Sections are unbranched lengths of continuous cable connected together to form a neuron. Sections can be connected to form any tree-shaped structure but loops are not permitted. (You may, however, develop membrane mechanisms, such as electrical gap junctions which do not have the loop restriction. But be aware that the electrical current flows through such connections are calculated by a modified euler method instead of the more numerically robust fully implicit/crank-nicholson methods)
Do not confuse sections with segments. Sections are divided into segments of equal length for numerical simulation purposes (see :hoc:data:`nseg`). NEURON uses segments to represent the electrical circuit shown below.
Ra
o/`--o--'\/\/`--o--'\/\/`--o--'\/\/`--o--'\o v
| | | |
--- --- --- ---
| | | | | | | |
--- --- --- ---
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-------------------------------------------- ground
Such segments are similar to compartments in compartmental modeling programs.
Section geometry is used to compute the area and axial resistance of each segment.
There are two ways to specify section geometry:
- The stylized method simply specifies parameters for length and diameter.
- The 3-D method specifies a section's shape, orientation, and location in three dimensions.
Choose the stylized method if the notions of cable length and diameter are authoritative and where 3-d shape is irrelevant. For plotting purposes, length and diameter will be used to generate 3-d info automatically for a stylized straight cylinder. (see :hoc:func:`define_shape`)
Choose the 3-D method if the shape comes from 3-d reconstruction data or if your 3-d visualization is paramount. This method makes the 3-d info authoritative and automatically determines the abstract cable's length and diameter. With this method, you may change a section's length/diameter only by changing its 3-d info. (but see :hoc:func:`pt3dconst`)
For simulations one needs to specify L, nseg, diam, Ra, and connectivity.
- L
- For each section, L is the length of the entire section in microns.
- nseg
- The section is divided into nseg compartments of length L/nseg. Membrane potential will be computed at the ends of the section and the middle of each compartment.
- diam
- The diameter in microns. Note that diam is a range variable and therefore must be respecified whenever :hoc:data:`nseg` is changed.
- Ra
- Axial resistivity in ohm-cm.
- connectivity
- This is established with the :ref:`connect <hoc_keyword_connect>` command and defines the parent of the section, which end of the section is attached to the parent, and where on the parent the attachment takes place. To avoid confusion, it is best to attach the 0 end of a section to the 1 end of its parent.
In the stylized specification, the shape model used for a section is
a sequence of right circular cylinders of length, L/nseg, with diameter
given by the diam range variable at the center of each segment.
The area of a segment is PI*diam*L/nseg (micron2) and the half-segment axial
resistance is .01*Ra*(L/2/nseg)/(PI*(diam/2)^2). The .01 factor is necessary
to convert ohm-cm micron/micron2 to MegOhms. Ends of cylinders are not
counted in the area and, in fact, the areas are very close to those of
truncated cones as long as the diameter does not change too much.
forall delete_section()
create a
access a
nseg = 10
Ra = 100
L=1000
diam(0:1)=10:100
for (x) print x, diam(x), area(x), PI*diam(x)*L/nseg, ri(x), .01*Ra*(L/2/nseg)/(PI*(diam(x)/2)^2)
Note that the area (and length) of the 0,1 terminal ends is equal to 0
and the axial resistance
is the sum of the adjacent half-segment resistances between segment and
parent segment. Such, niceties allow the spatial discretization error to
be proportional to (1/nseg)^2. However, for second order correctness,
all point processes must be located at the center of the segments or at the
ends and all branches should be connected at the ends or centers of segments.
Note that if one increases nseg by a factor of 3, old centers are preserved.
For single compartment simulations it is most convenient to choose a membrane area of 100 micron2 so that point process currents (nanoamps) are equivalent to density currents (milliamps/cm2).
Also note that a single compartment of length = diameter has the same effective area as that of a sphere of the same diameter.
- Example:
The following example demonstrates the automatic 3-d shape construction. The root section "a" is drawn with its 0 end (left) at the origin and is colored red.
