fig211

Fig. 211.—Dorsal (A) and ventral (B) views of the brain of Petromyzon marinus. ch.pl.1, Anterior choroid plexus forming the roof of the prosencephalon and thalamencephalon; ch.pl.2, aperture in the roof of the mid-brain exposed by the removal of the middle choroid plexus; ch.pl.3, the fourth ventricle exposed by the removal of the posterior plexus; cr.crb, crura cerebri; crb, cerebellum; crb.h, cerebral hemispheres; dien, thalamencephalon; inf, infundibulum; l.gn.hb, left ganglion habenulae; med.obl, medulla oblongata; nv.1, olfactory; nv.2, optic; nv.3, oculomotor; nv.5, trigeminal; and nv.8, auditory nerves; olf.l, olfactory lobes; opt.l, optic lobes; pn, pineal organ; r.gn.hb, right ganglion habenulae. (From Parker and Haswell, after Ahlborn.)

fig212

Fig. 212.—Dorsal view of the brain of Myxine. c.r, Corpora restiformia; m.o, medulla oblongata; n.p, naso-pituitary canal; ol.o, olfactory organ enclosed in its fenestrated cartilaginous capsule; op.l, optic lobes; pr, prosencephalon; s, s, dorsal roots of spinal nerves; sp.c, spinal cord; th, thalamencephalon. (From Wiedersheim, after Retzius.)

In the Cyclostome Petromyzon there is a small prosencephalon with an undivided prosocoele, and on each side of it a small cerebral hemisphere which appears as a mere appendage to the much larger olfactory lobe (Fig. 211). The prosocoele divides in front into two outwardly directed branches, and of the two diverticula into which each branch divides one extends as a lateral ventricle into the hemisphere of its side, and the other as a rhinocoele into the corresponding olfactory lobe. The ganglia habenulae are unusually large, the right one being larger than the left. The optic lobes are large, but not obviously double. So small is the cerebellum that it seems to be little more than a narrow transverse band crossing the fore-part of the fourth ventricle. The roof of the brain is largely epithelial, especially in the prosencephalon, the thalamencephalon, and the hind-brain. Over these epithelial areas the pia mater is unusually vascular and forms a series of "choroid plexuses." The ventricular system is complete and continuous. By contrast with the Lamprey the brain of Myxine[442] is very primitive, more so perhaps than in any other Craniate (Fig. 212). In a dorsal view the brain is divided into four pairs of laterally expanded and longitudinally compressed lobes by a median longitudinal fissure and three transverse fissures. The two anterior lobes are little more than the thickened anterior wall of the thalamencephalon, although, judging from their histological structure, they represent a very imperfectly differentiated prosencephalon and olfactory lobes. The second and largest pair constitute the thalamencephalon. The last two pairs of lobes represent a transversely divided pair of optic lobes, or "corpora quadrigemina." There is a large medulla oblongata with a pair of corpora restiformia, but the cerebellum is entirely absent. The ventricles are subject to some individual variation. Third and fourth ventricles are generally recognisable, either as isolated cavities or connected by a remnant of the mesocoele. In the feeble development of the prosencephalon, in the striking preponderance of the mid-brain over the rest of the brain, and in the absence of a cerebellum, Myxine is unique amongst Craniates.

fig213

Fig. 213.—The brain of a Dog-Fish (Scyllium canicula). A, dorsal view; B, ventral view. The choroid plexuses covering the roof of the third and fourth ventricles have been removed. b.o, Olfactory lobe; ep, origin of the stalk of the pineal body; f.b (in A), prosencephalon; f.b (in B), cerebral hemispheres; fr, fourth ventricle; h.b, cerebellum; h.p, pituitary body; i.f, lobi inferiores; m.b, optic lobes; m.d, medulla oblongata; sc, saccus vasculosus; th, thalamencephalon; t.o (i) olfactory peduncle; i.-x. cranial nerves. (From Wiedersheim.)

In Elasmobranchs among Fishes the brain attains a much higher grade of structure. In Scyllium (Fig. 213) there is a large prosencephalon, and directly in front of it a pair of imperfectly differentiated cerebral hemispheres, while from its antero-lateral regions the large olfactory lobes arise. The prosocoele divides in front into four diverticula, of which the two inner ones extend into the hemispheres as lateral ventricles, and the two outer as rhinocoeles into the olfactory lobes (Fig. 214). In connexion with the infundibulum there is a pair of sacci vasculosi, consisting mainly of gland-tubules, opening into the infundibular cavity.[443] The cerebellum is exceptionally large, but it does not form a "valvula cerebelli." Large ear-like corpora restiformia are present. The third and fourth ventricles alone retain an epithelial roof in relation with choroid plexuses.

