Affinage

Showing ASZ1 — GASZ is a alias.

ASZ1

Ankyrin repeat, SAM and basic leucine zipper domain-containing protein 1 · UniProt Q8WWH4

Length
475 aa
Mass
53.5 kDa
Annotated
2026-06-09
100 papers in source corpus 5 papers cited in narrative 5 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 5/5 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

ASZ1 (GASZ) is a germ cell-specific protein essential for piRNA biogenesis, transposon silencing, and fertility (PMID:19730684). It localizes to the outer mitochondrial membrane through a functional mitochondrial targeting signal and self-associates there, promoting mitofusin-dependent mitochondrial fusion and clustering; this mitochondrial localization is required for nuage (intermitochondrial cement) formation, transposon repression, and spermatogenesis (PMID:26711429). Within nuage, ASZ1 co-localizes with and stabilizes the piRNA pathway protein MILI, and its loss causes MILI downregulation, global piRNA depletion, retrotransposon derepression and hypomethylation, and a zygotene-pachytene spermatocyte arrest (PMID:19730684). Mechanistically, ASZ1 acts as a mitochondrial anchoring platform: together with its binding partner Daedalus it recruits and retains the piRNA biogenesis factor Armitage near the Zucchini processing machinery, making it essential for Zucchini-mediated piRNA production (PMID:31123065). The protein architecture comprises ankyrin repeats, a SAM domain, and a bZIP domain, consistent with a scaffolding role in protein-protein interactions (PMID:12040005).

Mechanistic history

Synthesis pass · year-by-year structured walk · 5 steps
  1. 2002 Medium

    Established ASZ1/GASZ as a previously unknown germ cell-specific cytoplasmic protein with a domain architecture (ankyrin/SAM/bZIP) suited to mediating protein-protein interactions during germ cell maturation.

    Evidence In silico subtraction with Northern blot, RT-PCR, in situ hybridization, and immunohistochemistry in germ cells

    PMID:12040005

    Open questions at the time
    • No functional loss-of-function tested
    • No binding partners identified
    • Subcellular localization defined only as cytoplasmic
  2. 2004 Low

    Showed the germ cell-specific expression and cytoplasmic localization are evolutionarily conserved across vertebrates, hinting at a conserved germline function.

    Evidence cDNA cloning, Northern/Western blot, ISH and IHC across pufferfish, zebrafish, and frog

    PMID:14766731

    Open questions at the time
    • Descriptive localization only, no functional consequence established
    • Balbiani body association not mechanistically linked to any pathway
  3. 2009 High

    Defined ASZ1's essential function in the piRNA pathway by showing it stabilizes MILI in nuage and is required for transposon silencing and male fertility.

    Evidence Gasz knockout mouse with co-localization, Western blot, small RNA-seq, and bisulfite sequencing

    PMID:19730684

    Open questions at the time
    • Did not establish how ASZ1 physically stabilizes MILI
    • Subcellular anchoring mechanism not yet resolved
  4. 2015 High

    Resolved the subcellular basis of ASZ1 function, showing it targets the outer mitochondrial membrane, self-associates, drives mitofusin-dependent mitochondrial fusion, and that this localization is required for nuage formation and spermatogenesis.

    Evidence Ectopic expression with live imaging/fractionation, self co-IP, and a mitochondrial targeting signal deletion mouse model with confocal imaging and transposon assays

    PMID:26711429

    Open questions at the time
    • Direct molecular link between mitochondrial fusion and piRNA processing not detailed
    • Partners bridging the mitochondrial platform to processing enzymes not identified in this study
  5. 2019 High

    Identified the molecular mechanism by which ASZ1 organizes piRNA biogenesis: with Daedalus it forms a mitochondrial anchoring platform recruiting Armitage to the Zucchini processing machinery.

