Licklider [Expcrientia 7, 128-133 (1951 } ] presented a theory of pitch highlighting the role of auditory-nerve interspike-interval timing information in the process of pitch extraction. His theory is simplified and amended and presented here as a computer implementation. This implementation has been successfully tested using simulations of a wide range of classical demonstrations of pitch phenomena including the missing fundamental, ambiguous pitch, pitch shift of equally spaced, inharmonic components, musical chords, repetition pitch, the pitch of interrupted noise, the existence region, and the dominance region for pitch. The theory is compared with a number of alternative theories and the physiological plausibility of a temporal model is considered.
A model of the mechanism of residue pitch perception is revisited. It is evaluated in the context of some new empirical results, and it is proposed that the model is able to reconcile a number of differing approaches in the history of theories of pitch perception. The model consists of four sequential processing stages: peripheral frequency selectivity, within-channel half-wave rectification and low-pass filtering, within-channel periodicity extraction, and cross-channel aggregation of the output. The pitch percept is represented by the aggregated periodicity function. Using autocorrelation as the periodicity extraction method and the summary autocorrelation function (SACF) as the method for representing pitch information, it is shown that the model can simulate new experimental results that show how the quality of the pitch percept is influenced by the resolvability of the harmonic components of the stimulus complex. These include: (i) the pitch of harmonic stimuli whose components alternate in phase; (ii) the increased frequency difference limen of tones consisting of higher harmonics; and (iii) the influence of a mistuned harmonic on the pitch of the complex as a function of its harmonic number. To accommodate these paradigms, it was necessary to compare stimuli along the length of the SACF rather than relying upon the highest peak alone. These new results demonstrate that the model responds differently to complexes consisting of low and high harmonics. As a consequence, it is not necessary to postulate two separate mechanisms to explain different pitch percepts associated with resolved and unresolved harmonics.
A probabilistic model is described for transmitter release from hair cells, auditory neuron EPSP's, and discharge patterns. The model assumes that the release fraction of the transmitter is a function of stimulus intensity. It further assumes that some of this transmitter substance is taken back into the cell while some is irretrievably lost from the cleft. These assumptions differ from other recent models which propose multiple release sites, fixed release fractions, and no transmitter reuptake. The model produces realistic mammalian rate intensity functions, interval and period histograms, incremental responses, and adaptation effects. It mimics successfully the adaptation of successive EPSP amplitudes of the afferent neuron of the goldfish sacculus and offers a reinterpretation of the implications of these studies for hair cell synaptic mechanism.
Cochlear nonlinearity was estimated over a wide range of center frequencies and levels in listeners with normal hearing, using a forward-masking method. For a fixed low-level probe, the masker level required to mask the probe was measured as a function of the masker-probe interval, to produce a temporal masking curve (TMC). TMCs were measured for probe frequencies of 500, 1000, 2000, 4000, and 8000 Hz, and for masker frequencies 0.5, 0.7, 0.9, 1.0 (on frequency), 1.1, and 1.6 times the probe frequency. Across the range of probe frequencies, the TMCs for on-frequency maskers showed two or three segments with clearly distinct slopes. If it is assumed that the rate of decay of the internal effect of the masker is constant across level and frequency, the variations in the slopes of the TMCs can be attributed to variations in cochlear compression. Compression-ratio estimates for on-frequency maskers were between 3:1 and 5:1 across the range of probe frequencies. Compression did not decrease at low frequencies. The slopes of the TMCs for the lowest frequency probe (500 Hz) did not change with masker frequency. This suggests that compression extends over a wide range of stimulus frequencies relative to characteristic frequency in the apical region of the cochlea.
A revised computational model of the inner-hair cell (IHC) and auditory-nerve (AN) complex is presented and evaluated. Building on previous models, the algorithm is intended as a component for use in more comprehensive models of the auditory periphery. It combines smaller components that aim to be faithful to physiology in so far as is practicable and known. Transduction between cochlear mechanical motion and IHC receptor potential (RP) is simulated using a modification of an existing biophysical IHC model. Changes in RP control the opening of calcium ion channels near the synapse, and local calcium levels determine the probability of the release of neurotransmitter. AN adaptation results from transmitter depletion. The exact timing of AN action potentials is determined by the quantal and stochastic release of neurotransmitter into the cleft. The model reproduces a wide range of animal RP and AN observations. When the input to the model is taken from a suitably nonlinear simulation of the motion of the cochlear partition, the new algorithm is able to simulate the rate-intensity functions of low-, medium-, and high-spontaneous rate AN fibers in response to stimulation both at best frequency and at other frequencies. The variation in fiber type arises in large part from the manipulation of a single parameter in the model: maximum calcium conductance. The model also reproduces quantitatively phase-locking characteristics, relative refractory effects, mean-to-variance ratio, and first- and second-order discharge history effects.
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