from __future__ import print_function from neuron import h from .cell import Cell # from .. import synapses from ..util import nstomho from ..util import Params import numpy as np from .. import data __all__ = ["MSO"] class MSO(Cell): type = "mso" @classmethod def create(cls, model="MSO-principal", **kwds): if model == "MSO-principal": return MSOPrincipal(**kwds) else: raise ValueError("MSO cell model %s is unknown", model) def make_psd(self, terminal, psd_type, **kwds): """ Connect a presynaptic terminal to one post section at the specified location, with the fraction of the "standard" conductance determined by gbar. The default condition is designed to pass the unit test (loc=0.5) Parameters ---------- terminal : Presynaptic terminal (NEURON object) psd_type : either simple or multisite PSD for MSO cell kwds: dictionary of options. Two are currently handled: postsite : expect a list consisting of [sectionno, location (float)] AMPAScale : float to scale the ampa currents """ if ( "postsite" in kwds ): # use a defined location instead of the default (soma(0.5) postsite = kwds["postsite"] loc = postsite[1] # where on the section? uname = ( "sections[%d]" % postsite[0] ) # make a name to look up the neuron section object post_sec = self.hr.get_section(uname) # Tell us where to put the synapse. else: loc = 0.5 post_sec = self.soma if psd_type == "simple": return self.make_exp2_psd(post_sec, terminal, loc=loc) elif psd_type == "multisite": if terminal.cell.type == "bushy": # Max conductances for the glu mechanisms are calibrated by # running `synapses/tests/test_psd.py`. The test should fail # if these values are incorrect self.AMPAR_gmax = ( data.get( "bushy_synapse", species=self.species, post_type=self.type, field="AMPAR_gmax", ) * 1e3 ) self.NMDAR_gmax = ( data.get( "bushy_synapse", species=self.species, post_type=self.type, field="NMDAR_gmax", ) * 1e3 ) self.Pr = data.get( "bushy_synapse", species=self.species, post_type=self.type, field="Pr", ) # adjust gmax to correct for initial Pr self.AMPAR_gmax = self.AMPAR_gmax / self.Pr self.NMDAR_gmax = self.NMDAR_gmax / self.Pr if "AMPAScale" in kwds: # normally, this should not be done! self.AMPAR_gmax = ( self.AMPAR_gmax * kwds["AMPAScale"] ) # allow scaling of AMPA conductances if "NMDAScale" in kwds: self.NMDAR_gmax = self.NMDAR_gmax * kwds["NMDAScale"] # and NMDA... return self.make_glu_psd( post_sec, terminal, self.AMPAR_gmax, self.NMDAR_gmax, loc=loc ) else: raise TypeError( "Cannot make PSD for %s => %s" % (terminal.cell.type, self.type) ) else: raise ValueError("Unsupported psd type %s" % psd_type) class MSOPrincipal(MSO): """ VCN MSO cell models. Using Rothman and Manis, 2003abc (Type II) MSO principal cell type """ def __init__( self, morphology=None, decorator=None, nach=None, ttx=False, species="guineapig", modelType=None, debug=False, temperature=None, ): """ Create a MSO principal cell, using the default parameters for guinea pig from R&M2003, as a type II cell. Additional modifications to the cell can be made by calling methods below. Parameters ---------- morphology : string (default: None) Name of a .hoc file representing the morphology. This file is used to constructe an electrotonic (cable) model. If None (default), then a "point" (really, single cylinder) model is made, exactly according to RM03. decorator : Python function (default: None) decorator is a function that "decorates" the morphology with ion channels according to a set of rules. If None, a default set of channels is inserted into the first soma section, and the rest of the structure is "bare". nach : string (default: None) nach selects the type of sodium channel that will be used in the model. A channel mechanism by that