Paper - The Development of the Scala Tympani, Scala Vestibuli and Perioticular Cistern in the Human Embryo
Embryology - 19 Apr 2024    Expand to Translate
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Streeter G.L. The Development of the Scala Tympani, Scala Vestibuli and Perioticular Cistern in the Human Embryo
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The Development of the Scala Tympani, Scala Vestibuli and Perioticular Cistern in the Human Embryo
Department of Embryology, Carnegie Institution of Washington, Johns Hopokins Medical School, Baltimore, Maryland
Nine Figures
The study of the development of the large walled—off connective tissue spaces that surround the membranous labyrinth
is particularly interesting in that it shows that they have a very
definite morphological individuality. It is evident at least
that they are not to be considered as insignificant accessories
that merely fill in the waste intervals between the membranous
labyrinth and the surrounding cartilage or bone. On the contrary, they have characteristics which are in many respects
as definite and constant as ‘those of the ossicles themselves. The
individuality of these spaces in all respects is most marked.
They make their appearance at a definite stage in the development of the embryo; they are formed at definite places; they
pass through a series of definite histogenetic processes; they
spread in a definite order and manner and eventually they attain a definite form and structure. The general morphology
and relations of these‘ spaces during their developmental period
will be described in the following paper, and the opportunity
will be taken to point out in the course of the description some
of these individualistic features.
Instead of designating the large spaces surrounding the membranous labyrinth as perilymphatic spaces, as has been the
general custom since the time of Breschet 1833, they will here
be spoken of as perioticular or periotic spaces. The use of the
term ‘periotic’ avoids the confusion arising from the incorporation
of the Word ‘lymphatic’ in the terminology. The present tendency is to restrict the use of the word ‘lymphatic’ to the lymphatic vascular system and its associated structures, with which
these particular spaces have no known connection, either in
their origin or in their ultimate relations.1 We shall therefore
speak of a periotic connective tissue that everywhere surrounds
the epithelial portion of the labyrinth. This connective tissue
includes, in part the fine-meshed periotic reticulum, and in
part the large walled-off perioticular spaces to which belong the
vestibular cistern, the scala Vestibuli and the scala tympani
with whose development we are primarily concerned. V
Material and Methods
The observations that are recorded in this paper are all based on human embryos and cover the period included between embryos 35 mm. and 130 mm. CR length, which is approximately equivalent to the period between the ninth and sixteenth week of fetal life.
To facilitate the determination of the form and relations of the spaces, wax-plate models of the membranous labyrinth and the surrounding spaces were reconstructed after -the Born method. Advantage was taken of the improvements in the method recently devised by Lewis 1915.9 The serial sections were photographed at a suitable enlargement on bromide paper. By means of a preliminary model of the membranous labyrinth, the necessary reconstruction lines were established and inscribed on the bromide prints. From these prints then the membranous labyrinth and the perioticular spaces were traced on waxplates. After cutting out from the plates the areas corresponding to these structures, the plates were piled and the resultant cavities were filled with plaster of Paris. The wax was finally melted off and there was left then a permanent plaster cast of the objects desired at a definite enlargement. Views of these models are shown in figures 4 to 9.
‘Sabin, F. R. Harvey Society Address. Science, vol. 44, 1916, p. 145.
9 Lewis, W. H. The use of guide planes and plaster of Paris for reconstructions
from serial sections. Anat. Rec., vol. 9, 1915.
In outlining the periotic spaces it was found necessary to make an arbitrary rule as to how much should be included in
the model. The smaller spaces of the reticulum that surrounds
the ‘main cavities can be seen coalescing to form larger spaces
and these in turn coalesce with the main cavity as it advances
into new territory. Thus in a given section there is a considerable range in the size and completeness of the spaces. The
main spaces and the larger adjacent ones that communicate with
them are outlined by a membrane—like border. This characteristic was utilized as the guide for determining which spaces to
admit into the model; only those possessing a more or less complete border of this kind were included.
Histogenesis of the Periotic Reticulum
Although this communication is more concerned with the process of conversion of the periotic reticular tissue into the larger walled—off spaces, yet for the purpose of completeness a brief survey will be taken of the earlier history of this tissue and the nature of its histogenesis.
