Denham Fens 2006

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A computational model of the differential suppression of AI transient

responses to acoustic sequences


S Denham (sdenham@plymouth.ac.uk)
Centre for Theoretical and Computational Neuroscience, University of Plymouth
Abstract

In recent studies of population responses in auditory cortex of awake monkey


to alternating tone sequences it was found that non best frequency tones
were suppressed by alternating best frequency tones, and the suppression
increased with increasing frequency difference and increasing presentation
rate [1, 2].
Since the characteristics of the temporal and frequency relationships closely
paralleled those found in two-tone streaming experiments [3], it was claimed
that the observed neural behaviour was an analogue of primitive auditory
streaming.
Furthermore, since the timing and frequency distribution of suppression
seemed to be consistent with forward masking studies [4] it was suggested
that there may be a common mechanism underlying both phenomena.
The present computational modelling study investigated the hypothesis that
the depression of thalamocortical synapses could be that mechanism.
Using a model of synaptic depression which we previously showed could
account for many of the temporal responses properties in primary auditory
cortex [5], we found that the responses to two tone streaming stimuli could
similarly be modelled.

Sustained activation at 20Hz reveals a progressive depression of unitary


EPSCs [10].

AB

Forward suppression in the model


Model
A
20 Hz

B
40 Hz

60 Hz

Cortex

10 Hz

Model

10 Hz

Although capable of very precisely timed responses to stimulus onsets,


auditory cortex tends not respond to rapid envelop fluctuations, or rapidly
repeating stimuli with temporally precise phase-locked activity [6].
Neither thalamic response properties, nor inhibition in cortex appear to
account for this forward suppression [7].

AB

5. Build up of streaming

Cortex

Auditory Streaming Paradigm

Thalamus

Change in perceptual organisation

20 Hz

Excitatory (green) and inhibitory (red)


conductances evoked by pairs of clicks
40 Hz

4. Dependence on tone duration


Monkey [4]

Monkey [1]

Forward Suppression

1. Differences between same frequency and


alternating frequency responses

Dependence on F and T

Model

Frequency
ratio B/A

Frequency

Thalamus

2. Response ratios
Monkey [1]

Segregated

Horse

A
60 Hz

1.5

Morse

From [6]

Time
From [7]

Slow

loss

z (t ) = I (t ). f [g , y ( t ) ]
dx
= z (t ) .x (t ) .x (t )
dt
dy
= .w (t ) z (t ) + .[1 y ( t )]
dt
dw
= . x ( t ) .w ( t )
dt
de
= [ . x ( t ) e (t ) ] / EPSP
dt
f [ g , y ( t )] = ( p event > r ). g . y (t ).(1 + rn )
p event = (1 g ) y ( t )

Paired pulse ratio A2/A1 versus probability of the first EPSC [9].

Presentation Rate Fast

From these simulations it appears that synaptic depression at thalamocortical


synapses may provide a relatively simple account for both forward
suppression [7] and the differential suppression of non best frequency tones
by best frequency tones in alternating tone sequences [1].
In addition, the model response replicate the dependence of differential
suppression on frequency difference, presentation rate, and tone duration [2];
and the build up of streaming [11].
However, the recent finding that auditory streaming is bi-stable, and has very
similar characteristics to bi-stabilities in visual perception [14], imply that
auditory streaming is a more complex process.
We suggest that the stimulus-driven stage of processing modelled here
results in the context-dependent segregation of activity belonging to
potentially independent sound sources, on the basis of which further stages
of perceptual organisation, in which predictive models compete to arrive at a
coherent interpretation of the world, operate.

The dynamical properties of cortical synapses can influence the temporal


sensitivity of cortical circuitry [8]. Synaptic responses are context dependent,
and may develop depression or facilitation, depending on the cells involved.
Synaptic model parameters were adjusted so as to replicate EPSC rise and
fall times and paired pulse ratios for low and high probability synapses [9],
and typical PP stochasticity and long term drift [10].
replenishment

Integrated

Build up of streaming [11]

Synaptic Depression

Discussion

Ambiguous

3. Dependence on F and T

Monkey [2]

Neural correlates of auditory streaming

Responses to alternating tone sequences ABABAB measured in primary


auditory cortex of awake monkey [1,2,11].
A is at the best frequency of the recording site.
Model consists of:

Peripheral processing: using the nonlinear DRNL model [12]

Central processing: within-channel transient enhancement [13], spike


train generation

Neural array (AI) with depressing synapses, convergent/divergent


connectivity

References
1.
2.

3.
4.
5.

Model

6.
7.
8.
9.

10.
11.
12.
13.
14.

Acknowledgements: Supported by European Research Area Specific Targeted Project, EmCAP (IST-FP6-013123).

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