Many Fishes can jump out of the water, either in pursuit of insect food, like the Trout, or to enable them to escape the pursuit of their foes, like the Flying-Fish (Exocoetus), by means of a single forcible stroke of the tail, when the Fish is in a nearly vertical position close to the surface of the water. It is thus that the Salmon executes its remarkable leaps over weirs or up salmon-ladders when ascending rivers for spawning.

The tail is also used for steering. If kept bent to one side when the Fish is moving the tail acts like a rudder, and the course of the Fish is deflected to that side; or the direction may be altered by single strokes of the tail to the right or left, according to the course which the Fish desires to pursue.

In the majority of Fishes the paired fins are probably of little use for propulsion, and their action in this as in other functions is not always clear. In the Sharks and Dog-Fishes as well as in some Teleosts their planes are nearly horizontal when the fins are extended from the body; in others they are more oblique, so that the surfaces of the fins look upwards and backwards, and downwards and forwards; and in others again their surfaces are so nearly vertical that their strokes will be backwards and forwards. The pectoral fins also vary in their position on the sides of the body, being much more dorsal in some Fishes than in others. The paired fins may act as lateral keels in steadying the course of the Fish especially when the fins are extended and their planes are horizontal. They certainly seem to act as balancers in keeping the Fish on an even keel, and in counteracting the tendency of the Fish to turn belly upwards—a result which is attained by a slight upward and downward movement of the fins, and particularly of the pectoral fins. A Fish deprived of its pectoral members sinks downwards at the head and assumes an oblique position in the water. Removal of both the pectoral and pelvic fins of one side causes the Fish to roll over to that side; and if the fins are removed from both sides the animal turns belly upwards like a dead Fish. The pectoral fins may also be used for steering: a backward stroke of one fin while the other is kept folded back against the body will wheel the Fish round to the opposite side. From the ventral position of its mouth a Shark is forced to turn over to one side in order to seize its prey, and this movement of rotation is probably produced by the down strokes of the pectoral fin of one side. In some Fishes it would seem that the pectoral fins may assist locomotion by acting as paddles. The 15-spined Stickleback (Gastrosteus spinosus) frequently progresses by their aid alone; and, as their action can be reversed at pleasure, it is not unusual to see this Fish move backwards. The fins appear to be rotated or twisted in spiral movements like the tail when used for swimming, or like the wings of Insects in flying.

It has been mentioned that the function of the median fins (dorsal and anal) is to give stability to the Fish by acting as dorsal and ventral keels. This is certainly the case in the generality of Fishes. Nevertheless, there are exceptional instances in which one, or even both, of these fins are important swimming organs, acting either as a substitute for a tail which has become adapted for other uses, or as supplementary to that organ. Thus, in some of the Syngnathidae (Pipe-Fishes and Sea-Horses) the small size or absence of the caudal fin, and its use as a prehensile organ, renders the tail of little or no value as a propelling organ: hence it is that these Fishes swim by a lateral undulating movement of the dorsal fin. To enable them to do this the supporting skeleton presents certain interesting modifications. In the majority of Teleosts the arrangement of the fin-muscles, and the nature of the articulation between the dermal fin-rays and their basal radial supports, which is generally some form of a hinge-joint, are such as to limit the motion of the rays to simple elevation or depression in the vertical plane, and no lateral motion of the fin is possible. But in the Syngnathidae, as in the Pipe-Fish (Siphonostoma typhle), there is an exceptionally mobile articulation between the dermal fin-rays and the distal radial nodules which their cleft bases embrace and the bony proximal or basal radials, so that the fin can be flexed or bent to the right or to the left. In addition to this, by a change in the insertion of their tendons, the muscles corresponding to the ordinary elevator and depressor muscles of the fin-rays in other Fishes are capable of producing extensive lateral movements of the fin, or, by contracting in orderly sequence, of bringing about the characteristic undulating motion of the fin. A similar mechanism exists in many Plectognathi (e.g. species of Balistes, Monacanthus, Diodon, Tetrodon and Orthagoriscus)^[416] in connexion with both the dorsal and anal fins, but in these Fishes the action of the median fins in swimming must be regarded as supplementary to that of the tail.

