Chapter VII: , is termed the endoskeleton. The latter consists (i.) of an (7)

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, {386}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.

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 {387}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, {388}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.

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.

{389}

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] {390}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]

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.

{391}

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.

{392}

The apparently monorhinal condition of the Cyclostomes is probably a secondary acquisition. At the earliest embryonic stage at which any trace of an olfactory organ is apparent, there is a median thickening of the epidermis, possibly a vestige of some older sensory organ comparable, it may be, to the so-called olfactory organ of Amphioxus; on each side of it there is a lateral thickening, the rudiments of the paired organs.[465] The three thickenings, or "plakodes," then sink inwards to form an olfactory pit. The partial subdivision of the adult organ by a vertical septum, and the presence of two olfactory nerves, point to the same conclusion.[466] All Fishes possess olfactory organs which are obviously paired. In Elasmobranchs and Dipnoi they retain their primitive ventral position. Many Sharks and Dog-Fishes possess an oro-nasal groove leading from each olfactory organ to the corresponding angle of the mouth. The Dipnoi proceed a stage farther, and, by the conversion of the grooves into short canals, the olfactory pits communicate with the mouth by true internal nostrils, as in the higher {393}Vertebrates. In the adults of existing Teleostomi the orifice of each organ is usually divided into two by the growth of a fold of skin across it, and the two apertures rotate outwards and upwards on to the lateral or the upper surface of the snout. Of the two nostrils the posterior one probably corresponds to an external nostril, and the anterior one to the internal nostril. Occasionally each olfactory organ has only a single orifice. In the Crossopterygii and in some Teleostei the nostrils become tubular. The lining epithelium of the olfactory pits is usually produced into ridges, disposed longitudinally or transversely, or in the form of radii from a central point in the roof. Many Teleosts have each olfactory organ prolonged backwards into one or two sacs, the nasal sacs, which are either simple reservoirs, or glandular and mucus-secreting. In a species of Chinese Sole (_Cynoglossus semilaevis_) the two sacs, one from each olfactory organ, unite over the roof of the mouth in a common median sac, and in one unique specimen the latter communicated with the mouth by a large naso-pharyngeal aperture.[467]

THE EYES.—In essential structure the eyes of Cyclostomes and Fishes resemble those of the higher Craniates. As a rule, in Fishes they are relatively larger, however, and the lens is globular and the cornea somewhat flatter. Ciliary processes and ciliary muscles are absent. As the eyes are nearly always lateral in position it is probable that monocular vision is the rule. In Teleosts and in _Amia_ a "choroid gland," consisting of a mass of capillary blood-vessels, surrounds the optic nerve externally to the retina, and derives its blood from the efferent artery of the pseudobranch (Fig. 226). In most Teleostomi, but not in Cyclostomes, Elasmobranchs, and Dipnoi, there is a singular prolongation of the choroid coat, known as the "processus falciformis," which extends across the vitreous humour to the inner face of the lens, where it ends in an expansion, the "campanula Halleri" (Fig. 226). Accommodation to vision at different distances is not effected by alterations in the convexity of the lens, but by a change in its position with regard to the retina, apparently brought about by the contraction of a special retractor muscle.[468] Some oceanic pelagic Teleosts are remarkable for their curious telescopic eyes in the shape of short protruding {394}cylinders, each terminating in a strongly convex cornea (Fig. 227).[469] The eyes are directed either upwards or forwards, and, as their long axes are parallel in either position, it is probable that these Fishes are capable of binocular vision. In the young of certain Teleosts occurring in the Antarctic and Indian Oceans the large eyes are situated at the extremities of extraordinary long stalks extending from the sides of the head.

In the quasi-parasitic Cyclostome, _Myxine_, and in many Teleosts belonging to widely different families, which live at great depths in the sea or inhabit subterranean waters, the eyes suffer from disuse and degenerate in structure. The influence of a deep-sea habitat on the eyes of Fishes is somewhat varied. The eyes are often small. A few abyssal Fishes are totally blind, and no external trace of eyes can be seen (Fig. 430). In such Fishes compensation is often afforded by an extraordinary development of tactile organs in the form of long barbels, or of trailing filaments derived from the median or the paired fins (Fig. 371, B). Many deep-sea forms possess eyes of the normal size, or even exceptionally large eyes, probably because either they occasionally migrate towards the surface, or else they possess phosphorescent organs and are able to see by the aid of the light they themselves emit. A blind Siluroid (_Amiurus nigrilabris_) frequents the cave streams of Pennsylvania, and many members of the same family which live in muddy waters have very small or even minute eyes. One of the Gobies (_Typhlogobius_),[470] which buries itself in the sand, or is found under stones in the holes of a burrowing Crab on the coast of California, is also {395}blind. Amongst other blind Fishes _Amblyopsis_ and _Typhlichthys_ (Amblyopsidae)[471] and _Lucifuga_ (Zoarcidae) may be mentioned, the first two inhabiting the cave streams of North America, while the third has a similar habitat in Cuba. When the eyes degenerate they dwindle in size and recede from the surface. The lens and the iris wholly or partially disappear, and although it is generally recognisable the retina loses certain of its characteristic layers, or the latter are but imperfectly formed. In _Myxine_ even the eye-muscles are absent.

