Cell and Development Biology

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  • Genge, M.G. and Mokranjac, D. (2022) Coordinated Translocation of Presequence-Containing Precursor Proteins Across Two Mitochondrial Membranes: Knowns and Unknowns of How TOM and TIM23 Complexes Cooperate With Each Other. Front. Physiol. https://doi.org/10.3389/fphys.2021.806426


  • Arduino, D.M., Goh, V., Mokranjac, D., and Perocchi F. (2021) Drug Discovery Assay to Identify Modulators of the Mitochondrial Ca2+ Uniporter. Methods Mol. Biol. 2277:69-89.


  • Günsel, U., Paz, E., Gupta, R., Mathes, I., Azem, A., and Mokranjac, D. (2020) In Vivo Dissection of the Intrinsically Disordered Receptor Domain of Tim23. J. Mol. Biol. 432(10):3326-3337.
  • Mokranjac, D. (2020) How to get to the other side of the mitochondrial inner membrane - the protein import motor. Biol. Chem. 401(6-7):723-736.
  • Günsel, U., and Mokranjac, D. (2019) A journey along the TIM23 complex, the major protein translocase of the mitochondrial inner membrane. Biologia Serbica 41(2): 27-35.


  • Rolland, S.G., Schneid, S., Schwarz, M., Rackles, E., Fischer, C., Haeussler, S., Regmi, S.G., Yeroslaviz, A., Habermann, B., , Lambie, E., and Conradt, B. (2019) Compromised Mitochondrial Protein Import Acts as a Signal for UPRmt. Cell Rep. 28(7): 1659-1669.
  • Dayan, D., Bandel, M., Günsel, U., Nussbaum, I., Prag, G., Mokranjac, D., Neupert, W., and Azem, A. (2019) A mutagenesis analysis of Tim50, the major receptor of the TIM23 complex, identifies regions that affect its interaction with Tim23. Sci. Rep. 9(1): 2012.


  • Wettmarshausen, J., Goh, V., Huang, K.T., Arduino, D.M., Tripathi, U., Leimpek, A., Cheng, Y., Pittis, A.A., Gabaldón, T., Mokranjac, D., Hajnóczky, G, and Perocchi, F. (2018) MICU1 confers protection from MCU-dependent manganese toxicity. Cell Rep. 25(6): 1425-1435.


  • Malhotra, K., Modak, A., Nangia, S., Daman, T.H., Günsel, U., Robinson, V., Mokranjac, D., May, E.R. and Alder, N. (2017) Cardiolipin mediates membrane and channel interactions of the mitochondrial TIM23 protein import complex receptor Tim50. Sci. Adv. 3: e1700532.
  • Arduino, D.M., Wettmarshausen, J., Vais, H., Navas-Navarro, P., Cheng, Y., Leimpek, A., Ma, Z., Delrio-Lorenzo, A., Giordano, A., Garcia-Perez, C., Médard, G., Kuster, B., García-Sancho, J., Mokranjac, D., Foskett, J.K., Alonso, M.T., and Perocchi F. (2017) Systematic identification of MCU modulators by orthogonal interspecies chemical screening. Mol. Cell 67(4): 711-723.
  • Banerjee, R.**, Günsel, U** and Mokranjac D. (2017) Chemical Crosslinking in intact mitochondria. Methods Mol. Biol. 1567: 139-154.
  • Demishtein-Zohary, K.**, Günsel, U.**, Marom, M., Banerjee, R., Neupert, W., Azem, A.* and Mokranjac, D.* (2017) Role of Tim17 in coupling the import motor to the translocation channel of the mitochondrial presequence translocase. eLife e22696.


  • Mokranjac, D. (2016) Mitochondrial protein import: an unexpected disulfide bond. J. Cell Biol. 214(4): 363-365.


  • Banerjee, R., Gladkova, C., Mapa, K., Witte, G., and Mokranjac, D. (2015) Protein translocation channel of mitochondrial inner membrane and matrix-exposed import motor communicate via two-domain coupling protein. eLife e11897.
  • Mokranjac, D.*, and Neupert W.* (2015) Cell biology: Architecture of a protein entry gate. Nature 528(7581): 201-202.
  • Bauer, D., Merz, D.R., Pelz, B., Theisen, K.E., Yacyshyn, G., Mokranjac, D., Dima, R.I., Rief, M., and Žoldák, G. (2015) Nucleotides regulate the mechanical hierarchy between subdomains of the nucleotide binding domain of the Hsp70 chaperone DnaK. Proc. Natl. Acad. Sci. U S A 112(33): 10389-10394.
  • Demishtein-Zohary, K., Marom, M., Neupert, W., Mokranjac, D.*, and Azem, A.* (2015) GxxxG motifs hold the TIM23 complex together. FEBS J. 2824(11): 2178-2186.
  • Mokranjac, D.,* and High, S.* (2015) Editorial overview: elucidation of protein translocation pathways, part II. J. Mol. Biol. 427(6 Pt A): 1133-1134.
  • Weems, E., Singha, U.K., Hamilton, V., Smith, J.T., Waegemann, K., Mokranjac, D., and Chaudhuri, M. (2015) Functional complementation analyses reveal that the single PRAT family protein of Trypanosoma brucei is a divergent homolog of Tim17 in Saccharomyces cerevisiae. Eukaryot. Cell. 14(3): 286-296.
  • Waegemann, K., Popov-Čeleketić, D., Neupert, W., Azem, A., and Mokranjac, D., (2015) Cooperation of TOM and TIM23 complexes during translocation of proteins into mitochondria. J. Mol. Biol. 427(5): 1075-1084.


