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The ultrastructure of normal and abnormal oligodendroglia.

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The Ultrastructure of Normal and Abnormal
Oligodendroglia’
SARAH A. LUSE
Departments of Anatomy and Pathology, and the Beaumont-May Institute
of Neurology, Washington University School of Medicine,
Saint Louis, Missouri
This review of the oligodendroglia is disposed principally in the white matter.
presented as a tribute to Professor Nor- In Hortega’s opinion, oligodendrocytes
mand Hoerr, under whose guidance the were analogous to the Schwann cells of the
author received her basic training in the peripheral nerves and related in some way
discipline and technics of neurohistology. to the formation and maintenance of the
By means of metallic impregnations, myelin sheath, irrespective of what other
Ramon y Cajal (’13) separated from the functions they might possess.
astrocytic glia a group of small cells that
Participation of the oligodendroglia in
had not previously been stained selectively neoplasia was first suggested by Bailey and
by any method then known. These aden- Hiller (’24) and two years later confirmed
dritic cells he called the “third element” in by Bailey and Cushing (’26). Bailey and
the central nervous system. Del Rio- Bucy (’29) were able to prove by the
Hortega (’19, ’21) devised a silver carbo- silver impregnation methods of Hortega
nate method that selectively stained the that these tumors actually were formed of
“third element of Cajal” and demonstrated oligodendrocytes. Little can be added to
that it was composed of two distinct cell their description. In hematoxylin and
types : oligodendroglia and microglia. eosin or similar stains, they are characterHortega credited Robertson (1899, ’00) ized by numerous peculiarly uniform cells
with the first description of the oligoden- with ovoid nuclei centrally placed in clear
droglia and also equated them with the cytoplasmic compartments. The margins
adendritic cells of Cajal. The historical of the clear halos about the nuclei stain
developments in this field have been re- densely and have been considered fine
viewed by both del Rio-Hortega (’28) and processes of either the oligodendroglia or
of stromal astrocytes. Bailey and Bucy
Glees (’55).
Del Rio-Hortega (’21, ’28) described the (’29) and other observers have pointed out
oligodendroglia as of variable size, and of the astrocytic stroma of these tumors as
round, polyhedral or pyriform shape. Their well as the transitional neoplastic forms
nuclei were round, and usually eccentri- that have both astrocytic and oligodendrogcally located with an accumulation of lial characteristics.
The role of the oligodendroglia in cerecytoplasm at the pole from which the
thickest process arose. Although Hortega bral edema has been more gradually
stressed the paucity of both cytoplasm and recognized. Although Weed and McKibben
processes, Penfield (’24, ’30) demonstrated (’19) described the accentuation of perithat this had been due to incomplete im- vascular spaces in experimental swelling
pregnation. Both authors described the of the brain, they did not equate these
oligodendrocytes as arranged in rows be- changes with an alteration in oligodendrogtween the nerve fibers of the white matter lial cells. Penfield and Cone (’26a, ’26b)
(fig. 1); as satellites to neurons (fig. 2 ) ; and Cone (’28) studied the oligodendrogand as vascular satellites (fig. 3). Their lia in a variety of experimental and cliniubiquitous distribution, however, was in
1This work was supported in part by grants
contrast to the protoplasmic astrocytes B-1539
and B-425 from the United States Public
which were restricted to the gray matter Health Service, the National Institutes of Health,
and to the fibrous astrocytes which were Bethesda, Maryland.
461
462
SARAH A. LUSE
cal conditions and observed that the earliest manifestations of anoxia or toxicity in
the central nervous system was an acute
swelling of the oligodendrocytes. Unfortunately, these cells responded in a similar
manner to autolysis. Courville ('42), Perret and Kernohan ('43) and Reid ('43),
among others, have all ascribed at least
the early changes in edema of the brain to
swelling of the oligodendrocytes, although
they could not exclude the possibility of an
additional accumulation of extracellular
fluid. Recently electron microscopy has
shown that edema of the brain is associated with an increase in volume of the
oligodendroglial cells without evidence of
increased extracellular fluid (Luse and
Harris, '59, '60; Torack, Terry and Zimmerman, '59, '60). Gerschenfeld, Wald,
Zadunaisky and de Robertis ('59) also
demonstrated an increase in volume of
the pale cells in edema, although they
identified these cells as astrocytes.
In contrast to neoplasia and edema, the
role of oligodendroglia in dehydration,
metabolic alterations and demyelination
has remained little understood.
MATERIALS AND METHODS
The tissues on which the present study
is based were obtained from a variety of
sources. Blocks were prepared from the
cerebral cortex and cerebellum of adult
cats; the spinal cord and cerebral cortex
of adult rabbits; the spinal cord, medulla
and cortex of young and adult mice and
rats and from the spinal cord of chicks.
Specimens from human cerebral cortex
and from oligodendrogliomas were obtained at the time of surgery. Rabbit brains
in which edema or dehydration had been
produced experimentally were also examined, as were the spinal cords of rabbits with experimental allergic encephalomyelitis.
Small blocks of tissue for electron microscopy were fixed one to two hours in a
1 % solution of osmium tetroxide in potassium dichromate, adjusted to a pH of 7.4
as recommended by Dalton ('55). Occasionally tissue was fixed in potassium permanganate as described by Luft ('56).
