Internalization of membrane-bound forms of Shh. Throughout, the Shh
form is visualized in red and PtcHA in green; colocalization is seen as
yellow. (A) A PtcHA-expressing cell, whose borders are
indicated by the dashed line, is shown contacting a wtShh-expressing
cell. PtcHA immunofluorescence is not observed where the filopodium of
the wtShh-expressing cell crosses the PtcHA-expressing cell, yet the
cell contains multiple PtcHA+/Shh+ vesicles.
(B) A PtcHA-expressing cell not in contact with
wtShh-expressing cells shows Shh+/PtcHA+
vesicles. The PtcHA+ cell is surrounded by untransfected
cells in a near-confluent field; the nearest wtShh-expressing cell is
out of the field of view. Nonspecific background stippling and
autofluorescence is evident in untransfected cells. (C)
PtcHA immunoreactivity colocalizing with wtShh at points of contact
between cells at 4°C. Processes of a PtcHA-expressing cell (arrows)
are shown contacting a filopodium (arrowheads) of a wtShh-expressing
cell. Thirty-three cell pairs were analyzed, which included 10 pairs
that were not in direct contact; all showed evidence of Shh
internalization. (D) A PtcHA-expressing cell is shown in
contact with filopodia of two ShhGPI-expressing cells (lower left).
Shh+/PtcHA+ vesicles are yellow.
(E) Shh immunoreactivity is not observed within a
PtcHA+ cell that is in close proximity, but not in contact
with a filopodium of a ShhGPI-expressing cell. For ShhGPI, 24 cell
pairs in contact were analyzed and all showed evidence of ShhGPI
internalization. (F) A PtcHA-expressing cell is shown in
contact with a ShhCD4-expressing cell (upper right).
Shh+/PtcHA+ vesicles are yellow.
(G) Two PtcHA-expressing cells are shown in contact with
a cell expressing ShhCD4F (upper right), which contains a cytoplasmic
Flag epitope. Flag+/PtcHA+ vesicles are
yellow. (H) Colocalization of PtcHA and ShhCD4 at points
of contact (arrowheads) between cells was observed frequently in cells
incubated at 37°C. For ShhCD4, a total of 32 cell pairs in contact
were analyzed, and all showed evidence of ShhCD4 internalization. In
control experiments, untransfected KNRK cells showed no evidence of Shh
internalization when cocultured with COS cells expressing any of the
Shh forms. Scale bars = 10 μm (the bar in E applies to D as
well).