Olfactory bulb microcircuits model with dual-layer inhibition (Gilra & Bhalla 2015)

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Accession:153574
A detailed network model of the dual-layer dendro-dendritic inhibitory microcircuits in the rat olfactory bulb comprising compartmental mitral, granule and PG cells developed by Aditya Gilra, Upinder S. Bhalla (2015). All cell morphologies and network connections are in NeuroML v1.8.0. PG and granule cell channels and synapses are also in NeuroML v1.8.0. Mitral cell channels and synapses are in native python.
Reference:
1 . Gilra A, Bhalla US (2015) Bulbar microcircuit model predicts connectivity and roles of interneurons in odor coding. PLoS One 10:e0098045 [PubMed]
Model Information (Click on a link to find other models with that property)
Model Type: Realistic Network;
Brain Region(s)/Organism: Olfactory bulb;
Cell Type(s): Olfactory bulb main mitral GLU cell; Olfactory bulb main interneuron periglomerular GABA cell; Olfactory bulb main interneuron granule MC GABA cell;
Channel(s): I A; I h; I K,Ca; I Sodium; I Calcium; I Potassium;
Gap Junctions:
Receptor(s): AMPA; NMDA; Gaba;
Gene(s):
Transmitter(s): Gaba; Glutamate;
Simulation Environment: Python; MOOSE/PyMOOSE;
Model Concept(s): Sensory processing; Sensory coding; Markov-type model; Olfaction;
Implementer(s): Bhalla, Upinder S [bhalla at ncbs.res.in]; Gilra, Aditya [aditya_gilra -at- yahoo -period- com];
Search NeuronDB for information about:  Olfactory bulb main mitral GLU cell; Olfactory bulb main interneuron periglomerular GABA cell; Olfactory bulb main interneuron granule MC GABA cell; AMPA; NMDA; Gaba; I A; I h; I K,Ca; I Sodium; I Calcium; I Potassium; Gaba; Glutamate;
#!/usr/bin/env python
# -*- coding: utf-8 -*-

import os
import sys
import math
import pickle
import datetime

sys.path.extend(["..","../networks","../generators","../simulations"])

from OBNetwork import *

from stimuliConstants import * # has SETTLETIME
from simset_activinhibition import * # has REALRUNTIME
from sim_utils import * # has build_tweaks()
from data_utils import *

## constrate firefiles start firing immediately (no SETTLETIME)
## Hence just have 20ms as SETTLETIME. overrides that from stimuliConstants.py.
## But PGs have an initial spike on a high initial settling transient, so 100ms:
SETTLETIME = 100e-3#20e-3 # s
RUNTIME = REALRUNTIME + SETTLETIME

from pylab import * # part of matplotlib that depends on numpy but not scipy

## set lateral_mitnum to 1 for 2MITS / 2 for 2GLOMS option set in generate_neuroML.py
## Note that for directed connectivity, mit 3 is used for directed conn to mit 0 in generate_neuroml.py,
## thus mit 2 represents a non-directed conn cell.
lateral_mitnum = 3#2#3
if REVERSED_ADI:
    mitralmainidx = lateral_mitnum
    mitralsidekickidx = 0
else:
    mitralmainidx = 0
    mitralsidekickidx = lateral_mitnum

########## You need to run:
## From gj:
## ./restart_mpd_static
## The 0th (boss process) will always be node000 as it is the first node in ~/hostfile.
## HENCE FROM node000: cd to the working directory simulations/ (so that sys.path has accurate relative paths)
## mpiexec -machinefile ~/hostfile -n <numplotpts*2+1> ~/Python-2.6.4/bin/python2.6 inhibition_tuftinput.py
## I typically take numplots = 20 (generate_firefiles_constrate.py should also be called accordingly first)
## nohup mpiexec -machinefile ~/hostfile -n 41 ~/Python-2.6.4/bin/python2.6 inhibition_tuftinput.py &> nohup_tuftinh.out < /dev/null &
## For not showing the plots, append NOSHOW as a commandline argument.

