ON THE MECHANISM OF TONIC IMMOBILITY IN VERTEBRATES.
1. The durations of successive periods of induced tonic immobility in the lizard Anolis carolinensis was examined as a function of temperature. An automatic recording method was employed and observations were made of 12,000 to 15,000 immobilizations with six animals over a temperature range of 5 degrees to 35 degrees C. during 5 months. 2. The durations of the immobile periods were found to vary rhythmically in most cases. The reciprocal of the duration of the rhythm, i.e., the rate of change of the process underlying the rhythms, when plotted as a function of temperature according to the Arrhenius equation show distributions of points in two straight line groups. One of these groups or bands of points extends throughout the entire temperature range with a temperature characteristic of approximately micro = 31,000 calories, and the other covers the range of 20 degrees to 35 degrees C. with micro equal to approximately 9,000 calories. 3. The initial stimulus in a series of inductions of immobility appears to set off a mechanism which determines the duration of the state of quiescence. Succeeding forced recoveries seem to have no effect on the normal duration of the rhythm. 4. These results are interpreted by assuming the release, through reflex stimulation, of hormonal substances, one effective between 5 degrees and 35 degrees C. and the other effective between 20 degrees and 35 degrees C. These substances are assumed to act as selective inhibitors of impulses from so called "higher centers," allowing impulses from tonic centers to pass to the muscles. 5. In some experiments a progressive lengthening in successively induced periods of immobility was observed. The logarithm of the frequency of recovery when plotted against time in most of these cases (i.e., except for a few in which irregularities occurred) gave a linear function of negative slope which was substantially unaffected by temperature. In these cases it is assumed that a diffusion process is controlling the amount of available A substance. 6. The results are similar to those obtained by Crozier with Cylisticus convexus. The duration of tonic immobility seems to be maintained in both arthropod and vertebrate by the chemical activity of "hormonal" selective inhibitors. The details of the mechanisms differ, but there is basic similarity. 7. Injections of small amounts of adrenalin above a threshold value are found to prolong the durations of tonic immobility of Anolis, by an amount which is a logarithmic function of the "dose." It is possible that internally secreted adrenalin, above a threshold amount, may be involved in the maintenance of tonic immobility. 8. The production of tonic immobility reflexly is a problem distinct from that of the duration of immobility. It is suggested that the onset may be induced by "shock" to the centers of reflex tonus causing promiscuous discharge of these centers with accompanying inhibition of the higher centers. Such a condition may result when an animal is suddenly lifted from the substratum and overturned, or when, as in the case of Anolis, it struggles with dorsum down. This reaction of the "tonic centers" may at the same time lead to discharge of the adrenal glands by way of their spinal connections thus prolonging the state.
Hoagland
H
H
Laboratory of General Physiology, Harvard University, Cambridge.