Fig. 97.—Development of Sea-bass,Centropristes striatus(Linnæus).a, egg prior to germination;b, germ-disk after first cleavage;c, germ-disk after third cleavage;d, embryo just before hatching. (After H. V. Wilson.)
Fig. 97.—Development of Sea-bass,Centropristes striatus(Linnæus).a, egg prior to germination;b, germ-disk after first cleavage;c, germ-disk after third cleavage;d, embryo just before hatching. (After H. V. Wilson.)
Fig. 97.—Development of Sea-bass,Centropristes striatus(Linnæus).a, egg prior to germination;b, germ-disk after first cleavage;c, germ-disk after third cleavage;d, embryo just before hatching. (After H. V. Wilson.)
In the early segmentation of the germ the first cleavage plane is established, and the nuclear divisions have taken place for the second; in the latter the third cleavage has been completed. As in other fishes these cleavages are vertical, the third parallel to the first. A segmentation cavity occurs as a central space between the blastomeres, as it does in the sturgeon and garpike.
In stages of late segmentation the segmentation cavity isgreatly flattened, but extends to the marginal cells of the germ-disk; its roof consists of two tiers of blastomeres, its floor of a thin film of the unsegmented substance of the germ; the marginal blastomeres are continuous with both roof and floor of the cavity, and are produced into a thin film which passes downward, around the sides of the yolk. Later the segmentation cavity is still further flattened; its roof is now a dome-shaped mass of blastomeres; the marginal cells have multiplied, and their nuclei are seen in the layer of the germ, below the plane of the segmentation cavity. These are seen in the surface view of the marginal cells of this stage; they are separated by cell boundaries only at the sides; below they are continuous in the superficial down-reaching layer of the germ. The marginal cells shortly lose all traces of having been separate; their nuclei, by continued division, spread into the layer of germ flooring the segmentation cavity, and into the delicate film of germ which now surrounds the entire yolk. Thus is formed theperiblastof the Teleost development, which from this point onward is to separate the embryo from the yolk; it is clearly the specialized inner part of the germ, which, becoming fluid-like, loses its cell-walls, although retaining and multiplying its nuclei. Later the periblast comes into intimate relations with the growing embryo; it lies directly against it, and appears to receive cell increments from it at various regions; on the other hand, the nuclei of the periblast, from their intimate relations with the yolk, are supposed to subserve some function in its assimilation.
Aside from the question of periblast, the growth of the blastoderm appears not unlike that of the sturgeon. From the blastula stage to that of the early gastrula, the changes have been but slight; the blastoderm has greatly flattened out as its margins grow downward, leaving the segmentation cavity apparent. The rim of the blastoderm has become thickened as the 'germ-ring'; and immediately in front of the dorsal lip of the blastopore its thickening marks the appearance of the embryo. The germ-ring continues to grow downward, and shows more prominently the outline of the embryo; this now terminates at the head region; while on either side of this pointspreads out tail-ward on either side the indefinite layer of outgrowingmesoderm. In the next stage the closure of the blastoporeis rapidly becoming completed; in front of it stretches the widened and elongated form of the embryo. The yolk-plug is next replaced by periblast, the dorsal lip by the tail-mass, or more accurately the dorsal section of the germ-rim; the cœlenteron under the dorsal lip has here disappeared, on account of the close approximation of the embryo to the periblast; its last remnant, the Kupffer's vesicle, is shortly to disappear. The germ-layers become confluent, but, unlike the sturgeon, the flattening of the dorsal germ-ring does not permit the formation of a neurenteric canal.
Fig. 98.Sea-bass,Centropristes striatus, natural size. (From life, by R. W. Shufeldt.)—Page 137.
Fig. 98.Sea-bass,Centropristes striatus, natural size. (From life, by R. W. Shufeldt.)—Page 137.
Fig. 98.Sea-bass,Centropristes striatus, natural size. (From life, by R. W. Shufeldt.)—Page 137.
The process of the development of the germ-layers in Teleosts appears as an abbreviated one, although in many of its details it is but imperfectly known. In the development of the medullary groove, as an example, the following peculiarities exist: the medullary region is but an insunken mass of cells without a trace of the groove-like surface indentation. It is only later, when becoming separate from the ectoderm, that it acquires its rounded character; its cellular elements then group themselves symmetrically with reference to a sagittal plane, where later, by their dissociation, the canal of the spinal cord is formed. The growth of the entoderm is another instance of specialized development. In an early stage the entoderm exists in the axial region, its thickness tapering away abruptly on either side; its lower surface is closely apposed to the periblast; its dorsal thickening will shortly become separate as the notochord. In a following stage of development the entoderm is seen to arch upward in the median line as a preliminary stage in the formation of the cavity of the gut. Later, by the approximation of the entoderm-cells in the median ventral line, the condition is reached where the completed gut-cavity exists.
