Text Book of Biology Part 8

Section 5. The lungs of the frog are bag-like; shelves and spongy part.i.tions 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 ma.s.s.

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 ma.s.s. 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 [furnis.h.i.+ng 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 r.e.t.a.r.ds 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 ma.s.s 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 pa.s.ses 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 pa.s.ses 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 ca.n.a.l]. 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 pa.s.ses 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 pa.s.ses 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 h.o.m.ologue 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 const.i.tuents 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 p.u.b.es 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 s.e.x 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, s.h.i.+ft back into the scrotal sacs, and the ureters s.h.i.+ft round the r.e.c.t.u.m 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 v.a.g.i.n.a as they do in the rabbit. In front of the genital organ in both s.e.xes 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 p.e.n.i.s 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 distinguis.h.i.+ng the s.e.x.

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 const.i.tute 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 a.s.sumed 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.

2. _The Skull of the Frog (and the vertebrate skull generally)_

Section 22. We have already given a description of the mammalian skull, and we have stated where the origin of the several bones was in membrane, and where in cartilage; but a more complete comprehension of the mammalian skull becomes possible with the handling of a lower type. We propose now, first to give some short account of the development and structure of the skull of the frog, and then to show briefly how its development and adult arrangement demonstrate the mammalian skull to be a fundamentally similar structure, complicated and disguised by further development and re-adjustment.

Section 23. Figure 1,I. Sheet 14, shows a dorsal view of a young tadpole cranium; the brain has been removed, and it is seen that it was supported simply upon two cartilaginous rods, the trabeculae cranii (tr.c.). Behind these trabeculae comes the notochord (n.c.), and around its anterior extremity is a paired tract of cartilage, the parachordals (p.c.). These structures, underlying the skull, are all that appear[s] at first of the brain box. In front, and separate from the cranium, are the nasal organs (n.c.); the eyes lie laterally to the trabeculae, and laterally to the parachordals are two tracts of cartilage enclosing the internal ear, the otic capsules.

Section 24. Figure 1, II., is a more advanced, phase of the same structures. The trabeculae have met in front and sent forward a median (c.t.) and lateral parts (a.o.) to support the nasal organs. They have also flattened, out very considerably, and have sent up walls on either side of the brain to meet above it and form an incomplete roof (t.) over it. The parachordals have similarly grown up round, the hind-brain and formed a complete ring, the roof of which is indicated, by b. Further, the otic capsules are fusing with the brain-case. With certain differences of form these elements-- the trabeculae, the parachordals, and the otic capsules, are also the first formed structures of the mammalian cranium.

Section 25. In Figures 1,I. and II., there appears beneath the eye a bar of cartilage (p.p.), the palato-pterygoid cartilage, which is also to be seen from the side in Figures 8,I. and III. It will be learnt from these latter that this bar is joined in front to the cranium behind the nasal organ, and behind to the otic capsule. The cartilaginous bar from the palato-pterygoid to the otic capsule is called the quadrate, and at the point of junction, at the postero-ventral angle of the palato-pterygoid, articulates with the cartilaginous bar which is destined to form the substratum of the lower jaw-- Meckel's cartilage (M.c. in Figure 8,I.).

Section 26. Figure 2 shows a dorsal view of these structures in a young frog. The parts corresponding to these in 1,II. will be easily made out, but now ossification has set in at various points of this cartilaginous cranium. In front of the otic capsule is the paired pro-otic bone (p.o.); behind it at the sides of the parachordal ring is the paired ex-occipital (e.o.); in front of the cranium box, and behind the nasal capsules, is a ring of bone, the (median, but originally paired) sphenethmoid (s.e.). -A paired ossification appears in the palato-pterygoid cartilage the pterygoid bone (pt.), while- A splint of bone, the quadrato-jugal, appears at the angle of articulation with the lower jaw. These are all the cartilage bones that appear in the cranium and upper jaw of the frog.

