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~ Structural overview:
~ The wall of the alimentary canal consists of 4 layers: (from outside inwards)
[Figure: The layers of the alimentary canal.]
~ Innervation of G.I.T. –
- Except in esophagus and proximal stomach, the smooth muscle of G.I. tract
shows spontaneous waves of electrical activity: the slow waves.
- Frequency of slow waves determines the basic electrical rhythm (BER).
- BER is generated by pacemaker cells: the interstitial cells of Cajal.
- Slow waves or BER does not cause muscle contraction. Spike potentials
superimposed on depolarizing portions of slow waves cause contraction.
- Amount of tension developed by smooth muscle: proportional to number
of spikes
- Rate of BER: stomach ~ 4/min., duodenum ~ 12/min., terminal ileum ~
8/min. In the colon: cecum ~ 8/min., sigmoid colon ~ 16/min. {Note that: In
the small intestine – Rate of BER decreases from upper to lower segments.
This would cause the peristalsis to propel food in the aboral direction. In
the colon – There is reverse gradient for BER; it favors mixing of contents
and extraction of water and electrolytes.}
The motor impulses for this stage are transmitted via the IX
and X cranial nerves. The reflex activity is co-ordinated at
the brain-stem swallowing center.
esophagus
fundus
body
pylorus
duodenum
Anatomical regions of the stomach are fundus, body, antrum, and pylorus.
Two functional divisions of the stomach: proximal reservoir (fundus and
proximal 1/3rd of the body) and distal antral pump (distal 2/3rd of the body,
antrum, pylorus). The proximal stomach can maintain “tonic contraction”;
distal stomach shows “phasic contractions” and propulsive activity. Reason
for these phasic contractions:- A pacemaker complex located near the
midregion of the body; it initiates the electrical and phasic contractile
activity.
Motor functions of stomach:
(i) Storage:
Large quantities of food can be stored in the stomach until the food
can be passed on and processed in the duodenum.
Receptive relaxation: Without a rise in intragastric pressure, stomach
can accommodate about 1.5 L of volume; due to receptive relaxation.
It is initiated by the act of swallowing food; it occurs in the proximal
body region. Distension of stomach by food initiates a “vago-vagal
reflex” that causes this relaxation.
(ii) Mixing:
Weak peristaltic constrictions, called ‘mixing waves’, begin in the
midportion of the stomach and move toward the antrum. As the
constrictor waves progress toward the antrum, they become more
intense.
~ From the pyloric sphincter, chyme enters small intestine. The 3 parts of the
small intestine are – duodenum, jejunum, and ileum. Ileum empties its contents
into the large intestine (colon); it begins with cecum and ascending colon,
transverse colon, descending colon. Then there is sigmoid colon that leads to
rectum, and finally the anal canal.
By the time chyme enters the colon, the digestion and absorption of the
nutrients in the foodstuffs has already occurred (in the small intestine). The
principal functions of the colon are: (i) absorption of water and electrolytes
from the chyme to form solid feces, and (ii) storage of fecal matter until it
can be expelled.
Colon is roughly divided into two segments – 1. Proximal colon: It includes
the cecum, ascending colon, and proximal half of transverse colon. 2. Distal
colon: Distal half of the transverse colon, descending colon, and sigmoid
colon. Sigmoid colon eventually leads to rectum and anal canal.
Proximal colon is the absorptive colon. Absorption of water and
electrolytes from the chyme converts it into a solid fecal matter. Distal
colon is the storage colon. Fecal material is stored temporarily until it can
be excreted. Because intense colon movements are not required for these
functions, the movements of the colon are normally very sluggish.
(I) Mixing movements: Large circular constrictions occur in the large
intestine; about 2.5 cm of the circular muscle contracts. The
longitudinal muscle of the colon is aggregated into 3 longitudinal
strips, called the teniae coli. There are combined contractions of the
circular and longitudinal strips of muscle. This causes the
unstimulated portion of the colon to bulge outward into baglike sacs
called “haustrations”.
(II) Propulsive movements – “Mass movements”: They are the
modified type of peristalsis. From the beginning of transverse colon
to the sigmoid, the “mass movements” take over the propulsive role.
They occur 1 to 3 times per day. They cause colonic contents to move
distally for long distances.
~ Defecation: {Excretion of the feces out from the rectum through the anal canal
to the exterior.}
When a mass movement forces feces into the rectum, the desire for
defecation is normally initiated. Once the rectum is filled to about 25% of
its capacity, there is an urge to defecate.