The formation of the mesoderm in Teleosts is not definitely understood. It is usually said to arise as a process of 'delamination,' i.e., detaching itself in a mass from the entoderm. Its origin is, however, looked upon generally as of a specialized and secondary character.
The mode of formation of the gill-slit of the Teleost does not differ from that in other groups; an evagination of the entoderm coming in contact with an invaginated tract ofectoderm fuses, and at this point an opening is later established.
The late embryo of the Teleost, though of rounded form, is the more deeply implanted in the yolk-sac than that of the sturgeon; it is transparent, allowing notochord, primitive segments, heart, and sense-organs to be readily distinguished; at about this stage both anus and mouth are making their appearance.
Fig. 99.—Young Sword-fish,Xiphias gladius(Linnæus). (After Lütken.)
Fig. 99.—Young Sword-fish,Xiphias gladius(Linnæus). (After Lütken.)
Fig. 99.—Young Sword-fish,Xiphias gladius(Linnæus). (After Lütken.)
The Larval Development of Fishes.[10]—"When the young fish has freed itself from its egg-membranes it gives but little suggestion of its adult form. It enters upon a larval existence, which continues until maturity. The period of change of form varies widely in the different groups of fishes, from a few weeks' to longer than a year's duration; and the extent of the changes that the larva undergoes are often surprisingly broad, investing every organ and tissue of the body, the immature fish passing through a series of form stages which differ one from the other in a way strongly contrasting with the mode of growth of amniotes; since the chick, reptile, or mammal emerges from its embryonic membranes in nearly its adult form.
Fig. 100.—Sword-fish,Xiphias gladius(Linnæus). (After Day.)
Fig. 100.—Sword-fish,Xiphias gladius(Linnæus). (After Day.)
Fig. 100.—Sword-fish,Xiphias gladius(Linnæus). (After Day.)
The fish may, in general, be said to begin its existence asa larva as soon as it emerges from its egg-membranes. In some instances, however, it is difficult to decide at what point the larval stage is actually initiated: thus in sharks the excessive amount of yolk material which has been provided for the growth of the larva renders unnecessary the emerging from the egg at an early stage; and the larval period is accordingly to be traced back to stages that are still enclosed in the egg-membranes. In all cases the larval life may be said to begin when the following conditions have been fulfilled: the outward form of the larva must be well defined, separating it from the mass of yolk, its motions must be active, it must possess a continuous vertical fin-fold passing dorsally from the head region to the body terminal, and thence ventrally as far as the yolk region; and the following structures, characteristic in outward appearance, must also be established: the sense-organs—eye, ear, and nose—mouth and anus, and one or more gill-clefts.
Fig. 101.—Larva of the Sail-fish,Istiophorus, very young. (After Lütken.)
Fig. 101.—Larva of the Sail-fish,Istiophorus, very young. (After Lütken.)
Fig. 101.—Larva of the Sail-fish,Istiophorus, very young. (After Lütken.)
Fig. 102.—Larva of Brook Lamprey,Lampetra wilderi, before transformation, being as large as the adult, toothless, and more distinctly segmented.
Fig. 102.—Larva of Brook Lamprey,Lampetra wilderi, before transformation, being as large as the adult, toothless, and more distinctly segmented.
Fig. 102.—Larva of Brook Lamprey,Lampetra wilderi, before transformation, being as large as the adult, toothless, and more distinctly segmented.
Fig. 103.—Common Eel.Anguilla chrisypaRafinesque. FamilyAnguillidæ.
Fig. 103.—Common Eel.Anguilla chrisypaRafinesque. FamilyAnguillidæ.
Fig. 103.—Common Eel.Anguilla chrisypaRafinesque. FamilyAnguillidæ.
Among the different groups of fishes the larval changes are brought about in widely different ways. These larval peculiarities appear at first of far-reaching significance, but may ultimately be attributed, the writer believes, to changed environmental conditions, wherein one process may be lengthened, another shortened. So, too, the changes from one stage to another may occur with surprising abruptness. As a rule, it may be said the larval stage is of longest duration in the Cyclostomes, and thence diminished in length in sharks, lung-fishes, Ganoids, and Teleosts; in the last-named group a very much curtailed (i.e., precocious) larval life may often occur.
Fig. 104.—Larva of Common Eel,Anguilla chrisypa(Rafinesque), calledLeptocephalus grassii. (After Eigenmann.)
Fig. 104.—Larva of Common Eel,Anguilla chrisypa(Rafinesque), calledLeptocephalus grassii. (After Eigenmann.)
Fig. 104.—Larva of Common Eel,Anguilla chrisypa(Rafinesque), calledLeptocephalus grassii. (After Eigenmann.)
The metamorphoses of the newly hatched Teleost must finally be reviewed; they are certainly the most varied and striking of all larval fishes, and, singularly enough, appear to be crowded into the briefest space of time; the young fish, hatched often as early as on the fourth day, is then of the most immature character; it is transparent, delicate, easily injured, inactive; within a month, however, it may have assumed almost every detail of its mature form. A form hatching three millimeters in length may acquire the adult form before it becomes much longer than a centimeter.
