Auditory Forebrain
Auditory Thalamus Auditory Cortex
Auditory Projections to Polysensory Cortex
Plasticity
Parallel Processing Pathways in the Subcortical System
Tonotopic/core: ICC MGv A1
(core) Non-tonotopic/diffuse:
ICX MGm/MGd Belt/parabelt).
Auditory Cortex is Comprised of Multiple Fields
Pro
TS1
TS2
Tpt
sts
ls
TS3
AI12vl
8a46dr
10
8b
46r
ps
946vr
asd
asv
cs
Ins45
TS3
Cat
Human
Bat
Macaque
Human
Topographic Organization
Primary auditory cortical areas (“core” areas) respond best to tones, and are tonotopically organized.
Borders between fields marked by reversal in tonotopic gradient.
“Belt” fields less responsive to tones, respond best to NB noise, yet share tonotopy of adjacent core fields.
Adjacent fields interconnect more extensively than non-adjacent fields.
Macaque
Hackett, Stepniewska, Kaas (1998) JCN 394: 475-495
Functional Organization of Isofrequency Slabs in A1: Evidence from Anaesthetized Cat
As in the ICC, receptive field properties are systematically organized in “hypercolumns” within iso-CF slabs: Spectral selectivity (“integration”; i.e., tuning curve
width, Q) Threshold Latency FM sweep direction, sweep rate selectivity Binaural Interaction
Functional Organization of Isofrequency Slabs in A1: Evidence from Anaesthetized Cat
Spectral selectivity or “integration” (tuning width, Q)
NB = Narrow bandBB = Broad band
Q40dB
FrequencyHigh (rostral)
Read et al. (2002)
Low (caudal)
Functional Organization of Isofrequency Slabs in Cat A1
… FM sweep direction preference
and rate selectivity
Up
Down
Fast
Slow
Up Fast
Mendelson et al. (1993)
Functional Organization of Isofrequency Slabs in Cat A1
…Binaural Interactions: Represented in elongated bands in cat AC
Bands with similar binaural properties are interconnected across fields ipsilaterally, and contralaterally.
EE: Binaural Summation (ITD)EI: Binaural Suppression (IID)
Space Map in Auditory Cortex?
Spatial selectivity is relatively poor.
No evidence for systematic map of space.
Alternate theory: Location encoded by populations of neurons with modest spatial selectivity.
Functional Organization of Primate A1
In awake behaving monkeys, evidence for internal organization is not as strong as that in anaesthetized cats (methodological?).
Primate rostral field neurons have longest latencies, and narrowest frequency and intensity tuning
A1 has shortest latencies, moderate tuning. Caudomedial field has broadest frequency tuning Lateral field neurons have monotonic R/L
functions.
Spectral Domain Properties
Classically-defined receptive fields resemble those at thalamic and IC levels.
More “multi-peaked” response areas are found.
Recanzone et al. (2000)
Spectrotemporal Response Areas
Response areas are the result of interactions between excitation and inhibition.
However, the time course of excitation and inhibition may vary.
STRA’s try to capture the time-dependency of E – I interactions to reveal dynamic spectral filtering.
“Classical” Frequency Response Area
Spectrotemporal Response Areas
Reverse correlation technique: Find which stimulus feature correlated most strongly with the response.
STRAs can be used to design “optimal stimuli”. DeCharms et al. (1998)
Spectrotemporal Response Areas
“Edge detection” (I.e., response to low pass or high pass noise).
“Orientation and direction specificity”: response to FM sweeps with particular modulation direction and speed.
“Optimal” stimuli generate much higher response rates than tonal stimuli.
Spectrotemporal Response Areas
“Edge detection” (i.e., response to low pass or high pass noise).
“Orientation and direction specificity”: response to FM sweeps with particular modulation direction and speed.
“Optimal” stimuli generate much higher response rates than tonal stimuli.
Spectro-temporal Facilitation (Combination Sensitivity)
Combination sensitivity may underlie rapid discrimination of vowels.
Kent & Read (1991)
Combination Sensitivity in Primates
66% of macaque A1 neurons are enhanced by presentation of tone combinations of different frequencies.
Brosch et al. (1999)
Responses to Complex Sounds and Vocalizations
Many neurons show specificity for sets of complex sounds, e.g., certain vocalizations, even specific combinations of utterances.
But there’s little evidence for specificity to a unique sound (e.g., Grandma’s name cell).
Klug et al. (2002)
Processing Signals in Noise
In auditory nerve fibers, background noise raises tonal thresholds, shifts rate level functions to the right, compresses dynamic range.
In A1, same stimuli generate increase in threshold, shift to right, but without compression.
Outputs of AC
A1 projects subcortically to ICdc, MGB, pontine gray.
AC belt and parabelt regions project to Superior temporal gyrus and sulcus Ventral prefrontal (cognition; saccade initiation) insular (multimodal: limbic, hippocampus)
Caudal AC belt regions project to LIP, lateral interparietal area (spatial; projects to
dorsal premotor cortex.
Projections to Prefrontal Cortex
Belt and Parabelt AC project to rostral STS/STG.
…and ventral prefrontal cortex.
Auditory responses prevalent in ventrolateral PFC (areas 12, 45): corresponds to Broca’s area in humans.
Rostro-caudal gradient of projections
Pro
TS1
TS2
Tpt
sts
ls
TS3
AI12vl
8a46dr
10
8b
46r
ps
946vr
asd
asv
cs
Ins45
TS3
Romanski et al. (1999)
Prefrontal Connections
Hackett and Kaas
Cortical Plasticity
Functional organization is maintained by experience.
E.g., Representation of frequency is massively altered by pairing stimulation with cholinergic nucleus basalis of the forebrain.
9 kHz (± 1/3 octave)
Kilgard and Merzenich (1998)
Before After
250 ms tones paired with nucleus basalis stimulation.
Cortical Plasticity
Experience-dependent plasticity develops over time…
…and dramatically increases with temporal complexity.
Conclusion: auditory cortical organization is strongly influenced by behavioral significance of acoustic stimulus.
Train of 15 ms tones paired with nucleus basalis stimulation.
Kilgard and Merzenich (1998)
Control
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