Localized Activation of Bending in Proximal, Medial and Distal Regions of Sea-Urchin Sperm Flagella

Spermatozoa from the sea urchin, Colobocentrotus atratus, were partially demembranated by extraction with solutions containing Triton X-100 at a concentration which was insufficient to solubilize the membranes completely. The resulting suspension was a mixture containing some spermatozoa in which a proximal, medial, or distal portion of the flagellum was membrane-covered, while the remaining portion was naked axoneme. In reactivating solutions containing 12 µM ATP, only the naked portions of the flagellum became motile. In reactivating solutions containing 0.8 mM ADP, the membrane-covered regions became motile and beat at 6-10 beats/s, while the naked regions remained immobile, or beat very slowly at about 0.3 beat/s. Activation of membrane-covered regions in ADP solutions probably results from the membrane restricting the diffusion of ATP which is formed from ADP by the axonemal adenylate kinase. The results indicate that any region of the flagellum has the capacity for autonomous beating, and that special properties of the basal end of the flagellum are not required for bend initiation. However, the beating of different regions of the flagellum is not completely independent, for in a fair number of spermatozoa the beating of the distal, membrane-covered region in 0.8 mM ADP was intermittent, and was turned on and off in phase with the much slower bending cycle in the proximal region of naked axoneme.