This study examined the perception and cortical representation of harmonic complex tones, from the perspective of the spectral fusion evoked by such sounds. Experiment 1 tested whether ferrets spontaneously distinguish harmonic from inharmonic tones. In baseline sessions, ferrets detected a pure tone terminating a sequence of inharmonic tones. After they reached proficiency, a small fraction of the inharmonic tones were replaced with harmonic tones. Some of the animals confused the harmonic tones with the pure tones at twice the false-alarm rate. Experiment 2 sought correlates of harmonic fusion in single neurons of primary auditory cortex and anterior auditory field, by comparing responses to harmonic tones with those to inharmonic tones in the awake alert ferret. The effects of spectro-temporal filtering were accounted for by using the measured spectrotemporal receptive field to predict responses and by seeking correlates of fusion in the predictability of responses. Only 12% of units sampled distinguished harmonic tones from inharmonic tones, a small percentage that is consistent with the relatively weak ability of the ferrets to spontaneously discriminate harmonic tones from inharmonic tones in Experiment 1.
REFERENCES
1.
Alain
, C.
, Schuler
, B. M.
, and McDonald
, K. L.
(2002
). “Neural activity associated with distinguishing concurrent auditory objects
,” J. Acoust. Soc. Am.
111
, 990
–995
.2.
Bendor
, D. A.
, and Wang
, X.
(2005
). “The neuronal representation of pitch in primate auditory cortex
,” Nature (London)
435
, 341
–346
.3.
Bieser
, A.
, and Muller-Preuss
, P.
(1996
). “Auditory responsive cortex in the squirrel monkey: Neural responses to amplitude-modulated sounds
,” Exp. Brain Res.
108
, 273
–284
.4.
Bizley
, J. K.
, Nodal
, F. R.
, Nelken
, I.
, and King
, A. J.
(2005
). “Functional organization of ferret auditory cortex
,” Cereb. Cortex
15
, 1637
–1653
.5.
Bregman
, A. S.
(1990
). Auditory Scene Analysis
(MIT Press
, Cambridge, MA
).6.
Brokx
, J.
, and Nooteboom
, S.
(1980
). “Intonation and perceptual separation of simultaneous voices
,” J. Phonetics
10
, 23
–36
.7.
Brunstrom
, J. M.
, and Roberts
, B.
(1998
). “Profiling the perceptual suppression of partials in periodic complex tones: Further evidence for a harmonic template
,” J. Acoust. Soc. Am.
104
, 3511
–3519
.8.
Brunstrom
, J. M.
, and Roberts
, B.
(2000
). “Separate mechanisms govern the selection of spectral components for perceptual fusion and for the computation of global pitch
,” J. Acoust. Soc. Am.
107
, 1566
–1577
.9.
Ciocca
, V.
, and Darwin
, C.
(1993
). “Effects of onset asynchrony on pitch perception: Adaptation or grouping?
,” J. Acoust. Soc. Am.
93
, 2870
–2878
.10.
Clarey
, J.
, Barone
, P.
, and Imig
, T.
(1992
). “Physiology of thalamus and cortex
,” in The Mammalian Auditory Pathway: Neurophysiology
, edited by D.
Webster
, A.
Popper
, and R.
Fay
(Springer-Verlag
, New York
), pp. 232
–334
.11.
Darwin
, C.
, and Ciocca
, V.
(1992
). “Grouping in pitch perception: Effects of onset asynchrony and ear of presentation of a mistuned component
,” J. Acoust. Soc. Am.
91
, 3381
–3390
.12.
de Boer
, E.
(1956
). “On the ‘residue’ in hearing
,” Ph.D. thesis, University of Amsterdam
.13.
de Boer
, E.
(1976
). “On the ‘residue’ and auditory pitch perception
,” in Handbook of Sensory Physiology
, edited by W.
Keidel
and W.
Neff
(Springer
, Verlag, Berlin
), Vol. 3
, pp. 479
–583
.14.
de Boer
, E.
