Tiago T, Hummel B, Morelli FF, Basile V, Vinet J, Galli V, Mediani L, Antoniani F, Pomella S, Cassandri M, Garone MG, Silvestri B, Cimino M, Cenacchi G, Costa R, Mouly V, Poser I, Yeger-Lotem E, Rosa A, Alberti S, Rota R, Ben-Zvi A, Sawarkar R, Carra S. Small heat-shock protein HSPB3 promotes myogenesis by regulating the lamin B receptor. Cell Death Dis. 2021 May 6;12(5):452. doi: 10.1038/s41419-021-03737-1.

Shemesh N, Jubran J, Dror S, Simonovsky E, Basha O, Argov C, Hekselman I, Abu-Qarn M, Vinogradov E, Mauer O, Tiago T, Carra S, Ben-Zvi A, Yeger-Lotem E. The landscape of molecular chaperones across human tissues reveals a layered architecture of core and variable chaperones. Nat Commun. 2021 Apr 12;12(1):2180. doi: 10.1038/s41467-021-22369-9.

Mediani L, Antoniani F, Galli V, Vinet J, Carrà AD, Bigi I, Tripathy V, Tiago T, Cimino M, Leo G, Amen T, Kaganovich D, Cereda C, Pansarasa O, Mandrioli J, Tripathi P, Troost D, Aronica E, Buchner J, Goswami A, Sterneckert J, Alberti S, Carra S. Hsp90-mediated regulation of DYRK3 couples stress granule disassembly and growth via mTORC1 signaling. EMBO Rep. 2021 Mar 19:e51740. doi: 10.15252/embr.202051740.


Mediani L, Galli V, Carrà AD, Bigi I, Vinet J, Ganassi M, Antoniani F, Tiago T, Cimino M, Mateju D, Cereda C, Pansarasa O, Alberti S, Mandrioli J, Carra S. BAG3 and BAG6 differentially affect the dynamics of stress granules by targeting distinct subsets of defective polypeptides released from ribosomes. Cell Stress Chaperones. 2020 Jul 21. doi: 10.1007/s12192-020-01141-w.

Adriaenssens E, Tedesco B, Mediani L, Asselbergh B, Crippa V, Antoniani F, Carra S*, Poletti A*, Timmerman V*. BAG3 Pro209 mutants associated with myopathy and neuropathy relocate chaperones of the CASA-complex to aggresomes. Sci Rep. 2020 May 29;10(1):8755. doi: 10.1038/s41598-020-65664-z.

Choudhary D, Mediani L, Carra S, Cecconi C. Studying heat shock proteins through single-molecule mechanical manipulation.  Cell Stress Chaperones. 2020 Apr 6. doi: 10.1007/s12192-020-01096-y.


Alberti S, Carra S. Nucleolus: A Liquid Droplet Compartment for Misbehaving Proteins. Curr. Biol., 2019. 29(19):R930-R932.

Mediani L, Guillén-Boixet J, Alberti S, Carra S. Nucleoli and Promyelocytic Leukemia Protein (PML) bodies are phase separated nuclear protein quality control compartments for misfolded proteins. Molecular & Cellular Oncology, https://doi.org/10.1080/23723556.2019.1652519

Cicardi ME, Cristofani R, Crippa V, Ferrari V, Tedesco B, Casarotto E, Chierichetti M, Galbiati M, Piccolella M, Messi E, Carra S, Pennuto M, Rusmini P, Poletti A. Autophagic and Proteasomal Mediated Removal of Mutant Androgen Receptor in Muscle Models of Spinal and Bulbar Muscular Atrophy. Front. Endocrinol. (Lausanne), 2019. 10:569.

Cristofani R, Rusmini P, Galbiati M, Cicardi ME, Ferrari V, Tedesco B, Casarotto E, Chierichetti M, Messi E, Piccolella M, Carra S, Crippa V, Poletti A. The Regulation of the Small Heat Shock Protein B8 in Misfolding Protein Diseases Causing Motoneuronal and Muscle Cell Death. Front. Neurosci., 2019. 13:796.

Mediani L, Guillén-Boixet J, Vinet J, Franzmann TM, Bigi I, Mateju D, Carrà AD, Morelli FF, Tiago T, Poser I, Alberti S, Carra S. Defective ribosomal products challenge nuclear function by impairing nuclear condensate dynamics and immobilizing ubiquitin. EMBO J., 2019. 38(15):e101341.

Mandrioli J, Mediani L, Alberti S, Carra S. ALS and FTD: where RNA metabolism meets protein quality control. Semin. Cell Dev. Biol., 2019. Jun 26. pii: S1084-9521(18)30200-3. doi: 10.1016/j.semcdb.2019.06.003. Review.

