On the Temperature Behavior of Pulse Propagation and Relaxation in Worms, Nerves and Gels

The effect of temperature on pulse propagation in biological systems has been an important field of research. Environmental temperature not only affects a host of physiological processes e.g. in poikilotherms but also provides an experimental means to investigate the thermodynamic phenomenology of nerves and muscle. In the present work, the temperature dependence of blood vessel pulsation velocity and frequency was studied in the annelid Lumbriculus variegatus. The pulse velocity was found to vary linearily between 0°C and 30°C. In contrast, the pulse frequency increased non-linearly in the same temperature range. A heat block ultimately resulted in complete cessation of vessel pulsations at 37.2±2.7°C (lowest: 33°C, highest: 43°C). However, quick cooling of the animal led to restoration of regularly propagating pulses. This experimentally observed phenomenology of pulse propagation and frequency is interpreted without any assumptions about molecules in the excitable membrane (e.g. ion channels) or their temperature-dependent behaviour. By following Einstein's approach to thermodynamics and diffusion, a relation between relaxation time τ and compressibility κ of the excitable medium is derived that can be tested experimentally (for κT ∼ κS). Without fitting parameters this theory predicts the temperature dependence of the limiting (i.e. highest) pulse frequency in good agreement with experimental data. The thermodynamic approach presented herein is neither limited to temperature nor to worms nor to living systems. It describes the coupling between pulse propagation and relaxation equally well in nerves and gels. The inherent consistency and universality of the concept underline its potential to explain the dependence of pulse propagation and relaxation on any thermodynamic observable.