Equations are used both in NeuronGroup and Synapses to:
- define state variables
- define continuous-updates on these variables, through differential equations
Note
Brian models are defined by systems of first order ordinary differential equations, but you might see the integrated form of synapses in some textbooks and papers. See :doc:`converting_from_integrated_form` for details on how to convert between these representations.
Equations are defined by multiline strings.
An Equation is a set of single lines in a string:
dx/dt = f : unit
(differential equation)x = f : unit
(subexpression)x : unit
(parameter)
Each equation may be spread out over multiple lines to improve formatting.
Comments using #
may also be included. Subunits are not allowed, i.e., one must write volt
, not mV
. This is
to make it clear that the values are internally always saved in the basic units, so no confusion can arise when getting
the values out of a NeuronGroup and discarding the units. Compound units are of course allowed as well (e.g. farad/meter**2
).
There are also three special "units" that can be used: 1
denotes a dimensionless floating point variable,
boolean
and integer
denote dimensionless variables of the respective kind.
Note
For molar concentration, the base unit that has to be used in the equations is mmolar
(or mM
), not
molar
. This is because 1 molar is 10³ mol/m³ in SI units (i.e., it has a "scale" of 10³), whereas
1 millimolar corresponds to 1 mol/m³.
Some special variables are defined: t, dt (time) and xi (white noise).
Variable names starting with an underscore and a couple of other names that have special meanings under certain
circumstances (e.g. names ending in _pre
or _post
) are forbidden.
For stochastic equations with several xi
values it is necessary to make clear whether they correspond to the same
or different noise instantiations. To make this distinction, an arbitrary suffix can be used, e.g. using xi_1
several times
refers to the same variable, xi_2
(or xi_inh
, xi_alpha
, etc.) refers to another. An error will be raised if
you use more than one plain xi
. Note that noise is always independent across neurons, you can only work around this
restriction by defining your noise variable as a shared parameter and update it using a user-defined function (e.g. with ~Group.run_regularly),
or create a group that models the noise and link to its variable (see :ref:`linked_variables`).
Equations defining neuronal or synaptic equations can contain references to
external parameters or functions. These references are looked up at the time
that the simulation is run. If you don't specify where to look them up, it
will look in the Python local/global namespace (i.e. the block of code where
you call run). If you want to override this, you can specify an explicit
"namespace". This is a Python dictionary with keys being variable names as
they appear in the equations, and values being the desired value of that
variable. This namespace can be specified either in the creation of the group
or when you can the run function using the namespace
keyword argument.
The following three examples show the different ways of providing external variable values, all having the same effect in this case:
# Explicit argument to the NeuronGroup G = NeuronGroup(1, 'dv/dt = -v / tau : 1', namespace={'tau': 10*ms}) net = Network(G) net.run(10*ms) # Explicit argument to the run function G = NeuronGroup(1, 'dv/dt = -v / tau : 1') net = Network(G) net.run(10*ms, namespace={'tau': 10*ms}) # Implicit namespace from the context G = NeuronGroup(1, 'dv/dt = -v / tau : 1') net = Network(G) tau = 10*ms net.run(10*ms)
See :doc:`../advanced/namespaces` for more details.
The following topics are not essential for beginners.
A flag is a keyword in parentheses at the end of the line, which qualifies the equations. There are several keywords:
- event-driven
- this is only used in Synapses, and means that the differential equation should be updated only at the times of events. This implies that the equation is taken out of the continuous state update, and instead a event-based state update statement is generated and inserted into event codes (pre and post). This can only qualify differential equations of synapses. Currently, only one-dimensional linear equations can be handled (see below).
- unless refractory
- this means the variable is not updated during the refractory period. This can only qualify differential equations of neuron groups.
- constant
- this means the parameter will not be changed during a run. This allows optimizations in state updaters. This can only qualify parameters.
