The Works of Francis Balfour 3-17

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Foster M. and Sedgwick A. The Works of Francis Balfour Vol. III. A Treatise on Comparative Embryology 2 (1885) MacMillan and Co., London.

Cephalochorda | Urochorda | Elasmobranchii | Teleostei | Cyclostomata | Ganoidei | Amphibia | Aves | Reptilia | Mammalia | Comparison of the Formation of Germinal Layers and Early Stages in Vertebrate Development | Ancestral form of the Chordata | General Conclusions | Epidermis and Derivatives | The Nervous System | Organs of Vision | Auditory, Olfactory, and Lateral Line Sense Organs | Notochord, Vertebral Column, Ribs, and Sternum | The Skull | Pectoral and Pelvic Girdles and Limb Skeleton | Body Cavity, Vascular System and Glands | The Muscular System | Excretory Organs | Generative Organs and Genital Ducts | The Alimentary Canal and Appendages in Chordata
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This historic 1885 book edited by Foster and Sedgwick is the third of Francis Balfour's collected works published in four editions. Francis (Frank) Maitland Balfour, known as F. M. Balfour, (November 10, 1851 - July 19, 1882) was a British biologist who co-authored embryology textbooks.



Foster M. and Sedgwick A. The Works of Francis Balfour Vol. I. Separate Memoirs (1885) MacMillan and Co., London.

Foster M. and Sedgwick A. The Works of Francis Balfour Vol. II. A Treatise on Comparative Embryology 1. (1885) MacMillan and Co., London.

Foster M. and Sedgwick A. The Works of Francis Balfour Vol. III. A Treatise on Comparative Embryology 2 (1885) MacMillan and Co., London.

Foster M. and Sedgwick A. The Works of Francis Balfour Vol. IV. Plates (1885) MacMillan and Co., London.
Modern Notes:

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Pages where the terms "Historic Textbook" and "Historic Embryology" appear on this site, and sections within pages where this disclaimer appears, indicate that the content and scientific understanding are specific to the time of publication. This means that while some scientific descriptions are still accurate, the terminology and interpretation of the developmental mechanisms reflect the understanding at the time of original publication and those of the preceding periods, these terms and interpretations may not reflect our current scientific understanding.     (More? Embryology History | Historic Embryology Papers)


Draft Version - Notice removed when completed.

Vol. III. A Treatise on Comparative Embryology 2 (1885)

Chapter XVII. Auditory Organ, Olfactory Organ and Sense Organs of the Lateral Line

Auditory Organs

A great variety of organs, very widely distributed amongst aquatic forms, and also found, though less universally, in land forms, are usually classed together as auditory organs.

In the case of all aquatic forms, or of forms which have directly inherited their auditory organs from aquatic forms, these organs are built upon a common type ; although in the majority of instances the auditory organs of the several groups have no genetic relations. All the organs have their origin in specialized portions of the epidermis. Some of the cells of a special region become provided at their free extremities with peculiar hairs, known as auditory hairs; while in other cells concretions, known as otoliths, are formed, which appear often to be sufficiently free to be acted upon by vibrations of the surrounding medium, and to be so placed as to be able in their turn to transmit their vibrations to the cells with auditory hairs 1 . The auditory regions of the epidermis are usually shut off from the surface in special sacks.

The actual function of these organs is no doubt correctly described, in the majority of instances, as being auditory; but it appears to me very possible that in some cases their function may be to enable the animals provided with them to detect the presence of other animals in their neighbourhood, through the unclulatory movements in the water, caused by the swimming of the latter.

1 The function of the otoliths is not always clear. There is evidence to shew that they sometimes act as dampers.


Auditory organs with the above characters, sometimes freely open to the external medium, but more often closed, are found in various Ccelenterata, Vermes and Crustacea, and universally or all but universally in the Mollusca and Vertebrata.

In many terrestrial Insects a different type of auditory organ has been met with, consisting of a portion of the integument modified to form a tympanum or drum, and supported at its edge by a chitinous ring. The vibrations set up in the membranous tympanum stimulate terminal nerve organs at the ends of chitinous processes, placed in a cavity bounded externally by the tympanic membrane.

The tympanum of Amphibia and Amniota is an accessory organ added, in terrestrial Vertebrata, to an organ of hearing primitively adapted to an aquatic mode of life ; and it is interesting to notice the presence of a more or less similar membrane in the two great groups of terrestrial forms, i.e. terrestrial Vertebrata and Insecta.

Nothing is known with reference to the mode of development or evolution of the tympanic type of auditory organ found in Insects, and, except in the case of Vertebrates, but little is known with reference to the development of what may be called the vesicular type of auditory organ found in aquatic forms. Some very interesting facts with reference to the evolution of such organs have however been brought to light by the brothers Hertwig in their investigations on the Ccelenterata; and I propose to commence my account of the development of the auditory organs in the animal kingdom by a short statement of the results of their researches.

Ccelenterata. Three distinct types of auditory organ have been recognised in the Medusae ; two of them resulting from the differentiation of a tentacle-like organ, and one from ectoderm cells on the under surface of the velum. We may commence with the latter as the simplest. It is found in the Medusae known as the Vesiculata. The least differentiated form of this organ, so far discovered, is present in Mitrotrocha, Tiaropsis and other genera. It has the form of an open pit ; and a series of such organs are situated along the attached edge of the velum with their apertures directed downwards. The majority of the cells lining the outer, i.e. peripheral side of the



FIG. 297. AUDITORY VESICLE OF PHIALIDIUM AFTER TREATMENT WITH DILUTE OSMIC ACID. (From Lankester; after O. and R. Hertwig.)

d l . epithelium of the upper surface of the velum; d 2 . epithelium of the under surface of the velum ; r. circular canal at the edge of the velum ; nr l . upper nervering ; h. auditory cells ; hh. auditory hairs ; np. nervous cushion formed of a prolongation of the lower nerve-ring. Close to the nerve-ring is seen a cell, shewn as black, containing an otolith.

pit, contain an otolith, while a row of the cells on the inner, i.e. central side, are modified as auditory cells. The auditory cells are somewhat strap-shaped, their inner ends being continuous with the fibres of the lower nerve-ring, and their free ends being provided with bent auditory hairs, which lie in contact with the convex surfaces of the cells containing the otoliths.

By the conversion of such open pits into closed sacks a more complicated type of auditory organ, which is present in many of the Vesiculata, viz. ^Equorea, Octorchis, Phialidium, &c., is produced. A closed vesicle of this type is shewn in fig. 297. Such organs form projections on the upper surface of the velum. They are covered by a layer of the epithelium (d 1 } of the upper surface of the velum, but the lining of the vesicle (d*} is derived from what was originally part of the epithelium of the lower surface of the velum, homologous with that lining the open pits in the type already described. The general arrangement of the cells lining such vesicles is the same as that of the cells lining the open pits.

A second type of auditory organ, found in the Trachymeclusa,", appears in its simplest condition as a modified tentacle.


