-polyacrylamide
gels and transferred to nitrocellulose using standard tech-