Hippocampal CA1 NN with spontaneous theta, gamma: full scale & network clamp (Bezaire et al 2016)

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Accession:187604
This model is a full-scale, biologically constrained rodent hippocampal CA1 network model that includes 9 cells types (pyramidal cells and 8 interneurons) with realistic proportions of each and realistic connectivity between the cells. In addition, the model receives realistic numbers of afferents from artificial cells representing hippocampal CA3 and entorhinal cortical layer III. The model is fully scaleable and parallelized so that it can be run at small scale on a personal computer or large scale on a supercomputer. The model network exhibits spontaneous theta and gamma rhythms without any rhythmic input. The model network can be perturbed in a variety of ways to better study the mechanisms of CA1 network dynamics. Also see online code at http://bitbucket.org/mbezaire/ca1 and further information at http://mariannebezaire.com/models/ca1
References:
1 . Bezaire MJ, Raikov I, Burk K, Vyas D, Soltesz I (2016) Interneuronal mechanisms of hippocampal theta oscillations in a full-scale model of the rodent CA1 circuit. Elife [PubMed]
2 . Bezaire M, Raikov I, Burk K, Armstrong C, Soltesz I (2016) SimTracker tool and code template to design, manage and analyze neural network model simulations in parallel NEURON bioRxiv
Model Information (Click on a link to find other models with that property)
Model Type: Realistic Network;
Brain Region(s)/Organism: Hippocampus;
Cell Type(s): Hippocampus CA1 pyramidal cell; Hippocampus CA1 interneuron oriens alveus cell; Hippocampus CA1 basket cell; Hippocampus CA1 stratum radiatum interneuron; Hippocampus CA1 bistratified cell; Hippocampus CA1 axo-axonic cell; Hippocampus CA1 PV+ fast-firing interneuron;
Channel(s): I Na,t; I K; I K,leak; I h; I K,Ca; I Calcium;
Gap Junctions:
Receptor(s): GabaA; GabaB; Glutamate; Gaba;
Gene(s):
Transmitter(s): Gaba; Glutamate;
Simulation Environment: NEURON; NEURON (web link to model);
Model Concept(s): Oscillations; Methods; Connectivity matrix; Laminar Connectivity; Gamma oscillations;
Implementer(s): Bezaire, Marianne [mariannejcase at gmail.com]; Raikov, Ivan [ivan.g.raikov at gmail.com];
Search NeuronDB for information about:  Hippocampus CA1 pyramidal cell; Hippocampus CA1 interneuron oriens alveus cell; GabaA; GabaB; Glutamate; Gaba; I Na,t; I K; I K,leak; I h; I K,Ca; I Calcium; Gaba; Glutamate;
/************************************************************
'ca1' model code repository
Written by Marianne Bezaire, marianne.bezaire@gmail.com, www.mariannebezaire.com
and Ivan Raikov, ivan.g.raikov@gmail.com
Last updated on April 10, 2015
Latest versions of this code are available online at:
ModelDB: 
Open Source Brain: http://www.opensourcebrain.org/projects/nc_ca1

This code is associated with the publications:

This code can be used with the SimTracker tool:
http://senselab.med.yale.edu/SimToolDB/ShowTool.asp?Tool=153281

If you do not want to run this code, but want to analyze the results
obtained by running this code, the results are available upon request by
emailing marianne.bezaire@gmail.com

If you want to view a graphical description of the model network built
by this code, please see: 
http://www.ivansolteszlab.org/models/ca1/graphicalmodel.html

For more information about this code, see:
- ModelDB Quick Start Guide.pdf (included with model code)
- Lab website: http://www.ivansolteszlab.org/models/ca1.html

To run this model from the command line, enter:
nrniv main.hoc
************************************************************
This main code file has the following structure:
I.  LOAD LIBRARIES
II. SET MODEL SIZE, CELL DEFINITIONS
III. SET UP PARALLEL CAPABILITY AND MEMORY USAGE RECORDER, AND WRITE OUT RUN RECEIPT
IV. CREATE, UNIQUELY ID, AND POSITION CELLS
V.	CONNECT THE MODEL CELLS AND CONNECT THE STIMULATION CELLS TO SOME MODEL CELLS
VI.	INITIALIZE AND RUN NETWORK, OUTPUT RESULT FILES
************************************************************/

loadstart = startsw()					// record the start time of the set up
/***********************************************************************************************
I.  LOAD LIBRARIES
***********************************************************************************************/
{load_file("nrngui.hoc")}				// Standard definitions - NEURON library file

