Gene Symbol: TIM9
Description: protein transporter TIM9
Alias: protein transporter TIM9
Species: Saccharomyces cerevisiae S288c

Top Publications

  1. Allen S, Lu H, Thornton D, Tokatlidis K. Juxtaposition of the two distal CX3C motifs via intrachain disulfide bonding is essential for the folding of Tim10. J Biol Chem. 2003;278:38505-13 pubmed
    The TIM10 complex, composed of the homologous proteins Tim10 and Tim9, chaperones hydrophobic proteins inserted at the mitochondrial inner membrane...
  2. Baker M, Mooga V, Guiard B, Langer T, Ryan M, Stojanovski D. Impaired folding of the mitochondrial small TIM chaperones induces clearance by the i-AAA protease. J Mol Biol. 2012;424:227-39 pubmed publisher
    ..We show that no individual cysteine residue is required for the function of Tim9 or Tim10 in yeast and that defective assembly of the small TIMs induces their proteolytic clearance from ..
  3. Sideris D, Petrakis N, Katrakili N, Mikropoulou D, Gallo A, Ciofi Baffoni S, et al. A novel intermembrane space-targeting signal docks cysteines onto Mia40 during mitochondrial oxidative folding. J Cell Biol. 2009;187:1007-22 pubmed publisher
  4. Milenkovic D, Ramming T, Müller J, Wenz L, Gebert N, Schulze Specking A, et al. Identification of the signal directing Tim9 and Tim10 into the intermembrane space of mitochondria. Mol Biol Cell. 2009;20:2530-9 pubmed publisher
    ..In this study, we found that a region within the Tim9 and Tim10 precursors, consisting of only nine amino acid residues, functions as a signal for the engagement of ..
  5. Morgan B, Ang S, Yan G, Lu H. Zinc can play chaperone-like and inhibitor roles during import of mitochondrial small Tim proteins. J Biol Chem. 2009;284:6818-25 pubmed publisher
    ..We propose that zinc plays a chaperone-like role in the cytosol during biogenesis of the small Tim proteins and that the proteins are imported into mitochondria through the apo-forms. ..
  6. Müller J, Milenkovic D, Guiard B, Pfanner N, Chacinska A. Precursor oxidation by Mia40 and Erv1 promotes vectorial transport of proteins into the mitochondrial intermembrane space. Mol Biol Cell. 2008;19:226-36 pubmed
    ..Thus, oxidation driven by Mia40 and Erv1 determines vectorial transport of the precursors into the mitochondrial intermembrane space. ..
  7. Sideris D, Tokatlidis K. Oxidative folding of small Tims is mediated by site-specific docking onto Mia40 in the mitochondrial intermembrane space. Mol Microbiol. 2007;65:1360-73 pubmed
    ..The subunits of the chaperone complex Tim9-Tim10 require Mia40 for their biogenesis...
  8. Curran S, Leuenberger D, Oppliger W, Koehler C. The Tim9p-Tim10p complex binds to the transmembrane domains of the ADP/ATP carrier. EMBO J. 2002;21:942-53 pubmed
    ..The chaperone-like Tim9p-Tim10p complex thus may prevent aggregation of the unfolded carrier proteins in the aqueous intermembrane space. ..
  9. Baker M, Webb C, Stroud D, Palmer C, Frazier A, Guiard B, et al. Structural and functional requirements for activity of the Tim9-Tim10 complex in mitochondrial protein import. Mol Biol Cell. 2009;20:769-79 pubmed publisher
    The Tim9-Tim10 complex plays an essential role in mitochondrial protein import by chaperoning select hydrophobic precursor proteins across the intermembrane space...

More Information


  1. Chacinska A, Pfannschmidt S, Wiedemann N, Kozjak V, Sanjuán Szklarz L, Schulze Specking A, et al. Essential role of Mia40 in import and assembly of mitochondrial intermembrane space proteins. EMBO J. 2004;23:3735-46 pubmed
    ..of Mia40 are selectively inhibited in the import of several small IMS proteins, including the essential proteins Tim9 and Tim10. The import of proteins to the other mitochondrial subcompartments does not depend on functional Mia40...
  2. Rissler M, Wiedemann N, Pfannschmidt S, Gabriel K, Guiard B, Pfanner N, et al. The essential mitochondrial protein Erv1 cooperates with Mia40 in biogenesis of intermembrane space proteins. J Mol Biol. 2005;353:485-92 pubmed
    ..Moreover, Erv1 associated with Mia40 in a reductant-sensitive manner. We conclude that two essential proteins, Mia40 and Erv1, cooperate in the assembly pathway of small proteins of the mitochondrial IMS. ..
