Note | PPP6R3 is involved in many and heterogeneous genomic translocations in different kind of tumors (Table. 3) and it has been proposed that this happens because it possess a potent promotor activity (Guo et al., 2016). Name | 5' end | 3' end | Loc1 | Loc2 | Description | Type | Disease | Organ | Code | Ref. | C11orf80/PPP6R3 | C11orf80 | PPP6R3 | 11q13.2 | 11q13.2 | t(11;11)(q13;q13) | Fusion gene | Adenocarcinoma | Breast | BRCA | Hu et al., 2018 | CCDC132/PPP6R3 | CCDC132 | PPP6R3 | 7q21.2 | 11q13.2 | t(7;11)(q21;q13) | Translocation | Adenocarcinoma | Lung | LUAD | - | CTTN/PPP6R3 | CTTN | PPP6R3 | 17q23.3 | 11q13.2 | t(11;11)(q13;q13) | Fusion gene | Malignant melanoma | Skin | SKCM | Hu et al., 2018 | CSHL1/PPP6R3 | CSHL1 | PPP6R3 | 17q23.3 | 11q13.2 | t(11:17)(q13;q23) | Translocation | (?) | (?) | (?) | - | DPP9/PPP6R3 | DPP9 | PPP6R3 | 19p13.3 | 11q13.2 | t(11;19)(q13;p13) | Translocation | Adenocarcinoma | Ovary | OVCA | Smebye et al., 2017 | EEF1G/PPP6R3 | EEF1G | PPP6R3 | 11q12.3 | 11q13.2 | t(11;11)(q12;q13) | Fusion gene | Squamous cell carcinoma | Lung | LUSC | Hammerman et al., 2012 | ITGB5 /PPP6R3 | ITGB5 | PPP6R3 | 3q21.2 | 11q13.2 | t(3;11)(q21;q13) | Translocation | (?) | (?) | (?) | - | KDM2A/PPP6R3 | KDM2A | PPP6R3 | 11q13.2 | 11q13.2 | t(11;11)(q13;q13) | Fusion gene | Adenocarcinoma | Breast | BRCA | Yoshihara et al., 2015 | KRT5/PPP6R3 | KRT5 | PPP6R3 | 12q13.13 | 11q13.2 | t(11;12)(q13;q13) | Translocation | (?) | (?) | (?) | - | MKL2/PPP6R3 | MKL2 | PPP6R3 | 16p13.12 | 11q13.2 | t(11;16)(q13;p13) | Translocation | (?) | (?) | (?) | - | NADSYN1/PPP6R3 | NADSYN1 | PPP6R3 | 11q13.4 | 11q13.2 | t(11;11)(q13;q13) | Fusion gene | Malignant melanoma | Skin | SKCM | Hu et al., 2018; Klijn et al., 2015 | PC/PPP6R3 | PC | PPP6R3 | 11q13.2 | 11q13.2 | t(11;11)(q13;q13) | Fusion gene | Adenocarcinoma | Breast | BRCA | Yoshihara et al., 2015 | PPP6R3/ACER3 | PPP6R3 | ACER3 | 11q13.2 | 11q13.5 | t(11;11)(q13;q13) | Fusion gene | Malignant melanoma | Skin | SKCM | Hu et al., 2018 | PPP6R3/ARHGAP1 | PPP6R3 | ARHGAP1 | 11q13.2 | 11p11.2 | t(11;11)(p11;q13) | Fusion gene | Squamous cell carcinoma | Lung | LUSC | Yoshihara et al., 2015 | PPP6R3/ATP6V0D2 | PPP6R3 | ATP6V0D2 | 11q13.2 | 8q21.3 | t(8;11)(q21;q13) | Translocation | Sarcoma | Soft tissue | SARC | - | PPP6R3/CNTN5 | PPP6R3 | CNTN5 | 11q13.2 | 11q22.1 | t(11;11)(q13;q22) | Fusion gene | Squamous cell carcinoma | Lung | LUSC | Hammerman et al., 2012 | PPP6R3/DPP3 | PPP6R3 | DPP3 | 11q13.2 | 11q13.2 | t(11;11)(q13;q13) | Fusion gene | Germ cell tumor, NOS | Testis | TGCT | Hoff et al., 2016 | PPP6R3/FADD | PPP6R3 | FADD | 11q13.2 | 11q13.3 | t(11;11)(q13;q13) | Fusion gene | Adenocarcinoma | Breast | BRCA | Yoshihara et al., 2015 | PPP6R3/EIF4G3 | PPP6R3 | EIF4G3 | 11q13.2 | 1p36.12 | t(1;11)(p36;q13) | Translocation | Malignant melanoma | Skin | SKCM | - | PPP6R3/INTS4 | PPP6R3 | INTS4 | 11q13.2 | 11q14.1 | t(11;11)(q13;q14) | Fusion gene | Acute lymphoblastic leukemia/lymphoblastic lymphoma (T-Lineage) | Blood | T-ALL | Liu