Читать книгу Mammalia - Frank E. Beddard - Страница 6

Fig. 2.—Four diagrams of stages in the development of a hair. A, Earliest stage in one of those mammals in which the dermal papilla appears first; B, C, D, three stages in the development of the hair in the human embryo. blb, Hair-bulb; crn, horny layer of the epidermis; foll, hair-follicle; grm, hair-germ; h, hair, in D, projecting on the surface; muc, Malpighian layer of epidermis; pp, dermal papilla; seb, developing sebaceous glands; sh.1, sh.2, inner and outer root-sheaths. (After Hertwig.)

Dr. Meijerle^[2] has lately described in some detail the particular arrangement of the individual hairs among mammals; they are not by any manner of means scattered without order, but show a definite and regular arrangement, which varies with the animal. For instance, in an American Monkey (Midas), the hairs arise in threes—three hairs of equal size springing from the epidermis close together; in the Paca (Coelogenys) there are in each group three stout hairs alternating with three slender hairs. In some forms a number of hairs spring from a common point: in the Jerboa (Dipus) twelve or thirteen arise from a single hole; in Ursus arctos there is the same general plan, but there is one stout hair and four or five slender ones. There are numerous other complications and modifications, but the facts, although interesting, do not appear to throw any light upon the mutual affinities of the animals. Allied forms may have a very different arrangement, while in forms which have no near relationship the plan may be very similar, as is shown by the examples cited from Dr. Meijerle's paper. The groups of hairs, moreover, have themselves a definite placing, which the same anatomist has compared with the disposition of the bundles of hairs behind and between the scales of the Armadillo, and which has led him to the view that the ancestors of mammals were scaly creatures—a view also supported by Professor Max Weber,^[3]and not in itself unreasonable when we consider the numerous points of affinity between the primitive Mammalia and certain extinct forms of reptiles.^[4]

The hairs are greatly modified in form in different mammals and in different parts of their bodies. It is very commonly the case that a soft under-fur can be distinguished from the longer and coarser hairs, which to some extent hide the latter. Thus the "sealskin" of commerce is the under-fur of the Otaria ursina of the North. The coarser hairs may be further differentiated into bristles; these again into spines, such as those of the Hedgehog and of the Porcupine. Again, the flattening and agglutination of hairs seems to be responsible for the scales of the Manis and for the horns of the Rhinoceros. It is a matter of common knowledge that upon the head of various animals, e.g. the Domestic Cat, long and sensitive hairs are developed, which are connected with the terminations of nerves, and perform a sensory, probably tactile function. These occur on the snout, above the eyes, and in the neighbourhood of the ears. It is an interesting fact that a tuft of quite similar hairs occurs on the hand of many mammals close to the wrist, which, at least in the case of Bassaricyon, are connected with a strong branch from the arm-nerve. These tufts also occur in Lemurs, in the Cat, various Rodents and Marsupials, and are probably quite general in mammals who "feel" with their fore-limbs;—in which, in fact, the fore-limbs are not exclusively running organs. That the last remaining hairs of the Cetacea are found upon the muzzle, is perhaps significant of the importance of these sensory bristles. The entire absence of hairs is quite common in this order, although traces of them are sometimes found in the embryo. The Sirenia, too, are comparatively hairless, as are also many Ungulates. Whether the presence of blubber in the former case and the existence of a very thick skin in the latter animals are facts which have had anything to do with the disappearance of hair or not, is a matter for further inquiry.