Section 27. But another series of bones, developed first chiefly in dermal connective tissue, and coming to plate over the cranium of cartilage, are not shown in Figure 2. They are, however, in Figure 3.

These membrane bones are: along the dorsal middle line, the parieto-frontals (p.f.), originally two pairs of bones which fuse in development, and the nasals (na.). Round the edge of the jaw, and bearing the teeth, are pre-maxillae (p.m.), and maxillae (mx.), and overlying the quadrate cartilage and lateral to the otic capsules are the T-shaped squamosal bones (sq.). In the ventral view of the skull (Figure 4) we see a pair of vomers (vo.) bearing teeth, a pair of palatines (pal.), [and a pair of pterygoids (pt.)] (which [palatines and pterygoids, we may note,] unlike those of the rabbit, are -stated to be- membrane bones), and a great median dagger-shaped para-sphenoid (p.sp.). These two Figures, and 5, which shows the same bones in side view, should be carefully mastered before the student proceeds with this chapter. The cartilage bones are distinguished from membrane bones by cross-shading.

Section 28. Turning now to Figure 8,I., we have a side view of a tadpole's skull. On the ventral side of the head is a series of vertical cartilaginous bars, the visceral arches supporting the walls of the tadpole's gill slits. The first of these is called the hyoid arch (c.h.), and the four following this, the first (br.1), second, third, and fourth (br.4), branchial arches. Altogether there are four gill slits and between the hyoid arch and the jaw arch, as it is called (= Meckel's cartilage + the palato-pterygoid), is "an imperforate slit," which becomes the ear-drum.* The frog no longer breathes by gills, but by lungs, and the gills are lost, the gill slits closed, and the branchial arches consequently much reduced. Figures 8, II., and 8, III., show stages in this reduction. The hyoid arch becomes attached, to the otic capsule, and its median ventral plate, including also the vestiges of the first, second, and fourth branchial arches, is called the hyoid apparatus. In Figure 5, the apparatus is seen from the side; c.h. is called the (right) anterior cornu** of the hyoid. The function of the hyoid apparatus in the frog is to furnish, a basis of attachment to the tongue muscles; it remains cartilaginous, with the exception of the relic of one branchial arch, which ossifies as the thyro-hyal (Figure 7 th.h.). It will be noted that, as development proceeds, the angle of the jaw swings backward, and the hyoid apparatus, s.h.i.+fts relatively forward. These changes of position are indicated in Figure 8, III., by little arrow-heads.

* We may note here that, comparing the ear of the frog with that of the rabbit, there is no external ear. There is, moreover, no bulla supporting the middle ear, and the tympanic membrane stretches between the squamosal in front and the anterior cornu of the hyoid behind. A rod-like columella auris replaces the chain of ear ossicles, and may, or may not, answer to the stapes alone, or even possibly to the entire series. In the internal ear there is no cochlea, and the otic ma.s.s is largely cartilaginous instead of entirely bony.

** Plural cornua.

Section 29. Before proceeding to the comparison of the mammalian skull with this, we would strongly recommend the student thoroughly to master this portion of the work, and in no way can he do this more thoroughly and quickly than by taking a parboiled frog, picking off the skin, muscle, and connective tissue from its skull, and making out the various bones with the help of our diagrams.

Section 30. Figure 9 represents, in the most diagrammatic way, the main changes in form of the essential const.i.tuents of the cranio-facial apparatus, as we pa.s.s from the amphibian to the mammalian skull. F.

is the frog from the side and behind; b.c. is the brain-case, o.c. the otic capsule, e. the eye, n.c. the nasal capsule, p.p. the palato-pterygoid cartilage, mx. the maxillary membrane bones, sq.

the squamosal, and mb. the mandible. The student should compare with Figure 5, and convince himself that he appreciates the diagrammatic rendering of these parts. Now all the distinctive differences in form, from this of the dog's skull (D.), are reducible to two primary causes--

(1) The brain is enormously larger, and the brain-case is vastly inflated, so that--