Clopath neuron models

class aeif_psc_delta_clopath : public Clopath_Archiving_Node
#include <aeif_psc_delta_clopath.h>

Name: aeif_psc_delta_clopath - Exponential integrate-and-fire neuron model according to Clopath et al. (2010).

Description:

aeif_psc_delta_clopath is an implementation of the neuron model as it is used in [1]. It is an extension of the aeif_psc_delta model and capable of connecting to a Clopath synapse.

Note that there are two points that are not mentioned in the paper but present in a MATLAB implementation by Claudia Clopath [3]. The first one is the clamping of the membrane potential to a fixed value after a spike occured to mimik a real spike and not just the upswing. This is important since the finite duration of the spike influences the evolution of the convolved versions (u_bar_[plus/minus]) of the membrane potential and thus the change of the synaptic weight. Secondly, there is a delay with which u_bar_[plus/minus] are used to compute the change of the synaptic weight.

Parameters:

The following parameters can be set in the status dictionary.

Dynamic state variables

V_m

mV

Membrane potential

w

pA

Spike-adaptation current

z

pA

Spike-adaptation current

V_th

mV

Adaptive spike initiation threshold

u_bar_plus

mV

Low-pass filtered Membrane potential

u_bar_minus

mV

Low-pass filtered Membrane potential

u_bar_bar

mV

Low-pass filtered u_bar_minus

Membrane Parameters

C_m

pF

Capacity of the membrane

t_ref

ms

Duration of refractory period

V_reset

mV

Reset value for V_m after a spike

E_L

mV

Leak reversal potential

g_L

nS

Leak conductance

I_e

pA

Constant external input current

tau_plus

ms

Time constant of u_bar_plus

tau_minus

ms

Time constant of u_bar_minus

tau_bar_bar

ms

Time constant of u_bar_bar

Spike adaptation parameters

a

nS

Subthreshold adaptation

b

pA

Spike-triggered adaptation

Delta_T

mV

Slope factor

tau_w

ms

Adaptation time constant

V_peak

mV

Spike detection threshold

V_th_max

mV

Value of V_th afer a spike

V_th_rest

mV

Resting value of V_th

Clopath rule parameters

A_LTD

1/mV

Amplitude of depression

A_LTP

1/mV^2

Amplitude of facilitation

theta_plus

mV

Threshold for u

theta_minus

mV

Threshold for u_bar_[plus/minus]

A_LTD_const

boolean

Flag that indicates whether A_LTD_ should be constant (true, default) or multiplied by u_bar_bar^2 / u_ref_squared (false).

delay_u_bars

real

Delay with which u_bar_[plus/minus] are processed to compute the synaptic weights.

U_ref_squared

real

Reference value for u_bar_bar_^2.

Other parameters

t_clamp

ms

Duration of clamping of Membrane potential after a spike

V_clamp

mV

Value to which the Membrane potential is clamped

Integration parameters

gsl_error_tol

real

This parameter controls the admissible error of the GSL integrator. Reduce it if NEST complains about numerical instabilities.

Note:

Neither the clamping nor the delayed processing of u_bar_[plus/minus] are mentioned in [1]. However, they are part of an reference implementation by Claudia Clopath et al. that can be found on ModelDB [3]. The clamping is important to mimic a spike which is otherwise not described by the aeif neuron model.

Author: Jonas Stapmanns, David Dahmen, Jan Hahne

Sends: SpikeEvent

Receives: SpikeEvent, CurrentEvent, DataLoggingRequest

References:

1

Clopath et al. (2010). Connectivity reflects coding: a model of voltage-based STDP with homeostasis. Nature Neuroscience 13(3):344-352. DOI: https://doi.org/10.1038/nn.2479

2

Clopath and Gerstner (2010). Voltage and spike timing interact in STDP – a unified model. Frontiers in Synaptic Neuroscience. 2:25 DOI: https://doi.org/10.3389/fnsyn.2010.00025

3

Voltage-based STDP synapse (Clopath et al. 2010) on ModelDB https://senselab.med.yale.edu/ModelDB/showmodel.cshtml?model=144566&file=%2f modeldb_package%2fVoTriCode%2faEIF.m

SeeAlso: aeif_psc_delta, clopath_synapse, hh_psc_alpha_clopath

class hh_psc_alpha_clopath : public Clopath_Archiving_Node
#include <hh_psc_alpha_clopath.h>

Name: hh_psc_alpha_clopath - Hodgkin-Huxley neuron model with support for the Clopath synapse.

