UNIT – I

DIGESTION
Definition - Digestion refers to the breakdown of food into smaller components that can be absorbed into the
bloodstream.
Digestion or catabolism is divided into two types – the mechanical digestion of food that occurs in the mouth
when it is physically broken up into smaller pieces and the chemical digestion that occurs in the
gastrointestinal tract when the food is broken down into small molecules by digestive enzymes.
Intracellular and Extracellular Digestion
Heterotrophic organisms obtain energy through the ingestion of energy-rich food. The ingested food should
be digested into small compounds to absorb as nutrients.

There are two methods of digestion namely – intracellular and extracellular digestion
Intracellular Digestion - Intracellular digestion refers to a form of digestion where the breakdown of
materials into small components takes place inside the cell.

Place of Occurrence - Intracellular digestion occurs inside food vacuoles within the cell. The hydrolytic
enzymes stored in the lysosomes are responsible for the chemical digestion of the food particle.
Intracellular digestion can be categorized into two types as heterophagic digestion and autophagic digestion.

Heterophagic Digestion - The heterophagic digestion is the breaking down of molecules brought into the cell
by endocytosis. The degradation of ingested food during intracellular digestion occurs in a process known as
phagotrophy. The endocytic vesicle or the food vacuole is fused with a lysosome and the chemical digestion
occurs inside the food vacuole. The nutrients diffuse to the cytoplasm through the walls of the vesicle. The
indigestible materials are excreted through exocytosis.

Amoeba Phagocytosis

Autophagic Digestion - Autophagic digestion occurs inside the cell to digest internal molecules and
organelles. Autophagy maintains energy sources in the cell by recycling the damaged proteins, aggregates,

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and organelles in the cell. The end products of the degradation can be used as building blocks for the
replacement of the depleted cellular components. Thereby, autophagy promotes the survival of the cell during
stress by balancing the cellular energy levels. It allows the clearing of unwanted components fromthe cell
as well. Hence, autophagy is pro-survival and capable of undergoing cellular stress like nutrient deprivation.
But, autophagy lets the cell die by destroying active organelles in it like mitochondria.

Extracellular Digestion - Extracellular digestion refers to a form of digestion where the breakdown of
materials into smaller components takes place outside the cell. Thus, the hydrolytic enzymes are secreted on
the food materials via the cell membrane.

Place of occurrence - In animals, extracellular digestion occurs inside the lumen of the alimentary canal.
• Ingestion occurs through the mouth in extracellular digestion.
• Glands of the alimentary canal secrete digestive enzymes into the lumen.
• Nutrients are absorbed into the blood through the gut epithelia
• Indigestible materials are excreted through the anus.

The alimentary canal of animals is differentiated into different regions such as mouth, esophagus, stomach,
small intestine, large intestine, and anus. Different regions possess different functions during the digestion of
food. The initial regions are involved in the mechanical digestion of food while the lattermost regions are
involved in the chemical digestion as well as the absorption of nutrients. The salivary, gastric, pancreatic, and
intestinal glands secrete digestive enzymes to the lumen.

Conclusion - Intracellular and extracellular digestion are the two types of digestion. In protozoans, the
ingested food particles are digested inside a food vacuole by intracellular digestion. Besides in animals with
an alimentary canal, the digestion occurs within the lumen of the alimentary canal by extracellular digestion.
The main difference between intracellular and extracellular digestion is the location and complexity of each
type of digestion mechanisms.

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 CHEMICAL DIGESTION

• In chemical digestion, food is broken down by the action of enzymes

• Large food molecules (for example, proteins, lipids, nucleic acids, and starches) must be broken down
into subunits that are small enough to be absorbed by the lining of the alimentary canal. This is
accomplished by enzymes through hydrolysis.

Place of Occurrence
• Amylase catalyses the breakdown of starch into maltose in the mouth and small intestine.
• Proteases catalyse the breakdown of proteins into amino acids in the stomach and small intestine.
• Lipases catalyse the breakdown of fats and oils into fatty acids and glycerol in the small intestine.
• Maltase catalyses the breakdown of maltose into glucose in the small intestine.

The chemical process of digestion occurs in different regions of the alimentary canal. Based on this, it is
grouped into three types:
1. Buccal digestion
2. Gastric digestion
3. Intestinal digestion

1. Digestion in the Buccal Cavity - Buccal Digestion or Salivary Digestion

Digestion of food occurring inside the buccal cavity is called Buccal digestion. As the salivary enzymes are
involved, it is also called as salivary digestion.
The oral cavity or the mouth contains salivary gland that secretes a wide range of enzymes to aid the first
step of metabolism.
Salivary gland - The salivary glands contain 3 pairs of salivary glands.

• The parotid gland is present on each side of the face below and in front of the ears; they open
into the mouth through duct of stensen. They are serous type containing enzymes
• The submandibular glands are present in the angles of the lower jaw, they open into the mouth
though Whartons duct. They are mixed type secrete both serous and mucous.
• The sublingual glands are located below the tongue. They are purely mucous type

The salivary glands are racemose glands in which glandular cells are arranged in acini or alveoli, each of
which consists of a single layer of cells around a central cavity into which these cells discharge their
secretions. Ducts (intercalated ducts) arise from each alveoli unite to form large ducts which open into the
main duct, and are poured over the food in the buccal cavity.
The food (bolus) is forced into the pharynx by the tongue. As the food is swallowed, it moves into the
esophagus, which essentially provides a passageway from the pharynx to the stomach. The mucous glands
of the esophagus secrete mucus to aid in moistening and lubricating the bolus.

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Composition of Saliva

• In man, the saliva contains two types of secretions: 1. Serous secretion containing ptyalin (ii)
mucous secretion for lubricating the food.
• It is colorless, slightly cloudy due to the presence of cells and mucin.
• It is slightly acidic with a pH of 6 -7. It has inorganic salts like sodium chloride, potassium chloride,
sodium phosphate, calcium carbonate and calcium phosphate.
• It also has organic substances like ureas, amino acids, cholesterol and vitamins.
• It secretes enzyme lysozyme which has antibacterial action.

Mechanical and Chemical Digestion in the Mouth

Mechanical Digestion
Mechanical digestion in the mouth results from chewing, or mastication, in which food is manipulated by the tongue, ground
by the teeth, and mixed with saliva. As a result, the food is reduced to a soft, flexible, easily swallowed mass calleda bolus.

Chemical Digestion
Two enzymes, salivary amylase and lingual lipase, contribute to chemical digestion in the mouth.
Salivary Amylase
Salivary amylase, also known as ptyalin, which is secreted by the salivary glands, initiates the breakdown of
starch. Dietary carbohydrates are either monosaccharide and disaccharide sugars or complex polysaccharides
such as starches. Most of the carbohydrates we eat are starches, but only monosaccharides can be absorbed
into the bloodstream. Thus, ingested disaccharides and starches must be broken down into monosaccharides.
The function of salivary amylase is to begin starch digestion by breaking down starch into smaller molecules
such as the disaccharide maltose, the trisaccharide maltotriose, and short-chain glucose polymers called α-
dextrins. Even though food is usually swallowed too quickly for all starches to be broken down in the mouth,
salivary amylase in the swallowed food continues to act on the starches for about another hour, at which time
stomach acids inactivate it.
Ptyalin (pH 6.8)
Starch →Dextrins→ Maltotriose → Maltose
Lingual Lipase
Saliva also contains lingual lipase, which is secreted by lingual glands in the tongue. This enzyme becomes
activated in the acidic environment of the stomach and thus starts to work after food is swallowed. It breaks
down dietary triglycerides (fats and oils) into fatty acids and diglycerides. A diglyceride consists of a glycerol
molecule that is attached to two fatty acids

Stimulation and Control of Salivary Secretion: Reflex salivation is normally is induced by the food inthe
mouth which in turn stimulates the salivary center of brain. Both sympathetic (vasoconstriction) and
parasympathetic (vasodilation) supply though VII and IX cranial nerves innervate the salivary glands.

Pharynx and Esophagus

• When food is first swallowed, it passes from the mouth into the pharynx Swallowed food passes from
the mouth into the oropharynx and laryngopharynx; the muscular contractions of these areas help
propel food into the esophagus and then into the stomach. The esophagus secretes mucus and transports
food into the stomach. It does not produce digestive enzymes, and it does not carry on absorption.
• The movement of food from the mouth into the stomach is achieved by the act of swallowing, or
deglutition
• Swallowing occurs in three stages: (1) the voluntary stage, in which the bolus is passed into the
oropharynx; (2) the pharyngeal stage, the involuntary passage of the bolus through the pharynx into

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 the esophagus; and (3) the esophageal stage, the involuntary passage of the bolus through the
esophagus into the stomach

2 Digestion in Stomach – Gastric Digestion

After the bolus has been swallowed, it passes through the Oesophagus to reach the stomach. Peristaltic action
in the stomach consists of a churning movement which is brought about the contraction and relaxation of
the gastric muscle layers. This further causes mechanical and chemical breakdown of the food, mixing of
the food with gastric juices and then its onward movement into the tract.
Stomach
It is a J-shaped organ. It has four regions: the cardia, fundus, body and pyloric part. When the stomach is
empty, the mucosa lies in large folds or rugae. The pylorus communicates with the duodenum of the small
intestine through pyloric sphincter. The concave side of the stomach is called the lesser curvature and the
convex lateral border is called the greater curvature.

