Digestion & Absorption

of Macronutrients
By; Abdur Rehman

PhD Scholar

Biochemistry
 Digestion
The process by which large and complex dietary molecules
converted to small absorbable forms in the GIT

Digestion

Carbohydrate
Proteins Lipids
s

Monosaccharides Amino Acids Glycerol

Fatty Acids

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Cholesterol
 GIT Major function in digestion &
Mouth:
Production of
Absorption
Saliva, alpha
amylase:
Partial Stomach:
Digestion of Gastric juice
Polysaccharides
Small
with HCL &
Proteases; Intestine:
partial Final Digestion Large
& Absorption
digestion of
Pancreas: Intestine
protein Absorption of
NaHCO3 &
enzyme release electrolytes,
to intestine bacterial
Liver/Gall utilization non
Bladder: digested food
Synthesis/
Storage of bile
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 GIT Major function in digestion &
Mouth: Absorption
Production of
Saliva, alpha
amylase:
Partial
Stomach:
Digestion of
Gastric juice
Small
Polysaccharides
with HCL & Intestine:
Proteases; Final Digestion Large
partial digestion & Absorption
of protein Pancreas: Intestine
NaHCO3 & Absorption of
enzyme release electrolytes,
bacterial
Liver/Gall to intestine
utilization non
Bladder: digested food
Synthesis/
Storage of bile
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 Digestion of carbohydrates

• Carbohydrates occurs briefly in mouth and largely in the

intestine
­ Glycosidases hydrolyze glycosidic bond, specific to the bond,
structure and configuration of monosaccharide units

• Digestion in the mouth: Carbohydrates,the only nutrients for

which the digestion begins in the mouth to a significant extent

• During the process of mastication, salivary -amylase (ptyalin)

acts on starch randomly and
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Abdur Rehman
KMU
Lecturer - 1,4-glycosidic bonds5
• The products formed include -limit dextrins, (containing about 8
glucose units with one or more -1,6-glycosidic bonds) maltotriose
and maltose

• Carbohydrates not digested in the stomach: The salivary amylase is

inactivated by low stomach pH and degradation of starch is stopped

• Digestion in the small intestine: The acidic dietary contents of the

stomach, on reaching small intestine, are neutralized by bicarbonate
produced by pancreas
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•The pancreatic -amylase acts on starch and continues the
digestion process
•Amylase specifically acts on -1,4-glycosidic bonds and not on
-1,6-bonds and release disaccharides (maltose, isomaltose)
and oligosaccharides

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 Digestion of Carbohydrates
• The final digestion of di- and oligo-saccharides to
monosaccharides primarily occurs at the mucosal lining of the
upper jejunum

• Oligosaccharidases (e.g. glucoamylase acting on amylose)

and disaccharidases(e.g. maltase, sucrase, lactase)
­ Sucrase hydrolyze a large quantity of table sugar (sucrose)
­ Lactase (β-galactosidase) is the rate-limiting, and, consequently, the
utilization of milk sugar (lactose) is limited in humans
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 Absorption of Monosaccharides
• Glucose & galactose absorbed by a sodium-dependent
process in the small intestine

• Carried by the same transport protein (SGLT1) and compete for

intestinal absorption
­ Other monosaccharides are absorbed by carrier-mediated diffusion

• Fructose and sugar alcohols are only absorbed down their

concentration gradient because of no active transport,
­ Fructose may remain in the intestinal lumen, acting as a substrate for
bacterial fermentation
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Transport of
glucose, fructose,
and galactose
• The SGLT1 transporter is coupled to
the Na+-K+ pump, allowing glucose
and galactose to be transported
against their concentration
gradients
• The GLUT5 Na+-independent
facilitative transporter allows
fructose as well as glucose and
galactose to be transported with their
concentration gradients
• Exit from the cell for all the sugars is
via the GLUT2 facilitative
transporter

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 Glucose transporters
Plasma membrane is impermeable to glucose
Cellular uptake is mediated by two distinct processes via specific
glucose transporter proteins that belong to the family of solute carriers
(SLC);
the SLC2 family members, GLUTs (glucose transporters), are sodium
independent facilitators of the glucose transport,
whereas the SLC5 family members, SGLTs (sodium and glucose
transporters) mediate the secondary-active sodium-glucose cotransport

Until now, 14 GLUTs and 12 SGLTs isoforms have been identified in

humans of which 5 GLUTs and none SGLTs were detected in the
mammalian blood cells
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 Insulin receptors and glucose
transporters
• To initiate its effects on target tissues, insulin binds to and activates a membrane
receptor.

