Chloroplasts have evolved an elaborate system of membrane and soluble subco
mpartments to organize and regulate photosynthesis and essential aspects of
amino acid and lipid metabolism. The biogenesis and maintenance of organel
lar architecture rely on protein subunits encoded by both nuclear and plast
id genomes. Import of nuclear-encoded proteins is mediated by interactions
between the intrinsic N-terminal transit sequence of the nuclear-encoded pr
eprotein and a common import machinery at the chloroplast envelope. Recent
investigations have shown that there are two unique membrane-bound transloc
ation systems, in the outer and inner envelope membranes, which physically
associate during import to transport preproteins from the cytoplasm to the
internal stromal compartment. This review discusses current understanding o
f these translocation systems and models for the way in which they might fu
nction.