It is formed of a basal portion, covered by auditory cells with long stiff auditory hairs, supporting at its apex a club-shaped body, attached to it by a delicate stalk. An endodermal axis is continued through the whole structure, and in one or more of the endoderm cells of the club-shaped body otoliths are always present. The tails of the auditory cells are directly continued into the upper nerve-ring.

In more complicated forms of this organ the tentacle becomes enclosed in a kind of cup, by a wall-like upgrowth of the surrounding parts (fig. 298) ; and in some forms, e.g. Geryonia, by the closure of the cup, the whole structure takes the form of a completely closed vesicle, in the cavity of which the original tentacle forms an otolith-bearing projection.



FIG. 298. AUDITORY ORGAN OF RHOPALONEMA. (From Lankester; after O. and R. Hertwig.)

The organ consists of a modified tentacle (hk) with auditory cells and concretions, partially enclosed in a cup.


The auditory organs found in the Acraspedote Medusae approach in many respects to the type of organ found in the Trachymedusse. They consist of tentacular organs placed in grooves on the under surface of the disc. They have a swollen extremity, and are provided with an endodermal axis for half the length of which there is a diverticulum of the gastrovascular canal system. The terminal portion of the endoderm is solid, and contains calcareous concretions. The ectodermal cells at the base of these organs have the form of auditory cells.

Mollusca. Auditory vesicles are found in almost all Mollusca on the ventral side of the body in close juxtaposition to the pedal ganglia. Except possibly in some Cephalopods, these vesicles are closed. They are provided with free otoliths, supported by the cilia of the walls of the sack, but in addition some of the cells of the sack are provided with stiff auditory hairs.

In many forms these sacks have been observed to originate by an invagination of the epiblast of the foot (Pahtdina, Nassa, Heteropoda, Limax, Clio, Cephalopoda and Lamellibranchiata). In other instances (some Pteropods, Lymnaeus, &c.) they appear, by a secondary modification in the development, to originate by a differentiation of a solid mass of epiblast.

According to Fol the otocysts in Gasteropods are formed by cells of the wall of the auditory sacks ; and the same appears to hold good for Cephalopoda (Grenacher) 1 shewing that free otoliths have in these instances originated from otoliths originally placed in cells.

Crustacea. In the decapodous Crustacea organs, which have been experimentally proved to be true organs of hearing, are usually present on the basal joint of the anterior antennae. They may have (Hensen, No. 384) the form either of closed or of open sacks, lined by an invagination of the epidermis. They are provided with chitinous auditory hairs and free otoliths. In the case of the open sacks the otoliths appear to be simply stones transported into the interior of the sacks, but in the closed sacks the otoliths, though free, are no doubt developed within the sacks.

The Schizopods, which, as mentioned in the last chapter, are remarkable as containing a genus (Euphausia) with abnormally situated eyes, distinguish themselves again with reference to their auditory organs, in that another genus (Mysis) is characterized by the presence of a pair of auditory sacks in the inner plates of the tail. These sacks have curved auditory hairs supporting an otolith at their extremity.

The development of the auditory organs in the Crustacea has not been investigated.

The Vertebrata. The Cephalochorda are without organs of hearing, and the auditory organ of the Urochorda is constructed on a special type of its own. The primitive auditory organs of the true Vertebrata have the same fundamental characters as those of the majority of aquatic invertebrate forms. They consist of a vesicle, formed by the invagination of a patch of epiblast, and usually shut off from the exterior, but occasionally (Elasmo branchii) remaining open. The walls of this vesicle are always much complicated and otoliths of various forms are present in its cavity. To this vesicle accessory structures, derived from the walls of the hyomandibular cleft, are added in the majority of terrestrial Vertebrata.


1 For the somewhat complicated details as to the development of the auditory sacks of Cephalopods I must refer the reader to Vol. II., pp. 278, 279, and to Grenacher (Vol. i., No. 280).


The development of the true auditory vesicle will be considered separately from that of the accessory structures derived from the hyomandibular cleft.

In all Vertebrata the development of the auditory vesicle commences with the formation of a thickened patch of epiblast, at the side of the hind-brain, on the level of the second visceral cleft.

t.v.v

This patch soon becomes invaginated in the form of a pit (fig. 299, aup), to the inner side of which the ganglion of the auditory nerve (ami), which as shewn in a previous chapter is primitively a branch of the seventh nerve, closely applies itself.

In those Vertebrata (viz. Teleostei, Lepidosteus and Amphibia) in which the epiblast is early divided into a nervous and epidermic stratum, the auditory pit arises as an invagination of the nervous stratum only, and the mouth of the auditory pit is always closed -(fig. 300) by the epidermic stratum of the skin. Since the opening of the pit is retained through life in Elasmobranchii the closed form of pit in the above forms is clearly secondary.

In Teleostei the auditory pit arises as a solid invagination of the epiblast.


FIG. 299. SECTION THROUGH THE HEAD OF AN ELASMOBRANCH EMBRYO, AT THE LEVEL OF THE AUDITORY INVOLUTION.

aup. auditory pit; aun. ganglion of auditory nerve ; iv.v. roof of fourth ventricle; a.c.v. anterior cardinal vein; aa. aorta; I.aa. aortic trunk of mandibular arch ; pp. head cavity of mandibular arch ; Ivc. alimentary pouch which will form the first visceral cleft; 77?. rudiment of thyroid body.


The mouth of the auditory vesicle gradually narrows, and in most forms soon becomes closed, though in Elasmobranchii it remains permanently open. In any case the vesicle is gradually removed from the surface, remaining connected with it by an elongated duct, either opening on the dorsal aspect of the head (Elasmobranchii), or ending blindly close beneath the skin.

In all Vertebrata the auditory vesicle undergoes further changes of a complicated kind. In the Cyclostomata these changes are less complicated than in other forms, though whether this is due to degeneration, or to the retention of a primitive of the epiblast, which soon becomes vesicular, and for a considerable period retains a simple character. As pointed out by Max Schultze, a number of otoliths appears in the vesicle during larval life, and, although such otoliths are stated by J. Miiller to be absent both in the full-grown Ammoccete and in the adult, they have since been found by Ketel (No. 387). The formation of the two semicircular canals has not been investigated.



FIG. 300. SECTION THROUGH THE HEAD OK A LEPIUOSTEUS EMBRYO ON THE SIXTH DAY AFTER IMPREGNATION. au.v. auditory vesicle ; au.n. auditory nerve ; ch. notochord ; hy. hypoblast.

state of the auditory organ, is not known. In the Lamprey the auditory vesicle is formed in the usual way by an invagination


FIG. 301. SECTION THROUGH THE HIND-BRAIN OK A CHICK AT THE END OF THE THIRD DAY OF INCUBATION.

IV. fourth ventricle. The section shews the very thin roof and thicker sides of the ventricle. Ch. notochord ; C V. anterior cardinal vein; CC. involuted auditory vesicle (CC points to the end which will form the cochlear canal) ; RL. recessus labyrinthi (remains of passage connecting the vesicle with the exterior) ; hy. hypoblast lining the alimentary canal; AO., AO.A. aorta, and aortic arch.


