FOOTNOTES:

Fig. 62.—Polypterus congicus, aCrossopterygianfish from the Congo River. Young, with external gills. (After Boulenger.)

"I have, therefore, to suggest that the more ancient Gnathostomata possessed a series of potentially motor, potentially supporting structures projecting from their visceral arches; it was inherently extremely probable that these should be made use of when actual supporting, and motor appendages had to be developed in connection with clambering about a solid substratum. If this had been so, we should look upon the limb as a modified external gill; the limb-girdle, with Gegenbaur, as a modified branchial arch.

"This theory of the vertebrate paired limb seems to me, I confess, to be a more plausible one on the face of it than either of the two which at present hold the field. If untrue, it is so dangerously plausible as to surely deserve more consideration than it appears to have had. One of the main differences between it and the other two hypotheses is that, instead of derivingthe swimming-fin from the walking and supporting limb, it goes the other way about. That this is the safer line to take seems to me to be shown by the consideration that a very small and rudimentary limb couldonlybe of use if provided with a fixedpoint d'appui. Also on this view, the pentadactyle limb and the swimming-fin would probably be evolved independently from a simple form of limb. This would evade the great difficulties which have beset those who have endeavored to establish the homologies of the elements of the pentadactyle limb with those of any type of fully formed fin."

Uncertain Conclusions.—In conclusion we may say that the evidence of embryology in this matter is inadequate, though possibly favoring on the whole the fin-fold theory; that of morphology is inconclusive, and probably the final answer may be given by paleontology. If the records of the rocks were complete, they would be decisive. At present we have to decide which is the more primitive of two forms of pectoral fin actually known among fossils. That ofCladoselacheis a low, horizontal fold of skin, with feeble rays, called by Copeptychopterygium. That ofPleuracanthusis a jointed paddle-shaped appendage with a fringe of rays on either side. In the theory of Gegenbaur and KerrPleuracanthusmust be, so far as the limbs are concerned, the form nearest the primitive limb-bearing vertebrate. In Balfour's theoryCladoselacheis nearest the primitive type from which the other and with it the archipterygium of later forms may be derived.

Boulenger and others question even this, believing that the archipterygium inPleuracanthusand other primitive sharks and that inNeoceratodusand its Dipnoan and Crossopterygian allies and ancestors have been derived independently, not the latter from the former. In this view there is no real homology between the archipterygium in the sharks possessing it and that in theDipnoansandCrossopterygians. In the one theory the type ofPleuracanthuswould be ancestral to the other sharks on the one hand, and to Crossopterygians and all higher vertebrates on the other. With the theory of the origin of the pectoral from a lateral fold,Pleuracanthuswould be merely a curious specialized offshoot from the primitive sharks, without descendants and without special significance in phylogeny.

As elements bearing on this decision we may note that the tapering unspecialized diphycercal tail ofPleuracanthusseems very primitive in comparison with the short heterocercal tail ofCladoselache. This evidence, perhaps deceptive, is balanced by the presence on the head ofPleuracanthusof a highly specialized serrated spine, evidence of a far from primitive structure. Certainly neither the one genus nor the other actually represents the primitive shark. But asCladoselacheappears in geological time, long beforePleuracanthus,Cladodus, or any other shark with a jointed, archipterygial fin, the burden of proof, according to Dean, rests with the followers of Gegenbaur. If the remains found in the Ordovician at Cañon City referred to Crossopterygians are correctly interpreted, we must regard the shark ancestry as lost in pre-Silurian darkness, for in sharks of some sort the Crossopterygians apparently must find their remote ancestry.

Fig. 63.—Heterocercal tail of Sturgeon,Acipenser sturio(Linnæus). (After Zittel.)

Forms of the Tail in Fishes.—In the process of development the median or vertical fins are, as above stated, older than the paired fins or limbs, whatever be the origin of the latter. They arise in a dermal keel, its membranes fitting and accentuating the undulatory motion of the body.

In this elementary fin-fold slender supports (actinotrichia), the rudiments of fin-rays, appear at intervals. In those fins of most service in the movement of the fish, the fin-rays are strengthened, and their basal supports specialized.

Dean calls attention to the fact that in fishes which swim,when adult, by an undulatory motion, the paired fins tend to disappear, as in the eel and in all eel-like fishes, as blennies and eel-pouts.

The form of the tail at the base of the caudal fin varies in the different groups. In most primitive types, as in most embryonic fishes, the vertebræ grow smaller to the last (diphycercal). In others, also primitive, the end of the tail is directed upward, and the most of the caudal fin is below it. Such a tail is seen in most sharks, in the sturgeon, garpike, bowfin, and in the Ganoid fishes. It is known as heterocercal, and finally in ordinary fishes the tail becomes homocercal or fan-shaped, although usually some trace of the heterocercal condition is traceable, gradually growing less with the process of development.

Since Professor Agassiz first recognized, in 1833, the distinction between the heterocercal and homocercal tail, this matter has been the subject of elaborate investigation and a number of additional terms have been proposed, some of which are in common use.

A detailed discussion of these is found in a paper by Dr. John A. Ryder "On the Origin of Heterocercy" in the Report of the U. S. Fish Commissioner for 1884. In this paper a dynamic or mechanical theory of the causes of change of form is set forth, parts of this having a hypothetical and somewhat uncertain basis.

Dr. Ryder proposes the namearchicercalto denote the cylindroidal worm-like caudal end of the larva of fishes and amphibians before they acquire median fin-folds. The termlophocercalis proposed by Ryder for the form of caudal fin which consists of a rayless fold of skin continuous with the skin of the tail, the inner surfaces of this fold being more or less nearly in contact. To the same type of tail Dr. Jeffries Wyman in 1864 gave the nameprotocercal. This name was used for the tail of the larval ray when it acquires median fin-folds. The term implies, what cannot be far from true, that this form of tail is the first in the stages of evolution of the caudal fin.

To the same type of tail Mr. Alexander Agassiz gave, in 1877, the name ofleptocardial, on the supposition that it represented the adult condition of the lancelet. In this creature,however, rudimentary basal rays are present, a condition differing from that of the early embryos.