name must exist. The default channel is set to 'nacn' (R&M03) temperature : float (default: 22) temperature to run the cell at. ttx : Boolean (default: False) If ttx is True, then the sodium channel conductance is set to 0 everywhere in the cell. This flag duplicates the effects of tetrodotoxin in the model. Currently, the flag is not implemented. temperature : float (default: None, sets to model default of 22) temperature (deg C) to run the cell at. Must be a valid temperature for the model. species: string (default 'guineapig') species defines the pattern of ion channel densities that will be inserted, according to prior measurements in various species. Note that if a decorator function is specified, this argument is ignored as the decorator will specify the channel density. modelType: string (default: None) modelType specifies the subtype of the cell model that will be used (e.g., "II", "II-I", etc). modelType is passed to the decorator, or to species_scaling to adjust point (single cylinder) models. debug: boolean (default: False) When True, there will be multiple printouts of progress and parameters. Returns ------- Nothing """ super(MSO, self).__init__() self.i_test_range = { "pulse": (-1, 1, 0.05) } # note that this gets reset with decorator according to channels # Changing the default values will cause the unit tests to fail! if modelType == None: modelType = "principal" if nach == None and species == "guineapig": nach = "na" if nach == None and species == "mouse": nach = "na" self.i_test_range = {"pulse": (-1, 1.2, 0.05)} self.status = { "soma": True, "axon": False, "dendrites": False, "pumps": False, "hillock": False, "initialsegment": False, "myelinatedaxon": False, "unmyelinatedaxon": False, "na": nach, "species": species, "modelType": modelType, "ttx": ttx, "name": "MSO", "morphology": morphology, "decorator": decorator, "temperature": temperature, } self.spike_threshold = -40 self.vrange = [-70.0, -55.0] # set a default vrange for searching for rmp print("model type, species: ", modelType, species, nach) if morphology is None: """ instantiate a basic soma-only ("point") model """ print("<< MSO model: Creating point principal cell >>") soma = h.Section( name="MSO_Soma_%x" % id(self) ) # one compartment of about 29000 um2 soma.nseg = 1 self.add_section(soma, "soma") else: """ instantiate a structured model with the morphology as specified by the morphology file """ print( "<< MSO principal cell model: Creating cell with morphology from %s >>" % morphology ) self.set_morphology(morphology_file=morphology) # decorate the morphology with ion channels if decorator is None: # basic model, only on the soma self.mechanisms = ["klt", "kht", "ihvcn", "leak", nach] for mech in self.mechanisms: self.soma.insert(mech) self.soma.ena = self.e_na self.soma.ek = self.e_k self.soma().ihvcn.eh = self.e_h self.soma().leak.erev = self.e_leak self.c_m = 0.9 self.species_scaling( silent=True, species=species, modelType=modelType ) # set the default type II cell parameters else: # decorate according to a defined set of rules on all cell compartments self.decorate() self.save_all_mechs() # save all mechanisms inserted, location and gbar values... self.get_mechs(self.soma) if debug: print(" << Created cell >>") def get_cellpars(self, dataset, species="guineapig", celltype="principal"): cellcap = data.get( dataset, species=species, cell_type=celltype, field="soma_Cap" ) chtype = data.get( dataset, species=species, cell_type=celltype, field="soma_na_type" ) pars = Params(cap=cellcap, natype=chtype) for g in ["soma_kht_gbar", "soma_klt_gbar", "soma_ih_gbar", "soma_leak_gbar"]: pars.additem( g, data.get(dataset, species=species, cell_type=celltype, field=g) ) return