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specific_ion_channels.h #ifndef __SPECIFIC_ION_CHANNELS_H_ #define __SPECIFIC_ION_CHANNELS_H_ #include "viongate.h" #include "conciongate.h" #include "activechannel.h" #include "activecachannel.h" #include "csimerror.h" // parameters for voltage range transformations #define V_REST_BIOL_MV (-70) // [mV] (general in publications used unit) #define V_THRESH_BIOL_MV (-40) // [mV] #define V_REST_SIM (0.0) // [V] (all simulation parameters are in SI units) #define V_THRESH_SIM (15e-3) // [V] #define A_MEMBRANE (0.01e-2) // [cm2] /* Transform biological Erev value to model voltage range */ /* Note that in the ActiveChannel constructor actual Neuron voltages are unknown */ #define TRANSFORM_EREV Erev = ((Erev - V_REST_BIOL_MV)*(V_THRESH_SIM - V_REST_SIM)/(V_THRESH_BIOL_MV-V_REST_BIOL_MV) + V_REST_SIM); #define TRANSFORM_V /* Transform model voltage value to biological voltage range */ \ V = (V - *Vresting)*(V_THRESH_BIOL_MV - V_REST_BIOL_MV)/(*VmScale) + V_REST_BIOL_MV #define TSCALE 1000 #define VSCALE 1000 //============================================================ #define CASCALE 1.1180e+07 #define IONGATE_CA_MIN 0.0 #define IONGATE_CA_MAX 1000e-9 #define IONGATE_CA_INC 5.0e-10 #define IONGATE_CA_TABLE_SIZE ((int)((IONGATE_CA_MAX - IONGATE_CA_MIN) / IONGATE_CA_INC + 1)) class CaGate_Yamada98 : public VIonGate { DO_REGISTERING public: CaGate_Yamada98(void) { k=2; Ca=0; Ts=1; C1=0; C2=0; } virtual ~CaGate_Yamada98(void) { if (getC1()) { free(getC1()); setC1(0); } if (getC2()) { free(getC2()); setC2(0); }} virtual double tau(double C) { return Ts/( pow(CASCALE*C,2) + 2.5); } virtual double infty(double C) { return pow(CASCALE*C,2)/( pow(CASCALE*C,2) + 2.5); } virtual double pInfty(MembranePatchSimple *) { return 0;} virtual void reset(void); virtual void setCa(double *C) { Ca = C; } virtual int updateInternal(void); virtual int advance(void); virtual int addIncoming(Advancable *Incoming); double Ts; protected: double *Ca; virtual double *getC1(void) { return C1; } virtual double *getC2(void) { return C2; } virtual void setC1(double *p) { C1=p; } virtual void setC2(double *p) { C2=p; } double *C1; double *C2; }; //== Channels used by James Maciokas / Reno ========================================================== class CaChannel_Yamada98 : public ActiveChannel { DO_REGISTERING public: CaChannel_Yamada98(void) { Erev = -5e-6 * VSCALE; // assumed resting potential Vresting = 0 u = 200e-9; // Mol Ts = 70e-3; // Sec Ca = 0; // Mol Gbar = 54e-9; // S } virtual void membraneSpikeNotify(double ) { Ca = Ca + u; }; virtual void reset(void); virtual int updateInternal(void); virtual int advance(void); float u; float Ts; double Ca; protected: float C1; }; class MmGate_Wang98 : public VIonGate { public: MmGate_Wang98(void) { k=1; } double tau(double V) { return TSCALE * exp(+( VSCALE * (V - *Vresting) - 12.3)/9.15) / 3800; } double infty(double V) { return 1/(1 + exp(-( VSCALE * (V - *Vresting) - 13)/4.4)); } IONGATE_TABLES(MmGate_Wang98); }; class MChannel_Wang98 : public ActiveChannel { DO_REGISTERING public: MChannel_Wang98(void) { addGate(new MmGate_Wang98); Erev = -80; // [mV] biological value // Transform to model voltage range TRANSFORM_EREV; } virtual ~MChannel_Wang98(void) { for(int