Mandrioli J, Crippa V, Cereda C, Bonetto V, Zucchi E, Gessani A, Ceroni M, Chio A, D’Amico R, Monsurrò MR, Riva N, Sabatelli M, Silani V, Simone IL, Sorarù G, Provenzani A, D’Agostino VG, Carra S*, Poletti A*. Proteostasis and ALS: protocol for a phase II, randomised, double-blind, placebo-controlled, multicentre clinical trial for colchicine in ALS (Co-ALS). BMJ Open, 2019. 9(5):e028486.*: co-last authors

Carra S, Alberti S, Benesch JLP, Boelens W, Buchner J, Carver JA, Cecconi C, Ecroyd H, Gusev N, Hightower LE, Klevit RE, Lee HO, Liberek K, Lockwood B, Poletti A, Timmerman V, Toth ME, Vierling E, Wu T, Tanguay RM. Small heat shock proteins: multifaceted proteins with important implications for life. Cell Stress Chaperones, 2019. 24(2):295-308.



Meister-Broekema M, Freilich R, Jagadeesan C, Rauch JN, Bengoechea R, Motley WW, Kuiper EFE, Minoia M, Furtado GV, van Waarde MAWH, Bird SJ, Rebelo A, Zuchner S, Pytel P, Scherer SS, Morelli FF, Carra S, Weihl CC, Bergink S, Gestwicki JE, Kampinga HH. Myopathy associated BAG3 mutations lead to protein aggregation by stalling Hsp70 networks.  Nat. Commun., 2018. 9(1):5342.

Weihl CC, Udd B, Hanna M; ENMC workshop study group. Collaborators: Ben-Zvi A, Blaettler T, Bryson-Richardson R, Carra S, Dimachkie M, Findlay A, Greensmith L, Greenspan S, Hanna M, Höhfled J, Jonson PH, Kampinga HH, Larsson L, Linke W, Lynch G, Machado P, Orlando L, Richard I, Roos A, Sarparanta J, Timmerman V, Udd B, Weihl C, Zah L. 234th ENMC International Workshop: Chaperone dysfunction in muscle disease Naarden, The Netherlands, 8-10 December 2017. Neuromuscul. Disord., 2018. 28(12): 1022-30.

Alberti S, Carra S. Quality Control of Membraneless Organelles.  J. Mol. Biol., 2018. 430(23):4711-4729.

Cicardi ME, Cristofani R, Rusmini P, Meroni M, Ferrari V, Vezzoli G, Tedesco B, Piccolella M, Messi E, Galbiati M, Boncoraglio A, Carra S, Crippa V, Poletti A. Tdp-25 Routing to Autophagy and Proteasome Ameliorates its Aggregation in Amyotrophic Lateral Sclerosis Target Cells. Sci. Rep., 2018. 8(1):12390.



Morelli FF, Verbeek S, Bertacchini J, Vinet J, Mediani L, Marmiroli S, Cenacchi G, Nasi M, De Biasi S, Brunsting JF, Lammerding J, Pegoraro E, Angelini C, Tupler R, Alberti S, Carra S. Aberrant compartment formation by HSPB2 mislocalizes lamin A and compromises nuclear integrity and function. Cell Rep., 2017. 20(9):2100-2115.

Rusmini P, Cristofani R, Galbiati M, Cicardi ME, Meroni M, Ferrari V, Vezzoli G, Tedesco B, Messi E, Piccolella M, Carra S, Crippa V, Poletti A. The Role of the Heat Shock Protein B8 (HSPB8) in Motoneuron Diseases. Front. Mol. Neurosci., 2017. 10:176.

Cristofani R, Crippa V, Vezzoli G, Rusmini P, Galbiati M, Cicardi ME, Meroni M, Ferrari V, Tedesco B, Piccolella M, Messi E, Carra S, Poletti A. The small heat shock protein B8 (HSPB8) efficiently removes aggregating species of dipeptides produced in C9ORF72-related neurodegenerative diseases. Cell Stress Chaperones, 2017. 23(1):1-12.

Alberti S, Mateju D, Mediani L and Carra S. Granulostasis: protein quality control of RNP granules. Front. Mol. Neurosci., 2017. 10:84.

Mateju D, Franzmann TM, Patel A, Kopach A, Boczek EE, Maharana S, Lee HO, Carra S, Hyman AA, Alberti S. An aberrant phase transition of stress granules triggered by misfolded protein and prevented by chaperone function. EMBO J., 2017. 36(12):1669-87.

