Chapter XX: Section V (2)

Klein describes a second system of ganglion-cells, situated in meshes, which are formed by the nerve-trunks of the plexus itself. These ganglion-cells are much larger than the former, and are multipolar; their protoplasm, which is distinctly fibrillar, with granules between the fibrils, is provided with one or two long, thick processes and several short and thin ones; generally the processes are branched. The general shape of the cells is oblong, the thick, long processes being commonly at the two opposite poles. The cells are generally isolated, sometimes situated in the centre of a mesh, or more commonly near a nerve-trunk that borders the mesh on one side. Each ganglion-cell is connected with a nerve-trunk of the plexus by at least one process. In a few of the nerve-trunks of the general plexus, isolated medullated nerve-fibrils are seen to pursue an almost straight course from one trunk into another and divide into two. There is no connection between these medullated fibres and the ganglion-cells.

*Auerbach’s plexus* is a much finer plexus than that just described, and the ganglia are much smaller. It supplies twigs to the _Muscularis mucosa_; these break up into fine fibrils, which follow the direction of the muscle-fibres; other twigs supply the blood-vessels, with which they can be traced into the bases of the folds of mucous membrane. Thanhoffer has recently (_l. c._) described nerve-fibrils, which terminate in the mucous epithelium.]

D. The _Large Intestine_ (Fig. 184 _R_).

From a transverse section of the large intestine of _Rana temporaria_, moderately distended.--G. H.

_A_ Denser portion of submucous coat.
_B_ Looser portion of submucous coat.
_C_ Artery, cut obliquely.
_D_ Circular muscle-layer.
_E_ Longitudinal muscle-layer.
]

*a.* *General description.* The large intestine is a flask-shaped viscus, lying in the median line. The small intestine opens into it by an abrupt curve at its anterior end. The large intestine is the widest part of the alimentary canal, is thin-walled, and diminishes in width towards its hinder end, where it opens into the cloaca above the opening of the bladder.

*b.* [*Minute structure.*

(1) The *serous coat* resembles that of the small intestine, etc.

(2) The *muscular coats* resemble those of the small intestine, but are thinner; the longitudinal layer is, however, proportionally thicker (Wiedersheim).

(3) The *submucous coat* (Fig. 192) resembles that of the small intestine in its general structure; the portion lying immediately beneath the mucous membrane (_B_) is denser in structure.

This layer has no _Muscularis mucosa_.

(4) The *mucous coat* (_A_) is a simple layer of large, columnar, epithelial cells, with large oval nuclei. The cells have a hyaline free border, but this possesses no striation. The mucous membrane is usually described as possessing numerous simple follicles (glands of Lieberkühn); in those specimens which I have examined they have been entirely absent (Figs. 192, 193). At six to eight points in a transverse section of a rectum moderately distended the submucous coat is thinner, and so throws the mucous coat into slight, longitudinal grooves; but these do not in the least resemble the glands of Lieberkühn.

Transverse section of large intestine of _Rana esculenta_; the mucous membrane thrown into longitudinal folds in consequence of the organ being contracted. Arteries injected with carmine.--G. H.

_A_ Large circular vessels within the muscular coats.
_B_ Fine anastomosis to the mucous coat.
]

(5) The epithelium possesses goblet- or chalice-cells (Hoffmann), the number of which probably depends upon the period of the year and the state of digestion. In the rectum, from which the section for the figures 192 and 193 were cut, not one goblet-cell was found in a complete series of sections.

If the rectum be contracted, the mucous membrane is thrown into longitudinal folds.

(6) The *blood-vessels* (Fig. 193) have a simple arrangement. The arteries are large, and form oblique loops around the intestine, lying under the peritoneum; from these branches are given off to form a series of rings in the submucous membrane (Fig. 193 _A_); from these fine twigs are given off to form a fine anastomosis under the mucous membrane (_B_).

(7) The *lymphatics* of the large intestine are arranged in two chief systems: one under the serous coat resembles the corresponding system of the small intestine. The second set forms a network of rounded loops in the submucous coat, which give rise to a secondary set of smaller vessels towards the mucous membrane; this secondary system forms a sort of trellis-work standing on the rounded loops, and so maps out small blocks of the thick submucous coat. Towards the cloaca the arrangement is simpler; the secondary lymphatics are lost, while the primary lymphatics tend to form elongated, longitudinal loops (Langer).]