The eyelids of Fishes are little more than marginal folds of skin, capable of little if any movement, and leave the eyes largely uncovered. Some Sharks have a third eyelid or "membrana nictitans" at the anterior corner of the eye. Lachrymal glands are unknown.

THE PARIETAL EYE.—It is only in the Cyclostomes that this structure can have any claim to be regarded as a visual organ. In the Lamprey (Fig. 228) the parietal eye is a slightly flattened vesicle lying directly over the pineal vesicle, and connected by a slender stalk or nerve with the right ganglion habenulae. The dorsal or more external half of the vesicle is bi-convex, and forms the "pellucida," while the inner half or retina is said to consist of supporting cells with interspersed deeply pigmented sense-cells and ganglion cells.[472] The external skin over the parietal eye is partially transparent in the living animal.

In many of the oldest known Fishes, such as the Ostracodermi, the Antiarchi, and the Crossopterygian Osteolepida, there are indications of the existence either of one or of two median sense-organs on the upper surface of the skull, in the shape of one or two foramina, or hollow protuberances, or pit-like grooves or {396}depressions, but, as a rule, when one of them is present the other is absent. It is probable that both these structures were associated with sensory organs, of which one may have been a parietal eye and the other a pineal eye. Some Teleosts (_e.g._ many deep-sea Scopelidae) have a transparent, convex, cornea-like prominence on the upper surface of the head which may be related to one of these singular organs.[473]

{397}CHAPTER XV

THE KIDNEYS AND THE REPRODUCTIVE ORGANS—BREEDING

The kidneys and the reproductive organs are so intimately connected that it is necessary to deal with them together. Both organs are specialised portions of the coelom and its epithelial lining. The KIDNEYS are essentially a series of tubular and at first segmentally-disposed outgrowths from the coelom (urocoeles) which acquire a connexion with the exterior, while the gonads have their origin from local modifications of the coelomic epithelium. At a very early embryonic stage each lateral half of the coelom presents three well-marked divisions: (1) a series of dorsal portions ("myocoeles"), the cavities of the myotomes or muscle-segments; (2) a longitudinally continuous unsegmented portion extending round the alimentary canal, the "ventral coelom"; and (3) a series of intermediate tubular portions or "nephrotomes," each of which leads from a myocoele to the ventral coelom (Fig. 229, A). The essential components of the kidneys, the urocoeles or renal tubules, are derived from the nephrotomes. In its typical condition each kidney consists of three portions, which, in accordance with their embryological and evolutionary sequence, are termed the "pronephros," the "mesonephros," and the "metanephros." The pronephros, the larval or provisional kidney, is formed from a limited number of the nephrotomes immediately behind the head. From each nephrotome a hollow tubular outgrowth is formed, which grows towards the lateral surface of the body, and then unites with its fellows of the same side to form a main longitudinal duct—the "archinephric" or "pronephric duct" (Fig. 229, A, Fig. 230, A). This duct grows backwards until it opens into the cloaca.[474]

{398}

At the same time the nephrotomes lose their connexion with the myocoeles, although they still retain their "nephrostomes" or apertures through which they communicate with the ventral coelom. When fully developed the pronephros consists of a few tubules, more or less convoluted, opening at their inner extremities into the coelom by means of their ciliated nephrostomes, and at their outer ends communicating with the exterior through the archinephric duct. In relation with the pronephros a branch from the dorsal aorta forms a tuft of capillary blood-vessels or "glomus," opposite the nephrostomes, which projects into the ventral coelom on each side. Later, a second series of much more numerous tubules is formed behind the pronephros, which constitute the mesonephros. In forming mesonephric tubules the nephrotomes become disconnected from the myotomes and their myocoeles, and curving outwards they {399}come to open into the archinephric duct, although they do not in any way contribute to its formation (Fig. 229, B). Segmentally-arranged twigs from the dorsal aorta end in tufts of capillaries or glomeruli, each of which projects into a small sac-like enlargement of a mesonephric tubule, pushing before it the wall of the sac. In this way a double-walled "Malpighian body," containing a "glomerulus," is formed in connexion with each tubule. Subsequently, the mesonephric tubules increase in number by budding. New nephrostomes and Malpighian bodies are developed on the secondary branches, and the original segmental arrangement of the tubules becomes obscured. With the growth of new tubules, and the formation of blood-vessels and of connective and lymphoid tissues between them, each mesonephros finally assumes the condition of a compact gland imbedded in the dorsal wall of the coelom, with its ventral surface invested by the peritoneum. A "metanephros," which in the higher Vertebrates replaces the mesonephros as the functional kidney, is perhaps not represented in Fishes.