Carra S, Alberti S, Arrigo PA, Benesch JL, Benjamin IJ, Boelens W, Bartelt-Kirbach B, Brundel BJ, Buchner J, Bukau B, Carver JA, Ecroyd H, Emanuelsson C, Finet S, Golenhofen N, Goloubinoff P, Gusev N, Haslbeck M, Hightower LE, Kampinga HH, Klevit RE, Liberek K, Mchaourab HS, McMenimen KA, Poletti A, Quinlan R, Strelkov SV, Toth ME, Vierling E, Tanguay RM. The growing world of small heat shock proteins: from structure to functions. Cell Stress Chaperones, 2017. 22(4):601-11.

Morelli FF, Mediani L, Heldens L, Bertacchini J, Bigi I, Carrà AD, Vinet J, Carra S. An interaction study in mammalian cells demonstrates weak binding of HSPB2 to BAG3, which is regulated by HSPB3 and abrogated by HSPB8. Cell Stress Chaperones, 2017. 22(4): 531-40.

Cristofani R, Crippa V, Rusmini P, Cicardi ME, Meroni M, Giorgetti E, Sala G, Galbiati M, Piccolella M, Messi E, Ferrarese C, Carra S, Poletti A. Inhibition of retrograde transport modulates misfolded protein accumulation and clearance in motoneuron diseases. Autophagy, 2017. 13(8): 1280-1303.

Piccolella M, Crippa V, Cristofani R, Rusmini P, Galbiati M, Elena Cicardi M, Meroni M, Ferri N, Morelli FF, Carra S, Messi E, Poletti A. The small heat shock protein B8 (HSPB8) modulates proliferation and migration of breast cancer cells. Oncotarget, 2017. 8(6):10400-10415.



Ganassi M, Mateju D, Bigi I, Mediani L, Poser I, Lee HO, Seguin SJ, Morelli FF, Vinet J, Leo G, Pansarasa O, Cereda C, Poletti A, Alberti S, Carra S. A Surveillance Function of the HSPB8-BAG3-HSP70 Chaperone Complex Ensures Stress Granule Integrity and Dynamism. Mol. Cell, 2016. 63(5):796-810. 

Crippa V, Cicardi ME, Ramesh N, Seguin SJ, Ganassi M, Bigi I, Diacci C, Zelotti E, Baratashvili M, Gregory JM, Dobson CM, Cereda C, Pandey UB, Poletti A, Carra S. The chaperone HSPB8 reduces the accumulation of truncated TDP-43 species in cells and protects against TDP-43-mediated toxicity.  Hum. Mol. Genet., 2016. 25(18):3908-24.

Crippa V, D’Agostino VG, Cristofani R, Rusmini P, Cicardi ME, Messi E, Loffredo R, Pancher M, Piccolella M, Galbiati M, Meroni M, Cereda C, Carra S, Provenzani A, Poletti A. Transcriptional induction of the heat shock protein B8 mediates the clearance of misfolded proteins responsible for motor neuron diseases.  Sci. Rep., 2016. 6:22827.

Rusmini P, Crippa V, Cristofani R, Rinaldi C., Cicardi ME, Galbiati M, Carra S, Bilal M, Greensmith L, Poletti A. The Role of the Protein Quality Control System in SBMA. J. Mol. Neurosci., 2016. 58(3):348-64.

Klionsky DJ, Abdelmohsen K, Abe A, Abedin MJ, Abeliovich H, Acevedo Arozena A, Adachi H, et al., Guidelines for the use and interpretation of assays for monitoring autophagy (3rd edition). Autophagy, 12(1):1-222.

Vos MJ, Carra S., Kanon B., Bosveld F., Klauke K., Sibon O.C.M., Kampinga H.H. Specific protein homeostatic functions of small heat shock proteins increase lifespan. Aging Cell 2016. 15(2):217-26.



Poletti A and Carra S. Role of HSPB8 in the proteostasis network: from protein synthesis to protein degradation and beyond. Series Title: Heat Shock Proteins; book title: “The Big Book of Small Heat Shock Proteins”, Eds RM Tanguay and LE Hightower, Springer International Publishing AG, ISBN:  978-3-319-16076-4.



Seguin SJ, Morelli FF, Vinet J, Amore D, De Biasi S, Poletti A, Rubinsztein DC, Carra S. Inhibition of autophagy, lysosome and VCP function impairs stress granule assembly. Cell Death Differ., 2014. 21(12):1838-51.