II. THE GLANDS CONNECTED WITH THE INTESTINAL CANAL.

A. The *Liver* and *Gall-bladder*.

*a.* *External form.*

(1) The liver (Figs. 185, 194) is a large, reddish-brown organ, occupying a large part of the anterior abdominal region. It consists of three or more lobes, which present many individual variations; as a rule there are two larger lateral lobes (_L_ and _L^1_), and a smaller median lobe (_L^2_). Each lobe has a superficial or ventral surface, which is convex, and a deep or dorsal surface, which is concave and directed towards the other abdominal viscera lying above the liver. The two surfaces of each lobe meet to form a sharp border around the lobe, except where the three lobes are more intimately attached, opposite the apex of the heart (Figs. 185, 194); at this place each lobe possesses a small, flat, or slightly concave anterior surface.

The left lobe (_L_) hides the greater portion of the stomach, and has near its inner border a deep fissure, which runs forwards and so marks off a more or less well-marked fourth lobe. The anterior portion of the left lobe is attached to the corresponding portion of the right lobe by a narrow commissure.

The liver, seen from the ventral surface.

_Du_ Duodenum.
_H_ Heart.
_L_ Left lobe of liver.
_L^1_ Right lobe of liver.
_L^2_ Middle lobe of liver.
_M_ Stomach.
]

The median lobe (_L^2_) extends backwards as far as the pylorus, and covers the commencement of the intestine together with the pancreas; these organs can, therefore, not be seen until this lobe is displaced. A fifth lobe is sometimes found on the dorsal surface of the median lobe, and to it or to the dorsal surface of the median lobe the small intestine is attached by the _Ligamentum hepato-duodenale_ (Fig. 195 _Lhp_). The _Vena portarum_ enters the liver behind this ligament.

The right lobe extends much further dorsalwards than the left lobe, and even comes in contact with the lung, the vessels of which sometimes indent its surface in spirit specimens. This lobe is also in contact with the base of the fat-body, and in females with the oviduct.

By drawing the lobes of the liver to either side and displacing the heart towards the head, the posterior caval vein is seen passing from the liver to the heart, and the hepatic commissure joining the lateral lobes is brought into view.

(2) The *gall-bladder* (Fig. 195 _G_) is placed on the dorsal surface of the liver in the deep niche between the right and left lobes; it is attached to the liver by connective-tissue and peritoneum.

The gall-bladder is round or oval in form; when moderately full it has a smooth, outer wall, which is thin and allows the green colour of the bile to be seen. It possesses a duct (_Dcy_), the cystic duct (_Ductus cysticus_), which bifurcates near its origin.

The two cystic ducts (Fig. 195 _Dcy_) join the larger hepatic ducts, as shown in the figure, and so form a simple anastomosis, from which three branches (3) of varying size unite at the anterior extremity of the pancreas to form the common bile-duct (_Ductus choledochus_, _Dc_). The common bile-duct runs through the whole length of the pancreas, receiving near its origin additional hepatic ducts (_Dh^1_) from the middle lobe of the liver. In this course the duct lies either on the ventral surface of the pancreas or under a thin layer of the glandular tissue; it receives the ducts of the pancreas and leaves that organ at its posterior border as a round and strong canal (_Dc^1_). The duct courses in the gastro-duodenal ligament, and reaches the dorsal surface of the duodenum at a very acute angle; it then pierces the wall obliquely and terminates with a slit-like or elongated oval opening.

The pancreas and bile-canals. The liver has been displaced towards the head.

_Dc_ Common bile-duct.
_Dc^1_ Common bile-duct after leaving the pancreas.
_Dc^2_ Opening of the common bile-duct into the duodenum.
_Dcy_ Cystic ducts.
_Dh_ Hepatic ducts.
_Dh^1_ Supplementary hepatic ducts from the middle lobe
of the liver.
_Du_ Duodenum.
_G_ Gall-bladder.
_L_ Left lobe of the liver.
_L^1_ Right lobe of the liver.
_L^2_ Middle lobe of the liver.
_L^3_ Fourth lobe of the liver.
_Lhp_ Gastro-hepatic ligament.
_M_ Stomach.
_P_ Pancreas.
_Py_ Pylorus.
_P^1_ Ducts of the pancreas.
]

*b.* [*Minute structure.*

(1) The *liver* is composed of various tissues: it possesses a peritoneal covering, a fibrous covering, which supplies trabeculae to support the various other tissues, blood-vessels, hepatic cells or true liver parenchyma, and bile-canals.

