Abstract
There is ample experimental evidence describing changes of tonotopic organisation in the auditory cortex due to environmental factors. In order to uncover the underlying mechanisms, we designed a large-scale computational model of the auditory cortex. The model has up to 100 000 Izhikevich’s spiking neurons of 17 different types, almost 21 million synapses, which are evolved according to Spike-Timing-Dependent Plasticity (STDP) and have an architecture akin to existing observations. Validation of the model revealed alternating synchronised/desynchronised states and different modes of oscillatory activity. We provide insight into these phenomena via analysing the activity of neuronal subtypes and testing different causal interventions into the simulation. Our model is able to produce experimental predictions on a cell type basis. To study the influence of environmental factors on the tonotopy, different types of auditory stimulations during the evolution of the network were modelled and compared. We found that strong white noise resulted in completely disrupted tonotopy, which is consistent with in vivo experimental observations. Stimulation with pure tones or spontaneous activity led to a similar degree of tonotopy as in the initial state of the network. Interestingly, weak white noise led to a substantial increase in tonotopy. As the STDP was the only mechanism of plasticity in our model, our results suggest that STDP is a sufficient condition for the emergence and disruption of tonotopy under various types of stimuli. The presented large-scale model of the auditory cortex and the core simulator, SUSNOIMAC, have been made publicly available.









Similar content being viewed by others
Notes
The scalingFactor is equal to 20 and corresponds to scaled density of synapses in comparison to the real cortex and is adopted from (Izhikevich and Edelman 2008).
Licenced under Creative Commons Attribution-NonCommercial-ShareAlike 4.0 International Public Licence. The code is at https://sites.google.com/site/susnoimac, last accessed: July 6, 2015.
Processor: Intel(R) Core(TM) i7 930 @ 2.80 GHz, RAM: 16 GB, OS: Windows 7.
The minute-long window contains sixty values of firing rate, one for each second in the window. The mean and standard deviation are computed from these sixty values.
We focused only on desynchronised states and nearly desynchronised states in this analysis in order to have the largest oscillations in the gamma band.
Input neurons (i.e., neurons from layer L4) are not accounted because they are trivially tonotopic and would skew the results by enforcing that every network has a large quantity of perfectly tonotopic neurons.
This means intensity corresponding to the probability 0.5, i.e., each neuron in the input layer has, in each time step, a probability 0.5 to be externally stimulated, as defined in Section 2.2.
Note that computation of a single 12 h development of a 50k network, along with the measurement of tonotopy, takes circa 10 days; we therefore could not compute more repetitions with different seeds.
The reason for using Kruskal-Wallis ANOVA (instead of standard ANOVA) is the fact that it does not require normally distributed data.
References
Arieli, A, Shoham, D, Hildesheim, R, & Grinvald, A (1995). Coherent spatiotemporal patterns of ongoing activity revealed by real-time optical imaging coupled with single-unit recording in the cat visual cortex. Journal of Neurophysiology, 73(5), 2072–2093.
Bandyopadhyay, S, Shamma, SA, & Kanold, PO (2010). Dichotomy of functional organization in the mouse auditory cortex. Nature Neuroscience, 13(3), 361–368.
Barbour, B, Brunel, N, Hakim, V, & Nadal, JP (2007). What can we learn from synaptic weight distributions? Trends in Neuroscience, 30(12), 622–629. doi:10.1016/j.tins.2007.09.005. http://www.ncbi.nlm.nih.gov/pubmed/17983670.
Barbour, DL, & Callaway, EM (2008). Excitatory local connections of superficial neurons in rat auditory cortex. The Journal of Neuroscience, 28(44), 11,174–11,185.
Bartos, M, Vida, I, Frotscher, M, Meyer, A, Monyer, H, Geiger, JRP, & Jonas, P (2002). Fast synaptic inhibition promotes synchronized gamma oscillations in hippocampal interneuron networks. Proceedings of the National Academy of Sciences of the United States of America, 99(20), 13,222–13,227.