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Structure of Gastric Glands
Gastric gland is the basic secretory unit of the stomach. The wall of the stomach is lined with many exocrine
glands (gastric glands), that secrete gastric juice into the stomach lumen. The mucosa of stomach has gastric
glands. Several gastric glands open into the bottom of narrow channels called gastric pits. secretions from the
gastric glands flow into each gastric pit and into the lumen of the stomach. Gastric glands have three major
types of glands namely:
• Cardiac glands – it is columnar glandular cells found in the cardiac part of the stomach.
• Fundal glands – they are tubular columnar glandular cells
• Pyloric glands – these are convoluted, tubule-alveolar glands.

These glands consist of the following types of cells:

i. Mucous neck cells

These cells secrete a sticky mucus substance composed of glycoprotein, water, ions and other
things.
The mucus lubricates the lining of the stomach and also protects the lining of the stomach
from degradation due to the highly acidic environment.
ii. Peptic or Zymogenic or chief cells
These cells release the chief zymogen (proenzyme) of the stomach pepsinogen
Pepsinogen is transferred into an active enzyme called pepsin by a low pH 2. Pepsin cleaves
proteins into smaller polypeptides
These cells are found deeper within the gastric glands of the stomach
iii. Parietal or Oxyntic Cells
Which secrete HCl and intrinsic factor (factor essential for absorption of vitamin B12).
The parietal cells located throughout the gland are responsible for the production of
hydrochloric acid. HCl converts pepsinogen into its active form called pepsin. It denatures
proteins which allow cleaving of proteins.
iv. G Cells
▪ The G cells are mostly found in pyloric glands in the antrum of the pylorus; some are found
in the duodenum and other tissues. The G cells secretes gastrin. They stimulate the parietal
cells to release HCl.
v. Enterochromaffin like cells (ECL)
▪ These release histamine which stimulates parietal cells.

Composition of Gastric Secretions (Gastric juice)

• Approximately 1200 to 1500ml of gastric juice is released in a single day. It contains 97-99% water
and roughly 0.6% solids.
• It is a transparent yellow fluid
• It is highly acidic in nature (pH of 0.9-1.5)
• Other constituents of gastric secretions are HCl, gastric enzymes (pepsin, rennin, gastric lipase and
gastric amylase); mucous and inorganic salts (chlorides of sodium, potassium, and calcium and certain
phosphates and bicarbonates)

Mechanical and Chemical Digestion in Stomach

Mechanical Digestion
Several minutes after food enters the stomach, waves of peristalsis pass over the stomach every 15 to 25
seconds. Each peristaltic wave moves gastric contents from the body of the stomach down into the antrum, a
process known as propulsion. The pyloric sphincter normally remains almost, but not completely, closed.
Because most food particles in the stomach initially are too large to fit through the narrow pyloric sphincter,
they are forced back into the body of the stomach, a process referred to as retropulsion. Another round of
propulsion then occurs, moving the food particles back down into the antrum. If the food particles are still too
large to pass through the pyloric sphincter, retropulsion occurs again as the particles are squeezed back
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 into the body of the stomach. Then yet another round of propulsion occurs and the cycle continues to repeat.
The net result of these movements is that gastric contents are mixed with gastric juice, eventually becoming
reduced to a soupy liquid called chyme. Once the food particles in chyme are small enough, they can pass
through the pyloric sphincter, a phenomenon known as gastric emptying.

Chemical Digestion
1. Pepsin – The main digestive function of the stomach is the partial breakdown of proteins into proteoses
and peptones. Gastric pepsin is produced by the chief cells as an inactive zymogen called pepsinogen.
It is activated by gastric HCl to form active pepsin. Pepsin breaks the polypeptide into peptones,
proteoses and polypeptides, which are latter broken down to amino acids by the proteolyticenzymes
of the pancreas and intestine.

Pepsinogen HCl Pepsin

Proteins Pepsin Proteoses, Peptones and polypeptides

2. Rennin – It is present basically in the sucking mammals. It is also secreted as an inactive prorennin
and is converted into active rennin by HCl. These enzymes bring about the coagulation of milk. This
process is called as curdling of milk.
Activated rennin splits soluble milk protein caseinogen into insoluble casein, paracasein and then
calcium paracaseinate. Calcium paracaseinate form the curd and can be further digested by gastric
pepsin to form proteoses and peptones

Prorennin HCl Rennin

Rennin + Caesinogen Casein
Rennin + Casein Paracasein
Paracasein + Calcium Calcium Paracaseinate (curdles)
Calcium Paracaseinate Pepsin Protesoses, Peptones, Polypeptides

3. Gastric lipase – Gastric lipase splits triglycerides (fats and oils) in fat molecules (such as those found
in milk) into fatty acids and monoglycerides. It is found in the gastric juice as a weak fat- splitting
enzyme which works in an acidic medium. It is a tributyrase and is meant to digest trybutyrin, the
common butter fat into fatty acids and glycerol.

Stimulation and Control of Gastric Digestion

The gastric glands pour out their secretions into the lumen of stomach in response to the reflexes set in by
the presence of food in the buccal cavity. The gastric glands are under both nervous and hormonal control.
During the gastric phase, the hormone gastrin is secreted by G cells in the stomach in response to the presence
of proteins. Gastrin stimulates the release of stomach acid, or hydrochloric acid (HCl) which aidsin the
digestion of the proteins. However, when the stomach is emptied, the acidic environment need not be
maintained and a hormone called somatostatin (D cells) stops the release of hydrochloric acid. This is
controlled by a negative feedback mechanism.
Nervous Mechanism (Additional Information)
Local myenteric reflex - The food particles stimulate the local nerve plexus(myenteric) present in the wall of the stomach. (These nerve fibers
release acetylcholine which stimulates the gastric glands to secrete large quantity of gastric juice. Simultaneously, acetylcholine stimulates G cells
to secrete gastrin).
Vagovagal reflex is the reflex which involves both afferent and efferent vagal fibers. (Entrance of bolus into the stomach stimulates the sensory
(afferent) nerve endings of vagus and generates sensory impulses, the impulses are transmitted to dorsal nucleus of vagus, located in medulla of
brainstem. This nucleus in turn, sends efferent impulses through the motor (efferent) fibers of vagus, back to stomach and cause secretion of gastric
juice. Since, both afferent and efferent impulses pass through vagus, this reflex is called vagovagal reflex).
Hormonal Mechanism – Gastrin
Gastrin is a gastrointestinal hormone secreted by the G cells which are present in the pyloric glands of stomach. Small amount of gastrin is also
secreted in mucosa of upper small intestine. Gastrin is released when food enters stomach. The release of gastrin may be the local nervous reflex
or vagovagal reflex. Nerve endings release the neurotransmitter called gastrin releasing peptide, which stimulates the G cells to secrete gastrin.
Gastrin stimulates the secretion hydrochloric acid and pepsinogen by the gastric glands.

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 3 Digestion in Intestine – Intestinal Digestion

From the stomach, the semi- digested food, called chyme, is passed down through pyloric valve into the small
intestine. Here it is further acted upon by bile and pancreatic secretion and then by intestinal juices called
Succus entericus
Hepatobilary system: The hepatobiliary system is essential for digestion and includes: the liver, pancreas,
bile ducts and the gallbladder.
Pancreas
The pancreas is an organ located in the abdomen. It is spongy, about six to ten inches long, and is shaped
like a flat pear or a fish extended horizontally across the abdomen. The wide part, called the head of the
pancreas, is positioned toward the center of the abdomen. The head of the pancreas is located at the juncture
where the stomach meets the first part of the small intestine. This is where the stomach empties partially
digested food into the intestine, and the pancreas releases digestive enzymes into these contents. The central
section of the pancreas is called the neck and body. The thin end is called the tail and extends to the left side.

The pancreas has two main functions: an exocrine function that helps in digestion and an endocrine
function that regulates blood sugar.

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The exocrine gland is composed of finer tubules ending into the pancreatic lobules or acini. The main duct
of the gland called duct of wirsung communicates with the bile duct at the ampulla of vater and discharges
its secretion into the duodenum through the sphincter of oddi.