The two anal sphincters are – (i) the internal anal sphincter, and (ii) the
external anal sphincter. The external sphincter is composed of striated
voluntary muscle, and it is controlled by nerve fibers in the pudendal nerve.
It exerts voluntary, conscious control over the process of defecation. As
long as the external sphincter is constricted, defecation if not possible. And,
when defecation is desired, the external sphincter will be relaxed, under
voluntary control, so as to allow passage of fecal material to the exterior.
Man being the social animal, time and place for defecation is chosen
appropriately. Until then, the process will be voluntarily withheld.
Defecation reflex: When feces enter the rectum, distension of the rectal
wall initiates afferent signals that spread through the myenteric plexus to
initiate peristaltic waves in the descending colon, sigmoid, and rectum.
Fecal material forced toward the anus. Internal anal sphincter is relaxed as
the peristaltic wave approaches downward. If the external anal sphincter is
also consciously, voluntarily relaxed at the same time, defecation occurs.
The pelvic parasympathetic nerves carry nerve fibers that send impulses
for the contractions of the sigmoid and rectum, for the process of
defecation.
The effects associated with defecation, which force the fecal contents
downward, are – closure of the glottis, and contraction of the abdominal
wall muscles.
SECRETIONS OF THE G.I.TRACT
Blood vessel
Primary secretion
Blood vessel
[Figure: The primary salivary secretion is rich in Na + & water. As it flows through the salivary
duct, “ductular modification” causes Na+ to be reabsorbed and K+ to be secreted. However,
Na+ reabsorption is excess over K+ secretion; and the duct is relatively impermeable to water.
Hence, the final secretion is hypotonic. HCO3- is present in the primary secretion as well.
Increased rate of secretion will therefore increase the pH of saliva correspondingly.]
Saliva is secreted from the blood into the acinus of the salivary gland.
Primary salivary secretion is isotonic to plasma; it is rich in Na +. However, as
it flows through the ducts, it gets modified. Na+ and Cl- are reabsorbed, and
K+ and HCO3- are added by the ducts. The final saliva reaching mouth is
hypotonic since Na+ was removed from it but water is not reabsorbed
(ducts are impermeable to water). It is rich in K+ and HCO3-, and has low
Na+ and Cl-.
If the salivary flow is rapid, there is less time for ductular modification.
Then, it will be rich in Na+ and Cl-; and K+ and HCO3- will be low.
Enzymes in saliva:
(i) Ptyalin (an -amylase) – for carbohydrate digestion; inactivated in
stomach (acidic pH);
(ii) Lingual lipase – can digest up to 30% of dietary triglycerides; active
in stomach.
Saliva also contains:
- lysozyme (proteolytic enzyme that attacks bacteria),
- thiocyanate ions (which can kill bacteria in the presence of
lysozyme),
- mucin (lubricating material; makes the food slippery)
- antibodies (immunoglobulins; they can destroy oral bacteria)
- blood group antigens (in some people, called “secretors”, blood
group antigens are secreted in the saliva)
- kallikrein (an enzyme that acts on the globulins of local interstitial
fluid to produce ‘bradykinin’ which is a powerful vasodilator)
Functions of saliva:
1. Digestive function –
(a) Carbohydrate digestion:
The -amylase (ptyalin) present in the saliva initiates starch digestion
in the mouth. Big polysaccharide molecules are broken down into
maltose and triose.
[Note: As the food mixed with saliva is swallowed and enters
stomach, the starch digestion stops. Stomach pH is acidic (around 2
to 3); at this pH the amylase enzyme cannot remain active.]
(b) Fat digestion:
The lingual lipase present in saliva initiates digestion of triglycerides
(TGs) in the ingested food. [Lingual lipase continues to act in the
stomach pH.]
2. Lubrication –
Mucin in the saliva helps in the lubrication of the ingested foodstuffs.
This facilitates the swallowing of food.
3. Helps in speech –
Saliva lubricates oral cavity. This facilitates articulation of organs of
mouth (tongue, lips, and palate) involved in speech. Speaking becomes
easier.
4. Appreciation of taste –
Saliva acts as a solvent for many substances. When such substances
dissolve in mouth and come in contact with the ‘taste buds’ (of tongue),
it produces the sensation of taste.