In all essentials the brain of the Holocephali is a repetition of the Elasmobranch type, more especially of the elongated form seen in Notidanus. Indications of a higher grade of structure are, however, to be seen in the reduction of the prosencephalon which, with its prosocoele, is now scarcely distinguishable from the thalamencephalon and its ventricle; and in the more complete differentiation of the cerebral hemispheres from one another and from the rest of the brain. Large frilled corpora restiformia are conspicuous structures on each side of the medulla oblongata. Besides the usual intra-cranial pituitary body, there is also a separate extra-cranial portion lodged in a pit on the ventral surface of the basis cranii: in the embryo the two are continuous.

fig214

Fig. 214.—Horizontal longitudinal section of brain of Chiloscyllium, to show the ventricles; semi-diagrammatic. cer, Origin of cerebellar ventricle or epicoele; dia, third ventricle; iter, mesocoele; meta, fourth ventricle; opt, optocoele, or cavity of an optic lobe; para, lateral ventricles; pros, prosocoele; rh, rhinocoele. (From Parker and Haswell.)

In the Teleostomi the brain is distinctly of a more primitive type than in any other Fishes (Fig. 215).[444] The most striking feature is the absence of cerebral hemispheres, the evolution of the primary fore-brain proceeding no farther than the formation of an undivided prosencephalon with a non-nervous roof, and a prosocoele which forms a continuous cavity with the third ventricle, or at the most is only separated from it by an infolding of the epithelial roof or velum transversum.

fig215

Fig. 215.A, dorsal view of the brain of a Trout (Salmo fario); B, a vertical longitudinal section. c.il, Commissura interlobularis; g.h, ganglion habenulae; h.c, habenular commissure; i.c, inferior commissure; l.i, lobus inferior; myc, myelocoele; p.c, posterior commissure; v.o, valvula cerebelli; v.t, velum transversum; ii., optic nerve; v.iii., v.iv., third and fourth ventricles; v, vii, viii, ix, x, fifth, seventh, eighth, ninth, and tenth cranial nerves; remaining reference letters as in Fig. 210. (A, From Wiedersheim; B, after Haller.)

Amongst other diagnostic characters may be mentioned the predominance of the mid-brain over the other divisions, the anterior extension of the large cerebellum into the mesocoele as a "valvula cerebelli," and the absence of corpora restiformia. This type of brain is most strongly marked in the Teleostei, but in other Teleostomes some, like Acipenser,[445] are typically Teleostean in this respect (Fig. 216), while others, such as Lepidosteus, have small cerebral hemispheres with lateral ventricles as well as a prosencephalon.

fig216

Fig. 216.—Vertical longitudinal section of the brain of a Sturgeon (Acipenser ruthenus.) c.p, Posterior commissure; c.r, cranial roof; mc, mesocoele; op.ch, optic chiasma; p.ch.p, posterior choroid plexus; v.c, valvula cerebelli; v.t, velum transversum; v.iii, v.iv., the third and fourth ventricles; other lettering as in Fig. 210. (From Goronowitsch.)

The most obvious feature in the brain of the Dipnoi is the great development of the cerebral hemispheres. In this respect these Fishes approach the Amphibia, but in other features of brain-structure they present points of agreement with most other groups of Fishes without being closely related to any one of them. In Protopterus[446] (Fig. 217) the hemispheres are quite distinct except behind, and the walls of their spacious lateral ventricles are entirely nervous. Olfactory lobes are sessile on their anterior extremities, and behind and below they enlarge into ventral lobes which probably represent the hippocampal lobes of the higher Vertebrates. A vesicular pineal body at the end of a slender stalk overlies a singular conical projection from the roof of the thalamencephalon or "pineal pillow."

fig217

Fig. 217.—Dorsal (A), ventral (B), and lateral (C) views of the brain of Protopterus annectens. C, Cerebellum; C.H, cerebral hemisphere; D.S.E, branches of the sinus endolymphaticus; In, infundibulum; L.I,, lobi inferiores; M.O, medulla oblongata; O.L, olfactory lobe; Op.L, optic lobe; P, pituitary body; P.B, "pineal pillow"; S.E, sinus endolymphaticus; Sp.c, spinal cord; Sp.n, spinal nerve; Vel, velum transversum; Z, pineal body; IV.V., fourth ventricle; ii., iii., iv., v., vi., vii., viii.1, viii.2, viii.3.4, ix., and x., roots of the cranial nerves. (From Burckhardt.)

