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LDOC1 (leucine zipper, down-regulated in cancer 1)

Written2014-12Jenn-Ren Hsiao; Jang-Yang Chang
Department of Otolaryngology, Head, Neck Collaborative Oncology Group, National Cheng Kung University Hospital, College of Medicine, National Cheng Kung University, Tainan, Taiwan; hsiaojr@mail.ncku.edu.tw (JRH); Department of Medicine, National Cheng Kung University Hospital, College of Medicine, National Cheng Kung University,, National Institute of Cancer Research, National Health Research Institutes, Tainan, Taiwan; E-mail: jychang@nhri.org.tw (JYC).

Abstract LDOC1 (leucine zipper, down-regulated in cancer 1) is expressed by a wide variety of normal human tissues but is downregulated in various kinds of human cancers. Epigenetic silencing by promoter hypermethylation was shown to be an important cause of LDOC1 down-regulation in oral, cervical and ovarian cancers. The normal physiological function of LDOC1 is still not clear. But the ability of LDOC1 to induce apoptosis in various kinds of human cancer cells suggests this gene may act as a tumor suppressor.

Keywords LDOC1, cancer, methylation

(Note : for Links provided by Atlas : click)

Identity

Alias_namesBCUR1
Alias_symbol (synonym)Mar7
Mart7
Other alias
HGNC (Hugo) LDOC1
LocusID (NCBI) 23641
Atlas_Id 41137
Location Xq27  [Link to chromosome band Xq27]
Location_base_pair Starts at 140269931 and ends at 140271399 bp from pter ( according to hg19-Feb_2009)  [Mapping LDOC1.png]
Local_order from centromere to telomere: SPANXB1, LDOC1, SPANXC
Fusion genes
(updated 2016)
LDOC1 (Xq27.1) / DYNC1H1 (14q32.31)

DNA/RNA

Description The LDOC1 gene includes only one exon.
Transcription A longer wild type 1376-bp transcript (Nagasaki et al., 1999), and a much shorter 165-bp splice variant LDOC1S (Duzakale et al. 2014) are reported. The cDNA sequence of wild type LDOC1 contains a 104-bp 5' untranslated region (5'-UTR), a 438-bp open reading frame (ORF), and a 834-bp 3' untranslated region (3'-UTR) including a consensus polyadenylation signal (Nagasaki et al., 1999).
Pseudogene Not reported.

