In his seminal paper, Damerau stated that in an investigation of spelling errors for an information-retrieval system, more than 80% were a result of a single error of one of the four types. Damerau's paper considered only misspellings that could be corrected with at most one edit operation. While the original motivation was to measure distance between human misspellings to improve applications such as spell checkers, Damerau–Levenshtein distance has also seen uses in biology to measure the variation between protein sequences.