Recording and spectrum analysis of the planarian electroencephalogram

Many animals produce continuous brainwaves, known as the electroencephalogram (EEG), but it is not known at what point in evolution the EEG developed. Planarians possess the most primitive form of brain, but still exhibit learning and memory behaviors. Here, we observed and characterized the EEG waveform of the planarian. We inserted a monopole electrode into the head of a planarian on a cold stage, and were able to observe the EEG at sub-microvolt amplitudes. The EEG had a continuous waveform, similar to that of evolutionarily advanced animals with more developed brains. Occasional myogenic potential spikes were observed in the EEG due to sticking of the electrode, but this was markedly diminished by cooling the sample, which enabled us to investigate the intrinsic character of the continuous EEG waveform. The frequency spectrum of the EEG was observed in the range of 0.1-5 Hz, showing a broad rise below 0.5 Hz and a monotonic decrease above 1 Hz, apparently following the 1/f law. The intensity of the total EEG diminished during anesthesia by cooling to 2-3 degrees C, and recovered when the sample was warmed to about 10 degrees C. The EEG signal was sustained for 30-40 min, and gradually weakened as the animal died. Stimulation of the planarian with water vibration at 0.5-2 Hz induced chaotic resonance with a broad peak spectrum of around the stimulation frequency. Strong illumination suppressed the EEG signals for several minutes, with the degree of suppression positively correlating with the intensity of the light. This provides evidence that the EEG responds to optical signals, although there were no synchronous reactions to light flashes. The continuous EEG waveform suggests the existence of feedback loop circuits in the neural network of the planarian, which was supposed in electric shock memory experiments [McConnell JV, Cornwell P, Clay M (1960) An apparatus for conditioning planaria. Am J Psychol 73:618-622]. However, because of the broad band character of chaotic resonance observed, these loops appear to be loose couplings between ganglia.