Gene Symbol: Pen
Description: Pendulin
Alias: CG4799, DPend, Dalpha2, Dimp-alpha2, Dmel\CG4799, IMPalpha2, Imp-alpha2, Kap alpha2, Kap-alpha2, Kpna2, OHO31, Oho31, Rch1, alpha2, alpha2A-Kap, alphaKap2, anon-WO0140519.258, bs29g06.y1, imp alpha2, imp-alpha2, impalpha2, importin alpha2/pendulin, l(2)144/1, l(2)k06324, l(2)k14401, oho-31, oho31, pen, pen-2, pendulin, CG4799-PA, Importin-a2, Importin-alpha, Importin-alpha2, Pen-PA, alpha-importin, importin alpha, importin alpha2, lethal (2) k06324
Species: fruit fly

Top Publications

  1. Mosca T, Schwarz T. The nuclear import of Frizzled2-C by Importins-beta11 and alpha2 promotes postsynaptic development. Nat Neurosci. 2010;13:935-43 pubmed publisher
    ..We found that Fz2-C localization to muscle nuclei required the nuclear import factors Importin-beta11 and Importin-alpha2 and that this pathway promoted the postsynaptic development of the subsynaptic reticulum (SSR), an elaboration of ..
  2. Giarre M, Török I, Schmitt R, Gorjanacz M, Kiss I, Mechler B. Patterns of importin-alpha expression during Drosophila spermatogenesis. J Struct Biol. 2002;140:279-90 pubmed
    ..antibodies raised against each of the three Importin-alpha proteins identified in Drosophila, namely, Imp-alpha1, -alpha2, and -alpha3. We found that each Imp-alpha is expressed during a specific and limited period of spermatogenesis...
  3. Mason D, Goldfarb D. The nuclear transport machinery as a regulator of Drosophila development. Semin Cell Dev Biol. 2009;20:582-9 pubmed publisher
    ..Rather, they support the idea that the basal nuclear transport machinery has adapted during the evolution of the metazoan body plan to play important regulatory roles in key developmental events. ..
  4. Malik H, Eickbush T, Goldfarb D. Evolutionary specialization of the nuclear targeting apparatus. Proc Natl Acad Sci U S A. 1997;94:13738-42 pubmed
    ..Phylogenetic analysis suggests that the various alpha-Kaps fall into two groups, alpha1 and alpha2. Whereas animals encode both types, the yeast genome encodes only an alpha1-Kap...
  5. Mason D, Fleming R, Goldfarb D. Drosophila melanogaster importin alpha1 and alpha3 can replace importin alpha2 during spermatogenesis but not oogenesis. Genetics. 2002;161:157-70 pubmed
    ..among conventional importin alpha genes by studying their capacity to rescue the male and female sterility of importin alpha2 null flies...
  6. Mathe E, Bates H, Huikeshoven H, Deak P, Glover D, Cotterill S. Importin-alpha3 is required at multiple stages of Drosophila development and has a role in the completion of oogenesis. Dev Biol. 2000;223:307-22 pubmed
    ..7% identical to that of Importin-alpha2 (Oho31/Pendulin), the previously reported Drosophila homologue...
  7. Torok I, Strand D, Schmitt R, Tick G, Torok T, Kiss I, et al. The overgrown hematopoietic organs-31 tumor suppressor gene of Drosophila encodes an Importin-like protein accumulating in the nucleus at the onset of mitosis. J Cell Biol. 1995;129:1473-89 pubmed
    The tumor suppressor gene overgrown hematopoietic organs-31 (oho31) of Drosophila encodes a protein with extensive homology to the Importin protein of Xenopus (50% identity), the related yeast SRP1 protein, and the mammalian hSRP1 and ..
  8. Küssel P, Frasch M. Pendulin, a Drosophila protein with cell cycle-dependent nuclear localization, is required for normal cell proliferation. J Cell Biol. 1995;129:1491-507 pubmed
    We describe the dynamic intracellular localization of Drosophila Pendulin and its role in the control of cell proliferation...
  9. Gorjanacz M, Adam G, Török I, Mechler B, Szlanka T, Kiss I. Importin-alpha 2 is critically required for the assembly of ring canals during Drosophila oogenesis. Dev Biol. 2002;251:271-82 pubmed
    ..Immunohistochemistry shows that, although the Imp-alpha2 protein cannot be detected on the ring canals, the Kelch protein, a known ring canal component, fails to bind to ..

