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Text Book of Biology, Part 1: Vertebrata by H. G. Wells, is part of the HackerNoon Books Series. You can jump to any chapter in this book here. General Anatomy
Section 1. We will now study the adult anatomy of the frog, and throughout we shall make constant comparisons with that of the rabbit. In the rabbit we have a distinctly land-loving, burrowing animal; it eats purely vegetable food, and drinks but little. In the frog we have a mainly insectivorous type, living much in the water. This involves the moister skin, the shorter alimentary canal, and the abbreviated neck (Rabbit, Section 2) of the frog; the tail is absent-- in a fish it would do the work the frog accomplishes with his hind legs-- and the apertures which are posterior in the rabbit, run together into one dorsal opening, the cloaca. There is, of course (Rabbit, Section 4), no hair the skin is smooth, and an external ear is also absent. The remarkable looseness of the frog's skin is due to great lymph spaces between it and the body wall.
Section 2. If we now compare the general anatomy of the frog (vide Sheet 11) with that of the rabbit, we notice that the diaphragm is absent (Rabbit, Section 4), and the body cavity, or coelom, is, with the exception of the small bag of the pericardium round the heart, one continuous space. The forked tongue is attached in front of the lower jaw, and can be flicked out and back with great rapidity in the capture of the small insects upon which the frog lives. The posterior nares open into the front of the mouth-- there is no long nasal chamber, and no palate, and there is no long trachea between the epiglottis and the lungs. The oesophagus is less distinct, and passes gradually, so far as external appearances go, into the bag-like stomach, which is much less inflated and transverse than that of the rabbit. The duodenum is not a U-shaped loop, but makes one together with the stomach; the pancreas lies between it and the stomach, and is more compact than the rabbit's. There is no separate pancreatic duct, but the bile duct runs through the pancreas, and receives a series of ducts from that gland as it does so. The ileum is shorter, there is no sacculus rotundus, and the large intestine has no caecum, none of the characteristic sacculations of the rabbit's colon, and does not loop back to the stomach before the rectum section commences. The anus opens not upon the exterior, but into a cloacal chamber. The urinary and genital ducts open separately into this cloaca, and dorsally and posteriorly to the anus. The so-called urinary bladder is ventral to the intestine, in a position answering to that of the rabbit, but it has no connection with the ureters, and it is two-horned.
Section 3. The spleen is a small, round body, not so intimately bound to the stomach as in the rabbit, but in essentially the same position.
Section 4. Much that we knew of the physiology of the frog is arrived at mainly by inferences from our mammalian knowledge. Its histology is essentially similar. Ciliated epithelium is commoner and occurs more abundantly than in the rabbit, in the roof of the mouth for instance, and its red blood corpuscles are much larger, oval, and nucleated.
Section 5. The lungs of the frog are bag-like; shelves and spongy partitions project into their cavities, but this structure is much simpler than that of the rabbit's lung, in which the branching bronchi, the imperfect cartilaginous rings supporting them, alveoli, arteries and veins, form together a quasi-solid mass.
Section 6. The mechanism of respiration is fundamentally different from that of the mammal. The method is as follows:-- The frog opens its anterior nares, and depresses the floor of the mouth, which therefore fills with air. The anterior nares are then closed, and the floor mouth rises and forces the air into the lungs-- the frog, therefore, swallows its air rather than inhales it. The respiratory instrument of the rabbit is a suction pump, while that of the frog is a "buccal force pump."
Section 7. The heart is not quadrilocular (i.e., of four chambers), but trilocular (of three), and two structures, not seen in Lepus, the truncus arteriosus and the sinus venosus, into the latter of which the venous blood runs before entering the right auricle, are to be noted. The single ventricle is blocked with bars of tissue that render its interior, not an open cavity, but a spongy mass. Figure 2, Sheet 11, shows the heart opened; l.au. and r.au. are the left and right auricles respectively; the truncus arteriosus is seen to be imperfectly divided by a great longitudino-spiral valve (l.s.v.); p.c. is the pulmo-cutaneous artery -going to the lungs- [supplying skin and lungs]; d.ao., the dorsal aorta [furnishing the supply of the body and limbs]; and c.a. the carotid artery going to the head; all of which vessels (compare Figure 1) are paired.
