Dual rate codes for the pitch of harmonic complex tones in the auditory midbrain

2017 ◽  
Vol 141 (5) ◽  
pp. 3891-3891
Author(s):  
Bertrand Delgutte ◽  
Yaqing Su
2019 ◽  
Author(s):  
Yaqing Su ◽  
Bertrand Delgutte

AbstractHarmonic complex tones (HCT) commonly occurring in speech and music evoke a strong pitch at their fundamental frequency (F0), especially when they contain harmonics individually resolved by the cochlea. When all frequency components of an HCT are shifted by the same amount, the pitch of the resulting inharmonic tone (IHCT) also shifts although the envelope repetition rate is unchanged. A rate-place code whereby resolved harmonics are represented by local maxima in firing rates along the tonotopic axis has been characterized in the auditory nerve and primary auditory cortex, but little is known about intermediate processing stages. We recorded single neuron responses to HCT and IHCT with varying F0 and sound level in the inferior colliculus (IC) of unanesthetized rabbits. Many neurons showed peaks in firing rates when a low-numbered harmonic aligned with the neuron’s characteristic frequency, demonstrating “rate-place” coding. The IC rate-place code was most prevalent for F0>800 Hz, was only moderately dependent on sound level over a 40 dB range, and was not sensitive to stimulus harmonicity. A spectral receptive-field model incorporating broadband inhibition better predicted the neural responses than a purely excitatory model, suggesting an enhancement of the rate-place representation by inhibition. Some IC neurons showed facilitation in response to HCT, similar to cortical “harmonic template neurons” (Feng and Wang 2017), but to a lesser degree. Our findings shed light on the transformation of rate-place coding of resolved harmonics along the auditory pathway, and suggest a gradual emergence of harmonic templates from low to high processing centers.Significance statementHarmonic complex tones are ubiquitous in speech and music and produce strong pitch percepts in human listeners when they contain frequency components that are individually resolved by the cochlea. Here, we characterize a “rate-place” code for resolved harmonics in the auditory midbrain that is more robust across sound levels than the peripheral rate-place code and insensitive to the harmonic relationships among frequency components. We use a computational model to show that inhibition may play an important role in shaping the rate-place code. We also show that midbrain auditory neurons can demonstrate similar properties as cortical harmonic template neurons. Our study fills a gap in understanding the transformation in neural representations of resolved harmonics along the auditory pathway.


Author(s):  
Joseph D Wagner ◽  
Alice Gelman ◽  
Kenneth E. Hancock ◽  
Yoojin Chung ◽  
Bertrand Delgutte

The pitch of harmonic complex tones (HCT) common in speech, music and animal vocalizations plays a key role in the perceptual organization of sound. Unraveling the neural mechanisms of pitch perception requires animal models but little is known about complex pitch perception by animals, and some species appear to use different pitch mechanisms than humans. Here, we tested rabbits' ability to discriminate the fundamental frequency (F0) of HCTs with missing fundamentals using a behavioral paradigm inspired by foraging behavior in which rabbits learned to harness a spatial gradient in F0 to find the location of a virtual target within a room for a food reward. Rabbits were initially trained to discriminate HCTs with F0s in the range 400-800 Hz and with harmonics covering a wide frequency range (800-16,000 Hz), and then tested with stimuli differing either in spectral composition to test the role of harmonic resolvability (Experiment 1), or in F0 range (Experiment 2), or both F0 and spectral content (Experiment 3). Together, these experiments show that rabbits can discriminate HCTs over a wide F0 range (200-1600 Hz) encompassing the range of conspecific vocalizations, and can use either the spectral pattern of harmonics resolved by the cochlea for higher F0s or temporal envelope cues resulting from interaction between unresolved harmonics for lower F0s. The qualitative similarity of these results to human performance supports using rabbits as an animal model for studies of pitch mechanisms providing species differences in cochlear frequency selectivity and F0 range of vocalizations are taken into account.


2000 ◽  
Vol 108 (1) ◽  
pp. 263-271 ◽  
Author(s):  
Nicolas Grimault ◽  
Christophe Micheyl ◽  
Robert P. Carlyon ◽  
Patrick Arthaud ◽  
Lionel Collet

1983 ◽  
Vol 73 (5) ◽  
pp. 1682-1685 ◽  
Author(s):  
Brian C. J. Moore ◽  
Brian R. Glasberg

2012 ◽  
Vol 292 (1-2) ◽  
pp. 26-34 ◽  
Author(s):  
Christopher J. Smalt ◽  
Ananthanarayan Krishnan ◽  
Gavin M. Bidelman ◽  
Saradha Ananthakrishnan ◽  
Jackson T. Gandour

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