More Information


  1. Mason D, Mathe E, Fleming R, Goldfarb D. The Drosophila melanogaster importin alpha3 locus encodes an essential gene required for the development of both larval and adult tissues. Genetics. 2003;165:1943-58 pubmed
    ..transport of classical nuclear localization signal (cNLS)-containing proteins is mediated by the cNLS receptor importin alpha. The conventional importin alpha gene family in metazoan animals is composed of three clades that are ..
  2. Fang X, Chen T, Tran K, Parker C. Developmental regulation of the heat shock response by nuclear transport factor karyopherin-alpha3. Development. 2001;128:3349-58 pubmed
    ..From cycle 13 onward the transport factor is present and the dHSF is localized within the nucleus thus allowing the embryo to respond to heat shock. ..
  3. Ratan R, Mason D, Sinnot B, Goldfarb D, Fleming R. Drosophila importin alpha1 performs paralog-specific functions essential for gametogenesis. Genetics. 2008;178:839-50 pubmed publisher
    b>Importin alpha's mediate nuclear transport by linking nuclear localization signal (NLS)-containing proteins to importin beta1...
  4. Chan C, Zhang S, Rousset R, Wharton K. Drosophila Naked cuticle (Nkd) engages the nuclear import adaptor Importin-alpha3 to antagonize Wnt/beta-catenin signaling. Dev Biol. 2008;318:17-28 pubmed publisher
    ..Our results provide further support for the hypothesis that Nkd inhibits nuclear step(s) in Wnt/beta-catenin signaling and broaden our understanding of signaling pathways that engage the nuclear import machinery. ..
  5. Mason D, Stage D, Goldfarb D. Evolution of the metazoan-specific importin alpha gene family. J Mol Evol. 2009;68:351-65 pubmed publisher
    ..Studies in Drosophila melanogaster, Caenorhabditis elegans, and mouse suggest that the animal importin alpha gene family evolved from ancestral plant-like genes to serve paralog-specific roles in gametogenesis...
  6. Torok T, Tick G, Alvarado M, Kiss I. P-lacW insertional mutagenesis on the second chromosome of Drosophila melanogaster: isolation of lethals with different overgrowth phenotypes. Genetics. 1993;135:71-80 pubmed
    ..In this paper, 16 overgrowth mutants are described. As revealed by in situ hybridization, none of the mutations corresponds to any of the previously known overgrowth mutations on the second chromosome. ..
  7. Mosca T, Schwarz T. Drosophila Importin-?2 is involved in synapse, axon and muscle development. PLoS ONE. 2010;5:e15223 pubmed publisher
    ..Thus, Importin-?2 is needed for some processes in the development of both the nervous system and the larval musculature. ..
  8. Samsonova A, Niranjan M, Russell S, Brazma A. Prediction of gene expression in embryonic structures of Drosophila melanogaster. PLoS Comput Biol. 2007;3:e144 pubmed
    ..Our novel organism-independent method will be of considerable utility in enriching the annotation of gene function and expression in complex multicellular organisms. ..
  9. Gorjanacz M, Török I, Pomozi I, Garab G, Szlanka T, Kiss I, et al. Domains of Importin-alpha2 required for ring canal assembly during Drosophila oogenesis. J Struct Biol. 2006;154:27-41 pubmed
    Null-mutation in Drosophila importin-alpha2, such as the deficiency imp-alpha2(D14), causes recessive female sterility with the formation of dumpless eggs...
  10. Mensch J, Serra F, Lavagnino N, Dopazo H, Hasson E. Positive selection in nucleoporins challenges constraints on early expressed genes in Drosophila development. Genome Biol Evol. 2013;5:2231-41 pubmed publisher
    ..Understanding these exceptions to the broad evolutionary conservation of early expressed developmental genes can shed light into relevant processes driving the evolution of species divergence. ..
  11. Cryderman D, Grade S, Li Y, Fanti L, Pimpinelli S, Wallrath L. Role of Drosophila HP1 in euchromatic gene expression. Dev Dyn. 2005;232:767-74 pubmed
    ..Collectively, these data demonstrate multiple mechanisms for HP1 localization within euchromatin and show that some genes associated with HP1 are not affected by alterations in Su(var)3-9 dosage. ..
  12. de Belle J. Unifying the genetics of behavior. Nat Genet. 2002;31:329-30 pubmed
  13. Greenspan R. The varieties of selectional experience in behavioral genetics. J Neurogenet. 2003;17:241-70 pubmed
  14. Toma D, White K, Hirsch J, Greenspan R. Identification of genes involved in Drosophila melanogaster geotaxis, a complex behavioral trait. Nat Genet. 2002;31:349-53 pubmed
    ..We have thus determined the identities of several genes that contribute to the complex, polygenic behavior of geotaxis. ..
