Читать книгу The Vertebrate Skeleton - Sidney H. Reynolds - Страница 14

Table of Contents

EXOSKELETON.

The most primitive type of exoskeleton is that found in Elasmobranchs and formed of placoid scales; these are tooth-like structures consisting of dentine and bone capped with enamel, and have been already described (p. 4). In most Elasmobranchs they are small and their distribution is fairly uniform, but in the Thornback skate, Raia clavata, they have the form of larger, more scattered spines. In adult Holocephali and in Polyodon and Torpedo there is no exoskeleton, in young Holocephali, however, there are a few small dorsal ossifications.

The plates or scales of many Ganoids may have been formed by the gradual fusion of elements similar to these placoid scales, and often bear a number of little tooth-like processes. In Lepidosteus, Polypterus, and many extinct species, these ganoid scales, which are rhomboidal in form and united to one another by a peg and socket articulation, enclose the body in a complete armour. In Trissolepis part of the tail is covered by rhomboidal scales, while rounded scales cover the trunk and remainder of the tail. Acipenser and Scaphirhynchus have large dermal bony plates which are not rhomboidal in shape and do not cover the whole body. In Acipenser a single row extends along the middle of the back and two along each side.

The majority of Teleosteans have thin flattened scales which differ from those of Ganoids in being entirely mesodermal in origin, containing no enamel. There are two principal types of Teleostean scales, the cycloid and ctenoid. A cycloid scale is a flat thin scale with concentric markings and an entire posterior margin. A ctenoid scale differs in having its posterior margin pectinate. The Dipnoi have overlapping cycloid scales. The rounded scales of Amia and of many fossil ganoids such as Holoptychius are shaped like cycloid scales, but differ from them in being more or less coated with enamel. In Eels and some other Teleosteans the scales are completely degenerate and have almost disappeared. Some Teleosteans, like Diodon hystrix, have scales with triradiate roots from which arise long sharp spines directed backwards. These scales, which resemble teeth, contain no enamel; they become erect when the fish inflates its body into a globular form. Many Siluroids have dermal armour in the form of large bony plates which are confined to the anterior part of the body. In Ostracion the whole body is covered by hexagonal plates, closely united together.

The fin-rays are structures of dermal origin which entirely or partially support the unpaired fins, and assist the bony or cartilaginous endoskeleton in the support of the paired fins.

In Elasmobranchs, Dipnoi, and Chondrosteous ganoids the skeletons of the fins are, as a rule, about half of exoskeletal, half of endoskeletal origin, the proximal and inner portion being cartilaginous and endoskeletal, the distal and outer portion being exoskeletal, and consisting of horny or of more or less calcified fin-rays. In bony Ganoids and Teleosteans the endoskeletal parts are greatly reduced and the fins come to consist mainly of the fin-rays, which are ossified and frequently become flattened at their distal ends.

The fin-rays of the ventral part of the caudal fin are carried by the haemal arches; those of the dorsal and anal fins and of the dorsal part of the caudal fin generally by interspinous bones, which in adult Teleosteans alternate with the neural and haemal spines. In Dipnoi these interspinous bones articulate with the neural and haemal spines. In many Siluroids the anterior rays of the dorsal and pectoral fins are developed into large spines which often articulate with the endoskeleton, or are sometimes fused with the dermal armour plates. Similar spines may occur in Ganoids in front of both the dorsal and anal fins. Polypterus has a small spine or fulcrum in front of each segment of the dorsal fin. Such spines are often found fossilised, and are known as ichthyodorulites.

Similar spines are found in many Elasmobranchs, but they are simply inserted in the flesh, not articulated to the endoskeleton. They also differ from the spines of Teleosteans and Ganoids in the fact that they are covered with enamel, and often have their edges serrated like teeth. In the extinct Acanthodii they generally occur in front of all the fins, paired and unpaired.

In Trygon, the Sting-ray, the tail bears a serrated spine which is used for purposes of offence and defence. Many ichthyodorulites may have been spines of this nature fixed to the tail, rather than spines situated in front of the fins. The spines, which are always found in front of the dorsal fin in Holocephali, agree with those of Elasmobranchs in containing enamel, and with those of Teleosteans in being articulated to the endoskeleton.

