Gene Symbol: TER94
Description: TER94
Alias: 22.26, 7.12, Apple, BcDNA.GM02885, BcDNA:GM02885, CDC48, CG2331, DmTER94, Dmel\CG2331, Ter94, Tera, VCP, VCP ATPase, VCP/p97, Vcp, anon-WO2004063362.65, anon-WO2004063362.67, cdc48, dTERA, dVCP, dmTERA, dvcp, l(2)03775, l(2)46CFf, l(2)46CFs, l(2)46Ch, p97, pTER94, ter94, vcp, TER94, CG2331-PA, CG2331-PC, CG2331-PD, CG2331-PE, TER94-PA, TER94-PC, TER94-PD, TER94-PE, TRANSITIONAL ENDOPLASMIC RETICULUM ATPASE TER94, Ter94, Valosin-Containing protein, complementation group I, p97 ATPase, transitional endoplasmic reticulum 94, transitional endoplasmic reticulum ATPase, valosin containing protein
Species: fruit fly

Top Publications

  1. Van Bortle K, Ramos E, Takenaka N, Yang J, Wahi J, Corces V. Drosophila CTCF tandemly aligns with other insulator proteins at the borders of H3K27me3 domains. Genome Res. 2012;22:2176-87 pubmed publisher
    ..We reveal an important relationship between dCTCF and other Drosophila insulator proteins and speculate that vertebrate CTCF may also align with other nuclear proteins to accomplish similar functions. ..
  2. Lundgren J, Masson P, Mirzaei Z, Young P. Identification and characterization of a Drosophila proteasome regulatory network. Mol Cell Biol. 2005;25:4662-75 pubmed
    ..functions, including Rad23, isopeptidase T, sequestosome, and the genes for the segregase complex TER94/VCP-Ufd1-Npl4 were also found to be up-regulated...
  3. Chang Y, Hung W, Chang Y, Chang H, Wu C, Chiang A, et al. Pathogenic VCP/TER94 alleles are dominant actives and contribute to neurodegeneration by altering cellular ATP level in a Drosophila IBMPFD model. PLoS Genet. 2011;7:e1001288 pubmed publisher
    ..To understand how VCP mutations cause IBMPFD, we have established a Drosophila model by overexpressing TER94 (the sole Drosophila VCP ortholog) carrying mutations analogous to those implicated in IBMPFD...
  4. Pinter M, Jekely G, Szepesi R, Farkas A, Theopold U, Meyer H, et al. TER94, a Drosophila homolog of the membrane fusion protein CDC48/p97, is accumulated in nonproliferating cells: in the reproductive organs and in the brain of the imago. Insect Biochem Mol Biol. 1998;28:91-8 pubmed
    ..The open reading frame of the Drosophila homolog encodes an 801 amino acid long protein (TER94), which shows high similarity to the known CDC48/p97 sequences. The chromosomal position of TER94 is 46 C/D...
  5. Leon A, McKearin D. Identification of TER94, an AAA ATPase protein, as a Bam-dependent component of the Drosophila fusome. Mol Biol Cell. 1999;10:3825-34 pubmed
    ..In this article we present evidence that the fly homologue of the transitional endoplasmic reticulum ATPase (TER94) is one such protein...
  6. Ruden D, Sollars V, Wang X, Mori D, Alterman M, Lu X. Membrane fusion proteins are required for oskar mRNA localization in the Drosophila egg chamber. Dev Biol. 2000;218:314-25 pubmed
    ..We identified mutations in ter94 which disrupt the localization of oskar mRNA to the posterior pole of the oocyte...
  7. Beskow A, Grimberg K, Bott L, Salomons F, Dantuma N, Young P. A conserved unfoldase activity for the p97 AAA-ATPase in proteasomal degradation. J Mol Biol. 2009;394:732-46 pubmed publisher
    The multifunctional AAA-ATPase p97 is one of the most abundant and conserved proteins in eukaryotic cells...
  8. Fujioka M, Wu X, Jaynes J. A chromatin insulator mediates transgene homing and very long-range enhancer-promoter communication. Development. 2009;136:3077-87 pubmed publisher
    ..Enhancers that normally drive Drosophila even skipped (eve) expression are located closer to the TER94 transcription start site than to that of eve...
  9. Zhang Z, Lv X, Yin W, Zhang X, Feng J, Wu W, et al. Ter94 ATPase complex targets k11-linked ubiquitinated ci to proteasomes for partial degradation. Dev Cell. 2013;25:636-44 pubmed publisher
    ..Here, we show that Ter94 ATPase and its mammalian counterpart, p97, are involved in processing Ci and Gli3 into Ci75 and Gli3(R), ..

