The Caulobacter crescentus flagellar filament is assembled from multiple flagellin proteins that are encoded by six genes. The amino acid sequences of the FljJ and FljL flagellins are divergent from those of the other four flagellins. Since these flagellins are the first to be assembled in the flagellar filament, one or both might have specialized to facilitate the initiation of filament assembly.Caulobacter crescentus is a dimorphic bacterium that differentiates as part of its normal cell cycle. Each cell division results in a stalked cell and a motile swarmer cell. The swarmer cell has a single flagellum at one pole that is responsible for motility. The flagellum is similar in structure to enteric flagella except that the filament is composed of multiple flagellin proteins (6,10,21,35,38). The hook-proximal portion of the flagellar filament consists of a 60-nm segment containing a 29-kDa flagellin (6). The next segment is 1 to 2 m in length and consists of a 27-kDa flagellin along with increasing amounts of a 25-kDa flagellin at its distal end. The remaining 2 to 10 m of the filament contains the 25-kDa flagellin. Thus, the flagellar filament consists of at least three different flagellar proteins in a precise arrangement.In addition to the three flagellins found in the filament, stationary-phase wild-type cells and some flagellar mutants synthesize a 22-kDa flagellin (15, 34). The 22-kDa flagellin is not found in functional flagellar filaments and may result from improper processing of the 25-kDa flagellin (22,34).Multiple flagellin genes have been identified in two unlinked clusters in the C. crescentus genome (11, 34). The alpha gene cluster includes the fljJ, fljK, and fljL genes that encode the 29-, 25-, and 27-kDa flagellins, respectively. The gene order in this region is flaY-flaE-fljJ-fljK-fljL-flaF-flbT-flBA-flaG (30, 32). Thus, the alpha cluster of flagellin genes is embedded in a cluster of other flagellar genes. The beta flagellin gene cluster is approximately 1,000 kb from the alpha cluster (8). No other flagellar genes are present at this locus.The flagellin gene family. We determined the nucleotide sequences of five of the six flagellin genes in the alpha (fljK and fljL) and beta (fljM, fljN, and fljO) regions using the plasmids described in Table 1. The nucleotide sequence of the sixth gene, fljJ, had been determined previously (11), and errors in this sequence were corrected using nucleotide sequence information produced by The Institute for Genome Research. Each region encodes a set of three independently transcribed flagellin genes. A comparison of these sequences to the C. crescentus genome sequence being produced by The Institute for Genome Research indicated that no additional flagellin genes were present elsewhere in the genome. Thus, C. crescentus has six flagellin genes in two clusters of three genes.To facilitate comparison of members of the flagellin gene family, the nucleotide sequences of the flagellin genes were aligned. There were 455 variable sites in the 822-bp sequence.Most of...