Attached to the basal bodies of Naegleria gruberi flagellates is a striated rootlet or rhizoplast . The rootlet-basal body complex has been isolated by Triton X-100 lysis of deflagellated cells and differential centrifugation through a 25% glycerol medium . Rootlets isolated from mature flagellates are -13 p,m long but vary from 8 to 15 ftm in length : they taper at both ends from a maximum width of -0 .25 p,m in the vicinity of the basal bodies . They are highly stable during isolation but can be solubilized by urea, high salt, low pH, or detergent (Sarkosyl) . Partial dissociation of rootlets with 1 M urea reveals that they are composed of filaments, -5 nm diameter, associated in linear fashion to yield the characteristic 21-nm cross-banded appearance . Differential solubilization of rootlets and their associated contaminants allowed identification of a major rootlet protein, comprising at least 50% of any purified rootlet preparation, with an apparent subunit molecular weight of 170,000. The localization of rootlets in situ by indirect immunofluorescence using a specific antibody directed against the purified rootlet protein demonstrated unequivocally that this 170,000-dalton protein is an organelle component.The flagella of eukaryotic cells are generally found as part of a flagellar apparatus-an interconnected assemblage of basal body, flagellum, and a striated rootlet fiber or rhizoplast . Likewise, cilia, basal bodies, and one or more rootlets often combine to constitute what could be called a "ciliary apparatus" in ciliated cells . Although flagellated and ciliated cells lacking rootlets are known, and even common (e .g., sperm cells, certain mammalian ciliated epithelia), some form of rootlet structure is attached to the basal bodies of most cilia and flagella (7,8,30) . Despite intense current interest in the structure, function, and morphogenesis of flagella and their basal bodies, virtually no attention has been paid to the other major component of the flagellar or ciliary apparatus, the rootlet. References to flagellar rootlets have most often been restricted to ultrastructural descriptions and to speculation about possible roles in motility (5,9,15,16,19,26,28,34,38) .Electron micrographs of rootlets generally reveal bundles of 4-to 5-nm-diameter filaments aggregated in a paracrystalline array to produce fibrous, cross-striated organelles reminiscent of collagen fibrils (9, 26, 28, 37) . The cross-striations form one major repeating unit that may be further divided into -13 intraperiod sub-bands (28, 38) . Reported widths for this major repeat unit range from 12 to 78 nm (34,35) . Though most authors describe a constant banding pattern, at least three reports have commented upon the variations in rootlet period-424 icity (15,32,34) and have inferred a possible physiological role for the altered states . The idea that the rootlet is a contractile element of the flagellar apparatus which may play an active role in modifying the flagellar beat is supported by the following : the recent demonst...