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Significance The plague bacillusYersinia pestis is transmitted by fleas but evolved very recently fromYersinia pseudotuberculosis , a closely related food- and water-borne enteric pathogen. Identifying the specific genetic changes underlying the switch to arthropod-borne transmission is essential to understanding the evolution ofY. pestis and the emergence of plague. We have identified an important early step in this process. The urease enzyme, beneficial for an enteric bacterium, causes acute oral toxicity and significant mortality to fleas.Y. pestis is not toxic to fleas because of a loss-of-function mutation in one of the genes essential for urease activity. Epidemiologic modeling suggests that this mutation was positively selected during evolution because it resulted in a significantly higher probability ofY. pestis transmission.

Silencing urease: A key evolutionary step that facilitated the adaptation of Yersinia pestis to the flea-borne transmission route