, and de Jongh
, H. R.
(1978
). “On cochlear encoding: Potentialities and limitations of the reverse-correlation technique
,” J. Acoust. Soc. Am.
63
, 115
–135
.15.
de Cheveigne
, A.
(2005
). “Pitch perception models
,” in Pitch: Neural Coding and Perception
, edited by C. J.
Plack
, A. J.
Oxenham
, R. R.
Fay
, and A. N.
Popper
, Springer Handbook of Auditory Research
(Springer
, New York
).16.
de Ribaupierre
, F.
, Goldstein
, M. H.
, and Yeni-Komshian
, G.
(1972
). “Cortical coding of repetitive acoustic pulses
,” Brain Res.
48
, 205
–225
.17.
Depireux
, D. A.
, Simon
, J. Z.
, Klein
, D. J.
, and Shamma
, S. A.
(2001
). “Spectro-temporal response field characterization with moving ripples in ferret primary auditory cortex
,” J. Neurophysiol.
85
, 1220
–1234
.18.
Dobbins
, H. D.
, Marvit
, P.
, Ji
, Y.
, and Depireux
, D. A.
(2007
). “Chronically recording with a multi-electrode array device in the auditory cortex of an awake ferret
,” J. Neurosci. Methods
161
, 101
–111
.19.
Duifhuis
, H.
, Willems
, L.
, and Sluyter
, R.
(1982
). “Measurement of pitch in speech: An implementation of Goldstein’s theory of pitch perception
,” J. Acoust. Soc. Am.
71
, 1568
–1580
.20.
Dyson
, B.
, and Alain
, C.
(2004
). “Representation of concurrent acoustic objects in primary auditory cortex
,” J. Acoust. Soc. Am.
115
, 280
–288
.21.
Efron
, B.
, and Tibshirani
, R.
(1993
). An Introduction to the Bootstrap
(Chapman and Hall
, New York
).22.
Elhilali
, M.
, Fritz
, J.
, Klein
, D.
, Simon
, J.
, and Shamma
, S.
(2004
). “Dynamics of precise spiking in primary auditory cortex
,” J. Neurosci.
24
, 1159
–1172
.23.
Fishman
, Y.
, Reser
, D. H.
, Arezzo
, J. C.
, and Steinschneider
, M.
(1998
). “Pitch versus spectral encoding of harmonic complex tones in primary auditory cortex of the awake monkey
,” Brain Res.
786
, 18
–30
.24.
Fishman
, Y.
, Reser
, D. H.
, Arezzo
, J. C.
, and Steinschneider
, M.
(2000
). “Complex tone processing in primary auditory cortex of the awake monkey. I. Neural ensemble correlates of roughness
,” J. Acoust. Soc. Am.
108
, 247
–262
.25.
Flanagan
, J. L.
, and Gutman
, N.
(1960
). “Pitch of periodic pulses without fundamental component
,” J. Acoust. Soc. Am.
32
, 1319
–1328
.26.
Fritz
, J.
, Shamma
, S.
, ElHilali
, M.
, and Klein
, D.
(2003
). “Rapid task-related plasticity of spectro-temporal receptive fields in primary auditory cortex
,” Nat. Neurosci.
6
, 1216
–1223
.27.
Goldstein
, J.
(1973
). “An optimum processor theory for the central formation of the pitch of complex tones
,” J. Acoust. Soc. Am.
54
, 1496
–1516
.28.
Hall
, D. A.
, and Plack
, C. J.
(2006
). “Searching for a pitch center in human auditory cortex
,” in Hearing — From Sensory Processing to Perception
, edited by B.
Kollmeier
(Springer Verlag
, Berlin Heidelberg
).29.
Hall
, D. A.
, and Plack
, C. J.
(2007
). “The human ‘pitch center’ responds differently to iterated noise and huggins pitch
,” NeuroReport
18
, 323
–330
.30.