Beierlein, M, Gibson, JR, & Connors, BW (2003). Two dynamically distinct inhibitory networks in layer 4 of the neocortex. Journal of Neurophysiology, 90(5), 2987–3000.
Binzegger, T, Douglas, RJ, & Martin, KAC (2004). A quantitative map of the circuit of cat primary visual cortex. The Journal of Neuroscience, 24(39), 8441–8453.
Bollimunta, A, Mo, J, Schroeder, CE, & Ding, M (2011). Neuronal mechanisms and attentional modulation of corticothalamic α oscillations. The Journal of Neuroscience, 24(39), 8441–8453.
Buonomano, DV, & Maass, W (2009). State-dependent computations: spatiotemporal processing in cortical networks. Nature Reviews Neuroscience, 10(2), 113–125.
Buzsáki, G, & Wang, XJ (2012). Mechanisms of gamma oscillations. Annual review of neuroscience, 35, 203–225.
Cardin, JA, Carlén, M, Meletis, K, Knoblich, U, Zhang, F, Deisseroth, K, Tsai, LH, & Moore, CI (2009). Driving fast-spiking cells induces gamma rhythm and controls sensory responses. Nature, 459 (7247), 663–667. doi:10.1038/nature08002. http://www.ncbi.nlm.nih.gov/pubmed/19396156.
Chang, EF, & Merzenich, MM (2003). Environmental noise retards auditory cortical development. Science, 300(5618), 498–502.
Christophe, E, Roebuck, A, Staiger, JF, Lavery, DJ, Charpak, S, & Audinat, E (2002). Two types of nicotinic receptors mediate an excitation of neocortical layer I interneurons. Journal of Neurophysiology, 88(3), 1318–1327. http://www.ncbi.nlm.nih.gov/pubmed/12205153.
Chrostowski, M, Yang, L, Wilson, HR, Bruce, IC, & Becker, S (2011). Can homeostatic plasticity in deafferented primary auditory cortex lead to travelling waves of excitation? Journal of Computational Neuroscience, 30(2), 279–299.
Clement, EA, Richard, A, Thwaites, M, Ailon, J, Peters, S, & Dickson, CT (2008). Cyclic and sleep-like spontaneous alternations of brain state under urethane anaesthesia. PLoS One, 3(4), e2004.
Cottam, JC, Smith, SL, & Häusser, M (2013). Target-specific effects of somatostatin-expressing interneurons on neocortical visual processing. The Journal of Neuroscience, 33(50), 19567–19578.
Cruikshank, SJ, Rose, HJ, & Metherate, R (2002). Auditory thalamocortical synaptic transmission in vitro. Journal of Neurophysiology, 87(1), 361–384.
de Pinho, M, & Roque-da Silva, AC (1999). A realistic computational model of formation and variability of tonotopic maps in the auditory cortex. Neurocomputing, 26, 355–359.
de Pinho, M, Mazza, M, & Roque, AC (2006). A computational model of the primary auditory cortex exhibiting plasticity in the frequency representation. Neurocomputing, 70(1), 3–8.
de la Rocha, J, Marchetti, C, Schiff, M, & Reyes, AD (2008). Linking the response properties of cells in auditory cortex with network architecture: Cotuning versus lateral inhibition. The Journal of Neuroscience, 28(37), 9151–9163.
de Villers-Sidani, E, Chang, EF, Bao, S, & Merzenich, MM (2007). Critical period window for spectral tuning defined in the primary auditory cortex (A1) in the rat. The Journal of Neuroscience, 27(1), 180–189.
DeFelipe, J, Alonso-Nanclares, L, & Arellano, JI (2002). Microstructure of the neocortex: comparative aspects. Journal of Neurocytology, 31(3–5), 299–316.
Feldmeyer, D, Egger, V, Lubke, J, & Sakmann, B (1999). Reliable synaptic connections between pairs of excitatory layer 4 neurones within a single barrel of developing rat somatosensory cortex. Journal of Physiology, 521 Pt 1, 169–190. http://www.ncbi.nlm.nih.gov/pubmed/10562343.
