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THE ANATOMICAL RECORD 2 4 4 5 7 2 4 7 8 (1996)
Posterior “Septum” of Human Spinal Cord: Normal Developmental
Variations, Composition, and Terminology
D. PARKINSON AND M.R. DEL BIG10
Departments of Anatomy and Pathology, Faculty of Medicine, University of Manitoba,
Winnipeg, Manitoba, Canada
ABSTRACT
Background: The boundary separating the posterior columns of the spinal cord is formed by the lateral margins of the neural
groove approximating to form the neural canal. In anatomy texts this line
is usually drawn as continuous, uniform, centered, and straight. It is universally termed posterior or dorsal, median “septum.”
Methods: Sections from the cervical and lumbar enlargements and the
mid-thoracic region were examined from 35 human autopsy specimens
from 20 weeks gestation to 70 years with no history of spinal cord disease
or trauma. They were stained with Masson’s trichrome, and by immunohistochemistry for collagen types 1 and 4, and for glial fibrillary acidic
protein (GFAP).
Results: One or more variations were found in the position character,
shape, or extent of the line at one or more levels in every case. There was
no midline staining for collagen other than that associated with blood vessels. There is intense immunoreactivity for GFAP from 20 weeks gestation
to 35 weeks diminishing thereafter. When the posterior columns are separated the “septum” divides.
Conclusion: In the absence of any collagen this line of separation is more
akin to a raphe than a septum. Inasmuch as there is an immediately adjacent subarachnoid posterior median septum it would be advantageous to
re-name this intraspinal structure “dorsal” or “posterior” median raphe.
0 1996 Wiley-Liss, Inc.
Key words: Human, Spinal cord developmental variations, Septum,
“Raphe”
Credit for the first observation and naming of the
posterior spinal cord sulcus and “septum” goes wanting. There is no evidence that Leonard0 observed these
features (Todd, 1983) and neither Vesalius (1543) nor
Willis (1680) drew the sulci or fissures or septa in their
depictions of the upper cord. According to Spillane
(1981),in 1774 Charles Bell noted “the shallow posterior and deep anterior median Sulci.” These were probably noted by many of his contemporary anatomists.
By 1889 standard anatomy texts (Macallister 1889)
described andlor drew in cross-sections of the spinal
cord, a straight midline separation between the posterior columns, extending from the cord surface to approximate the posterior grey commissure. The terms
“posterior septum” and “posterior fissure” were applied
to this structure. [We suspect branching clefts seen in
some of our specimens and in some text illustrations
are drying artifacts.] Subsequent standard texts (Table
1)use the word “septum” with posterior or dorsal, several with addition of “median,” for this midline separation of the posterior columns. “Sulcus” or “furrow” is
used for the surface midline indentation or those at the
dorsal root entrance. These are without bibliographical
credit references and do not change their original de0 1996 WILEY-LISS, INC.
scription in subsequent editions. Many of these same
anatomy texts describing the pia-arachnoid septum in
the posterior midline subarachnoid space also use the
term “septum posticus” or “posterior septum.” This
term is credited to Schwalbe by Morris and McMurrich
(1907) and to Magendie by Schafer et al. (19081, without reference in either case.
Medical dictionaries (Becker, 1989; Dorland, 1985;
Wiley, 1986) under the term “posterior [or “posticum”1
septum” state, “a glial-pia mater septum which dips
into the posterior median sulcus of the spinal cord and
separates the posterior funiculi.” The other they term
“subarachnoidal septum.” One dictionary, Stedman’s
(Hensyl, 19901, has no reference to either spinal septum.
Valentino et al. (1983) reported intense glial fibrillary acidic protein (GFAP) immunoreactivity in the
dorsal median septum of the rat spinal cord on day one
Received May 8, 1995; accepted October 2, 1995.
Address reprint requests to Dwight Parkinson, M.D., #128 Basic
Sciences Building, 730 William Avenue, Winnipeg, Manitoba, Canada R3E OW3.
POSTERIOR MEDIAN “SEPTUM OR “RAPHE”?
TABLE 1. Representative texts that use the term
posterior (median) septum
Austin, G.W. 1984 The Spinal Cord, 3rd ed. Igaku-Shoin,
New York.
Barr, M.L, and J.A. Kiernan 1988 The Human Nervous
System. An Anatomical Viewpoint. J. Lippincott Co.,
Philadelphia, pp. 64.
Brash, J.C. 1951 Cunningham’s Text Book of Anatomy,
9th ed. Oxford University Press, London, pp. 859-866.
Carpenter, M.B. 1984 Core Text of Neuroanatomy, 3rd
ed. William and Wilkins, Baltimore, pp. 59.
