Wang J , Fresquez T , Kandachar V , Deretic D .

R01 EY012421 NEI NIH HHS

	Fig. 1. The Arf GEF GBF1 is localized at the trans-Golgi and interacts with Arf4, the Arf GAP ASAP1 and the ciliary cargo, rhodopsin. (A–D) Retinas were labeled with rabbit anti-GBF1 and mouse anti-GM130, followed by Cy3- and Cy5-conjugated secondary antibodies, followed by rabbit anti-Rab6 conjugated to Alexa Fluor 488. An individual optical section is shown. G, Golgi; N, nucleus. Scale bar: 3 µm. GBF1 (red) overlaps with the trans-Golgi marker Rab6 (green) (arrows), significantly better than with the cis-Golgi marker GM130 (blue), as per pixel colocalization analysis performed within the Golgi and expressed as the Pearson’s coefficient (***P=3.42×10−5 ) (n=5 cells). (E) Schematic of a photoreceptor cell. The ROS is an elaborate primary cilium. Golgi and the TGN are localized in the myoid region (M) of the RIS. RTCs bud from the TGN and travel to the cilium (arrow), through the ellipsoid region (E) packed with mitochondria. Adherens junctions (AJ) form the outer limiting membrane (OLM) throughout the retina. (F) GBF1+Rab6 interaction sites (red dots) detected by PLA using mouse (m) anti-GBF1 and rabbit (r) anti-Rab6. Following the detection of interaction sites by PLA (arrows), sections were subsequently stained with an antibody against Rab6 conjugated to Alexa Fluor 488 (green). Nuclei were stained with TO-PRO-3 (blue). (G,H) PLA for GM130(m)+GBF1(r) (G) and P115(m)+GBF1(r) (H). (I) Golgi staining with Rab6(Alexa Fluor 488), p115(r) and GM130(m). (J–M) PLA for GBF1(m)+Arf4(r) (J), rhodopsin(m)+GBF1(r) (K), rhodopsin(m)+Arf4(r) (L) and ASAP1(m)+GBF1(r) (M). Scale bar: 5 µm. (N) Red dots were counted for the PLA pairs shown in panels J–M (30 cells each) in three separate experiments. The data from a representative experiment were expressed as a percentage of total interaction sites within the RIS, analyzed using Student’s t-test (n=30) and presented as mean±s.e.m. (O) PNS (0.1 retina), or T/G/E, RTC and cytosolic fractions (0.25 retina each), were analyzed by immunoblotting (IB), as indicated. All antibodies were tested on a single blot.
	Fig. 2. GCA, a selective inhibitor of GBF1, significantly disrupts rhodopsin-GBF1- Arf4-ASAP1 interactions. (A) Rhodopsin(m) +GBF1(r) interaction sites (red dots) detected by PLA in control retinas. Retinal sections were visualized by DIC. (B–E) The same was repeated for rhodopsin(m)+Arf4(r) (B), GBF1 (m)+Arf4(r) (C), rhodopsin(m)+ASAP1(r) (D) and ASAP1(m)+GBF1(r) (E). (F–J) PLA of GCA-treated retinas for rhodopsin(m)+GBF1 (r) (F), rhodopsin(m)+Arf4(r) (G), GBF1(m) +Arf4(r) (H), rhodopsin(m)+ASAP1(r) (I) and ASAP1(m)+GBF1(r) (J). Scale bar: 5 µm. (K–O) Experiments were repeated three times and >10 Z-stacks containing ≥10 confocal optical sections were generated for each PLA pair. From these Z-stacks, two representative confocal sections encompassing at least five photoreceptors with clearly visible RIS demarcations were selected from each experiment. Interaction sites were counted (10 photoreceptors×3 experiments) for each PLA pair, analyzed using Student’s t-test (n=30) and presented as in Fig. 1 (***P<2.2×10−6 ).
	Fig. 3. GBF1 regulates Golgi-to-cilia transport of rhodopsin. (A) Control, GCA- and BFA-treated retinas were labeled with antibody to Rab6(r) (green) and GM130(m) (red). Scale bars: 5 µm (1 µm in inset). (B) Isolated retinas were incubated in the presence or absence of GCA during the pulse-chase experiment. Following treatment, T/G/E, RTC and ROS fractions were analyzed by SDS-PAGE and autoradiography. (C) Autoradiogram of the gel shown in B. (D) Radiolabeled rhodopsin was quantified in three separate experiments. The data were analyzed using Student’s t-test (n=3) and presented as mean±s.e.m. (*P=0.01). (E) Control and GCA-treated retinas were labeled with anti-Rab6 (green), GBF1 (red) and GM130 (blue), as in Fig. 1. Rab6 and GBF1 colocalize (arrows) in control and GCA-treated retinas. Labeling with anti-GBF1(m) and anti-ASAP1(r) in control and GCA-treated retinas shows Golgi and RTC colocalization (arrows), whereas colocalization detected with anti-GBF1(m) and anti-Arf4(r) in controls is lost upon GCA treatment (arrows). Scale bar: 5 µm. (F) Following the pulse-chase experiment, subcellular fractions of control or GCA-treated retinas were separated by SDS-PAGE and immunoblotted as indicated. All antibodies were tested on a single blot. ASAP1 detected in the crude ROS fraction originates from a minor contamination with RIS proteins. (G) The distribution of GBF1, Arf4 and ASAP1 in the T/G/E fraction, RTCs and the cytosol was quantified in three separate experiments and presented as mean±s.e.m.
