(Padova, Italy)

Department of Biomedical Sciences, University of Padova, Via Ugo Bassi 58b, I-35131 Padova, Italy

Place and date of birth Cividale del Friuli (Italy) November 21st, 1953

Phone +39 049 827 6365     Fax +39 049 827 6049


Citizenship Italian

Marital Status Married, one son


High School Diploma Liceo Classico “Paolo Diacono”, Cividale del Friuli (Italy) 1971

M.D. Degree University of Padova (Italy), cum laude 1978


1979 – 1987: Assistant Professor, University of Padova Medical School

1988 – 1999: Associate Professor, University of Padova Medical School

2000 – present: Full Professor, University of Padova Medical School

2001 – 2004: Deputy Dean of the Medical Faculty, University of Padova

2003 – 2009: Director, Department of Biomedical Sciences, University of Padova

2012 – present: Coordinator, Ph.D. Program in Biomedical Sciences, University of Padova

2012 – present: Director, Postgraduate School of Clinical Pathology, University of Padova


1984: EMBO Fellow, University of Helsinki, Finland

1985 – 87: Fogarty Fellow, Massachusetts Institute of Technology, Whitehead Institute for Biomedical Research, Cambridge, Massachusetts, USA

2006 – 2010 and 2014-present: Council Member, Bioenergetics Subgroup, Biophysical Society USA

2006 – present: Socio Corrispondente Residente, Istituto Veneto di Scienze, Lettere ed Arti, Venice

2008: Visiting Professor, Oregon Clinical and Translational Research Institute, Portland, Oregon, USA

2009 – 2011: Council Member, Associazione di Biologia Cellulare e del Differenziamento (ABCD)

2009 – present: Member, Confrérie des Chevaliers de Tastevin, Château Clos Vougeot, Bourgogne, France

2010 – 2013: President, Italian Group of Biomembranes and Bioenergetics

2010 – present: Member, Academia Europaea, Section for Biochemistry and Molecular Biology

2012: Prize for Physiology and Pathology, Ministro Beni e Attività Culturali, Accademia Nazionale dei Lincei, Roma


Paolo Bernardi began his studies on mitochondrial physiology and ion transport under the guidance of Giovanni Felice Azzone, one of the founding Fathers of Bioenergetics. His education in Cellular and Molecular Biology was completed with a long-term stay at the Whitehead Institute for Biomedical Research – Massachusetts Institute of Technology, where he worked under the supervision of Harvey F. Lodish. He pioneered the field of mitochondrial channels and their role in cellular pathophysiology. In particular, he focused on the permeability transition pore (PTP), a high conductance channel that is increasingly recognized as a key player in cell death. During the early 1990s he defined key points of regulation of the PTP in isolated mitochondria (membrane potential, matrix pH, Me2+-binding sites, specific redox-sensitive sites). He then developed tools to reliably monitor mitochondrial function in situ, and addressed mechanistic questions on the PTP as a target in degenerative diseases and cancer. His studies have been extended to in vivo models, and led to the demonstration that early mitochondrial adaptation plays a key role in hepatocarcinogenesis [Klöhn et al. (2003) Proc Natl Acad Sci USA 100, 10014-10019] and in onset of the Warburg effect [Sciacovelli et al. (2013) Cell Metab 17, 988-999]; and that mitochondrial dysfunction mediated by the PTP unexpectedly causes muscular dystrophy in collagen VI deficiency [Irwin et al. (2003) Nat Genet 35, 367-371; Angelin et al. (2007) Proc Natl Acad Sci USA 104, 991-996; Merlini et al. (2008) Proc Natl Acad Sci USA 105, 5225-5229]. These studies paved the way to a potential therapy of Ullrich Congenital Muscular Dystrophy and Bethlem Myopathy with NIM811, a non immunosuppressive analog of cyclosporin A [Zulian et al. (2014) Hum Mol Genet 23, 5353-5363]. The recent identification of the PTP, which appears to originate from dimers of the FOF1 ATP synthase [Giorgio et al. (2013) Proc Natl Acad Sci USA 110, 5887-5892; Carraro et al. (2014) J Biol Chem 23, 15980-15985; von Stockum et al (2015) J Biol Chem 290, 4537-4544], offers great promise for further molecular definition of the pore and of its function in health and disease. Development of novel chemical inhibitors of the PTP with potential use in degenerative diseases developed with NIH funding is currently one of the most successful research programs of the Bernardi lab.