In all the higher Vertebrates the changes of the auditory sacks are more complicated. The ventral end of the sack is produced into a short process (fig. 301, CC}\ while at the dorsal end there is the canal-like prolongation of the lumen of the sack (RL}, derived from the duct which primitively opened to the exterior, and which in most cases persists as a blind diverticulum of the auditory sack, known as the recessus labyrinthi or aqueductus vestibuli. The parts thus indicated give rise to the whole of the membranous labyrinth of the ear. The main body of the vesicle becomes the utriculus and semicircular canals, while the ventral process forms the sacculus hemisphericus and cochlear canal.

The growth of these parts has been most fully studied in Mammalia, where they reach their greatest complexity, and it will be convenient to describe their development in this group, pointing out how they present, during some of the stages in their growth, a form permanently retained in lower types.

The auditory vesicle in Mammalia is at first nearly spherical, and is imbedded in the mesoblast at the side of the hind-brain. It soon becomes triangular in section, with the apex of the triangle pointing inwards and downwards. This apex gradually elongates to form the rudiment of the cochlear canal and sacculus hemisphericus (fig. 302, CC). At the same time the recessus labyrinthi (R.L) becomes distinctly marked, and the outer wall of the main body of the vesicle grows out into two protuberances, which form the rudiments of the vertical semicircular canals ( V.B}. In the lower forms (fig. 305) the cochlear process of the vestibule hardly reaches a higher stage of development than that found at this stage in Mammalia.

The parts of the auditory labyrinth thus established soon increase in distinctness (fig. 303) ; the cochlear canal (CC} becomes longer and curved ; its inner and concave surface being lined by a thick layer of columnar epiblast. The recessus labyrinthi also increases in length, and just below the point where the bulgings to form the vertical semicircular canals are situated, there is formed a fresh protuberance for the horizontal semi



FIG. 302. TRANSVERSE SECTION OF THE HEAD OF A FCETAL SHEEP (16 MM. IN LENGTH) IN THE REGION OF THE HIND-BRAIN. (After Bottcher.)

HB. the hind -brain.

The section is somewhat oblique, hence while on the right side the connections of the recessus vestibuli R.L., and of the commencing vertical semicircular canal V.B., and of the ductus cochlearis CC., with the cavity of the primary otic vesicle are seen : on the left side, only the extreme end of the ductus cochlearis CC, and of the semicircular canal V.B. are shewn.

Lying close to the inner side of the otic vesicle is seen the cochlear ganglion GC ; on the left side the auditory nerve G and its connection N with the hind-brain are also shewn.

Below the otic vesicle on either side lies the jugular vein.

circular canal. At the same time the central parts of the walls of the flat bulgings of the vertical canals grow together, obliterating this part of the lumen, but leaving a canal round the periphery ; and, on the absorption of their central parts, each of the original simple bulgings of the wall of the vesicle becomes converted into a true semicircular canal, opening at its two extremities into the auditory vesicle. The vertical canals are first established and then the horizontal canal.


Shortly after the formation of the rudiment of the horizontal semicircular canal a slight protuberance becomes apparent on the inner commencement of the cochlear canal. A constriction arises on each side of the protuberance, converting it into a prominent hemispherical projection, the sacculus hemisphericus (fig. 304, S.R\



FIG. 303. SECTION OF THE HEAD OF A FCETAL SHEEP 20 MM. IN LENGTH. (After Bottcher.)

R. V. recessus labyrinthi ; V.B. vertical semicircular canal ; H.B. horizontal semicircular canal; C.C. cochlear canal ; G. cochlear ganglion.


The constrictions are so deep that the sacculus is only connected with the cochlear canal on the one hand, and with the general cavity of the auditory vesicle on the other, by, in each case, a narrow though short canal.

The former of these canals (fig. 304, b) is known as the canalis reuniens. At this stage we may call the remaining cavity of the original otic vesicle, into which all the above parts open, the utriculus.

Soon after the formation of the sacculus hemisphericus, the cochlear canal and the semicircular canals become invested with cartilage. The recessus labyrinthi remains however still enclosed in undifferentiated mesoblast

Between the cartilage and the parts which it surrounds there remains a certain amount of indifferent connective tissue, which is more abundant around the cochlear canal than around the semicircular canals.

As soon as they have acquired a distinct connective-tissue coat, the semicircular canals begin to be dilated at one of their terminations to form the ampullae. At about the same time a constriction appears opposite the mouth of the recessus labyrinthi, which causes its opening to be divided into two branches one towards the utriculus and the other towards the sacculus hemisphericus ; and the relations of the parts become so altered that communication between the sacculus and utriculus can only take place through the mouth of the recessus labyrinthi (fig. 305).

When the cochlear canal has come to consist of two and a half coils, the thickened epithelium which lines the lower surface of the canal forms a double ridge from which the organ of Corti is subsequently developed. Above the ridge there appears a delicate cuticular membrane, the membrane of Corti or membrana tectoria.

The epithelial walls of the utricle, the recessus labyrinthi, the semicircular canals, and the cochlear canal constitute together the highly complicated product of the original auditory vesicle. The whole structure forms a closed cavity, the various parts of which are in free communication. In the adult the fluid present in this cavity is known as the endolymph.

In the mesoblast lying between these parts and the cartilage, which at this period envelopes them, lymphatic spaces become established, which are partially developed in the Sauropsida, but become in Mammals very important structures.

They consist in Mammals partly of a space surrounding the utricle and semicircular canals, and partly of two very definite channels, which largely embrace between them the cochlear canal. The latter channels form the scala vestibuli on the upper side of the cochlear canal and the scala tympani on the lower. The scala vestibuli is in free communication with the lymphatic cavity surrounding the vestibule, and opens at the apex of the cochlea into the scala tympani. The latter ends blindly at the fenestra rotunda.

The fluid contained in the two scalae, and in the remaining lymphatic cavities of the auditory labyrinth, is known as perilymph.

The cavities just spoken of are formed by an absorption of parts of the embryonic mucous tissue between the perichondrium and the walls of the membranous labyrinth.



FIG. 304. SECTION THROUGH THE INTERNAL EAR OF AN EMBRYONIC SHEEP 28 MM. IN LENGTH. (After Bottcher.)

D.M. dura mater; R. V. recessus labyrinthi ; H.V.B. posterior vertical semicircular canal ; U. utriculus ; H.B. horizontal semicircular canal; b. canalis reuniens ; a. constriction by means of which the sacculus hemisphericus S.R. is formed ; f. narrowed opening between sacculus hemisphericus and utriculus ; C. C. cochlea ; C.C. lumen of cochlea; K.K. cartilaginous capsule of cochlea; K.B. basilar plate; Ch. notochord.


The scala vestibuli is formed before the scala tympani, and both scalae begin to be developed at the basal end of the cochlea : the cavity of each is continually being carried forwards towards the apex of the cochlear canal by a progressive absorption of the mesoblast. At first both scalae are somewhat narrow, but they soon increase in size and distinctness.

