Trevor M. Shackleton

The role of resolved and unresolved harmonics in pitch perception and frequency modulation discrimination

A series of experiments investigated the influence of harmonic resolvability on the pitch of, and the discriminability of differences in fundamental frequency (F0) between, frequency-modulated (FM) harmonic complexes. Both F0 (62.5 to 250 Hz) and spectral region (LOW: 125-625 Hz, MID: 1375-1875 Hz, and HIGH: 3900-5400 Hz) were varied orthogonally. The harmonics that comprised each complex could be summed in either sine (0 degree) phase (SINE) or alternating sine-cosine (0 degree-90 degrees) phase (ALT). Stimuli were presented in a continuous pink-noise background. Pitch-matching experiments revealed that the pitch of ALT-phase stimuli, relative to SINE-phase stimuli, was increased by an octave in the HIGH region, for all F0s, but was the same as that of SINE-phase stimuli when presented in the LOW region. In the MID region, the pitch of ALT-phase relative to SINE-phase stimuli depended on F0, being an octave higher at low F0s, equal at high F0s, and unclear at intermediate F0s. The same stimuli were then used in three measures of discriminability: FM detection thresholds (FMTs), frequency difference limens (FDLs), and FM direction discrimination thresholds (FMDDTs, defined as the minimum FM depth necessary for listeners to discriminate between two complexes modulated 180 degrees out of phase with each other). For all three measures, at all F0s, thresholds were low (< 4% for FMTs, < 5% for FMDDTs, and < 1.5% for FDLs) when stimuli were presented in the LOW region, and high (> 10% for FMTs, > 7% for FMDDTs, and > 2.5% for FDLs) when presented in the HIGH region. When stimuli were presented in the MID region, thresholds were low for low F0s, and high for high F0s. Performance was not markedly affected by the phase relationship between the components of a complex, except for stimuli with intermediate F0s in the MID spectral region, where FDLs and FMDDTs were much higher for ALT-phase stimuli than for SINE-phase stimuli, consistent with their unclear pitch. This difference was much smaller when FMTs were measured. The interaction between F0 and spectral region for both sets of experiments can be accounted for by a single definition of resolvability.

Implementation details of a computation model of the inner hair‐cell auditory‐nerve synapse

A simple and computationally efficient model of auditory-neural transduction at the inner hair cell has recently been described, (Meddis, 1986a nd 1988). This paper briefly presents a short computer program to implement the model, an exploration of the effects of modifying the parameters of the model, anew set of parameters for simulating an auditory nerve fiber showing amedium rate of spontaneous activity with extended ynamic range, and some methods of quickly estimating some of the characteristics. It is intended as advice for researchers who wish to implement he model as part of a speech recognition device or as input to another model of more centrally located neurophysiological functions.

Comparing the fundamental frequencies of resolved and unresolved harmonics: Evidence for two pitch mechanisms?

Four experiments measured sensitivity (d′) to differences in fundamental frequency (F0) between two simultaneously presented groups of frequency‐modulated harmonics. Each group was passed through a bandpass filter in either a LOW (125–625 Hz), MID (1375–1875 Hz), or HIGH (3900–5400 Hz) frequency region. In the first two experiments, a dynamic F0 difference (ΔF0) was created by introducing a 180° disparity between the frequency modulations imposed on the two groups. Experiment 1 measured sensitivity to such ΔF0’s between a MID group with a baseline F0 of 125 Hz and all components summed in sine phase, and a HIGH group, in four conditions. When the baseline F0 of the HIGH group was also 125 Hz, performance was good when its components were summed in sine phase and bad when they were in alternating phase. Conversely, when the HIGH F0 was 62.5 Hz, performance was better for alternating phase than for sine phase, consistent with alternating phase doubling the internal representation of HIGH group’s F0. Similar...

Across frequency integration in a model of lateralization

A computational model of binaural lateralization is described. An accurate model of the auditory periphery feeds a tonotopically organized multichannel cross‐correlation mechanism. Lateralization predictions are made on the basis of the integrated activity across frequency channels. The model explicitly weights cross‐correlation peaks closer to the center preferentially, and effectively weights information that is consistent across frequencies more heavily because they have a greater impact in the across frequency integration. This model is complementary to the weighted‐image model of Stern et al. [J. Acoust. Soc. Am. 84, 156–165 (1988)], although the model described in this paper is physiologically more plausible, is simpler, and is more versatile in the range of input stimuli that are possible.

The ability of inferior colliculus neurons to signal differences in interaural delay

Sound localization in humans depends largely on interaural time delay (ITD). The ability to discriminate differences in ITD is highly accurate. ITD discrimination (Δ ITD) thresholds, under some circumstances, are as low as 10–20 μs. It has been assumed that thresholds this low could only be obtained if the outputs from many neurons were combined. Here we use Receiver Operating Characteristic analysis to compute neuronal Δ ITD thresholds from 53 cells in the inferior colliculus in guinea pigs. The Δ ITD thresholds of single neurons range from several hundreds of μs down to 20–30 μs. The lowest single-cell thresholds are comparable to human thresholds determined with similar stimuli. This finding suggests that the highly accurate sound localization of human observers is consistent with the resolution of single cells and need not reflect the combined activity of many neurons.

