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MAZ (MYC Associated Zinc Finger Protein)

Written2017-11Burcu Karakaya, Mesut Muyan
Middle East Technical University, Department of Biological Sciences, ankaya 06800, Ankara, Turkey.;

Abstract Myc-associated zinc finger protein (MAZ), also known as serum amyloid A-activating factor 1 (SAF1), Pur-1 or Zif87, is ubiquitously expressed in various tissues. MAZ is a transcription factor with six Cys2His2-type zinc finger motifs at the carboxyl-terminus that interact with a permutation of the GGGAGGG sequence motif present in GC-rich promoter regions of target genes, likely through DNA unfolding of G-quadruplex structures to modulate gene expressions. MAZ is also suggested to participate in transcription termination and polyadenylation. Deregulated expression of MAZ is reported to correlate with various tissue malignancies that include the breast, thyroid, hepatocellular and urothelial cancers.

Keywords MAZ; transcription factor; Zinc finger; DNA binding; Purine binding; breast cancer; thyroid cancer; hepatocellular cancer; urothelial cancer.

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Alias_namesMYC-associated zinc finger protein (purine-binding transcription factor)
Alias_symbol (synonym)ZF87
Other aliasPUR1 (Purine-Binding Transcription Factor)
SAF-1 (Serum Amyloid A Activating Factor 1)
SAF-2 (Serum Amyloid A Activating Factor 2)
SAF-3 (Serum Amyloid A Activating Factor 3)
ZF87 (Transcription Factor Zif87)
ZNF801 (Zinc Finger Protein 801)
LocusID (NCBI) 4150
Atlas_Id 41307
Location 16p11.2  [Link to chromosome band 16p11]
Location_base_pair Starts at 29806537 and ends at 29811183 bp from pter ( according to hg19-Feb_2009)  [Mapping MAZ.png]
  UCSC representation of the gene on chromosome 16. RefSeq sequence shows introns as lines, exons as boxes and encoding exons as thicker boxes. Retrieved from: on November 6, 2017.
Fusion genes
(updated 2017)
Data from Atlas, Mitelman, Cosmic Fusion, Fusion Cancer, TCGA fusion databases with official HUGO symbols (see references in chromosomal bands)


  The human MAZ consists of six exons, the first five of which are encoding exons; total exon length is 4.57 kb (Song et al., 1998).
Description The human MAZ contains six exons; the encoding sequence consists of 1431 bases (Song et al., 1998).
Transcription The human gene encoding for MAZ is located on chromosome 16p11.2 and is transcribed as an mRNA of 2.7 kilobases (kb). The primary transcript encodes a 477 amino-acid long MAZ-1 protein with a 60-kDa molecular mass that contains six C2H2-type zinc-finger domains responsible for DNA binding. MAZ protein has two additional isoforms: MAZ-2 and MAZ-3. The MAZ-2 transcript is generated by an alternative splicing that results in the insertion of a new exon originating from the non-coding sequences of the intron 4. This transcript gives rise to the MAZ-2 isoform, which is a 493 amino-acids long protein with distinct carboxyl-terminus which contains two additional zinc-finger domains (Ray et al., 2002). The MAZ-2 isoform is reported to have a higher DNA-binding activity and to act as a negative regulator of MAZ-1 function (Ray et al., 2002). The MAZ-3 transcript is expressed at very low levels under normal physiological condition in various tissues, but is highly expressed during inflammation. The MAZ-3 transcript is transcribed from a distinct upstream promoter and is processed with alternative splicing. The MAZ-3 transcript is translated from a different starting codon that gives rise to the MAZ-3 isoform of 455 amino-acids (Ray et al., 2009).
Pseudogene No reported pseudogenes are found.