The cochlear canal, which is often known as the scala media of the cochlea, becomes compressed on the formation of the scalae so as to be triangular in section, with the base of the triangle outwards. This base is only separated from the surrounding cartilage by a narrow strip of firm mesoblast, which becomes the stria vascularis, etc. At the angle opposite the base the canal is joined to the cartilage by a narrow isthmus of firm material, which contains nerves and vessels. This isthmus subsequently forms the lamina spiralis, separating the scala vestibuli from the scala tympani.

The scala vestibuli lies on the upper border of the cochlear canal, and is separated from it by a very thin layer of mesoblast, bordered on the cochlear aspect by flat epiblast cells. This membrane is called the membrane ofReissner. The scala tympani is separated from the cochlear canal by a thicker sheet of mesoblast, called the basilar membrane, which supports the organ of Corti and the epithelium adjoining it. The upper extremity of the cochlear canal ends in a blind extremity called the cupola, to which the two scalae do not for some time extend. This condition is permanent in Birds, where the cupola is represented by a structure known as the lagena (fig. 305, II. L}. Subsequently the two scalae join at the extremity of the cochlear canal ; the point of the cupola still however remains in contact with the bone, which has now replaced the cartilage, but at a still later period the scala vestibuli, growing further round, separates the cupola from the adjoining osseous tissue.

The ossification around the internal ear is at first confined to the cartilage, but afterwards extends into the thick periosteum between the cartilage and the internal ear, and thus eventually makes its way into the lamina spiralis, etc.

The organ of Corti. In Mammalia there is formed from the epithelium of the cochlear canal a very remarkable organ known as the organ of Corti, the development of which is of sufficient importance to merit a brief description. A short account of this organ in the adult state may facilitate the understanding of its development.

The cochlear canal is bounded by three walls, the outer one being the osseous wall of the cochlea. The membrane of Reissner bounds it towards



FIG. 305. DIAGRAMS OF THE MEMBRANOUS LABYRINTH. (From Gegenbaur.)

I. Fish. II. Bird. III. Mammal.

U. utriculus ; S. sacculus ; US. utriculus and sacculus ; Cr. canalis reuniens ; R. recessus labyrinthi ; UC. commencement of cochlea ; C. cochlear canal ; L. lagena ; PC. cupola at apex of cochlear canal; V. csecal sack of the vestibulum of the cochlear canal.

the scala vestibuli, and the basilar membrane towards the scala tympani. This membrane stretches from the margin of the lamina spiralis to the ligamentum spirale ; the latter being merely an expanded portion of the connective tissue lining the osseous cochlea.

The lamina spiralis is produced into two lips, called respectively the labium tympanicum and labium vestibulare ; it is to the former and longer of these that the basilar membrane is attached. At the margin of the junction of the labium tympanicum with the basilar membrane the former is perforated for the passage of the nervous fibres, and this region is called the habenula perforata.

The labium vestibulare, so called from its position, is shorter than the labium tympanicum and is raised above into numerous blunt teeth. Partly springing out from the labium vestibulare, and passing from near the inner attachment of the membrane of Reissner towards the outer wall of the cochlea, is an elastic membrane, the membrana tectoria. Resting on the basilar membrane is the organ of Corti.

Considering for the moment that a transverse section of the cochlear canal only one cell deep is being dealt with, the organ of Corti will be found to consist of a central part composed of two peculiarly shaped rods widely separated below, but in contact above. These are the rods or fibres of Corti. On their outer side, i.e. on the side towards the osseous wall of the canal, is a reticulate membrane which passes from the inner rod of Corti towards the osseous wall of the canal. With their upper extremities fixed in that membrane, and their lower resting on the basilar membrane are three (four in man) cells with auditory hairs known as the outer 'hair cells,' which alternate with three other cells known as Deiters' cells. Between these and the outer attachment of the basilar membrane is a series of cells gradually diminishing in height in passing outwards. On the inner side of the rods of Corti is one hair cell, and then a number of peculiarly modified cells which fill up the space between the two lips of the lamina spiralis.

It will not be necessary to say much in reference to the development of the labium tympanicum and the labium vestibulare.

The labium vestibulare is formed by a growth of the connective tissue which fuses with and passes up between the epithelial cells. The epithelial cells which line its upper (vestibular) border become modified, and remain as its teeth.

The labium tympanicum is formed by the coalescence of the connective tissue layer separating the scala tympani from the cochlear canal with part of the connective tissue of the lamina spiralis. At first these two layers are separate, and the nerve fibres to the organ of Corti pass between them. Subsequently however they coalesce, and the region where they are penetrated by the nervous fibres becomes the habenula perforata.

The organ of Corti itself is derived from the epiblast cells lining the cochlear canal, and consists in the first instance of two epithelial ridges or projections. The larger of them forms the cells on the inner side of the organ of Corti, and the smaller the rods of Corti together with the inner and outer hair cells and Deiters' cells.

At first both these ridges are composed of simple elongated epithelial cells one row deep. The smaller ridge is the first to shew any change. The cells adjoining the larger ridge acquire auditory hairs at their free extremities, and form the row of inner hair cells ; the next row of cells acquires a broad attachment to the basilar membrane, and gives origin to the inner and outer rods of Corti.

Outside the latter come several rows of cells adhering together so as to form a compact mass which is quadrilateral in section. This mass is composed of three upper cells with nuclei at the same level, which form the outer hair cells, each of them ending above in auditory hairs, and three lower cells which form the cells of Deiters. Beyond this the cells gradually pass into ordinary cubical epithelial cells.

As just mentioned, the cells of the second row, resting with their broad bases on the basilar membrane, give rise to the rods of Corti. The breadth of the bases of these cells rapidly increases, and important changes take place in the structure of the cells themselves.


The nucleus of each cell divides ; so that there come to be two nuclei or sometimes three which lie close together near the base of the cell. Outside the nuclei on each side a fibrous cuticular band appears. The two bands pass from the base of the cell to its apex, and there meet though widely separated below. The remaining contents of the cell, between the two fibrous bands, become granular, and are soon to a great extent absorbed ; leaving at first a round, and then a triangular space between the two fibres. The two nuclei, surrounded by a small amount of granular matter, come to lie, each at one of the angles between the fibrous bands and the basilar membrane.

The two fibrous bands become, by changes which need not be described in detail, converted into the rods of Corti each of their upper ends growing outwards into the processes which the adult rods possess.

Each pair of rods of Corti is thus (Bottcher) to be considered as the product of one cell ; and the nuclei embedded in the granular mass between them are merely the remains of the two nuclei formed by the division of the original nucleus of that cell 1 . The larger ridge is for the most part not permanent, and from being the most conspicuous part of the organ of Corti comes to be far less important than the smaller ridge. Its cells undergo a partial degeneration ; so that the epithelium in the hollow between the two lips of the lamina spiralis, which is derived from the larger ridge, comes to be composed of a single row of short and broad cells. In the immediate neighbourhood however of the inner hair cell, one or two of the cells derived from the larger ridge are very much elongated.

