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Figure 1.

Analysis of the human 5′-UTR database, overview of the approach, and construction of the in-frame cDNA expression library.

(A) Analysis of the human 5′-UTR database (http://utrdb.ba.itb.cnr.it/) to predict their effects on expressed sequences following translation with a YFP1 tag peptide as fusion proteins during the construction of a prey cDNA library. (B) Overview of the screening procedure. (C) For the construction of the in-frame cDNA expression library, mRNA was isolated from normal human urothelial cells and was used as a template for first-strand cDNA synthesis using polyT primer. Double-stranded cDNAs without 5′-UTRs were synthesized using primers 1 and 2 (representing approximately 40% of the Kozak sequences that are present in vertebrate genomes) complemented with primer mixes 3 and 4 (representing the remaining 60% of the Kozak sequence combinations in vertebrates). In primer mixes 3 and 4, the combination of sequences referred to as “D” is an equal mixture of A, G and T, “H” is an equal mixture of A, C and T, “K” is an equal mixture of G and T, and “W” is an equal mixture of A and T. There are 19,683 and 157,464 possible sequence combinations in primer mixes 3 and 4, respectively. (D) Sequence analysis of the in-frame cDNA library was performed on 198 representative plasmids isolated from random colonies of the library.

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Figure 2.

Predicted expression of CRABP2 and PGAM1 proteins by the constructs with and without 5′-UTRs.

(A) Comparison of CRABP2 expression constructs with and without the CRABP2 5′-UTR. (B) Comparison of PGAM1 expression constructs with and without the PGAM1 5′-UTR.

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Figure 3.

Identification of the CRABP2 and PGAM1 proteins as ARL11-binding partners using the in-frame cDNA expression library.

(A) Western blot analysis of N-terminal YFP1-tagged fusion proteins expressed by the constructs with and without 5′-UTRs in HEK-293T cells. Expression of a construct containing only YFP1 was used as a control. An anti-GFP N-terminal antibody was used to visualize the expressed tagged proteins. (B) YFP fluorescence in HEK-293T cells after the co-transfection of YFP1-CRABP2 with ARL11-YFP2 and YFP1-PGAM1 with ARL11-YFP2. Nuclei were counterstained with DAPI. (C) Confirmation of the interaction between ARL11 and CRABP2 by western blotting and co-immunoprecipitation. HEK-293T cells were transfected with HA-tagged ARL11 and FLAG-tagged CRABP2 constructs without its 5′-UTR. Protein expression was verified by immunoblotting using anti-ARL11 and anti-CRABP2 antibodies in direct western blots (DWB). Immunoprecipitation with western blotting (IPWB) was performed by anti-HA antibody pull-down of ARL11 to detect CRABP2 binding (top panel). Results were confirmed using a complementary approach (HA-tagged CRABP2, anti-HA antibody immunoprecipitation, and anti-ARL11 immunoblotting (bottom panel). (D) Confirmation of ARL 11 and PGAM1 binding by IPWB. HEK-293T cells were transfected with HA-tagged ARL11 and flag-tagged PGAM1 constructs as indicated. Protein expression verified by immunoblotting with anti-PGAM1 (top panel) or anti-ARL11 (bottom panel) antibodies (DWBs). IPWBs were performed by anti-HA immunoprecipiation of ARL11 followed by immunoblotting with anti-PGAM1 (Top panel). Alternatively, immunoprecipitation was performed using HA-tagged PGAM1 followed by immunoblotting with anti-ARL11 (Bottom panel). (E) The in-frame cDNA library prevented interference caused by the CRABP2 5′-UTR that inhibits its binding to ARL11. HEK-293T cells were transfected with HA-ARL11 and with YFP1-CRABP2 or YFP1-5′-UTR-CRABP2 as indicated. Protein expression was confirmed by immunoblotting with anti-CRABP2 antibody (top panel) or anti-YFP1 antibody (bottom panel). Alternatively, ARL11 was immunoprecipitated using the anti-HA antibody, and bound proteins were detected by immunoblotting with anti-CRABP2 (top panel) or anti-YFP1 (bottom panel) antibody. (F) The in-frame cDNA library prevents interference caused by the PGAM1 5′-UTR that prevents its binding to ARL11. HEK-293T cells were transfected with HA-ARL11 and YFP1-PGAM1, or YFP1-5′-UTR-PGAM1. Protein expression was confirmed using anti-PGAM1 (top panel) or anti-YFP1 (bottom panel) antibodies (DWB). To identify ARL11-associated proteins (IPWB), ARL11 was immunoprecipitated using the anti-HA antibody and bound proteins were detected using either an anti-PGM1 (top panel) or anti-YFP (bottom panel) antibody.

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Table 1.

Sequences of forward primers used to construct in-frame cDNA library.

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