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Computational Modeling of the Auditory Periphery: From Soundfiles to Spiketrains Marcos Cantu Center for Computational Neuroscience and Neural Technology (CompNet Graduate Program for Neuroscience (GPN) Boston University [email protected] MCN 2013

Computational Modeling of the Auditory Periphery: From Soundfiles to Spiketrains Marcos Cantu Center for Computational Neuroscience and Neural Technology

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Computational Modeling of the Auditory Periphery:From Soundfiles to Spiketrains

Marcos CantuCenter for Computational Neuroscience and Neural Technology (CompNet)Graduate Program for Neuroscience (GPN)Boston [email protected]

MCN 2013Figure 11. Circuit model of the DCN that includes all the effects described above. Shaded neurons are the primarily auditory neurons described in Fig. 6. The granule cell and cartwheel cell of the superficial layer and a GABAergic neuron of unknown identity have been added. The relative size of synaptic terminals corresponds roughly to the relative synaptic strength needed to account for the effects described above. (Redrawn from Davis and Young 2000 with permission.)1Project Aims:- Create Large Scale Spiking Network with several Cell Types- Model Sensory Transduction at the Auditory Periphery- Use Natural Ethologically Relevant Stimuli (i.e. Vocalizations)I want to see how the circuit reacts to natural sound. 2Basilar Membrane model(Bank of Bandpass Filters)Meddis Hair Cell ModelAuditory Nerve FiberSpiketrainsAcoustic StimulusAcoustic StimulusBasilar Membrane model(Bank of Bandpass Filters)Auditory Nerve FiberSpiketrainsAuditory Toolboxby Malcolm SlaneyPatterson-HoldsworthERB Filter Bank Ray Meddis, 1986 and 1990Auditory Toolboxby Malcolm SlaneyMeddis Hair Cell ModelEugene Izhikevich Spiking Neuron (2003)

Meddis Hair Cell Model Gammatone Filterbank Spiking Izhikevich Neurons .WAV audiofileFrom Soundfiles to Spiketrains

Getting TO the cochlear nucleus.

Before I get to the cochlear nucleus, I should talk about how I got to the cochlear nucleus. That is, how I created the input vector for each timestep of the network.

From sound to spiketrain5

Sound to spiketrain Meddis Hair Cell Model(actually Sound to Probability)

q = free transmitter pool c = transmitter at synaptic cleft w = reprocessed transmitter The higher the stimulus, the higher the permeability (k)output:prob(event) = hc(t) dt

Inner hair cellAuditory nerve fiberK is driven by filter respone at frequency6Spiking neurons were modeled with the system of ordinary differential equations developed by Eugene M. Izhikevich : v = 0.04v2 + 5v + 140 - u + I u = a(bv - u) with the auxiliary after-spike resetting: if v >= 30 mV then v c and u u + dv corresponds to the membrane potential of the neuron u represents a membrane recovery variable

a, b, c and d are dimensionless parameters

a time scale of recovery, b the sensitivity of recovery, c the after-spike reset value of the membrane potential d the after-spike reset of the recovery variable. Izhikevich, EM: Simple model of spiking neurons. IEEE Transactions on Neural Networks 2003, 14(6):1569-1572 (2003)Auditory Nerve Fibers (ANFs)K is driven by filter respone at frequency7

Meddis Hair Cell Model Gammatone Filterbank Spiking Izhikevich Neurons .WAV audiofileFrom Soundfiles to Spiketrains

Getting TO the cochlear nucleus.

Before I get to the cochlear nucleus, I should talk about how I got to the cochlear nucleus. That is, how I created the input vector for each timestep of the network.

From sound to spiketrain8Spiking Neuron Model of Dorsal Cochlear Nucleus (DCN): 4 cell types2 excitatory: Auditory Nerve Fiber (Input to DCN)Type IV cell (Output Cell of DCN)2 inhibitory: Type II cell (Low to High Frequency Inhibition)WBI cell (Wide Band Inhibition)the responses to sound of DCN neurons display substantial inhibitory influencesthe responses to sound of DCN neurons display substantial inhibitory influences9

ouYoung and Davis 2001Wiring Diagram of Principal Cell Types in the Dorsal Cochlear Nucleus:All three cell types receive feedforward input from frequency tuned Auditory Nerve Fibers (ANFs)This is the circuitry in functional space, not in physical space.

Output CellFigure 11. Circuit model of the DCN that includes all the effects described above. Shaded neurons are the primarily auditory neurons described in Fig. 6. The granule cell and cartwheel cell of the superficial layer and a GABAergic neuron of unknown identity have been added. The relative size of synaptic terminals corresponds roughly to the relative synaptic strength needed to account for the effects described above. (Redrawn from Davis and Young 2000 with permission.)

This is the circuitry in functional space, not in physical space. 10

Figure 11. Circuit model of the DCN that includes all the effects described above. Shaded neurons are the primarily auditory neurons described in Fig. 6. The granule cell and cartwheel cell of the superficial layer and a GABAergic neuron of unknown identity have been added. The relative size of synaptic terminals corresponds roughly to the relative synaptic strength needed to account for the effects described above. (Redrawn from Davis and Young 2000 with permission.)11

Recovering the Missing Fundamental with a Feedforward NetworkSpiking Network (1000 frequency tuned neurons in each layer)12

Spiking Network (1000 frequency tuned neurons in each layer)Recovering the Missing Fundamental with a Feedforward Network13

Projection to 1st Population1st 2nd2nd 3rd3rd 4thFeed-Forward Connectivity Matrices14

Spike Timing Dependent Plasticity (STDP) Connectivity Matrix (previous work)15

1st 2nd2nd 3rd3rd 4thFeed-Forward Connectivity Matrices

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Fundamental Frequency (500Hz) Absent in Input Spectrum Larger Spiking Network (4000 frequency tuned neurons in each layer)17

Fundamental Frequency (500Hz) Present in Input Spectrum Larger Spiking Network (4000 frequency tuned neurons in each layer)18Future Directions:Adding Various Forms of Inhibition to NetworkAdding Plasticity and Training the Network on Natural SoundAdding Top-Down Recurrent Connectivity19