-The Rabbit._
1. _External Form and General Considerations._
Section 1. It is unnecessary to enter upon a description of the appearance of this familiar type, but it is not perhaps superfluous, as we proceed to consider its anatomy, to call attention to one or two points in its external, or externally apparent structure. Most of our readers know that it belongs to that one of two primary animal divisions which is called the vertebrata, and that the distinctive feature which place it in this division is the possession of a spinal column or backbone, really a series of small ring-like bones, the vertebrae ([Figure 1 v.b.]) strung together, as it were, on the main nerve axis, the spinal cord ([Figure 1 s.c.]). This spinal column can be felt along the neck and back to the tail. This tail is small, tilted up, and conspicuously white beneath, and it serves as a "recognition mark" to guide the young when, during feeding, an alarm is given and a bolt is made for the burrows. In those more primitive (older and simpler-fashioned) vertebrata, the fishes, the tail is much large and far more important, as compared with the rest of the body, than it is in most of the air-inhabiting vertebrates. In the former it is invariably a great muscular mass to propel the body forward; in the latter it may disappear, as in the frog, be simply a feather-bearing stump, as in the pigeon, a fly flicker, as in the cow or horse, a fur cape in squirrel, or be otherwise reduced and modified to meet special requirements.
Section 2. At the fore end, or as English zoologists prefer to say, anterior end, of the vertebral column of the rabbit, is of course the skull, containing the anterior portion of the nerve axis, the brain ([Figure 1 br.]). Between the head and what is called "the body," in the more restricted sense of the word, is the neck. The neck gives freedom of movement to the head, enables the animal to look this way and that, to turn its ears about to determine the direction of a sound, and to perform endless motions in connexion with biting and so forth easily. We may note that in types which swim through the water, the neck dose not appear-- in the fish and frog, for instance-- and the head simply widens out as one passes back to the body. The high resistance offered by water necessitates this tendency to a cigar or ship outline, just as it has determined the cigar shape of the ordinary fish torpedo.
Section 3. In the body of the rabbit, as examined from the outside, we can make out by feeling two distinct regions, just as we might in the body of a man; anteriorly a bony cage, having the ribs at the sides, a rod-like bone in the front, the sternum ([Figure 1 -st.-, [stm.]), and the backbone behind, and called the chest or thorax; and posteriorly a part called the abdomen, which has no bony protection over its belly, or ventral surface. These parts together with the neck constitute the trunk. As a consequence of these things, in the backbone of the rabbit there are four regions: the neck, or cervical part, consisting of seven vertebrae, the thoracic part of twelve joined to ribs, the abdominal (also called the lumbar) region of seven without ribs, and the tail or caudal of about fifteen. Between the lumbar and caudal come four vertebrae, the sacral, which tend to run together into a bony mass as the animal grows old, and which form a firm attachment for the base of the hind limb.
Section 4. The thorax and abdomen are separated by a partition, the diaphragm ([Figure 1 dia.]). This structure is distinctive of that class of the vertebrata called mammals, and which includes man, most of the larger and commoner land animals, and whales and manatee. We shall find later that it is essentially connected with the perfection of the air breathing to which this group has attained. Another characteristic shared by all mammals, and by no other creature, is the presence of hair. In birds we have an equally characteristic cover in the feathers, the frog is naked, and the fishes we find either naked skins or scales.
Section 5. The short strong fore limbs are adapted to the burrowing habit, and have five digits; the hind limbs are very much longer and muscular, enable the animal to progress rapidly by short leaps, and they have four toes. If the student thinks it worth while to attempt to remember the number of digits-- it is the fault of examiners if any value dose attach to such intrinsically valueless facts-- he should associate the number 54 (5 in front, 4 behind) with the rabbit, and observe that with the frog the reverse is the case.
Section 6. We may note here the meaning of certain terms we shall be constantly employing. The head end of the rabbit is anterior, the tail end posterior, the backbone side of the body-- the upper side in life-- is dorsal, the breast and belly side, the lower side of the animal, is ventral. If we imagine the rabbit sawn asunder, as it were, by a plane passing through the head and tail, that would be the median plane, and parts on either side of it are lateral, and left or right according as they lie to the animal's left or right. In a limb, or in the internal organs, the part nearest the central organ, or axis, is proximal, the more remote or terminal parts are distal. For instance, the mouth is anteriorly placed, the tongue on its ventral wall; the tongue is median, the eyes are lateral, and the fingers are distal to the elbow. The student must accustom himself to these words, and avoid, in his descriptions, the use of such terms as "above," "below," "outside," which vary with the position in which we conceive the animal placed.
Section 7. So much for the general form; we may note a few facts of general knowledge, in connection with the rabbit's life-activity. In a day of the rabbit's life a considerable amount of work is done-- the animal runs hither and thither, for instance; in other words, a certain mass of matter is moved through space, and for that we know force must be exerted. Whence comes the force?
Section 8. We find the rabbit occupies a considerable amount of its time in taking in vegetable matter, consisting chiefly of more or less complex combustible and unstable organic compounds. It is a pure vegetarian, and a remarkably moderate drinker. Some but only a small proportion, of the vegetable matter it eats, leaves its body comparatively unchanged, in little pellets, the faeces, in the process of defaecation. For the rest we have to account.
Section 9. We find, also, that the rabbit breathes air into its lungs, which is returned to the atmosphere with a lessened amount of oxygen, and the addition of a perceptible amount of carbon dioxide. The rabbit also throws off, or excretes, a fluid, the urine, which consists of water with a certain partially oxydised substance containing nitrogen, and called urea, and other less important salts. The organs within the body, by which the urine is separated, are called the kidneys.
Section 10. Repeating these facts in other words, the rabbit takes into its body complex and unstable organic compounds containing nitrogen, carbon, hydrogen, a certain amount of oxygen, a small quantity of sulphur, and still smaller amounts of other elements. It also breathes in oxygen.
Section 11. It returns a certain rejected part of its food comparatively unchanged. Besides this, it returns carbon dioxide and water, which are completely oxydised, and very simple and stable bodies, and urea-- a less completely oxydised compound, but a very simple one compared with the food constituents.
Section 12. Now the chemist tells us that when a stable body is formed, or when an unstable compound decomposes into simpler stable ones, force is evolved. The oxydation of carbon, for instance, in the fireplace, is the formation of the stable compound called carbon dioxide, and light and heat are evolved. The explosion of dynamite, again is the decomposition of an unstable compound. Hence, we begin to perceive that force-- the vital force-- which keeps the rabbit moving, is supplied by the decomposition and partial oxydation of compounds continued in its food, to carbon dioxide, water, urea, and smaller quantities of other substances.
Section 13. This is the roughest statement of the case possible, but it will give the general idea underlying our next chapters. We shall consider how the food enters the body and is taken up into the system, how it is conveyed to the muscles in the limbs, to the nerve centres, and to wherever work is done, to be there decomposed and partially oxydised, and finally how the products of its activity-- the katastases, of which the three principal are carbon dioxide, water, and urea-- are removed from the body.
Section 14. There are one or two comparatively modern terms that we may note here. This decomposition of unstable chemical compounds, releasing energy, is called kataboly. A reverse process, which has a less conspicuous part in our first view of the animal's life action, by which unstable compounds are built up and energy stored, is called anaboly. The katastases are the products of kataboly.
Section 15. In an ordinary animal, locomotion and other activity predominate over nutritive processes, which fact we may express, in the terms just given, by saying that kataboly prevails over anaboly. An animal, as we have just explained, is an apparatus for the decomposition and partial oxydation of certain compounds, and these are obtained either directly or indirectly-- through other animals, in the case of meat-eaters-- from the vegetable kingdom. As the student will learn early in his botanical reading, the typical plant has, in its green colouring matter, chlorophyll, a trap to catch the radiating energy of the sun, and to accomplish, by the absorption of that energy, the synthesis (building up) of those organic compounds which the animal destroys. The typical plant is, on whole, passive and synthetic, or anabolic; the typical animal, active and katabolic; and the excess of kataboly over anaboly in the animal is compensated for by the anabolic work stored up, as it were, by the plant, which is, directly or indirectly, the animal's food.
2. _The Alimentary Canal of the Rabbit_
Section 16. [Figure 1] represents the general anatomy of the rabbit, but is especially intended to show the alimentary (= food) canal, shortened to a certain extent, and with the proportions altered, in order to avoid any confusing complications. It is evidently simply a coiled tube-- coiled for the sake of packing-- with occasional dilatations, and with one side-shunt, the caecum (cae.), into which the food enters, and is returned to the main line, after probably absorbent action, imperfectly understood at present. A spiral fold in this cul-de-sac {bottom-of-sack}, which is marked externally by constrictions, has a directive influence on the circulation of its contents. The student should sketch Figure 1 once or twice, and make himself familiar with the order and names of the parts before proceeding. We have, in succession, the mouth (M.), separated from the nasal passage (Na.) above the palate; the pharynx (ph.), where the right and left nasal passages open by the posterior nares into the mouth; the oesophagus (oes.); the bag-like stomach, its left ([Section 6]) end being called the cardiac (cd.st.), and its right the pyloric end (py.); the U-shaped duodenum (ddnm.) and the very long and greatly coiled ileum (il.). The duodenum and ileum together form the small intestine; and the ileum is dilated at its distal end into a thick-walled sacculus rotundus (s.r.), beyond which point comes the large intestine. The colon (co.) and rectum (r.) continue the main line of the alimentary canal; but, at the beginning of the large intestine, there is also inserted a great side-shunt, the caecum (cae.), ending blindly in a fleshy vermiform appendix (v.ap.). The figure will indicate how the parts are related better than any verbal description can. Between the coiling alimentary tube and the body walls is a space, into which the student cuts when he begins dissecting; this is the peritoneal cavity (pt.). A thin, transparent membrane, the mesentery, holds the intestines in place, and binds them to the dorsal wall of this peritoneal space.
Section 17. The food stuffs of an animal, the unstable compounds destined ultimately to be worked into its life, and to leave it again in the form of katastases ([Section 13]), fall into two main divisions. The first of these includes the non-nitrogenous food stuffs, containing either carbon together with hydrogen and oxygen in the proportion of H2O (the carbo-hydrates), or carbon and hydrogen without oxygen (the hydrocarbons). The second division consists of the nitrogenous materials, containing also carbon, hydrogen, a certain amount of oxygen, sulphur, and possibly other elements. Among the carbohydrates, the commonest are starch and cellulose, which are insoluble bodies, and sugar, which is soluble. The hydrocarbons, fats, oils, and so on, form a comparatively small proportion of the rabbit's diet; the proverb of "oil and water" will remind the student that these are insoluble. The nitrogenous bodies have their type in the albumen of an egg; and muscle substance and the less modified living "protoplasm" of plants, a considerable proportion of the substance of seeds, bulbs, and so on, are albuminous bodies, or proteids. These also are insoluble bodies, or when soluble, will not diffuse easily through animal membranes.
Section 18. Now the essential problem which the digestive canal of the rabbit solves is to get these insoluble, or quasi-insoluble, bodies into its blood and system. They have to pass somehow into the circulation through the walls of the alimentary canal. In order that a compound should diffuse through a membrane, it must be both soluble and diffusible, and therefore an essential preliminary to the absorption of nutritive matter is its conversion into a diffusible soluble form. This is effected by certain fluids, formed either by the walls of the alimentary canal or by certain organs called glands, which open by ducts into it; all these fluids contain small quantities of organic compounds of the class called ferments, and these are the active agents in the change. The soluble form of the carbohydrates is sugar; proteids can be changed into the, of course, chemically equivalent but soluble and diffusible the peptones; and fats and oils undergo a more complicated, but finally similar change.
Section 19. We shall discuss the structure and action of -a gland- [glands] a little more fully in a subsequent chapter. Here we will simply say that they are organs forming each its characteristic fluid or secretion, and sending it by a conduit, the duct, to the point where its presence is required. The saliva in our mouths, tears, and perspiration, are examples of the secretions of glands.
Section 20. In the month of the rabbit the food is acted upon by the teeth and saliva. The saliva contains ptyalin, a ferment converting starch into sugar, and it also serves to moisten the food as it is ground up by the cheek teeth. It does not act on fat to any appreciable extent. The teeth of the rabbit are shown in Figure XVIII., [Sheet 4]. The incisor teeth in front, two pairs above and one pair below (i.), are simply employed in grasping the food; the cheek teeth-- the premolars (pm.) and molars (m.) behind-- triturate the food by a complicated motion over each. Their crowns are flat for this purpose, with harder ridges running across them.
Section 21. This grinding up of the food in the mouth invariably occurs in herbivorous animals, where there is a considerable amount of starch and comparatively little hydrocarbon in the food. By finely dividing the food, it ensures its intimate contact with the digestive ferment, ptyalin. In such meat-eaters as the cat and dog, where little starchy matter and much fat is taken, the saliva is, of course, of less importance, and this mastication does not occur. The cheek teeth of a dog ({[Section 91]}), and more so of a cat, are sharp, and used for gnawing off fragments of food, which are swallowed at once. Between the incisors and premolars of a dog come the characteristic biting teeth, or canines, absent in the rabbit.