## for lateral propagation of AP :
## You need to set seed,mitdist,mitdiststr (simset_activinhibition_minimal.py)
## and segnames (in code below), and run :
## python2.6 inhibition_tuftinput.py

##### 0 rank process is for collating all jobs. (rank starts from 0)
##### I assume rank 0 process always runs on the machine whose X window system has a Display connected
##### and can show the graphs!!!!!!
##### The rank 0 stdout is always directed to the terminal from which mpiexec was run.
##### I hope X output also works the same way.
##### For long simulations save results in a text file for replotting later and avoid above ambiguity.
from mpi4py import MPI

mpicomm = MPI.COMM_WORLD
mpisize = mpicomm.Get_size() # Total number of processes
mpirank = mpicomm.Get_rank() # Number of my process
mpiname = MPI.Get_processor_name() # Name of my node
# The 0th process is the boss who collates/receives all data from workers
boss = 0
print 'Process '+str(mpirank)+' on '+mpiname+'.'

if PLOT_EXTRAS and mpisize>11:
    print "You want to plot Vm-s of mitrals, singles and granules for",mpisize,"processes."
    print "To avoid you the embarrassment of lots of figures, I'm aborting."
    print "Check if PLOT_EXTRAS in simset_activinhibition.py is True."
    sys.exit(1)

## half jobs run with mitral B off, half with on, hence twice the step
if mpisize>1: Ainjectarray = arange(0.0,ORNmax,ORNmax/float(mpisize-1)*2)
else: Ainjectarray = [15.0]
numpts = len(Ainjectarray)
onInject = oninject_ext # Hz # from simset_activinhibition_minimal.py
ORN_constrate_str = '../firefiles/firefiles_constrate/firetimes_constrate'

## Output file name
today = datetime.date.today()
if NO_SINGLES: singles_str = '_NOSINGLES'
else: singles_str = '_SINGLES'
if NO_JOINTS: joints_str = '_NOJOINTS'
else: joints_str = '_JOINTS'
if NO_PGS: pgs_str = '_NOPGS'
else: pgs_str = '_PGS'
if IN_VIVO: invivo_str = '_invivo'
else: invivo_str = ''
if DIRECTED: dirt_str = '_directed'+str(FRAC_DIRECTED)
else: dirt_str = ''
if REVERSED_ADI: rev_str = '_reversed'
else: rev_str = ''
if ASYM_TEST: asym_str = '_asym' ## asym test acts as same ORN frate into A and B
else: asym_str = ''
if ODORINH: odorinh_str = '_odorinh'
else: odorinh_str = '_airinh'
#now =  datetime.datetime.now().strftime("%Y_%m_%d_%H_%M")+'_'
now = '' # stable enough to not bother about the date-time of simulation
outfilename = '../results/tuftADI/'+now+'tuftADI_'+str(lateral_mitnum)+'_seed'+netseedstr+mitdistancestr+\
    singles_str+joints_str+pgs_str+invivo_str+dirt_str+rev_str+asym_str+odorinh_str+'.pickle'

#-----------------------------------------------------------

def setup_stim(network,mpirank_given):
    iAinject = Ainjectarray[(mpirank_given-1)%numpts]
    ## half jobs run with mitral B off, half with on
    if ASYM_TEST: # for asymmetry, inject same current in mitA and mitB
        iBinject = ((mpirank_given-1)/numpts) * iAinject # integer division
    else: # for ADI, inject current to get 80Hz in mitB
        iBinject = ((mpirank_given-1)/numpts) * onInject # integer division
    ipulse_duration = REALRUNTIME # seconds

    setup_tuftinject(network, mitralmainidx, iAinject, mpirank_given)        
    setup_tuftinject(network, mitralsidekickidx, iBinject, mpirank_given)        
    print 'Tuft ORN injecting mitral A with '+str(iAinject)+' Hz and B with '+\
        str(iBinject)+' Hz at process = '+str(mpirank_given)+'.'

