TMEM126A is a mitochondrial located mRNA (MLR) protein of the mitochondrial inner membrane

Biochimica et Biophysica Acta - General Subjects

Volumn 1830, Issue 6, 2013, Pages 3719-3733

  Hanein, Sylvain ^a   Garcia, Mathilde ^b   Fares Taie, Lucas ^a   Serre, Valérie ^a,c   De Keyzer, Yves ^a   Delaveau, Thierry ^b   Perrault, Isabelle ^a   Delphin, Nathalie ^a   Gerber, Sylvie ^a   Schmitt, Alain ^c,d,e   Masse, Jean Marc ^c,d,e   Munnich, Arnold ^a   Kaplan, Josseline ^a   Devaux, Frédéric ^b   Rozet, Jean Michel ^a  

^a INSERM   (France)

^b UMR 7238   (France)

^c UNIVERSITÉ PARIS DESCARTES   (France)

^d INSTITUT COCHIN   (France)

^e CNRS   (France)

Author keywords

Cristae; Mitochondria localizedmRNA; Mitochondrial inner membrane; MLR; Optic neuropathy; TMEM126A

Indexed keywords

MEMBRANE PROTEIN; MESSENGER RNA; TRANSMEMBRANE PROTEIN 126A; UNCLASSIFIED DRUG;

ARTICLE; CELL ASSAY; CELL COMPARTMENTALIZATION; CELL FRACTIONATION; CELL SPECIFICITY; CELL SURFACE; CELLULAR DISTRIBUTION; CONFOCAL MICROSCOPY; CONTROLLED STUDY; FLUORESCENCE IN SITU HYBRIDIZATION; GENE EXPRESSION; GENE FUNCTION; GENE IDENTIFICATION; IMMUNOFLUORESCENCE TEST; MITOCHONDRIAL GENE; MITOCHONDRIAL MEMBRANE; MOLECULAR DYNAMICS; PRIORITY JOURNAL; PROTEIN DEGRADATION; PROTEIN DETERMINATION; PROTEIN EXPRESSION; PROTEIN FUNCTION; PROTEIN LOCALIZATION; QUANTITATIVE ANALYSIS; TRANSMEMBRANE PROTEIN 126A GENE; TRANSMISSION ELECTRON MICROSCOPY;

ANIMALS; CERCOPITHECUS AETHIOPS; COS CELLS; GENETIC DISEASES, INBORN; GTP PHOSPHOHYDROLASES; HUMANS; MEMBRANE PROTEINS; MITOCHONDRIAL MEMBRANES; MITOCHONDRIAL PROTEINS; MUTATION; OPTIC NERVE DISEASES; PROTEIN BIOSYNTHESIS; PROTEIN STRUCTURE, TERTIARY; RETINAL GANGLION CELLS; RNA, MESSENGER;

EID: 84876011799     PISSN: 03044165     EISSN: 18728006     Source Type: Journal    
DOI: 10.1016/j.bbagen.2013.02.025     Document Type: Article

References (85)