The diphycercal tail, as usually understood, is one in which the end of the vertebral column bears "not only hypural but also epural intermediary pieces which support rays." The term is used for the primitive type of tail in which the vertebræ, lying horizontally, grow progressively smaller, as inNeoceratodus,Protopterus, and other Dipnoans and Crossopterygians. The term was first applied by McCoy to the tails of the Dipnoan generaDiplopterusandGyroptychius, and for tails of this type it should be reserved.

Fig. 64.—Heterocercal tail of Bowfin,Amia calva(Linnæus). (After Zittel.)

Fig. 65.—Heterocercal tail of Garpike,Lepisosteus osseus(Linnæus).

The heterocercal tail is one in which the hindmost vertebræ are bent upwards. The term is generally applied to those fishes only in which this bending is considerable and is externally evident, as in the sharks and Ganoids. The character disappears by degrees, changing sometimes to diphycercal or leptocercal by a process of degeneration, or in ordinary fishes becominghomocercal. Dr. Ryder uses the term heterocercal for all cases in which any up-bending of the axis takes place, even though it involves the modification of but a single vertebra. With this definition, the tail of salmon, herring, and even of most bony fishes would be considered heterocercal, and most or all of these pass through a heterocercal stage in the course of development. The term is, however, usually restricted to those forms in which the curving of the axis is evident without dissection.

Fig. 66.—Coryphænoides carapinus(Goode and Bean), showing leptocercal tail. Gulf Stream.

The homocercal tail is the fan-shaped or symmetrical tail common among the Teleosts, or bony fishes. In its process of development the individual tail is first archicercal, then lophocercal, then diphycercal, then heterocercal, and lastly homocercal. A similar order is indicated by the sequence of fossil fishes in the rocks, although some forms of diphycercal tail may be produced by degeneration of the heterocercal tail, as suggested by Dr. Dollo and Dr. Boulenger, who divide diphycercal tails into primitive and secondary.

The peculiar tapering tail of the cod, the vertebræ growing progressively smaller behind, is termedisocercalby Professor Cope. This form differs little from diphycercal, except in its supposed derivation from the homocercal type. A similar form is seen in eels.

Fig. 67.—Heterocercal tail of Young Trout,Salmo fario(Linnæus). (After Parker and Haswell.)

The termleptocercalhas been suggested by Gaudry, 1883, for those tails in which the vertebral column ends in a point. We may, perhaps, use it for all such as are attenuate, ending in a long point or whip, as in theMacrouridæ, or grenadiers, the sting-rays, and in various degenerate members of almost every large group.

The termgephyrocercalis devised by Ryder for fishes in which the end of the vertebral axis is aborted in the adult, leaving the caudal elements to be inserted on the end of this axis, thus bridging over the interval between the vertical fins,as the name (γεφύρος, bridge; κέρκος, tail) is intended to indicate. Such a tail has been recognized in four genera only,Mola,Ranzania,Fierasfer, andEchiodon, the head-fishes and the pearl-fishes.

Fig. 68.—Isocercal tail of Hake,Merluccius productus(Ayres).

Fig. 69.—Homocercal tail of a Flounder,Paralichthys californicus.

The part of the body of the fish which lies behind the vent is known as the urosome. The urostyle is the name given to a modified bony structure, originally the end of the notochord, turned upward in most fishes. The termopisthureis suggested by Ryder for the exserted tip of the vertebral column, which in some larvæ (Lepisosteus) and in some adult fishes (Fistularia,Chimæra) projects beyond the caudal fin. The urosome, or posterior part of the body, must be regarded as a product of evolution and specialization, its function being largely that of locomotion. In the theoretically primitive fish there is no urosome, the alimentary canal, as in the worm, beginning at one end of the body and terminating at the other.

Fig. 70.—Gephyrocercal tail ofMola mola(Linnæus). (After Ryder.)

Homologies of the Pectoral Limb.—Dr. Gill has made an elaborate attempt to work out the homologies of the bones of the pectoral limb.[4]From his thesis we take the following:

"The following are assumed as premises that will be granted by all zootomists:

"1. Homologies of parts are best determinable,ceteris paribus, in the most nearly related forms.

"2. Identification should proceed from a central or determinate point outwards.

"The applications of these principles are embodied in the following conclusions:

"1. The forms that are best comparable and that are most nearly related to each other are the Dipnoi, an order of fishes at present represented byLepidosiren,Protopterus, andCeratodus,and the Batrachians as represented by theGanocephala, Salamanders, and Salamander-like animals.

"2. The articulation of the anterior member with the shoulder-girdle forms the most obvious and determinable point for comparison in the representatives of the respective classes.

Fig. 71.—Shoulder-girdle ofAmia calva(Linnæus).

Fig. 72.—Shoulder-girdle of a Sea Catfish,Selenaspis dowi.

The Girdle in Dipnoans.—"The proximal element of the anterior limb in the Dipnoi has almost by common consent been regarded as homologous with thehumerusof the higher vertebrates.

"The humerus of Urodele Batrachians, as well as the extinct Ganocephala and Labyrinthodontia, is articulated chiefly with the coracoid. Therefore the element of the shoulder-girdle with which the humerus of the Dipnoi is articulated must also be regarded as thecoracoid(subject to the proviso hereinafter stated), unless some specific evidence can be shown to the contrary. No such evidence has been produced.

"The scapula in the Urodele and other Batrachians is entirely or almost wholly excluded from the glenoid foramen, and abovethe coracoid. Therefore the corresponding element in Dipnoi must be thescapula.

"The other elements must be determined by their relation to the preceding, or to those parts from or in connection with which they originate. All those elements inimmediateconnection with the pectoral fin and the scapula must be homologous as a whole with the coraco-scapular plate of the Batrachians; that is, it is infinitely more probable that they represent, as a whole or as dismemberments therefrom, the coraco-scapular element than that they independently originated. But the homogeneity of that coraco-scapular element forbids the identification of the several elements of the fish's shoulder-girdle with regions of the Batrachian's coraco-scapular plate.

