GASTROINTESTINAL TRACT
The gastro intestinal tract (GIT) comprises the stomach, duodenum, jejunum, ileum, colon, rectum and anal canal. The GIT and oesophagus form the alimentary canal. The basic structure of the walls of the alimentary canal has been described already (see HA235 - Histology - Oral Cavity, Oesophagus).
The stomach functions both as a reservoir and as a digestive organ. It empties its contents in small portions (suitable for continued digestion) into the small intestine.
Anatomically, the stomach is divided into
- a cardiac part,
- fundus,
- body or corpus, and
- a pyloric part (pyloric antrum and pyloric canal)
Histologically, most of the layers of the wall of the stomach appear similar in its different parts. Regional differences are mainly restricted to the appearance of the gastric mucosa.
The Mucosa (epithelium, lamina propria, muscularis mucosae)
The mucosa is thrown into longitudinal folds (gastric folds or rugae), which disappear when the stomach is fully distended. A network of shallow grooves divides the mucosa into gastric areas (1-5 mm). On the mucosal surface we see small, funnel-shaped depressions (gastric pits). Almost the entire mucosa is occupied by simple, tubular gastric glands which open into the bottom of the gastric pits.
The structure and cellular composition of the surface epithelium (simple, tall columnar) does not change throughout the stomach. It contains mucus-producing cells, which form a secretory sheath (glandular epithelium). The mucus is alkaline and adheres to the epithelium. The mucus forms an ~ 1 mm thick layer, which protects the mucosa from the acidic contents of the stomach. The surface epithelium is renewed approximately every third day. The source of the new cells is the isthmus, i.e. the upper part of the neck, of the gastric glands, where cells divide and then migrate towards the surface epithelium and differentiate into mature
epithelial cells.
In contrast to the surface epithelium, cellular composition and function of the gastric glands are specialized in the different parts of the stomach.
Cardiac glands
Cardiac glands are heavily branched tubular glands (similar to the cardiac glands of the esophagus), which contain mainly mucus-producing cells. A few of the secretory cells characteristic for the corpus-fundic glands (see below) may be present.
Corpus-fundic (or principal) glands
Each glandular tubule (oriented more or less perpendicular to the surface of the epithelium) consists of three parts: a deep body, an intermediary neck and an upper isthmus. In corpus-fundic glands we find four cell types: chief cells, parietal cells, mucous neck cells and endocrine cells.
- Chief cells (or zymogenic cells)
-
are the most numerous of the four types. They occur primarily in the
body of the glands. They produce pepsinogen, which is a precursor of the
proteolytic enzyme pepsin.
The pH optimum of of pepsin is about 2. This enzyme
is able to break down collagen.
- Parietal cells (or oxyntic cells)
-
occur most frequently in the neck of the glands, where they reach the
lumen of the gland. They are situated deeper, between and below chief
cells, in lower parts of the gland. Parietal cells secrete the hydrochloric
acid of the gastric juice. Aside from activating the pepsinogen the hydrochloric
acid also effectively sterilizes the contents of the stomach.
Note that so far only one type of bacteria has found
which can live happily in the stomach - Helicobacter
pylori. Unfortunately these bacteria are involved in the pathogenesis
of gastritis and gastric ulcers.
Parietal cell also secrete intrinsic factor,
which is necessary for the resorption of vitamin B12.
Vitamin B12 is a cofactor of enzymes which synthesise
tetrahydrofolic acid, which, in turn, is needed for the synthesis of DNA
components. An impairment of DNA synthesis will affect rapidly dividing
cell populations, among them the
haematopoietic
cells of the bone marrow, which may result in pernicious
anemia. This condition may result from a destruction of the gastric
mucosa by e.g. autoimmune gastritis or
the resection of large parts of the lower ileum, which is the main site
of vitamin B12 absorption, or of the stomach.
- Mucous neck cells
-
are found between the parietal cells in the neck of the gland.