The tissue in which the perioticular spaces develop is derived
from the condensed mesenchyme that establishes itself as an
encapsulating mass around the otic vesicle in embryos between
4 mm. and 10 mm. long. This condensed mesenchyme is subsequently differentiated into the cartilagenous capsule that
completely invests the epithelial labyrinth excepting for the
three openings that persist in the adult as the internal auditory
meatus, the aqueaductus cochleae and the aquaeductus vestibuli,
which openings are present in the very earliest stages.
Originally the cartilagenous capsule abuts directly against
the epithelial wall of the labyrinth. In embryos about 14 mm.
long, however, the cartilage-forming tissue in the immediate
neighborhood of the epithelium undergoes a dedifferentiation,
so that an area is established all around the membranous labyrinth, and conforming to it in shape, that is less like cartilage
and more like embryonic connective tissue. It is this that constitutes the foundation for the open—meshed periotic reticulum
which in embryos 30 mm. long everywhere bridges the space
existing between the membranous labyrinth and the surrounding cartilage. The membrana propria that supports the epithelial
part of the labyrinth and the perichondrium lining the cartilage
are both derived from this periotic reticulum. It i.s also a
modification of the meshes of this same reticulum that results
in the formation of the perioticular spaces in a manner that will
now be outlined.
Unmodified periotic reticulum is characterized by a rather
uniform narrow mesh. The essential change which it undergoes in the process of space formation consists in the disappearance of some of the trabeculae of the mesh followed by the coalescence of the corresponding adjacent spaces. The trabeculae
consist of the protoplasmic processes of the constituent cells
of the reticulum and their disappearance is probably to be
explained, not by a dissolution or liquefaction of these cellprocesses but by an alteration in their form. It apparently is
the result of an active motility of the cell protoplasm involving
the successive detachment and retraction of the trabeculae.
When a trabecula becomes detached it gradually retracts and
adapts itself to the formation of a larger space, reshaping itself
either as a smooth border or as a constituent part of another
trabecula. As spaces become larger they require longer trabeculae, and as trabeculae become longer they also tend to
become thicker.
The differentiation of the margin of the periotic spaces constitutes the final feature in their maturation. During the period
in which the enlargement of an individual space is actively
going on, the margins of the main cavity consist of smooth
delicate strands of nucleated protoplasm that resemble the
trabeculae between the large reticular spaces. These linear
margins are interrupted here and there by openings into adjacent
spaces. They tend, however, to form a continuous line that
definitely marks off the space from the adjacent reticulum.
As the space becomes more mature, the membrane-like border
becomes thicker until it reaches a state that will probably not
admit of any further opening-up for the coalescence of additional
spaces. Any further growth is thereafter limited to simple
distention of the wall of the space with the consequent adjustment of its constituent cells. In its final form the margin of the
space roughly resembles an endothelial membrane. Immediately
lining the space is a thin membrane with flattened nuclei which
is supported underneath by a thin coat of nucleated protoplasm
that has the form of fibrous connective tissue. The former,
judging only from its final appearance, could be designated as
endothelium, thus making a distinction between it and the
underlying tissue. In its histogenesis, however, it differs in
no way from the rest of the Wall and the difference that exists
later seems to be merely the result of its adaptation to the existing physical conditions. Its early behavior is entirely different
from that of vascular endothelium. Therefore if one uses the
term endothelium for its designation this must be done with a
considerable amount of reservation.
These phenomena can be particularly well studied in the
scalae While they are in the process of spreading and enlarging.
As we shall see, the scalae are more mature in their proximal
portions and are progressively less mature as one approaches
the apex of the cochlea. Thus any one specimen shows several
stages in the development. Typical views showing some of
the steps in this process are represented in figures 1 to 3. Figure
1 represents a section through the second turn of the cochlea in
ahuman fetus 130 mm. CR length (Carnegie Collection, No. 1018).
