Vesicle-associated Membrane Protein-associated Protein-A (VAP-A) Interacts with the Oxysterol-binding Protein to Modify Export from the Endoplasmic Reticulum

Oxysterol-binding protein (OSBP) is 1 of 12 related proteins implicated in the regulation of vesicle transport and sterol homeostasis. A yeast two-hybrid screen using full-length OSBP as bait was undertaken to identify partner proteins that would provide clues to the function of OSBP. This resulted in the cloning of vesicle-associated membrane protein-associated protein-A (VAP-A), a syntaxin-like protein implicated in endoplasmic reticulum (ER)/Golgi vesicle transport, and phospholipid regulation in mammalian cells and yeast, respectively. By using a combination of yeast two-hybrid, glutathione S-transferase pull-down and immunoprecipitation experiments, the VAP-A-binding region in OSBP was localized to amino acids 351–442. This region did not include the pleckstrin homology (PH) domain but overlapped with the N terminus of the oxysterol binding and OSBP homology domains. C- and N-terminal truncations or deletions of VAP prevented interaction with OSBP but did not affect VAP multimerization. Although the OSBP PH domain was not necessary for VAP-A binding in vitro, interaction with VAP-A was enhanced in cells by mutation of the conserved PH domain tryptophan (OSBP W174A) or deletion of the C-terminal half of the PH domain (OSBP Δ132–182). OSBP W174A retained oxysterol binding activity, association with phospholipid vesicles via the PH domain, and localized with VAP in unusual ER-associated structures. At 40 °C, misfolded ts045-vesicular stomatitis virus G protein fused to green fluorescent protein was co-localized with VAP-A/OSBP W174A structures on the ER but was exported to the Golgi when folded normally at 32 °C. A fluorescent ceramide analogue also accumulated in these ER inclusions, and export to the Golgi was partially inhibited as indicated by decreased Golgi staining and a 30% reduction in sphingomyelin synthesis. These studies show that OSBP binding to the ER and Golgi apparatus is regulated by its PH domain and VAP interactions, and the complex is involved at a stage of protein and ceramide transport from the ER. Oxysterol-binding protein (OSBP) is 1 of 12 related proteins implicated in the regulation of vesicle transport and sterol homeostasis. A yeast two-hybrid screen using full-length OSBP as bait was undertaken to identify partner proteins that would provide clues to the function of OSBP. This resulted in the cloning of vesicle-associated membrane protein-associated protein-A (VAP-A), a syntaxin-like protein implicated in endoplasmic reticulum (ER)/Golgi vesicle transport, and phospholipid regulation in mammalian cells and yeast, respectively. By using a combination of yeast two-hybrid, glutathione S-transferase pull-down and immunoprecipitation experiments, the VAP-A-binding region in OSBP was localized to amino acids 351–442. This region did not include the pleckstrin homology (PH) domain but overlapped with the N terminus of the oxysterol binding and OSBP homology domains. C- and N-terminal truncations or deletions of VAP prevented interaction with OSBP but did not affect VAP multimerization. Although the OSBP PH domain was not necessary for VAP-A binding in vitro, interaction with VAP-A was enhanced in cells by mutation of the conserved PH domain tryptophan (OSBP W174A) or deletion of the C-terminal half of the PH domain (OSBP Δ132–182). OSBP W174A retained oxysterol binding activity, association with phospholipid vesicles via the PH domain, and localized with VAP in unusual ER-associated structures. At 40 °C, misfolded ts045-vesicular stomatitis virus G protein fused to green fluorescent protein was co-localized with VAP-A/OSBP W174A structures on the ER but was exported to the Golgi when folded normally at 32 °C. A fluorescent ceramide analogue also accumulated in these ER inclusions, and export to the Golgi was