α. The *peritoneal covering* of the liver encloses the organ almost completely, the only exceptions being where the various attachments of the liver are found (see peritoneum). The peritoneal covering of the liver is for the most part composed of flattened, ciliated cells (Neumann and Grunau); on the middle lobe these are, however, more or less replaced by non-ciliated cells. The thickness of these cells varies considerably, according to the amount of distension to which the liver is subjected.

β. The *fibrous covering* of the liver is very thin and very difficult to demonstrate; it consists of connective-tissue fibres with very few corpuscles. This covering is prolonged into the liver along the portal canals, where traces of connective-tissue can always be made out.

From these processes and from the whole of the inner surface of the general connective-tissue capsule are given off fine trabeculae, in which it is very difficult to find any nuclei; these trabeculae are everywhere extremely delicate and difficult to demonstrate; nowhere do they form distinct boundaries between lobuli: the structure closely resembles the sustentacular tissue of a lymphatic gland (Eberth).

γ. The *blood-vessels* to the liver are the portal vessels (p. 249) the hepatic veins (p. 247), and the hepatic artery[74] (p. 233)

[Footnote 74: See Plate II, Fig. 196.

I. Partial injection of the liver from the portal vein (blue): _Rana esculenta_.--G. H.

II. Partial injection of the liver from the hepatic vein (red): _Rana esculenta_.--G. H.

III. Complete injection of the liver from the hepatic artery (red) and from the portal vein (blue): _Rana esculenta_.--G. H.

_A_ Portal (interlobular) veins and their branches.
_B_ Hepatic (intralobular) veins and their branches.
_C_ Hepatic arteries and their branches.
]

(1) The portal vessels pass into the liver on its ventral surface; they divide into branches which course along the middle parts of each lobe and give off smaller branches in all directions towards the periphery; the interlobular branches (Fig. 196 I) forming a very complex capillary network. As compared with the hepatic veins (Fig. 196 II), the interlobular veins do not give off their capillaries so abruptly, but tend to supply these from small lateral branches. The portal veins are accompanied in their course by branches of the hepatic artery, and often by larger bile-ducts, and thus form portal canals. In no part do the portal (interlobular) veins or their branches limit the lobules by distinct rings of vessels, as seen in many higher animals. The interlobular veins and intralobular veins simply interdigitate with each other.

(2) The hepatic veins (Fig. 196 I and II) also course chiefly in the middle parts of the lobes of the liver; they branch, and ultimately supply intralobular veins which interdigitate with the interlobular veins (I and II). The capillaries arise very abruptly from an intralobular vein, and form a network of vessels communicating very freely with the corresponding capillaries of the interlobular veins.

(3) The hepatic arteries (Fig. 196 III) break up into small branches which, as a rule, course along the portal canal until near the surface of the liver, when they leave the portal veins and pass to the surface to supply the coverings of the liver. In their course along the portal canals they supply a few very small twigs to the structures forming the canals. At the surface of the liver the branches of the hepatic artery form capillaries, which empty themselves into the general capillary anastomosis beneath (Fig. 196 III).

δ. The *liver-cells* (Figs. 197, 198) are large, and of compressed spheroidal or polygonal shape. They possess no cell-wall, have usually one but sometimes two large nuclei, each with a distinct nucleolus. The cells often contain granules of glycogen or fat-globules. The main fibrillae of the intercellular network are arranged so as to extend between a bile-capillary and a blood-capillary (Fig. 197).

ε. The *bile-ducts* (Fig. 198) commence as fine tubes between the liver-cells, where they are simply small spaces enclosed by the hepatic cells; they are usually enclosed by three or four cells, possibly sometimes by only two cells (Hering). Such bile-capillaries are usually separated from a blood-capillary by the thickness of one liver-cell only (Fig. 198).