1. V. Carelli, F.N. Ross-Cisneros, and A.A. Sadun Mitochondrial dysfunction as a cause of optic neuropathies Prog. Retin. Eye Res. 23 2004 53 89
2. V. Carelli, C. La Morgia, M.L. Valentino, P. Barboni, F.N. Ross-Cisneros, and A.A. Sadun Retinal ganglion cell neurodegeneration in mitochondrial inherited disorders Biochim. Biophys. Acta 1787 2009 518 528
3. P. Yu-Wai-Man, P.G. Griffiths, and P.F. Chinnery Mitochondrial optic neuropathies - disease mechanisms and therapeutic strategies Prog. Retin. Eye Res. 30 2011 81 114
4. P. Amati-Bonneau, D. Milea, D. Bonneau, A. Chevrollier, M. Ferré, V. Guillet, N. Gueguen, D. Loiseau, M.A. de Crescenzo, C. Verny, V. Procaccio, G. Lenaers, and P. Reynier OPA1-associated disorders: phenotypes and pathophysiology Int. J. Biochem. Cell Biol. 41 2009 1855 1865
5. S. Hanein, I. Perrault, O. Roche, S. Gerber, N. Khadom, M. Rio, N. Boddaert, M. Jean-Pierre, N. Brahimi, V. Serre, D. Chretien, N. Delphin, L. Fares-Taie, S. Lachheb, A. Rotig, F. Meire, A. Munnich, J.L. Dufier, J. Kaplan, and J.M. Rozet TMEM126A, encoding a mitochondrial protein, is mutated in autosomal-recessive nonsyndromic optic atrophy Am. J. Hum. Genet. 84 2009 493 498
6. E. Meyer, M. Michaelides, L.J. Tee, A.G. Robson, F. Rahman, S. Pasha, L.M. Luxon, A.T. Moore, and E.R. Maher Nonsense mutation in TMEM126A causing autosomal recessive optic atrophy and auditory neuropathy Mol. Vis. 16 2010 650 664
7. J. Désir, F. Coppieters, N. Van Regemorter, E. De Baere, M. Abramowicz, and M. Cordonnier TMEM126A mutation in a Moroccan family with autosomal recessive optic atrophy Mol. Vis. 18 2012 1849 1857
8. J.J. Assink, N.T. Tijmes, J.B. ten Brink, R.J. Oostra, F.C. Riemslag, P.T. de Jong, and A.A. Bergen A gene for X-linked optic atrophy is closely linked to the Xp11.4-Xp11.2 region of the X chromosome Am. J. Hum. Genet. 61 1997 934 939
9. J.B. Kerrison, V.J. Arnould, J.M. Ferraz Sallum, M.R. Vagefi, M.M. Barmada, Y. Li, D. Zhu, and I.H. Maumenee Genetic heterogeneity of dominant optic atrophy, Kjer type: identification of a second locus on chromosome 18q12.2-12.3 Arch. Ophthalmol. 117 1999 805 810
10. F. Barbet, S. Hakiki, C. Orssaud, S. Gerber, I. Perrault, S. Hanein, D. Ducroq, J.L. Dufier, A. Munnich, J. Kaplan, and J.M. Rozet A third locus for dominant optic atrophy on chromosome 22q J. Med. Genet. 42 2005 e1
11. F. Barbet, S. Gerber, S. Hakiki, I. Perrault, S. Hanein, D. Ducroq, G. Tanguy, J.L. Dufier, A. Munnich, J.M. Rozet, and J. Kaplan A first locus for isolated autosomal recessive optic atrophy (ROA1) maps to chromosome 8q Eur. J. Hum. Genet. 11 2003 966 971
12. V. Carelli, S. Schimpf, N. Fuhrmann, M.L. Valentino, C. Zanna, L. Iommarini, M. Papke, S. Schaich, S. Tippmann, B. Baumann, P. Barboni, L. Longanesi, M. Rugolo, A. Ghelli, M.V. Alavi, R.J. Youle, L. Bucchi, R. Carroccia, M.P. Giannoccaro, C. Tonon, R. Lodi, G. Cenacchi, P. Montagna, R. Liguori, and B. Wissinger A clinically complex form of dominant optic atrophy (OPA8) maps on chromosome 16 Hum. Mol. Genet. 20 2011 1893 1905
13. A. Olichon, L.J. Emorine, E. Descoins, L. Pelloquin, L. Brichese, N. Gas, E. Guillou, C. Delettre, A. Valette, C.P. Hamel, B. Ducommun, G. Lenaers, and P. Belenguer The human dynamin-related protein OPA1 is anchored to the mitochondrial inner membrane facing the inter-membrane space FEBS Lett. 523 2002 171 176
14. G. Lenaers, P. Reynier, G. Elachouri, C. Soukkarieh, A. Olichon, P. Belenguer, L. Baricault, B. Ducommun, C. Hamel, and C. Delettre OPA1 functions in mitochondria and dysfunctions in optic nerve Int. J. Biochem. Cell Biol. 41 2009 1866 1874