partially inhibited as indicated by decreased Golgi staining and a 30% reduction in sphingomyelin synthesis. These studies show that OSBP binding to the ER and Golgi apparatus is regulated by its PH domain and VAP interactions, and the complex is involved at a stage of protein and ceramide transport from the ER. oxysterol-binding protein N-(4,4-difluoro-5-(2-thienyl)-4-boro-3a,4a-diaza-s-indacene-3-yl)phenoxy)acetyl)sphingosine endoplasmic reticulum fetal calf serum fluorescein isothiocyanate green fluorescent protein OSBP-related proteins OSBP homology domain pleckstrin homology phosphatidylinositol 4-phosphate 5-P2, phosphatidylinositol 4,5-bisphosphate phosphatidylserine, PtdCho, phosphatidylcholine phosphatidylethanolamine protein-disulfide isomerase sphingomyelin vesicle-associated membrane protein VAMP-associated protein vesicular stomatitis virus G glutathione S-transferase phosphate-buffered saline Chinese hamster ovary tetracycline doxycycline soluble N-ethylmaleimide-sensitive factor attachment protein receptors Oxysterol-binding protein (OSBP)1 and 11 other related proteins constitute the recently identified OSBP-related (ORP) gene family. Family members share amino acid sequence identity in a 350-amino acid C-terminal OSBP homology (OH) domain that is encompassed in the oxysterol-binding region of OSBP. Most ORPs also contain an N-terminal pleckstrin homology (PH) domain (1Lehto M. Laitinen S. Chinetti G. Johansson M. Ehnholm C. Staels B. Ikonen E. Olkkonen V.M. J. Lipid Res. 2001; 42: 1203-1213Abstract Full Text Full Text PDF PubMed Google Scholar, 2Fournier M.V. Guimaraes da Costa F. Paschoal M.E. Ronco L.V. Carvalho M.G. Pardee A.B. Cancer Res. 1999; 59: 3748-3753PubMed Google Scholar, 3Xu J. 2001; Full Text Full Text PDF PubMed Google Scholar, J. 2001; Full Text Full Text PDF PubMed Google Scholar, C. J. PubMed Google of the of gene that ORPs and OSBP share related in cells or of these is but in yeast and mammalian cells a in vesicular and and sterol regulation J. 2001; Full Text Full Text PDF PubMed Google Scholar, J. 2001; Full Text Full Text PDF PubMed Google Scholar, C. J. PubMed Google Scholar, J. PubMed Google Scholar, J. J. 2001; PubMed Google the of the gene to was identified by and J. Full Text PDF PubMed Google as a for a of oxysterol of as on a the of to and for was that OSBP a in the of on J. Full Text PDF PubMed Google and studies that OSBP was a soluble protein that from a to the Golgi apparatus when cells with J. PubMed Google with the Golgi apparatus to the OSBP PH domain and is by phosphatidylinositol 4,5-bisphosphate and other protein or J. PubMed Google Scholar, S. Full Text Full Text PDF PubMed Google interaction of OSBP with the Golgi apparatus via the PH domain is for function of OSBP with deletions of the PH domain did not in that with protein J. PubMed Google a for binding to the C-terminal region of OSBP the PH domain and binding to or protein in the Golgi of the OSBP PH domain was to Golgi and with transport of vesicular stomatitis virus G protein in the Golgi apparatus S. Full Text Full Text PDF PubMed Google This that an in vesicle transport is regulated by and or sterol also to the Golgi in to other of with J. Full Text Full Text PDF PubMed Google by in cells J. PubMed Google or the J. Lipid Res. 2001; 42: Full Text Full Text PDF PubMed Google OSBP to the of these was that oxysterol for of membrane sphingomyelin and phosphatidylcholine with also of OSBP to the Golgi apparatus J. Full Text Full Text PDF PubMed Google This the that OSBP is to the of the ceramide and yeast, a and Golgi vesicle by the protein M. S. PubMed Google Scholar, S. PubMed Google Scholar, S. J. Full Text Full Text PDF PubMed Google of the yeast OSBP the in regulation of Golgi vesicular transport, Golgi function in to these M. S. M. J. PubMed Google Although of the yeast proteins deletion of the gene was J. J. 2001; PubMed Google of or of was by in and membrane function J. J. 