Composition of Pancreatic juice – It is a clear, watery but potent digestive juice. About 500-800ml of it is
secreted per day which contains 98.5% water, 1.5% solids containing many enzymes, sodium bicarbonate and
many other salts. It is alkaline with a pH of 7.5 to 8.0. The most important pancreatic enzymes are trypsin,
chymotrypsin, carboxypeptidase, pancreopeptidase, amylase, lipase, maltase and esterase.
Chemical Digestion – The pancreatic juice has the following enzymes
1. Trypsin - It is secreted by the pancreas in an inactive form called trypsinogen. It is activated by the
other enzyme called enterokinase (enteropeptidase – this is produced and secreted by the duodenal
mucosa as the food comes in contact with it). It converts proteins, proteoses, and peptones into
polypeptides and dipeptides.

Trypsinogen (inactive) Enteropeptidase Trypsin(active)

Proteins + Proteoses + Peptones Trypsin Tripeptides and Dipeptides

2. Chymotrypsin - It is secreted in pancreas in an inactive form called chymotrypsinogen. It is activated

by trypsin. It acts on native proteins, proteoses, and peptones and gives polypeptides and
dipeptides. Chymotrypsin also acts on milk.

Chymotrypsinogen (inactive)→Chymotrypsin(active)
Proteins + Proteoses + Peptones chymotrypsin Tripeptides and Dipeptides

3. Carboxypeptidase - Inactive form of procarboxypeptidase are activated by trypsin. It is another

proteolytic enzyme which hydrolyze the amino acids one by one from the end of a polypeptide chain
have a free carboxyl group. (exopeptidase)
4. Pancreopeptidase or Elastase – it is also called as pancreatic endopeptidase. Elastases are
synthesized as inactive precursors called proelastases which are stored in the acinar cells of the
pancreas. Trypsin-induced activation takes place in the duodenum. Pancreatic elastase is a protein-
digesting enzyme, and serves as a non-invasive stool biomarker of pancreatic exocrine function.
5. Pancreatic amylase - Amylopsin - Its action is like that of salivary amylase. It digests cooked starch
more actively than raw starch. It requires chloride ions for normal activity. The end products of
amylopsin are maltose and small amount of glucose.

Starch Amylase Maltose

6. Pancreatic lipase - Steapsin or lipase - It is a fat splitting enzyme. Its activity is increased in the
presence of a number of substances like calcium, bile acids, certain amino acids and peptides. It is a
glyceride hydrolyzing enzyme. The end product of the hydrolysis consists of diglycerides,
monoglycerides and fatty acids.

Fats Lipase Fatty acid + Glycerol

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 7. Pancreatic Nuclease – It includes ribonuclease or RNAase and deoxyribonuclease or DNA ase. They
act on the polynucleotide chains of RNA and DNA leading to the formation of nucleotides of different
length.
Liver and Gallbladder
The liver is the heaviest gland of the body, weighing about 1.4 kg in an average adult. The liver is located
in the upper right-hand portion of the abdominal cavity, beneath the diaphragm, and on top of the stomach,
right kidney, and intestines. It is cone-shaped. The liver is a dark reddish-brown.
The gallbladder, is a pear-shaped sac that is located in a depression of the posterior surface of the liver. It is
7–10 cm long and typically hangs from the anterior inferior margin of the liver.
The liver is covered by visceral peritoneum and by a dense irregular connective tissue layer that lies deep
to the peritoneum. The liver is divided into two main lobes – a large right lobe and a smaller left lobe by
the falciform ligament. (right lobe is considered to include an inferior quadrate lobe and a posterior caudate
lobe by anatomist).
The part of the gallbladder includes the broad fundus, the central portion – body, and the tapered portion –
neck. A cystic duct arises from the gall bladder. It opens into the hepatic duct. The two ducts form a common
bile duct which opens into the duodenum along with the pancreatic duct
• Liver is made up of many lobes called hepatic lobes.
• Each consists of many lobules called hepatic lobules
• Hepatic lobule is the structural and functional unit of liver.
• There are about 50,000 to 100,000 lobules in the liver.
• The lobules are made up of liver cells called hepatocytes.
• Hepatocytes are arranged in columns, which form the hepatic plates. Each plate is made up of two columns
of cells. In between the two columns of each plate lies a bile canaliculus.
• A blood space, called sinusoids is present in between the neighboring plates,
• In between the endothelial cells, Kupffer cells (macrophages) are present.
• Each lobule is surrounded by many portal triads (consists of hepatic artery, portal vein and tributary of bile
duct)
• Branches of hepatic artery and portal vein opens into the sinusoids, which then opens into the central vein and
this empties into hepatic vein.
• Bile, secreted by hepatic cells (liver cells) are emptied into bile canaliculus. From here, the bile enters the
tributary of bile duct. Tributaries of bile duct from canaliculi of neighboring lobules unite to form small bile
ducts.
• These small bile ducts join together and finally form left and right hepatic ducts, which emerge out of liver.
• Biliary system – is formed by gallbladder and extrahepatic bile ducts.
• Right and left hepatic bile ducts, which comes out of the liver join to form common hepatic duct. It unites
with the cystic duct from gallbladder to form common bile duct.
• Common bile duct unites with the pancreatic duct to form the common hepatopancreatic duct to form the
common hepatopancreatic duct or ampulla of vater, which opens into the duodenum.
• Sphincter of Oddi at the lower portion of the common bile duct, is opened upon appropriate stimulation and
allows the flow of bile from gallbladder into the intestine.
• Blood supply to Liver - Liver receives blood from two sources, namely the hepatic artery and portal vein.
• Hepatic artery arises from aorta and supplies oxygenated blood to liver. After entering the liver, hepatic
artery divides into many branches. Each branch enters a portal triad.
• Portal vein is formed by superior mesenteric vein and splenic vein. It brings deoxygenated blood from
stomach, intestine, spleen and pancreas.
• Portal blood is rich in monosaccharides and amino acids. It also contains bile salts, bilirubin, urobilinogen and
GI hormones. However, the oxygen content is less in portal blood.
• Flow of blood from intestine to liver through portal vein is known as enterohepatic circulation
• The blood from hepatic artery mixes with blood from portal vein in hepatic sinusoids. Hepatic cells obtain
oxygen and nutrients from the sinusoid.
• Substances synthesized by hepatic cells, waste products and carbon dioxide are discharged into sinusoids.

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• Sinusoids drain them into central vein of the lobule. Central veins from many lobules unite to form
bigger veins, which ultimately form hepatic veins (right and left) which open into inferior vena cava.

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Schematic diagram of blood flow through liver Gallbladder

Functions of the liver- Some of the more well-known functions include the following:
In addition to secreting bile, which is needed for absorption of dietary fats, the liver performs many other vital functions:
• Carbohydrate metabolism - The liver is especially important in maintaining a normal blood glucose level. When
blood glucose is low, the liver can break down glycogen to glucose and release the glucose into the bloodstream.
The liver can also convert certain amino acids and lactic acid to glucose, and it can convert othersugars, such as
fructose and galactose, into glucose. When blood glucose is high, as occurs just after eating a meal, the liver
converts glucose to glycogen and triglycerides for storage.
• Lipid metabolism - Hepatocytes store some triglycerides; breakdown fatty acids to generate ATP; synthesize
lipoproteins, which transport fatty acids, triglycerides, and cholesterol to and from body cells; synthesize
cholesterol; and use cholesterol to make bile salts.
• Protein metabolism - Hepatocytes deaminate (remove the amino group, NH2, from) amino acids so that the
amino acids can be used for ATP production or converted to carbohydrates or fats. The resulting toxic ammonia
(NH3) is then converted into the much less toxic urea, which is excreted in urine. Hepatocytes also synthesize
most plasma proteins, such as alpha and beta globulins, albumin, prothrombin, and fibrinogen.
• Processing of drugs and hormones - The liver can detoxify substances such as alcohol and excrete drugs such
as penicillin, erythromycin, and sulfonamides into bile. It can also chemically alter or excrete thyroid hormones
and steroid hormones such as estrogens and aldosterone.
• Excretion of bilirubin - Bilirubin, derived from the heme of aged red blood cells, is absorbed by the liver from
the blood and secreted into bile. Most of the bilirubin in bile is metabolized in the small intestine by bacteria
and eliminated in feces.
• Synthesis of bile salts - Bile salts are used in the small intestine for the emulsification and absorption of lipids.
• Phagocytosis - The stellate reticuloendothelial (Kupffer) cells of the liver phagocytize aged red blood
cells,white blood cells, and some bacteria.
• Activation of vitamin D - The skin, liver, and kidneys participate in synthesizing the active form of vitamin D.