Sections connected to its 1 end (sections b, c, d) get drawn from left to right. Sections descended from the 0 end (section e) of the root get drawn from right to left.
Especially note the diameter pattern of section c whose "1" end is connected to the "b" parent. You don't have to understand this if you always connect the "0" end to the parent.
forall delete_section() create a, b, c, d, e connect b(0), a(1) connect c(1), b(1) connect d(0), b(1) connect e(0), a(0) forall nseg=20 forall L=100 forall diam(0:1) = 10:40 objref s s = new Shape() s.show(0) a s.color(2) topology() finitialize() forall { print secname() for i=0,n3d()-1 print i, x3d(i), y3d(i), z3d(i), diam3d(i) }If you change the diameter or length, the Shape instances are automatically redrawn or when :hoc:func:`doNotify` is called. Segment area and axial resistance will be automatically recomputed prior to their use.
Under some circumstances, involving nonlinearly varying diameters across a section, at first sight surprising results can occur when the stylized method is used and a Shape instance is created. This is because under a define_shape() with no pre-existing 3-d points in a section, a number of 3-d points is created equal to the number of segments plus the end areas. When 3-d points exist, they determine the calculation of L, diam, area, and ri. Thus diam can change slightly merely due to shape creation. When L and diam are changed, there is first a change to the 3-d points and then L and diam are updated to reflect the actual values of these 3-d points. Due to multiple interpolation effects, specifying a nonlinearly varying diam will, in general, not give exactly the same diameter values as the case where no 3-d information exists. This effect is illustrated in the following example
forall delete_section() objref s proc pr() { pt3dclear() nseg = $1 diam = 10 diam(.34:.66) = 20:20 define_shape() for(x) print x*L, diam(x), area(x), ri(x) } xpanel("change nseg") xradiobutton("nseg = 3", "pr(3)") xradiobutton("nseg = 11", "pr(11)") xradiobutton("nseg = 101", "pr(101)") xpanel() create a access a nseg=3 {Ra=100 L=100} diam=10 diam(.34:.66) = 20:20 for(x) print x*L, diam(x), area(x), ri(x) s = new Shape() s.show(0) for i=0, n3d()-1 print i, arc3d(i), diam3d(i) print "L=", L for(x) print x*L, diam(x), area(x), ri(x)
The difference is that the 3-d points define a series of truncated cones instead of a series of right circular cylinders. The difference is reduced with larger nseg. With the stylized method, abrupt changes in diameter should only take place at the boundaries of sections if you wish to view shape and also make use of the fewest possible number of segments. But remember, end area of the abrupt changes is not calculated. For that, you need an explicit pair of 3-d points with the same location and different diameters.
3-d information for a section is kept in a list of (x,y,z,diam) "points". The first point is associated with the end of the section that is connected to the parent (NB: Not necessarily the 0 end) and the last point is associated with the opposite end. There must be at least two points and they should be ordered in terms of monotonically increasing arc length.
The root section is treated as the origin of the cell with respect to 3-d position. When any section's 3-d shape or length changes, all the sections in the child trees have their 3-d information translated to correspond to the new position. So, assuming the soma is the root section, to translate an entire cell to another location it suffices to change only the location of the soma. It will avoid confusion if, except for good reason, one attaches only the 0 end of a child section to a parent. This will ensure that the diam(x) as x ranges from 0 to 1 has the same sense as diam3d(i) as i ranges from 0 to n3d()-1.
The shape model used for a section when the pt3d list is non-empty
is that of a sequence of truncated cones in which the pt3d points define
the location and diameter of the ends. From this sequence of points,
the effective area, diameter, and resistance is computed for each segment
via a trapezoidal integration across the segment length. This takes
into account the extra area due to sqrt(dx^2 + dy^2) for fast changing
diameters (even degenerate cones of 0 length can be specified, ie. two
points with same coordinates but different diameters)
but no attempt is made to deal with centroid curvature effects
on the area. Note that the number of 3d points used to describe a shape
has nothing to do with nseg and does not affect simulation speed.