i=0;i<nGates;i++) delete gates[i]; } }; class AmGate_Hoffman97 : public VIonGate { public: AmGate_Hoffman97(void) { k=1; } double tau(double V){ return TSCALE*0.2e-6; } double infty(double V) { TRANSFORM_V; return 1/(1+exp(-(V-11)/18)); } /* Directly transformed parameters double tau(double ) { return TSCALE*0.2e-6; } double infty(double V) { return 1/(1 + exp(-( VSCALE * (V - *Vresting) - 40.5)/9)); } */ IONGATE_TABLES(AmGate_Hoffman97); }; class AhGate_Hoffman97 : public VIonGate { public: AhGate_Hoffman97(void) { k=1; } double tau(double V) { TRANSFORM_V; return (V < -20) ? 5e-3 : 1e-3 * (5+2.6*(V+20)/10); } double infty(double V) { TRANSFORM_V; return 1/(1+exp((V+56)/8)); } /* Directly transformed parameters double tau(double ) { return TSCALE*5e-6; } double infty(double V) { return 1/(1 + exp(( VSCALE * (V - *Vresting) - 7)/4)); } */ IONGATE_TABLES(AhGate_Hoffman97); }; class AChannel_Hoffman97 : public ActiveChannel { DO_REGISTERING public: AChannel_Hoffman97(void) { addGate(new AmGate_Hoffman97); addGate(new AhGate_Hoffman97); Erev = -80; // [mV] biological value // Transform to model voltage range TRANSFORM_EREV; } virtual ~AChannel_Hoffman97(void) { for(int i=0;i<nGates;i++) delete gates[i]; } }; class SICmGate_Maciokas02 : public VIonGate { public: SICmGate_Maciokas02(void) { k=1; } double tau(double ) { return TSCALE*0.2e-6; } double infty(double V) { return 1/(1 + exp(-( VSCALE * (V - *Vresting) - 20)/2.5)); } IONGATE_TABLES(SICmGate_Maciokas02); }; class SIChGate_Maciokas02 : public VIonGate { public: SIChGate_Maciokas02(void) { k=1; } double tau(double ) { return TSCALE*5e-6; } double infty(double V) { return 1/(1 + exp(( VSCALE * (V - *Vresting) - 15)/2.5)); } IONGATE_TABLES(SIChGate_Maciokas02); }; class SICChannel_Maciokas02 : public ActiveChannel { DO_REGISTERING public: SICChannel_Maciokas02(void) { addGate(new SICmGate_Maciokas02); addGate(new SIChGate_Maciokas02); Erev = -80; // [mV] biological value // Transform to model voltage range TRANSFORM_EREV; } virtual ~SICChannel_Maciokas02(void) { for(int i=0;i<nGates;i++) delete gates[i]; } }; //== Channels of neocortical pyramidal neurons from the SenseLab ModelDB archiv ======== class AnGate_Korngreen02 : public VIonGate { public: AnGate_Korngreen02(void) { k=2;} double tau(double V) { TRANSFORM_V; return (V < -50) ? 1e-3 * (1.25+175.03*exp(V*0.026)) : 1e-3 * (1.25+13*exp(-V*0.026)); } double infty(double V) { TRANSFORM_V; return 1/(1+exp(-(V+14)/14.6)); } IONGATE_TABLES(AnGate_Korngreen02); }; class AlGate_Korngreen02 : public VIonGate { public: AlGate_Korngreen02(void) { k=1; } double tau(double V) { TRANSFORM_V; return 1e-3 * (360+(1010+24*(V+55))*exp(-pow((V+75)/48,2))); } double infty(double V) { TRANSFORM_V; return 1/(1+exp((V+54)/11)); } IONGATE_TABLES(AlGate_Korngreen02); }; class AChannel_Korngreen02 : public ActiveChannel { DO_REGISTERING public: AChannel_Korngreen02(void) { addGate(new AnGate_Korngreen02); addGate(new AlGate_Korngreen02); Ts = 1; Gbar = 8e-3*A_MEMBRANE; // [S/cm2]*[cm2] Erev = -80; // (original -90) [mV] biological value // Transform biological Erev value to model voltage range TRANSFORM_EREV; } virtual ~AChannel_Korngreen02(void) { for(int i=0;i<nGates;i++) delete gates[i]; } virtual int updateInternal(void); double