A more or less well-developed pronephros is present in the embryos or larvae of the Cyclostomes and of all Fishes, but as a rule it completely disappears at an early period and is replaced by the mesonephros. It is retained throughout life, however, in the Myxinoid Cyclostomes (Fig. 230, B), and has its persistent nephrostomes opening into the pericardial cavity.[475] In a few Teleosts the pronephros is also persistent, as in _Fierasfer_ and _Dactylopterus_, and in others the organ may not completely disappear until the approach of sexual maturity. But with these exceptions the mesonephros is the sole functional kidney in the adults of the Cyclostomes and of all Fishes. As regards the nature of the duct by which the excretion of the mesonephros is conveyed outwards, there are notable differences in different Craniates. The Cyclostomes and the Teleostomi retain that part of the archinephric duct into which the mesonephric tubules open, and which remains after the atrophy of the pronephros (Fig. 230, B, E, F). In Elasmobranchs, and probably also in the Dipnoi, a special mesonephric duct is developed in a way which will be described later (Fig. 230, C, D).

{400}

{401}In the males of Elasmobranchs some of the hinder mesonephric tubules unite to form a single main duct opening into the terminal part of the mesonephric duct, and these tubules and their separate duct are sometimes regarded as a metanephros and a metanephric duct. The mesonephric nephrostomes are persistent throughout life in a few Elasmobranchs (_e.g._ Notidanidae, Heterodontidae, Rhinidae, and some Scylliidae), and also in _Amia_:[476] in all other Fishes as well as in the Cyclostomes they become closed in early life.

In many Fishes the hinder extremity of the coelom communicates directly with the exterior through "abdominal pores," of which there is usually a pair, rarely a single pore, situated close to the cloacal or the anal aperture.[477] Elasmobranchs usually have a pair, often at the extremities of a pair of cloacal papillae (Fig. 231), but they are absent in some families (_e.g._ Heterodontidae and Rhinidae); and in some Scylliidae (e.g. _Scyllium canicula_) they are very variable, being either present or absent on both sides, or an open pore is present on one side only. Pores are present and paired in the Crossopterygii, the Chondrostei, and the Holostei. Amongst the Dipnoi _Neoceratodus_ has a pair of pores. _Protopterus_ sometimes has two pores opening into the cloaca, but as a rule the two become confluent and have a single external aperture. In _Lepidosiren_ pores are wanting. Abdominal pores are rarely present in Teleostei. They exist, however, in the Mormyridae (_Gymnarchus_ and several species of _Mormyrus_), and also in the {402}Salmonidae,[478] where they are as singularly variable in different species and individuals as in the Elasmobranch Scylliidae. The use of abdominal pores is not certainly known, unless the coelom of those Fishes which possess them continues to retain some measure of its primitive excretory function, and the pores act as excretory ducts. That the nephrostomes are excretory organs has been shown by experiment, and it is worthy of note that there exists a reciprocal relation between these structures and abdominal pores, to the extent that while there are a few Fishes (_e.g._ certain Elasmobranchs and _Amia_) in which both coexist, there are many others in which the presence of nephrostomes is correlated with the absence of pores and _vice versâ_.

The male and female GONADS, testes and ovaries, are derived from the coelomic epithelium near the inner or median aspect of the nephrotomes (Fig. 229, B). Here the epithelium remains columnar, and soon projects into the ventral coelom as a continuous longitudinal ridge. It is probable that at first the modified epithelium is segmented as a series of "gonotomes," but if so, the latter must soon coalesce into a continuous ridge. Some of the epithelial cells enlarge to form the primitive sex-cells. In the development of an ovary, portions of the epithelium sink inwards, carrying with them the primitive ova. Certain of the cells form the epithelial walls of a number of ovisacs, each of which encloses an ovum. As the ovisacs increase in number and size the germinal ridges project more and more into the coelom until, as ripe ovaries, they become suspended from its dorsal wall by a double peritoneal fold, the "mesovarium" (Fig. 156). The testes develop in a similar fashion except that the primitive sex-cells, which later give rise to spermatozoa, form the lining of a number of simple or ampulla-like tubules, the seminiferous tubules, and the suspensory fold is termed the "mesorchium."