15. J. Stone, D. van Driel, K. Valter, S. Rees, and J. Provis The locations of mitochondria in mammalian photoreceptors: relation to retinal vasculature Brain Res. 1189 2008 58 69
16. P. Marc, A. Margeot, F. Devaux, C. Blugeon, M. Corral-Debrinski, and C. Jacq Genome-wide analysis of mRNAs targeted to yeast mitochondria EMBO Rep. 3 2002 159 164
17. A. Margeot, M. Garcia, W. Wang, E. Tetaud, J.P. di Rago, and C. Jacq Why are many mRNAs translated to the vicinity of mitochondria: a role in protein complex assembly? Gene 354 2005 64 71
18. M. Garcia, X. Darzacq, T. Delaveau, L. Jourdren, R.H. Singer, and C. Jacq Mitochondria-associated yeast mRNAs and the biogenesis of molecular complexes Mol. Biol. Cell 18 2007 362 368
19. Y. Saint-Georges, M. Garcia, T. Delaveau, L. Jourdren, S. Le Crom, S. Lemoine, V. Tanty, F. Devaux, and C. Jacq Yeast mitochondrial biogenesis: a role for the PUF RNA-binding protein Puf3p in mRNA localization PLoS One 3 2008 e2293
20. G. Egea, J.M. Izquierdo, J. Ricart, C. San Martín, and J.M. Cuezva mRNA encoding the beta-subunit of the mitochondrial F1-ATPase complex is a localized mRNA in rat hepatocytes Biochem. J. 322 1997 557 565
21. J. Ricart, G. Egea, J.M. Izquierdo, C. San Martín, and J.M. Cuezva Subcellular structure containing mRNA for beta subunit of mitochondrial H + -ATP synthase in rat hepatocytes is translationally active Biochem. J. 324 1997 635 643
22. T. Lithgow, J.M. Cuezva, and P.A. Silver Highways for protein delivery to the mitochondria Trends Biochem. Sci. 22 1997 110 113
23. A.E. Gioio, M. Eyman, H. Zhang, Z.S. Lavina, A. Giuditta, and B.B. Kaplan Local synthesis of nuclear-encoded mitochondrial proteins in the presynaptic nerve terminal J. Neurosci. Res. 64 2001 447 453
24. J. Sylvestre, S. Vialette, M. Corral Debrinski, and C. Jacq Long mRNAs coding for yeast mitochondrial proteins of prokaryotic origin preferentially localize to the vicinity of mitochondria Genome Biol. 4 2003 R44
25. J. Sylvestre, A. Margeot, C. Jacq, G. Dujardin, and M. Corral-Debrinski The role of the 3′ untranslated region in mRNA sorting to the vicinity of mitochondria is conserved from yeast to human cells Mol. Biol. Cell 14 2003 3848 3856
26. S. Matsumoto, T. Uchiumi, T. Saito, M. Yagi, S. Takazaki, T. Kanki, and D. Kang Localization of mRNAs encoding human mitochondrial oxidative phosphorylation proteins Mitochondrion 12 2012 391 398
27. M. Garcia, X. Darzacq, F. Devaux, R.H. Singers, and C. Jacq Yeast mitochondrial transcriptomics Methods Mol. Biol. 372 2007 505 528
28. L. Jourdren, T. Delaveau, E. Marquenet, C. Jacq, and M. Garcia CORSEN, a new software dedicated to microscope-based 3D distance measurements: mRNA-mitochondria distance, from single-cell to population analyses RNA 16 2010 1301 1307
29. E. Lécuyer, H. Yoshida, N. Parthasarathy, C. Alm, T. Babak, T. Cerovina, T.R. Hughes, P. Tomancak, and H.M. Krause Global analysis of mRNA localization reveals a prominent role in organizing cellular architecture and function Cell 131 2007 174 187
30. B. Rost, G. Yachdav, and J. Liu The PredictProtein server Nucleic Acids Res. 32 2004 W321 326
31. K. Hofmann, and W. Stoffel TMbase - a database of membrane spanning proteins segments Biol. Chem. Hoppe-Seyler 374 1993 166
32. G.E. Tusnády, and I. Simon Principles governing amino acid composition of integral membrane proteins: application to topology prediction J. Mol. Biol. 283 1998 489 506
33. M. Cserzö, E. Wallin, I. Simon, G. von Heijne, and A. Elofsson Prediction of transmembrane alpha-helices in prokaryotic membrane proteins: the dense alignment surface method Protein Eng. 10 1997 673 676