(Although, of course, it does affect how fast one can draw the shape)
- Example:
The following illustrates the notion of the 3-d points as describing a sequence of cones. Note that the segment area and resistance is different than the simplistic calculation used in the stylized method. In this case the area of the segment has very little to do with the diameter of the center of the segment.
forall delete_section() create a access a Ra=100 nseg = 10 pt3dclear() for i=0,30 { x = PI*i/30 pt3dadd(200*sin(x), 200*cos(x), 0, 100*sin(4*x)) } objref s s = new Shape() s.show(0) print L for (x) print x, diam(x), area(x), PI*diam(x)*L/nseg, ri(x), .01*Ra*(L/2/nseg)/(PI*(diam(x)/2)^2)Note that at one point the diameter is numerically 0 and the axial resistance becomes essentially infinite thus decoupling the adjacent segments. Take care to avoid constructing spheres with a beginning and ending diameter of 0. No current would flow from the end to a connecting section. The end diameter should be the diameter of the end of the connecting section.
The following loads the pyramidal cell 3-d reconstruction from the demo directory of your neuron system. Notice that you can modify the length only if the pt3dconst mode is 1.
forall delete_section() xopen("$(NEURONHOME)/demo/pyramid.nrn") mode = 1 pt3dconst(mode) objref s s = new Shape() s.action("dendrite_1[8] s.select()") dendrite_1[8] s.color(2) xpanel("Change Length") xvalue("dendrite_1[8].L", "dendrite_1[8].L", 1) xcheckbox("Can't change length", &mode, "pt3dconst(mode)") xpanel()
.. seealso::
:hoc:func:`pt3dclear`, :hoc:func:`pt3dadd`, :hoc:func:`pt3dconst`, :hoc:func:`pt3dstyle`, :hoc:func:`n3d`, :hoc:func:`x3d`, :hoc:func:`y3d`, :hoc:func:`z3d`, :hoc:func:`diam3d`, :hoc:func:`arc3d`
:hoc:func:`getSpineArea`, :hoc:func:`setSpineArea`, :hoc:func:`spine3d`
.. seealso::
:hoc:func:`define_shape`, :hoc:func:`pt3dconst`
If 3-D shape is not an issue it is sufficient to specify the section variables L (length in microns), Ra (axial resistivity in ohm-cm), and the range variable diam (diameter in microns).
A list of 3-D points with corresponding diameters describes the geometry of a given section.
.. hoc:function:: pt3dclear
Syntax:
``buffersize = pt3dclear()``
``buffersize = pt3dclear(buffersize)``
Description:
Destroy the 3d location info in the currently accessed section.
With an argument, that amount of space is allocated for storage of
3-d points in that section.
.. hoc:function:: pt3dadd
Syntax:
``pt3dadd(x,y,z,d)``
Description:
Add the 3d location and diameter point at the end of the current pt3d
list. Assume that successive additions increase the arc length
monotonically. When pt3d points exist in a section they are used
to compute *diam* and *L*. When *diam* or *L* are changed and \ ``pt3dconst()==0``
the 3-d info is changed to be consistent with the new values of
*L* and *diam*. (Note: When *L* is changed, \ ``diam_shape()`` should be executed
to adjust the 3-d info so that branches appear connected.)
The existence of a spine at this point is signaled
by a negative value for *d*.
.. hoc:function:: pt3dconst
Syntax:
``pt3dconst(0)``
``pt3dconst(1)``
Description:
If \ ``pt3dconst`` is set at 0, newly assigned values for *d* and *L* will
automatically update pre-existing 3d information.
\ ``pt3dconst`` returns its previous state on each call. Its original value is 0.
Note that the *diam* information transferred to the 3d point information
comes from the current diameter of the segments and does not change
the number of 3d points. Thus if there are a lot of 3d points the
shape will appear as a string of uniform diameter cylinders each of
length L/nseg. ie. after transfer \ ``diam3d(i) == diam(arc3d(i))``.