The optic lobes form a single oval body, and, as in Petromyzon and the Amphibia, the cerebellum is very small. A posterior choroid plexus covers the roof of the fourth ventricle, and an anterior plexus in connexion with the roof of the thalamencephalon projects downwards into the third ventricle, and is also prolonged forwards into each lateral ventricle. In Neoceratodus[447] the brain is certainly more primitive and distinctly less Amphibian. As compared with Protopterus the olfactory lobes and the cerebellum are larger, and the optic lobes are paired. The smaller hemispheres are non-nervous dorsally and medianly, the roof and inner wall of each being formed by an extension of the thick, glandular choroid plexus which forms the roof of the thalamencephalon.

The Spinal Nerves.—The spinal nerves of Cyclostomes (e.g. Petromyzon) consist of a series of dorsal nerves arising on each side from the dorsal surface of the spinal cord, and of a similar double series arising from the ventral surface, the dorsal nerves regularly alternating with the ventral nerves. Each myotome is supplied by a dorsal and a ventral nerve which pass separately to their peripheral distribution in the skin and muscles. In Fishes, as in the higher Vertebrates, each dorsal nerve, now termed a dorsal root, enlarges into a ganglion and then unites, either before or directly after issuing from the neural canal, with the next ventral nerve or ventral root in front to form a main spinal nerve. At the same time the spinal nerves of opposite sides tend to form pairs in the same transverse plane. After the union of the two roots the spinal nerve divides into three typical branches: a dorsal nerve (ramus dorsalis), and a ventral nerve (ramus ventralis), both of which include somatic sensory or afferent fibres, and somatic motor or efferent fibres, for the innervation of the skin and muscles of the dorsal and lateral portions of a myotome; and a visceral branch (ramus visceralis), composed of afferent and efferent visceral fibres, which supplies the adjacent viscera (alimentary canal and its glands and blood-vessels), and helps to form the sympathetic nervous system.[448] The somatic afferent and the visceral afferent fibres enter the spinal cord by the dorsal roots, the somatic efferent leaving the cord through the ventral roots, although the visceral efferent fibres traverse both roots. In the region of the paired fins more or fewer of the rami ventrales unite to form a plexus, the brachial or the pelvic plexus, from which the nerves to the fins take their origin.

The Cranial Nerves.—It is usual to describe the cranial nerves of Cyclostomes and Fishes as consisting of ten serially disposed pairs, viz.: the olfactory (i.), optic (ii.), oculomotor (iii.), trochlear (iv.), trigeminal (v.), abducens (vi.), facial (vii.), auditory (viii.), glossopharyngeal (ix.), and the vagus (x.) Like the spinal nerves, the cranial nerves collectively include somatic sensory (general cutaneous) and motor fibres, and also visceral sensory and motor fibres, all of which have their own special centres in the brain, but the proportions of these nerve components differ greatly in different nerves. Certain preoral nerves (iii., iv., and vi.) are exclusively somatic motor; others (i. and ii.) are special sensory nerves for the olfactory and visual organs; but most of the other cranial nerves include several components, and are therefore "mixed" nerves. Besides these components some cranial nerves include also a quasi-independent system of nerve-fibres, which converge from certain cutaneous sense-organs to an independent centre in the medulla oblongata, the tuber acusticum,[449] and is probably derived from the general cutaneous system of nerve components. Such nerve fibres, including also the auditory nerve, which has its origin from the same centre, constitute the lateralis system. Perhaps the most striking feature in the postoral cranial nerves is the predominance of the visceralis or sympathetic system over the somatic. Omitting the lateralis fibres and a relatively few somatic sensory fibres, visceral fibres, sensory and motor, are the principal components of all these nerves, including v. but excluding viii. The reason for this is to be found in the fact that splanchnic or visceral muscles in relation with the jaws and branchial arches have usurped the place of somatic muscles in the muscular system of the head. For developmental and other reasons the olfactory and optic nerves stand in a category of their own, and the same may be said of the third, fourth, and sixth nerves, which innervate the muscles of the eyeball. The remaining nerves, all of which have their origin in the medulla oblongata, possess certain features in common, and as they are related to the gill-clefts in such a way that each forks over a cleft, they may be conveniently distinguished as "branchial" or "branchiomeric nerves." A typical branchial nerve consists of (1) a principal ganglion near the origin of the nerve from the brain; (2) a main trunk which gives off (3) a somatic sensory branch or dorsal nerve to the skin; (4) a palatine nerve (visceral sensory) to the oral or pharyngeal mucous membrane; (5) an epibranchial ganglion which is associated with a transitory embryonic epibranchial sensory organ at the dorsal border of a branchial cleft; (6) a pre-branchial nerve (visceral sensory), skirting the anterior margin of a cleft in its ventral course; and (7) a post-branchial branch (visceral motor) similarly related to the hinder margin.