Minoia M, Boncoraglio A, Vinet J, Morelli FF, Brunsting JF, Poletti A, Krom S, Reits E, Kampinga HH, Carra S. BAG3 induces the sequestration of proteasomal clients into cytoplasmic puncta: implication for a proteasome-to-autophagy switch. Autophagy, 2014. 10(9):163-21.

Kakkar V, Meister-Broekema M, Minoia M, Carra S, Kampinga HH. Barcoding Heat Shock Proteins to Human Diseases: Looking Beyond The Heat Shock Response. Dis. Model Mech., 2014. 7(4):421-34.



Yang J, Carra S, Zhu WG, Kampinga HH. The regulation of the autophagic network and its implications for human disease. Int. J. Biol. Sci., 2013. 9(10):1121-33.

Crippa V, Boncoraglio A, Galbiati M, Aggarwal T, Rusmini P, Giorgetti E, Cristofani R, Carra S, Pennuto M, Poletti A. Differential autophagy power in the spinal cord and muscle of transgenic ALS mice. Front. Cell. Neurosci., 2013. 7:234.

Rusmini P, Crippa V, Giorgetti E, Boncoraglio A, Cristofani R, Carra S, Poletti A. Clearance of the mutant androgen receptor in motoneuronal models of spinal and bulbar muscular atrophy. Neurobiol. Aging, 2013. 34(11):2585-603.

Carra S, Rusmini P, Crippa V, Giorgetti E, Boncoraglio A, Cristofani R, Naujock M, Meister M, Minoia M, Kampinga HH and Poletti A. Different anti-aggregation and pro-degradative functions of the members of the mammalian sHSP family in neurological disorders. Phil. Trans. R. Soc. B, 2013. 368(1617):20110409.



Klionsky DJ, Abdalla FC, Abeliovich H, Abraham RT, Acevedo-Arozena A, Adeli K, Agholme L, Agnello M, Agostinis P, Aguirre-Ghiso JA,  et al., Guidelines for the use and interpretation of assays for monitoring autophagy. Autophagy, 2012. 8(4):445-544.

Boncoraglio A, Minoia M, Carra S. The family of mammalian small heat shock proteins (HSPBs): Implications in protein deposit diseases and motor neuropathies. Int. J. Biochem. Cell Biol., 2012. 44(10):1657-69.

Carra S, Crippa V, Rusmini P, Boncoraglio A, Minoia M, Giorgetti E, Kampinga HH, Poletti A. Alteration of protein folding and degradation in motor neuron diseases: Implications and protective functions of small heat shock proteins. Prog. Neurobiol., 2012. 97(2):83-100.

Seidel K, Vinet J, den Dunnen WFA, Brunt ER, Meister M, Boncoraglio A, Zijlstra MP, Boddeke HWGM, Rüb U, Kampinga HH and Carra S. The HSPB8-BAG3 chaperone complex is upregulated in astrocytes in the human brain affected by protein aggregation diseases. Neuropathol. Appl. Neurobiol., 2012. (1):39-53.



Gamerdinger M, Carra S, Behl C. Emerging roles of molecular chaperones and co-chaperones in selective autophagy: focus on BAG proteins. J. Mol. Med. (Berl)., 2011. (12):1175-82.

Hishiya A, Salman MN, Carra S, Kampinga HH and Takayama S. BAG3 directly interacts with mutated alphaB-crystallin to suppress its aggregation and toxicity. PLOS ONE, 2011. 6(3):e16828.

Vos MJ, Zijlstra MP, Carra S, Sibon OC, Kampinga HH. Small heat shock proteins, protein degradation and protein aggregation diseases. Autophagy, 2011. 7(1):101-103.



Carra S, Boncoraglio A, Kanon B, Brunsting JF, Minoia M, Rana A, Vos MJ, Seidel K, Sibon OC, Kampinga HH. Identification of the Drosophila ortholog of HSPB8: implication of HSPB8 loss of function in protein folding diseases. J. Biol. Chem., 2010. 285(48):37811-22.

Vos MJ, Zijlstra MP, Kanon B, van Waarde-Verhagen MA, Brunt ER, Oosterveld-Hut HM, Carra S, Sibon OC, Kampinga HH. HSPB7 is the most potent polyQ aggregation suppressor within the HSPB family of molecular chaperones. Hum. Mol. Genet., 2010. 19(23):4677-93.

Crippa V, Carra S, Rusmini P, Sau D, Bolzoni E, Bendotti C, De Biasi S, Poletti A. A role of small heat shock protein B8 (HSPB8) in the autophagic removal of misfolded proteins responsible for neurodegenerative diseases. Autophagy, 2010. 6(7):958-60.