Description:

hh_psc_alpha_clopath is an implementation of a spiking neuron using the Hodgkin-Huxley formalism and that is capable of connecting to a Clopath synapse.

(1) Post-synaptic currents Incoming spike events induce a post-synaptic change of current modelled by an alpha function. The alpha function is normalised such that an event of weight 1.0 results in a peak current of 1 pA.

(2) Spike Detection Spike detection is done by a combined threshold-and-local-maximum search: if there is a local maximum above a certain threshold of the membrane potential, it is considered a spike.

Parameters:

The following parameters can be set in the status dictionary.

Dynamic state variables

V_m

mV

Membrane potential

u_bar_plus

mV

Low-pass filtered Membrane potential

u_bar_minus

mV

Low-pass filtered Membrane potential

u_bar_bar

mV

Low-pass filtered u_bar_minus

Membrane Parameters

E_L

mV

Leak reversal potential

C_m

pF

Capacity of the membrane

g_L

nS

Leak conductance

tau_ex

ms

Rise time of the excitatory synaptic alpha function

tau_in

ms

Rise time of the inhibitory synaptic alpha function

E_Na

mV

Sodium reversal potential

g_Na

nS

Sodium peak conductance

E_K

mV

Potassium reversal potential

g_K

nS

Potassium peak conductance

Act_m

real

Activation variable m

Inact_h

real

Inactivation variable h

Act_n

real

Activation variable n

I_e

pA

External input current

Clopath rule parameters

A_LTD

1/mV

Amplitude of depression

A_LTP

1/mV^2

Amplitude of facilitation

theta_plus

mV

Threshold for u

theta_minus

mV

Threshold for u_bar_[plus/minus]

A_LTD_const

boolean

Flag that indicates whether A_LTD_ should be constant (true, default) or multiplied by u_bar_bar^2 / u_ref_squared (false).

delay_u_bars

real

Delay with which u_bar_[plus/minus] are processed to compute the synaptic weights.

U_ref_squared

real

Reference value for u_bar_bar_^2.

Problems/Todo:

better spike detection initial wavelet/spike at simulation onset

References:

1

Gerstner W and Kistler WM (2002). Spiking neuron models: Single neurons, populations, plasticity. New York: Cambridge university press.

2

Dayan P and Abbott L (2001). Theoretical Neuroscience: Computational and Mathematical Modeling of Neural Systems. Cambridge, MA: MIT Press. https://pure.mpg.de/pubman/faces/ViewItemOverviewPage.jsp?itemId=item_3006127

3

Hodgkin AL and Huxley A F (1952). A quantitative description of membrane current and its application to conduction and excitation in nerve. The Journal of Physiology 117. DOI: https://doi.org/10.1113/jphysiol.1952.sp004764

4

Clopath et al. (2010). Connectivity reflects coding: a model of voltage-based STDP with homeostasis. Nature Neuroscience 13(3):344-352. DOI: https://doi.org/10.1038/nn.2479

5

Clopath and Gerstner (2010). Voltage and spike timing interact in STDP – a unified model. Frontiers in Synaptic Neuroscience. 2:25 DOI: https://doi.org/10.3389/fnsyn.2010.00025

6

Voltage-based STDP synapse (Clopath et al. 2010) connected to a Hodgkin-Huxley neuron on ModelDB: https://senselab.med.yale.edu/ModelDB/showmodel.cshtml?model=144566&file =%2fmodeldb_package%2fstdp_cc.mod

Sends: SpikeEvent

Receives: SpikeEvent, CurrentEvent, DataLoggingRequest

Author: Jonas Stapmanns, David Dahmen, Jan Hahne (adapted from hh_psc_alpha by Schrader)

SeeAlso: hh_psc_alpha, clopath_synapse, aeif_psc_delta_clopath