• It is the activated receptor that is responsible for the cellular effects of insulin.
• Within seconds after insulin binds to its membrane receptor, the membranes of
about 80% of body tissues increase their uptake of glucose by means of
special glucose transporters (especially skeletal muscles and adipose tissue).

• GLUT-4 is the insulin-dependent glucose transporter for skeletal muscle and

adipose tissue.

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 Non-digestible carbohydrates

• The plant foods are rich in fibrous material which cannot be

digested either by the human enzymes or intestinal bacteria

• The fibers are chemically complex carbohydrates which include

cellulose, hemicellulose, pectins, lignin and gums

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 Abnormalities of carbohydrate
• In
digestion
general, humans possess an efficient system of carbohydrate
digestion and absorption

• Since only monosaccharides are absorbed, any defect in the

activities of disaccharidases results in the passage of
undigested disaccharides into the large intestine
­ The disaccharides draw water from intestinal mucosa by osmosis and
cause diarrhea

• Further, bacterial action on undigested carbohydrates leads to

flatulence
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 Lactose intolerance
• Due to a defect in the enzyme lactase (β-galactosidase)
• Lactose intolerance may be primary (congenital) or secondary
(acquired)

• Acquired lactose intolerance may occur due to a sudden and high

intake of milk-based diets

• For lactose intolerant people, consumption of curd is beneficial,

since lactobacilli present in curd contain the enzyme lactase

• Further, yeast rich in lactase, can also be used for treatment of

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 The problem of flatulence
• Flatulence, characterized by increased intestinal motility, cramps
and irritation
­ The carbohydrates (di-, oligo-, and polysaccharides) not hydrolyzed by -
amylase and other intestinal enzymes cannot be absorbed
­ Lactose is not hydrolyzed in some individuals due to the deficiency of
lactase

• The di-, and oligosaccharides can be degraded by the bacteria

present in ileum to liberate monosaccharides

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• As the monosaccharides are utilized by the intestinal bacteria,
gases such as hydrogen, methane and carbon dioxide—
besides, lactate and short chain fatty acids—are released
which cause flatulence

• The occurrence of flatulence after the ingestion of leguminous

seeds (Bengal gram, red gram, beans, peas, soya bean) is
very common

• Raffinose containing galactose, glucose and fructose is a

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 Protein Digestion & Absorption
• The intake of dietary protein is in the range of 50-100 g/day
• About 30-100 g/day of endogenous protein is derived form the
digestive enzymes and worn out cells of the digestive tract

• The digestion & absorption of proteins is very efficient in healthy

humans,
­ very little protein (about 5-10 g/day) is lost through feces

• Proteins degraded hydrolases—which specifically cleave the

peptide bonds, hence known as peptidases
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Digestion of
Protein:
• Endopeptidases attack the internal peptide
Proteolytic
enzymes bonds
responsible for • e.g. pepsin, trypsin
the digestion • Exopeptidases act on the peptide bonds of
of proteins and terminal amino acids
are produced • carboxypeptidases (act on C-terminal
by the amino acid)
stomach, the • aminopeptidases (act on N-terminal amino
pancreas and acid)
the small
intestine

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• Proteins are not digested in the mouth due to the absence of
proteases in saliva

• Gastric juice contains hydrochloric acid and a proenzyme

pepsinogen

• Hydrochloric acid: HCl is secreted by parietal (oxyntic) cells of

gastric glands

• HCL in the stomach performs two important functions-

denaturation
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of proteins and
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killing of certain
23
 Proteases as proenzymes
• Pepsin: Pepsin (Greek: pepsis—digestion) produced by the serous cells of
the stomach as pepsinogen, the inactive zymogen or proenzyme

• Pepsinogen is converted to active pepsin either by autocatalysis, brought

about by other pepsin molecules or by gastric HCl (pH < 2).

