INTRODUCTION
Colon cancer is a consequence of the accumulation of
genetic alterations that lead to an abnormal balance in the
events that regulate intestinal epithelial cell growth and
differentiation (Fearon and Vogelstein, 1990). In vitro, the
study of these processes has been hampered by the lack of
reproducible methods for the culture of normal intestinal
epithelium and the undifferentiated phenotype of most col-
orectal cancer cell lines (Neutra and Louvard, 1991;
Zweibaum et al., 1991). Recently, intestinal epithelial cell
lines that are able to differentiate in culture into goblet or
absorptive cells have been described (Neutra and Louvard,
1991; Zweibaum et al., 1991). HT-29 M6 is a subpopula-
tion of the HT-29 cell line that is isolated by its ability to
grow in medium supplemented with 10
-6
methotrexate
(Lesuffleur et al., 1990)*. After confluence, HT-29 M6 cells
develop a polarized phenotype, with a moderately well
defined apical brush border and a cytoplasm essentially full
of mucous droplets (Lesuffleur et al., 1990, 1991). Prolif-
erating HT-29 M6 cells resemble pre-differentiated colon
crypt cells that are committed to differentiate into goblet
cells (Lesuffleur et al., 1990, 1991). Thus, although HT-29
M6 cells are derived from a tumour, the characteristics
described above make them a suitable model with which to
identify the molecular mechanisms controlling epithelial
cell proliferation and differentiation.
The phorbol ester 12-O-tetradecanoylphorbol-13-acetate
(TPA) is a potent activator of protein kinase C (PK-C), its
specific cellular receptor (Blumberg, 1988; Jaken, 1990).
Great interest in this phorbol ester has evolved from its use
in carcinogenesis models, in particular in skin and in the
gastrointestinal tract (Boutwell, 1974). In vivo carcinogen-
esis models have shown that tumour development proceeds
in two distinct stages: initiation, which involves a genetic
change caused by a mutagen; and promotion, an amplifi-
cation of the mutated cell population as the result of the
stimulation of cell proliferation by a ‘promoter’. TPA falls
in this second category of molecules (Goerttler et al., 1979).
Moreover, a role for other activators of PK-C in the devel-
opment and/or progression of colon cancer has been
suggested (Weinstein, 1991). To characterize further the
role of PK-C in intestinal epithelial cells, we have investi-
513
Journal of Cell Science 106, 513-522 (1993)
Printed in Great Britain © The Company of Biologists Limited 1993
The effects of tumor promoter 12-O-tetradecanoylphor-
bol-13-acetate (TPA) on the growth characteristics of
the colon cancer cell line HT-29 M6 were studied. TPA
induced the scattering of proliferative HT-29 M6 cells:
in the presence of the phorbol ester, HT-29 M6 colonies
scattered and the cells acquired a flatter aspect with
diminished cell-cell contacts. This effect of TPA
required a persistent activation of PK-C and was acom-
panied by a slight decrease (30%) in the growth rate.
Modifications by TPA of two scattering associated prop-
erties of these cells were also detected: TPA decreased
cell-to-cell aggregation and enhanced the cellular
attachment to matrix substrata (collagen, laminin). The
decrease in cell-to-cell adhesion was correlated with a
loss of cellular E-cadherin as evidenced by immunoflu-
orescence or immunoblotting with a specific monoclonal
antibody. Cell scattering was dependent on the extra-
cellular concentration of Ca
2+
; an increase from 1.6 to
10 mM in the concentration of this ion completely
blocked the morphological effects of TPA as well as its
action on cell aggregation. This high concentration of
Ca
2+
also prevented the down modulation of E-cadherin
as determined by immunofluorescence. However, the
TPA-induced increase in cell attachment to the matrix
was not affected by high calcium. These findings sup-
port the importance of altered cell-cell adhesion in the
process of scattering and provide a good system for the
study of down modulation of E-cadherin, a protein
involved in the control of cell growth, differentiation and
invasion of epithelial cells.
Key words: HT-29 cells, TPA tumor promoter, E-cadherin, cell
scattering, calcium
SUMMARY
Phorbol ester-induced scattering of HT-29 human intestinal cancer cells is
associated with down-modulation of E-cadherin
Myriam Fabre and Antonio García de Herreros*
Departament d’Immunologia, Institut Municipal d’Investigació Mèdica, Universitat Autònoma de Barcelona, Spain
*Author for correspondence
*For the sake of brevity we have given the name of HT-29 M6 to the cell line
previously named HT-29 (10-6 MTX) (Lesuffleur et al., 1990)