Vögtle, F. N. et al. Global analysis of the mitochondrial N-proteome identifies a processing peptidase critical for protein stability. Cell 139, 428–439 (2009).

Article  PubMed  Google Scholar 

Endo, T. & Kohda, D. Functions of outer membrane receptors in mitochondrial protein import. Biochim. Biophys. Acta 1592, 3–14 (2002).

Article  CAS  PubMed  Google Scholar 

Lee, W. et al. Molecular basis for maternal inheritance of human mitochondrial DNA. Nat. Genet. 55, 1632–1639 (2023).

Article  PubMed  PubMed Central  Google Scholar 

Backes, S. et al. Tom70 enhances mitochondrial preprotein import efficiency by binding to internal targeting sequences. J. Cell Biol. 217, 1369–1382 (2018).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Kreimendahl, S., Schwichtenberg, J., Günnewig, K., Brandherm, L. & Rassow, J. The selectivity filter of the mitochondrial protein import machinery. BMC Biol. 18, 156 (2020).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Chang, Y.-T. et al. Cytoplasmic ribosomes on mitochondria alter the local membrane environment for protein import. J. Cell Biol. 224, e202407110 (2025).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Gold, V. A., Chroscicki, P., Bragoszewski, P. & Chacinska, A. Visualization of cytosolic ribosomes on the surface of mitochondria by electron cryo‐tomography. EMBO Rep. 18, 1786–1800 (2017).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Krämer, L. et al. MitoStores: chaperone-controlled protein granules store mitochondrial precursors in the cytosol. EMBO J. 42, e112309 (2023).

Article  PubMed  PubMed Central  Google Scholar 

Qin, W., Myers, S. A., Carey, D. K., Carr, S. A. & Ting, A. Y. Spatiotemporally-resolved mapping of RNA binding proteins via functional proximity labeling reveals a mitochondrial mRNA anchor promoting stress recovery. Nat. Commun. 12, 4980 (2021).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Kuzniewska, B. et al. Mitochondrial protein biogenesis in the synapse is supported by local translation. EMBO Rep. 21, e48882 (2020).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Williams, C. C., Jan, C. H. & Weissman, J. S. Targeting and plasticity of mitochondrial proteins revealed by proximity-specific ribosome profiling. Science 346, 748–751 (2014).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Tsuboi, T. et al. Mitochondrial volume fraction and translation duration impact mitochondrial mRNA localization and protein synthesis. eLife 9, e57814 (2020).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Barba-Aliaga, M. et al. eIF5A controls mitoprotein import by relieving ribosome stalling at TIM50 translocase mRNA. J. Cell Biol. 223, e202404094 (2024).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Arceo, X. G., Koslover, E. F., Zid, B. M. & Brown, A. I. Mitochondrial mRNA localization is governed by translation kinetics and spatial transport. PLoS Comput. Biol. 18, e1010413 (2022).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Eliyahu, E. et al. Tom20 mediates localization of mRNAs to mitochondria in a translation-dependent manner. Mol. Cell Biol. 30, 284–294 (2010).

Article  CAS  PubMed  Google Scholar 

Hemono, M. et al. FRIENDLY (FMT) is an RNA binding protein associated with cytosolic ribosomes at the mitochondrial surface. Plant J. 112, 309–321 (2022).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Lesnik, C., Cohen, Y., Atir-Lande, A., Schuldiner, M. & Arava, Y. OM14 is a mitochondrial receptor for cytosolic ribosomes that supports co-translational import into mitochondria. Nat. Commun. 5, 5711 (2014).

Article  CAS  PubMed  Google Scholar 

Saint-Georges, Y. et al. Yeast mitochondrial biogenesis: a role for the PUF RNA-binding protein Puf3p in mRNA localization. PLoS ONE 3, e2293 (2008).

Article  PubMed  PubMed Central  Google Scholar 

Bykov, Y. S., Rapaport, D., Herrmann, J. M. & Schuldiner, M. Cytosolic events in the biogenesis of mitochondrial proteins. Trends Biochem. Sci. 45, 650–667 (2020).

Article  CAS  PubMed  Google Scholar 

Lewis, B. M. et al. LARP4 is an RNA-binding protein that binds nuclear-encoded mitochondrial mRNAs to promote mitochondrial function. RNA 30, 223–239 (2024).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Brar, K. K. et al. PERK–ATAD3A interaction provides a subcellular safe haven for protein synthesis during ER stress. Science 385, eadp7114 (2024).

Article  CAS  PubMed  Google Scholar 

Muthukumar, G. et al. Triaging of α-helical proteins to the mitochondrial outer membrane by distinct chaperone machinery based on substrate topology. Mol. Cell 84, 1101–1119.e9 (2024).

Article  CAS  PubMed  Google Scholar 

Kampinga, H. H. & Craig, E. A. The HSP70 chaperone machinery: J proteins as drivers of functional specificity. Nat. Rev. Mol. Cell Biol. 11, 579–592 (2010).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Döring, K. et al. Profiling Ssb–Nascent chain interactions reveals principles of Hsp70-assisted folding. Cell 170, 298–311.e20 (2017).

Article  PubMed  PubMed Central  Google Scholar 

Caplan, A. J., Cyr, D. M. & Douglas, M. G. YDJ1p facilitates polypeptide translocation across different intracellular membranes by a conserved mechanism. Cell 71, 1143–1155 (1992).

Article  CAS  PubMed  Google Scholar 

Deshaies, R. J., Koch, B. D., Werner-Washburne, M., Craig, E. A. & Schekman, R. A subfamily of stress proteins facilitates translocation of secretory and mitochondrial precursor polypeptides. Nature 332, 800–805 (1988).

Article  CAS  PubMed  Google Scholar 

Drwesh, L. et al. A network of cytosolic (co)chaperones promotes the biogenesis of mitochondrial signal-anchored outer membrane proteins. eLife 11, e77706 (2022).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Jores, T. et al. Cytosolic Hsp70 and Hsp40 chaperones enable the biogenesis of mitochondrial β-barrel proteins. J. Cell Biol. 217, 3091–3108 (2018).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Young, J. C., Hoogenraad, N. J. & Hartl, F. U. Molecular chaperones Hsp90 and Hsp70 deliver preproteins to the mitochondrial import receptor Tom70. Cell 112, 41–50 (2003).

Article  CAS  PubMed  Google Scholar 

Opaliński, Ł. et al. Recruitment of cytosolic J-proteins by TOM receptors promotes mitochondrial protein biogenesis. Cell Rep. 25, 2036–2043.e5 (2018).

Article