Onset Neurones in the Anteroventral Cochlear Nucleus Project to the Dorsal Cochlear Nucleus

Considerable circumstantial evidence suggests that cells in the ventral cochlear nucleus, that respond predominantly to the onset of pure tone bursts, have a stellate morphology and project, among other places, to the dorsal cochlear nucleus. The characteristics of such cells make them leading candidates for providing the so-called “wideband inhibitory input” which is an essential part of the processing machinery of the dorsal cochlear nucleus. Here we use juxtacellular labeling with biocytin to demonstrate directly that large stellate cells, with onset responses, terminate profusely in the dorsal cochlear nucleus. They also provide widespread local innervation of the anteroventral cochlear nucleus and a small innervation of the posteroventral cochlear nucleus. In addition, some onset cells project to the contralateral dorsal cochlear nucleus.

Organisation of binaural interactions in the primary and dorsocaudal fields of the guinea pig auditory cortex

This study investigated the nature and topography of binaural interactions in the primary auditory field (AI) and dorsocaudal field (DC) of the urethane anaesthetised guinea pig auditory cortex. Single and multi-units were classified by their responses to monaural and binaural stimulation. In both AI and DC, units displayed binaural facilitation, binaural inhibition, or a level dependent mixture of facilitation and inhibition. There was a significant difference in the distribution of binaural response types between the two fields. Facilitated units predominated in DC (facilitated: 58%; inhibited: 24%; mixed: 6%; non-interacting: 12%), while inhibited units were the most common class in AI (facilitated: 15%; inhibited: 44%; mixed: 18%; non-interacting: 22%). It has previously been suggested that inhibited and facilitated units are concerned with processing different areas of space suggesting a possible separation of function between the two core fields. Topographically, the binaural response properties in AI and DC varied along isofrequency bands, with neurones displaying similar interactions aggregating in clusters. These clusters were similar in size for the two fields and often overlapped neighbouring isofrequency bands. However, their shape and position varied between different animals. This clustered organisation of binaural interactions is similar to that reported in recent studies of AI in other mammals.

Phase-locked responses to pure tones in the primary auditory cortex.

At the level of the brainstem, precise temporal information is essential for some aspects of binaural processing, while at the level of the cortex, rate and place mechanisms for neural coding seem to predominate. However, we now show that precise timing of steady-state responses to pure tones occurs in the primary auditory cortex (AI). Recordings were made from 163 multi-units in guinea pig AI. All units increased their firing rate in response to pure tones at 100 Hz and 46 (28%) gave sustained responses which were synchronised with the stimulus waveform (phase-locking). The phase-locking units were clustered together in columns. Phase-locking was generally strongest in layers III and IV but was also recorded in layers I, II and V. Good phase-locking was observed over a range of 60-250 Hz: some units (30%) were narrow band while others (37%) were low-pass (33% were not determined). Phase-locking strength was also influenced by sound level: some units showed monotonic increases in strength with level and others were non-monotonic. Ten of the units provided a good temporal representation of the fundamental frequency (270 Hz) of a guinea pig vocalisation (rumble) and may be involved in analysing communication calls.

Sensitivity to Interaural Correlation of Single Neurons in the Inferior Colliculusof Guinea Pigs

Sensitivity to changes in the interaural correlation of 50-ms bursts of narrowband or broadband noise was measured in single neurons in the inferior colliculus of urethane-anaesthetized guinea pigs. Rate vs. interaural correlation functions (rICFs) were measured using two methods. These methods compensated in different ways for the inherent variance in interaural correlation between tokens with the same expected correlation. The shape of all rICFs could be best described by power functions allowing them to be summarized by two parameters. Most rICFs were best fit by a power below 2, indicating that they were only slightly nonlinear. However, there were a few fitted functions that had a power of 3–6, indicating marked curvature. Modeling results indicate that the nonlinearity of the majority of rICFs was explicable in terms of the monaural transduction stages; however, some of the rICFs with power greater than 2 require either multiple inputs to the coincidence detector or additional nonlinearities to be included in the model. Discrimination thresholds were estimated at reference correlations of −1, 0, and +1 using receiver operating characteristic (ROC) analysis of the spike-count distribution at each correlation. Thresholds spanned the full possible range, from a minimum of 0.1 to the maximum possible of 2. Thresholds were generally highest with a reference correlation of −1, intermediate with a reference of 0, and lowest with a reference correlation of +1. Thresholds were lowest for the most steeply sloped rICFs, but thresholds were not strongly correlated to the spike rate variance. The lowest thresholds occurred using narrowband noise that was compensated for internal delays, but they were still about three times larger than human psychophysical thresholds measured using similar stimuli. The data suggest that, unlike pure tone interaural time difference, discrimination of a population measure is required to account for behavioral interaural correlation discrimination performance.

Perceived contrast of gratings and plaids: Non-linear summation across oriented filters

Plaids composed of two orthogonal sine-wave gratings appeared to be of lower contrast than single gratings of the same Michelson luminance contrast. This effect for plaids was obtained at all spatial frequencies (1-16 c/deg) and contrast levels (2-32%). Contrast-matching data plotted as a function of the angle between plaid components (0-90 deg) and as a function of spatial frequency and standard contrast level were consistent with a model in which the response of each orientation-tuned spatial frequency channel is a threshold-corrected power function of contrast, and is followed by quadratic summation of responses across all channels. The best-fitting contrast-response exponent in the main experiment was 0.63. Analysis of several other data-sets suggested a slightly higher value, 0.80. The same model gave a good account of contrast-matching between simple and compound (two-component) one dimensional gratings, accounting in particular for the apparent increase in contrast summation exponent at low contrasts reported by Quick, Hamerly and Reichert [(1976) Vision Research, 16, 351-355]. The model can, with one further assumption, account for the finding that contrast-matching between sine-wave and square-wave gratings depended only on the amplitude at the fundamental frequency. Comparison with contrast discrimination studies suggests that internal noise (variance of a channels contrast-response) is not constant, but increases approximately in proportion to the mean response.