    Evidence Reciprocal co-IP, Drosophila genetic loss-of-function, immunofluorescence localization, and small RNA sequencing

    PMID:31123065

    Open questions at the time
    • A direct mammalian counterpart of the Gasz-Daedalus-Armitage axis not confirmed in mouse
    • Structural basis of the anchoring platform unknown

Open questions

Synthesis pass · forward-looking unresolved questions
  • How the ASZ1 mitochondrial scaffold mechanistically couples mitochondrial fusion to nuage assembly and piRNA precursor presentation at the molecular and structural level remains unresolved.
  • No structural model of the ASZ1 anchoring platform
  • Mechanism by which mitochondrial fusion supports piRNA processing unclear

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0060090 molecular adaptor activity 2
Localization
GO:0005739 mitochondrion 2 GO:0005829 cytosol 1
Pathway
R-HSA-1474165 Reproduction 2 R-HSA-8953854 Metabolism of RNA 2
Partners
ARMITAGEDAEDALUSMFNMILI
Complex memberships
intermitochondrial cement (nuage)

Evidence

Reading pass · 5 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2002 GASZ (ASZ1) was identified as a novel germ cell-specific protein containing four ankyrin repeats, a sterile-alpha motif (SAM), and a basic leucine zipper (bZIP) domain, localizing to the cytoplasm of pachytene spermatocytes, early spermatids, oocytes, and early preimplantation embryos, consistent with a role as a cytoplasmic signal transducer mediating protein-protein interactions during germ cell maturation. In silico subtraction, Northern blot, RT-PCR, in situ hybridization, immunohistochemistry Molecular endocrinology (Baltimore, Md.) Medium 12040005
2004 GASZ (ASZ1) orthologs are evolutionarily conserved across vertebrates (pufferfish, zebrafish, frog), retaining germ cell-specific expression and cytoplasmic localization in pachytene spermatocytes and oocytes; in frog oocytes, GASZ protein localizes to a cytoplasmic structure resembling the Balbiani body. cDNA cloning, Northern blot, Western blot, in situ hybridization, immunohistochemistry Biology of reproduction Low 14766731
2009 GASZ (ASZ1) co-localizes with MILI in intermitochondrial cement (nuage) in male germ cells; knockout of Gasz in mice causes dramatic downregulation of MILI protein levels, phenocopies the zygotene-pachytene spermatocyte block and male sterility of MILI-null mice, and results in increased retrotransposon expression, hypomethylation of retrotransposons, and global downregulation of piRNAs (repeat-associated, known, and novel), establishing an essential structural role for GASZ in stabilizing MILI in nuage. Co-localization by immunofluorescence, Gasz knockout mouse generation, Western blot, small RNA sequencing, bisulfite sequencing, RT-PCR PLoS genetics High 19730684
2015 GASZ (ASZ1) contains a functional mitochondrial targeting signal and localizes predominantly to mitochondria in germ cells endogenously and in somatic cells when ectopically expressed; GASZ interacts with itself at the outer mitochondrial membrane and promotes mitofusion in a mitofusin/MFN-dependent manner; deletion of the mitochondrial targeting signal in mice reveals that mitochondrial localization of GASZ is essential for nuage formation, mitochondrial clustering, transposon repression, and spermatogenesis. Ectopic expression with live imaging/fractionation, co-immunoprecipitation (self-interaction), mitochondrial targeting signal deletion mouse model, confocal microscopy, transposon expression assays EMBO reports High 26711429
2019 Drosophila Gasz (ortholog of ASZ1) interacts with Daedalus (Daed) on the outer mitochondrial membrane and together they form a mitochondrial anchoring platform that recruits and retains the piRNA biogenesis factor Armitage (Armi) proximal to Zucchini for piRNA processing; Gasz is essential for Zucchini-mediated piRNA production and correct localization of Armi. Co-immunoprecipitation, genetic loss-of-function (Drosophila), immunofluorescence localization, small RNA sequencing Genes & development High 31123065