Fig. 105.—Larva of Sturgeon,Acipenser sturio(Linnæus). (After Kupffer, per Dean.)
Fig. 105.—Larva of Sturgeon,Acipenser sturio(Linnæus). (After Kupffer, per Dean.)
Fig. 105.—Larva of Sturgeon,Acipenser sturio(Linnæus). (After Kupffer, per Dean.)
Fig. 106.—Larva (calledTholichthys) ofChætodon sedentarius(Poey). Cuba. (After Lütken,)
Fig. 106.—Larva (calledTholichthys) ofChætodon sedentarius(Poey). Cuba. (After Lütken,)
Fig. 106.—Larva (calledTholichthys) ofChætodon sedentarius(Poey). Cuba. (After Lütken,)
Fig. 107.—Butterfly-fish,Chætodon capistratusLinnæus. Jamaica.
Fig. 107.—Butterfly-fish,Chætodon capistratusLinnæus. Jamaica.
Fig. 107.—Butterfly-fish,Chætodon capistratusLinnæus. Jamaica.
Peculiar Larval Forms.—The young fish usually differs from the adult mainly in size and proportions. The head is larger in the young, the fins are lower, the appendages less developed, and the body more slender in the young than in the adult. But to most of these distinctions there are numerous exceptions, and in some fish there is a change so marked as to be fairly called a metamorphosis. In such cases the young fish in its first condition is properly called a larva. The larva of the lamprey (Petromyzon) is nearly blind and toothless, with slender head, and was long supposed to belong to a different genus (Ammocœtes) from the adult. The larva of sharks and rays, and also of Dipnoans and Crossopterygians, are provided with bushy external gills, which disappear in the process of development. In most soft-rayed fishes the embryonic fringe which precedes thedevelopment of the vertical fins persists for a considerable time. In many young fishes, especially theChætodontidæand their allies (butterfly-fishes), the young fish has the head armed with broad plates formed by the backward extension of certain membrane-bones. In other forms the bones of the head are in the young provided with long spines or with serrations, which vanish totally with age. Such a change is noticeable in the swordfish. In this species the production of the bones of the snout and upper jaw into a long bony sword, or weapon of offense, takes place only with age. The young fish have jaws more normally formed, and armed with ordinary teeth. In the headfish (Mola mola) large changes take place in the course of growth, and the young have been taken for a different type of fishes. Among certain soft-rayed fishes and eels the young is often developed in a peculiar way, being very soft, translucent, or band-like, and formed of large or loosely aggregated cells. These peculiar organisms, long known as leptocephali, have been shown to be the normal young of fishes when mature very different. In the ladyfish (Albula) Dr. Gilbert has shown, by a full series of specimens, that in their further growth these pellucid fishes shrink in size, acquiring greater compactness of body, until finally reaching about half their maximum length as larvæ. After this, acquiring essentially the form of the adult fish, they begin a process of regular growth. This leptocephalous condition is thought by Günther to be due to arrest of growth in abnormal individuals, but this is not the case inAlbula, and it is probably fully normal in the conger and other eels. In the surf-fishes the larvæ have their vertical fins greatly elevated, much higher than in the adult, while the body is much more closely compressed. In the deal-fish (Trachypterus) the form of the body and fins changes greatly with age, the body becoming more elongate and the fins lower. The differences between different stages of the same fish seem greater than the differences between distinct species. In fact with this and with other forms which change with age, almost the only test of species is found in the count of the fin-rays. So far as known the numbers of these structures do not change. In the moonfishes (Carangidæ) the changes with age are often very considerable. We copy Lütken's figure of the changes in the genusSelene(fig. 113). Similar changes take place inAlectis,Vomer, and other genera.
Fig. 108.—Mola mola(Linnæus). Very early larval stage of the Headfish, calledCentaurus boöps. (After Richardson.)
Fig. 108.—Mola mola(Linnæus). Very early larval stage of the Headfish, calledCentaurus boöps. (After Richardson.)
Fig. 108.—Mola mola(Linnæus). Very early larval stage of the Headfish, calledCentaurus boöps. (After Richardson.)
Fig. 109.—Mola mola(Linnæus). Early larval stage, calledMolacanthus nummularis. (After Ryder.)
Fig. 109.—Mola mola(Linnæus). Early larval stage, calledMolacanthus nummularis. (After Ryder.)
Fig. 109.—Mola mola(Linnæus). Early larval stage, calledMolacanthus nummularis. (After Ryder.)
Fig. 110.—Mola mola(Linnæus). Advanced larval stage. (After Ryder.)
Fig. 110.—Mola mola(Linnæus). Advanced larval stage. (After Ryder.)
Fig. 110.—Mola mola(Linnæus). Advanced larval stage. (After Ryder.)