Protein

Description The wild type LDOC1 is a 17 kDa protein containing 146 amino acids, with a leucine-zipper like motif at the N-terminal and a proline-rich region that similarity to an Src homolog 3 (SH3)-binding domain (Nagasaki et al., 1999). A shorter 165-bp splice variant is predicted to translate into a truncated protein of 44 amino acids that contains only the leucine-zipper region of wide type protein (Duzakale et al. 2014).
Expression LDOC1 is expressed in a wide variety of normal tissues including heart, brain, lung, skeletal muscles, kidney, pancreas, spleen, thymus, prostate, testis, ovary, stomach, small intestine, colon, adrenal medulla, thyroid, adrenal cortex, thymus (Nagasaki et al., 1999). Down-regulation of LDOC1 have been reported in various kinds of human cancer tissues, including oral cancer (Lee et al., 2013), hepatocellular carcinoma (Riordan and Dupuy, 2013), Chronic lymphocytic leukemia (Duzakale et al., 2014), prostate cancer (Camões et al., 2012), as well as in a variety of human cancer cell lines, including pancreatic cancer cell line, gastric cancer cell lines, some breast cancer cell lines (Nagasaki et al., 1999), ovarian cancer cell lines (Buchholtz et al., 2014), cervical cancer cell lines (Buchholtz et al., 2013) and oral cancer cell lines (Lee et al., 2013). Epigenetic silencing by promoter hypermethylation was shown to be an important cause of LDOC1 down-regulation in oral cancer (Lee et al., 2013), cervical cancer (Buchholtz et al., 2013), and ovarian cancer cells (Buchholtz et al., 2014).
Localisation LDOC1 localizes predominantly in the nucleus (Nagasaki et al., 1999). It may also be retained in cytoplasm through interaction with WAVE3 (Mizutani et al., 2005).
Function Most reports demonstrated that the biological function of LDOC1 is closely related to modulation of apoptosis. Forced expression of LDOC1 induced apoptosis in Jurkat lymphoma cells, K562 leukemia cells (Inoue et al., 2015), Hela cervical cancer cells (Buchholtz et al., 2013) and oral cancer cells (Lee et al., 2013). Expression of LDOC1 also enhanced TNF-? or phorbol 12-myristate 13-acetate (PMA)-induced anti-proliferative effects in pancreatic cancer cells (Nagasaki et al., 2003). A hematopoietic transcriptional factor (MZF-1) important in regulating myeloid cell differentiation and proliferation may interact with LDOC1 and enhance it apoptosis-inducing function in K562 leukemia cells (Inoue et al., 2015). LDOC1 also induced apoptosis and inhibited degradation of p53, resulting in p53 accumulation in MDCK cells. WAVE3 could interact with LDOC1 with its C-terminal verprolin homolog domain and act as a negative regulator of LDOC1. Forced expression of WAVE3 retained LDOC1 in cytoplasm and attenuated the apoptosis-inducing effect of LDOC1 (Mizutani et al., 2005). Expression of LDOC1 also significantly inhibited the NF-?B reporter activity in LDOC1-negative BxPC-3 pancreatic cancer cells (Nagasaki et al., 2003) and suppressed TNF-? induced activation of NF-?B (Lee et al., 2013). LDOC1 may also interact with CCAAT-enhancer binding protein-? (C/EBP?) (Chih et al., 2004). However, in human intrahepatic biliary epithelial cells, over expression LDOC1 increased the expression of NF-?B (p65), promoted IL-2 and TNF-? secretion and inhibited apoptosis (Song et al., 2013). LDOC1 has recently been reported as a eutherian-specific acquired gene that regulates placental endocrine function (Naruse et al., 2014).
Homology HomoloGene:32153

Implicated in

Note
  
Entity Chronic lymphocytic leukemia
Prognosis Expression of LDOC1 mRNA correlated with prognostic cytogenetic markers of chronic lymphocytic leukemia patients. In primary patients, LDOC1 mRNA-positive group of patients had significantly poorer overall survival compared to LDOC1 mRNA-negative group of patients (Duzakale et al., 2014).
  
  
Entity Lymphoma
Oncogenesis Forced expression of LDOC1 induced apoptosis in Jurkat lymphoma cell lines (Inoue et al., 2015).
  
  
Entity Leukemia
Oncogenesis Forced expression of LDOC1 induced apoptosis in K562 leukemia cell lines (Inoue et al., 2015).
  
  
Entity Ovarian cancer
Cytogenetics No gene deletion was detected in the 7 ovarian cancer cell lines.
Oncogenesis Three of the 7 ovarian cancer cell lines showed complete loss of LDOC1 transcription, which may due to hypermethylation and inactivation of LDOC1 promoter. Treatment of de-methylation agent (5-aza-2'-deoxycytidine) on ovarian cancer cells induced transcriptional reactivation of LDOC1 gene (Buchholtz et al., 2014).
  
  
Entity Cervical cancer
Cytogenetics No gene deletion was detected in the 6 cervical cancer cell lines.
Oncogenesis LDOC1 was silenced in 4 of the 6 cervical cancer cell lines. An association of promoter hypermethylation and gene silencing was noted. Reactivation of LDOC1 gene was noted in Me180 and SiHa cervical cancer cell lines by demethylation agent treatment (5-aza-2'-deoxycytidine). Re-expression of LDOC1induced apoptosis-like cell death in Hela cervical cancer cells (Buchholtz et al., 2013).
  
