Plasticity in sensory systems
Jan Schnupp on the monocycle
Activity and size of auditory cortex… Schneider et al. Nat. Neurosci. 2003
…Are correlated…
…and correlated with musical abilities
Is musical practice increasing the size of auditory cortex, or do people with large auditory cortex become musicians?
What do we learn when we learn a new skill?
Nat. Neurosci. 2006
Human psychoacoustical performance Frequency differences Pressure ratio between softest and loudest sounds… Hair motion at absolute threshold…
Learning protocol
Perceptual learning Partially non-specific –Playing tetris improves frequency discrimination Partially due to passive exposure But also to some extent requires active task performance
Animal models of auditory plasticity Classical conditioning –Fear conditioning: associating a sound with a foot shock Environmental enrichment and relatives –Manipulating the environment can have both beneficial and disruptive effects on the auditory system Spatial hearing
Nat. Rev. Neurosci. 2004
Fear conditioning…
…changes cortical neurons
Brain Research 2007
Environmental enrichment…
Plasticity in auditory enriched environments
Auditory plasticity requires stimuli but not interaction
Just noticeable differences in azimuth at the center, tone stimuli
time amplitude Interaural Time Differences (ITDs) Interaural Level Differences (ILDs) Binaural Cues for Localising Sounds in Space
Interaural Time Difference (ITD) Cues ITD ITDs are powerful cues to sound source direction, but they are ambiguous (“cones of confusion”)
Binaural disparities in humans ITD ILD
Disambiguating the cone of confusion Sounds on the median plane (azimuth 0, different elevations) have zero binaural disparities This is a special case of the cone of confusion Nevertheless, humans and other animals can determine the elevation of a sound source
Spectral information about space
The barn owl…
Binaural Cues in the Barn Owl Barn owls have highly asymmetric outer ears, with one ear pointing up, the other down. Consequently, at high frequencies, barn owl ILDs vary with elevation, rather than with azimuth (D). Consequently ITD and ILD cues together form a grid specifying azimuth and elevation respectively.
Phase locking at high frequencies in the barn owl C. Köppl, 1997
Processing of Interaural Time Differences Interaural time difference MSO neuron response Sound on the ipsilateral side Contra- lateral side Medial superior olive To the Inferior Colliculus
Preservation of Time Cues in AVCN Auditory Nerve Fibers connect to spherical and globular bushy cells in the antero- ventral cochlear nucleus (AVCN) via large, fast and secure synapses known as “endbulbs of Held”. Phase locking in bushy cells is even more precise than in the afferent nerve fibers. Bushy cells project to the superior olivary complex. Auditory Nerve Fibers connect to spherical and globular bushy cells in the antero- ventral cochlear nucleus (AVCN) via large, fast and secure synapses known as “endbulbs of Held”. Phase locking in bushy cells is even more precise than in the afferent nerve fibers. Bushy cells project to the superior olivary complex. spherical bushy cell endbulb of Held VIII nerve fiber
The coincidence detection model of Jeffress (1948) is the widely accepted model for low-frequency sound localisation
Response Interaural Time Difference 0
Response Interaural Time Difference 0
Interaural Phase Sensitivity in the MSO to 1000 Hz Yin and Chan (1988) 1 ms
Processing of Interaural Level Differences Interaural intensity difference LSO neuron response Sound on the ipsilateral side Contralateral side C > II > C Lateral superior olive To the Inferior Colliculus
The Calyx of Held MNTB relay neurons receive their input via very large calyx of Held synapses. These secure synapses would not be needed if the MNTB only fed into “ILD pathway” in the LSO. MNTB also provides precisely timed inhibition to MSO.
Caird and Klinke 1983 Frequency (kHz) Sound level (dB SPL) IpsilateralContralateral
The Superior Olivary Nuclei – a Summary Most neurons in the MSO respond best to sounds that occur earlier in the contralateral ear. Most neurons in the LSO respond best to sounds that are louder in the ipsilateral ear. Space representation is crossed, and therefore LSO projects mostly contralaterally and MSO ipsilaterally. MNTB MSO LSO CN Midline Inhibitory Connection Excitatory Connection IC
Spatial hearing is plastic
Plasticity in adults Nat. Neurosci. 1998
New ears…
Sound localization by humans
Sound localization by humans
Effect of modifying the ear
Learning the new ears
Knowing both ears
Plasticity of the space map Knudsen, Nature 2002
Orientation responses to auditory and visual stimuli are congruent… Auditory orientation response Visual orientation response
Prisms that shift the visual scene
Auditory responses adapt to the visual shift
The brain of the barn owl
The ICC, ICX and the Superior Collicullus (Optic Tectum)
Point-to-point correspondence between ICX and OT
Neural correlate of the shift of auditory responses
Shift in ITD sensitivity occurs first in ICX
Axonal sprouting cause shift of ITD sensitivity in ICX
Time course of ITD shift
Cellular mechanisms of ITD shift
Anatomy of the instructive signal
Visual activity in ICX uncovered by removing inhibition in OT
Cellular mechanisms of ITD shift
NMDA receptors are present at the transition stage…
…but not when the shift is complete
Cellular mechanisms of ITD shift
GABA participates in the suppression of the normal responses Control Bicuculline
Plasticity and age
Old animals cannot change
A sensitive period…
During the sensitive period, plasticity potential is very large
The normal map is robust and can be recovered at any age
Recovery of the normal map requires rich environment
Adult plasticity is possible after juvenile experience
Time course of adult adjustment