Then, after a call to an internal function such as \ ``area()`` or
\ ``finitialize()``, the 3d point info will be used to determine the values
of the segment diameters.
Because of the three separate interpolations:
hoc range spec -> segment diameter -> 3d point diam -> segment diameter,
the final values of the segment diameter may be different from the
case where 3d info does not exist.
Because of the surprises noted above, when using 3d points
consider treating them as the authoritative diameter info and set
\ ``pt3dconst(1)``.
3d points are automatically generated when one uses
the nrniv Shape class. If you want the flexibility of being able
to specify 3d diameter using range variable notation
(eg diam(0:1) = 10:20) you will need to experiment with \ ``nseg`` and
\ ``n3d()`` in order to understand the exact consequences of interpolation.
.. seealso::
:hoc:func:`pt3dstyle`
.. hoc:function:: pt3dstyle
Syntax:
``style = pt3dstyle()``
``style = pt3dstyle(0)``
``style = pt3dstyle(1, x, y, z)``
``style = pt3dstyle(1, &x, &y, &z)``
Description:
With no args, returns 1 if using a logical connection point.
With a first arg of 0, then style is NO logical connection point
and (with :hoc:func:`pt3dconst` == 0 and define_shape is executed)
the 3-d location info is translated so the first 3-d point coincides with
the parent connection location. This is the classical and default behavior.
With a first arg of 1 and x,y,z value arguments, those values are used
to define a logical connection point relative to the first 3-d point.
When :hoc:func:`pt3dconst` == 0 and define_shape is executed, the 3-d location
info is translated so that the logical connection point coincides
with the parent connection location. Note that logical connection points
have absolutely no effect on the electrical properties of the structure since
they do not affect the length or area of a section.
They are useful mostly for accurate visualization of a dendrite connected
to the large diameter edge of a soma that happens to be far from the
soma centroid. The logical connection point should be set to the location
of the parent centroid connection, i.e. most often the 0.5 location
of the soma. Note, that under translation and scaling,
the relative position between
the logical connection point and the first 3-d point is preserved.
With a first arg of 1 and x,y,z reference arguments, the x,y,z variables
are assigned the values of the logical connection point (if the style
in fact was 1).
.. seealso::
:hoc:func:`pt3dconst`, :hoc:func:`define_shape`
.. hoc:function:: pt3dinsert
Syntax:
``pt3dinsert(i, x, y, z, diam)``
Description:
Insert the point (so it becomes the i'th point). If i is equal to
:hoc:func:`n3d` the point is appended (equivalent to :hoc:func:`pt3dadd`).
.. hoc:function:: pt3dremove
Syntax:
``pt3dremove(i)``
Description:
Remove the i'th point.
.. hoc:function:: pt3dchange
Syntax:
``pt3dchange(i, x, y, z, diam)``
``pt3dchange(i, diam)``
Description:
Change the i'th 3-d point info. If only two args then the second arg
is the diameter and the location is unchanged.
.. code-block::
none
pt3dchange(5, x3d(5), y3d(5), z3d(5), (spine3d(5)+1)/2 * diam3d(5))
leaves the pt3d info unchanged.
.. hoc:function:: n3d
Syntax:
``n3d()``
Description:
Return the number of 3d locations stored in the currently accessed section.
.. hoc:function:: x3d
Syntax:
``x3d(i)``
Description:
Returns the x coordinate of the ith point in the 3-d list of the
currently accessed section.
.. seealso::
:hoc:func:`y3d`, :hoc:func:`z3d`, :hoc:func:`arc3d`, :hoc:func:`diam3d`
.. hoc:function:: y3d
Syntax:
``y3d(i)``
.. seealso::
:hoc:func:`x3d`
.. hoc:function:: z3d
Syntax:
``z3d(i)``
.. seealso::
:hoc:func:`x3d`
.. hoc:function:: diam3d
Syntax:
``diam3d(i)``
Description:
Returns the diameter of the ith 3d point of the currently accessed
section.
