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The neuronal representation of pitch in primate auditory cortex


Pitch perception is critical for identifying and segregating auditory objects1, 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 species2,3. Pitch is the subjective attribute of a sound's fundamental frequency (f0) 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 f0. Even when the acoustic energy at f0 is removed (‘missing fundamental’) the same pitch is still perceived1. 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 f0, providing a neural correlate for pitch constancy1. 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 humans4,5.

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Figure 1: An example of a pitch-selective neuron (unit M36n-532).
Figure 2: Location and characteristic frequency distribution of the pitch area in marmoset auditory cortex.
Figure 3: Pitch-selective neurons share a similar tuning for pure tones and MFs.
Figure 4: Pitch-selective neurons are sensitive to pitch salience.
Figure 5: MF responses are not caused by combination tones.


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This work was supported by NIH grants to X.W. and D.B. We thank B. Delgutte, D. McAlpine, E. Young, B. Moore and members of the Laboratory of Auditory Neurophysiology for their comments and suggestions related to this manuscript, and A. Pistorio, E. Bartlett and E. Issa for assistance with animal care. E. Issa contributed data to the characteristic frequency maps.Author Contributions D.B. and X.W. designed the experiment and co-wrote the paper. D.B. carried out the electrophysiological recordings and data analysis.

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Correspondence to Daniel Bendor or Xiaoqin Wang.

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Supplementary information

Supplementary Figure S1

This shows responses of pitch-selective and non-pitch neurons to pure tone MFs. (PDF 876 kb)

Supplementary Figure S2

This figure shows the cortical frequency maps and location of pitch-selective neurons in two of the subjects as well as the distribution of standard deviation of CF across the maps of all three subjects (PDF 770 kb)

Supplementary Figure S3

This figure shows two examples of a pitch-selective neuron’s response to pure tone and MFs (PDF 407 kb)

Supplementary Figure S4

This figure shows three examples of a pitch-selective neuron's response to complex sounds (irregular click trains and iterated ripple noise) varying in their pitch salience (PDF 619 kb)

Supplementary Figure S5

This figure shows the response of pitch selective neurons to the noise masker played alone, as well as an example of a pitch selective neuron's response to pure tones and MFs in the presence of a noise masker. (PDF 402 kb)

Supplementary Figure Legends

Text to accompany the above Supplementary Figures. (DOC 26 kb)

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Bendor, D., Wang, X. The neuronal representation of pitch in primate auditory cortex. Nature 436, 1161–1165 (2005).

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