It shows the topography of the cochlear duct and the general
character of the perioticular spaces that are developing along
its inner margins. The upper one or scala Vestibuli is in a more
mature condition. The lower one or scala tympani is less mature and along its peripheral (right) margin, it is in the act of
spreading so as to underlie, as it eventually will do, the future
basilar membrane. Thescala tympani finally reaches the peripheral margin of the cochlear duct, and it does this by the coalescence of the enlarging reticular spaces which become incorporated with the main cavity of the scala.
This area is a particularly good one for studying the histogenesis of these spaces. It is shown under higher magnification
in figure 2, which is a detail of the same section. By comparing
this figure with figure 1, the exact location can be readily made out. That portion of the cochlear duct that is to form the organ
of Corti can be recognized by the characteristic form and grouping of its cells. A portion of the main cavity of the scala tympani is indicated and to the right of this are a few enlarged reticular spaces that are uniting with each other subjacent to the
organ of Corti and the basilar membrane. These will in the
end become part of the main space. They are here just in the process of coalescence, the histological features of which procedure are Well illustrated in this figure. The trabeculae are
stretched out in long strands and in many cases are detached
and project into the spaces as free ends. The detached trabeculae are seen in different degrees of retraction as their constituent protoplasm reshapes itself in adaptation to the new
boundaries. It is only at the margins of the larger spaces that
the cell-processes exhibit the characteristic flattened appearance, which is the first indication of the formation of the marginal
membrane. The residual undifferentiated reticulum that does
not enter into the direct formation of the larger spaces constitutes
the tissue from which is derived the adventitial coat of the completed scala.
Fig. 1 Section through the second turn of the cochlea in a human fetus 130 mm. CR length (Carnegie Collection, No. 1018). Enlarged 60 diameters. This section shows the topography of the cochlear duct and the general character of the periotic spaces that are developing along its inner margin. Details of this same section as seen under higher magnification are shown in figures 2 and 3.
Fig. 2 Detail of the section shown in figure 1, enlarged 278 diameters. This
figure shows the part of the cochlear duct that is to form the organ of Corti,
and the adjacent tissue that becomes incorporated in the basilar membrane.
Below this is the periotic reticulum whose spaces are in the process of enlarging.
By repeated coalescence these spaces finally unite with the large space that
constitutes the scala tympani. This figure shows the histological appearance of
the reticulum where the formation of tissue spaces is in active operation.
The appearance of the marginal membrane as seen in a more mature space is shown in figure 3, being a detail of the margin of the same scala Vestibuli that is shown in figure 1. Here we have a firm membrane that forms a complete barrier between the periotic reticulum and the lumen of the scala. After reaching this degree of development there is no evidence of any further coalescence of the surrounding reticular spaces with the main cavity. The membrane itself as seen in cross section consists of rather compact nucleated strands of protoplasm, which cannot as yet be separated into the so-called endothelial coat and the supporting fibrous coat. However, a comparison of the coagulated elements of the fluid seen in the reticular spaces with those seen in the scala would indicate a difference between the two and therefore it is probable that the membrane is already partially impervious.
Fig. 3 Detail of the section shown in figure 1, enlarged 400 diameters. It
shows the character of the margin of the scala vestibuli in a more mature condition. The scala vestibuli is inclosed by a membrane consisting of the cells
that had previously constituted the reticulum occupying this area and which
have been modified in form in adaptation to the formation of this large tissue.
space, closing it off from the surrounding tissue.
Development of the Periotic Cistern of the Vestibule
Aside from the scala vestibuli and the scala tympani, the largest of the periotic spaces is the large reservoir situated between the tympanic wall of the bony vestibuli with its articulated stapes, and the vestibular chambers of the membraneous labyrinth. This is the spatium perilymphaticum vestibuli (BNA) or the cysterna perilymphatica (Retzius). In order to eliminate the word lymphatic from the terminology it will be designated here as the Cisterna periotica vestibuli or less formally the periotic cistern. In this manner the descriptive term introduced by Retzius is retained.