Fig. 73.—Clavicles of a Sea Catfish,Selenaspis dowi(Gill).

"And it is equally impossible to identify the fish's elements with those of the higher reptiles or other vertebrates which have developed from the Batrachians. The elements in the shoulder-girdles of the distantly separated classesmaybe (to use the terms introduced by Dr. Lankester) homoplastic, but theyare nothomogenetic. Therefore they must be named accordingly. The element of the Dipnoan's shoulder-girdle, continuous downward from the scapula, and to which the coracoid is closely applied, may be namedectocoracoid.

"Neither the scapula in Batrachians nor the cartilaginous extension thereof, designated suprascapula, is dissevered from the coracoid. Therefore there is ana prioriimprobabilityagainst the homology with the scapula of any part having a distant and merely ligamentous connection with the humerus-bearing element. Consequently, as an element better representing the scapula exists, the element named scapula (by Owen, Günther, etc.) cannot be the homologue of the scapula of Batrachians. On the other hand, its more intimate relations with the skull and the mode of development indicate that it is rather an element originating and developed in more intimate connection with the skull. It may therefore be considered, with Parker, as apost-temporal.

"The shoulder-girdle in the Dipnoi is connected by an azygous differentiated cartilage, swollen backwards. It is more probable that this is the homologue of thesternumof Batrachians, and that in the latter that element has been still more differentiated and specialized than that it should have originatedde novofrom an independently developed nucleus."

Fig. 74.—Shoulder-girdle of a Batfish,Ogcocephalus radiatus(Mitchill).

The Girdle in Fishes Other than Dipnoans.—"Proceeding from the basis now obtained, a comparative examination of other types of fishes successively removed by their affinities from the Lepidosirenids may be instituted.

"With the humerus of the Dipnoans, the element of the Polypterids (single at the base, but immediately divaricating and with its limbs bordering an intervening cartilage which supports the pectoral and its basilar ossicles) must be homologous. But it is evident that the external elements of theso-called carpus of the teleosteoid Ganoids are homologous with that element in Polypterids. Therefore those elements cannot be carpal, but must represent the humerus.

Fig. 75.—Shoulder-girdle of a Threadfin,Polydactylus approximans(Lay and Bennett).

"The element with which the homologue of the humerus, in Polypterids, is articulated must be homologous with the analogous element in Dipnoans, and therefore with thecoracoid. The coracoid of Polypterids is also evidently homologous with the corresponding element in the other Ganoids, and the latter consequently must be alsocoracoid. It is equally evident, after a detailed comparison, that the single coracoid element of the Ganoids represents the three elements developed in the generalized Teleosts (Cyprinids, etc.) in connection with the basis of the pectoral fin, and, such being the case, the nomenclature should correspond. Therefore the upper element may be namedhypercoracoid; the lower,hypocoracoid; and the transverse or median,mesocoracoid.

"The two elements of the arch named by Parker, inLepidosiren, 'supraclavicle' (scapula) and 'clavicle' (ectocoracoid) seem to be comparable together, and as a whole, with the single element carrying the humerus and pectoral fin in the Crossopterygians (PolypterusandCalamoichthys) and other fishes, and therefore not identical respectively with the 'supraclavicle' and 'clavicle' (except in part) recognized by him in other fishes. As this compound bone, composed of the scapula and ectocoracoid fused together, has received no name which is not ambiguous or deceptive in its homologous allusions, it may be designated asproscapula.

"The post-temporal of the Dipnoans is evidently represented by the analogous element in the Ganoids generally, as well as in the typical fishes. The succeeding elements (outside those already alluded to) appear from their relations to be developed from or in connection with the post-temporal, and not from the true scapular apparatus; they may therefore be namedpost-temporal,posterotemporal, andteleo-temporal. It will be thus seen that the determinations here adopted depend mainly (1) on the interpretation of the homologies of the elements with which the pectoral limbs are articulated, and (2) on the application of the term 'coracoid.' The name 'coracoid,' originally applied to the process so called in the human scapula and subsequently extended to the independent element homologous with it in birds and other vertebrates, has been more especially retained (e.g., by Parker in mammals, etc.) for the region including the glenoid cavity. On the assumption that this may be preferred by some zootomists, the preceding terms have been applied. But if the name should be restricted to the proximal element, nearest the glenoid cavity, in which ossification commences, the nameparaglenalgiven by Dugès to the cartilaginous glenoid region can be adopted, and the coracoid would then be represented (in part) rather by the element so named by Owen. That eminent anatomist, however, reached his conclusion (only in part the same as that here adopted) by an entirely different course of reasoning, and by a process, as it may be called, of elimination; that is, recognizing first the so-called 'radius' and 'ulna,' the 'humerus,' the 'scapula,' and the 'coracoid' were successively identified from their relations to the elements thus determined and because they were numerically similar to the homonymous parts among higher vertebrates."

FOOTNOTES:[4]Catalogue of the Families of Fishes, 1872.

[4]Catalogue of the Families of Fishes, 1872.

HowFishes Breathe.—The fish breathes the air which is dissolved in water. It cannot use the oxygen which is a component part of water, nor can it, as a rule, make use of atmospheric air. The amount of oxygen required for the low vegetative processes of the fish is comparatively small. According to Dr. Günther, a man consumes 50,000 times as much oxygen as a tench. But some fishes demand more oxygen than others. Some, like the catfish or the loach, will survive long out of water, while others die almost instantly if removed from their element or if the water is allowed to become foul. In most cases the temperature of the blood of the fish is but little above that of the water in which they live, but in the mackerel and other muscular fishes the temperature of the body may be somewhat higher.

Some fishes which live in mud, especially in places which become dry in summer, have special contrivances by which they can make use of atmospheric air. In a few primitive fishes (Dipnoans, Crossopterygians, Ganoids) the air-bladder retains its original function of a lung. In other cases some peculiar structure exists in connection with the gills. Such a contrivance for holding water above the gills is seen in the climbing perch of India (Anabas scandens) and other members of the group called Labyrinthici.