They are difficult to distinguish from chief cells
in plain H&E stained section.
- Endocrine cells
-
Endocrine cells are scattered, usually solitary, throughout the epithelium
of the gastro-intestinal tract. They are part of the gastro-entero-pancreatic
(GEP) endocrine system. The best characterized endocrine cells in the
gastric mucosa are gastrin-producing cells (G
cells) and somatostatin-producing cells (D
cells). G cells are most frequent in the middle third of the glands.
They stimulate the secretion of acid and pepsinogen. G cell function is
stimulated by nervous input, the distension of the stomach or secretagogues.
D cells are found mainly in glands of the pyloric antrum. They inhibit
G cells and thereby acid production. D cell function is stimulated by
acid in the lumen of the stomach and duodenum.
Other types of endocrine cells encompass VIP-producing
cells (or D1 cells; vasoactive intestinal
peptide) and serotonin-containing cells
(enterochromaffin cells). Endocrine cells in the gastrointestinal tract
are alternatively named APUD-cells: amine
precursor uptake
and decarboxylation cells.
Pyloric glands
Pyloric glands are more coiled than corpus-fundic glands, and they may be branched. Endocrine cells, in particular gastrin-producing cells, are more frequent than in corpus-fundic glands. A few parietal cells may be present but chief cells are usually absent.
The lamina propria is formed by a very cell-rich loose connective tissue (fibroblasts, lymphocytes, plasma cells, macrophages, eosinophilic leucocytes and mast cells). The muscularis mucosae of the stomach contains both circular and longitudinal layers of muscle cells. Its organization is somewhat variable depending on the location in the stomach.
Large blood vessels, lymph vessels and nerves are located in the submucosa which consists of loose connective tissue.
Note that the muscularis externa consists of three layers of muscles: an inner oblique layer, a middle circular layer and an outer longitudinal layer.
The small intestine is divided into duodenum (25-30 cm), jejunum (about first two-fifths of the rest) and ileum. The three segments merge imperceptibly and have the same basic histological organization.
The Mucosa
The mucosa of the small intestine has various structural features which considerably increase the luminal surface area and consequently support the main function of the small intestine - the absorption of the degraded components of the food.
Plicae circulares (of Kerkering) are macroscopically visible, crescent-shaped folds of the mucosa and submucosa. Plicae circulares extend around one-half to two-thirds of the circumference of the lumen of the small intestine.
Plicae circulares
- are permanent structures, i.e. their presence does not depend on the state of distension of the small intestine.
- are absent from the first few centimetres of the duodenum and the distal part of the ileum.
- are particularly well developed in the jejunum.
- increase the surface area of the mucosa by a factor of ~ three.
The entire intestinal mucosa forms intestinal villi (about one mm long), which increase the surface area by a factor of ~ ten. The surface of the villi is formed by a simple columnar epithelium. Each absorptive cell, or enterocyte, of the epithelium forms numerous microvilli (1 µm long and about 0.1 µm wide), which increase the surface area by a factor of ~ 20.
Between
the intestinal villi we see the openings of simple tubular glands, the crypts
of Lieberkühn. They extend through the lamina propria down to
the muscularis mucosae. Undifferentiated cells close to the bottom of the
crypts regenerate the epithelium (epithelial cell
turnover time is less than one week). Other epithelial cells in the
crypts correspond largely to those in the epithelium of the intestinal villi.
One exception are Paneth cells which are located
at the bottom of the crypts. They release a number of anti-bacterial substances,
among them lysozyme, and are thought to be involved in the control of infections.
One function of the crypts of Lieberkühn is the secretion of "intestinal juice" (about 2 liter/day), which in its composition closely resembles extracellular fluid and which is rapidly reabsorbed. The only enzymes which can be demonstrated in the intestinal juice are enteropeptidase (or enterokinase), which activates the pancreatic enzyme trypsin, and small amounts of amylase. In addition to enterocytes, the epithelium is composed of mucus-secreting goblet cells and endocrine cells.