Froemke, RC, Merzenich, MM, & Schreiner, CE (2007). A synaptic memory trace for cortical receptive field plasticity. Nature, 450(7168), 425–429.
Froemke, RC, & Jones, BJ (2011). Development of auditory cortical synaptic receptive fields. Neuroscience & Biobehavioral Reviews, 35(10), 2105–2113.
Galarreta, M, & Hestrin, S (1999). A network of fast-spiking cells in the neocortex connected by electrical synapses. Nature, 402(6757), 72–75.
Gasparini, S, Migliore, M, & Magee, JC (2004). On the initiation and propagation of dendritic spikes in Ca1 pyramidal neurons. The Journal of Neuroscience, 24(49), 11,046–11,056.
Grossman, RL, Greenway, M, Heath, AP, Powell, R, Suarez, R, Wells, W, White, KP, Atkinson, M, Klampanos, I, Alvarez, H, Harvey, C, & Mambretti, J. (2012). The design of a community science cloud: The open science data cloud perspective. https://www.opensciencedatacloud.org/.
Happel, MF, Jeschke, M, & Ohl, FW (2010). Spectral integration in primary auditory cortex attributable to temporally precise convergence of thalamocortical and intracortical input. The Journal of Neuroscience, 30(33), 11,114–11,127.
Haeusler, S, Schuch, K, & Maass, W (2009). Motif distribution, dynamical properties, and computational performance of two data-based cortical microcircuit templates. Journal of Physiology Paris, 103(1-2), 73–87.
Harris, KD, & Shepherd, GM (2015). The neocortical circuit: themes and variations. Nature neuroscience, 18(2), 170–181.
Hestrin, S, & Armstrong, WE (1996). Morphology and physiology of cortical neurons in layer I. The Journal of Neuroscience, 16(17), 5290–5300.
Hodgkin, AL, & Huxley, AF (1952). A quantitative description of membrane current and its application to conduction and excitation in nerve. The Journal of Physiology, 117(4), 500.
Holmgren, C, Harkany, T, Svennenfors, B, & Zilberter, Y (2003). Pyramidal cell communication within local networks in layer 2/3 of rat neocortex. Journal of Physiology, 551(Pt 1), 139–53. doi:10.1113/jphysiol.2003.044784. http://www.ncbi.nlm.nih.gov/pubmed/12813147.
Huang, CL, & Winer, JA (2000). Auditory thalamocortical projections in the cat: laminar and areal patterns of input. Journal of Comparative Neurology, 427(2), 302–331.
Huxter, J, Burgess, N, & O’Keefe, J (2003). Independent rate and temporal coding in hippocampal pyramidal cells. Nature, 425(6960), 828–832.
Insanally, MN, Albanna, BF, & Bao, S (2010). Pulsed noise experience disrupts complex sound representations. Journal of Neurophysiology, 103(5), 2611.
Issa, JB, Haeffele, BD, Agarwal, A, Bergles, DE, Young, ED, & Yue, DT (2014). Multiscale optical Ca 2+ imaging of tonal organization in mouse auditory cortex. Neuron, 83(4), 944– 959.
Iurilli, G, Ghezzi, D, Olcese, U, Lassi, G, Nazzaro, C, Tonini, R, Tucci, V, Benfenati, F, & Medini, P (2012). Sound-driven synaptic inhibition in primary visual cortex. Neuron, 73(4), 814–828.
Izhikevich, EM (2003). Simple model of spiking neurons. IEEE Transactions on Neural Networks, 14(6), 1569–1572.
Izhikevich, EM (2004). Which model to use for cortical spiking neurons? IEEE Transactions on Neural Networks, 15(5), 1063–1070.
Izhikevich, EM (2006). Polychronization: Computation with spikes. Neural Computation, 18(2), 245–282.
Izhikevich, EM (2007). Solving the distal reward problem through linkage of STDP and dopamine signaling. Cerebral Cortex, 17(10), 2443–2452.
Izhikevich, EM, & Edelman, GM (2008). Large-scale model of mammalian thalamocortical systems. Proceedings of the National Academy of Sciences, 105(9), 3593–3598.