The intimate structure of the hair varies considerably. The variations concern the form of the hair, which may be round in transverse section, or so oval as to appear quite flat when the hair is examined in its entirety. The substance of the hair is made up of a central medulla or pith with a peripheral cortex; the latter is scaled, and the scales are often imbricated and with prominent edges. The amount of the two constituents also differs, and the cortex may be reduced to a series of bands surrounding only tracts of the enclosed pith. In the hair is contained the pigment to which the colour of mammals is chiefly due. Tracts of brightly-coloured skin may exist, as in the Apes of certain genera; but such structures are not general. The pigment of the hair seems to consist of those pigmentary substances known as melanins. It is remarkable to find such a uniform cause of coloration, when we consider the great variety of feather-pigments found in birds. The variations of colour of the hair of mammals are due to the unequal distribution of these brown pigments. There are very few mammals which can be called brightly coloured. The Bats of the genus Kerivoula have been compared to large butterflies, and some of the Flying Squirrels have strongly-marked contrasts of reddish brown, white, and yellow. The same may be said of the spines of certain Porcupines. But we find in the hair no bright blues, greens, and reds such as are common among birds.

There are certain general facts about the coloration of mammals which require some notice here. Next to the usually sombre hues of these animals the general absence of secondary sexual coloration is noteworthy. In but a few cases among the Lemurs and Bats do we find any marked divergences in hues between males and females. Secondary sexual characters in mammals are, it is true, often exhibited by the great length of certain hair-tracts in the male, such as the mane of the Lion, the throat- and leg-tufts of the Barbary Sheep, and so forth; but apart from these, the secondary sexual characters of mammals are chiefly shown in size, e.g. the Gorilla, or in the presence of tusks, e.g. various Boars, or of horns, as in the Deer, etc. The coloration of mammals frequently exhibits conspicuous patterns of marking. These are in the form of longitudinal stripes, of cross-stripes, or of spots; the latter may be "solid" spots, or broken up, as in the Leopard and Jaguar, into groups of smaller spots arranged in a rosette-fashion. We never find in mammals the complicated "eyes" and other markings which occur in so many birds and in other lower Vertebrates. It is important to note that in the Mammalia whose sense of sight is quite keen there should be a practical absence of secondary sexual colours. As to the relationship of the various forms of marking that do occur, it seems clear that there has been a progression from a striped or spotted condition to uniform coloration. For we find that many Deer have spotted young; that the young Tapir of the New World is spotted, while its parents are uniform blackish brown; the strongly-marked spotting of the young Puma contrasts with the uniform brown of the adult; and the Lion cub, as every one knows, is also spotted, the adult lioness showing considerable traces of the spots.

The seasonal change in the colours of certain mammals is a subject upon which much has been written. The extreme of this is seen in those creatures, such as the Polar Hare and the Arctic Fox, which become entirely blanched in the winter, recovering their darker coat in the spring. This is, however, only an extreme case of a change which is general. Most animals get a thicker fur in winter and exchange it for a lighter one in summer. And the hues of the coat change in correspondence.

Glands of the Skin.—The great variety of integumental glands possessed by the Mammalia distinguishes them from any group of lower Vertebrates. This variability, however, only concerns the anatomical structure of the glands in question. Histologically they are all of them apparently to be referred to one of two types, the sudoriparous or sweat gland and the sebaceous gland. Simple sweat and sebaceous glands are abundant in mammals, with but a few exceptions. The structures that we are now concerned with are agglomerations of these glands. The mammary glands will be treated of in connexion with the marsupium; they are either masses of sweat glands, or of sebaceous glands whose secretion has been converted into milk.

Many Carnivora possess glands opening to the exterior, near the anus, by a large orifice. These secrete various odoriferous substances, of which the well-known "civet" is an example. Other odoriferous glands are the musk glands of the Musk-deer and of the Beaver; the suborbital gland of many Antelopes; the dorsal gland of the Peccary, which has given the name of Dicotyles to the genus on account of its resemblance in form to a navel. This gland may be seen to secrete a clear watery fluid. The Elephant has a gland situated on the temple, which is said to secrete during certain periods only, and to be a warning to leave the animal alone. Very remarkable are the foot glands of certain species of Rhinoceros; they are not universally present in those animals, and are therefore useful as specific distinctions. On the back of the root of the tail in many Dogs are similar glands. The Gentle Lemur (Hapalemur) has a peculiar gland upon the arm, about the size of an almond, which in the male underlies a patch of spiny outgrowths. In Lemur varius is a hard patch of black skin which may be the remnants of such a gland. It is thought that the callosities on the legs of Horses and Asses are remnants of glands.

