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Zum Mechanismus der biologischen 24-Stunden-Periodik

Research paper by Wever Von Rütger

Indexed on: 01 May '63Published on: 01 May '63Published in: Biological Cybernetics



Abstract

The level (=arithmetic average of all instantaneous values)of a self-sustained oscillation in general influences all properties of the oscillation, including period, amplitude and shape of the oscillation, and the rate of exchange of energy between the oscillator and its environment. Only when the non-linear damping factor does not depend on the instantaneous value of the oscillating function, but only on the amplitude of the oscillation, are the other properties independent of the average level. The differential equations describing self-sustained oscillations cannot be solved exactly, but methods of approximation are applicable. Numerical solutions to several different forms of the equations will be discussed.In the simplest case (van der Pol equation) all properties of the self-sustained oscillation (e.g. period, amplitude) are extreme when the level is zero. The oscillation continues only within a given range of levels (oscillating range); outside this range, the oscillation damps out. In other modifications of the equation, the oscillating function cannot assume a zero value. In all cases, the extent to which the average level influences the different properties depends on the factor ɛ, which describes the position of the oscillation within the range between harmonic and relaxation types of oscillation.In the elementary van der Pol equation, the correlation between level and frequency changes sign within the oscillating range; that is, the circadian rule, demanding an always positive correlation between level and frequency, cannot be fulfilled. Only with an additional non-linearity in the energy of recoil does the correlation remain unchanged in sign throughout the oscillating range. A stability condition demands a positive sign for this non-linearity, and hence, for the correlation (fulfilling the circadian rule); if the sign is negative (violating the circadian rule), the oscillation becomes unstable. With an additional term of the third order, the oscillation acquires a two-peaked shape typical of many circadian oscillations.A simple differential equation describing all general properties of the circadian periodicity must fulfil these conditions: the oscillation must be self-sustained and limited to positive values; and the energy of recoil must be non-linear with a positive coefficient to obtain the appropriate correlation between level and frequency. In the equations here developed the environment directly influences only one parameter of the oscillation, i.e. the level. In addition to the circadian periodicity, the differential equations here examined describe the behavior of several other biological oscillations.