Kawaguchi, Y (1995). Physiological subgroups of nonpyramidal cells with specific morphological characteristics in layer II/III of rat frontal cortex. The Journal of Neuroscience, 15(4), 2638–2655.
Kawaguchi, Y, & Kubota, Y (1997). GABAergic cell subtypes and their synaptic connections in rat frontal cortex. Cerebral Cortex, 7(6), 476–486.
Kim, H, & Bao, S (2009). Selective increase in representations of sounds repeated at an ethological rate. The Journal of Neuroscience, 29(16), 5163–5169.
Kimura, A, Donishi, T, Sakoda, T, Hazama, M, & Tamai, Y (2003). Auditory thalamic nuclei projections to the temporal cortex in the rat. Neuroscience, 117(4), 1003–1016.
Knight, BW (1972). Dynamics of encoding in a population of neurons. The Journal of General Physiology, 59 (6), 734–766.
Kotak, VC, Fujisawa, S, Lee, FA, Karthikeyan, O, Aoki, C, & Sanes, DH (2005). Hearing loss raises excitability in the auditory cortex. The Journal of Neuroscience, 25(15), 3908–3918.
Kral, A, Tillein, J, Heid, S, Klinke, R, & Hartmann, R (2006). Cochlear implants: cortical plasticity in congenital deprivation. Progress in Brain Research, 157, 283–313.
Lapicque, L (1907). Recherches quantitatives sur l’excitation électrique des nerfs traitée comme une polarisation. Journal de Physiologie et de Pathologie Générale, 9(1), 620–635.
Larson, E, Billimoria, CP, & Sen, K (2009). A biologically plausible computational model for auditory object recognition. Journal of Neurophysiology, 101(1), 323–331.
Larson, E, Perrone, BP, Sen, K, & Billimoria, CP (2010). A robust and biologically plausible spike pattern recognition network. The Journal of Neuroscience, 30(46), 15,566–15,572.
Lee, CC, & Winer, JA (2008). Connections of cat auditory cortex: I. Thalamocortical system. Journal of Comparative Neurology, 507(6), 1879–1900.
Letzkus, JJ, Wolff, SBE, Meyer, EMM, Tovote, P, Courtin, J, Herry, C, & Luethi, A (2011). A disinhibitory microcircuit for associative fear learning in the auditory cortex. Nature, 480(7377), 331–335.
Levy, RB, & Reyes, AD (2012). Spatial profile of excitatory and inhibitory synaptic connectivity in mouse primary auditory cortex. Journal of Neuroscience, 32(16), 5609–5619.
Li, LY, Ji, XY, Liang, FX, Li, YT, Xiao, ZJ, Tao, HZW, & Zhang, LI (2014a). A feedforward inhibitory circuit mediates lateral refinement of sensory representation in upper layer 2/3 of mouse primary auditory cortex. Journal of Neuroscience, 34(41), 13670–13683.
Li, LY, Xiong, XR, Ibrahim, LA, Yan, W, Tao, H, & Zhang, LI (2014b). Differential receptive field properties of parvalbumin and somatostatin inhibitory neurons in mouse auditory cortex. Cerebral Cortex. Epub ahead of print.
Liebe, S, Hoerzer, GM, Logothetis, NK, & Rainer, G (2012). Theta coupling between V4 and prefrontal cortex predicts visual short-term memory performance. Nature Neuroscience, 15(3), 456–462.
Markram, H, Lübke, J, Frotscher, M, & Sakmann, B (1997). Regulation of synaptic efficacy by coincidence of postsynaptic APs and EPSPs. Science, 275(5297), 213–215.
Markram, H, Toledo-Rodriguez, M, Wang, Y, Gupta, A, Silberberg, G, & Wu, C (2004). Interneurons of the neocortical inhibitory system. Nature Reviews Neuroscience, 5(10), 793–807.
Matsumoto, M, & Nishimura, T (1998). Mersenne twister: A 623-dimensionally equidistributed uniform pseudo-random number generator. ACM Transactions on Modeling and Computer Simulation (TOMACS), 8(1), 3–30.