fig218

Fig. 218.—Diagram showing the principal branches of the cranial nerves in a Fish, mk.c, Meckel's cartilage; ol.o, olfactory organ; p.q, palato-quadrate; s, spiracle; i-v, branchial clefts; i, ii, iii, iv, vi, the first, second, third, fourth, and sixth cranial nerves. The remaining nerves are differently shaded. Black.—The trigeminal nerve: g.g, Gasserian ganglion; md, mandibularis; mx, maxillaris; op.p, ophthalmicus profundus. Oblique shading.—The lateralis system and its centre (t.a), the tuber acusticum: bucc.vii, buccalis branch of vii; md.ex, external mandibular branches of vii; l.n.x, lateralis nerve, with its supra-temporal branch (s.t), and its commissural connexion (c) with op.s.vii, the ophthalmicus superficialis of vii; viii, auditory nerve. Dotted.—The facialis proper, including c.t, chorda tympani; gn.g, geniculate ganglion; hy, hyomandibularis, with its motor branches m, m, m; p.n.vii., palatine. Dark grey.—The glossopharyngeal (ix), with its pre- and post-branchial branches and its palatine nerve, p.n.ix; anastomosing with the palatine branch of vii (Jacobson's anastomosis). White.—The vagus: x1-4, the branchial nerves, ganglionated and forking over clefts ii-v; v.n.x, visceral nerve; oc, occipito-spinal nerves; d.r and v.r, the dorsal and ventral roots of the first two spinal nerves. (Slightly modified after Wiedersheim.)