  15. Starz Gaiano M, Lehmann R. Moving towards the next generation. Mech Dev. 2001;105:5-18 pubmed
    ..Here we review findings from Drosophila, zebrafish, and mouse; each organism provides unique insight into the mechanisms that determine germ cell fate and the cues that may guide their migration. ..
  16. Dearolf C. Fruit fly "leukemia". Biochim Biophys Acta. 1998;1377:M13-23 pubmed
  17. Phadnis N, Hsieh E, Malik H. Birth, death, and replacement of karyopherins in Drosophila. Mol Biol Evol. 2012;29:1429-40 pubmed publisher
  18. Taniue K, Nishida A, Hamada F, Sugie A, Oda T, Ui Tei K, et al. Sunspot, a link between Wingless signaling and endoreplication in Drosophila. Development. 2010;137:1755-64 pubmed publisher
    ..In addition, Ssp was found not to be involved in the signaling pathway mediated by Arm associated with dTCF. Our findings indicate that Arm controls development in part by regulating the function of Ssp. ..
  19. Gibson G, Wemple M, van Helden S. Potential variance affecting homeotic Ultrabithorax and Antennapedia phenotypes in Drosophila melanogaster. Genetics. 1999;151:1081-91 pubmed
    ..It is argued that the existence of "potential" variance, which is genetic variation whose effects are not observable in wild-type individuals, is a prerequisite for the uncoupling of genetic from phenotypic divergence. ..
  20. Cserpan I, Mathe E, Patthy A, Udvardy A. Characterization of a Drosophila phosphorylation-dependent nuclear-localization-signal-binding protein. Biochem J. 1997;328 ( Pt 3):821-6 pubmed
    ..In agreement with the immunoprecipitation results, cross-linking experiments demonstrated the interaction of p94 with three additional proteins. These protein-protein interactions were also phosphorylation-dependent. ..
  21. Mikhaylova L, Nguyen K, Nurminsky D. Analysis of the Drosophila melanogaster testes transcriptome reveals coordinate regulation of paralogous genes. Genetics. 2008;179:305-15 pubmed publisher
  22. Amaral A, Brito F, Chobanyan T, Yoshikawa S, Yokokura T, Van Vactor D, et al. Quality assessment and control of tissue specific RNA-seq libraries of Drosophila transgenic RNAi models. Front Genet. 2014;5:43 pubmed publisher
    ..Finally, we showed that trimming 5' ends of reads decreases nucleotide frequency biases, increasing the coverage of protein coding genes with a potential positive impact in the incurrence of systematic technical errors. ..
  23. Kim Y, Baker B. Isolation of RRM-type RNA-binding protein genes and the analysis of their relatedness by using a numerical approach. Mol Cell Biol. 1993;13:174-83 pubmed
    ..This result suggests that we have isolated good candidates for both housekeeping and developmentally important genes involved in RNA metabolism. ..
  24. Bienz M, Clevers H. Armadillo/beta-catenin signals in the nucleus--proof beyond a reasonable doubt?. Nat Cell Biol. 2003;5:179-82 pubmed
    ..Plausible explanations allow these observations to be reconciled with the large body of evidence supporting a nuclear function of Arm/beta-catenin. ..
  25. Sabri N, Roth P, Xylourgidis N, Sadeghifar F, Adler J, Samakovlis C. Distinct functions of the Drosophila Nup153 and Nup214 FG domains in nuclear protein transport. J Cell Biol. 2007;178:557-65 pubmed
    ..nucleoporins in cultured Drosophila melanogaster S2 cells by RNA interference and analyzed the phenotypes on importin alpha/beta-mediated import and CRM1-dependent protein export...
  26. Belacortu Y, Weiss R, Kadener S, Paricio N. Transcriptional activity and nuclear localization of Cabut, the Drosophila ortholog of vertebrate TGF-?-inducible early-response gene (TIEG) proteins. PLoS ONE. 2012;7:e32004 pubmed publisher
  27. Jagut M, Mihaila Bodart L, Molla Herman A, Alin M, Lepesant J, Huynh J. A mosaic genetic screen for genes involved in the early steps of Drosophila oogenesis. G3 (Bethesda). 2013;3:409-25 pubmed publisher
    ..This collection of mutants will be useful to investigate further the early steps of Drosophila oogenesis at a genetic level. ..