The Cyclostomes have gonads in the shape of unpaired organs extending nearly the whole length of the coelom, but in all Fishes the organs are primarily paired, although by fusion, or by the absorption of one gonad, the ovaries or the testes sometimes appear as if single. The ovaries may either be naked, as in Elasmobranchs, Dipnoi, Crossopterygii, and Chondrostei, and in _Amia_ amongst the Holostei; or, as in _Lepidosteus_ and most Teleosts, they become enclosed in coelomic sacs. The {403}former, or "gymnoarian," condition is primitive; the latter, or "cystoarian," is secondary, and is brought about by the growth of two peritoneal folds round the ovary and the union of their margins. Into these coelomic sacs the egg-bearing or real ovarian tissue projects either in the form of processes or of transversely- or longitudinally-arranged plates or folds (Fig. 232, B). The testes are composed of seminal ampullae, as in Elasmobranchs, or of radially-arranged and sometimes plexiform tubules opening into the gonoduct, as in nearly all other Fishes (Fig. 232, A).

In the Cyclostomes (e.g. _Petromyzon_) the eggs and spermatozoa are discharged from the gonads into the coelom, whence they reach the exterior through a pair of "genital pores" leading from the hinder end of the coelom into a urinogenital sinus formed by the united extremities of the two archinephric ducts.[479] _Myxine_ has, however, but a single median pore, opening into an integumentary cloaca, which also receives the rectal and urinary orifices. _Bdellostoma_ has two such pores communicating with a similar cloaca.[480]

{404}The nature and homologies of the genital ducts in the different groups of Fishes are amongst the most puzzling of the many problems which vex the soul of the Vertebrate morphologist, and although there is a fairly general agreement on some points, there are others of great importance of which it may be said _quot homines, tot sententiae_.

Broadly speaking, there are two types of genital ducts in Fishes: (1) those which are obviously derived from some part of the kidney system; and (2) those which are special ducts and appear to have no connexion with kidney-ducts.

The Elasmobranchs offer a typical example of gonoducts of the first kind. At an early embryonic period in both sexes each archinephric duct becomes longitudinally split into two ducts, of which one continues to receive the openings of the mesonephric tubules and remains as a mesonephric duct (Fig. 229, B).[481] The other, which has no connexion with the mesonephros, opens anteriorly into the coelom by means of the united nephrostomes of the pronephros, and is known as the "Müllerian duct" (Fig. 230, C and D). In the adult male the Müllerian ducts are useless vestiges, but in the female they persist and act as oviducts, receiving the eggs set free from the ovarian ovisacs through their coelomic apertures, and thence conveying them to the cloaca. In the male, certain of the anterior mesonephric tubules become connected with the testicular ampullae by means of a network of slender tubules, the "vasa efferentia" or testicular network, and through the latter the spermatozoa pass from the testes to the mesonephric duct (Fig. 230, C). Consequently, the mesonephric duct conveys both spermatozoa and the kidney excretion to the cloaca. It is obvious, therefore, that both the male and female gonoducts are derived from kidney-ducts.

The Teleostei afford an equally typical illustration of the second type. Each female gonoduct (oviduct) is formed by a backward growth of the same two peritoneal folds which enclose the ovary; these are converted into a "peritoneal tube" or canal by the union of their margins. The male gonoducts are also formed in continuity with the testes, that is, as backward prolongations from the latter. Each duct, male or {405}female, seems to be a duct _sui generis_ and to have no connexion whatever with the kidney system (Fig. 230, E). In the Salmonidae, Anguillidae, Galaxiidae, Hyodontidae, Notopteridae, and Osteoglossidae, and also in _Misgurnus_, the oviducts lose their continuity with ovaries and degenerate to an extent which differs greatly in different families. Thus in some Salmonidae, as in the Smelt (_Osmerus eperlanus_),[482] the oviducts end anteriorly in wide funnel-like coelomic apertures after the fashion of Müllerian ducts, and do not embrace the ovaries: hence the ovaries are naked and not cystoarian, and their ducts are not peritoneal tubes but "peritoneal funnels" (Fig. 230, F). In other Salmonidae and in the Anguillidae the oviducts appear to have so far degenerated that they are represented either by a pair of very short funnels or by a pair of genital pores, which, as in the Salmon, have a common external aperture behind the anus and in front of the single orifice of the united archinephric ducts (Fig. 233, A). In all such instances the eggs are set free from the ovaries into the coelom, from whence they escape through the peritoneal funnels or genital pores. In the Eels the male gonoducts also degenerate, and, losing all connexion with the testes, they become reduced to genital pores as in the female.

The Holocephali and probably the Dipnoi conform to the Elasmobranch type in the nature of their male and female gonoducts. In the Crossopterygii[483] each testis has its own proper duct, which has no connexion with the kidney system and apparently belongs to the Teleostean type, while the oviduct, which is almost certainly not a Müllerian duct, is probably a peritoneal funnel. On the other hand, the Chondrostei and the Holostei are in the interesting transitional condition of possessing male ducts of the Elasmobranch type and female ducts of the Teleostean type, the latter being either ducts directly continuous with the ovaries, as in _Lepidosteus_, or of the nature of peritoneal funnels, as in _Acipenser_, _Polyodon_, and _Amia_ (Fig. 230, E and F).

