Excitability of nerve-free hydra.
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Biomedical subjects
Publications and source records attributed to R K Josephson.
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The wing muscles used in singing by the katydid, Neoconocephalus robustus, are extraordinarily fast. At 35 degrees C, the animal's thoracic temperature during singing, an isometric twitch lasts only five to eight msec (onset to 50% relaxation) and the fusion frequency of these muscles is greater than 400 Hz. Stimulating the motornerve to a singing muscle initiates a short (2.5 msec at 35 degrees C), sometimes overshooting depolarization of the muscle fibers. Despite their spike-like appearance, the electrical responses are largely synaptic potentials. The muscle membrane appears to be capable of only weak, electrically-excitable, depolarizing electrogenesis. The short synaptic potentials result in part from rapidly-developing delayed rectification, in part from a low resting membrane resistance (Rm = 162 omega cm2) and a concomitantly short membrane time constant (about 1.5 msec).
Striated muscle is obviously a versatile tissue, one which has been malleable to selective pressure and has become modified for many specific tasks. The variations which adapt striated muscle to particular functions involve both changes in its structural organization and changes in the chemical nature of its components. Although a number of factors have been identified which contribute to the diversity of muscle performance, it is not yet possible to account adequately for the wide range in muscle performance throughout the animal kingdom. While not a new direction in comparative physiology, developing quantitative explanations for the diversity of muscle performance is still an obvious, remaining task.
Singing muscles of the katydid, Neoconocephalus robustus (Insecta, Tettigoniidae) are neurogenic, yet perform at contraction-relaxation frequencies as high as 212 Hz (Josephson and Halverson, '71). The mechanical and electrical responses of different bands of one of these muscles (the dorsal longitudinal muscle, DLM) has been examined with respect to ultrastructural features of each part which may be related to muscle performance. The DLM is composed of three bands and is innervated by four motoneurones. The cell bodies of three of these motoneurones occur ipsilaterally in the prothroracic ganglion; the cell body of the other motoneurone is contralateral in the mesothoracic ganglion. Three of the motoneurones (as yet unidentified fast axons) initiate extraordinarily fast twitches (rise time equal 7.3 msec, half duration equals 14.3 msec, 25 C), the fourth (an unidentified slower axon) evokes twitches which are considerably slower (rise time equals 18.9 msec, half duration equals 5.10 msec). Whereas the ventral and medial bands of the muscle are innervated only by fast axons (some fibers of the medial band are doubly innervated), the dorsal band is innervated by both a fast axon and the slower axon. A few fibers of the dorsal band are doubly innervated. The structure of fibers from the ventral and medial bands is very similar, with short sarcomeres (4.0 and 4.3 mum, respectively) and thin strap-like myofibrils delineated by well-developed sarcoplasmic reticulum (SR). Twenty-four percent of the volume of ventral band fibers is SR and the diffusion distance from SR to the center of the adjacent myofibril averages 0.083 mum. Twenty percent of the medial band fiber volume is SR, with a diffusion distance of 0.118 mum. Ventral and medial band fibers contain about 40% mitochondria, and 33% myofibrils. The dorsal band fibers have longer sarcomeres (9.5 mum), and only 10% of the fiber volume is SR. The muscle fibrils of the dorsal band are larger and consequently the diffusion distance is greater (0.227 mum) than in the ventral and medial bands. Mitochondria comprise 23% of the volume of dorsal band fibers. Most dorsal band mitochondria are aggregated into distinct clumps. Although some dorsal band fibers are innervated by a fast axon and some by the slower axon, the dorsal band fibers are structurally homogeneous, suggesting that neurotrophic effects are not important in maintaining the structure of dorsal band fibers. The mechanical-electrical performance and ultrastructure of the ventral and medial bands suggest their roll as fast, metabolically active but weak muscles, used in singing; the dorsal band as a slower but stronger muscle, perhaps involved in postural movements of the wing during singing.
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There is a maintained electrical potential of 15 to 40 millivolts across the two epithelial layers forming the body wall of Hydra, the inside of the animal being positive. Negativegoing (depolarizing) spikes are recorded spontaneously and sometimes in response to depolarizing current pulses. These spikes usually overshoot the zero potential level. The large size of the spikes and the orientation of the potential difference across the body wall indicate that this electrical activity is epithelial rather than nervous in origin.
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