Before there is any trace of the scalae the initial steps in the
formation of the cistern can be seen. This is well illustrated in
an embryo 35 mm. long (Carnegie Collection, No. 199). This
particular embryo is cut into a sagittal series and the sections
on slides 53 and 54 show the periotic cistern in its most rudimentary form. It consists of an area of reticulum bounded by
the utricle, saccule, ductus reuniens, the proximal endof the
cochlear duct and the ampulla of the posterior canal. The reticulum here is of the type seen along the semicircular canals in
considerably older embryos. Whereas the reticulum elsewhere
in this 35 mm. embryo presents a uniformly narrow mesh that
is interrupted only by the numerous capillaries branching
through it, this particular field gives the appearance of spaces
which are more open and which are irregular both in shape and
in size. From the very first the increase in the size of the mesh
seems to be attained by the detachment and retraction of its
constituent protoplasmic bridges, thereby allowing adjacent
spaces to unite in the formation of composite larger spaces. Thus
in the above section a few irregular protoplasmic free-ends are
seen still projecting into the newly enlarged spaces. The area
of this rudimentary periotic cistern is as yet very small and
merges indefinitely into the adjoining reticulum. It is not
until we come to fetuses about 40 mm. long that it develops
spaces of any considerable size, and it is not until we come to
fetuses about 50 mm. long that we find a single large space with
walls that are definitely outlined so that it can be satisfactorily
modelled.
In a fetus 43 mm. long (Carnegie Collection, No. 886), the
spaces forming the rudimentary cistern stand out much more
definitely than is the case in the 35 mm. embryo that has just
been referred to. There is now just opposite the stapes one space
which is much larger than the adjoining spaces. On part of its
margin the protoplasmic bridges are stretched along so as to
form a smoothly curved continuous boundary. This boundary
is defective in some portions and at such places the space merges with the adjoining secondary spaces. Within the space are
some faintly refractive branching threads of coagulated plasma.
The scala Vestibuli is not yet laid down and the scala tympani is
only represented by a moderate widening of the meshes of the
reticulum in the neighborhood of the fenestra cochleae (rotundum), along the basal border of the first turn of the cochlear
duct.
In fetuses 50 mm. long the outlines of the cistern become very
distinct due to the marked increase in the size of its main cavity
and to the more definite membrane at its junction with the
rest of the reticulum. Its form and relations are shown in
figures 4 and 5. They represent a median and a lateral view of
a waxplate reconstruction of this region in a human fetus 50 mm.
long (Carnegie Collection, No. 84). Only the main cavity is
shown in the model. At certain places around its borders the
meshes of the reticulum are uniting into larger spaces and these
in turn are taken up by the main cavity as it advances into the
new territory. These smaller incomplete spaces were omitted
in constructing the plates of the model.
It will be seen then from figure 4 and 5 that the periotic
cistern in 50 mm. embryos consists of a flattened rounded bursalike cavity that intervenes between the stapes and the lateral
surface of the saccule and adjoining utricle. It extends forward
to the ampulla of the lateral canal and upward to the beginning
of the crus commune. Posteriorly it crowds backward against
the ductus reuniens filling in the space between the utricle, saccule
and the proximal end of the cochlear duct. Both on its median
and lateral surfaces there is no further opportunity for expansion
except as the vestibule itself enlarges. The delicate membranelike wall of the cistern hugs closely against the parts of the membranous labyrinth on the one side and the tympanic wall of the cartilagenous vestibule on the other, being separated from them
only by a thin layer of the original reticulum. Along the dorsal
margin of the cistern, however, there is room for expansion and
the reticulum in this region shows enlarging spaces in the process of uniting with the main cavity. On its ventral margin
near the cochlea and extending along the apical surface of the latter there is a definite row of reticular spaces actively coalescing
and constituting the beginning of the scala vestibuli. The
scala tympani is already well started at this time, but its development is quite independent of the cistern. Within the cistern can be seen scattered clumps of faintly refractive granular
threads of what seems to be a coagulated constituent of the
plasma.
Fig. 4 The figures 4 to 9 represent a series of median and lateral views of wax-plate reconstructions of the membranous labyrinth and the surrounding perioticular tissue spaces, illustrating under the same scale of enlargement three typical stages in the development of these spaces. This figure shows a lateral view of a model reconstructed from a human fetus 50 mm. CR length (Carnegie Collection, No. 84). The scala vestibuli is in the first stage of its development and consists of a row of large reticular spaces which extend from the ventral margin of the cistern downward along the apical surface of the cochlear duct. The scala tympani is more advanced and shows more complete coalescence of its constituent spaces. Enlarged 9 diameters.