Teeth.

The teeth of fish[40] are subject to a very large amount of variation, perhaps to more variation than are those of any other class of animals. Sometimes, as in adult Sturgeons, they are entirely absent, sometimes they are found on all the bones of the mouth, and also on the hyoid and branchial arches. The teeth are all originally developed in the mucous membrane of the mouth, but they afterwards generally become attached to firmer structures, especially to the jaws. In Elasmobranchs, however, they are generally simply imbedded in the tough fibrous integument of the mouth. Their attachment to the jaws may take place in three different ways.

Fig. 14. Diagram of a section through the jaw of a Shark (Odontaspis americanus) showing the succession of teeth (Brit. Mus. from specimen and diagram).

1. teeth in use.	cartilage.
2. teeth in reserve.	6. connective tissue.
3. skin.	7. mucous membrane of the
4. cartilage of the jaw.	mouth.
5. encrusting calcification of

(1) By an elastic hinge-joint, as in the Angler (Lophius), and the Pike (Esox lucius). In the Angler the tooth is held by a fibrous band attaching its posterior end to the subjacent bone, in the Pike by uncalcified elastic rods in the pulp cavity.

(2) By ankylosis, i.e. by the complete union of the calcified tooth substance with the subjacent bone. This is the commonest method among fish.

(3) By implantation in sockets. This method is not very common among fish. The teeth are sometimes, as in Lepidosteus, ankylosed to the base of the socket. In this genus there is along each ramus of the mandible a median row of large teeth placed in perfect sockets, and two irregular lateral rows of small teeth ankylosed to the jaw.

Dentine, enamel and cement are all represented in the teeth of fishes, but the enamel is generally very thin, and cement is but rarely developed. Dentine forms the main bulk of the teeth; it is sometimes of the normal type, but generally differs from that in higher vertebrates in being vascular, and is known as vasodentine. A third type occurs, known as osteodentine; it is traversed by canals occupied by marrow, and is closely allied to bone.

Fig. 15. Part of the lower jaw of a Shark (Galeus) (from Owen after André).

1. teeth in use.	a Sting-ray (Trygon) which has
2. reserve teeth folded back.	pierced the jaw and affected the
3. part of the caudal spine of	growth of the teeth.

The teeth are generally continually renewed throughout life, but sometimes one set persists.

The teeth of Selachii are fundamentally identical with placoid scales. They are developed from a layer of dental germs which occurs all over the surface of the skin, except in the region of the lips. At this point the layer of tooth-producing germs extends back into the mouth, being projected by a fold of the mucous membrane (fig. 14, 7). Here new teeth are successively formed, and as they grow each is gradually brought into a position to take the place of its predecessor by the shifting outwards of the gum over the jaw. Owing to this arrangement sharks have practically an unlimited supply of teeth (figs. 14 and 15).

Two principal types of teeth are found in Elasmobranchs. In Sharks and Dogfish, on the one hand, the teeth are very numerous, simple, and sharp-pointed, and are with or without serrations and lateral cusps. Many Rays and fossil Elasmobranchs, on the other hand, have broad flattened teeth adapted for crushing shells. Intermediate conditions occur between these two extremes. Thus in Cestracion and many extinct sharks, such as Acrodus, while the median teeth are sharp, the lateral teeth are more or less flattened and adapted for crushing. In various species belonging to the genus Raia the teeth of the male are sharp, while those of the female are blunt. A very specialised dentition is met with in the Eagle-rays (Myliobatidae), in which the jaws are armed with flattened angular tooth-plates, arranged in seven rows, forming a compact pavement; the plates of the middle row are very wide and rectangular, those of the other rows are much smaller and hexagonal. Lastly, in Cochliodus the individual crushing teeth are fused, forming two pairs of spirally-coiled dental plates on each side of each jaw. Pristis, the Saw-fish, has a long flat cartilaginous snout, bearing a double row of persistently-growing teeth planted in sockets along its sides. Each tooth consists of a number of parallel dentinal columns, united at the base, but elsewhere distinct.

