é, Mouth of ductus choledochus; f, stomach; i, adipose agglomeration; l, first compartment of intestinal spire; m, spleen; oe, lower part of œsophagus, opened; p, double pyloric fold; q q, glandular patches.
CHAPTER IX.
ORGANS OF RESPIRATION.
Fishes breathe the air dissolved in water by means of gills or branchiæ. The oxygen consumed by them is not that which forms the chemical constituent of the water, but that contained in the air which is dissolved in water. Hence fishes transferred into water from which the air has been driven out by a high temperature, or in which the air absorbed by them is not replaced, are speedily suffocated. The absorption of oxygen by fishes is comparatively small, and it has been calculated that a man consumes 50,000 times more than is required by a Tench. However, some fishes evidently require a much larger supply of oxygen than others: Eels and Carps, and other fishes of similar low vitality, can survive the removal out of their elements for days, the small quantity of moisture retained in their gill-cavity being sufficient to sustain life, whilst other fishes, especially such as have very wide gill-openings, are immediately suffocated after being taken out of the water. In some fishes noted for their muscular activity, like the Scombridæ, the respiratory process is so energetic as to raise the temperature of their blood far beyond that of the medium in which they live. A few fishes, especially such as are periodically compelled to live in water thickened into mud by desiccation and vitiated by decomposing substances, breathe atmospheric air, and have generally special contrivances for this purpose. These are so much habituated to breathing air that many of them, even when brought into pure water of normal condition, are obliged to rise to the surface at frequent intervals to take in a quantity of air, and if they be kept beneath the surface by means of a gauze net, they perish from suffocation. The special contrivances consist of additional respiratory organs, lodged in cavities either adjoining the gill-cavity or communicating with the ventral side of the œsophagus, or of the air-bladder which enters upon respiratory functions (Dipnoi, Lepidosteus, Amia).
The water used by fishes for respiration is received by the mouth, and by an action similar to that of swallowing driven to the gills, and expelled by the gill-openings, of which there may be one or several on each side behind the head; rarely one only in the median line of the ventral surface.
Fig. 58.—Fore-part of the body of an embryon of Carcharias, showing the branchial filaments (natural size).
The gills or branchiæ consist essentially of folds of the mucous membrane of the gill-cavity (laminæ branchiales), in which the capillary vessels are distributed. In all fishes the gills are lodged in a cavity, but during the embryonic stage the Chondropterygians have the gill-laminæ prolonged into long filaments projecting beyond the gill-cavity (Fig. [58]), and in a few young Ganoids external gills are superadded to the internal.
In Branchiostoma the dilated pharynx is perforated by numerous clefts, supported by cartilaginous rods (Fig. [29], h). The water passes between these clefts into the peritoneal cavity, and makes its exit by the porus abdominalis situated considerably in advance of the vent. The water is propelled by cilia.
In the Cyclostomes the gills of each side are lodged in a series of six or more antero-posteriorly compressed sacs, separated from each other by intervening septa. Each sac communicates by an inner duct with the œsophagus, the water being expelled by an outer duct. In Bdellostoma each outer duct has a separate opening, but in Myxine all the outer ducts pass outwards by one common gill-opening on each side. In the Lampreys the ducts are short, the outer ones having separate openings (Fig. [2], p. 39). The inner ducts lead into a single diverticulum or bronchus, blind behind, situated below the œsophagus, and communicating in front with the pharynx, where it is provided with two valves by which the regurgitation of the water into the buccal cavity is prevented.
The same type of branchial organs persists in Chondropterygians, which possess five, rarely six or seven, flattened pouches with transversely plaited walls. The septa between them are supported by cartilaginous filaments rising from the hyoidean and branchial arches. Each pouch opens by a cleft outwards, and by an aperture into the pharynx, without intervening ducts. The anterior wall of the first pouch is supported by the hyoidean arch. Between the posterior wall of the first and the anterior of the second sac, and between the adjacent walls of the succeeding, a branchial arch with its two series of radiating cartilaginous filaments is interposed. Consequently the first and last pouch have one set of gill-laminæ only, viz. the first on its posterior and the last on its anterior wall. The so-called spiracles on the upper surface of the head of Chondropterygians are to be referred to in connection with the respiratory organs. They are the external openings of a canal leading on each side into the pharynx, and situated generally close to and behind the orbit. They frequently possess valves or an irregularly indented margin, and are found in all species during the embryonic stage, but remaining persistent in a part only. The spiracles are the remains of the first visceral cleft of the embryo, and in the fœtal state long branchial filaments have been observed to protrude, as from the other branchial clefts.