Although nuclear envelope (NE) assembly is known to require the GTPase Ran,
the membrane fusion machinery involved is uncharacterized. NE assembly inv
olves formation of a reticular network on chromatin, fusion of this network
into a closed NE and subsequent expansion. Here we show that p97, an AAA-A
TPase previously implicated in fusion of Golgi and transitional endoplasmic
reticulum (ER) membranes together with the adaptor p47, has two discrete f
unctions in NE assembly. Formation of a closed NE requires the p97-Ufd1-Npl
4 complex, not previously implicated in membrane fusion. Subsequent NE grow
th involves a p97-p47 complex. This study provides the first insights into
the molecular mechanisms and specificity of fusion events involved in NE fo
rmation.