Table 2.

Localization of TORC1 to other organelles

Pathway Mechanism of localization Cell type Technique Reference 
mTORC1 Rheb is at the ER and the Golgi HEK293 Confocal imaging; expression of EGFP-Rheb Buerger et al., 2006  
mTORC1 Rheb and the TSC complex are at the peroxisome FAO; HepG2; MEF; HeLa Confocal imaging and subcellular fractionation; quantification of colocalization; endogenous TSC1, TSC2, and Rheb Zhang et al., 2013  
mTORC1 The TSC complex is mainly cytoplasmic COS; HeLa Wide-field imaging and subcellular fractionation; PFA fixation; expression of TSC1 and TSC2 van Slegtenhorst et al., 1998; Nellist et al., 1999  
mTORC1 The TSC complex is cytoplasmic and nuclear; Akt stimulates TSC translocation from the cytoplasm to the nucleus NIH3T3; HeLa; HEK293; Rat-1 Subcellular fractionation Rosner and Hengstschläger, 2007; Rosner et al., 2007, 
mTORC1 mTOR is at ER, Golgi, and in the nucleus; Rheb is cytoplasmic and nuclear; mTORC1 (mTOR, raptor) is in punctate structures upon amino acid starvation HEK293; CHO; HeLa Single photon FRET-FLIM on live cells; expression of mTOR-EGFP, Ds-Red-raptor, and Rheb-EGFP Yadav et al., 2013  
mTORC1 mTOR and raptor are highly abundant in the nucleus but mTORC1 integrity is higher in the cytoplasm HEK293; NIH3T3; IMR-90; MRC-5; WI-38 Subcellular fractionation Rosner and Hengstschläger, 2008  
mTORC1 mTORC1 (mTOR, raptor) is in the nucleus and the cytoplasm; mTORC1 phosphorylates NFACTc4 in the nucleus; mTOR binds to rDNA, rDNA, and tRNA genes in a rapamycin-sensitive manner; serum stimulates mTOR binding to hnRNPs COS; HEK293; HeLa Subcellular fractionation; ChIP Yang et al., 2008; Goh et al., 2010; Kantidakis et al., 2010; Shor et al., 2010; Tsang et al., 2010  
mTORC1 During hypoxia, PML inhibits mTORC1 (mTOR) by sequestering it in the nucleus away from cytoplasmic Rheb MEF; HEK293 Subcellular fractionation; confocal imaging; PFA fixation Bernardi et al., 2006  
mTORC1/2 mTOR shuttles from the cytoplasm to the nucleus HEK293; CV-1 Wide-field imaging and subcellular fractionation; PFA fixation; expression of mTOR-FLAG Kim and Chen, 2000  
mTORC1/2 mTOR is predominantly nuclear except in HEK293, where it is excluded from the nucleus HEK293; Rh30; Rh41; IMR90; HCT8; HCT29; HCT116 Confocal imaging and subcellular fractionation; PFA fixation; endogenous mTOR Zhang et al., 2002  
mTORC1 mTOR and raptor are associated with mitochondria; mTOR is associated with the mitochondrial channel VDAC Jurkat; HEK293 Subcellular fractionation Schieke et al., 2006; Ramanathan and Schreiber, 2009  
Yeast TORC1 TOR1 is mainly nuclear, and nutrient starvation or rapamycin treatment induce translocation to the cytoplasm S. cerevisiae Wide-field imaging and subcellular fractionation; endogenous TOR1 Li et al., 2006  
Pathway Mechanism of localization Cell type Technique Reference 
mTORC1 Rheb is at the ER and the Golgi HEK293 Confocal imaging; expression of EGFP-Rheb Buerger et al., 2006  
mTORC1 Rheb and the TSC complex are at the peroxisome FAO; HepG2; MEF; HeLa Confocal imaging and subcellular fractionation; quantification of colocalization; endogenous TSC1, TSC2, and Rheb Zhang et al., 2013  
mTORC1 The TSC complex is mainly cytoplasmic COS; HeLa Wide-field imaging and subcellular fractionation; PFA fixation; expression of TSC1 and TSC2 van Slegtenhorst et al., 1998; Nellist et al., 1999  
mTORC1 The TSC complex is cytoplasmic and nuclear; Akt stimulates TSC translocation from the cytoplasm to the nucleus NIH3T3; HeLa; HEK293; Rat-1 Subcellular fractionation Rosner and Hengstschläger, 2007; Rosner et al., 2007, 
mTORC1 mTOR is at ER, Golgi, and in the nucleus; Rheb is cytoplasmic and nuclear; mTORC1 (mTOR, raptor) is in punctate structures upon amino acid starvation HEK293; CHO; HeLa Single photon FRET-FLIM on live cells; expression of mTOR-EGFP, Ds-Red-raptor, and Rheb-EGFP Yadav et al., 2013  
mTORC1 mTOR and raptor are highly abundant in the nucleus but mTORC1 integrity is higher in the cytoplasm HEK293; NIH3T3; IMR-90; MRC-5; WI-38 Subcellular fractionation Rosner and Hengstschläger, 2008  
mTORC1 mTORC1 (mTOR, raptor) is in the nucleus and the cytoplasm; mTORC1 phosphorylates NFACTc4 in the nucleus; mTOR binds to rDNA, rDNA, and tRNA genes in a rapamycin-sensitive manner; serum stimulates mTOR binding to hnRNPs COS; HEK293; HeLa Subcellular fractionation; ChIP Yang et al., 2008; Goh et al., 2010; Kantidakis et al., 2010; Shor et al., 2010; Tsang et al., 2010  
mTORC1 During hypoxia, PML inhibits mTORC1 (mTOR) by sequestering it in the nucleus away from cytoplasmic Rheb MEF; HEK293 Subcellular fractionation; confocal imaging; PFA fixation Bernardi et al., 2006  
mTORC1/2 mTOR shuttles from the cytoplasm to the nucleus HEK293; CV-1 Wide-field imaging and subcellular fractionation; PFA fixation; expression of mTOR-FLAG Kim and Chen, 2000  
mTORC1/2 mTOR is predominantly nuclear except in HEK293, where it is excluded from the nucleus HEK293; Rh30; Rh41; IMR90; HCT8; HCT29; HCT116 Confocal imaging and subcellular fractionation; PFA fixation; endogenous mTOR Zhang et al., 2002  
mTORC1 mTOR and raptor are associated with mitochondria; mTOR is associated with the mitochondrial channel VDAC Jurkat; HEK293 Subcellular fractionation Schieke et al., 2006; Ramanathan and Schreiber, 2009  
Yeast TORC1 TOR1 is mainly nuclear, and nutrient starvation or rapamycin treatment induce translocation to the cytoplasm S. cerevisiae Wide-field imaging and subcellular fractionation; endogenous TOR1 Li et al., 2006  

This table groups the most important reports that link TORC1 and mTORC1 to sites apart from the lysosome. By using comparable techniques to those that revealed lysosomal localization of mTORC1 (Table 1), distinct components of TORC1 have also been identified at other sites including the nucleus and mitochondria. ChIP, chromatin immunoprecipitation; PFA, paraformaldehyde.

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