Section 22. The student will probably ask why the cheek teeth, which are all similar in appearance, are divided into premolars and molars. The rabbit has a set of milk molars-- a milk dentition-- which are followed by the permanent teeth, just as in man. Those cheek teeth of the second set, which have predecessors in the first series, are called premolars; the ones posterior to these are the molars.
Section 23. After mastication, the food is worked by the tongue and cheeks into a saliva-soaked "bolus" and swallowed. The passage down the oesophagus is called deglutition. In the stomach it comes under the influence of the gastric juice, formed in little glandular pits in the stomach wall-- the gastric (Figure VIII. [Sheet 3]) and pyloric glands. This fluid is distinctly acid, its acidity being due to about one-tenth per cent {of a hundred} of hydrochloric acid, and it therefore stops any further action of the ptyalin, which can act only on neutral or slightly alkaline fluids. The gastric juice does not act on carbo-hydrates or hydrocarbons to any very noticeable degree. Its essential property is the conversion of proteids into peptones, and the ferment by which this is effected is called pepsin. Milk contains a peculiar soluble proteid, called casein, which is precipitated by a special ferment, the rennet-ferment, and the insoluble proteid, the curd, thus obtained is then acted on by the pepsin. In the manufacture of cheese, the rennetferment obtained, from the stomach of a calf is used to curdle the milk.
Section 24. After the food has undergone digestion in the stomach it passes into the duodenum, the U-shaped loop of intestine immediately succeeding the stomach. The duodenum is separated from the stomach by a ring-like muscular valve, the pylorus; this valve belongs to the class of muscles called sphincters, which, under ordinary circumstances, are closed, but which relax to open the circular central aperture. The valve at the anus, which retains the faeces, is another instance of a sphincter.
Section 25. The food at this stage is called chyme; it is an acid and soup-like fluid-- acid through the influence of the gastric juice. The temperature of the animal's body is sufficiently high to keep most of the fat in the food melted and floating in oily drops; much of the starch, has been changed to sugar, and the solid proteids to soluble peptones, but many fragments of material still float unchanged.
Section 26. It meets now with the bile, a greenish fluid secreted by that large and conspicuous gland the liver. The bile is not simply a digestive secretion, like the saliva or the gastric juice; it contains matters destined to mix in, and after a certain amount of change to be passed out of the body with, the faeces; among these substances, of which some portion is doubtless excretory, are compounds containing sulphur-- the bile salts. There is also a colouring matter, bili verdin, which may possibly also be excretory. If the student will compare Sections [10] and [11], he will notice that in those paragraphs no account is taken of the sulphur among the katastases, the account does not balance, and he will at once see that here probably is the missing item on the outgoing side. The bile, through the presence of these salts, is strongly alkaline, and so stops the action of the gastric juice, and prepares for that of the pancreas, which can act only in an alkaline medium. The fermentive action of the bile is trifling; it dissolves fats, to a certain extent, and is antiseptic, that is, it prevents putrefaction to which the chyme might be liable; it also seems to act as a natural purgative.
Section 27. The bile, as we shall see later, is by no means the sole product of the liver.
Section 28. The pancreatic juice, the secretion of the pancreas is remarkable as acting on all the food stuffs that have not already become soluble. It emulsifies fats, that is, it breaks, the drops up into extremely small globules, forming a milky fluid, and it furthermore has a fermentive action upon them; it splits them up into fatty acids, and the soluble body glycerine. The fatty acids combine with alkaline substances ([Section 26]) to form bodies which belong to the chemical group of Soaps, and which are soluble also. The pancreatic juice also attacks any proteids that have escaped the gastric juice, and converts them into peptones, and any residual starch into sugar. Hence by this stage, in the duodenum, all the food constituents noticed in [Section 17] are changed into soluble forms. There are probably, three distinct ferments in the pancreatic juice acting respectively on starch, fat, and proteid, but they have not been isolated, and the term pancreatin is sometimes used to suggest the three together.
Section 29. A succus entericus, a saliva-like fluid secreted by numerous small glands in the intestine wall (Brunner's glands, Lieberkuhnian follicles), probably aids, to an unknown but comparatively small extent, in the digestive processes.
Section 30. The walls of the whole of the small intestine are engaged in the absorption of the soluble results of digestion. In the duodenum, especially, small processes, the villi project into the cavity, and being, like the small hairs of velvet pile, and as thickly set, give its inner coat a velvety appearance. In a villus we find (Figure IX., [Sheet 3]) a series of small blood-vessels and with it another vessel called a lacteal. The lacteals run together into larger and larger branches until they form a main trunk, the thoracic duct, which opens into the blood circulation at a point near the heart; but of this we shall speak further later. They contain, after a meal, a fluid called chyle.
Section 31. Emulsified fats pass into the chyle. Water and diffusible salts certainly pass into the vein. The course taken by the peptones is uncertain, but Professor Foster favours the chyle in the case of the rabbit-- the student should read his Text-book of Physiology, Part 2, Chapter 1, Section 11, if interested in the further discussion of this question.
Section 32. The processes that occur in the remaining portions of the alimentary canal are imperfectly understood. The caecum is so large in the rabbit that it must almost certainly be of considerable importance. In carnivorous animals it may be so much reduced as to be practically absent. An important factor in the diet of the herbivorous animals, and one absent from the food of the carnivora, is that carbohydrate, the building material of all green-meat- [food], cellulose, and there is some ground for thinking that the caecum is probably a region of special fermentive action upon it. The pancreatic juice, it may be noted, exercises a slight digestive activity upon this substance.
Section 33. Water is most largely absorbed in the large intestine, and in it the rejected (mainly insoluble) portion of the food gradually acquires its dark colour and other faecal characteristics.
3. _The Circulation_
Section 34. The next thing to consider is the distribution of the food material absorbed through the walls of the alimentary canal to the living and active parts of the body. This is one of the functions of the series of structures-- heart and blood-vessels, called the circulation, circulatory system, or vascular system. It is not the only function. The blood also carries the oxygen from the lungs to the various parts where work is done and kataboly occurs, and it carries away the katastases to the points where they are excreted-- the carbon dioxide and some water to the lungs, water and urea to the kidneys, sulphur compounds of some kind to the liver.
Section 35. The blood (Figure 4, [Sheet 2]) is not homogeneous; under the low power of the microscope it may be seen to consist of--
(1.) a clear fluid, the plasma, in which float--
(2.) a few transparent colourless bodies of indefinite and changing shape, and having a central brighter portion, the nucleus with a still brighter dot therein the nucleolus-- the white corpuscles (w.c.), and
(3.) flat round discs, without a nucleus, the red corpuscles (r.c.), greatly more numerous than the white.
Section 36. The chyle of the lacteals passes, as we have said, by the thoracic duct directly into the circulation. It enters the left vena cava superior (l.v.c.s.) near where this joins the jugular vein (ex.j.) (see Figure 1, [Sheet 2], th.d.) and goes on at once with the rest of the blood to the heart. The small veins of the villi, however, which also help suck up the soluble nutritive material, are not directly continuous with the other body veins, the systemic veins; they belong to a special system, and, running together into larger and larger branches, form the lieno gastric (l.g.v.) and mesenteric (m.v.) veins, which unite to form the portal vein (p.v.) which enters the liver (l.v.) and there breaks up again into smaller and smaller branches. The very finest ramifications of this spreading network are called the (liver) capillaries, and these again unite to form at last the hepatic vein (h.v.) which enters the vena cava inferior (v.c.i.), a median vessel, running directly to the heart. This capillary network in the liver is probably connected with changes requisite before the recently absorbed materials can enter the general blood current.
Section 37. The student has probably already heard the terms vein and artery employed. In the rabbit a vein is a vessel bringing blood towards the heart, while an artery is a vessel conducting it away. Veins are thin-walled, and therefore flabby, a conspicuous purple when full of blood, and when empty through bleeding and collapsed sometimes difficult to make out in dissection. They are formed by the union of lesser factors. The portal breaks up into lesser branches within the liver. Arteries have thick muscular and elastic walls, thick enough to prevent the blood showing through, and are therefore pale pink or white and keep their round shape.
Section 38. The heart of the rabbit is divided by partitions into four chambers: two upper thin-walled ones, the auricles (au.), and two lower ones, both, and especially the left, with very muscular walls, the ventricles (vn.). The right ventricle (r.vn.) and auricle (r.au.) communicate, and the left ventricle (l.vn.) and auricle (l.au.).
Section 39. The blood coming from all parts of the body, partly robbed of its oxygen and containing much carbon dioxide and other katastases, enters the right auricle of the heart through three great veins, the median vena cava inferior from the posterior parts of the body, and the paired venae cavae superiores from the anterior. With the beating of the heart, described below, it is forced into the right ventricle and from there through the pulmonary artery (p.a.) seen in the figure passing under the loop of the aorta (ao.) to the lungs.
Section 40. The lungs (lg. Figure 1, [Sheet 1]) are moulded to the shape of the thoracic cavity and heart; they communicate with the pharynx by the trachea (tr. in Figure 1, [Sheet 1]) or windpipe, and are made up of a tissue of continually branching and diminishing air-tubes, which end at last in small air-sacs, the alveoli. The final branches of the pulmonary arteries, the lung capillaries, lie in the walls of these air-sacs, and are separated from the air by an extremely thin membrane through which the oxygen diffuses into, and the carbon dioxide escapes from, the blood.
Section 41. The mechanism of respiration will be understood by reference to Figure 3, [Sheet 2]. It will be noted, in dissecting that the lungs have shrunk away from the walls of the thorax; this collapse occurs directly an aperture is made in the thorax wall, and is in part due to their extreme elasticity. In life the cavity of the thorax forms an air-tight box, between which and the lungs is a slight space, the pleural cavity (pl.c.) lined by a moist membrane, which is also reflected, over the lungs. The thorax wall is muscular and bony, and resists the atmospheric pressure on its outer side, so that the lungs before this is cut through are kept distended to the size of the thoracic cavity by the pressure of the air within them. In inspiration (or breathing-in) the ribs are raised by the external intercostal (Anglice, between-ribs, e.i.c.m.) and other allied muscles, and the diaphragm (dia.) contracts and becomes flatter; the air is consequently sucked, in as the lungs follow the movement of the thorax wall. In expiration the intercostals and diaphragm relax and allow the elastic recoil of the lungs to come into play. The thoracic wall is simultaneously depressed by the muscles of the abdominal area, the diaphragm thrust forwards, as the result of the displacement and compression of the alimentary viscera thus brought about. (r.r.r. in the [Figure] mark ribs.)
Section 42. The oxygen and carbon dioxide are not carried in exactly the same way by the blood. The student will know from his chemical reading that neither of these gases is very soluble, but carbon dioxide is sufficiently so in an alkaline fluid to be conveyed by the liquid plasma. The oxygen however, needs a special portative mechanism in the colouring matter of the red corpuscles, the haemoglobin, with which it combines weakly to form oxy-haemoglobin of a bright red colour, and decomposing easily in the capillaries (the finest vessels between the arteries and veins), to release the oxygen again. The same compound occurs in all true vertebrata, and in the blood-fluid of the worm; in the crayfish a similar substance, haemocyanin, which when oxygenated is blue, and when deoxydized colourless, discharges the same function.
Section 43. The blood returns from the lungs to the left auricle (l.au.) by the pulmonary veins, hidden in the [Figure] by the heart, passes thence to the thick-walled left ventricle (l.vn.), and on into the aorta (ao.).
Section 44. The beating of the heart is, of course, a succession of contractions and expansions of its muscular wall. The contraction, or systole, commences at the base of the venae cavae and passes to the auricles, driving the blood before it into the ventricles, which then contract sharply and drive it on into the aorta or pulmonary artery; a pause and then a dilatation, the diastole follows. The flow of the blood is determined in one direction by the various valves of the heart. No valves occur in the opening of the superior cavae but an imperfect one, the Eustachian valve, protects the inferior cava; the direction of the heart's contraction prevents any excessive back-flow into the veins, and the onward, tendency is encouraged by the suck of the diastole of the ventricles. Between the left ventricle and auricle is a valve made up of two flaps of skin, the mitral valve, the edges of the flaps being connected with the walls of the ventricle through the intermediation of small muscular threads, the chordae tendinae, which stretch across its cavity to little muscular pillars, the papillary muscles; these attachments prevent the mitral valve from flapping back into the auricle, and as the blood flows into and accumulates in the ventricle it gets behind the flaps of the valve and presses its edges together. When the systole of the ventricle occurs, the increased, tension of the blood only closes the aperture the tighter, and the current passes on into the aorta, where we find three watch-pocket valves, with the pocket turned away from the heart, which are also closed and tightened by any attempt at regurgitation (back-flow). A similar process occurs on the right side of the heart, but here, instead of a mitral valve of two flaps between auricle and ventricle, we have a tricuspid valve with three. The thickness of the muscular walls, in view of the lesser distance through which it has to force the blood, -are- [is] less for the right ventricle than the left.