  Domains of MAZ are depicted with vertical colored lines; Blacks are Poly-Alanine repeats; Green is Poly-Proline tract; Red column is Poly-Glycine repeat. C2H2-type zinc finger domains of MAZ-1 are represented in dark blue-green vertical lines.
Description The human MAZ protein contains three Poly alanine, one poly-proline and one poly-glycine domains (Song et al., 1998). Poly-alanine repeats considered to have role in cellular localization of the protein; the alteration in the intracellular distribution may contribute to diseases, including muscular dystrophy (OPMD) (Oma et al., 2004). Similarly, poly-glycine repeats are responsible in protein targeting (Uthayakumar et al., 2012). Poly-proline tracks, on the other hand, generates structures that are predicted to have important roles in protein-protein interactions (Williamson, 1994). The human MAZ-1 contains six C2H2-type zinc finger domains (Song et al., 1998), which are frequently occurring in proteins involved in transcriptional regulation.
  Expression and synthesis of MAZ in various cancerous tissues. Retrieved from: on November 2, 2017.
Expression MAZ is expressed in the human heart, brain, lungs, liver, skeletal muscle, pancreas, and prostate (Jiao et al., 2013; Dudas et al., 2008). The synthesis of MAZ protein is observed to occur at high levels in breast, thyroid and urothelial cancers as well as in melanoma (Ugai et al., 2001)
  Immunofluorescent staining of human cell line MCF7. Retrieved from: on November 2, 2017. Immunofluorescent staining of MCF7 cells derived from breast adenocarcinoma shows that MAZ localizes to the nucleus.
Localisation MAZ is located in the nucleus (Jordan-Sciutto et al, 2000).
Function MAZ as a transcription factor interacts with a permutation of the GGGAGGG sequence motif present in GC-rich promoter regions of target genes by unfolding of G-quadruplex structures of DNA (Cogoi et al., 2014) to activate or repress transcription. MAZ is also suggested to participate in transcription termination and polyadenylation. Several oncogenes, including MYC, HRAS, PPARG, TSG101, VEGFA, CAV1, PTHR1, NOS3, MYB, and hTER, are transcriptionally regulated by MAZ (Jun Song et al., 2001; Lee et al., 2016; Ray et al., 2002). Deregulated expression of MAZ appears to participate in the development and/or progress various tissue malignancies including the breast, thyroid, hepatocellular and urothelial cancers (Jiao et al., 2013; Dudas et al., 2008; Yu et al., 2017; Ray, 2011; Zhu et al., 2016)
Homology The human MAZ protein is conserved 100% in chimpanzee (P.troglodytes), 98.4% in mouse (M.musculus), and 98.4% in rat (R.norvegicus); with conserved DNA of 99.8%, 93.2%, and 92.7%, respectively (Retrieved from: November 2, 2017).


Note Genetic mutations are not described for MAZ.

Implicated in

Entity Prostate cancer
Note It was reported that the MAZ expression is higher in clinical prostate cancer (PCa) specimens than in benign prostatic hyperplasia (BPH) and adjacent normal tissues (Jiao et al., 2013). Moreover, the MAZ expression appears to be positively correlated with the expression of androgen receptor ( AR), which is critical for the initiation and development of androgen-dependent PCa (Jiao et al., 2013). Extending these findings, experimental studies in cell models derived from PCa indicated that MAZ is involved in the phenotypic manifestation of PCa cell models as siRNA knockdown of MAZ levels reduces cell proliferation, migration, and invasion through mechanisms involve the expression of AR (Jiao et al., 2013).
Entity Hepatocellular carcinoma
Note The expression of MAZ was reported to be upregulated in the majority (78.94%) of hepatocellular carcinoma (HCC) samples compared to normal liver samples (Dudas et al., 2008). Experimental studies using cell lines derived from HCC further suggest that MAZ-mediated regulation of PROX1, which is a transcription factor critical for the expression of a number of genes involved in hepatic metabolic functions, contributes to the progression of HCC (Dudas et al., 2008).
Entity Breast cancer
Note Based on data sets in Gene expression-based Outcome for Breast Cancer Online (GOBO,, the expression of MAZ is found to be correlated with distant metastasis-free survival (DMFS) in basal-like breast cancer (BLBC) patients and that the under-expression of MAZ is involved in the metastatic spread of BLBC (Yu et al., 2017). Based on these finding, it was suggested that MAZ plays dual roles in basal-like breast cancer (BLBC): it suppresses cancer progression but promotes cellular proliferation (Yu et al., 2017). Experimental studies using model cell lines derived from breast cancer indeed suggest that MAZ promotes cell proliferation yet it suppresses the aggressiveness of BLBC by controlling the transition toward a more mesenchymal phenotype (Yu et al., 2017; Ray, 2011).
Entity Pancreatic carcinoma
Note Based on samples from pancreatic carcinoma patients, it was reported that the expression of MAZ is significantly higher in PC tissue compared to the adjacent non-tumor tissues (Zhu et al., 2016). Moreover, it appears that the over-expression of MAZ is associated with poor prognosis of PC patients (Zhu et al., 2016).
Entity Hodgkin's Disease and Paraneoplastic Cerebellar Dysfunction
Note In neuronal cells, MAZ interacts with the Deleted in Colorectal Cancer product ( DCC), the receptor for NTN1 netrin-1 which plays a central role in axonal guidance and neuronal migration as well as survival during development. Analyses of sera from patients with HD and PCD Hodgkin's disease and paraneoplastic cerebellar degeneration indicated that patient sera contain auto-antigens directed against the MAZ-DCC complex. Based on these observations, it was speculated that auto-antigens could interfere with neuronal function resulting in neuronal degeneration (Bataller and Wade, 2002).