  • Burbulla, L.F., Fitzgerald, J.C., Stegen, K., Westermeier, .J, Thost, A.K., Kato, H., Mokranjac, D., Sauerwald, J., Martins, L.M., Woitalla, D., Rapaport, D., Riess, O., Proikas-Cezanne, T., Rasse, T.M., Krüger, R. (2014) Mitochondrial proteolytic stress induced by loss of mortalin function is rescued by Parkin and PINK1. Cell Death Dis. 5: e1180.

  • Sikor, M., Mapa, K., von Voithenberg, L.V., Mokranjac, D.*, and Lamb, D.C*. (2013) Real-time observation of the conformational dynamics of mitochondrial Hsp70 by spFRET. EMBO J. 32(11): 1639-1649.

  • Harner, M., Mokranjac, D., and Neupert, W. (2012) Die molekulare Architektur der Mitochondrien. BIOspektrum 06.12: 576-581.
  • Schreiner, B., Westerburg, H., Forné, I., Imhof, A., Neupert, W., and Mokranjac, D. (2012) Role of the AAA protease Yme1 in folding of proteins in the intermembrane space of mitochondria. Mol. Biol. Cell 23(22): 4335-4346.
  • Kudryavtsev, V., Sikor, M., Kalinin, S., Mokranjac, D., Seidel, C.A., and Lamb, D.C. (2012) Combining MFD and PIE for accurate single-pair Förster resonance energy transfer measurements. Chemphyschem. 13(4): 1060-1078.

  • Harner, M., Körner, C., Walther, D., Mokranjac, D., Kaesmacher, J., Welsch, U., Griffith, J., Mann, M., Reggiori, F., and Neupert, W. (2011) The mitochondrial contact site complex, a determinant of mitochondrial architecture. EMBO J. 30(21): 4356-4370.
  • Marom, M., Dayan, D., Demishtein-Zohary, K., Mokranjac, D., Neupert, W., and Azem, A. (2011) Direct interaction of mitochondrial targeting presequences with purified components of the TIM23 complex. J. Biol. Chem. 286(51): 43809-43815.
  • Popov-Čeleketić, D., Waegemann, K., Mapa, K., Neupert, W., and Mokranjac, D. (2011) Role of the import motor in insertion of transmembrane segments by the mitochondrial TIM23 complex. EMBO Reports 12(6): 542-548.
  • Marom, M., Azem. A.*, and Mokranjac, D.* (2011) Understanding the molecular mechanism of protein translocation across the mitochondrial inner membrane: Still a long way to go. Biochim. Biophys. Acta. 1808(3): 990-1001.

  • Mapa, K., Sikor, M., Kudryavtsev, V., Waegemann, K., Kalinin, S., Seidel, C.A.M., Neupert, W., Lamb, D.*, and Mokranjac, D.* (2010) The conformational dynamics of the mitochondrial Hsp70 chaperone. Mol. Cell 38: 89-100.
  • Mokranjac, D.*, and Neupert, W.* (2010) The many faces of the mitochondrial TIM23 complex. Biochim. Biophys. Acta 1797(6-7): 1045-1054.
  • Blamowska, M., Sichting, M., Mapa, K., Mokranjac, D., Neupert, W., and Hell, K. (2010) ATPase domain and interdomain linker play a key role in aggregation of mitochondrial Hsp70 chaperone Ssc1. J. Biol. Chem. 285(7): 4423-4431.

