The centriolar apparatus in the flagellate genera Trichonympha and Pseudotrichonympha is located at the anterior end of the cell, in the rostrum. It forms part of a complex system of structures which includes the rostral tube, inner and outer caps, and the rostral flagella. The fine structure of these organelles is described in detail on the basis of an electron-microscope study of sectioned and negatively stained material. In Trichonympha the rostral tube is a hollow cylinder, made of a cross-striated protein with a periodicity of about 450 Å. This is organized into radially arranged lamellae, which continue posteriorly as the parabasal filaments. The tube is continuous anteriorly with two finely striated crescentic bodies, which correspond to the so-called short centrioles of some previous workers. There is no evidence that they are centriolar in function. In the interphase cell the centriolar apparatus consists principally of a long centriolar rod, of complex fine structure, lying in the anterior end of the rostral tube. There is no evidence of typical centriolar structure in this, but at division an aster forms around one end of it. Surmounting the apex of the rostral tube is a dense, finely fibrous body, the inner cap. Lying within this there is a typical centriole (similar in structure to a basal body), and also the basal body of one flagellum, which appears to be distinct from all the rest. The functions of these two structures are not known. The margin of the inner cap connects with the complex system of delicate fibres which links the basal bodies of the rostral flagella. The function of the fibres, and possibly also of the inner cap, may be to coordinate the activities of the rostral flagella. The outer cap is composed mainly of tubules, 250 Å in diameter, but shows variations in structure in different species. The structures in Pseudotrichonympha which presumably serve similar functions are in many respects very differently organized. The rostral tube is more complex, with distinct inner and outer walls of different fine structure. There are also complex inner and outer surface layers. A striking feature is that although the various components of the tube are quite different in structure, they display a common periodicity in their organization. The centriolar apparatus appears to consist principally of two dense bands running along the inner wall of the tube, connecting anteriorly to an extended layer of centriolar material to which spindle fibres are attached in radially polarized fashion throughout interphase. There is no centriolar rod or typical centriole, such as is found in Trichonympha . Very elaborate systems of fibres are associated with the inner cap and the anterior end of the rostral tube. The two genera are compared, and the findings related to knowledge of centriolar structures in other types of cell. Possible evolutionary explanations for the complexity and variation in fine structure in these flagellates are considered.