et al., 2017 | PPP6R3/LRP5 | PPP6R3 | LRP5 | 11q13.2 | 11q13.2 | t(11;11)(q13;q13) | Fusion gene | Squamous cell carcinoma | Lung | LUSC | Yoshihara et al., 2015 | Adenocarcinoma | Pancreas | PAAD | - | PPP6R3/MGAM | PPP6R3 | MGAM | 11q13.2 | 7q34 | t(7;11)(q34;q13) | Translocation | Squamous cell carcinoma | Lung | LUSC | - | PPP6R3/MTL5 | PPP6R3 | MTL5 | 11q13.2 | 11q13.3 | t(11;11)(q13;q13) | Fusion gene | Adenocarcinoma | Breast | BRCA | Yoshihara et al., 2015 | PPP6R3/POLD3 | PPP6R3 | POLD3 | 11q13.2 | 11q13.4 | t(11;11)(q13;q13) | Fusion gene | Adenocarcinoma | Breast | BRCA | Yoshihara et al., 2015 | PPP6R3/PREPL | PPP6R3 | PREPL | 11q13.2 | 2p21 | t(2;11)(p21;q13) | Translocation | (?) | (?) | (?) | - | PPP6R3/RHOD | PPP6R3 | RHOD | 11q13.2 | 11q13.2 | t(11;11)(q13;q13) | Fusion gene | Adenocarcinoma | Stomach | STAD | Hu et al., 2018 | PPP6R3/SHANK2 | PPP6R3 | SHANK2 | 11q13.2 | 11q13.3 | t(11;11)(q13;q13) | Fusion gene | Transitional cell carcinoma | Bladder | BLCA | Hu et al., 2018 | PPP6R3/SPTBN2 | PPP6R3 | SPTBN2 | 11q13.2 | 11q13.2 | t(11;11)(q13;q13) | Fusion gene | Adenocarcinoma | Breast | BRCA | Yoshihara et al., 2015 | PPP6R3/SSH2 | PPP6R3 | SSH2 | 11q13.2 | 17q11.2 | t(11;17)(q13;q11) | Translocation | Adenocarcinoma | Breast | BRCA | Yoshihara et al., 2015 | Neuroblastoma | Soft tissue | NBL | McRobert et al., 1992 | Acute lymphoblastic leukemia/lymphoblastic lymphoma | Blood | ALL | Schmiegelow et al., 2012 | PPP6R3/SSH3 | PPP6R3 | SSH3 | 11q13.2 | 11q13.2 | t(11;11)(q13;q13) | Fusion gene | Adenocarcinoma | Pancreas | PAAD | Hu et al., 2018 | PPP6R3/USP6 | PPP6R3 | USP6 | 11q13.2 | 17p13.2 | t(11;17)(q13;p13) | Translocation | Nodular fasciitis | Soft tissue | NF | Guo et al., 2016 | Hairy cell leukemia | Blood | HCL | Nacheva et al., 1992 | Multiple myeloma | Blood | MM | Sawyer et al., 1995; Sawyer et al., 2014 | Chronic myeloid leukemia | Blood | CML | Stopera et al., 1990 | PPP6R3/YAP1 | PPP6R3 | YAP1 | 11q13.2 | 11q22.1 | t(11;11)(q13;q22) | Fusion gene | Transitional cell carcinoma | Bladder | BLCA | Yoshihara et al., 2015 | RAB11A/PPP6R3 | RAB11A | PPP6R3 | 15q22.31 | 11q13.2 | t(11;15)(q13;q22) | Translocation | Cell carcinoma and endocervical adenocarcinoma | Cervical | CESC | - | RNF121/PPP6R3 | RNF121 | PPP6R3 | 11q13.4 | 11q13.2 | t(11;11)(q13;q13) | Fusion gene | Adenocarcinoma | Breast | BRCA | Yoshihara et al., 2015 | SH3RF1/PPP6R3 | SH3RF1 | PPP6R3 | 4q32.3 | 11q13.2 | t(4;11)(q32;q13) | Translocation | (?) | (?) | (?) | - | SMAD2/PPP6R3 | SMAD2 | PPP6R3 | 18q21.1 | 11q13.2 | t(11;18)(q13;q21) | Translocation | (?) | (?) | (?) | - | SSBP1/PPP6R3 | SSBP1 | PPP6R3 | 7q34 | 11q13.2 | t(7;11)(q34;q13) | Translocation | Squamous cell carcinoma | Lung | LUSC | - | STRN3/PPP6R3 | STRN3 | PPP6R3 | 14q12 | 11q13.2 | t(11;14)(q13;q12) | Translocation | Adenocarcinoma | Breast | BRCA | Yoshihara et al., 2015 | TMPRSS6/PPP6R3 | TMPRSS6 | PPP6R3 | 22q12.3 | 11q13.2 | t(11;22)(q13;q12) | Translocation | Liver hepatocellular carcinoma | Liver | LIHC | - | TNFRSF21/PPP6R3 | TNFRSF21 | PPP6R3 | 6p12.3 | 11q13.2 | t(6;11)(p12;q13) | Translocation | Adenocarcinoma | Breast | BRCA | Banerji et al., 2012 | VPS50/PPP6R3 | VPS50 | PPP6R3 | 7q21.3 | 11q13.2 | t(7;11)(q21;q13) | Translocation | (?) | (?) | (?) | - |
