parasympathetic)
Sympathetic and Parasympathetic divisions typically function in opposition
to each other. The gastrointestinal tract is regulated, in part, by the
autonomic nervous system.
This highly sophisticated regulatory system requires coordination of
motor, secretory, digestive and absorptive function.
Parasympathetic supply:
Parasympathetic nerves that supply the oesophagus, stomach, small
intestine, ascending colon, liver, gall bladder and pancrease arise from the
medulla and conducted through the vagus nerve.
Parasympathetic nerves to the rest of the colon arise in sacral spinal
cord and conducted through the pelvic nerves.
In general stimulatory
Sympathetic Supply:
Arise in the spinal cord
Postganglionic fibres from the prevertebral ganglia innervate the GI
tract
In general inhibitory
The sympathetic, parasympathetic, and ENS divisions of the autonomic
nervous system (ANS) innervate the digestive tract. Neural control of the
gut is hierarchical, with five basic levels of integrative organization Level 1
is the ENS, which behaves like an independent integrative nervous system
(minibrain) inside the walls of the gut. Level 2 consists of the prevertebral
ganglia of the sympathetic nervous system. Levels 3, 4, and 5 are within
the CNS. Sympathetic and parasympathetic signals to the digestive tract
originate at levels 3 and 4 (central sympathetic and parasympathetic
centers) in the medulla oblongata and represent the final common
pathways for the outflow of information from the brain to the gut. Level 5
includes higher brain centers that provide input for integrative functions
at levels 3 and 4.
Autonomic signals to the gut are carried from the brain and spinal cord by
sympathetic and parasympathetic nervous pathways, which represent the
extrinsic component of innervation. Neurons of the ENS form the local
intramural control networks that make up the intrinsic component of the
autonomic innervation. The parasympathetic and sympathetic
subdivisions are identified by the positions of the ganglia that contain the
cell bodies of the postganglionic neurons and by the point of outflow from
the CNS. Comprehensive autonomic innervation of the digestive tract
consists of interactive interconnections between the brain, the spinal cord,
and the ENS.
Water Absorption
In human adults, the average daily intake of water is about 2 L. secretions
from the salivary glands, pancreas, liver, and GI tract make up most of the
fluid entering the GI tract (about 7 L). Despite this large volume of fluid,
only 100 mL is lost in the feces. Therefore, the GI tract is extremely
entry of Cl into the enterocytes. The Cl then leaves the cell through
facilitated transport. This mode of Na+ uptake is called Na+/H+
Cl/HCO3 counter transport.
In the colon, the mechanisms for Na+ absorption are mostly similar to
those described for the ileum. There is no sugar-coupled or amino acidcoupled Na+ transport because most sugars and amino acids have
already been absorbed.
Sodium is also absorbed here via Na+-selective ion channels in the apical
cell membrane (electrogenic Na+ absorption).
Potassium
The average daily intake of K+ is about 4 g. Absorption takes place
throughout the intestine by passive diffusion through the tight junctions
and lateral intercellular spaces of the enterocytes. The driving force for K+
absorption is the difference between luminal and blood K+ concentration.
The absorption of water results in an increase in luminal K+ concentration,
resulting in K+ absorption by the intestine. In the colon, K+ can be
absorbed or secreted depending on the luminal K+ concentration. With
diarrhea, considerable K+ can be lost. Prolonged diarrhea can be lifethreatening, because the dramatic fall in extracellular K+ concentration
can cause complications such as cardiac arrhythmias.
Chloride
Most of the Cl ions added to the GI tract from the diet and from the
various secretions of the GI system are absorbed.
Intestinal chloride absorption involves both passive and active
processes. In the jejunum, active Na+ absorption generates a potential
difference across the small intestinal mucosa, with the serosal side more
positive than the lumen.
Chloride ions follow this potential difference and enter the bloodstream via
the tight junctions and lateral intercellular spaces. In the ileum and colon,
enterocytes actively take up Cl via Cl/HCO3 exchange, as discussed
above. The presence of other halides inhibits this absorption of Cl.