The first six cranial nerves resemble those of the higher Craniates in their mode of origin from the brain, in the physiological nature of their component fibres, and in their peripheral distribution, and therefore they need not be specially referred to here. The principal branches of the fifth or trigeminal nerve are shown in Fig. 218. Comparing this nerve with a typical branchial nerve it would seem that the profundus and superficialis ophthalmic nerves are dorsal nerves; the maxillaris and mandibularis, pre- and post-branchial branches, respectively, in relation with the modified gill-cleft which forms the mouth, while the branch to the oral surface represents a palatine nerve. The most important of the distinctive features in the cranial nerves of Fishes are to be found in the relations of nerves vii., ix., and x. to branchial clefts, and in the lateralis system of nerve components and its association with the lateral line sensory organs. The seventh or facial nerve is an exceptionally interesting nerve. Besides the usual components of a typical branchial nerve certain of its so-called branches are wholly or largely derived from the lateralis system. For this reason the nerve may be said to consist of two portions, the facial proper, or those fibres which constitute the facial nerve in air-breathing Craniates, and the lateralis branches which solely innervate lateral line sense-organs, and are therefore peculiar to aquatic forms. The facial proper has a ganglion (the facial or geniculate ganglion) on its root, and on entering the orbit after traversing the cranial wall it gives off a palatine nerve. Just over the spiracle a pre-branchial nerve, the representative of the chorda tympani of Mammals, leaves the main trunk, and passes ventrally in relation with the anterior wall of the spiracle to its ultimate distribution in the walls of the mouth cavity. The main trunk, now called the ramus hyomandibularis, then pursues a ventral course behind the spiracle as a post-branchial nerve, and certain of its mainly motor branches which pass downwards in connexion with the hyoid arch supply the muscles of that arch, and, if an operculum is present, the opercular muscles as well. The lateralis portion of the facial includes the following principal branches, each of which may have a ganglion on its root: (1) an ophthalmicus superficialis; (2) a buccalis nerve with its ramus oticus; and (3) external mandibular nerves which course in the ramus hyomandibularis. The addition of the great lateralis nerve, which is usually described as the lateral branch of the tenth nerve, and of the eighth or auditory nerve which supplies the auditory organ, completes the enumeration of the main factors of the lateralis system. The ninth or glosso-pharyngeal nerve, perhaps the most typical of all the branchial nerves, has pre- and post-branchial branches which enclose the hyo-branchial cleft. Its palatine nerve usually extends forwards and anastomoses with the corresponding branch of the seventh, thus forming a connexion (Jacobson's anastomosis) between the two cranial nerves. In some Elasmobranchs and Teleosts fibres derived from the dorsal branch of the ninth nerve innervate a few sense-organs of the lateral sensory canal of the head, and hence that nerve sometimes contains lateralis fibres. The tenth or vagus is a compound nerve. Besides the great lateralis nerve generally associated with it, the vagus includes as many typical branchial nerves as there are branchial clefts behind the hyo-branchial cleft, and in Elasmobranchs and in Chimaera these nerves have independent origins from the medulla oblongata. Each nerve has the typical structure, a ganglionated trunk which forks over a gill-cleft into the usual pre- and post-branchial branches, and palatine branches to the pharyngeal walls. In the Dipnoi the lateralis nerve is connected with the superficial ophthalmic branch of the seventh nerve by a commisural nerve which curves across the outer face of the auditory capsule. A somewhat similar anastomosis is also present in Petromyzon. The vagus also includes a large ramus intestinalis, which in Elasmobranchs, at all events, has a distinct ganglionated root. The nerve forms characteristic plexuses on the oesophagus and stomach, and in Cyclostomes its branches may extend nearly the whole length of the intestine. In Ganoids and Teleosts there is an interesting nerve known as the "lateralis accessorius." It is a compound nerve, and owes its formation to the union of somatic sensory fibres derived in succession from dorsal branches of the v., vii., ix., and x. nerves, and also from the corresponding branches of a variable number of spinal nerves. The finer branches of the nerve are distributed to the skin of one or more of the fins, or even, as in Gadus, to all the fins, especially to the numerous "end-buds" which are present on those organs. In many Fishes a variable number of the anterior spinal nerves (spino-occipital) perforate the occipital region of the skull. They probably represent the ventral roots only of the ordinary spinal nerves of this region.

Sense-Organs

The Cutaneous Sense-Organs.—These organs, the most remarkable and certainly the most characteristic of the sense-organs of Cyclostomes and Fishes, are bud-like groups of epidermic cells in relation with the ends of sensory nerve fibres. Each consists of a central core of sensory cells, provided with terminal cuticular sensory hairs, and surrounded by a zone of supporting and mucus-secreting cells which leave the hairs exposed at the apex of the bud. Two kinds of these organs can be distinguished, which differ in their innervation and in their position in the skin. Of the two, the so-called end-buds are the more primitive. They occupy a superficial position in the epidermis, and their sense-cells are as long as the supporting cells. They are present in Cyclostomes and Elasmobranchs, and especially in Teleosts, where they are irregularly distributed over the surface of the body, on the fins, lips, and barbels, and also in the epithelium of the mouth and pharynx. In the Dipnoi they are limited to the oral cavity, and in the higher Craniates they become taste-buds.[450] Their somatic sensory nerves[451] are derived from the vii., ix., and x. cranial nerves, and the lateralis accessorius. In the second type, usually called "nerve-eminences," the sensory cells are shorter than the supporting cells, and they are always innervated by the lateralis system. When first developed in the embryo they are quite superficial, like end-buds, but later the epidermis in which they lie sinks inwards so as to line a series of pits, closed sacs, tubes, open grooves, or closed canals. Pit-organs, so abundant on the head and trunk of Teleosts (Fig. 220), are simple epidermic pits with insunken nerve-eminences, disposed in groups or in lines (accessory lateral lines) or irregularly distributed. The "Spalt-papillen" of Elasmobranchs are pit-organs in which the orifice of the pit is reduced to a slit. The more deeply-seated Savi's vesicles on the ventral surface of the Torpedo, and the nerve-sacs of Ganoids, are similar organs converted into closed sacs and pinched off from the rest of the epidermis. Lorenzini's ampullae or mucus canals, which are found in definitely located groups on the lateral and upper surfaces of the head in Elasmobranchs, may perhaps be compared to pit-organs prolonged inwards to form subcutaneous tubes, each of which terminates in a radially-septate, chambered dilatation or ampulla, containing groups of sensory cells.