• Removal of a fragment of polypeptide chain (44 amino acids in case of pig

enzyme) makes the inactive enzyme active after attaining a proper
conformation.

• Pepsin is an acid-stable endopeptidase optimally active at a very low pH

(2.0)
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 Renin

• Rennin: also called chymosin, found in the stomach of infants &

children
­ Involved in the curdling of milk

• converts milk protein casein to calcium para caseinate which can

be effectively digested by pepsin
­ Rennin is absent in adults.

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Digestion of
proteins by
pancreatic
proteases

•The proteases of pancreatic juice

are secreted as zymogens
(proenzymes) and then converted
to active forms
•These processes are initiated by
the release of two polypeptide
hormones, cholecystokinin and
secretin from the intestine

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 Release and activation of zymogens
• The key enzyme for activation of zymogen is enteropeptidase (formerly
enterokinase) produced by intestinal (mostly duodenal) mucosal cells

• Enteropeptidase cleaves off a hexapeptide from the N-terminal end of

trypsinogen to produce trypsin, the active enzyme

• Trypsin, in turn, activates other trypsinogen molecules (autocatalysis).

• Further, trypsin is the common activator of all other pancreatic
zymogens to produce the active proteases, chymotrypsin, elastase
and carboxypeptidases (A and B)

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 Specificity and action of pancreatic
proteases
• Trypsin, chymotrypsin and elastase are endopeptidases active at
neutral pH

• Gastric HCl is neutralized by pancreatic NaHCO3 in the intestine and

this creates favorable pH for the action of proteases

• The substrate specificity of pancreatic proteases is depicted in Fig.8.7.

• For instance, trypsin cleaves the peptide bonds, the carbonyl ( CO )
group of which is contributed by arginine or lysine
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 Action of carboxypeptidases

• The pancreatic carboxypeptidases (A and B) are

metalloenzymes that are dependent on Zn2+ for their catalytic
activity, hence they are sometimes called Zn-proteases

• They also possess certain degree of substrate specificity in their

action

• For example, carboxypeptidase B acts on peptide bonds of

COOH-terminal amino acid, the amino group of which is
contributed by arginine or lysineKMU(Fig.8.7)
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Digestion of proteins by small intestinal enzymes

• The luminal surface of intestinal epithelial cells contains

aminopeptidases and dipeptidases

• Aminopeptidase is a non-specific exopeptidase which repeatedly

cleaves N-terminal amino acids one by one to produce free amino
acids and smaller peptides

• The dipeptidases act on different dipeptides to liberate amino

acids (Fig.8.8)
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Absorption of Free Amino Acids & Small Peptides

• The end product of the action of endopeptidases and exopeptidases is a

mixture of free amino acids, di- and tripeptides, and oligopeptides, all
of which are absorbed

• Free amino acids are absorbed across the intestinal mucosa by

sodium-dependent active transport

• There are several different amino acid transporters, with specificity for
the nature of the amino acid side chain (large or small; neutral, acidic,
or basic)
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 Amino acid transporters
• Amino acids are transported by specific active transporters showing
mechanisms which are similar to ones active in glucose transport

• These Na dependant symporters are located at the brush- border membrane

• At least six specific symporter systems have been identified as follows:
1. Neutral amino acid symporter for amino acids with short or polar side- chains
(Ser, Thr, Ala)

2. Neutral amino acid symporter for aromatic or hydrophobic side chains (Phe,
Tyr, Tryp, Met, Val, Leu, Ileu)

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3. Imino acid symporter (Pro, OH-Pro)
4. Basic amino acid symporter (Lys, Arg, Cys)
5. Acidic amino acid symporter ( Asp, Glu)
6. β - amino acid symporter ( β- Ala, Tau)

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Absorption of Free Amino Acids & Small Peptides

• Dipeptides and tripeptides enter the brush border of the

intestinal mucosal cells, where they are hydrolyzed to free
amino acids, which are then transported into the hepatic
portal vein