{load_file("netparmpi.hoc")}			// Contains the template that defines the properties of
										//  the ParallelNetManager class, which is used to set
										//   up a network that runs on parallel processors
										
{load_file("./setupfiles/ranstream.hoc")}	// Contains the template that defines a RandomStream
											//  class used to produce random numbers
											
{load_file("./setupfiles/CellCategoryInfo.hoc")}	// Contains the template that defines a 
													//  CellCategoryInfo class used to store
													// 	celltype-specific parameters

{load_file("./setupfiles/SynStore.hoc")}	// Contains the template that defines a holder of
											//  synapse type info
													
{load_file("./setupfiles/defaultvar.hoc")}	// Contains the proc definition for the default_var proc

{load_file("./setupfiles/parameters.hoc")}	// Loads in default values for operational and model
											// parameters whose values were not specified ahead of
											// time, ie by passing them in at the command line or
											// in a wrapper hoc file						
			
{load_file("./setupfiles/set_other_parameters.hoc")}// Loads in operational and model parameters
													//  that we would not want to define ahead of
													// time (either because they depend on ones that
													// *are* specified ahead of time or because they
													// don't usually change

//{load_file("./setupfiles/check_for_files.hoc")}	// Checks that the necessary chosen files exist
													//  (datasets, connectivity, and stimulation)
                                                        
                                                        
                                                            
/***********************************************************************************************
II. SET MODEL SIZE, CELL DEFINITIONS
***********************************************************************************************/

{load_file("./setupfiles/load_cell_category_info.hoc")}	// Loads cell-specific information into one
														// 'CellCategoryInfo' object for each cell
														// type. Info includes number of cells,
														// gid ranges, type of cell, name of cell 
														// Also, defines 'numCellTypes' used below

{load_file("./setupfiles/load_cell_conns.hoc")}	// Load in the cell connectivity info
{load_file("./setupfiles/load_cell_syns.hoc")}	// Load in the synapse info

strdef tempFileStr						// Define a string reference to store the name of the
										//  current cell template file 

proc loadCellTemplates(){local i		// Proc to load the template that defines (each) cell class

	for i=0, numCellTypes-1 {			// Iterate over each cell type
	
		sprint(tempFileStr,"./cells/class_%s.hoc",cellType[i].technicalType)	// Concatenate the
																				//  path and file
																				
		load_file(tempFileStr)			// Load the file with the template that defines the class
										//  for each cell type
	}
}	
loadCellTemplates()

proc calcNetSize(){local i				// Calculate the final network size (after any cell death)

	totalCells = 0						// Initialize totalCells (which counts the number of 'real'
										//  cells) so we can add to it iteratively in the 'for' loop
										
	ncell = 0							// Initialize ncell (which counts all 'real' and 'artificial'
										//  cells) so we can add to it iteratively in the 'for' loop
										
	prevgid = -1						// Initialize gid setter
	
	for i=0,numCellTypes-1 {			// For each cell type
	
		if (cellType[i].layerflag==0 && cellType[i].is_art==0) {	// For cell types in layer 0, which are susceptible to death
		
			cellType[i].numCells = int(cellType[i].numCells * ((100-PercentCellDeath)/100))
										// Calculate the number of cells surviving after cell loss
			
			if (cellType[i].numCells == 0) {cellType[i].numCells = 1}	// If all cells of one type
																		//  are killed, let 1 live
		}
		
		cellType[i].updateGidRange(prevgid+1)	// Update the gid range for each
												//  cell type

		if (cellType[i].is_art==0) {
			totalCells = totalCells + cellType[i].numCells		// Update the total number of cells
																//   after cell death, not including
																//   artificial cells
		}
		