\ ``diam3d(i)`` will always be positive even
if there is a spine at the ith point.
.. seealso::
:hoc:func:`spine3d`
.. hoc:function:: arc3d
Syntax:
``arc3d(i)``
Description:
This is the arc length position of the ith point in the 3d list.
\ ``arc3d(n3d()-1) == L``
.. hoc:function:: spine3d
Syntax:
``spine3d(i)``
Description:
Return 0 or 1 depending on whether a spine exists at this point.
.. hoc:function:: setSpineArea
Syntax:
``setSpineArea(area)``
Description:
The area of an average spine in um2. \ ``setSpineArea`` merely adds to
the total area of a segment.
.. hoc:function:: getSpineArea
Syntax:
``getSpineArea()``
Description:
Return the area of the average spine.
.. hoc:function:: define_shape
Syntax:
``define_shape()``
Description:
Fill in empty pt3d information with a naive algorithm based on current
values for *L* and *diam*. Sections that already have pt3d info are
translated to ensure that their first point is at the same location
as the parent. But see :hoc:func:`pt3dstyle` with regard to the use of
a logical connection point if the translation ruins the
visualization.
Note: This may not work right when a branch is connected to
the interior of a parent section \ ``0 < x < 1``,
rather only when it is connected to the parent at 0 or 1.
.. hoc:function:: area
Syntax:
``area(x)``
Description:
Return the area (in square microns) of the segment which contains *x*.
\ ``area(0)`` and \ ``area(1)`` = 0
.. hoc:function:: ri
Syntax:
``ri(x)``
Description:
Return the resistance (in megohms) between the center of the segment containing x
and its parent segment. This can be used to compute axial current
given the voltage at two adjacent points. If there is no parent
the "infinite" resistance returned is 1e30.
Example:
.. code-block::
none
for (x) print x, area(x), ri(x)
will print the arc length, the segment area at that arc length, and the resistance along that length
for the currently accessed section.
.. hoc:function:: distance
Syntax:
``distance() or distance(0, x)``
``len = distance(x) or len = distance(1, x)``
Description:
Compute the path distance between two points on a neuron.
If a continuous path does not exist the return value is 1e20.
\ ``distance()`` with no arguments
specifies the origin as location 0
of the currently accessed section.
\ ``distance(x) (0<=x<=1)``
returns the distance (in microns) from the origin to
this point on the currently accessed section.
To overcome the
old initialization restriction, distance(0, x) can be used to set the
origin. Note that distance is measured from the centers of
segments.
Example:
.. code-block::
none
create a, b
connect b(0), a(1)
a { L = 1000 nseg = 5 }
b { L = 200 nseg = 5 }
{ a distance(0, 0.5) } // origin is center of a
b print distance(0.0) // 500
b print distance(0.5) // 600
b print distance(1.0) // 700
.. warning::
When subtrees are connected by :hoc:meth:`ParallelContext.multisplit` , the
distance function returns 1e20 if the path spans the split location.
.. seealso::
:hoc:class:`RangeVarPlot`
.. hoc:data:: diam_changed
Syntax:
``diam_changed``
Description:
Signals the system that the coefficient matrix needs to be
recalculated.
This is not needed since \ ``Ra`` is now a section variable
and automatically sets diam_changed whenever any sections Ra is
changed.
Changing diam or any pt3d value will cause it to be set automatically.
.. hoc:data:: L
Length of a section in microns.
.. hoc:data:: diam
Diameter range variable of a section in microns.
.. hoc:data:: Ra
Syntax:
``Ra``
Description:
Axial resistivity in ohm-cm. This used to be a global variable
so that it was the same for all sections. Now, it is a section
variable and must be set individually for each section. A simple
way to set its value is
\ ``forall Ra=35.4``
Prior to 1/6/95 the default value for Ra was 34.5. Presently it is
35.4.