5. Defense mechanisms/oral hygiene –
- Flow of saliva helps to wash out bacteria of mouth.
- Lysozyme, thiocyanate ions, and IgA antibodies (immunoglobulins)
present in saliva can attack and kill bacteria that cause dental caries.
6. Regulation of water content/osmolarity of the body fluids – (an
indirect function)
When body water content is decreased (or plasma becomes hypertonic),
saliva dries up. Water loss via saliva is minimized. In addition to the
stimulation of the thirst center, it is an indication to drink fluids; the
water content will normalize.
7. Temperature regulation – (not important in humans)
(In some animals) a mechanism of “panting” helps reduce body
temperature when it is increased. Heavy breathing by a dog causes
evaporation of saliva from the tongue, reducing the temperature in the
oral region. Then, heat from the other parts of the body is carried via
blood to the oral region and lost from there, thus reducing body
temperature.
Regulation of salivary secretion:
[All other digestive tract secretions are regulated by neural and hormonal
mechanisms. Salivary secretion is regulated only by neural mechanism.]
o Salivary secretion is regulated by parasympathetic and sympathetic
nervous systems.
o Saliva production is unique in that it is increased by both
parasympathetic and sympathetic activity. {Generally, these two
systems have opposite effects on the effector organs.} However,
parasympathetic regulation is more important.
o Parasympathetic stimulation (impulses carried by VII & IX cranial
nerves) – increases salivary secretion by increasing transport
processes in the acinar cells of the salivary glands and cells lining
their ducts. Acetyl choline is the transmitter released by the
parasympathetic nerves.
o Parasympathetic nerve signals arise from the superior and inferior
salivatory nuclei in the brain stem. They are stimulated by taste
stimuli from tongue, especially the sour taste. Smell and sight of food
also stimulate nerve signals.
o Parasympathetic nerve signals increase salivary secretion by another
(indirect) mechanism. These signals cause vasodilatation near the
salivary glands. Increased blood flow increases the salivary secretion.
[Partly, the vasodilator effect is due to kallikrein secreted by the
activated salivary cells. It acts as an enzyme that forms bradykinin in
blood. Bradykinin is a strong vasodilator.]
o Fear, dehydration, sleep inhibit parasympathetic signals, and thus
reduce salivary secretion.
o Sympathetic stimulation can also increase salivation, but only to a
lesser extent. Sympathetic nerves arise from the superior cervical
ganglion and reach the salivary glands.
Salivary reflexes:
Reflex salivary secretion occurs with two types of reflexes ~ (1)
unconditioned reflex, and (2) conditioned reflex.
It was demonstrated by the Russian physiologist Pavlov. Pavlov’s
experiment on dog was aimed to prove the effects of training or
“conditioning”.
Unconditioned reflexes are inborn reflexes. {When the dog was presented
with some food, there was reflex salivation. Or, if a sour or acidic food is
placed on the tongue of a person, there is reflex salivation. This type of
“unconditioned” reflex is present naturally since birth; no training
required.}
Conditioned reflexes are not inborn; they are acquired after training
(“conditioning”). With previous experience of sour or acidic taste, when the
person only sees the same food, there is reflex salivation. This occurs due
to the process of learning or training. {In the dog’s experiment – a bell is
rung first and then food is presented. This is repeated several times. The
dog “learns” that every time the bell is rung, food is served. Then, reflex
salivation occurs only on ringing of bell.}
~ Applied physiology –
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~ Gastric juice: (LAQ – Composition, functions, and regulation of secretion of the
gastric juice) (short note – mechanism of HCl secretion in stomach)
Gastric juice has acidic pH. Enzyme present in the gastric juice can be active at
acidic pH only.
The stomach consists of 2 types of glands – (1) gastric glands, and (2)
pyloric glands.
1. The gastric glands has the following cell types (with their secretions
mentioned:
a. Mucus secreting cells – mucus
b. Parietal/oxyntic cells – (i) HCl, (ii) intrinsic factor of Castle
c. Peptic/chief cells – pepsinogen
2. The pyloric gland has following cell types (and secretions):
a. Mucous cells – mucus
b. “G” cells – gastrin
c. Enterochromaffin-like (ECL) cells – histamine
3. Gastric juice also contains these enzymes – rennin, gastric lipase.
- HCl is secreted by the parietal cell located in the stomach wall. Parietal cell
shows a canaliculus where H+ ions are secreted actively.