The Development of Flounders.—In the great group of flounders and soles (Heterosomata) the body is greatly compressed and the species swim on one side or lie flat on the bottom, with one side uppermost. This upper side is colored like the bottom, sand-color, gray, or brown, while the lower side is mostly white. Both eyes are brought around to the upper side by a twisting of the cranium and a modification or division of the frontal bones. When the young flounder is hatched it is translucent and symmetrical, swimming vertically in the water, with one eye on either side of the head. After a little the young fish rests the ventral edge on the bottom. It then leans to one side, and as its position gradually becomes horizontal the eye on the lower side moves across with its frontal and other bones to the other side. In most species it passes directly under the first interneurals of the dorsal fin. These changes are best observed in the genusPlatophrys.
Hybridism.—Hybridism is very rare among fishes in a state of nature. Two or three peculiar forms among the snappers (Lutianus) in Cuba seem fairly attributable to hybridism, the single specimen of each showing a remarkable mixture of characters belonging to two other common species. Hybrids may be readily made in artificial impregnation among those fishes with which this process is practicable. Hybrids of the different salmon or trout usually share nearly equally the traits of the parent species.
The Age of Fishes.—The age of fishes is seldom measured by a definite period of years. Most of them grow as long as they live, and apparently live until they fall victims to some stronger species. It is reputed that carp and pike have lived for a century, but the evidence needs verification. Some fishes, as the salmon of the Pacific (Oncorhynchus), have a definite periodof growth (usually four years) before spawning. After this act all the individuals die so far as known. In Japan and China the Ice-fish (Salanx), a very long, slender, transparent fish allied to the trout, may possibly be annual in habit, all the individuals perhaps dying in the fall to be reproduced from eggs in the spring. But this alleged habit needs verification.
Fig. 111.—Headfish (adult),Mola mola(Linnæus). Virginia.
Fig. 111.—Headfish (adult),Mola mola(Linnæus). Virginia.
Fig. 111.—Headfish (adult),Mola mola(Linnæus). Virginia.
Tenacity of Life.—Fishes differ greatly in tenacity of life. In general, fishes of the deep seas die at once if brought near the surface. This is due to the reduction of external pressure. The internal pressure forces the stomach out through the mouth and may burst the air-bladder and the large blood-vessels. Marine fishes usually die very soon after being drawn out from the sea.
Fig. 112.—Albula vulpes(Linnæus). Transformation of the Ladyfish, from the translucent, loosely compacted larva to the smaller, firm-bodied young. Gulf of California. (After Gilbert.)
Fig. 112.—Albula vulpes(Linnæus). Transformation of the Ladyfish, from the translucent, loosely compacted larva to the smaller, firm-bodied young. Gulf of California. (After Gilbert.)
Fig. 112.—Albula vulpes(Linnæus). Transformation of the Ladyfish, from the translucent, loosely compacted larva to the smaller, firm-bodied young. Gulf of California. (After Gilbert.)
Fig. 113.—Development of the Horsehead-fish,Selene vomer(Linnæus). FamilyCarangidæ. (After Lütken.)
Fig. 113.—Development of the Horsehead-fish,Selene vomer(Linnæus). FamilyCarangidæ. (After Lütken.)
Fig. 113.—Development of the Horsehead-fish,Selene vomer(Linnæus). FamilyCarangidæ. (After Lütken.)
Some fresh-water fishes are very fragile, dying soon in the air, often with injured air-bladder or blood-vessels. They will die even sooner in foul water. Other fishes are extremely tenacious of life. The mud-minnow (Umbra) is sometimes ploughed up in the half-dried mud of Wisconsin prairies. The related Alaskan blackfish (Dallia) has been fed frozen to dogs, escaping alive from their stomachs after being thawed out. Many of the catfishes (Siluridæ) will live after lying half-dried in the dust for hours. The Dipnoan,Lepidosiren, lives in a ball of half-dried mud during the arid season, and certain fishes, mostly Asiatic, belonging to the groupLabyrinthici, with accessory breathing organ can long maintain themselves out of water. Among these is the China-fish (Ophiocephalus), often kept alive in the Chinese settlements in California and Hawaii. Some fishes can readily endure prolonged hunger, while others succumb as readily as a bird or a mammal.
Fig. 114.—Ice-fish,Salanx hyalocraniusAbbott. FamilySalangidæ. Tientsin, China.
Fig. 114.—Ice-fish,Salanx hyalocraniusAbbott. FamilySalangidæ. Tientsin, China.
Fig. 114.—Ice-fish,Salanx hyalocraniusAbbott. FamilySalangidæ. Tientsin, China.
Fig. 115.—Alaska Blackfish,Dallia pectoralis(Bean). St. Michaels, Alaska.
Fig. 115.—Alaska Blackfish,Dallia pectoralis(Bean). St. Michaels, Alaska.
Fig. 115.—Alaska Blackfish,Dallia pectoralis(Bean). St. Michaels, Alaska.
