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Update documentation for iaf_psc_exp #3084

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135 changes: 98 additions & 37 deletions models/iaf_psc_exp.h
Original file line number Diff line number Diff line change
Expand Up @@ -34,43 +34,93 @@

namespace nest
{

// Disable clang-formatting for documentation due to over-wide table.
// clang-format off
/* BeginUserDocs: neuron, integrate-and-fire, current-based

Short description
+++++++++++++++++

Leaky integrate-and-fire neuron model with exponential PSCs
Leaky integrate-and-fire neuron model with exponential shaped input currents.
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Suggested change
Leaky integrate-and-fire neuron model with exponential shaped input currents.
Leaky integrate-and-fire neuron model with exponential-shaped input currents


Description
+++++++++++

``iaf_psc_exp`` is an implementation of a leaky integrate-and-fire model
with exponential shaped postsynaptic currents (PSCs) according to [1]_.
Thus, postsynaptic currents have an infinitely short rise time.
``iaf_psc_exp`` a leaky integrate-and-fire model with

* a hard threshold,
* a fixed refractory period,
* no adaptation mechanisms,
* exponential shaped synaptic input currents according to [1]_
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* exponential shaped synaptic input currents according to [1]_
* exponential-shaped synaptic input currents according to [1]_.


Membrane potential evolution, spike emission, and refractoriness
................................................................

The membrane potential evolves according to

.. math::

\frac{dV_\text{m}}{dt} = -\frac{V_{\text{m}} - E_\text{L}}{\tau_{\text{m}}} + \frac{I_{\text{syn}} + I_\text{e}}{C_{\text{m}}}

where the synaptic input current :math:`I_{\text{syn}}(t)` is discussed below and :math:`I_\text{e}` is
a constant input current set as a model parameter.

A spike is emitted at time step :math:`t^*=t_{k+1}` if

.. math::

V_\text{m}(t_k) < V_{th} \quad\text{and}\quad V_\text{m}(t_{k+1})\geq V_\text{th} \;.

Subsequently,

.. math::

V_\text{m}(t) = V_{\text{reset}} \quad\text{for}\quad t^* \leq t < t^* + t_{\text{ref}} \;,

that is, the membrane potential is clamped to :math:`V_{\text{reset}}` during the refractory period.

The threshold crossing is followed by an absolute refractory period (``t_ref``)
during which the membrane potential is clamped to the resting potential
and spiking is prohibited.
Synaptic input
..............

The neuron dynamics is solved on the time grid given by the computation step
size. Incoming as well as emitted spikes are forced to that grid.
The synaptic input current has an excitatory and an inhibitory component

The linear subthreshold dynamics is integrated by the Exact
Integration scheme [2]_, which is more precise, but different from the
implementation in [1]_, which uses the forward Euler integration scheme.
This precludes an exact numerical reproduction of the results from [1]_.
.. math::

An additional state variable and the corresponding differential
equation represents a piecewise constant external current.
I_{\text{syn}}(t) = I_{\text{syn, ex}}(t) + I_{\text{syn, in}}(t)

The general framework for the consistent formulation of systems with
neuron like dynamics interacting by point events is described in
[2]_. A flow chart can be found in [3]_.
where

Spiking in this model can be either deterministic (delta=0) or stochastic (delta
> 0). In the stochastic case this model implements a type of spike response
model with escape noise [4]_.
.. math::

I_{\text{syn, X}}(t) = \sum_{j} w_j \sum_k i_{\text{syn, X}}(t-t_j^k-d_j) \;,

where :math:`j` indexes either excitatory (:math:`\text{X} = \text{ex}`)
or inhibitory (:math:`\text{X} = \text{in}`) presynaptic neurons,
:math:`k` indexes the spike times of neuron :math:`j`, and :math:`d_j`
is the delay from neuron :math:`j`.