Section 45. The following are the chief branches of the aorta. The student should be able to follow them with certainty in dissection; they are all displayed in the [Figure]; but it must not be imagined for a moment that familiarity with this diagram will obviate the necessity for the practical work; (in.) is the innominate artery; it forks into (s.cl.a.) the right subclavian, and (r.c.c.) the right common carotid. Each carotid splits at the angle of the jaw into an internal and an external branch. The left common carotid, (l.c.c.) arises from the base of the innominate,* (l.s.cl.a.) the left subclavian, directly from the aorta. The aorta now curves round to the dorsal middle line, and runs down as seen in Figure 1, [Sheet 1] (d.ao.) and Figure 1, [Sheet 2] (d.ao.). Small branches are given off to the ribs, and then comes the median coeliac (coe.a.) to the stomach and spleen, the median superior mesenteric (s.mes.a.) to the main portion of the intestine, and the inferior mesenteric (p.m.a.) to the rectum. Note that no veins to the inferior vena cava correspond to these arteries-- the blood they supply going back by the portal vein (p.v.). The paired renal arteries (r.a.) supply the kidneys, and the common iliacs (c.il.a.) the hind legs, splitting into the internal iliacs (i.il.a.) and the femoral (f.).
{Lines from Second Edition only.}
[The student should note that the only arteries in the middle line are those supplying the alimentary canal.]
{Lines from First Edition only.}
* -The figure is inaccurate, and represents the left common carotid as arising from the aortic arch.-
Section 46. The distribution of the veins of the rabbit has only a superficial parallelism with arteries. The chief factors of vena cava inferior are the hepatic vein (h.v.), which receives the liver blood, the renal veins (r.v.), from the kidneys, the ilaeo-lumbar, from the abdominal wall, and the external (e.il.v.) and internal ilias (i.il.v.); with the exception of the renal veins none of these run side by side with arteries. The superior cavae (r. and l.v.c.s.) are formed by the union of internal (i.j.) and external jugular (e.j.) veins with a subclavian (s.cl.v.) from the fore limb. The term pre-caval vein is sometimes used for superior cava. The attention, of the student is called to the small azygos vein (az.) running into the right vena cava superior, and forming the only asymmetrical (not-balancing) feature of the veins in front of the heart; it brings blood back from the ribs of the thorax wall, and is of interest mainly because it answers to an enormous main vessel, the right post-cardinal sinus, in fishes. There are spermatic arteries and veins (s.v. and a.) to the genital organs. All these vessels should be patiently dissected out by the student, and drawn.
Section 47. Between the final branches of the arteries and the first fine factors of the veins, and joining them, come the systemic capillaries. These smallest and ultimate ramifications of the circulation penetrate every living part of the animal, so that if we could isolate the vascular system we should have the complete form of the rabbit in a closely-meshed network. It is in the capillaries that the exchange of gases occurs and that nutritive material passes out to the tissues and katastases in from them; they are the essential factor in the circulatory system of the mammal-- veins, arteries, and heart simply exist to remove and replace their contents. The details of the branching of the pulmonary artery and the pulmonary veins need not detain us now.
Section 48. Summarising the course of the circulation, starting from the right ventricle, we have-- pulmonary artery, pulmonary capillaries, pulmonary vein, left auricle, left ventricle, aorta, arteries, and systemic capillaries. After this, from all parts except the spleen and alimentary canal, the blood returns to systemic veins, superior or inferior cavae, right auricle, and right ventricle. The blood from the stomach spleen, and intestines however, passes via {through} the portal vein to the liver capillaries and then through the hepatic vein to inferior cava, and so on. Material leaves the blood to be excreted in lungs, kidneys, by the skin (as perspiration), and elsewhere. New material enters most conspicuously;
(a) by the portal veins portal veins and
(b) by the thoracic duct and left superior cava.
Section 49. The following table summarises what we have learnt up to the present of the physiology of the Rabbit, considered as a mechanism using up food and oxygen and disengaging energy:--
-Air_ {Nitrogen... returned unchanged.}
{Oxygen... through Pulmonary Vein to--} {see 3.}
-Food_ {Carbo-Hydrates (Starch, Sugar, Cellulose.)} Sugar.
{Protein.} {Peptones.}
{Fat (little in Rabbit.)} {Glycerine, and fatty acids in soups.}
{Rejected matter got rid of in Defaecation.}
1a. {Chyle in Lacteals going via {through} Thoracic Duct and Left
Superior Cava to--} {see 2.}
1b. {Veins of Villi--}
{Portal Vein--}
{Liver--}
{Hepatic Vein and Inferior Cava to--} {see 2.}
2. {Right side of heart; then to lungs, and then to--} {see 3.}
3. {Left side of heart; whence to Systemic Arteries and Capillaries.}
4. {The tissues and -Kataboly_.}
5. {Urea (?Liver) Kidney and Sweat Glands}
{CO2} {Lungs}
{H2O} {Lungs, Kidney, Sweat Glands}
{Other Substances} {Mainly by [Kidney,] Liver and Alimentary Canal}
4. _The Amoeba. Cells, and Tissue_
Section 50. We have thus seen how the nutritive material is taken into the animal's system and distributed over its body, and incidentally, we have noted how the resultant products of the creature's activity are removed. The essence of the whole process, as we have already stated, is the decomposition and partial oxydation of certain complex chemical compounds to water, carbon dioxide, a low nitrogenous body, which finally takes the form of urea, and other substances. We may now go on to a more detailed study, the microscopic study, or histology, of the tissues in which metaboly and kataboly occur, but before we do this it will be convenient to glance for a moment at another of our animal types-- the Amoeba, the lowest as the rabbit is the highest, in our series.
Section 51. This is shown in Figure III., [Sheet 3], as it would appear under the low power of the microscope. We have a mass of a clear, transparent, greyish substance called protoplasm, granular in places, and with a clearer border; within this is a denser portion called the nucleus, or endoplast (n.), which, under the microscope, by transmitted light, appear brighter, and within that a still denser spot, the nucleolus (ns.) or endoplastule. The protoplasm is more or less extensively excavated by fluid spaces, vacuoles; one clearer circular space or vacuole, which is invariably present, appears at intervals, enlarges gradually, and then vanishes abruptly, to reappear after a brief interval; this is called the contractile vacuole (c.v.). The amoeba is constantly changing its shape, whence its older name of the Proteus animalcule, thrusting out masses of its substance in one direction, and withdrawing from another, and hence slowly creeping about. These thrust-out parts, in its outline, are called pseudopodia (ps.). By means of them it gradually creeps round and encloses its food. Little particles of nutritive matter are usually to be detected in the homogeneous protoplasm of its body; commonly these are surrounded by a drop of water taken in with them, and the drop of water is then called a food vacuole. The process of taking in food is called ingestion. The amoeba, in all probability, performs essentially the same chemical process as we have summarised in Sections [10], [11], [12]; it ingests food, digests it in the food vacuoles and builds it up into its body protoplasm, to undergo kataboly and furnish the force of its motion-- the contractile vacuole, is probably respiratory and perhaps excretory, accumulating and then, by its "systole" (compare [Section 44]), forcing out of its body, the water, carbon dioxide, urea, and other katastases, which are formed concomitantly with its activity. The amoeba reproduces itself in the simplest way; the nucleus occasionally divides into two portions and a widening fissure in the protoplasm of the animal's body separates one from the other. It is impossible to say that one is the parent cell, and the other the offspring; the amoeba we merely perceive, was one and is now two. It is curious to note, therefore, that the amoeba is, in a sense, immortal-- that the living nucleus of one of these minute creatures that we examine to-day under a microscope may have conceivably drawn, out an unbroken thread of life since the remotest epochs of the world's history. Although no sexual intercourse can be observed, there is reason to believe that a process of supposed "cannabalism," in which a larger amoeba may occasionally engulph a smaller one, is really a conjugative reproductive process, and followed by increased vitality and division.
Section 52. Now if the student will compare [Section 35], he will see that in the white blood corpuscles we have a very remarkable resemblance to the amoeba; the contractile vacuole is absent, but we have the protoplasmic body, the nucleus and nucleolus, and those creeping fluctuations of shape through the thrusting out and withdrawal of pseudopodia, which constitute "amoeboid" motion. They also multiply, in the same way, by division.
Section 53. It is not only in the white corpuscle of the blood that we find this resemblance; in all the firmer parts of the body we find, on microscopic examination, similar little blebs of protoplasm, and at an early stage of development the young rabbit is simply one mass of these protoplasmic bodies. Their division and multiplication is an essential condition, of growth. Through an unfortunate accident, these protoplasmic blebs, which constitute the living basis of the animal body, have come to be styled "cells," though the term "corpuscles" is far more appropriate.
Section 54. The word is "cell" suggests something enclosed by firm and definite walls, and it was first employed in vegetable histology. Unlike the typical cells of animals, the cells of most plants are not naked protoplasm, but protoplasm enclosed in a wall of substance (cell wall) called cellulose. The presence of this cellulose cell wall, and the consequent necessity of feeding entirely upon liquids and gases that soak through it instead of being able to ingest a portion of solid food is indeed, the primary distinction between the vegetable and the animal kingdoms, as ordinarily considered.
Section 55. Throughout life, millions of these cells retain their primary characters, and constitute the white corpuscles of blood, "phagocytes," and connective tissue corpuscles; others again, engage in the formation of material round themselves, and lie, in such cases, as gristle and bone, embedded in the substance they have formed; others again, undergo great changes in form and internal structure, and become permanently modified into, for instance, nerve fibres and muscle substance. The various substances arising in this way through the activity of cells are called tissues, the building materials of that complex thing, the animal body. Since such a creature as the rabbit is formed through the co-operation of a vast multitude of cells, it is called multicellular; the amoeba, on the other hand, is unicellular. The rabbit may be thus regarded as a vast community of amoeboid creatures and their products.
Section 56. Figure IV., [Sheet 3] represents, diagrammatically, embryonic tissue, of which, to begin with, the whole animal consists. The cells are all living, capable of dividing and similar, but as development proceeds, they differentiate, some take on one kind of duty (function), and some another, like boys taking to different trades on leaving school, and wide differences in structure and interdependence become apparent.
Section 57. It is convenient to divide tissues into three classes, though the divisions are by no means clearly marked, nor have they any scientific value. The first of these comprises tissues composed wholly, or with the exception of an almost imperceptible cementing substance, of cells; the second division includes the skeletal tissues, the tissue of mesentery, and the connective and basement tissue of most of the organs, tissues which, generally speaking, consist of a matrix or embedding substance, formed by the cells and outside of them, as well as the cells themselves; and, thirdly, muscular and nervous tissue. We shall study the former two in this chapter, and defer the third division until later.
Section 58. The outer layer of the skin (the epidermis), the inmost lining of the alimentary canal, the lining of the body cavity, and the inner linings of blood-vessels, glands, and various ducts constitute our first division. The general name for such tissues is epithelium. When the cells are more or less flattened, they form squamous epithelium ([Figure VI.]) such as we find lining the inside of a man's cheek (from which the cells sq.ep. were taken) or covering the mesentery of various types-- sq.end. are from the mesentery ([Section 16]) of a frog. A short cylindroidal form of cell makes up columnar epithelium, seen typically in the cells covering the villi of the duodenum ([Figure V.]). This epithelium of the villi has the outer border curiously striated, and this is usually spoken of as leading towards "ciliated" epithelium, to be described immediately. The epithelium of the epidermis is stratified-- that is to say, has many thicknesses of cells; the deeper layers are alive and dividing (stratum mucosum), while the more superficial are increasingly flattened and drier as the surface is approached (stratum corneum) and are continually being rubbed off and replaced from below.
Section 59. In the branching air-tubes of the lung, the central canal of the spinal cord, and in the ureters of the rabbit, and in most other types, in various organs, we find ciliated epithelium ([Figure VII.]). This is columnar or cubical in form, and with the free edge curiously modified and beset with a number of hair-like processes, the cilia, by which, during the life of the cell, a waving motion is sustained in one direction. This motion assists in maintaining a current in the contents of ducts which are lined with this tissue. The motion is independent of the general life of the animal, so long as the constituent cell still lives, and so it is easy for the student to witness it himself with a microscope having a 1/4-inch or 1/6-inch objective. Very fine cilia may be seen by gently scraping the roof of a frog's mouth (the cells figured are from this source), or the gill of a recently killed mussel, and mounting at once in water, or, better, in a very weak solution of common salt.
Section 60. The lining of glands is secretory epithelium; the cells are usually cubical or polygonal (8, g.ep.), and they display in the most characteristic form what is called metabolism. Anaboly (see [Section 14]) we have defined, as a chemical change in an upward direction-- less stable and more complex compounds are built up in the processes of vegetable and animal activity towards protoplasm; kataboly is a chemical running down; metaboly is a more general term, covering all vital chemical changes. The products of the action of a glandular epithelium are metabolic products, material derived from the blood is worked, up within the cell, not necessarily with conspicuous gain or loss of energy, and discharged into the gland space. The most striking case of this action is in the "goblet cells" that are found among the villi; these are simply glands of one cell, unicellular glands, and in [Figure V.] we see three stages in their action: at g.c.1 material (secretion) is seen forming in the cell, at g.c.2 it approaches the outer border, and at g.c.3 it has been discharged, leaving a hollowed cell. Usually however, the escape of secreted matter is not so conspicuous, and the gland-cells are collected as the lining of pits, simple, as in the gastric, pyloric, and Lieberkuhnian glands (Figure VIII., Sections [23], [29]), or branching like a tree or a bunch of grapes (Figure r.g.), as in Brunner's glands ([Section 29]) the pancreas, and the salivary glands. The salivary glands, we may mention, are a pair internal to the posterior ventral angle of the jaw, the sub-maxillary; a pair anterior to these, the sub-lingual; a pair posterior to the jaw beneath the ear, the parotid, and a pair beneath the eye, the infra orbital.
