A final subset was characterized as ‘hybrid’ neutrophils due to their macrophage-like characteristics, i.e., expression of MHC class II and complement activation genes (C1QB, C1QC, CD74), cathepsins (CTSB, CTSL) and APOE. All neutrophil subclusters were more frequent in COVID-19 than non-COVID-19, but most significant changes were noticed for the ‘progenitor’ and ‘inflammatory mature’ neutrophils (Fig. 6d), both for the mild and critical COVID-19 vs non-COVID-19 comparison, albeit not always significantly (Fig. 6e; Supplementary information, Fig. S6g). Here, CTSL is linked to COVID-19.