To be noted

Expression of the MAZ gene is found to be regulated by MIR-125B, which is suggested to affect VEGF-induced angiogenesis in glioblastoma (Vandertop et al., 2017).
MIR449A targets MAZ transcripts, the down-regulation of which is reported to contribute to glioblastoma (Chen et al., 2015; Zhao et al., 2014; Yao et al., 2015).
MIR34C is also reported to target MAZ. Decrease levels of MAZ by miR-34c are suggested to impair the integrity and increased the permeability of blood-tumor barrier (Zhao et al., 2014).


The MAZ Protein is an Autoantigen of Hodgkin's Disease and Paraneoplastic Cerebellar Dysfunction
Bataller L, Wade DF, Graus F, Rosenfeld MR, Dalmau J.
Ann Neurol. 2003 Jan;53(1):123-7.
PMID 12509857
MAZ, a zinc finger protein, binds to c-MYC and C2 gene sequences regulating transcriptional initiation and termination
Bossone, S. A., Asselin, C., Patel, A. J., & Marcu, K. B.
Proc Natl Acad Sci U S A. 1992 Aug 15;89(16):7452-6.
PMID 1502157
HRAS is silenced by two neighboring G-quadruplexes and activated by MAZ, a zinc-finger transcription factor with DNA unfolding property.
Cogoi, S., Shchekotikhin, A. E., & Xodo, L. E.
Nucleic Acids Research, 42(13), 8379-8388.
PMID 25013182
MiR-449a suppresses the epithelial- mesenchymal transition and metastasis of hepatocellular carcinoma by multiple targets.
Chen, S., Liu, B., Xu, J., Pei, X., Liao, Y., Yuan, F., & Zheng, F.
BMC Cancer 2015; 1-13.
PMID 26471185
Altered regulation of Prox1-gene-expression in liver tumors
Dudas, J., Mansuroglu, T., Moriconi, F., Haller, F., Wilting, J., Lorf, T., Füzesi L & Ramadori, G.
BMC Cancer 2008 ; 8: 92
PMID 18400094
MAZ-dependent termination between closely spaced human complement genes
Ashfield, R., Patel, A. J., Bossone, S. A., Brown, H., Campbell, R. D., Marcu, K. B., & Proudfoot, N. J.
EMBO J. 1994 Dec 1;13(23):5656-67.
PMID 7988563
The prostate cancer-up-regulated myc-associated zinc-finger protein (MAZ) modulates proliferation and metastasis through reciprocal regulation of androgen receptor.
Jiao, L., Li, Y., Shen, D., Xu, C., Wang, L., Huang, G., Chen L, Yang Y, Yang C, Yu Y & Sun, Y.
Medical Oncology 2013; 30(2)
PMID 23609189
Fetal Alz-50 clone 1 (FAC1) protein interacts with the myc-associated zinc finger protein (ZF87/MAZ) and alters its transcriptional activity.
Jordan-Sciutto, K. L., Dragich, J. M., Caltagarone, J., Hall, D. J., & Bowser, R.
Biochemistry 2000 ; 39(12): 3206-3215
PMID 10727212
A prospero-related homeodomain protein is a novel co-regulator of hepatocyte nuclear factor 4 ? that regulates the cholesterol 7?-hydroxylase gene
Kwang-Hoon Song, Tiangang Li & Chiang J. Y. L.
NIH Public Access 2007; 281(15) : 10081-10088.
PMID 16488887
Akt phosphorylates myc-associated zinc finger protein (MAZ), releases P-MAZ from the p53 promoter, and activates p53 transcription.
Lee, W. P., Lan, K. H., Li, C. P., Chao, Y., Lin, H. C., & Lee, S. D.
Cancer Letters, 2016; 375(1): 9-19
PMID 26902421
Control of VEGF Expression in Triple-Negative Breast Carcinoma Cells by Suppression of SAF-1 Transcription Factor Activity
Ray, A., Dhar, S., & Ray, B. K.