		ncell = ncell + cellType[i].numCells 					// Update the total number of cells
																//   after cell death, including
																//   artificial cells
		prevgid = cellType[i].cellEndGid						// Track the prevgid so the subsequent
																//  gid ranges can be updated
	}
}
calcNetSize()

proc calcBinSize(){

	for i=0, numCellTypes-1 {		// Using the specified dimensions of the network (in um) and
									//  the total number of cells of each type, set the number
									//  of bins in X, Y, Z dimensions such that the cells will be
									//  evenly distributed throughout the allotted volume
	
		cellType[i].setBins(LongitudinalLength,TransverseLength,LayerVector.x[cellType[i].layerflag])  
									// For the z length, use the height of the layer in which the
									// cell somata are found for this cell type
	}
}
calcBinSize()

/***********************************************************************************************
III. SET UP PARALLEL CAPABILITY AND MEMORY USAGE RECORDER, AND WRITE OUT RUN RECEIPT
***********************************************************************************************/
objref pnm, pc, nc, nil
proc parallelizer() {
	pnm = new ParallelNetManager(ncell)	// Set up a parallel net manager for all the cells
	pc = pnm.pc
	pnm.round_robin()					// Incorporate all processors - cells 0 through ncell-1
										//	are distributed throughout the hosts
										//	(cell 0 goes to host 0, cell 1 to host 1, etc, like
										//	dealing a deck of cards)
}
parallelizer()

iterator pcitr() {local i1, i2, gid, startgid	// Create an iterator for use instead of standard 'for' 
												//  loop. This provides an easy way for each processor 
												//  to do a 'for' loop only over cells that it owns. 
												//  This particular iterator code is slightly more 
												//  complex than Michael Hines' usual example, as it 
												//  allows for getting the index of any cell on a 
												//  processor relative to all cells on a processor or 
												//  just that cell type.
												//
												//  usage:
												//  for pcitr(&i1, &i2, &gid, it_start, it_end) {do stuff}
												//   the first three arguments are pointers that pcitr will
												//	  fill for you:
												//    - i1: index of the cell on the cell list for that host
												//    - i2: index of the cell for that cell type on that host
												//    - gid: index of the cell in the whole network
												//   it_start and it_end let you define the gid range over
												//     which to iterate, usually one of the following:
												//		- the gid range for a cell type
												//		- the gid range for the whole network
										
	numcycles = int($4/pc.nhost)
	extra = $4%pc.nhost
	addcycle=0
	if (extra>pc.id) {addcycle=1}
	startgid=(numcycles+addcycle)*pc.nhost+pc.id
	i1 = numcycles+addcycle // the index into the cell # on this host.
	i2 = 0 // the index of the cell in that cell type's list on that host
	if (startgid<=$5) {
		for (gid=startgid; gid <= $5; gid += pc.nhost) {	// Just iterate through the cells on
														//  this host(this simple statement
														//  iterates through all the cells on
														//  this host and only these cells because 
														//  the roundrobin call made earlier dealt
														//  the cells among the processors in an
														//  orderly manner (like a deck of cards)
				$&1 = i1
				$&2 = i2
				$&3 = gid
				iterator_statement						// this is the code that you put inside  
														//  your pcitr loop when you call the 
														//  pcitr iterator
				i1 += 1
				i2 += 1
		}
	}
}

objref  strobj
strobj = new StringFunctions()
strdef direx
if (strcmp(UID,"0")==0 && pc.id==0) {					// if a unique ID (UID) was not already 
														//  specified for this run, then generate 
														//  it now. Only one per run is needed, so
														//  only host 0 should generate it. A unique 
														//  ID could have been generated outside of 
														//  NEURON (say, by using the SimTracker) and 
														//  then passed in to a wrapper hoc code file
														//  that calls this file, or passed in at the 
														//  command line. If one was not already passed 
														//  in, then the UID defaults to 0 and the code
														//  below runs
														
	type = unix_mac_pc() 		// get the operating system type:
								//  1 if unix, 2 if mac, 3 if mswin, or 4 if mac osx darwin
	
