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The PM in ltc1Δ cells over-accumulates sterols and upon Arp2/3 inhibition forms offered ER-interacting invaginations, suggesting that sterol transfer plays a role in PM size homeostasis. Anterograde sterol activity from STRIC is independent of canonical vesicular trafficking but calls for Arp2/3, suggesting a novel role because of this complex. Therefore, transfer roads orthogonal to vesicular trafficking govern the flow of sterols in the cell. © 2020 Marek et al.Cotton leaf curl i