  
Entity Oral cancer
Oncogenesis Hypermethylation and down-regulation of LDOC1 were consistently noted in oral cancer samples and oral cancer cell lines. Reactivation of LDOC1 gene was noted in oral cancer cells by demethylation agent treatment (5-aza-2'-deoxycytidine). Re-expression of LDOC1suppressed TNF-? induced activation of NF-?B and suppressed the colony forming and in vivo tumorigenic abilities of oral cancer cells. Epigenetic silencing of LDOC1 may contribute to alcohol, betel quid and smoking-associated oral carcinogenesis (Lee et al., 2013).
  
  
Entity Prostate cancer
Oncogenesis LDOC1 was one of the two transcriptional units (LDOC1, HPANXC) mapped in a critical region of HPCX locus linked to prostate cancer susceptibility in Finland (Baffoe-Bonnie et al., 2005). In addition, compared to non-malignant prostate tissue, LDOC1 was under-expressed (1.8 fold decrease) in prostate cancer tissues, although no methylation of LDOC1 promoter was noted in prostate tumor samples (Camões et al., 2012).
  
  
Entity Hepato cellular carcinoma
Oncogenesis LDOC1 was downregulated in 60% (29/48) of hepatic cellular carcinoma samples compared to adjacent non-tumor liver tissues (Riordan and Dupuy, 2013).
  
  
Entity Pancreatic cancer
Oncogenesis Down-regulation of LDOC1 was noted in 8 of 9 pancreatic cancer cell lines except CAPAN2 cell line (Nagasaki et al., 1999). LDOC1 was expressed in normal pancreatic tissue (Nagasaki et al., 2003). Expression of LDOC1 significantly inhibited the NF-?B reporter activity but does not affect p53, AP1 and CRE-dependent reporter gene expression in LDOC1-negative BxPC-3 pancreatic cancer cell line. LDOC1 also enhanced TNF-? or phorbol 12-myristate 13-acetate (PMA)-induced anti-proliferative effects in pancreatic cancer cells (Nagasaki et al., 2003).
  
  
Entity Gastric cancer
Oncogenesis Using northern blot, down-regulation of LDOC1 was noted in 6 (100%) of 6 gastric cancer cell lines. LDOC1 was normally expressed in stomach tissue (Nagasaki et al., 1999).
  
  
Entity Lung cancer
Oncogenesis LDOC1 was differentially expressed in MASPIN-overexpression lung carcinoma cells (LC5) compared to MASPIN-knockdown LC5 cells (Liu et al., 2012).
  
  
Entity Esophageal cancer
Oncogenesis LDOC1 was down-regulated in a radio-sensitive esophageal cancer cell line (TE-11) compared to other relative radio-resistant esophageal cancer cell lines (Ogawa et al., 2008).
  
  
Entity Cutaneous malignant melanoma
Oncogenesis LDOC1 was downregulated in peripheral blood leukocytes from a cutaneous malignant melanoma patient with multiple in-transient metastases, compared to 3 normal controls (Salemi et al., 2010).
  
  
Entity Hypertension
Note LDOC1 gene was consistently influential in a three-endophenotype model for Taiwanese hypertensive males (Lynn et al., 2009).
  
  
Entity Premature ovarian failure/insufficiency (POF/POI)
Note Array comparative genomic hybridization identified a cryptic deletion region of Xq27.2 containing LDOC1 and SPANX genes in a 23-year old woman with premature ovarian failure/insufficiency (Vitek et al., 2012).
  
  
Entity Down's syndrome
Note Gene expression of LDOC1 was increased in peripheral blood leukocytes from 15 (75%) of 20 patients Down's syndrome, as compared to that of 20 age-, sex-matched normal controls (Salemi et al., 2012).
  
  
Entity Biliary atresia
Note Over expression LDOC1 in human intrahepatic biliary epithelial cells increased the expression of NF-?B (p65), promoted IL-2 and TNF-? secretion and inhibited apoptosis of human intrahepatic biliary epithelial cells, which may induce intrahepatic biliary epithelial cell hyperplasia and trigger biliary atresia, bile duct stenosis and hepatic fibrosis (Song et al., 2013).
  

