Last changed: 04 January 2020

Proteases cut polypeptide chains and in this way modulate activity of target proteins and downstream pathways. We are curious about how caspase-fold proteases are able to perform multiple and often antagonistic roles, such as in the regulation of cell death and cell division and establishment of cell polarity.

Tudor staphylococcal nuclease (TSN) is an evolutionarily conserved substrate of caspase-fold proteases.

Selected publications

Minina EA, Moschou PN, and Bozhkov PV (2017) Limited and digestive proteolysis: crosstalk between evolutionary conserved pathways​. New Phytologist 215, 958–964.

Minina EA, Coll NS, Tuominen H and Bozhkov PV (2017) Metacaspases versus caspases in development and cell fate regulation. Cell Death and Differentiation 24, 1314–1325.

Chen L, Stael S, Gevaert K, Van Breusegem F, Bozhkov P, Moschou P (2017) The Proteolytic Landscape Of An Arabidopsis Separase-Deficient Mutant Reveals Novel Substrates Associated With Plant Development. bioRxiv doi:

Moschou PN, Gutierrez-Beltran E, Bozhkov PV (co-senior author),Smertenko AP (2016) Separase Promotes Microtubule Polymerization by Activating CENP-E-Related Kinesin Kin7. Developmental Cell 37: 350-361.

Moschou PN, Savenkov EI, Minina EA, Fukada K, Reza SH, Gutierrez-Beltran E, Sanchez-Vera V, Suarez MF, Hussey P, Smertenko AP and Bozhkov PV (2016) EXTRA SPINDLE POLES (Separase) controls anisotropic cell expansion in Norway spruce (Picea abies) embryos independently from its role in anaphase progression. New Phytologist 212: 232-243.

Gutierrez-BeltranE, Denisenko TV, Zhivotovsky B and Bozhkov PV (2016) Tudor Staphylococcal Nuclease: biochemistry and functions. Cell Death and Differentiation 23, 1739-1748.

P. Bozhkov and G. Salvesen (Editors). Caspases, Paracaspases and Metacaspases. Series: Methods in Molecular Biology, Vol. 1133 Humana Press. 2014. ISBN 978-1-4939-0356-6

Minina EA, Smertenko AP and Bozhkov PV (2014) Vacuolar cell death in plants: metacaspase releases the brakes on autophagy. Autophagy 10: 928-929.

Minina E, Filonova L, Fukada K, Savenkov EI, Sanchez-Vera V, Suarez MF, Daniel G, Clapham D, Gogvadze V, Zhivotovsky B, Smertenko AP and Bozhkov PV (2013) Metacaspase and autophagy determine the mode of plant cell death. J Cell Biol. 203: 917-927.

Peng Zhao, Xue-mei Zhou, Li-yao Zhang, Wei Wang, Li-gang Ma, Li-bo Yang, Xiong-bo Peng, Bozhkov PV, Meng-xiang Sun. (2013) A bipartite molecular module controls cell death activation in the basal cell lineage of plant embryos. PLOS Biol 11(9): e1001655. doi:10.1371/journal.pbio.1001655.

Moschou PN, Smertenko AP, Fukada K, Savenkov EI, Minina EA, Robert S, Hussey PJ, Bozhkov PV (2013) The Caspase-Related Protease Separase (EXTRA SPINDLE POLES) Regulates Cell Polarity and Cytokinesis in Arabidopsis. Plant Cell 25: 2171-2186.

Moschou PN and Bozhkov PV (2012) Separases: biochemistry and function. Physiologia Plantarum 145: 67-76.

Tsiatsiani L, Van Breusegem F, Gallois P, Zavialov A, Lam E and Bozhkov PV (2011) Metacaspases. Cell Death and Differentiation 18: 1279-1288.

Bozhkov PV, Smertenko AP and Zhivotovsky B (2010) Aspasing out metacaspases and caspases: proteases of many trades. Science Signaling, 3: pe48.

Sundström J, Vaculova A, Smertenko AP, Savenkov EI, Golovko A, Minina E, Tiwari BS, Rodriguez-Nieto S, Zamyatnin AAJr, Välineva T, Saarikettu J, Frilander MJ, Suarez MF, Zavialov A, Ståhl U, Hussey PJ, Silvennoinen O, Sundberg E, Zhivotovsky B, and Bozhkov PV (2009) Tudor staphylococcal nuclease is an evolutionarily conserved component of the programmed cell death degradome. Nature Cell Biology 11: 1347-1354.

Helmersson A, von Arnold S and Bozhkov PV (2008) The level of free intracellular zinc mediates programmed cell death/cell survival decisions in plant embryos. Plant Physiology 147: 1158-1167.

Bozhkov PV and Jansson C. (2007) Autophagy and cell-death proteases in plants: Two wheels of a funeral cart. Autophagy 3: 136-138.

Boren M, Höglund A-S, Bozhkov P and Jansson C (2006) Developmental regulation of a VEIDase caspase-like proteolytic activity in barley caryopsis. J. Exp. Bot 57: 3747-3753.

Bozhkov PV, Suarez MF, Filonova LH, Daniel G, Zamyatnin AA Jr, Rodriguez-Nieto S, Zhivotovsky B and Smertenko A. (2005) Cysteine protease mcII-Pa executes programmed cell death during plant embryogenesis. Proc Natl Acad Sci USA 102: 14463-14468.

Suarez MF, Filonova LH, Smertenko A, Savenkov EI, Clapham DH, von Arnold S, Zhivotovsky B and Bozhkov PV. (2004) Metacaspase-dependent programmed cell death is essential for plant embryogenesis. Current Biology 14: R339-R340.

Bozhkov PV, Filonova LH, Suarez MF, Helmersson A, Smertenko AP, Zhivotovsky B and von Arnold S. (2004) VEIDase is a principal caspase-like activity involved in plant programmed cell death and essential for embryonic pattern formation. Cell Death and Differentiation. 11: 175-182.


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