// Reinoud Maex 20 June 2007 // The AMPA and GABAA channels are copied from De Schutter & Bower // Purkinje cell model, modified by Sergio Solinas et al. (EJN 2006). // The NMDA channel is copied from the Maex & De Schutter (J Neurophysiol 1998) // granule cell model. // Hence this script merges the files Purk_syn37dC.g and Gran_synchan.g //genesis - Purkinje cell M9 genesis2.1 script /* Copyright E. De Schutter (Caltech and BBF-UIA) */ /********************************************************************** ** Sets of synapse objects developed for rat cerebellum Purkinje ** E. De Schutter, Caltech, 1991-1992 **********************************************************************/ /* Reference: ** E. De Schutter and J.M. Bower: An active membrane model of the ** cerebellar Purkinje cell: II. Simulation of synaptic responses. ** Journal of Neurophysiology 71: 401-419 (1994). ** http://bbf-www.uia.ac.be/TNB/TNB_pub7.html ** Consult this reference for sources of experimental data. */ include ../L5P37C/L5P_const+axon+syn.g // CONSTANTS /* should be defined by calling routine (all correctly scaled): ** E_non_NMDA ** E_GABA, G_GABA */ // factor Q10 of 1.5: Regehr et al. J. Neuroscience 1996 16(18):5661-5671 float Q10 = 1.5 function Calc_tau(Q10,T1,T2,tau1) return {tau1 / { Q10**{{T2 -T1}/10}}} end float temp = 37 // Celsius degrees float Q12_non_NMDA = 1.68 // for 10 1.4 // correctio for low input resistance float Rin_corr = 1 /********************************************************************* ** The synaptic conductance equations *********************************************************************/ function make_GABAA_channels /* GABA channel, made by SS */ /* Reference: current clamp data from ** Pouzat C. and Hestrin S. J. Neuroscince 1997 ** V_drive 60 mV, I_syn 20 pA -> G_peak = 333 pS ** (that is 35.3 pA at 37 C which should be recorded in voltage clamp) ** T_rise: 2.6 +- 0.5 ms, Thalf-width = 16.7 +- 2.7 ms ** room temperature: we assume 23 Celsius degrees ** Found T_on = 1.65 ms and T_off = 9.3 ms by hand (MATLAB) ** Found with c++ program /bbf/milaan/sergio/Work/C++/Rise2Tau/rise2tau: ** Found T_on = 2.1 ms and T_off = 15.3 ms by hand (c++) */ float PC_GABAA_factor = 5.8 // Stell & Stell1 channels: average conpartment surface 4.020627647e-11 m2 float PC_GABAA_gmax = {333e-12 / 4.020627647e-11 * PC_GABAA_factor} // Stell3 channels: average conpartment surface 2.247033843e-10 m2 float PC_GABAAm_gmax = {333e-12 / 2.49670427e-10 * PC_GABAA_factor} // Stell4 channels: somatic surface m2 float PC_GABAAs_gmax = {3.5e-9 / 2.789857497e-09 } float PC_GABAA_t_on = 1.65e-3 float PC_GABAA_t_off = 9.3e-3 /* Synaptic channel for Stellate connections */ if (!({exists GABA})) create synchan GABA end setfield GABA Ek {E_GABA} \ tau1 {Calc_tau {Q10} 23 {temp} {PC_GABAA_t_on}} \ tau2 {Calc_tau {Q10} 23 {temp} {PC_GABAA_t_off}} \ gmax {G_GABA} frequency 0.0 /* Synaptic channel for Basket connections on PC main dendrite */ if (!({exists GABA2})) create synchan GABA2 end setfield GABA2 Ek {E_GABA} \ tau1 {Calc_tau {Q10} 23 {temp} {PC_GABAA_t_on}} \ tau2 {Calc_tau {Q10} 23 {temp} {PC_GABAA_t_off}} \ gmax {G_GABA} frequency 0.0 /* Synaptic channel for Basket connections PC soma */ if (!({exists GABA3})) create synchan GABA3 end // SS We use 37 degrees here since it's not specified in the article // RM This does not make sense because then the somatic GABA is slower than the dendritic; // changed to 37 to 23 setfield GABA3 Ek {E_GABA} \ tau1 {Calc_tau {Q10} 23 {temp} {PC_GABAA_t_on}} \ tau2 {Calc_tau {Q10} 23 {temp} {PC_GABAA_t_off}} \ gmax {G_GABA} frequency 0.0 // set the right GABAA density value to get gmax from Hausser M. and Clark B. A. Neuron 1997 G_GABA = {{PC_GABAA_gmax} * {Q10**{{{temp} - 23}/10}}} echo GABAA gmax {G_GABA} G_GABAm = {{PC_GABAAm_gmax} * {Q10**{{{temp} - 23}/10}}} echo GABAA2 gmax {GB_GABA} G_GABAs = {{PC_GABAAs_gmax} * {Q10**{{{temp} - 