Paper - 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
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. (2021, June 23) 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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