Neural temporal coding of low pitch. I. Human frequency-following responses to complex tones

The neural basis of low pitch was investigated in the present study by recording a brainstem potential from the scalp of human subjects during presentation of complex tones which evoke a variable sensation of pitch. The potential recorded, the frequency-following response (FFR), reflects the temporal discharge activity of auditory neurons in the upper brainstem pathway. It was used as an index of neural periodicity in order to determine the extent to which the low pitch of complex tones is encoded in the temporal discharge activity of auditory brainstem neurons. A tone composed of harmonics of a common fundamental produces a sensation of pitch equal to that of the 'missing' fundamental. Such signals generate brainstem potentials which are spectrally similar to FFR recorded in response to sinusoidal signals equal in frequency to the missing fundamental. Both types of signals generate FFR which are periodic, with a frequency similar to the perceived pitch of the stimuli. It is shown that the FFR to the missing fundamental is not the result of a distortion product by recording FFR to a complex signal in the presence of low-frequency bandpass noise. Neither is the FFR the result of neural synchronization to the waveform envelope modulation pattern. This was determined by recording FFR to inharmonic and quasi-frequency-modulated signals. It was also determined that the 'existence region' for FFR to the missing fundamental lies below 2 kHz and that the most favorable spectral region for FFR to complex tones is between 0.5 and 1.0 kHz. These results are consistent with the hypothesis that far-field-recorded FFR does reflect neural activity germane to the processing of low pitch and that such pitch-relevant activity is based on the temporal discharge patterns of neurons in the upper auditory brainstem pathway.