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
1995 Targeting of adenovirus E1A and E4-ORF3 proteins to nuclear matrix-associated PML bodies. The Journal of cell biology 248 7559785
1999 Functional assignment of the ORF2-iscS-iscU-iscA-hscB-hscA-fdx-ORF3 gene cluster involved in the assembly of Fe-S clusters in Escherichia coli. Journal of biochemistry 228 10544286
2006 A bicistronic subgenomic mRNA encodes both the ORF2 and ORF3 proteins of hepatitis E virus. Journal of virology 220 16731930
1997 The ORF3 protein of hepatitis E virus is a phosphoprotein that associates with the cytoskeleton. Journal of virology 207 9371561
2017 Hepatitis E virus ORF3 is a functional ion channel required for release of infectious particles. Proceedings of the National Academy of Sciences of the United States of America 192 28096411
2009 ORF3 protein of hepatitis E virus is essential for virion release from infected cells. The Journal of general virology 190 19339479
1999 Hyperproduction of recombinant ferredoxins in escherichia coli by coexpression of the ORF1-ORF2-iscS-iscU-iscA-hscB-hs cA-fdx-ORF3 gene cluster. Journal of biochemistry 173 10393315
1990 Cytochrome oxidase assembly in yeast requires the product of COX11, a homolog of the P. denitrificans protein encoded by ORF3. The EMBO journal 154 2167832
2009 GASZ is essential for male meiosis and suppression of retrotransposon expression in the male germline. PLoS genetics 149 19730684
2012 PEDV ORF3 encodes an ion channel protein and regulates virus production. FEBS letters 146 22245155
2008 Monoclonal antibodies raised against the ORF3 protein of hepatitis E virus (HEV) can capture HEV particles in culture supernatant and serum but not those in feces. Archives of virology 144 18679765
2010 A PSAP motif in the ORF3 protein of hepatitis E virus is necessary for virion release from infected cells. The Journal of general virology 139 21068219
2006 The ORF3 protein of porcine circovirus type 2 is involved in viral pathogenesis in vivo. Journal of virology 136 16641298
2005 Relocalization of the Mre11-Rad50-Nbs1 complex by the adenovirus E4 ORF3 protein is required for viral replication. Journal of virology 126 15858005
2001 The ORF3 protein of hepatitis E virus binds to Src homology 3 domains and activates MAPK. The Journal of biological chemistry 121 11518702
2002 The phosphorylated form of the ORF3 protein of hepatitis E virus interacts with its non-glycosylated form of the major capsid protein, ORF2. The Journal of biological chemistry 104 11934888
1997 Baculovirus expression of proteins of porcine reproductive and respiratory syndrome virus strain Olot/91. Involvement of ORF3 and ORF5 proteins in protection. Virus genes 104 9208452
2006 ORF3 protein of hepatitis E virus is not required for replication, virion assembly, or infection of hepatoma cells in vitro. Journal of virology 102 16928762
1999 The adenovirus type 5 E1b 55K and E4 Orf3 proteins associate in infected cells and affect ND10 components. The Journal of general virology 93 10211970
2008 The hepatitis E virus ORF3 protein modulates epidermal growth factor receptor trafficking, STAT3 translocation, and the acute-phase response. Journal of virology 86 18448545
2007 Cloning and further sequence analysis of the ORF3 gene of wild- and attenuated-type porcine epidemic diarrhea viruses. Virus genes 86 17932736
2005 Enteropathogenic Escherichia coli EspG disrupts microtubules and in conjunction with Orf3 enhances perturbation of the tight junction barrier. Molecular microbiology 82 15813736
2000 Identification and genomic mapping of the ORF3 and VPg proteins in feline calicivirus virions. Virology 81 11062050
2018 Palmitoylation mediates membrane association of hepatitis E virus ORF3 protein and is required for infectious particle secretion. PLoS pathogens 80 30532200
2008 Cellular proteins PML and Daxx mediate an innate antiviral defense antagonized by the adenovirus E4 ORF3 protein. Journal of virology 80 18480450
2003 Distinct roles of the Adenovirus E4 ORF3 protein in viral DNA replication and inhibition of genome concatenation. Journal of virology 76 12692231