Ts; }; class KnGate_Korngreen02 : public VIonGate { public: KnGate_Korngreen02(void) { k=4;} double tau(double V) { TRANSFORM_V; return 1e-3 * (0.34 + 0.92*exp(-pow((V+71)/59,2))); } double infty(double V) { TRANSFORM_V; return 1/(1+exp(-(V+47)/29)); } IONGATE_TABLES(KnGate_Korngreen02); }; class KlGate_Korngreen02 : public VIonGate { public: KlGate_Korngreen02(void) { k=1; } double tau(double V) { TRANSFORM_V; return 1e-3 * (8 + 49*exp(-pow((V+73)/23,2))); } double infty(double V) { TRANSFORM_V; return 1/(1+exp((V+66)/10)); } IONGATE_TABLES(KlGate_Korngreen02); }; class KChannel_Korngreen02 : public ActiveChannel { DO_REGISTERING public: KChannel_Korngreen02(void) { addGate(new KnGate_Korngreen02); addGate(new KlGate_Korngreen02); Ts = 1; Gbar = 8e-3*A_MEMBRANE; // [S/cm2]*[cm2] Erev = -80; // (original -90) [mV] biological value // Transform biological Erev value to model voltage range TRANSFORM_EREV; } virtual ~KChannel_Korngreen02(void) { for(int i=0;i<nGates;i++) delete gates[i]; } virtual int updateInternal(void); double Ts; }; class NPmGate_McCormick92 : public VIonGate { public: NPmGate_McCormick92(void) { k=1; } double alpha(double V) { TRANSFORM_V; return 1e3 * (0.091*(V+48)/(1.0-exp(-(V+48.0)/5.0))); } double beta(double V) { TRANSFORM_V; return 1e3 * (-0.062*(V+48.0)/(1.0-exp((V+48.0)/5.0))); } double infty(double V) { TRANSFORM_V; return 1.0/(1.0+exp(-(49.0+V)/5)); } IONGATE_TABLES(NPmGate_McCormick92); }; class NPChannel_McCormick02 : public ActiveChannel { DO_REGISTERING public: NPChannel_McCormick02(void) { addGate(new NPmGate_McCormick92); Ts = 1; Gbar = 2.2e-3*A_MEMBRANE; // [S/cm2]*[cm2] Erev = 55.0; // [mV] biological value // Transform biological Erev value to model voltage range TRANSFORM_EREV; } virtual ~NPChannel_McCormick02(void) { for(int i=0;i<nGates;i++) delete gates[i]; } virtual int updateInternal(void); double Ts; }; class MnGate_Mainen96 : public VIonGate { public: MnGate_Mainen96(void) { k=1; } double alpha(double V) { TRANSFORM_V; return 1e3 * (0.001*(V+30)/(1-exp(-(V+30)/9))); } double beta(double V) { TRANSFORM_V; return 1e3 * (-0.001*(V+30)/(1-exp((V+30)/9))); } IONGATE_TABLES(MnGate_Mainen96); }; class MChannel_Mainen96 : public ActiveChannel { DO_REGISTERING public: MChannel_Mainen96(void) { addGate(new MnGate_Mainen96); Ts = 1; Gbar = 1e-3*A_MEMBRANE; // [S/cm2]*[cm2] Erev = -80.0; // (original -90) [mV] biological value // Transform biological Erev value to model voltage range TRANSFORM_EREV; } virtual ~MChannel_Mainen96(void) { for(int i=0;i<nGates;i++) delete gates[i]; } virtual int updateInternal(void); double Ts; }; #define CELSIUS 30 class HnGate_Stuart98 : public VIonGate { public: HnGate_Stuart98(void) { k=1; } double alpha(double V) { return exp(1e-3*3*(V+88)*9.648e4/(8.315*(273.16+CELSIUS))); } double beta(double V) { return exp(1e-3*3*0.4*(V+88)*9.648e4/(8.315*(273.16+CELSIUS))); } double tau(double V) { TRANSFORM_V; return 1e-3 * (beta(V)/(0.00057+0.00057*alpha(V))); } double infty(double V) { TRANSFORM_V; return 1/(1+alpha(V)); } IONGATE_TABLES(HnGate_Stuart98); }; class HChannel_Stuart98 : public ActiveChannel { DO_REGISTERING public: HChannel_Stuart98(void) { addGate(new HnGate_Stuart98); Ts = 1; Gbar = 3e-3*A_MEMBRANE; // [S/cm2]*[cm2] Erev = -35.0; // [mV] biological