To identify cells and extract their properties, the CellProfiler program (Carpenter et al., 2006) was used with custom modifications. First, image intensities were rescaled. Then, cell nuclei were identified based on the Hoechst signal using Otsu thresholding and the Watershed algorithm for closely spaced nuclei. The cytoplasm was approximated as a ring around the cell nucleus (width of ~ 6.5 µm). Using the detected objects (nucleus and cytoplasm) as masks, cell features such as intensity values, textural properties, and morphological descriptors were extracted (Wild et al., 2010).