PHOTOTROPIC CIRCUS MOVEMENTS OF LIMAX AS AFFECTED BY TEMPERATURE

1. The theory of animal phototropism requires for particular instances a knowledge of the action of light as exerted through each of two bilaterally located receptors functioning singly. The measurement of "circus movements" which this involves must be concerned with such aspects of the reaction as are demonstrably dependent upon the effect of light. 2. The negatively phototropic slug Limax maximus exhibits very definite and continuous circus movement under vertical illumination when one eye-tentacle has been removed. The amplitude of the circling movement, measured in degrees deflection per cm. of path as an index of maintained differential tonus, is intimately related to the concurrent velocity of creeping. Analysis of the orienting mechanism is facilitated by the fact that in gasteropods such as Limax the animal creeps by means of the pedal organ, but orients (turns) by a totally distinct set of muscles in the dorsal and lateral regions of the body wall. 3. The expression of the phototropic orienting tendency, with illumination constant, is greatly influenced by the temperature. Above a zone centering at 15°, the amplitude of turning (degrees per cm. of path) is determined by the temperature in accurate agreement with Arrhenius' equation for chemical reaction velocity, with the critical increment µ = 16,820; and the rate of creeping is progressively less as the temperature rises, µ for its reciprocal being 10,900. Below 15°, the velocity of creeping becomes less the the lower the temperature, µ being again 16,800; while the amplitude of orientation is limited merely by the velocity of creeping, its reciprocal being directly proportional thereto. 4. Measurements of Limax circus movements in terms of turning deflection as function of light intensity must therefore be carried out at a temperature well above 15°. 5. The analysis provides a gross physical model of how an end-result may be influenced by temperature according to the effect of temperature upon each of several interconnected processes when the "temperature vs. effect" curves for these processes dynamically intersect. 6. It is pointed out that a certain type of unpredictability (quantative variability) in animal behavior under "normal" natural conditions probably results from dynamic equilibrium there obtaining between diverse mechanisms competing for effector control (in the present case, the creeping mechanism and that for turning, in the range 14–16°C.). It follows that the unraveling of the elements of conduct necessitates experimentation under diverse abnormal conditions favoring individual mechanism of response.