More Information


  1. Ritson G, Custer S, Freibaum B, Guinto J, Geffel D, Moore J, et al. TDP-43 mediates degeneration in a novel Drosophila model of disease caused by mutations in VCP/p97. J Neurosci. 2010;30:7729-39 pubmed publisher
    ..VCP (p97 in mouse, TER94 in Drosophila melanogaster, and CDC48 in Saccharomyces cerevisiae) is a highly conserved AAA(+) (ATPases ..
  2. Kim N, Tresse E, Kolaitis R, Molliex A, Thomas R, Alami N, et al. VCP is essential for mitochondrial quality control by PINK1/Parkin and this function is impaired by VCP mutations. Neuron. 2013;78:65-80 pubmed publisher
    Mutations in VCP cause multisystem degeneration impacting the nervous system, muscle, and/or bone. Patients may present with ALS, Parkinsonism, frontotemporal dementia, myopathy, Paget's disease, or a combination of these...
  3. Manno A, Noguchi M, Fukushi J, Motohashi Y, Kakizuka A. Enhanced ATPase activities as a primary defect of mutant valosin-containing proteins that cause inclusion body myopathy associated with Paget disease of bone and frontotemporal dementia. Genes Cells. 2010;15:911-22 pubmed publisher
    Valosin-containing protein (VCP) has been shown to colocalize with abnormal protein aggregates, such as nuclear inclusions of Huntington disease and Machado-Joseph disease, Lewy bodies in Parkinson disease...
  4. Higashiyama H, Hirose F, Yamaguchi M, Inoue Y, Fujikake N, Matsukage A, et al. Identification of ter94, Drosophila VCP, as a modulator of polyglutamine-induced neurodegeneration. Cell Death Differ. 2002;9:264-73 pubmed
    ..Through genetic screening, using this fly model, we identified loss-of-function mutants of the ter94 gene that encodes the Drosophila homolog of VCP/CDC48, a member of the AAA+ class of the ATPase protein family, as ..
  5. Sackerson C, Fujioka M, Goto T. The even-skipped locus is contained in a 16-kb chromatin domain. Dev Biol. 1999;211:39-52 pubmed
    ..4) Expression in different sites in the central nervous system is driven by separable elements widely dispersed throughout 8 kb 3' of the gene. ..
  6. Hare E, Peterson B, Iyer V, Meier R, Eisen M. Sepsid even-skipped enhancers are functionally conserved in Drosophila despite lack of sequence conservation. PLoS Genet. 2008;4:e1000106 pubmed publisher
    ..Together, these observations suggest that the local arrangement of binding sites relative to each other is more important than their overall arrangement into larger units of cis-regulatory function...
  7. Fujita K, Nakamura Y, Oka T, Ito H, Tamura T, Tagawa K, et al. A functional deficiency of TERA/VCP/p97 contributes to impaired DNA repair in multiple polyglutamine diseases. Nat Commun. 2013;4:1816 pubmed publisher
    ..Here we show that transitional endoplasmic reticulum ATPase (TERA)/valosin-containing protein (VCP)/p97 directly binds to multiple polyglutamine disease ..
  8. Bour B, O Brien M, Lockwood W, Goldstein E, Bodmer R, Taghert P, et al. Drosophila MEF2, a transcription factor that is essential for myogenesis. Genes Dev. 1995;9:730-41 pubmed
    ..These results indicate that MEF2 is required for later aspects of differentiation of the three major types of musculature, which include body wall muscles, gut musculature, and the heart, in the Drosophila embryo. ..
  9. McKearin D. The Drosophila fusome, organelle biogenesis and germ cell differentiation: if you build it. Bioessays. 1997;19:147-52 pubmed
    ..Future efforts will consider to what extent an organelle assembly-dependent model for differentiation is heuristic and whether the Drosophila fusome represents a homolog of a similar organelle in vertebrate lymphocytes. ..
  10. Lighthouse D, Buszczak M, Spradling A. New components of the Drosophila fusome suggest it plays novel roles in signaling and transport. Dev Biol. 2008;317:59-71 pubmed publisher
    ..In contrast, rab11 is required to maintain the germline stem cells, and to maintain the vesicle content of the spectrosome, suggesting that the fusome mediates intercellular signals that depend on the recycling endosome. ..