  3. Milenkovic D, Gabriel K, Guiard B, Schulze Specking A, Pfanner N, Chacinska A. Biogenesis of the essential Tim9-Tim10 chaperone complex of mitochondria: site-specific recognition of cysteine residues by the intermembrane space receptor Mia40. J Biol Chem. 2007;282:22472-80 pubmed
    ..we systematically analyzed the role of cysteine residues in the biogenesis of the essential IMS chaperone complex Tim9-Tim10...
  4. Peleh V, Cordat E, Herrmann J. Mia40 is a trans-site receptor that drives protein import into the mitochondrial intermembrane space by hydrophobic substrate binding. elife. 2016;5: pubmed publisher
  5. Ivanova E, Jowitt T, Lu H. Assembly of the mitochondrial Tim9-Tim10 complex: a multi-step reaction with novel intermediates. J Mol Biol. 2008;375:229-39 pubmed
    ..The mitochondrial Tim9-Tim10 translocase complex, located in the mitochondrial intermembrane space, plays an essential chaperone-like role ..
  6. Ivanova E, Pang J, Jowitt T, Yan G, Warwicker J, Sutcliffe M, et al. Temperature-dependent study reveals that dynamics of hydrophobic residues plays an important functional role in the mitochondrial Tim9-Tim10 complex. Proteins. 2012;80:602-15 pubmed publisher
    Protein-protein interaction is a fundamental process in all major biological processes. The hexameric Tim9-Tim10 (translocase of inner membrane) complex of the mitochondrial intermembrane space plays an essential chaperone-like role ..
  7. Adam A, Endres M, Sirrenberg C, Lottspeich F, Neupert W, Brunner M. Tim9, a new component of the TIM22.54 translocase in mitochondria. EMBO J. 1999;18:313-9 pubmed
    We have identified Tim9, a new component of the TIM22.54 import machinery, which mediates transport of proteins into the inner membrane of mitochondria...
  8. Kawano S, Yamano K, Naoé M, Momose T, Terao K, Nishikawa S, et al. Structural basis of yeast Tim40/Mia40 as an oxidative translocator in the mitochondrial intermembrane space. Proc Natl Acad Sci U S A. 2009;106:14403-7 pubmed publisher
    ..These results are consistent with the growth phenotypes of Mia40 mutant cells containing Ser replacement of the conserved cysteine residues. ..
  9. Rampello A, Glynn S. Identification of a Degradation Signal Sequence within Substrates of the Mitochondrial i-AAA Protease. J Mol Biol. 2017;429:873-885 pubmed publisher
    ..the yeast i-AAA protease, Yme1p, to examine the in vitro degradation of two intermembrane space chaperone subunits, Tim9 and Tim10...
  10. Grossmann J, Callaghan A, Marcaida M, Luisi B, Alcock F, Tokatlidis K, et al. Complementing structural information of modular proteins with small angle neutron scattering and contrast variation. Eur Biophys J. 2008;37:603-11 pubmed publisher
    ..contrast variation of the multi-protein TIM10 complex, a mitochondrial chaperone assembly comprising the subunits Tim9 and Tim10, has been used to determine a low-resolution shape reconstruction of the complex, highlighting the ..
  11. Morgan B, Lu H. Oxidative folding competes with mitochondrial import of the small Tim proteins. Biochem J. 2008;411:115-22 pubmed
    ..In the present study, the effect of glutathione on the cysteine-redox state of Tim9 was investigated, and the standard redox potential of Tim9 was determined to be approx. -0.31 V at pH 7...
  12. Vergnolle M, Baud C, Golovanov A, Alcock F, Luciano P, Lian L, et al. Distinct domains of small Tims involved in subunit interaction and substrate recognition. J Mol Biol. 2005;351:839-49 pubmed
    b>Tim9 and Tim10 belong to the small Tim family of mitochondrial ATP-independent chaperones. They are organised in a specific hetero-oligomeric complex (TIM10) that escorts polytopic proteins into the mitochondrial inner membrane...
  13. Koehler C, Murphy M, Bally N, Leuenberger D, Oppliger W, Dolfini L, et al. Tim18p, a new subunit of the TIM22 complex that mediates insertion of imported proteins into the yeast mitochondrial inner membrane. Mol Cell Biol. 2000;20:1187-93 pubmed
    ..We suggest that Tim18p functions in the assembly and stabilization of the TIM22 complex but does not directly participate in protein insertion into the inner membrane. ..
  14. Baud C, De Marcos Lousa C, Tokatlidis K. Molecular interactions of the mitochondrial Tim12 translocase subunit. Protein Pept Lett. 2007;14:597-600 pubmed
    ..We purified Tim12 and found it forms dimers that bind to Tim9. In this interaction, Tim12 undergoes structural changes that may be important for transport of its substrates in ..