	if (type<3 || type==4) { // unix or mac
		if (OK2executeSysCmds==1) {
			{system("uuidgen", direx)} // use a built in system command to generate the UID
			strobj.left(direx, strobj.len(direx)-1) // remove a newline character from the end
		} else { // Can't access system commands so can't get a UID
			direx = "unknown"
		}
	} else { // pc
		{system("cscript //NoLogo setupfiles/uuid.vbs", direx)} // use a custom VBS (Visual Basic script)
																//  to generate the UID. This VBS file is
																//  included in the code repository
	}
	UID = direx
}

if (OK2executeSysCmds==1) {
	{load_file("./setupfiles/save_run_info.hoc")}		// This file will ensure a unique RunName (if 
														//  the specified RunName was already used, it 
														//  will append numbers to it like "_00", "_01" 
														//  to create a unique RunName). Then it will 
														//  create a RunName directory within the results 
														//  folder, where all the results will be stored. 
														//  It will also write out a receipt of the run 
														//  parameters and other administrative sorts of 
														//  run information.
	// NOTE: If you don't like the NEURON code to make new directories, concatenate files, or delete files, 
	//  set the OK2executeSysCmds parameter to 0 in "set_other_parameters.hoc" file. Watch out though, if 
	//  OK2executeSysCmds is set to 0, the code will use whatever RunName you supply or 'none' if you do not
	//  supply a RunName. This means:
	//  - the program will expect a directory named 'RunName' to be in the results directory
	//  - the program will overwrite any existing files in the ./results/RunName directory
}

objref  memfile, topfile

{zzz=0} // zzz contains the last memory usage from the previous call to mallinfo

if (pc.id==0) {
	{sprint(cmd,"./results/%s/memory.dat", RunName)} 	// Host 0 will make repeated calls to the 
	{memfile = new File(cmd)}							//  mallinfo function defined below and 
	{memfile.wopen()}									//  record the nrn_mallinfo output here

	{sprint(cmd,"./results/%s/topoutput.dat", RunName)}	// Host 0 will make repeated calls to the 
	{topfile = new File(cmd)}							//  mallinfo function defined below and  
	{topfile.wopen()}									//  record the top command output here
}

{load_file("./setupfiles/memory_fcns.hoc")} 	// Defines a memory recording function that calls
											//  mallinfo and also top to track memory usage by
											//  the code

{zzz = mallinfo(zzz, "Prior to creating cells")}	// Memory usage prior to creating the cells

loadtime = startsw() - loadstart		// Calculate the time used for the set up phase (now - recorded start time) in seconds
if (pc.id == 0) {printf("\nTIME HOST 0: %g seconds (set up)\n************\n", loadtime)}
createstart = startsw()					// Record the start time of the cell creation phase

/***********************************************************************************************
IV. CREATE, UNIQUELY ID, AND POSITION CELLS
***********************************************************************************************/
objref cells, ransynlist, ranstimlist, raninitlist, ranwgtlist, ranlfplist
cells = new List()			// a list that will contain an entry for each cell in the network						
ransynlist = new List()		// a list that will contain an entry for each random number generator
							//  used by the synapse chooser (during the connectivity part of the 
							//  code) each cell will have its own random number generator for 
							//  synapse choosing
							
ranwgtlist = new List()		// a list that will contain an entry for each random number generator
							//  used during the connectivity part of the 
							//  code to randomize the synapse weights of the connections
							
ranstimlist = new List()	// a list that will contain an entry for each random number generator
							//  used by the stimulation generator (during the actual simulation)
							//  each cell will have its own random number generator for stimulation
							//  specification. But only artificial cells will actually use their 
							//  generators
							
raninitlist = new List()	// a list that will contain an entry for each random number generator
							//  used by the voltage initializer (when designing the simulation)
							//  each cell will have its own random number generator that will be
							//  called once per point per section within the cell
							
ranlfplist = new List()	// a list that will contain an entry for each random number generator
							//  used by the LFP sampling algorithm (when designing the simulation)
							//  if the cell type's contribution is included in the LFP calculation
							//  and the cell is further away from the electrode point than MaxEDist
							//  (see npole_lfp.hoc for more detail)
													