value // Transform biological Erev value to model voltage range TRANSFORM_EREV; } virtual ~HChannel_Stuart98(void) { for(int i=0;i<nGates;i++) delete gates[i]; } virtual int updateInternal(void); double Ts; }; #define SECH(_V_) 2/(exp(2*_V_)+exp(-2*_V_)) class HVACAuGate_Brown93 : public VIonGate { public: HVACAuGate_Brown93(void) { k=2;} double tau(double V) { TRANSFORM_V; return 1e-3 * (1.25*SECH(-0.031*(V+37.1))); } double infty(double V) { TRANSFORM_V; return 1/(1+exp(-(V+24.6)/11.3)); } IONGATE_TABLES(HVACAuGate_Brown93); }; class HVACAvGate_Brown93 : public VIonGate { public: HVACAvGate_Brown93(void) { k=1; } double tau(double V) { TRANSFORM_V; return 1e-3 * 420; } double infty(double V) { TRANSFORM_V; return 1/(1+exp((V+12.6)/18.9)); } IONGATE_TABLES(HVACAvGate_Brown93); }; class HVACAChannel_Brown93 : public ActiveCaChannel { DO_REGISTERING public: HVACAChannel_Brown93(void) { addGate(new HVACAuGate_Brown93); addGate(new HVACAvGate_Brown93); Ts = 1; Gbar = 0.3e-3*A_MEMBRANE; // [S/cm2]*[cm2] Erev = 140; // [mV] biological value // Transform biological Erev value to model voltage range TRANSFORM_EREV; } virtual ~HVACAChannel_Brown93(void) { for(int i=0;i<nGates;i++) delete gates[i]; } virtual int updateInternal(void); double Ts; }; class CALmGate_Destexhe98 : public VIonGate { public: CALmGate_Destexhe98(void) { k=2;} double tau(double V) { TRANSFORM_V; return 1e-3 * 0.1; } /* Originally 0 */ double infty(double V) { TRANSFORM_V; return 1.0/(1+exp(-(V+57)/6.2)); } IONGATE_TABLES(CALmGate_Destexhe98); }; class CALhGate_Destexhe98 : public VIonGate { public: CALhGate_Destexhe98(void) { k=1; } double tau(double V) { TRANSFORM_V; return 1e-3 * (30.8 + (211.4 + exp((V+113.2)/5))/(1+exp((V+84)/3.2))); } double infty(double V) { TRANSFORM_V; return 1.0/(1+exp((V+81)/4.0)); } IONGATE_TABLES(CALhGate_Destexhe98); }; class CALChannel_Destexhe98 : public ActiveChannel { DO_REGISTERING public: CALChannel_Destexhe98(void) { addGate(new CALmGate_Destexhe98); addGate(new CALhGate_Destexhe98); Ts = 1; Gbar = 2e-3*A_MEMBRANE; // [S/cm2]*[cm2] Erev = 140; // [mV] biological value // Transform biological Erev value to model voltage range TRANSFORM_EREV; } virtual ~CALChannel_Destexhe98(void) { for(int i=0;i<nGates;i++) delete gates[i]; } virtual int updateInternal(void); double Ts; }; class KCAnGate_Mainen96 : public ConcIonGate { public: KCAnGate_Mainen96(void) { k=1; ConcType=CA;} double alpha(double C) { return 0.01*C*1e3; } double beta(double ) { return 0.02; } double tau(double C) { return 1e-3/(alpha(C) + beta(C)); } double infty(double C) { return alpha(C)/(alpha(C) + beta(C)); } IONGATE_TABLES(KCAnGate_Mainen96); }; class KCAChannel_Mainen96 : public ActiveChannel { DO_REGISTERING public: KCAChannel_Mainen96(void) { addGate(new KCAnGate_Mainen96); Ts = 1; Gbar = 1e-3*A_MEMBRANE; // [S/cm2]*[cm2] Erev = -80; // (original -90) [mV] biological value // Transform biological Erev value to model voltage range TRANSFORM_EREV; } virtual ~KCAChannel_Mainen96(void) { for(int i=0;i<nGates;i++) delete gates[i]; } virtual int updateInternal(void); double Ts; }; //== Channels used by Destexhe take from Traub et al. and Mainen et al. ========================================================== #define Amembrane 34636e-12 // [m2] // #define