Graeme Clark Collection

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    Calculation of interspike intervals for integrate-and-fire neurons with Poisson distribution of synaptic inputs
    Burkitt, A. N. ; Clark, Graeme M. ( 2000)
    We present a new technique for calculating the interspike intervals of integrate-and-fire neurons. There are two new components to this technique. First, the probability density of the summed potential is calculated by integrating over the distribution of arrival times of the afferent postsynaptic potentials (PSPs), rather than using conventional stochastic differential equation techniques. A general formulation of this technique is given in terms of the probability distribution of the inputs and the time course of the postsynaptic response. The expressions are evaluated in the gaussian approximation, which gives results that become more accurate for large numbers of small-amplitude PSPs. Second, the probability density of output spikes, which are generated when the potential reaches threshold, is given in terms of an integral involving a conditional probability density. A.N. Burkitt and G.M. Clark, 'Calculation of Interspike Intervals for Integrate and Fire Neurons with Poisson Distribution of Synaptic Inputs ', Neural Computation, 12:8 (August, 2000), pp. 1789-1820. © 2000 by the Massachusetts Institute of Technology. http://www.mitpressjournals.org.ezp.lib.unimelb.edu.au/toc/neco/12/8
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    Analysis of synchronization in response of neurons to noisy periodic synaptic input
    Burkitt, A. N. ; Clark, Graeme M. ( 2000)
    Unavailable due to copyright.
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    Ventral cochlear nucleus coding of voice onset time in naturally spoken syllables
    Clarey, Janine C. ; Paolini, Antonio, G. ; Grayden, David B. ; Burkitt, Anthony, N. ; Clark, Graeme M. ( 2004)
    Unavailable due to copyright.
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    Summation of spatiotemporal input patterns in leaky integrate-and-fire neurons: Application to neurons in the cochlear nucleus receiving converging auditory nerve fiber input
    Kuhlmann, L ; Burkitt, AN ; Paolini, A ; Clark, GM (SPRINGER, 2002)
    The response of leaky integrate-and-fire neurons is analyzed for periodic inputs whose phases vary with their spatial location. The model gives the relationship between the spatial summation distance and the degree of phase locking of the output spikes (i.e., locking to the periodic stochastic inputs, measured by the synchronization index). The synaptic inputs are modeled as an inhomogeneous Poisson process, and the analysis is carried out in the Gaussian approximation. The model has been applied to globular bushy cells of the cochlear nucleus, which receive converging inputs from auditory nerve fibers that originate at neighboring sites in the cochlea. The model elucidates the roles played by spatial summation and coincidence detection, showing how synchronization decreases with an increase in both frequency and spatial spread of inputs. It also shows under what conditions an enhancement of synchronization of the output relative to the input takes place.
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    Temporal processing from the auditory nerve to the medial nucleus of the trapezoid body in the rat
    Paolini, AG ; FitzGerald, JV ; Burkitt, AN ; Clark, GM (ELSEVIER SCIENCE BV, 2001-09)
    This investigation examines temporal processing through successive sites in the rat auditory pathway: auditory nerve (AN), anteroventral cochlear nucleus (AVCN) and the medial nucleus of the trapezoid body (MNTB). The degree of phase-locking, measured as vector strength, varied with intensity relative to the cell's threshold, and saturated at a value that depended upon stimulus frequency. A typical pattern showed decline in the saturated vector strength from approximately 0.8 at 400 Hz to about 0.3 at 2000 Hz, with similar profiles in units with a range of characteristic frequencies (480-32,000 Hz). A new expression for temporal dispersion indicates that this variation corresponds to a limiting degree of temporal imprecision, which is relatively consistent between different cells. From AN to AVCN, an increase in vector strength was seen for frequencies below 1000 Hz. At higher frequencies, a decrease in vector strength was observed. From AVCN to MNTB a tendency for temporal coding to be improved below 800 Hz and degraded further above 1500 Hz was seen. This change in temporal processing ability could be attributed to units classified as primary-like with notch (PL(N)). PL(N) MNTB units showed a similar vector strength distribution to PL(N) AVCN units. Our results suggest that AVCN PL(N) units, representing globular bushy cells, are specialised for enhancing the temporal code at low frequencies and relaying this information to principal cells of the MNTB.
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    Delay analysis in the auditory brainstem of the rat: comparison with the click latency
    FitzGerald, John V. ; Burkitt, Anthony N. ; Clark, Graeme M. ; Paolini, Antonio G. ( 2001)
    Unavailable due to copyright.
