Ion Channels
The KSChannel object encompasses all the allowed formats for ion channel definitions. The "KS" prefix indicates that it is focused on kinetic schemes, but it also allows multi-complex channels which enables Hodgkin-Huxley style models to be included within the same formalism. A single complex scheme is one where any state is connected to all other states directly or indirectly by channel transitions. A multi-complex scheme is one where the states fall into disjoint groups, or gating complexes. Each gating complex behaves independently and the relative conductance of the whole channel is the product of the relative conductances of each complex. A typical Hodgkin-Huxley model has three or four independent two-state complexes.
For stochastic calculations, multi-complex schemes are internally converted to the equivalent single-complex scheme. For deterministic calculations either form may be used, depending which is more efficient. Further details on the stochastic channel algorithm can be found in the numerical methods section of the user guide.
A channel is specified as a collection of states and transitions. There are two types of state, OpenState or ClosedState, and rather more types of transition. Each state should have an id that is unique within the channel definition. The transitions then refer to states by this id in their from and to attributes specifying the start and end states. There are a number of different parameterizations provided for the transitions themselves, but in general models should use the VHalfTransition for all voltage-dependent transitions. The other transition types are provided so that PSICS can exactly express certain legacy channel models for validation purposes but do not necessarily have much biophysical justification.
For multi-complex channels, the states and transitions for each complex should be embedded within KSComplex objects. This is not strictly required (PSICS can work out what the sub-complexes are just from the transition start and end points) but is good practice. It is required if there are multiple instances of a certain complex and you do not want to list all the states and transitions explicitly. In that case you can define them once and set the instances attribute to say how many instances of the complex there are in the channel. This is primarily useful for representing Hodgkin-Huxley style models where the powers on the gating variables can be interpreted as the number of instances of a two-state gate.
KSChannel |
An ion channel represented by a kinetic scheme |
Standalone model |
Also known as a Markov model,the schem defines the possible states (configurations) of the channel and the transitions between them. If the scheme falls into disjoint sub-schemes these are interpreted as independent serial gating complexes and the effective relative conductance os the product of the relative conductances of separate complexes. This enables Hodgkin Huxley styleto be economically represented as a collection of separate two-state complexes
Attributes
Name | Type | Definition | Units | Range | Required |
---|---|---|---|---|---|
id | identifier | Identifier (name) for the element; unique within the model | yes | ||
permeantIon | identifier reference | The permeant ion (Ion) | yes | ||
gSingle | Floating point value | Single channel conductance. Separate open states canspecify their relative conductance as a fraction of this quantity. | pS | (0.1, 100) | yes |
Elements
Element type | Role |
---|---|
OpenState, ClosedState | list of states |
FixedRateTransition, VHalfTransition, VRateTransition, ExpLinearTransition | list of transitions |
KSComplex | gating complexes |
OhmicConductanceModel | Conductance model - just one per channel |
CodedTransitionFunction | (deprecated) Additional coded functions, if any, needed for coded transitions. |
Function | Additional functions, if any, needed for transitions. |
About | Extended textual information about the model |
Parameter | Parameters that can be used within the component |
ClosedState |
non-permeable configuration of an ion channel |
within: KSChannel |
Attributes
Name | Type | Definition | Units | Range | Required |
---|---|---|---|---|---|
id | identifier | Identifier (name) for the state; unique within this channel | yes | ||
x | Floating point value | position for visualization of state diagram | none | (-1, 1) | |
y | Floating point value | position for visualization of state diagram | none | (-1, 1) |
Elements - No child elements are allowed
Example
OpenState |
Permeable configuration of an ion channel |
within: KSChannel |
The conductance is specified relative to the maximum conductance for the channel. Different states can have different conductances, but other permeation properties are assumed to be the same for all open states.
Attributes
Name | Type | Definition | Units | Range | Required |
---|---|---|---|---|---|
id | identifier | Identifier (name) for the state; unique within this channel | yes | ||
gRel | Floating point value | relative conductance for this state compared to the channel conductance | none | (0, 1) | yes |
x | Floating point value | position for visualization of state diagram | none | (-1, 1) | |
y | Floating point value | position for visualization of state diagram | none | (-1, 1) |
Elements - No child elements are allowed
Example
FixedRateTransition |
A transition between states of a kinetic scheme with fixed forward and reverse rates |
within: KSChannel |
A fixed-rate transition, sometimes known as a time-dependent transition, is one where the probability of the transition occuring in a given time interal is always the same, independent of external factors such as the membrane potential.