Bibliography

A major locus for hereditary prostate cancer in Finland: localization by linkage disequilibrium of a haplotype in the HPCX region
Baffoe-Bonnie AB, Smith JR, Stephan DA, Schleutker J, Carpten JD, Kainu T, Gillanders EM, Matikainen M, Teslovich TM, Tammela T, Sood R, Balshem AM, Scarborough SD, Xu J, Isaacs WB, Trent JM, Kallioniemi OP, Bailey-Wilson JE
Hum Genet 2005 Aug;117(4):307-16
PMID 15906096
 
Epigenetic silencing of the LDOC1 tumor suppressor gene in ovarian cancer cells
Buchholtz ML, Brüning A, Mylonas I, Jückstock J
Arch Gynecol Obstet 2014 Jul;290(1):149-54
PMID 24554348
 
Potential downstream target genes of aberrant ETS transcription factors are differentially affected in Ewing's sarcoma and prostate carcinoma
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Exp Hematol 2004 Dec;32(12):1173-81
PMID 15588942
 
LDOC1 mRNA is differentially expressed in chronic lymphocytic leukemia and predicts overall survival in untreated patients
Duzkale H, Schweighofer CD, Coombes KR, Barron LL, Ferrajoli A, O'Brien S, Wierda WG, Pfeifer J, Majewski T, Czerniak BA, Jorgensen JL, Medeiros LJ, Freireich EJ, Keating MJ, Abruzzo LV
Blood 2011 Apr 14;117(15):4076-84
PMID 21310924
 
LDOC1, a novel MZF-1-interacting protein, induces apoptosis
Inoue M, Takahashi K, Niide O, Shibata M, Fukuzawa M, Ra C
FEBS Lett 2005 Jan 31;579(3):604-8
PMID 15670815
 
Epigenetic regulation of the X-linked tumour suppressors BEX1 and LDOC1 in oral squamous cell carcinoma
Lee CH, Wong TS, Chan JY, Lu SC, Lin P, Cheng AJ, Chen YJ, Chang JS, Hsiao SH, Leu YW, Li CI, Hsiao JR, Chang JY
J Pathol 2013 Jul;230(3):298-309
PMID 23362108
 
Comparative proteomic analysis of the function and network mechanisms of MASPIN in human lung cells
Liu Y, Geng Y, Li K, Wang F, Zhou H, Wang W, Hou J, Liu W
Exp Ther Med 2012 Mar;3(3):470-474
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A neural network model for constructing endophenotypes of common complex diseases: an application to male young-onset hypertension microarray data
Lynn KS, Li LL, Lin YJ, Wang CH, Sheng SH, Lin JH, Liao W, Hsu WL, Pan WH
Bioinformatics 2009 Apr 15;25(8):981-8
PMID 19237446
 
WAVE3 functions as a negative regulator of LDOC1
Mizutani K, Koike D, Suetsugu S, Takenawa T
J Biochem 2005 Nov;138(5):639-46
PMID 16272576
 
Leucine-zipper protein, LDOC1, inhibits NF-kappaB activation and sensitizes pancreatic cancer cells to apoptosis
Nagasaki K, Schem C, von Kaisenberg C, Biallek M, Rösel F, Jonat W, Maass N
Int J Cancer 2003 Jul 1;105(4):454-8
PMID 12712434
 
Sirh7/Ldoc1 knockout mice exhibit placental P4 overproduction and delayed parturition
Naruse M, Ono R, Irie M, Nakamura K, Furuse T, Hino T, Oda K, Kashimura M, Yamada I, Wakana S, Yokoyama M, Ishino F, Kaneko-Ishino T
Development 2014 Dec;141(24):4763-71
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Identification of candidate genes involved in the radiosensitivity of esophageal cancer cells by microarray analysis
Ogawa R, Ishiguro H, Kuwabara Y, Kimura M, Mitsui A, Mori Y, Mori R, Tomoda K, Katada T, Harada K, Fujii Y
Dis Esophagus 2008;21(4):288-97
PMID 18477249
 