Paolo Bernardi was a major actor in the Mitochondrial Renaissance of the 1990s. As early as 1992 he was one of the few to realise the importance of mitochondria in cell death well before the role of cytochrome c release was shown to be a key event in apoptosis. He pioneered the field rapidly reaching international recognition, as testified by 199 invited lectures at meetings and 129 seminars at prestigious Institutions worldwide. He has organized or coorganized key meetings on mitochondrial pathophysiology that significantly contributed to the continuing success of the field. His achievements were possible through the work and training of junior personnel (from 1991 Prof. Bernardi has supervised the work of 27 Graduate Students from the University of Padova, 4 visiting Graduate Students from other Universities and 20 postdoctoral Fellows).


1. Chief Specialty Editor, Frontiers in Mitochondrial Research (from 2012).

2. Editorial Board Member (current): Biochimica et Biophysica Acta (from 1999), Pharmacological Research (from 2015), Current Opinion in Physiology (from 2017).

3. Editorial Board Member (past): Journal of Biological Chemistry (1997-2002, 2003-2008, 2010-2015), Archives of Biochemistry and Biophysics (1998-2003), IUBMB Life (2002-2007).

4. Guest Editor: Biochimica et Biophysica Acta “Mitochondria in Cell Death” (1998); BioFactors “New Perspectives in Mitochondrial Research” (1998); IUBMB Life Special Issue “Perspectives in Mitochondrial Research” (2001); Biochimica et Biophysica Acta EBEC Special Issue (2016).

5. Section Editor, Pathobiology of Human Disease: A Dynamic Encyclopedia of Disease Mechanisms, Elsevier (2014).

6. Member of the Education Committee, International Union of Biochemistry and Molecular Biology (1998-2003).


Ministry for the University and Scientific Research – Italy; University of Padova; Telethon – Italy; National Institutes of Health – Public Health Service (USA); AIRC (Italian Association on Cancer Research); Fondazione Cassa di Risparmio di Padova e Rovigo; Leducq Foundation.


As of December, 2017 Prof. Bernardi has published 233 peer-reviewed articles which have received 28,313 citations with an H index of 87 (Google Scholar). His most quoted paper [Bernardi P (1999) Mitochondrial transport of cations: Channels, exchangers and permeability transition. Physiol Rev 79, 1127-1155] has 1,493 citations; his most quoted experimental paper [Basso, E., Fante, L., Fowlkes, J., Petronilli, V. Forte, M.A. and Bernardi, P. (2005) Properties of the Permeability Transition Pore in Mitochondria Devoid of Cyclophilin D, J Biol Chem 280, 18558-18561] ranks 4th with 726 quotes. His “top ten” list gathers over 7,300 quotes.

PUBLICATIONS (last 5 years)

1. Bernardi, P. (2013) The mitochondrial permeability transition pore, Encyclopedia of Biological Chemistry 3, 162-167

2. Pellegrini, C. , Zulian, A., Gualandi, F., Manzati, E., Merlini, L., Michelini, M.E., Benassi, L., Ferlini, A., Sabatelli, P., Bernardi, P., and Maraldi, N.M. (2013) Melanocytes – A novel tool to study mitochondrial dysfunction as a biomarker of Duchenne Muscular Dystrophy, J. Cell. Physiol. 228,1323-1331

3. Bernardi, P. and Giorgio, V. (2013) EBEC2012 – An energetic time in Freiburg, EMBO Rep. 14, 7-9

4. Bernardi, P. and Bonaldo, P. (2013) Mitochondrial Dysfunction and Defective Autophagy in the Pathogenesis of Collagen VI Muscular Dystrophies, Cold Spring Harb. Perspect. Biol. 5, a011387

5. De Palma, S., Leone, R., Grumati, P., Vasso, M., Polishchuk, R., Capitanio, D., Braghetta, P., Bernardi, P., Bonaldo, P. and Gelfi, G. (2013) Changes in Muscle Cell Metabolism and Mechanotransduction are Associated with Myopathic Phenotype in a Mouse Model of Collagen VI Deficiency, PloS One 8, e56716