Section 61. The liver is the most complicated gland in the body ([Figure X.]). The bile duct (b.d.) branches again and again, and ends at last in the final pits, the lobuli (lb.), which are lined with secretory epithelium, and tightly packed, and squeeze each other into polygonal forms. The blood supply from which the bile would appear to be mainly extracted, is brought by the portal vein, but this blood is altogether unfit for the nutrition of the liver tissue; for this latter purpose a branch of the coeliac artery, the hepatic serves. Hence in the tissue of the liver we have, branching and interweaving among the lobuli, the small branches of the bile duct (b.d.), which carries away the bile formed, the portal vein (p.v.), the hepatic artery (h.a.), and the hepatic vein (h.v.). (Compare [Section 45].) Figure X.b shows a lobule; the portal vein and the artery ramify round the lobules-- are inter-lobular, that is (inter, between); the hepatic vein begins in the middle of the lobules (intra-lobular), and receives their blood. (Compare X.a.) Besides its function in the manufacture of the excretory, digestive, and auxiliary bile, the liver performs other duties. It appears to act as an inspector of the assimilation material brought in by the portal vein. The villi, for instance, will absorb arsenic, but this is arrested and thrown down in the liver. A third function is the formation of what would seem to be a store of carbo-hydrate, glycogen, mainly it would appear, from the sugar in the portal vein, though also, very probably, from nitrogenous material, though this may occur only under exceptional conditions. Finally, the nitrogenous katastases, formed in the working of muscle and nerve, and returned by them to the blood for excretion, are not at that stage in the form of urea. Whatever form they assume, they undergo a further metabolism into urea before leaving the body, and the presence of considerable quantities of this latter substance in the liver suggests this as a fourth function of this organ-- the elaboration of urea.
Section 62. Similar from a physiological point of view, to the secretory glands which form the digestive fluids are those which furnish lubricating fluids, the lachrymal gland, and Harderian glands in the orbit internally to the eye, and posterior and anterior to it respectively, the sebaceous glands (oil glands) connected with the hair, and the anal and perineal glands. The secretions of excretory glands are removed from the body; chief among them are the sweat glands and kidneys. The sweat glands are microscopic tubular glands, terminating internally in a small coil ([Figure VIII.] s.g.) and are scattered thickly over the body, the water of their secretion being constantly removed by evaporation, and the small percentage of salt and urea remaining to accumulate as dirt, and the chief reasonable excuse for washing. The kidney structure is shown diagrammatically in Figure 5, [Sheet 7]. A great number of branching and straight looped, tubuli (little tubes) converge on an open space, the pelvis. Towards the outer layers (cortex) of the kidney, these tubuli terminate in little dilatations into which tangled knots of blood-vessels project: the dilatations are called Bowman's capsules (B.c.), and each coil of bloodvessel a glomerulus (gl.). In the capsules, water is drained from the blood; in the tubuli, urea and other salts in the urine are secreted from a branching network of vessels.
Section 63. In all the epithelial tissues that we have considered we have one feature in common: they are cells, each equivalent to the amoeba, that have taken on special duties-- in a word, they are specialists. The amoeba is Jack of all trades and a free lance; the protective epidermal cell, the current-making ciliated cell, the bile or urea-making secretory cell, is master of one trade, and a soldier in a vast and wonderfully organized host. We will now consider our second kind of cell in this organization, the cell of which the especial aim is the building round it of a tissue.
Section 64. The simplest variety in this group is hyaline (i.e. glassy) cartilage (gristle). In this the formative cells (the cartilage corpuscles) are enjellied in a clear structureless matrix ([Figure XII.]), consisting entirely of organic compounds accumulated by their activity. Immediately round the cell lies a capsule of newer material. Some of the cells have recently divided (1); others have done so less recently, and there has been time for the interpolation of matrix, as at 2. In this way the tissue grows and is repaired. A thin layer of connective tissue (see below), the perichondrium, clothes the cartilaginous structure.
Section 65. Connective tissue ([Figure XIII]) is a general name for a group of tissues of very variable character. It is usually described as consisting typically in the mammals of three chief elements felted together; of comparatively unmodified corpuscles (c.c.), more or less amoeboid, and of fibres which are elongated, altered, and distorted cells. The fibres are of two kinds: yellow, branching, and highly elastic (y.e.f.), in consequence of which they fall into sinuous lines in a preparation, and white and inelastic ones (w.i.f.), lying in parallel bundles. Where the latter element is entirely dominant, the connective tissue is tendon, found especially at the point of attachment of muscles to the parts they work. Some elastic ligaments are almost purely yellow fibrous tissue. A loose interweaving of the three elements is areolar tissue, the chief fabric of mesentery, membrane, and the dermis (beneath the epidermis). With muscle it is the material of the walls of the alimentary canal and bloodvessels, and generally it enters into, binds together, and holds in place other tissue. The connective tissue of fishes displays the differentiation of fibres in a far less distinct manner.
Section 66. Through connective tissues wander the phagocytes, cells that are difficult to distinguish, if really distinct, from the white blood corpuscles. These cells possess a remarkable freedom; they show an initiative of their own, and seem endowed with a subordinate individuality. They occur in great numbers in a tissue called, botryoidal tissue ([Figure XIV.]), which occurs especially in masses and patches along the course of the alimentary canal, in its walls. The tonsils, swellings on either side of the throat, are such masses, and aggregates occur as visible patches, the Peyer's patches, on the ileum. It also constitutes the mass of the vermiform appendix and the wall of the sacculus rotundus; and in the young animal the "thymus gland," ventral to the heart, and less entirely, the "thyroid gland," ventral to the larynx, are similar structures, which are reduced or disappear as development proceeds. It is evident that in these two latter cases the term "gland" is somewhat of a misnomer. The matrix of botryoidal tissue is a network of stretched and hollowed connective tissue cells-- it is not a secretion, as cartilage matrix appears to be. During digestion, the phagocytes prowl into the intestine, and ingest and devour bacteria, that might otherwise give rise to disease. In inflammation, we may note here, they converge from all directions upon the point wounded or irritated. They appear to be the active agents in all processes of absorption (see osteoclasts under bone), and for instance, migrate into and devour the tissue of the tadpole's tail, during its metamorphosis to the adult frog.
Section 67. Within the connective tissue cells fat drops may be formed, as in [Figure XV.] Adipose tissue is simply connective tissue loaded with fat-distended cells. The tissue is, of course, a store form of hydro-carbon ([Section 17]) provided against the possible misadventure of starvation. With the exception of some hybernating animals, such store forms would seem to be of accidental importance only among animals, whereas among plants they are of invariable and necessary occurrence.
Section 68. We now come to Bone, a tissue confined to the vertebrata, and typically shown only in the higher types. As we descend in the scale from birds and mammals to lizards, amphibia (frogs and toads) and fish, we find cartilage continually more important, and the bony constituent of the skeleton correspondingly less so. In such a type as the dog-fish, the skeleton is entirely cartilaginous, bone only occurs in connection with the animal's scales; it must have been in connection with scales that bone first appeared in the vertebrate sub-kingdom. In the frog we have a cartilaginous skeleton overlaid by numerous bony scutes (shield-like plates) which, when the student comes to study that type, he will perceive are equivalent to the bony parts of such scales as occur in the dog-fish, sunk inward, and plating over the cartilage; and in the frog the cartilage also is itself, in a few places, replaced by bony tissue. In the adult rabbit these two kinds of bone, the bone overlying what was originally cartilage (membrane bone), and the bone replacing the cartilage (cartilage bone) have, between them, practically superseded the cartilage altogether. The structure of the most characteristic kind of bone will be understood by reference to [Figure XVI]. It is a simplified diagram of the transverse section of such a bone as the thigh bone. M.C. is the central marrow cavity, H.v., H.v. are cross sections of small bloodvessels, the Haversian vessels running more or less longitudinally through, the bone in canals, the Haversian canals. Arranged round these vessels are circles of the formative elements, the bone corpuscles or osteoblasts (b.c.) each embedded in bony matrix in a little bed, the lacuna, and communicating one with another by fine processes through canaliculi in the matrix, which processes are only to be seen clearly in decalcified bone (See [Section 70]). The osteoblasts are arranged in concentric series, and the matrix is therefore in concentric layers, or lamellae (c.l.). Without and within the zone of Haversian systems are (o.l. and i.l.), the outer and inner lamellae. The bone is surrounded by connective tissue, the periosteum. In addition to this compact bone, there is a lighter and looser variety in which spicules and bars of bony tissue are loosely interwoven. Many flat bones, the bones of the skull, for instance, consist of this spongy bone, plated (as an electro spoon is plated) with compact bone.
Section 69. Among the bony bars and spicules of spongy bone occurs the red marrow-- which must not be confused with the yellow marrow, the fatty substance in the central cavity of long bones. In this red marrow are numerous large colourless cells, which appear to form within their substance and then liberate red blood corpuscles. This occurs especially in the spongy bone within the ribs.
Section 70. The matrix of bone differs from that of cartilage or of most other tissues in consisting chiefly of inorganic salts. The chief of these is calcium phosphate, with which much smaller quantities of calcium carbonate, and magnesium phosphate and carbonate occur. These inorganic salts can be removed by immersion of the bone in weak hydrochloric acid, and a flexible network of connecting tissue, Haversian vessels, bone corpuscles, and their processes remains. This is decalcified bone alluded to above.
Section 71. In the very young rabbit, the limb bones, vertebral column, and many of the skull bones are simply plates and bars of cartilage; the future membrane bones, however are planned out in connective tissue. The development of the latter is simple, the connective tissue corpuscles functioning by a simple change of product as osteoblast. The development of the cartilage bones, however, is more complicated. [Figure XVII.], represents, in a diagrammatic way, the stages in the conversion of a cartilaginous bar to bone. To begin with, the previously sporadically-arranged (scattered anyhow) corpuscles (u.c.c.) are gathered into groups in single file, or in other words, into "columnar" groups (as at c.c.). The matrix becomes clouded with inorganic salts of lime, and it is then said to be calcified. This calcified cartilage then undergoes absorption-- it must not be imagined for a moment that bone is calcified cartilage. Simultaneous with the formation of the cavities (s.) due to this absorption, connective tissue (p.c.i.) from the surrounding perichondrium (p.c.) grows into the ossifying* bar. It is from this connective tissue that the osteoblasts (o.b.) arise, and bone is built up. Throughout life a bone is continually being absorbed and reformed by the activity of the osteoblasts. An osteoblast engaged in the absorption instead of the formation of bone is called an osteoclast.
* The formation of bone is called ossification. To ossify is to become bony.
Section 72. The great thing to notice about this is that cartilage does not become bone, but is eaten into and ousted by it; the osteoblasts and osteoclasts replace entirely the cartilage corpuscles, and are not derived from them.
Section 73. We may mention here the structure of the spleen (Figure 1, [Sheet 1]). It consists of a connective tissue and muscular coating, with an internal soft matrix much resembling botryoidal tissue, traversed by fibrous trabeculae (= beams, planks) containing blood-vessels, and the whole organ is gorged with blood, particularly after meals. The consideration of its function the student may conveniently defer for the present.
Section 74. Here also, we may notice the lymphatics, a series of small vessels which return the overflow of the blood serum from the capillaries, in the nutrition of the tissues in all parts of the body, to the thoracic duct (see [Section 36]), and the general circulation. At intervals their course is interrupted by gland-like dilatations, the lymphatic glands, in which masses of rapidly dividing and growing (proliferating) cells occur, of which, doubtless, many are detached and become, first "lymph corpuscles," and, when they reach the veins, white blood corpuscles.
5. _The Skeleton_
Section 75. We are now in a position to study the rabbit's skeleton. We strongly recommend the student to do this with the actual bones at hand-- they may be cleared very easily in a well-boiled rabbit. This recommendation may appear superfluous to some readers, but, as a matter of fact, the marked proclivity of the average schoolmaster for mere book-work has put such a stamp on study, that, in nine cases out of ten, a student, unless he is expressly instructed to the contrary, will go to the tortuous, and possibly inexact, descriptions of a book for a knowledge of things that lie at his very finger-tips. We have not written, this chapter to give a complete knowledge of the skeleton, but simply as an aid in the actual examination of the bones.
Section 76. We may take the skeleton under five headings. There is the central axis, the chain of little bones, the vertebrae, threaded on the spinal cord (see Figure 1 and [Section 1]); the thorax, the box enclosed by ribs and sternum; the fore-limb and bones connected with it (pectoral girdle and limb), and the hind-limb and bones connected with it (pelvic girdle). Finally there is the skull, but following the London University syllabus, we shall substitute the skull of the dog for of that of the rabbit, as more typically mammalian ([Section 4]).
