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      The Frequency Following Response (FFR) May Reflect Pitch-Bearing Information But is Not a Direct Representation of Pitch

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          Abstract

          The frequency following response (FFR), a scalp-recorded measure of phase-locked brainstem activity, is often assumed to reflect the pitch of sounds as perceived by humans. In two experiments, we investigated the characteristics of the FFR evoked by complex tones. FFR waveforms to alternating-polarity stimuli were averaged for each polarity and added, to enhance envelope, or subtracted, to enhance temporal fine structure information. In experiment 1, frequency-shifted complex tones, with all harmonics shifted by the same amount in Hertz, were presented diotically. Only the autocorrelation functions (ACFs) of the subtraction-FFR waveforms showed a peak at a delay shifted in the direction of the expected pitch shifts. This expected pitch shift was also present in the ACFs of the output of an auditory nerve model. In experiment 2, the components of a harmonic complex with harmonic numbers 2, 3, and 4 were presented either to the same ear (“mono”) or the third harmonic was presented contralaterally to the ear receiving the even harmonics (“dichotic”). In the latter case, a pitch corresponding to the missing fundamental was still perceived. Monaural control conditions presenting only the even harmonics (“2 + 4”) or only the third harmonic (“3”) were also tested. Both the subtraction and the addition waveforms showed that (1) the FFR magnitude spectra for “dichotic” were similar to the sum of the spectra for the two monaural control conditions and lacked peaks at the fundamental frequency and other distortion products visible for “mono” and (2) ACFs for “dichotic” were similar to those for “2 + 4” and dissimilar to those for “mono.” The results indicate that the neural responses reflected in the FFR preserve monaural temporal information that may be important for pitch, but provide no evidence for any additional processing over and above that already present in the auditory periphery, and do not directly represent the pitch of dichotic stimuli.

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          Most cited references39

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          A cochlear frequency-position function for several species--29 years later.

          Accurate cochlear frequency-position functions based on physiological data would facilitate the interpretation of physiological and psychoacoustic data within and across species. Such functions might aid in developing cochlear models, and cochlear coordinates could provide potentially useful spectral transforms of speech and other acoustic signals. In 1961, an almost-exponential function was developed (Greenwood, 1961b, 1974) by integrating an exponential function fitted to a subset of frequency resolution-integration estimates (critical bandwidths). The resulting frequency-position function was found to fit cochlear observations on human cadaver ears quite well and, with changes of constants, those on elephant, cow, guinea pig, rat, mouse, and chicken (Békésy, 1960), as well as in vivo (behavioral-anatomical) data on cats (Schucknecht, 1953). Since 1961, new mechanical and other physiological data have appeared on the human, cat, guinea pig, chinchilla, monkey, and gerbil. It is shown here that the newer extended data on human cadaver ears and from living animal preparations are quite well fit by the same basic function. The function essentially requires only empirical adjustment of a single parameter to set an upper frequency limit, while a "slope" parameter can be left constant if cochlear partition length is normalized to 1 or scaled if distance is specified in physical units. Constancy of slope and form in dead and living ears and across species increases the probability that the function fitting human cadaver data may apply as well to the living human ear. This prospect increases the function's value in plotting auditory data and in modeling concerned with speech and other bioacoustic signals, since it fits the available physiological data well and, consequently (if those data are correct), remains independent of, and an appropriate means to examine, psychoacoustic data and assumptions.
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            The processing of temporal pitch and melody information in auditory cortex.

            An fMRI experiment was performed to identify the main stages of melody processing in the auditory pathway. Spectrally matched sounds that produce no pitch, fixed pitch, or melody were all found to activate Heschl's gyrus (HG) and planum temporale (PT). Within this region, sounds with pitch produced more activation than those without pitch only in the lateral half of HG. When the pitch was varied to produce a melody, there was activation in regions beyond HG and PT, specifically in the superior temporal gyrus (STG) and planum polare (PP). The results support the view that there is hierarchy of pitch processing in which the center of activity moves anterolaterally away from primary auditory cortex as the processing of melodic sounds proceeds.
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              The neuronal representation of pitch in primate auditory cortex.

              Pitch perception is critical for identifying and segregating auditory objects, especially in the context of music and speech. The perception of pitch is not unique to humans and has been experimentally demonstrated in several animal species. Pitch is the subjective attribute of a sound's fundamental frequency (f(0)) that is determined by both the temporal regularity and average repetition rate of its acoustic waveform. Spectrally dissimilar sounds can have the same pitch if they share a common f(0). Even when the acoustic energy at f(0) is removed ('missing fundamental') the same pitch is still perceived. Despite its importance for hearing, how pitch is represented in the cerebral cortex is unknown. Here we show the existence of neurons in the auditory cortex of marmoset monkeys that respond to both pure tones and missing fundamental harmonic complex sounds with the same f(0), providing a neural correlate for pitch constancy. These pitch-selective neurons are located in a restricted low-frequency cortical region near the anterolateral border of the primary auditory cortex, and is consistent with the location of a pitch-selective area identified in recent imaging studies in humans.
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                Author and article information

                Contributors
                +44-1223-273697 , +44-1223-359062 , hedwig.gockel@mrc-cbu.cam.ac.uk
                Journal
                J Assoc Res Otolaryngol
                JARO: Journal of the Association for Research in Otolaryngology
                Springer-Verlag (New York )
                1525-3961
                1438-7573
                9 August 2011
                9 August 2011
                December 2011
                : 12
                : 6
                : 767-782
                Affiliations
                [1 ]MRC-Cognition and Brain Sciences Unit, 15 Chaucer Road, Cambridge, CB2 7EF UK
                [2 ]Division of Human Communication and Deafness, The University of Manchester, Manchester, M13 9PL UK
                Article
                284
                10.1007/s10162-011-0284-1
                3214239
                21826534
                f4fc3980-94f0-4722-9e51-3a22cdf53c00
                © The Author(s) 2011
                History
                : 23 March 2011
                : 18 July 2011
                Categories
                Article
                Custom metadata
                © Association for Research in Otolaryngology 2011

                Otolaryngology
                dichotic presentation,complex tones,monaural temporal information
                Otolaryngology
                dichotic presentation, complex tones, monaural temporal information

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