Hartmann
, W.
, McAdams
, S.
, and Smith
, B.
(1993
). “Hearing a mistuned harmonic in an otherwise periodic complex tone
,” J. Acoust. Soc. Am.
88
, 1712
–1724
.31.
Heffner
, H.
, and Whitfield
, I.
(1976
). “Perception of the missing fundamental by cats
,” J. Acoust. Soc. Am.
59
, 915
–919
.32.
Heffner
, H. E.
, and Heffner
, R. S.
(1995
). “Conditioned avoidance for animal psychoacoustics
,” in Methods in Comparative Psychoacoustics
, edited by G.
Klump
(Birkhauser Verlag
, Basel
).33.
Hiraumi
, H.
, Nagamine
, T.
, Morita
, T.
, Naito
, Y.
, Fukuyama
, H.
, and Ito
, J.
(2005
). “Right hemispheric predominance in the segregation of mistuned partials
,” Eur. J. Neurosci.
22
, 1821
–1824
.34.
Houtsma
, A.
, and Smurzynski
, J.
(1990
). “Pitch identification and discrimination for complex tones with many harmonics
,” J. Acoust. Soc. Am.
87
, 304
–310
.35.
Kadia
, S.
, and Wang
, X.
(2003
). “Spectral integration in A1 of awake primates: Neurons with single- and multipeaked tuning characteristics
,” J. Neurophysiol.
89
, 1603
–1622
.36.
Klein
, D. J.
, Depireux
, D. A.
, Simon
, J. Z.
, and Shamma
, S. A.
(2000
). “Robust spectro-temporal reverse correlation for the auditory system: Optimizing stimulus design
,” J. Comput. Neurosci.
9
, 85
–111
.37.
Klein
, D. J.
, Simon
, J. Z.
, Depireux
, D. A.
, and Shamma
, S. A.
(2006
). “Stimulus-invariant processing and spectro-temporal reverse correlation in primary auditory cortex
,” J. Comput. Neurosci.
20
, 111
–136
.38.
Kowalski
, N.
, Depireux
, D.
, and Shamma
, S. A.
(1996
). “Analysis of dynamic spectra in ferret primary auditory cortex. I. Characteristics of single unit responses to moving ripple spectra
,” J. Neurophysiol.
76
, 3503
–3523
.39.
Kowalski
, N.
, Vernsel
, H.
, and Shamma
, S. A.
(1995
). “Comparison of responses in the anterior and primary auditory fields of the ferret cortex
,” J. Neurophysiol.
73
, 1513
–1523
.40.
Krumbholz
, K.
, Patterson
, R.
, Seither-Preisler
, A.
, Lammertmann
, C.
, and Lutkenhoner
, B.
(2003
). “Neuromagnetic evidence for a pitch processing center in heschl’s gyrus
,” Cereb. Cortex
13
, 765
–772
.41.
Langner
, G.
, Sams
, M.
, Heil
, P.
, and Schulze
, H.
(1997
). “Frequency and periodicity are represented in orthogonal maps in the human auditory cortex: Evidence from magnetoencephalography
,” J. Comp. Physiol., A
181
, 665
–676
.42.
Lewicki
, M.
(1994
). “Bayesian modeling and classification of neural signals
,” Neural Comput.
6
, 1005
–1030
.43.
Liang
, L.
, Lu
, T.
, and Wang
, X.
(2001
). “Neural representations of sinusoidal amplitude and frequency modulations in the primary auditory cortex of awake primates
,” J. Neurophysiol.
87
, 2237
–2261
.44.
Lin
, J.
, and Hartmann
, W.
(1998
). “The pitch of a mistuned harmonic: Evidence for a template model
,” J. Acoust. Soc. Am.
103
, 2608
–2617
.45.
McAlpine
, D.
(2002
). “Are pitch neurones the result of difference tones on the basilar membrane?