  11. Huang Z, Ren S, Jiang Y, Wang T. PINK1 and Parkin cooperatively protect neurons against constitutively active TRP channel-induced retinal degeneration in Drosophila. Cell Death Dis. 2016;7:e2179 pubmed publisher
    ..In addition, the results demonstrated that maintaining mitochondrial homeostasis via PINK1/Parkin-dependent mitochondrial quality control can potentially alleviate cell death in a wide range of neurodegenerative diseases. ..
  12. Fujioka M, Mistry H, Schedl P, Jaynes J. Determinants of Chromosome Architecture: Insulator Pairing in cis and in trans. PLoS Genet. 2016;12:e1005889 pubmed publisher
    ..These pairing principles provide a mechanistic underpinning for the observed topologies within and between chromosomes. ..
  13. Wang T, Xu W, Qin M, Yang Y, Bao P, Shen F, et al. Pathogenic Mutations in the Valosin-containing Protein/p97(VCP) N-domain Inhibit the SUMOylation of VCP and Lead to Impaired Stress Response. J Biol Chem. 2016;291:14373-84 pubmed publisher
    Valosin-containing protein/p97(VCP) is a hexameric ATPase vital to protein degradation during endoplasmic reticulum stress. It regulates diverse cellular functions including autophagy, chromatin remodeling, and DNA repair...
  14. Koike M, Fukushi J, Ichinohe Y, Higashimae N, Fujishiro M, Sasaki C, et al. Valosin-containing protein (VCP) in novel feedback machinery between abnormal protein accumulation and transcriptional suppression. J Biol Chem. 2010;285:21736-49 pubmed publisher
    ..Genetic and biochemical analyses indicate that valosin-containing protein (VCP) is a crucial molecule in the pathogenesis of human neurodegenerative disorders...
  15. Goldstein E, Treadway S, Stephenson A, Gramstad G, Keilty A, Kirsch L, et al. A genetic analysis of the cytological region 46C-F containing the Drosophila melanogaster homolog of the jun proto-oncogene. Mol Genet Genomics. 2001;266:695-700 pubmed
    ..There are twelve embryonic lethal groups and seven larval lethal groups. Three lines show visible abnormalities in gut and tracheal development prior to death. ..
  16. Delabaere L, Orsi G, Sapey Triomphe L, Horard B, Couble P, Loppin B. The Spartan ortholog maternal haploid is required for paternal chromosome integrity in the Drosophila zygote. Curr Biol. 2014;24:2281-7 pubmed publisher
    ..Like Spartan, MH protein is involved in the resistance to UV radiation and recruits the p97/TER94 segregase to stalled DNA replication forks in somatic cells...
  17. Li Y, Zhang Q, Carreira Rosario A, Maines J, McKearin D, Buszczak M. Mei-p26 cooperates with Bam, Bgcn and Sxl to promote early germline development in the Drosophila ovary. PLoS ONE. 2013;8:e58301 pubmed publisher
    ..Together with recently published studies, these data suggest that Mei-P26 mediates both GSC self-renewal and germline differentiation through distinct modes of translational repression depending on the presence of Bam...
  18. Casci I, Pandey U. A fruitful endeavor: modeling ALS in the fruit fly. Brain Res. 2015;1607:47-74 pubmed publisher
    ..Our primary goal in this review is to highlight the lessons we have learned using Drosophila models pertaining to ALS research. This article is part of a Special Issue entitled ALS complex pathogenesis. ..
  19. Wang Z, Clark C, Geisbrecht E. Drosophila clueless is involved in Parkin-dependent mitophagy by promoting VCP-mediated Marf degradation. Hum Mol Genet. 2016;25:1946-1964 pubmed
    ..Loss of clu leads to the recruitment of Parkin, VCP/p97, p62/Ref(2)P and Atg8a to depolarized swollen mitochondria. However, clearance of damaged mitochondria is impeded...
  20. Rode S, Ohm H, Zipfel J, Rumpf S. The spliceosome-associated protein Mfap1 binds to VCP in Drosophila. PLoS ONE. 2017;12:e0183733 pubmed publisher
    ..Previously, we found that the AAA chaperone Valosin-Containing Protein (VCP) regulates ecdysone-dependent dendrite pruning of Drosophila class IV dendritic arborization (c4da) neurons via an ..
  21. Reim G, Hruzova M, Goetze S, Basler K. Protection of armadillo/β-Catenin by armless, a novel positive regulator of wingless signaling. PLoS Biol. 2014;12:e1001988 pubmed publisher
    ..We show by biochemical and in vivo studies that Als interacts directly with Ter94, an AAA ATPase known to associate with E3 ligases and to drive protein turnover...