Besides the more diffusely scattered sense-organs there are others which become disposed in definite lines along the sides of the body and on the head, and, enclosed in grooves or closed canals, constitute the highly characteristic lateral line system of Cyclostomes and Fishes.[452] The auditory organ must also be included as a specialised portion of this system. Both organs are innervated by the lateralis system, and both arise from a common rudiment in the embryonic epidermis in the position of the future auditory organ. This rudiment grows backwards along the side of the body in the form of a cord of cells differentiated from the epidermis, and also forwards, where it soon divides into the rudiments of future supra-orbital and infra-orbital canals. Sense-organs are differentiated at intervals along the line of the cord; and in the body, but not on the head, they frequently exhibit a segmental disposition. Each sensory organ then sinks down into a short epidermic groove, which by the subsequent meeting of its lips becomes a canal detached from the epidermis. The short canals then become continuous, leaving, however, an externally opening primary pore between every two consecutive canals, and the result is a continuous canal having sense-organs imbedded in its epidermic lining and connected with the exterior by pores at intervals (Fig. 219).[453] The enclosure of the canals in the scales of the lateral line of the trunk or in special drain-pipe ossicles on the head, and the dichotomous subdivision of the primary pores into groups of surface-pores, complete the evolution of the system in its more advanced condition.

fig219

Fig. 219.—Vertical longitudinal section through the lateral canal of Amia calva. l.n, Lateralis nerve with its branches, n, n, to the sensory organs, s.o, s.o; p, p, p, external pores; s.c, sensory canal; s, s, scales of the lateral line. (From Wiedersheim, after Allis.)

Typically, the lateral line system consists of certain canals or grooves, usually but not invariably continuous, and defined by their innervation, (i.) a lateral canal extending along the side of the body and the hinder part of the head, and having its sensory organs supplied by the great lateralis nerve (Fig. 220); (ii.) a supra-orbital canal passing forwards over the eye and innervated by the superficial ophthalmic branch of the facial nerve; (iii.) an infra-orbital canal supplied by the buccalis and otic branches of the same nerve; and (iv.) a hyo-mandibular or operculo-mandibular canal, situated on the outer side of the hyoid region, and thence prolonged downward and forward in relation with the lower jaw, and innervated by the external mandibular branches of the facial nerve. The hyo-mandibular canal is sometimes distinct from the other canals, as in Elasmobranchs and some Teleosts (Fig. 220); and in certain North American Siluroids the same may be said of the supra-orbital. But, as a rule, the infra-orbital is continuous behind both with the lateral and the supra-orbital canals, while the hyo-mandibular canal joins the infra-orbital, or, exceptionally, the supra-orbital canal. Transverse commissural canals often connect the lateral and supra-orbital canals of opposite sides across the dorsal surface of the head, and the corresponding infra-orbital and hyo-mandibular canals may also be continuous at the extremity of the snout or at the mandibular symphysis.

fig220

Fig. 220.—Sensory canals of the left side of the head of Gadus virens. e, Eye; i.o, infra-orbital canal (dotted); l.c, lateral canal (oblique shading); n, nasal apertures; op, operculum; op.m, operculo-mandibular canal (longitudinal shading); p.o, pit-organs; s.o, supra-orbital canal (cross-hatched); s.o.c, supra-orbital commissure; s.t, supra-temporal branch; t.t, tubuli by which the canals communicate with the exterior. (From Cole.)

Throughout their extent the canals communicate with the exterior by pores, or short canals terminating in pores, or by branched canals ending in groups of pores. In Cyclostomes[454] the lateral line system is represented by pit-organs disposed as in Fishes, and innervated by a true lateralis nerve. Some Elasmobranchs have the lateral canal of the trunk represented by an open groove protected by marginal denticles. Chimaera is more primitive still in this respect, for on the head as well as on the body the sensory organs are in open grooves. Amongst Fishes these organs are most primitive in the Dipnoi, where they retain their superficial position in the epidermis. In Teleostomes the lateral canals perforate the scales of the lateral line, and at intervals they open externally by simple or multiple pores which perforate the scales. On the head they are more or less completely enclosed in special ossicles which either remain distinct or fuse with certain of the adjacent dermal or cartilage bones of the skull. The use of the lateral line organs is not certainly known. They occur only in Fishes and Amphibia, and as blind Fishes are able to avoid obstacles with the greatest ease when swimming, it is possible that these organs enable their possessors to appreciate undulatory movements in water in the shape of reflex waves from contiguous surfaces or objects.[455] Their great antiquity is shown by their existence in most of the Heterostraci, and in the Antiarchi and Arthrodira, although they have not yet been discovered in the Osteostraci.