The membrana reticularis is a cuticular structure derived from the parts to which it is attached. .

Accessory structures connected with the organ of hearing- in Terrestrial Vertebrata.

In all the Amphibia, Sauropsida and Mammalia, except the Urodela and a few Anura and Reptilia, the first visceral or hyomandibular cleft enters into intimate relations with the organs of hearing, and from it and the adjoining parts are formed the tympanic cavity, the Eustachian tube, the tympanic membrane and the meatus auditorius externus. The tympanic membrane serves to receive from the air the sound vibrations, which are communicated to fluids contained in the true auditory labyrinth by one ossicle or by a chain of auditory ossicles.

The addition to the organ of hearing of a tympanic membrane to receive aerial sound vibrations is an interesting case of the adaptation of a structure, originally required for hearing in water, to serve for hearing in air ; and as already pointed out, the similarity of this membrane to the tympanic membrane of some Insects is also striking.


1 It is not clear from Bottcher's description how it comes about that the inner rods of Corti are more numerous than the outer.


There is much that is obscure with reference to' the actual development of the above parts of the ear, which has moreover only been carefully studied in Birds and Mammals.

The Eustachian tube and tympanic cavity seem to be derived from the inner part of the first visceral or hyomandibular cleft, the external opening of which becomes soon obliterated. Kolliker holds that the tympanic cavity is simply a dorsally and posteriorly directed outgrowth of the median part of the inner section of this cleft; while Moldenhauer (No. 392) holds, if I understand him rightly, that it is formed as an outgrowth of a cavity called by him the sulcus tubo-tympanicus, derived from the inner aperture of the first visceral cleft together with the groove of the pharynx into which it opens ; and Moldenhauer is of opinion that the greater part of the original cleft atrophies.

The meatus auditorius externus is formed at the region of a shallow depression where the closure of the first visceral cleft takes place. It is in part formed by the tissue surrounding this depression growing up in the form of a wall, and Moldenhauer believes that this is the whole process. Kolliker states however that the blind end of the meatus becomes actually pushed in towards the tympanic cavity.

The tympanic membrane is derived from the tissue which separates the meatus auditorius externus from the tympanic cavity. This tissue is obviously constituted of an hypoblastic epithelium on its inner aspect, an epiblastic epithelium on its outer aspect, and a layer of mesoblast between them, and these three layers give rise to the three layers of which this membrane is formed in the adult. During the greater part of fcetal life it is relatively very thick, and presents a structure bearing but little resemblance to that in the adult.

A proliferation of the connective tissue-cells in the vicinity of the tympanic cavity causes in Mammalia the complete or nearly complete obliteration of the cavity during fcetal life.

The tympanic cavity is bounded on its inner aspect by the osseous investment of the internal ear, but at one point, known as the fenestra ovalis, the bone is deficient in the Amphibia, Sauropsida and Mammalia, and its place is taken by a membrane ; while in Mammalia and Sauropsida a second opening, the fenestra rotunda, is also present.

These two fenestrae appear early, but whether they are formed by an absorption of the cartilage, or by the nonchondrification of a small area, is not certainly known. The upper of the two, or fenestra ovalis, contains the base of a bone, known in the Sauropsida and Amphibia as the columella. The main part of the columella is formed of a stalk which is held by Parker to be derived from part of the skeleton of the visceral arches, but its nature is discussed in connection with the skeleton, while the base, forming the stapes, appears to be derived from the wall of the periotic cartilage.

In all Amphibia and Sauropsida with a tympanic cavity, the stalk of the columella extends to the tympanic membrane ; its outer end becoming imbedded in this membrane, and serving to transmit the vibrations of the membrane to the fluid in the internal ear. In Mammalia there is a stapes not directly attached to the tympanic membrane by a stalk, and two additional auditory ossicles, derived from parts of the skeleton of the visceral arches, are placed between the stapes and the tympanic membrane. These ossicles are known as the malleus and incus, and the chain of the three ossicles replaces physiologically the single ossicle of the lower forms.

These ossicles are at first imbedded in the connective tissue in the neighbourhood of the tympanic cavity, but on the full development of this cavity, become apparently placed within it ; though really enveloped in the mucous membrane lining it.

The fenestra ovalis is in immediate contiguity with the walls of the utricle, while the fenestra rotunda adjoins the scala tympani.

Hunt (No. 391) holds, from his investigations on the embryology of the pig, that " the Eustachian tube is an involution of the pharyngeal mucous membrane ;" and that "the meatus is an involution of the integument " while " the drum is formed by the Eustachian tube overlapping the extremity of the meatus." Urbantschitsch also holds that the first visceral cleft has nothing to do with the formation of the tympanic cavity and Eustachian tube, and that these parts are derived from lateral outgrowths of the oral cavity.

The evolution of the accessory parts of the ear would be very difficult to explain on Darwinian principles if the views of Hunt and Urbantschitsch were correct ; and the accepted doctrine, originally proposed by Huschke (No. 389), according to which these structures have originated by a ' change of function' of the parts of the first visceral cleft, may fairly be held till more conclusive evidence has been brought against it than has yet been done.

Tunicata. The auditory organ of the Tunicata (fig. 306) is placed on the under surface of the anterior vesicle of the brain.



FIG. 306. LARVA OF ASCIDIA MENTULA. (From Gegenbaur ; after Kupffer.)

Only the anterior part of the tail is represented.

N'. anterior swelling of neural tube ; IV. anterior swelling of spinal portion of neural tube; n. hinder part of neural tube; ch. notochord ; A", branchial region of alimentary tract ; d. cesophageal and gastric region of alimentary tract ; O. eye ; a, otolith ; o. mouth ; s. papilla for attachment.

It consists of two parts (i) a prominence of the cells of the floor of the brain forming a crista acustica, and (2) an otolith projecting into the cavity of the brain, and attached to the crista by delicate hairs.

The crista acustica is formed of very delicate cylindrical cells, and in its most projecting part is placed a vesicle with clear contents. The otolith is an oval body with its dorsal half pigmented, and its ventral half clear and highly refractive. It is balanced on the highest point of the crista.

The crista acustica would seem to be developed from the cells of the lower part of the front vesicle of the brain. The otolith however is developed from a single cell on the dorsal and right side of the brain. This cell commences to project into the cavity of the brain and its free end becomes pigmented. It gradually grows inwards till it forms a spherical prominence in the cavity of the brain, to the wall of which it is attached by a stalk. At the same time it travels round the right side of the vesicle of the brain (in a way not fully explained) till it reaches the summit of the crista, which has become in the meantime established.

The auditory organ of the simple Ascidians can hardly be brought into relation with that of the other Chordata, and has most probably been evolved within the Tunicate phylum.

Bibliography

Invertebrata.

(384) V. Hensen. "Studien lib. d. Gehororgan d. Decapoden." Zeit.f. wiss. ZooL, Vol. xm. 1863.