2001; PubMed Google Scholar, M. S. M. J. PubMed Google Scholar, B. J. PubMed Google to an for OSBP and related members in and in the and of the of proteins that with OSBP using the yeast two-hybrid protein identified in screen was vesicle-associated membrane protein protein-A (VAP-A), of a and of related membrane proteins localized to the endoplasmic reticulum (ER)/Golgi PubMed Google M. J. 1999; PubMed Google of VAP proteins with and other of the vesicle Res. 2001; PubMed Google and and to the J. PubMed Google a in vesicle of VAP proteins to a of related to vesicle transport regulation of vesicle transport in the M. J. 1999; PubMed Google PubMed Google at the membrane PubMed Google and interaction with the S. PubMed Google and J. J. 1999; PubMed Google the the of VAP-A and OSBP that interaction in the Golgi and show that the VAP-A/OSBP interaction in is regulated by the PH domain and involved at a stage of protein and export from the of its oxysterol binding and and in to and transport, OSBP implicated in regulation of and sterol homeostasis. This the of interaction with of the vesicle transport OSBP localized to the Golgi apparatus in an J. PubMed Google and the OSBP PH domain was to function when in cells S. Full Text Full Text PDF PubMed Google of that OSBP with VAP-A and that OSBP PH domain with VAP in the ER in and export of and is a membrane protein with a N-terminal domain the and a acid terminus M. J. 1999; PubMed Google VAP as an acid to the N terminus that is conserved S. M. J. J. PubMed Google a and a region at the function of these is but deletion OSBP but not VAP as by yeast two-hybrid interaction M. J. 1999; PubMed Google mammalian VAP implicated in vesicle transport but at stage is PubMed Google Scholar, M. J. 1999; PubMed Google Scholar, PubMed Google the of as as to in transport M. J. 1999; PubMed Google of vesicles in the of VAP that VAP was involved in the of vesicles transport the Golgi apparatus and ER. or is involved in these is and VAP did not in and with other of the vesicle transport in Res. 2001; PubMed Google J. PubMed Google of OSBP by yeast two-hybrid and pull-down was to identify a region amino acids and that VAP-A This region overlapped with the N terminus of the oxysterol binding domain and a of the domain amino acids and that is by OSBP members (1Lehto M. Laitinen S. Chinetti G. Johansson M. Ehnholm C. Staels B. Ikonen E. Olkkonen V.M. J. Lipid Res. 2001; 42: 1203-1213Abstract Full Text Full Text PDF PubMed Google that other OSBP members with VAP in yeast two-hybrid and by immunoprecipitation the that the region is involved and that VAP is a partner for other OSBP did not J. PubMed Google but a with that oxysterol binding is not necessary for VAP-A VAP-A/OSBP interaction did not the PH of domain enhanced to ER-associated structures in A for is that the PH domain is for of OSBP to the Golgi VAP-A is involved in OSBP interaction at the ER. of the OSBP PH localized to the Golgi apparatus in cells and to Golgi in S. Full Text Full Text PDF PubMed Google VAP co-localized with OSBP in a but also in a region that to the Golgi the OSBP PH domain was that the VAP-A interaction was and OSBP was localized with VAP-A in ER structures. This is not a of for OSBP binding to receptors deletion of the OSBP PH domain the protein J. PubMed Google and the W174A retained binding to a in the that with the PH domain is involved in the interaction with This protein or partner is by the W174A mutation or of the PH a the OSBP PH domain in 1999; PubMed Google and the OSBP PH domain is to other G. B. J. Full Text Full Text PDF PubMed Google OSBP W174A is not to or of the protein as indicated by of the VAP-A and interaction of the PH domain with vesicles that fluorescent ceramide accumulated in the W174A ER and was partially was not OSBP and to J. Full Text Full Text PDF PubMed Google Scholar, J. Lipid Res. 1999; Full Text Full Text PDF PubMed Google Scholar, J. Lipid Res. Full Text PDF