2003 Pseudomonas syringae pv. tomato DC3000 HopPtoM (CEL ORF3) is important for lesion formation but not growth in tomato and is secreted and translocated by the Hrp type III secretion system in a chaperone-dependent manner. Molecular microbiology 75 12940984
2007 The ORF3 protein of porcine circovirus type 2 interacts with porcine ubiquitin E3 ligase Pirh2 and facilitates p53 expression in viral infection. Journal of virology 73 17581998
2019 Porcine epidemic diarrhea virus ORF3 protein causes endoplasmic reticulum stress to facilitate autophagy. Veterinary microbiology 70 31383304
2002 Identification of Gasz, an evolutionarily conserved gene expressed exclusively in germ cells and encoding a protein with four ankyrin repeats, a sterile-alpha motif, and a basic leucine zipper. Molecular endocrinology (Baltimore, Md.) 70 12040005
2007 Adenovirus E4 ORF3 protein inhibits the interferon-mediated antiviral response. Journal of virology 69 17301128
2007 The hepatitis E virus Orf3 protein protects cells from mitochondrial depolarization and death. The Journal of biological chemistry 66 17488721
2020 Porcine Epidemic Diarrhea Virus (PEDV) ORF3 Enhances Viral Proliferation by Inhibiting Apoptosis of Infected Cells. Viruses 65 32075094
2004 The ORF3 protein of hepatitis E virus interacts with liver-specific alpha1-microglobulin and its precursor alpha1-microglobulin/bikunin precursor (AMBP) and expedites their export from the hepatocyte. The Journal of biological chemistry 65 15037615
1996 Detection of the ORF3 polypeptide of feline calicivirus in infected cells and evidence for its expression from a single, functionally bicistronic, subgenomic mRNA. The Journal of general virology 65 8558120
2014 Enhancement of interferon induction by ORF3 product of hepatitis E virus. Journal of virology 59 24850742
2006 Enhanced alpha1 microglobulin secretion from Hepatitis E virus ORF3-expressing human hepatoma cells is mediated by the tumor susceptibility gene 101. The Journal of biological chemistry 57 16407257
2006 Interaction of the adenovirus type 5 E4 Orf3 protein with promyelocytic leukemia protein isoform II is required for ND10 disruption. Journal of virology 55 16501113
1999 Roles for the E4 orf6, orf3, and E1B 55-kilodalton proteins in cell cycle-independent adenovirus replication. Journal of virology 55 10438837
1989 Analysis of the cis-acting requirements for germ-line-specific splicing of the P-element ORF2-ORF3 intron. Genes & development 55 2545526
2013 Genetic diversity of ORF3 and spike genes of porcine epidemic diarrhea virus in Thailand. Infection, genetics and evolution : journal of molecular epidemiology and evolutionary genetics in infectious diseases 54 24291225
2015 GASZ and mitofusin-mediated mitochondrial functions are crucial for spermatogenesis. EMBO reports 53 26711429
2009 The hepatitis E virus ORF3 protein stabilizes HIF-1alpha and enhances HIF-1-mediated transcriptional activity through p300/CBP. Cellular microbiology 53 19489936
2016 The ORF3 Protein of Genotype 1 Hepatitis E Virus Suppresses TLR3-induced NF-κB Signaling via TRADD and RIP1. Scientific reports 51 27270888
2015 Porcine epidemic diarrhea virus ORF3 gene prolongs S-phase, facilitates formation of vesicles and promotes the proliferation of attenuated PEDV. Virus genes 49 26531166
2007 T-cell epitope mapping of ORF2 and ORF3 proteins of human hepatitis E virus. Journal of viral hepatitis 47 17381721
2012 The PSAP motif within the ORF3 protein of an avian strain of the hepatitis E virus is not critical for viral infectivity in vivo but plays a role in virus release. Journal of virology 46 22438540
2010 The ORF3 protein of hepatitis E virus delays degradation of activated growth factor receptors by interacting with CIN85 and blocking formation of the Cbl-CIN85 complex. Journal of virology 46 20130058
2012 ORF3 of duck circovirus: a novel protein with apoptotic activity. Veterinary microbiology 45 22537707
2009 The open reading frame 3 (ORF3) of porcine circovirus type 2 (PCV2) is dispensable for virus infection but evidence of reduced pathogenicity is limited in pigs infected by an ORF3-null PCV2 mutant. Virus research 45 19852989