In respiration, in fishes generally, the water is swallowed through the mouth and allowed to pass out through the gill-openings, thus bathing the gills. In a few of the lower types a breathing-pore takes the place of the gill-openings.

The gills, or branchiæ, are primarily folds of the skin lining the branchial cavity. In most fishes they form fleshy fringes or laminæ throughout which the capillaries are distributed. In the embryos of sharks, skates, chimæras, lung-fishes,and Crossopterygians external gills are developed, but in the more specialized forms these do not appear outside the gill-cavity. In some of the sharks, and especially the rays, a spiracle or open foramen remains behind the eye. Through this spiracle, leading from the outside into the cavity of the mouth, water is drawn downwards to pass outward over the gills. The presence of this breathing-hole permits these animals to lie on the bottom without danger of inhaling sand.

Fig. 76.—Gill-basket of Lamprey. (After Dean.)

The Gill-structures.—The three main types of gills among fishes are the following: (a) the purse-shaped gills found in the hagfishes and lampreys, known as a class as Marsipobranchs, or purse-gills. These have a number (5 to 12) of sac-like depressions on the side of the body, lined with gill-fringes and capillaries, the whole supported by an elaborate branchial basket formed of cartilage. (b) The plate-gills, found among the sharks, rays, and chimæras, thence called Elasmobranchs, or plate-gills. In these the gill-structures are flat laminæ, attached by one side to the gill-arches. (c) The fringe-gills found in ordinary fishes, in which the gill-filaments containing the capillaries are attached in two rows to the outer edge of each gill-arch. The so-called tuft-gills (Lophobranchs) of the sea-horse and pipefish are like these in structure, but the filaments are long, while the arches are very short. In most of the higher fishes a small accessory gill (pseudobranchia) is developed in the skin of the inner side of the opercle.

The Air-bladder.—The air-bladder, or swim-bladder, must be classed among the organs of respiration, although in the higher fishes its functions in this regard are rudimentary, and in some cases it has taken collateral functions (as a hydrostaticorgan of equilibrium, or perhaps as an organ of hearing) which have no relation to its original purpose.

Fig. 77.—Weberian apparatus and air-bladder of Carp. (From Günther, after Weber.)

The air-bladder is an internal sac possessed by many fishes, but not by all. It lies in the dorsal part of the abdominal cavity above the intestines and below the kidneys. In some cases it is closely adherent to the surrounding tissues. In others it is almost entirely free, lying almost loose in the cavity of the body. In some cases it is enclosed in a bony capsule. In the allies of the carp and catfish, which form the majority of fresh-water fishes, its anterior end is connected through a chain of modified vertebræ to the ear. Sometimes its posterior end fits into an enlarged and hollow interhæmal bone. Sometimes, again, a mass of muscle lies in front of it or is otherwise attached to it. Sometimes it is divided into two or three parts by crosswise constrictions. Sometimes it is constricted longitudinally, and at other times it has attached to it a complication of supplemental tubes of the same character as the air-bladder itself. In still other cases it is divided by many internal partitions into a cellular body, similar to the lung of the higher vertebrates, though the cells are coarser and less intricate. This condition is evidently more primitive than that of the empty sac.

The homology of the air-bladder with the lung is evident. This is often expressed in the phrase that the lung is a developed air-bladder. This is by no means true. To say that the air-bladder is a modified and degenerate lung is much nearer thetruth, although we should express the fact more exactly to say that both air-bladder and lung are developed from a primitive cellular breathing-sac, originally a diverticulum from the ventral walls of the œsophagus.

The air-bladder varies in size as much as in form. In some fishes it extends from the head to the tail, while in others it is so minute as to be scarcely traceable. It often varies greatly in closely related species. The common mackerel (Scomber scombrus) has no air-bladder, while in the closely related colias or chub mackerel (Scomber japonicus) the organ is very evident. In other families, as the rockfishes (Scorpænidæ), genera with and those without the air-bladder are scarcely distinguishable externally. In general, fishes which lie on the bottom, those which inhabit great depths, and those which swim freely in the open sea, as sharks and mackerel, lack the air-bladder. In the sharks, rays, and chimæras there is no trace of an air-bladder. In the mackerel and other bony fishes without it, it is lost in the process of development.

The air-bladder is composed of two layers of membrane, the outer one shining, silvery in color, with muscular fibres, the inner well supplied by blood-vessels. The gas within the air-bladder must be in most cases secreted from the blood-vessels. In river fishes it is said to be nearly pure nitrogen. In marine fishes it is mostly oxygen, with from 6 to 10 per cent of carbonic-acid gas, while in the deep-sea fishes oxygen is greatly in excess. InLopholatilus, a deep-sea fish, Professor R. W. Tower finds 66 to 69 per cent of oxygen. InTrigla lyraBiot records 87 per cent. InDentex dentex, a shore fish of Europe, 40 per cent of oxygen was found in the air-bladder. Fifty per cent is recorded from the European porgy,Pagrus pagrus. In a fish dying from suffocation the amount of carbonic-acid gas (CO2) is greatly increased, amounting, according to recent researches of Professor Tower on the weak-fish,Cynoscion regalis, to 24 to 29 per cent. This shows conclusively that the air-bladder is to some degree a reservoir of oxygen secreted from the blood, to which channel it may return through a kind of respiration.

The other functions of the air-bladder have been subject to much question and are still far from understood. The following summary of the various views in this regard we copyfrom Professor Tower's paper on "The Gas in the Swim-bladder of Fishes":

"The function of the swim-bladder of fishes has attracted the attention of scientists for many centuries. The rôle that this structure plays in the life of the animal has been interpreted in almost as many ways as there have been investigators, and even now there is apparently much doubt as to the true functions of the swim-bladder. Consequently any additional data concerning this organ are of immediate scientific value.

"Aristotle, writing about the noises made by fishes, states that 'some produce it by rubbing the gill-arches ...; others by means of the air-bladder. Each of these fishes contains air, by rubbing and moving of which the noise is produced.' The bladder is thus considered a sound-producing organ, and it is probable that he arrived at this result by his own investigations.