The individual post-synaptic currents (PSCs) are given by

.. math::

i_{\text{syn, X}}(t) = e^{-\frac{t}{\tau_{\text{syn, X}}}} \Theta(t)
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I would include the weight w here and in the following equations to ensure that units work out, in the same corresponding way as for iaf_psc_delta.


where :math:`\Theta(x)` is the Heaviside step function. The PSCs are normalized to unit maximum, that is,

.. math::

i_{\text{syn, X}}(t= 0) = 1 \;.

As a consequence, the total charge :math:`q` transferred by a single PSC depends
on the synaptic time constant according to

.. math::

q = \int_0^{\infty} i_{\text{syn, X}}(t) dt = \tau_{\text{syn, X}} \;.

By default, :math:`V_\text{m}` is not bounded from below. To limit
hyperpolarization to biophysically plausible values, set parameter
:math:`V_{\text{min}}` as lower bound of :math:`V_\text{m}`.
Comment on lines +121 to +123
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iaf_psc_exp does not seem to have this ...

Suggested change
By default, :math:`V_\text{m}` is not bounded from below. To limit
hyperpolarization to biophysically plausible values, set parameter
:math:`V_{\text{min}}` as lower bound of :math:`V_\text{m}`.


.. note::

Expand Down Expand Up @@ -108,21 +158,32 @@ Parameters

The following parameters can be set in the status dictionary.

=========== ======= ========================================================
E_L mV Resting membrane potential
C_m pF Capacity of the membrane
tau_m ms Membrane time constant
tau_syn_ex ms Exponential decay time constant of excitatory synaptic
current kernel
tau_syn_in ms Exponential decay time constant of inhibitory synaptic
current kernel
t_ref ms Duration of refractory period (V_m = V_reset)
V_m mV Membrane potential in mV
V_th mV Spike threshold in mV
V_reset mV Reset membrane potential after a spike
I_e pA Constant input current
t_spike ms Point in time of last spike
=========== ======= ========================================================

=============== ================== =============================== ========================================================================
**Parameter** **Default** **Math equivalent** **Description**
=============== ================== =============================== ========================================================================
``E_L`` -70 mV :math:`E_\text{L}` Resting membrane potential
``C_m`` 250 pF :math:`C_{\text{m}}` Capacity of the membrane
``tau_m`` 10 ms :math:`\tau_{\text{m}}` Membrane time constant
``t_ref`` 2 ms :math:`t_{\text{ref}}` Duration of refractory period
``V_th`` -55 mV :math:`V_{\text{th}}` Spike threshold
``V_reset`` -70 mV :math:`V_{\text{reset}}` Reset potential of the membrane
``tau_syn_ex`` 2 ms :math:`\tau_{\text{syn, ex}}` Rise time of the excitatory synaptic alpha function
``tau_syn_in`` 2 ms :math:`\tau_{\text{syn, in}}` Rise time of the inhibitory synaptic alpha function
``I_e`` 0 pA :math:`I_\text{e}` Constant input current
``V_min`` :math:`-\infty` mV :math:`V_{\text{min}}` Absolute lower value for the membrane potential
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A bit surprising, but iaf_psc_exp does not seem to have V_min. Adding it would be a different issue.

Suggested change
``V_min`` :math:`-\infty` mV :math:`V_{\text{min}}` Absolute lower value for the membrane potential

=============== ================== =============================== ========================================================================

The following state variables evolve during simulation and are available either as neuron properties or as recordables.

================== ================= ========================== =================================
**State variable** **Initial value** **Math equivalent** **Description**
================== ================= ========================== =================================
``V_m`` -70 mV :math:`V_{\text{m}}` Membrane potential
``I_syn_ex`` 0 pA :math:`I_{\text{syn, ex}}` Excitatory synaptic input current
``I_syn_in`` 0 pA :math:`I_{\text{syn, in}}` Inhibitory synaptic input current
================== ================= ========================== =================================


References
++++++++++
Expand Down