Negri was convinced the inclusions were a parasitic protozoon and the etiologic agent of rabies. Later that same year, however, Paul Remlinger and Rifat-Bey Frasheri in Constantinople and, separately, Alfonso di Vestea in Naples showed that the etiologic agent of rabies is a filterable virus. Negri continued until 1909 to try to prove that the intraneuronal inclusions named after him corresponded to steps in the developmental cycle of a protozoan.