One of the most complex of these structures which has been examined microscopically exists in the Marsupial Myrmecobius.^[5] On the skin of the anterior part of the chest, just in front of the sternum, is a naked patch of skin which is seen to be perforated by numerous pores. Besides the ordinary sebaceous and sweat glands there are a series of masses of glands, opening by larger orifices, which present the appearance of groups of sebaceous glands, and are of a racemose character; but the existence of muscular fibres in their coats seems to show that they should be referred rather to the sudoriparous series. Beneath the integument is a large compound tubular gland quite half an inch in diameter.

In Didelphys dimidiata there is a precisely similar glandular area and large underlying gland, the correspondence being remarkable in two Marsupials so distant in geographical position and affinities. Even among the Diprotodont genera there is something of the kind; for in Dorcopsis luctuosa and D. muelleri is a collection of four unusually large sebaceous follicles upon the throat, and in the Tree Kangaroo (Dendrolagus bennettii) there is the same collection of enlarged hair-follicles, though they are apparently somewhat reduced as compared with those of Dorcopsis. These are of course a few examples out of many.

It seems to be possible that the functions of these various glands is at least twofold. In the first place, they may serve, where predominant in one sex, to attract the sexes together. In the second place, the glands may be useful to enable a strayed animal of a gregarious species to regain the herd. It is perfectly conceivable too that in other cases the glands may be a protection, as they most undoubtedly are in the Skunk, from attacks. In connexion with the first, and more especially the second, of the possible uses of these glands, it is interesting to note that in purely terrestrial creatures, such as the Rhinoceros, the glands are situated on the feet, and would therefore taint the grass and herbage as the animal passed, and thus leave a track for the benefit of its mate. The same may be said of the rudimentary glands of Horses if they are really glands. The secretion of the "crumen" of Antelopes is sometimes deposited deliberately by Oreotragus upon surrounding objects, a proceeding which would attain the same end. One may even perhaps detect "mimicry" in the similar odours of certain animals. Prey may be lured to their destruction, or enemies frightened away. The defenceless Musk-deer may escape its foes by the suggestion of the musky odour of a crocodile. It is at any rate perfectly conceivable that the variety of odours among mammals may play a very important part in their life, and it is perhaps worthy of note that birds with highly-variegated plumage are provided only with the uropygial gland, while mammals with usually dull and similar coloration have a great variety of skin glands. Scent is no doubt a sense of higher importance in mammals than in birds. The subject is one which will bear further study.

Nails and Claws.—Except for the Cetacea (where rudiments have been found in the foetus), the extremities of the fingers and of the toes of mammals are covered by, or encased in, horny epidermic plates, known as nails, claws, and hoofs.

The variety in the shape and development of these corneous sheaths to the digits is highly characteristic of mammals as opposed to lower Vertebrates. If we take extreme cases, such as the nail of the thumb in Man, the hoof of a Horse, and the claw of a Cat, it is easy to distinguish the three kinds of phalangeal horny coverings. But the differences become extinguished as we pass from these to related types. The nail of the little finger in Man approaches the claw-like form; and the hoofs of the Lama are almost claws in the sharpness of their extremities. On the whole it may be said that claws and hoofs embrace the bone which they cover, while nails lie only upon its dorsal surface. The form of the distal phalanx which bears the nail shows, however, two kinds of modification which do not support such a classification. When those phalanges are clad with hoofs or covered by a nail they end in a rounded and flattened termination. On the other hand, when they bear a claw they are themselves sharpened at the extremity and often grooved above.