"Borelli (De Motu Animalium, 1680) attributed to the air-bladder a hydrostatic function which enabled the fish to rise and fall in the water by simply distending or compressing the air-bladder. This hypothesis, which gives to the fish a volitional control over the air-bladder—it being able to compress or distend the bladder at pleasure—has prevailed, to a greater or less degree, from the time of Borelli to the present. To my knowledge, however, there are no investigations which warrant such a theory, while, on the other hand, there are many facts, as shown by Moreau's experiment, which distinctly contradict this belief. Delaroche (Annales du Mus. d'Hist. Nat., tome XIV, 1807-1809) decidedly opposed the ideas of Borelli, and yet advanced an hypothesis similar to it in many respects. Like Borelli, he said that the fish could compress or dilate the bladder by means of certain muscles, but this was to enable the fish to keep the same specific gravity as the surrounding medium, and thus be able to remain at any desired depth (and not to rise or sink). This was also disproved later by Moreau. Delaroche proved that there existed a constant exchange between the air in the air-bladder and the air in the blood, although he did not consider the swim-bladder an organ of respiration.

"Biot (1807), Provençal and Humboldt (1809), and others made chemical analyses of the gas in the swim-bladder, andfound 1 to 5 per cent of CO2, 1 to 87 per cent of O2, and the remainder nitrogen. The most remarkable fact discovered about this mixture was that it frequently consisted almost entirely of oxygen, the per cent of oxygen increasing with the depth of the water inhabited by the fish. The reasons for this phenomenon have never been satisfactorily explained.

"In 1820 Weber described a series of paired ossicles which he erroneously called stapes, malleus, and incus, and which connected the air-bladder in certain fishes with a part of the ear—the atrium sinus imparis. Weber considered the swim-bladder to be an organ by which sounds striking the body from the outside are intensified, and these sounds are then transmitted to the ear by means of the ossicles. The entire apparatus would thus function as an organ of hearing. Weber's views remained practically uncontested for half a century, but recently much has been written both for and against this theory. Whatever the virtues of the case may be, there is certainly an inviting field for further physiological investigations regarding this subject, and more especially on the phenomena of hearing in fishes.

"Twenty years later Johannes Müller described, in certain Siluroid fishes, a mechanism, the so-called 'elastic-spring' apparatus, attached to the anterior portion of the air-bladder, which served to aid the fish in rising and sinking in the water according as the muscles of this apparatus were relaxed or contracted to a greater or lesser degree. This interpretation of the function of the 'elastic-spring' mechanism was shown by Sörensen to be untenable. Müller also stated that in some fish, at least, there was an exchange of gas between blood and air-bladder—the latter having a respiratory function—and regarded the gas in the air-bladder as the result of active secretion. InMalapterurus(Torpedo electricus) he stated that it is a sound-producing organ.

"Hasse, in 1873, published the results of his investigations on the functions of the ossicles of Weber, stating that their action was that of a manometer, acquainting the animal with the degree of pressure that is exerted by the gases in the air-bladder against its walls. This pressure necessarily varies with the different depths of water which the fish occupies. Hassedid not agree with Weber that the ear is affected by the movements of these ossicles.

"One year later Dufosse described in some fishes an air-bladder provided with extrinsic muscles by whose vibration sound was produced, the sound being intensified by the air-bladder, which acted as a resonator. He also believed that certain species produced a noise by forcing the gas from the air-bladder through a pneumatic duct.

"At about the same time Moreau published his classical work on the functions of the air-bladder. He proved by ingenious experiments that many of the prevailing ideas about the action of the air-bladder were erroneous, and that this organ serves to equilibrate the body of the fish with the water at any level. This is not accomplished quickly, but only after sufficient time for the air in the bladder to become adjusted to the increase or decrease in external pressure that has taken place. The fish, therefore, makes no use of any muscles in regulating the volume of its air-bladder. The animal can accommodate itself only gradually to considerable changes in depth of water, but can live equally comfortably at different depths, provided that the change has been gradual enough. Moreau's experiments also convinced him that the gas is actually secreted into the air-bladder, and that there is a constant exchange of gas between it and the blood. In these investigations he has also noticed that section of the sympathetic-nerve fibres supplying the walls of the air-bladder hastens the secreting of the gas into the empty bladder. Since then Bohr has shown that section of the vagus nerve causes the secretion to cease. Moreau noticed in one fish (Trigla) having an air-bladder supplied with muscles that the latter served to make the air-bladder produce sound.

"Again, in 1885, the Weberian mechanism was brought to our attention with a new function attributed to it by Sagemehl who stated that this mechanism exists not for any auditory purposes, nor to tell the fish at what level of the water it is swimming, but to indicate to the fish the variations in the atmospheric pressure. Sörensen tersely contrasts the views of Hasse and Sagemehl by saying that 'Hasse considers the air-bladder with the Weberian mechanism as a manometer; Sagemehl regards it as a barometer.' The theory of Sagemehl has, naturallyenough, met with little favor. Sörensen (1895) held that there is but little evidence for attributing to the air-bladder the function of a lung. It is to be remembered, however, that, according to Sörensen's criterion no matter what exchange of gases takes place between blood and air-bladder, it cannot be considered an organ of respiration, 'unless its air is renewed by mechanical respiration.'

"Sörensen also refutes, from anatomical and experimental grounds, the many objections to Weber's theory of the function of the ossicles. He would thus attribute to the air-bladder the function of hearing; indeed in certain species the only reason for the survival of the air-bladder is that 'the organ is still of acoustic importance; that it acts as a resonator.' This idea, Sörensen states, is borne out by the anatomical structure found inMisgurnusandChlarias, which resembles the celebrated 'Colladon resonator.' This author attributes to the air-bladder with its 'elastic spring' and various muscular mechanisms the production of sound as its chief function."

