Small Intestine Most digestion and absorption occurs in small intestine Small intestine is about 10 feet long. Duodenum (about 5 feet) Jejunum (about 2.5 feet) Ileum (about 2.5 feet)

Small Intestinal Motility Segmentation contractions (unique to small intestine: Mostly mixing of contents and bringing contents into contact with mucosal surface. Minor propulsion because frequency is greater in proximal end and is reduced as we go further down the small intestine.

Peristalsis in Small Intestine Peristalsis is major propulsion mechanism in small intestine. Peristaltic waves in small intestine typically travel only a few inches, then end. But when there’s an irritant in the small intestine, a single peristaltic wave can traverse the entire 10 foot length in a few seconds. This tends to rapidly force the irritant out of the small intestine. This is known as a peristaltic rush.

Surface Specializations in Small Intestine

Muscularis Mucosae in Small Intestine Muscularis mucosae is just under the mucosa. When it contracts, villus shortens slightly. This mixes the content that is in contact with the villus, important for absorption and digestion. It also forces fluid from the villus interstitial space into the lymphatics.

Ileocecal Sphincter and Valve Sphincter at distal end of ileum is normally closed, preventing reflux from colon. When peristaltic waves reach it, it relaxes and allows flow from ileum to colon. Ileocecal valve, normally closed, opens passively. It’s a one-way valve, much like those in veins and in the heart.

Cells in small intestinal mucosa secrete 1 to 2 liters/day of water into lumen osmotically, hydrolyzed food being the solutes that create the osmotic gradient. How much fluid enters the gut? 1. Everything we drink (about 1 liter/day) 2. Saliva (1 to 2 liters/day) 3. Gastric secretions (2 to 3 liters/day) 4. Pancreatic and hepatic secretions (2 to 3 liters/day) 5. Secretions of small intestine (1 to 2 liters/day) Nearly none is absorbed in mouth, esophagus or stomach, so the volume entering the small intestine is 7 to 11 liters/day (let’s use 10 liters/day as an approximation) About 500 ml/day enters the large intestine. Therefore, small intestine must absorb about 95% of the water that enters it.

Absorption in Small Intestine For all practical purposes, all digested nutrients are absorbed in the small intestine. Except for bile salts and vitamin B12, absorption of everything except electrolytes is complete before the gut contents reach the ileum.

Surface Specializations in Small Intestine These greatly increase (500 to 1,000 fold) the surface area of mucosa in small intestine.

Enzymes of Brush Border Membranes Proteases digest proteins mostly to the level of dipeptides. Dipeptidases in brush border convert them to amino acids. Pancreatic amylase converts starch to disaccharides (chains of two sugars). Disaccharidases in brush border membranes convert them to simple sugars (monosaccharides). Brush border membranes contain active transport systems that absorb amino acids and peptides. Note that they are absorbed as they are formed because the enzymes that generate them and the active transport systems that absorb them are on the same membranes. This maintains very high concentration gradients across the membranes, which osmotically absorb water. Sodium is also actively absorbed through the brush border membranes, which drives most of the osmotic absorption of water. Active transport of sugars and amino acids depend on simultaneous sodium transport.

Disaccharidases Starch gets hydrolyzed down to the disaccharide level (maltose) before being hydrolyzed all the way to glucose. We also ingest some glucose as well as some disaccharides: Maltose (= glucose + glucose) Sucrose (= glucose + fructose) Lactose (= glucose + galactose) These are all hydrolyzed by brush border enzymes (maltase, sucrase and lactase), and the products are absorbed. Fructose is absorbed by simple diffusion. Some people have a lactase deficiency (used to be called milk allergy), and can’t digest lactase. A dairy meal results in undigested lactose in the gut, which osmotically causes diarrhea.

Dipeptidases There are at least three categories of brush border dipeptidases: one for neutral amino acids, one for acidic amino acids, one for basic amino acids A very small percentage of peptides up to four amino acids long are actively absorbed as well. Newborns absorb very large proteins (antibodies) from mother’s milk by endocytosis. Protease inhibitors in mother’s milk prevent digestion of antibodies.