Swimming is by no means the only form of locomotion in vogue amongst Fishes. A few, like the Angler-Fishes (Lophius), habitually use the pectoral fins for crawling about the sea-bottom. The East Indian Goby, Periophthalmus, uses its pectoral fins, which are bent at an angle like an elbow-joint, for hopping over sandy flats left bare by the retreating tide. The Flying-Fish (Exocoetus), when projected from the water by a stroke of its powerful tail, expands its large pectoral fins, and, using them after the fashion of a parachute, floats through the air for considerable distances before returning to its natural medium. The "Flying Gurnards" (Dactylopterus) are also capable of short aerial excursions in a similar fashion. Nor is tree-climbing beyond the province of a Fish, if credit be given to the assertion that the Indian "Climbing-Perch" (Anabas scandens) uses its opercular spines for ascending trees. Many freshwater Fishes are known to migrate across land from one pool or river to another, usually during the night. Eels do so by a serpentine or wriggling motion of their long bodies, but in others the pectoral fins seem to be the principal organs used for the purpose, aided, it may be, by a perverted use of the tail.

Sound-producing Organs.—Contrary to popular belief sound-producing or vocal organs are by no means uncommon in Fishes, especially in certain families of Teleosts. It is not always easy, however, to discriminate between involuntary, abnormal, or accidental sounds, and those due to the action of special vocal organs. There are, moreover, some Fishes which observations have shown to utter highly characteristic sounds, although the precise nature of the sound-producing mechanism is at present unknown; while other Fishes appear to possess organs which, on anatomical grounds, are perhaps vocal in function, although nothing is known of the nature of the sounds they emit. Here those organs only will be considered which, either with certainty or with some degree of probability, may be regarded as vocal structures. For most of our knowledge of these interesting structures we are indebted to the researches of Sörensen and Dufossé.^[417]

(a) Stridulation.—Stridulation as a method of sound-production has been recorded in many Teleosts, and one of the most interesting examples occurs in the singular Indian Siluroid, (Callomystax gagata).^[418] In this Fish (Fig. 205) the first five vertebrae are rigidly connected with one another and with the skull, mainly through the union of the neural spines of the third, fourth, and fifth vertebrae, and their articulation with the supra-occipital bone. The united spines together form a high, laterally-compressed lamina of bone, the hinder portion of which is vertically cleft into two thin plates separated by an interval sufficiently wide to receive the first interspinous bone of the dorsal fin. The inner surface of each of the two plates is traversed by a series of about thirty parallel, close-set, vertical ridges, while the first interspinous bone is similarly ridged on both its faces like a double file. Lastly, it may be mentioned that owing to the width of the intervertebral ligament between them the fifth and sixth vertebral centra are articulated by a joint of unusual mobility. The action of the mechanism is simple. By the vertical movements of the sixth and succeeding trunk vertebrae, with the interspinous bones which they support, on the rigid structure formed by the head and first five vertebrae, the file-like first interspinous bone moves backwards and forwards, and, by scraping against the ridges on the inner surfaces of the cleft neural spines, gives rise to a harsh grating noise, which is particularly unpleasant when artificially produced. The lateral movements of the trunk in ordinary locomotion do not affect the mechanism: it is only when the trunk is alternately flexed and extended in the vertical plane that the mechanism comes into play and a noise is produced. In the Bull-head (Cottus scorpius) the preoperculum is modified for stridulation, and in Dactylopterus the hyomandibular bone; in other Fishes, as in some Siluroids (e.g. species of Doras), stridulation takes place between a basal process from the great spine of the pectoral fin and the wall of a socket in the cleithrum into which the process is received, or between the small first spine of the dorsal fin and a roof-like process at the upper extremity of the first interspinous bone; also, in a somewhat similar fashion in the anterior dorsal fin of such widely different Fishes as certain Trigger-Fishes (Sclerodermi) pertaining to the genera Balistes, Monacanthus, and Triacanthus, Acanthurus chirurgus (Acanthuridae), the Boar-Fish (Capros aper), Centriscus scolopax (Centriscidae), and the Three-spined Stickleback (Gastrosteus aculeatus); and even between the spinose ray of the pelvic fin and the basipterygium in Triacanthus, Capros, and Gastrosteus.

In the "Drumming" Trigger-Fish (Balistes aculeatus),^[419] which frequents the coral-reefs off the Island of Mauritius, stridulation takes place between the postclavicles and a longitudinally grooved area on the inner surface of each cleithrum. Both the cleithra and postclavicles are in intimate relation with the air-bladder, and the sound produced by friction is apparently strengthened by the transference of the vibrations to the walls and gaseous contents of that organ. The passage of the sound-vibrations to the surrounding medium is facilitated by the fact that for a portion of their extent the lateral walls of the air-bladder are in contact with the superficial skin, which visibly shares in the vibratory movement of the bladder when the characteristic drumming sounds of Balistes are being emitted.