Fig. 5 Median View of the same model shown in figure 4. This view shows the topography of the scala tympani. Its large proximal end lies opposite the
fenestra cochleac and corresponds to the focus at which its development originates. Distally it tapers off rapidly, where the spaces are smaller and their
coalescence less complete. Enlarged 9 diameters.
The subsequent growth of the cistern is shown in figures 6
to 9. Figures 6 and 7 show respectively a median and lateral
View of a Wax-plate reconstruction of the membranous labyrinth
and its perioticular spaces in a human fetus 85 mm. long (Carnegie Collection, No. 140030). The growth of the cistern here
has kept pace with the increase in size of the labyrinth and
maintains the same general relations as regards the stapes and
the parts of the membranous labyrinth. The View of the cis-
tern in figure 6 is an oblique one Which would tend to mislead
one as to its width. In reality it is relatively a little wider.
Fig. 6 Lateral View of reconstruction of the left membranous labyrinth
and the periotic spaces in a human fetus 85 mm. CR length (Carnegie Collection,
No. 1400-30) enlarged 9 diameters. Although the greater part of the cistern
abuts against the stapes it will be noted that it also is beginning to spread over
the dorsal surface of the utricle and along the inner border of the lateral canal.
The scala vestibuli communicates freely with the cistern and extends downward
along the apical surface of the cochlear duct throughout nearly two turns, show-
ing the characteristic sacciilatcd appearance near its tip where the coalescence
of the spaces is less complete.
Fig. 7 Median view of same model shown in figure 6. The oval indentation
in the proximal end of the scala tympani corresponds to the fenestra cochleae.
This space extends along the cochlear duct about the same distance as the scala
vestibuli; the two however do not communicate with each other as yet. The
peripheral border of the scala tympani is characterized by sacculations cor-
responding to spaces that are coalescing with the main space. This indicates
the direction of the growth of the" scala at this time.
It has also extended upward on the dorsal surface of the utricle
and is beginning to creep along the inner side of the posterior
end of the lateral canal. Ventrally it communicates freely
with the scala vestibuli which now extends well down along the
cochlear duct.
The oldest stage studied is shown in figures 8 and 9. These
show two views of a wax-plate reconstruction of these structures
in a human fetus 130 mm. long (Carnegie Collection, No. 1018).
At this time the periotic cistern has spread over the vestibular
part of the membranous labyrinth, covering it nearly everywhere
excepting at the macular portions where the nerves terminate.
In figure 9 it can be seen that the mesial surface of the saccule
is not covered; this lies closely against the wall of the cartilagenous vestibule. The uppermost division of the cistern situated
between the crus commune and the ampulla of the posterior
canal does not yet open into the general cavity. It has formed
separately and owing to the position in which it lies its coalescence with the other parts of the cistern is retarded. Otherwise
free communication exists between all divisions of the cistern.
Development of the Scala Tympani and Scala Vestibuli
The scala vestibuli may be regarded as an extension downward of the cistern into the region of the cochlea and as such its growth starts from a focus opposite the fenestra vestibuli (ovale). The scala tympani in a similar way makes its first appearance opposite the fenestra cochleae. From these two foci the scalae extend gradually downward along the cochlear duct as two separate spaces which do not communicate with each other until they reach the tip of the duct, where there is finally developed a free opening between them known as the helicotrema.
In their formation they go through a series of histogenetic changes in essentially the same manner that has been followed in the case of the formation of the cistern. This as we shall see consists of the enlargement of the spaces of the periotic reticulum that originally occupies this region, the enlargement being a result of the disappearance of the protoplasmic bridges of the:
Cite this page: Hill, M.A. (2024, April 19) Embryology Paper - The Development of the Scala Tympani, Scala Vestibuli and Perioticular Cistern in the Human Embryo. Retrieved from https://embryology.med.unsw.edu.au/embryology/index.php/Paper_-_The_Development_of_the_Scala_Tympani,_Scala_Vestibuli_and_Perioticular_Cistern_in_the_Human_Embryo
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