  15. Senapin S, Chen X, Clark Walker G. Transcription of TIM9, a new factor required for the petite-positive phenotype of Saccharomyces cerevisiae, is defective in spt7 mutants. Curr Genet. 2003;44:202-10 pubmed
    b>TIM9 has been identified as an additional novel gene required for the petite-positive phenotype in Saccharomyces cerevisiae...
  16. Klöppel C, Suzuki Y, Kojer K, Petrungaro C, Longen S, Fiedler S, et al. Mia40-dependent oxidation of cysteines in domain I of Ccs1 controls its distribution between mitochondria and the cytosol. Mol Biol Cell. 2011;22:3749-57 pubmed publisher
    ..We propose a model in which the Mia40-mediated oxidative folding of domain I controls the cellular distribution of Ccs1 and, consequently, active Sod1. ..
  17. Leuenberger D, Bally N, Schatz G, Koehler C. Different import pathways through the mitochondrial intermembrane space for inner membrane proteins. EMBO J. 1999;18:4816-22 pubmed
    ..Yeast mitochondria thus use multiple pathways for escorting hydrophobic inner membrane proteins across the aqueous intermembrane space. ..
  18. Luciano P, Vial S, Vergnolle M, Dyall S, Robinson D, Tokatlidis K. Functional reconstitution of the import of the yeast ADP/ATP carrier mediated by the TIM10 complex. EMBO J. 2001;20:4099-106 pubmed
    ..We report here that Tim9 and Tim10 purified from Escherichia coli can form a complex of the same size as the endogenous complex from yeast ..
  19. Hasson S, Damoiseaux R, Glavin J, Dabir D, Walker S, Koehler C. Substrate specificity of the TIM22 mitochondrial import pathway revealed with small molecule inhibitor of protein translocation. Proc Natl Acad Sci U S A. 2010;107:9578-83 pubmed publisher
    ..proteins into the mitochondrial inner membrane and consists of two intermembrane space chaperone complexes, the Tim9-Tim10 and Tim8-Tim13 complexes...
  20. Gebert N, Gebert M, Oeljeklaus S, von der Malsburg K, Stroud D, Kulawiak B, et al. Dual function of Sdh3 in the respiratory chain and TIM22 protein translocase of the mitochondrial inner membrane. Mol Cell. 2011;44:811-8 pubmed publisher
    ..We conclude that the assembly of Sdh3 with different partner proteins, Sdh4 and Tim18, recruits it to two different mitochondrial membrane complexes with functions in bioenergetics and protein biogenesis, respectively. ..
  21. Spiller M, Guo L, Wang Q, Tran P, Lu H. Mitochondrial Tim9 protects Tim10 from degradation by the protease Yme1. Biosci Rep. 2015;35: pubmed publisher
    ..Tim10 are essential homologue proteins of the mitochondrial intermembrane space (IMS) and form a stable hexameric Tim9-Tim10 complex there...
  22. Koehler C, Merchant S, Oppliger W, Schmid K, Jarosch E, Dolfini L, et al. Tim9p, an essential partner subunit of Tim10p for the import of mitochondrial carrier proteins. EMBO J. 1998;17:6477-86 pubmed
    ..Tim9p is a new subunit of the TIM machinery that guides hydrophobic inner membrane proteins across the aqueous intermembrane space. ..
  23. Sirrenberg C, Endres M, FOLSCH H, Stuart R, Neupert W, Brunner M. Carrier protein import into mitochondria mediated by the intermembrane proteins Tim10/Mrs11 and Tim12/Mrs5. Nature. 1998;391:912-5 pubmed
    ..This interaction of Tim10 and Tim12 with the precursors depends on the presence of divalent metal ions. Both proteins contain a zinc-finger-like motif with four cysteines and bind equimolar amounts of zinc ions. ..
  24. Gebert N, Chacinska A, Wagner K, Guiard B, Koehler C, Rehling P, et al. Assembly of the three small Tim proteins precedes docking to the mitochondrial carrier translocase. EMBO Rep. 2008;9:548-54 pubmed publisher
    ..The soluble Tim9-Tim10 complex transfers hydrophobic precursor proteins through the aqueous intermembrane space to the carrier ..
  25. Stojanovski D, Milenkovic D, Müller J, Gabriel K, Schulze Specking A, Baker M, et al. Mitochondrial protein import: precursor oxidation in a ternary complex with disulfide carrier and sulfhydryl oxidase. J Cell Biol. 2008;183:195-202 pubmed publisher
  26. Vial S, Lu H, Allen S, Savory P, Thornton D, Sheehan J, et al. Assembly of Tim9 and Tim10 into a functional chaperone. J Biol Chem. 2002;277:36100-8 pubmed
    ..We have characterized TIM10 assembly and analyzed the structural properties of its subunits, Tim9 and Tim10...