Stridulating sounds may also be produced by the friction of the upper and lower pharyngeal teeth, as in a species of Mackerel (Scomber brachyurus). By the grating of its teeth the Sun-Fish (Orthagoriscus mola) is said to emit sounds similar to those produced by the grinding of the teeth in Pigs and Ruminants; and Moseley^[420] has remarked of a species of Balistes that the "living Fish when held in the hand makes a curious metallic clicking noise by grating its teeth."

(b) Breathing sounds.—Characteristic breathing or murmuring sounds, or "bruits de souffle" as Dufossé terms them, are produced by a few Teleosts, among which may be mentioned the Eels, certain Cyprinidae, as, for example, the Carp (Cyprinus carpio), several species of Loaches (e.g. Misgurnus fossilis and Cobitis taenia), and the European Siluroid, Silurus glanis. According to Dufossé these sounds originate in some cases from the expulsion of gas from the air-bladder through the ductus pneumaticus and mouth, and in others, as in Misgurnus fossilis, they are produced by the rapid ejection through the anus of bubbles of air previously taken in at the mouth.

(c) Sounds produced through the agency of muscles connected with the air-bladder.—In addition to its usual function as a hydrostatic organ or "float" the air-bladder is often modified in various ways in different Teleosts, and adapted for use as a sound-producing organ.

In the South American Siluroid, Auchenipterus nodosus, the transverse processes of the fourth vertebra are bent downwards and backwards, and at the same time become converted into flexible and highly elastic springs (Fig. 206, B). Their distal extremities expand into oval bony plates which are imbedded in the anterior wall of the air-bladder, and often cause the latter to bulge inwards (Fig. 206, A). From the occipital region of the skull arise two powerful muscles which pass backwards to their insertion into the anterior faces of the two springs. By the contraction of these muscles the springs, and consequently also the front wall of the bladder, are drawn forwards; but directly the muscles relax, the elasticity of the springs causes them to move backwards to their former position, carrying with them the wall of the air-bladder. Hence it follows that the rapid alternating contraction and relaxation of the muscles will impart a vibratory movement to the anterior wall of the bladder and to the gaseous contents of that organ, with the result that a sound is produced. As a rule, those Fishes in which an elastic-spring-mechanism is present have the air-bladder subdivided by internal septa into a series of chambers freely communicating with one another; and no doubt the intensity of the sound is greatly increased by the vibratory movements of the gases across the free edges of the septa, and from one chamber to another. The elastic-spring type of vocal organ is apparently restricted to the Siluridae,^[421] and besides occurring in Auchenipterus is found also in the South American genera Doras, Oxydoras, Rhinodoras, and Euanemus; in the African genera Synodontis and Malopterurus; and in at least four species of the Indian genus Pangasius.^[422] There are also a few Teleosts in which the air-bladder is provided with special muscles, but, instead of being connected with elastic springs, the muscles extend from the skull, and are inserted directly into the wall of the bladder (Fig. 207); or, without being in any way attached to the skeleton, the muscles simply invest some portion of the surface of the air-bladder. In other Fishes the air-bladder, without possessing special muscles of its own, may, nevertheless, be partially invested by tendinous, or partly muscular and partly tendinous, extensions from the muscles of the body-wall (Fig. 208), or may be intimately related to certain muscles connected with the pectoral girdle.

Whatever the precise relation of the air-bladder to its muscles it is probable that the physiological effect is in most cases the same. By the rapid alternating contraction and relaxation of the muscles, some part of the wall of the bladder becomes alternately compressed and relaxed in such a way as to initiate a series of vibratory movements in the gases of that organ, and so produce definite sounds. In not a few of the Fishes the cavity of the bladder is subdivided by external constrictions or by internal septa, or is complicated by the development of lateral, tubular, caecal branches; and hence the vibratory movements of the gases will be greatly strengthened by their passage across the edges of the septa, or the apertures of the caeca, and the intensity of the resultant sounds also increased. It will be readily understood that the nature and quality of the sounds emitted by different Fishes will necessarily vary with the shape of the air-bladder, the number and arrangement of the internal septa and the caeca, and the strength and disposition of the contracting muscles. In a few Teleosts (Triglidae and Zeidae) sounds are said to be produced by the rapid vibration of an annular, or centrally-perforated, muscular diaphragm, which stretches across the cavity of the air-bladder.^[423] Nevertheless, it must be strongly emphasised that, while in some Fishes the air-bladder and its muscles undoubtedly constitute a vocal organ, there are many others in which the bladder can only be inferred to be sound-producing from its general agreement in anatomical structure with the same organ in Fishes where its vocal function has been clearly proved.