Daube, J.A., and B.A. Sandok 1978 Medical
Neurosciences. Little, Brown and Company, Boston,
pp. 302.
Everett, N.B. 1972 Functional Neuroanatomy. Lea and
Febiger, Philadelphia, pp. 31-33.
Haines, D.E. 1983 Neuroanatomy: An atlas of
structures, sections and systems. Urban and
Schwarzenberg, Baltimore, pp. 65, 67, 137.
Martin, J.H. 1989 Neuroanatomy Text and Atlas.
Elsevier, New York, pp. 116.
Macallister, A. 1989 Text Book of Human Anatomy.
Philadelphia, P. Blakiston & Son and Co.,
Philadelphia, pp. 743, 745.
Marinekovic, S.S., A. Ilic, M. Milisavjevic, and V. Kostic
1989 Funkcionalna I Topografska Neuroanatomija.
Savremena Administrackja Beograd, pp. 265.
Montemurro, D.G., and J.E. Bruni 1981 The Human
Brain in Dissection. W.B. Saunders Co., Philadelphia,
pp. 115.
Moore K.L. 1992 Clinically Oriented Anatomy, 3rd ed.
William and Wilkins, Baltimore, pp. 361.
Romero-Scierra, R. 1986 Neuroanatomy. A Conceptual
Approach. Churchill Livingstone Inc., New York, pp.
79.
Wilson, D.B., and W.J. Wilson 1983 Human Anatomy.
Oxford University Press, New York, pp. 256.
post partum. Bohme (19881, using cats, postulated that
the increasing bulk of the dorsal funiculi causes the
elongation of the ependymal cells, the basal process of
which remain to form the dorsal glial septum. Thus
until 2 or 3 months after birth (in cats) there is a small
wedge-shaped area in the dorsal wall of the central
canal consisting of fetal matrix cells with long tapering
basal processes extending into the glial septum. After
this date the matrix is exhausted and the ependyma
forms the complete lining of the canal (see Figs. 1-3).
Joosten and Gribnau (1989) reported a prominent
glial septum in the midline raphe of the medulla oblongata and in the spinal cord whereas it was absent in
the decussation area of the corticospinal tract in rats.
After the first postnatal week, the major vimentin-immunoreactive glial barrier completely disappears in
the medullary levels and reduces to a minor GFAP
immunoreactive line in the spinal cord for the rat.
Sarnat (1992)found that cells in the dorsal roof plate
of spinal cords in humans were immunoreactive for
GFAP as early as 8 weeks gestation and that the expression of GFAP diminished by 34 weeks. He believed,
as did Snow et al. (1990) working with rats, that this
midline collection of parallel fibers was important in
fetal development through its potential for repelling
growing axons and thus preventing aberrant decussation of neuronal pathways from the posterior columns.
During our earlier study of the midline subarachnoid
posterior septum o f the human spinal cord [Septum pos-
573
ticum, posterior septum] (Parkinson, 19911, it was
noted that the configuration and structure of the midline separation of the posterior columns within the spinal cord exhibited considerable variation. The present
study enlarges on these earlier observations, and considers the structure of, and terminology for, what has
classically been called the posterior (median) septum.
MATERIALS AND METHODS
Thirty-five human fresh autopsy spinal cords and
coverings that ranged from 20 weeks gestation to 70
years were examined. None had a history of cord disease or injury. Ten 6 pm cross-sections were taken from
paraffin blocks at the mid-cervical enlargement, midthoracic region, and the mid-lumbar enlargement and
mounted; a total of thirty for each of the thirty-five
cords. These were stained with Masson’s trichrome
(Masson, 1929; Lillie, 19481, and, in addition, rabbit
See figures on following pages
Figs. 1-6. Chronologically left to right. Bottom, ventral; top, dorsal.
Fig. 1. Twenty week female. Lumbar level. Early phase of approximation with incomplete closure of canal consisting of ependymal and
matrix cells. x 40.
Fig. 2. Twenty-two week male. Thoracic level. Early tenting of fibers dwindling into one to two fibers forming wavy line. The two
circular areas of loose tissue just below the canal are associated with
blood vessels. x 80.
Fig. 3. Twenty-two week male. Thoracic level. Completion of canal
lining with ependyma. Note “wavy” nature of Zine in Figures 2 and 3.
Fig. 4. Twenty-five week male. Thoracic level. Early interposition of
transverse fibers in region of posterior commissure. Note “reverse
flow” connection of tributary vessel. x 40.
Fig. 5. Thirty-seven week female. Cervical level. Tenting fibers into
one or two strands forming separating Zine. Note nest of closely
packed nuclei within tent. x 40.
