Gene Symbol: TOMM22
Description: translocase of outer mitochondrial membrane 22
Alias: 1C9-2, MST065, MSTP065, TOM22, mitochondrial import receptor subunit TOM22 homolog, mitochondrial import receptor Tom22, translocase of outer membrane 22 kDa subunit homolog, translocase of outer mitochondrial membrane 22 homolog
Species: human
Products:     TOMM22

Top Publications

  1. Yano M, Terada K, Mori M. AIP is a mitochondrial import mediator that binds to both import receptor Tom20 and preproteins. J Cell Biol. 2003;163:45-56 pubmed
    ..These results suggest that AIP functions as a cytosolic factor that mediates preprotein import into mitochondria. ..
  2. Yano M, Hoogenraad N, Terada K, Mori M. Identification and functional analysis of human Tom22 for protein import into mitochondria. Mol Cell Biol. 2000;20:7205-13 pubmed
    ..We report here the identification and functional analysis of human Tom22 (hTom22)...
  3. Sakaue H, Shiota T, Ishizaka N, Kawano S, Tamura Y, Tan K, et al. Porin Associates with Tom22 to Regulate the Mitochondrial Protein Gate Assembly. Mol Cell. 2019;73:1044-1055.e8 pubmed publisher
    ..between the majority population of a three-channel gateway ("trimer") and the minor population that lacks Tom22 and has only two Tom40 channels ("dimer")...
  4. Opaliński Ł, Song J, Priesnitz C, Wenz L, Oeljeklaus S, Warscheid B, et al. Recruitment of Cytosolic J-Proteins by TOM Receptors Promotes Mitochondrial Protein Biogenesis. Cell Rep. 2018;25:2036-2043.e5 pubmed publisher
    ..We identify the cytosolic J-protein Xdj1 as a specific interaction partner of the central receptor Tom22. Tom22 recruits Xdj1 to the mitochondrial surface to promote import of preproteins and assembly of the TOM complex...
  5. Alles J, Hasler D, Kazmi S, Tesson M, Hamilton A, Schlegel L, et al. Epstein-Barr Virus EBER Transcripts Affect miRNA-Mediated Regulation of Specific Targets and Are Processed to Small RNA Species. Noncoding RNA. 2015;1:170-191 pubmed publisher
    ..However, the EBERs have no effect upon another target of miR-142-3p, ADCY9, nor on TOMM22, a target of ebv-miR-BART16, indicative of selective modulation of gene expression by the EBERs.
  6. Prasad M, Kaur J, Pawlak K, Bose M, Whittal R, Bose H. Mitochondria-associated endoplasmic reticulum membrane (MAM) regulates steroidogenic activity via steroidogenic acute regulatory protein (StAR)-voltage-dependent anion channel 2 (VDAC2) interaction. J Biol Chem. 2015;290:2604-16 pubmed publisher
    ..In the MAM, StAR interacts with mitochondrial proteins Tom22 and VDAC2. However, Tom22 knockdown by siRNA had no effect on pregnenolone synthesis...
  7. Asmarinah A, Paradowska Dogan A, Kodariah R, Tanuhardja B, Waliszewski P, Mochtar C, et al. Expression of the Bcl-2 family genes and complexes involved in the mitochondrial transport in prostate cancer cells. Int J Oncol. 2014;45:1489-96 pubmed
    ..e., VDAC1, VDAC2 and VDAC3; and TOMM genes, i.e., TOMM20, TOMM22 and TOMM40. This study was performed at the mRNA and the protein level...
  8. Frank D, Dengjel J, Wilfling F, Kozjak Pavlovic V, Hacker G, Weber A. The pro-apoptotic BH3-only protein Bim interacts with components of the translocase of the outer mitochondrial membrane (TOM). PLoS ONE. 2015;10:e0123341 pubmed publisher
    ..However, RNAi against components of TOM (Tom40, Tom70, Tom22 or Tom20) by siRNA, individually or in combination, did not consistently change the amount of Bim on HeLa ..
  9. Nyindodo Ogari L, Schwartzbach S, Estraño C. Giardia mitosomal protein import machinery differentially recognizes mitochondrial targeting signals. Infect Disord Drug Targets. 2014;14:23-9 pubmed
    ..These recombinant proteins include Tom20 and Tom22; two components of the mitochondrial protein import machinery...