The lamina propria is, similar to the lamina propria of the stomach, unusually cell rich. Lymphocytes often invade the epithelium or form solitary lymphoid nodules in the lamina propria. Lymph nodules may form longitudinal aggregations of 30-50 nodules in the lamina propria of the ileum. These large arggregations are called Peyer's patches.
The muscularis mucosae has two layers and extends into the intestinal villi, where the smooth muscle cells form a longitudinal bundle in the centre of the villi.
The Submucosa
The submucosa contains glands only in the duodenum. Submucosal glands of the duodenum are also called Brunner's glands. Their secretion is mucous and slightly alkaline due to bicarbonate ions (pH 7-8). The amount of bicarbonate is however too low to neutralize the acidic contents of the duodenal lumen. Instead, the secretion of Brunner's glands protects the duodenal mucosa - similar to the mucus which protects the gastric mucosa.
In addition to gastrin- and somatostatin-producing cells, we also find endocrine cells secreting cholecystokinin and secretin. Cholecystokinin stimulates the secretion of digestive enzymes in the pancreas and the contraction of the gall bladder. Secretin stimulates the pancreas to release "pancreatic juice", which is rich in bicarbonate ions. Secretin also amplifies the effects of cholecystokinin.
The large intestine constitutes the terminal part of the digestive system. It is divided into three main sections: cecum including the appendix, colon, and rectum with the anal canal. The primary function of the large intestine is the reabsorption of water and inorganic salts. The only secretion of any importance is mucus, which acts as a lubricant during the transport of the intestinal contents.
The surface of the mucosa is relatively smooth as there are no plicae circulares or intestinal villi. Crypts of Lieberkühn are present and usually longer and straighter than those of the small intestine. Goblet cells account for more of the epithelial cells than in the small intestine.
There is only little lamina propria squeezed between the glands. The muscularis mucosae again forms two layers.
Considerable amounts of fat may be found in the submucosa.
The appearance of the muscularis externa is different from that of the small intestine. The inner circular layer of muscle forms the usual sheath around the large intestine, but the outer longitudinal muscle layer forms three flattened strands, the taenia coli. Only a thin layer of longitudinal muscle surrounds the inner circular muscle layer between the taenia coli .
The adventitia forms small pouches (appendices epiploicae) filled with fatty tissue along the large intestine.
Specialized Sections of the Large Intestine
- The vermiform appendix
-
is a small blind-ending diverticulum from the cecum. The most important of its features is the thickening of its walls primarily due to large accumulations of lymphoid nodules in the lamina propria. Intestinal villi are usually absent, and crypts are comparatively rare. There is often fatty tissue in the submucosa. The muscularis externa is thinner than in the remainder of the large intestine.
- The anal canal
-
is the terminal part of the digestive tract (2.5-4 cm long). The mucosa has a characteristic surface relief of 5-10 longitudinal folds, the anal columns, each of which contains a terminal branch of the superior rectal artery and vein.
Small folds between the anal columns (anal valves) form the pectinate line. This line defines sections of the anal canal with different arterial and nerve supplies, different venous and lymphatic drainages and different embryological origins. Crypts disappear below the pectinate line and the epithelium changes from the tall, columnar type seen in other parts of the large intestine to a stratified squamous epithelium. The muscularis externa gradually becomes thicker and forms the involuntary internal anal sphincter.
Junctions between the major parts of the alimentary canal share a rapid transition from tissues characteristic of one part to those characteristic for the next part, e.g. the transition from the stratified squamous epithelium of the oesophagus to the glandular epithelium of the stomach. Many junctions are in additon accompanied by morphological specializations, e.g. the pylorus at the gastro-duodenal junctions or the ileo-caecal valve at the ileo-caecal junction. You will have a chance to examine these specializations in the DR.
Compared with these rapid changes, "junctions" between parts of the small intestine are very gradual morphological transitions.