Fig. 6. Two month post partum male. Lumbar level. Tenting into
line of two or three fibers. Again note nest of closely packed nuclei
with sparse cytoplasm and cell processes. Wider separation of line and
canal. x40.
Figs. 7-12. Chronologically left to right. Bottom, ventral; top, dor-
sal.
Fig. 7. Four month male. Thoracic level. Very narrow posterior commissure. Incomplete alar plate approximation with pia covering
smooth shoulders to bottom of cleft. Note arachnoid septum. Collagen
staining ends a t base of valley. Normal tributary branching. X 40.
Fig. 8. Eighteen month male. Thoracic level. Forked or split line at
arrows. x40.
Fig. 9. Two and one-half year female. Thoracic level. Gently curved
line from off center tented takes off becoming straight thereafter.
Usual vessel branching. x 20.
Fig. 10. Two and one-half year female. Thoracic level. Line absent or
indistinct from commissure to lower arrow. Single fibre from there on.
x 40.
Fig. 11. Four year male. Cervical level. Off center tenting origin
(arrow) quickly tapering to single fiber line to vessel and beyond.
x 40.
Fig. 12. Five year male. Lumbar level. Tenting origin dwindles to
2-3 fibers. Nest of 14 nuclei within tent with sparse cytoplasm and
processes. Posterior column deeper into commissure region on left
than right. x 60.
574
D. PARKINSON AND M.R. DEL BIG10
Figs. 1-6.
POSTERIOR MEDIAN “SEPTUM” OR “RAPHE”?
Figs. 7-1 2.
575
576
D. PARKINSON AND M.R. DEL BIG10
polyclonal anti-collagen type I (Monosan: M O O dilu- presumably venous as Herren and Alexander (1939)
tion), mouse monoclonal anti-collagen type IV (Dako, state that there are only veins posteriorly in the midCarpinteria, CA: 1/100 dilution), and rabbit polyclonal line. Craigie (1944a,b) and Gray (Williams and Waranti GFAP (BioGenix, San Ramon, CA: 1/100 dilution). rick, 1980) draw both arteries and veins in the postePrimary antibodies were detected using biotinylated rior midline. The usual branching configuration would
secondary antibodies followed by avidin-biotin peroxi- fit with either an artery running centripetally from the
dase method and diaminobenzidine reaction. The ex- surface or a vein running centrifugally from the inteaminations were done under an Olympus BH2 micro- rior, i.e., the branches are at an acute angle towards
scope and the photographs taken on Pantomatic-X film the center. However, several of our specimens clearly
depict a “reverse” branching, i.e., acutely away from
100 with an Olympus camera.
the center of the cord (Fig. 4).
RESULTS
Collagens type I and IV were detected in arachnoid,
From twenty to twenty-five weeks gestation ependy- pia, and the connective tissue associated with the
ma1 cells are seen (Figs. 1-4) approximating dorsal to larger blood vessels. In addition, small blood vessels
the central canal. These cells give off processes extend- exhibited collagen type IV immunoreactivity. There
ing centrifugally along the dorsal midline. The central was no immunoreactivity in the posterior midline nor
canal origin of the line is evident up to about 25 weeks, collagen staining with Masson’s unless a blood vessel
after which there are transversely arranged strands was present. The line did, however, exhibit strong iminterspersed in the region of the posterior grey com- munoreactivity for GFAP at 20 weeks gradually diminmissure (Fig. 4). The posterior grey commissure attains ishing to 35 weeks and persisting faintly into adultadult proportions by about 2 months post partum (Fig. hood, corresponding t o Sarnat’s (1992) finding in the
6). Beyond this period no age related differences were human roof plate ependymal cells.
If the surface pia-arachnoid is opened, the posterior
found (Figs. 6-18). The mature line (2 months or older)
usually arises from a “tenting” of fibers at the posterior columns can be separated evenly with a relatively
grey commissure (Figs. 2-6, 9, 11, 12, 14, 16). The smooth plane of cleavage. Fine filaments and occasionnumber of these “tenting” centrifugal fibers diminishes ally narrow ribbons or a vessel can be picked up from
with increasing distance from the central canal. The the exposed surfaces, but on neither side is a septum
fibers are replaced at varying intervals by additional demonstrable. The “septum” itself divides when the
cells, rarely more than two at any one level (Figs. 2-14, two sides are pulled apart.