Bicarbonate
Bicarbonate ions are absorbed in the jejunum together with Na+. The
absorption of HCO3 by the jejunum stimulates the absorption of
Na+ and H2O
Through an Na+/H+ exchanger, H+ is secreted into the intestinal lumen,
where H+ and HCO3 react to form H2CO3, which then dissociates to
form CO2 and H2O. The CO2 diff uses into the enterocytes, where it reacts
with H2O to form H2CO3 (catalyzed by carbonic anhydrase). H2CO3
dissociates into
HCO3 and H+, and the HCO3 then diffuses into the blood.
In the ileum and colon, HCO3 is actively secreted into the lumen in
exchange for Cl. This secretion of HCO3 is important in buffering the
decrease in pH resulting from the short-chain fatty acids produced by
bacteria in the distal ileum and colon.
Calcium
The amount of Ca2+ entering the GI tract is about 1 g/day, approximately
half of which is derived from the diet. Most dietary Ca2+ is derived from
meat and dairy products. Of the Ca2+ presented to the GI tract, about
40% is absorbed. Several factors affect Ca2+ absorption. For instance, the
presence of fatty acid can retard Ca2+ absorption by the formation of
Ca2+ soap. In contrast, bile salt molecules form complexes with Ca2+
ions, which facilitate Ca2+ absorption.
Calcium absorption takes place predominantly in the duodenum and
jejunum, is mainly active, and involves three steps:
(1) Enterocytes take up calcium by passive diffusion through a Ca2+
channel, because there is a large Ca2+ concentration gradient; the
luminal Ca2+ is about 5 to 10 mM,whereas free intracellular Ca2+ is
about 100 nM.
(2) Once inside the cell, Ca2+ is complexed with Ca2+-binding protein,
calbindin D (CaBP).
(3) At the basolateral membrane, Ca2+ is extruded from the enterocytes
via the Ca2+-ATPase pump. Calcium uptake by enterocytes, the level of
CaBP in the cells, and transport by Ca2+-ATPase pumps are increased by
1,25-dihydroxyvitamin D3. Once inside the cell, the Ca2+ ions are
sequestered in the ER and Golgi membranes by binding to the CaBP in
these organelles.
Calcium absorption by the small intestine is regulated by the circulating
plasma Ca2+ concentration. Lowering of the Ca2+ concentration
stimulates the release of parathyroid hormone, which stimulates the
conversion of vitamin D to its active metabolite, 1,25-dihydroxyvitamin
D3, in the kidney.
This, in turn, stimulates the synthesis of CaBP and the Ca2+-ATPase by
the enterocytes (Fig. 26.26). Because protein synthesis is involved in the
stimulation of Ca2+ uptake by parathyroid hormone, a lapse of a few
hours usually occurs between the release of parathyroid hormone and the
increase in Ca2+ absorption by the enterocytes.
Magnesium
Humans ingest about 0.4 to 0.5 g/d of Mg2+. The absorption of Mg2+
seems to take place along the entire small intestine, and the mechanism
involved seems to be passive.
Vitamin Absorption
An organic compound is called a vitamin because it cannot be synthesized
by the body and must be obtained from diet
For absorption, vitamins are classified in two ways:
1. Water soluble
2. Fat-soluble
13 vitamins are universally recognized
9 water soluble and 4 fat soluble
to lipid peroxidation, which may explain why the red cells are more fragile
in people with vitamin E defi ciency than in healthy people.
Vitamin K can be derived from green vegetables in the diet or the gut
flora. The vitamin K derived from green vegetables is in the form of
phylloquinones. Vitamin K derived from bacteria in the small intestine is
in menaquinones.
The small intestine takes up phylloquinones via an energy-dependent
process from the proximal small intestine.
In contrast, menaquinones are absorbed from the small intestine
passively, depending only on the micellar solubilization of these
compounds by bile salts. Vitamin K is incorporated into chylomicrons. It is
rapidly taken up by the liver and secreted together with VLDLs. No carrier
protein for vitamin K has been identified.
Vitamin K is essential for the synthesis of various clotting factors by the
liver. Vitamin K deficiency is associated with bleeding disorders.