  28. Erdelyi M, Mathe E, Szabad J. Genetic and developmental analysis of mutant Ketel alleles that identify the Drosophila importin-beta homologue. Acta Biol Hung. 1997;48:323-38 pubmed
    ..Molecular analysis (to be published elsewhere) confirmed the genetic data and revealed that the Ketel gene encodes the Drosophila homologue of importin-beta, an essential component of nuclear protein import. ..
  29. Jans D, Xiao C, Lam M. Nuclear targeting signal recognition: a key control point in nuclear transport?. Bioessays. 2000;22:532-44 pubmed
    ..Targeting sequence recognition thus appears to represent a key control point in the regulation of nuclear transport. BioEssays 22:532-544, 2000. ..
  30. Jang A, Moravcevic K, Saez L, Young M, Sehgal A. Drosophila TIM binds importin α1, and acts as an adapter to transport PER to the nucleus. PLoS Genet. 2015;11:e1004974 pubmed publisher
    ..Thus, the circadian clock uses specific components of the importin pathway with a novel twist in that TIM serves a karyopherin-like role for PER. ..
  31. Hogarth C, Itman C, Jans D, Loveland K. Regulated nucleocytoplasmic transport in spermatogenesis: a driver of cellular differentiation?. Bioessays. 2005;27:1011-25 pubmed
  32. Liu J, Lee D, Yu C, Angers S, Harris T. Stepping stone: a cytohesin adaptor for membrane cytoskeleton restraint in the syncytial Drosophila embryo. Mol Biol Cell. 2015;26:711-25 pubmed publisher
    ..Overall we propose that Sstn acts as a cytohesin adaptor that promotes Steppke activity for localized membrane cytoskeleton restraint in the syncytial Drosophila embryo. ..
  33. Amcheslavsky A, Nie Y, Li Q, He F, Tsuda L, Markstein M, et al. Gene expression profiling identifies the zinc-finger protein Charlatan as a regulator of intestinal stem cells in Drosophila. Development. 2014;141:2621-32 pubmed publisher
    ..The results together suggest that Charlatan does not simply act as an anti-differentiation factor but instead functions to maintain a chromatin structure that is compatible with stem cell properties, including proliferation. ..
  34. Kirkpatrick C, Peifer M. Not just glue: cell-cell junctions as cellular signaling centers. Curr Opin Genet Dev. 1995;5:56-65 pubmed
  35. Sachan N, Mishra A, Mutsuddi M, Mukherjee A. The Drosophila importin-?3 is required for nuclear import of notch in vivo and it displays synergistic effects with notch receptor on cell proliferation. PLoS ONE. 2013;8:e68247 pubmed publisher
    ..Taken together, our results suggest that Importin-?3 mediated nuclear import of Notch-ICD may play important role in regulation of Notch signaling. ..
  36. Viragh E, Gorjanacz M, Török I, Eichhorn T, Kallakuri S, Szlanka T, et al. Specific Cooperation Between Imp-?2 and Imp-?/Ketel in Spindle Assembly During Drosophila Early Nuclear Divisions. G3 (Bethesda). 2012;2:1-14 pubmed publisher
    ..These data suggest that a fine balance among Imp-?2, Imp-?, RanGTP, and the NLS cargos is critical for mitotic progression during early embryonic development. ..
  37. Graham A, Davis S, Andrulis E. Interdependent nucleocytoplasmic trafficking and interactions of Dis3 with Rrp6, the core exosome and importin-alpha3. Traffic. 2009;10:499-513 pubmed publisher
    ..We propose that the movement and deposition of this complex is important for the subcellular compartmentalization and regulation of the exosome core. ..
  38. Davis I. Nuclear polarity and nuclear trafficking in Drosophila. Semin Cell Dev Biol. 1997;8:91-7 pubmed
  39. Küssel P, Frasch M. Yeast Srp1, a nuclear protein related to Drosophila and mouse pendulin, is required for normal migration, division, and integrity of nuclei during mitosis. Mol Gen Genet. 1995;248:351-63 pubmed
    ..mouse with significant sequence similarities to a Drosophila gene that encodes the blood cell tumor suppressor pendulin. The protein encoded by the yeast gene, Srp1p, and mouse pendulin share 42% and 51% amino acid identity with ..
  40. Wang S, Lu Y, Yin M, Wang C, Wu W, Li J, et al. Importin α1 Mediates Yorkie Nuclear Import via an N-terminal Non-canonical Nuclear Localization Signal. J Biol Chem. 2016;291:7926-37 pubmed publisher
    ..Taken together, we identified a potential nuclear localization signal at the N-terminal end of Yorkie as well as a critical role for Importin α1 in Yorkie nuclear import. ..