The Effects of Temperature on Fish.—The limits of distribution of many fishes are marked by changes in temperature. Few marine fishes can endure any sudden or great change in this regard, although fresh-water fishes adapt themselves to the seasons. I have seen the cutlass-fish (Trichiurus) benumbed with cold off the coast of Florida while the temperature was still above the frost-line. Those fishes which are tenacious of life and little sensitive to changes in climate and food are most successfully acclimatized or domesticated. The Chinese carp(Cyprinus carpio) and the Japanese goldfish (Carassius auratus) have been naturalized in almost all temperate and tropical river basins. Within the limits of clear, cold waters most of the salmon and trout are readily transplanted. But some similar fishes (as the grayling) are very sensitive to the least change in conditions. Most of the catfish (Siluridæ) will thrive in almost any fresh waters except those which are very cold.
Fig. 116.—Snake-headed China-fish,Ophiocephalus barca. India. (After Day.)
Fig. 116.—Snake-headed China-fish,Ophiocephalus barca. India. (After Day.)
Fig. 116.—Snake-headed China-fish,Ophiocephalus barca. India. (After Day.)
Transportation of Fishes.—The eggs of species of salmon, placed in ice to retard their development, have been successfully transplanted to great distances. The quinnat-salmon has been thus transferred from California to Australia. It has been found possible to stock rivers and lakes with desirable species, or to restock those in which the fish-supply has been partly destroyed, through the means of artificially impregnated eggs.
The method still followed is said to be the discovery of J. L. Jacobi of Westphalia (about 1760). This process permits the saving of nearly all the eggs produced by the individuals taken. In a condition of nature very many of these eggs would be left unfertilized, or be destroyed by other animals. Fishes are readily kept in captivity in properly constructed aquaria. Unless injured in capture or transportation, there are few species outside the deep seas which cannot adapt themselves to life in a well-constructed aquarium.
Reproduction of Lost Parts.—Fishes have little power to reproduce lost parts. Only the tips of fleshy structures are thus restored after injury. Sometimes a fish in which the tail has been bitten off will survive the injury. The wound will heal, leaving the animal with a truncate body, fin-rays sometimes arising from the scars.
Fig. 117.—Monstrous Goldfish (bred in Japan),Carassius auratus(Linnæus). (After Günther.)
Fig. 117.—Monstrous Goldfish (bred in Japan),Carassius auratus(Linnæus). (After Günther.)
Fig. 117.—Monstrous Goldfish (bred in Japan),Carassius auratus(Linnæus). (After Günther.)
Monstrosities among Fishes.—Monstrosities are rare among fishes in a state of nature. Two-headed young are frequently seen at salmon-hatcheries, and other abnormally divided or united young are not infrequent. Among domesticated species monstrosities are not infrequent, and sometimes, as in the goldfish, these have been perpetuated to become distinct breeds or races. Goldfishes with telescopic eyes and fantastic fins, and with the green coloration changed to orange, are reared in Japan, and are often seen in other countries. The carp has also been largely modified, the changes taking place chiefly in the scales. Some are naked (leather-carp), others (mirror-carp) have a few large scales arranged in series.
FOOTNOTES:[9]This account of the normal development of the Teleost fishes is condensed from Dr. Dean's "Fishes Living and Fossil," in which work the details of growth in the Teleost are contrasted with those of other types of fishes.[10]This paragraph is condensed from Dean's "Fishes Living and Fossil."
[9]This account of the normal development of the Teleost fishes is condensed from Dr. Dean's "Fishes Living and Fossil," in which work the details of growth in the Teleost are contrasted with those of other types of fishes.
[9]This account of the normal development of the Teleost fishes is condensed from Dr. Dean's "Fishes Living and Fossil," in which work the details of growth in the Teleost are contrasted with those of other types of fishes.
[10]This paragraph is condensed from Dean's "Fishes Living and Fossil."
[10]This paragraph is condensed from Dean's "Fishes Living and Fossil."
TheHabits of Fishes.—The habits of fishes can hardly be summarized in any simple mode of classification. In the usual course of fish-life the egg is laid in the early spring, in water shallower than that in which the parents spend their lives. In most cases it is hatched as the water grows warmer. The eggs of the members of the salmon and cod families are, however, mostly hatched in cooling waters. The young fish gathers with others of its species in little schools, feeds on smaller fishes of other species or of its own, grows and changes until maturity, deposits its eggs, and the cycle of life begins again, while the old fish ultimately dies or is devoured.
Irritability of Animals.—All animals, of whatever degree of organization, show in life the quality of irritability or response to external stimulus. Contact with external things produces some effect on each of them, and this effect is something more than the mere mechanical effect on the matter of which the animal is composed. In the one-celled animals the functions of response to external stimulus are not localized. They are the property of any part of the protoplasm of the body. In the higher or many-celled animals each of these functions is specialized and localized. A certain set of cells is set apart for each function, and each organ or series of cells is released from all functions save its own.