• Many such peptides are large enough to stimulate antibody

formation—this is the basis of allergic reactions to foods

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Abnormalities of protein digestion and amino acid
absorption

• Any defect in the pancreatic secretion impairs protein and fat

digestion

• This causes the loss of undigested protein in the feces along with
the abnormal appearance of lipids

• Deficiency of pancreatic secretion may be due to pancreatitis,

cystic fibrosis or surgical removal of pancreas

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 Hartnup’s disease (neutral amino
aciduria)
• Hartnup's is the name of the family in whom this disease was first
discovered

• It is characterized by the inability of intestinal and renal epithelial cells to

absorb neutral amino acids (tryptophan, alanine, serine, threonine, valine)

• Tryptophan absorption is most severely affected with a result that typical

symptoms of pellagra are observed in the patients of Hartnup’s disease

• This is related to the impairment in the conversion of tryptophan to NAD+

and NADP+, the coenzymes of niacin

• The treatment includes high protein diet and nicotinamide supplementation

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 Coeliac Disease

• Celiac disease is a serious autoimmune disorder that can occur in

genetically predisposed people where the ingestion of gluten
leads to damage in the small intestine

• Characterized by severe malabsorption and specific

diagnostic features exhibited by the intestinal mucosa

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 Coeliac Disease

• The deficit is located within the mucosal cells, and permits

certain polypeptides; resulting from digestion of gluten; to exert
local effects within the intestine, and to be absorbed, inducing an
antibody response

• The intolerance is for gliadin portion of gluten (found in wheat and

several other grains)
• Diarrhoea, Abdominal bloating , Weight loss
• Gluten free diet
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 Lipids
• There is considerable variation in the daily consumption of lipids which
mostly depends on the economic status and dietary habits

• The intake of lipids is much less (often < 60 g/day) in poorer sections of the
society, particularly in the less developed countries

• In the developed countries, an adult ingests about 60-150 g of lipids per day
• Of this, more than 90% is fat (triacylglycerol)
• The rest of the dietary lipid is made up of phospholipids, cholesterol,
cholesteryl esters and free fatty acids

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 Lipids

• Lipids are insoluble or sparingly soluble in aqueous solution

• The digestive enzymes, however, are present in aqueous medium
• This poses certain problems for the digestion and absorption of
lipids

• Fortunately, the digestive tract possesses specialized machinery

to

1. Increase the surface area of lipids for digestion

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 Minor digestion of lipids in the stomach
• Stomach contains gastric lipase which can degrade fat containing short chain
fatty acids at neutral pH

• The digestion of lipids in the stomach of an adult is almost negligible, since

lipids are not emulsified and made ready for lipase action

• Further, the low pH in the stomach is unfavorable for the action of gastric lipase
• In case of infants, the milk fat (with short chain fatty acids) can be hydrolyzed
by gastric lipase to some extent

• Lingual lipase secreted by Ebner's gland in the tongue

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Minor digestion of lipids in the stomach
• In humans, lingual lipase makes little or no contribution to the
preduodenal lipase activity

• The pancreas secretes a 50 kDa lipase and a 10 kDa colipase.

• The pancreatic colipase binds to the triglycerides surface with
the help of the bile acids and anchors pancreatic lipase

• In the absence of colipase, bile acids remove pancreatic lipase

from the lipid surface and thereby inhibit lipolysis

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 Emulsification of lipids in the small
intestine
• Emulsification occurs by three complementary mechanisms
1. Detergent action of bile salts
2. Surfactant action of degraded lipids
3. Mechanical mixing due to peristalsis

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 Detergent action of bile salts
• Bile salts are the biological detergents synthesized from cholesterol in
the liver

• They are secreted with bile into the duodenum

• Bile salts possess steroid nucleus, the side chain of which is attached
to either glycine (glycocholic acid) or taurine (taurocholic acid)

• Bile salts are the most effective biological emulsifying agents

• They interact with lipid particles and the aqueous duodenal contents
and convert them into smaller particles (emulsified droplets)
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 Surfactant action of degraded lipids
• The initial digestive products of lipids- free fatty acids, and
monoacylglycerols promote emulsification