2016 Molecular Characterization of the ORF3 and S1 Genes of Porcine Epidemic Diarrhea Virus Non S-INDEL Strains in Seven Regions of China, 2015. PloS one 44 27494026
1998 A nonstructural and antigenic glycoprotein is encoded by ORF3 of the IAF-Klop strain of porcine reproductive and respiratory syndrome virus. Archives of virology 43 9856081
2018 The Accessory Protein ORF3 Contributes to Porcine Epidemic Diarrhea Virus Replication by Direct Binding to the Spike Protein. Viruses 42 30060558
2005 The 41-amino-acid C-terminal region of the hepatitis E virus ORF3 protein interacts with bikunin, a kunitz-type serine protease inhibitor. Journal of virology 40 16140784
2019 Daedalus and Gasz recruit Armitage to mitochondria, bringing piRNA precursors to the biogenesis machinery. Genes & development 38 31123065
2010 ORF3 of porcine circovirus 2 enhances the in vitro and in vivo spread of the of the virus. Virology 38 21144544
1999 Characterization of Moraxella (Branhamella) catarrhalis lbpB, lbpA, and lactoferrin receptor orf3 isogenic mutants. Infection and immunity 38 10024604
2015 Replacement of the hepatitis E virus ORF3 protein PxxP motif with heterologous late domain motifs affects virus release via interaction with TSG101. Virology 36 26457367
2011 The hepatitis E virus ORF3 protein regulates the expression of liver-specific genes by modulating localization of hepatocyte nuclear factor 4. PloS one 36 21799848
2019 Porcine Circovirus Type 2 Induces ORF3-Independent Mitochondrial Apoptosis via PERK Activation and Elevation of Cytosolic Calcium. Journal of virology 35 30651358
2012 Herpesvirus saimiri antagonizes nuclear domain 10-instituted intrinsic immunity via an ORF3-mediated selective degradation of cellular protein Sp100. Journal of virology 35 22278248
2008 The ORF3 protein of hepatitis E virus interacts with hemopexin by means of its 26 amino acid N-terminal hydrophobic domain II. Biochemistry 35 18211098
2001 Self-association and mapping of the interaction domain of hepatitis E virus ORF3 protein. Journal of virology 35 11160756
2016 Detection and phylogenetic analysis of porcine epidemic diarrhea virus in central China based on the ORF3 gene and the S1 gene. Virology journal 34 27887624
2008 Attenuation of porcine circovirus 2 in SPF piglets by abrogation of ORF3 function. Virology 33 19012942
2009 Porcine circovirus type 2 ORF3 protein competes with p53 in binding to Pirh2 and mediates the deregulation of p53 homeostasis. Virology 31 20004925
2013 Phylogenetic analysis of porcine epidemic diarrhea virus (PEDV) field strains in central China based on the ORF3 gene and the main neutralization epitopes. Archives of virology 30 24292967
1992 Identification of the three-dimensional thioredoxin motif: related structure in the ORF3 protein of the Staphylococcus aureus mer operon. Biochemistry 30 1591248
2018 HEV ORF3 downregulates TLR7 to inhibit the generation of type I interferon via impairment of multiple signaling pathways. Scientific reports 28 29872132
2020 The accessory protein ORF3 of porcine epidemic diarrhea virus inhibits cellular interleukin-6 and interleukin-8 productions by blocking the nuclear factor-κB p65 activation. Veterinary microbiology 27 33126183
2010 Cytotoxicity of ORF3 proteins from a nonpathogenic and a pathogenic porcine circovirus. Journal of virology 27 20810737
2001 Efficacy of a transmissible gastroenteritis coronavirus with an altered ORF-3 gene. Canadian journal of veterinary research = Revue canadienne de recherche veterinaire 27 11227191
1994 Sequence of the rec-2 locus of Haemophilus influenzae: homologies to comE-ORF3 of Bacillus subtilis and msbA of Escherichia coli. Gene 27 8063112
2016 Identification of critical residues in Hepatitis E virus macro domain involved in its interaction with viral methyltransferase and ORF3 proteins. Scientific reports 26 27113483
2015 The ORF3 protein of porcine circovirus type 2 promotes secretion of IL-6 and IL-8 in porcine epithelial cells by facilitating proteasomal degradation of regulator of G protein signalling 16 through physical interaction. The Journal of general virology 26 25575706