Crippa V, Sau D, Rusmini P, Boncoraglio A, Onesto E, Bolzoni E, Galbiati M, Fontana E, Marino M, Carra S, Bendotti C, De Biasi S, Poletti A. The small heat shock protein B8 (HSPB8) promotes autophagic removal of misfolded proteins involved in amyotrophic lateral sclerosis (ALS). Hum. Mol. Genet., 2010. 19(17):3440-56.

Sun X, Fontaine JM, Hoppe AD, Carra S, DeGuzman C, Martin JL, Simon S, Vicart P, Welsh MJ, Landry J, Benndorf R. Abnormal interaction of motor neuropathy-associated mutant HspB8 (Hsp22) forms with the RNA helicase Ddx20 (gemin3). Cell Stress & Chaperones, 2010. 15(5):567-582.

Carra S and Kampinga HH. Cytoprotective function of small stress proteins in conformational disorders. In Review Book: Small Stress Proteins and Human Diseases. Chapter 1.1, 2010, Nova Publishers.



Fuchs M, Poirier DJ, Seguin S, Lambert H, Carra S, Charrette SJ, Landry J. Identification of the key structural motifs involved in HspB8/HspB6-Bag3 interaction. Biochem. J., 2009. 425(1):245-55.

Carra S. The stress-inducible HspB8-Bag3 complex induces the eIF2alpha kinase pathway: Implications for protein quality control and viral factory degradation? Autophagy, 2009. 5(3):428-9.



Carra S, Brunsting JF, Lambert H, Landry J, Kampinga HH. HspB8 participates in protein quality control by a non chaperone-like mechanism that requires eIF2alpha phosphorylation. J. Biol. Chem., 2008. 284(9): 5523-32.

Hageman J, Vos MJ,Carra S, Kampinga HH. Structural and functional homologies and diversities between members of the human HspH (Hsp110), HspA (Hsp70), DnaJ (Hsp40) and HspB (Hsp27) families. Biochemistry, 2008. 47(27):7001-11.

Carra S, Seguin SJ and Landry J. HspB8 and Bag3: a new chaperone complex targeting misfolded proteins to macroautophagy. Autophagy. 2008. 4(2):237-9.

Carra S, Seguin SJ, Lambert H and Landry J. HspB8 chaperone activity toward poly(Q)-containing proteins depends on its association with Bag3, a stimulator of macroautophagy. J. Biol. Chem., 2008. 283(3):1437-44.

Carra S & Landry J. Role of HspB1 and HspB8 in hereditary peripheral neuropathies: beyond the chaperone function. In Review Book: Heat shock proteins and the brain: implications for neurodegenerative diseases and neuroprotection; Chapter 7, 139-155, 2008, Editors: Asea AAA, Brown IR.



Carra S & Landry J. Small Heat Shock Proteins in Neurodegenerative Diseases. In Review Book: Heat Shock Proteins in Biology and Medicine; Section VI, Chapter 18, 331-352, 2006, Editors: Jurgen Radons and Gabriele Multhoff.



Carra S, Sivilotti M, Chavez-Zobel AT, Lambert H, Landry J. HspB8, a small heat shock protein mutated in human neuromuscular disorders, has in vivo chaperone activity in cultured cells. Hum. Mol. Genetics, 2005. 14(12):1659-69.



Carra S,Vinet J, Blom JM, Brunello N, Barden N, Tascedda F. Chronic treatment with desipramine and fluoxetine modulate BDNF, CaMKKalpha and CaMKKbeta mRNA levels in the hippocampus of transgenic mice expressing antisense RNA against the glucocorticoid receptor. Neuropharmacology. 2004. 47(7):1062-9.



Vinet J, Carra S, Blom JM, Harvey M, Brunello N, Barden N, Tascedda F. Cloning of mouse Ca2+/calmodulin-dependent protein kinase kinase beta (CaMKKβ) and characterization of CaMKKβ and CaMKKα distribution in the adult mouse brain. Mol. Brain Research, 2003. 111(1-2):216-21.



Blom JM, Tascedda F, Carra S, Ferraguti C, Barden N, Brunello N. Altered regulation expression of CREB by chronic antidepressant administration in the brain of transgenic mice with impaired glucocorticoid receptor function. Neuropsychopharmacology, 2002. 26(5):605-14.



Tascedda F, Blom JM, Carra S, Brunello N, Racagni G, Riva MA. Modulation of glutamate receptors in response to the novel antipsychotic olanzapine in rats. Biol. Psychiatry, 2001. 50(2):117-22.