Vectorial Cholesterol Transport by STARD4 is Mediated by Specific PIP2 Membrane Composition

STARD4 (steroidogenic acute regulatory protein-related lipid-transfer domain containing 4) is known to transport cholesterol from the plasma membrane (PM) to intracellular organelles, such as the endoplasmic reticulum and endocytic recycling compartment, but it has not been known if the transport is selective. Identification of a surface-exposed basic region encompassing the C-terminal helix in the crystal structure of STARD4 suggested the possibility that anionic lipids confer membrane selectivity.