C. elegans: Sensing the low-frequency profile of amplitude-modulated ultrasound

Several research groups have demonstrated that C. elegans can respond to pulsed ultrasound stimuli, and elucidating the underlying mechanisms is necessary to develop ultrasound neuromodulation. Here, amplitude-modulated (AM) ultrasound is applied to C. elegans, and its behavioral responses are investigated in detail. By loading surface acoustic waves (SAWs) onto free-moving worms on an agar surface, a carrier wave with a frequency of 8.80 MHz is selected. The signal is modulated by a rectangular or sinusoidal profile. It is demonstrated that sinusoidal modulation can produce similar responses in worms to rectangular modulation, with the strongest responses occurring at modulation frequencies of around 1.00 kHz. Meanwhile, the behavioral response is relatively weak when the ultrasonic signal is unmodulated, that is, when only the carrier wave is applied. At modulation frequencies other than 100.00 Hz to 10.00 kHz, the worms respond weakly, but when a second modulation frequency of 1.00 kHz is introduced, an improvement in response can be observed. These results suggest that C. elegans may sense the low-frequency envelope and respond to amplitude-modulated ultrasonic stimuli like an amplitude demodulator. MEC-4, an ion channel for touch sensing, is involved in the behavioral response of C. elegans to ultrasound in the present setup.