Ripple Analysis in Ferret Primary Auditory Cortex. II. Topographic and Columnar Distribution of Ripple Response Parameters

Abstract

We examined the columnar and topographic distribution of response parameters using spectral ripples and tonal stimuli in the primary auditory cortex (AI) of the barbiturate-anesthetized ferret. The ripple stimuli consisted of broadband stimuli (1-20 kHz) with sinusoidally modulated spectral envelopes.Responses to ripples were parametrized in terms of characteristic ripple Wo(ripple frequency where the magnitude of the ripple transfer function is maximal, i.e., where the cell responds best) and characteristic phase Fo (intercept of the phase of the ripple transfer function, i.e., phase where the cell responds best). The response area (measured with tones) was parametrized in terms of its excitatory bandwidth at 20 dB above threshold (BW20), and its asymmetry as reflected by the directional sensitivity index (C) to frequency-modulated (FM) tones. Columnar organization for the above four parameters was investigated in 66 single units from 23 penetrations. It was confirmed for Wo, Fo, and the C index, but it appeared to be ambiguous for BW20. The response parameters measured from multiunit recordings corresponded closely to those obtained from single units in the same cluster. In a local region, most cells exhibited closely matched, response fields (RFs, inverse Fourier transformed ripple transfer function) and response areas (measured with two-tone stimuli), and had correspondingly similar response parameters to ripples and tones. The topographic distribution of the response parameters across the surface of AI was studied with multiunit recordings in four animals. In all maps, systematic patterns or clustering of, response parameters could be discerned along the isofrequency planes.The distribution of the characteristic ripple Wo exhibited two trends. First, along the isofrequency planes, it was largest near the center of AI, gradually decreasing towards the edges of the field where often a secondary maximum was found.The second trend occurred along the tonotopic axis where the maximum Wo found in an isofrequency range increases with increasing BF. The tonal bandwidth BW20, which was inversely correlated with Wo, exhibited a similar topographic distribution along the tonotopic axis and the isofrequency planes. The distribution of the characteristic ripple phase, Fo which reflects the asymmetry in the response field, showed a systematic order along the isofrequency axis. At the center of AI symmetric responses (Fo 0) predominated. Towards the edges, the RFs became more asymmetric with Fo < 0 caudally, and Fo > 0 rostrally. The asymmetric response types tended to cluster along repeated bands that paralleled the tonotopic axis. The FM directional sensitivity (C index, reflecting asymmetry of tonal response areas) tends to have similar trends along the isofrequency axis as Fo.