Mol Cancer Res. 2011 Aug;9(8):1030-41.
PMID 21665940
Activation of Sp1 and Its Functional Co-operation with Serum Amyloid A-activating Sequence Binding Factor in Synoviocyte Cells Trigger Synergistic Action of Interleukin-1 and Interleukin-6 in Serum Amyloid A Gene Expression
Ray, A., Schatten, H., & Ray, B. K.
J Biol Chem.1999; 274(7): 4300-4308.
PMID 9933631
SAF-2, a splice variant of SAF-1, acts as a negative regulator of transcription.
Ray, B. K., Murphy, R., Ray, P., & Ray, A.
Journal of Biological Chemistry, 277(48), 46822-46830.
PMID 12270922
Myc-associated zinc finger protein ( MAZ ) is regulated by miR-125b and mediates VEGF- induced angiogenesis in glioblastoma
Smits M, Wurdinger T, van het Hof B, Drexhage JA, Geerts D, Wesseling P, Noske DP, Vandertop WP, de Vries HE, & Reijerkerk A.
FASEB J 2012; 26(6): 2639-47
PMID 22415301
Independent Repression of a GC-rich Housekeeping Gene by Sp1 and MAZ Involves the Same cis -Elements
Song J., Ugai H., Kanazawa I, Sun, K. & Yokoyama, K. K.
J Biol Chem 2001; 276(23): 19897-19904.
PMID 11259406
Two Consecutive Zinc Fingers in Sp1 and in MAZ Are Essential for Interactions with cis-Elements.
Song, J., Ugai, H., Ogawa, K., Wang, Y., Sarai, A., Obata, Y., Kanazawa I, Sun K, Itakura, K & Yokoyama, K. K.
Journal of Biological Chemistry 2001; 276(32): 30429-30434.
PMID 11395515
Interaction of Myc-Associated Zinc Finger Protein with DCC , the Product of a Tumor-Suppressor Gene , during the Neural Differentiation of P19
Ugai, H., Li, H., Komatsu, M., Tsutsui, H., Song, J., & Shiga, T.
EC Cells 2001; 1097: 1087-1097
PMID 11527412
Myc-associated zinc finger protein ( MAZ ) is regulated by miR-125b and mediates VEGF- induced angiogenesis in glioblastoma Myc-associated zinc finger protein ( MAZ ) is regulated by miR-125b and mediates VEGF-induced angiogenesis in glioblastoma.
Vandertop, W. P., Smits, M., Wurdinger, T., Hof, B. Van, Drexhage, J. A. R., Geerts, D., Reijerkerk, A
The FASEB Journal 2012; 2639-2647.
PMID 22415301
Overexpression and clinical significance of MYC- associated zinc finger protein in pancreatic carcinoma.
X, A. Z., Luo, W., Liang, W., Tang, F., Bei, C., Ren, Y., Tan, C.
Onco Targets Ther. 2016 Dec 12;9:7493-7501.
PMID 28008270
MiR-449a exerts tumor-suppressive functions in human glioblastoma by targeting Myc-associated zinc-finger protein.
Yao, Y., Ma, J., Xue, Y., Wang, P., Li, Z., Li, Z., Hu Y, Shang X, & Liu, Y.
Molecular Oncology 2015; 9(3): 640-656.
PMID 25487955
Specific Transcriptional Pausing Activates Polyadenylation in a Coupled In Vitro System.
Yonaha, M., Proudfoot, N. J., & William, S.
Mol Cell. 1999 May;3(5):593-600.
PMID 10360175
Dual function of MAZ mediated by FOXF2 in basal-like breast cancer?: Promotion of proliferation and suppression of progression.
Yu, Z., Lun, S., He, R., Tian, H., Huang, H., Wang, Q., Li XQ, & Feng, Y.
Cancer Letters 2017; 402: 142-152.
PMID 28577976
miR-34c Regulates the Permeability of Blood - Tumor Barrier via MAZ-Mediated Expression Changes of ZO-1 , Occludin , and Claudin-5
Zhao, L., Wang, P., Liu, Y., Ma, J. U. N., & Xue, Y.
J Cell Physiol 2014;716-731.
PMID 25201524