In the Holocephali—Chimaera, Hariotta and Callorhynchus—only three pairs of teeth or dental plates occur, two pairs in the upper jaw, one in the lower. These structures persist throughout life and grow continuously. The upper tooth structures are attached respectively to the ethmoid or vomerine region of the skull, and to the palato-pterygoids. The vomerine teeth are small, while those attached to the mandible and the palato-pterygoid region are large and bear several roughened ridges adapted for grinding food. The teeth of the two opposite sides of the jaw meet in a median symphysis. The teeth of Chimaera are more adapted for cutting, those of Callorhynchus for crushing. Many extinct forms are known, some of whose teeth are intermediate in structure between those of Chimaera and Callorhynchus.

The teeth of Ganoids are also extremely variable. Among living forms, the Holostei are more richly provided with teeth than are any other fishes, as they may occur on the premaxillae, maxillae, palatines, pterygoids, parasphenoid, vomers, dentaries, and splenials. Among the Chondrostei, on the other hand, the adult Acipenseridae are toothless; small teeth however occur in the larval sturgeon, and in Polyodon many small teeth are found attached merely to the mucous membrane of the jaws. Many fossil Ganoids have numerous flattened or knob-like teeth, borne on the maxillae, palatines, vomers and dentaries. Others have a distinctly heterodont dentition. Thus in Lepidotus the premaxillae bear chisel-like teeth, while knob-like teeth occur on the maxillae, palatines and vomers. In Rhizodus all the teeth are pointed, but while the majority are small a few very large ones are interspersed.

In Teleosteans, too, the teeth are eminently variable both in form and mode of arrangement. They may be simple and isolated, or compound, and may be borne on almost any of the bones bounding the mouth cavity, and also as in the Pike, on the hyoid and branchial arches. The splenial however never bears teeth and the pterygoid and parasphenoid only rarely, thus differing from the arrangement in the Holostei.

The isolated teeth are generally conical in form and are ankylosed to the bone that bears them. Such teeth are, with a few exceptions such as Balistes, not imbedded in sockets nor replaced vertically.

In some fish beak-like structures occur, formed partly of teeth, partly of the underlying jaw bones. These beaks are of two kinds: (1) In Scarus, the parrot fish, the premaxillae and dentaries bear numerous small, separately developed teeth, which are closely packed together and attached by their proximal ends to the bone, while their distal ends form a mosaic. Not only the teeth but the jaws which bear them are gradually worn away at the margins, while both grow continuously along their attached edge. (2) In Gymnodonts, e.g. Diodon, the beaks are formed by the coalescence of broad calcified horizontal plates, which when young are free and separated from one another by a considerable interval.

In some Teleosteans the differentiation of the teeth into biting teeth and crushing teeth is as complete as in Lepidosteus. Thus in the Wrasse (Labrus), the jaws bear conical slightly recurved teeth arranged in one or two rows, with some of the anterior ones much larger than the rest. The bones of the palate are toothless, while both upper and lower pharyngeal bones are paved with knob-like crushing teeth; such pharyngeal teeth occur also in the Carp but are attached only to the lower pharyngeal bone, the jaw bones proper being toothless.

In Dipnoi the arrangement of the teeth is very similar to that in Holocephali. The mandible bears a single pair of grinding teeth attached to the splenials, and a corresponding pair occur on the palato-pterygoids. In front of these there are a pair of small conical vomerine teeth loosely attached to the ethmoid cartilage. The palato-pterygoid teeth of Ceratodus are roughly semicircular in shape with a smooth convex inner border, and an outer border bearing a number of strongly marked ridges. The teeth of the extinct Dipteridae resemble those of Ceratodus but are more complicated.

ENDOSKELETON.

Spinal column[41].

The spinal column of fishes is divisible into only two regions, a caudal region in which the haemal arches or ribs meet one another ventrally, and a precaudal region in which they do not meet.

The various modifications of the spinal column in fishes can be best understood by comparing them with the arrangement in the simplest type known, namely Amphioxus. In Amphioxus the notochord is immediately surrounded by a structureless cuticular layer, the chordal sheath. Outside this is the skeletogenous layer, which in addition to surrounding the notochord and chordal sheath embraces the nerve cord dorsally, and laterally sends out septa forming the myomeres.

