Neural simulations

Overview

A simulation of a network is like an experiment with the difference that it takes place inside the computer's memory rather than in the physical world.

Like in a real experiment, you need a system which you want to investigate. Moreover, you need a clear idea of what you want to learn from the experiment. In the context of a network simulation this means that you have to know which input you want to give to your network and which output you expect.

The next section will illustrate the main concepts of NEST simulations, using a simple simulation. The following sections will then give a step-by-step introduction to the main concepts of NEST simulations. Finally, we will discuss a complex example.

A simple simulation

The simplest simulation in NEST is that of a network which contains just one neuron:

SLI ] /iaf_psc_alpha Create /neuron Set

We are going to simulate a standard integrate and fire model with resting potential at -70 mV and spike threshold at -55 mV. In this line, we use the model iaf_psc_alpha to create a neuron. The command Create returns a handle to the created neuron, which we store in the variable neuron.

Next, we would like to add a stimulation and a recording device to the neuron, so that we will see something during the simulation.

In our little example, we want to inject a current into the neuron and record its membrane potential. Thus, we need to create the corresponding devices and connect them to the neuron:

SLI ] /voltmeter Create  /vm Set

This command creates a voltmeter node. The handle to the new voltmeter is stored in the variable vm.

By default, the voltmeter will only record membrane potential values, so we configure it to show us the time stamp of each value as well. We also set it to print the time and potential to the screen and we set the recording interval to 0.1 ms. The default is 1.0 ms.

SLI ] vm << /withtime true /to_screen true /interval 0.1 >> SetStatus

The double angled brackets << and >>delimit a dictionary definition which consists of successive /key value pairs.

After setting the parameters of the voltmeter, we connect it to the neuron, using the handles we have created above:

SLI ] vm neuron Connect

Now we create a DC generator which will supply a constant current to our neuron:

SLI ] /dc_generator Create /stim Set

Next, we want to set the amplitude of the DC generator such that it delivers enough current to elicit a spike in the neuron:

SLI ] stim << /amplitude 600.0 >> SetStatus

Here, we only adjust the strength of the input, we set its new value to 600.0, which is the amplitude in pA.

We have to connect the DC generator to the neuron:

SLI ] stim neuron Connect

We can now run the simulation and expect to see some results:

SLI ] 15.0 Simulate

The command Simulate runs the simulation for the specified number of milliseconds. Below, you see a transcript of the simulation:

Nov 10 08:57:51 Simulate [Info]:
    Simulating 15 ms.
Nov 10 08:57:51 Scheduler::prepare_nodes [Info]:
    Please wait. Preparing elements.
Nov 10 08:57:51 Scheduler::prepare_nodes [Info]:
    Simulating 3 local nodes.
Nov 10 08:57:51 Scheduler::update [Info]:
    Simulating using OpenMP.

1   0.1 -70
1   0.2 -70
1   0.3 -70
1   0.4 -70
1   0.5 -70
1   0.6 -70
1   0.7 -70
1   0.8 -70
1   0.9 -70
1   1   -70
1   1.1 -70
1   1.2 -69.7612    
1   1.3 -69.5248    
1   1.4 -69.2907    
1   1.5 -69.0589    

:

1   10.5    -55.3751    
1   10.6    -55.2818    
1   10.7    -55.1894    
1   10.8    -55.098
1   10.9    -55.0075    
1   11  -70
1   11.1    -70
1   11.2    -70
1   11.3    -70

:

1   12.7    -70
1   12.8    -70
1   12.9    -70
1   13  -70
1   13.1    -69.7612    
1   13.2    -69.5248    
1   13.3    -69.2907    
1   13.4    -69.0589    
1   13.5    -68.8295    
1   13.6    -68.6023    
1   13.7    -68.3775    
1   13.8    -68.1548    
1   13.9    -67.9343    
1   14  -67.7161    

Nov 10 08:57:51 Scheduler::resume [Info]:
    Simulation finished.
Nov 10 08:57:51 Scheduler::finalize_nodes() [Info]:
     using OpenMP.

After some initial messages from the simulation scheduler, we see the output from the voltmeter. The number in the left column represents the global ID (GID) of the model neuron and the center column the network time in milliseconds. The right column contains the values of the membrane potential at that time. The potential is given in mV.

By default, NEST uses a simulation stepsize of 0.1 ms. With a simulation time of 15.0 ms, we have 150 simulation steps.