• These compounds along with phospholipids are known as

surfactants

• Surfactants get absorbed to the water-lipid interfaces and increase

the interfacial area of lipid droplets

• Thus, the initial action of the enzyme lipase helps in further

digestion of lipids
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 Degradation of triacylglycerols (fat)
• Pancreatic lipase is the major enzyme that digests dietary fats
• This enzyme preferentially cleaves fatty acids (particularly long chain, above 10
carbons) at position 1 and 3 of triacylglycerols

• The products are 2-monoacylglycerol (formerly 2-monoglyceride) and free fatty acids
(Fig.8.9)

• The activity of pancreatic lipase is inhibited by bile acids which are present along with
the enzyme in the small intestine

• This problem is overcome by a small protein, colipase

• It is also secreted by pancreas as pro-colipase and converted to active form by trypsin
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Colipase binds at the lipid-aqueousAbdur
interface and helps to anchor and stabilize lipase49
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 Degradation of triacylglycerols (fat)

• Lipid esterase is a less specific enzyme present in pancreatic

juice

• It acts on monoacylglycerols, cholesteryl esters etc. to liberate

free fatty acids

• The presence of bile acids is essential for the activity of lipid

esterase.

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Degradation of cholesteryl esters

•Pancreatic cholesterol esterase

(cholesteryl ester hydrolase) cleaves
cholesteryl esters to produce
cholesterol and free fatty acids
(Fig.8.10)

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 Degradation of phospholipids

• Phospholipases are responsible for the hydrolysis of

phospholipids

• Pancreatic juice is rich in phospholipase A2 which cleaves the

fatty acid at the 2nd position of phospholipids

• The products are a free fatty acid and a lysophospholipid.

• Phospholipase A2 is secreted as a zymogen which is activated in
the intestine by the action of trypsin
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 Absorption of lipids
• The primary products obtained from the lipid digestion are 2-
monoacylglycerol, free fatty acids and free cholesterol

• Bile salts form mixed micelles with lipids

• The micelles have a disk like shape with lipids (monoacylglycerol, fatty
acids, cholesterol and phospholipids) at the interior and bile salts at the
periphery

• The hydrophilic groups of the lipids are oriented to the outside (close to the
aqueous environment) and the hydrophobic groups to the inside

• In this fashion, the bile salt micelles exert a solubilizing effect on the lipids
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 Mechanism of lipid absorption
• The mixed micelles serve as the major vehicles for the transport of lipids
from the intestinal lumen to the membrane of the intestinal mucosal cells,
the site of lipid absorption

• The lipid components pass through the unstirred fluid layer and are
absorbed through the plasma membrane by diffusion (Fig.8.12).

• Absorption is almost complete for monoacylglycerols and free fatty acids

which are slightly water soluble

• However, for water insoluble lipids, the absorption is incomplete

• For instance, less than 40% of the dietary cholesterol is absorbed.
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 Mechanism of lipid absorption
• The micelle formation is also essential for the absorption of fat-soluble vitamins,
particularly vitamins A and K.

• The efficiency of lipid absorption is dependent on the quantity of bile salts to

solubilize digested lipids in the mixed micelles

• It may, however, be noted that in the absence of bile salts, the lipid absorption
occurs to a minor extent

• This is mostly due to the slightly water-soluble nature of monoacylglycerols and

free fatty acids.

• Further, short and medium chain fatty acids are not dependent on micelle
formation for the absorption
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Synthesis of lipids in the intestinal mucosal cells

• The fatty acids of short and medium chain length (< 10 carbons), after their
absorption into the intestinal cells, do not undergo any modification

• They enter the portal circulation, bound to albumin and are transported to the
liver

• The long chain fatty acids are activated by thiokinase (fatty acyl CoA
synthetase) in the intestinal cells

• The acyl CoA derivatives so formed combine with 2-monoacylglycerols

to produce triacylglycerols
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Synthesis of lipids in the intestinal mucosal cells

• These reactions are catalyzed by a group of enzymes, namely

acyltransferases

• Further, within the intestinal cells, cholesterol is converted to

cholesteryl ester, and phospholipids are regenerated from the
absorbed lysophospholipids

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 Secretion of lipids from the intestinal mucosal
cells
• The lipids that are resynthesized (described above) in the intestinal cells are hydrophobic
in nature

• They are put together as lipid droplets and surrounded by a thin layer of apolipoproteins
(AI and B-48) and phospholipids.