Gall bladder
• The gallbladder has a storage capacity of 30-50ml
• Its primary function is to store and concentrate bile, a yellow-brown digestive enzyme produced by
the liver.
• When food is not eaten, the bile passes into the gall bladder from the liver and the sphincter of oddi
remains closed.

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 • When food is eaten, the gall bladder contracts, the sphincter of oddi opens and the bile passes
through the cystic duct into the duodenum.
Bile
The bile is an alkaline fluid. It contains about 88.9% - 97.5% water, bile salts, pigments, fatty acids, cholesterol
and many inorganic constituents. It is slightly alkaline in gall bladder (pH 7.0 to 7.6) but the hepatic bile has
a pH range of 8.0 to 8.6.
Composition of bile

Bile salts - Bile salts are the sodium and potassium salts of bile acids, which are conjugated with glycine or taurine.

Formation of Bile Salts: Bile salts are formed from bile acids. The two primary bile acids namely cholic acid and chenodeoxycholic
acid which are formed in the liver and then enters the intestine. Due to the bacterial action in the intestine, the primary bile acids are converted
into secondary bile acids.
Cholic acid → Deoxycholic acid
Chenodeoxycholic acid → lithocholic acid

The secondary bile acids from intestine are transported back to liver through enterohepatic circulation. In liver, these acids are conjugated
with glycine (amino acid) or taurine (derivative of an amino acid) and form conjugated bile acids namely glycocholic acid and taurocholic
acids. These bile acids combine with sodium or potassium ions to form the salts, sodium or potassium glycocholate and sodium or
potassium taurocholate.

Enterohepatic circulation is the transport of substances from small intestine to liver through portal vein. About 90 to 95% of bile salts from
intestine are transported to liver through enterohepatic circulation. The remaining 5 to 10% of the bile salts enter large intestine. Here
the bile salts are converted into deoxycholate and lithocholate and excreted in feces.

Functions of Bile salts - They are required for digestion and absorption of fats in the intestine.
Emulsification of Fats - Emulsification is the process by which the fat globules are broken down into minute droplets and made
in the form of a milky fluid called emulsion in small intestine, by the action of bile salts. Unemulsified fat usually passes through
the intestine and then it is eliminated in feces.
(Lipolytic enzymes of GI tract cannot digest the fats directly because the fats are insoluble in water due to the surface tension. Bile salts emulsify the fats by reducing the surface tension
due to their detergent action. Now the fats can be easily digested by lipolytic enzymes).
Absorption of Fats - Bile salts help in the absorption of digested fats from intestine into blood. Bile salts combine with fats and
make complexes of fats called micelles. The fats in the form of micelles can be absorbed easily.
Prevention of Gallstone Formation - Bile salts prevent the formation of gallstone by keeping the cholesterol and lecithin in
solution. In the absence of bile salts, cholesterol precipitates along with lecithin and forms gallstone.

Bile Pigments - Bile pigments are the excretory products in bile. Bilirubin and biliverdin are the two bile pigments and bilirubin
are the major bile pigment in human beings.
Additional Information (Formation of Bile pigments -Bile pigments are formed during the breakdown of hemoglobin, which is released from the destroyed RBCs in the reticuloendothelial system. Senile
erythrocytes are destroyed in reticuloendothelial system and hemoglobin is released from them. Hemoglobin is broken into globin and heme. Heme is split into iron and the pigment biliverdin. Iron goes to iron
pool and is reused. First formed pigment biliverdin is reduced to bilirubin. Bilirubin is released into blood from the reticuloendothelial cells. In blood, the bilirubin is transported by the plasma protein, albumin.
Bilirubin circulating in the blood is called free bilirubin or unconjugated bilirubin. Within few hours after entering the circulation, the free bilirubin is taken up by the liver cells. In the liver, it is conjugated with
glucuronic acid to form conjugated bilirubin. Conjugated bilirubin is then excreted into intestine through bile.

Excretion of Bile pigments - In intestine, 50% the conjugated bilirubin is converted into urobilinogen by intestinal bacteria. First the conjugated bilirubin is deconjugated into free bilirubin, which is later reduced into
urobilinogen. Remaining 50% of conjugated bilirubin from intestine is absorbed into blood and enters the liver through portal vein (enterohepatic circulation). From liver, it is re-excreted in bile. Most of the urobilinogen
from intestine enters liver via enterohepatic circulation. Later, it is re- excreted through bile. About 5% of urobilinogen is excreted by kidney through urine. In urine, due to exposure to air, the urobilinogen is converted
into urobilin by oxidation. Some of the urobilinogen is excreted in feces as stercobilinogen. In feces, stercobilinogen is oxidized to stercobilin)

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 Formation & Circulation of Bile pigments
• They activate the enzymes cholesterol esterase and pancreatic lipase.
• They activate the inactive lipase
• Certain wastes are excreted through bile.

Control of Pancreatic and Bile Secretion

The duodenal secretion of pancreas and liver are regulated by stimulation of vagus nerves leads to the
secretion of pancreatic secretion. Two hormones such as secretin and pancreozymin controls the enzyme
producing functions of the pancreas.
• Secretin is produced by the enteroendocrine cells of the duodenal mucosa and it has the effect of
stimulating the pancreas to produce alkaline secretions as well as slowing the emptying of the
stomach.
• Cholecystokinin (CCK) is produced in the duodenum. It stimulates the release of bile from the gall
bladder, stimulation of pancreatic juice secretion.

Small Intestine
Most digestion and absorption of nutrients occur in a long tube called the small intestine. Its length alone provides a large
surface area for digestion and absorption, and that area is further increased by circular folds, villi, and microvilli. The small
intestine is divided into 3 regions – duodenum. jejunum and ileum.

Intestinal villi
Mucous membrane of small intestine is covered by minute projections called villi. Villi are lined by columnar cells, which are
called enterocytes. Each enterocyte gives rise to hair-like projections called microvilli. Villi and microvilli increase the surface
area of mucous membrane by many folds. Within each villus, there is a central channel called lacteal, which opens into lymphatic
vessels. It contains blood vessels also.
When viewed through microscope, the microvilli are too small to be seen individually, instead they form a fuzzy line, called the
brush border, extending into the lumen of the small intestine. There are an estimated 200 million microvilli per square millimeter
of small intestine. The brush border also contains several brush border enzymes that have digestive functions. Secretion from
small intestine is called succus entericus. The food entering the intestine is called as the Chyle.

The epithelial layer of the small intestinal mucosa consists of simple columnar epithelium that contains many types of cells.
• Absorptive cells of the epithelium release enzymes that digest food and contain microvilli that absorb nutrients in
small intestinal chyme.
• Goblet cells, which secrete mucus.
• Paneth cells – secrete cytokines called defensins; secret lysozyme, a bactericidal enzyme and are capable of
phagocytosis.
• Three types of enteroendocrine cells are found – S cells (secrete secretin), CCK cells (cholecystokinin) and K
cells (secrete glucose – dependent insulinotropic peptide)

Intestinal glands, or crypts of Lieberkühn - The small intestinal mucosa contains many deep crevices lined with glandular
epithelium. Cells lining the crevices form the intestinal glands, or crypts of Lieberkühn and secrete intestinal juices.
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Duodenal glands – the submucosa of the duodenum contains duodenal glands or Brunner’s gland, which secrete an
alkaline mucus that help neutralize gastric acid in the chyme.
Circular folds or Plicae circulares are folds of the mucosa and submucosa facilitate the process of digestion and
absorption.
Role of intestinal juice and brush-border enzyme
Together, pancreatic and intestinal juices provide a liquid medium that aids the absorption of substances
from chyme in the small intestine. The absorptive cells of the small intestine synthesize several digestive
enzymes, called brush-border enzymes, and insert them in the plasma membrane of the microvilli. Thus,
some enzymatic digestion occurs at the surface of the absorptive cells that line the villi, rather than in the
lumen exclusively, as occurs in other parts of the GI tract.
Among the brush-border enzymes are four carbohydrate-digesting enzymes called α-dextrinase, maltase,
sucrase, and lactase; protein-digesting enzymes called peptidases (aminopeptidase and dipeptidase); and
two types of nucleotide-digesting enzymes, nucleosidases and phosphatases.

Composition of Succus Entericus

They produce intestinal juice at the rate of 1800 ml per day. It has a pH of about 8.3. about 98.5% of it is a
water and the organic and the inorganic constituents are 0.6 to 1.0% respectively.