More Information


  1. Youcef G, Belaidi E, Waeckel L, Fazal L, Clemessy M, Vincent M, et al. Tissue kallikrein is required for the cardioprotective effect of cyclosporin A in myocardial ischemia in the mouse. Biochem Pharmacol. 2015;94:22-9 pubmed publisher
    ..By proximity ligation assay TK was found in close proximity to the mitochondrial membrane proteins VDAC and Tom22, and CypD...
  2. Li Q, Vande Velde C, Israelson A, Xie J, Bailey A, Dong M, et al. ALS-linked mutant superoxide dismutase 1 (SOD1) alters mitochondrial protein composition and decreases protein import. Proc Natl Acad Sci U S A. 2010;107:21146-51 pubmed publisher
    ..protein import only in spinal cord mitochondria, despite an increase in the mitochondrial import components TOM20, TOM22, and TOM40...
  3. Hu W, Wang Z, Zheng H. Mitochondrial accumulation of amyloid β (Aβ) peptides requires TOMM22 as a main Aβ receptor in yeast. J Biol Chem. 2018;293:12681-12689 pubmed publisher
    ..that Aβ is specifically recognized by translocase of outer mitochondrial membrane subunit 22 (Tom22 in yeast; TOMM22 in human), a noncanonical receptor within the mitochondrial protein import machinery, and that this recognition is ..
  4. Bajaj R, Jaremko Å, Jaremko M, Becker S, Zweckstetter M. Molecular basis of the dynamic structure of the TIM23 complex in the mitochondrial intermembrane space. Structure. 2014;22:1501-11 pubmed publisher
    ..we characterized the dynamic interaction network of the intermembrane space domains of Tim23, Tim21, Tim50, and Tom22 at single-residue level...
  5. Gornicka A, Bragoszewski P, Chrościcki P, Wenz L, Schulz C, Rehling P, et al. A discrete pathway for the transfer of intermembrane space proteins across the outer membrane of mitochondria. Mol Biol Cell. 2014;25:3999-4009 pubmed publisher
    ..outer membrane translocation did not directly involve other core components of the TOM complex, including Tom22. Thus MIA-dependent proteins take another route across the outer mitochondrial membrane that involves Tom40 in a ..
  6. Parvin N, Carrie C, Pabst I, Läßer A, Laha D, Paul M, et al. TOM9.2 Is a Calmodulin-Binding Protein Critical for TOM Complex Assembly but Not for Mitochondrial Protein Import in Arabidopsis thaliana. Mol Plant. 2017;10:575-589 pubmed publisher
    ..TOM9, the plant homolog of yeast Tom22, is significantly smaller due to a truncation in the cytosolic receptor domain, and its precise function is not ..
  7. Prasad M, Walker A, Kaur J, Thomas J, Powell S, Pandey A, et al. Endoplasmic Reticulum Stress Enhances Mitochondrial Metabolic Activity in Mammalian Adrenals and Gonads. Mol Cell Biol. 2016;36:3058-3074 pubmed
    ..of a complex consisting of the mitochondrial translocase, translocase of outer mitochondrial membrane 22 (Tom22), steroidogenic acute regulatory protein (StAR), and 3?-hydroxysteroid dehydrogenase type 2 (3?HSD2) via its ..
  8. Way L, Faktor J, Dvořáková P, Nicholson J, Vojtesek B, Graham D, et al. Rearrangement of mitochondrial pyruvate dehydrogenase subunit dihydrolipoamide dehydrogenase protein-protein interactions by the MDM2 ligand nutlin-3. Proteomics. 2016;16:2327-44 pubmed publisher
    ..Mitotracker confirmed that Nutlin-3 inhibits mitochondrial activity. Enrichment of mitochondria using TOM22+ immunobeads and TMT labeling defined key changes in the mitochondrial proteome after Nutlin-3 treatment...
  9. Backes S, Hess S, Boos F, Woellhaf M, Gödel S, Jung M, et al. Tom70 enhances mitochondrial preprotein import efficiency by binding to internal targeting sequences. J Cell Biol. 2018;217:1369-1382 pubmed publisher
    ..N-terminal matrix-targeting signals (MTSs) direct preproteins to the surface receptors Tom20, Tom22, and Tom70...
  10. Peng C, Yang Q, Wei B, Liu Y, Li Y, Gu D, et al. Identification of potential target genes and related regulatory transcription factors in spontaneous hairline fracture induced by hypervitaminosis A. Injury. 2017;48:1475-1479 pubmed publisher
    ..Key TFs including E2F1, GABP, NRF2 and ELK1 and their targets genes such as Ube2d3, Uba1, Phb2 and Tomm22 may play potential key roles in spontaneous fracture induced by hypervitaminosis A...
