Cytoplasmic TDP-25 aggregates were co-labelled with antibody to TDP-43 phosphorylated at serine 409/410 (pTDP-43), appearing as an outer shell, and were also labelled with ubiquitin and p62 (Fig. 4a), recapitulating hallmarks of TDP-43 pathology in ALS/FTD. Here, TARDBP is linked to frontotemporal dementia.