def setup_tuftinject(network, mitnum, frate, mpirank_given=mpirank):
    ## first figure out those PGs connected to this mitral mitnum
    PGs_to_thismit = []
    for projname in network.projectionDict.keys():
        if 'PG_mitral' in projname:
            for i,proj in enumerate(network.projectionDict[projname][2]):
                ## get the glomnum from the post path proj[2]
                ## name of the mitral cell from '/mitrals_2/...'
                mitname = string.split(proj[2],'/')[1]
                ## mit number from 'mitrals_2'
                conn_mitnum = int(string.split(mitname,'_')[1])
                if conn_mitnum == mitnum:
                    ## name of the PG cell from '/PGs_2/...'
                    pgname = string.split(proj[1],'/')[1]
                    PGs_to_thismit.append(pgname)
    ## Now connect the ORNs
    ORNfilebase = ORN_constrate_str + str(frate)+'_trial'+str(mpirank_given-1)
    for projname in network.projectionDict.keys():
        #### Calling attach_spikes() for each projection,
        #### would reconnect files to the same segment multiple times.
        #### But attach_files_uniquely() checks whether timetable.tableSize is zero or not
        #### i.e. files already attached or not.
        ## connect ORNs to mitrals
        if 'ORN_mitral' in projname:
            print "connecting ORN files to mitrals for mitnum",mitnum
            for i,proj in enumerate(network.projectionDict[projname][2]):
                ## get the mitnum from the post path proj[2]
                ## name of the mitral cell from '/mitrals_2/...'
                mitname = string.split(proj[2],'/')[1]
                ## mit number from 'mitrals_2'
                conn_mitnum = int(string.split(mitname,'_')[1])
                if conn_mitnum == mitnum:
                    attach_files_uniquely(ORNfilebase,proj[0],proj[2])
        ## connect ORNs to PG
        if 'ORN_PG' in projname:
            print "connecting ORN files to PGs for mitnum",mitnum
            for i,proj in enumerate(network.projectionDict[projname][2]):
                pgname = string.split(proj[2],'/')[1] # name of the PG cell from '/PGs_2/...'
                if pgname in PGs_to_thismit:
                    attach_files_uniquely(ORNfilebase,proj[0],proj[2])

def attach_files_uniquely(filebase,synname,postsegpath):
    ttpath = postsegpath+'/'+synname+'_tt'
    if moose.context.exists(ttpath):
        # timetable already created by networkML reader - just wrap it below.
        tt = moose.TimeTable(ttpath) # post_segment_path+'/'+syn_name+'_tt'
    else:
        ## if timetable was not already created by networkML reader,
        ## it means that the synaptic weights must be zero!
        ## (no extra inhibition - only main inhibition)
        ## hence do not attach spikefiles
        return
    if tt.tableSize != 0: return # if files are already attached, do nothing!
    ## attach_spikes() accesses filenumbers to this segment
    ## from 'fileNumbers' field (of the timetable object in MOOSE)
    ## which is created while reading in networkML.
    attach_spikes(filebase, tt, uniquestr+str(mpirank))

def run_inhibition(network, tables):
    resetSim(network.context, SIMDT, PLOTDT) # from moose_utils.py sets clocks and resets
    network.context.step(RUNTIME)
    # get mitral A's firing rate
    oneoverISI, meanfreq, events = \
        calcFreq(network.mitralTable[mitralmainidx]._vmTableSoma,\
            RUNTIME, SETTLETIME, PLOTDT, THRESHOLD, SPIKETABLE)
    mpicomm.send( meanfreq, dest=boss, tag=0 ) # frequency tag
    oneoverISI, meanfreq, events = \
        calcFreq(network.mitralTable[mitralsidekickidx]._vmTableSoma,\
            RUNTIME, SETTLETIME, PLOTDT, THRESHOLD, SPIKETABLE)
    mpicomm.send( meanfreq, dest=boss, tag=1 ) # frequency of mit B tag
    #print 'Firing rate = '+str(result[3])+'Hz on injecting mitral A with '+str(iAinject)+\
    #    ' and B with '+str(iBinject)+' at process = '+str(mpirank)+'.'
    ##mpicomm.send( array(network.mitralTable[mitralmainidx]._vmTableSoma), dest=boss, tag=1 )
    ##mpicomm.send( array(network.mitralTable[mitralsidekickidx]._vmTableSoma), dest=boss, tag=2 )
    mpicomm.send( numpy_convert_tables(tables), dest=boss, tag=3 ) # extra tables tag   
    print 'Sent output from process '+str(mpirank)+'.'