  11. Mani J, Meisinger C, Schneider A. Peeping at TOMs-Diverse Entry Gates to Mitochondria Provide Insights into the Evolution of Eukaryotes. Mol Biol Evol. 2016;33:337-51 pubmed publisher
    ..Only two of its subunits, Tom40 and Tom22, are conserved and thus likely were present in the last eukaryotic common ancestor...
  12. Wu J, Choi T, Shin D. tomm22 Knockdown-Mediated Hepatocyte Damages Elicit Both the Formation of Hybrid Hepatocytes and Biliary Conversion to Hepatocytes in Zebrafish Larvae. Gene Expr. 2017;17:237-249 pubmed publisher
    ..might occur in another zebrafish liver injury model in which temporary knockdown of the mitochondrial import gene tomm22 by morpholino antisense oligonucleotides (MO) induces hepatocyte death...
  13. Joseph A, Ljubicic V, Adhihetty P, Hood D. Biogenesis of the mitochondrial Tom40 channel in skeletal muscle from aged animals and its adaptability to chronic contractile activity. Am J Physiol Cell Physiol. 2010;298:C1308-14 pubmed publisher
    ..05) in mitochondria from aged compared with young animals. This was associated with a higher abundance of Tom22, a protein vital for the assembly process...
  14. Samanta L, Agarwal A, Swain N, Sharma R, Gopalan B, Esteves S, et al. Proteomic Signatures of Sperm Mitochondria in Varicocele: Clinical Use as Biomarkers of Varicocele Associated Infertility. J Urol. 2018;200:414-422 pubmed publisher
    ..differentially expressed proteins related to mitochondrial structure (LETM1, EFHC, MIC60, PGAM5, ISOC2 and import TOM22) and function (NDFSU1, UQCRC2 and COX5B, and the core enzymes of carbohydrate and lipid metabolism)...
  15. Joseph A, Adhihetty P, Wawrzyniak N, Wohlgemuth S, Picca A, Kujoth G, et al. Dysregulation of mitochondrial quality control processes contribute to sarcopenia in a mouse model of premature aging. PLoS ONE. 2013;8:e69327 pubmed publisher
    ..05). Additionally, levels of the Tom22 import protein were higher in PolG animals when compared to WT (P<0.05)...
  16. Große L, Wurm C, Brüser C, Neumann D, Jans D, Jakobs S. Bax assembles into large ring-like structures remodeling the mitochondrial outer membrane in apoptosis. EMBO J. 2016;35:402-13 pubmed publisher
    ..indicates that the area enclosed by a Bax ring is devoid of mitochondrial outer membrane proteins such as Tom20, Tom22, and Sam50...
  17. Mani J, Rout S, Desy S, Schneider A. Mitochondrial protein import - Functional analysis of the highly diverged Tom22 orthologue of Trypanosoma brucei. Sci Rep. 2017;7:40738 pubmed publisher
    The β-barrel protein Tom40 and the α-helically anchored membrane protein Tom22 are the only universally conserved subunits of the protein translocase of the mitochondrial outer membrane (TOM)...
  18. Wenz L, Ellenrieder L, Qiu J, Bohnert M, Zufall N, van der Laan M, et al. Sam37 is crucial for formation of the mitochondrial TOM-SAM supercomplex, thereby promoting β-barrel biogenesis. J Cell Biol. 2015;210:1047-54 pubmed publisher
    ..Sam37 interacts with the receptor domain of Tom22 on the cytosolic side of the mitochondrial outer membrane and links TOM and SAM complexes...
  19. Wojtkowska M, Buczek D, Stobienia O, Karachitos A, Antoniewicz M, Slocinska M, et al. The TOM Complex of Amoebozoans: the Cases of the Amoeba Acanthamoeba castellanii and the Slime Mold Dictyostelium discoideum. Protist. 2015;166:349-62 pubmed publisher
    ..castellanii and D. discoideum TOM complex, while the presence of Tom22 and Tom20 is less supported...
  20. van Dooren G, Yeoh L, Striepen B, McFadden G. The Import of Proteins into the Mitochondrion of Toxoplasma gondii. J Biol Chem. 2016;291:19335-50 pubmed publisher
    ..We demonstrate that a T. gondii homologue of Tom22 (TgTom22), a central component of the translocon of the outer mitochondrial membrane (TOM) complex, is essential ..