Table.3 PPP6R3 rearrangements: translocations and fusion genes (reworked from ps://www.ncbi.nlm.nih.gov/homologene; http://www.tumorfusions.org; https://cgap.nci.nih.gov/Chromosomes; http://quiver.archerdx.com) [ (?) ] unknown; [ - ] no reference |
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Entity | Acute lymphoblastic leukemia (ALL) |
Hybrid/Mutated Gene | The fusion gene PPP6R3/ INTS4 has been found in T-ALL (Liu et al., 2017), while a t(11;17)(q13;q11) PPP6R3/ SSH2 was discovered in a childhood T/BCP (T/ B-cell precursor) ALL (Schmiegelow et al., 2012). |
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Entity | Bladder cancer |
Hybrid/Mutated Gene | The fusion genes PPP6R3/ SHANK2 and PPP6R3/ YAP1 have been found in transitional cell carcinoma of the bladder (Hu et al., 2018; Yoshihara et al., 2015; http://www.tumorfusions.org). |
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Entity | Breast cancer |
Hybrid/Mutated Gene | Many fusion genes and abnormal translocation was discovered. C11orf80 /PPP6R3, KDM2A/PPP6R3, PC/PPP6R3, PPP6R3/ FADD, PPP6R3/ MTL5, PPP6R3/ POLD3, PPP6R3/ SPTBN2, t(11;17)(q13;q11) PPP6R3/SSH2, RNF121/PPP6R3, t(11;14)(q13;q12) STRN3/PPP6R3 and t(6;11)(p12;q13) TNFRSF21/PPP6R3 have been found in breast adenocarcinoma (Hu et al., 2018; Yoshihara et al., 2015; http://www.tumorfusions.org) |
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Entity | Cervical carcinoma |
Hybrid/Mutated Gene | A t(11;15)(q13;q22) RAB11A/PPP6R3 has been found in cervical carcinoma (http://www.tumorfusions.org). |
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Entity | Gastric cancer |
Hybrid/Mutated Gene | The fusion gene PPP6R3/ RHOD has been found in gastric adenocarcinoma (Hu et al., 2018; http://www.tumorfusions.org) |
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Entity | t(11;17)(q13;p13)PPP6R3/USP6 in hematological malignancies. |
Hybrid/Mutated Gene | A t(11;17)(q13;p13)PPP6R3/ USP6 has been found in Hairy cell leukemia, Multiple myeloma, and Chronic myeloid leukemia (Sawyer et al., 2014 ; Sawyer et al., 1995 ; Nacheva et al., 1992; Stopera et al., 1990 http://www.tumorfusions.org) |
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Entity | Hepatocellular carcinoma |
Hybrid/Mutated Gene | A t(11;22)(q13;q12) TMPRSS6/PPP6R3 has been found in hepatocellular carcinoma (http://www.tumorfusions.org). |
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Entity | Lung squamous cell carcinoma |
