Hormones and Heredity / A Discussion of the Evolution of Adaptations and the Evolution of Species

by J. T. Cunningham

P. limanda, the Dab: scales uniform all over the body, with spinules on the projecting edges, making the skin rough; lateral line with a semicircular curve above the pectoral fin.

P. microcephalus, the Lemon-dab: scales small, smooth, and imbedded; skin slimy, head and mouth very small, colour yellowish brown with large round darker marks.

P. cynoglossus, the Witch or Pole-dab: head and mouth smaller than in the Plaice, eyes rather larger; scales all alike and uniformly distributed, slightly spinulate on upper side, smooth on the lower; blister-like cavities beneath the skin of the head on the lower side.

With regard to the generic characters, it is difficult to give any reason why the mouth should be at the end of the head instead of behind the apex of the snout as in the genus Solea, but, as we have seen already, the small size of the mouth and the greater development of teeth on the lower side are adapted to the food and mode of feeding. It is impossible to say why one genus of Flat-fishes should have the right side uppermost and others, e.g. Sole and Turbot, the left; it would almost seem to have been a matter of chance at the commencement of the evolution: reversed specimens occur as variations in most of the species.

When we consider the specific differences, we find very definite characters in the structure and distribution of the scales, and no evidence has yet been discovered that these differences are related to external conditions. There are, of course, slight differences in habits and habitat, but no constant relation between these and the structural differences of the scales. Plaice and Dab are taken together on the same ground, and nothing has been discovered to indicate that the spinulate scales of the Dab are adapted to one peculiarity in habits or conditions, the spineless scales of the Plaice to another. In comparing certain geographical races of Plaice and Flounder the facts seem to suggest that differences of habitat may have something to do with the development of the scales. In the Baltic the Flounders are as large as those on our own coasts, but the thorny tubercles are much more developed, nearly the whole of the upper surface being covered with them. The Plaice, on the other hand, are smaller than those of the North Sea, and the males have the scales spinulate over a considerable portion of the upper side. The chief difference between the Baltic and the North Sea is the reduced salinity of the former, so that it might be supposed that fresher water caused the greater development of the dermal skeleton. On the other hand, a species or geographical variety of the Plaice, whose proper is P. glacialis, is found on the Arctic coasts of Asia and America, on both sides of the extreme North Pacific, and on the east coast of North America. In this form the bony tubercles on the head in the Plaice are replaced by a continuous rough osseous ridge, and the scales are as much spinulated as in the Plaice of the Baltic. On the east coast of North America the males in this form are more spinulated than the females; on the Alaskan coast, and apparently the Arctic coast, the females are spinulated, and the sexual difference in this respect is slight or absent. Lower salinity cannot be the cause of greater spinulation in this case, and thus it might be suggested that the condition was due to lower temperature. But we do not find that northern or Arctic species of fish in general have the scales more developed than southern species.

The Dab, which occurs in the same waters as the Plaice, has the spines more spinulated than any of the forms of plaice above mentioned, therefore the absence or slight development of spinules in the typical Plaice is not explained by physical conditions alone. Freshness of water again will not explain the difference of the structure and distribution of scales in Flounder and Plaice, considering the variety of squamation in fishes confined to fresh water. Still less can we attribute any of the peculiarities of scales to utility. We can discover no possible benefit of the condition in one species which would be absent in the case of other species. We can go much further than this, and maintain that there is no reason to believe that scales in general in Teleosteans, or any of their various modifications, are of special utility: they are not adaptive structures at all, although of great importance as diagnostic characters. It may be urged that in some cases, such as the little Agonus cataphractus or the Seahorse among the Syngnathidae, the body is protected by a complete suit of bony armour; but accompanying these in the littoral region are numerous other species such as the Gobies, and even other species of Syngnathidae which have soft unprotected skins.

Similarly with colour characters: the power of changing the colour so as to harmonize with the ground is obviously beneficial and adaptive, but in each species there is a specific pattern or marking which remains constant throughout life and has nothing to do with protective resemblance, variable or permanent. The red spots of the Plaice are specific and diagnostic, but they confer no advantage over the Dab or the Lemon-dab, in which they are absent, nor can any relation be discovered between these spots and mode of life or habits.

The function of the lateral line organs is still somewhat obscure. The theory that they are sensitive to differences of hydrostatic pressure as the fish moves from one depth to another rests on no foundation, since it has yet to be shown how a change of pressure within the limits of the incompressibility of water can produce a sensation in an organ permeated throughout with water. It is more probable that the organs are affected by vibrations in the water, but we are unable to understand how a difference in the anterior curvature of the lateral line would make a difference in the function in any way related to the difference in conditions of life between Plaice and Dab. There is, however, reason to conclude that the organs, especially on the head, are more important and larger in deeper water, and thus the enlargement of the sensory canals in the head of the Witch, which lives in deeper water than other species, may be an adaptive character.