6. Giorgio, V., von Stockum, S., Antoniel, M., Fabbro, A., Fogolari, F., Forte, M., Glick, G.D., Petronilli, V., Zoratti, M., Szabó, I., Lippe, G., and Bernardi, P. (2013) Dimers of Mitochondrial ATP Synthase Form the Permeability Transition Pore, Proc. Natl. Acad. Sci. USA 110, 5887-5892

7. Sassi, N., Mattarei, A., Azzolini, M., Bernardi, P., Szabó, I., Paradisi, C., Zoratti, M., Biasutto, L. (2013) Mitochondria-targeted resveratrol derivatives act as cytotoxic pro-oxidants, Current Pharm. Des. 20, 172-179

8. Sciacovelli, M., Guzzo, G., Morello, V., Frezza, C., Nannini, N., Calabrese, F., Laudiero, G., Esposito, F., Landriscina, M., Gottlieb, E., Defilippi, P., Bernardi, P.* and Rasola, A.* (2013) The mitochondrial chaperone TRAP1 promotes neoplastic growth by inhibiting succinate dehydrogenase, *Corresponding Authors Cell Metab. 17, 988-999

9. Bernardi, P. (2013) The mitochondrial permeability transition pore: A mystery solved? Front.Physiol. 4, 95.

10. Battigelli, A., Russier, J., Venturelli, E., Fabbro, C., Petronilli, V., Bernardi, P., Da Ros, T., Prato, M. and Bianco, A. (2013) Peptide-based Carbon Nanotubes for Mitochondrial Targeting, Nanoscale 5, 9110-9117

11. Pantic, B., Trevisan, E., Citta, A., Rigobello, M.P., Marin, O., Bernardi, P., Salvatori, S. and Rasola, A. (2013) Myotonic dystrophy protein kinase (DMPK) prevents ROS-induced cell death by assembling a hexokinase II-Src complex on the mitochondrial surface, Cell Death Dis. 4, e858

12. Trinei M., Migliaccio, E., Bernardi, P., Paolucci, F., Pelicci, P, and Giorgio, M. (2013) p66Shc, mitochondria, and the generation of reactive oxygen species. Methods Enzymol. 528, 99-110

13. Bernardi, P. and Rasola, A. (2014) Inner Membrane Permeabilization – The Permeability Transition, Pathobiology of Human Disease (McManus, Linda M. and Mitchell, Richard N., Eds.) Academic Press, San Diego pp. 162-169

14. Ciscato, F., Sciacovelli, M., Villano, G., Turato, C., Bernardi, P., Rasola, A. and Pontisso, P. (2014) SERPINB3 protects from oxidative damage by chemotherapeutics through inhibition of mitochondrial respiratory Complex I, Oncotarget 5, 2418-2427

15. Da-Rè, C., Franzolin, E., Biscontin, A., Piazzesi, A., Pacchioni, B., Gagliani, C., Mazzotta, G., Tacchetti, C., Zordan, M.A., Zeviani, M., Bernardi, P., Bianchi, V., De Pittà, C. and Costa, R. (2014) Functional characterization of drim2, the Drosophila melanogaster homolog of the yeast mitochondrial deoxynucleotide transporter, J. Biol. Chem. 289, 7448-7459

16. Nowikovsky, K. and Bernardi, P. (2014) LETM1 in mitochondrial cation transport, Front. Physiol., 5:83

17. Da-Rè, C., De Pittà, C., Zordan, M.A., Zeviani, M., Teza, G., Nestola, F., Costa, R., and Bernardi, P. (2014) UCP4C mediates uncoupled respiration in larvae of Drosophila melanogaster, EMBO Rep. 15, 586–591

18. Villano, G., Turato, C., Quarta, S., Ruvoletto, M., Ciscato, F., Terrin, L., Semeraro, R., Paternostro, C., Parola, M., Alvaro, D., Bernardi, P., Gatta, A. and Pontisso, P. (2014) Hepatic progenitor cells express SerpinB3. BMC Cell Biol. 15, 5

19. Šileikytė, J., Blachly-Dyson, E., Sewell, R., Ricchelli, F., Bernardi, P.* and Forte, M.* (2014) Regulation of the Mitochondrial Permeability Transition Pore by the Outer Membrane does not Involve the Peripheral Benzodiazepine Receptor (TSPO), *Corresponding Authors, J. Biol. Chem. 289, 13769-13781