Mechanical and Chemical Digestion in the Small Intestine

Mechanical Digestion
The two types of movements of the small intestine—segmentations and a type of peristalsis called migrating motility
complexes – are governed mainly by the myenteric plexus.
Segmentations are localized, mixing contractions that occur in portions of intestine distended by a large volume of chyme.
Segmentations mix chyme with the digestive juices and bring the particles of food into contact with the mucosa for
absorption; they do not push the intestinal contents along the tract.
A segmentation starts with the contractions of circular muscle fibers in a portion of the small intestine, an action that constricts
the intestine into segments. Next, muscle fibers that encircle the middle of each segment also contract, dividing each segment
again. Finally, the fibers that first contracted relax, and each small segment unites with an adjoining small segment so that
large segments are formed again. As this sequence of events repeats, the chyme moves back and forth. Segmentations occur
most rapidly in the duodenum, about 12 times per minute, and progressively slow to about 8 times per minute in the ileum.
After most of a meal has been absorbed, which lessens distension of the wall of the small intestine, segmentation stops and
peristalsis begins.

The type of peristalsis that occurs in the small intestine, termed a migrating motility complex (MMC), begins in the lower
portion of the stomach and pushes chyme forward along a short stretch of small intestine before dying out. The MMC
slowly migrates down the small intestine, reaching the end of the ileum in 90–120 minutes. Then another MMC begins in
the stomach. Altogether, chyme remains in the small intestine for 3–5 hours.

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Chemical Digestion
In the mouth, salivary amylase converts starch (a polysaccharide) to maltose (a disaccharide), maltotriose (a trisaccharide),
and α-dextrins (short-chain, branched fragments of starch with 5–10 glucose units). In the stomach, pepsin converts proteins
to peptides (small fragments of proteins), and lingual and gastric lipases convert some triglycerides into fatty acids,
diglycerides, and monoglycerides. Thus, chyme entering the small intestine contains partially digested carbohydrates,
proteins, and lipids. The completion of the digestion of carbohydrates, proteins, and lipids is a collective effort of pancreatic
juice, bile, and intestinal juice in the small intestine.
The enteric enzymes bring about the following digestive changes in the chyle

Erepsin – it is a complex of enzymes namely aminopeptidase, dipepetidase, carboxypeptidase and

polypeptidase. The proteins enter the intestine mainly as polypeptides; thus, carboxypeptidase breaks-up the
peptide linkage next to carboxyl group; while the aminopeptidase acts on a peptide linkage next to an amino
group, which are finally hydrolyzed into simple amino acids:
Polypeptides + Proteases Erepsin Amino acids
Intestinal Lipase – The intestinal lipase together with the pancreatic lipase splits the remaining fats into fatty
acid and glycerol.
Emulsified fats Lipase fatty acid and glycerol

Intestinal maltase – in cooperation with pancreatic maltase, changes maltose into glucose
Maltose maltase glucose
Lactase – converts milk sugar into galactose and glucose
Invertase – acts on the disaccharide sucrose and change into glucose and fructose.
Nucleases – acts upon the nucleic acids and hydrolyze them into simple nucleotides.

Control of Intestinal Secretions

Nervous Regulation
Stimulation of parasympathetic nerves causes vasodilatation and increases the secretion of succus entericus. Stimulation of
sympathetic nerves causes vasoconstriction and decreases the secretion of succus entericus. But, the role of these nerves in the
regulation of intestinal secretion in physiological conditions is uncertain. However, the local nervous reflexes play an important
role in increasing the secretion of intestinal juice. When chyme enters the small intestine, the mucosa is stimulated by tactile
stimuli or irritation. It causes the development of local nervous reflexes, which stimulate the glands of intestine.

Hormonal Regulation
When chyme enters the small intestine, intestinal mucosa secretes Enterocrinin, secretin and cholecystokinin, which promote
the secretion of succus entericus by stimulating the intestinal glands.
(GhrelinA hormone produced in the stomach that boosts appetite, slows metabolism and reduces fat burning. It may be involved in the
development of obesity. is produced in the stomach, and its function is to tell the brain that the body has to be fed. It increases appetite).

Functions of Small Intestine

• Segmentations mix chyme with digestive juices and bring food into contact with mucosa for absorption; peristalsis propels chyme through small
intestine.
• Completes digestion of carbohydrates, proteins, and lipids; begins and completes digestion of nucleic acids.
• Absorbs about 90% of nutrients and water that pass through digestive system

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 Digestion in the Large Intestine

The large intestine is the terminal portion of the GI tract. The overall functions of the large intestine are the
completion of absorption, the production of certain vitamins, the formation of feces, and the expulsion
of feces from the body.
The large intestine is about 1.5m long. It extends from the ileum to the anus. Structurally, the four major
regions of the large intestine are the cecum, colon, rectum, and anal canal. The opening from the ileum into
the large intestine is guarded by the ileocecal sphincter (valve), which allows materials from the small
intestine to pass into the large intestine. Hanging inferior to the ileocecal valve is the cecum, a small pouch.
Attached to the cecum is a twisted, coiled tube, called the appendix or vermiform appendix.
The open end of the cecum merges with a long tube called the colon, which is divided into ascending,
transverse, descending, and sigmoid portions. The sigmoid colon terminates as the rectum. The rectum is
about 15 cm in length and the terminal 2–3 cm of the large intestine is called the anal canal. The opening of
the anal canal to the exterior, called the anus, is guarded by an internal anal sphincter of smooth muscle
(involuntary) and an external anal sphincter of skeletal muscle (voluntary). Normally these sphincters keep
the anus closed except during the elimination of feces.
The epithelium contains mostly absorptive and goblet cells. The absorptive cells function primarily in
water absorption; the goblet cells secrete mucus that lubricates the passage of the colonic contents.

Mechanical Digestion in the Large Intestine

The passage of chyme from the ileum into the cecum is regulated by the action of the ileocecal sphincter.
Normally, the valve remains partially closed so that the passage of chyme into the cecum usually occurs
slowly. Immediately after a meal, a gastroileal reflex intensifies peristalsis in the ileum and forces any chyme
into the cecum. Whenever the cecum is distended, the degree of contraction of the ileocecal sphincter
intensifies.
Movements of the colon begin when substances pass the ileocecal sphincter. Because chyme moves through
the small intestine at a fairly constant rate, the time required for a meal to pass into the colon is determined by
gastric emptying time. As food passes through the ileocecal sphincter, it fills the cecum and accumulates in
the ascending colon.
Haustral churning - In this process, the haustra remain relaxed and become distended while they fill up.
When the distension reaches a certain point, the walls contract and squeeze the contents into the next haustrum.
Peristalsis also occurs, although at a slower rate (3–12 contractions per minute) than in more proximal portions
of the tract.
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A final type of movement is mass peristalsis, a strong peristaltic wave that begins at about the middle of the
transverse colon and quickly drives the contents of the colon into the rectum. Because food in the stomach
initiates this gastrocolic reflex in the colon, mass peristalsis usually takes place three or four times a day.

Chemical Digestion in the Large Intestine

The final stage of digestion occurs in the colon through the activity of bacteria that inhabit the lumen. Mucus
is secreted by the glands of the large intestine, but no enzymes are secreted. Chyme is prepared for elimination
by the action of bacteria, which ferment any remaining carbohydrates and release hydrogen, carbon dioxide,
and methane gases. These gases contribute to flatus (gas) in the colon, termed flatulence when it is excessive.
Bacteria also convert any remaining proteins to amino acids and break down the amino acids into simpler
substances: indole, skatole, hydrogen sulfide, and fatty acids. Some of the indole and skatole is eliminated in
the feces and contributes to their odor; the rest is absorbed and transported to the liver, where these compounds
are converted to less toxic compounds and excreted in the urine. Bacteria also decompose bilirubin to simpler
pigments, including stercobilin, which gives feces their brown color. Bacterial products that are absorbed in
the colon include several vitamins needed for normal metabolism, among them some B vitamins and vitamin
K.
The Defecation Reflex
Mass peristaltic movements push fecal material from the sigmoid colon into the rectum. The resulting
distension of the rectal wall stimulates stretch receptors, which initiates a defecation reflex that results in
defecation, the elimination of feces from the rectum through the anus.

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 DIGESTIVE ENZYMES

Digestive enzymes are enzymes that break down polymeric macromolecules into smaller building blocks,
in order to facilitate their absorption by the body to maintain a healthy life. Most digestive enzymes are
hydrolases and produce their action by the process of hydrolysis.

Classification of Digestive Enzymes

The enzymes are classified on the nature of substrates they work. Digestive enzymes are broadly
classified into four groups. They are
1. Proteolytic Enzyme - It splits protein into amino acids.
2. Amylolytic Enzyme - It splits carbohydrate and starch into simple sugars.
3. Lipolytic Enzyme - It splits fats into fatty acid and glycerol
4. Nucleolytic Enzyme - It splits nucleic acids to nucleotides

Chemical digestion of food – Chemical digestion s a process of hydrolysis in which a complex food molecule
combines chemically with water in the presence of an enzyme and split into two or more simpler molecules.