PubMed Google and protein transport from the ER to S. M. M. M. S. J. 2001; Full Text Full Text PDF PubMed Google Scholar, M. M. J. 1999; PubMed Google is ceramide and would the in OSBP is that the of W174A OSBP was on ceramide transport, and was localized to the ER. was also in the and from W174A structures at the This was not the for was retained in these structures and to export to the Golgi and of misfolded and ceramide in ER by interaction VAP-A and OSBP a in an ER export or protein This would at a stage these did not ER export or the of misfolded that is involved in a protein misfolded was in the as and other soluble not in these structures and not from the ER. a the yeast VAP and the protein that mammalian VAP and OSBP a related function J. E. J. PubMed Google of resulted in °C, a that was by phospholipid and the protein indicated by the of in by J. E. J. PubMed Google is a protein that the for the protein from the ER to the by in of the factor in protein and 1999; PubMed Google the ER to the of its in vesicle transport in mammalian affect the protein of by or export from the ER a to the is that the interaction OSBP and VAP a that the export of proteins from the ER to the Golgi to J. J. PubMed Google and sterol synthesis. a export of the sterol protein complex from the ER is regulated by Full Text Full Text PDF PubMed Google but is VAP or involved in a of yeast of a complex that was using as bait M. M. J. M. C. M. M. M. S. B. C. E. G. M. B. B. G. G. PubMed Google This resulted in of that and members of the yeast OSBP that contain N-terminal PH and J. J. 2001; PubMed Google Although members of complex to is that the complex also a of J. Full Text Full Text PDF PubMed Google and a phosphatidylinositol J. M.E. J. Full Text Full Text PDF PubMed Google These that and involved at the regulation and the protein in yeast, and with that VAP and the is conserved of OSBP members that with VAP that and related regulated by Oxysterol-binding protein (OSBP)1 and 11 other related proteins constitute the recently identified OSBP-related (ORP) gene family. Family members share amino acid sequence identity in a 350-amino acid C-terminal OSBP homology (OH) domain that is encompassed in the oxysterol-binding region of OSBP. Most ORPs also contain an N-terminal pleckstrin homology (PH) domain (1Lehto M. Laitinen S. Chinetti G. Johansson M. Ehnholm C. Staels B. Ikonen E. Olkkonen V.M. J. Lipid Res. 2001; 42: 1203-1213Abstract Full Text Full Text PDF PubMed Google Scholar, 2Fournier M.V. Guimaraes da Costa F. Paschoal M.E. Ronco L.V. Carvalho M.G. Pardee A.B. Cancer Res. 1999; 59: 3748-3753PubMed Google Scholar, 3Xu J. 2001; Full Text Full Text PDF PubMed Google Scholar, J. 2001; Full Text Full Text PDF PubMed Google Scholar, C. J. PubMed Google of the of gene that ORPs and OSBP share related in cells or of these is but in yeast and mammalian cells a in vesicular and and sterol regulation J. 2001; Full Text Full Text PDF PubMed Google Scholar, J. 2001; Full Text Full Text PDF PubMed Google Scholar, C. J. PubMed Google Scholar, J. PubMed Google Scholar, J. J. 2001; PubMed Google the of the gene to was identified by and J. Full Text PDF PubMed Google as a for a of oxysterol of as on a the of to and for was that OSBP a in the of on J. Full Text PDF PubMed Google and studies that OSBP was a soluble protein that from a to the Golgi apparatus when cells with J. PubMed Google with the Golgi apparatus to the OSBP PH domain and is by phosphatidylinositol 4,5-bisphosphate and other protein or J. PubMed Google Scholar, S. Full Text Full Text PDF PubMed Google interaction of OSBP with the Golgi apparatus via the PH domain is for function of OSBP with deletions of the PH domain did not in that with protein J. PubMed Google a for binding to the C-terminal region of OSBP the PH domain and binding to or protein in the Golgi of the OSBP PH domain was to Golgi and with transport of vesicular stomatitis virus G protein in the