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. 2022 Sep 21;221(10):e202111076. doi: 10.1083/jcb.202111076

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© 2022 Liu et al.

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Figure 7. — Hypothetical model for how Chlamydomonas ARL3 promotes diffusion of the retrograde IFT train-shed and PLD-laden BBSome through the TZ for ciliary retrieval. IFT22/BBS3, when both in a GTP-bound state (IFT22^GTP/BBS3^GTP), recruits the BBSome from cytoplasm to the basal body for integrating into anterograde IFT trains (Xue et al., 2020). Upon reaching the ciliary tip via anterograde IFT, the BBSome remodels with the aid of the IFT-B1-shed IFT25/27 for promoting its reassembly (Sun et al., 2021). The reassembled BBSome interacts with its cargo PLD in a BBS3-dependent manner (Liu et al., 2021). This enables the PLD-laden BBSome to U turn in cilia for transporting to the ciliary base via retrograde IFT. ARL3^GDP binds to the ciliary membrane for diffusing into cilia and is activated to become ARL3^GTP by an unknown mechanism (?). ARL3^GTP detaches from the ciliary membrane and resides in the ciliary matrix by concentrating at the proximal ciliary region right above the TZ. Following the transportation from the ciliary tip to base, the PLD-laden BBSome separates from retrograde IFT trains at the proximal ciliary region right above the TZ and is bound to ARL3^GTP as an ARL3 effector. ARL3^GTP then facilitates the retrograde IFT train-shed and PLD-laden BBSome to diffuse through the TZ for ciliary retrieval.