Domesticated transposable element gene products in human cancer
Riordan JD, Dupuy AJ
Mob Genet Elements 2013 Sep 1;3(5):e26693
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Expression of LDOC1 mRNA in leucocytes of patients with Down's syndrome
Salemi M, Barone C, Romano C, Ridolfo F, Salluzzo R, Scillato F, Scavuzzo C, Caraci F, Calogero AE, Romano C, Bosco P
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Two proapoptotic genes are downregulated in a patient with melanoma and repeated in-transit metastases
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Overexpression of LDOC1 in human biliary epithelial cells inhibits apoptosis through NF-B signaling
Song Z, Dong R, Zhao R, Zheng S
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Xq;autosome translocation in POF: Xq27
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Citation

This paper should be referenced as such :
Hsiao JR, Chang JY
LDOC1 (leucine zipper, down-regulated in cancer 1);
Atlas Genet Cytogenet Oncol Haematol. in press
On line version : http://AtlasGeneticsOncology.org/Genes/LDOC1ID41137chXq27.html


External links

Nomenclature
HGNC (Hugo)LDOC1   6548
Cards
AtlasLDOC1ID41137chXq27
Entrez_Gene (NCBI)LDOC1  23641  leucine zipper, down-regulated in cancer 1
AliasesBCUR1; Mar7; Mart7
GeneCards (Weizmann)LDOC1
Ensembl hg19 (Hinxton)ENSG00000182195 [Gene_View]  chrX:140269931-140271399 [Contig_View]  LDOC1 [Vega]
Ensembl hg38 (Hinxton)ENSG00000182195 [Gene_View]  chrX:140269931-140271399 [Contig_View]  LDOC1 [Vega]
ICGC DataPortalENSG00000182195
TCGA cBioPortalLDOC1
AceView (NCBI)LDOC1
Genatlas (Paris)LDOC1
WikiGenes23641
SOURCE (Princeton)LDOC1
Genetics Home Reference (NIH)LDOC1
Genomic and cartography
GoldenPath hg19 (UCSC)LDOC1  -     chrX:140269931-140271399 -  Xq27   [Description]    (hg19-Feb_2009)
GoldenPath hg38 (UCSC)LDOC1  -     Xq27   [Description]    (hg38-Dec_2013)
EnsemblLDOC1 - Xq27 [CytoView hg19]  LDOC1 - Xq27 [CytoView hg38]
Mapping of homologs : NCBILDOC1 [Mapview hg19]  LDOC1 [Mapview hg38]
OMIM300402   
Gene and transcription
Genbank (Entrez)AB019527 AF086537 BC003104 CR457088 EU446786
RefSeq transcript (Entrez)NM_012317
RefSeq genomic (Entrez)NC_000023 NC_018934 NT_011786 NW_004929446
Consensus coding sequences : CCDS (NCBI)LDOC1
Cluster EST : UnigeneHs.45231 [ NCBI ]
CGAP (NCI)Hs.45231
Alternative Splicing GalleryENSG00000182195
Gene ExpressionLDOC1 [ NCBI-GEO ]   LDOC1 [ EBI - ARRAY_EXPRESS ]   LDOC1 [ SEEK ]   LDOC1 [ MEM ]
Gene Expression Viewer (FireBrowse)LDOC1 [ Firebrowse - Broad ]
SOURCE (Princeton)Expression in : [Datasets]   [Normal Tissue Atlas]  [carcinoma Classsification]  [NCI60]
GenevisibleExpression in : [tissues]  [cell-lines]  [cancer]  [perturbations]  
BioGPS (Tissue expression)23641
GTEX Portal (Tissue expression)LDOC1
Protein : pattern, domain, 3D structure
UniProt/SwissProtO95751   [function]  [subcellular_location]  [family_and_domains]  [pathology_and_biotech]  [ptm_processing]  [expression]  [interaction]
NextProtO95751  [Sequence]  [Exons]  [Medical]  [Publications]
With graphics : InterProO95751
Splice isoforms : SwissVarO95751