As a number of such bile-capillaries run together to form a larger duct, the cells enclosing them change their character, becoming flattened and broader; these cells may, however, be traced continuously into the true hepatic cells (Hering and Eberth). The bile-duct so formed then obtains a slight covering of fibrous tissue, which rapidly increases in quantity, courses along a portal canal, and receives other ducts on the way; the epithelium lining it gradually becomes more elongated, and ultimately resembles that found in the common bile-duct or the gall-bladder.

In the larger ducts the epithelium is, according to v. Brunn, ciliated; they also possess a layer of unstriated muscle-fibre (Eberth).]

Liver-cells, after Kupffer.]

ζ. The *pigment* of the liver varies very much in amount and character, according to the time of the year and state of health of the animal. According to Eberth the pigmentary masses are of about the same size as the white blood-corpuscles, and are possessed of the power of amoeboid movement (in young animals). The cells possess two to seven nuclei and vary much in colour and distribution. As a rule the larger the amount of pigment in a given liver the smaller is the number of fat-globules found in the individual liver-cells (Eberth).]

The bile-capillaries; natural injection with sulphindigotate of sodium: v v v represent blood-capillaries.--G. H.]

(2) [The *gall-bladder* and *bile-ducts*. The gall-bladder has four coverings.

α. A serous coat of peritoneal endothelium.

β. A muscular coat, containing unstriated muscle-fibres and connective-tissue.

γ. A sub-mucous coat of areolar-tissue.

δ. An internal lining of columnar epithelium.

ε. The walls of the gall-bladder are richly supplied with blood-vessels from the cystic arteries (p. 233); these form a close network in the submucous coat.

ζ. The muscular and submucous coats also possess a rich nervous plexus, which contains ganglia and resembles Auerbach’s plexus of the intestine (Popoff, Gerlach).]

B. The *Pancreas* (Fig. 199 _P_).

*a.* *General description.* The pancreas is a flattened, light yellowish-brown organ, placed in the loop of the duodenum between this latter and the stomach. The whole organ is within the gastro-duodenal ligament, and is attached to the liver; hence it is little influenced by changes in the amount of distension of the stomach or intestine.

The pancreas and bile-canals. The liver has been displaced towards the head.

_Dc_ Common bile-duct.
_Dc^1_ Common bile-duct after leaving the pancreas.
_Dc^2_ Opening of the common bile-duct into the duodenum.
_Dcy_ Cystic ducts.
_Dh_ Hepatic ducts.
_Dh^1_ Supplementary hepatic ducts from
the middle lobe of the liver.
_Du_ Duodenum.
_G_ Gall-bladder.
_L_ Left lobe of the liver.
_L^1_ Right lobe of the liver.
_L^2_ Middle lobe of the liver.
_L^3_ Fourth lobe of the liver.
_Lhp_ Gastro-hepatic ligament.
_M_ Stomach.
_P_ Pancreas.
_P^1_ Ducts of the pancreas.
_Py_ Pylorus.
]

The organ may be completely exposed by either of two methods: in the former, the liver, stomach, and duodenum are drawn towards the head and the dorsal surface of the organ so exposed. In the second method the liver is drawn backwards, the various peritoneal folds which connect the duodenum with the posterior border of the liver cut through, and the three organs then separated; the pancreas can then be conveniently examined.

The size and shape of the pancreas are subject to great variations in different specimens. The usual shape of the organ is somewhat triangular (Fig. 199), the left border being usually unbroken, while the other two shorter borders are broken into lobes. The longest lobe stretches as far as the pylorus (_Py_), to which it is attached by connective-tissue: the opposite extremity of the gland is attached to the liver.

The excretory duct of the pancreas (_Ductus Wirsurgianus_) opens into the common bile-duct at about the middle of the pancreas (_P^1_); other smaller ducts may open into the same canal.

*b.* *Minute structure.* [The pancreas consists of a number of lobes loosely held together by connective-tissue; each lobe is composed of a number of lobules attached to each other much more intimately. The lobules are made up of tubes lined by a single layer of glandular epithelium. This epithelium is, as a rule, broadly columnar, but in the smaller tubes may be cubical or polygonal. Each cell has a cell-wall, nucleus, and very granular protoplasm; the latter shows two zones (in the inactive condition), a granular zone near the lumen, and an outer, clear, and finely striated zone (Nussbaum). The lumen of each alveolus is very small, and in many cases difficult to make out.