(385) O. and R. Hertwig. Das Nervensystem u. d. Sinnesorgane d. Medusen. Leipzig, 1878.

Vertebrata.

(386) A. Boettcher. "Bau u. Entwicklung d. Schnecke." Denkschriften d. kaiserl. Leop. Carol. Akad. d. Wissenschaft., Vol. xxxv.

(387) C. H asse. Die vergleich. Morphologic u. Histologied. hiiutigen Gehororgane d. Wirbelthiere. Leipzig, 1873.

(388) V. Hensen. "Zur Morphologic d. Schnecke." Zeit. f. wiss. ZooL, Vol. XIII. 1863.

(389) E. Huschke. "Ueb. d. erste Bildungsgeschichte d. Auges u. Ohres beim bebriiteten Kiichlein." Isis von Oken, 1831, and Meckel's Archiv, Vol. vi.

(390) Reissner. De Auris internes formatione. Inaug. Diss. Dorpat, 1851.

Accessory parts of Vertebrate Ear.

(391) David Hunt. "A comparative sketch of the development of the ear and eye in the Pig. " Transactions of the International Otological Congress, \ 876.

(392) W. Moldenhaueir. "Zur Entwick. d. mittleren u. ausseren Ohres." Morphol. Jahrbuch) Vol. III. 1877.

(393) V. Urbantschitsch. " Ueb. d. erste Anlage d. Mittelohres u. d. Trommelfelles." Mittheil. a. d. embryol. Instit. Wien, Heft i. 1877.

Olfactory organ

Amongst the Invertebrata numerous sense organs have been described under the title of olfactory organs. In aquatic animals they often have the form of ciliated pits or grooves, while in the Insects and Crustacea delicate hairs and other structures present on the antennae are usually believed to be organs of smell. Our knowledge of all these organs is however so vague that it would not be profitable to deal with them more fully in this place. Amongst the Chordata there are usually well developed olfactory organs.

Amongst the Urochorda (Tunicata) it is still uncertain what organs (if any) deserve this appellation. The organ on the dorsal side of the opening of the respiratory pharynx may very possibly have an olfactory function, but it is certainly not homologous with the olfactory pits of the true Vertebrata, and as mentioned above (pp. 436 and 437), may perhaps be homologous with the pituitary body.

In the Cephalochorda (Amphioxus) there is a shallow ciliated pit, discovered by Kolliker, which is situated on the left side of the head, and is closely connected with a special process of the front end of the brain. It is most probably the homologue of the olfactory pits of the true Vertebrata.



FlG. 307. VIEWS OF THE HEAD OF ELASMOBRANCH EMBRYOS AT TWO STAGES AS TRANSPARENT OBJECTS.

A. Pristiurus embryo of the same stage as fig. 28 F.

B. Somewhat older Scyllium embryo.

III. third nerve ; V. fifth nerve ; VII. seventh nerve ; au.n. auditory nerve ; gl. glossopharyngeal nerve; Vg. vagus nerve; fb. fore-brain; pn. pineal gland; mb. midbrain; hb. hind-brain; iv.v. fourth ventricle; cb. cerebellum; ol. olfactory pit; op. eye; au.V. auditory vesicle; m. mesoblast at base of brain; ch. notochord; kt. heart; Vc. visceral clefts; eg. external gills; //. sections of body cavity in the head.


In the true Vertebrata the olfactory organ has usually the form of a pair of pits, though in the Cyclostomata the organ is unpaired.

In all the Vertebrata with two olfactory pits these organs are formed from a pair of thickened patches of the epiblast, on the under side of the fore-brain, immediately in front of the mouth (fig. 307, ol). Each thickened patch of epiblast soon becomes involuted as a pit (fig. 308, N), the lining cells of which become the olfactory or Schneiderian epithelium. The surface of this epithelium is usually much increased by various foldings, which in the Elasmobranchii arise very early, and are bilaterally symmetrical, diverging on each side like the barbs of a feather from the median line. They subsequently become very pronounced (fig. 309), serving greatly to increase the surface of the olfactory epithelium. At a very early stage the olfactory nerve attaches itself to the olfactory epithelium.

In Petromyzon the olfactory organ arises as an unpaired thickening of the epiblast, which in the just hatched larva forms a shallow pit, on the ventral side of the head, immediately in front of the mouth. This pit rapidly deepens, and soon extends itself backwards nearly as far as the infundibulum (fig. 310, 0!}. By the development of the upper lip the opening of the olfactory pit is gradually carried to the dorsal surface of the head, and becomes at the same time narrowed and ciliated (fig. 47, ol). The whole organ forms an elongated sack, and in later stages becomes nearly divided by a median fold into two halves.

It is probable that the unpaired condition of the olfactory organ in the Lamprey has arisen from the fusion of two pits into one ; there is however no evidence of this in the early development ; but the division of the sack into two halves by a median fold may be regarded as an indication of such a paired character in the later stages.

In Myxine the olfactory organ communicates with the mouth through the palate, but the meaning of this communication, which does not appear to be of the same nature as the communication between the olfactory pits and the mouth by the posterior nares in the higher types, is not known.

The opening of the olfactory pit does not retain its embryonic characters. In Elasmobranchii and Chimaera it becomes enclosed by a wall of integument, often deficient on the side of the mouth, so that there is formed a groove leading from the nasal pit towards the angle of the mouth. This groove is usually constricted in the middle, and the original single opening of the nasal sack thus becomes nearly divided into two. In Teleostei and Ganoids the division of the nasal opening into two parts becomes complete, but the ventral opening is generally carried off some distance from the mouth, and placed, by the growth of the snout, on the upper surface of the head (figs. 54 and 68). In all these instances it is probable that the dorsal opening of the nasal sack is homologous with the external nares, and the ventral opening with the posterior nares of higher types. Thus the posterior nares would in fact seem to be represented in all Fishes by a ventral part of the opening of the original nasal pit which either adjoins the border of the mouth (many Elasmobranchii) or is quite separate from the mouth (Teleostei and Ganoidei). In the Dipnoi, Amphibia and all the higher types the oral region becomes extended so as to enclose the posterior nares, and then each nasal pit acquires two openings ; viz. one outside the mouth, the external nares, and one within the mouth, the internal or posterior nares. In the Dipnoi the two nasal openings are very similar to those in Ganoidei and Teleostei, but both are placed on the under surface of the head, the inner one being within the mouth, and the external one is so close to the outer border of the upper lip that it also has been considered by some anatomists to lie within the mouth.


EXTERNAL AND INTERNAL NARES.

In all the higher types the nasal pits have originally only a single opening, and the ontogenetic process by which the posterior nasal opening is formed has been studied in the Amniota and Amphibia. Amongst the Amniota we may take the Chick as representing the process in a very simple form. The general history of the process was first made out by Kolliker.


FIG. 308. SIDE VIEW OF THE HEAD OF AN EMBRYO CHICK OF THE THIRD DAY AS AN OPAQUE OBJECT. (Chromic acid preparation.)