ROLE OF ENZYMES IN CARBOHYDRATE DIGESTION

Carbohydrates are hydrates of carbon. They contain carbon, hydrogen and oxygen in the ratio 1:2:1. They
are defined as an organic compound that contain carbonyl group, namely aldehyde or ketone in addition
to two or more alcohol groups or that yields such compounds on hydrolysis. The basic unit of carbohydrate
molecule is known as monosaccharide. It is the main source of energy and plays a main role in metabolism.
Example – glucose, fructose, sucrose, lactose, cellulose, starch, etc. Human diet contains 60 to 83%
carbohydrates
Definition: The breakdown of complex carbohydrates into monosaccharides is called carbohydrate digestion.
The chemical digestion or hydrolysis of carbohydrates occurs in the presence of differentenzymes known as
glycosidases.
Places of carbohydrate digestion - The digestion of carbohydrates occurs in the buccal cavity, and
intestine.

Glands involved in Carbohydrate Digestion

1. Salivary glands, 2. Pancreas and 3. Intestinal glands

Enzymes involved in Carbohydrate Digestion

The enzymes involved in the digestion of carbohydrates are called as Carbohydrases. They are: a)
Ptyalin or salivary amylase, b) Amylopsin or Pancreatic Amylase, c) Maltase, d) Lactase, e) Sucrose or
Invertase and f) Cellulase
By Salivary Gland
Digestion of carbohydrate begins in the mouth, with the secretion of the enzyme salivary amylase from the
serous cells (opaque zymogen granules) of the salivary gland. This enzyme breaks starch and glycogen into
disaccharides. The mucous cells of the salivary gland secrete mucus, which causes the food to stick together,
and acts as a lubricant to aid in swallowing.

The salivary glands are grouped into three categories: the parotid gland is present on each side of the face
below and in front of the ears, the submandibular glands are present in the angles of the lower jaw, and the
sublingual glands are located below the tongue. The salivary glands are glandular and are made up of many
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sac-like alveoli or acini which form lobules. Ducts arise from each alveolus unite to form large ducts which
open into the main duct.
The food (bolus) is forced into the pharynx by the tongue. As the food is swallowed, it moves into the
esophagus, which essentially provides a passageway from the pharynx to the stomach. The mucous glands
of the esophagus secrete mucus to aid in moistening and lubricating the bolus.

Salivary Amylase (Digestion of Starch)

Salivary amylase, also known as ptyalin, which is secreted by the salivary glands, initiates the breakdown of
starch. Dietary carbohydrates are either monosaccharide and disaccharide sugars or complex polysaccharides
such as starches. Most of the carbohydrates we eat are starches, but only monosaccharides can be absorbed
into the bloodstream. Thus, ingested disaccharides and starches must be broken down into monosaccharides.
The function of salivary amylase is to begin starch digestion by breaking down starch into smaller molecules
such as the disaccharide maltose, the trisaccharide maltotriose, and short-chain glucose polymers called
α-dextrins. Even though food is usually swallowed too quickly for all starches to be broken down in the
mouth, salivary amylase in the swallowed food continues to act on the starches for about another hour, at which
time stomach acids inactivate it.
Ptyalin (pH 6.8)
Starch →Dextrins→ Maltotriose → Maltose
Lingual Lipase
Saliva also contains lingual lipase, which is secreted by lingual glands in the tongue. This enzyme becomes
activated in the acidic environment of the stomach and thus starts to work after food is swallowed. It breaks
down dietary triglycerides (fats and oils) into fatty acids and diglycerides. A diglyceride consists of a glycerol
molecule that is attached to two fatty acids

By Gastric glands
No digestion of carbohydrates takes place in the stomach. The bolus passes first section of the stomach. The
stomach is divided into several regions: the cardiac region, body region, fundic region, and pyloric region.
The stomach works to mix and churn the food, which aids in further digestion of carbohydrates. At this point,
the bolus is converted into a semi fluid paste of bolus and gastric juices called chyme
By Intestinal Gland

The chyme then travels through the first section of the small intestines. The small intestine is divided into
three sections: the duodenum, jejunum, and ileum. The majority of digestion of carbohydrates takes place
in the small intestines. As the chyme moves into the duodenum, an enzyme called pancreatic amylase is
released through the pancreatic duct. This enzyme splits molecules of starch and glycogen into maltose
(disaccharides).

The interior wall of the small intestines is covered with tiny projections called the villi. These projections
increase the surface area of the intestines and play an important part in the process of absorption of the
nutrients. The epithelial cells of the villi contain even smaller extensions, called microvilli. Embedded in the
microvilli are digestive enzymes which are needed to further break down carbohydrates. These include
sucrase, maltase, and lactase, which break down the disaccharides into monosaccharide (glucose,
galactose and fructose). This monosaccharide is then absorbed by the villi and enters the blood capillaries
to be transported to other parts of the body.
Pancreatic amylase - Amylopsin
Its action is like that of salivary amylase. It digests cooked starch more actively than raw starch. It requires
chloride ions for normal activity. The end products of amylopsin are maltose and small amount of glucose.

Starch Amylase Maltose

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Intestinal maltase – in cooperation with pancreatic maltase, changes maltose into glucose
Maltose Intestinal maltase Glucose

Lactase – Converts milk sugar into galactose and glucose

Invertase – Acts on the disaccharide sucrose and change into glucose and fructose.

Digestion of Cellulose
Cellulose is polysaccharide found abundantly in the cell wall of plants. Cellulose is digested by cellulase
into glucose. Man cannot produce cellulase. In herbivorous animas, the alimentary canal contains numerous
bacteria and protozoans.
In ruminant mammals, cellulose is digested by the bacteria and protozoan ciliates living inside the rumen. In
the rumen, bacteria digest the cellulose into acetic acid, butyric acid, propionic acid, etc. when first ingested,
food enters into the anterior dorsal sac of the rumen, where mixing and grinding occurs. Some food also
goes into the reticulum. Portion of the ingested food in the rumen and reticulum are sent back to the mouth by
periodic contraction of the diaphragm. The food reaches the mouth where it is well masticated and then
returned to the rumen, from where it passes into the reticulum and then into the psalterium. Here water and
fatty acids are absorbed and the food content pass to the true stomach or abomasum. During the stay of food
in the rumen, fermentation of the cellulose and other carbohydrates occurs by the action of bacterialenzymes
In rabbit, the plant material swallowed enters the caecum. They remain for one or two days and are
fermented. The fermented plant materials are expelled out as soft faeces. These are again eaten by the rabbits
(Coprophagy) and they reach the cardiac stomach. After digestion and absorption, the waste is excreted as
hard pellets.

ROLE OF ENZYMES IN PROTEIN DIGESTION

Protein (Greek: Protes- meaning primary or holding first place)

Protein is a macromolecule composed of one or more polypeptide chains possessing a characteristic
amino acid sequence. It is a polymer of amino acids.
Protein contains carbon, nitrogen, hydrogen, oxygen, and sulphur. They are made up of amino
acids. They are organic nitrogenous compounds. Protein is an essential constituent of living cells. They are
responsible for comprising the structure of the cell but are also concerned with function of the cell including
catalyzing of enzyme reaction, regulation of metabolism, movement and defense reaction. They are the body-
builders.
Example – keratin in hair, casein in milk, lipoprotein of blood, haemoglobin, myoglobin in muscle
tissue, ovalbumin and ovaglobulin in egg, etc.
Place of protein digestion - Digestion of protein occurs in the stomach and intestine
Glands involved in protein digestion - Digestion of protein includes the following digestive glands
1. Pancreas, 2. Gastric glands and 3. Intestinal glands

Step wise break down of protein

The digestion of the proteins involves the stepwise breakdown of the complex molecules intoproteases
and peptones, then into polypeptides, which are further broken down into dipeptides and ultimately into amino
acids.
Proteins Proteases Peptones Polypeptides Amino acids

Enzymes involved in protein digestion: The enzymes responsible for the digestion of proteins are called
proteases or peptidases or proteolytic enzymes.
The proteolytic enzymes are classified into two types –

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 1. Endopeptidase – they cut the polypeptide chains at peptide bonds away from the ends in the protein
chain. Example – pepsin, trypsin, chymotrypsin, etc.
2. Exopeptidase – they catalyze or bring about the removal of terminal amino acids. Example –
carboxypeptidase, aminopeptidase etc.
The following are proteases: a) Pepsin, b) Trypsin, c) Chymotrypsin, d) Carboxypeptidase, e) Erepsin f)
Renin

By Gastric glands: As food containing protein enters the mouth, mucus is secreted to bind the particles
together. The food then travels down the esophagus and more mucus is secreted by the wall of the esophagus.
The chief cells located in the stomach secrete gastric juices containing an enzyme precursor called pepsinogen.
As it comes in contact with hydrochloric acid, it is converted into an active form referredto as pepsin. This
begins the process of chemical digestion of dietary protein.