By one or other of these various methods the air-bladder is either known to be sound-producing, or is believed with good reason to be such, in the following Teleosts,^[424] and many others:—Certain species of the South-American genera of Siluridae, Pimelodus, Sorubim, Platystoma, Piratinga, Centromochlus, and Trachelyopterus; species of the South-American family Characinidae; Amblyopsis spelaea, the blind Fish from the Mammoth Cave of Kentucky (Amblyopsidae); among the Syngnathidae, the short-snouted Sea-Horse (Hippocampus brevirostris) of the British Coasts; certain Sclerodermi, such as the Trigger-Fishes, Batistes vetula, Triacanthus brevirostris, T. biaculeatus, and Monacanthus pardalis, and also some "Coffer Fishes" (e.g. species of Ostracion); some Gymnodontes (species of Diodon and Tetrodon); a few Serranidae (e.g. species of Therapon and Pristipoma); species of Holacanthus (Chaetodontidae) and in Holocentrum sogho (Berycidae); such Sciaenidae as the "Drum" (Pogonias chromis), the "Maigre" (Sciaena aquila), which has sometimes been taken in British waters, Umbrina cirrhosa, Otolithus regalis, and Micropogon undulatus, and, with more or less probability, many other species of the same family; one species of Zeidae, the John Dory (Zeus faber); Batrachus tau among the Batrachidae; several species of Gurnards (Triglidae) belonging to the genera Prionotus and Trigla; the so-called Flying Gurnard, Dactylopterus volitans (Dactylopteridae); the Indian species Ophiocephalus marulius and O. gachua (Ophiocephalidae); amongst the Gadidae, the Cod (Gadus morrhua) and the Haddock (G. aeglefinus); in such Zoarcidae as the blind Fish (Lucifuga subterranea) from the subterranean waters of the caves of Cuba, and also in some Ophidiidae (e.g. species of Ophidium).

In Fishes other than Teleosts, instances of normal sound-production by special vocal structures are rare. No recorded instances are known in the Cyclostomes or the Elasmobranchs,^[425] but there is evidence that sounds are emitted by Polypterus among the Crossopterygii, and by the Dipnoids Neoceratodus,^[426] Protopterus, and Lepidosiren, although it is not certainly known how they are produced, or that they may not be the accidental concomitants of the inspiratory or expiratory action of the lungs in breathing.

As to the nature of the sounds produced by the air-bladder and its muscles in different Teleosts, a few examples may be given.