Attributes
Name | Type | Definition | Units | Range | Required |
---|---|---|---|---|---|
id | identifier | Identifier (name) for the transition; unique within this channel | yes | ||
from | identifier reference | The originating state of the transition (ClosedState, OpenState) | yes | ||
to | identifier reference | The destination state of the transition (ClosedState, OpenState) | yes | ||
forward | Floating point value | Forward rate, transitions per ms | per_ms | (0.01, 1000) | yes |
reverse | Floating point value | Reverse rate, transitions per ms | per_ms | (0.01, 1000) | yes |
baseTemperature | Floating point value | Base temperature for Q10 | Celsius | (5,40) | |
q10 | Floating point value | Temperature dependence of rates: rate change for a rise of ten degrees | none | (0,4) |
Elements - No child elements are allowed
Example
VHalfTransition |
Voltage dependent transition defined by its midpoint |
within: KSChannel |
This is one of two ways of expressing a first order Boltzmann transition with one or two extra term(s) to make the rates saturate rather than become indefinitely large for extreme potentials. This form uses the mid-pint potential, vHalf at which forward and reverse rates are equal, and the timescale (reciprocol of the rate) at that point. The saturation timescale can be expressed either as tauMin, or separately for the forward and reverse rates as tauMinFwd and tauMinRev.
Attributes
Name | Type | Definition | Units | Range | Required |
---|---|---|---|---|---|
id | identifier | Identifier (name) for the transition; unique within this channel | yes | ||
from | identifier reference | The originating state of the transition (ClosedState, OpenState) | yes | ||
to | identifier reference | The destination state of the transition (ClosedState, OpenState) | yes | ||
vHalf | Floating point value | Potential at which the forward and reverse rates are equal | mV | (-80, -20) | yes |
z | Floating point value | Equivalent gating charg in electronic charge units | e | (-4, 4) | yes |
gamma | Floating point value | Gating assymetry: relative position of potential peak in the transit of gating particle | none | [0,1] | yes |
tau | Floating point value | transition timescale (reciprocol ofthe rate at vHalf) | ms | (0.001, 1) | yes |
tauMin | Floating point value | minimum transition timescale (saturation of rate for extreme potentials) - must either set tauMin or the forward and reverse timescales separately | ms | (1.e-5, 1.) | |
tauMinFwd | Floating point value | minimum transition timescale for forward transition (if tauMin is not set) | ms | (1.e-5, 1.) | |
tauMinRev | Floating point value | minimum transition timescale for reverse transition (if tauMin is not set) | ms | (1.e-5, 1.) | |
baseTemperature | Floating point value | Base temperature for Q10 | Celsius | (5,40) | |
q10 | Floating point value | Temperature dependence of rates: rate change for a rise of ten degrees | none | (0,4) |
Elements - No child elements are allowed
Example
VRateTransition |
Voltaqe dependent transition defined by forward and reverse rates are v=0 |
within: KSChannel |
This is one of two ways of expressing a first order Boltzmann transition with one or two extra term(s) to make the rates saturate rather than become indefinitely large for extreme potentials. This version requires the forward and reverse rates for a membrane potential of zero. Internally, these transitions are converted into exactly the same form as is used for the VHalfTransition.
Attributes
Name | Type | Definition | Units | Range | Required |
---|---|---|---|---|---|
id | identifier | Identifier (name) for the transition; unique within this channel | yes | ||
from | identifier reference | The originating state of the transition (ClosedState, OpenState) | yes | ||
to | identifier reference | The destination state of the transition (ClosedState, OpenState) | yes | ||
forward | Floating point value | Forward rate at zero potential difference | per_ms | (0.01, 1000) | yes |
reverse | Floating point value | Reverse rate at zero potential difference | per_ms | (0.01, 1000) | yes |
z | Floating point value | Equivalent gating charge | e | (-5, 5) | yes |
gamma | Floating point value | Gating assymetry | none | [0., 1.] | yes |
tauMin | Floating point value | Saturation time constant | ms | (0.0001, 1) | yes |
baseTemperature | Floating point value | Base temperature for Q10 | Celsius | (5,40) | |
q10 | Floating point value | Temperature dependence of rates: rate change for a rise of ten degrees | none | (0,4) |
Elements - No child elements are allowed
Example
ExpLinearTransition |
HH style one-way exponential-linera transition |
within: KSChannel, KSComplex |
A one-way transition expressed in the form A x / (1 - exp(-x)) where x = (v - v0) / B. The parameters are the rate, A, the modpoint, v0, where x = 0, and the voltage scale, scale, which divides both the midpoint and v0. Note that this expression has numerous variants, so published parameters may not map directly onto the rate, midpoint and scale. Most often the signs of the rate and scale are reversed or the scale dependence of the numerator is wrapped into the rate, in a form sucha as alpha (v - v0) / (exp (beta (v - v0)) - 1).