The neuron that we have simulated was a standard integrate-and-fire neuron [Tuckwell91] with a resting potential of -70 mV and a threshold at -55.0 mV. We see the first effect of the DC input current at 1.2 ms. This time delay is due to several reasons: The dc_generator emits the first current output at the end of the first time step, i.e., at 0.1 ms. Since the connection between generator and neuron was created with the default delay of 1 ms, the current signal arrives at the neuron at 1.1 ms and thus affects the membrane potential for the first time during the time step from 1.1 ms to 1.2 ms.

During the time step from 10.9 ms to 11.0 ms, the membrane potential crosses the threshold value -55.0 mV. Thus, the neuron emits an output spike at 11.0 ms and the membrane potential is then reset to -70.0 mV and clamped to the resting value for 2 ms, the refractory period of the neuron. After the refractory period, the membrane continues to depolarize due to the continuing input current.

Nodes and Models

In NEST, the neural system is a collection of nodes and their interactions. Nodes correspond to things like neurons, synapses, and devices, and are implemented in C++. The network and its configuration are defined at the level of the simulation language interpreter.

Nodes are created from a set of prescribed models which are stored in the dictionary modeldict. The most important neuron models are:

Model name Description iaf_psc_alpha Simple integrate-and-fire neuron with alpha-function PSCs. iaf_psc_delta Integrate-and-fire neuron with delta-function PSCs. iaf_cond_alpha Conductance-based integrate-and-fire neuron with alpha-function synapses. iaf_cond_exp Conductance-based integrate-and-fire neuron with exp-function synapses. hh_psc_alpha hh_cond_exp_traub In order to make the models visible to the interpreter, the model dictionary has to be opened.

Creating nodes

Before continuing with the example, we reset NEST, to clear all nodes that we have created before.

SLI ] ResetKernel
Sep 21 10:13:39 Network::clear_models [Info]:
 Models will be cleared and parameters reset.

Nodes are created from a model, using the command Create.

SLI ] /iaf_psc_alpha Create ==
1

In the fist line, we create one integrate and fire neuron from the model iaf_psc_alpha.

The return value of Create is an integer that identifies the last node that was created in the network (note that this can be different from 1 if you have not called ResetKernel before). This integer is called the node's global id (the network as a whole owns the global id 0, therefore the ids of user-created nodes start with 1). Often, it is neccessary to have a large number of nodes of the same type. The command Create can also be used for this purpose. The following line of code create 10 integrate and fire neurons:

SLI ] /iaf_psc_alpha 10 Create ==
11

Status information

Nodes have a state which can be extracted and modified. In the follwing example, we display the status information of one the neurons in the layer we have created above:

SLI ] 1 ShowStatus
--------------------------------------------------                                                                                
Name                     Type                Value                                                                                
--------------------------------------------------                                                                                
archiver_length          integertype         0                                                                                    
C_m                      doubletype          250                                                                                  
E_L                      doubletype          -70                                                                                  
frozen                   booltype            false
global_id                integertype         1
I_e                      doubletype          0
local                    booltype            true
local_id                 integertype         1
model                    literaltype         iaf_neuron
node_type                literaltype         neuron
parent                   integertype         0
recordables              arraytype           <arraytype>
state                    integertype         0
tau_m                    doubletype          10
tau_minus                doubletype          20
tau_minus_triplet        doubletype          110
tau_syn                  doubletype          2
thread                   integertype         0
t_ref                    doubletype          2
t_spike                  doubletype          -1
vp                       integertype         0
V_m                      doubletype          -70
V_reset                  doubletype          -70
V_th                     doubletype          -55
--------------------------------------------------
Total number of entries: 24

Using the command SetStatus, it is possible to change the entries of this so called status dictionary. The following lines of code change the threshold value V_th to -60 mV:

SLI ] 1 << /V_th -60.0 >> SetStatus
SLI ] 1 GetStatus /V_th get =
-60

Please note, that SetStatus checks if a property really exists in a node and will issue an error if it doesn't. This behavior can be changed by the following command:

0 << /dict_miss_is_error false >> SetStatus

Then, NEST is very tolerant with respect to the property that you are trying to change: If it does not know the property, or if the property cannot be changed, there will be no error, but only a warning. In any case, SetStatus does complain if the new value does not match in the expected type:

SLI ] 1 << /V_th (60) >> SetStatus


Dec 01 15:33:54 SetStatus_ad [Error]: TypeMismatch
    Expected datatype: doubletype
    Provided datatype: stringtype

In order to find out, which properties of a given model can be changed an which not, you have to refer to the model's documentation.

Connections

Connections between nodes define possible channels for interactions between them. A connection between two nodes is established, using the command Connect.