{load_file("./setupfiles/create_cells_pos.hoc")}	// Creates each cell on its assigned host
													//  and sets its position

strdef cmd
{load_file("./setupfiles/write_positions.hoc")}		// Defines the proc to write the position
													//  of each cell to file
													
if (PrintCellPositions>0) {posout()}	// Calls the proc that writes the position of each cell 
										//  to the file "position.dat" in the format: gid, x, y, z

createtime = startsw() - createstart	// Calculate time taken to create the cells
if (pc.id == 0) {printf("\nTIME HOST 0: %g seconds (created cells)\n************\n", createtime)}
connectstart = startsw()				// Grab start time of cell connection

oldtimeout = pc.timeout(0)	// Addresses timeout error that was occuring on some computers

zzz = mallinfo(zzz, "After creating cells") // Record the memory usage after cells are created

// Define the position of an arbitrary point within the network. This
//  point can then be used to set regional stimulation or to limit the
//  pyramidal cells from which the average LFP is recorded to a specific
//  location, etc.

zzz = mallinfo(zzz, "After creating cells")

/***********************************************************************************************
V.	CONNECT THE MODEL CELLS AND CONNECT THE STIMULATION CELLS TO SOME MODEL CELLS
***********************************************************************************************/

{load_file("./setupfiles/nc_append_functions.hoc")}	// Defines nc_append and nc_appendo, which 
													//  are procs used to create the netcon
													//  object associated with each synapse
													//  between cells

MakeArtConns=1
{sprint(cmd,"./connectivity/%s_connections.hoc", Connectivity)}
{load_file(cmd)}			// Loads in a particular algorithm to connect the cells in the network.
							//  Each algorithm is specified by a different file in the connectivity
							//  folder, and the file names are all of the format:
							//  [Connectivity]_connections.hoc, where [Connectivity] is the value in
							//  the Connectivity parameter that was set in the parameters.hoc file
							//  loaded above, or passed to this code file via the command line or a
							//  wrapper hoc file. The program only loads one connectivity file for
							//  each simulation run.
							// Some connectivity algorithms connect all network cells, including the
							//  artificial cells, while others only make the connections between
							//  real cells. Which connectivity algorithm you use will depend on the
							//  stimulation algorithms you use (as some stimulation algorithms specify
							//  the connectivity of the artificial cells onto the real cells).							

zzz = mallinfo(zzz, "After connecting cells")	// Record the memory usage after creating the connections

{sprint(cmd,"./stimulation/%s_stimulation.hoc", Stimulation)}
{load_file(cmd)}			// Loads in a particular algorithm to stimulation the network. The algorithm
							//  will always specify the spike properties (or actual spike trains) of the
							//  artificial cells in the model, and for some cases will also specify the
							//  connectivity from the artificial (stimulating) cells to the real cells of
							//  the model.

zzz = mallinfo(zzz, "After defining stimulation")	// Record the memory usage after defining stimulation

{load_file("./setupfiles/write_conns_functions.hoc")}	// Defines procedures that can be used to write out
														//  connectivity information for the model network.
														//  The procedures are not called by this file, but
														//  may be called later on in this code.
														
if (PrintConnSummary==1) {printNumConFile()}	// Write "numcons.dat": a QUICK, SMALL summary file of the # of
												//  connections by pre and post cell types
										
if (PrintConnDetails==2) {allCellConnsParallel()}// Write "connections.dat": a VERY SLOW, LARGE file of all cell
												//// Write "connections.dat": a VERY SLOW, LARGE file of all cell
												//	connections (pre- and post-synaptic cell gids, synapse type,
												//  host on which the postsynaptic cell exists). Do NOT do this
												//  for large networks. Just print out a few of the connections
												//  instead, using recordedCellConnsParallel()

if (PrintVoltage>0) {load_file("./setupfiles/recordvoltageauto.hoc")}	// This file contains a function to
																		//  record intracellular somatic
																		//  voltage traces of a few cells
																		//  of each type in your model.
																		//  Alternatively, you can save only
																		//  the connectivity information for
																		//  a few cells and combine that
																		//  information with the spike raster
																		//  to regenerate the voltage trace of
																		//  any cell for which you saved the 
																		//  connectivity information.

if ((PrintConnDetails==0) && (PrintVoltage>0)) {recordedCellConnsParallel()}// If detailed connections are not being
																			//  written out, at least write them out 
																			//  for cells that are being traced
																			