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    Synchronization of the neural response to noisy periodic synaptic input
    Burkitt, AN ; Clark, GM (M I T PRESS, 2001-12)
    The timing information contained in the response of a neuron to noisy periodic synaptic input is analyzed for the leaky integrate-and-fire neural model. We address the question of the relationship between the timing of the synaptic inputs and the output spikes. This requires an analysis of the interspike interval distribution of the output spikes, which is obtained in the gaussian approximation. The conditional output spike density in response to noisy periodic input is evaluated as a function of the initial phase of the inputs. This enables the phase transition matrix to be calculated, which relates the phase at which the output spike is generated to the initial phase of the inputs. The interspike interval histogram and the period histogram for the neural response to ongoing periodic input are then evaluated by using the leading eigenvector of this phase transition matrix. The synchronization index of the output spikes is found to increase sharply as the inputs become synchronized. This enhancement of synchronization is most pronounced for large numbers of inputs and lower frequencies of modulation and also for rates of input near the critical input rate. However, the mutual information between the input phase of the stimulus and the timing of output spikes is found to decrease at low input rates as the number of inputs increases. The results show close agreement with those obtained from numerical simulations for large numbers of inputs.
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    Peak-Splitting in the Response of the Leaky Integrate-and-Fire Neuron Model to Low-Frequency Periodic Inputs
    KUHLMANN, L ; BURKITT, AN ; CLARK, GM (Monash University Press, 2001)
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    The relationship between the output synchrony of cochlear nucleus neurons and the site of stimulation in the cochlea
    Kuhlmann, L. ; Burkitt, A. N. ; Paolini, A. G. ; Clark, Graeme M. ( 2001)
    A model has been developed to determine the relationship between the output synchrony of cochlear nucleus neurons and the site of stimulation in the cochlea. This is an Integrate and Fire Neuron Model in which noisy periodic synaptic inputs to the neuron are summed and a spike is generated when the membrane potential reaches threshold. The model describes the stochastic input that auditory nerve fibres provide to a cochlear nucleus neuron and the corresponding stochastic output. To investigate the relationship between the output synchrony of cochlear nucleus neurons (namely globular bushy cells) and the site of stimulation in the cochlea, phase differences between the periodic inputs of the model were incorporated, in order to mimic how the travelling wave consecutively activates auditory nerve fibres originating over a spatial spread of the basilar membrane. Analysis of the model found that output synchrony decreased with an increase in frequency and spatial spread. Furthermore, enhancement of the output synchrony relative to the input synchrony occurred for small spatial spreads of the basilar membrane over which input primary afferent fibres originate. Adding noise helped to make the model more realistic. As a result enhancement of synchrony occurred with a spatial spread of less than 1.25 mm and 0.75 mm for 0.5 kHz and I kHz respectively, while for the higher frequencies analysed (2 kHz and 5 kHz) enhancement of synchrony did not occur. This research has implications for the design of electrode arrays in cochlear implants. The number and geometry of the electrodes and the stimulus patterns to be used will depend on the degree of convergence of fibres and how phase information is processed by neurons in the brainstem.
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    Effects of sensorineural hearing loss on the refractory properties of auditory nerve fibers
    Roberts, L. A. ; Shepherd, R. K. ; Paolini, A. G. ; Clark, Graeme M. ; Burkitt, A. N. ( 2000)
    We hypothesised that the loss of the peripheral processes and the partial demyelination of auditory nerve fibres (ANFs) following a sensorineural hearing loss would increase their refractory properties. Normal control, and long-term (2.5 months) systemically deafened rats were anaesthetised (urethane, 1.3 g/kg i.p.), a bipolar stimulating electrode was implanted into the scala tympani and glass microelectrodes (30-80 MΩ) used to record single ANF activity. Stimuli (pairs of 100 µs/phase charge balanced biphasic pulses with interpulse intervals (IPIs) of 0.34-10 ms) were presented at 6 dB above threshold using a repetition interval of 250 ms. Absolute refractory period (ARP) was defined as the IPI at which the probability of eliciting a spike to the second stimulus was 0.1. In the present results, based on recordings from 62 fibres, ANFs were distinguished from cochlear nucleus (CN) neurones by their significantly shorter median latencies (AN: 0.575ms vs CN: 1.137ms; Whitney-Mann Rank Sum, p<0.0001). There were no significant differences between minimum ANF latencies from normal and deafened animals. Although the median ARP was greater in deafened versus normal animals, this difference was not statistically significant (normals: median0.658ms, interquartile range 0.554-0.913ms; deafened: 0.772ms and 0.616-1.073ms; p=0.16). Finally, the spike latency associated with the second pulse of a pair systematically increased with decreasing IPI, contrasting with the stable latency of the response to the leading pulse. Although pathological changes to ANFs may increase their refractory properties, at this duration of deafness these changes were not significant.