HEK293 is a cell line derived from human embryonic kidney cells grown in tissue culture. The cell line was generated in 1973 by transfection of normal healthy human embryonic kidney (HEK) cells with adenovirus 5 DNA by Alex Van der Eb in Leiden, Netherlands. The use of 293 refers to the experiment number referenced in his laboratory notebook. The original kidney cell culture that served as the source of the cell line was heterogeneous, and so evidence suggests fibroblast, endothelial or epithelial cells as possible lineages. HEK293 cells are hypotriploid human cell lines, meaning they possess an abnormal chromosome number. They carry 64 chromosomes, three X chromosome copies, and an incorporation of 4 kbp Ad5 to chromosome 19 of the HEK293 genome indicating cytogenetic instability. HEK293 cells are commonly used in cancer research, biotechnology, and basic medical research. They are considered a low-maintenance and robust cell line, and their culturing requirement is generally uncomplicated. HEK293 cells are exceptionally efficient at yielding recombinant proteins in large amounts, and several vaccines and biotherapeutic proteins have been already developed using HEK293 cells.