Origin of the Air-bladder.—In the more primitive forms, and probably in the embryos of all species, the air-bladder is joined to the œsophagus by an air-duct. This duct is lost entirely in the adult of all or nearly all of the thoracic and jugular fishes, and in some of the abdominal forms. The lancelets, lampreys, sharks, rays, and chimæras have no air-bladder, but in the most primitive forms of true fishes (Dipnoans and Crossopterygians), having the air-bladder cellular or lung-like, the duct is well developed, freely admitting the external air which the fish may rise to the surface to swallow. In most fishes the duct opens into the œsophagus from the dorsal side, but in the more primitive forms it enters from the ventral side, like the windpipe of the higher vertebrates. In some of the Dipnoans the air-bladder divides into two parts, in further resemblance to the true lungs.

The Origin of the Lungs.—The following account of the function of the air-bladder and of its development and decline is condensed from an article by Mr. Charles Morris:[5]

"If now we seek to discover the original purpose of thisorgan, there is abundant reason to believe that it had nothing to do with swimming. Certainly the great family of the sharks, which have no bladder, are at no disadvantage in changing their depth or position in the water. Yet if the bladder is necessary to any fish as an aid in swimming, why not to all? And if this were its primary purpose, how shall we explain its remarkable variability? No animal organ with a function of essential importance presents such extraordinary modifications in related species and genera. In the heart, brain, and other organs there is one shape, position, and condition of greatest efficiency, and throughout the lower forms we find a steady advance towards this condition. Great variation, on the other hand, usually indicates that the organ is of little functional importance, or that it has lost its original function. Such we conceive to be the case with the air-bladder. The fact of its absence from some and its presence in other fishes of closely related species goes far to prove that it is a degenerating organ; and the same is shown by the fact that it is useless in some species for the purpose to which it is applied in others. That it had, at some time in the past, a function of essential importance there can be no question. That it exists at all is proof of this. But its modern variations strongly indicate that it has lost this function and is on the road towards extinction. Larval conditions show that it had originally a pneumatic duct as one of its essential parts, but this has in most cases disappeared. The bladder itself has in many cases partly or wholly disappeared. Where preserved, it seems to be through its utility for some secondary purpose, such as an aid in swimming or in hearing. That its evolution began very long ago there can be no question; and the indications are that it began long ago to degenerate, through the loss of its primitive function.

"What was this primitive function? In attempting to answer this question we must first consider the air-bladder in relation to the fish tribe as a whole. No shark or ray possesses the air-bladder. In some few sharks, indeed, there is a diverticulum of the pharynx which may be a rudimentary approach to the air-bladder; but this is very questionable. The conditions of its occurrence in the main body of modern fishes, the Teleostean, we have already considered. But in the most ancient living ordersof fishes it exists in an interesting condition. In every modern Dipnoan, Crossopterygian, and Ganoid the air-bladder has an effective pneumatic duct. This in the Ganoids opens into the dorsal side of the œsophagus, but in the Dipnoans and Crossopterygians, like the windpipe of lung-breathers, it opens into the ventral side. In the Dipnoans, also survivors from the remote past, the duct not only opens ventrally into the œsophagus, but the air-bladder does duty as a lung. Externally it differs in no particular from an air-bladder; but internally it presents a cellular structure which nearly approaches that of the lung of the batrachians. There are three existing representatives of the Dipnoans. One of these, the Australian lung-fish (Neoceratodus) has a single bladder, which, however, is provided with breathing-pouches having a symmetrical lateral arrangement. It has no pulmonary artery, but receives branches from thearteria cœliaca. In the other two forms,LepidosirenandProtopterus, the kindred 'mudfishes' of the Amazon basin and tropical Africa, the bladder or lung is divided into two lateral chambers, as in the land animals, and is provided with a separate pulmonary artery.

"The opinion seems to have been tacitly entertained by physiologists that this employment of the air-bladder by the Dipnoans as a lung is a secondary adaptation, a side issue from its original purpose. It is more likely that this is the original purpose, and that its degeneration is due to the disappearance of the necessity of such a function. As regards the gravitative employment of the bladder, the Teleostean fishes, to which this function is confined, are of comparatively modern origin; while the Dipnoans are surviving representatives of a very ancient order of fishes, which flourished in the Devonian age of geology, and in all probability breathed air then as now; and the Crossopterygians and Ganoids, which approach them in this particular, are similarly ancient in origin, and were the ancestors of the Teleosteans. The natural presumption, therefore, is that the duty which it subserved in the most ancient fishes was its primitive function.

"The facts of embryology lend strong support to this hypothesis. For the air-bladder is found to arise in a manner very similar to the development of the lung. They each begin as anoutgrowth from the fore part of the alimentary tract, the only difference being that the air-bladder usually rises dorsally and the lung ventrally. The fact already cited, that the pneumatic duct is always present in the larval form in fishes that possess a bladder, is equally significant. All the facts go to show that the introduction of external air into the body was a former function of the air-bladder, and that the atrophy of the duct in many cases, and the disappearance of the bladder in others, are results of the loss of this function.

"Such an elaborate arrangement for the introduction of air into the body could have, if we may judge from analogy, but one purpose, that of breathing, to which purpose the muscular and other apparatus for compressing and dilating the bladder, now seemingly adapted to gravitative uses, may have been originally applied. The same may be said of the great development of blood-capillaries in the inner tunic of the bladder. These may now be used only for the secretion of gas into its interior, but were perhaps originally employed in the respiratory secretion of oxygen. In fact all the circumstances mentioned—the similarity in larval development between the bladder and lung, the larval existence of the pneumatic duct, the arrangements for compressing and dilating the bladder, and the capillary vessels on its inner tunic—point to the breathing of air as its original purpose.