The smallest ducts have no special lining, and are therefore bounded by the glandular cells; the larger ducts have a flattened, cubical epithelium, which when seen from the surface has an appearance as if the individual cells were widely separated from one another; these ducts have a considerable layer of connective-tissue around them. The largest ducts are lined with a layer of columnar, ciliated epithelium; the cilia are very long, usually of about the same length as the cell, sometimes appearing to be even longer. This epithelium is continuous with the ciliated, epithelial layer of the common bile-duct, which it resembles. These largest ducts have an extremely thick layer of connective-tissue around them.

The pancreas and its ducts are very richly supplied with vessels and nerves; the nerves forming everywhere a fine plexus, the larger strands of which usually, but not always, course with the larger blood-vessels. The larger ducts have an especially rich supply of nerves, of which fibrils may be traced towards the ciliated epithelium; an anatomical connection between the two has not, however, been made out.]

III. THE SPLEEN.

*a.* *General description.* Although the spleen belongs to the lymphatic system, it is considered here in order to complete the description of the abdominal viscera. It is a small, rounded-oval body, of a reddish-brown colour, suspended in the mesentery near the anterior end of the large intestine (Fig. 184 _Mz_). In medium-sized animals the longer diameter is about 6 mm., and is parallel with the long axis of the body; the shorter diameter is about 5 mm., and the thickness varies from about 3 mm. to 4 mm. The dorsal surface is flat or slightly concave (_Hilus lienis_), and receives the relatively large afferent and efferent vessels; the rest of the organ is smooth and rounded, and with its greatest convexity directed towards the left side.

*b.* [*Minute structure.* The structure of the spleen resembles that of higher animals. It possesses a *serous coat* of peritoneum, under which is a *fibrous coat*; the latter sends in trabeculae, which divide and form a fine meshwork of supporting-tissue; the finest trabeculae are formed by the processes of the connective-cells of the sustentacular structure. According to Hoffmann, the thickness of the fibres averages 0.001 to 0.011 mm.; the intervening spaces measure 0.002 to 0.012 mm. The spaces are filled by the spleen pulp, which consists of true spleen-corpuscles, blood-corpuscles, and pigment-corpuscles.

The *spleen-corpuscles* have an average diameter of 0.006 mm., and are round or of a rounded oval form. Each consists of a nucleus, with a very small amount of adherent protoplasm; the nucleus possesses one or two nucleoli. Some of these cells contain a brownish or black pigment in granules, but most of them are colourless (Hoffmann).

The *pigment-cells* equal the white blood-corpuscles in size, and exactly resemble the corresponding pigment-cells of the liver.

The *blood-corpuscles* are found in various stages of disintegration and regeneration.

The *arteries*, on entering the spleen, at once break up into branches which pass in all directions, giving off twigs on all sides and at varying angles; from these capillaries are supplied, which traverse the parenchyma in all directions. The capillaries empty themselves partly into veins, partly into the splenic spaces.

The *veins* commence either as capillaries in connection with the arteries or by communicating with the splenic spaces. This communication is brought about by small twigs of about 0.015 mm. diameter, which have incomplete walls, and so open into the splenic spaces (Hoffmann).

*Malpighian bodies* are represented by collections of splenic cells on various arterial twigs; they are, however, not so sharply defined as is the case in some higher animals.]

IV. THE PERITONEUM.

*a.* *General description.* The *peritoneum* is a thin, pigmented membrane lining the abdominal cavity. Tracing it forwards from the ventral wall of the abdomen (_Peritoneum parietale_), it can be followed along the deeper surface of the muscles to the pericardium. The middle portion leaves the abdominal wall by accompanying the anterior abdominal vein; the lateral portions are continued further forwards, and then ascend on the pericardium and the deeper surfaces of the lateral walls in the thoracic region.

The peritoneum passes thence to the ventral surface of the liver (_Ligamentum coronarium_), covers this surface and passes on to the dorsal surface of the organ, which, together with the gall-bladder, it completely encloses. The membrane thus reaches the dorsal wall (_Ligamentum suspensorium hepatis et pericardii_); from the lateral borders of the liver it passes upwards to the dorsal wall, and thus forms a pocket-like pleuro-peritoneal cavity on either side.