	Fig. 4. GBF1 directly interacts with Arf4 and the ciliary cargo, rhodopsin. (A) Schematic of GBF1. DCB-HUS (AA 1–710) and Sec7-HDS1 (AA 695–1066) are indicated. (B) GST-DCB-HUS, GST-Sec7-HDS1 or GST were incubated with purified bovine rhodopsin, with or without recombinant human Arf4 bound to GDPβS or GTPγS. Rhodopsin and Arf4 were detected by immunoblotting. The GST fusion proteins were detected with anti-GST antibody. Arrowheads point to the GST fusion proteins used in pulldowns. Breakdown products of GBF1 DCB-HUS were also observed by Galindo et al. (2016). Sec7-HDS1 is partially obscured by the BSA present in all samples. Rhodopsin and Arf4 were quantified in three separate experiments. The data were analyzed using Student’s t-test (n=3) and presented as mean±s.e.m. (**P<0.005). (C) Comparable amounts of bovine rhodopsin and human Arl1Q71L were subjected to pulldowns by GSTDCB-HUS. Bound proteins and GST fusion proteins were detected by specific antibodies. Rhodopsin and Arl1 were quantified in two separate experiments and presented as mean±range (for two experiments). (D) GST pulldown of human Arf4, or Δ17Arf1, bound to GDPβS or GTPγS. Bound Arfs and GST fusion proteins were detected by immunoblotting, and Arf4 was quantified in three separate experiments and presented as mean±s.e.m.
	Fig. 5. Influx of rhodopsin provides a signal crucial to Arf4 interaction with GBF1. (A–H) PLA was performed to detect various interaction sites in control (A, B,C,D) and cycloheximide-treated (E,F,G,H) retinas: rhodopsin(m)+GBF1(r) (A,E), rhodopsin(m)+Arf4(r) (B,F), GBF1(m)+Arf4(r) (C,G) and ASAP1(m)+GBF1(r) (D,H). Scale bar: 5 µm. (I) Interaction sites were analyzed and presented as in Fig. 2 (***P<2.0×10−13). (J) Isolated retinas were incubated in the presence or absence of cycloheximide. Following a pulse-chase experiment, T/G/E, RTCs, ROS and cytosol were analyzed by SDS-PAGE. (K) Distribution of Arf4 and ASAP1 among subcellular fractions in control and cycloheximide-treated cells was determined by immunoblotting, as indicated. (L) Autoradiogram of the gel shown in J. (M) Golgi localization (arrows) of GBF1 (red) and Arf4 (green) in control and cycloheximide-treated retinas. (N,O) Retinas were labeled with antibody to Rab6(r) (green) and rhodopsin(m) (red). Arrows indicate Golgi-localized rhodopsin in the control (N), but not in cycloheximide-treated retinas (O). Scale bars: represents 8 µm for M–O; 5 µm for insets in N and O. (P) Schematic of the apparent sequence of events in ciliary trafficking leading to the formation of the complex consisting of the cargo, the cognate Arf and an Arf GEF. Cytosolic Arf4 becomes membrane associated and activated, either through random activation events, or by an Arf-GEF. Through interactions with Rab6, GBF1 is positioned at the trans-Golgi membranes, where it associates with rhodopsin and Arf4. This process is inhibited by cycloheximide, which chiefly blocks the influx of rhodopsin into the endomembrane system. GBF1, stimulated by the cargo and Arf4 binding to the regulatory DCB-HUS domain, quickly builds up levels of active Arf4, a process inhibited by GCA. Activated Arf4 recognizes and directly binds to the VxPx C-terminal signal of incoming rhodopsin, which leads to the assembly of the ciliary targeting complex, starting with the Arf GAP ASAP1. GTP hydrolysis on Arf4, catalyzed by ASAP1, releases inactive Arf4 into the cytosol and directs rhodopsin into the nascent RTCs that contain both ASAP1 and GBF1

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