34. M.G. Claros, and G. von Heijne TopPred II: an improved software for membrane protein structure predictions CABIOS 10 1994 685 686
35. L. Käll, A. Krogh, and E.L. Sonnhammer A combined transmembrane topology and signal peptide prediction method J. Mol. Biol. 338 2004 1027 1036
36. D.W. Buchan, S.M. Ward, A.E. Lobley, T.C. Nugent, K. Bryson, and D.T. Jones Protein annotation and modelling servers at University College London Nucleic Acids Res. 38 2010 W563 568
37. R.J. Janssen, L.G. Nijtmans, L.P. van den Heuvel, and J.A. Smeitink Mitochondrial complex I: structure, function and pathology J. Inherit. Metab. Dis. 29 2006 499 515
38. E. Blachly-Dyson, E.B. Zambronicz, W.H. Yu, V. Adams, E.R. McCabe, J. Adelman, M. Colombini, and M. Forte Cloning and functional expression in yeast of two human isoforms of the outer mitochondrial membrane channel, the voltage-dependent anion channel J. Biol. Chem. 268 1993 1835 1841
39. V. Sampson, and T. Alleyne Cytochrome c/cytochrome c oxidase interaction. Direct structural evidence for conformational changes during enzyme turnover Eur. J. Biochem. 268 2001 6534 6544
40. H.H. Dahl, S.M. Hunt, W.M. Hutchison, and G.K. Brown The human pyruvate dehydrogenase complex. Isolation of cDNA clones for the E1 alpha subunit, sequence analysis, and characterization of the mRNA J. Biol. Chem. 262 1987 7398 7403
41. J. Yang, X. Liu, K. Bhalla, C.N. Kim, A.M. Ibrado, J. Cai, T.I. Peng, D.P. Jones, and X. Wang Prevention of apoptosis by Bcl-2: release of cytochrome c from mitochondria blocked Science 275 1997 1129 1132
42. C.M. Ott, and V.R. Lingappa Integral membrane protein biosynthesis: why topology is hard to predict J. Cell Sci. 115 2002 2003 2009
43. L.A. Kelley, and M.J. Sternberg Protein structure prediction on the Web: a case study using the Phyre server Nat. Protoc. 4 2009 363 371
44. K. Karplus SAM-T08, HMM-based protein structure prediction Nucleic Acids Res. 37 2009 W492 497
45. C. Cole, J.D. Barber, and G.J. Barton The Jpred 3 secondary structure prediction server Nucleic Acids Res. 36 2008 W197 201
46. R. Gautier, D. Douguet, B. Antonny, and G. Drin HELIQUEST: a web server to screen sequences with specific alpha-helical properties Bioinformatics 24 2008 2101 2102
47. S.J. Habib, W. Neupert, and D. Rapaport Analysis and prediction of mitochondrial targeting signals Methods Cell Biol. 80 2007 761 781
48. M.G. Claros, and P. Vincens Computational method to predict mitochondrially imported proteins and their targeting sequences Eur. J. Biochem. 241 1996 779 786
49. O. Emanuelsson, S. Brunak, G. von Heijne, and H. Nielsen Locating proteins in the cell using TargetP, SignalP and related tools Nat. Protoc. 2 2007 953 971
50. S. Bolte, and F.P. Cordelières A guided tour into subcellular colocalization analysis in light microscopy J. Microsc. 224 2006 213 232
51. S. Di Donato Multisystem manifestations of mitochondrial disorders J. Neurol. 256 2009 693 710
52. A.H. Schapira Mitochondrial diseases Lancet 379 2012 1825 1834
53. K.S. Sitarz, P.F. Chinnery, and P. Yu-Wai-Man Disorders of the optic nerve in mitochondrial cytopathies: new ideas on pathogenesis and therapeutic targets Curr. Neurol. Neurosci. Rep. 12 2012 308 317
