Review: The Biological Basis of Audition
Recanzone and Sutter
Presented by Joseph Schilz
Outline
Introduction Organization of Audition Auditory Spatial Processing
Interactions with visual stimuli Ventriloquism effect Ventriloquism after-effect
Auditory Temporal ProcessingTemporal integrationForward maskingGap detection
Introduction
Audition and vision major sensory systems Audition does not have the significant history
of research that vision does Differences between audition and vision Tasks in decoding audition:
Where a sound came fromSpectral propertiesTemporal propertiesIdentifying what sound represents
Review's focus: location and temporal properties
Organization of Audition
Fig 1. Recanzone and Sutter
Organization of Audition
Fig 2.Recanzone & Sutter
Organization of Audition
Fig 2. Kalatsky et al.
Tonotopy
Organization of Audition
Fig 2:Bitterman et al.
Tonal Tuning
Organization of Audition
Fig 2.Bizley & King
Tonal/Spatial Tuning
Organization of Audition
Fig 1: DesignAhveninen et al.
“What” and “Where” Paths?
Organization of Audition
Fig 2: ResultsAhveninen et al.
“What” and “Where” Paths?
Auditory Spatial Processing
Cues Interactions with visual stimuli
Ventriloquism effect Ventriloquism after-effect
Auditory Spatial Processing
Fig 3. Recanzone and Sutter
Cues
Auditory Spatial Processing
Fig 1. Yu and Young
Cues
Auditory Spatial ProcessingVentriloquism effect
Definition
Cognitive factors “unity assumption”
Non-cognitive factors Timing Compellingness Spatial discrepancy/agreement
Auditory Spatial ProcessingVentriloquism effect
Early studies assumed that the more precise modality would “capture” the less precise modality.
Later studies showed a near optimal “mixing” of modality reports, respecting the measure-error variance of each modality.
Auditory Spatial ProcessingVentriloquism effect
Fig 2: Design and ResultsKording et al.
Auditory Spatial ProcessingVentriloquism after-effect
If subject is presented with audio/visual stimuli of a consistent spatial disparity, subjects spatial perception of acoustic space shifted after
Long lasting Does not transfer across frequencies Different from other adaption illusions: lasts
tens of minutes, does not transfer across frequencies, in the same direction of adapting stimulus.
Auditory Spatial ProcessingVentriloquism after-effect
Fig 4: Design and ResultsRecanzone
Auditory Spatial ProcessingVentriloquism after-effect
Difficulty of single unit recording in illusions.
Direct projection from auditory to visual (observed in primates)
Direct projection from visual to auditory (observed in ferrets)
Several areas in brain with multisensory response
Ghanzafar study
Auditory Temporal Processing
Definition Temporal integration Forward masking Gap detection
Auditory Temporal ProcessingDefinition
Can mean processing of temporal aspects of stimuli or ability of neurons to encode stimulus by temporal aspects of firing
We refer to former
Temporal processing could be interpreted to include spectral processing
We don't consider spectral processing here
Auditory Temporal ProcessingTemporal integration
Our environments are noisy; audition might wait a bit and let noise average itself out before passing on a percept
On the other hand, some decisions need to be made quickly; audition shouldn't hold onto information for too long
How to assess? At what levels is this occurring?
Auditory Temporal ProcessingTemporal integration
Fig 4. Dallos and Olsen
Auditory Temporal ProcessingTemporal integration
Varies with loudness, frequency, duration
Bloch's Law (Loudness x Duration = k) Leaky integrator model
Mean integration times of 30-40msec in humans
Auditory Temporal Processing
Clock et al. found similar constants of integration in chinchilla cochlear nucleus neurons and chinchilla behavior
Exponential leaky integrator fit model well
But, auditory nerve neurons had time constants much larger
Explanation
Temporal integration
Auditory Temporal ProcessingForward masking
Two sounds presented sequentially, with some gap, sometimes subject will not perceive second sound.
Depends on many factors Generally measured as a
function of first sound's duration.
Fig 4. Recanzone and Sutter
Auditory Temporal ProcessingForward masking
Likely a result of adaptation.
Auditory Temporal ProcessingGap detection
Temporal resolution vs. temporal integration One paradigm for temporal resolution: gap
detection
Humans able to detect gaps in noise as small as 1-2msec
Auditory nerve firing shows gaping pattern Likely some role of cortex in detecting gaps,
as shown by lesion, deactivation studies
Auditory Temporal ProcessingGap detection
Fig 2. Zhang et al.
Additional Works ReferencedAhveninen et al. Task-modulated “what” and “where” pathways in human auditory
cortex PNAS 2006 103 (39) 14608-14613; published ahead of print September 18, 2006, doi:10.1073/pnas.0510480103
Y. Bitterman, R. Mukamel, R. Malach, I. Fried, & I. Nelken Ultra-fine frequency
tuning revealed in single neurons of human auditory cortex Nature 451, 197-201 (10 January 2008)
Jennifer K. Bizley, Andrew J. King, Visual-auditory spatial processing in auditory cortical neurons, Brain Research, Volume 1242, Multisensory Integration, 25 November 2008, Pages 24-36, ISSN 0006-8993, DOI: 10.1016/j.brainres.2008.02.087.
P. Dallos, W. Olsen, Integration of energy at threshold with gradual rise-fall tone pips. Journal of the Acoustical Soc. of America.Vol. 36, pp. 743-751, April 1964
V Kalatsky, D Polley, M Merzenich, C Schreiner, Mstryker, Fine functional organization of auditory cortex revealed by Fourier optical imaging PNAS 2005 102 (37) 13325-13330; published ahead of print September 1, 2005, doi:10.1073/pnas.0505592102
Additional Works Referenced
Kording KP, Beierholm U, Ma WJ, Quartz S, Tenenbaum JB, et al (2007) Causal Inference in Multisensory Perception. PLoS ONE 2(9): e943.
doi:10.1371/journal.pone.0000943
Recanzone G, Rapidly induced auditory plasticity: The ventriloquism aftereffect. Proc. Natl. Acad. Sci. USA Vol. 95, pp. 869–875, February 1998
J Yu, E Young, Linear and nonlinear pathways of spectral information transmission in the cochlear nucleus PNAS 2000 97 (22) 11780-11786
W. Zhang, R.J. Salvi, S.S. Saunders, Neural correlates of gap detection in auditory nerve fibers of the chinchilla, Hearing Research, Volume 46, Issue 3, July 1990, Pages 181-200, ISSN 0378-5955, DOI: 10.1016/0378-5955(90)90001-6.
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