Dehydration was in graded concentrations
of ethanol (10% to 100% ) and infiltration was with a 7: 1 mixture of
butyl and methyl methacrylates. Tissues
were embedded in the partially prepolymerized methacrylate mixture to which
benzoyl peroxide had been added as a
catalyst, and then were hardened at 60°C
overnight. Some tissues were embedded
in epoxy resins (Araldite) according to the
procedure of Richardson, Jarrett and Finke
('60). Thin sections were cut on a PorterBlum microtome ('53) using glass knives,
and were mounted on collodion-covered
copper grids. Sections were examined in
RCA electron microscopes (models EMU2E, 3B, or 3C) without removing the
plastic. Original micrographs were made
at 1000 to 10,000 diameters and subsequently enlarged photographically. For
light microscopy, tissue was fixed in formalin and embedded in paraffin for staining. Thick sections of osmium fixed plastic
embedded material were treated with 5%
periodic acid to remove the osmium tetroxide and then stained with a solution of
1% azure A and 1% methylene blue at
an alkaline pH.
OBSERVATIONS
N o r m a l oligodendroglia
A d u l t . Interfascicular oligodendrocytes
in the white matter are arranged in short
or long rows between the myelinated fibers,
and tend to be smaller than either the
vascular or neuronal satellite cells. Although usually round, their cytoplasm is
plastic and may vary in shape depending
on the pressures from adjacent cells. Their
plasma membranes are smooth and usually
delicate so that they may be broken during
dehydration or embedding. Cytoplasm of
the interfascicular oligodendrocytes is of
two types. A majority have a pale cytoplasm (fig. 4 ) similar to that of satellite
cells and have scant small mitochondria,
scant Golgi membranes and only a small
amount of ergastoplasm. Interspersed between these pale cells are others with
similar rounded nuclei and outlines but
with a more dense cytoplasm (figs. 5 and
6 ) . Mitochondria and the Golgi apparatus
are similar in both types of oligodendroglia,
the difference in the two being that the
darker cells possess numerous small intracytoplasmic vesicles, some of which have
attached RNA granules. Ocasionally, entirely different cells with dense cytoplasmic
ULTRASTRUCTURE OF OLIGODENDROGLIA
fibrils, the fibrous astrocytes, are intercalated in the rows of oligodendrocytes but
are readily distinguished from the smoothy
outlined oligodendroglia such as those
demonstrated in figure 6.
In both the white and gray matter, oligodendrocytes are arranged so that either
their soma or processes are in intimate
contact with vessels, forming vascular
satellites. Hortega demonstrated their perivascular arrangement by the classic gold
and silver impregnations (figs. 2 and 3).
In electron micrographs pale oligodendroglial processes almost invariably contact
the capillary wall (fig. 14) and occasionally as in figure 9 practically surround it.
Between these processes that seemingly
only touch the vascular wall are the firmly
adherent astrocytic processes, the so-called
astrocytic feet (fig. 14). In fortuitous sections one can actually demonstrate that
the same cell is a satellite both to a vessel
and to a neuron (fig. 9).
A majority of the neuronal satellites
are oligodendroglia, although protoplasmic
astrocytes and microglia may also occur
in this position. Satellite oligodendrocytes
have uniformly round or oval nuclei with
a single small nucleolus. Their most characteristic feature, however, is abundant
pale cytoplasm with scant ergastoplasm
and a paucity of Golgi membranes or
mitochondria. Their plasma membranes
are sharply outlined and most cells tend
to be round in section with a single stout
process that may extend from the cell body
(fig. 9) or may branch dichotomously as
in figure 8. In the neuropil the round or
ovoid pale processes of oligodendrocytes
are, for the most part, readily distinguished
from other structures, but may be confused with unmyelinated axons if neurofilaments are not seen.
Not all oligodendrocytes of the gray matter have their cell bodies adjacent to either
vessels or neurons. Some may be isolated,
or arranged in short rows (fig. 7).
Whether or not their processes touch those
of other oligodendroglial cells, vessels, or
neurons, cannot be determined in thin
sections and can only be surmised.
The gliosomes of light microscopy apparently correspond to mitochondria since
no other organelles of the proper size are
present and since the gliosomes are stained
463
by specific mitochondrial stains. During
myelin formation electron opaque cytoplasmic structures occur that are not
mitochondria, but which may have constituted some of the gliosomes of light
microscopy (fig. 11 ). In addition, in adult
animals of all species examined, and
especially in the human being, there are
accumulations of dense irregularly outlined particles that we have equated with
the lipochrome pigment granules associated with aging (fig. 21).
Immature: during myelination. The
brains and spinal cords of rats and mice
one to 20 days of age were examined. No
attempt was made to investigate the development of oligodendroglia prior to the
time of birth or to identify their precursors.
During the fust three weeks of life
oligodendrocytes were more numerous
than in the adult and somewhat larger
with an irregular cytoplasmic contour.