The Auditory Organs.—In its more typical condition each auditory organ consists of a membranous sac or vestibule, partially constricted into an upper portion or utriculus and a lower or sacculus (Fig. 221, A). Three semicircular canals are connected with the utriculus, of which two are vertical and at right angles to one another, and the third is horizontal. One end of each canal is dilated into an ampulla. A slender tube, the ductus endolymphaticus, leaves the sacculus, and ends in a sac-like swelling, the sinus endolymphaticus, which apparently represents a portion of the embryonic epidermic involution from which the auditory organ is formed. A smaller sac-like outgrowth from the sacculus, the lagena, corresponds to the cochlea of the higher Vertebrates. The epidermic lining of this system of cavities is differentiated into patches or ridges of sense-cells (maculae or cristae), separated by supporting cells and innervated by the terminal branches of the auditory nerve. There is a crista acustica in each ampulla; and maculae acusticae are present in the utriculus, sacculus, and lagena. A fluid, the endolymph, fills all the cavities, and a similar fluid or perilymph occupies the spaces in the periotic capsule in which the various chambers are lodged. Among the more notable deviations from this type of auditory organ the Cyclostome Myxine, apparently, has but a single semicircular canal with an ampulla at each end, and the vestibule is a simple sac (Fig. 221, B). Petromyzon has two canals, but lacks the horizontal canal. In Elasmobranchs, including Chimaera (C), the ductus endolymphaticus retains its primitive connexion with the exterior by means of a pore on the dorsal surface of the head. In the Dipnoi (e.g. Protopterus) the paired endolymphatic sinuses divide into a number of caecal branches containing otoliths, which meet and interlace over the fourth ventricle (Fig. 217).[456] Otoliths, either in the form of fine, mucus-connected, calcareous particles, as in Elasmobranchs, or as massive solid concretions in Teleosts, are present in relation with the sensory areas of the utriculus, sacculus, and lagena.

fig221

Fig. 221.—Auditory organs of Fishes. A, of a typical Fish; B, of Myxine; C, of Chimaera; and D, of Perca. a.c, Anterior canal; am', am", am'", ampullae; am.n, nerves to ampullae; c, semicircular canal (in Myxine); d.e, ductus endo-lymphaticus; h.c. horizontal canal; l, lagena; mc, macula acustica; m.s, macula acustica of the sacculus; n, nerves to ampullae; o, external aperture of the ductus endo-lymphaticus; p.c, posterior canal; s, sacculus; s.e, sinus endo-lymphaticus; sk, superficial skin; s.s, sinus superior; u, utriculus; viii, auditory nerve. (From Wiedersheim, after Retzius.)

In a few marine and in a large number of freshwater Teleosts the auditory organ enters into a more or less intimate connexion with the air-bladder by one of three different methods.

fig222

Fig. 222.—Cavity of the air-bladder of a Siluroid (Macrones nemurus) exposed by the removal of its ventral wall. a.c, Anterior chamber; b.o, basioccipital; b.w, body wall, here reduced to the external skin; cl, clavicle; l.c, lateral chamber; l.s, longitudinal septum; pt, post-temporal; tr.a, anterior portion of the tripus; tr.c, crescentic portion of the tripus; t.s, transverse septum; t.s', shorter transverse septum. (From Bridge and Haddon.)

