/* Parses a cell specification file implemented as a hoc template.
It is assumed that dendritic sections are named "dend[n]". If the global
flag_spines is set to 1, then any explicit spines (whose names match the
pattern "spine[n]") are counted on each section, and then used to adjust the
section's length and diameter according to the normalization procedure
discussed in Guy Major's PhD thesis.
Regardless of the setting of flag_spines, all explicit spines are then
deleted, and tree properties set according to the passive properties
defined at the top of this file.
Arguments:
$1: the index of the cell to read.
*/
/* Passive properties. These are mapped to NEURON's definitions of the
same properties below. */
/* The values we used in the study */
CM=0.966 // average of Hanbing's WTD1 result.
RM=1/G_PAS
RA=100 // used by default for Wolf, Evans MSN studies
/* Traub values
* CM = 0.9
* RM = 50000
* RA = 250
*/
G_PAS=1/RM
ApicalHeadDiam = .47
ApicalHeadLen = .71
ApicalNeckDiam = .19
ApicalNeckLen = .44
BasalHeadDiam = .56
BasalHeadLen = .82
BasalNeckDiam = .16
BasalNeckLen = .54
SurfaceAreaOneApicalSpine = (ApicalNeckDiam * PI * ApicalNeckLen + \
ApicalHeadDiam * PI * ApicalHeadLen)
SurfaceAreaOneBasalSpine = (BasalNeckDiam * PI * BasalNeckLen + \
BasalHeadDiam * PI * BasalHeadLen)
VolumeOneApicalSpine = \
PI * (ApicalNeckDiam/2.0) * (ApicalNeckDiam/2.0) * ApicalNeckLen + \
PI * (ApicalHeadDiam/2.0) * (ApicalHeadDiam/2.0) * ApicalHeadLen
VolumeOneBasalSpine = \
PI * (BasalNeckDiam/2.0) * (BasalNeckDiam/2.0) * BasalNeckLen + \
PI * (BasalHeadDiam/2.0) * (BasalHeadDiam/2.0) * BasalHeadLen
/*
* Applies spines to a cell on all dendrites matching the provided forsec
* pattern. The
* global variable flag_spines is ignored, since this method only makes sense
* to call when spine processing is desired.
*
* Arguments:
* $1: "forsec" pattern describing the dendrites in the tree.
*/
proc applySubtreeSpecificSpines() { local total_surface_area, dend_surface_area, \
surface_area_one_spine, surface_area_all_spines \
localobj dendrite_pattern, each_section
dendrite_pattern = new String($s1)
surface_area_one_spine = $2
dendrite_count = 0
total_surface_area = 0
forsec dendrite_pattern.s {
each_section = new SectionRef()
dendrite_count = dendrite_count + 1
temp = area(0.5)
num_spines = 0
for i = 0, (each_section.nchild - 1) each_section.child[i] {
if (issection("spine.*")) {
num_spines = num_spines + 1
}
}
dend_surface_area = 0
for (x) {
dend_surface_area = dend_surface_area + area(x)
}
total_surface_area = total_surface_area + dend_surface_area
if (dend_surface_area > 0 && num_spines > 0) {
surface_area_all_spines = (surface_area_one_spine * num_spines)
factor = (dend_surface_area + surface_area_all_spines) / dend_surface_area
L = L * (factor^(2/3))
diam = diam * (factor^(1/3))
}
}
printf("Dendrite_count: %d\n", dendrite_count)
printf("total surface area before spine correction: %f\n", total_surface_area)
}
/*
* Loads a cell, replaces the spines with the Major spine correction factor.
*
* Arguments:
* $1: Path to neuron
* $2: 1 if apical spine parameters should be assumed, 2 if basal spines,
* 3 if both should be present, depending on whether the sections are
* named dend_apical* or dend_basal*.
*/
proc readcell() { localobj sref
strdef neuron_name
neuron_name = $s1
printf("Loading neuron: %s\n", neuron_name)
spine_type = $2
// Before this point, a "create soma" call will have been needed to declare the
// soma. However, we don't want that soma to prevent the new soma from loading.
// So delete it here. (Yes, it's awkward, but the earlier create seems to be
// required by NEURON.)
forall { delete_section() }
xopen(neuron_name)
// Ensure that NEURON evaluates the cell in 3D mode when calling diam(), by
// using a side effect of the area() call. It doesn't matter which section
// is used for the call, and the return value of area() can be discarded.
forall {
area(0.5)
}
// This used to be at the end of the function. I'm trying to move it to the
// top, where it makes more sense, since the for(x) construct gets used to
// do the spine adjustment.
