The variant surface glycoproteins
(VSG) of Trypanosoma brucei are anchored to the cell surface via a
glycosylphosphatidylinositol (GPI) anchor. All GPI-anchored proteins
are synthesized with a C-terminal signal sequence, which is replaced
by a GPI-anchor in a rapid post-translational transamidation
reaction. VSG GPI signal sequences are extraordinarily conserved.
They contain either 23 or 17 amino acids, a difference that
distinguishes the two major VSG classes, and consist of a spacer
sequence followed by a more hydrophobic region. The w-position, to
which GPI is transferred, is either Ser, Asp or Asn, the w+2-position
is always Ser, and the w+7 position is almost always Lys. In order to
determine whether this high conservation is necessary for
GPI-anchoring, we introduced several mutations into the signal
peptide. Surprisingly, changing the most conserved amino acids, at
positions w+1, w+2 and w+7, had no detectable effect on the
efficiency of GPI-anchoring or on protein abundance. Several more
extensive changes also had no discernable impact on GPI-anchoring.
Deleting the entire 23 amino-acid signal sequence or the 15
amino-acid hydrophobic region generated proteins that were not
anchored. Instead of being secreted, these truncated proteins
accumulated in the endoplasmic reticulum, prior to lysosomal
degradation. Replacing the GPI signal sequence with a proven
cell-surface membrane-spanning domain reduced expression by about 99%
and resulted not in cell-surface expression but in accumulation close
to the flagellar pocket and in non-lysosomal compartments. These
results indicate that the high conservation of the VSG GPI signal
sequence is unnecessary for efficient expression and GPI attachment,
but that the GPI anchor is essential for surface expression of VSG.
However, because the VSG is a major virulence factor, it is possible
that small changes in the efficiency of GPI anchoring, undetectable
in our experiments, might have influenced the evolution of VSG GPI
signal sequences.