How far the distinction between the two types of gonoduct holds good in the case of the male is not quite clear, and it has recently been argued that the Dipnoi offer a connecting link between the two.[484]

{406}

In _Protopterus_ each testis is divided into an anterior sperm-producing part and a posterior tubular portion which has lost the capacity of producing sex-cells. The testicular network is greatly reduced, and forms but a limited connexion between the tubular portion of the testes and the mesonephric duct (Fig. 233, B). If it be supposed that the testicular network became still further reduced so that the connexion between the testes and the kidney-duct took place directly through a single channel instead of through several, the result would be a gonoduct essentially similar to the male duct of an ordinary Teleost. Should this view prove to be correct, it will follow that the male gonoducts of _all_ Fishes are differently-modified examples of the Elasmobranch type. But there will still remain the female gonoducts of Ganoids and Teleosts, which must be regarded as distinct from Müllerian ducts unless it can be shown that their different methods of development are not necessarily fatal to their homology with Müllerian ducts, or that both types of gonoduct can be derived from some intermediate type. Assuming that some Fishes do possess male or female ducts which have not been derived from the kidney system, but have been independently acquired, there is still the question, which of the two types is the more primitive, or, in other words, has the Elasmobranch type superseded the Teleostean, or _vice {407}versa?_ To this question no decisive answer can at present be given.

The terminal relations of the kidney-ducts and the gonoducts, and the presence of accessory or of vestigial organs in connexion with them, will now be briefly dealt with. In the males of the Elasmobranchs the mesonephric ducts which, as already pointed out, act both as kidney-ducts and gonoducts, dilate posteriorly to form a pair of vesiculae seminales, and then unite to form a urinogenital sinus, opening into the cloaca at the extremity of a median papilla (Fig. 230, C). The sinus also receives ducts from the hinder part of the mesonephros, either separately, as in the female, or by a common duct on each side—the so-called metanephric duct—as in the male. Two tubular caecal outgrowths from the sinus form two sperm sacs. Only the anterior portions of the Müllerian ducts with their coelomic apertures are retained in the adult. In the female the mesonephric ducts are purely excretory, but otherwise they are similar, and the oviducts (Müllerian ducts) open into the cloaca separately or by a common orifice (Fig. 230, D). A glandular dilatation of each oviduct forms the oviducal or shell gland by which the horny egg-cases are secreted. In the males of the Holocephali the gonoducts open into a urinogenital sinus with an external orifice distinct from and behind the anus; but the female has separate apertures for the rectum, the conjoined oviducts, and the united mesonephric ducts. Both sexes have complete Müllerian ducts communicating with the coelom in front, and behind with the exterior. The Dipnoi of both sexes essentially resemble the Elasmobranchs in the general relations of their ducts, but the Müllerian ducts of the male exhibit marked differences in the three genera.[485] In _Neoceratodus_ the ducts are as complete as their functional representatives in the female. _Protopterus_ retains anterior vestiges and the coelomic apertures, and also vestiges of the hinder portions which unite and end blindly in the urinogenital papilla, but the middle sections of the two ducts are suppressed (Fig. 233, B). In the Teleostomi there is a general similarity in the terminal relations of the gonoducts and kidney-ducts. In the Ganoids the archinephric ducts unite and then expand into a urinary sinus or bladder, and the gonoducts of the female, or of both sexes in _Lepidosteus_, open either into the archinephric {408}ducts or into the common sinus, and therefore both ducts communicate with the exterior by a urinogenital orifice behind the anus. Peritoneal funnels, similar to the functional oviducts of the female, are present in the males of the Chrondrostei and of _Amia_. In Teleosts the terminal connexions of the ducts tend to become less intimate. The archinephric ducts often dilate into a urinary bladder either before or after their union, and the common duct joins the united gonoducts to form a short urinogenital sinus which opens externally, or the confluent gonoducts have an independent genital orifice between the anus and the urinary aperture. Not rarely the genital or the urinogenital orifice is prolonged into a tubular papilla, which in the male acts as an intromittent organ, or, as in the females of the Cyprinoid _Rhodeus amarus_, the long oviducal tube serves the purpose of an ovipositor. The males and females of the Siluroid _Plotosus_ have a remarkable vascular and glandular arborescent appendage just behind the urinogenital papilla, the use of which is unknown.[486]

The EGGS of different Fishes[487] exhibit considerable diversity in size and shape as well as in the nature of their external coverings and their mode of deposition.[488] The size of the eggs largely depends on the quantity of food-yolk stored up in their substance for the nutrition of the embryo: hence the eggs of Elasmobranchs, which resemble Fowls' eggs in the superabundance of their yolk, are by far the largest. Teleostomi have much smaller eggs. The largest Teleostean ova are those which are heavy and sink (demersal ova); the smallest, those which are buoyant and float (pelagic ova). Of the former, the eggs of _Gymnarchus_ are about 10 mm. in diameter; those of the Salmon about 5 mm.; and those of some species of _Arius_, 5 to 10 mm. The eggs of the Wolf-Fish (_Anarrhichas lupus_) are about 6 mm. Smaller demersal ova are those of the Lump-sucker (_Cyclopterus_) and _Heterotis_, which are 2.6 and 2.5 mm. respectively. Pelagic {409}eggs are very small, those of the Plaice, which are exceptionally large, varying from 1.65 to 1.95 mm.

