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. Author manuscript; available in PMC: 2020 Jan 1.
Published in final edited form as: Neuroscience. 2019 Jan 1;396:A3–A20. doi: 10.1016/j.neuroscience.2018.10.033

Table 1.

Genetic modifiers that may modify multiple ALS genes. This list includes the human orthologs that were identified as modifiers of more than one ALS-causal genes (e.g. SRRT orthologs were reported as modifiers in both SOD1 and TDP43 models). Additionally, human genes reported as modifiers through GWAS or linkage analysis studies are also included in this list.

Human Gene Organism Modifier for Reference
ATXN2 Yeast, Human TDP43, Human (Elden et al., 2010; Figley and Gitler, 2013; Lee T. et al., 2011)
BMP2 Nematode, Fly SOD1, C9orf72 (Wang J. et al., 2009; Zhang et al., 2015)
CCNB1 Yeast, Fly FUS, VAPB (Sanhueza et al., 2015; Sun Z. et al., 2011)
CCT4 Nematode, Fly SOD1, C9orf72 (Lee K. H. et al., 2016; Wang J. et al., 2009)
CDC6 Yeast, Fly C9orf72, VAPB (Deivasigamani et al., 2014; Jovicic et al., 2015)
CHGB Mouse, Human SOD1, Human (Gros-Louis et al., 2009; Ohta et al., 2016)
COA4 Nematode, Yeast SOD1, FUS (Sun Z. et al., 2011; Wang J. et al., 2009)
DAZAP1 Fly C9orf72, VCP (Ritson et al., 2010; Zhang et al., 2015)
DBR1 Yeast TDP43, FUS (Armakola et al., 2012; Figley and Gitler, 2013)
DNAJA1 Yeast, Fly SOD1, C9orf72 (Lee K. H. et al., 2016; Strain et al., 1998)
FBXW7 Nematode, Yeast SOD1, FUS (Sun Z. et al., 2011; Wang J. et al., 2009)
GNAQ Fly C9orf72, VAPB (Deivasigamani et al., 2014; Zhang et al., 2015)
HNRNPC Yeast, Fly C9orf72, FUS (Lee K. H. et al., 2016; Sun Z. et al., 2011)
IMPA1 Yeast, Fly FUS, VAPB (Deivasigamani et al., 2014; Sun Z. et al., 2011)
IPO5 Fly C9orf72, VAPB (Deivasigamani et al., 2014; Lee K. H. et al., 2016)
KPNB1 HeLa cells, Fly TDP43, C9orf72, VAPB (Boeynaems et al., 2016; Kim S. H. et al., 2012; Sanhueza et al., 2015)
MMP9 Mouse, Human SOD1, Human (Kaplan et al., 2014; Lorenzl et al., 2006)
MRPL33 Yeast TDP43, FUS (Armakola et al., 2012; Sun Z. et al., 2011)
NAA20 Fly C9orf72, VAPB (Deivasigamani et al., 2014; Zhang et al., 2015)
NCL Yeast, Fly C9orf72, FUS (Lee K. H. et al., 2016; Sun Z. et al., 2011)
NOB1 Nematode, Yeast SOD1, C9orf72 (Jovicic et al., 2015; Wang J. et al., 2009)
NUP50 Fly TDP43, C9orf72 (Boeynaems et al., 2016; Freibaum et al., 2015; Zhan et al., 2013)
NUP107 Fly FUS, C9orf72 (Boeynaems et al., 2016; Sun Z. et al., 2011)
PDE6D Nematode, Fly SOD1, VAPB (Deivasigamani et al., 2014; Silva et al., 2011)
PELO Yeast, Fly TDP43, C9orf72 (Armakola et al., 2012; Jovicic et al., 2015)
PGM1 Yeast, Fly TDP43, C9orf72 (Kim H. J. et al., 2014; Zhang et al., 2015)
PIAS1 Nematode, Fly SOD1, VAPB (Sanhueza et al., 2015; Wang J. et al., 2009)
POLD3 Yeast, Fly C9orf72, VAPB (Deivasigamani et al., 2014; Jovicic et al., 2015)
PPP1R3C Fly, Yeast C9orf72, FUS (Sun Z. et al., 2011; Zhang et al., 2015)
RAN Human cells, Fly TDP43, C9orf72 (Freibaum et al., 2015; Kim S. H. et al., 2012)
RBM10 Fly C9orf72, VAPB (Lee K. H. et al., 2016; Sanhueza et al., 2015)
SAMD4A Yeast TDP43, C9orf72 (Jovicic et al., 2015; Kim H. J. et al., 2014)
SCARB1 Fly C9orf72, VAPB (Sanhueza et al., 2015; Zhang et al., 2015)
SIRT1 Fly C9orf72, VAPB (Sanhueza et al., 2015; Zhang et al., 2015)
SLC17A5 Nematode, Fly SOD1, C9orf72 (Wang J. et al., 2009; Zhang et al., 2015)
SOD1 Fly SOD1, VAPB (Deivasigamani et al., 2014; Kumimoto et al., 2013)
SRRT Fly C9orf72, VAPB (Deivasigamani et al., 2014; Freibaum et al., 2015)
TARDBP Fly TDP43, C9orf72, VAPB, VCP (Deivasigamani et al., 2014; Lee K. H. et al., 2016; Ritson et al., 2010; Sreedharan et al., 2015; Wang J. W. et al., 2011)
TNPO1 Fly C9orf72, VAPB (Boeynaems et al., 2016; Deivasigamani et al., 2014)
TOP1 Nematode, Fly SOD1, C9orf72 (Lee K. H. et al., 2016; Wang J. et al., 2009)
UBR5 Yeast, Fly TDP43, C9orf72 (Kim H. J. et al., 2014; Lee K. H. et al., 2016)
UQCRC2 Nematode, Fly SOD1, VAPB (Deivasigamani et al., 2014; Silva et al., 2011)
XPO1 Nematode, Fly SOD1, C9orf72 (Silva et al., 2011; Zhang et al., 2015)