Their abundant cytoplasm, although pale,
differed from that of the adult in their
cytoplasmic components. Mitochondria
were numerous and sometimes large. The
ergastoplasm (endoplasmic reticulum) in
some cells was arranged in distinct parallel
arrays, whereas in others it was as inconspicuous as in adult cells. Occasionally
large round to oval dense bodies were
present as cytoplasmic inclusions, but their
significance is unknown (fig. 11). Rare
cytoplasmic lipid droplets occurred. Expansions arising from oligodendroglia were
broad based, but distant from the cell body
it was difficult if not impossible to distinguish them from unmyelinated axons in
the one- to three-day-old animal. As myelination proceeded, this distinction was
more readily made. In the 5- to 14-day-old
animal, the voluminous pale expansions
were often in close apposition to or surrounded the myelinating axon as has been
described previously (Luse, '56b, '59).
Vesicles were numerous in the terminal
portions of oligodendroglial processes surrounding myelinating axons (fig. 12). Although the story of myelin formation is
still incomplete, the presence of vesicles
adjacent to myelinating fibers, both centrally and peripherally, suggests that the
myelin lamellae are formed, at least in
part, by fusion of cytoplasmic membranesurrounded vesicles which later become
464
SARAH A. LUSE
organized to form uniform myelin membranes. The possibility of overlapping glial
processes and their membranes contributing to the sheath has also been postulated
(Luse, '56b).
A b n m a l oligodendroglia
Neoplasia. The tumors which we examined were all well differentiated oligodendrogliomas of the classical light microscopic appearance (fig. 18). At low magnifications, electron micrographs presented
a picture strikingly remininscent of that
seen with hematoxylin and eosin stains:
rather uniform cells containing round to
oval nuclei set in voluminous pale cytoplasm (figs. 18 and 19). The nuclear
chromatin varied from finely granular to
coarsely clumped and often formed a dense
rim at the inner margin of the nuclear
membrane. Although most nuclei had
round or oval outlines, some were bizarre.
Mitotic figures occurred. Nucleoli of tumor
cells were large, sometimes multiple, and
frequently had a unique punched out appearance (fig. 20).
Cytoplasmic membranes of tumor cells
were variable. Some were smooth and
sharply delineated while others had numerous short, delicate expansions that interlocked with those of adjacent cells (fig.
24). Tumor cells often were clumped, the
entire group being surrounded by stromalastrocytic processes or cells. Cytoplasm of
the neoplastic oligodendroglia was abundant and pale due to the paucity of ergastoplasm, which was only slightly more prominent than in normal cells. The Golgi
apparatus was rarely evident. Mitochondria of these cells often were vastly altered. Sometimes they were so numerous
that they actually filled the cytoplasm
(fig. 24), measuring as much as three
CI in width. In addition there were
cells with characteristic oligodendroglial
nuclei that had cytoplasmic fibrils or even
processes resembling those of astrocytes.
Other cells with reniform or indented nuclei and astrocyte-like processes possessed
the clear cytoplasm and numerous mitochondria of neoplastic oligodendroglial
cells. The end result is that with the resolution and magnifications of electron microscopy, one still finds a few cells in oligodendrogliomas that defy attempts to classify
them. Indeed, they possess characteristics
of both the oligodendrocyte and of the
astrocyte. This same dilemma was faced
by Hortega ('21) and by Penfield ('32),
who decided that there existed forms intermediate between oligodendrocytes and
astrocytes.
Dense irregular lipid droplets occurred
in some tumor cells. An occasional cell
was almost filled with fine or coarse extremely electron dense granules (fig. 22)
which corresponded with the presence of
calcium in contiguous thicker sections.
Mitotic figures were not infrequent (fig.
23).
Edema and dehydration
Electron microscopy of the swollen
brains of rabbits given intravenous distilled water revealed a marked increase in
volume of oligodendroglial cytoplasm, both
in the cell body and in its processes. This
was particularly distinct about the vessels,
where the distended perivascular space of
light microscopy proved to be dilated oligodendroglial processes (fig. 15). With more
severe swelling, cytoplasmic expansion of
oligodendrocytes was sufficient to rupture
cell membranes and to compress both
astrocytic and neural processes. Spontaneous edema at the margins of metastatic
tumors in the human being showed the
same picture of oligodendroglial swelling
without extracellular accumulation of
fluid.
With injection of hypertonic solutions
('50% sucrose, 25% glucose, 30% urea or
50% sorbitol), the cytoplasm of oligodendroglia was markedly shrunken (fig. 16).
Nuclei of oligodendrocytes apparently were
unaltered in dehydration, but their cytoplasm was reduced to a mere perinuclear
rim. Cytoplasmic organelles were concentrated in the residual cytoplasm so that it
was no longer pale. Similarly reduced in
volume were the oligodendroglial processes
about vessels and in the neuropil. With
rehydration the cytoplasm of oligodendrocytes re-expanded.
Injection of 5 % glucose intravenously
produced little, if any, change in the appearance of the brain. In contrast excess
intravenous 0.9% saline produced an unequivocal change in oligodendroglial cells.
Numerous fine cytoplasmic tonofibrils ap-
ULTRASTRUCTURE OF OLIGODENDROGLIA
peared in the cytoplasm of almost all
oligodendrocytes (fig. 17). The significance of this observation is not known,
nor has the reaction of oligodendroglia to
other salts yet been investigated.