The Cartilaginous ganoids[42] Acipenser, Polyodon and Scaphirhynchus are the simplest fishes as regards their spinal column. The notochord remains permanently unconstricted and is enclosed in a chordal sheath, external to which is the skeletogenous layer. In this layer the development of cartilaginous elements has taken place. In connection with each neuromere, or segment as determined by the points of exit of the spinal nerves, there are developed two pairs of ventral cartilages, the ventral arches (basiventralia) and intercalary pieces (interventralia); and at least two pairs of dorsal pieces, the neural arches (basidorsalia) and intercalary pieces (interdorsalia). The lateral parts of the skeletogenous layer do not become converted into cartilage, so there are no traces of vertebral centra. The ventral or haemal arches meet one another ventrally and send out processes to protect the ventral vessels. The neural arches do not meet, but are united by a longitudinal elastic band.

In Cartilaginous ganoids the only indications of metameric segmentation are found in the neural and haemal arches. The case is somewhat similar with the Holocephali and Dipnoi.

In the Holocephali the notochord grows persistently throughout life, and is of uniform diameter throughout the whole body except in the cervical region and in the gradually tapering tail. The chordal sheath is very thick and includes a well-marked zone of calcification which separates an outer zone of hyaline cartilage from an inner zone. There are also a number of cartilaginous pieces derived from the skeletogenous layer which are arranged in two series, a dorsal series forming the neural arches and a ventral series forming the haemal arches. These do not, except in the cervical region, meet one another laterally round the notochord and form centra. To each neuromere there occur a pair of basidorsals, a pair of interdorsals, and one or two supradorsals. In the tail the arrangement is irregular.

In the Dipnoi as in the Holocephali the notochord grows persistently and uniformly, and the chordal sheath is thick and cartilaginous though there are no metamerically arranged centra. The neural and haemal arches and spines are cartilaginous and interbasalia (intercalary pieces) are present. The basidorsalia and basiventralia do not in Ceratodus meet round the notochord and enclose it except in the anterior part of the cervical and posterior part of the caudal region.

In Elasmobranchii the chordal sheath is weak and the skeletogenous layer strong. Biconcave cartilaginous vertebrae are developed, and as is the case in most fishes, constrict the notochord vertebrally.

Two distinct types of vertebral column can be distinguished in Elasmobranchs[43]:

1. In many extinct forms and in the living Notidanidae, Cestracion, and Squatina, the dorsal and ventral arches do not meet one another laterally round the centrum, and consequently readily come away from it.

2. In most living Elasmobranchs the arches meet laterally round the centrum.

The vertebrae are never ossified but endochondral calcification nearly always takes place, though it very rarely reaches the outer surface of the vertebrae. Elasmobranchs are sometimes subdivided into three groups according to the method in which this calcification takes place:

1. Cyclospondyli (Scymnus, Acanthias), in which the calcified matter is deposited as one ring in each vertebra.

2. Tectospondyli (Squatina, Raia, Trygon), in which there are several concentric rings of calcification.

3. Asterospondyli (Notidanidae, Scyllium, Cestracion), in which the calcified material instead of forming one simple ring, extends out in a more or less star-shaped manner.

In Heptanchus the length of the vertebral centra in the middle of the trunk is double that in the anterior and posterior portions, and as the length of the arches does not vary, the long centra carry more of them than do the short centra.

In many Rays the skull articulates with the vertebral column by distinct occipital condyles.

In Bony Ganoids the skeletogenous layer becomes calcified ectochondrally in such a way that the notochord is pinched in at intervals, and distinct vertebrae are produced. Ossification of the calcified cartilage rapidly follows. In Amia the vertebrae are biconcave, in Lepidosteus they are opisthocoelous, cup and ball joints being developed between the vertebrae in a manner unique among fishes. The notochord entirely disappears in the adult Lepidosteus, but at one stage in larval life it is expanded vertebrally and constricted intervertebrally in the manner usual in the higher vertebrata, but unknown elsewhere among fishes.

The tail of Amia is remarkable from the fact that as a rule to each neuromere, as determined by the exit of the spinal nerves, there are two centra, a posterior one which bears nothing, and an anterior one which bears the neural and haemal arches, these being throughout the vertebral column connected with the centra by cartilaginous discs.

In most Teleosteans but not in the Plectognathi the neural arches are continuous with the centra, which are nearly always deeply biconcave.