An egg-cell consists of living protoplasm and a nucleus, a variable quantity of non-living food-yolk, and of certain enveloping and protective egg-membranes. The ova of Fishes differ principally in the amount and disposition of the food-yolk, in the character of the egg-membranes, and in the presence or absence of special perforations in the egg-membranes for the entrance of spermatozoa into the eggs. In the small ova of some of the lower Chordata (_e.g._ Amphioxus), where the very small quantity of food-yolk is uniformly distributed, and its presence affects all parts of the egg alike, the process of segmentation which follows fertilisation results in the transformation of the entire egg into a mass of approximately equal-sized cells or blastomeres (Fig. 82). {410}The eggs are therefore described as "alecithal," and the segmentation as being "holoblastic" and "equal." On the other hand, all Fishes possess "telolecithal" eggs, that is, ova in which the food-yolk is more or less abundant, and tends to accumulate at one pole of the egg ("vegetative pole"), while the opposite or "animal pole" consists of protoplasm, comparatively free from yolk granules and containing the nucleus (Fig. 234, A). The term telolecithal is, however, a somewhat comprehensive one, and covers important variations in the relations of the inert food-yolk and the living protoplasm in different Fishes, which greatly modify the process of segmentation. Thus there are some Fishes in which the amount of food-yolk at the vegetative pole is sufficient to retard segmentation in that part of the egg without actually preventing it, and consequently segmentation begins in the animal pole, and takes place more rapidly there than it does when it extends into the vegetative pole. Hence it follows that although the entire egg is segmented the blastomeres are of unequal size, the animal pole giving rise to a large number of small cells or micromeres, and the vegetative pole to a smaller number of much larger cells or macromeres. The segmentation of such an egg is said to be holoblastic but unequal (Fig. 234, B and C). This type of egg is characteristic of the Chondrostei, the Holostei, and the Dipnoi. In other Fishes, like the Elasmobranchs and the Teleostei, the food-yolk so greatly preponderates that it entirely prevents segmentation in the vegetative part of the egg, and segmentation is restricted to the small mass of protoplasm (germinal disc) at the animal pole, in which the nucleus is situated (Fig. 234, D). Eggs undergoing partial segmentation in this way are termed "meroblastic." No hard and fast line can be drawn between the two types, and in the Chondrostei and Holostei an interesting transition between the holoblastic and meroblastic ova may be observed. The egg-membranes are formed either by the egg itself or by the epithelium of the ovarian ovisacs, and, as will shortly be seen, the character of the outer egg-membrane greatly influences the mode of deposition of the eggs and their location afterwards. In Elasmobranchs the egg is enclosed in a stout horny egg-shell, secreted by the oviducal shell gland.[489] In many Fishes, as in the Chondrostei, Holostei, and Teleostei, the egg-membranes {411}are perforated at the animal pole of the egg by a small aperture or "micropyle," which is only large enough to admit of the entrance of a single spermatozoon at a time (Fig. 235). Generally, there is only a single micropyle, but, according to Salensky, the Sturgeon (_A. sturio_) has from 3 to 9, and the Sterlet (_A. ruthenus_) from 5 to 13.

