It has been reported that phosphorylated TDP-43 aggregates were observed along with CDC7, Tau-tubulin kinase 1 and 2 (TTBK1 and TTBK2) in frontal cortex sections from ALS and frontotemporal lobar degeneration (FTLD-TDP) cases, and TTBK1/2 also directly phosphorylated TDP-43 at S409/410 residues in vitro [39]. Here, TARDBP is linked to amyotrophic lateral sclerosis.