The Marsupium.—It may appear to be unnecessary at this juncture to speak of the marsupial pouch, which is so usually believed to be a characteristic of the group Marsupialia. Rudiments of this structure have, however, been recently discovered in the higher mammals, and, as Dr. Klaatsch^[6] has remarked, all researches into the "history of the mammals culminate in the question whether the placental mammals pass through a marsupial stage or not." We cannot, therefore, look upon the marsupial pouch as a matter affecting only the Marsupials, though it is true that this organ is at present functional only in them and in the Monotremata.

Fig. 3.—Echidna hystrix. A, Lower surface of brooding female; B; dissection showing a dorsal view of the pouch and mammary glands; ††, the two tufts of hair in the lateral folds of the mammary pouch from which the secretion flows, b.m, Pouch; cl, cloaca; g.m, groups of mammary glands. (From Wiedersheim's Comparative Anatomy, after W. Haacke.)

In the Marsupials the pouch shelters the young, which are born in an exceedingly imperfect state, minute, nude, and blind, with a "larval" mouth fitted only to grasp in a permanent fashion the teat, upon which they are carefully fixed by the parent. But even later the pouch is made use of as a temporary harbour of refuge: from the pouch of female Kangaroos at the Zoological Gardens may frequently be observed to protrude the tail and hind-legs of a young Kangaroo as big as a Cat, and perfectly well able to take care of itself.

In the Monotremata (in Echidna) there is a deep fold of the skin which lodges the unhatched egg, and into which the mammary glands open, one on either side. This structure is only periodically developed, and arises from two rudiments, one corresponding to each mammary area; but in the female with eggs or young there is but a single deep depression, which occupies the same region of the body as the marsupial pouch of the Marsupials.^[7] It is usually held that this structure is not of precisely the same morphological value as the pouch of the Marsupial; and the difference is expressed by terming the one (that of Echidna) the mammary pouch, and the other the marsupium. At first sight it may appear to be an unnecessary refinement to separate two structures which have so many and such obvious likenesses. It is not quite certain, however, that the difference is not even more profound than later opinions seem to indicate. The Monotremata not only have no teats, as has already been pointed out, but the mammary glands themselves are of a perfectly different nature to those of the higher mammals, including the Marsupials. There is therefore no a priori objection to the view that the accessory parts developed in connexion with the mammary glands should also be different. The teat of the higher Mammalia grows up round the area upon which the ducts of the mammary glands open; it is a fold of skin which eventually assumes the cylindrical form of the adult teat, and which includes the ducts of the milk glands. It has been suggested that the two folds of skin which form the mammary pouch of Echidna are to be looked upon as the equivalent of the commencing teat of the higher mammal.^[8] In this case it is clear that the marsupial folds of the Marsupial cannot correspond accurately with the apparently similar folds of Echidna, because there are teats as well. It is the teats which correspond to the marsupial folds of Echidna. This view is in apparent contradiction to an interesting discovery in a specimen of a Phalanger by Dr. Klaatsch.^[9] This Marsupial, like most others, has a well-developed marsupial pouch, in which the young are lodged at birth; but round two of the teats is another distinct fold on either side, the outer wall of which forms the general wall of the pouch. Dr. Klaatsch thinks that these smaller and included pouches are the equivalents of the mammary pouches of Echidna. They contain teats, but this comparison does not do away with the validity of Gegenbaur's suggestion already referred to, because the teats are (see above) secondary. If this fact be fairly to be interpreted in the sense which Dr. Klaatsch attaches to it, we have an interesting case of the growth of a new organ out of and partly replacing an old organ. In the Monotremes there is a pouch which facilitates or performs both nutritive and protective functions; in the Phalanger these two functions are carried on in separate pouches; finally, in other Marsupials, there is a return to the undifferentiated state of affairs found in the Monotremata, but with the help of a new organ not found in them.

Fig. 4.—Diagram of the development of the nipple (in vertical section). A, Indifferent stage, glandular area flat; B, elevation of the glandular area with the nipple; C, elevation of the periphery of the glandular area into the false teat, a, Periphery of the glandular area; b, glandular area; gl, glands. (From Gegenbaur.)