NUMMETHOD 0 #define NUMMETHOD 1 // #define GSCALE 0.4 #define GSCALE 1 #define ABSCALE 1e3 #define TADJM 2.9529 // = 2.3 ^ ((CELSIUS-23)/10), CELSIUS = 36 #define VT -63 + 1e-5 // [mV], -58 mV needed for Vthresh adjustment // #define VT -58 + 1e-5 // [mV], -58 mV needed for Vthresh adjustment class NAmGate_Traub91 : public VIonGate { public: NAmGate_Traub91(void) { k=3; nummethod = NUMMETHOD;} double alpha(double V) { V = V*1e3; return ABSCALE*-0.32 * (V - VT - 13)/(exp(-(V-VT-13)/4)-1); } double beta(double V) { V = V*1e3; return ABSCALE*0.28*(V-VT-40)/(exp((V-VT-40)/5)-1); } IONGATE_TABLES(NAmGate_Traub91); }; class NAhGate_Traub91 : public VIonGate { public: NAhGate_Traub91(void) { k=1; nummethod = NUMMETHOD;} double alpha(double V) { V = V*1e3; return ABSCALE*0.128 * exp(-(V-VT+10-17)/18); } double beta(double V) { V = V*1e3; return ABSCALE*4 / (1+exp(-(V-VT+10-40)/5)); } IONGATE_TABLES(NAhGate_Traub91); }; class NAChannel_Traub91 : public ActiveChannel { DO_REGISTERING public: NAChannel_Traub91(void) { addGate(new NAmGate_Traub91); addGate(new NAhGate_Traub91); Ts = 1; // Gbar = GSCALE * 30 * Amembrane; // [S/m2]*[m2] // Gbar = GSCALE * 360 * Amembrane; // [S/m2]*[m2] // Gbar = GSCALE * 120 * Amembrane; // [S/m2]*[m2] Gbar = 4.3* GSCALE * 120 * Amembrane; // [S/m2]*[m2] // Gbar = 1.67* GSCALE * 120 * Amembrane; // [S/m2]*[m2] Erev = 50e-3; // [mV] } virtual ~NAChannel_Traub91(void) { for(int i=0;i<nGates;i++) delete gates[i]; } virtual int updateInternal(void); double Ts; }; class KDnGate_Traub91 : public VIonGate { public: KDnGate_Traub91(void) { k=4; nummethod = NUMMETHOD;} double alpha(double V) { V = V*1e3; return ABSCALE*-0.032*(V-VT-15)/(exp(-(V-VT-15)/5)-1); } double beta(double V) { V = V*1e3; return ABSCALE*0.5*exp(-(V-VT-10)/40); } IONGATE_TABLES(KDnGate_Traub91); }; class KDChannel_Traub91 : public ActiveChannel { DO_REGISTERING public: // original model has Vrest = -80 mV, Vwork = -65 mV, Vthresh = -55 mV // shift V by +10 results in Vrest = -70 mV, Vwork = -55 mV, Vthresh = -45 mV // what is approx. the above defined biological working range needed for parameter rescaling KDChannel_Traub91(void) { addGate(new KDnGate_Traub91); Ts = 1; // Gbar = GSCALE * 50 * Amembrane; // [S/m2]*[m2] // Gbar = GSCALE * 70 * Amembrane; // [S/m2]*[m2] Gbar = GSCALE * 100 * Amembrane; // [S/m2]*[m2] Erev = -90e-3; // [mV] } virtual ~KDChannel_Traub91(void) { for(int i=0;i<nGates;i++) delete gates[i]; } virtual int updateInternal(void); double Ts; }; class MpGate_Mainen96orig : public VIonGate { public: MpGate_Mainen96orig(void) { k=1; nummethod = NUMMETHOD;} double alpha(double V) { V = V*1e3; return TADJM*ABSCALE*0.0001*(V+30)/(1-exp(-(V+30)/9)); } double beta(double V) { V = V*1e3; return TADJM*ABSCALE*-0.0001*(V+30)/(1-exp((V+30)/9)); } IONGATE_TABLES(MpGate_Mainen96orig); }; class MChannel_Mainen96orig : public ActiveChannel { DO_REGISTERING public: MChannel_Mainen96orig(void) { addGate(new MpGate_Mainen96orig); Ts = 1; // Gbar = 0 * Amembrane; // [S/m2]*[m2] Gbar = GSCALE * 5 * Amembrane; // [S/m2]*[m2] Erev = -90e-3; // [V] } virtual ~MChannel_Mainen96orig(void) { for(int i=0;i<nGates;i++) delete gates[i]; } virtual int updateInternal(void); double Ts; }; #endif

(C) 2003, Thomas Natschläger

last modified 07/10/2006