def collate():
    if not NOSHOW:
        mainfig = figure(facecolor='w')
        mainaxes = mainfig.add_subplot(111)
    dual_firingratearray = []
    for mitralBinject in [0,1]:
        firingratearray = []
        for A,mitralAinject in enumerate(Ainjectarray):
            procnum = mitralBinject*numpts + A + 1
            print 'waiting for process '+str(procnum)+'.'
            Afiringrate = mpicomm.recv(source=procnum, tag=0)
            firingratearray.append(Afiringrate)
            Bfiringrate = mpicomm.recv(source=procnum, tag=1)
            print "mitral B firing at",Bfiringrate
            ##mitA = mpicomm.recv(source=procnum, tag=1)
            ##mitB = mpicomm.recv(source=procnum, tag=2)
            tables = mpicomm.recv(source=procnum, tag=3)
            if PLOT_EXTRAS:
                timevec = arange(0.0,RUNTIME+1e-12,PLOTDT)
                titlestr = 'Ainject='+str(mitralAinject)+'Hz Binject='+str(mitralBinject*onInject)+'Hz'
                if not NOSHOW:
                    #figure()
                    #title('red:mitA, green:mitB, '+titlestr)
                    #plot(timevec, mitA, 'r,')
                    #plot(timevec, mitB, 'g,')
                    plot_extras(timevec, tables, NO_PGS, NO_SINGLES, NO_JOINTS, titlestr)
            else:
                iAinject = mitralAinject
                iBinject = mitralBinject * onInject
                activity_table = print_extras_activity(tables, NO_PGS, NO_SINGLES, NO_JOINTS, NO_MULTIS,\
                    'I_A='+str(iAinject)+'Hz & I_B='+str(iBinject)+'Hz.')
                ## annotate each point with #joints;#singles firing
                firingcells = ''
                if not NO_JOINTS:
                    firingcells += str(activity_table['joints'][0])
                    if not NO_SINGLES: firingcells += ';'+str(activity_table['singles'][0])
                elif not NO_SINGLES: firingcells += str(activity_table['singles'][0])
                if not NOSHOW:
                    mainaxes.annotate(firingcells,xy=(mitralAinject,Afiringrate))
        if not NOSHOW:
            mainaxes.plot(Ainjectarray, firingratearray, color=(mitralBinject,1-mitralBinject,0),\
                marker=['+','x'][mitralBinject], label="mitral B ORNs = "+str(mitralBinject*onInject)+" Hz.")
        dual_firingratearray.append(firingratearray)

    fvsifile = open(outfilename,'w')
    pickle.dump((Ainjectarray, dual_firingratearray), fvsifile)
    fvsifile.close()
    print "Wrote",outfilename

    if not NOSHOW:
        mainaxes.legend(loc="lower right")
        mainaxes.set_xlabel("mit A tuft ORNs mean firing rate (Hz)",fontsize=16)
        mainaxes.set_ylabel("mitral A firing rate (Hz)",fontsize=16)
        show()

#----------------------------------------------------

def varyVrest_inject(network,popname,raiseRMP,inject):
    for cell in network.populationDict[popname][1].values():
        tweak_field(cell.path+'/##[TYPE=Compartment]', 'Em', 'Em+'+str(raiseRMP))
        tweak_field(cell.path+'/##[TYPE=Compartment]', 'inject', str(inject))

if __name__ == "__main__":
    if 'NOSHOW' in sys.argv:
        NOSHOW = True
        ## If NOSHOW, then automatic mode, hence don't overwrite resultfile, if exists beforehand.
        if os.path.exists(outfilename):
            ## activdep_inhibition_repeats.py searches for Wrote in first word,
            ## and filename in second word. so output that even if not simulating.
            if mpirank==boss:
                for procnum in range(1,mpisize):
                    mpicomm.recv(source=procnum,tag=10)
                print "ExistsSoNotWrote",outfilename
            else:
                mpicomm.send('done',dest=boss,tag=10)
            sys.exit()
    else: NOSHOW = False
    if mpisize == 1:
        if len(sys.argv)>2: uniquestr = sys.argv[2]+'_' # _ necessary, else say 'adi2'+mpirank is screwed
        else: uniquestr = 'tuft_'
        ## includeProjections gets used only if ONLY_TWO_MITS is True:
        ## Keep below projections to 'second order cells'
        ## i.e. to cells (granules) connected to mits0&1.
        ## The connections between second order cell
        ## and mits0&1 are automatically retained of course.
        ## important to include 'PG' below as 'ORN_PG' is needed,
        ## though 'PG_mitral', 'mitral_PG' connections to/from mits0&1 are kept automatically.
        includeProjections = ['granule_baseline','PG']
        tweaks = build_tweaks( CLUB_MITRALS, NO_SPINE_INH,\
            NO_SINGLES, NO_JOINTS, NO_MULTIS, NO_PGS, ONLY_TWO_MITS,\
            includeProjections, (mitralmainidx,mitralsidekickidx) )
        ## send mpirank to put in ORN filenames
        ## so they do not clash between processes - not needed here
        network = OBNetwork(OBNet_file, synchan_activation_correction,\
            tweaks, mpirank, uniquestr, granfilebase, spiketable=SPIKETABLE)
        #printNetTree() # from moose_utils.py

