A B S T R A C TBridges between microtubules have been studied with the electron microscope in the axostyle of Saccinobaculus and in various tubule systems of chicken testis, including the helix of tubules surrounding the elongating spermatid nucleus and the flagellum of the sperm tail. In addition to the previously described periodic bridges, evidence is presented that nonperiodic bridges exist between certain tubules. An analysis of axial spacing between adjacent nonperiodic bridges suggests that these structures are attached to periodic binding sites on the microtubule wall, but that not all the binding sites are filled. The bridges appear nonperiodic as a result of random occupancy of some fraction of the periodic sites. The distribution of these binding sites is related to the substructure of the microtubule wall as seen with negative staining and optical diffraction.
I N T R O D U C T I O NThere are numerous examples in the literature of highly ordered arrays of microtubules. Reported studies include: the axostyle of certain flagellates (25, 26, 8, 29, 41, and 39); the axopods of heliozoa (34,31, 35,58,57,59,56,48,49,52,55); the tentacles of suctoria (50, 3); the cytopharyngial basket of some ciliates (60,61,62); the cortical fibers in other ciliates (2, 35 a, 32); transient structures in a variety of spermatids (10,40,46,18); permanent structures in other sperm (5,44,45,47); and the ubiquitous 9 -t-2 structure of cilia and flagella (19, 23, i, 51, 43, 30, 63, 64, 65). Studies on each of these systems have revealed thin bridges between the component microtubules. Several investigators have suggested that the bridges are important for maintaining the order of the microtubule arrays (34,26,40,55,48,4,56,39). Evidence supporting this conjecture includes the observation that the interrelationship of the outer doublets in the 9 -I-2 array can be maintained during solubilization of the other structural components of a flagellum until only the A subtubules and periodic links between them remain (53). From this evidence and our general knowledge of how macromolecular aggregates are put together in biological systems (1 I, 12), it seems reasonable to conclude that bridges are important for the establishment of specific intertubule geometries in ordered rnicrotubule arrays.Intertubule bridges have attracted additional attention because of the possibility that some of them may be functionally analogous to myosin, and may play a direct role in certain examples of cell motility. A few arrays of microtubules are motile organelles (e.g., cilia and flagella, the axostyle, coccid sperm tubule bundles [44], and the mitotic spindle), and others have motion associated with them (e.g., nuclear migration in virus-induced syncytia [29 a]; granule motion in chromatophores [6 a], in the arms of heliozoa [58], and of suctoria [50]; and the elongation of certain spermatid nuclei [40]). It has been postulated that intertubule bridges and projections from tubule surfaces are transducers which hydrolyze 166