Section 8. It might be inferred from this that pure and impure blood mix in the ventricle, and that a blood of uniform quality flows to lungs, head, and extremities; but this is not so. The spongy nature of the ventricle sufficiently retards this mixing. It will be noted that the opening of pulmonary arteries lies nearest to the heart, next come the aortic and carotid arches, which have a common opening at A. Furthermore, at c.g.l. [the carotid artery, repeatedly divides to form a close meshwork of arterioles, the carotid gland, forming a sponge-like plug in this vessel.] is a spongy mass of matter, the carotid gland inserted upon the carotid. Hence the pulmonary arteries yawn nearest for the blood, and, being short, wide vessels, present the least resistance to the first rush of blood-- mainly venous blood for the right auricle. As they fill up, the back resistance in them becomes equal, and then greater, than the resistance at A, and the rush of blood, now of a mixed quality passes through that aperture. It selects the dorsal aorta, because the carotid arch, plugged by the carotid gland, offers the greater resistance. Presently, however, the back resistance of the filled dorsal aorta rises above this, and the last flow of blood, from the ventricular systole-- almost purely oxygenated blood for the left auricle-- goes on towards the head.
Section 9. At the carotid gland the carotid artery splits into -an- [a] -external carotid- [lingual] (e.c.), and a deeper internal carotid. The dorsal aorta passes round on each side of the oesophagus, as indicated by the dotted lines in Figure 2, Sheet 11, and meets its fellow dorsal to the liver. Each arch gives off subclavian arteries to the limbs, and the left, immediately before meeting the right, gives off the coeliaco-mesenteric artery [to the alimentary canal]. This origin of the coeliaco-mesenteric artery a little to the left, is the only asymmetry (want of balance) in the arterial system of the frog, as contrasted with the very extensive asymmetry of the great vessels near the heart of the rabbit. [Posteriorly the dorsal aorta forks into two common iliac arteries (right and left) supplying the hind limbs.]
Section 10. Figure 3 gives a side view of the frog, to display the circulation.
{Lines from Second Edition only.}
[The venous return to the heart, as in the rabbit, is by paired venae cavae anteriores and by a single vena cava inferior. The factors of the anterior cava on either side are an external jugular (ex.j.) an innominate vein (in.v.) and subclavian (scl.v.). The last receives not only the brachial vein (b.v.) from the fore limb, but also a large vein bringing blood for the skin, the cutaneous (p.v.). The innominate vein has also two chief factors, the internal jugular (l.i.j.v.) and the subscapular (s.s.v.). The blood returns from each hind limb by a sciatic (l.sc.) or femoral (f.m.) vein, and either passes to a renal portal vein (l.r.p.), which breaks into capillaries in the kidney, or by a paired pelvic vein (l.p.v. in Figures 1 and 3) which meets its fellow in the middle line to form the anterior abdominal vein (a.ab.v.) going forward and uniting with the (median) portal vein (p.v.) to enter the liver.]
-The vessels are named in the references to the figure, which should be carefully copied and mastered. Here we need only- [Comparing with the rabbit, we would especially] call attention to the fact that the vena cava inferior extends posteriorly only to the kidney, and that there is a renal portal system. The blood from the hind limbs either flows by the anterior abdominal vein to the portal vein and liver, or it passes by the renal portal vein to the kidney. There the vein breaks up, and we find in the frog's kidney, just as we find in the frog's and rabbit's liver, a triple system of (a) nutritive arterial, (b) afferent* venous and (c) efferent** venous vessels.