geom_nseg(100, 0.1) // nseg according to frequency
forall {
nseg *= 9
}
surface_area_one_spine = -1
volume_one_spine = -1
if (spine_type == 3) {
applySubtreeSpecificSpines("dend_apical", SurfaceAreaOneApicalSpine)
applySubtreeSpecificSpines("dend_basal", SurfaceAreaOneBasalSpine)
} else {
if (spine_type == 1) { // Apical
printf("using apical spines\n")
surface_area_one_spine = SurfaceAreaOneApicalSpine
volume_one_spine = VolumeOneApicalSpine
} else if (spine_type == 2) {
printf("using basal spines\n")
surface_area_one_spine = SurfaceAreaOneBasalSpine
volume_one_spine = VolumeOneBasalSpine
}
totalSurfaceArea = 0
spineSurfaceArea = 0
spineVolume = 0
if (flag_spines == 1) { // Global
forsec "dend" {
temp = area(0.5)
sref = new SectionRef()
num_spines = 0
for j = 0, sref.nchild-1 sref.child[j] {
if (issection("spine.*")) {
num_spines = num_spines + 1
}
}
SurfaceAreaDend = 0
volumeDend = 0
for (x) {
SurfaceAreaDend = SurfaceAreaDend + area(x)
}
totalSurfaceArea = totalSurfaceArea + SurfaceAreaDend
if (SurfaceAreaDend > 0 && num_spines > 0) {
SurfaceAreaAllSpines = (surface_area_one_spine * num_spines)
factor = (SurfaceAreaDend + SurfaceAreaAllSpines) / SurfaceAreaDend
L = L * (factor^(2/3))
diam = diam * (factor^(1/3))
}
spineSurfaceArea = spineSurfaceArea + SurfaceAreaAllSpines
spineVolume = spineVolume + (volume_one_spine * num_spines)
}
}
// Print some summary info for tables 1 and 2
printf("\n")
// Technically, this is the surface area *before* spine correction.
printf("surface area: %g\n", totalSurfaceArea)
} // spine_type != 3
forsec "soma" {
insert hh
/* Units are S/cm^2, values are taken from Traub 2003 */
gnabar_hh = 0.1875
gkbar_hh = 0.125
/* As far as I can find, Traub 2003 doesn't set a value for the leak
conductance, and the Traub-approved model on ModelDB doesn't include
one. */
gl_hh = 0
el_hh = -70
}
forsec "spine" { delete_section() }
forall {
Ra = RA
cm = CM
insert pas g_pas = G_PAS e_pas=E_PAS
//insert max
v = E_PAS
}
celsius = 21
}
/*
* Adds spines to a cell on all dendrites that are part of the specified SectionList.
* pattern. The
* global variable flag_spines is ignored, since this method only makes sense
* to call when spine processing is desired.
*
* Arguments:
* $o1: SectionList to loop over
* $2: Surface area of a single spine
* $3: spine density for branches in the SectionList
*
* written by Christina Weaver, Jan 2012
*/
proc applySubtreeConstantSpineDensity() { local total_surface_area, dend_surface_area, \
surface_area_one_spine, surface_area_all_spines, spine_dens, mean_diam
print "readcell: top of Subtree proc."
// Ensure that NEURON evaluates the cell in 3D mode when calling diam(), by
// using a side effect of the area() call. It doesn't matter which section
// is used for the call, and the return value of area() can be discarded.
forall {
area(0.5)
}
// This used to be at the end of the function. I'm trying to move it to the
// top, where it makes more sense, since the for(x) construct gets used to
// do the spine adjustment.
// geom_nseg(100, 0.1) // nseg according to frequency
geom_nseg(500, 0.1) // nseg according to frequency
forall {
nseg *= 9
}
surface_area_one_spine = $2
spine_dens = $3
printf("Adding spines to %s, density %.3f, spine SA %g\n",$o1,$3,$2)
dendrite_count = 0
total_surface_area = 0
forsec $o1 {
printf("\t%s\n",secname() )
dendrite_count = dendrite_count + 1
temp = area(0.5)
num_spines = L * spine_dens
dend_surface_area = 0
mean_diam = 0
for (x) {
dend_surface_area = dend_surface_area + area(x)
if( x > 0 && x < 1 ) mean_diam += diam(x)
}
mean_diam /= nseg
total_surface_area = total_surface_area + dend_surface_area
// adjusted by Christina Weaver, 5 Jan 12. Still some error, but better than using
// Patrick's method which sets the diam throughout the section to whatever it is in the
// middle of the section. See SpineDensFix.hoc for other attempts to fix this.
//
if (dend_surface_area > 0 && num_spines > 0) {
surface_area_all_spines = (surface_area_one_spine * num_spines)
factor = (dend_surface_area + surface_area_all_spines) / dend_surface_area
L = L * (factor^(2/3))
diam = mean_diam * (factor^(1/3))
}
}
printf("Dendrite_count: %d\n", dendrite_count)
printf("total surface area before spine correction: %f\n", total_surface_area)
}
/************************************************************************************
find_totLen_byDist()
DEPRECATED. Use find_totLen() below instead.
May 2012, cmw
Does NOT calculate the total length accurately. It misses half a section
length, equal to L/(2*nseg). The find_totLen() function below calculates it
properly. This code is kept here as legacy.