C.H. cerebral hemispheres ; F.B. vesicle of third ventricle; M.B. mid-brain; Cb. cerebellum; H.B. medulla oblongata; N. nasal pit ; ot. auditory vesicle in the stage of a pit with the opening not yet closed up; op. optic vesicle, with /. lens and ch.f. choroidal fissure.

i F. The first visceral fold ; above it is seen the superior maxillary process.

2, 3, 4 F. Second, third and fourth visceral folds, with the visceral clefts between them.


The opening of the nasal pit becomes surrounded by a ridge except on its oral side. The deficiency of this ridge on the side of the mouth gives rise to a kind of shallow groove leading from the nasal pit to the mouth. The ridge enveloping the opening of the nasal pit next becomes prolonged along the sides of this groove, especially on its inner one; and at the same time the superior maxillary process grows forwards so as to bound the lower part of its outer side. The inner and outer ridges, together with the superior maxillary process, enclose a deep groove, connecting the original opening of the nasal pit with the mouth. The process just described is illustrated by fig. 311 A, and it may be seen that the ridge on the inner side of the groove forms the edge of the fronto-nasal process (k).



FIG. 309. SECTION THROUGH THE BRAIN AND OLFACTORY ORGAN OF AN EMBRYO OF SCYLLIUM. (Modified from figures by Marshall and myself.)

c.h. cerebral hemispheres; oLv. olfactory vesicle; olf, olfactory pit ; Seh. Schneiderian folds ; /. olfactory nerve. The reference line has been accidentally taken through the nerve to the brain.



FIG. 310. DIAGRAMMATIC VERTICAL SECTION THROUGH THE HEAD OF A LARVA OF PETROMYZON.

The larva had been hatched three days, and was 4-8 mm. in length. The optic and auditory vesicles are supposed to be seen through the tissues.

c.h. cerebral hemisphere; th. optic thalamus; in. infundibulum ; pn. pineal gland; mb. mid-brain; cb. cerebellum; md. medulla oblongata; au.v. auditory vesicle; op. optic vesicle; ol. olfactory pit; m. mouth; br.c. branchial pouches; th. thyroid involution; v. ao. ventral aorta ; ht. ventricle of heart ; ch. notochord.


On the sixth day (Born, 394) the sides of this groove unite together in the middle, and convert it into a canal open at both ends the ventral openings of the canals of the two sides being placed just within the border of the mouth, and forming the posterior nares ; while the external openings form the anterior nares. The upper part of the canal, together with the original



FIG. 311. HEAD OF A CHICK FROM BELOW ON THE SIXTH AND SEVENTH DAYS OF INCUBATION. (From Huxley.)

/". cerebral vesicles ; a. eye, in which the remains of the choroid slit can still be seen in A ; g. nasal pits ; k. fronto-nasal process ; /. superior maxillary process ; i. inferior maxillary process or first visceral arch; 2. second visceral arch; x. first visceral cleft.

In A the cavity of the mouth is seen enclosed by the fronto-nasal process, the superior maxillary processes and the first pair of visceral arches. At the back of it is seen the opening leading into the throat. The nasal grooves leading from the nasal pits to the mouth are already closed over.

In B the external opening of the mouth has become much constricted, but it is still enclosed by the fronto-nasal process and superior maxillary processes above, and by the inferior maxillary processes (first pair of visceral arches) below.

The superior maxillary processes have united with the fronto nasal process, along nearly the whole length of the latter.

nasal pit, is alone lined by olfactory epithelium ; the remaining epithelium of the nasal cavity being indifferent epiblastic epi


OLFACTORY ORGANS.


537


thelium. Further changes subsequently take place in connection with the posterior nares, but these are described in the section dealing with the mouth.

In Mammalia the general formation of the anterior and posterior nares is the same as in Birds ; but, as shewn by Dursy and Kolliker, an outgrowth from the inner side of the canal between the two openings arises at an early period ; and becoming separate from the posterior nares and provided with a special opening into the mouth, forms the organ of Jacobson. The general relations of this organ when fully formed are shewn in fig. 312.

In Lacertilia the formation of the posterior nares differs in some particulars from that in Birds (Born). A groove is formed leading from the primitive nasal pit to the mouth, bordered on its inner side by the swollen edge of the fronto-nasal process, and on its outer by an outernasal process ; while the superior maxillary process does not assist in bounding it. On the inner side of the narrowest part of this groove there is formed a large lateral diverticulum, which is lined by a continuation of the Schneiderian epithelium, and forms the rudiment of Jacobson's organ. The nasal groove continues to grow in length, but soon becomes converted into a canal by the junction of the outer-nasal process with the fronto-nasal process. This canal is open at both ends : at its dorsal end is placed the original opening of the nasal pit, and its ventral opening is situated within the cavity of the mouth. The latter forms the primitive posterior nares. The superior maxillary process soon grows inwards on the under side of the posterior part of the nasal passage, and assists in forming its under wall. This ingrowth of the superior maxillary process is the rudiment of the hard palate.

On the conversion of the nasal groove into a closed passage, the opening of Jacobson's organ into the groove becomes concealed ; and at a later period Jacobson's organ becomes completely shut off from the nasal cavity, and opens into the mouth at the front end of an elongated groove leading back to the posterior nares.

In Amphibia the posterior nares are formed in a manner very different from that of the Amniota. At an early stage a shallow groove is formed leading from the nasal pit to the mouth ; but this groove instead of forming the posterior nares soon vanishes, and by the growth of the front of the head the nasal pits are carried farther away from the mouth.


FIG. 312. SECTION THROUGH THE NASAL CAVITY AND JA COBSON'S ORGAN. (From Gegenbaur.)

sn. septum nasi ; en. nasal cavity ; y. Jacobson's organ ; d, edge of upper jaw.



The actual posterior nares are formed by a perforation in the palate, opening into the blind end of the original nasal pit.

Considering that the various stages in the formation of the posterior nares of the Amniota are so many repetitions of the adult states of lower forms, it may probably be assumed that the mode of formation of the posterior nares in Amphibia is secondary, as compared with that in the Amniota.

A diverticulum of the front part of the nasal cavity of the Anura is probably to be regarded as a rudimentary form of Jacobson's organ.

Bibliography

(394) G. Born. "Die Nasenhohlen u. d. Thranennasengang d. amnioten Wirbelthiere." Parts I. and II. Morphologisches Jahrbuch, Bd. V., 1879.

(395) A. Kollicker. " Ueber die Jacobson'schen Organe des Menschen." Festschrift f. Rienecker, 1877.

(396) A. M. Marshall. "Morphology of the Vertebrate Olfactory Organ." Quart. Journ. of Micr. Science, Vol. xix., 1879.

Sense organs of the lateral line.

Although I do not propose dealing with the general development of various sense organs of the skin, there is one set of organs, viz. that of the lateral line, which, both from its wide extension amongst the Ichthyopsida and from the similarity of some of its parts to certain organs found amongst the Chastopoda 1 , has a great morphological importance.