        ## Vary the resting potential of PGs
        if not NO_PGS: varyVrest_inject(network,'PGs',PG_raiseRMP,PG_inject)
        ## Vary the resting potential of granules
        if not NO_SINGLES: varyVrest_inject(network,'granules_singles',granule_raiseRMP,granule_inject)
        if not NO_JOINTS: varyVrest_inject(network,'granules_joints',granule_raiseRMP,granule_inject)
        if not NO_MULTIS: varyVrest_inject(network,'granules_multis',granule_raiseRMP,granule_inject)

        setup_stim(network,2) # 1 to give no i/p, 2 to give oninject_ext to mitB.
        
        ## setup a separate Vm table, since the one setup by OBNetwork() above stores only spikes.
        mitAsoma_Vm = setupTable('mitAsomaVm',network.mitralTable[mitralmainidx].soma,'Vm')
        #mitBsoma_Vm = setupTable('mitBsomaVm',network.mitralTable[mitralsidekickidx].soma,'Vm')
        mitBsoma_Vm = setupTable('mitBsomaVm',moose.Compartment('/mitrals_3/Seg0_soma_0'),'Vm')
        ## get the nearA and afterA ids for each netfile instance -- see my notes of 17Apr2013 and 5Feb2013.
        mitB_nearA_Vm = setupTable('mitB_nearA_Vm',moose.Compartment('/mitrals_3/Seg0_sec_dendd4_0_261'),'Vm')
        mitB_afterA_Vm = setupTable('mitB_afterA_Vm',moose.Compartment('/mitrals_3/Seg0_sec_dendd4_1_265'),'Vm')

        ## if SPIKETABLE (in simset_activinhibition.py): record the Vm-s of a few interneurons
        ## else: record spiketimes of all interneurons
        tables = setupTables(network, NO_PGS, NO_SINGLES, NO_JOINTS, NO_MULTIS,
            args={'mitrals':(mitralmainidx,)}, spikes=SPIKETABLE)

        resetSim(network.context, SIMDT, PLOTDT) # from moose_utils.py sets clocks and resets
        network.context.step(RUNTIME)

        mitAsoma_Vm = array(mitAsoma_Vm)
        mitBsoma_Vm = array(mitBsoma_Vm)
        timevec_unbinned = arange(0.0,RUNTIME+1e-12,PLOTDT)
        ## plot mitral A/B soma Vm
        figure(facecolor='w')
        title('Vm in mitral A soma')
        plot(timevec_unbinned,mitAsoma_Vm,label='Vm',color='k')
        axes_labels(gca(),"time (s)","Vm (V)",fontsize=label_fontsize+2)
        figure(facecolor='w')
        title('Vm in mitral B soma')
        plot(timevec_unbinned,mitBsoma_Vm,label='Vm',color='k')
        plot(timevec_unbinned,mitB_nearA_Vm,label='Vm',color='b')
        plot(timevec_unbinned,mitB_afterA_Vm,label='Vm',color='r')
        axes_labels(gca(),"time (s)","Vm (V)",fontsize=label_fontsize+2)
        