Nerve-cells and Fibres.—In the development of the individual animal certain cells from the primitive external layer or ectoblast of the embryo are set apart to preside over the relations of the creature to its environment. These cells are highly specialized, and while some of them are highly sensitive, others are adapted for carrying or transmitting the stimuli received by the sensitive cells, and still others have the function of receiving sense-impressions and of translating them into impulses of motion. The nerve-cells are receivers of impressions. These are gathered together in nerve-masses or ganglia, the largest of these being known as the brain, the ganglia in general being known as nerve-centres. The nerves are of two classes. The one class, called sensory nerves, extends from the skin or other organ of sensation to the nerve-centre. The nerves of the other class, motor nerves, carry impulses to motion.
The Brain, or Sensorium.—The brain or other nerve-centre sits in darkness, surrounded by a bony protecting box. To this main nerve-centre, orsensorium, come the nerves from all parts of the body that have sensation, the external skin as well as the special organs of sight, hearing, taste, and smell. With these come nerves bearing sensations of pain, temperature, muscular effort—all kinds of sensation which the brain can receive. These nerves are the sole sources of knowledge to any animal organism. Whatever idea its brain may contain must be built up through these nerve-impressions. The aggregate of these impressions constitute the world as the organism knows it. All sensation is related to action. If an organism is not to act, it cannot feel, and the intensity of its feeling is related to its power to act.
Reflex Action.—These impressions brought to the brain by the sensory nerves represent in some degree the facts in the animal's environment. They teach something as to its food or its safety. The power of locomotion is characteristic of animals. If they move, their actions must depend on the indications carried to the nerve-centre from the outside; if they feed on living organisms, they must seek their food; if, as in many cases, other living organisms prey on them, they must bestir themselves to escape. The impulse of hunger on the one hand and of fear on the other are elemental. The sensorium receives an impression that food exists in a certain direction. At once an impulse to motion is sent out from it to the muscles necessary to move the body in that direction. In the higher animals these movements are more rapid and more exact. This is because organs of sense, muscles, nerve-fibres, and the nerve-cells are all alike highly specialized. In the fish the sensation is slow, the muscular response sluggish, but the method remains the same. This is simple reflex action, an impulse from theenvironment carried to the brain and then unconsciously reflected back as motion. The impulse of fear is of the same nature. Reflex action is in general unconscious, but with animals, as with man, it shades by degrees into conscious action, and into volition or action "done on purpose."
Instinct.—Different animals show differences in method or degree of response to external influences. Fishes will pursue their prey, flee from a threatening motion, or disgorge sand or gravel swallowed with their food. Such peculiarities of different forms of life constitute the basis of instinct.
Instinct is automatic obedience to the demands of conditions external to the nervous system. As these conditions vary with each kind of animal, so must the demands vary, and from this arises the great variety actually seen in the instincts of different animals. As the demands of life become complex, so do the instincts. The greater the stress of environment, the more perfect the automatism, for impulses to safe action are necessarily adequate to the duty they have to perform. If the instinct were inadequate, the species would have become extinct. The fact that its individuals persist shows that they are provided with the instincts necessary to that end. Instinct differs from other allied forms of response to external condition in being hereditary, continuous from generation to generation. This sufficiently distinguishes it from reason, but the line between instinct and reason and other forms of reflex action cannot be sharply drawn.
It is not necessary to consider here the question of the origin of instincts. Some writers regard them as "inherited habits," while others, with apparent justice, doubt if mere habits or voluntary actions repeated till they become a "second nature" ever leave a trace upon heredity. Such investigators regard instinct as the natural survival of those methods of automatic response which were most useful to the life of the animal, the individual having less effective methods of reflex action perishing, leaving no posterity.
Classification of Instincts.—The instincts of fishes may be roughly classified as to their relation to the individual into egoistic and altruistic instincts.
Egoistic instinctsare those which concern chiefly the individual animal itself. To this class belong the instincts of feeding, those of self-defense and of strife, the instincts of play, the climatic instincts, and environmental instincts, those which direct the animal's mode of life.
Altruistic instinctsare those which relate to parenthood and those which are concerned with the mass of individuals of the same species. The latter may be called the social instincts. In the former class, the instincts of parenthood, may be included the instinct of courtship, reproduction, home-making, nest-building, and care for the young. Most of these are feebly developed among fishes.
The instincts of feeding are primitively simple, growing complex through complex conditions. The fish seizes its prey by direct motion, but the conditions of life modify this simple action to a very great degree.
The instinct of self-defense is even more varied in its manifestations. It may show itself either in the impulse to make war on an intruder or in the desire to flee from its enemies. Among carnivorous forms fierceness of demeanor serves at once in attack and in defense.
Herbivorous fishes, as a rule, make little direct resistance to their enemies, depending rather on swiftness of movement, or in some cases on simple insignificance. To the latter cause the abundance of minnows, anchovies, and other small or feeble fishes may be attributed, for all are the prey of carnivorous fishes, which they far exceed in number.