- constant over dt
- this means that the subexpression will be only evaluated once at the beginning
of the time step. This can be useful to e.g. approximate a non-linear term as
constant over a time step in order to use the
linear
numerical integration algorithm. It is also mandatory for subexpressions that refer to stateful functions likerand()
to make sure that they are only evaluated once (otherwise e.g. recording the value with a StateMonitor would re-evaluate it and therefore not record the same values that are used in other places). This can only qualify subexpressions. - shared
- this means that a parameter or subexpression is not neuron-/synapse-specific but rather a single value for the whole NeuronGroup or Synapses. A shared subexpression can only refer to other shared variables.
- linked
- this means that a parameter refers to a parameter in another NeuronGroup. See :ref:`linked_variables` for more details.
Multiple flags may be specified as follows:
dx/dt = f : unit (flag1,flag2)
The following lists all of the special symbols that Brian uses in equations and code blocks, and their meanings.
- dt
- Time step width
- i
- Index of a neuron (NeuronGroup) or the pre-synaptic neuron of a synapse (Synapses)
- j
- Index of a post-synaptic neuron of a synapse
- lastspike
- Last time that the neuron spiked (for refractoriness)
- lastupdate
- Time of the last update of synaptic variables in event-driven equations.
- N
- Number of neurons (NeuronGroup) or synapses (Synapses). Use
N_pre
orN_post
for the number of presynaptic or postsynaptic neurons in the context of Synapses. - not_refractory
- Boolean variable that is normally true, and false if the neuron is currently in a refractory state
- t
- Current time
- xi, xi_*
- Stochastic differential in equations
Equations defined as event-driven are completely ignored in the state update. They are only defined as variables that can be externally accessed. There are additional constraints:
- An event-driven variable cannot be used by any other equation that is not also event-driven.
- An event-driven equation cannot depend on a differential equation that is not event-driven (directly, or indirectly through subexpressions). It can depend on a constant parameter.
Currently, automatic event-driven updates are only possible for one-dimensional linear equations, but this may be extended in the future.
The model definitions for NeuronGroup and Synapses can be simple strings or Equations objects. Such objects can be combined using the add operator:
eqs = Equations('dx/dt = (y-x)/tau : volt') eqs += Equations('dy/dt = -y/tau: volt')
Equations allow for the specification of values in the strings, but does this by simple string replacement, e.g. you can do:
eqs = Equations('dx/dt = x/tau : volt', tau=10*ms)
but this is exactly equivalent to:
eqs = Equations('dx/dt = x/(10*ms) : volt')
The Equations object does some basic syntax checking and will raise an error if two equations defining the same variable are combined. It does not however do unit checking, checking for unknown identifiers or incorrect flags -- all this will be done during the instantiation of a NeuronGroup or Synapses object.
Concatenating equations
>>> membrane_eqs = Equations('dv/dt = -(v + I)/ tau : volt')
>>> eqs1 = membrane_eqs + Equations('''I = sin(2*pi*freq*t) : volt
... freq : Hz''')
>>> eqs2 = membrane_eqs + Equations('''I : volt''')
>>> print(eqs1)
I = sin(2*pi*freq*t) : V
dv/dt = -(v + I)/ tau : V
freq : Hz
>>> print(eqs2)
dv/dt = -(v + I)/ tau : V
I : V
Substituting variable names
>>> general_equation = 'dg/dt = -g / tau : siemens'
>>> eqs_exc = Equations(general_equation, g='g_e', tau='tau_e')
>>> eqs_inh = Equations(general_equation, g='g_i', tau='tau_i')
>>> print(eqs_exc)
dg_e/dt = -g_e / tau_e : S
>>> print(eqs_inh)
dg_i/dt = -g_i / tau_i : S
Inserting values
>>> eqs = Equations('dv/dt = mu/tau + sigma/tau**.5*xi : volt',
... mu=-65*mV, sigma=3*mV, tau=10*ms)
>>> print(eqs)
dv/dt = (-65. * mvolt)/(10. * msecond) + (3. * mvolt)/(10. * msecond)**.5*xi : V