The organs of the lateral line consist as a rule of canals, partly situated in the head, and partly in the trunk. These canals open at intervals on the surface, and their walls contain a series of nerve-endings. The branches of the canal in the head are innervated for the most part by the fifth pair, and those of the trunk by the nervus lateralis of the vagus nerve. There is typically but a single canal in the trunk, the openings and nerve-endings of which are segmentally arranged.

Two types of development of these organs have been found. One of these is characteristic of Teleostei ; the other of Elasmobranchii.

In just hatched Teleostei, Schulze (No. 402) found that instead of the normal canals there was present a series of sense bulbs, projecting freely on the surface and partly composed of cells with stiff hairs. In most cases each bulb is enclosed in a delicate tube open at its free extremity ; while the bulbs correspond in number with the myotomes. In some Teleostei (Gobius, Esox, etc.) such sense organs persist through life ; in most forms however each organ becomes covered by a pair of lobes of the adjacent tissue, one formed above and the other below it. The two lobes of each pair then unite and form a tube open at both ends. The linear series of tubes so formed is the commencement of the adult canal ; while the primitive sense bulbs form the sensory organs of the tubes. The adjacent tubes partially unite into a continuous canal, but at their points of apposition pores are left, which place the canal in communication with the exterior.

1 The organs which resemble those of the lateral line are the remarkable sense organs found by Eisig in the Capitellidse (Mittheil. a. d. ZooL Station zu Neapel, Vol. I.) ; but I am not inclined to think that there is a true homology between these organs and the lateral line of Vertebrata. It seems to me probable that the segmentally arranged optic organs of Polyophthalmus are a special modification of the more indifferent sense organs of the Capitellidse. The close affinity of these two types of Chsetopods is favourable to this view.



Besides these parts, I have found that there is present in the just hatched Salmon a linear streak of modified epidermis on the level of the lateral nerve, and from the analogy of the process described below for Elasmobranchii it appears to me probable that these streaks play some part in the formation of the canal of the lateral line.

In Elasmobranchii (Scyllium) the lateral line is formed as a linear thickening of the mucous layer of the epidermis. This thickening is at first very short, but gradually grows backwards, its hinder end forming a kind of enlarged growing point. The lateral nerve is formed shortly after the lateral line, and by the time that the lateral line has reached the level of the anus the lateral nerve has grown back for about two-thirds of that distance. The lateral nerve would seem to be formed as a branch of the vagus, but is at first half enclosed in the modified cells of the lateral line (fig. 275, nl) 1 , though it soon assumes a deeper position.

A permanent stage, more or less corresponding to the stage just described in Elasmobranchii, is retained in Chimasra, and Echinorhinus spinosus, where the lateral line has the form of an open groove (Solger, No. 404).

The epidermic thickening, which forms the lateral line, is converted into a canal, not as in Teleostei by the folding over of the sides, but by the formation of a cavity between the mucous and epidermic layers of the epiblast, and the subsequent enclosure of this cavity by the modified cells of the mucous layer of the epiblast which constitute the lateral line. The cavity first appears at the hind end of the organ, and thence extends forwards.

After its conversion into a canal the lateral line gradually recedes from the surface ; remaining however connected with the epidermis at a series of points corresponding with the segments, and at these points perforations are eventually formed to constitute the segmental apertures of the system.

The manner in which the lumen of the canal is formed in Elasmobranchs bears the same relation to the ordinary process of conversion of a groove into a canal that the formation of the auditory involution

1 Gotte and Semper both hold that the lateral nerve, instead of growing in a centrifugal manner like other nerves, is directly derived from the epiblast of the lateral line. For the reasons which prevent me accepting this view I must refer the reader to my Monograph on Elasmobranch Fishes, pp. 141 146.

in Amphibia does to the same process in Birds. In both Elasmobranchii and Amphibia the mucous layer of the epiblast behaves exactly as does the whole epiblast in the other types, but is shut off from the surface by the passive epidermic layer of the epiblast.

The mucous canals of the head and the ampullae are formed from the mucous layer of the epidermis in a manner very similar to the lateral line ; but the nerves to them arise as simple branches of the fifth and seventh nerves, which unite with them at a series of points, but do not follow their course like the lateral nerve.

It is clear that the canal of the lateral line is secondary, as compared with the open groove of Chimaera or the segmentally arranged sense bulbs of young Teleostei ; and it is also clear that the phylogenetic mode of formation of the canal consisted in the closure of a primitively open groove. The abbreviation of this process in Elasmobranchii was probably acquired after the appearance of food-yolk in the egg, and the consequent disappearance of a free larval stage.

While the above points are fairly obvious it does not seem easy to decide a priori whether a continuous sense groove or isolated sense bulbs were the primitive structures from which the canals of the lateral line took their origin. It is equally easy to picture the evolution of the canal of the lateral line either from (i) a continuous unsegmented sense line, certain points of which became segmentally differentiated into special sense bulbs, while the whole subsequently formed a groove and then a canal ; or from (2) a series of isolated sense bulbs, for each of which a protective groove was developed ; and from the linear fusion of which a continuous canal became formed.

From the presence however of a linear streak of modified epidermis in larval Teleostei, as well as in Elasmobranchii, it appears to me more probable that a linear sense streak was the primitive structure from which all the modifications of the lateral line took their origin, and that the segmentally arranged sense bulbs of Teleostei are secondary differentiations of this primitive structure.

The, at first sight remarkable, distribution of the vagus nerve to the lateral line is probably to be explained in connection with the evolution of this organ. As is indicated both by its innervation from the vagus, as also from the region where it first becomes developed, the lateral line was probably originally restricted to the anterior part of the body. As it became prolonged backwards it naturally carried with it the vagus nerve, and thus a sensory branch of this nerve has come to innervate a region which is far beyond the limits of its original distribution.

Bibliography

(397) F. M. Balfour. A Monograph onthe development of Elasnwbranch Fishes, pp. 141 146. London, 1878.

(398) H. Eisig. "Die Segmentalorgane cl. Capitelliden." Mitthcil. a. d. zool. Station zu Neapel> Vol. I. 1879.

(399) A. Gotte. Entwicklungsgeschichte d. Unke. Leipzig, 1875.

(400) Fr. Leydig. Lehrbuch d. Histologie des Memchen u. d. Thiere. Hamm.

1857 (401) Fr. Leydig. Neue Beitrdge z. anat. Kenntniss d. Hautdecke u. Hautsinnesorgane d. Fische. Halle, 1879.

(402) F. E. Schulze. " Ueb. d. Sinnesorgane d. Seitenlinie bei Fischen und Amphibien." Archiv f. mikr. Anat., Vol. vi. 1870.

(403) C. Semper. "Das Urogenitalsy stem d. Selachier." Arbeit, a. d. zoo!.zoot. Instil. Wiirzburg, Vol. II.

(404) B. Solger. "Neue Untersuchungen zur Anat. d. Seitenorgane d. Fische." Archiv f. mikr. Anat., Vol. xvn. and xvm. 1879 an< * !88o.