The Problem of Apoptotic Processes Reversibility
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REVIEW
The Problem of Apoptotic Processes Reversibility I. I. Zakharov1, M. A. Savitskaya1, and G. E. Onishchenko1,a* 1
Faculty of Biology, Lomonosov Moscow State University, 119234 Moscow, Russia a email: [email protected] Received July 10, 2020 Revised July 23, 2020 Accepted July 23, 2020
Abstract—Apoptosis is the best understood variant of regulated cell death, which has been considered irreversible for a long time. To date, an increasing amount of data has been accumulating indicating that key events of apoptosis, such as the exter nalization of phosphatidylserine, mitochondrial outer membrane permeabilization, caspase activation, DNA damage, and cytoplasmic blebbing are not irreversible and can be involved in the normal cell functioning not associated with the induc tion of apoptosis. Anastasis – cell recovery after induction of apoptosis – can occur following elimination of proapoptotic stimuli. This can facilitate survival of damaged or tumor cells. This review describes key processes of apoptosis, which do not necessarily lead to cell death during normal cell activity as well as anastasis. Understanding mechanisms and conse quences of apoptotic processes reversibility, on the one hand, could contribute to the improvement of existing therapeutic approaches for various diseases, including malignant neoplasms, and, on the other hand, could open up new possibilities for protecting cellular elements of tissues and organs from death during treatment of degenerative pathologies. DOI: 10.1134/S000629792010003X Keywords: apoptosis, apoptosis reversibility, anastasis, caspase activation, MOMP, PS externalization, blebbing
INTRODUCTION Apoptosis is a type of regulated cell death [1] wide spread in the course of ontogenesis of multicellular organisms. Apoptosis comprises a sequence of molecular, structural, and physiological changes of a cell, resulting in its fragmentation and loss of viability, i.e., death. This type of cell death plays an important role in many physi ological processes, including embryogenesis, mainte nance of cell homeostasis, differentiation and tissue mor phogenesis, while its disruptions could be a sign of many pathologies, from degenerative syndromes, to tumors, and to autoimmune diseases. The sequence of apoptotic processes, the mecha nisms of its induction and regulation have been studied for fifty years. There are several apoptosis initiation path Abbreviations: AIF, apoptosisinducing factor; BCLXL, Bcell lymphoma extralarge; BCL2, Bcell lymphoma 2; BID, BH3 interactingdomain death agonist; CAD, caspaseactivated DNase; ENDOG, endonuclease G; ER, endoplasmic reticu lum; FASLG, FASligand; MOMP, mitochondrial outer mem brane permeabilization; PARP, poly(ADPribose) polymerase; PS, phosphatidylserine; TNFα, tumor necrosis factor α; TRAIL, TNFrelated apoptosisinducing ligand; XIAP, Xchromosomelinked inhibitor of apoptosis. * To whom correspondence should be addressed.
ways including the receptordependent pathways [TNF related apoptosisinducing ligand (TRAIL), tumor n
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