Neural representations of complex temporal modulations in the human auditory cortex.

Natural sounds such as speech contain multiple levels and multiple types of temporal modulations. Because of nonlinearities of the auditory system, however, the neural response to multiple, simultaneous temporal modulations cannot be predicted from the neural responses to single modulations. Here we show the cortical neural representation of an auditory stimulus simultaneously frequency modulated (FM) at a high rate, f(FM) approximately 40 Hz, and amplitude modulation (AM) at a slow rate, f(AM) <15 Hz. Magnetoencephalography recordings show fast FM and slow AM stimulus features evoke two separate but not independent auditory steady-state responses (aSSR) at f(FM) and f(AM), respectively. The power, rather than phase locking, of the aSSR of both decreases with increasing stimulus f(AM). The aSSR at f(FM) is itself simultaneously amplitude modulated and phase modulated with fundamental frequency f(AM), showing that the slow stimulus AM is not only encoded in the neural response at f(AM) but also encoded in the instantaneous amplitude and phase of the neural response at f(FM). Both the amplitude modulation and phase modulation of the aSSR at f(FM) are most salient for low stimulus f(AM) but remain observable at the highest tested f(AM) (13.8 Hz). The instantaneous amplitude of the aSSR at f(FM) is successfully predicted by a model containing temporal integration on two time scales, approximately 25 and approximately 200 ms, followed by a static compression nonlinearity.