  21. Yano M, Terada K, Mori M. Mitochondrial import receptors Tom20 and Tom22 have chaperone-like activity. J Biol Chem. 2004;279:10808-13 pubmed
    ..with presequences are initially recognized by Tom20 (translocase of the outer membrane) and, subsequently, by Tom22, whereas hydrophobic preproteins with internal targeting signals are first recognized by Tom70...
  22. Ott M, Norberg E, Walter K, Schreiner P, Kemper C, Rapaport D, et al. The mitochondrial TOM complex is required for tBid/Bax-induced cytochrome c release. J Biol Chem. 2007;282:27633-9 pubmed
    ..Here, we show that the protein translocase of the outer mitochondrial membrane is required for Bax insertion and cytochrome c release. ..
  23. Kravic B, Harbauer A, Romanello V, Simeone L, Vögtle F, Kaiser T, et al. In mammalian skeletal muscle, phosphorylation of TOMM22 by protein kinase CSNK2/CK2 controls mitophagy. Autophagy. 2018;14:311-335 pubmed publisher
    ..Enzymatically, active muscle lysates from skeletal muscle Csnk2b cKO mice phosphorylate murine TOMM22, the mammalian ortholog of yeast Tom22, to a lower extent than lysates prepared from controls...
  24. Bellot G, Cartron P, Er E, Oliver L, Juin P, Armstrong L, et al. TOM22, a core component of the mitochondria outer membrane protein translocation pore, is a mitochondrial receptor for the proapoptotic protein Bax. Cell Death Differ. 2007;14:785-94 pubmed
    ..By using a bacterial two-hybrid assay and crosslinking strategies, we have identified TOM22, a component of the translocase of the outer mitochondrial membrane (TOM), as a mitochondrial receptor of Bax...
  25. Suzuki H, Okazawa Y, Komiya T, Saeki K, Mekada E, Kitada S, et al. Characterization of rat TOM40, a central component of the preprotein translocase of the mitochondrial outer membrane. J Biol Chem. 2000;275:37930-6 pubmed
    ..Taken together, these results indicate that OM38 is a structural and functional homolog of fungal Tom40 and functions as a component of the preprotein import machinery of the rat mitochondrial outer membrane. ..
  26. Renault T, Grandier Vazeille X, Arokium H, Velours G, Camougrand N, Priault M, et al. The cytosolic domain of human Tom22 modulates human Bax mitochondrial translocation and conformation in yeast. FEBS Lett. 2012;586:116-21 pubmed publisher
    ..Expression of the cytosolic receptor domain of human Tom22 increased Bax mitochondrial localization, but decreased the proportion of active Bax...
  27. Bausewein T, Mills D, Langer J, Nitschke B, Nussberger S, Kuhlbrandt W. Cryo-EM Structure of the TOM Core Complex from Neurospora crassa. Cell. 2017;170:693-700.e7 pubmed publisher
    ..Each Tom40 pore is surrounded by the transmembrane segments of the ?-helical subunits Tom5, Tom6, and Tom7. Tom22, the central preprotein receptor, connects the two Tom40 pores at the dimer interface...
  28. Bertolin G, Ferrando Miguel R, Jacoupy M, Traver S, Grenier K, Greene A, et al. The TOMM machinery is a molecular switch in PINK1 and PARK2/PARKIN-dependent mitochondrial clearance. Autophagy. 2013;9:1801-17 pubmed publisher
    ..The downregulation of TOMM40 or its associated core subunit, TOMM22, was sufficient to trigger OMM protein clearance in the absence of PINK1 or PARK2...
  29. Melin J, Kilisch M, Neumann P, Lytovchenko O, Gomkale R, Schendzielorz A, et al. A presequence-binding groove in Tom70 supports import of Mdl1 into mitochondria. Biochim Biophys Acta. 2015;1853:1850-9 pubmed publisher
    ..Aside from the well-characterized presequence receptors Tom20 and Tom22 a third TOM receptor, Tom70, binds proteins of the carrier family containing multiple transmembrane segments...
  30. Marquitz A, Raab Traub N. The role of miRNAs and EBV BARTs in NPC. Semin Cancer Biol. 2012;22:166-72 pubmed publisher
    ..This article will focus on the nature of this complicated set of transcripts and the evidence that they contribute to the development of nasopharyngeal carcinoma. ..