PhosPhoSitePlusO95751
Domains : Interpro (EBI)DUF4939    LDOC1    LDOC1-rel   
Domain families : Pfam (Sanger)DUF4939 (PF16297)   
Domain families : Pfam (NCBI)pfam16297   
Conserved Domain (NCBI)LDOC1
DMDM Disease mutations23641
Blocks (Seattle)LDOC1
SuperfamilyO95751
Human Protein AtlasENSG00000182195
Peptide AtlasO95751
HPRD02322
IPIIPI00032909   
Protein Interaction databases
DIP (DOE-UCLA)O95751
IntAct (EBI)O95751
FunCoupENSG00000182195
BioGRIDLDOC1
STRING (EMBL)LDOC1
ZODIACLDOC1
Ontologies - Pathways
QuickGOO95751
Ontology : AmiGOprotein binding  nucleus  nucleolus  negative regulation of cell proliferation  
Ontology : EGO-EBIprotein binding  nucleus  nucleolus  negative regulation of cell proliferation  
NDEx NetworkLDOC1
Atlas of Cancer Signalling NetworkLDOC1
Wikipedia pathwaysLDOC1
Orthology - Evolution
OrthoDB23641
GeneTree (enSembl)ENSG00000182195
Phylogenetic Trees/Animal Genes : TreeFamLDOC1
HOVERGENO95751
HOGENOMO95751
Homologs : HomoloGeneLDOC1
Homology/Alignments : Family Browser (UCSC)LDOC1
Gene fusions - Rearrangements
Polymorphisms : SNP and Copy number variants
NCBI Variation ViewerLDOC1 [hg38]
dbSNP Single Nucleotide Polymorphism (NCBI)LDOC1
dbVarLDOC1
ClinVarLDOC1
1000_GenomesLDOC1 
Exome Variant ServerLDOC1
ExAC (Exome Aggregation Consortium)LDOC1 (select the gene name)
Genetic variants : HAPMAP23641
Genomic Variants (DGV)LDOC1 [DGVbeta]
DECIPHER (Syndromes)X:140269931-140271399  ENSG00000182195
CONAN: Copy Number AnalysisLDOC1 
Mutations
ICGC Data PortalLDOC1 
TCGA Data PortalLDOC1 
Broad Tumor PortalLDOC1
OASIS PortalLDOC1 [ Somatic mutations - Copy number]
Somatic Mutations in Cancer : COSMICLDOC1  [overview]  [genome browser]  [tissue]  [distribution]  
Mutations and Diseases : HGMDLDOC1
LOVD (Leiden Open Variation Database)Whole genome datasets
LOVD (Leiden Open Variation Database)LOVD - Leiden Open Variation Database
LOVD (Leiden Open Variation Database)LOVD 3.0 shared installation
BioMutasearch LDOC1
DgiDB (Drug Gene Interaction Database)LDOC1
DoCM (Curated mutations)LDOC1 (select the gene name)
CIViC (Clinical Interpretations of Variants in Cancer)LDOC1 (select a term)
intoGenLDOC1
NCG5 (London)LDOC1
Cancer3DLDOC1(select the gene name)
Impact of mutations[PolyPhen2] [SIFT Human Coding SNP] [Buck Institute : MutDB] [Mutation Assessor] [Mutanalyser]
Diseases
OMIM300402   
Orphanet
MedgenLDOC1
Genetic Testing Registry LDOC1
NextProtO95751 [Medical]
TSGene23641
GENETestsLDOC1
Huge Navigator LDOC1 [HugePedia]
snp3D : Map Gene to Disease23641
BioCentury BCIQLDOC1
ClinGenLDOC1
Clinical trials, drugs, therapy
Chemical/Protein Interactions : CTD23641
Chemical/Pharm GKB GenePA30329
Clinical trialLDOC1
Miscellaneous
canSAR (ICR)LDOC1 (select the gene name)
Probes
Litterature
PubMed24 Pubmed reference(s) in Entrez
GeneRIFsGene References Into Functions (Entrez)
CoreMineLDOC1
EVEXLDOC1
GoPubMedLDOC1
iHOPLDOC1
REVIEW articlesautomatic search in PubMed
Last year publicationsautomatic search in PubMed

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indexed on : Wed Apr 12 11:34:18 CEST 2017

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