Section 77. In [Section 3] (which the student should refer to) we have a division of the vertebrae into four varieties. Of these most representative is the thoracic. A thoracic vertebra (Figure 4, [Sheet 5], T.V.) consists of a central bony mass, the body or centrum (b.), from which there arises dorsally an arch, the neural arch (n.a.), completed by a keystone, the neural spine (n.s.); and coming off laterally from the arch is the transverse process (tr.p.). Looking at the vertebra sideways, we see that the arch is notched, for the exit of nerves. Jointed to the thoracic vertebrae on either side are the ribs (r.). Each rib has a process, the tuberculum, going up to articulate with the transverse process, and one, the capitulum articulating between the bodies of two contiguous vertebrae. The facets for the articulation of the capitulum are indicated in the side view by shading. At either end of the body of a vertebra of a young rabbit are bony caps, the epiphyses (ep.), separated from the body by a plane of unossified cartilage (indicated, by the dots). These epiphyses to the vertebral bodies occur only among mammals, and are even absent in some cases within the class. In the adult rabbit they have ossified continuously with the rest of the body.
Section 78. A cervical vertebra (C.V.) seems, upon cursory inspection, to have no rib. The transverse processes differ from those of thoracic series in having a perforation, the vertebrarterial canal, through which the vertebral artery runs up the neck. A study of the development of these bones shows that the part marked f.r. ossifies separately from the rest of the transverse process; and the form of the equivalent structures in certain peculiar lower mammals and in reptiles leaves no doubt that f.r. is really an abbreviated rib; fused up with the transverse process and body. The two anterior cervical vertebrae are peculiar. The first (at.) is called the Atlas-- the figure shows the anterior view-- and has great articular faces for the condyles ([Section 86]) of the skull, and a deficient centrum. The next is the axis, and it is distinguished by an odontoid peg (od.p.), which fits into the space where the body of the atlas is deficient. In development the centrum of the axis ossifies from one centre, and the odontoid, peg from another, which at that time occupies the position of centrum of the atlas. So that it would seem that the atlas is a vertebra minus a centrum, and the axis is a vertebra plus a centrum, added at the expense of the atlas.
Section 79. The lumbar vertebrae (l.v.) are larger, and have cleft transverse processes, each giving rise to an ascending limb, the metapophyses, and a descending one. The latter (generally spoken of as the transverse processes) point steeply downward, and are considerably longer than those of thoracic series. The sacral vertebrae (s.v.) have great flattened transverse processes for articulation with the ilia. The caudal vertebrae (c.v.) are gradually reduced to the mere elongated centra, as we proceed, towards the tip of the tail.
Section 80. All the vertebrae join with their adjacent fellows through the intermediation of certain intervertebral pads, and also articulate by small processes at either end at the upper side of the arch, the zygapophyses. The normals to the polished facets of these point, in the case of the anterior zygapophyses, up and in (mnemonic: ant-up-in), and in the case of the posterior, down and out. The student should make this, and the other features of vertebrae, out upon actual specimens.
Section 81. The thorax is bounded dorsally by the vertebral column, and ventrally by the sternum. The sternum consists of segments, the sternebrae (st.); anteriorly there is a bony manubrium (mb.), posteriorly a thin cartilaginous plate, the xiphisternum (xi.). Seven pairs of ribs articulate by cartilaginous ends (sternal ribs) with the sternum directly, as indicated in the figure; five (false) ribs are joined, to each other and to the seventh, and not to the sternum directly. The last four ribs have no tuberculum ([Section 77]).
Section 82. The fore-limb (pectoral limb) consists of an upper arm bone, the humerus (hum.) the distal end of which is deeply excavated by the olecranon fossa (o.f.) as indicated by the dotted lines; of two bones, the ulna (u.) and radius (r.) which are firmly bound by ligament in the position of the figure (i.e., with the palm of the hand downward, "prone"); of a number of small bones (carpalia), the carpus (c.); of a series of metacarpals (mc.); and of three digits (= fingers) each, except the first, or pollex, of three small bones-- the phalanges, only the proximal of which appear in the figure. The ulna has a hook-like head, the olecranon (o.) which distinguishes it easily from the distally thickened radius. The limb is attached to the body through the intermediation of the shoulder-blade (scapula, sc.) a flattened bone with a median external ridge with a hook-like termination, the acromion (acr.). There is also a process overhanging the glenoid cavity (g.) wherein the humerus articulates, which process is called coracoid (co.); it is ossified from two separate centres, and represents a very considerable bone in the bird, reptile, and frog. Along the dorsal edge of the scapula of the rabbit is unossified cartilage, which is called the supra-scapula (s.sc.). In man there runs from the acromion to the manubrium of the sternum a bone, the collar-bone or clavicle. This is represented by a very imperfectly ossified rudiment in the rabbit. The scapula and clavicle, the bones of the body connected with the fore-limb, are frequently styled the pectoral girdle, or shoulder-girdle; this name of girdle will appear less of a misnomer when lower vertebrate types are studied.
Section 83. The hind limb and its body bones-- pelvic limb and girdle-- are shown in [Figure 2.] The limb skeleton corresponds closely with that of the fore-limb. The femur (fe.) answers to the humerus, and is to be distinguished from it by the greater distinctness of its proximal head (hd.) and by the absence of an olecranon fossa from its distal end. The tibia (ti = the radius) is fused for the distal half of its length with the fibula (fb. = ulna). A tarsus (tarsalia) equals the carpus.* Two of the proximal tarsalia may be noted: one working like a pulley under the tibia, is the astragalus (as.); one forming the bony support of the heel, is the calcaneum (ca.). There is a series of metatarsals, and then come four digits of three phalanges each.
* Such a resemblance as exists between one vertebra and another in the rabbit, or between the humerus and the femur, is called serial homology; the two things correspond with each other to the extent of imperfect reduplication. "Homology" simply is commonly used to indicate the resemblance between any two structures in different animals, in origin and position as regards other parts. Thus the heart of the rabbit and of the frog are homologous structures, corresponding in position, and resembling each other much as two memory sketches of one picture might do.
Section 84. The pelvic girdle differs from the pectoral in most land vertebrata in being articulated with the vertebral column. This difference does not exist in fishes. It consist in the rabbit of four bones; the ilium (i.), the ischium (is.), the pubis (pb.), and the small cotyloid bone-- the first two and the latter one meeting in the acetabular fossa (ac.) in which the head of the femur works. The pubes and ischia are fused along the mid-ventral line. Many morphologists regard, the ilium as equivalent to, that is, strictly corresponding in its relation, to the scapula, the pubis to the cartilaginous substratum of the clavicle, and the ischium to the coracoid.
Section 85. These bones will be studied at the greatest advantage when dissected out from a boiled rabbit. Prepared and wired skeletons, disarticulated skeletons, plates of figures, and written descriptions are in succession more tedious and less satisfactory ways to a real comprehension, of this matter. This chapter directs the student's attention to the most important points in the study of the skeleton, but it is in no way intended to mitigate the necessity of practical work. It is a guide simply.
Section 86. The mammalian skull will be better understood after the study of that of some lower vertebrate. We shall describe its main features now, but their meaning will be much clearer after the lower type is read. Our figures are of Canis. In section (Figure VI., [Sheet 6]), we perceive a brain case (cranium) opening behind by a large aperture, the foramen magnum (F.M.). In front of this is an extensive passage, the nasal passage (E.N. to P.N.) which is divided from the mouth by a bony floor, the palate, and which opens into the pharynx behind at the posterior nares (P.N.) and to the exterior by the anterior or external nares (E.N.). It is divided into right and left passages by a middle partition, the nasal septum. Outside the skull, on its wings, is a flask-like bone, the bulla tympani (b. in [Figures 2 and 3]), protecting the middle ear, and from above this there passes an arch, the cheek bone (ju. in [Figures 1, 2, and 3]), to the upper jaw, forming in front the bony lower protection of the cavity containing the eye, the orbit. The cheek arch, nasal passage, and jaws, form collectively the "facial apparatus," as distinguished from the cranium, and the whole skull is sometimes referred to as, the "cranio-facial apparatus." Two eminences for articulation with the atlas vertebra, the condyles (c.), lie one on each side of the lower boundary of the foramen magnum.
Section 87. The floor of the cranium consists of a series of cartilage bones, the basi-occipital (b.o.), basi-sphenoid (b.sp.), pre-sphenoid (p.sp.), and in front, the ethmoid (eth.), which sends down a median plate, not shown, in the figure, to form the nasal septum between right and left nasal passages. Like extended wings on either side of the basi-occipital are the ex-occipital (e.o.) (the bone is marked in [Figure 4], but the letters are a little obscured by shading). Similarly the ali-sphenoids (a.s.), are wings to the basi-, and the orbito-sphenoids (o.s.), to the pre-sphenoid bone (p.sp.). Between the ex-occipital and ali-sphenoid there is wedged in a bone, the periotic (p.o.) containing the internal ear ([Section 115]). Above the foramen magnum the median supra-occipital bone completes what is called the occipital arch. A pair of parietals (pa.) come above the ali-sphenoids, and a pair of frontals (f.) above the orbito-sphenoids. At the side the brain case is still incomplete, and here the squamosal (sq.) enters into its wall. In the external view ([Figure 3]) the bulla hides the periotic bone from without. The student should examine all four figures for these bones before proceeding.
Section 88. The outer edge of the upper jaw and the cheek arch are made up of three paired bones. First comes the premaxilla (p.m.) (not p.m.1 or p.m.4), containing in the dog, the three incisors of either side. Then comes the maxilla, bearing the rest of the teeth.* The jugal or malar (ju.) reaches over from the maxilla to meet a zygomatic process (= connecting outgrowth) (z.p.) of the squamosal bone.
* In the dog a sabre-like canine (c.), four premolars (p.m.1 and p.m.4) and two molars (m.1 and m.2).
Section 89. In the under view of the skull ([Figure 2]) it will be seen that the maxilla sends in a plate to form the front part of the hard palate. Behind, the hard palate is completed by the pair of palatine bones (pal.), which conceal much of the pre- and orbito-sphenoid in the ventral view, and which run back as ridges to terminate in two small angular bones, the pterygoids (pt.) which we shall find represent much more important structures in the lower vertebrata.
Section 90. The pre-maxillae and maxillae bound the sides of the nasal passage, and it is completed above by a pair of splints, the nasals. Along the floor of the nasal passage, on the middle line, lies a splint of bone formed by the coalescence of two halves. It embraces in a V-like groove the mesethmoid (nasal septum) above, and lies on the palate.
{Lines from First Edition only.}
-Its position is indicated by a heavy black line in 4, and it is
called, the vomer bone (vo.).-
{Lines from Second Edition only.}
[In the frog it is represented by two laterally situated bones. This is
the vomer bone (vo.).]
The nasal passages are partially blocked by foliated bony outgrowths, from the inner aspect of their walls, which in life are covered with mucous membrane, and increase the surface sensitive to smell. The ethmoid ends in the ethmo-turbinal (e.t.); the nasal, the naso-turbinal (n.t.); and the maxilla, the maxillo-turbinal (m.t.). In the anterior corner of the orbit there is a bone, the lachrymal (lc. [Figure 1]), which is hidden by the maxilla in the side view of the skull.
Section 91. The lower jaw (mandible) is one continuous bone in the mammal. Three incisors bite against the three of the upper jaw. Then comes a canine, four premolars, and three molars, the first of which is blade-like (sectorial tooth), and bites against the similar sectorial tooth (last premolar) of the upper jaw. The third molar is small. The arrangement of tooth is indicated in the following dental formula:-- I. 3.3/3.3, C. 1.1/1.1, P.M. 4.4/4.4, M. 2.2/3.3
Section 92. Attached just behind the bulla above, and passing round on either side of the throat to meet at the base of the tongue, is the hyoid apparatus ([Figure 6]). The stylohyal (s.h.), epihyal (e.h.), and ceratohyal (c.h.) form the anterior cornu of the hyoid. The body of the hyoid (b.h.) forms a basis for the tongue. The posterior coruna (t.h.) of the hyoid are also called the thyrohyals.
Section 93. The following table presents these bones in something like their relative positions. A closer approximation to the state of the case will be reached if the student will imagine the maxilla raised up so as to overlie and hide the palatine and presphenoid, the squamosal similarly overlying the periotic bone, and the jugal reaching between them. Membrane bones are distinguished by capital letters.
-Cranium_
-Nasal_ (paired), Ethmoid Bone (median), -Vomer_
-Upper Jaw_
-Frontal_ (paired), -Lachrymal_ (paired), Orbito-sphenoid (paired),
Pre-sphenoid (median), Eye
-Parietal_ (Paired), Ali-sphenoid (paired), Basi-sphenoid (median)*,
Periotic Bone (paired)
-Bulla_ (paired)
Supra-occipital (median), Ex-occipital (paired), Basi-occipital (median)-Pre-Maxilla_ (paired)
Palatine (paired)
Pterygoid (paired)
-Lower Jaw_-Maxilla_ (paired)
-Jugal_ (paired)
-Squamosal_ (paired)*In this table the small bones of the ear are simply indicated by an asterisk.