"It is probable that the Ganoid, as well as the Dipnoan, air-bladder is to some extent still used in breathing. The Dipnoans have both lungs and gills, and probably breathe with the latter in ordinary cases, but use their lungs when the inland waters in which they live become thick and muddy, or are charged with gases from decomposing organic matter. The Ganoid fishes to some extent breathe the air. InPolypterusthe air-bladder resembles the Dipnoan lung in having lateral divisions and a ventral connection with the œsophagus, while inLepisosteus(the American garpike) it is cellular and lung-like. This fish keeps near the surface, and may be seen to emit air-bubbles, probably taking in a fresh supply of air. The American bowfin, or mudfish (Amia), has a bladder of the same lung-like character, and has been seen to come to the surface, open its jaws widely, and apparently swallow a large quantity of air. Heconsiders that bothLepisosteusandAmiainhale and exhale air at somewhat regular intervals, resembling in this the salamanders and tadpoles, 'which, as the gills shrink and the lungs increase, come more frequently to the surface for air.'

"As the facts stand there is no evident line of demarcation between the gas-containing bladders of many of the Teleosteans, the air-containing bladders of the others and the Ganoids, and the lung of the Dipnoans, and the indications are in favor of their having originally had the same function, and of this being the breathing of air.

"If now we ask what were the conditions of life under which this organ was developed, and what the later conditions which rendered it of no utility as a lung, some definite answer may be given. The question takes us back to the Devonian and Silurian geological periods, during which the original development of the bladder probably took place. In this era the seas were thronged with fishes of several classes, the Elasmobranchs among others, followed by the Dipnoi and Crossopterygians. The sharks were without, the Dipnoans and Crossopterygians doubtless with, an air-bladder—a difference in organization which was most likely due to some marked difference in their life-habits. The Elasmobranchs were the monarchs of the seas, against whose incursions the others put on a thick protective armor, and probably sought the shallow shore waters, while their foes held chief possession of the deeper waters without.

"We seem, then, to perceive the lung-bearing fishes, driven by their foes into bays and estuaries, and the waters of shallow coasts, ascending streams and dwelling in inland waters. Here two influences probably acted on them. The waters they dwelt in were often thick with sediment, and were doubtless in many instances poorly aerated, rendering gill-breathing difficult. And the land presented conditions likely to serve as a strong inducement to fishes to venture on shore. Its plant-life was abundant, while its only animal inhabitants seem to have been insects, worms, and snails. There can be little doubt that the active fish forms of that period, having no enemies to fear on the land, and much to gain, made active efforts to obtain a share of this vegetable and animal food. Even to-day, when they have numerous foes to fear, many fishes seek food on the shore, andsome even climb trees for this purpose. Under the conditions of the period mentioned there was a powerful inducement for them to assume this habit.

"Such conditions must have strongly tended to induce fishes to breathe the air, and have acted to develop an organ for this purpose. In addition to the influences of foul or muddy water and of visits to land may be named that of the drying-out of pools, by which fishes are sometimes left in the moist mud till the recurrence of rains, or are even buried in the dried mud during the rainless season. This is the case with the modern Dipnoi, which use their lungs under such circumstances. In certain other fresh-water fishes, of the family Ophiocephalidæ, air is breathed while the mud continues soft enough for the fish to come to the surface, but during the dry period the animal remains in a torpid state. These fishes have no lungs, but breathe the air into a simple cavity in the pharynx, whose opening is partly closed by a fold of the mucous membrane. Other Labyrinthici, of similar habits, possess a more developed breathing organ. This is a cavity formed by the walls of the pharynx, in which are thin laminæ, or plates, which undoubtedly perform an oxygenating function. The most interesting member of this family isAnabas scandens, the climbing perch. In this fish, which not only leaves the water, but is said to climb trees, the air-breathing organ is greatly developed. The labyrinthici, moreover, have usually large air-bladders. As regards the occasional breathing of air by fishes, even in species which do not leave the water, it is quite common, particularly among fresh-water species. Cuvier remarks that air is perhaps necessary to every kind of fish; and that, particularly when the atmosphere is warm, most of our lacustrine species sport on the surface for no other purpose.

"It is not difficult to draw a hypothetical plan of the development of the air-bladder as a breathing organ. In the two families of fishes just mentioned, whose air-bladders indicate that they once possessed the air-breathing function and have lost it, we perceive the process of formation of an air-breathing organ beginning over again under stress of similar circumstances. The larval development of the air-bladder points significantly in the same direction. In fact we have strong reason to believe thatair-breathing in fishes was originally performed, as it probably often is now, by the unchanged walls of the œsophagus. Then these walls expanded inwardly, forming a simple cavity, partly closed by a fold of membrane, like that of the Ophiocephalidæ. A step further reduced this membranous fold to a narrow opening, leading to an inner pouch. As the air-breathing function developed, the opening became a tube, and the pouch a simple lung, with compressing muscles and capillary vessels. By a continuation of the process the smooth-walled pouch became sacculated, its surface being increased by folding into breathing cells. Finally, a longitudinal constriction divided it into two lateral pouches, such as we find in the lung of the Dipnoans. This brings us to the verge of the lung of the amphibians, which is but a step in advance, and from that the line of progress is unbroken to the more intricate lung of the higher land animals.

"The dorsal position of the bladder and its duct would be a difficulty in this inquiry, but for the fact that the duct is occasionally ventral. This dorsal position may have arisen from the upward pressure of air in the swimming fish, which would tend to lift the original pouch. But in the case of fishes which made frequent visits to the shore new influences must have come into play. The effect of gravity tended to draw the organ and its duct downward, as we find in the Crossopterygians and in all the Dipnoans, and its increased use in breathing required a more extended surface. Through this requirement came the pouched and cellular lung of the Dipnoans. Of every stage of the process here outlined examples exist, and there is great reason to believe that the development of the lung followed the path above pointed out.

"When the carboniferous era opened there may have been many lung- and gill-breathing fishes which spent much of their time on land, and some of which, by a gradual improvement of their organs of locomotion, changed into batrachians. But with the appearance of the latter, and of their successors, the reptiles, the relations of the fish to the land radically changed. The fin, or the simple locomotor organ, of the Dipnoans could not compete with the leg and foot as organs of land locomotion, and the fish tribe ceased to be lords of the land, where, instead of feeble prey, they now found powerful foes, and were driven back totheir native habitat, the water. Nor did the change end here. In time the waters were invaded by the reptiles, numerous swimming forms appearing, which it is likely were abundant in the shallower shore-line of the ocean, while they sent many representatives far out to sea. These were actively carnivorous, making the fish their prey, the great mass of whom were doubtless driven into the deeper waters, beyond the reach of their air-breathing foes.

