The scala media contains the essential organ of hearing or organ of Corti (fig. 4, oc), which lies upon the inner part of the basilar membrane; it consists of a tunnel bounded on each side of the inner and outer rods of Corti; on each side of these are the inner and outer hair cells, between the latter of which are found the supporting cells of Deiters. Most externally are the large cells of Hensen. A delicate membrane called the lamina reticularis covers the top of all these, and is pierced by the hairs of the hair cells, while above this is the loose membrana tectoria attached to the periosteum of the lamina spiralis, near its tip, internally, and possibly to some of Deiter’s cells externally. The cochlear branch of the auditory nerve enters the lamina spiralis, where a spiral ganglion (fig. 4, sg) is developed on it; after this it is distributed to the inner and outer hair cells.

(From R. Howden—Cunningham’s Text-Book of Anatomy.)
Fig. 5.—Transverse Section of Corti’s Organ from the Central Coil of Cochlea (Retzius).

For further details see Text-Book of Anatomy, edited by D.J. Cunningham (Edinburgh, 1906); Quain’s Elements of Anatomy (London, 1893); Gray’s Anatomy (London, 1905); A Treatise on Anatomy, edited by H. Morris (London, 1902); A Text-Book of Human Anatomy, by A. Macalister (London, 1889).

Embryology.—The pinna is formed from six tubercles which appear round the dorsal end of the hyomandibular cleft or, more strictly speaking, pouch. Those for the tragus and anterior part of the helix belong to the first or mandibular arch, while those for the antitragus, antihelix and lobule come from the second or hyoid arch. The tubercle for the helix is dorsal to the end of the cleft where the two arches join. The external auditory meatus, tympanum and Eustachian tube are remains of the hyomandibular cleft, the membrana tympani being a remnant of the cleft membrane and therefore lined by ectoderm outside and entoderm inside. The origin of the ossicles is very doubtful. H. Gadow’s view, which is one of the latest, is that all three are derived from the hyomandibular plate which connects the dorsal ends of the hyoid and mandibular bars (Anatomischer Anzeiger, Bd. xix., 1901, p. 396). Other papers which should be consulted are those of E. Gaupp, Anatom. Hefte, Ergebnisse, Bd. 8, 1898, p. 991, and J.A. Hammar, Archiv f. mikr. Anat. lix., 1902. These papers will give a clue to the immense literature of the subject. The internal ear first appears as a pit from the cephalic ectoderm, the mouth of which in Man and other mammals closes up, so that a pear-shaped cavity is left. The stalk of the pear which is nearest the point of invagination is called the recessus labyrinthi, and this, after losing its connexion with the surface of the embryo, grows backward toward the posterior cranial fossa and becomes the ductus endolymphaticus. The lower part of the vesicle grows forward and becomes the cochlea, while from the upper part three hollow circular plates grow out, the central parts of which disappear, leaving the margin as the semicircular canals. Subsequently constrictions appear in the vesicle marking off the saccule and utricle. From the surrounding mesoderm the petrous bone is formed by a process of chondrification and ossification.

See W. His, Junr., Archiv f. Anat. und Phys., 1889, supplement, p. 1; also Streeter, Am. Journ. of Anat. vi., 1907.

Comparative Anatomy.—The ectodermal inpushing of the internal ear has probably a common origin with the organs of the lateral line of fish. In the lower forms the ductus endolymphaticus retains its communication with the exterior on the dorsum of the head, and in some Elasmobranchs the opening is wide enough to allow the passage of particles of sand into the saccule. It is probable that this duct is the same which, taking a different direction and losing its communication with the skin, abuts on the posterior cranial fossa of higher forms (see Rudolf Krause, “Die Entwickelung des Aq. vestibuli seu d. Endelymphaticus,” Anat. Anzeiger, Bd. xix., 1901, p. 49). In certain Teleostean fishes the swim bladder forms a secondary communication with the internal ear by means of special ossicles (see G. Ridewood, Journ. Anat. & Phys. vol. xxvi.). Among the Cyclostomata the external semicircular canals are wanting; Petromyzon has the superior and posterior only, while in Myxine these two appear to be fused so that only one is seen. In higher types the three canals are constant. Concretions of carbonate of lime are present in the internal ears of almost all vertebrates; when these are very small they are called otoconia, but when, as in most of the teleostean fishes, they form huge concretions, they are spoken of as otoliths. One shark, Squatina, has sand instead of otoconia (C. Stewart, Journ. Linn. Society, xxix. 409). The utricle, saccule, semicircular canals, ductus endolymphaticus and a short lagena are the only parts of the ear present in fish.

The Amphibia have an important sensory area at the base of the lagena known as the macula acustica basilaris, which is probably the first rudiment of a true cochlea. The ductus endolymphaticus has lost its communication with the skin, but it is frequently prolonged into the skull and along the spinal canal, from which it protrudes, through the intervertebral foramina, bulging into the coelom. This is the case in the common frog (A. Coggi, Anat. Anz. 5. Jahrg., 1890, p. 177). In this class the tympanum and Eustachian tube are first developed; the membrana tympani lies flush with the skin of the side of the head, and the sound-waves are transmitted from it to the internal ear by a single bony rod—the columella.

In the Reptilia the internal ear passes through a great range of development. In the Chelonia and Ophidia the cochlea is as rudimentary as in the Amphibia, but in the higher forms (Crocodilia) there is a lengthened and slightly twisted cochlea, at the end of which the lagena forms a minute terminal appendage. At the same time indications of the scalae tympani and vestibuli appear. As in the Amphibia the ductus endolymphaticus sometimes extends into the cranial cavity and on into other parts of the body. Snakes have no tympanic membrane. In the birds the cochlea resembles that of the crocodiles, but the posterior semicircular canal is above the superior where they join one another. In certain lizards and birds (owls) a small fold of skin represents the first appearance of an external ear. In the monotremes the internal ear is reptilian in its arrangement, but above them the mammals always have a spirally twisted cochlea, the number of turns varying from one and a half in the Cetacea to nearly five in the rodent Coelogenys. The lagena is reduced to a mere vestige. The organ of Corti is peculiar to mammals, and the single columella of the middle ear is replaced by the three ossicles already described in Man (see Alban Doran, “Morphology of the Mammalian Ossicula auditus,” Proc. Linn. Soc., 1876-1877, xiii. 185; also Trans. Linn. Soc. 2nd Ser. Zool. i. 371). In some mammals, especially Carnivora, the middle ear is enlarged to form the tympanic bulla, but the mastoid cells are peculiar to Man.

For further details see G. Retzius, Das Gehörorgan der Wirbelthiere (Stockholm, 1881-1884); Catalogue of the Museum of the R. College of Surgeons—Physiological Series, vol. iii. (London, 1906); R. Wiedersheim’s Vergleichende Anatomie der Wirbeltiere (Jena, 1902).

(F. G. P.)