20. Carraro, M, Giorgio, V., Šileikytė, J., Sartori, G., Forte, M., Lippe, G., Zoratti, M, Szabò, I. and Bernardi, P. (2014) Channel Formation by Yeast F-ATP Synthase and the Role of Dimerization in the Mitochondrial Permeability Transition, J. Biol. Chem. 289, 15980-15985 – paper of the year in Bioenergetics

21. Zulian, A., Rizzo, E., Schiavone, M., Palma, E., Tagliavini, F., Blaauw, B., Merlini, L., Maraldi, N.M., Sabatelli, P., Braghetta, P., Bonaldo, P., Argenton, F. and Bernardi, P. (2014) NIM811, a cyclophilin inhibitor without immunosuppressive activity, is beneficial in collagen VI congenital muscular dystrophy models, Hum. Mol. Genet. 23, 5353-5363

22. Antoniel,M., Giorgio, V., Fogolari, F., Glick, G.D., Bernardi, P. and Lippe, G. (2014) The oligomycinsensitivity conferring protein of mitochondrial ATP synthase: Emerging new roles in mitochondrial pathophysiology, Int. J. Mol. Sci. 15, 7513-7536

23. Fancelli, D., Abate, A., Amici, R., Bernardi, P., Ballarini, M., Cappa, A., Carenzi, G., Colombo, A., Contursi, C., Di Lisa, F., Dondio, G., Gagliardi, S., Milanesi, E., Minucci, S., Pain, G., Pelicci, P.G., Saccani, A., Storto, M., Thaler, F., Varasi, M., Villa, M., Plyte, S. (2014) Cinnamic Anilides as New Mitochondrial Permeability Transition Pore Inhibitors Endowed with Ischemia-Reperfusion Injury Protective Effect in Vivo, J. Med. Chem. 57, 5333-5347

24. Sorato, E., Menazza, S., Zulian, A., Sabatelli, P., Gualandi, F., Merlini, L., Bonaldo, P., Canton, M., Bernardi, P. and Di Lisa, F. (2014) Monoamine oxidase inhibition prevents mitochondrial dysfunction and apoptosis in myoblasts from patients with collagen VI myopathies, Free Radic. Biol. Med. 75C, 40-47

25. Gibellini, L., Pinti, M., Boraldi, F., Giorgio, V., Bernardi, P., Bartolomeo, R., Nasi, M., De Biasi, S., Missiroli, S., Carnevale, G., Losi, L., Tesei, A., Pinton, P., Quaglino, D., and Cossarizza, A. (2014) Silencing of mitochondrial Lon protease deeply impairs mitochondrial proteome and function in colon cancer cells, FASEB J. 28, 5122-5135

26. Da-Rè, C., von Stockum, S., Biscontin, A., Millino, C., Cisotto, P., Zordan, M.A., Zeviani, M., Bernardi, P., De Pittà, C., and Costa, R. (2014) Leigh Syndrome in Drosophila melanogaster: Morphological and Biochemical Characterization of Surf1 Post-transcriptional Silencing, J. Biol. Chem. 289, 29235-29246

27. Guzzo, G., Sciacovelli, M., Bernardi, P. and Rasola, A. (2014) Inhibition of succinate dehydrogenase by the mitochondrial chaperone TRAP1 has anti-oxidant and anti-apoptotic effects on tumor cells, Oncotarget 5, 11897-11908

28. Zulian, A., Tagliavini, F., Rizzo, E. Pellegrini, C., Sardone, F., Zini, N., Maraldi, N.M., Faldini, C., Merlini, L., Bernardi, P.* and Sabatelli, P.* (2014) Melanocytes from patients affected by Ullrich congenital muscular dystrophy and Bethlem myopathy have dysfunctional mitochondria that can be rescued with cyclophilin inhibitors, *Corresponding Authors, Front. Aging Neurosci. 6, 324

29. Rasola, A. and Bernardi, P. (2014) The mitochondrial permeability transition pore and its adaptive responses in tumor cells, Cell Calcium 56, 437-445

30. Bernardi, P. and Di Lisa, F. (2015) The mitochondrial permeability transition pore: Molecular nature and role as a target in cardioprotection, J. Mol. Cell. Cardiol. 78, 100-106

