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      The harmonic organization of auditory cortex

      review-article
      1 , 2
      Frontiers in Systems Neuroscience
      Frontiers Media S.A.
      harmonicity, auditory cortex, marmoset, pitch, music

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          Abstract

          A fundamental structure of sounds encountered in the natural environment is the harmonicity. Harmonicity is an essential component of music found in all cultures. It is also a unique feature of vocal communication sounds such as human speech and animal vocalizations. Harmonics in sounds are produced by a variety of acoustic generators and reflectors in the natural environment, including vocal apparatuses of humans and animal species as well as music instruments of many types. We live in an acoustic world full of harmonicity. Given the widespread existence of the harmonicity in many aspects of the hearing environment, it is natural to expect that it be reflected in the evolution and development of the auditory systems of both humans and animals, in particular the auditory cortex. Recent neuroimaging and neurophysiology experiments have identified regions of non-primary auditory cortex in humans and non-human primates that have selective responses to harmonic pitches. Accumulating evidence has also shown that neurons in many regions of the auditory cortex exhibit characteristic responses to harmonically related frequencies beyond the range of pitch. Together, these findings suggest that a fundamental organizational principle of auditory cortex is based on the harmonicity. Such an organization likely plays an important role in music processing by the brain. It may also form the basis of the preference for particular classes of music and voice sounds.

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          Most cited references83

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          Subdivisions of auditory cortex and processing streams in primates.

          The auditory system of monkeys includes a large number of interconnected subcortical nuclei and cortical areas. At subcortical levels, the structural components of the auditory system of monkeys resemble those of nonprimates, but the organization at cortical levels is different. In monkeys, the ventral nucleus of the medial geniculate complex projects in parallel to a core of three primary-like auditory areas, AI, R, and RT, constituting the first stage of cortical processing. These areas interconnect and project to the homotopic and other locations in the opposite cerebral hemisphere and to a surrounding array of eight proposed belt areas as a second stage of cortical processing. The belt areas in turn project in overlapping patterns to a lateral parabelt region with at least rostral and caudal subdivisions as a third stage of cortical processing. The divisions of the parabelt distribute to adjoining auditory and multimodal regions of the temporal lobe and to four functionally distinct regions of the frontal lobe. Histochemically, chimpanzees and humans have an auditory core that closely resembles that of monkeys. The challenge for future researchers is to understand how this complex system in monkeys analyzes and utilizes auditory information.
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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

                Journal
                Front Syst Neurosci
                Front Syst Neurosci
                Front. Syst. Neurosci.
                Frontiers in Systems Neuroscience
                Frontiers Media S.A.
                1662-5137
                17 December 2013
                2013
                : 7
                : 114
                Affiliations
                [1] 1Department of Biomedical Engineering, Johns Hopkins University School of Medicine Baltimore, MD, USA
                [2] 2Tsinghua-Johns Hopkins Joint Center for Biomedical Engineering Research and Department of Biomedical Engineering, Tsinghua University Beijing, China
                Author notes

                Edited by: Jonathan B. Fritz, University of Maryland, USA

                Reviewed by: Preston E. Garraghty, Indiana University, USA; Michael Brosch, Leibniz Institute for Neurobiology, Germany; Jean-Marc Edeline, Université de Paris, France

                *Correspondence: Xiaoqin Wang, Department of Biomedical Engineering, Johns Hopkins University School of Medicine, 720 Rutland Avenue, Traylor 410, Baltimore, MD 21025, USA e-mail: xiaoqin.wang@ 123456jhu.edu

                This article was submitted to the journal Frontiers in Systems Neuroscience.

                Article
                10.3389/fnsys.2013.00114
                3865599
                24381544
                a2617648-bdfb-42d0-ada8-83f38483b58e
                Copyright © 2013 Wang.

                This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) or licensor are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.

                History
                : 16 July 2013
                : 01 December 2013
                Page count
                Figures: 5, Tables: 0, Equations: 0, References: 96, Pages: 11, Words: 9370
                Categories
                Neuroscience
                Review Article

                Neurosciences
                harmonicity,auditory cortex,marmoset,pitch,music
                Neurosciences
                harmonicity, auditory cortex, marmoset, pitch, music

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