Located over the entire surface of the small intestine are small pits called crypts of Lieberkühn, one of which is illustrated in Figure 1. These crypts lie between the intestinal villi. The surfaces of both the crypts and the villi are covered by an epithelium composed of two types of cells: (1) a moderate number of goblet cells, which secrete mucus that lubricates and protects the intestinal surfaces, and (2) a large number of enterocytes, which, in the crypts, secrete large quantities of water and electrolytes and, over the surfaces of adjacent villi, reabsorb the water and electrolytes along with the end products of digestion.

Fig1. A crypt of Lieberkühn, found in all parts of the small
intestine between the villi, which secretes almost pure extracellular
fluid.
The intestinal secretions are formed by the enterocytes of the crypts at a rate of about 1800 ml/day. These secretions are almost pure extracellular fluid and have a slightly alkaline pH in the range of 7.5 to 8.0. The secretions are also rapidly reabsorbed by the villi. This flow of fluid from the crypts into the villi supplies a watery vehicle for absorption of substances from chyme when it comes in contact with the villi. Thus, the primary function of the small intestine is to absorb nutrients and their digestive products into the blood.
Mechanism of Secretion of the Watery Fluid. The exact mechanism that controls the marked secretion of watery fluid by the crypts of Lieberkühn is still unclear, but it is believed to involve at least two active secretory processes: (1) active secretion of chloride ions into the crypts and (2) active secretion of bicarbonate ions. The secretion of both ions causes electrical drag of positively charged sodium ions through the membrane and into the secreted fluid as well. Finally, all these ions together cause osmotic movement of water.
Digestive Enzymes in the Small Intestinal Secretion. When secretions of the small intestine are collected without cellular debris, they have almost no enzymes. The enterocytes of the mucosa, especially those that cover the villi, contain digestive enzymes that digest specific food substances while they are being absorbed through the epithelium. These enzymes are the following: (1) several peptidases for splitting small peptides into amino acids; (2) four enzymes—sucrase, maltase, isomaltase, and lactase—for splitting disaccharides into monosaccharides; and (3) small amounts of intestinal lipase for split ting neutral fats into glycerol and fatty acids.
The epithelial cells deep in the crypts of Lieberkühn continually undergo mitosis, and new cells migrate along the basement membrane upward out of the crypts toward the tips of the villi, thus continually replacing the villus epithelium and also forming new digestive enzymes. As the villus cells age, they are finally shed into the intestinal secretions. The life cycle of an intestinal epithelial cell is about 5 days. This rapid growth of new cells also allows rapid repair of excoriations that occur in the mucosa.