54. P. Yu-Wai-Man, P.G. Griffiths, G.S. Gorman, C.M. Lourenco, A.F. Wright, M. Auer-Grumbach, A. Toscano, O. Musumeci, M.L. Valentino, L. Caporali, C. Lamperti, C.M. Tallaksen, P. Duffey, J. Miller, R.G. Whittaker, M.R. Baker, M.J. Jackson, M.P. Clarke, B. Dhillon, B. Czermin, J.D. Stewart, G. Hudson, P. Reynier, D. Bonneau, W. Marques Jr., G. Lenaers, R. McFarland, R.W. Taylor, D.M. Turnbull, M. Votruba, M. Zeviani, V. Carelli, L.A. Bindoff, R. Horvath, P. Amati-Bonneau, and P.F. Chinnery Multi-system neurological disease is common in patients with OPA1 mutations Brain 133 2010 771 786
55. G. Lelandais, Y. Saint-Georges, C. Geneix, L. Al-Shikhley, G. Dujardin, and C. Jacq Spatio-temporal dynamics of yeast mitochondrial biogenesis: transcriptional and post-transcriptional mRNA oscillatory modules PLoS Comput. Biol. 5 2009 e1000409
56. A.P. Gerber, D. Herschlag, and P.O. Brown Extensive association of functionally and cytotopically related mRNAs with Puf family RNA-binding proteins in yeast PLoS Biol. 2 2004 E79
57. M. Wickens, D.S. Bernstein, J. Kimble, and R. Parker A PUF family portrait: 3′UTR regulation as a way of life Trends Genet. 18 2002 150 157
58. L.J. Garcia-Rodriguez, A.C. Gay, and L.A. Pon Puf3p, a Pumilio family RNA binding protein, localizes to mitochondria and regulates mitochondrial biogenesis and motility in budding yeast J. Cell Biol. 176 2007 197 207
59. A. Galgano, M. Forrer, L. Jaskiewicz, A. Kanitz, M. Zavolan, and A.P. Gerber Comparative analysis of mRNA targets for human PUF-family proteins suggests extensive interaction with the miRNA regulatory system PLoS One 3 2008 e3164
60. A.D. Ortega, I.M. Willers, S. Sala, and J.M. Cuezva Human G3BP1 interacts with beta-F1-ATPase mRNA and inhibits its translation J. Cell Sci. 123 2010 2685 2696
61. R.A. Stuart, and W. Neupert Topogenesis of inner membrane proteins of mitochondria Trends Biochem. Sci. 21 1996 261 267
62. B. Martoglio, and B. Dobberstein Signal sequences: more than just greasy peptides Trends Cell Biol. 8 1998 410 415
63. K. Diekert, G. Kispal, B. Guiard, and R. Lill An internal targeting signal directing proteins into the mitochondrial intermembrane space Proc. Natl. Acad. Sci. U. S. A. 12 1999 11752 11757
64. C.M. Lee, J. Sedman, W. Neupert, and R.A. Stuart The DNA helicase, Hmi1p, is transported into mitochondria by a C-terminal cleavable targeting signal J. Biol. Chem. 274 1999 20937 20942
65. A. Olichon, L. Baricault, N. Gas, E. Guillou, A. Valette, P. Belenguer, and G. Lenaers Loss of OPA1 perturbates the mitochondrial inner membrane structure and integrity, leading to cytochrome c release and apoptosis J. Biol. Chem. 278 2003 7743 7746
66. C. Frezza, S. Cipolat, O. Martins de Brito, M. Micaroni, G.V. Beznoussenko, T. Rudka, D. Bartoli, R.S. Polishuck, N.N. Danial, B. De Strooper, and L. Scorrano OPA1 controls apoptotic cristae remodeling independently from mitochondrial fusion Cell 126 2006 177 189
67. C. Zanna, A. Ghelli, A.M. Porcelli, M. Karbowski, R.J. Youle, S. Schimpf, B. Wissinger, M. Pinti, A. Cossarizza, S. Vidoni, M.L. Valentino, M. Rugolo, and V. Carelli OPA1 mutations associated with dominant optic atrophy impair oxidative phosphorylation and mitochondrial fusion Brain 131 2008 352 367
68. M.V. Alavi, S. Bette, S. Schimpf, F. Schuettauf, U. Schraermeyer, H.F. Wehrl, L. Ruttiger, S.C. Beck, F. Tonagel, B.J. Pichler, M. Knipper, T. Peters, J. Laufs, and B. Wissinger A splice site mutation in the murine Opa1 gene features pathology of autosomal dominant optic atrophy Brain 130 2007 1029 1042
69. K.E. White, V.J. Davies, V.E. Hogan, M.J. Piechota, P.P. Nichols, D.M. Turnbull, and M. Votruba OPA1 deficiency associated with increased autophagy in retinal ganglion cells in a murine model of dominant optic atrophy Invest. Ophthalmol. Vis. Sci. 50 2009 2567 2571
