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Many studies have demonstrated changes in the levels of several ions during apoptosis, but a few recent studies have reported conflicting results concerning the changes in water content in apoptotic cells. We used a correlative light and cryo-scanning transmission electron microscopy method to quantify water and ion/element contents simultaneously at a nanoscale resolution in the various compartments of cells, from the onset to the end of apoptosis. We used stably transfected HeLa cells producing H2B-GFP to identify the stages of apoptosis in cells and for a targeted elemental analysis within condensed chromatin, nucleoplasm, mitochondria and the cytosol. We found that the compartments of apoptotic cells contained, on average, 10% more water than control cells. During mitochondrial outer membrane permeabilization, we observed a strong increase in the Na +  and Cl -  contents of the mitochondria and a strong decrease in mitochondrial K +  content. During the first step in apoptotic volume decrease (AVD), Na +  and Cl -  levels decreased in all cell compartments, but remained higher than those in control cells. Conversely, during the second step of AVD, Na +  and Cl -  levels increased considerably in the nucleus and mitochondria. During these two steps of AVD, K +  content decreased steadily in all cell compartments. We also determined  in vivo  ion status during caspase-3 activity and chromatin condensation. Finally, we found that actinomycin D-tolerant cells had water and K +  contents similar to those of cells entering apoptosis but lower Na +  and Cl -  contents than both cells entering apoptosis and control cells.

plos one

Stage-Specific Changes in the Water, Na+, Cl- and K+ Contents of Organelles during Apoptosis, Demonstrated by a Targeted Cryo Correlative Analytical Approach