This paper should be referenced as such :
Karakaya, B.; Muyan, M
MAZ (MYC Associated Zinc Finger Protein);
Atlas Genet Cytogenet Oncol Haematol. in press
On line version :

External links

HGNC (Hugo)MAZ   6914
Entrez_Gene (NCBI)MAZ  4150  MYC associated zinc finger protein
AliasesPUR1; Pur-1; SAF-1; SAF-2; 
SAF-3; ZF87; ZNF801; Zif87
GeneCards (Weizmann)MAZ
Ensembl hg19 (Hinxton)ENSG00000103495 [Gene_View]
Ensembl hg38 (Hinxton)ENSG00000103495 [Gene_View]  chr16:29806537-29811183 [Contig_View]  MAZ [Vega]
ICGC DataPortalENSG00000103495
TCGA cBioPortalMAZ
Genatlas (Paris)MAZ
SOURCE (Princeton)MAZ
Genetics Home Reference (NIH)MAZ
Genomic and cartography
GoldenPath hg38 (UCSC)MAZ  -     chr16:29806537-29811183 +  16p11.2   [Description]    (hg38-Dec_2013)
GoldenPath hg19 (UCSC)MAZ  -     16p11.2   [Description]    (hg19-Feb_2009)
EnsemblMAZ - 16p11.2 [CytoView hg19]  MAZ - 16p11.2 [CytoView hg38]
Mapping of homologs : NCBIMAZ [Mapview hg19]  MAZ [Mapview hg38]
Gene and transcription
Genbank (Entrez)AA588396 AB209061 AF489858 AI079560 AI360548
RefSeq transcript (Entrez)NM_001042539 NM_001276275 NM_001276276 NM_002383
RefSeq genomic (Entrez)
Consensus coding sequences : CCDS (NCBI)MAZ
Cluster EST : UnigeneHs.23650 [ NCBI ]
CGAP (NCI)Hs.23650
Alternative Splicing GalleryENSG00000103495
Gene ExpressionMAZ [ NCBI-GEO ]   MAZ [ EBI - ARRAY_EXPRESS ]   MAZ [ SEEK ]   MAZ [ MEM ]
Gene Expression Viewer (FireBrowse)MAZ [ Firebrowse - Broad ]
SOURCE (Princeton)Expression in : [Datasets]   [Normal Tissue Atlas]  [carcinoma Classsification]  [NCI60]
GenevestigatorExpression in : [tissues]  [cell-lines]  [cancer]  [perturbations]  
BioGPS (Tissue expression)4150
GTEX Portal (Tissue expression)MAZ
Human Protein AtlasENSG00000103495-MAZ [pathology]   [cell]   [tissue]
Protein : pattern, domain, 3D structure
UniProt/SwissProtP56270   [function]  [subcellular_location]  [family_and_domains]  [pathology_and_biotech]  [ptm_processing]  [expression]  [interaction]
NextProtP56270  [Sequence]  [Exons]  [Medical]  [Publications]
With graphics : InterProP56270
Splice isoforms : SwissVarP56270
Domaine pattern : Prosite (Expaxy)ZINC_FINGER_C2H2_1 (PS00028)    ZINC_FINGER_C2H2_2 (PS50157)   
Domains : Interpro (EBI)Znf_C2H2    Znf_C2H2-like    Znf_C2H2/integrase_DNA-bd   
Domain families : Pfam (Sanger)zf-C2H2 (PF00096)   
Domain families : Pfam (NCBI)pfam00096   
Domain families : Smart (EMBL)ZnF_C2H2 (SM00355)  
Conserved Domain (NCBI)MAZ
DMDM Disease mutations4150
Blocks (Seattle)MAZ
Human Protein Atlas [tissue]ENSG00000103495-MAZ [tissue]
Peptide AtlasP56270
