A chromosomal region present in Salmonella typhimurium but absent from related species was identified by hybridization. A DNA probe originating from 78 min on the S. typhimurium chromosome hybridized with DNA from Salmonella enteritidis, Salmonella heidelberg, and Salmonella dublin but not with DNA from Salmonella typhi, Salmonella arizonae, Escherichia coli, and Shigella serotypes. Cloning and sequence analysis revealed that the corresponding region of the S. typhimurium chromosome encodes a fimbrial operon. Long fimbriae inserted at the poles of the bacterium were observed by electron microscopy when this fimbrial operon was introduced into a nonpiliated E. coli strain. The genes encoding these fimbriae were therefore termed lpfABCDE, for long polar fimbriae. Genetically, the lpf operon was found to be most closely related to the fim operon of S. typhimurium, both in gene order and in conservation of the deduced amino acid sequences.All members of the family Enterobacteriaceae express fimbriae, or pili, which allow them to attach to host surfaces. This attachment is the first step in the colonization of their preferred host niche. The various fimbrial adhesins were originally distinguished by their receptor specificity. Hemagglutination of erythrocytes was used as a criterion to define three fimbrial classes (for a recent review, see reference 14). Fimbriae whose agglutination is prevented by coincubation with D-mannose were designated type 1 fimbriae (41). Type 2 fimbriae, a nonhemagglutinating fimbrial class, were later found to resemble nonadhesive variants of type 1 fimbriae (20,36,43). Finally, pili able to agglutinate animal erythrocytes only after the erythrocytes have been treated with tannic acid were designated type 3 fimbriae (22, 23). This early classification was supported by data on the morphological characteristics of these fimbriae. Type 1 and type 2 fimbriae have a width of 7 to 8 nm and display a channeled rodlike appearance due to the arrangement of subunits around a hollow core (43). Type 3 fimbriae are 4 to 5 nm in diameter and lack an axial hole on the basis of electron microscopic examination (42). Recently, thin fimbriae (3 to 4 nm) with a unique curly morphology in members of the Enterobacteriaceae have been described (4,44). A comparison of sequence data obtained from several fimbrial systems with similar morphologies revealed that the major fimbrial subunits of curly fimbriae show a high degree of sequence conservation. These fimbriae are therefore now considered to form a distinct fimbrial class. The designation GVVPQ has been proposed, in reference to a short amino acid sequence conserved at the N terminus of their major fimbrial subunit (19). Type 1, type 2, type 3, and GVVPQ fimbriae have the common characteristic that their fimbrial subunits contain signal sequences which are cleaved by signal peptidase I. In contrast, a fifth enterobacterial class of pili, termed type 4 fimbriae, has been shown to use a different export system for translocation of its fimbrial subunits across t...