• This package of lipids enveloped in the layer stabilizes the droplets and increases their
solubility

• These particles are known as chylomicrons.

• Chylomicrons migrate to the plasma membrane of intestinal mucosal cells
• They are released into the lymphatic vessels by exocytosis
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 Secretion of lipids from the intestinal mucosal
cells
• The presence of chylomicrons (Greek: chylos–juice) gives the lymph a milky appearance,
which is observed after a lipid-rich meal.

• Chylomicrons enter the large body veins via the thoracic duct
• Blood from here flows to the heart and then to the peripheral tissues (muscle, adipose
tissue) and, finally, to the liver

• Adipose tissue and muscle take up a large proportion of dietary lipids from chylomicrons
for storage and transport

• It is believed that this bypass arrangement (passage of chylomicrons through peripheral

tissues) protects the liver from a lipid overload after a meal.

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 Abnormalities of lipid digestion and absorption
• The gastrointestinal tract possesses an efficient system for digestion and
absorption of lipids

• It can comfortably handle as much as 4 times the normal daily intake of lipids
• Steatorrhea: It is a condition characterized by the loss of lipids in the feces.
Steatorrhea may be due to

1. A defect in the secretion of bile or pancreatic juice into the intestine;

2. Impairment in the lipid absorption by the intestinal cells.
3. Steatorrhea is commonly seen in disorders associated with pancreas, biliary
obstruction, severe liver dysfunction etc
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 Cholesterol stones

• Cholesterol stone formation in gall-bladder (gall stones) is a

frequent health complication

• It is found more frequently in females than in males often in

association with obesity

• Cholesterol gall stones are formed when liver secretes bile

(containing phospholipids, bile acids etc.), supersaturated with
respect to cholesterol
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 Absorption of vitamins
• The fat soluble vitamins (A, D, E, and K) are absorbed with fats.
• Their absorption is reduced in biliary tract obstruction. Most vitamins are absorbed in
the upper small intestine.

• The water-soluble vitamins –thiamin, riboflavin, niacin, pyridoxine, pantothenate, biotin

and ascorbic acid – are absorbed in the small intestine by carriers that are sodium
co-transporters.

• Absorption of Vitamin B 12 and folate is sodium independent.

• The water-soluble vitamin B 12 is absorbed in the ileum.
• This vitamin binds to intrinsic factor before the complex is transported across the ileal
mucosa
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 Absorption of calcium

• Calcium is absorbed from the intestine with the help of the

steroid hormone vitamin D (1, 25 dihydroxy cholecalciferol)

• When the levels of ionized calcium in blood is high, the levels of

1, 25 dihydroxy cholecalciferol levels fall and vice-versa

• Calcium absorption is also decreased by substances, which form

insoluble salts with calcium like phytates and oxalates

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 Absorption of iron
• Most of the iron in the diet is in the ferric form whereas it is absorbed in the
ferrous form.

• The gastric secretions dissolve the iron and it is reduced by ascorbic acid
and other reducing substances in the stomach.

• Also, the iron transporters in the brush border of the enterocytes have ferric
reductase activity associated with them.

• Almost all iron absorption occurs in duodenum.

• Intestinal absorption of iron is regulated by state of iron stores in the body
and state of erythropoietin in the bone marrow.
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 Absorption of water and electrolytes
• The gastrointestinal tract absorbs as well as secretes electrolytes and
water throughout the intestine.

• There are two routes of transport. One is the paracellular route and the
other is the transcellular route.

• In the paracellular route the flow is through the tight junctions between
cells because of osmotic, hydrostatic or electrical gradients.

• In the transcellular route the water and electrolytes cross apical and
basolateral membranes by energy requiring specific active transport carriers

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