The sound produced by the elastic-spring-apparatus of a recently caught Doras maculatus, has been described as a "deep growling tone," which may be distinctly heard at a distance of 100 feet when the Fish is out of the water. Under like conditions the air-bladder and its muscles, in a species of Platystoma, emit a similar sound. On the other hand, the sound produced by the elastic springs of the Electric Siluroid (Malopterurus electricus) has been compared to the hissing of a cat. The Sea-Horse (Hippocampus brevirostris) utters a monotonous sound analogous to that of a tambour, which is characteristic of both sexes, but is more intense and frequent in the breeding season. The "Coffer Fish" (Ostracion trigonus) emits a growling sound, as also does the "Globe Fish" (Tetrodon honckenii) when taken out of the water.^[427] The air-bladder and its muscles in the "Drum" (Pogonias chromis), constitute the most powerful sound-producing organ yet found in any Fish. The sounds emitted by the "Drum" are better expressed by the word drumming than by any other, and have frequently been heard by persons in vessels lying at anchor on the coasts of the United States, where these Fishes abound.^[428] The "Drum" begins its drumming noise in the spawning season in April, but is rarely heard afterwards. The "Maigre" (Sciaena aquila), whose musical performances are perhaps responsible for the Homeric fable of the song of the Sirens, is remarkable among Fishes for the variety of its sounds, which have been compared to bellowing, purring, buzzing, and whistling.^[429] The sound is often so intense that it may be heard when the Fish is at a depth of 18 metres, and the ear of the observer two metres above the water; and it has been recorded that by listening for these sounds, shoals of Maigres have been successfully netted. They rarely emit sounds when isolated; but in shoals, during the breeding season, they do not cease to make sounds with a vigour and a persistency which apparently must soon wear out their strength. One of the Indian Horse-Mackerels (Caranx hippos) grunts like a young Pig when captured, and the sound is repeated whenever it is moved, as long as vitality remains. A West Indian species of the same family (Argyriosus vomer) has been observed to produce a like sound, while an Egyptian Caranx (C. rhonchus) is known to the Arabs as the "Chakoura" or "Snorter."^[430] The sounds produced by the different British Gurnards, such as the Grey Gurnard (Trigla gurnardus), the Piper (T. lyra), the Elleck or Cuckoo Gurnard (T. cuculus), and the Tub-Fish (T. hirundo), have been compared to snoring, a sonorous and prolonged grunting, crooning (whence, perhaps, the term "crooner," by which the Grey Gurnard is known in Ireland), and croaking. The John Dory (Zeus faber)^[431] also utters sounds analogous to those of the Gurnards. Among the Dipnoi Lepidosiren is said to make a growling sound, and Neoceratodus a grunting noise which may be heard at night for some distance.

Whatever the nature of the vocal mechanism, it is highly probable that the sounds produced by Fishes travel to considerable distances in the water, inasmuch as the latter medium is a far better conductor of sound than air, and, moreover, the transmission of sound-vibrations from the air-bladder to the water is facilitated in many Fishes by the fact that, for a portion of its extent on each side the bladder is in direct contact with the superficial skin behind the pectoral girdle.

From the by no means exhaustive list of examples given above, it is obvious that in some form or other vocal organs are present in a considerable number of Fishes, both freshwater and marine, belonging to widely different groups; and further, that even in the same species (e.g. Doras maculatus and other Siluridae), both stridulation and the action of extrinsic muscles on the air-bladder may be utilised as a means of sound-production. Certain Teleostean families like the Siluridae, the Sciaenidae, and the Triglidae, seem to be distinguished above all others by the prevalence of some form of vocal organ. According to Sörensen, the first mentioned of the three families includes no less than 68 species, which utilise the air-bladder alone as a sound-producing organ. Nevertheless, there still remain many Teleostean families, rich in genera and species, and with an almost world-wide geographical distribution, in which such organs have not yet been found.

The advantages which Fishes derive from the possession of sound-producing organs are sufficiently obvious.

A characteristic feature in the reproduction of most Fishes is the general absence of any process of conjugation between the sexes, the eggs being fertilised in the water after their extrusion from the body of the female, and, consequently, any device which will facilitate the formation of shoals during the breeding season must be of great advantage to the species by largely increasing the chances that the ova will be fertilised, and thus secure the more successful propagation of the race. Hence it may be concluded that the vocal organs of Fishes are a means to this end, and that the sounds they produce are in fact recognition-sounds which enable Fishes of the same species to congregate together at periods when reproductive activity is greatest. This view is in harmony with much that is known of the habits of these Fishes, especially with the fact that particular sounds are often characteristic of particular species, and that the sounds are produced most frequently and with greater intensity during the breeding season than at any other time. While useful to all Fishes that possess them, vocal organs are, no doubt, specially serviceable to those Fishes which, from the nature of their habitat, can make but little use of their eyes; and this fact may perhaps explain the prevalence of such organs in the Siluridae, which are frequently bottom- or ground-feeding Fishes, and often live in muddy waters.

The sounds emitted by Fishes may also, in some instances at least, be warning sounds. Many of the sound-producing Fishes are provided with exceptionally strong spines either in connexion with the median and paired fins, as in many Siluridae, or on the general surface of the body, as in Diodon hystrix. Such spines are very effective weapons for offensive or defensive purposes, and are capable of inflicting very severe wounds. The natural enemies of these Fishes learn by experience or instinct to associate particular sounds with the possession of dangerous spines, and warned by the sounds, they refrain from attacking the owner of the spines, to the mutual advantage of both.