  • Mokranjac, D.*, Sichting, M., Popov-Čeleketić, D., Mapa, K., Gevorkyan-Airapetov, L., Zohary, K., Hell, K., Azem, A., and Neupert, W.* (2009) Role of Tim50 in the transfer of precursor proteins from the outer to the inner membrane of mitochondria. Mol. Biol. Cell 20(05): 1400-1407.
  • Gevorkyan-Airapetov, L., Zohary, K., Popov-Čeleketić, D., Mapa, K., Hell, K., Neupert, W., Azem, A., and Mokranjac, D. (2009) Interaction of Tim23 with Tim50 is essential for protein translocation by the mitochondrial TIM23 complex. J. Biol. Chem. 284(8): 4865-4872.
  • Mokranjac, D., and Neupert, W. (2009) Thirty years of protein translocation into mitochondria: unexpectedly complex and still puzzling. Biochem. Biophys. Acta 1793(1): 33-41.

  • Mokranjac, D., and Neupert, W. (2008) Energetics of protein translocation into mitochondria. Biochem. Biophys. Acta 1777(7-8): 758-762.

  • Mokranjac, D., Berg, A., Adam, A., Neupert, W., and Hell, K. (2007) Association of the Tim14*Tim16 subcomplex with the TIM23 translocase is crucial for function of the mitochondrial protein import motor. J. Biol. Chem. 282(25): 18037-18045.
  • Mokranjac, D., and W. Neupert (2007) Protein import into isolated mitochondria. Methods Mol. Biol. 372: 277-286.
  • Murcha, M.W., Elhafez, D., Lister, R., Tonti-Filippini, J., Baumgartner, M., Philippar, K., Carrie, C., Mokranjac, D., Soll, J., and Whelan, J. (2007) Characterisation of the Preprotein and Amino Acids Transporter Gene Family in Arabidopsis. Plant Physiol. 143(1): 199-212.

  • Mokranjac, D., Bourenkov, G., Hell, K., Neupert, W., and Groll, M. (2006) Structure and function of Tim14 and Tim16, the J and J-like components of the mitochondrial protein import motor. EMBO J. 25(19): 4675-4685.

  • Reichert, A. S., Mokranjac, D., Neupert, W., and Hell, K. (2005) Analysis of Protein-Protein Interactions by Chemical Crosslinking. In: Celis, J. (ed). Cell Biology: Laboratory manual 3rd edition, Academic Press, New York
  • Mokranjac, D., and Neupert, W. (2005) Protein import into mitochondria. Biochem. Soc. Trans. 33(5): 1019-1023.
  • Mokranjac, D., Sichting, M., Popov-Čeleketic, D., Berg, A., Hell, K., and Neupert, W. (2005) The import motor of the yeast mitochondrial TIM23 preprotein translocase contains two different J proteins, Tim14 and Mdj2. J. Biol. Chem. 280(36): 31608-31614.
  • Mokranjac, D.**, Popov- Čeleketić, D.**, Hell, K., and Neupert, W. (2005) Role of Tim21 in mitochondrial translocation contact sites. J. Biol. Chem. 280(25): 23437-23340.
  • Sichting, M., Mokranjac, D., Azem, A., Neupert, W., and Hell, K. (2005) Maintenance of structure and function of mitochondrial Hsp70 chaperones requires the chaperone Hep1. EMBO J. 24(5): 1046-1056.
  • Terziyska, N., Lutz, T., Kozany, C., Mokranjac, D., Mesecke, N., Neupert. W., Herrmann, J.M., and Hell, K. (2005) Mia40, a novel factor for protein import into the intermembrane space of mitochondria is able to bind metal ions. FEBS Lett. 579(1): 179-184.

  • Waizenegger, T., Habib, S.J., Lech, M., Mokranjac, D., Paschen, S.A., Hell, K., Neupert, W., and Rapaport, D. (2004) Tob38, a novel essential component in the biogenesis of -barrel proteins in mitochondria. EMBO Reports 5(7): 704-709.
  • Kozany, C.**, Mokranjac, D.**, Sichting, M., Neupert, W., and Hell, K. (2004) The J-domain related co-chaperone Tim16 is a constituent of the mitochondrial TIM23 preprotein translocase. Nat. Struct. Mol. Biol. 11: 234-241.

  • Mokranjac, D., Sichting, M., Neupert, W., and Hell, K. (2003) Tim14, a novel key component of the import motor of the TIM23 protein translocase of mitochondria. EMBO J. 22: 4945-4956.
  • Mokranjac, D., Paschen, S.A., Kozany, C., Prokisch, H., Hoppins, S.C., Nargang, F.E., Neupert, W. and Hell, K. (2003) Tim50, a novel component of the TIM23 preprotein translocase of mitochondria. EMBO J. 22: 816-825.

  • Steinmetz, L.M., Scharfe, C., Deutschbauer, A.M., Mokranjac, D., Herman, Z.S., Jones, T., Chu, A.M., Giaever, G., Prokisch, H., Oefner, P.J., and Davis, R.W. (2002) Systematic screen for human disease genes in yeast. Nat. Genet. 31: 400-404.