70. F. Ferreirinha, A. Quattrini, M. Pirozzi, V. Valsecchi, G. Dina, V. Broccoli, A. Auricchio, F. Piemonte, G. Tozzi, L. Gaeta, G. Casari, A. Ballabio, and E.I. Rugarli Axonal degeneration in paraplegin-deficient mice is associated with abnormal mitochondria and impairment of axonal transport J. Clin. Invest. 113 2004 231 242
71. G. Casari, M. De Fusco, S. Ciarmatori, M. Zeviani, M. Mora, P. Fernandez, G. De Michele, A. Filla, S. Cocozza, R. Marconi, A. Dürr, B. Fontaine, and A. Ballabio Spastic paraplegia and OXPHOS impairment caused by mutations in paraplegin, a nuclear-encoded mitochondrial metalloprotease Cell 93 1998 973 983
72. S. Klebe, C. Depienne, S. Gerber, G. Challe, M. Anheim, P. Charles, E. Fedirko, E. Lejeune, J. Cottineau, A. Brusco, H. Dollfus, P.F. Chinnery, C. Mancini, X. Ferrer, G. Sole, A. Destée, J.M. Mayer, B. Fontaine, J. de Seze, M. Clanet, E. Ollagnon, P. Busson, C. Cazeneuve, G. Stevanin, J. Kaplan, J.M. Rozet, A. Brice, and A. Durr Spastic paraplegia gene 7 in patients with spasticity and/or optic neuropathy Brain 135 2012 2980 2993
73. K.L. van Gassen, C.D. van der Heijden, S.T. de Bot, W.F. den Dunnen, L.H. van den Berg, C.C. Verschuuren-Bemelmans, H.P. Kremer, J.H. Veldink, E.J. Kamsteeg, H. Scheffer, and B.P. van de Warrenburg Genotype-phenotype correlations in spastic paraplegia type 7: a study in a large Dutch cohort Brain 135 2012 2994 3004
74. M.T. Ryan, and N.J. Hoogenraad Mitochondrial-nuclear communications Annu. Rev. Biochem. 76 2007 701 722
75. J.S. Bae, J.K. Choi, J.H. Moon, E.C. Kim, M. Croft, and H.W. Lee Novel transmembrane protein 126A (TMEM126A) couples with CD137L reverse signals in myeloid cells Cell. Signal. 24 2012 2227 2236
76. N. Korfali, G.S. Wilkie, S.K. Swanson, V. Srsen, D.G. Batrakou, E.A. Fairley, P. Malik, N. Zuleger, A. Goncharevich, J. de Las Heras, D.A. Kelly, A.R. Kerr, L. Florens, and E.C. Schirmer The leukocyte nuclear envelope proteome varies with cell activation and contains novel transmembrane proteins that affect genome architecture Mol. Cell Proteomics 9 2010 2571 2585
77. P.J. Hollenbeck, and W.M. Saxton The axonal transport of mitochondria J. Cell Sci. 118 2005 5411 5419
78. M.A. Sutton, and E.M. Schuman Local translational control in dendrites and its role in long-term synaptic plasticity J. Neurobiol. 64 2005 116 131
79. C.R. Bramham, and D.G. Wells Dendritic mRNA: transport, translation and function Nat. Rev. Neurosci. 8 2007 776 789
80. M. Hillefors, A.E. Gioio, M.G. Mameza, and B.B. Kaplan Axon viability and mitochondrial function are dependent on local protein synthesis in sympathetic neurons Cell. Mol. Neurobiol. 27 2007 701 716
81. L. Wang, J. Dong, G. Cull, B. Fortune, and G.A. Cioffi Varicosities of intraretinal ganglion cell axons in human and nonhuman primates Invest. Ophthalmol. Vis. Sci. 44 2003 2 9
82. R.M. Andrews, P.G. Griffiths, M.A. Johnson, and D.M. Turnbull Histochemical localisation of mitochondrial enzyme activity in human optic nerve and retina Br. J. Ophthalmol. 83 1999 231 235
83. E.A. Bristow, P.G. Griffiths, R.M. Andrews, M.A. Johnson, and D.M. Turnbull The distribution of mitochondrial activity in relation to optic nerve structure Arch. Ophthalmol. 120 2002 791 796
84. E. Lécuyer, H. Yoshida, and H.M. Krause Global implications of mRNA localization pathways in cellular organization Curr. Opin. Cell Biol. 21 2009 409 415
85. R. Ben-Menachem, M. Tal, T. Shadur, and O. Pines A third of the yeast mitochondrial proteome is dual localized: a question of evolution Proteomics 11 2011 4468 4476