Kosmidis et al., (2010) showed that the expression of a human 4R tau phospho-mutant (2N4RSTA which cannot be phosphorylated at Ser238 and Thr245) which is highly phosphorylated at the AT100, AT8, pS262, and pS356, leads to profound memory impairments in the absence of mushroom body degeneration [59]. This evidence concerns the gene MAPT and memory impairment.