Demyelination: (experimental allergic
encephalomyelitis)
Allergic encephalomyelits is a disease in
which demyelination occurs with sparing
of the axis cylinders. Oligodendrocytes are
distinctly altered in this disease. They
are somewhat increased in volume. Their
mitochondria are swollen, whereas mitochondria of adjacent microglial cells or
axons are normal (fig. 26). Ergastoplasmic sacs of the oligodendroglia may also be
distended. It appears that the primary
insult in allergic encephalomyelitis may
well be to the oligodendrocyte, and that
the myelin disintegration is secondary to
the oligodendroglial lesion (Luse and McDougal, '60).
DISCUSSION
The importance of oligodendroglia in
the maintenance of the integrity of the
nervous system has become increasingly
apparent in recent years. Although Cajal
postulated that they were in symbiosis
with neurons, Hortega believed them to be
the Schwann cells of the central nervous
system and related to formation and maintenance of the myelin sheath. Both views
have by now been amply verified.
Penfield ('24, '30, '32) in particular has
amplified the work of Hortega and substantiated his observations on the role of
the oligodendrocyte in myelin formation;
while at the same time Bailey extended
the silver technics to neoplasia (Bailey and
Hiller, '24; Bailey and Cushing, '26; and
Bailey and Bucy, '29).
Electron microscopy has opened a new
field in the area of morphology. The
identification of the various cellular components of the nervous system at the
magnifications now available should be
relatively simple. The reverse, however,
appears to be the case, and a controversy
has arisen over the identification of glia,
cf. Farqhuar and Hartmann ('57), Gerschenfeld et al. ('59), de Robertis et al.
('60) and Schultz, Maynard and Pease
('57).
465
The cells with folded membranes and
dense cytoplasm have been identified by
Luse ('56a, '58), Palay ('57) and Dempsey
and Luse ('57) as astrocytes. They bear
a close resemblance to: ( 1 ) the cells in
astrocytic tumors; and (2) cells proliferating at the margins of stab wounds, abscesses or injections of alumina gel, cells
that may also be impregnated by the gold
chloride-sublimate method of Cajal; and
( 3 ) many of the cells forming the pia-glial
membrane of the mouse optic nerve and
of the cat cortex. Furthermore, the cells
equated by Luse ('56a, '56b, '59) and
Palay ('57) with oligodendroglia (the
satellite cells of the neurons and vessels
and the rows of cells in the white matter)
are strikingly similar to the cells of the
5 clear-cut oligodendroglial tumors that
we have examined. The close investment
of myelinating axons by pale cell cytoplasm (Luse '56b, '59); the degenerative
changes in these same cells in allergic
encephalomyelitis; and their increase in
volume in cerebral edema further point
to their being oligodendroglia.
An exactly opposite evaluation of the
glial cells has been arrived at by Farqhuar
and Hartmann ('57), de Robertis, Gerschenfeld and Wald ('60), Gerschenfeld et
al. ('59) and Schultz, Maynard and Pease
('57). Their opinions have been based in
part on the abundance of pale processes
that abut upon the blood vessels and that
they have interpreted as astrocytic feet.
Actually, oligodendroglia make as many, if
not more, contacts with vessels than
astrocytes, as Hortega ('21, '28) pointed
out. Hortega did not consider these contacts of oligodendroglia and vessels to be
as firmly adherent as the astrocytic "sucker
feet" although Ferraro and Davidoff ('28)
apparently did. Furthermore, these investigators have considered the pale processes at the pia-glial membrane as necessarily being astrocytic feet. However, their
work has mostly been with the rat, rather
than the cat, monkey or man. The piaglial membrane in the latter has a higher
density of definite astrocytes than in the
mouse, rat or rabbit, excepting the optic
nerve. The astrocytes forming the pial
surface of the brain are specialized in type
and less complex than elsewhere, nor are
all cells reaching the surface of the brain
466
SARAH A. LUSE
necessarily astrocytes. Indeed, both Hortega ('21, '28) and Bailey and Schaltenbrandt ('27) have contended that the
oligodendrocyte extends to and contributes
to the pia-glial membrane.
In edema of the brain, experimental or
spontaneous, (Luse and Harris, ' 6 0 ) , the
pale cells considered to be oligodendrocytes by us, have been massively increased
in volume. Gerschenfeld et al. ('59) and
de Robertis, Gerschenfeld and Wald ('60)
have also seen swelling of pale cells in
edema of the brain, but have identified
them as astrocytes basing this in part on
the recent work of Klatzo et al. ('57).
Klatzo's identification of edema as being
an astrocytic change is at variance with
an extensive literature on this subject:
Courville ('42), Hassin ('28), Scheinker
('41, '47), Perret and Kernochan ('43)
Evans and Scheinker ('45), as well as the
experimental work of Reid ('43), Penfield
and Cone ('26a, '26b) and Cone ('28).
All of these investigators demonstrated
swelling of oligodendrocytes in cerebral
edema and Reid ('43) and Cone ('28) also
demonstrated the swollen oligodendrocytes
in edema by the silver carbonate stains of
Hortega.
Electron microscopy has revealed the
lack of an extracellular fluid compartment
in edema of the brain (Luse and Harris,
'59, '60; Gerschenfeld et al., '59; Torack,
Terry and Zimmerman, '59, '60). Even
more remarkable is the fact that the only
cellular element that undergoes detectable
change in edema or dehydration is the
oligodendrocyte.
