Absorption of Water and ions in alimentary canal
Absorption of Water by Osmosis
Isosmotic
Absorption
Water is transported through the intestinal membrane
entirely by diffusion. Furthermore, this diffusion obeys the usual laws of
osmosis. Therefore, when the chyme is dilute enough, water is absorbed through
the intestinal mucosa into the blood of the villi almost entirely by osmosis.
Conversely, water can also be transported in the
opposite direction from plasma into the chyme. This occurs especially when
hyperosmotic solutions are discharged from the stomach into the duodenum.
Within minutes, sufficient water usually will be transferred by osmosis to make
the chyme isosmotic with the plasma.
Absorption
of Ions
Sodium is actively transported through the
intestinal membrane. Twenty to 30 grams of sodium are secreted in the
intestinal secretions each day. In addition, the average person eats 5 to 8
grams of sodium each day. Therefore, to prevent net loss of sodium into the
feces, the intestines must absorb 25 to 35 grams of sodium each day, which is
equal to about one seventh of all the sodium present in the body. Whenever
significant amounts of intestinal secretions are lost to the exterior, as in
extreme diarrhea, the sodium reserves of the body can sometimes be depleted to
lethal levels within hours.
Normally, however, less than 0.5 percent of the
intestinal sodium is lost in the feces each day because it is rapidly absorbed
through the intestinal mucosa. Sodium also plays an important role in helping
to absorb sugars and amino acids, as subsequent discussions reveal.
The principle of this mechanism is also essentially
the same as for absorption of sodium from the gallbladder and renal tubules.
The motive power for sodium absorption is provided by active transport of
sodium from inside the epithelial cells through the basal and lateral walls of
these cells into paracellular spaces.
This active transport obeys the usual laws of active
transport: It requires energy, and the energy process is catalyzed by
appropriate adenosine triphosphatase (ATP) enzymes in the cell membrane. Part
of the sodium is absorbed along with chloride ions; in fact, the negatively
charged chloride ions are mainly passively dragged by the positive electrical
charges of the sodium ions.
Active transport of sodium through the basolateral
membranes of the cell reduces the sodium concentration inside the cell to a low
value (≈50 mEq/L). Because the sodium concentration in the chyme is normally
about 142 mEq/L (i.e., about equal to that in plasma), sodium moves down this
steep electrochemical gradient from the chyme through the brush border of the
epithelial cell into the epithelial cell cytoplasm. Sodium is also
co-transported through the brush border membrane by several specific carrier
proteins, including
·
Sodium-glucose co-transporters,
·
Sodium amino acid co-transporters,
·
Sodium-hydrogen exchanger.
These transporters function similarly as in the
renal tubules, and provide still more sodium ions to be transported by the
epithelial cells into the paracellular spaces. At the same time they also
provide secondary active absorption of glucose and amino acids, powered by the
active Na+-K+ ATPase pump on the basolateral membrane.
Osmosis
of the Water
The next step in the transport process is osmosis of
water by Transcellular and Para cellular pathways. This occurs because a large
osmotic gradient has been created by the elevated concentration of ions in the
para cellular space. Much of this osmosis occurs through the tight junctions
between the apical borders of the epithelial cells (para cellular pathway) but
much also occurs through the cells themselves (Transcellular pathway).
Osmotic movement of water creates flow of fluid into
and through the para cellular spaces and, finally, into the circulating blood
of the villus.
When a person becomes dehydrated, large amounts of
aldosterone almost always are secreted by the cortices of the adrenal glands.
Within 1 to 3 hours this aldosterone causes increased activation of the enzyme
and transport mechanisms for all aspects of sodium absorption by the intestinal
epithelium. And the increased sodium absorption in turn causes secondary
increases in absorption of chloride ions, water, and some other substances.
This effect of Aldosterone is especially important
in the colon because it allows virtually no loss of sodium chloride in the
feces and also little water loss. Thus, the function of aldosterone in the
intestinal tract is the same as that achieved by aldosterone in the renal
tubules, which also serves to conserve sodium chloride and water in the body
when a person becomes dehydrated.
Absorption
of Chloride Ions in the Small Intestine
In the upper part of the small intestine, chloride
ion absorption is rapid and occurs mainly by diffusion (i.e., absorption of
sodium ions through the epithelium creates electro negativity in the chyme and
electro positivity in the para cellular spaces between the epithelial cells).
Then chloride ions move along this electrical gradient to follow the sodium
ions.
Chloride is also absorbed across the brush border
membrane of parts of the ileum and large intestine by a brush border membrane
chloride bicarbonate exchanger; chloride exits the cell on the basolateral
membrane through chloride channels.
Absorption
of Bicarbonate Ions in the Duodenum and Jejunum
Often large quantities of bicarbonate ions must be
reabsorbed from the upper small intestine because large amounts of bicarbonate
ions have been secreted into the duodenum in both pancreatic secretion and
bile. The bicarbonate ion is absorbed in an indirect way as follows:
When sodium ions are absorbed, moderate amounts of
hydrogen ions are secreted into the lumen of the gut in exchange for some of
the sodium. These hydrogen ions in turn combine with the bicarbonate ions to
form carbonic acid (H2CO3), which then dissociates to
form water and carbon dioxide.
The water remains as part of the chyme in the
intestines, but the carbon dioxide is readily absorbed into the blood and
subsequently expired through the lungs. Thus, this is so-called active absorption
of bicarbonate ions.
Secretion
of Bicarbonate Ions in the Ileum and Large Intestine
Simultaneous
Absorption of Chloride Ions
The epithelial cells on the surfaces of the villi in
the ileum, as well as on all surfaces of the large intestine, have a special
capability of secreting bicarbonate ions in exchange for absorption of chloride
ions. This is important because it provides alkaline bicarbonate ions that
neutralize acid products formed by bacteria in the large intestine.
Deep in the spaces between the intestinal epithelial
folds are immature epithelial cells that continually Na divide to form new
epithelial cells. These in turn spread outward over the luminal surfaces of the
intestines. While still in the deep folds, the epithelial cells secrete sodium
chloride and water into the intestinal lumen. This secretion in turn is
reabsorbed by the older epithelial cells outside the folds, thus providing flow
of water for absorbing intestinal digestates.
The toxins of cholera and of some other types of
diarrheal bacteria can stimulate the epithelial fold secretion so greatly that
this secretion often becomes much greater than can be reabsorbed, thus
sometimes causing loss of 5 to 10 liters of water and sodium chloride as
diarrhea each day. Extreme diarrheal secretion is initiated by entry of a
subunit of cholera toxin into the epithelial cells.
This stimulates formation of excess cyclic adenosine
monophosphate, which opens tremendous numbers of chloride channels, allowing
chloride ions to flow rapidly from inside the cell into the intestinal crypts.
In turn, this is believed to activate a sodium pump
that pumps sodium ions into the crypts to go along with the chloride ions.
Finally, all this extra sodium chloride causes extreme osmosis of water from
the blood, thus providing rapid flow of fluid along with the salt. All this
excess fluid washes away most of the bacteria and is of value in combating the
disease, but too much of it can be lethal because of serious dehydration of the
whole body that might ensue.
No comments:
Post a Comment