{load_file("./setupfiles/dipole_lfp.hoc")}	// Defines the code to calculate the dipole LFP
{load_file("./setupfiles/npole_lfp.hoc")}	// Defines the code to calculate the n-pole LFP
{load_file("./setupfiles/randomVinit.hoc")}	// Random voltage initialization

connecttime = startsw() - connectstart			// Calculate time taken to connect the cells
if (pc.id == 0) {printf("\nTIME HOST 0: %g seconds (connected cells)\n************\n", connecttime)}
simstart = startsw()

/***********************************************************************************************
VI.	INITIALIZE AND RUN NETWORK, OUTPUT RESULT FILES
***********************************************************************************************/
{load_file("./setupfiles/custom_init.hoc")} 	// Redefines the init proc that initializes the
												//  simulation so that it includes a "pre-run" of the
												//  simulation to reach steady state prior to the start
												//  of the simulation at time t = 0 ms.
												// This function will be called later, just before the
												//  simulation starts.
												//
												// WARNING: Any time you redefine the init function, you
												//  risk not completely initializing everything that needs
												//  to be initialized. Read the Initialization section of
												//  the NEURON Book and see this NEURON forum thread for an
												//  example of how custom initialization can be mishandled:
												//  https://www.neuron.yale.edu/phpBB/viewtopic.php?f=8&t=3127

{load_file("./setupfiles/write_results_functions.hoc")}		// Defines procedures to write out various
															//  simulation results, but does not call
															//  the procedures yet. They will be called
															//  later on.

{load_file("./setupfiles/simulation_optimization.hoc")}		// Sets various simulation conditions to
															//  make the simulation more efficient. Note
															//  that the settings in here will need to be
															//  tested and possibly altered for each
															//  computer that the code is run on, and
															//  the ideal value for some settings may
															//  also change if your network model changes
															//  significantly (in number of cells, activity
															//  level of cells, number of processors you
															//  usually use, etc).

if (get_spike_hist==1) {		// If you use spike compression to speed up the spike exchange part of the
								//  simulation (see the simulation_optimization file), this setting is
								//  relevant. It allows you to obtain a histogram of how many spikes are 
								//  transferred at each exchange. That knowledge can then be used to set the 
								//  spike compression settings appropriately.
								// When running a significantly changed network or running it using a
								//  different supercomputer or configuration, you may want to generate a new
								//  histogram to get a sense of the new spike exchange structure so you can
								//  again find the most efficient setting for the spike compression
	setupSpikeHistogram()
}
															
{load_file("./setupfiles/sim_execution_functions.hoc")}		// Defines a procedure to run the simulation and another one
															//  to periodically check how long the simulation is taking
															//  so as to end it early enough to write out results if
															//  there is limited time on the supercomputer

zzz = mallinfo(zzz, "Before running simulation")	// Get memory usage before running simulation

{rrun()}	// Run the network simulation using a custom run procedure and write out results

zzz = mallinfo(zzz, "After running simulation")		// Get memory usage after running simulation

if (pc.id==0) {			// Close the files used to record the memory usage as no more data points will be written
	{memfile.close()}
	{topfile.close()}
}

if (pc.id==0 && get_spike_hist==1) {	// If a spike histogram was generated, print it out.
	printSpikeHistogram()
}

// New quit commands, for use with NEURON development version 91f44d14afd5 or higher.
//  Use these commands with this NEURON version 91f44d14afd5 or higher if the original
//  quit commands don't stop the code when you run the program on a supercomputer. This
//  NEURON version, according to Michael Hines, "allows one to safely quit without a
//  prior {pc.runworker() pc.done()}".

{pc.barrier}
{quit()}

// Original quit commands, which usually work on most computers and supercomputers:
/*{pc.runworker()} 	// Everything after this line is executed only by the host node
					//  The NEURON website describes this as "The master host returns immediately. Worker hosts
					//  start an infinite loop of requesting tasks for execution." 
					
{pc.done()}			// Sends the quit message to the worker processors, which then quit NEURON

quit()	// Sends the quit message to the host processor, which then quits NEURON
*/

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