"In this change of conditions we seem to perceive an adequate cause for the loss of air-breathing habits in those fishes in which the lung development had not far progressed. It may indeed have been a leading influence in the development of the Teleostean or bony fishes, as it doubtless was in the loss of its primitive function by, and the subsequent changes of, the air-bladder.

"Such of the Crossopterygians and Dipnoans as survived in their old condition had to contend with adverse circumstances. Most of them in time vanished, while their descendants which still exist have lost in great measure their air-breathing powers, and the Dipnoans, in which the development of the lung had gone too far for reversal, have degenerated into eel-like, mud-haunting creatures, in which the organs of locomotion have become converted into the feeble paddle-like limbs of Neoceratodus and the filamentary appendages of the other species.

"As regards the presence of a large quantity of oxygen in the bladders of deep-swimming marine fishes, it not unlikely has a respiratory purpose, the bladder being, as suggested by Semper, used as a reservoir for oxygen, to serve the fish when sleeping, or when, from any cause, not actively breathing. The excess of oxygen is not due to any like excess in the gaseous contents of sea-water, for the percentage of oxygen decreases from the surface downward, while that of nitrogen remains nearly unchanged. In all cases, indeed, the bladder may preserve a share of its old function, and act as an aid in respiration. Speaking of this, Cuvier says: 'With regard to the presumed assistance which the swim-bladder affords in respiration, it is a fact that when a fish is deprived of that organ, the production of carbonic acid by the branchiæ is very trifling,' thus strongly indicating that the bladder still plays a part in the oxygenation of the blood.

"Under the hypothesis here presented the process of evolution involved may be thus summed up. Air-breathing in fishes was originally performed by the unchanged walls of the œsophagus perhaps at specially vascular localities. Then the wall folded inward, and a pouch was finally formed, opening to the air. The pouch next became constricted off, with a duct of connection. Then the pouch became an air-bladder with respiratory function, and finally developed into a simple lung. These air-breathing fishes haunted the shores, their fins becoming converted into limbs suitable for land locomotion, and in time developed into the lung- and gill-breathing batrachia, and these in their turn into the lung-breathing reptilia, the locomotor organs gradually increasing in efficiency. Of these pre-batrachia we have existing representatives in the mud-haunting Dipnoi, with their feeble limbs. In the great majority of the Ganoid fishes the bladder served but a minor purpose as a breathing organ, the gills doing the bulk of the work. In the Teleostean descendants of the Ganoids the respiratory function of the bladder in great measure or wholly ceased, in the majority of cases the duct closing up or disappearing, leaving the pouch as a closed internal sac, far removed from its place of origin. In this condition it served as an aid in swimming, perhaps as a survival of one of its ancient uses. It gained also in certain cases some connection with the organ of hearing. But these were makeshift and unimportant functions, as we may gather from the fact that many fishes found no need for them, the bladder, in these cases, decreasing in size until too small to be of use in swimming, and in other cases completely disappearing after having travelled far from its point of origin. In some other cases, above cited, the process seems to have begun again, in modern times, in an eversion of the wall of the œsophagus for respiratory purposes. The whole process, if I have correctly conceived it, certainly forms a remarkable organic cycle of development and degeneration, which perhaps has no counterpart of similarly striking character in the whole range of organic life."

The Heart of the Fish.—The heart of the fish is simple in structure, small in size, and usually placed far forward, just behind the branchial cavity, and separated from the abdominalcavity by a sort of "diaphragm" formed of thickened peritoneum. In certain eels the heart is remote from the head.

The heart consists of four parts, the sinus venosus, into which the veins enter, the auricle or atrium, the ventricle, and the arterial bulb at the base of the great artery which carries the blood to the gills. Of these parts the ventricle is deepest in color and with thickest walls. The arterial bulb varies greatly in structure, being in the sharks, rays, Ganoids, and Dipnoans muscular and provided with a large number of internal valves, and contracting rhythmically like the ventricle. In the higher fishes these structures are lost, the walls of the arterial bulb are not contractile, and the interior is without valves, except the pair that separate it from the ventricle.

In the lancelet there is no proper heart, the function of the heart being taken by a contractile blood-vessel situated on the ventral side of the alimentary canal. In the Dipnoans, which are allied to the ancestors of the higher vertebrates, there is the beginning of a division of the ventricle, and sometimes of the auricle, into parts by a median septum. In the higher vertebrates this septum becomes more and more specialized, separating auricle and ventricle into right and left cavities. The blood in the fish is not returned to the heart after purification, but is sent directly over the body.

The Flow of Blood.—The blood in fishes is thin and pale red (colorless in the lancelet) and with elliptical blood-corpuscles. It enters thesinus venosusfrom the head through the jugular vein, from the kidney and body walls through the cardinal vein, and from the liver through the hepatic veins. Hence it passes to the auricle and ventricle, and from the ventricle through the arterial bulb, or conus arteriosus to the ventral aorta. Thence it flows to the gills, where it is purified. After passing through the capillaries of the gill-filaments it is collected in paired arteries from each pair of gills. These vessels unite to form the dorsal aorta, which extends the length of the body just below the back-bone. From the dorsal aorta the subclavian arteries branch off toward the pectoral fins. From a point farther back arise the mesenteric arteries carrying blood to the stomach, intestine, liver, and spleen. In the tail the caudal vein carries blood to the kidneys. These secrete impurities arising fromwaste of tissues, after which the blood again passes to the heart through thecardinal vein. From the intestine the blood, charged with nutritive materials in solution, is carried by theportal veinto the liver. Here it again passes by thehepatic sinusto thesinus venosusand the heart.

The details of the circulatory system vary a good deal in the different groups, and a comparative study of the direction of veins and arteries is instructive and interesting.

The movement of the blood in fishes is relatively slow, and its temperature is raised but little above that of the surrounding water.

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