The instincts of courtship relate chiefly to the male, the female being more or less passive. Among many fishes the male makes himself conspicuous in the breeding season, spreading his fins, intensifying his pigmented colors through muscular tension, all this supposedly to attract the attention of the female. That this purpose is actually accomplished by such display is not, however, easily proved. In the little brooks in spring, male minnows can be found with warts on the nose or head, with crimson pigment on the fins, or blue pigment on the back, or jet-black pigment all over the head, or with varied combination of all these. Their instinct is to display all these to the best advantage, even though the conspicuous hues lead to their own destruction.
The movements of many migratory animals are mainly controlled by the impulse to reproduce. Some pelagic fishes, especially flying fishes and fishes allied to the mackerel, swim long distances to a region favorable for a deposition of spawn. Some species are known only in the waters they make their breeding homes, the individuals being scattered through the wide seas at other times. Many fresh-water fishes, as trout, suckers, etc., forsake the large streams in the spring, ascending the small brooks where they can rear their young in greater safety. Still others, known as anadromous fishes, feed and mature in the sea, but ascend the rivers as the impulse of reproduction grows strong. An account of these is given in a subsequent paragraph.
Fig. 118.—Jaws ofNemichthys avocetta. Jordan and Gilbert.
Fig. 118.—Jaws ofNemichthys avocetta. Jordan and Gilbert.
Fig. 118.—Jaws ofNemichthys avocetta. Jordan and Gilbert.
Variability of Instincts.—When we study instincts of animals with care and in detail, we find that their regularity is much less than has been supposed. There is as much variation in regard to instinct among individuals as there is with regard to other characters of the species. Some power of choice is found in almost every operation of instinct. Even the most machine-like instinct shows some degree of adaptability to new conditions. On the other hand, in no animal does reason show entire freedom from automatism or reflex action. "The fundamental identity of instinct with intelligence," says Dr. Charles O. Whitman, "is shown in their dependence upon the same structural mechanism (the brain and nerves) and in their responsive adaptability."
Adaptation to Environment.—In general food-securing structures are connected with the mouth, or, as in the anglers, are hung as lures above it; spines of offense and defense, electric organs, poison-glands, and the like are used in self-protection; the bright nuptial colors and adornments of the breeding season are doubtfully classed as useful in rivalry; the egg-sacs, nests, and other structures or habits may serve to defend the young, while skinny flaps, sand or weed-like markings, andmany other features of mimicry serve as concessions to the environment.
Each kind of fishes has its own ways of life, fitted to the conditions of environment. Some species lie on the bottom, flat, as a flounder, or prone on their lower fins, as a darter or a stone-roller. Some swim freely in the depths, others at the surface of the depths. Some leap out of the water from time to time, as the mullet (Mugil) or the tarpon (Tarpon atlanticus).
Fig. 119.—Catalina Flying Fish,Cypsilurus californicus(Cooper). Santa Barbara.
Fig. 119.—Catalina Flying Fish,Cypsilurus californicus(Cooper). Santa Barbara.
Fig. 119.—Catalina Flying Fish,Cypsilurus californicus(Cooper). Santa Barbara.
Flight of Fishes.—Some fishes called the flying-fishes sail through the air with a grasshopper-like motion that closely imitates true flight. The long pectoral fins, wing-like in form, cannot, however, be flapped by the fish, the muscles serving only to expand or fold them. These fishes live in the open sea or open channel, swimming in large schools. The small species fly for a few feet only, the large ones for more than an eighth of a mile. These may rise five to twenty feet above the water.
The flight of one of the largest flying fishes (Cypsilurus californicus) has been carefully studied by Dr. Charles H. Gilbert and the writer. The movements of the fish in the water are extremely rapid. The sole motive power is the action under the water of the strong tail. No force can be acquired while the fish is in the air. On rising from the water the movementsof the tail are continued until the whole body is out of the water. When the tail is in motion the pectorals seem in a state of rapid vibration. This is not produced by muscular action on the fins themselves. It is the body of the fish which vibrates, the pectorals projecting farthest having the greatest amplitude of movement. While the tail is in the water the ventral fins are folded. When the action of the tail ceases the pectorals and ventrals are spread out wide and held at rest. They are not used as true wings, but are held out firmly, acting as parachutes, enabling the body to skim through the air. When the fish begins to fall the tail touches the water. As soon as it is in the water it begins its motion, and the body with the pectorals again begins to vibrate. The fish may, by skimming the water, regain motion once or twice, but it finally falls into the water with a splash. While in the air it suggests a large dragon-fly. The motion is very swift, at first in a straight line, but is later deflected in a curve, the direction bearing little or no relation to that of the wind. When a vessel passes through a school of these fishes, they spring up before it, moving in all directions, as grasshoppers in a meadow.
Fig. 120.—Sand-darter,Ammocrypta clara(Jordan and Meek). Des Moines River.
