Deciphering the role of USH1G/SANS in proteinprotein interactions and nuclear shuttling.

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Abstract

In my thesis I investigated the molecular mechanisms underlying Human Usher syndrome (USH), particularly focusing on the protein USH1G/SANS. Usher syndrome, a clinically and genetically heterogeneous disorder that leads to hearing and vision loss. The USH1G gene encodes the scaffold protein SANS, which is highly expressed in the eye and ear. Recently, SANS has also been found in the cell nucleus, where it participates in the regulation of pre-mRNA splicing. In my thesis I aimed to elucidate the interactions of SANS with splicing-related proteins PRPF31 and PRPF6, as well as the mechanism of SANS’ nuclear-cytoplasmic shuttling. Additionally, I established a method to monitor the nuclear transfer of PRPF31 by live cell imaging. I show that SANS interacts with PRPF31 and PRPF6 at specific sites within its CENTn domain. Using FRET-based interaction assays and AlphaFold2, we identified for PRPF31 and PRPF6 specific binding sites in the CENTn domain of SANS, namely binding to the structured CENTn1 and to the unstructured CENTn2, respectively. In addition, we found evidence for sequential binding of PRPF31 and PRPF6 to the SANS molecule, which might be crucial for role of SANS in splicing processes in the nucleus. To fulfill its nuclear functions, SANS needs to be transported into the nucleus. In my thesis I identified two nuclear localization sequences (NLSs) and two nuclear export sequences (NESs). I highlighted their critical roles in SANS subcellular localization and in nuclear–cytoplasmic shuttling. Comparative analysis with SANS’ paralogue ANKS4B revealed distinct localization patterns, with ANKS4B more confined to the cytoplasm due to a lack of NLS. Using pathogenic variants of USH1G/SANS, we demonstrated altered interactions with the splicing proteins PRPF31 and PRPF6. Additionally, these pathogenic variants displayed subcellular mislocalization, likely impacting SANS’ function in splicing regulation and in the development of USH. Further, I provide data on the nuclear mobility of photoactivatable fluorescencetagged PRPF31 and the rapid transfer of splicing components between nuclear compartments. With these results, I could lay the groundwork for future studies on SANS-dependent molecular dynamics. Collectively, I enlighten the role of SANS in the nucleus. My findings advance the understanding of the molecular functions of SANS and its broader implications for USH1G pathogenesis.

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