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Crumbs2 is a slit diaphragm protein, organized in clusters and able to interact with Nephrin. In mice, loss of Crumbs2 results in heavy proteinuria, accompanied by progressive podocyte foot process effacement, injury of renal cells, and inflammation, indicating that Crb2 is of similar relevance for the GFB as Nephrin.\n                      <jats:italic toggle=\"yes\">In vitro<\/jats:italic>\n                      , CRB2 mutant proteins accumulate in the ER and trigger ER stress. Thus, in addition to loss of Crumbs2 at the slit diaphragm, ER stress induction could play a pivotal role in onset and disease progression in patients with SRNS.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Background<\/jats:title>\n                    <jats:p>\n                      Crumbs2 is expressed at embryonic stages as well as in the retina, brain, and glomerular podocytes. Recent studies identified\n                      <jats:italic toggle=\"yes\">CRB2<\/jats:italic>\n                      mutations as a novel cause of steroid-resistant nephrotic syndrome (SRNS).\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Methods<\/jats:title>\n                    <jats:p>\n                      To study the function of Crb2 at the renal filtration barrier, mice lacking Crb2 exclusively in podocytes were generated. Gene expression and histologic studies as well as transmission and scanning electron microscopy were used to analyze these\n                      <jats:italic toggle=\"yes\">Crb2<\/jats:italic>\n                      <jats:sup>podKO<\/jats:sup>\n                      knockout mice and their littermate controls. Furthermore, high-resolution expansion microscopy was used to investigate Crb2 distribution in murine glomeruli. For pull-down experiments, live cell imaging, and transcriptome analyses, cell lines were applied that inducibly express fluorescent protein\u2013tagged CRB2 wild type and mutants.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Results<\/jats:title>\n                    <jats:p>\n                      <jats:italic toggle=\"yes\">Crb2<\/jats:italic>\n                      <jats:sup>podKO<\/jats:sup>\n                      mice developed proteinuria directly after birth that preceded a prominent development of disordered and effaced foot processes, upregulation of renal injury and inflammatory markers, and glomerulosclerosis. Pull-down assays revealed an interaction of CRB2 with Nephrin, mediated by their extracellular domains. Expansion microscopy showed that in mice glomeruli, Crb2 and Nephrin are organized in adjacent clusters. SRNS-associated CRB2 protein variants and a mutant that lacks a putative conserved\n                      <jats:italic toggle=\"yes\">O<\/jats:italic>\n                      -glycosylation site were not transported to the cell surface. Instead, mutants accumulated in the ER, showed altered glycosylation pattern, and triggered an ER stress response.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Conclusions<\/jats:title>\n                    <jats:p>Crb2 is an essential component of the podocyte\u2019s slit diaphragm, interacting with Nephrin. Loss of slit diaphragm targeting and increasing ER stress are pivotal factors for onset and progression of CRB2-related SRNS.<\/jats:p>\n                  <\/jats:sec>","DOI":"10.1681\/asn.2020040501","type":"journal-article","created":{"date-parts":[[2021,3,9]],"date-time":"2021-03-09T11:25:15Z","timestamp":1615289115000},"page":"1053-1070","update-policy":"https:\/\/doi.org\/10.1097\/lww.0000000000001000","source":"Crossref","is-referenced-by-count":38,"title":["Crumbs2 Is an Essential Slit Diaphragm Protein of the Renal Filtration Barrier"],"prefix":"10.1681","volume":"32","author":[{"given":"Annika","family":"M\u00f6ller-Kerutt","sequence":"first","affiliation":[{"name":"Internal Medicine D, Department of Molecular Nephrology, University Hospital of Muenster, Muenster, Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Juan E.","family":"Rodriguez-Gatica","sequence":"additional","affiliation":[{"name":"Institute of Physical and Theoretical Chemistry, 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