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Auditory Neuroscience 1 Spatial Hearing. Systems Biology Doctoral Training Program Physiology course Prof. Jan Schnupp Hearing: an impossible task!. ITD. Interaural Time Difference (ITD) Cues.

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Auditory Neuroscience 1

Spatial Hearing

Systems Biology Doctoral Training Program

Physiology course

Prof. Jan Schnupp

interaural time difference itd cues


Interaural Time Difference (ITD) Cues

ITDs are powerful cues to sound source direction, but they are ambiguous (“cones of confusion”)

front back ambiguity and phase ambiguity
Front-Back Ambiguity and Phase Ambiguity

interaural level cues ilds
Interaural Level Cues (ILDs)

Unlike ITDs, ILDs are highly frequency dependent. At higher sound frequencies ILDs tend to become larger, more complex, and hence potentially more informative.

ILD at 700 Hz

ILD at 11000 Hz

adapting to changes in spectral cues
Adapting to Changes in Spectral Cues

Hofman et al. made human volunteers localize sounds in the dark, then introduced plastic molds to change the shape of the concha. This disrupted spectral cues and led to poor localization, particularly in elevation.

Over a prolonged period of wearing the molds, (up to 3 weeks) localization accuracy improved.


Phase locking improves in the cochlear nucleus


Endbulbof Held

Auditory nervefiber


The Jeffress model: mapping ITDs in the brain?


McAlpine and colleagues

ITD tuning varies with sound frequency: no map?


The Auditory Pathway

CN, cochlear nuclei;

SOC, superior olivary complex; NLL, nuclei of the lateral lemniscus; IC, inferior colliculus; MGB, medial geniculate body.

lesion studies suggest important role for a1
Lesion Studies Suggest Important Role for A1

Jenkins & Merzenich, J. Neurophysiol, 1984


Linear Prediction of Responses


“w matrix”

Input“i vector”

r(t) = i1(t-1) w1(1) + i1(t-2) w1(2)+ ...+ i2(t-1) w2(1) + i2(t-2) w2(2)+ ...

+ i3(t-1) w3(1) + i2(t-2) w3(2)+ ...

Frequency [kHz]



Predicting Space from Spectrum


Left and Right Ear Frequency-Time Response Fields

Virtual Acoustic Space Stimuli


Frequency [kHz]

Elev [deg]





Schnupp et al Nature 2001

higher order cortical areas
“Higher Order” Cortical Areas
  • In the macaque, primary auditory cortex(A1) is surrounded by rostral (R), lateral (L), caudo-medial (CM) and medial “belt areas”.
  • L can be further subdivided into anterior, medial and caudal subfields (AL, ML, CL)
are there what and where streams in auditory cortex
Are there “What” and “Where” Streams in Auditory Cortex?


  • Some reports suggest that anterior cortical belt areas may more selective for sound identity and less for sound source location, while caudal belt areas are more location specific.
  • It has been hypothesized that these may be the starting positions for a ventral “what” stream heading for inferotemporal cortex and a dorsal “where” stream which heads for postero-parietal cortex.


a panoramic code for auditory space
A “Panoramic” Code for Auditory Space?
  • Middlebrooks et al.found neural spike patterns to vary systematically with sound source direction in a number cortical areas of the cat (AES, A1, A2, PAF).
  • Artificial neural networks can be trained to estimate sound source azimuth from the neural spike pattern.
  • Spike trains in PAF carry more spatial information than other areas, but in principle spatial information is available in all auditory cortical areas tested so far.
artificial vowel sounds
Artificial Vowel Sounds
  • Bizley et al J Neurosci 2009 29:2064
responses to artificial vowels in space

Vowel type (timbre)

Pitch (Hz)

Responses to Artificial Vowels in Space
  • Bizley et al J Neurosci 2009 29:2064
azimuth pitch and timbre sensitivity in ferret auditory cortex
Azimuth, Pitch and Timbre Sensitivity in Ferret Auditory Cortex
  • Bizley et al J Neurosci 2009 29:2064