IPIIPI01014257   IPI01009451   IPI00869267   IPI00168859   IPI00008800   
Protein Interaction databases
IntAct (EBI)P56270
Ontologies - Pathways
Ontology : AmiGORNA polymerase II core promoter sequence-specific DNA binding  RNA binding  protein binding  nucleus  regulation of transcription, DNA-templated  transcription initiation from RNA polymerase II promoter  termination of RNA polymerase II transcription  metal ion binding  
Ontology : EGO-EBIRNA polymerase II core promoter sequence-specific DNA binding  RNA binding  protein binding  nucleus  regulation of transcription, DNA-templated  transcription initiation from RNA polymerase II promoter  termination of RNA polymerase II transcription  metal ion binding  
NDEx NetworkMAZ
Atlas of Cancer Signalling NetworkMAZ
Wikipedia pathwaysMAZ
Orthology - Evolution
GeneTree (enSembl)ENSG00000103495
Phylogenetic Trees/Animal Genes : TreeFamMAZ
Homologs : HomoloGeneMAZ
Homology/Alignments : Family Browser (UCSC)MAZ
Gene fusions - Rearrangements
Fusion : QuiverMAZ
Polymorphisms : SNP and Copy number variants
NCBI Variation ViewerMAZ [hg38]
dbSNP Single Nucleotide Polymorphism (NCBI)MAZ
Exome Variant ServerMAZ
ExAC (Exome Aggregation Consortium)ENSG00000103495
GNOMAD BrowserENSG00000103495
Genetic variants : HAPMAP4150
Genomic Variants (DGV)MAZ [DGVbeta]
DECIPHERMAZ [patients]   [syndromes]   [variants]   [genes]  
CONAN: Copy Number AnalysisMAZ 
ICGC Data PortalMAZ 
TCGA Data PortalMAZ 
Broad Tumor PortalMAZ
OASIS PortalMAZ [ Somatic mutations - Copy number]
Somatic Mutations in Cancer : COSMICMAZ  [overview]  [genome browser]  [tissue]  [distribution]  
Mutations and Diseases : HGMDMAZ
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 MAZ
DgiDB (Drug Gene Interaction Database)MAZ
DoCM (Curated mutations)MAZ (select the gene name)
CIViC (Clinical Interpretations of Variants in Cancer)MAZ (select a term)
NCG5 (London)MAZ
Cancer3DMAZ(select the gene name)
Impact of mutations[PolyPhen2] [SIFT Human Coding SNP] [Buck Institute : MutDB] [Mutation Assessor] [Mutanalyser]
Genetic Testing Registry MAZ
NextProtP56270 [Medical]
Target ValidationMAZ
Huge Navigator MAZ [HugePedia]
snp3D : Map Gene to Disease4150
BioCentury BCIQMAZ
Clinical trials, drugs, therapy
Chemical/Protein Interactions : CTD4150
Chemical/Pharm GKB GenePA30657
Clinical trialMAZ
canSAR (ICR)MAZ (select the gene name)
PubMed48 Pubmed reference(s) in Entrez
GeneRIFsGene References Into Functions (Entrez)
REVIEW articlesautomatic search in PubMed
Last year publicationsautomatic search in PubMed

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