31. von Stockum, S., Giorgio, V., Trevisan, E., Lippe, G., Glick, G.D., Forte, M., Da-Rè, C., Checchetto, V., Mazzotta, G., Costa, R., Szabò, I., and Bernardi, P. (2015) F-ATPase of D. melanogaster Forms 53-Picosiemen (53-pS) Channels Responsible for Mitochondrial Ca2+-induced Ca2+ Release, J. Biol. Chem. 290, 4537-4544

32. Di Lisa, F. and Bernardi, P. (2015) Modulation of Mitochondrial Permeability Transition Prevents Energetic Failure in Ischemia-Reperfusion Injury of The Heart. Advantages and Limitations, Curr. Med. Chem. 22, 2480-2487

33. Bernardi, P., Di Lisa, F., Fogolari, F. and Lippe, G. (2015) From ATP to PTP and back. A dual function for the mitochondrial ATP synthase, Circ. Res. 116, 1850-1862

34. Bernardi, P., Rasola, A., Forte, M. and Lippe, G. (2015) The Mitochondrial Permeability Transition Pore: Channel Formation by F-ATP Synthase, Integration in Signal Transduction and Role in Pathophysiology, Physiol. Rev. 95, 1111-1155

35. Roy, S., Šileikytė, J., Schiavone, M., Neuenswander, B., Argenton, F., Aubé, J., Hedrick, M.P., Chung, T.D.Y., Forte, M.A.*, Bernardi, P.* and Schoenen F.J.* (2015) Discovery, Synthesis, and Optimization of Diarylisoxazole-3-carboxamides as Potent Inhibitors of the Mitochondrial Permeability Transition Pore, *corresponding Authors, ChemMedChem 10, 1655-1671

36. Šileikytė, J., Roy, S., Porubsky, P., Neuenswander, B., Wang, J., Hedrick, M., Pinkerton, A.B., Salaniwal, S., Kung, P., Mangravita-Novo, A., Smith, L.H., Bourdette, D.N., Jackson, M.R., Aubé, J., Chung, T.D.Y., Schoenen, F.J., Forte, M.A. and Bernardi, P. (2015) Small Molecules Targeting the Mitochondrial Permeability Transition. Updated 2015 Jan 16, In: Probe Reports from the NIH Molecular Libraries Program [Internet]. Bethesda (MD): National Center for Biotechnology Information (US); 2010-

37. Bernardi, P. and Forte, M. (2015) Commentary: SPG7 is an essential and conserved component of the mitochondrial permeability transition pore. Front. Physiol. 6, 320

38. Granatiero, V., Giorgio, V., Calì, T., Patron, M., Brini, M., Bernardi, P., Tiranti, V., Zeviani, M., Pallafacchina, G., De Stefani, D. and Rizzuto, R. (2016) Reduced mitochondrial Ca2+ transients stimulate autophagy in human fibroblasts carrying the 13514A>G mutation of the ND5 subunit of NADH dehydrogenase, Cell Death Diff. 23, 231-241

39. Roy, S., Šileikytė, J., Neuenswander, B., Hedrick, M.P., Chung, T.D.Y., Aubé, J., Schoenen, F.J., Forte, M.A. and Bernardi, P. (2016) N-Phenylbenzamides as Potent Inhibitors of the Mitochondrial Permeability Transition Pore, ChemMedChem 11, 283-288

40. Bernardi, P. and Di Lisa, F. (2016) Correspondence on article “Cyclosporine before PCI in Patients with Acute Myocardial Infarction” N. Engl. J. Med. 374, 88-90

41. Fontaine, E. and Bernardi, P. (2016) Lethal and non-lethal functions of the permeability transition pore. Mitochondria and Cell Death (D.M. Hockenbery ed.) Springer, New York 2016, pp. 1-15

42. Carraro M. and Bernardi, P. (2016) Calcium and reactive oxygen species in regulation of the mitochondrial permeability transition and of programmed cell death in yeast, Cell Calcium 60, 102-107