The first and simplest is by the apposition of the extremities of a pair of caecal tubular prolongations from the air-bladder to the outer surfaces of the fibrous membranes which close a pair of vacuities in the outer bony walls of the periotic capsules, the inner surfaces being bathed by the perilymph surrounding the auditory organs. This method is characteristic of certain Serranidae, Berycidae, Sparidae, Gadidae, and Notopteridae,[457] and probably in the Hyodontidae. In the second method, of which several Clupeidae (e.g. Herring, Pilchard, etc.) furnish examples, the periotic vacuities are open instead of closed, and the sac-like ends of the tubular extensions from the air-bladder are in actual contact with protruding outgrowths from the utriculus.[458] The third method, by far the most elaborate, is by the intervention of a series of movably connected "Weberian" ossicles, of which the most posterior on each side (the tripus) is inserted into the dorsal wall of the air-bladder (Fig. 223), while the anterior one (scaphium) forms the outer wall of a median backward prolongation (sinus impar) of the perilymph-containing spaces surrounding the two auditory organs. This in turn encloses a similar median prolongation (sinus endolymphaticus) from the two sub-cerebrally united endolymphatic ducts (Fig. 223).[459] This complex mechanism is present in the Cyprinidae, Siluridae, Characinidae, and Gymnotidae; and hence the term "Ostariophysi"[460] as a collective name for these families.[461] The physiological raison d'être of the connexion between the air-bladder and the auditory organ cannot yet be regarded as satisfactorily determined. It is possible, as Weber thought, that it may be an auxiliary to the function of hearing by transmitting to the ear sound-waves impinging on the surface of the body and affecting the gases in the air-bladder.[462] On the other hand, it may be urged with perhaps greater probability that the connexion exists for the purpose of conveying to the ear stimuli due to the varying degrees of distension of the air-bladder, such as, it may be presumed, are naturally brought about by the variations of hydrostatic pressure which a Fish encounters in the course of its ascent or descent in the water.[463] Whether regarded as an accessory to hearing, or as a means of regulating the distension of the air-bladder, the physiological value of the connexion must be considerable, and on this point it is at least significant that the Weberian mechanism is characteristic of the dominant families of freshwater Teleosts at the present day.[464]

fig223

Fig. 223.—Diagram to show the Weberian ossicles and their relations to the ear and the air-bladder. at, Atrium, an extension of the sinus impar; a.v.c, anterior vertical canal; b.w, bony wall of the periotic capsule; d.e, the medianly-united endolymphatic ducts; h.c, horizontal canal; in, intercalarium, a third ossicle imbedded in the ligament (i.lg) connecting the scaphium with the tripus; n, bony nodules on the sides of the complex vertebral centrum; p.v.c, posterior vertical canal; s, sacculus; sc, scaphium; s.e, sinus endolymphaticus; s.i, sinus impar; tr.a, tr.c, the anterior and crescentic parts of the tripus; ut, utriculus. The radial lines represent the fibres of the dorsal wall of the air-bladder. (From Bridge and Haddon.)

The Olfactory Organs.—These organs are essentially a pair of pit-like inpushings of the skin of the ventral side of the head in front of the mouth, with their lining epidermis differentiated into sensory cells separated by supporting cells, and connected with the olfactory lobes of the brain by olfactory nerves.

fig224

Fig. 224.—Two stages in the development of the olfactory organ and the pituitary involution in Petromyzon. A is the earlier, B a much later stage. br, Brain; in, infundibulum; l.lp, lower lip; ms, mesenteron; n, notochord; ol.o, olfactory organ; pn, pineal body; pt.s, pituitary sac; st, stomodaeum; u.lp, upper lip. (From Parker and Haswell, after Dohrn.)

The Cyclostomata are unique amongst Craniates in the apparently unpaired condition of the olfactory organ, and in its remarkable relation to the pituitary involution. In the embryo Lamprey the median and ventral olfactory pit is carried inwards with the pituitary invagination, so that the former appears as a dorsal outgrowth from the latter, and the two have a common external opening, the naso-pituitary aperture (Fig. 224). Later the extraordinary forward growth of the upper lip to form the roof of the buccal funnel has the effect of shifting the naso-pituitary involution and its aperture to a final position on the dorsal side of the head. It is due to this dorsal displacement that, as we shall see, the pituitary caecum reaches the ventral surface of the brain by perforating the basis cranii from above, instead of from below as in all other Craniates. The pituitary body is pinched off from the dorsal side of the naso-pituitary involution. In the adult Lamprey the olfactory organ appears as a round sac divided by a median septum into two lateral chambers (Fig. 225), the lining epithelium of which is raised into prominent ridges. Behind the sac the pituitary involution is prolonged backwards beneath the brain, and, after traversing the basi-cranial fontanelle, it widens out into a spacious cul-de-sac and terminates on the dorsal side of the pharynx, beneath the anterior end of the notochord. In Myxine the pituitary involution ends by opening into the pharynx.