Golgi apparatus S. Full Text Full Text PDF PubMed Google This that an in vesicle transport is regulated by and or sterol OSBP also to the Golgi in to other of with J. Full Text Full Text PDF PubMed Google by in cells J. PubMed Google or the J. Lipid Res. 2001; 42: Full Text Full Text PDF PubMed Google OSBP to the of these was that oxysterol for of membrane sphingomyelin and phosphatidylcholine with also of OSBP to the Golgi apparatus J. Full Text Full Text PDF PubMed Google This the that OSBP is to the of the ceramide and yeast, a and Golgi vesicle by the protein M. S. PubMed Google Scholar, S. PubMed Google Scholar, S. J. Full Text Full Text PDF PubMed Google of the yeast OSBP the in regulation of Golgi vesicular transport, Golgi function in to these M. S. M. J. PubMed Google Although of the yeast proteins deletion of the gene was J. J. 2001; PubMed Google of or of was by in and membrane function J. J. 2001; PubMed Google Scholar, M. S. M. J. PubMed Google Scholar, B. J. PubMed Google Although to an for OSBP and related members in and in the and of the of proteins that with OSBP using the yeast two-hybrid protein identified in screen was vesicle-associated membrane protein protein-A (VAP-A), of a and of related membrane proteins localized to the endoplasmic reticulum (ER)/Golgi PubMed Google M. J. 1999; PubMed Google of VAP proteins with and other of the vesicle Res. 2001; PubMed Google and and to the J. PubMed Google a in vesicle of VAP proteins to a of related to vesicle transport regulation of vesicle transport in the M. J. 1999; PubMed Google PubMed Google at the membrane PubMed Google and interaction with the S. PubMed Google and J. J. 1999; PubMed Google the the of VAP-A and OSBP that interaction in the Golgi and show that the VAP-A/OSBP interaction in is regulated by the PH domain and involved at a stage of protein and export from the ER. of its oxysterol binding and and in to and transport, OSBP implicated in regulation of and sterol homeostasis. This the of interaction with of the vesicle transport OSBP localized to the Golgi apparatus in an J. PubMed Google and the OSBP PH domain was to function when in cells S. Full Text Full Text PDF PubMed Google of that OSBP with VAP-A and that OSBP PH domain with VAP in the ER in and export of and is a membrane protein with a N-terminal domain the and a acid terminus M. J. 1999; PubMed Google VAP as an acid to the N terminus that is conserved S. M. J. J. PubMed Google a and a region at the function of these is but deletion OSBP but not VAP as by yeast two-hybrid interaction M. J. 1999; PubMed Google mammalian VAP implicated in vesicle transport but at stage is PubMed Google Scholar, M. J. 1999; PubMed Google Scholar, PubMed Google the of as as to in transport M. J. 1999; PubMed Google of vesicles in the of VAP that VAP was involved in the of vesicles transport the Golgi apparatus and ER. or is involved in these is and VAP did not in and with other of the vesicle transport in Res. 2001; PubMed Google J. PubMed Google of OSBP by yeast two-hybrid and pull-down was to identify a region amino acids and that VAP-A This region overlapped with the N terminus of the oxysterol binding domain and a of the domain amino acids and that is by OSBP members (1Lehto M. Laitinen S. Chinetti G. Johansson M. Ehnholm C. Staels B. Ikonen E. Olkkonen V.M. J. Lipid Res. 2001; 42: 1203-1213Abstract Full Text Full Text PDF PubMed Google that other OSBP members with VAP in yeast two-hybrid and by immunoprecipitation the that the region is involved and that VAP is a partner for other OSBP did not J. PubMed Google but a with that oxysterol binding is not necessary for VAP-A VAP-A/OSBP interaction did not the PH of domain enhanced to ER-associated structures in A for is that the PH domain is for of OSBP to the Golgi VAP-A is involved in OSBP interaction at the ER. of the OSBP PH localized to the Golgi apparatus in cells and to Golgi in S. Full Text Full Text PDF PubMed