16-18). These “tenting” fibers are not compacted as
DISCUSSION
though being pushed dorsally but rather are progressively more separated as though pulled from the cenThe term “septum” denotes a free-standing structure
tral canal (Figs. 6, 12, 14). These linear arranged cen- (Hensyl, 1990) which could be removed intact without
trifugal fibers originating near the canal and the disturbing the separated structures as for instance the
additional cell processes along the way do not look to be atrial septum, septum pellucidurn, or the nasal septum.
separable from the posterior columns; in fact some fi- Even muscle septa with muscle attachments are freebers appear to be entering or leaving this linear bundle standing separable structures. Our sections of the hufrom or into the adjacent posterior columns. These fi- man spinal cord from 20 weeks gestation to 70 years do
bers do not stain for collagen with Masson’s stain. not show such a structure except for an occasional short
Within the tented or wedge-shaped origin of the line extension of the arachnoid into the spinal cord, prethere are frequently clusters of small dense nuclei. sumably indicating incomplete approximation of the
(Figs. 5 , 6, 12, 16).
neural plate dorsally (Fig. 7).
At any section there may be one or more of eight
The constituents of the line separating the posterior
major variations. Rarely is the line classic: i.e., cen- columns begin as the neural plate folds on either side of
tered, straight, and complete from commissure to pial
surface (Type I) (Figs. 5-7). More often it is missing in
part (Fig. 10) (Type 11),or missing completely (Fig. 15)
Figs. 13-18. Chronologically left to right. Bottom, ventral; top,
(Type 111). It may have a single curve (Figs. 9, 16, 17) dorsal.
(Type IV), or multiple curves (Fig. 13)with long (Type
Fig. 13. Seven year female. Thoracic level. Gently curved line from
Va) or short (Type Vb) (Figs. 2,3) wavelengths. It may
arise off center (Figs. 9,11,16,17) (Type VI). It may be blood vessel to pial surface. x 40.
forked and/or divided with one or more components
Fig. 14. Nine year male. Lumbar level. Open tent containing vessel
(Fig. 8) (Type VII). The duplicated or dividing lines do and loose collagen staining reticulum leading to multiple fiber line.
not suggest enclosure of the septo-marginal bundle. x 60.
There may, or may not, be a pial surface indentation
Fig. 15. Nine year male. Cervical level. Absent line. x 40.
(Figs. 9, 18) with any of these variations. When the
approximation is incomplete (Type VIII), a valley or
Fig. 16. Nine year female. Thoracic level. Tenting origin off center
sulcus of varying depth remains with smooth rounded to left tapers to single cell curving line. Arrows: Cluster of 5 nuclei
pial shoulders and usually a pia-arachnoid extension of with sparse cytoplasm in tent. x 40.
the subarachnoid dorsal median septum into the defect
Fig. 17. Eleven year male. Lumbar level. Off center curving origin.
(Fig. 7). Any collagen staining ends at the bottom of x 40.
such a sulcus (Fig. 7). The line separating the posterior
Fig. 18. Thirty year female. Thoracic level. Multiple parallel similar
columns may be occupied by a blood vessel (Suh and lines
all blood vessels which stained for collagen below arrows. No
Carpenter, 1939)for part or all of the distance from the staining for collagen above arrows including pale wedge of cells at the
surface to the commissure (Figs. 4, 9, 13, 18). This is pial margin. x 40.
POSTERIOR MEDIAN “SEPTUM” OR “RAPHE”?
Figs. 13-18.
577
578
D. PARKINSON AND M.R. DEL BIG10
the neural groove approximate to form and enclose
the central canal. At this time in development there
is no collagen included on, or between, the approaching surfaces (Rutka et al., 1988; Sarnat, 1992; Joosten
and Gribnau, 1989). In the absence of any collagen or
any semblance of a separable structure, this line of
separation, which apparently arises from the ependyma of the central canal early in development and is
subsequently modified during maturation, is more
akin to a ruphd (Parkinson, 1995) than a septum. One
can only speculate as to whether the occasional asymmetry andlor curvature of the line is due t o chance
during embryonic approximation or differential development of various portions of the posterior columns.
Examination of 35 normal cords from 20 weeks gestation to age 70 reveals a great variation in the line
separating the posterior columns of the cord. There is
no evidence of a true septum, or even of any midline
condensation of collagen other than that seen in blood
vessel walls. Whether called posterior or dorsal, (central or medial), ‘‘raph6” more accurately defines its
true nature and corresponds with the terminology used
more cephalad in the brain stem. This has the added
benefit of distinguishing this structure from the true
septum given the same name immediately adjacent
posteriorly in the sub-arachnoid space.
ACKNOWLEDGMENTS
The cooperation of Dr. Wm. Halliday, neuropathologist, in providing the specimens, of Roy Simpson of the
photography department, and of Fran Thompson in editorial assistance is gratefully acknowledged.