pars def species_scaling(self, species="guineapig", modelType="principal", silent=True): """ Adjust all of the conductances and the cell size according to the species requested. This scaling should be used ONLY for point models, as no other compartments are scaled. This scaling routine also sets the temperature for the model to a default value. Some models can be run at multiple temperatures, and so a default from one of the temperatures is used. The calling cell.set_temperature(newtemp) will change the conductances and reinitialize the cell to the new temperature settings. Parameters ---------- species : string (default: 'guineapig') name of the species to use for scaling the conductances in the base point model Must be one of mouse, cat, guineapig modelType: string (default: 'principal') definition of model type from RM03 models, principal cell for mso silent : boolean (default: True) run silently (True) or verbosely (False) """ # print '\nSpecies scaling: %s %s' % (species, type) knownspecies = ["guineapig"] soma = self.soma if modelType == "principal": celltype = ( "MSO-principal" ) # There are other possiblities in the literature - this is just the main one else: raise ValueError("model type not recognized") if species == "guineapig": print( " Setting conductances for guinea pig %s MSO cell, based on Rothman and Manis, 2003 bushy cell" % modelType ) self._valid_temperatures = (22.0, 38.0) if self.status["temperature"] is None: self.status["temperature"] = 22.0 self.i_test_range = {"pulse": (-0.4, 0.4, 0.02)} sf = 1.0 if ( self.status["temperature"] == 38.0 ): # adjust for 2003 model conductance levels at 38 sf = 2 # Q10 of 2, 22->38C. (p3106, R&M2003c) # note that kinetics are scaled in the mod file. dataset = "MSO_principal_channels" pars = self.get_cellpars(dataset, species=species, celltype=celltype) self.set_soma_size_from_Cm(pars.cap) self.status["na"] = pars.natype self.adjust_na_chans(soma, sf=sf) soma().kht.gbar = nstomho(pars.soma_kht_gbar, self.somaarea) soma().klt.gbar = nstomho(pars.soma_klt_gbar, self.somaarea) soma().ihvcn.gbar = nstomho(pars.soma_ih_gbar, self.somaarea) soma().leak.gbar = nstomho(pars.soma_leak_gbar, self.somaarea) self.axonsf = 0.57 else: errmsg = ( 'Species "%s" or model type "%s" is not recognized for MSO cells.' % (species, modelType) ) errmsg += "\n Valid species are: \n" for s in knownspecies: errmsg += " %s\n" % s errmsg += "-" * 40 raise ValueError(errmsg) self.status["species"] = species self.status["modelType"] = modelType self.check_temperature() # self.cell_initialize(vrange=self.vrange) # no need to do this just yet. if not silent: print(" set cell as: ", species) print(" with Vm rest = %6.3f" % self.vm0) def channel_manager(self, modelType="MSO-principal"): """ This routine defines channel density maps and distance map patterns for each type of compartment in the cell. The maps are used by the ChannelDecorator class (specifically, its private \_biophys function) to decorate the cell membrane. These settings are only used if the decorator is called; otherwise for point cells, the species_scaling routine defines the channel densities. Parameters ---------- modelType : string (default: 'RM03') A string that defines the type of the model. Currently, only 1 type is implemented: RM03: Rothman and Manis, 2003 somatic densities based on guinea pig bushy cell Returns ------- Nothing Notes ----- This routine defines the following variables for the class: * conductances (gBar) * a channelMap (dictonary of channel densities in defined anatomical compartments) * a current injection range for IV's (used for testing) * a distance map, which defines how each conductance in a selected compartment changes with