Absorption of Nutrients
Absorption
of Carbohydrates
Essentially all the carbohydrates in the food are absorbed in the
form of monosaccharides; only a small fraction are absorbed as disaccharides
and almost none as larger carbohydrate compounds. By far the most abundant of
the absorbed monosaccharides is glucose,
usually accounting for more than 80 per cent of carbohydrate calories absorbed.
The reason for this is that glucose is the final digestion product of our most
abundant carbohydrate food, the starches. The remaining 20 per cent of absorbed
monosaccharides are composed almost entirely of galactose and fruc-tose,
the galactose derived from milk and the fructoseas one of the monosaccharides
digested from cane sugar.
Virtually all the monosaccharides are absorbed by an active
transport process. Let us first discuss the absorption of glucose.
Glucose Is Transported by a Sodium Co-Transport
Mechanism.
In the absence of sodium transport through the intes-tinal
membrane, virtually no glucose can be absorbed. The reason is that glucose
absorption occurs in a co-transport mode with active transport of sodium.
There are two stages in the transport of sodium through the intestinal membrane. First is active trans-port
of sodium ions through the basolateral mem-branes of the intestinal epithelial
cells into the blood, thereby depleting sodium inside the epithelial cells.
Second, decrease of sodium inside the cells causes sodium from the intestinal
lumen to move through the brush border of the epithelial cells to the cell
interiors by a process of facilitated
diffusion. That is, a sodium ion combines with a transport protein, but the trans-port protein will not transport
the sodium to the inte-rior of the cell until the protein itself also combines
with some other appropriate substance such as glucose. Fortunately, intestinal
glucose also combines simultaneously with the same transport protein, and then
both the sodium ion and glucose molecule are transported together to the
interior of the cell. Thus, the low concentration of sodium inside the cell
liter-ally “drags” sodium to the interior of the cell and along with it the
glucose at the same time. Once inside the epithelial cell, other transport
proteins and enzymes cause facilitated diffusion of the glucose through the
cell’s basolateral membrane into the paracellular space and from there into the
blood.
To summarize, it is the initial active transport of sodium through
the basolateral membranes of the intestinal epithelial cells that provides the
eventual motive force for moving glucose also through the membranes.
Absorption
of Other Monosaccharides. Galactose is trans-ported by almost exactly the same mechanism as
glucose. Conversely, fructose transport does not occur by the sodium
co-transport mechanism. Instead, fruc-tose is transported by facilitated
diffusion all the way through the intestinal epithelium but not coupled with
sodium transport.
Much of the fructose, on entering the cell, becomes phosphorylated,
then converted to glucose, and finally transported in the form of glucose the
rest of the way into the blood. Because fructose is not co-transported with
sodium, its overall rate of transport is only about one half that of glucose or
galactose.
Absorption
of Proteins
Most proteins, after digestion, are absorbed through the luminal
membranes of the intestinal epithelial cells in the form of dipeptides,
tripeptides, and a few free amino acids. The energy for most of this transport
is supplied by a sodium co-transport mechanism in the same way that sodium
co-transport of glucose occurs. That is, most peptide or amino acid molecules
bind in the cell’s microvillus membrane with a specific transport protein that
requires sodium binding before transport can occur. After binding, the sodium
ion then moves down its electrochemical gradient to the interior of the cell
and pulls the amino acid or peptide along with it. This is called co-transport (or secondary active transport) of
the amino acids and peptides. A few amino acids donot require this sodium
co-transport mechanism but instead are transported by special membrane trans-port
proteins in the same way that fructose is trans-ported, by facilitated
diffusion.
At least five types of transport proteins for trans-porting amino
acids and peptides have been found in the luminal membranes of intestinal
epithelial cells. This multiplicity of transport proteins is required because
of the diverse binding properties of different amino acids and peptides.
Absorption
of Fats
Earlier, it was pointed out that when fats are digested to form
monoglycerides and free fatty acids, both of these digestive end products first
become dissolved in the central lipid portions of bile micelles. Because the molecular dimensions of these micelles
are only 3 to 6 nanometers in diameter, and because of their highly charged
exterior, they are soluble in chyme. In this form, the monoglycerides and free
fatty acids are carried to the surfaces of the microvilli of the intestinal
cell brush border and then penetrate into the recesses among the moving,
agitating microvilli. Here, both the monoglycerides and fatty acids diffuse
imme-diately out of the micelles and into the interior of the epithelial cells,
which is possible because the lipids are also soluble in the epithelial cell
membrane. This leaves the bile micelles still in the chyme, where they function
again and again to help absorb still more monoglycerides and fatty acids.
Thus, the micelles perform a “ferrying” function that is highly
important for fat absorption. In the presence of an abundance of bile micelles,
about 97 per cent of the fat is absorbed; in the absence of the bile micelles,
only 40 to 50 per cent can be absorbed.
After entering the epithelial cell, the fatty acids and
monoglycerides are taken up by the cell’s smooth endoplasmic reticulum; here,
they are mainly used to form new triglycerides that are subsequently released
in the form of chylomicrons through
the base of the epithelial cell, to flow upward through the thoracic lymph duct
and empty into the circulating blood.
Direct
Absorption of Fatty Acids into the Portal Blood. Smallquantities of short- and
medium-chain fatty acids, such as those from butterfat, are absorbed directly
into the portal blood rather than being converted into triglyc-erides and
absorbed by way of the lymphatics. The cause of this difference between short-
and long-chain fatty acid absorption is that the short-chain fatty acids are
more water-soluble and mostly are not recon-verted into triglycerides by the
endoplasmic reticulum. This allows direct diffusion of these short-chain fatty
acids from the intestinal epithelial cells directly into the capillary blood of
the intestinal villi.
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