The function of the neuroglial cells has
tantalized both the morphologist and the
chemist. The supportive functions of the
fibrous astrocyte are clear enough; but
little is known of the role of the protoplasmic astrocyte, nor of the oligodendroglial cells. Electron microscopy has confirmed the beliefs and observations of del
Rio-Hortega and of Penfield as to the close
association of oligodendrocytes with the
formation of myelin (figs. 12 and 13).
Electron microscopy of the spinal cord in
allergic encephalomyelitis has further
strengthened this relation by showing that
the first lesion in myelin destruction is a
degenerative alteration in the oligodendrocyte. Greenfield ('33) and Adams and
Kubik ('52), among others, had suggested
that some clinical types of demyelinating
diseases might be related to changes in the
interfascicular cells.
Cajal's concept of symbiosis of oligodendrocytes and neurons also has been amplified by electron microscopy. In the central
nervous system, cells and cellular processes are so closely wedged together that
the distances between any two are of the
order of 100 to 200 Angstrom units, with
a resulting extracellular space calculated
by Horstmann and Meves ('57) as not
exceeding 3 to 5% ; whereas the chloride
space had been calculated as approximately 30 to 35% (Woodbury, '57). Thus,
no extracellular space is available in the
central nervous system for transport of
fluid or electrolytes, the most likely site of
such transport being the oligodendroglial
cell. The oligodendroglial cell expands in
edema and contracts with dehydration;
and since it makes both vascular and neuronal contacts (fig. 9 ) forms a possible
channel for cellular exchange by neurons
otherwise isolated from the blood vascular
system.
Although little has yet been done in the
areas of enzyme or microchemistry of the
glial cells, some interesting differences
have been suggested. Kluver ('55) has
described a high porphyrin content in the
nervous system, apparently located in the
oligodendrocytes. C avanaugh, Thompson
and Webster ('54) have demonstrated
large amounts of pseudocholinesterase in
the white matter which they belive to be in
oligodendrocytes; and Ashby and coworkers ('52) an elevated carbonic anhydrase. These changes could indicate a
role in the carbon dioxide or oxygen transport systems of the cell or in membrane
permeability. In addition, Pope, Hess and
Allen ('55), Victor and Wolf ('37) and
Heller and Elliott ('55) have shown that
the oxygen consumption of oligodendroglial tumors is considerably higher than
that of astrocytic tumors and that mixed
tumors with an increased number of
oligodendrocytes have an increased level
of cytochrome oxidase.
The findings in tissue culture of both
normal and neoplastic oligodendroglial
cells has also been informative. Canti,
Bland and Russell ('37) first cultured the
ULTRASTRUCTURE OF OLIGODENDROGLIA
oligodendroglioma. Normal oligodendrocytes have shown a similar morphology
and a similar pulsatile activity which
distinguishes them from the astrocytes
(Lumsden and Pomerat, '51; Pomerat, '57).
Benitez, Murray and Wooley ('55) and
Murray ('57) have added to this their
demonstration, in vitro, of the tonic contraction response of oligodendrocytes to
the addition of serotonin, which suggests
that the metabolically active oligodendrocyte may have a pumping action which is
stimulated by serotonin.
SUMMARY
Normal and pathologic oligodendroglia
have been studied. They are usually pale
round cells arranged in rows in the white
matter or as satellites to neurons and
vessels. During the time of myelin formation and in the immature animal, they are
more voluminous than in the adult and are
intimately associated with the rnyelinating
axons. In experimental or spontaneous
cerebral edema, oligodendroglial cytoplasm
is dilated and conversely in dehydration
the oligodendroglial cytoplasm shrinks. In
allergic encephalomyelitis the earliest lesion apparently is a degenerative change
in the oligodendrocyte with swelling of
both mitochondria and ergastoplasm, and
secondarily degeneration of myelin. Oligodendroglial tumors are composed of cells
reminiscent of normal oligodendrocytes
with voluminous pale cytoplasm and round
or bizarre nuclei. It is suggested that the
oligodendrocytes subserve at least two
functions: (1) formation of the myelin
sheath, and (2) a pathway for fluid and
electrolyte transport within the nervous
system in lieu of an extraceUuIar space.
ACKNOWLEDGMENTS
It is a pleasure to acknowledge the collaboration of Dr. Basil Harris in the studies on edema and dehydration of the brain.
I am also indebted to Miss Ann Jones and
Miss Helen Ferguson for their technical
assistance.
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Biochem. Physiol., 33: 395-403.
Pomerat, C. M. 1957 Functional concepts based
Horstmann, E., and H. Meves 1957 Die Feinon tissue culture studies of neuroglia cells.
struktur des molekularen Rindengraves und
In: Biology of Neuroglia, ed. W. F. Windle.
ihre physiologische Bedeutung. Ztschr. ZellCharles C Thomas, Springfield, Ill., pp. 167forsch., 49: 569-604.
175.
Klatzo, Igor, A. Piraux and E. J. Laskowski 1958 Pope, A., H. H. Hess and J. N. Allen 1957
The relationship between edema, blood-brain
Quantitative histochemistry of proteolytic and
barrier and tissue elements in a local brain
oxidative enzymes in human cerebral cortex
injury. J. Neuropath. Exp. Neur., 17: 548-564.
and brain tumors. Chapter 10 in: Progress in
Kliiver, H. 1955 Porphyrins in relation to the
Neurobiology, vol. 2, “Ultrastructure and Celludevelopment of the nervous system. In: Biolar Chemistry of Neural Tissue,” ed. H.
chemistry of the Developing Nervous System,
Waelsch. P. B. Hoeber, New York.
