less efficiency of the archipterygium and its predominance amongst the more ancient forms of fishes point to its being the more archaic of these two types.
In the less highly specialized groups of fishes the pectoral fins are close behind the head, the pelvic fins in the region of the cloacal opening. In the more specialized forms the pelvic fins frequently show a more or less extensive shifting towards the head, so that their position is described as thoracic (fig. 4) or jugular (Gadus—cod, haddock, &c., fig. 5).
Fig. 5.—Burbot (Lota vulgaris), with jugular ventral fins.
The median fin, especially in its caudal section, is the main propelling organ: the paired fins in the majority of fishes serve for balancing. In the Dipneusti the paired fins are used for clambering about amidst vegetation, much in the same fashion as the limbs of Urodeles. In Ceratodus they also function as paddles. In various Teleosts the pectoral fins have acquired secondarily a leg-like function, being used for creeping or skipping over the mud (Periophthalmus; cf. also Trigloids, Scorpaenids and Pediculati). In the “flying” fishes the pectoral fins are greatly enlarged and are used as aeroplanes, their quivering movements frequently giving a (probably erroneous) impression of voluntary flapping movements. In the gobies and lumpsuckers (Cyclopteridae) the pelvic fins are fused to form an adhesive sucker; in the Gobiesocidae they take part in the formation of a somewhat similar sucker.
The evolutionary history of the paired limbs forms a fascinating chapter in vertebrate morphology. As regards their origin two hypotheses have attracted special attention: (1) that enunciated by Gegenbaur, according to which the limb is a modified gill septum, and (2) that supported by James K. Thacher, F. M. Balfour, St George Mivart and others, that the paired fins are persisting and modified portions of a once continuous fin-fold on each side of the body. The majority of morphologists are now inclined to accept the second of these views. Each has been supported by plausible arguments, for which reference must be made to the literature of the subject.[1] Both views rest upon the assumed occurrence of stages for the existence of which there is no direct evidence, viz. in the case of (1) transitional stages between gill septum and limb, and in the case of (2) a continuous lateral fin-fold. (There is no evidence that the lateral row of spines in the acanthodian Climatius has any other than a defensive significance.) In the opinion of the writer of this article, such assumptions are without justification, now that our knowledge of Dipnoan and Crossopterygian and Urodele embryology points towards the former possession by the primitive vertebrate of a series of projecting, voluntarily movable, and hence potentially motor structure on each side of the body. It must be emphasized that these—the true external gills—are the only organs known actually to exist in vertebrates which might readily be transformed into limbs. When insuperable objections are adduced to this having actually taken place in the course of evolution, it will be time enough to fall back upon purely hypothetical ancestral structures on which to base the evolutionary history of the limbs.
The ectoderm covering the general surface is highly glandular. In the case of the Dipneusti, flask-shaped multicellular glands like those of Amphibians occur in addition to the scattered gland cells.
A characteristic feature of glandular activity is the production of a slight electrical disturbance. In the case of Malopterurus this elsewhere subsidiary function of the skin has become so exaggerated as to lead to the conversion of the skin of each side of the body into a powerful electrical organ.[2] Each of these consists of some two million small chambers, each containing an electric disk and all deriving their nerve supply from the branches of a single enormous axis cylinder. This takes its origin from a gigantic ganglion cell situated latero-dorsally in the spinal cord between the levels of the first and second spinal nerves.
Cement Organs.—The larvae of certain Teleostomes and Dipnoans possess special glandular organs in the head region for the secretion of a sticky cement by which the young fish is able to attach itself to water-plants or other objects. As a rule these are ectodermal in origin; e.g. in Lepidosiren and Protopterus[3] the crescentic cement organ lying ventrally behind the mouth consists of a glandular thickening of the deep layer of the ectoderm. In young ganoid fishes preoral cement organs occur. In Crossopterygians there is one cup-shaped structure on each side immediately in front of the mouth. Here the glandular epithelium is endodermal, developed[4] as an outgrowth from the wall of the alimentary canal, closely resembling a gill pouch. In Amia[5] the same appears to be the case. In a few Teleosts similar organs occur, e.g. Sarcodaces, Hyperopisus,[6] where so far as is known they are ectodermal.
Photogenic Organs.—The slimy secretion produced by the epidermal glands of fishes contains in some cases substances which apparently readily undergo a slow process of oxidation, giving out light of low wave-length in the process and so giving rise to a phosphorescent appearance. In many deep-sea fishes this property of producing light-emitting secretion has undergone great development, leading to the existence of definite photogenic organs. These vary much in character, and much remains to be done in working out their minute structure. Good examples are seen in the Teleostean family Scopelidae, where they form brightly shining eye-like spots scattered about the surface of the body, especially towards the ventral side.
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| From Trans. Zool. Soc. of London. |
| Fig. 6.—Larva of Polypterus. (After Budgett.) |
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| From Phil. Transactions, Royal Society of London. |
| Fig. 7.—Thirty Days’ Larval Lepidosiren. (After Graham Kerr.) |
External Gills.—In young Crossopterygians and in the young Protopterus and Lepidosiren true external gills occur of the same morphological nature as those of Urodele amphibians. In Crossopterygians a single one is present on each side on the hyoid arch; in the two Dipnoans mentioned four are present on each side—on visceral arches III., IV., V. and VI. (It may be recalled that in Urodeles they occur on arches III., IV. and V., with vestiges[7] on arches I. and II.). Each external gill develops as a projection of ectoderm with mesodermal core near the upper end of its visceral arch; the main aortic arch is prolonged into it as a loop. When fully developed it is pinnate, and is provided with voluntary muscles by which it can be moved freely to renew the water in contact with its respiratory surface. In the case of Polypterus a short rod of cartilage projects from the hyoid arch into the base of the external gill. Their occurrence with identical main features in the three groups mentioned indicates that the external gills are important and archaic organs of the vertebrata. Their non-occurrence in at least some of the groups where they are absent is to be explained by the presence of a large vascular yolk sac, which necessarily fulfils in a very efficient way the respiratory function.
Alimentary Canal.—The alimentary canal forms a tube traversing the body from mouth to cloacal opening. Corresponding with structural and functional differences it is for descriptive
- ↑ Cf. J. Graham Kerr, Proc. Camb. Phil. Soc. x. 227.
- ↑ For electric organs see W. Biedermann, Electro-Physiology.
- ↑ J. Graham Kerr, Quart. Journ. Micr. Sci. vol. xlvi.
- ↑ J. Graham Kerr, The Budgett Memorial Volume.
- ↑ J. Phelps, Science, vol. N.S. ix. p. 366; J. Eycleshymer and Wilson, Amer. Journ. Anat. v. (1906) p. 154.
- ↑ J. S. Budgett, Trans. Zool. Soc. Lond. xvi., 1901, p. 130.
- ↑ L. Drüner, Zool. Jahrbücher Anat. Band xix. (1904), S. 434.