Hybrid/Mutated Gene | In squamous cell carcinoma of the lung was discovered the fusion gene EEF1G/PPP6R3 (Hammerman et al., 2012). Other detected are: a t(7;11)(q21;q13) CCDC132/PPP6R3, PPP6R3/ ARHGAP1 fusion gene, PPP6R3/ CNTN5 fusion gene, PPP6R3/ LRP5 fusion gene, a t(7;11)(q13;q34)PPP6R3/ MGAM and a t(7;11)(q34;q13) SSBP1/PPP6R3 (Yoshihara et al., 2015; Hammerman et al., 2012; http://www.tumorfusions.org) In squamous cell carcinoma of the lung was found the presence of the fusion gene 5' EEF1G- 3' PPP6R3 deriving by the genomic translocation and fusion of a part of EEF1G gene with a portion of PPP6R3 gene, both located on chromosome 11 (Hammerman et al., 2012). |
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Entity | Malignant melanoma |
Hybrid/Mutated Gene | The fusion genes CTTN/PPP6R3, NADSYN1/PPP6R3 and PPP6R3/ ACER3 have been found in malignant melanoma. In addition, a t(1;11)( p36;q13) PPP6R3/ EIF4G3 has also been found (Hu et al., 2018; Klijn et al., 2015; http://www.tumorfusions.org) |
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Entity | Malignant Nodular Fasciitis |
Disease | Nodular fasciitis (NF) occurs in subcutaneous tissue and it is generally a benign self-limiting myofibroblastic proliferation (Bernstein and Lattes, 1982). Sometimes was found USP6 gene rearrangements (Erickson-Johnson et al., 2011) and surprendently in one case was revealed the presence of PPP6R3/USP6 fusion gene with both genomic amplification and overexpression (Guo et al., 2016) that could explain the more aggressive behavior seen in malignant NF case report. |
Hybrid/Mutated Gene | It was described the fusion gene 5' PPP6R3 / 3' USP6 deriving by the genomic translocation and fusion of a part of PPP6R3 gene, situated on chromosome 11, with a portion of USP6 gene, instead located on chromosome 17. In particular, the breakpoint reside at the end of exon 1 of PPP6R3 (non-coding) and 155 bp upstream of coding sequence of USP6 gene (Guo et al., 2016). |
Oncogenesis | The presence of the novel PPP6R3/USP6 fusion gene was linked to the increase of malignancy of the nodular faciitis (NF). Other tumors were tested for the presence of this chimeric gene, such as aggressive angiomyxoma , malignant peripheral nerve sheath tumor, dermatofibroma/ benign fibrous histiocytoma and melanoma. In that cases the results were negatives (Guo et al., 2016). Some authors detect PPP6R3/USP6 in hairy cell leukemia, multiple myeloma and chronic myeloid leukemia (Sawyer et al., 2014; Sawyer et al., 1995; Nacheva et al., 1992 ; Stopera et al., 1990). |
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Entity | Nervous system tumors |
Disease | Malignant peripheral nerve sheath tumors (MPNSTs), also called neurofibrosarcomas, are a rare aggressive, metastatic, nerve-associated cancer type. It was found that in about 10% of this malignancy there are mutations on PPP6R3 gene that cause disrupt gene transcription (Rahrmann et al., 2013). In addition a t(8;11)(q21;q13)PPP6R3/ ATP6V0D2 is found in sarcoma (http://www.tumorfusions.org) and a t(11;17)(q13;q11)PPP6R3/SSH2 is detected in neuroblastoma (McRobert et al., 1992). |
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Entity | Ovarian carcinoma |