* a, ad = to;
** e, ex = out of.
{This Section missing from Second Edition.}
-Section 11. It is not very improbable that the kidney of the frog shares, or performs, some of the functions of the rabbit's liver, or parallel duties, in addition to the simply excretory function. Since specialization of cells must be mainly the relatively excessive exaggeration of some one of the general properties of the undifferentiated cell, it is not a difficult thing to imagine a gradual transition, as we move from one organism to another, of the functions of glands and other cellular organs. It is probable that the mammalian kidney is, physiologically, a much less important (though still quite essential) organ than the structures which correspond to it in position and development in the lower vertebrate types.-
Section 12. The lymphatic system is extensively developed in the frog, but, in the place of a complete system of distinctly organized vessels, there are great lymph sinuses (compare Section 1). In Figure 5, Sheet 12, the position of two lymph hearts (l.h., l.h.) which pump lymph into the adjacent veins, is shown.
Section 13. The skull of the frog will repay a full treatment, and will be dealt with by itself later. The vertebral column (Sheet 12) consists of nine vertebrae, the centra of which have faces, not flat, but hollow in front (pro-coelous), and evidently without epiphyses (compare the Rabbit). The anterior is sometimes called the atlas, but it is evidently not the homologue of the atlas of the rabbit, since the first spinal nerve has a corresponding distribution to the twelfth cranial of the mammal, and since, therefore, it is probable that the mammalian skull = the frog's skull + one (or more) vertebrae incorporated with it. Posteriorly the vertebral column terminates in the urostyle, a calcified unsegmented rod. The vertebrae have transverse processes, but no ribs.
Section 14. The fore-limb (Figure 6, Sheet 12) consists of an upper segment of one bone, the humerus, as in the rabbit; a middle section, the radius and ulna, fused here into one bone, and not, as in the mammalian type, separable; of a carpus, and of five digits, of which the fourth is the longest. The shoulder girdle is more important and complete than that of the higher type. There is a scapula (sc.) with an unossified cartilaginous supra-scapula (s.sc.); the anterior border of the scapula answers to the acromion. On the ventral side a cartilaginous rod, embraced by the clavicle (cl.) (a membrane bone in this type), runs to the sternum, and answers to the clavicle of the rabbit. In the place of the rabbit's coracoid process, is a coracoid bone (co.), which reaches from the glenoid cavity to the sternum; it is hidden on the right side of Figure 6, which is a dorsal view of the shoulder girdle. There is a pre-omosternum (o.st.) and a post-omosternum, sometimes termed a xiphisternum (x.).
Section 15. Figure 7 shows the pelvic girdle and limb of the frog. There is a femur (f.); tibia and fibula (t. and f.) are completely fused; the proximal bones of the tarsus, the astragalus (as.), and calcaneum (cal.) are elongated, there are five long digits, and in the calcar (c.) an indication of a sixth. With considerable modifications of form, the three leading constituents of the rabbit's pelvic girdle occur in relatively identical positions. The greatly elongated ilium (il.) articulates with the single (compare Rabbit) sacral vertebra (s.v. in Figure 5). The ischium (is.) is relatively smaller than in the rabbit, and the pubis (pu.) is a ventral wedge of unossified cartilage. The shape of the pelvic girdle of the frog is a wide departure from that found among related forms. In connection with the leaping habit, the ilia are greatly elongated, and the pubes and ischia much reduced. Generally throughout the air-frequenting vertebrata, we find the same arrangement of these three bones, usually in the form of an inverted. Y-- the ilium above, the ischium and pubis below, and the acetabulum at the junction of the three.