2007 The adenovirus E4 ORF3 protein binds and reorganizes the TRIM family member transcriptional intermediary factor 1 alpha. Journal of virology 26 17287283
2018 An analysis of two open reading frames (ORF3 and ORF4) of rat hepatitis E virus genome using its infectious cDNA clones with mutations in ORF3 or ORF4. Virus research 25 29471051
2016 The adenovirus E4-ORF3 protein functions as a SUMO E3 ligase for TIF-1γ sumoylation and poly-SUMO chain elongation. Proceedings of the National Academy of Sciences of the United States of America 25 27247387
2009 Varying abilities of recombinant polypeptides from different regions of hepatitis E virus ORF2 and ORF3 to detect anti-HEV immunoglobulin M. Journal of medical virology 25 19382255
1994 Cloning and sequencing of the chicken anaemia virus (CAV) ORF-3 gene, and the development of an ELISA for the detection of serum antibody to CAV. Veterinary microbiology 25 8203122
2009 ORF3 protein of hepatitis E virus interacts with the Bbeta chain of fibrinogen resulting in decreased fibrinogen secretion from HuH-7 cells. The Journal of general virology 24 19264644
2023 Coronavirus accessory protein ORF3 biology and its contribution to viral behavior and pathogenesis. iScience 23 36945252
2017 The role of ORF3 accessory protein in replication of cell-adapted porcine epidemic diarrhea virus (PEDV). Archives of virology 23 28474223
1998 Kluyveromyces lactis killer plasmid pGKL2: evidence for a viral-like capping enzyme encoded by ORF3. Plasmid 23 9806862
2021 Comparative Characterization and Pathogenicity of a Novel Porcine Epidemic Diarrhea Virus (PEDV) with a Naturally Occurring Truncated ORF3 Gene Coinfected with PEDVs Possessing an Intact ORF3 Gene in Piglets. Viruses 22 34452427
2021 The Viral ORF3 Protein Is Required for Hepatitis E Virus Apical Release and Efficient Growth in Polarized Hepatocytes and Humanized Mice. Journal of virology 22 34523963
2020 Porcine Epidemic Diarrhea Virus ORF3 Protein Is Transported through the Exocytic Pathway. Journal of virology 22 32554695
2016 The Adenovirus E4-ORF3 Protein Stimulates SUMOylation of General Transcription Factor TFII-I to Direct Proteasomal Degradation. mBio 22 26814176
2014 Evidence of MAPK-JNK1/2 activation by hepatitis E virus ORF3 protein in cultured hepatoma cells. Cytotechnology 22 25280525
2009 Immunogenicity and efficacy of a bacterially expressed HEV ORF3 peptide, assessed by experimental infection of primates. Archives of virology 22 19763777
1997 Expression and characterization of the hepatitis E virus ORF3 protein in the methylotrophic yeast, Pichia pastoris. Gene 22 9185850
2015 Adenovirus E4-ORF3 Targets PIAS3 and Together with E1B-55K Remodels SUMO Interactions in the Nucleus and at Virus Genome Replication Domains. Journal of virology 21 26223632
2014 Genetic variability and phylogeny of current Chinese porcine epidemic diarrhea virus strains based on spike, ORF3, and membrane genes. TheScientificWorldJournal 21 24578626
2013 Sequence heterogeneity of the ORF3 gene of porcine epidemic diarrhea viruses field samples in Fujian, China, 2010-2012. Viruses 21 24084234
2003 Two nonoverlapping domains on the Norwalk virus open reading frame 3 (ORF3) protein are involved in the formation of the phosphorylated 35K protein and in ORF3-capsid protein interactions. Journal of virology 21 12610132
2022 Genetic Characteristics and Pathogenicity of a Novel Porcine Epidemic Diarrhea Virus with a Naturally Occurring Truncated ORF3 Gene. Viruses 20 35336894
2020 PEDV ORF3 Independently Regulates IκB Kinase β-Mediated NF-κB and IFN-β Promoter Activities. Pathogens (Basel, Switzerland) 20 32423021
2016 ORF3 of Hepatitis E Virus Inhibits the Expression of Proinflammatory Cytokines and Chemotactic Factors in LPS-Stimulated Human PMA-THP1 Cells by Inhibiting NF-κB Pathway. Viral immunology 20 26771290
2014 Proteomic analysis of ubiquitin-like posttranslational modifications induced by the adenovirus E4-ORF3 protein. Journal of virology 20 25410875
2004 Identification and characterization of evolutionarily conserved pufferfish, zebrafish, and frog orthologs of GASZ. Biology of reproduction 19 14766731

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