,” Assoc. Res. Otolaryngol. Abstr.
25
, 154
.46.
McAlpine
, D.
(2004
). “Neural sensitivity to periodicity in the inferior colliculus: Evidence for the role of cochlear distortions
,” J. Neurophysiol.
92
, 1295
–1311
.47.
McDonald
, K.
, and Alain
, C.
(2005
). “Contribution of harmonicity and location to auditory object formation in free field: Evidence from event-related brain potentials
,” J. Acoust. Soc. Am.
118
, 1593
–604
.48.
Moore
, B.
, and Glasberg
, B. R.
(1985
). “Relative dominance of individual partials in determining the pitch of complex tones
,” J. Acoust. Soc. Am.
77
, 1853
–1860
.49.
Moore
, B.
, Peters
, R.
, and Galsberg
, B.
(1986
). “Thresholds for hearing mistuned partials as separate tones in harmonic complexes
,” J. Acoust. Soc. Am.
80
, 479
–483
.50.
Moore
, D.
, and Kitzes
, L.
(1985
). “Projections from the cochlear nucleus to the inferior colluculus in normal and neonatally cochlea-ablated gerbils
,” J. Comp. Neurol.
240
, 180
–195
.51.
Patterson
, R.
, Uppenkamp
, S.
, Johnsrude
, I.
, and Griffiths
, T.
(2003
). “The processing of temporal pitch and melody information in auditory cortex
,” Neuron
36
, 767
–776
.52.
Penagos
, H.
, Melcher
, J. R.
, and Oxenham
, A. J.
(2004
). “A neural representation of pitch salience in nonprimary human auditory cortex revealed with functional magnetic resonance imaging
,” J. Neurosci.
24
, 6810
–6815
.53.
Plomp
, R.
, and Levelt
, W.
(1965
). “Tonal consonance and critical bandwidth
,” J. Acoust. Soc. Am.
38
, 548
–560
.54.
Riquimaroux
, H.
(1995
). “How to enjoy music by the brain not by the ears: Neuronal basis of pitch extraction
,” RIKEN Rev.
9
, 13
–14
.55.
Riquimaroux
, H.
, and Kosaki
, H.
(1995
). “Involvement of neurons in the primary auditory cortex in pitch extraction
,” Absr. Assoc. Res. Otolaryngol.
18
.56.
Ritsma
, R.
(1962
). “Existence region of the tonal residue i
,” J. Acoust. Soc. Am.
34
, 1224
–1229
.57.
Roberts
, B.
, and Brunstrom
, J. M.
(1998
). “Perceptual segregation and pitch shifts of mistuned components in harmonic complexes and in regular inharmonic complexes
,” J. Acoust. Soc. Am.
104
, 2326
–2338
.58.
Roberts
, B.
, and Brunstrom
, J. M.
(2001
). “Perceptual fusion and fragmentation of complex tones made inharmonic by applying different degrees of frequency shift and spectral stretch
,” J. Acoust. Soc. Am.
110
, 2479
–2490
.59.
Roberts
, B.
, and Brunstrom
, J. M.
(2003
). “Spectral pattern, harmonic relations, and the perceptual grouping of low-numbered components
,” J. Acoust. Soc. Am.
114
, 2118
–2134
.60.
Schechter
, B.
, and Depireux
, D. A.
(2007
). “Stability of spectro-temporal tuning over several sessions in primary auditory cortex of the awake ferret
,” Neuroscience
148
, 806
–14
.61.
Schreiner
, C. E.
and Calhoun
, B. M.
(1994
). “Spectral envelope coding in the cat primary auditory cortex: Properties of ripple transfer functions
,” Aud. Neurosci.
1
, 39
–61
.62.
Schulze
, H.
, Hess
, A.
, Ohl
, F.
, and Scheich
, H.
(2002
). “Superposition of horseshoe-like periodicity and linear tonotopic maps in auditory cortex of the Mongolian gerbil
,” Eur. J. Neurosci.