43. Scotton, C., Bovolenta, M., Schwartz, E., Falzarano, M.S., Martoni, E., Passarelli, C., Armaroli, A., Osman, H., Rodolico, C., Messina, S., Pegoraro, E., D’Amico, A., Bertini, E., Gualandi, F., Neri, M., Selvatici, R., Boffi, P., Maioli, M.A., Lochmuller, H., Straub, V., Bushby, K., Castrignanò, T., Pesole, G., Sabatelli, P., Merlini, L., Braghetta, P., Bonaldo, P., Bernardi, P., Reghan Foley, A., Cirak, S., Zaharieva, I., Muntoni, F., Capitanio, D., Gelfi, C., Kotelnikova, E., Yuryev, A., Lebowitz, M., Zhang, X., Hodge, B.A., Esser K. and Ferlini, A. (2016) Deep RNA profiling identified Clock and molecular clock genes as pathophysiological signatures in collagen VI myopathy J. Cell Sci. 129, 1671-1684

44. Bernardi, P. (2016) 19th European Bioenergetics ConferencePreface, Biochim. Biophys. Acta 1857, 1023-1026

45. Zulian, A., Schiavone, M., Giorgio, V. and Bernardi, P. (2016) Forty years later: mitochondria as therapeutic targets in muscle diseases, Pharmacol. Res. 113, 563-573

46. Bernardi, P. and Forte, M (2016) Commentary: The m-AAA protease associated with neurodegeneration limits MCU activity in mitochondria, Front. Physiol. 7, 583

47. Masgras, I., Ciscato, F., Brunati, A.M., Tibaldi, E., Indraccolo, S., Curtarello, M., Chiara, F., Cannino, G., Papaleo, E., Lambrughi, M., Guzzo, G., Gambalunga, A., Pizzi, M., Guzzardo, V., Rugge, M., Vuljan, S.E., Calabrese, F., Bernardi, P. and Rasola, A. (2017) Absence of neurofibromin induces an oncogenic metabolic switch via mitochondrial ERK-mediated phosphorylation of the chaperone TRAP1, Cell Rep. 18, 659-672

48. Giorgio, V., Guo, L., Bassot, C., Petronilli, V. and Bernardi, P. (2017) Calcium and regulation of the mitochondrial permeability transition, Cell Calcium, in press DOI: 10.1016/j.ceca.2017.05.004

49. Giorgio, V., Burchell, V., Schiavone, M., Bassot, C., Minervini, G., Petronilli, V., Argenton, F., Forte, M., Tosatto, S., Lippe, G. and Bernardi P. (2017) Ca2+ binding to F-ATP synthase  subunit triggers the mitochondrial permeability transition, EMBO Rep. 18, 1065-1076

50. Schiavone, M., Zulian, A., Menazza, S., Petronilli, V., Argenton, F., Merlini, L., Sabatelli, P. and Bernardi, P. (2017) Alisporivir rescues defective mitochondrial respiration in Duchenne muscular dystrophy, Pharmacol. Res. 125, 122-131

51. Connolly N.M.C., Theurey, P., Adam-Vizi, V., Bazan, N.G., Bernardi, P., Bolaños, J.P., Culmsee, C., Dawson, V.L., Deshmukh, M., Duchen, M.R., Düssmann, H., Fiskum, G.M., Galindo, M.F., Hardingham, G.E., Hardwick, J.M., Jekabsons, M.B., Jonas, E.A., Jordán, J., Lipton, S.A., Manfredi, G., Mattson, M.P., McLaughlin, B., Methner, A., Murphy, A.N., Murphy, M.P., Nicholls, D.G., Polster, B.M., Pozzan, T., Rizzuto, R., Slack, R.S., Satrústegui, J., Swanson, R.A., Swerdlow, R., Will, Y., Ying, Z., Joselin, A., Gioran, A., Moreira Pinho, C., Watters, O., Salvucci, M., Llorente-Folch, I., Park, D.S., Bano, D., Ankarcrona, M., Pizzo, P. and Prehn, J.H.M. (2017) Guidelines on experimental methods to assess mitochondrial dysfunction in cellular models of neurodegenerative diseases, Cell Death Differ. in press

52. Antoniel, M., Jones, K., Antonucci, S., Spolaore, B., Fogolari, F., Petronilli, V., Giorgio, V., Carraro, M., Di Lisa, F., Forte, M., Szabó, I., Lippe, G and Bernardi, P. (2017) The unique histidine in OSCP subunit of FATP synthase mediates inhibition of the permeability transition pore at acidic pH, EMBO Rep. in press DOI: 10.15252/embr.201744705 53. Bernardi, P. and Lippe, G. (2017) Channel Formation by F-ATP Synthase and the Permeability Transition Pore: An Update, Curr. Opin. Physiol. in press