Section 94. Hidden by the bulla, and just external to the periotic bone, are the auditory ossicles, the incus, malleus, os orbiculare, and stapes. These will be more explicitly treated when we discuss the ear.
Section 95. When we come to the study of the nerves, we shall revert to the skull, and treat of its perforations. The student should not fail, before proceeding, to copy and recopy our figures, and to make himself quite familiar with them, and he should also obtain and handle an actual skull. For all practical purposes the skull of a sheep or cat will be almost as useful as that of the dog.
6. _Muscle and Nerve_
Section 96. We have, in the skeleton, a complicated apparatus of parts hinged and movable upon one another; the agent moving these parts is the same agent that we find in the heart walls propelling the blood through the circulation, in the alimentary canal squeezing the food along its course, and universally in the body where motion occurs, except in the case of the creeping phagocytes, and the ciliary waving of ciliated epithelium. This agent is muscle. We have, in muscular tissue, a very wide departure from the structure of the primordial cell; to use a common biological expression, a very great amount of modification (= differentiation). [Sheet 7] represents the simpler kind of muscular tissue, unstriated muscle, in which the cell character is still fairly obvious. The cells are fusiform (spindle-shaped), have a distinct nucleus and faint longitudinal striations (striations along their length), but no transverse striations.
Section 97. In striated muscle extensive modifications mask the cell character. Under a 1/4 inch objective, transverse striations of the fibres are also distinctly visible, and under a much higher power we discern in a fibre ([Sheet 7]) transverse columns of rod-like sarcous elements (s.e.), the columns separated by lines of dots, the membranes of Krause (k.m.), and nuclei (n.), flattened and separated into portions, and lying, in some cases, close to the sarcolemma (sc.) the connective tissue enclosing the fibre, in others scattered throughout the substance of the fibre. The figure shows the fibre ruptured, in order to display the sarcolemma; e.p. is the end plate of a nerve (n.v.), and fb. are the fibrillae into which a fibre may be teased.
Section 98. In the heart we have an intermediate kind of muscle cardiac muscle ([Figure 2]), in which the muscle fibres branch; there is apparently no sarcolemma, and the undivided nuclei lie in the centre of the cell.
Section 99. Unstriated muscle is sometimes called involuntary, and striated, voluntary muscle; but there is really not the connexion with the will that these terms suggest. We have just mentioned that the heart-muscle is striated, but who can alter the beating of the heart by force of will? And the striated muscles of the limbs perform, endless involuntary acts. It would seem that unstriated muscle contracts slowly, and we find it especially among the viscera; in the intestine for instance, where it controls that "peristaltic" movement which pushes the food forward. Voluntary muscle, on the other hand, has a sharp contraction. The muscle of the slow-moving snails, slugs, and mussels is unstriated; all the muscle of the active insects and crustacea (crabs, lobsters, and crayfish) is striated. Still if the student bears the exception of the heart in mind, and considers muscles as "voluntary" that his will can reach, the terms voluntary and involuntary will serve to give him an idea of the distribution of these two types of muscle in his own body, and in that of the rabbit.
Section 100. Muscular contraction, and generally all activity in the body is accompanied by kataboly. The medium by which these katabolic changes are set going and controlled is the nervous system. The nervous system holds the whole body together in one harmonious whole; it is the governing organization of the multicellular community ([Section 55]), and the supreme head of the government resides in the brain, and is called the mind. But just as in a political state only the most important and most exceptional duties are performed by the imperial body, and minor matters and questions of routine are referred to boards and local authorities, so the mind takes cognisance only of a few of the higher concerns of the animal, and a large amount of the work of the nervous system goes on insensibly, in a perfectly automatic way-- even much that occurs in the brain.
Section 101. The primary elements in the tissue of the nervous system are three; nerve fibres, which are simply conducting threads, telegraph wires; ganglion cells, which are the officials of the system; and neuroglia, a fine variety of connective tissue which holds these other elements together, and may also possibly exercise a function in affecting impressions. A message along a nerve to a ganglion cell is an afferent impression, from a cell to a muscle or other external end is an efferent impression. The passage of an impression may be defined as a flash of kataboly along the nerve, and so every feeling, thought, and determination involves the formation of a certain quantity of katastases, and the necessity for air and nutrition.
Section 102. Unlike telegraph wires, to which they are often compared, nervous fibres usually convey impressions only in one direction, either centrally (afferent or sensory nerve fibres), or outwardly (efferent or motor nerve fibres). But the so-called motor nerve fibres include not only those that set muscles in motion, but those that excite secretion, check impulsive movements, and govern nutrition.
Section 103. Figure 7, [Sheet 8], shows the typical structure of nervous tissues. The nerve fibres there figured are bound together by endoneurium into small ropes, the nerves, encased in perineurium. There is always a grey axis cylinder (a.c.), which may (in medullated nerves), or may not (in non-medullated or grey nerves) have a medullary sheath (s.S.) interrupted at intervals by the nodes of Ranvier (n.R.). Nuclei (n.) at intervals under the sheath indicate the cells from which nerve fibres are derived by a process of elongation. The nerves of invertebrata, where they possess nerves, are mostly grey, and so are those of the sympathetic system of vertebrata, to be presently described, g.c., g.c. are ganglion cells; they may have many hair-like processes, usually running into continuity with the axis cylinders of nerve fibres, in which case they are called multi-polar cells, or they may be uni- or bi-polar.
Section 104. The simplest example of the action of the nervous system is reflex action. For instance, when the foot of a frog, or the hand of a soundly sleeping person, is tickled very gently, the limb is moved away from the irritation, without any mental action, and entirely without will being exercised. And when we go from light into darkness, the pupil of the eye enlarges, without any direct consciousness of the change of its shape on our part. Similarly, the presence or food in the pharynx initiates a series of movements-- swallowing, the digestive movements, and so on-- which in health are entirely beyond our mental scope.
Section 105. A vast amount of our activities are reflex, and in such action an efferent stimulus follows an afferent promptly and quite mechanically. It is only where efferent stimuli do not immediately become entirely transmuted into outwardly moving impulses that mental action comes in and an animal feels. There appears to be a direct relation between sensation and motion. For instance, the shrieks and other instinctive violent motions produced by pain, "shunt off" a certain amount of nervous impression that would otherwise register itself as additional painful sensation. Similarly most women and children understand the comfort of a "good cry," and its benefit in shifting off a disagreeable mental state.
Section 106. The mind receives and stores impressions, and these accumulated experiences are the basis of memory, comparison, imagination, thought, and apparently spontaneous will. Voluntary actions differ from reflex by the interposition of this previously stored factor. For instance, when a frog sees a small object in front of him, that may or may not be an edible insect, the direct visual impression does not directly determine his subsequent action. It revives a number of previous experiences, an image already stored of similar insects and associated with painful or pleasurable gustatory experiences. With these arise an emotional effect of desire or repulsion which, passes into action of capture or the reverse.
Section 107. Voluntary actions may, by constant repetition, become quasi-reflex in character. The intellectual phase is abbreviated away. Habits are once voluntary and deliberated actions becoming mechanical in this way, and slipping out of the sphere of mind. For instance, many of the detailed movements of writing and walking are performed without any attention to the details. An excessive concentration of the attention upon one thing leads to absent-mindedness, and to its consequent absurdities of inappropriate, because imperfectly acquired, reflexes.
Section 108. This fluctuating scope of mind should be remembered, more especially when we are considering the probable mental states of the lower animals. An habitual or reflex action may have all the outward appearance of deliberate adjustment. We cannot tell in any particular case how far the mental comes in, or whether it comes in at all. Seeing that in our own case consciousness does not enter into our commonest and most necessary actions, into breathing and digestion, for instance, and scarcely at all in the details of such acts as walking and talking we might infer that nature was economical in its use, and that in the case of such an animal as the Rabbit, which follows a very limited routine, and in which scarcely any versatility in emergencies is evident, it must be relatively inconsiderable. Perhaps after all, pain is not scattered so needlessly and lavishly throughout the world as the enemies of the vivisectionist would have us believe.
7. _The Nervous System_
Section 109. A little more attention must now be given to the detailed anatomy of the peripheral and central nerve ends. A nerve, as we have pointed out, terminates centrally in some ganglion cell, either in a ganglion or in the spinal cord or brain; peripherally there is a much greater variety of ending. We may have tactile (touch) ends of various kinds, and the similar olfactory and gustatory end organs; or the nerve may conduct efferent impressions, and terminate in a gland which it excites to secretion, in a muscle end-plate, or in fact, anywhere, where kataboly can be set going and energy disengaged. We may now briefly advert to the receptive nerve ends.
Section 110. Many sensory nerves, doubtless, terminate in fine ends among the tissues. There are also special touch corpuscles, ovoid bodies, around which a nerve twines, or within which it terminates.
Section 111. The eye ([Figure 8]) has a tough, dense, outer coat, the sclerotic (sc.), within which is a highly vascular and internally pigmented layer, the choroid, upon which the percipient nervous layer, the retina (r.) rests. The chief chamber of the eye is filled with a transparent jelly, the vitreous humour (v.h.). In front of the eye, the white sclerotic passes into the transparent cornea (c.). The epidermis is continued over the outer face of this as a thin, transparent epithelium. The choroid coat is continued in front by a ring-shaped muscle, the iris (ir.) the coloured portion of the eyes. This iris enlarges or contracts its central aperture (the black pupil) by reflex action, as the amount of light diminishes or increases. Immediately behind this curtain is the crystalline lens (l.), the curvature of the anterior face or which is controlled by the ciliary muscle (c.m.). In front of the lens is the aqueous humour (a.h.). The description of the action of this apparatus involves the explanation of several of the elementary principles of optics, and will be found by the student in any text-book of that subject. Here it would have no very instructive bearing, either on general physiological considerations or upon anatomical fact.
Section 112. The structure of the retina demands fuller notice. [Figure 9] shows an enlarged, diagram of a small portion of this, the percipient part of the eye. The optic nerve (o.n. in [Figure 8]) enters the eye at a spot called the blind spot (B.S.), and the nerve fibres spread thence over the inner retinal surface. From this layer of nerve fibres (o.n. in [Figure 9]) threads run outward, through certain clear and granular layers, to an outermost stratum of little rods (r.) and fusiform bodies called cones (c.), lying side by side. The whole of the retina consists of quite transparent matter, and it is this outermost layer of rods and cones (r. and c.) that receives and records the visual impression. This turning of the recipient ends away from the light is hardly what one would at first expect-- it seems such a roundabout arrangement-- but it obtains in all vertebrata, and it is a striking point of comparison with the ordinary invertebrate eye.
Section 113. We may pause to call the student's attention to a little point in the physiology of nerves, very happily illustrated here. The function of a nerve fibre is the conduction of impressions pure and simple; the light radiates through the fibrous layer of the retina without producing the slightest impression, and at the blind spot, where the rods and cones are absent, and the nerve fibres are gathered together, no visual impressions are recorded. If there is any doubt as to the existence of a blind spot in the retinal picture, the proof is easy. Let the reader shut his left eye, and regard these two asterisks, fixing his gaze intently upon the left-hand one of them.
* *
At a distance of three or four inches from the paper, both spots will be focussed on his retina, the left one in the centre of vision, and the right one at some spot internal to this, and he will see them both distinctly. Now, if he withdraws his head slowly, the right spot will of course appear to approach the left, and at a distance of ten or twelve inches it will, in its approach, pass over the blind spot and vanish, to reappear as he continues to move his head away from the paper. The function of nerve fibres is simply conduction, and the nature of the impressions they convey is entirely determined by the nature of their distal and proximal terminations.
Section 114. Certain small muscles in the orbit (eye-socket) move the eye, and by their action contribute to our perception of the relative position of objects. There is a leash of four muscles rising from a spot behind the exit of the optic nerve from the cranium to the upper, under, anterior, and posterior sides of the eyeball. These are the superior, inferior, anterior, and posterior recti. Running from the front of the orbit obliquely to the underside of the eyeball is the inferior oblique muscle. Corresponding to it above is a superior oblique. A lachrymal gland lies in the postero-inferior angle of the orbit, and a Handerian gland in the corresponding position in front. In addition to the upper and lower eyelids of the human subject, the rabbit has a third, the nictitating lid, in the anterior corner of the eye.