Section 16. The uro-genital organs of the frog, and especially those of the male, correspond with embryonic stages of the rabbit. In this sex the testes (T., Sheet 13) lie in the body cavity, and are white bodies usually dappled with black pigment. Vasa efferentia (v.e.) run to the internal border of the anterior part of the kidney, which answers, therefore, to the rabbit's epididymis. The hinder part of the kidney is the predominant renal organ. There is a common uro-genital duct, into which a seminal vesicle, which is especially large in early spring, opens. This is the permanent condition of the frog. In the rabbit, for urogenital duct, we have ureter and vas deferens; the testes and that anterior part of the primitive kidney, the epididymis, shift back into the scrotal sacs, and the ureters shift round the rectum and establish a direct connection with the bladder, carrying the genital ducts looped over them. The oviducts of the female do not fuse distally to form a median vagina as they do in the rabbit. In front of the genital organ in both sexes is a corpus adiposum (c.ad.), which acts as a fat store, and is peculiar to the frogs and toads. The distal end of the oviduct of the female is in the breeding season (early March) enormously distended with ova, and the ovaries become then the mere vestiges of their former selves. The distal end of the oviduct is, therefore, not unfrequently styled the uterus. There is no penis in the male, fertilisation of the ova occurring as they are squeezed out of the female by the embracing fore limbs of the male. The male has a pad, black in winter, shown in Figure 1, which is closely pressed against the ventral surface of the female in copulation, and which serves as a ready means of distinguishing the sex.
Section 17. The spinal cord has a general similarity to that of the rabbit; the ratio of its size to that of the brain is larger, and the nerves number ten pairs altogether. The first of these (sp. 1, in Figure 2, Sheet -12- ) {First Edition error.} [13] corresponds in distribution with the rabbit's hypoglossal nerve, a point we shall refer to again when we speak of the skull. The second and third constitute the brachial plexus. The last three form the sciatic plexus going to the hind limb.
Section 18. The same essential parts are to be found in the brain of both frog and rabbit, but in the former the adult is not so widely modified from the primitive condition as in the latter. The fore-brain consists of a thalamencephalon (th.c. and 1), which is exposed in the dorsal view of the brain, and which has no middle commissure. The cerebral hemispheres (c.h.) are not convoluted, do not extend back to cover parts behind them, as they do in the rabbit, and are not connected above the roof of the thalamencephalon by a corpus callosum. Moreover, the parts usually regarded, as the olfactory lobes (rh.) fuse in the middle line. The mid-brain gives rise to the third nerve, and has the optic lobes on its dorsal side, but these are hollow, and they are not subdivided by a transverse groove into corpora quadrigemina, as in the rabbit. In the hind-brain the cerebellum is a mere band of tissue without lateral lobes or flocculi, and the medulla gives origin only to nerves four to ten; there is no eleventh nerve, and the hypoglossal is the first spinal-- from which it has been assumed that the rabbit's medulla equals that of the frog, plus a portion of the spinal cord incorporated with it. The hypoglossal is very distinctly seen on opening the skin beneath the hyoid plate.
Section 19. The first, second, third, and fourth cranial nerves of the frog correspond with those of the rabbit in origin and distribution. So do five, six and eight. The seventh nerve forks over the ear-drum-- the larger branch emerging behind it and running superficially, as shown in Figure 4. There is also a deeper palatine branch of VII. (P.) running under V2 and V3 below the orbit, and to be seen together with V1 and V2 after removal of the eyeball. The ninth nerve similarly forks over the first branchial slit of the tadpole, and evidence of the fork remains in the frog. It is seen curving round anterior to the hypoglossal nerve, and lying rather deeper in dissection. The vagus (tenth) nerve is distributed to heart, lungs, and viscera-- in the tadpole it also sends for forking branches over the second, third, and fourth branchial slits. It lies deeper than IX., and internal to the veins, and runs close beside the cutaneous artery. Most of these nerves are easily dissected and no student should rest satisfied until he has actually seen them.
Section 20. The sympathetic chain is closely connected with the aorta. It is, of course, paired, and is easily found in dissection by lifting the dorsal aorta and looking at its mesentery. In the presence of ganglia corresponding to the spinal nerves, and of rami communicantes, it resembles that of the rabbit.
Section 21. The whole of this chapter is simply a concise comparison, of frog and rabbit. In addition to reading it, the student should very carefully follow the annotations to the figures, and should copy and recopy these side by side with the corresponding diagrams of the other types.
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