H. Waelsch, ed. Academic Press, New York.
Porter, K. R., and J. Blum 1953 A study in
Luft, J. H. 1956 Permanganate-A new fixamicrotomy for electron microscopy. Anat. Rec.,
tive for electron microscopy. J. Biophys. Bio117: 685-710.
chem. Cytol., 2: 799-801.
Ramon y Cajal, S. 1913 Sobre u n nuevo proLumsden, C. E., and C. M. Pomerat 1951
ceder de impregnacion de la neuroglia y sus
Normal oligodendrocytes in tissue culture. Exp.
resultados en 10s centros nerviosos del hombre
Cell Res., 2: 103-114.
y animales. Trab. Lab. Inv. Biol. Univ. Madrid,
Luse, S. A. 1956a Electron microscopic observa11, 219.
tions of the central nervous system. J. Biophys.
Reid, W. L. 1943 Cerebral oedema. Austral.
Biochem. Cytol., 2: 531-541.
N. Zealand J. Surg., 17: 439446.
19561, Formation of myelin in the central nervous system of mice and rats, as stud- Richardson, K. C., L. 0. Jarrett and E. H. Finke
1960 Notes on the use of Araldite epoxy resins
ied with the electron microscope. Ibid., 2:
for ultra-thin sectioning in electron microscopy.
777-784.
Stain Tech., in press.
1958 Ultrastructure of reactive and
neoplastic astrocytes. Lab. Invest., 7: 401-417. del Rio-Hortega, P. 1919 El “tercer elements”
1959 The fine structure of the morphode 10s centros nerviosos. I. La microglia e n
genesis of myelin. Progress in Neurobiology,
estado normal. 11. Intervencion de la microg4: 59-81.
lia e n 10s procesos patologicos. 111. NaturLuse, S. A., and B. Harris 1959 Electron mialeza probable de l a microglia. Bol. SOC. Esp.
croscopy of the brain in experimental edema.
Biol., 9: 69.
Anat. Rec., 133: 305.
1921 Estudios sobre la neurolgia. La
1960 Electron microscopy of the brain
glia de escasas radiaciones (oligodendroglia).
in experimental edema. J. Neurosurg., 17:
Bol. Real SOC.Expan. Hist. Nat., 21: 63-92.
439-446.
1928 Tercera aportacion a1 conocimiLuse, S. A., and J. B. McDougal, Jr. 1960
ento morfologico e interpretacion funcional de
Electron microscopic observations on allergic
la oligodendroglia. Mem. Real SOC. Expan.
encephalomyelitis in the rabbit. J. Exp. Med.,
Hist. Nat., 14: 1-122.
in press.
de Robertis, E. D. P., H. &I. Gerschenfeld and
Murray, M. R. 1957 Response of oligodendroF. Wald 1960 Some aspects of glial function
cytes to serotonin. I n : Biology of Neuroglia,
as revealed by electron microscopy. In:
ed. W. F. Windle. Charles C Thomas, SpringFourth International Conference on Electron
field, Illinois, pp. 176-180.
Microscopy, Springer-Verlag, Berlin, pp. 443Palay, S. L. 1957 An electron microscopical
447.
study of neuroglia. Ibid., pp. 24-38.
Robertson, W. F. 1899 On a new method of
Penfield, W. 1924 Oligodendroglia and its relaobtaining a black reaction in certain tissuetion to classical neuroglia. Brain, 47: 430-452.
elements of the central nervous system (plat1930 Further modification of del Rioinum method). Scott. Med. Surg. J., 4: 23Hortega’s method of staining oligodendroglia.
1900 A microscopic demonstration of
Am. J. Path., 6: 445448.
the normal and pathologic histology of mesog1932 Neuroglia and microglia. The inlia cells. J. Ment. Sci., 46: 724.
terstitial tissue of the central nervous system.
In: Special Cytology, ed. E. C. Cowdry. Paul Scheinker, I. 1941 Cerebral swelling and edema
associated with cerebral tumor. A histogenetic
B. Hoeber, N. Y., 2nd ed., pp. 1447-1482.
and histopathologic study. Arch. Neur. PsyPenfield, W., and W. Cone 1926a Acute swellchiat., 45: 117-129.
ing of oligodendroglia. A specific type of neuroglia change. Arch. Neur. Psychiat., 16: 131- Scheinker, I. M. 1947 Cerebral swelling. Histopathology, classification and clinical signifi153.
cance of brain edema. J. Neurosurg., 4: 25519261, The acute regressive changes of
275.
neuroglia (ameboid glia and acute swelling of
oligodendroglia). J. f. Psych. Neur., 34: 204- Schultz, R. L., E. A. Maynard and D. C. Pease
224.
1957 Electron microscopy of neurons and
Perret, G. E., and J. W. Kernohan 1943 Histoneuroglia of cerebral cortex and corpus callopathologic changes of the brain caused by
sum. Am. J. Anat., 100: 368408.