LITERATURE CITED
Becker E.L. 1989 Churchill’s Medical Dictionary. Churchill Livingstone, New York.
Bohme, G. 1988 Formation of the central canal and dorsal glial septum in the spinal cord of the domestic cat. J. Anat., 159:37-47.
Chambers, W.W., and L. Chan-Nau Liu 1944 Developmental anatomy. In: The Spinal Cord, 3rd ed. Austin G.W., ed. Igaku-Shoin,
New York, pp. 8, 17.
Craigie, E.H. 1944a Extrinsic arteries. In: The Spinal Cord, 3rd. ed.
G.W. Austin, ed. Igaku-Shoin, New York. pp. 53, 62, 67.
Craigie, E.H. 1944b Intrinsic veins. In: The Spinal Cord, 3rd. ed. G.W.
Austin, ed. Igaku-Shoin, New York, pp. 53, 62, 67.
Dorland, W.A. 1985 Dorland’s Illustrated Medical Dictionary, 26th
ed. J.P. Friel, ed. W.B. Saunders, Philadelphia.
Hensyl W.R. 1990 Stedman’s Medical Dictionary, 25th ed. William
and Wilkins, Baltimore.
Herren, R.Y., and L. Alexander 1939 Sulcal and intrinsic blood vessels of the human spinal cord. Arch. Neurol. Psychiatry, 41:678687.
Joosten, E.A., and A.A. Gribnau 1989 Astrocytes and guidance of
outgrowing corticospinal tract axons in the rat. An immunocytochemical study using anti vimentin and anti glial fibrillary
acidic protein. Neuroscience, 31:439-452.
Lillie, R.D. 1948 Histopathological Technic. Philadelphia, Blakiston
Co., pp. 196.
Macallister, A. 1889 Text Book of Human Anatomy. P. Blakiston &
Son and Co., Philadelphia, pp. 743, 745.
Masson P.J. 1929 J . Tech. Methods, 12:75-90.
Morris, H., and J.P. McMurrich 1907 Morris’s Human Anatomy. P.
Blakiston & Son and Co., Philadelphia, pp. 297, 765, 793.
Parkinson D. 1991 Human arachnoid septa, trabeculae, and “rogue
strands.” Am. J . Anat., 192:498-509.
Parkinson, D. 1995 Posterior septum of spinal cord: Normal developmental variations. FASEB J., 9: Abstract 1249.
Rutka J.T., G. Apodaca, R. Stern, and M.K.M. Rosenblum 1988 The
extracellular matrix of the central and peripheral nervous systems: Structure and function. J . Neurosurg., 69:155-170.
Sarnat, H.B. 1992 Regional differentiation of the human fetal
ependyma: Immunocytochemical markers. J . Neuropathol. Exp.
Neurol., 51:58-75.
Schafer, E.A., J. Symington, and T.H. Bryce 1908 Quain’s Elements of
Anatomy. Longmans, Green and Co., London, pp. 472-473.
Snow, D.M., D.A. Steindler, and J . Silver 1990 Molecular and cellular
characterization of the glial roof plate of the spinal cord and optic
tectum: A possible role for a proteoglycan in the development of
an axon barrier. Dev. Biol., 138:359-376.
Spillane J.D. 1981 The Doctrine of the Nerves. The History of Neurology. Oxford University Press, Oxford, New York, pp. 118.
H. and L. Alexander 1939 Vascular system of the human spinal cord.
Arch. Neurol. Psychiatry, 41 :659-677.
Suh, T., and Carpenter, M.B. 1984 Core Text of Neuroanatomy, 3rd
ed. Williams and Wilkins, Baltimore, pp. 59.
Todd E.M. 1983 The Neuroanatomy of Leonard0 da Vinci. Capra
Press, Santa Barbara, pp. 113.
Valentino, K.L., E.G. Jones, and S.A. Kane 1983 Expression of GFAP
immunoreactivity during development of long fiber tracts in the
rat CNS. Brain Res., 285:317-336.
Vesalii A. 1543 Humani Corporis Fabrica. Johanes Oporinus Printer,
Basel, pp. 331.
Wiley, J . 1986 International Dictionary of Medicine and Biology. Vol.
3. A Wiley Publication. John Wiley and Sons, New York, pp.
2577.
Williams, P.L., and R. Wanvick 1980 Gray’s Anatomy, 36th ed.
Churchill Livingston, Edinburg, pp. 867, 896, 1050.
Willis T. 1680 Opera Omnia Samvelem de Tournes. Caput XIII, p. 10,
and Caput XXIX, p. 175.
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