1. Pepsin: The main digestive function of the stomach is the partial breakdown of proteins into proteoses
and peptones. Gastric pepsin is produced by the chief cells as an inactive zymogen called pepsinogen.
It is activated by gastric HCl to form active pepsin. Pepsin breaks the polypeptide into peptones,
proteoses and polypeptides, which are latter broken down to amino acids by the proteolyticenzymes
of the pancreas and intestine.
Pepsinogen HCl Pepsin
Proteins Pepsin Proteoses, Peptones and polypeptides
2. Rennin: It is present basically in the sucking mammals. It is also secreted as an inactive prorennin and
is converted into active rennin by HCl. These enzymes bring about the coagulation of milk. This
process is called as curdling of milk.
Activated rennin splits soluble milk protein caseinogen into insoluble casein, paracasein and then
calcium paracaseinate. Calcium paracaseinte form the curd and can be further digested by gastric
pepsin to form proteoses and peptones

Prorennin HCl Rennin

Rennin + Caesinogen Casein
Rennin + Casein Paracasein
Paracasein + Calcium Calcium Paracaseinate
Calcium paracaseinate Pepsin Protesoses, Peptones, Polypeptides

By Pancreas: The chyme moves into the duodenum and enzymes begin to work on the protein. The hormone
called intestinal gastrin stimulates the gastric glands to increase secretion. The pancreatic juice contains three
protein splitting enzymes in inactive forms called trypsinogen, chymotrypsinogen, and procarboxypeptidase.
Once trypsinogen comes in contact with an enzyme called enterokinase, which is secreted by the mucosal cells
of the small intestines, then trypsin is activated. The presence of trypsin then activates the inactive
procarboxypeptidase and chymotrypsinogen, and they are converted into chymotrypsin and carboxypeptidase.
The enzymes are then able to work on the protein.

1. Trypsin - It is secreted by the pancreas in an inactive form called trypsinogen. It is activated by the
other enzyme called enterokinase (enteropeptidase – this is produced and secreted by the duodenal
mucosa as the food comes in contact with it). It converts proteins, proteoses, and peptones into
polypeptides and dipeptides. Once formed, trypsin itself activates trypsinogen by means of
autocatalytic or autoactive action.

Trypsinogen (inactive) Enteropeptidase Trypsin(active)

Proteins + Proteoses + Peptones Trypsin Tripeptides and Dipeptides

2. Chymotrypsin - It is secreted in pancreas in an inactive form called chymotrypsinogen. It isactivated

by trypsin.
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 Action of Chymotrypsin
Digestion of Protein:
It is an endopeptidase. It acts on native proteins, proteoses, and peptones and gives polypeptides and
dipeptides.
Chymotrypsinogen (inactive) Chymotrypsin(active)

Proteins + Proteoses + Peptones chymotrypsin Tripeptides and Dipeptides

Digestion of Milk: Chymotrypsin digests caseinogen faster than trypsin. Combination of both
enzymes causes rapid digestion of milk.

3. Carboxypeptidase - Inactive form of procarboxypeptidase are activate by trypsin. It is another

proteolytic enzyme which hydrolyze the amino acids one by one from the end of a polypeptide chain
have a free carboxyl group.
4. Pancreopeptidase or Elastase – it is also called as pancreatic endopeptidase. Elastases are
synthesized as inactive precursors called proelastases which are stored in the acinar cells of the
pancreas. Trypsin-induced activation takes place in the duodenum. Pancreatic elastase is a protein-
digesting enzyme, and serves as a non-invasive stool biomarker of pancreatic exocrine function.
5. Nucleases - Nucleases of pancreatic juice are ribonuclease and deoxyribonuclease, which are
responsible for the digestion of nucleic acids. These enzymes convert the ribonucleic acid (RNA) and
deoxyribonucleic acid (DNA) into mononucleotides.

By Intestinal Juice
The intestinal juice or succus entericus is secreted by two different types of glands found in the small
intestine. They are
1) Duodenal glands of or glands Brunner – these are found in the sub mucosa of the duodenum. They have
an alkaline secretion containing mucin and a weak proteolytic enzyme. The principal function of this gland is
to protect the mucosa of the first part of the duodenum against damage by acid chime from the stomach.
2) Crypts of Liberkuhn – it is present throughout the small intestine and secretes mucous and large number
of enzymes. The mucosal cells secrete an enzyme called peptidase, which splits peptide bonds into amino
acids to allow for digestion. Protein digestion is completed in the small intestines. Smaller particles of amino
acids are absorbed into the villi, and are carried away by the blood.
3) Erepsin – it is a complex of enzymes namely Aminopeptidase, Dipepetidase, Carboxypeptidase and
Polypeptidase
The proteins enter the intestine mainly as polypeptides; thus, carboxypeptidase breaks-up the peptide linkage
next to carboxyl group; while the aminopeptidase acts on a peptide linkage next to an amino group, which are
finally hydrolyzed into simple amino acids:
Polypeptides + Proteases Erepsin Amino acids

ROLE OF ENZYMES IN LIPID DIGESTION

Lipids: Lipids are organic compounds insoluble in water but soluble in fat solvents like ether, chloroform,
boiling alcohol and benzene.
Definition: Lipids are defined as the ester (combination of alcohol with an acid) of fatty acid or substances
capable of forming such esters. It contains carbon, hydrogen, oxygen and other elements such as
phosphorous and nitrogen.
Lipids are composed of 3 fatty acids joined to a glycerol molecule. Fatty acid and alcohol are the
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building blocks components of lipids. They are the important constituent of diet due to their higher energy
value. Examples – fats, oils, steroids, waxes, some hormones and vitamins, carotenoids etc.

Digestion of Lipids
The dietary lipids include neutral fats, phospholipids, free cholesterol, fatty acid and glycerol. During
digestion, the lipids are split into fatty acids and glycerol.
Lipids → Fatty acids + Glycerol.
Emulsification: It is a process by which the insoluble lipids are converted into soluble milky liquid containing
drops of oil and fat. Emulsification is the first step in the digestion of fat. It occurs in the stomach. It is brought
about by the bile secreted by the liver. Only after emulsification, the lipases begin their digestion.
Place of Lipid Digestion - The lipid digestion occurs in the stomach and intestine.
Glands involved in Lipid Digestion: Gastric glands, 2. Liver 3. Pancreas.
Enzymes involved in Lipid Digestion: The breakdown of the lipids is done by a set of enzymes called lipases.
They are: a) Lingual lipase, b) Gastric lipase, c) Pancreatic lipase, d) Phospholipase, e) Phosphodiesterase, f)
Phosphatase, g) Cholesterol esterase

In Mouth
The first step in the digestion of triacylglycerol and phospholipids begins in the mouth as lipids encounter
saliva. It is secreted by the dorsal surface of the tongue which is active at pH 2.0-7.5ml. Next, the physical
action of chewing coupled with the action of emulsifiers enables the digestive enzymes to do their tasks. The
enzyme lingual lipase, along with a small amount of phospholipid as an emulsifier, initiates the process of
digestion. These actions cause the fats to become more accessible to the digestive enzymes. As a result, the
fats become tiny droplets and separate from the watery components.

a) Lingual Lipase
It helps in splitting of short chain such as triglycerides into free fatty acid and glycerol.
Triglycerides Lingual lipase free fatty acid + Glycerol

By Gastric Glands
As food containing fat travels through the digestive tract, digestion takes place in different locations. The food
containing fats essentially follow the same pathway as carbohydrate foods, with chemical digestion primarily
beginning in the stomach.
The surface of the inner lining of the stomach contains openings called gastric glands, which are made up of
three types of cells: mucous cells, chief cells, and parietal cells.
A hormone called gastrin stimulates the gastric glands to secrete their fluids. Gastric enzymes are secreted
by the chief cells, while hydrochloric acid is secreted by the parietal cells. The combination of mucus,
hydrochloric acid and enzymes is referred to as gastric juices. The gastric juices contain small amounts of
gastric lipase, which begin the breaking down of specific lipids.

Role of Gastric lipase

Digestion of Neutral fats are acted by gastric lipase and are split into fatty acids and glycerol. First, hydrolysis
of neutral fats leads to the formation of one molecule of fatty acid and another molecule of diglyceride. This
in turn is split into one molecule of catty acid and another molecule of monoglyceride. Themonoglyceride are
further hydrolyzed into one molecule of fatty acid and one molecule of glycerol
Neutral fat Gastric Lipase Diglyceride + Fatty acid
Diglyceride Gastric Lipase Monoglyceride + Fatty acid
Monoglyceride Gastric Lipase Glycerol + Fatty acid

By Pancreas
Pancreas is both an endocrine and an exocrine gland. The exocrine part of the pancreas produces many
enzymes which enter the duodenum via the pancreatic duct. The endocrine part produces insulin and
Glucagon, the blood sugar regulators.