Fig. 120.—Sand-darter,Ammocrypta clara(Jordan and Meek). Des Moines River.
Fig. 120.—Sand-darter,Ammocrypta clara(Jordan and Meek). Des Moines River.
Quiescent Fishes.—Some fishes, as the lancelet, lie buried in the sand all their lives. Others, as the sand-darter (Ammocrypta pellucida) and the hinalea (Julis gaimard), bury themselves in the sand at intervals or to escape from their enemies. Some live in the cavities of tunicates or sponges or holothurians or corals or oysters, often passing their whole lives inside the cavity of one animal. Many others hide themselves in the interstices of kelp or seaweeds. Some eels coil themselves in the crevices of rocks or coral masses, striking at their prey like snakes. Some sea-horses cling by their tails to gulfweed or sea-wrack. Manylittle fishes (Gobiomorus,Carangus,Psenes) cluster under the stinging tentacles of the Portuguese man-of-war or under ordinary jellyfishes. In the tide-pools, whether rock, coral, or mud, in all regions multitudes of little fishes abound. As these localities are neglected by most collectors, they have proved of late years a most prolific source of new species. The tide-pools of Cuba, Key West, Cape Flattery, Sitka, Unalaska, Monterey, San Diego, Mazatlan, Hilo, Kailua and Waiahæ in Hawaii, Apia and Pago-Pago in Samoa, the present writer has found peculiarly rich in rock-loving forms. Even richer are the pools of the promontories of Japan, Hakodate Head, Misaki, Awa, Izu, Waka, and Kagoshima, where a whole new fish fauna unknown to collectors in markets and sandy bays has been brought to light. Some of these rockfishes are left buried in the rock weeds as the tide flows, lying quietly until it returns. Others cling to the rocks by ventral suckers, while still others depend for their safety on their powers of leaping or on their quickness of their movements in the water. Those of the latter class are often brilliantly colored, but the others mimic closely the algæ or the rocks. Some fishes live in the sea only, some prefer brackish-water. Some are found only in the rivers, and a few pass more or less indiscriminately from one kind of water to another.
Fig. 121.—Pearl-fish,Fierasfer acus(Linnæus), issuing from aHolothurian. Coast of Italy. (After Emery.)
Fig. 121.—Pearl-fish,Fierasfer acus(Linnæus), issuing from aHolothurian. Coast of Italy. (After Emery.)
Fig. 121.—Pearl-fish,Fierasfer acus(Linnæus), issuing from aHolothurian. Coast of Italy. (After Emery.)
Fig. 122.—Portuguese Man-of-war Fish,Gobiomorus gronovii. FamilyStromateidæ.
Fig. 122.—Portuguese Man-of-war Fish,Gobiomorus gronovii. FamilyStromateidæ.
Fig. 122.—Portuguese Man-of-war Fish,Gobiomorus gronovii. FamilyStromateidæ.
Migratory Fishes.—The movements of migratory fishes are mainly controlled by the impulse of reproduction. Some pelagic fishes, especially those of the mackerel and flying-fish families, swim long distances to a region favorable for the deposition of spawn. Others pursue for equal distances the schools of menhaden or other fishes which serve as their prey. Some species are known mainly in the waters they make their breeding homes, as in Cuba, Southern California, Hawaii, or Japan, the individuals being scattered at other times through the wide seas.
Anadromous Fishes.—Many fresh-water fishes, as trout and suckers, forsake the large streams in the spring, ascending the small brooks where their young can be reared in greater safety. Still others, known asanadromousfishes, feed and mature in the sea, but ascend the rivers as the impulse of reproduction grows strong. Among such fishes are the salmon, shad, alewife, sturgeon, and striped bass in American waters. The most remarkable case of the anadromous instinct is found in the king salmon or quinnat (Oncorhynchus tschawytscha) of the Pacific Coast. This great fish spawns in November, at the age of four years and an average weight of twenty-two pounds. In the Columbia River it begins running with the spring freshets in March and April. It spends the whole summer, without feeding, in the ascent of the river. By autumn the individuals have reached the mountain streams of Idaho, greatly changed in appearance,discolored, worn, and distorted. The male is humpbacked, with sunken scales, and greatly enlarged, hooked, bent, or twisted jaws, with enlarged dog-like teeth. On reaching the spawning beds, which may be a thousand miles from the sea in the Columbia, over two thousand in the Yukon, the female deposits her eggs in the gravel of some shallow brook. The male covers them and scrapes the gravel over them. The female salmon does as much as the male in covering the eggs. Then both male and female drift tail foremost helplessly down the stream; none, so far as certainly known, ever survive the reproductive act. The same habits are found in the five other species of salmon in the Pacific, but in most cases the individuals do not start so early nor run so far. The blue-back salmon or redfish, however, does not fall far short in these regards. The salmon of the Atlantic has a similar habit, but the distance traveled is everywhere much less, and most of the hook-jawed males drop down to the sea and survive to repeat the acts of reproduction.