  22. Chan H, Lee T, Huang S, Lee H, Sang T, Chan H, et al. Proteomic analysis of a drosophila IBMPFD model reveals potential pathogenic mechanisms. Mol Biosyst. 2012;8:1730-41 pubmed publisher
    ..with the head proteome of transgenic mutant type VCPs that correspond to human IBMPFD disease alleles (TER94(A229E), TER94(R188Q), and TER94(R152H))...
  23. Liu L, Qi H, Wang J, Lin H. PAPI, a novel TUDOR-domain protein, complexes with AGO3, ME31B and TRAL in the nuage to silence transposition. Development. 2011;138:1863-73 pubmed publisher
    ..Overall, our study reveals a function of the nuage in safeguarding the germline genome against deleterious retrotransposition via the piRNA pathway. ..
  24. Dunne J, Kondylis V, Rabouille C. Ecdysone triggers the expression of Golgi genes in Drosophila imaginal discs via broad-complex. Dev Biol. 2002;245:172-86 pubmed
    ..The ecdysone-triggered biogenesis of Golgi stacks at the onset of imaginal disc elongation offers the exciting possibility of advancing our understanding of the relationship between gene expression and organelle biogenesis. ..
  25. Nahvi A, Shoemaker C, Green R. An expanded seed sequence definition accounts for full regulation of the hid 3' UTR by bantam miRNA. RNA. 2009;15:814-22 pubmed publisher
    ..Further support for the potential contribution of the 3-9 seed register to microRNA-mediated gene regulation is provided by the experimental validation of several novel bantam targets identified with a more relaxed search algorithm. ..
  26. Fujioka M, Sun G, Jaynes J. The Drosophila eve insulator Homie promotes eve expression and protects the adjacent gene from repression by polycomb spreading. PLoS Genet. 2013;9:e1003883 pubmed publisher
    ..It blocks the spreading of the eve Pc domain, preventing repression of the adjacent gene, TER94. It prevents activation of TER94 by eve regulatory DNA. It also facilitates normal eve expression...
  27. Thomson T, Liu N, Arkov A, Lehmann R, Lasko P. Isolation of new polar granule components in Drosophila reveals P body and ER associated proteins. Mech Dev. 2008;125:865-73 pubmed publisher
    ..We identified Maternal expression at 31B (ME31B), eIF4A, Aubergine (AUB) and Transitional Endoplasmic Reticulum 94 (TER94) as components of both VAS and TUD complexes and confirmed their localization to polar ..
  28. Avery M, Sheehan A, Kerr K, Wang J, Freeman M. Wld S requires Nmnat1 enzymatic activity and N16-VCP interactions to suppress Wallerian degeneration. J Cell Biol. 2009;184:501-13 pubmed publisher
    ..of Nmnat1-protective activity in Wld(S) requires the N16-binding protein valosin-containing protein (VCP)/TER94. Thus, Wld(S)-mediated suppression of Wallerian degeneration results from VCP-N16 interactions and Nmnat1 activity ..
  29. Panda D, Rose P, Hanna S, Gold B, Hopkins K, Lyde R, et al. Genome-wide RNAi screen identifies SEC61A and VCP as conserved regulators of Sindbis virus entry. Cell Rep. 2013;5:1737-48 pubmed publisher
    ..We identified a conserved role for SEC61A and valosin-containing protein (VCP) in facilitating SINV entry in insects and mammals...
  30. Santhanam A, Peng W, Yu Y, Sang T, Chen G, Meng T. Ecdysone-induced receptor tyrosine phosphatase PTP52F regulates Drosophila midgut histolysis by enhancement of autophagy and apoptosis. Mol Cell Biol. 2014;34:1594-606 pubmed publisher
    ..The mutation of the Ptp52F gene caused significant delay in larval midgut degradation. Transitional endoplasmic reticulum ATPase (TER94), a regulator of ubiquitin proteasome system, was identified as a substrate and ..
  31. Ristić G, Tsou W, Guzi E, Kanack A, SCAGLIONE K, Todi S. USP5 Is Dispensable for Monoubiquitin Maintenance in Drosophila. J Biol Chem. 2016;291:9161-72 pubmed publisher
    ..Our work indicates that a primary physiological role of USP5 is not to recycle monoubiquitin for reutilization, but that it may involve disassembly of conjugated ubiquitin to maintain proteasome function. ..
  32. Li M, Lai P, Chou Y, Chi A, Mi Y, Khoo K, et al. Protein tyrosine phosphatase PTPN3 inhibits lung cancer cell proliferation and migration by promoting EGFR endocytic degradation. Oncogene. 2015;34:3791-803 pubmed publisher
    ..Taken together, these results indicate that PTPN3 may act as a tumor suppressor in lung cancer through its modulation of EGFR signaling. ..