Moore, AK, & Wehr, M (2013). Parvalbumin-expressing inhibitory interneurons in auditory cortex are well-tuned for frequency. The Journal of Neuroscience, 33(34), 13,713–13,723.
Muresan, RC, & Savin, C (2007). Resonance or integration? Self-sustained dynamics and excitability of neural microcircuits. Journal of Neurophysiology, 97(3), 1911–1930.
Nahmani, M, & Turrigiano, GG (2014). Deprivation-induced strengthening of presynaptic and postsynaptic inhibitory transmission in layer 4 of visual cortex during the critical period. The Journal of Neuroscience, 34(7), 2571–2582.
Nelken, I (2014). Stimulus-specific adaptation and deviance detection in the auditory system: Experiments and models. Biological Cybernetics, 1–9. doi:10.1007/s00422-014-0585-7.
Nordlie, E, Gewaltig, MO, & Plesser, HE (2009). Towards reproducible descriptions of neuronal network models. PLoS Computational Biology, 5(8), 456.
Nunez, PL, & Srinivasan, R. (2006). Electric fields of the brain: The neurophysics of EEG. USA: Oxford University Press.
O’Keefe, J, & Recce, ML (1993). Phase relationship between hippocampal place units and the EEG theta rhythm. Hippocampus, 3(3), 317–330.
Oswald, AMM, Schiff, ML, & Reyes, AD (2006). Synaptic mechanisms underlying auditory processing. Current Opinion in Neurobiology, 16(4), 371–376.
Ouda, L, Druga, R, & Syka, J (2011). Distribution of SMI-32-immunoreactive neurons in the central auditory system of the rat. Brain Structure and Function, 217(1), 19–36.
Phoka, E, Wildie, M, Schultz, SR, & Barahona, M (2012). Sensory experience modifies spontaneous state dynamics in a large-scale barrel cortical model. Journal of Computational Neuroscience, 33(2), 323–339.
Popelová, M. (2013). Software tool for modelling coding and processing of information in auditory cortex of mice. Master Thesis, Charles University in Prague, Faculty of Mathematics and Physics. http://www.marketa.najevisti.info/dokumenty/master_thesis.pdf.
Raghavachari, S, Lisman, JE, Tully, M, Madsen, JR, Bromfield, EB, & Kahana, MJ (2006). Theta oscillations in human cortex during a working-memory task: Evidence for local generators. Journal of Neurophysiololgy, 95(3), 1630–1638. doi:10.1152/jn.00409.2005. http://www.ncbi.nlm.nih.gov/pubmed/16207788.
Reale, RA, Breugge, JF, & Chan, JC (1987). Maps of auditory cortex in cats reared after unilateral cochlear ablation in the neonatal period. Brain Research, 431(2), 281–290.
Richardson, RJ, Blundon, JA, Bayazitov, IT, & Zakharenko, SS (2009). Connectivity patterns revealed by mapping of active inputs on dendrites of thalamorecipient neurons in the auditory cortex. Journal of Neuroscience, 29(20), 6406–17. doi:10.1523/JNEUROSCI.0258-09.2009 10.1523/JNEUROSCI.0258-09.2009. http://www.ncbi.nlm.nih.gov/pubmed/19458212.
Romanski, LM, & LeDoux, JE (1993). Organization of rodent auditory cortex: anterograde transport of PHA-L from MGv to temporal neocortex. Cerebral Cortex, 3(6), 499–514.
Rothschild, G, Nelken, I, & Mizrahi, A (2010). Functional organization and population dynamics in the mouse primary auditory cortex. Nature Neuroscience, 13(3), 353–360.
Sakata, S, & Harris, KD (2009). Laminar structure of spontaneous and sensory-evoked population activity in auditory cortex. Neuron, 64(3), 404–418.
Sakata, S, & Harris, KD (2012). Laminar-dependent effects of cortical state on auditory cortical spontaneous activity. Frontiers in Neural Circuits, 6(109). doi:10.3389/fncir.2012.00109.
Schreiner, CE, Read, HL, & Sutter, ML (2000). Modular organization of frequency integration in primary auditory cortex. Annual Review of Neuroscience, 23(1), 501–529.