Section 115. The ear ([Sheet VII.]) consists of an essential organ of hearing, and of certain superadded parts. The essential part is called the internal ear, and is represented in all the true vertebrata (i.e., excluding the lancelet and its allies). In the lower forms it is a hollow membranous structure, embedded in a mass of cartilage, the otic capsule; in the mammal the latter is entirely ossified, to form the periotic bone. The internal ear consists of a central sac, from which three semicircular canals spring. The planes of the three canals are mutually at right angles; two are vertical, the anterior and posterior (p.v.c.) vertical canals, and one is horizontal, the horizontal canal (h.c.). There are dilatations, called ampullae, at the anterior base of the anterior, and at the posterior base of the posterior and horizontal canals. Indirectly connected with the main sac is a spirally-twisted portion, resembling a snail shell in form, the cochlea. This last part is distinctive of the mammalia, but the rest of the internal ear is represented in all vertebrata, with one or two exceptions. The whole of the labyrinth is membranous, and contains a fluid, the endolymph; between the membranous wall of the labyrinth and the enclosing bone is a space containing the perilymph. Strange as it may appear at first, the entire lining of the internal ear is, at an early stage, continuous with the general epidermis of the animal. It grows in just as a gland might grow in, and is finally cut off from the exterior; but a considerable relic of this former communication remains as a thin, vertical blind tube (not shown in the figure), the ductus endolymphaticus.
Section 116. The eighth nerve runs from the brain case (Cr.), into the periotic bone, and is distributed to the several portions of this labyrinth. In an ordinary fish this internal ear is the sole auditory organ we should find; the sound-waves would travel through the water to the elastic cranium and so reach and affect the nerves. But in all air-frequenting animals this original plan of an ear has to be added to, to fit it to the much fainter sound vibrations of the compressible and far less elastic air. A "receiving apparatus" is needed, and is supplied by the ear-drum, middle ear, or tympanic cavity (T.). In the mammal there is also a collecting ear trumpet (the ear commonly so-called), the external ear, and external auditory meatus (e.a.m.). A tightly stretched membrane, the tympanic membrane, separates this from the drum. A chain of small bones, the malleus (m.), the incus (i.), the os orbiculare (o.or.), a very small bone, and a stirrup-shaped stapes, swing across the tympanum, from the tympanic membrane to the internal ear. At two points the bony investment of this last is incomplete-- at the fenestra rotunda (f.r.), and at the fenestra ovalis, (f.o.), into which latter the end of the stapes fits, and so communicates the sound vibrations of the tympanic membrane to the endolymph. A passage, the Eustachian tube, communicates between the tympanic cavity and the pharynx (Ph.), and serves to equalize the pressure on either side of the drum-head. A comparative study of the ears of the vertebrata brings to light the fact that, as we descend in the animal scale, the four ear ossicles are replaced by large bones and cartilages connected with the jaw, and the drum and Eustachian tube by a gill slit. We have, in fact, in the ear, as the student will perceive in the sequel, an essentially aquatic auditory organ, added to and patched up to fit the new needs of a life out of water.
Section 117. The impressions of smell are conducted through the first nerve to the brain, and are first received by special hair-bearing cells in the olfactory mucous membrane of the upper part of the nasal passage. The sense of taste has a special nerve in the ninth, the fibres of which terminate in special cells and cell aggregates in the little papillae (velvet pile-like processes) that cover the tongue.
Section 118. At an early stage in development, the brain of a mammal consists of a linear arrangement of three hollow vesicles (Figure 5, [Sheet VIII.], 1, 2, and 3), which are the fore-, mid-, and hind-brain respectively. The cavities in these in these vesicles are continuous with a hollow running through the spinal cord. On the dorsal side of the fore-brain is a structure to be dealt with more fully later, the pineal gland (p.g.), while on its under surface is the pituitary body (pt.).
Section 119. The lower [figure] of (5) shows, in a diagrammatic manner, the derivation of the adult brain from this primitive state. From the fore-brain vesicle, a hollow outgrowth on either side gives rises to the (paired) cerebral hemisphere (c.h.), which is prolonged forward as the olfactory lobe (o.l.). From the fore-brain the retina of the eye and the optic nerve also originate as an, at first, hollow outgrowth (op.). The roof of the mid-brain is also thickened, and bulges up to form two pairs of thickenings, the corpora quadrigemina, (c.q.). The hind-brain sends up in front a median outgrowth, which develops lateral wings, the cerebellum (cbm.), behind which the remainder of the hind-brain is called the medulla oblongata, and passes without any very definite demarcation into the spinal cord.
Section 120. [Figure 1] is a corresponding figure of the actual state of affairs in the adult. The brain is seen in median vertical section. (ch.) is the right cerebral hemisphere, an inflated vesicle, which, in the mammal-- but not in our lower types-- reaches back over the rest of the fore-brain, and also over the mid-brain, and hides these and the pineal gland in the dorsal view of the brain ([Figure 2]). The hollow of the hemisphere on either side communicates with the third ventricle, the original cavity of the fore-brain (1 in [Figure 5]), by an aperture (the foramen of Monro), indicated by a black arrow (f.M.). Besides their original communication through the intermediation of the fore-brain, the hemispheres are also united above its roof by a broad bridge of fibre, the corpus callosum (c.c.), which is distinctive of the mammalian animals. The original fore-brain vesicle has its lateral walls thickened to form the optic thalami (o.th.), between which a middle commissure, (m.c.), absent in lower types, stretches like a great beam across the third ventricle. The original fore-brain is often called the thalamencephalon, the hemisphere, the prosencephalon, the olfactory lobes, the rhinencephalon.
Section 121. The parts of mid-brain (mesencephalon) will be easily recognised. Its cavity is in the adult mammal called the iter; its floor is differentiated into bundles of fibres, the crura cerebri (c.cb.), figured also in [Figure 4].
Section 122. The cerebellum (metencephalon) consists of a central mass, the vermis (v.cbm.), and it also has lateral lobes (l.l.), prolonged into flocculi (f.cbm.), which last are -em-bedded in pits, [in] the periotic bone, and on that account render the extraction of the brain from the cranium far more difficult than it would otherwise be. The roof of the hind-brain, before and behind the cerebellum, consists of extremely thin plates of nervous matter. Its floor is greatly thickened to form the mass of the medulla, and in front a great transverse track of fibres is specialized, the pons Varolii (p.V.). Its cavity is called, the fourth ventricle.
Section 123. [Figure 2] gives a dorsal view of the rabbit's brain; a horizontal slice has been taken at the level of the corpus callosum. The lateral ventricle (i.e., the hollows of the hemisphere) is not yet opened. A lower cut ([Figure 3]) exposes this (V.L.). The level of these slices is approximately indicated in [Figure 1] by the lines A and B. This latter figure will repay careful examination. The arrow, ar., plunges into the third ventricle, behind the great middle commissure (m.c.), and the barb is supposed to lie under the roof of the mid-brain, the corpora quadrigemina (c.q.). The position of ar. is also indicated in [Figure 1]. Before reading on, the beginner should stop a while here; he should carefully copy or trace our figures and, putting the book aside, name the parts, and he should then recopy, on an enlarged scale, and finally draw from memory, correct, and again draw. By doing this before the brain is dissected a considerable saving of time is possible.
Section 124. Proceeding from the brain are twelve pairs of cranial nerves. From the fore-brain spring two pairs, which differ from the rest of the cranial nerves in being, first of all, hollow outgrowths of the brain-- the others are from the beginning solid. The first nerve is the olfactory lobe, which sends numerous filaments through the ethmoid bone to the olfactory organ. The second is the optic nerve, the visual sensory nerve.
Section 125. The mid-brain gives rise to only one nerve, the third, which supplies all the small muscles of the eye (see [Section 114]), except the superior oblique and external rectus.
Section 126. The remainder of the nerves spring from the hind-brain. The fourth pair supply the superior obliques, and the sixth the external recti; so that III., IV., and VI. are alike purely motor nerves, small and distributed, to the orbit. The fifth nerve, the trigeminal, is a much larger and more important one; it is a mixed nerve, having three main branches, of which the first two are chiefly sensory, the third almost entirely motor; it lies deeply in the orbit. V1 (see [Sheet 9]) runs up over the recti behind the eyeball, it is the ophthalmic branch; V2, the maxillary branch, runs deeply under the eyeball and emerges in front of the malar, and V3, the mandibular branch, runs down on the inner side of the jaw-bone to the jaw muscles and tongue.
Section 127. If the student will now recur to the figures of the dog's skull ([Sheet 6]), he will see certain apertures indicated in the cranial wall. Of these, o.f. is the optic foramen for the exit of nerve II., perforating the orbito-sphenoid. Behind this there comes an irregular aperture, (f.l.a.), the foramen lacerum anterius, giving exit to III., IV., VI., and V1. V2 emerges from the foramen rotundum, and V3 from the foramen ovale, two apertures uniting behind a bony screen.* Just in front of the bulla is a foramen lacerum medium (f.l.M.), through which no nerve passes.
* In the rabbit's skull f.l. anterius, the foramen rotundum, and foramen ovale are not distinct, and there are two condylar foramina instead of one, through each of which, a moiety of XII. passes.
Section 128. The eighth nerve (auditory) is purely sensory, the nerve of the special sense of hearing; it runs into the periotic bone, and breaks up on the labyrinth. The seventh nerve (facial) is almost entirely motor; it passes through the periotic anterior to VIII., and emerges by the stylo-mastoid foramen (s.m.f.) behind the bulla, to run outside the great jaw muscle across the cheek immediately under the skin ([Figure 1]).
Section 129. The ninth (glossopharyngeal) nerve is chiefly sensory; it is the special nerve of taste, and is distributed to the tongue. The tenth nerve (vagus) arises by a number of roots, and passes out of the skull, together with IX and XI, by the foramen lacerum -posterium- [posterius] (f.l.p.). It is a conspicuous white nerve, and runs down the neck by the side of the common carotid artery. It sends a superior laryngeal branch (Xa) to the larynx. The left vagus passes ventral to the aortic arch, and sends a branch (l.x.b.) under this along the trachea to the larynx-- the recurrent laryngeal nerve. The corresponding nerve on the right (r.x.b.) loops under the subclavian artery. The main vagus, after this branching, passes behind the heart to the oesophagus and along it to the stomach. XI., the spinal accessory, supplies certain of the neck nerves. XII., the hypoglossal, runs out of the skull by the condylar foramen (c.f.), is motor, crosses the roots of XI., X., and IX., passes ventral to the carotid, and breaks up among the muscles of the tongue and neck.
Section 130. Of the functions of the several parts of the brain there is still very considerable doubt. With disease or willful destruction of the cerebral tissue the personal initiative is affected-- the animal becomes more distinctly a mechanism; the cerebellum is probably concerned in the coordination of muscular movements; and the medulla is a centre for the higher and more complicated respiratory reflexes, yawning, coughing, and so on. The great majority of reflex actions centre, however, in the spinal cord, and do not affect the brain.
Section 131. A cross section of the spinal cord is shown in Figure 6, [Sheet 8]. It is a cylinder, almost bisected by a dorsal (d.f.) and a ventral (v.f.) fissure. Through its centre runs a central canal (c.c.), continuous with the brain ventricles, and lined by ciliated epithelium. The spinal cord consists of an outer portion, mainly of nervous fibres, the white matter, and of inner, ganglionated, and more highly vascular grey matter. (In the cerebrum the grey matter is external, and the white internal.) The cord, like the brain, is surrounded by a vascular fibrous investment, and protected from concussion by a serous fluid. The nerves which emerge from the vertebral column between the vertebrae, arise, unlike the cranial nerves, by two roots. The dorsal of these, the sensory root (d.n.), has a swelling upon it, the dorsal ganglion, and-- by experiments upon living animals-- has been shown to contain only afferent fibres; the ventral, the motor root, is without a ganglion, and entirely or mainly motor. The two unite outside the cord, and thereafter the spinal nerves are both sensory and motor.
Section 132. Besides the great mass of brain and spinal cord (cerebro-spinal axis), there is, on either side of the dorsal wall of the body cavity, a sympathetic nervous chain. The nerve fibres of this system, like the nerve fibres of invertebrates, are non-medullated. It may be seen as a greyish thread running close by the common carotid in the neck (sym., [Figure 1]); it then runs over the heads of the ribs in the thorax and close beside the dorsal aorta in the abdominal region. In the anterior region of the neck it dilates to form a superior cervical ganglion, and opposite the first rib it forms an inferior cervical ganglion. Thence, backwards, there is a ganglion on each sympathetic chain opposite each spinal nerve, and the two exchange fibres through a thread, the ramus communicans. To the sympathetic chain is delegated much of the routine work of reflex control of the bloodvessels and other viscera, which would otherwise fall upon the spinal cord.
Section 133. There are eight cervical (spinal) nerves, one in front of the atlas, and one behind each of the cervical vertebrae. The last four and the first thoracic (spinal) contribute to a leash of nerves running out to the fore limb, the brachial plexus (plexus, literally network, but here meaning a plaited cord). The fourth cervical also sends down a phrenic nerve (p.n., [Figure 1]), along by the external jugular vein and the superior caval vein to the diaphragm. The last three lumbar and the sacral nerves form a sacral plexus, supplying the hind limb.
Section 134. From the sympathetic in the hinder region of the thorax a nerve, the great splanchnic nerve, arises, and runs, back to a ganglionated nervous network, just behind the coeliac artery, into which the vagus also enters; this is the coeliac ganglion, and together with a similar superior mesenteric ganglion around the corresponding artery, makes up a subsidiary visceral nervous network, the solar plexus. A similar and smaller nervous tangle, bearing an inferior mesenteric ganglion, lies near the inferior mesenteric artery.
