It is suggested that the reader review the basic principles of transport of substances through cell membranes dis cussed in Chapter 4. The following paragraphs present specialized applications of these transport processes during gastrointestinal absorption.
ANATOMICAL BASIS OF ABSORPTION
The total quantity of fluid that must be absorbed each day by the intestines is equal to the ingested fluid (about 1.5 liters) plus that secreted in the various gastrointestinal secretions (about 7 liters), which comes to a total of 8 to 9 liters. All but about 1.5 liters of this fluid is absorbed in the small intestine, leaving only 1.5 liters to pass through the ileocecal valve into the colon each day.
The stomach is a poor absorptive area of the gastrointestinal tract because it lacks the typical villus type of absorptive membrane, and also because the junctions between the epithelial cells are tight junctions. Only a few highly lipid-soluble substances, such as alcohol and some drugs like aspirin, can be absorbed in small quantities.
Folds of Kerckring, Villi, and Microvilli Increase the Mucosal Absorptive Area by Nearly 1000Fold. Figure 1 demonstrates the absorptive surface of the small intestinal mucosa, showing many folds called valvulae conniventes (or folds of Kerckring), which increase the surface area of the absorptive mucosa about threefold. These folds extend circularly most of the way around the intestine and are especially well developed in the duodenum and jejunum, where they often protrude up to 8 millimeters into the lumen.
Fig1. Longitudinal section of the small intestine, showing the valvulae conniventes covered by villi.
Also located on the epithelial surface of the small intestine all the way down to the ileocecal valve are millions of small villi. These villi project about 1 millimeter from the surface of the mucosa, as shown on the surfaces of the valvulae conniventes in Figure 1 and in individual detail in Figure 2. The villi lie so close to one another in the upper small intestine that they touch in most areas, but their distribution is less profuse in the distal small intestine. The presence of villi on the mucosal surface enhances the total absorptive area another 10-fold.
Fig2. Functional organization of the villus. A, Longitudinal section. B, Cross section showing a basement membrane beneath the epithelial cells and a brush border at the other ends of these cells.
Finally, each intestinal epithelial cell on each villus is characterized by a brush border, consisting of as many as 1000 microvilli that are 1 micrometer in length and 0.1 micrometer in diameter and protrude into the intestinal chyme. These microvilli are shown in the electron micro graph in Figure 3. This brush border increases the surface area exposed to the intestinal materials at least another 20-fold.
Fig3. Brush border of a gastrointestinal epithelial cell, also showing absorbed pinocytic vesicles, mitochondria, and endoplasmic reticulum lying immediately beneath the brush border. (Courtesy Dr. William Lockwood.)
Thus, the combination of the folds of Kerckring, the villi, and the microvilli increases the total absorptive area of the mucosa perhaps 1000-fold, making a tremendous total area of 250 or more square meters for the entire small intestine—about the surface area of a tennis court.
Figure 2A shows in longitudinal section the general organization of the villus, emphasizing (1) the advantageous arrangement of the vascular system for absorption of fluid and dissolved material into the portal blood and (2) the arrangement of the “central lacteal” lymph vessel for absorption into the lymph. Figure 2B shows a cross section of the villus, and Figure 3 shows many small pinocytic vesicles, which are pinched-off portions of infolded enterocyte membrane forming vesicles of absorbed fluids that have been entrapped. Small amounts of substances are absorbed by this physical process of pinocytosis.
Extending from the epithelial cell body into each microvillus of the brush border are multiple actin filaments that contract rhythmically to cause continual movement of the microvilli, keeping them constantly exposed to new quantities of intestinal fluid.