Though so characteristic of Marsupials, the marsupial pouch is not always developed in them. It is present in all the Kangaroos, Wallabies, and Wombats, in fact in the Diprotodonts. It is also present in a number of the carnivorous Polyprotodont Marsupials; but in Phascologale it is only present in rudiment, and in Myrmecobius it is entirely obsolete. In the American Opossums the state of the pouch is variable. "Generally absent, sometimes merely composed of two lateral folds of skin separate at each end, rarely complete," is Mr. Thomas' summary in his definition of the family Didelphyidae.^[10] Another curious feature of the pouch in the Marsupials is the variability in the position of the mouth of the pouch: in all the Diprotodonts it looks forward; but in many Polyprotodonts it looks backward. This, however, has some connexion with the habitual attitude of the possessor: in the Kangaroo, leaping along on its hind-legs, it is requisite that the pouch should open forwards; but in the dog-like Thylacine, going on all fours, the fact that the pouch opens backwards is less disadvantageous to the contained young.

The male Thylacine has a pouch which is quite or very nearly as well formed as in the female. There are also rudiments of a pouch in the male foetuses of many Marsupials, especially of those belonging to the Polyprotodont section of the order, though these rudiments are by no means confined to that subdivision. Up to so late a period as the age of four months (length 19.8 cm.) the male Dasyurus ursinus has a pouch.

We have now to consider the interesting series of facts relative to the permanence—in a rudimentary condition it is true—of the mammary pouch in the higher Mammalia, facts which seem to be an additional proof that they have been derived from an ancestor in which the pouch was an organ of functional importance. The first definite proof of the occurrence of a pouch in any mammal not a Marsupial or a Monotreme was made by Malkmus, who found this structure in a Sheep. It seems, however, that the structures found in the higher mammals are not always comparable to the marsupium of the Marsupials, but sometimes to the mammary pouch of the Monotreme. That the Marsupials are a side line, and not involved in the ancestry of the Eutheria, is an opinion which is at present widely held. At the same time it is reasonable to suppose that the original stock lying between the Prototheria and the Metatheria, whence the latter and the Eutheria have arisen, preserved both the mammary pouch of the lower mammal and the marsupium of the further-developed stage, as does Phalangista occasionally at the present day. Hence to find remnants of both structures in existing mammals would not so incredible. This is what Dr. Klaatsch believes to be the case. In certain Ungulates, including two species of Antelope, Dr. Klaatsch found very considerable rudiments of folds provided with unstriated muscular fibre; there were in the adult Cervicapra isabellina a pair of pouches, one on each side, and a rudiment of a second on either side; possibly this multiplication of the pouches has relation to the number of young. That there is more than one pouch makes a comparison with the mammary pouch rather than with the marsupium probable. The Ungulate teat, it must be remembered (see p. 16), is a secondary teat; hence there is no difficulty in the comparison from this point of view. A pouch containing a primary teat would of course be absolutely incomparable with a mammary pouch, because in that case the wall of the teat itself would be the pouch.

Mammals belonging to quite different Orders show traces more or less marked of a marsupium. In young Dogs the teats are borne upon an area where the skin is thinner, the covering of hair less dense than elsewhere—all points of resemblance to the inside of the pouch of a Marsupial; in addition to this there are traces of the sphincter marsupii muscle. In other Carnivora there are similar vestiges. In Lemur catta a more complete rudiment of a marsupial pouch is to be met with. In this Lemur the teats are both inguinal and pectoral; the skin in these regions is thin and but slightly hairy, and extends forwards as two bands of the same thinness and smoothness on each side of the densely hairy skin covering the sternum. This area is sharply separated from the rest of the integument by a fold which runs parallel to the longitudinal axis of the body, and can be comparable with nothing save the rudiment of the marsupial fold.

One is tempted to wonder how far the habit which certain Lemurs have of carrying their young across the abdomen with the tail wrapped round the body of the mother is a reminiscence of a marsupial pouch.