distance from the soma. The current implementation includes both linear and exponential gradients, the minimum conductance at the end of the gradient, and the space constant or slope for the gradient. """ self.c_m = 1e-6 # default in units of F/cm^2 if modelType == "MSO-principal": # # Create a model based on the Rothman and Manis 2003 conductance set from guinea pig # self.c_m = 0.9e-6 # default in units of F/cm^2 totcap = 12.0e-12 # in units of F, from Rothman and Manis, 2003. refarea = totcap / self.c_m # area is in cm^2 # MSO Rothman-Manis, guinea pig type II # model gave cell conductance in nS, but we want S/cm^2 for NEURON # so conversion is 1e-9*nS = uS, and refarea is already in cm2 self._valid_temperatures = (22.0, 38.0) sf = 1.0 if self.status["temperature"] == None: self.status["temperature"] = 22.0 if self.status["temperature"] == 38: sf = 3.03 self.gBar = Params( nabar=sf * 1000.0e-9 / refarea, khtbar=sf * 150.0e-9 / refarea, kltbar=sf * 200.0e-9 / refarea, ihbar=sf * 20.0e-9 / refarea, leakbar=sf * 2.0e-9 / refarea, ) print("MSO principal channels gbar:\n", self.gBar.show()) self.channelMap = { "axon": { "nacn": self.gBar.nabar, "klt": self.gBar.kltbar, "kht": self.gBar.khtbar, "ihvcn": 0.0, "leak": self.gBar.leakbar / 2.0, }, "hillock": { "nacn": self.gBar.nabar, "klt": self.gBar.kltbar, "kht": self.gBar.khtbar, "ihvcn": 0.0, "leak": self.gBar.leakbar, }, "initseg": { "nacn": self.gBar.nabar, "klt": self.gBar.kltbar, "kht": self.gBar.khtbar, "ihvcn": self.gBar.ihbar / 2.0, "leak": self.gBar.leakbar, }, "soma": { "nacn": self.gBar.nabar, "klt": self.gBar.kltbar, "kht": self.gBar.khtbar, "ihvcn": self.gBar.ihbar, "leak": self.gBar.leakbar, }, "dend": { "nacn": self.gBar.nabar, "klt": self.gBar.kltbar * 0.5, "kht": self.gBar.khtbar * 0.5, "ihvcn": self.gBar.ihbar / 3.0, "leak": self.gBar.leakbar * 0.5, }, "apic": { "nacn": self.gBar.nabar, "klt": self.gBar.kltbar * 0.2, "kht": self.gBar.khtbar * 0.2, "ihvcn": self.gBar.ihbar / 4.0, "leak": self.gBar.leakbar * 0.2, }, } # self.irange = np.linspace(-1., 1., 21) self.distMap = { "dend": { "klt": {"gradient": "linear", "gminf": 0.0, "lambda": 100.0}, "kht": {"gradient": "linear", "gminf": 0.0, "lambda": 100.0}, "nacn": {"gradient": "exp", "gminf": 0.0, "lambda": 100.0}, }, # linear with distance, gminf (factor) is multiplied by gbar "apic": { "klt": {"gradient": "linear", "gminf": 0.0, "lambda": 100.0}, "kht": {"gradient": "linear", "gminf": 0.0, "lambda": 100.0}, "nacn": {"gradient": "exp", "gminf": 0.0, "lambda": 100.0}, }, # gradients are: flat, linear, exponential } else: raise ValueError("model type %s is not implemented" % modelType) self.check_temperature() def adjust_na_chans(self, soma, sf=1.0, gbar=1000.0, debug=False): """ adjust the sodium channel conductance Parameters ---------- soma : neuron section object A soma object whose sodium channel complement will have its conductances adjusted depending on the channel type gbar : float (default: 1000.) The maximal conductance for the sodium channel debug : boolean (false): Verbose printing Returns ------- Nothing : """ if self.status["ttx"]: gnabar = 0.0 else: gnabar = nstomho(gbar, self.somaarea) * sf nach = self.status["na"] if nach == "jsrna": soma().jsrna.gbar = gnabar soma.ena = self.e_na if debug: print("jsrna gbar: ", soma().jsrna.gbar) elif nach == "nav11": soma().nav11.gbar = gnabar soma.ena = 50 # self.e_na # print('gnabar: ', soma().nav11.gbar, ' vs: 0.0192307692308') soma().nav11.vsna = 4.3 if debug: print("MSO using inva11") elif nach in ["na", "nacn"]: soma().na.gbar = gnabar soma.ena = self.e_na if debug: print("na gbar: ", soma().na.gbar) else: raise ValueError("Sodium channel %s is not recognized for MSO cells", nach)