37}/10}}} echo GABAAs soma gmax {GB_GABAs} end // make_GABAA_channels function make_AMPA_channels /* non-NMDA channel, made by SS */ /* Reference: Barbour B. 2002 (personal communication) ** room temp: 32 C ** somatic EPSC peak = 8.4 +- 7.1 pA ** (that is 10.2 pA at 37 C which should be recorded in voltage clamp) ** the driving force at the excitatory synapse is 70 mV ** during Voltage clamp at -70 mV ** thus the peak conductance G_par_syn = 120 pS ** t_on = 1.0 +- 0.7 ms ** the low-pass filtering effect of the large PC dendritic tree ** could slow down the fast rise EPSC fase. Since the effect of single vescicle release ** is kown to have an almost instantaneous effect on PSC we use a faster t_on ** t_on = 0.7 ms ** t_off = 11.1 +- 5.7 ms ** t_off is long and it light be due to the glutammate spillover ** activating the extrasynaptic AMPA receptors ** We use t_off = 1.2 ms */ /* Modified by SS 30/04/2002 ** We need to be able to set the AMPA receptor strength ** indipendently on previous settings ** since when the AMPA receptor is placed on the spine head ** it's scaled by its surface here we devide the gmax by the surface */ float dia = 0.54e-6 float surf = dia*dia*{PI} //- here we can keep control of the Gmax since the readcell // will add the spines (*rand_spines) without modify them float PC_AMPA_factor = 9 float PC_AMPA_gmax = {120e-12/surf * PC_AMPA_factor} float PC_AMPA_t_on = 0.7e-3 float PC_AMPA_t_off = 1.2e-3 /* asynchronously firing channel */ if (!({exists AMPA})) create synchan AMPA end setfield AMPA Ek {E_AMPA} \ tau1 {Calc_tau {Q10} 32 {temp} {PC_AMPA_t_on}} \ tau2 {Calc_tau {Q10} 32 {temp} {PC_AMPA_t_off}} \ frequency 0.0 G_par_syn = {{PC_AMPA_gmax} * {Q10**{{{temp} - 32}/10}} } echo AMPA gmax = {G_par_syn} setfield AMPA gmax {G_par_syn} /* synchronously firing channel removed, was for climbing fibre synapses on PC */ end // make_AMPA_channels function make_NMDA_channels /* NMDA channel made by CP */ /* From Gabbiani et al.(model) 1994, based on Jahr and Stevens */ float Q10_synapse = 3.0 //[Mg] in mM float CMg = 1.2 // per mM float eta = 0.2801 // per V float gamma = 62 float offset = - 0.01 echo eta = {eta} eta = eta * {exp {- gamma * offset}} echo new eta = {eta} if (!({exists NMDA})) create synchan2 NMDA end setfield NMDA Ek {E_NMDA} tau2 {3e-3 / Q10_synapse} \ tau1 {40e-3 / Q10_synapse} \ gmax {G_NMDA} // use the following value for synaptic activation when TEST.g is run // gmax {4.0 * G_NMDA} if (! {exists NMDA/Mg_BLOCK}) create Mg_block NMDA/Mg_BLOCK end setfield NMDA/Mg_BLOCK CMg {CMg} \ KMg_A {1/eta} \ \\ *({exp {EREST_ACT*gamma}})} \ KMg_B {1.0/gamma} end // make_NMDA_channels function make_GABAB_channels /* GABA_B channel, using a dual exponential function with time constants of 80 ** and 40 msec as in Suarez, Koch and Douglas 1995 (J. Neurosci. 15, ** 6700-1719; cat visual cortex). ** A more detailed model can be found in Otis, De Koninck and Mody 1993 ** (J. Physiol. 463, 391-407; rat hippocampal slices; this model uses 4th ** power exponential activation and dual exponential inactivation). ** See also Benardo 1994 (J. Physiol. 476.2, 203-215; slice rat neocortex) ** and Connors, Malenka and Silva 1988 (J. Physiol. 406, 443-468; slice ** rat and cat visual cortex. */ float Q10_synapse = 3.0 if (!({exists GABAB})) create synchan2 GABAB end setfield GABAB Ek {E_GABAB} tau1 {0.080 / Q10_synapse} \ tau2 {0.040 / Q10_synapse} \ gmax {G_GABAB} frequency 0.0 end // make_GABAB_channels function make_L5P_synchans echo making Purkinje synapse library... /* The conductance equations in this library are general and not ** specific to the Purkinje celxl */ // Make library protoptypes ***** if ({!{exists /library}}) create neutral /library disable /library end ce /library if ({!{exists L5P}}) create neutral L5P end ce L5P make_GABAA_channels make_AMPA_channels make_NMDA_channels make_GABAB_channels end // make_L5P_synchans make_L5P_synchans