In some cases many of the anterior vertebrae are ankylosed together and to the skull. The vertebrae often articulate with one another by means of obliquely placed flattened surfaces, the zygapophyses. The centrum in early stages of development is partially cartilaginous, but the neural arches and spines in the trunk at any rate, pass directly from the membranous to the osseous condition.

Fins.

The most primitive fins are undoubtedly the unpaired ones, which probably originally arose as ridges or folds of skin along the mid-dorsal line of the body, and passed thence round the posterior end on to the ventral surface, partially corresponding in position and function to the keel of a ship.

In long 'fish' which pass through the water with an undulating motion such simple continuous fins may be the only ones found, as in Myxine. To support these median fins skeletal structures came to be developed; these show two very distinct forms, viz. cartilaginous endoskeletal pieces, the radiale, and horny exoskeletal fibres, the fin-rays. Mechanical reasons caused the fin to become concentrated at certain points and reduced at intervening regions. Thus a terminal caudal fin arose and became the chief organ of propulsion, and the dorsal and ventral fins became specialised to act as balancing organs.

In some of the earlier Elasmobranchs, the Pleuracanthidae, the endoskeletal cartilaginous radiale are directly continuous with outgrowths from the dorsal and ventral arches of the vertebrae, and form the main part of the fin. In later types of Elasmobranchs the horny exoskeletal fin-rays have comparatively greater prominence. In bony fish, as has been already stated, the horny fibres are replaced by bony rays of dermal origin, and at the same time complete reduction and disappearance of the cartilaginous radiale takes place.

The Caudal fin.

The caudal region of the spinal column in fishes is of special importance. It is distinctly marked off from the rest of the spinal column by the fact that the ventral or haemal arches meet one another and are commonly prolonged into spines, while in the trunk region they do not meet but commonly diverge from one another.

In some fish the terminal part of the caudal region of the spinal column retains the same direction as the rest of the spinal column. The blade of the caudal fin is then divided into two nearly equal portions, and is said to be diphycercal. This condition is generally regarded as the most primitive one; it occurs in the Ichthyotomi, Holocephali, all living Dipnoi, Polypterus and some extinct Crossopterygii, and a few Selachii and Teleostei. It occurs also in deep-sea fish belonging to almost every group, and under these conditions obviously cannot be regarded as primitive, but must be looked on as a feature induced by the peculiar conditions of life.

In the great majority of fish the terminal part of the caudal region of the spinal column is bent dorsalwards, and the part of the blade of the caudal fin which arises on the dorsal surface is much smaller than is that arising on the ventral surface. Such a fin is said to be heterocercal.

Strictly speaking all fish whose tails are not diphycercal have heterocercal tails, but the term is commonly applied to two-bladed tails in which the spinal column forms a definite axis running through the dorsal blade, while the ventral blade is enlarged and generally forms the functional part of the tail. Such heterocercal tails are found in nearly all Elasmobranchii, together with the living cartilaginous Ganoidei, and many extinct forms belonging to the same order; Lepidosteus, Amia, and the Dipteridae among Dipnoi, have tails which, though obviously heterocercal, are not two-bladed.

The vast majority of the Teleostei and some extinct Ganoidei have heterocercal tails of the modified type to which the term homocercal is applied. The hypural bones which support the lower half of the tail fin become much enlarged, and frequently unite to form a wedge-shaped bone which becomes ankylosed to the last ossified vertebral centrum. The fin-rays then become arranged in such a way as to produce a secondary appearance of symmetry. Some homocercal fish such as the Perch have the end of the notochord protected by a calcified or completely ossified sheath, the urostyle, to which several neural and haemal arches may be attached, and which becomes united with the centrum of the last vertebra; in others such as the Salmon the end of the notochord is protected only by laterally placed bony plates.

The Skull.

It is often impossible to draw a hard and fast line between the cranium and the vertebral column. This is the case for instance in Acipenser (fig. 18, 16) among Chondrostei, in Amia among Holostei, and in Ceratodus and Protopterus among Dipnoi. The occipital region of the skull in Amia is clearly formed of three cervical vertebrae whose centra have become absorbed into the cranium, while the neural arches and spines are still distinguishable.