Attributes
Name | Type | Definition | Units | Range | Required |
---|---|---|---|---|---|
id | identifier | Identifier (name) for the transition; unique within this channel | yes | ||
from | identifier reference | The originating state of the transition (ClosedState, OpenState) | yes | ||
to | identifier reference | The destination state of the transition (ClosedState, OpenState) | yes | ||
rate | Floating point value | Rate at inflection | per_ms | (0.01, 1000) | yes |
midpoint | Floating point value | Potential of inflection between exponential and linear dependiecies | mV | (-80, -20) | yes |
scale | Floating point value | Potential scaling | mV | (-80, 40) | yes |
baseTemperature | Floating point value | Base temperature for Q10 | Celsius | (5,40) | |
q10 | Floating point value | Temperature dependence of rates: rate change for a rise of ten degrees | none | (0,4) |
Elements - No child elements are allowed
Example
ExpTransition |
Exponential HH style rate |
within: KSChannel |
A one-way transition as used in the original HH model, giving the rate, beta, as beta = rate exp((v - midpoint)/ vscale)
Attributes
Name | Type | Definition | Units | Range | Required |
---|---|---|---|---|---|
id | identifier | Identifier (name) for the transition; unique within this channel | yes | ||
from | identifier reference | The originating state of the transition (ClosedState, OpenState) | yes | ||
to | identifier reference | The destination state of the transition (ClosedState, OpenState) | yes | ||
rate | Floating point value | Rate at v = midpoint | per_ms | (0.01, 1000) | yes |
scale | Floating point value | Scale factor for voltage term | mV | (0.0, 100) | yes |
midpoint | Floating point value | Potential of inflection between exponential and linear dependiecies | mV | (-80, -20) | yes |
baseTemperature | Floating point value | Base temperature for Q10 | Celsius | (5,40) | |
q10 | Floating point value | Temperature dependence of rates: rate change for a rise of ten degrees | none | (0,4) |
Elements - No child elements are allowed
Example
SigmoidTransition |
Exponential HH style rate |
within: KSChannel |
A one-way transition as used in the original HH model, giving the rate, beta, as beta = rate / (1 + exp(-(v - midpoint)/ vscale))
Attributes
Name | Type | Definition | Units | Range | Required |
---|---|---|---|---|---|
id | identifier | Identifier (name) for the transition; unique within this channel | yes | ||
from | identifier reference | The originating state of the transition (ClosedState, OpenState) | yes | ||
to | identifier reference | The destination state of the transition (ClosedState, OpenState) | yes | ||
rate | Floating point value | Rate at v = midpoint | per_ms | (0.01, 1000) | yes |
scale | Floating point value | Scale factor for voltage term | mV | (0.0, 100) | yes |
midpoint | Floating point value | Potential of midpoint of curve, where the rate is half its maximum value | mV | (-80, -20) | yes |
baseTemperature | Floating point value | Base temperature for Q10 | Celsius | (5,40) | |
q10 | Floating point value | Temperature dependence of rates: rate change for a rise of ten degrees | none | (0,4) |
Elements - No child elements are allowed
Example
KSComplex |
a single complex within a kinetic scheme channel |
Standalone model or within: KSChannel |
KSComplexes provide an optional grouping of states within a channel. The effect is the same as specifying transitions between top-level that implicitly define separate gating complexes, with the exception that the complex also allows an instances attribute that can be used to specify that the channel has more tna one sequential instance of the same complex. This is primarily useful for the economical representation of Hodgkin-Huxley style channels that have multiple independent, but identically gated, two-state complexes.
Attributes
Name | Type | Definition | Units | Range | Required |
---|---|---|---|---|---|
id | identifier | Identifier (name) for the complex | yes | ||
instances | Whole number | number of complexes of this type in the channel, equivalentto the power used in HH models | [1,4) |
Elements
Element type | Role |
---|---|
OpenState, ClosedState | list of states |
FixedRateTransition, VHalfTransition, VRateTransition, ExpLinearTransition | list of transitions |
TauInfCodedTransition |
Voltaqe dependent transition defined by a fragment of code |
within: KSChannel |
This
Attributes
Name | Type | Definition | Units | Range | Required |
---|---|---|---|---|---|
id | identifier | Identifier (name) for the transition; unique within this channel | yes | ||
from | identifier reference | The originating state of the transition (ClosedState, OpenState) | yes | ||
to | identifier reference | The destination state of the transition (ClosedState, OpenState) | yes | ||
baseTemperature | Floating point value | Base temperature for Q10 | Celsius | (5,40) | |
q10 | Floating point value | Temperature dependence of rates: rate change for a rise of ten degrees | none | (0,4) | |
tauvar | plain text | The name of the output variable for the time constant. | |||
infvar | plain text | The name of the output variable for the equilbrium gate position. |
Elements
Element type | Role |
---|---|
Constant |
Example
TauInfTransition |
Voltaqe dependent transition defined by functions for Tau and m-infinity |
within: KSChannel |
Attributes
Name | Type | Definition | Units | Range | Required |
---|---|---|---|---|---|
id | identifier | Identifier (name) for the transition; unique within this channel | yes | ||
from | identifier reference | The originating state of the transition (ClosedState, OpenState) | yes | ||
to | identifier reference | The destination state of the transition (ClosedState, OpenState) | yes | ||
baseTemperature | Floating point value | Base temperature for Q10 | Celsius | (5,40) | |
q10 | Floating point value | Temperature dependence of rates: rate change for a rise of ten degrees | none | (0,4) |
Elements
Element type | Role |
---|---|
Constant |