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Table 2

Tumor-inhibiting and tumor-promoting genes induced in senescent tumor cells

GeneSenescent cells/inducerReference
Intracellular growth inhibitors   
p21Waf1/Cip1/Sdi1 (pleiotropic CDK inhibitor) Different tumor cell lines/various drugs and radiation (31, 42, 48, 49) 
p16Ink4A (CDK4/6 inhibitor; tumor suppressor) K562 leukemia/hydroxyurea; murine Eμ-myc lymphoma/cyclophosphamide (42, 60) 
p27Kip1 (CDK2 inhibitor) K562 leukemia/hydroxyurea  (42)  
BTG1 (tumor suppressor) HCT116 colon caa/doxorubicin  (49)  
BTG2 (BTG1-related) HCT116 colon ca/doxorubicin  (49)  
 EPLIN (actin-binding protein; putative tumor suppressor) HCT116 colon ca/doxorubicin (49, 56) 
 MCF-7 breast ca/retinoids  
FAT10 (ubiquitin family) MCF-7 breast ca/retinoids  (56)  
Secreted and cell-surface-associated tumor-inhibiting proteins   
IGFBP-3 (IGF-binding protein; cytostatic; apoptogenic) HeLa cervical ca/bromodeoxyuridine; MCF-7 breast ca/retinoids (48, 56) 
IGFBP-6 (IGF-binding protein; cytostatic) HCT116 colon ca/doxorubicin  (49)  
IGFBP-4 (IGF-binding protein; cytostatic) HeLa cervical ca/bromodeoxyuridine  (48)  
Maspin (serine protease inhibitor; inhibits invasion and angiogenesis; tumor suppressor) HCT116 colon ca/doxorubicin  (49)  
MIC-1 (TGF-β family) HCT116 colon ca/doxorubicin  (49)  
βig-H3 (ECM protein; inhibits tumorigenesis) HeLa cervical ca/bromodeoxyuridine; MCF-7 breast ca/retinoids (48, 56) 
Secreted and cell-surface-associated tumor-promoting proteins   
TGFα (mitogen; EGFR ligand) HCT116 colon ca/doxorubicin  (49)  
Cyr61 (angiogenic; mitogenic) HCT116 colon ca/doxorubicin  (49)  
CTGF (Cyr61-related; angiogenic; mitogenic) HT1080 fibrosarcoma/p21  (4)  
Epithelin/granulin (mitogenic) HT1080 fibrosarcoma/p21  (4)  
Galectin-3 (antiapoptotic) HT1080 fibrosarcoma/p21  (4)  
Prosaposin (antiapoptotic) HT1080 fibrosarcoma/p21 (4, 49) 
 HCT116 colon ca/doxorubicin  
 βAPP (Alzheimer’s β amyloid precursor; mitogenic) HT1080 fibrosarcoma/p21 (4, 49) 
 HCT116 colon ca/doxorubicin  
Cathepsin B (serine protease; promotes invasion) HT1080 fibrosarcoma/p21  (4)  
Plasminogen activator inhibitor-1 (serine protease inhibitor; promotes angiogenesis and invasion) HeLa cervical ca/bromodeoxyuridine; HT1080 fibrosarcoma/p21 (4, 48) 
Dual-function growth regulators   
Amphiregulin (EGF family; mitogenic in normal cells; antimitogenic in tumor cells) HCT116 colon ca/doxorubicin  (49)  
Activin A (TGF-β family) HT1080 fibrosarcoma/p21  (4)  
CD44 (cell adhesion molecule; pleiotropic growth modulator) HCT116 colon ca/doxorubicin  (49)  
Jagged-1 (Notch receptor; stem cell growth regulator; angiogenic) HCT116 colon ca/doxorubicin  (49)  
GeneSenescent cells/inducerReference
Intracellular growth inhibitors   
p21Waf1/Cip1/Sdi1 (pleiotropic CDK inhibitor) Different tumor cell lines/various drugs and radiation (31, 42, 48, 49) 
p16Ink4A (CDK4/6 inhibitor; tumor suppressor) K562 leukemia/hydroxyurea; murine Eμ-myc lymphoma/cyclophosphamide (42, 60) 
p27Kip1 (CDK2 inhibitor) K562 leukemia/hydroxyurea  (42)  
BTG1 (tumor suppressor) HCT116 colon caa/doxorubicin  (49)  
BTG2 (BTG1-related) HCT116 colon ca/doxorubicin  (49)  
 EPLIN (actin-binding protein; putative tumor suppressor) HCT116 colon ca/doxorubicin (49, 56) 
 MCF-7 breast ca/retinoids  
FAT10 (ubiquitin family) MCF-7 breast ca/retinoids  (56)  
Secreted and cell-surface-associated tumor-inhibiting proteins   
IGFBP-3 (IGF-binding protein; cytostatic; apoptogenic) HeLa cervical ca/bromodeoxyuridine; MCF-7 breast ca/retinoids (48, 56) 
IGFBP-6 (IGF-binding protein; cytostatic) HCT116 colon ca/doxorubicin  (49)  
IGFBP-4 (IGF-binding protein; cytostatic) HeLa cervical ca/bromodeoxyuridine  (48)  
Maspin (serine protease inhibitor; inhibits invasion and angiogenesis; tumor suppressor) HCT116 colon ca/doxorubicin  (49)  
MIC-1 (TGF-β family) HCT116 colon ca/doxorubicin  (49)  
βig-H3 (ECM protein; inhibits tumorigenesis) HeLa cervical ca/bromodeoxyuridine; MCF-7 breast ca/retinoids (48, 56) 
Secreted and cell-surface-associated tumor-promoting proteins   
TGFα (mitogen; EGFR ligand) HCT116 colon ca/doxorubicin  (49)  
Cyr61 (angiogenic; mitogenic) HCT116 colon ca/doxorubicin  (49)  
CTGF (Cyr61-related; angiogenic; mitogenic) HT1080 fibrosarcoma/p21  (4)  
Epithelin/granulin (mitogenic) HT1080 fibrosarcoma/p21  (4)  
Galectin-3 (antiapoptotic) HT1080 fibrosarcoma/p21  (4)  
Prosaposin (antiapoptotic) HT1080 fibrosarcoma/p21 (4, 49) 
 HCT116 colon ca/doxorubicin  
 βAPP (Alzheimer’s β amyloid precursor; mitogenic) HT1080 fibrosarcoma/p21 (4, 49) 
 HCT116 colon ca/doxorubicin  
Cathepsin B (serine protease; promotes invasion) HT1080 fibrosarcoma/p21  (4)  
Plasminogen activator inhibitor-1 (serine protease inhibitor; promotes angiogenesis and invasion) HeLa cervical ca/bromodeoxyuridine; HT1080 fibrosarcoma/p21 (4, 48) 
Dual-function growth regulators   
Amphiregulin (EGF family; mitogenic in normal cells; antimitogenic in tumor cells) HCT116 colon ca/doxorubicin  (49)  
Activin A (TGF-β family) HT1080 fibrosarcoma/p21  (4)  
CD44 (cell adhesion molecule; pleiotropic growth modulator) HCT116 colon ca/doxorubicin  (49)  
Jagged-1 (Notch receptor; stem cell growth regulator; angiogenic) HCT116 colon ca/doxorubicin  (49)  
a

ca, carcinoma; EGF, epidermal growth factor; EGFR, EGF receptor.

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