Schutter, ED. (2009). Computational modeling methods for neuroscientists. MIT Press.
Smith, P.H., & Populin, L.C. (2001). Fundamental differences between the thalamocortical recipient layers of the cat auditory and visual cortices. Journal of Comparative Neurology, 436(4), 508–519.
Smith, PH, Uhlrich, DJ, Manning, KA, & Banks, MI (2012). Thalamocortical projections to rat auditory cortex from the ventral and dorsal divisions of the medial geniculate nucleus. Journal of Comparative Neurology, 520 (1), 34–51. doi:10.1002/cne.22682. http://www.ncbi.nlm.nih.gov/pubmed/21618239.
Song, S, Miller, KD, & Abbott, LF (2000). Competitive Hebbian learning through spike-timing-dependent synaptic plasticity. Nature Neuroscience, 3(9), 919–926. doi:10.1038/78829. http://www.ncbi.nlm.nih.gov/pubmed/10966623.
Song, S, Sjöström, PJ, Reigl, M, Nelson, S, & Chklovskii, DB (2005). Highly nonrandom features of synaptic connectivity in local cortical circuits. PLoS Biology, 3(3), e68.
Stanton, SG, & Harrison, RV (2000). Projections from the medial geniculate body to primary auditory cortex in neonatally deafened cats. Journal of Comparative Neurology, 426, 117–129.
Steriade, M, McCormick, DA, & Sejnowski, TJ (1993). Thalamocortical oscillations in the sleeping and aroused brain. Science, 262(5134), 679–685.
Stiebler, I, Neulist, R, Fichtel, I, & Ehret, G (1997). The auditory cortex of the house mouse: left-right differences, tonotopic organization and quantitative analysis of frequency representation. Journal of Comparative Physiology A, 181(6), 559–571.
Sun, YJ, Wu, GK, Liu, Bh, Li, P, Zhou, M, Xiao, Z, Tao, HW, & Zhang, LI (2010). Fine-tuning of pre-balanced excitation and inhibition during auditory cortical development. Nature, 465(7300), 927–931.
Syka, J, Šuta, D, & Popelář, J (2005). Responses to species-specific vocalizations in the auditory cortex of awake and anesthetized guinea pigs. Hearing research, 206(1), 177–184.
Šuta, D, Kvašňák, E, Popelář, J, & Syka, J (2003). Representation of species-specific vocalizations in the inferior colliculus of the guinea pig. Journal of neurophysiology, 90(6), 3794–3808.
Šuta, D, Popelář, J, Kvašňák, E, & Syka, J (2007). Representation of species-specific vocalizations in the medial geniculate body of the guinea pig. Experimental brain research, 183(3), 377–388.
Šuta, D, Popelář, J, Burianová, J, & Syka, J (2013). Cortical representation of species-specific vocalizations in Guinea pig. PloS one, 8(6), e65432.
Timofeev, I, Grenier, F, Bazhenov, M, Sejnowski, TJ, & Steriade, M (2000). Origin of slow cortical oscillations in deafferented cortical slabs. Cerebral Cortex, 10(12), 1185–1199.
Traub, RD, Spruston, N, Soltesz, I, Konnerth, A, Whittington, MA, & Jefferys, GR (1998). Gamma-frequency oscillations: a neuronal population phenomenon, regulated by synaptic and intrinsic cellular processes, and inducing synaptic plasticity. Progress in Neurobiology, 55(6), 563–575.
Wagatsuma, N, Potjans, TC, Diesmann, M, & Fukai, T (2011). Layer-Dependent Attentional Processing by Top-down Signals in a Visual Cortical Microcircuit Model. Frontiers in Computational Neuroscience, 5(31), 1–15.
Watson, C. (2012). The mouse nervous system. Academic Press.
Wendykier, P (2013). Parallel Colt. https://sites.google.com/site/piotrwendykier/software/parallelcolt.
Wendykier, P, & Nagy, JG (2010). Parallel colt: A high-performance Java library for scientific computing and image processing. ACM Transactions on Mathematical Software (TOMS), 37(3), 31.