  33. Romano M, Feiguin F, Buratti E. Drosophila Answers to TDP-43 Proteinopathies. J Amino Acids. 2012;2012:356081 pubmed publisher
  34. Currie P, Sullivan D. Structure and expression of the gene encoding phosphofructokinase (PFK) in Drosophila melanogaster. J Biol Chem. 1994;269:24679-87 pubmed
    ..Deficiency analysis of the 46D/E region has identified a lethal complementation group associated with reduced PFK activity, thereby establishing the vital role of PFK function in Drosophila. ..
  35. Griciuc A, Aron L, Roux M, Klein R, Giangrande A, Ueffing M. Inactivation of VCP/ter94 suppresses retinal pathology caused by misfolded rhodopsin in Drosophila. PLoS Genet. 2010;6: pubmed publisher
    ..and we have recently shown that Rh(P23H) is a substrate of the ER-associated degradation (ERAD) effector VCP/ter94, a chaperone that extracts misfolded proteins from the ER (a process called retrotranslocation) and facilitates ..
  36. Rumpf S, Bagley J, Thompson Peer K, Zhu S, Gorczyca D, Beckstead R, et al. Drosophila Valosin-Containing Protein is required for dendrite pruning through a regulatory role in mRNA metabolism. Proc Natl Acad Sci U S A. 2014;111:7331-6 pubmed publisher
    ..Here, we show that mutation or inhibition of Valosin-Containing Protein (VCP), a ubiquitin-dependent ATPase whose human homolog is linked to neurodegenerative disease, leads to specific ..
  37. Johnson A, Shu H, Hauswirth A, Tong A, Davis G. VCP-dependent muscle degeneration is linked to defects in a dynamic tubular lysosomal network in vivo. elife. 2015;4: pubmed publisher
    ..The dynamics and integrity of this tubular lysosomal network requires VCP, an AAA-ATPase that, when mutated, causes degenerative diseases of muscle, bone and neurons...
  38. Rumpf S, Lee S, Jan L, Jan Y. Neuronal remodeling and apoptosis require VCP-dependent degradation of the apoptosis inhibitor DIAP1. Development. 2011;138:1153-60 pubmed publisher
    ..Here, we examined the function of Valosin-containing protein (VCP), a ubiquitin-selective AAA chaperone involved in endoplasmic reticulum-associated degradation, autophagy and ..
  39. Wojcik C, Yano M, Demartino G. RNA interference of valosin-containing protein (VCP/p97) reveals multiple cellular roles linked to ubiquitin/proteasome-dependent proteolysis. J Cell Sci. 2004;117:281-92 pubmed
    ..RNA interference (RNAi) to examine the functional relationship between valosin-containing protein (VCP/p97/Cdc48p/TER94) ATPase and the ubiquitin-proteasome system (UPS) in Drosophila S2 and human HeLa cells...
  40. Azuma Y, Tokuda T, Shimamura M, Kyotani A, Sasayama H, Yoshida T, et al. Identification of ter94, Drosophila VCP, as a strong modulator of motor neuron degeneration induced by knockdown of Caz, Drosophila FUS. Hum Mol Genet. 2014;23:3467-80 pubmed publisher
    ..We examined several previously reported ALS-related genes and found genetic links between Caz and ter94, the Drosophila ortholog of human Valosin-containing protein (VCP)...
  41. Zeng Z, de Gorter D, Kowalski M, Ten Dijke P, Shimmi O. Ter94/VCP is a novel component involved in BMP signaling. PLoS ONE. 2014;9:e114475 pubmed publisher
    ..b>Ter94, a member of the family of AAA ATPases, was identified as a novel mediator of BMP signaling, which is required for ..
  42. Hao Q, Jiao S, Shi Z, Li C, Meng X, Zhang Z, et al. A non-canonical role of the p97 complex in RIG-I antiviral signaling. EMBO J. 2015;34:2903-20 pubmed publisher
    ..Consistently, administration of compounds targeting p97 ATPase activity was shown to inhibit viral replication and protect mice from vesicular stomatitis virus (VSV) infection...
  43. Liang C, Chang Y, Chang H, Wang C, Hung Y, Lin Y, et al. Derlin-1 regulates mutant VCP-linked pathogenesis and endoplasmic reticulum stress-induced apoptosis. PLoS Genet. 2014;10:e1004675 pubmed publisher
    ..Using a Drosophila IBMPFD model, we have identified the ER protein Derlin-1 as a modifier of pathogenic TER94 (the fly VCP homolog) mutants...