15
, 1077
–1084
.63.
Schulze
, H.
, and Langner
, G.
(1997a
). “Periodicity coding in the primary auditory cortex of the Mongolian gerbil (Meriones unguiculatus): Two different strategies for coding pitch and rhythm
,” J. Comp. Physiol., A
181
, 651
–663
.64.
Schulze
, H.
and Langner
, G.
(1997b
). “Representation of periodicity pitch in the primary auditory cortex of the Mongolian gerbil
,” Acta Oto-Laryngol., Suppl.
532
, 89
–95
.65.
Schwarz
, D. W.
, and Tomlinson
, R. W.
(1990
). “Spectral response patterns of auditory cortex neurons to harmonic complex tones in alert monkey (macaca mulatta)
,” J. Neurophysiol.
64
, 282
–298
.66.
Shamma
, S. A.
, Versnel
, H.
, and Kowalski
, N.
(1995
). “Ripple analysis in ferret primary auditory cortex. I. Response characteristics of single units to sinusoidally rippled spectra
,” Aud. Neurosci.
1
, 233
–254
.67.
Sinex
, D.
, Henderson-Sabes
, J.
, and Li
, H.
(2002
). “Responses of inferior colliculus neurons to harmonic and mistuned complex tones
,” Hear. Res.
168
, 150
–162
.68.
Sinex
, D.
, Li
, H.
, and Velenovsky
, D.
(2005
). “Prevalence of stereotypical responses to mistuned complex tones in the inferior colliculus
,” J. Neurophysiol.
94
, 3523
–37
.69.
Srulovicz
, P.
, and Goldstein
, J.
(1983
). “A central spectrum model: A synthesis of auditory-nerve timing and place cues in monaural communication of frequency spectrum
,” J. Acoust. Soc. Am.
73
, 1266
–1276
.70.
Steinschneider
, M.
, Reser
, D.
, Fishman
, Y.
, Schroeder
, C.
, and Arezzo
, J.
(1998
). “Click train encoding in primary auditory cortex of the awake monkey: Evidence for two mechanisms subserving pitch perception
,” J. Acoust. Soc. Am.
104
, 2935
–2955
.71.
Suga
, N.
, O’Neill
, W.
, Kujirai
, K.
, and Manabe
, T.
(1983
). “Specificity of combination-sensitive neurons for processing of complex biosonar signals in auditory cortex of the mustached bat
,” J. Neurophysiol.
49
, 1573
–1626
.72.
Terhardt
, E.
(1974
). “Pitch, consonance, and harmony
,” J. Acoust. Soc. Am.
55
, 1061
–1069
.73.
Terhardt
, E.
(1979
). “Calculating virtual pitch
,” Hear. Res.
1
, 155
–182
.74.
Tomlinson
, R. W. W.
, and Schwartz
, D. W. F.
(1988
). “Perception of the missing fundamental in nonhuman primates
,” J. Acoust. Soc. Am.
84
, 560
–565
.75.
von Helmholtz
, H.
(1954
). On the Sensations of Tone as a Physiological Basis for the Theory of Music
(Dover
, New York
).76.
Warren
, J. D.
, Uppenkamp
, S.
, Patterson
, R. D.
, and Griffiths
, T. D.
(2003
). “Separating pitch chroma and pitch height in the human brain
,” Proc. Natl. Acad. Sci. U.S.A.
100
, 10038
–10042
.77.
Whitfield
, I. C.
(1980
). “Auditory cortex and the pitch of complex tones
,” J. Acoust. Soc. Am.
67
, 644
–647
.78.
Wightman
, F. L.
(1973
). “The pattern-transformation model of pitch
,” J. Acoust. Soc. Am.
54
, 407
–416
.79.
Zatorre
, R.
(1988
). “Pitch perception of complex tones and human temporal-lobe function
,” J. Acoust. Soc. Am.
84
, 566
–572
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