-
-
-
-
-
ULTRASTRUCTURE OF OLIGODENDROGLIA
Torack, R. M., R. D. Terry and H. M. Zimmerman
1959 The fine structure of cerebral fluid accumulation. I. Swelling secondary to cold
injury. Am. J. Path., 35: 1135-1147.
1960 The fine structure of cerebral
fluid accumulation. 11. Swelling produced by
triethyl tin poisoning and its comparison with
that in the human brain. Ibid., 36: 273-287.
469
Victor, J. V., and A. Wolf 1937 Metabolism of
brain tumors. Res. Publ. Assoc. Res. New.
Merit. Dis., 1 6 : 44-58.
Weed, L. H., and P. S. McKibben 1919 Experimental alteration of brain bulk. Am. J. Physiol., 48: 531-558.
Woodbury, D. M. 1957 I n : Biology of Neuroglia, ed. W. F. Windle. Charles C Thomas,
Springfield, Ill., pp. 120-126.
PLATE 1
EXPLANATION OF FIGURES
470
1
This is a copy of figure 2, Wortega (’21), “Neuroglia de la substancia blanca del cerebro
humano, tenida por el metodo aurjco de Cajal. A, glia interfascicular de escasas radiaciones; B, gliocio fibroso de largas radiaciones.” It demonstrates oligodendroglial cells
“A” arranged in rows in the white matter and their lack of processes in the gold
chloride-sublimate stain, whereas radiations of astrocyes “B” are depicted.
2
This is a copy of figure 1 from Hortega (’21). “Neuroglia de la corteza cerebral humana,
tenida por el metodo aurico de Capal: A oligodendroglia; B, gliocito protoplasmic0 o de
cortas radiaciones; C, nucleo de microglia.”
3
This a copy of figure 66 from Hortega (’21). “Substancia blance de una laminilla cerebeloso del mono. Junto a 10s vasos existen abundantes gliocitos con escasas radiaciones,
bastante largas y rainificadas.” This demonstrates the difference in staining with the
silver carbonate method.
ULTRASTRUCTURE OF OLIGODENDROGLIA
Sarah A. Luse
PLATE 1
471
PLATE 2
EXPLANATION O F FIGURES
4 Electron micrograph of a small oligodendroglial cell i n the white matter of the brain
of a n adult mouse. Numerous myelinated axons are present. The cytoplasm is pale
with scant mitochondria, ergastoplasm or Golgi membranes.
5
x
9,000.
Numerous large and small myelinated fibers surround this oligodendrocyte in the white
matter of the brain of a n adult albino rat. The oligodendrocyte is almost round with a
single process evident at its upper margin (arrow). The round nucleus is centrally
located. The cytoplasm of this cell is in contrast to figure 4, in that numerous vesicles,
some associated with RNA granules, are present. X 4,000.
6 Electron micrograph from spinal cord of a newborn chick. Parts of 4 cells are lined up
in a row between myelinated axons. A margin of myelin is evident at the arrows at
the bottom of the figure. Two of the cells, A and C, have dense cytoplasm, and two
others, B and D, have pale cytoplasm. The cytoplasm of C is of intermediate density
and is readily recognizable as olgiodendroglial, whereas that of A is so dense that its
possible oligodendroglial origin is suggested by position only. X 7,500.
4 72
ULTRASTRUCTURE OF OLIGODENDROGLIA
Sarah A. Luse
PLATE 2
473
PLATE 3
EXPLANATION OF FIGURE
7
474
Two oligodendroglial cells deep in the cortex of the brain of an adult rabbit. Part of a
neuron is present at the left. The oligodendroglial nuclei are almost round. The cytoplasm is abundant, pale, and has few organelles. The upper cell has two blunt processes
and the lower one a single process (arrows). X 10,000.
ULTRASTRUCTURE OF OLIGODENDROGLIA
Sarah A. Luse
PLATE 3
475
PLATE 4
EXPLANATION O F FIGURES
8
This is a n electron micrograph of a perineuronal satellite. The oligodendrocyte is surrounded by two dendrites ( D ) and is not separated from them by any other cell or cell
process. The oligodendroglial nucleus is round and its cytoplasm pale. The cytoplasmic
organelles are, for the most part, concentrated near the zone where the two oligodendroglial processes (arrows) are arising. Rabbit cortex. x 6,000.
9 Electron micrograph of oligodendrocyte (oligo) that is a satellite to a neuron and also
is i n contact with a blood vessel ( V ) . Rabbit cortex.
476
x
4,500.
ULTRASTRUCTURE OF OLIGODENDROGLIA
Sarah A. Luse
PLATE 4
477
PLATE 5
EXPLANATION O F FIGURES
10 Electron micrograph of oligodendrocyte i n the medulla of a mouse 6 days after birth.
The cytoplasm is more abundant than in similar cells from mature animals and contains
more cytoplasmic organelles. The number of cytoplasmic vesicles with or without RNA
granules is greater than in cells from adult animals. X 6,500.
11
478
Electron micrograph of oligodendroglial process from spinal cord of a one-day-old mouse.
The prominent granular component of the cytoplasm is evident. In addition, three dense
aggregates are present (arrows). Their significance is unknown, but they are seen
almost exclusively i n immature animals. These aggregates are readily distinguished
from the lipofuchsin granules seen i n aged animals or i n humans (fig. 21). >( 16,000.
