CHAPTER VIII.
THE ORGANS OF NUTRITION AND DIGESTION.
Fishes are either exclusively carnivorous or herbivorous, but not a few feed on vegetable substances as well as animal, or on mud containing alimentary substance in a living or decomposing state. Generally they are very voracious, especially the carnivorous kinds, and the rule of “eat or be eaten” applies to them with unusual force. They are almost constantly engaged in the pursuit and capture of their prey, the degree of their power in these respects depending on the dimensions of the mouth and gullet and the strength of the teeth and jaws. If the teeth are sharp and hooked, they are capable of securing the most slender and agile animals; if this kind of teeth is combined with a wide gullet and distensible stomach, they are able to overpower and swallow other fish larger than themselves; if the teeth are broad, strong molars, they are able to crush the hardest aliments; if they are feeble, they are only serviceable in procuring some small or inert and unresisting prey. Teeth may be wanting altogether. Whatever the prey, in the majority of cases it is swallowed whole; but some of the most voracious fishes, like some Sharks and Characinidæ, are provided with cutting teeth, which enable them to tear their prey to pieces if too large to be swallowed whole. Auxiliary organs for the purpose of overpowering their prey, which afterwards is seized or torn by the teeth, like the claws of some carnivorous mammals and birds, are not found in this class; but in a few fishes the jaws themselves are modified for that purpose. In the Sword-fishes the bones of the upper jaw form a long dagger-shaped weapon, with which they not only attack large animals, but also frequently kill fishes on which they feed. The Saw-fishes are armed with a similar but still more complicated weapon, the saw, which is armed on each side with large teeth implanted in deep sockets, specially adapted for killing and tearing the prey before it is seized and masticated by the small teeth within the mouth. Fishes show but little choice in the selection of their food, and some devour their own offspring indiscriminately with other fishes. Their digestive powers are strong and rapid, but subject in some degree to the temperature, which, when sinking below a certain point, lowers the vital powers of these cold-blooded animals. On the whole, marine fishes are more voracious than those inhabiting fresh waters; and whilst the latter may survive total abstinence from food for weeks or months, the marine species succumb to hunger within a few days. The growth of fishes depends greatly on the nature and supply of food, and different individuals of the same species may exhibit a great disparity in their respective dimensions. They grow less rapidly and to smaller dimensions in small ponds or shallow streams than in large lakes and deep rivers. The young of coast fishes, when driven out to sea, where they find a much smaller supply of food, remain in an undeveloped condition, assuming an hydropic appearance. The growth itself seems to continue in most fishes for a great length of time, and we can scarcely set bounds to—certainly we know not with precision—the utmost range of the specific size of fishes. Even among species in no way remarkable for their dimensions we sometimes meet with old individuals, favourably situated, which more or less exceed the ordinary weight and measurement of their kind. However, there are certain evidently short-lived species of fishes which attain a remarkably uniform size within a very short time; for instance, the Stickleback, many species of Gobius and Clupea.
The organs of nutrition, manducation, and deglutition, are lodged in two large cavities—an anterior (the mouth or buccal cavity), and a posterior (the abdominal cavity). In the former the alimentary organs are associated with those fulfilling the respiratory functions, the transmission of food to the stomach and of water to the gills being performed by similar acts of deglutition. The abdominal cavity commences immediately behind the head, so, however, that an extremely short thoracic cavity for the heart is partitioned off in front. Beside the alimentary organs it contains also those of the urogenital system and the air-bladder. The abdominal cavity is generally situated in the trunk only, but in numerous fishes it extends into the tail, being continued for some distance along each side of the hæmal apophyses.
In numerous fishes the abdominal cavity opens outwards by one or two openings. A single porus abdominalis in front of the vent is found in Lepidosiren and some Sturgeons; a paired one, one on each side of the vent, in Ceratodus, some species of Sturgeon, Lepidosteus, Polypterus, Amia, and all Chondropterygians. As in these fishes semen and ova are discharged by proper ducts, the abdominal openings may serve for the expulsion of semen, and those ova only which, having lost their way to the abdominal aperture of the oviduct, would be retained in the abdominal cavity. In those Teleosteans which lack an oviduct a single porus genitalis opens behind the vent.