Section 135. Finally, we may note the pineal gland and the pituitary body, as remarkable appendages above and below the thalamencephalon. Their function, if they have a function, is altogether unknown. Probably, they are inherited from ancestors to whom they were of value. Such structures are called reduced or vestigial structures, and among other instances are the clavicles of the rabbit, the hair on human limbs, the little pulpy nodule in the corner of the human eye, representing the rabbit's third eyelid, and the caudal vertebrae at the end of the human spinal column. In certain lowly reptiles, in the lampreys, and especially in a peculiar New Zealand lizard, the pineal gland has the most convincing resemblance to an eye, both in its general build and in the microscopic structure of its elements; and it seems now more than probable that this little vascular pimple in our brains is a relic of a third and median eye possessed by ancestral vertebrata. The pituitary body is probably equivalent to a ciliated pit we shall describe in the lancelet (Amphioxus).
8. _Renal and Reproductive Organs_
Section 136. We have now really completed our survey of the individual animal's mechanism. But no animal that was merely complete in itself would be long sanctioned by nature. For an animal species to survive, there must evidently, also, be proper provision for the production of young, and the preservation of the species as well as of the individual. Hence in an animal's physiology and psychology we meet with a vast amount of unselfish provision, and its structure and happiness are more essentially dependent on the good of its kind than on its narrow personal advantage. The mammalia probably owe their present dominant position in the animal kingdom to the exceptional sacrifices made by them for their young. Instead of laying eggs and abandoning them before or soon after hatching, the females retain the eggs within their bodies until the development of the young is complete, and thereafter associate with them for the purposes of nourishment, protection, and education. In the matter of the tail, for instance, already noted, the individual rabbit incurs the disadvantage of conspicuousness for the rear, in order to further the safety of the young.
Section 137. The female organs of reproduction are shown in [Sheet 10]. The essential organ is the ovary (ov.), in which the ova (eggs) are formed. [Figure 3] gives an enlarged and still more diagrammatic rendering of the ovary. There is a supporting ground mass, or stroma, into which numerous bloodvessels and nerves enter and break up. The ova appear first as small cells in the external substance of the ovary (as at 1), and move inward (2 and 3), surrounded by a number of sister cells, which afford them nourishment. At (4) an ovum with its surrounding group of cells is more distinct and near the centre of the ovary; a fluid is appearing within the ovisac as the development proceeds. (5) is a much more mature ovisac or Graafian follicle.
Section 138. The ovum (ov.), is now large, and its nucleus and nucleolus (the germinal vesicle and spot) are very distinct. The wall of the follicle consists, in the mammal, of several layers of cells, the membrana granulosa (or "granulosa" simply); the ovum lies on its outer side embedded in a mass of cells, discus proligerus, separated from actual contact with the ovum by a zona pellucida. The ripening follicle moves to the surface of the ovary and bursts, the ovum falls into the body cavity. In [Figure 2], a ripe Graafian follicle (G.F.), projects upon the ovary.
Section 139. The liberated ovum is caught up by the funnel-shaped opening of the Fallopian tube, which passes without any very conspicuous demarcation into the cornu uteri (c.ut.) of its side; the two uterine cornua meeting together in the middle line form the vagina (V.), which runs out into a vestibule (vb.) opening between tumid lips to the exterior. The urinary bladder (ur.b.) also opens into the vestibule, and receives the two ureters from the kidney.
Section 140. In the male we find, in the position of the female uterus, a uterus masculinus (u.m.). The essential sexual organ is the testis (T.), a compact mass of coiling tubuli, which opens by a number of ducts, the vasa efferentia, into a looser and softer epididymis (ep.), which sends the sexual product onward through a vas deferens (v.d.), to open at the base of the uterus masculinus. The urinary bladder and ureters correspond with those of the female, and the common urogenital duct (= vestibule), the urethra, is prolonged into an erectile penis (P.) surrounded by a fold of skin, the prepuce. A prostate gland (pr.), contributes to the male sexual fluid. The character of the essential male element, the spermatozoon, the general nature of the reproductive process, will be conveniently deferred until the chapters upon development are reached.
9. _Classificatory Points_
Section 141. The following facts of classificatory importance may now be considered, but their full force will be better appreciated after the study of other vertebrate types. They are such as come prominently forward in the comparison of the rabbit with other organisms.
Section 142. In the first place, the rabbit is a metazoon, one of the metazoa, i.e., a multicellular organism, as compared with the amoeba, which belongs to the protozoa or one-cell animals ([Section 55]). In the next place, it is externally bilaterally symmetrical, its parts balance, and where, in its internal anatomy, it departs from this symmetry (as in the case of the aorta, the stomach and intestines, and the kidneys), the departure has an appearance of being the results of partial reductions and distortions of an originally quite symmetrical plan. And the facts of development strengthen this idea; in the very earliest stages we have paired aortic arches, of which, the left only remains, a straight alimentary canal, and less asymmetrical kidneys. In the vast majority of animals the same bilateral symmetry is to be seen, but in the star-fish and sea-urchins, and in the jelly-fish, corals, sea anemones, and hydra, the general form of the animal is, instead, arranged round a centre, like a star and its rays, and the symmetry is called radial.
Section 143. We also see in various organs of the rabbit, and especially in the case of the limbs and vertebral column, what is called metameric segmentation, that is, a repetition of parts, one behind the other, along the axis of the body. Thus the bodies and arches of the vertebrae repeat each other, and so do the spinal nerves. The renal organ of the rabbit, some time before birth, displays a metameric arrangement of its parts; but this disappears, as development proceeds, into the compact kidney of the adult. But the metameric segmentation in the rabbit's organism is not nearly so marked as that of an earthworm, for instance, which is visibly a chain of rings. If the student wants a perfect figure of metameric segmentation he should think of a train of precisely similar carriages, or a string of beads. One bead, one carriage, one vertebra, would be a metamere.
Section 144. In contrast to metameric segmentation is the antimeric repetition of radial symmetry ([Section 142]), in which each ray of the star is called an antimere. It is possible to have bilateral symmetry without a metameric arrangement of parts, as in the mussel and the cuttle-fish; but metameric segmentation without complete or reduced bilateral symmetry does not occur.
Section 145. We are now in a position to appreciate the fact that the old and more popularly know division of animals into vertebrata and invertebrata scarcely represents the facts of the case, that the primary division should be into protozoa and metazoa, and that the vertebrata are one of several groups of metazoa with a fundamental bilateral symmetry and imperfect metameric segmentation.
The rabbit is one of the vertebrata, and, in common with all the other animals collected under this head, it has--
(a) A skeletal axis (the vertebral column) between its central nervous system and its body cavity. In the adult rabbit this consists of a chain of vertebrae, but in the embryo (i.e., the young rabbit before birth) it is represented by a continuous chord, the notochord, and it remains as such in some of the lowest vertebrata throughout life. In other words, in these lower vertebrata, the vertebral axis is not metameric.
(b) A dorsal and -Tubular_ nervous axis. ([Section 131], the central canal)
(c) It has, though in the embryo only, certain slits between the throat and the exterior, like the gill slits of a fish. Such slits are-- with one or two remarkable exceptions outside the sub-kingdom-- distinctly vertebrate features, and remain, of course, in fishes throughout life.
The presence of true cartilage and bone mark a vertebrate, but vertebrata occur in which -these tissues- [bone] -are- [is] absent.
Section 146. The rabbit shares the following features with all the vertebrata, except the true fishes, which do not possess any of them--
(a) Lungs (but many fish have a swimming bladder which answers to the lungs in its anatomical relations.)
(b) Limbs which consist of a proximal joint of one bone an intermediate part of two, and a distal portion which has five digits, or is evidently a reduced form of the five-digit limb.*
(c) The absence of a median fin supported by fin rays.**
* The frog shows indications of a sixth digit.
** The frog's tadpole has a median fin, but no fin rays.
Section 147. The rabbit shares the following features with all the vertebrata above the fishes and amphibia (= frogs, toads, newts, and etc.)--
(a) Absence of gills (not gill slits, note) at any stage in development.
(b) An amnion, and
(c) An allantois in development.
The meaning of (b) and (c) we shall explain to the student in the chapters on embryology. We simply mention them here to render our table complete.
Section 148. The rabbit shares with all mammals, and differs from all other vertebrata (i.e., birds, reptiles, amphibia, and fishes), in having--
(a) Hair.
(b) A diaphragm.
(c) Only one aortic arch, and that on the left side of the body.
(d) Its young born alive. (But two very reptile-like mammals of Australia, the duck-billed platypus and the echidna, lay eggs, and certain fish and reptiles bear living young.)
(e) Epiphyses to its vertebral -centre- [centra].*
(f) The cerebral hemispheres covering the mid-brain.
(g) Corpora quadrigemina instead of bigemina.
[(h) A corpus callosum.]
[(i) A spirally coiled cochlea to the internal ear.]
[(In respect to h and i also, the echidna and platypus are scarcely mammalian.)]
* But certain mammals have no such epiphyses.
Section 149. The rabbit, together with the hares and conies, rats and mice, voles, squirrels, beavers, cavies, guineapigs is included in that order of the class of mammals which is called the rodentia, and is distinguished by the character of the incisor teeth from other orders of the class.
10. _Questions and Exercises_
- Describe the venous circulation of the rabbit (with diagrams).
Compare a vein and artery. Compare the distribution of the great
venous trunks with that of the arterial system.
- Construct a general diagram of the circulation of the rabbit, to show
especially the relation of the portal system, the lymphatics and
lacteals, and the renal circulation to the main blood current.
- Draw the alimentary canal of the rabbit from memory.
- What is a villus? Describe its epithelium, and the vessels within it.
Write as explicit an account as you can of the absorbent action of a
villus.
- Tabulate the alimentary secretions, and their action on the food.
- What is botryoidal tissue? Where does it occur? What is known
of its functions?
- Copy Diagram I. (enlarged), and insert upon it the visceral nerves
as far as you can.
- What are the most characteristic points in the mammalian vertebral
column?
- Describe cartilage and bone, and compare them with one another.
- Give an account of the amoeba, and compare it with a typical
tissue cell in a metazoon (e.g., the rabbit).
- Give a general account of connective tissue. What is tendon?
- Trace, briefly, the increased modification of tissues in the
vertebrata.
- Describe, with diagrams, the structure of blood. State the function
of each factor you describe.
- Compare the pectoral with the pelvic limb and girdle. What other
structures of the adult rabbit display a similar repetition of similar
parts?
- Draw from memory typical vertebrae from each region of the
vertebral column.
- What are bilateral symmetry and metameric segmentation?
- Give a schedule of distinctive mammalian features.
- Describe the rabbit's brain (with diagrams).
- Give a list of the cranial nerves of the rabbit, and note their origin
in the brain.
- Give a list of the nerve apertures of the dog's skull.
- What are the chief anatomical differences between a typical
cranial, a spinal, and a sympathetic nerve?
- Describe and figure the distribution of nerves V., VII., IX., and X.
- Describe the muscles, glands, and nerves of the orbit of the
rabbit.
- Describe, with figures, the eye of the rabbit.
- Give a diagram of the rabbit's internal ear.
- Draw and describe the ear ossicles. What is their function?
- Draw and state the precise position of the hyoid bone, the
clavicle, the calcaneum, and the olecranon process.
- Describe, as accurately as possible, the position of palatine
bones, pterygoids, the ethmoid bone, the pre- and basi-sphenoids,
in the dog's skull.
- What is membrane bone? What is cartilage bone? Discuss their
mutual relationship.
- What is an excretion? What are the chief excretory products of an
animal? How are they removed?
- Describe the minute anatomy of the liver. Give a general account
of its functions.
- Describe the minute anatomy of the kidney, and the functions of
the several parts.
- What is ciliated epithelium? Where does it occur in the rabbit?
- Describe the mechanism of respiration. What is the relation of
respiration to the general life of the animal?
- What are the functions of the skin? Describe its structure.
- What is a secretion? Tabulate and classify secretary organs.
What is a goblet cell?
- Draw, from memory, the dorsal and ventral aspects of, and a
median section through, a dog's skull.
- Name any structures that appear to you to be vestiges or
rudiments, i.e., structures without adequate physiological reason, in
the rabbit's anatomy.
- How are such structures interpreted?
- Describe the structure of striated muscular fibre. Describe its
functions, and the various means by which they may be called into
activity.
- Describe the characters and structure of the blood of the rabbit.
What is the lymphatic system? Describe its relation to the blood
system in a mammal.
- Describe the structure of (a) blood, (b) hyaline cartilage, (c)
bone, in the rabbit; (d) point out the most important resemblances and
differences between these tissues; (e) state what you know of the
development of the same tissues.
- Draw diagrams, with the parts named, of the male and female
generative organs of the rabbit.
- In the rabbit provided dissect on one side and demonstrate by
means of flag-labels the main trunk of the vagus nerve, the phrenic
nerve, and the recurrent laryngeal nerve.
- Dissect the rabbit provided so as to expose the abdominal viscera. Mark with flag-labels the duct of the pancreas, the ureters, and the oviducts or the sperm ducts (as the case may be).
[Many of the above questions were actually set at London University Examinations in Biology.] {In Both Editions.}