Electric Organs.—Electric organs capable of generating more or less powerful electric discharges are present in certain Fishes, both marine and freshwater. They occur in a few Elasmobranchs (species of Raia, Torpedo, and Hypnos), in such Teleosts as the African Silurid Malopterurus the "Electric Eel" (Gymnotus), and in species of Mormyridae (e.g. Mormyrus). With one exception electric organs are composed of metamorphosed muscular fibres, and their nerve-endings or motor end-plates. The species of Raia have two small electric organs, one on each side of the terminal portion of the tail.^[432] In Gymnotus^[433] the organs are much larger, and extend the whole length of the tail, which is fully four-fifths of the total length of the Fish. The Mormyridae also have their feeble electric organs in the caudal region. In all these Fishes the electric organs are modified portions of the caudal muscles. In the Torpedo, however, these organs are two large oval masses, one on each side of the head, between the gills and the cephalic prolongation of the pectoral fin (Fig. 209). Malopterurus^[434] is exceptional in possessing an electric organ derived from the epidermis and not from the muscular system. In this Fish the organ envelops nearly the whole body like a mantle, between the skin and the subjacent muscles of the trunk and tail. An electric organ is composed of an immense number of "electric plates" (modified motor end-plates), abundantly supplied with nerves on one of their surfaces, and disposed in a series of vertical (Torpedo) or longitudinal (Gymnotus) columns, separated by septa of connective tissue. In the active state of the organ in the Torpedo^[435] the ventral surfaces of the plates, on which the nerves are distributed, become negative to the dorsal, and "the effect in all the plates of a column when summed up is, therefore, such that the dorsal end of a column becomes positive to the ventral end."^[436] Hence the current in the form of a succession of shocks passes from the ventral to the dorsal surface of the head. In Gymnotus, where the columns are longitudinally arranged, it is the anterior and posterior surfaces which become oppositely electrified, and the current passes from the tail to the head. The shock imparted by an electric discharge is most powerful in Gymnotus,^[437] Malopterurus, and Torpedo, in the order named, and relatively weak in the remaining genera. The strength of the shock increases with the number of electric plates included in the circuit. Thus in Gymnotus the maximum shock is given when the body of the Fish is so curved that the head and the tail are in contact with different points on the surface of some other Fish. The discharge may be reflex or voluntary. Repeated discharges induce fatigue and weaken the shocks. Electric organs are powerful offensive or defensive structures, enabling the Fish to repel the attacks of enemies, or to stun or kill their prey.

CHAPTER XIV

NERVOUS SYSTEM AND ORGANS OF SPECIAL SENSE

The nervous system consists of the brain and the spinal cord, and of the cranial and spinal nerves. The rudiment of the future brain and spinal cord first appears in the embryos of some Cyclostomes (e.g. Bdellostoma), of Elasmobranchs, and of Chondrostei (e.g. Acipenser), and of Neoceratodus among the Dipnoi, in the form of a tubular medullary canal pinched off from the epiblast of the dorsal surface of the body. By a somewhat different method, but with the same final result, a medullary canal is formed in other Cyclostomes (e.g. Petromyzon), in the Holostei and Teleostei, and in Lepidosiren,^[438] from a solid ingrowing keel of epiblast which subsequently becomes tubular. Later, the medullary canal in the head enlarges, and becomes divided by two transverse constrictions into three vesicles, the primary fore-, mid-, and hind-brain, leaving the rest of the canal to form the spinal cord.

The Spinal Cord.—This portion of the medullary canal retains a simpler and more uniform cylindrical structure. Its walls thicken and their component cells become converted into nerve cells and nerve fibres, but a remnant of the original cavity remains in the adult as a minute axial canal, with a ciliated epithelial lining, the central canal of the spinal cord or myelocoele. In most Fishes the spinal cord extends the whole length of the body, but in some Teleosts, especially in certain Plectognathi, it is remarkably short. In a Sun-Fish (Orthagoriscus), 2½ metres long, and weighing about a ton and a half, the cord was only 15 mm. in length, or shorter than the brain.