The mouth of fishes shows extreme variation with regard to form, extent, and position. Generally opening in front, it may be turned upwards, or may lie at the lower side of the snout, as in most Chondropterygians, Sturgeons, and some Teleosteans. Vogt regards this position as a persistent fœtal condition. In most fishes the jaws are covered by the skin, which, before passing over the jaws, is often folded, forming more or less fleshy lips. In the Sharks the skin retains its external character even within the teeth, but in other fishes it changes into a mucous membrane. A tongue may exist as a more or less free and short projection, formed by the glosso*-hyal and a soft covering, or may be entirely absent. Salivary glands and a velum palati are absent in fishes.
With regard to the dentition, the class of Fishes offers an amount of variation such as is not found in any of the other classes of Vertebrates. As the teeth form one of the most important elements in the classification of fishes, their special arrangement and form will be referred to in the account of the various families and genera. Whilst not a few fishes are entirely edentulous, in others most of the bones of the buccal cavity, or some of them, may be toothed, as the bones of the jaws, the palatines, pterygoids, vomers, basisphenoid, glossohyal, branchial arches, upper and lower pharyngeals. In others teeth may be found fixed in some portion of the buccal membrane without being supported by underlying bone or cartilage; or the teeth have been developed in membrane overlying one of the dentigerous bones mentioned, without having become anchylosed to the bone. When the tooth is fixed to the bone the attachment has generally been effected by the ossification of the bone of the tooth, but in some fishes a process of the bone projects into the cavity of the tooth; in others the teeth are implanted in alveoli. In these, again, frequently a process of bone rises from the bottom, on which the tooth rests.
Many fishes, especially predatory fishes with long, lancet-shaped teeth, have all or some of the teeth capable of being bent inwards towards the mouth. Such “hinged” teeth resume at once the upright position when pressure is removed from them. They are, however, depressible in one direction only, thus offering no obstacle to the ingress, but opposing the egress of prey. Mr. C. S. Tomes has shown that the means by which this mechanism is worked are different in different fishes; for whilst, in the Pediculati and Gadoids (Hake) the elasticity resides solely in the tissue of the hinge (the tooth being as resilient as ever after everything else is severed), in the Pike the hinge is not in the least endowed with elasticity, but the bundles of fibres proceeding from the interior of the dentine cap are exceedingly elastic.
As regards texture the teeth of fishes show considerable variation. The conical teeth of the Cyclostomes and the setiform teeth of many Teleosteans consist of a horny albuminous substance. The principal substance of the teeth of other fishes consists of dentine, with numerous dividing and anastomosing tubercles, sometimes covered by a stratum of unvascular dentine. An enamel-like substance has been observed on the crown of the teeth of Sargus and Balistes, and an ossification of the capsule of their matrix covers the enamel with a thin coating of cement. The teeth either possess a cavity in which the matrix is received, or, more frequently, they are solid, in which case vascular canals of the underlying bone are continued into the substance of the tooth. In the teeth of some fishes numerous sets of canals and tubes are so arranged that they do not anastomose with one another, each set being surrounded by a layer of dentine and cement. These apparently simple teeth are evidently composed of numerous small teeth, and called compound teeth.
The teeth may be, and generally are, very different as regards size or form in the different parts of the mouth; they may be also different according to the age or sex of the fish (Raja). The teeth may be few in number and isolated, or placed in a single, double, or triple series, distant from one another or closely set; they may form narrow or broad bands, or patches of various forms. As regards form, they may be cylindrical or conical, pointed, straight, or curved, with or without an angular bent near their base; some are compressed laterally or from the front backwards; the latter may be triangular in shape, or truncated at the top like incisors of mammals; they may have one apex (cusp) only, or be bi- or tri-lobate (bi- or tri-cuspid); or have the margins denticulated or serrated. Compressed teeth may be confluent, and form a cutting edge in both jaws, which assume the shape of a parrot’s beak (Fig. [53]). In some the apex is hooked or provided with barbs. Again, some teeth are broad, with flat or convex surface, like molar teeth. With regard to size, the finest teeth are like fine flexible bristles, ciliiform or setiform; or, if very short and anchylosed to the bone, they appear only as inconspicuous asperities of the bone. Very fine conical teeth arranged in a band are termed villiform teeth; when they are coarser, or mixed with coarser teeth, they are card-like (dents en rape or en cardes) (Fig. [54]); molar-like teeth of very small size are termed granular.