SNCA and multiple system atrophy: In healthy brain, only a small proportion of α-synuclein is phosphorylated (Gonzalez et al., 2019), whereas under pathological conditions phosphorylated α-synuclein -mostly at Ser129- is increased in the majority of pathological inclusions, including LBs in PD and DLB, GCIs in MSA and LB-type inclusions of AD (Fujiwara et al., 2002; Saito et al., 2003; Nishie et al., 2004; Waxman et al., 2009).