Google VAP co-localized with OSBP in a but also in a region that to the Golgi the OSBP PH domain was that the VAP-A interaction was and OSBP was localized with VAP-A in ER structures. This is not a of for OSBP binding to receptors deletion of the OSBP PH domain the protein J. PubMed Google and the W174A retained binding to a in the that with the PH domain is involved in the interaction with This protein or partner is by the W174A mutation or of the PH a the OSBP PH domain in 1999; PubMed Google and the OSBP PH domain is to other G. B. J. Full Text Full Text PDF PubMed Google OSBP W174A is not to or of the protein as indicated by of the VAP-A and interaction of the PH domain with vesicles that fluorescent ceramide accumulated in the W174A ER and was partially was not OSBP and to J. Full Text Full Text PDF PubMed Google Scholar, J. Lipid Res. 1999; Full Text Full Text PDF PubMed Google Scholar, J. Lipid Res. Full Text PDF PubMed Google and protein transport from the ER to S. M. M. M. S. J. 2001; Full Text Full Text PDF PubMed Google Scholar, M. M. J. 1999; PubMed Google is ceramide and would the in OSBP is that the of W174A OSBP was on ceramide transport, and was localized to the ER. was also in the and from W174A structures at the This was not the for was retained in these structures and to export to the Golgi and of misfolded and ceramide in ER by interaction VAP-A and OSBP a in an ER export or protein This would at a stage these did not ER export or the of misfolded that is involved in a protein misfolded was in the as and other soluble not in these structures and not from the ER. a the yeast VAP and the protein that mammalian VAP and OSBP a related function J. E. J. PubMed Google of resulted in °C, a that was by phospholipid and the protein indicated by the of in by J. E. J. PubMed Google is a protein that the for the protein from the ER to the by in of the factor in protein and 1999; PubMed Google the ER to the of its in vesicle transport in mammalian affect the protein of by or export from the ER a to the is that the interaction OSBP and VAP a that the export of proteins from the ER to the Golgi to J. J. PubMed Google and sterol synthesis. a export of the sterol protein complex from the ER is regulated by Full Text Full Text PDF PubMed Google but is VAP or involved in a of yeast of a complex that was using as bait M. M. J. M. C. M. M. M. S. B. C. E. G. M. B. B. G. G. PubMed Google This resulted in of that and members of the yeast OSBP that contain N-terminal PH and J. J. 2001; PubMed Google Although members of complex to is that the complex also a of J. Full Text Full Text PDF PubMed Google and a phosphatidylinositol J. M.E. J. Full Text Full Text PDF PubMed Google These that and involved at the regulation and the protein in yeast, and with that VAP and the is conserved of OSBP members that with VAP that and related regulated by By of its oxysterol binding and and in to and transport, OSBP implicated in regulation of and sterol homeostasis. This the of interaction with of the vesicle transport OSBP localized to the Golgi apparatus in an J. PubMed Google and the OSBP PH domain was to function when in cells S. Full Text Full Text PDF PubMed Google of that OSBP with VAP-A and that OSBP PH domain with VAP in the ER in and export of and VAP-A is a membrane protein with a N-terminal domain the and a acid terminus M. J. 1999; PubMed Google VAP as an acid to the N terminus that is conserved S. M. J. J. PubMed Google a and a region at the function of these is but deletion OSBP but not VAP as by yeast two-hybrid interaction M. J. 1999; PubMed Google mammalian VAP implicated in vesicle transport but at stage is PubMed Google Scholar, M. J. 1999; PubMed Google Scholar, PubMed Google the of as as to in transport M. J. 1999; PubMed Google of vesicles in the of VAP that VAP was involved in the of vesicles transport the Golgi apparatus and ER. or is involved in these is and VAP did not in and with other