Disease | Ovarian malignancies, a type of gynecological cancer, show high genomic instability and structural chromosomal aberrations which are believed to be common mechanisms for both the inactivation of tumour suppressor genes and the production of fusion genes useful for cancer vitality. The precise roles and functions of these chimeric gene are not always understood (Smebye et al., 2017). The fusion gene DPP9 /PPP6R3 play a role in tumorigenesis and/or cancer progression through the loss-of-function of the DPP9 protein, a serine protease that act as a tumour suppressor and is a inducer of apoptosis. |
Hybrid/Mutated Gene | A fusion gene 5' DPP9 / 3' PPP6R3 deriving from the genomic translocation and fusion of a part of DPP9 (dipeptidyl peptidase 9) gene, situated on chromosome 19 with a portion of PPP6R3 gene, located on chromosome 11 was found in a high-grade serous ovarian carcinoma. The fusion occured between DPP9 exon 11 and PPP6R3 exon 18. DPP9 is a serine protease that act as a tumor suppressor and as a inducer of apoptosis (Smebye et al., 2017). |
Abnormal Protein | The fusion between DPP9 and PPP6R3 leads to disruption and deregulation of expression of DPP9 gene due to the introduction of a stop codon (TAG) directly after the junction that leads to the production of truncated form of DPP9 protein. This protein loss some functional domains, i.e. the peptidase and esterase-lipase domains (Smebye et al., 2017) and this causes its loss-of-function. |
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Entity | Pancreatic cancer |
Hybrid/Mutated Gene | The fusion genes PPP6R3/LRP5 and PPP6R3/ SSH3 have been found in pancreas adenocarcinoma (Hu et al., 2018; http://www.tumorfusions.org) |
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Entity | Testicular germ cell tumors |
Disease | Testicular germ cell tumors (TGCTs) are the most frequently diagnosed solid malignancies in young males and men by 15 to 44 years old and that includes two main subtypes: seminomas and non-seminomas (Hoff et al., 2016; Znaor et al., 2014). Some authors found a novel fusion gene 5' PPP6R3 - 3' DPP3 in pluripotent embryonal carcinomas (EC) cell lines (Hoff et al., 2016). |
Hybrid/Mutated Gene | The fusion gene 5' PPP6R3 / 3' DPP3 resulting by the translocation t(11;11)(q13;q13) and PPP6R3/DPP3 is a novel genomic chimeric rearrangement occurs in vitro. However more studies are need to characterize it and to elucidate its exact role in testicular germ cell tumors. |
Abnormal Protein | PPP6R3-DPP3 fusion gene encodes a protein of 198 amino acids long (Hoff et al., 2016). The functions of this chimeric protein are unknown. |
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Entity | Type 1 diabetes mellitus 4 |
Note | The IDDM4 locus has been localized to chromosome 11q13 and PPP6R3 gene is linked with type 1 diabetes mellitus 4 (Twells et al., 2001). |
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Entity | Effects on HIV-1 replication |
Note | Knockdown of PPP6R3 by short interfering RNAs (siRNA) library screening inhibits HIV-1 replication in cultured Jurkat T-cells (Yeung et al., 2009). |
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