An important distinction may be made between the ova of different Teleostomi as regards their location after extrusion from the female. From this point of view two types of ova can be distinguished, _demersal_ and _pelagic_ ova. Demersal eggs are characterised by their larger size and greater weight, so that they always sink after extrusion; and by their opacity. They may either have an outer egg-membrane which is viscid and adhesive, so that the eggs readily adhere to one another or to foreign objects, or the membrane is smooth and non-adhesive. The Salmonidae, for example, produce non-adhesive demersal eggs, which remain separate after being deposited on the gravelly bed of a stream. Most freshwater and many marine shore Fishes have adhesive demersal eggs, which are deposited at the bottom of the water, generally adhering to one another in larger or smaller clumps, masses, or sheets, and attached to rocks, stones, or empty shells, like the eggs of many shore Fishes, or to aquatic plants after the fashion of the eggs of the Carp, Perch, and Pike, or even to branching zoophytes, as is the case with the eggs of the Sea-snail (_Liparis_). In some adhesive eggs the external egg-membrane forms threads for their attachment. The eggs of the Gar-Fish (_Belone_), and those of the Saury Pike (_Scombresox_) and of the Flying Fishes (_Exocoetus_), have viscid threads developed from opposite points on the surface, which are either attached to foreign objects or they become entangled with those of other eggs of the same species. The oval eggs of some of the Gobies have a bunch of fibres at one pole which serves to attach them. In the Smelt (_Osmerus eperlanus_) a portion of the outer egg-membrane breaks away from the rest and becomes turned back, inside out, but remains attached to the egg at one point. By means of this membrane the egg is attached to rocks or stones. Pelagic eggs are distinguished by their lightness and buoyancy, so that they always float near the surface of the water, and by their smaller size and remarkable transparency (Fig. 235). A conspicuous feature in many of them is the presence of a single {412}large oil globule on the surface of the yolk, and not infrequently the yolk becomes partially or completely broken up into small masses. Pelagic eggs are always non-adhesive and free, and they invariably belong to marine Fishes. Amongst the British food Fishes which produce pelagic ova may be mentioned the Gadidae (_e.g._ Cod, Whiting, Hake, Ling), the Pleuronectidae (_e.g._ Turbot, Brill, Sole, Plaice), Scombridae (_e.g._ Mackerel), Triglidae (_e.g._ the Gurnards), Percidae (_e.g._ the Bass), and Clupeidae like the Pilchard and Sprat, but not the Herring, whose adhesive demersal eggs are deposited in clumps on shingly banks in the sea at varying distances from the shore.

The eggs of Elasmobranchs are deposited singly or in pairs at considerable intervals, and the period of egg-laying is prolonged over a considerable part of the year. In most other Fishes, as in Teleosts, the period of spawning is limited to a few months, usually in the spring and summer in temperate latitudes; and in the case of a single Fish it may last only a few days or weeks, but the number of eggs produced is often enormous. Thus, in a Ling 61 inches long and weighing 54 pounds the ovaries contained 28,361,000 eggs. A Turbot, 17 pounds in weight, had 9,161,000 eggs; and a Cod of 21½ pounds 6,652,000. The least prolific of the British food Fishes is the Herring, in which the number of ovarian eggs varied from 21,000 to 47,000 in four specimens examined.[490] The extraordinary fecundity of many Fishes seems to bear no relation to the relative abundance of the Fishes themselves, but rather it is to be associated with certain {413}disadvantages attendant on the sexual relations of Fishes, involving a considerable waste of the sex-cells, while in many Fishes it no doubt helps to compensate for any subsequent mortality among the larvae, which may result from an uncertain and precarious food supply and from the attacks of enemies. Whenever internal fertilisation is the rule, or when, as in nest-building and marsupial Fishes, the propinquity of the sexes in the breeding season ensures the fertilisation of a larger proportion of the eggs and the protection of the young, the number of eggs produced is small.

The male sex-cells or spermatozoa are essentially similar to those of other Vertebrates, although in different Fishes they may vary in such details as length, and the shape and size of the head, which may be rod-like and wavy, elliptical or globular.

As a rule, in Fishes females are more numerous than males, and generally they are larger, but to both statements there are notable exceptions. The relations of the sexes in the breeding season are usually very promiscuous, especially in those Teleosts which discharge their sex-cells while swimming together in shoals. A female may, however, consort with several males (_polyandry_), or a male with several females (_polygamy_); or, as in some of the nest-building Fishes (e.g. _Gastrosteus_), there are not wanting examples of the pairing of one male with one female (_monogamy_).

Fishes often migrate at the commencement of the breeding-season to localities most suitable for the deposition of the eggs. Many marine species seek banks or shallower water near the shore, and some, like the Salmon and the Sturgeon, are _anadromous_, and ascend rivers for long distances to deposit their spawn.