Parietal cell
CO2
+ H2O CO2
Lumen
of
stomach H2CO3
Cl- H+
H+ HCO3- HCO3-
Cl- Cl-
Blood vessel
- CO2 from blood enters the parietal cell. There, it combines with water to
form H2CO3. {CO2 + H2O → H2CO3.} This reaction is catalyzed by the enzyme
carbonic anhydrase. H2CO3 then splits into H+ and HCO3-.
- H+ is then secreted into the parietal cell canaliculus by an active transporter
(ATPase) or pump. The H+-K+-ATPase is the active transporter(also called
“proton pump”) which actively secretes the H+ ions against their
concentration gradient.
- HCO3- is transported into blood and Cl- from blood is transported from
blood into the parietal cell by a transporter called chloride-bicarbonate
exchanger (Cl-/HCO3- exchanger).
- Chloride ions diffuse from the parietal cell into the lumen of the stomach,
combine with the H+ ions and form HCl.
[Also note: H+ is pumped against a concentration gradient that is about 1
million-fold: pH 7 in the parietal cell cytosol to about pH 1 in the lumen of
gastric gland.
As H+ is secreted in the lumen of stomach, HCO3- (alkali) is added to blood.
This bicarbonate will be excreted in urine (“alkaline tide”).]
~ Applied Physiology –
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chymotrypsinogen chymotrypsin
Once trypsin is formed, it acts on chymotrypsinogen to form
chymotrypsin. Trypsin also converts procarboxypeptidase into
carboxypeptidase. The action of proteolytic enzymes is inhibited by
“trypsin inhibitor” in the pancreatic juice.
~ Applied physiology:
1. Steatorrhea –
When there is deficiency of the pancreatic enzymes, fat digestion does not
occur. Unabsorbed fats then appear in feces; fecal fat content increases –
“steatorrhea”.
2. Autodigestion of pancreas and pancreatitis –
Pancreatic enzymes may get activated within the pancreas. They cause
autodigestion and damage to the pancreas. Resultant inflammation of the
gland is termed pancreatitis.
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Composition of bile:
- Daily secretion: about 1000 mL.
- Colour: yellowish green
- Reaction: alkaline; pH ~ 7-7.6
- Bile salts: sodium taurocholate & sodium glycocholate
- Bile pigments: bilirubin & biliverdin
- Cholesterol and phospholipids
- Inorganic salts: chlorides, carbonates, and phosphates of Na+,
K+, Ca++.
It is sac present underneath the liver. The capacity of the gall bladder is about 50
mL.
Bile synthesized by liver is flown to gall bladder where it is stored. When fats in
the diet reach duodenum, gall bladder contractions empty the bile into the
duodenum where it aids in digestion and absorption of fats.
1. Storage of bile:
Until it is required for digestion of fats, bile is stored in gall bladder.
2. Concentration of bile:
Gall bladder epithelium absorbs water from the bile. Bile is concentrated up
to 5 times (compared to the bile that was secreted by liver.)
sucrase
Sucrose glucose + fructose
lactase
Lactose glucose + galactose
maltase
Maltose glucose + glucose
(ii) Peptidases –
Once pancreatic peptidases have digested the proteins and
polypeptides in diet, further action of the peptidases in succus
entericus will digest them so as to form tri- and dipeptides and single
amino acids. These can then be absorbed from small intestine.
(iii) Intestinal lipase –
It causes further digestion of fats (after the action of pancreatic
lipases); free fatty acids and glycerol will be formed. These fat
digestion products will then be absorbed.
Regulation of intestinal secretion:
- Local stimulus: Chyme in the intestine increases the secretion
of small intestine.
- Hormonal regulation: Secretin & CCK increase small intestinal
secretion.
{The proximal colon includes ascending colon and first part of transverse colon.
Distal colon includes later part of transverse colon, descending colon, and sigmoid
colon. The proximal colon is also called “absorptive colon”. It has a capacity to
absorb large amount of water (and electrolytes). When un-digested and un-
absorbed fraction of food reaches colon, most of water & electrolytes are
absorbed from the proximal colon. The remaining portion will be the fecal
material to be excreted.
Distal colon is also called “storage colon”. It stores the fecal material temporarily,
until defecation process can be initiated. With the defecation process, fecal
matter will be moved through the sigmoid colon, rectum, and finally the anal
canal.}
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