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 Pancreatic Lipase
Pancreatic lipase breaks down triglycerides into free fatty acids and monoglycerides. Pancreatic lipase works
with the help of the salts from the bile secreted by the liver and the gall bladder. Bile salts attach to
triglycerides to help emulsify them, which aids access by pancreatic lipase. This occurs because the lipase is
water-soluble but the fatty triglycerides are hydrophobic and tend to orient towards each other and away from
the watery intestinal surroundings. The bile salts emulsify the triglycerides in the watery surroundings until
the lipase can break them into the smaller components that are able to enter the villi for absorption

Triglycerides Pancreatic Lipase free fatty acid + Glycerol (monoglycerides)

Activity of pancreatic lipase is accelerated in the presence of bile. Optimum pH required for activity of this
enzyme is 7 to 9. Digestion of fat by pancreatic lipase requires two more factors:
i. Bile salts, which are responsible for the emulsification of fat, prior to their digestion
ii. Colipase, which is a coenzyme necessary for the pancreatic lipase to digest the dietary lipids.
About 80% of the fat is digested by pancreatic lipase. Deficiency or absence of this enzyme leads to excretion
of undigested fat in feces

Cholesterol ester hydrolase: Cholesterol ester hydrolase or cholesterol esterase converts cholesterol ester
into free cholesterol and fatty acid by hydrolysis.

Steapsin or lipase
It is a fat splitting enzyme. Its activity is increased in the presence of a number of substances like
calcium, soaps, bile acids, certain amino acids and peptides. It is a glyceride hydrolyzing enzyme. The end
product of the hydrolysis consists of diglycerides, monoglycerides and fatty acids.
Triglyceride Steapsin Diglyceride + Fatty acid
Diglyceride Steapsin Monoglyceride + Fatty acid
Monoglyceride Steapsin Glycerol + Fatty acid

Phospholipase

Phospholipase A – It is activated by Trypsin. It digests phospholipids namely lecithin and cephalin and converts
Them into lysophospholipids (lysolecithin and lysocephalin)
Phospholipase B – activated by Trypsin. It converts lysolecithin and lysocephalin to phosphoryl choline and free
Fatty acids
Lecithin Phospholipase Lysolecithin + Fatty acid
Cephalin Phospholipase Lysocephalin + Fatty acid

Colipase – It is a small coenzyme secreted as inactive procolipase. It is activated by trypsin. It facilitates

digestive action of pancreatic lipase on fats.

Bile – Refer above in Liver and gallbladder section

Intestinal lipase
The cells of the small intestines release intestinal gastrin, which increases gastric secretions. The mucosal cells
release an enzyme called intestinal lipase, which splits fats into fatty acids and glycerol. The fatty acids are
then dissolved in the epithelial cell membranes of the villi and diffuse into them. Some fatty acids may be
absorbed directly into the blood capillary, without being converted back into fat, while others are incorporated
into chylomicrons (large molecules of lipoprotein) for transport.

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 INTESTINAL ABSORPTION

ABSORPTION OF CARBOHYDRATES

Carbohydrates are broken down into smaller polysaccharides in the mouth by the action of salivary amylase. Once in
the small intestine, pancreatic amylase further breaks the polysaccharides into disaccharides. The three most common
disaccharides are maltose, sucrose and lactose. These disaccharides are broken down into monosaccharides by the
digestive enzymes found at the brush border of the enterocytes.
Carbohydrates (mouth) → Polysaccharides → Lumen of small intestine Disaccharides (Maltose, Sucrose and Lactose) →
Brush Border (Glucose, Fructose, and Galactose)

Maltose is obtained via the digestion of starch by amylase. It consists of 2 glucose monomers, which are broken down
by the brush border enzyme called maltase.

Sucrose consists of glucose and fructose and is broken down by sucrase while lactose consists of glucose and galactose
and it is broken down by lactase.

The cells of the small intestine, called enterocytes, can only absorb the simplest of sugars, in their monomeric forms.
Therefore, they absorb glucose, galactose and fructose.

Glucose and galactose enter enterocytes through sodium –linked secondary active transport (SGLT sodium
dependent glucose transporters). ATP is used to establish an electrochemical gradient in which there is a higher
concentration of sodium on the lumen side. As the sodium travels into the cell via a membrane protein, it pulls glucose/
galactose with it. The simultaneous transport of sodium and glucose is called co-transport. Since sodium and glucose
are transported simultaneously in the same direction, it is called symport.
Fructose however enters the cell via passive transport.

*Enzymes on brush border (apical sides) break down disaccharides

*Na+/K+ ATPase found on the basolateral side use ATP to pump three sodium out of the cell and 2 potassium into the
cell. This establishes an electrochemical gradient with a lower sodium concentration inside.
*While inside, majority of the fructose is converted into glucose
*Fructose, glucose and galactose then leave via passive transport on the basolateral side and enter the blood and travel
to liver.

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 ABSORPTION OF PROTEINS

Although proteins undergo mechanical digestion in the mouth, they do not begin chemical digestion until they are in
the stomach. In the stomach, pepsin cleaves proteins and breaks them down into polypeptides. Once these polypeptides
reach the lumen of the small intestine, pancreatic peptidase (trypsin, chymotrypsin and carboxypeptidases) break down
these polypeptides into small peptides. These small peptides are ultimately broken into tripeptides, dipeptides and amino
acids by the brush border enzymes.
Proteins (stomach)→polypeptides → Lumen of small intestine (small peptides) → Brush border (tripeptides,
dipeptides and amino acids)

All proteins are digested by proteolytic enzymes into either amino acid constituents or dipeptides and tripeptides
before being absorbed by the cells of the small intestine (enterocytes)
Individual amino acids are absorbed by the enterocyte using sodium-dependent cotransport (secondary transport). ATP
is used to establish an electrochemical gradient and then the sodium moves down its electrochemical gradient and brings
the amino acid with it.
Dipeptides and tripeptides use the hydrogen ion dependent co-transporter system. Here, a sodium –hydrogen exchange
protein is used to set up a hydrogen gradient. Then the hydrogen ion enters the cell and brings the different peptides with
it.
The L form of amino acids are and actively transported. The D forms are transported very slowly and passively. There
are separate transport systems for neutral amino acids, more non-polar amino acids, less non-polar amino acids, basic
amino acids and for proline and hydroxyproline.

1. Hydrolysis of small peptides and brush border on apical side.

2. Creation of sodium gradient by Na+/K+ ATP ase on basolateral side
3. Na+/H+ exchange on apical side creates H+ electrochemical gradient
4. Co-transport of amino acids using sodium movement down gradient
5. Co-transport of dipeptide and tripeptides using H+ movement down gradient
6. Majority of the Dipeptides are broken down into amino acids.
7. Amino acids exit via basolateral side and enter the blood system, where they travel to the cells of the body.

ABSORPTION OF LIPIDS
• Dietary lipids are digested in the stomach first and later in the small intestine. Various lipid digestive enzymes
digest dietary lipids. Lipid digestive enzymes are lingual lipase, gastric lipase, pancreatic lipase along with
colipase, cholesterol esterase, phospholipase A2.
• All these lipid digestive enzymes act on dietary lipids within the micelles. Micelles are formed by emulsification
process in the presence of bile acids and bile salts. Once lipids are digested into their simple form, they will
undergo simple diffusion into enterocytes.
• Once inside the cell, the fatty acids are transformed back to triglycerides in the smooth endoplasmic reticulum.
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 • Inside the lumen of the Smooth Endoplasmic Reticulum (SER), many triglycerides aggregate together and
along with cholesterol and phospholipids, form spherical structure called Chylomicrons. Proteins are also
attached to these structures and so chylomicrons are large lipoprotein.
• The majority of the lipids in the enterocytes is packaged into chylomicrons and exit the cell via exocytosis on
the basolateral side of the cell.
• They then enter the lymphatic system through the lacteal, travel via the lymph vessels and eventually enter the
blood system via the thoracic duct
•

•
(Once the chylomicrons are in the blood system, they travel to the cells of the body, especially liver and fat cells. The chylomicrons use these
apoprotein extensions to bind to membrane of endothelial cells found on blood capillaries. The membrane contains a proteolytic enzyme called
lipoprotein lipase which breaks the triglycerides inside the chylomicron into fatty acids and glycerol, which is then quickly absorbed by the cell.
The cells can then resynthesize the triglycerides and store for later use. If energy is low, they can be broken down into fatty acids and released
into the blood plasma. Inside the blood plasma, fatty acids are transported using a protein called albumin.

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