In all Fishes except the Elasmobranchs and a few Teleosts the fertilisation of the eggs takes place in the water after their extrusion, the male depositing its seminal fluid over the eggs or in their neighbourhood. The waste of the sex-cells is often, no doubt, very considerable, especially when the eggs are adhesive and fixed, and the seminal fluid is liable to drift at the mercy of tides and currents. With pelagic ova the waste is perhaps not so great, inasmuch as the eggs as well as the spermatozoa would probably drift at the same rate and in the same direction. Liability to waste must also be greatly diminished in many Fishes by their habit of living in shoals, or of congregating {414}together in the breeding season, in which they are sometimes aided by their power of emitting characteristic sounds, and in the case of nest-building Fishes by the still more intimate relations of the sexes. Even when the liability to waste is very great, compensation may be afforded by exceptional fecundity. The copulation of the sexes and the internal fertilisation of the eggs occur only in Elasmobranchs and some Teleosts. The copulatory organs of Elasmobranchs are the so-called "claspers" with which the males are provided. Some form of copulation is probably the rule in the viviparous Teleosts, where the eggs are fertilised in the oviducts, or even while they are still in the ovaries, and the young are born alive. As mentioned above, an intromittent organ is often formed by the prolongation of the genital or the urinogenital orifice into a papilla, or a longer or shorter tube.[491] Some Cyprinodontidae[492] (e.g. _Anableps_) have the anterior part of the anal fin modified in the male to form an intromittent organ along which the urinogenital canal runs (Fig. 374). In the females the genital aperture is covered by a special scale, which is free on one side and not on the other. "The male organ in some individuals is turned to the right, in others to the left, and in some females the opening beneath the special scale is to the right, in others to the left. Copulation thus takes place sideways, a left-sided male pairing with a right-sided female, and _vice versa_."[493] The anal fin also forms an intromittent organ in the "Half-beak" (_Hemirhamphus_). In a genus (_Girardinus_) of the same family the anal fin is modified to form an apparatus for holding the female during sexual congress.[494] The singular method of fertilisation practised by the males and females of _Callichthys paleatus_ is referred to elsewhere.[495]

With the exception of the pelagic _Antennarius_, which builds its nest in the Sargasso weed in mid-ocean, nest-building and parental solicitude for the young are confined to freshwater Fishes and to marine forms with demersal ova. Pelagic ova must necessarily be beyond the scope of parental care. As a rule it is the male which acts as guardian nurse, the female troubling herself but little about the fate of her eggs or her offspring.

{415}

Perhaps the more primitive form of parental foresight is exhibited by those Fishes which, like the females of the Salmonidae, make a furrow in the gravelly bottom of a running stream for the reception of the eggs, and then cover them over with a layer of gravel, or like the Siluroid _Arius australis_, of the Burnett river in Queensland, which deposits its eggs in circular excavations in the sandy bed of the river and covers them with layers of large stones. But in neither case does it appear that either the male or the female takes any further interest in the eggs or in the young when hatched. Without actual nest-building, or even the preparation of a place for their reception, the eggs may be protected in various ways by the male. The common British Gunnel or Butter-Fish (_Pholis gunnellus_) rolls its eggs into a rounded mass by coiling its body round them, the male and female taking possession of them alternately. The little clumps of eggs are then deposited in holes made by the boring Mollusc, _Pholas_. Some British Blennies attach their eggs in a single layer to the sides of cavities in rocks, or between stones, where they are watched over by the male parent. The eggs of the Lump-Sucker (_Cyclopterus lumpus_) are attached in masses to rocks or to piles and guarded by the male, who aerates them by keeping up a flow of water over the spawn through the action of his pectoral fins. When hatched, the young fry cling to the body of their watchful parent by their suckers. A more decided approach to nest-building is exhibited by the Sand Goby (_Gobius minutus_). In this species the male scoops out the sand from beneath an empty shell, generally that of a Pecten, and the female deposits her adhesive eggs on the under surface of the shell.

{416}

The male remains on guard, and by the movements of its pectoral fins promotes the aeration of this rude form of nest. References to some of the more striking examples of true nest-building in Fishes will be found in the systematic part of this volume, especially in those chapters treating of the Dipnoi and Amiidae, and such Teleosts as the Mormyridae, Osteoglossidae, Siluridae, Gastrosteidae, Centrarchidae, Osphromenidae, Labridae, and Antennariidae. Other illustrations of parental care are to be found in the development of marsupial pouches or grooves for the reception of the eggs in the males of the Syngnathidae (Fig. 387) and the females of the Solenostomidae, and the use of the oral cavity for a similar purpose in the males, rarely in the females, of some Siluridae, and the males or females, according to the species, of the Cichlidae. The singular method by which the female _Aspredo_ safeguards both her eggs and her progeny is referred to on p. 596. The Cyprinoid, _Rhodeus amarus_ (the "Bitterling" of Central Europe), is unique in the means which it adopts to {417}secure the same result.[496] By means of its long ovipositor the female Fish deposits its eggs in the mantle cavity of a _Unio_, or of an Anodon. Here they are fertilised by spermatozoa carried in through the inhalent siphon of the Mollusc with the inspiratory water current, and they complete their development in the gill-cavities (Fig. 237).[497]

The time which elapses between the fertilisation of the egg and the hatching out of the young Fish varies greatly in different Teleosts. The eggs of some Clupeidae hatch in a very short time, two to three days in the Anchovy, and three to four days in the Sprat. In most of the British marine food Fishes the period rarely exceeds twelve to fourteen days. The larger demersal eggs with much food-yolk are longer in hatching; in the Salmon the time ranging from thirty-five to one hundred and forty-eight days. A low temperature lengthens the time. The eggs of the Herring which hatched in eight to nine days at a temperature of 52° to 58° F. took forty-seven days in water at 32° F.