EV-71 was inoculated in human intestinal organoids as a positive control and displayed a robust viral propagation (Fig. 4c, d), congruent with previous reports.21,26 Notably, CoV-HKU4 also replicated in human intestinal organoids (Fig. 4e), echoing the previous finding of its productive infection in human DPP4 transgenic mice.35 CoV-HKU4 infection in bat and human intestinal organoids was verified by immunofluorescence staining (Fig. 4f) using an antiserum against CoV-HKU4 nucleocapsid protein (NP). Here, DPP4 is linked to infection.