        ## Paper figure: mitB i.e. directed mitral 3's soma vs inhibitory sites
        fig2 = figure(figsize=(columnwidth/2.0,linfig_height/2.0),dpi=1200,facecolor='w') # 'none' is transparent
        ax = fig2.add_subplot(111)
        plot(timevec_unbinned*1000,array(mitBsoma_Vm)*1000,'-r', label='soma',\
            linewidth=0.5, linestyle='-') # ms and mV
        plot(timevec_unbinned*1000,array(mitB_nearA_Vm)*1000,'-b', label='at inh site',\
            linewidth=0.5, dashes=(1.5,0.5)) # ms and mV
        plot(timevec_unbinned*1000,array(mitB_afterA_Vm)*1000,'-k', label='after inh site',\
            linewidth=0.5, dashes=(0.5,0.5)) # ms and mV
        beautify_plot(ax,x0min=False,drawxaxis=True,drawyaxis=True,\
            xticks=[150,200],yticks=[-75,0,40])
        #add_scalebar(ax,matchx=False,matchy=False,hidex=True,hidey=False,\
        #    sizex=50,labelx='50 ms',sizey=20,labely='20 mV',label_fontsize=5,\
        #    bbox_to_anchor=[0.7,0.6],bbox_transform=ax.transAxes)
        ax.set_xlim(150,200)
        ax.set_xticklabels(['150','200'])
        ax.set_ylim(-75,40)
        axes_labels(ax,"time (ms)","Vm (mV)",fontsize=label_fontsize,xpad=-4,ypad=-3)
        fig2.tight_layout()
        fig2.savefig('../figures/connectivity/cells/mitral_spikeprop.png',dpi=fig2.dpi)
        fig2.savefig('../figures/connectivity/cells/mitral_spikeprop.svg',dpi=fig2.dpi)
        ## Paper figure done

        timevec = arange(0.0,RUNTIME+PLOTDT+1e-12,PLOTDT)
        if SPIKETABLE:
            ## 50ms bins for firingrate
            timevec = arange(50e-3/2.0,REALRUNTIME,50e-3)
            plot_extras_spikes(timevec, tables, NO_PGS, NO_SINGLES, \
                NO_JOINTS, NO_MULTIS, len(timevec), RUNTIME, SETTLETIME)
        else:
            plot_extras(timevec, tables, NO_PGS, NO_SINGLES, NO_JOINTS, NO_MULTIS)
        
        show()

    else:
        if mpirank==boss:
            collate()
        else:
            if len(sys.argv)>2: uniquestr = sys.argv[2]+'_' # _ necessary, else say 'adi2'+mpirank is screwed
            else: uniquestr = 'tuft_'
 
            ## If CLUB_MITRAL=False, then extra exc from mitral sisters
            ## (to certain connected granules as proxy for unmodelled sisters) does NOT get used.
            ## Instead, here I connect extra baseline excitation to ALL granules if odor is non-zero.
            ## This extra basline is only if ODORINH i.e. we are simulating odor inh and not air inh
            if ODORINH and not CLUB_MITRALS:
                granfilebase += '_extra'

            ## includeProjections gets used only if ONLY_TWO_MITS is True:
            ## Keep below projections to 'second order cells'
            ## i.e. to cells (granules) connected to mits0&1.
            ## The connections between second order cell
            ## and mits0&1 are automatically retained of course.
            ## important to include 'PG' below as 'ORN_PG' is needed,
            ## though 'PG_mitral', 'mitral_PG' connections to/from mits0&1 are kept automatically.
            includeProjections = ['granule_baseline','PG']
            tweaks = build_tweaks( CLUB_MITRALS, NO_SPINE_INH,\
                NO_SINGLES, NO_JOINTS, NO_MULTIS, NO_PGS, ONLY_TWO_MITS,\
                includeProjections, (mitralmainidx,mitralsidekickidx) )
            ## send mpirank to put in ORN filenames
            ## so they do not clash between processes - not needed here
            network = OBNetwork(OBNet_file, synchan_activation_correction,\
                tweaks, mpirank, uniquestr, granfilebase, spiketable=SPIKETABLE)
            #printNetTree() # from moose_utils.py

            ### Vary the resting potential of PGs
            #if not NO_PGS: varyVrest_inject(network,'PGs',PG_raiseRMP,PG_inject)
            ### Vary the resting potential of granules
            #if not NO_SINGLES: varyVrest_inject(network,'granules_singles',granule_raiseRMP,granule_inject)
            #if not NO_JOINTS: varyVrest_inject(network,'granules_joints',granule_raiseRMP,granule_inject)
            #if not NO_MULTIS: varyVrest_inject(network,'granules_multis',granule_raiseRMP,granule_inject)

            setup_stim(network, mpirank)
            ## if SPIKETABLE: record the Vm-s of a few interneurons
            ## else: record spiketimes of all interneurons
            tables = setupTables(network, NO_PGS, NO_SINGLES, NO_JOINTS, NO_MULTIS,
                args={'mitrals':(mitralmainidx,)}, spikes=SPIKETABLE)
            run_inhibition(network, tables)

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