Fat Digestion and Absorption Lipases only act on surfaces of fat droplets, making detergent action of bile salts very important. Triglycerides (most dietary fat) are hydrolyzed to free fatty acids, glycerol, and monoglycerides, all are absorbed through brush border membranes. Free fatty acids are also detergents, so they promote additional fat digestion along with the bile salts. Free fatty acids and monoglycerides aren’t very water soluble, and saturate the solution in the lumen of the small intestine. With bile salts, the undissolved fatty acids and monoglycerides aggregate to form micelles (about 20 lipid molecules per micelle), which become a reservoir for free fatty acids and monoglycerides – as they are absorbed, more leave the micelles and dissolve. Micelles also store vitamin D and other fat soluble compounds. Fat absorption is usually complete by the proximal part of the ileum. Bile salts are reabsorbed in the terminal ileum.

More on Absorption Most things absorbed by brush border cells diffuse through their basal borders into the capillaries. Fats are an exception. Fatty acids, glycerol and monoglycerides absorbed by brush border cells are resynthesized into triglycerides. A layer of lecithin (another kind of lipid) and protein is added within the cells. The resulting particles are chylomicrons. Chylomicrons are released into the interstitial fluid by exocytosis, but are too big to enter capillaries. Instead, they diffuse into the lymphatics via the central lacteal. Muscularis mucosae is important in forcing interstitial fluid into central lacteal.

Colon (= large intestine) Length = 4 feet. No enzymes or villi. About 500 ml enters every day, about 100 ml is lost in feces. Therefore, reabsorbs about 400 ml, about 80% of the volume that enters it. MUCH less than small intestine (9,500 ml and 95%, respectively). Some vitamins and nutrients produced by colon flora are absorbed, but are of little importance. They also produce nitrogen, CO2 and other gases from stuff that we can’t digest. Beans contain lots of undigestible (for us) stuff that colon bacteria can’t get enough of. Some of the gases produced by colon flora are absorbed through colon wall, but an average of more than 1 liter/day isn’t (lots more for your uncle or brother). That’s flatus. Colon secretes some mucous fluid and fairly small amounts of an alkaline fluid that neutralizes acid produced by colon flora.

Absorption in Colon Mainly, active transport of sodium, passively followed by chloride, creating osmotic gradient that absorbs water (about 400 ml/day). This is almost entirely in proximal half of colon, called absorbing colon for that reason. Distal half, which stores fecal material for long periods, is called storage colon.

Motility in Colon Longitudinal muscle in colon is arranged in three bands (taeniae coli). Circular muscle generates something similar to segmentation contractions, referred to as haustral contraction. All of these are for mixing contents.

Motility in Colon (continued) Stimuli to Mass Movements Peristalsis is rare, if it occurs at all. Propulsion in colon is mostly by mass movements – rings of contraction that extend in length. Very effective because of the high resistance to backward flow. Stimuli to Mass Movements Local distension of colon Gastrocolic reflex – food entering stomach elicits reflex (via vagus nerve) that initiates mass movements. Useful in housebreaking a puppy

Defecation Storage colon stores contents for hours, until mass movement propels it into rectum. This stretches rectal wall, initiating defecation reflex, which includes vagal afferents from CNS and via myenteric plexus. Motor response includes mass movements in storage colon, contraction of rectal muscle, relaxation of internal and external anal sphincters. Uh-oh!

Modulation of Defecation Reflex Rectal muscles are contracting, anal sphincters are relaxing. This spells trouble. Good news! External anal sphincter is skeletal muscle, so we can voluntarily contract it. Better news! Contracting the external anal sphincter inhibits the defecation reflex, usually for several hours. Toilet training is essentially learning to contract the external anal sphincter. The neural connections making it possible don’t develop in humans until after the second birthday, doesn’t develop in puppies until the third month. Many animals (most herbivores) don’t have an external anal sphincter, and can’t be housebroken. Don’t try to housebreak a horse. It’s a waste of your time and, sooner or later, it will annoy the horse.

Forces in Defecation Much comes from muscles of colon and rectum. Augmented by “straining movements” – contracting abdominal muscles with glottis closed. This causes very high abdominal and thoracic pressures. The abdominal pressure contributes to forcing stool out of the colon. The straining can cause very high arterial pressure, but the elevated thoracic pressure can close off the veins, bringing venous return to zero and, consequently, reducing cardiac output to zero. Many strokes and heart attacks occur during straining movements.