Since Andrewes (1922) described “specific” and “nonspecific” flagellar antigens in several
Salmonella types, the diphasic state has been recognized in hundreds of
Salmonella serotypes and has been recognized also in the Arizona group (Edwards and West, 1945) . Similar variations have been found in the
Escherichia freundii (Citobacter) group by Edwards (1946), in the Hafnia group by Deacon (1952), and in the
Aerobacter cloacae (Cloaca) group by Sakazaki and Namioka (1960) .
Salmonella types which had complex phases and which gave rise to simpler types through loss variation were reported. Edwards and Bruner (1942) described
S. salinatis (4, 12: d, e, h: d, e, n, z
15) which could be changed irreversibly to
S. san diego (4, 12: e, h: e, n, z
15) while Edwards, Kauffmann and Huey (1957) found that
S. montgomery (11: d, a: d, e, n, z
15) was changed to
S. luciana (11: a: e, n, z
15) . Both of the changes resulted from the permanent loss of the common major antigenic constituent of phases 1 and 2 through cultivation in semisolid medium which contained d serum. Further, Edwards (1950) observed spontaneous irreversible segregation of e, h phases from more complex d, e, h phases. “Artificial” or induced phases often have been obtained by cultivation of salmonellae in homologous flagellar antisera. Thus, Kauffmann (1936) obtained j phases by cultivation of
S, typhi in d serum and Edwards and Bruner (1939) and Bruner and Edwards (1941) obtained induced phases from
S. abortus equi and
S. paratyphi A. Such induced phases were reverted to naturally occurring antigens only with difficulty if, indeed, reversion was accomplished.
It is known that
Salmonella types with similar H antigens yield similar induced phases. For example, salmonellae which contain antigen d tend to produce j phases when cultivated in homologous H antisera, and organisms which contained b antigens tend to yield z
33 phases (Edwards and Moran, 1946; Lederberg and Edwards, 1953) . These induced antigens function as phase 1 if the organism which yielded them originally was diphasic.
For instance,
S. paratyphi B when cultivated in b and 1, 2 antisera yields a diphasic form having the H formula z
33: 1, 2. Further, such induced phases replace phase 1 of other types in transduction experiments (Lederberg and Edwards, unpublished data) . Thus, while various phases have been induced in enteric bacteria, these apparently were artifacts which occurred rarely, if at all, in nature and resulted from changes produced in the natural antigens. These changes were not additive changes but were substitutions and in no instance was the number of flagellar phases of the organisms increased. Diphasic organisms, after change of one of the phases to an induced antigen, still exhibited variation only between two points of reference, as did the original culture.
It was not until recently that
Enterobacteriaceae which possessed three naturally occurring H antigens were found. Edwards and Fife (1960) recognized an Arizona serotype which had the antigenic formula 5, 29: 33: 21: 40. The three H phases, 33, 21, and 40, were known to occur naturally in other serotypes and were completely reversible at will in all directions. Later, Taylor and Lee (1960) recognized a culture of
S. Worthington which had the antigenic formula 1, 13, 23: z: 1, w: z
43. The third phase was known to occur naturally in a number of cultures of
S. senftenberg. The z
43 phase could be changed to z or 1, w at will and natural phase variation occurred between the z and 1, w components but neither of these was induced to revert to z
43 which was irretrievably lost.
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