of the vesicle transport in Res. 2001; PubMed Google J. PubMed Google of OSBP by yeast two-hybrid and pull-down was to identify a region amino acids and that VAP-A This region overlapped with the N terminus of the oxysterol binding domain and a of the domain amino acids and that is by OSBP members (1Lehto M. Laitinen S. Chinetti G. Johansson M. Ehnholm C. Staels B. Ikonen E. Olkkonen V.M. J. Lipid Res. 2001; 42: 1203-1213Abstract Full Text Full Text PDF PubMed Google that other OSBP members with VAP in yeast two-hybrid and by immunoprecipitation the that the region is involved and that VAP is a partner for other OSBP did not J. PubMed Google but a with that oxysterol binding is not necessary for VAP-A VAP-A/OSBP interaction did not the PH of domain enhanced to ER-associated structures in A for is that the PH domain is for of OSBP to the Golgi VAP-A is involved in OSBP interaction at the ER. of the OSBP PH localized to the Golgi apparatus in cells and to Golgi in S. Full Text Full Text PDF PubMed Google VAP co-localized with OSBP in a but also in a region that to the Golgi the OSBP PH domain was that the VAP-A interaction was and OSBP was localized with VAP-A in ER structures. This is not a of for OSBP binding to receptors deletion of the OSBP PH domain the protein J. PubMed Google and the W174A retained binding to a in the that with the PH domain is involved in the interaction with This protein or partner is by the W174A mutation or of the PH a the OSBP PH domain in 1999; PubMed Google and the OSBP PH domain is to other G. B. J. Full Text Full Text PDF PubMed Google OSBP W174A is not to or of the protein as indicated by of the VAP-A and interaction of the PH domain with vesicles that fluorescent ceramide accumulated in the W174A ER and was partially was not OSBP and to J. Full Text Full Text PDF PubMed Google Scholar, J. Lipid Res. 1999; Full Text Full Text PDF PubMed Google Scholar, J. Lipid Res. Full Text PDF PubMed Google and protein transport from the ER to S. M. M. M. S. J. 2001; Full Text Full Text PDF PubMed Google Scholar, M. M. J. 1999; PubMed Google is ceramide and would the in OSBP is that the of W174A OSBP was on ceramide transport, and was localized to the ER. was also in the and from W174A structures at the This was not the for was retained in these structures and to export to the Golgi and of misfolded and ceramide in ER by interaction VAP-A and OSBP a in an ER export or protein This would at a stage these did not ER export or the of misfolded that is involved in a protein misfolded was in the as and other soluble not in these structures and not from the ER. a the yeast VAP and the protein that mammalian VAP and OSBP a related function J. E. J. PubMed Google of resulted in °C, a that was by phospholipid and the protein indicated by the of in by J. E. J. PubMed Google is a protein that the for the protein from the ER to the by in of the factor in protein and 1999; PubMed Google the ER to the of its in vesicle transport in mammalian affect the protein of by or export from the ER a to the is that the interaction OSBP and VAP a that the export of proteins from the ER to the Golgi to J. J. PubMed Google and sterol synthesis. a export of the sterol protein complex from the ER is regulated by Full Text Full Text PDF PubMed Google but is VAP or involved in a of yeast of a complex that was using as bait M. M. J. M. C. M. M. M. S. B. C. E. G. M. B. B. G. G. PubMed Google This resulted in of that and members of the yeast OSBP that contain N-terminal PH and J. J. 2001; PubMed Google Although members of complex to is that the complex also a of J. Full Text Full Text PDF PubMed Google and a phosphatidylinositol J. M.E. J. Full Text Full Text PDF PubMed Google These that and involved at the regulation and the protein in yeast, and with that VAP and the is conserved of OSBP members that with VAP that and related regulated by and in the

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