  31. Johnston A, Hoogenraad J, Dougan D, Truscott K, Yano M, Mori M, et al. Insertion and assembly of human tom7 into the preprotein translocase complex of the outer mitochondrial membrane. J Biol Chem. 2002;277:42197-204 pubmed
    ..intermediate can be chased to the stable approximately 380-kDa mammalian TOM complex that additionally contains Tom22. Overexpression of Tom22 in HeLa cells results in the rapid assembly of Tom7 into the 380-kDa complex indicating ..
  32. Bertolin G, Jacoupy M, Traver S, Ferrando Miguel R, Saint Georges T, Grenier K, et al. Parkin maintains mitochondrial levels of the protective Parkinson's disease-related enzyme 17-β hydroxysteroid dehydrogenase type 10. Cell Death Differ. 2015;22:1563-76 pubmed publisher
    ..The loss of this protective mechanism may contribute to mitochondrial dysfunction and neuronal degeneration in autosomal recessive PD. ..
  33. Zhang J, Li M, Zhang Z, Zhu R, Olcese R, Stefani E, et al. The mitochondrial BKCa channel cardiac interactome reveals BKCa association with the mitochondrial import receptor subunit Tom22, and the adenine nucleotide translocator. Mitochondrion. 2017;33:84-101 pubmed publisher
    ..A directed proteomic approach discovered the novel interaction of BKCa with Tom22, a component of the mitochondrion outer membrane import system, and the adenine nucleotide translocator (ANT)...
  34. Di Maio R, Barrett P, Hoffman E, Barrett C, Zharikov A, Borah A, et al. ?-Synuclein binds to TOM20 and inhibits mitochondrial protein import in Parkinson's disease. Sci Transl Med. 2016;8:342ra78 pubmed publisher
    ..This binding prevented the interaction of TOM20 with its co-receptor, TOM22, and impaired mitochondrial protein import...
  35. Saeki K, Suzuki H, Tsuneoka M, Maeda M, Iwamoto R, Hasuwa H, et al. Identification of mammalian TOM22 as a subunit of the preprotein translocase of the mitochondrial outer membrane. J Biol Chem. 2000;275:31996-2002 pubmed
    ..1C9-2 had 19-20% sequence identity to fungal Tom22, a component of the preprotein translocase of the outer membrane (the TOM complex) with receptor and organizer ..
  36. Rumlova M, Krízová I, Keprová A, Hadravová R, Dolezal M, Strohalmová K, et al. HIV-1 protease-induced apoptosis. Retrovirology. 2014;11:37 pubmed publisher
    ..In addition, in vitro data demonstrated that HIV-1 PR mediates cleavage of mitochondrial proteins Tom22, VDAC and ANT, leading to release of AIF and Hsp60 proteins...
  37. Lazarou M, Jin S, Kane L, Youle R. Role of PINK1 binding to the TOM complex and alternate intracellular membranes in recruitment and activation of the E3 ligase Parkin. Dev Cell. 2012;22:320-33 pubmed publisher
    ..We propose that the association of PINK1 with the TOM complex allows rapid reimport of PINK1 to rescue repolarized mitochondria from mitophagy, and discount mitochondrial-specific factors for Parkin translocation and activation. ..
  38. Rajapaksha M, Kaur J, Prasad M, Pawlak K, Marshall B, Perry E, et al. An Outer Mitochondrial Translocase, Tom22, Is Crucial for Inner Mitochondrial Steroidogenic Regulation in Adrenal and Gonadal Tissues. Mol Cell Biol. 2016;36:1032-47 pubmed publisher
    ..We report that the mitochondrial translocase Tom22 is essential for metabolic conversion, as its knockdown by small interfering RNA (siRNA) completely ablated ..
  39. Wu X, Li L, Jiang H. Mitochondrial inner-membrane protease Yme1 degrades outer-membrane proteins Tom22 and Om45. J Cell Biol. 2018;217:139-149 pubmed publisher
    ..Our findings indicate a proteolysis pathway monitoring MOM proteins from the IMS side and suggest that the MOM proteome is surveilled by mitochondrial and cytoplasmic quality control machineries in parallel. ..
  40. Humphries A, Streimann I, Stojanovski D, Johnston A, Yano M, Hoogenraad N, et al. Dissection of the mitochondrial import and assembly pathway for human Tom40. J Biol Chem. 2005;280:11535-43 pubmed
    ..Nevertheless, we show that Tom40 assembly is reduced in mitochondria depleted of human Sam50. These findings are discussed in context with current models from fungal studies. ..