************************************************************************************/
func find_totLen_byDist() { local a_prLen, a_totLen, b_prLen, b_totLen, prxCut, lastX
prxCut = $1
printf("\tProximal < %g um\n",prxCut)
soma { distance() }
a_prLen = 0
a_totLen = 0
b_prLen = 0
b_totLen = 0
//printf("APICAL\n")
forsec apical {
a_totLen += L
if( distance(1) < prxCut ) {
a_prLen += L
//printf("%s: **** 0 (%g) - 1 (%g): prL %g\n",secname(),distance(0),distance(1),a_prLen)
}
if( distance(0) < prxCut && distance(1) > prxCut ) {
lastX = 0
for(x) {
if( distance(x) < prxCut ) lastX = x
}
a_prLen += distance(lastX)-distance(0)
//printf("%s: ---- 0 (%g) - %g (%g) - 1 (%g): prL %g\n",secname(),distance(0),lastX,distance(lastX),distance(1),a_prLen)
}
}
//printf("BASAL\n")
forsec basal {
b_totLen += L
if( distance(1) < prxCut ) {
b_prLen += L
//printf("%s: **** 0 (%g) - 1 (%g): prL %g\n",secname(),distance(0),distance(1),b_prLen)
}
if( distance(0) < prxCut && distance(1) > prxCut ) {
lastX = 0
for(x) {
if( distance(x) < prxCut ) lastX = x
}
b_prLen += distance(lastX)-distance(0)
printf("%s: ---- 0 (%g) - lastX = %g (%g) - 1 (%g): prL %g\n",secname(),distance(0),lastX,distance(lastX),distance(1),b_prLen)
}
}
printf("Soma\t%g\n",soma.L)
printf("Apical\n\tprox\t%g\n\tdist\t%g\n\ttotl\t%g\n",a_prLen,a_totLen-a_prLen,a_totLen)
printf("Basal\n\tprox\t%g\n\tdist\t%g\n\ttotl\t%g\n",b_prLen,b_totLen-b_prLen,b_totLen)
return a_prLen + b_prLen + soma.L
}
func find_totLen() { local prLen, totLen, prxCut, lastX, dstChg, LChg, prDistLen
prxCut = $1
//printf("\tProximal < %g um\n",prxCut)
soma { distance() }
prLen = 0
totLen = 0
forsec "soma" {
for(x) {
if( x==0 || x == 1 ) continue
Lcnt += L / nseg
}
}
prLen = Lcnt
totLen = Lcnt
forsec dendritic {
lastX = 0
LChg = 0
for(x) {
if( x==0 || x==1 ) continue
totLen += L / nseg
if( distance(x) > prxCut ) {
continue
}
//lastX = x
LChg += (L / nseg)
}
prLen += LChg
/****
if( distance(0) < prxCut && distance(1) > prxCut ) {
dstChg = distance(lastX)-distance(0)
printf("%s: [L %g / nseg %d = %g; half = %g] ---- 0 (%g) - last X = %g (%g) - 1 (%g): dstL %g\tLChg %g\tDIFF %g\n",secname(),L,nseg, L/nseg,L/(2*nseg),distance(0),lastX,distance(lastX),distance(1),dstChg,LChg,dstChg-LChg)
}
****/
}
printf("Total length, including soma (computed with L/nseg):\n\tprox\t%g\ndist\t%g\ntotal\t%g\n",prLen,totLen-prLen,totLen)
return prLen
}
func find_totApicLen() { local prLen, totLen, prxCut, lastX, dstChg, LChg, prDistLen
prxCut = $1
soma { distance() }
prLen = 0
totLen = 0
forsec "soma" {
for(x) {
if( x==0 || x == 1 ) continue
Lcnt += L / nseg
}
}
prLen = Lcnt
totLen = Lcnt
forsec apical {
lastX = 0
LChg = 0
for(x) {
if( x==0 || x==1 ) continue
totLen += L / nseg
if( distance(x) > prxCut ) {
continue
}
//lastX = x
LChg += (L / nseg)
}
prLen += LChg
}
printf("Total apical length, including soma (computed with L/nseg):\n\tprox\t%g\ndist\t%g\ntotal\t%g\n",prLen,totLen-prLen,totLen)
return prLen
}
func find_totBasLen() { local prLen, totLen, prxCut, lastX, dstChg, LChg, prDistLen
prxCut = $1
soma { distance() }
prLen = 0
totLen = 0
forsec "soma" {
for(x) {
if( x==0 || x == 1 ) continue
Lcnt += L / nseg
}
}
prLen = Lcnt
totLen = Lcnt
forsec basal {
lastX = 0
LChg = 0
for(x) {
if( x==0 || x==1 ) continue
totLen += L / nseg
if( distance(x) > prxCut ) {
continue
}
//lastX = x
LChg += (L / nseg)
}
prLen += LChg
}
printf("Total basal length, including soma (computed with L/nseg):\n\tprox\t%g\ndist\t%g\ntotal\t%g\n",prLen,totLen-prLen,totLen)
return prLen
}