ULTRASTRUCTURE OF OLIGODENDROGLIA
Sarah A. Luse
PLATE 5
479
PLATE 6
EXPLANATION O F FIGURES
12 Electron micrograph of a n axon which is i n the process of myelination. Glial cytoplasm
(G) is loculated between the axolemma and the outer layers of myelin. Surrounding
the outermost myelin lamella are at least oligodendroglial processes, some of which
contain numerous vesicles. Their plasma membranes are evident at the arrows. An
oligodendroglial nucleus is present a t the left ( N ) . X 12,000.
13
Electron micrograph of medulla of an adult mouse. A myelinated axon is present in
the center of the field and is in close apposition to a large oligodendroglial process
(oligo). Numerous neuronal processes that are approaching their terminations are present ( N ) and are recognizable by their numerous vesicles. Embedded i n Araldite and
stained with potassium pcrmanganate. X 12,000.
ULTRASTRUCTURE OF OLIGODENDROGLIA
Sarah A. Luse
PLATE 6
481
PLATE 7
EXPLANATION
482
O F FIGURES
14
Capillary in the brain of a n adult rabbit. Much of the outer surface of the vessel is
covered by astrocytic process (arrows). Three oligodendroglial process ( O L ) are
present at the lower right. :
:9,000.
15
Cerebral edema i n a n adult rabbit given intravenous distilled water. The capillary is
surrounded by distended oligodendroglial processes. Mitochondria (M) are still present.
Plasma membranes ( P ) are evident. The neuropil actually has been compressed by the
distended oligodendroglial processes. X 6,000.
ULTRASTRUCTURE OF OLIGODENDROGLIA
Sarah A. Luse
PLATE 7
483
PLATE 8
EXPLANATION O F FIGURES
16 Electron micrograph of a n oligodendrocyte in the brain of a rabbit that had been given
intravenous 50% sucrose. The oligodendroglial cytoplasm is shrunk to only a narrow rim
about the nucleus. A process is extending out a t the lower left (PR) and most of the
organelles of the cell are concentrated in this region. x 12,000.
17 Oligodendrocyte from the brain of a rabbit that had received 300 cm3 of IV normal
saline. The cytoplasm (arrows) contains numerous delicate fibrils. X 12,000.
484
ULTRASTRUCTURE OF OLIGODENDROGLIA
Sarah A. Luse
PLATE 8
485
PLATE 9
EXPLANATION O F FIGURES
18 Electron micrograph of a n oligodendroglioma (light micrograph is shown i n insert at
upper left). The cells are irregular with delicate plasma membranes and ovoid to
round nuclei. The cytoplasm is somewhat pale with scant organelles. X 5,000.
19
486
Electron micrograph of another region from the same tumor as that in figure 18. The
pale cytoplasm is more voluminous in this group of cells. Part of a blood vessel is
evident at the upper right (vessel). X 5,000.
ULTRASTRUCTURE OF OLIGODENDROGLIA
Sarah A. Luse
PLATE 9
487
PLATE 10
EXPLANATION O F FIGURES
4aa
20
Electron micrograph of part of the nucleus of a tumor cell in another oligodendroglioma.
The nucleolus is abnormally large with a number of punctate pale zones. x 10,000.
21
Lipochrome pigment in the cytoplasm of a normal oligodendrocyte in the brain of a n
adult human being. X 10,000.
22
Cytoplasm of part of a neoplastic oligodendroglial cell from the same tumor as figures
18 and 19. Dense irregular aggregates i n the cytoplasm (arrows) are one of the manifestations of calcium deposition in these cells. X 10,000.
23
Mitotic figure occurring in an oligodendroglioma. The nuclear membrane has disappeared. The cytoplasm is more dense than usual. Mitochondria are not seen. The cell
is in metaphase, and has been sectioned equatorially through the metaphase plate.
x 10,000.
ULTRASTRUCTURE OF OLIGODENDROG1,IA
Sarah A. Luse
PLATE 10
489
PLATE 11
EXPLANATION OF FIGURES
490
24
Electron micrograph of parts of 4 cells i n a n oligodendroglioma. This figure illustrates two points: in the center the fine process of adjacent cells are interdigitated with
each other, and at the upper right the cytoplasm of one cell is almost filled with normal
sized mitochondria. X 12,000.
25
Electron micrograph of another oligodendroglial tumor cell in which some of the mitochondria are of normal size and others are abnormally large ( M ) . In the lower central
mitochondrion (arrow) there are a few cristae extending across the cell. In the two
upper large ones there are lipid inclusions. X 12,000.
ULTRASTRUCTURE OF OLIGODENDROGLIA
Sarah A. Luse
PLATE 11
491
ULTRASTRUCTURE OF OLIGODENDROGLIA
Sarah A. Luse
26
492
PLATE 12
Electron micrograph of a n oligodendroglial cell in allergic encephalomyelitis. The nucleus is normal. The cytoplasm is increased in amount. The mitochondria ( M ) and
ergastoplasmic sacs ( E ) are swollen. A few fragmented mitochondria1 cristae remain
(arrows). The microglial phagocytic cell at the upper right contains myelin debris and
mitochondria ( M ) that are normal. x 10,000.
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