Another genus of whose characters I once made a special study is that named Zeugopterus. The name was originally given by Gottsche to the largest species Z. punctatus, from the fact that the pelvic fins are united to the ventral, but this character does not occur in other species now included in the genus. There are three species, occurring only in European waters, which form this genus and agree in the following characters. The outline of the body is more nearly rectangular than in other Flat-fishes from the obtuseness of the snout and caudal end, and the somewhat uniform breadth of the body. The surface is rough from the presence of long slender spines on the scales. There is a large perforation in the septum between the gill cavities, but this occurs also in Arnoglossus megastoma, which is placed in another genus. But the generic character of Zeugopterus, which is most important for the present discussion, is the prolongation of the dorsal and ventral fins on to the lower of the body at the base of the tail, the attachments of these accessory portions being transverse to the axis of the body. These fishes have the peculiar habit of adhering to the vertical surfaces of sides of aquaria, even the smooth surfaces of slate or glass. In nature they are taken occasionally on gravelly or sandy ground, but probably live also among rocks and adhere to them in the same way as to vertical surfaces in captivity. Many years ago (Journ. Mar. Biol. Assn., vol. iii 1893-95) I made a careful investigation of the means by which these fishes were able to adhere to a smooth surface, at least in the case of the largest and commonest species Z. punctatus. It was observed that so long as the fish was clinging to a vertical surface the posterior parts of the fins were in rhythmical motion, undulations passing along them in succession from before backwards, the edge of the body to which they were attached moving with them. The effect of these movements was to pump out water backwards from the space between the body and the surface it was clinging to, and to cause water to flow into this space at the anterior edges of the head. The subcaudal flaps were perfectly motionless and tightly pressed between the base of the tail and the surface of support, so that any movement of them was impossible. The question arose, however, whether the tail and these flaps acted as a sucker which aided in the adhesion. The flaps were therefore cut off with scissors—an operation which caused practically no pain or injury to the fish—and it adhered afterwards quite as well as when the fin-flaps were intact. The subcaudal prolongations of the fins are therefore not necessary to the adhesion, nor to the pumping action, of the muscles and fins, which went on as before. It seemed probable, therefore, that the pumping action was itself the cause of the adhesion. But the difficulty in accepting this conclusion was that there was a distinct though gentle respiratory movement of the jaws and opercula; and if the pumping of the water from beneath the body caused a negative pressure there, and a positive pressure on the outer side of the body, it seemed equally certain that the respiratory movement must force water into the space beneath the body and so cause a positive pressure there which would tend to force the fish away from the surface with which it was in contact. Examination of the currents of water around the edges of the fish, by means of suspended carmine, showed that water passed in at the mouth and out at the lower respiratory orifice, but also into the space below the body at the upper and lower edges of the head, without passing through the respiratory channel. It was thus proved that the rate at which water was pumped out at the sides of the tail was greater than that at which it passed in by the respiratory movements, and consequently there a resultant negative pressure beneath the body. By means of a model made of a thin flexible sheet of rubber, at each end of which on one side was fastened a short piece of glass tube, I was able to imitate the physical action observed in the fish. A long piece of rubber tube was attached to one of the pieces of glass tube, and brought over the edge of the glass front of an aquarium. The long rubber tube was set in action as a siphon and the sheet of rubber placed against the glass. As long as water was running through the siphon the sheet of rubber remained pressed against the glass and supported. As soon as the current of water was stopped the apparatus fell to the bottom of the tank. In this model water passed out from beneath the rubber through the glass tube attached to the siphon and passed in by the opposite glass tube, and at the sides of it. The latter tube represented the respiratory channel of the fish, and the space between tube and rubber represented the spaces between the head of the fish and the vertical surface to which it clung.

In the fish the marginal fins not only extend to the base of the tail, but are broader at the posterior end than elsewhere, whereas in other Flat-fishes the posterior part of the marginal fins are the narrowest parts. The shape of the fins and the breadth of the body posteriorly, then, are adaptations which have a definite function, that of enabling the fish to adhere to vertical surfaces. But, on the other hand, the extension of the marginal fins in a transverse direction beneath the tail has no use in the process of adhesion, nor has any other use been found for it. It is a generic character, so far as we know, without utility. On the other hand, it is very probable that this subcaudal extension of the fins is merely a result of the posterior extension and enlargement of these fins which has taken place in the evolution of the adaptation. If the Lamarckian explanation of adaptation were true, it would be possible to understand that the constant movements of the fins and muscles by which the adhesion was effected caused a longitudinal growth of the fins in excess of the length actually required, and that this extra growth extended on to the body beneath the tail, although the small flaps on the lower side were not necessary to the new function which the fins performed.