From the dorsal wall and above the attachment of the coronary ligament of the bladder the peritoneum reaches the root of the lung on each side, and completely invests the organ: while in the middle line it covers the outer surface of the oesophagus and attaches it to the dorsal wall, thus forming the first part of the mesentery.

Just behind the root of the lung, the peritoneum, in female specimens, has an opening on each side (_Ostium abdominale tubae Fallopiae_), by which the oviduct communicates with the peritoneal cavity.

Tracing the peritoneum backwards, it passes over the ventral surface of the kidneys so as to exclude them from the peritoneal sac: in the middle line, between the kidneys, the peritoneum descends to form the mesentery for the small intestine. At the inner borders of the kidneys, the testes or ovaries are pushed into the abdominal sac, and so possess well-marked mesenteries; the mesovarium becomes longer towards the cloaca, but attains its greatest development in the breeding season, when it is arranged in numerous folds. Along the outer borders of the kidneys, in females, the peritoneum again descends into the abdominal cavities to enclose the oviducts, which have broad mesenteries. Towards the rectum these mesenteries are shorter and attached to either side of the bladder by a well-marked free border: as the middle line of the bladder is attached by the peritoneum to the rectum, two distinct pouches (_Cava recto-vesicalia_) are formed, which descend deeply into the pelvic cavity. The upper walls of these pouches are pushed in between the urostyle and the rectum, and together form a strong meso-rectum, which is longer near the _Valvula Bauhinii_ and continuous with the mesentery of the small intestine. The hinder portion of this mesentery is very short, and only covers the lateral walls of the rectum.

With the exception of a small portion of its dorsal surface, which is attached to the rectum, the whole surface of the bladder is covered with peritoneum.

The mesentery of the alimentary canal commences in connection with the oesophagus between the roots of the lungs; it is attached to the dorsal surface of the liver, covering the posterior caval vein, and is attached to the gall-bladder. From this point it extends, as a free, arched fold, to the concave right border of the stomach, which is completely surrounded by peritoneum.

The gastro-duodenal fold (_Ligamentum gastro-duodenale_) extends from the stomach to the pylorus and includes the pancreas. The hepato-duodenal fold (_Lig. hepato-duodenale_) extends from the portal fissure of the liver to the duodenum.

The mesentery of the small intestine is broad and arranged in folds, which follow the curves of the intestine; and is attached in the middle line, immediately beneath the vertebrae, where it encloses the aorta.

The various folds and mesenteries carry the blood-vessels and nerves to the different organs; in this course the vessels are surrounded by large lymphatics, which communicate with each other.

Preparations from the peritoneum of _Rana esculenta_.--G. H.

I. From peritoneum of the ventral wall of abdomen, stained with silver and logwood (Hartnack, Oc. I, Syst. 7).

II. From mesentery of small intestine of _Rana esculenta_, stained with silver (Hartnack, Oc. I, Syst. 7).

III. Preparation to show ciliated cells between non-ciliated cells; after Neumann. IV. Vertical section at border of liver to show ciliated epithelium; after Neumann.

_A_ Stoma.
_B_ Pigment-cells.
_C_ Ciliated cells.
_D_ Non-ciliated cells.
]

*b.* *Minute structure* (Fig. 201).

[The peritoneum is a serous membrane, formed for the most part of a layer of irregular endothelial cells, arranged on a thin layer of subserous, connective-tissue (Fig. 201 I, II, and III).

The endothelial cells are attached to each other by cement-substance, easily stained by silver nitrate. The cells covering the general surface of the peritoneal cavity are larger and broader than those covering the mesentery of the small intestine (compare I and II, Fig. 201).

At various points stomata are found, bordered by smaller and more deeply staining epithelium (I, _A_). The membrane covering the general cavity is also much more pigmented than that covering the mesentery (compare I and II).

Various portions of the peritoneal surface possess ciliated cells, and these cells are usually thicker than the surrounding non-ciliated cells. Such cells are found especially near the openings of the oviducts and on the liver. The dimensions of the cells vary; according to Neumann the average dimensions of ciliated cells on the liver are: 0.006 mm. depth (without the cilia); nucleus, 0.012 mm. long and 0.003 mm. broad. The cells are five- or six-sided and bounded by straight sides (Neumann).]