Wilson, NR, Runyan, CA, Wang, FL, & Sur, M (2012). Division and subtraction by distinct cortical inhibitory networks in vivo. Nature, 488(7411), 343–348.
Winer, JA (2006). Decoding the auditory corticofugal systems. Hearing Research, 212(1), 1–8.
Winer, JA, & Lee, CC (2007). The distributed auditory cortex. Hearing Research, 229(1-2), 3.
Wu, GK, Arbuckle, R, Liu, Bh, Tao, HW, & Zhang, LI (2008). Lateral sharpening of cortical frequency tuning by approximately balanced inhibition. Neuron, 58(1), 132–143.
Wu, GK, Tao, HW, & Zhang, LI (2011). From elementary synaptic circuits to information processing in primary auditory cortex. Neuroscience & Biobehavioral Reviews, 35(10), 2094– 2104.
Xu, S, Jiang, W, Poo, M, & Dan, Y (2012). Activity recall in a visual cortical ensemble. Nature Neuroscience, 15(3), 449–455.
Xu, H, Jeong, HY, Tremblay, R, & Rudy, B (2013). Neocortical somatostatin-expressing GABAergic interneurons disinhibit the thalamorecipient layer 4. Neuron, 77(1), 155–167.
Yuste, R, & Denk, W (1995). Dendritic spines as basic functional units of neuronal integration. Nature, 375 (6533), 682–684.
Zhang, LI, Bao, S, & Merzenich, MM (2001). Persistent and specific influences of early acoustic environments on primary auditory cortex. Nature Neuroscience, 4(11), 1123–1130.
Zhang, LI, Bao, S, & Merzenich, MM (2002). Disruption of primary auditory cortex by synchronous auditory inputs during a critical period. Proceedings of the National Academy of Sciences, 99(4), 2309–2314.
Zhou, X, Nagarajan, N, Mossop, BJ, & Merzenich, MM (2008). Influences of un-modulated acoustic inputs on functional maturation and critical-period plasticity of the primary auditory cortex. Neuroscience, 154(1), 390–396.
Zhou, Y, Mesik, L, Sun, YJ, Liang, F, Xiao, Z, Tao, HW, & Zhang, LI (2012). Generation of spike latency tuning by thalamocortical circuits in auditory cortex. The Journal of Neuroscience, 32(29), 9969–9980.
Zingg, B, Hintiryan, H, Gou, L, Song, MY, Bay, M, Bienkowski, M S, Foster, NN, Yamashita, S, Bowman, I, & Toga, AW (2014). Neural networks of the mouse neocortex. Cell, 156(5), 1096–1111.
Zucker, RS (1989). Short-term synaptic plasticity. Annual Review of Neuroscience, 12(1), 13–31.
Acknowledgments
We would like to thank Elisa Brann, Martin Popel and Felix Zhou for help with polishing the manuscript and The Bakala Foundation for financial support of the two first authors during their studies. This work was supported by the Grant Agency of the Czech Republic (P303/12/1347), by PRVOUK “P46 – Informatika”, and by Engineering and Physical Sciences Research Council (EP/F500394/1). This work made use of the Open Science Data Cloud (OSDC) which is an Open Cloud Consortium (OCC)-sponsored project (Grossman et al. 2012); which is a work supported in part by grants from Gordon and Betty Moore Foundation and the National Science Foundation and major contributions from OCC members like the University of Chicago.
Conflict of interest
The authors declare that they have no conflict of interest.
Author information
Authors and Affiliations
Corresponding authors
Additional information
Action Editor: Erik De Schutter
Electronic supplementary material
Below is the link to the electronic supplementary material.
Rights and permissions
About this article
Cite this article
Tomková, M., Tomek, J., Novák, O. et al. Formation and disruption of tonotopy in a large-scale model of the auditory cortex. J Comput Neurosci 39, 131–153 (2015). https://doi.org/10.1007/s10827-015-0568-2
Received:
Revised:
Accepted:
Published:
Issue Date:
DOI: https://doi.org/10.1007/s10827-015-0568-2