The Brain.—At an early stage in its embryonic history the brain consists of three simple vesicles, the fore-, the mid-, and the hind-brain, the first of which lies in front of the anterior end of the notochord and is therefore pre-chordal in position. As development proceeds the walls of the vesicles undergo local thickenings, or they give rise to hollow paired or median outgrowths, and by one or other of these methods the different parts of the complex adult brain are evolved, while the original cavities of the vesicles or of their outgrowths persist as a continuous system of epithelium-lined spaces or "ventricles."^[439] The fore-brain is remarkable for the number and importance of the parts to which it gives rise. First, it bulges out in front into a hollow vesicle, the prosencephalon, leaving the rest of the fore-brain as the thalamencephalon or diencephalon (Fig. 210). The cavity of the prosencephalon is the prosocoele, and a pair of thickenings in its floor form two basal ganglia or corpora striata. In many Fishes the prosencephalon retains this simple vesicular condition, in which case the roof or pallium is usually epithelial and non-nervous; but in others two hollow lobes grow out from it in front and give rise to two cerebral hemispheres or parencephala.^[440] Both contain extensions of the prosocoele, the paracoeles or lateral ventricles, from the floor of which the corpora striata now project. The prolongation of the pallium forming the roof of the lateral ventricles either remains partially epithelial, or it may acquire a wholly nervous structure and thicken to an extent which differs greatly in different Fishes. With the formation of the hemispheres the prosencephalon and its prosocoele become of secondary importance, and may cease to be recognisable as distinct from the thalamencephalon and its ventricle. The lateral ventricles then appear to communicate directly with the third ventricle by two apertures, the foramina of Munro. The forward growth of the brain is completed by the development of two hollow lobes, the olfactory lobes or rhinencephala, each of which contains a ventricle or rhinocoele communicating behind with the prosocoele, or, if hemispheres are present, with the corresponding lateral ventricle.

Scarcely less complicated, and perhaps even more interesting from a morphological standpoint, are the structures arising out of the thalamencephalon. By thickenings of its lateral walls two large ganglia, the optic thalami, are formed, and on the inner or dorsal aspect of each of these a ganglion habenulae is developed. From the sides of the thalamencephalon the primary optic vesicles are derived, which later become transformed into the retinal parts of the paired eyes and the optic nerves. Besides the optic vesicles there is a second pair of embryonic outgrowths which arise from the roof of the thalamencephalon. These outgrowths form stalked vesicles and represent a pair of degenerate visual organs. Usually they become so displaced that the left one lies in front of the right, and they appear as if median. The subsequent fate of the vesicles differs greatly in different Craniates. Both persist in the Lamprey, the right vesicle to some extent retaining its primitive visual function as a parietal eye and directly overlying the left or pineal vesicle. In Elasmobranchs the two unite to form a glandular organ, the so-called pineal body of the adult, and in Teleosts the left vesicle disappears, leaving the right as a pineal body.^[441] There is also an embryonic median outgrowth from the roof of the prosencephalon, the paraphysis, which soon disappears and whose significance is not known. A median hollow downgrowth from the floor of the thalamencephalon forms the infundibulum, which becomes attached to a caecal diverticulum from the roof of the mouth. With rare exceptions the diverticulum loses all connexion with the mouth, and, as the pituitary body or hypophysis, it appears as an appendage to the extremity of the infundibulum. In the Crossopterygii the connexion is retained even in the adult by means of a slender canal extending from the pituitary body and opening into the oral cavity. Laterally, the base of the infundibulum grows out into a pair of rounded lobes, the lobi inferiores, and distally into a thin-walled glandular sac, the saccus vasculosus, which lies just behind the pituitary body. The cavity of the thalamencephalon persists as the third ventricle or diacoele. The parts of the brain developed from the mid-brain and the hind-brain are much less complicated, and, except for variations in size, they present a fairly uniform character in most Fishes.

In the mid-brain the roof bulges out into a pair of optic lobes, and by the growth of lateral thickenings in its floor two thick strands of longitudinally disposed nerve fibres, the crura cerebri, are formed. The cavity of the mid-brain remains as the mesocoele, and from it an extension may be prolonged into each optic lobe.

From the hind-brain are formed the cerebellum or epencephalon and the medulla oblongata or metencephalon, the former as a dorsal bulging, the latter as a ventral thickening. Except where the cerebellum is developed the dorsal wall remains epithelial, and forms the roof of the persistent cavity of the hind-brain, the fourth ventricle or metacoele, which retains its primitive continuity with the central canal of the spinal cord. Lateral lobe-like outgrowths from the dorsal columns of the medulla are conspicuous structures in some Fishes, and are known as corpora restiformia. The paired portions of the brain are connected across the middle line by a series of transverse commissures. The more important modifications of the brain in Cyclostomes and Fishes will now be briefly dealt with.