Each connection has two basic parameters, weight and delay. The weight determines the strength of the connection, the delay determines how long an event needs to travel from the sending to the receiving node. The delay must be a positive number greater or equal to the simulation stepsize and is given in ms.

Example 1

SLI ] /iaf_psc_alpha Create /n1 Set
SLI ] /iaf_psc_alpha Create /n2 Set
SLI ] /iaf_psc_alpha Create /n3 Set
SLI ]
SLI ] n1 n2 Connect
SLI ] n1 n3 Connect

To inspect the parameters of a connection, one first needs to obtain a handle to the connection. This is done using the command GetConnections. It takes a dictionary that at least contains the id of the source node and will return a list of handles for all outgoing connections. The search can be restricted by using the optional parameters target and synapse_type.

Example 2

SLI ] << /source n1 >> GetConnections /c1 Set
SLI ] c1 length ==
2
SLI ] << /source n1 /target n2 >> GetConnections /c2 Set
SLI ] c2 length ==
1

To actually see the parameters of the connection, GetStatus is used, just like it is for nodes.

Example 3

SLI ] c1 0 get GetStatus info
--------------------------------------------------
Name                     Type                Value
--------------------------------------------------
delay                    doubletype          1
receptor                 integertype         0
sizeof                   integertype         32
source                   integertype         1
synapse_model            literaltype         static_synapse
target                   integertype         2
weight                   doubletype          1
--------------------------------------------------
Total number of entries: 7

To change the paramters of a connection, SetStatus is used, just like it is for nodes.

Example 4

SLI ] c1 0 get << /weight 2.0 >> SetStatus
SLI ] c1 0 get GetStatus /weight get ==
2.000000e+00

Devices

Devices are network nodes which provide input to the network or record its output. They encapsulate the stimulation and measurement process. If you want to extract certain information from a simulation, you need a device which is able to deliver this information. Likewise, if you want to send specific input to the network, you need a device which delivers this input.

Devices have a built-in timer which controls the period they are active. Outside this interval, a device will remain siltent. The timer can be configured using the command SetStatus.

By definition a device is active in the interval \((t_1,t_2)\) , if we can observe events \(E\) with time stamps \(t_E\) which obey \(t_1 <= t_E < t_2\) for all \(E\) . In other words, the interval during which the device is active corresponds to the range of time-stamps of the device's events.

Note that it is not possible to generate/observe an event with time stamp 0.

Device parameters

The following entries of the status dictionary are the same for all stimulation and recording devices:

Property Type Description /start double First time of activity, relative to the value of origin in ms. /stop double First time of inactivity, relative to the value of origin in ms. /origin double Origin of the device clock, relative to the network time in ms. In general, the following must hold:

  1. stop >= start
  2. If stop = start, the device is inactive.

Stimulating Devices

A range of devices is available for the stimulation of neurons. The most important ones are listed in the following table. For details, refer to the documentation of the respective decive.

Model name Description spike_generator Device to generate spikes at specific times. poisson_generator Device to generate poisson shotnoise. dc_generator Device to generate a constant current. ac_generator Device to generate an alternating (sine) current. step_current_generator Device to generate a step current with different amplitudes at different times.

Example 5

SLI ] /iaf_psc_alpha Create /n Set
SLI ] /poisson_generator Create /pg Set
SLI ] pg << /rate 220.0 Hz >> SetStatus
SLI ] pg n Connect

Recording devices

All devices which are used to observe the state of other network nodes are called recording devices. Examples are voltmeter and spike_detector.

Recording devices have properties which control the amount, the format, and the destination of their output. All recorders can either dump the recorded data to a file (property to_file), print it to the screen (property to_screen) or hold the data in memory (property to_memory). Data stored in memory can be retrieved after the simulation using GetStatus.

Device models are also stored in the dictionary modeldict. The most important devices are:

Model name Description voltmeter Device to observe membrane potentials. multimeter Device to observe arbitrary analog quantities. spike_detector Device to observe spike times. Please note that the connection direction for analog recorders (all except spike_detector in above list) is inverted with respect to other recorders.

Example 6

SLI ] /iaf_psc_alpha Create /n Set
SLI ] /voltmeter Create /vm Set
SLI ] /spike_detector Create /sd Set
SLI ] vm n Connect
SLI ] n sd Connect

Simulation

NEST simulations are time driven. The simulation time proceeds in discrete steps of size dt, set using the property resolution of the root node. In each time slice, all nodes in the system are updated and pending events are delivered.

The simulation is run by calling the command t Simulate, where t is the simulation time in milliseconds