UNIT 5 DIGESTION, ABSORPTION AND

TRANSPORT OF CARBOHYDRATES,
PROTEINS AND LIPIDS

Structure

5.1 Introduction
5.2 Digestion, Absorption and Transport - Basic Concept
5.3 Digestion
5.3.1 Digestion in the Mouth
5.3.2 Digestion in the Stomach
5.3.3 Role of Pancreas in Digestion
5.3.4 Role of Bile in Digestion
5.3.5 Digestion in the Intestine
5.4 Digestion of Food Materials
5.4.1 Digestion of Carbohydrates
5.4.2 Digestion of Proteins
5.4.3 Digestion of Lipids
5.4.4 Digestion of Nucleic Acids
5.5 Absorption and Transport
5.5.1 Absorption of Carbohydrates
5.5.2 Absorption of Proteins
5.5.3 Absorption of Lipids
5.6 Let Us Sum Up
5.7 Glossary
5.8 Answers to Check Your Progress Exercises

5.1 INTRODUCTION

Nutrition emphasizes the role of foodstuffs in meeting the energy and specific
requirements of the body for continuation of cellular activity. Although food assumes
a wide variety of forms, it is categorized into major six chemical forms i.e.
carbohydrates, proteins, lipids, vitamins, minerals and water and these are
collectively termed as ‘nutrients’. The first three units in this block focused on the
chemistry of these nutrients and their properties. In this unit, we shall deal with their
digestion, absorption and transportation.

Next, once ingested, what happens to these nutrients in our body? These nutrients, as
you may already know, are utilized to perform the various functions i.e. provide
energy, body building, protection against diseases etc. in our body. Of the major six
nutrients, only carbohydrates, proteins and lipids can give energy. The ingested food
material is broken down into smaller constituents which are then passed into the
gastrointestinal tract and then into the bloodstream. The process through which the
major nutrients namely carbohydrates, proteins and lipids are converted to simple
substances and then passed into the gastrointestinal tract and then into the
bloodstream is called digestion and absorption. In this unit we shall focus on the
processes of digestion, absorption and transport of nutrients in our body, with
emphasis on the role of various enzymes which facilitate the process.

Objectives
After studying this unit, you will be able to:
• recognize the organs of the digestive system and their functions,
• understand the process of digestion in stomach, and intestine,
• discuss the role of various enzymes in facilitating the digestion process,
• explain the hormonal and neural control of secretions of salivary glands, gastric
glands, pancreas, liver and intestine, and
123
Nutritional Biochemistry • differentiate between the types of absorption and transport of final end products
across the intestinal brush border into the portal and lymphatic system.

5.2 DIGESTION, ABSORPTION AND TRANSPORT -

BASIC CONCEPT
What is digestion? The ingested food material is broken down into smaller
constituents which are assimilable by the blood. The process through which the
major nutrients namely carbohydrates, proteins and lipids are converted to simple
sugars, amino acids, fatty acids and glycerol respectively, while passing from mouth
to small intestine, constitutes digestion. The process of digestion is facilitated with
the help of secretions from salivary glands, stomach, pancreas and liver. Both
hormonal and neural controls regulate these secretions. Microvilli of the small
intestine are the major site of absorption. So then what is absorption? Absorption
involves the transfer of materials through the mucosa of the alimentary tract into
blood and lymph vessels. The transfer comprises of different transport mechanisms.
In the next few sections we shall learn about each of these processes i.e. digestion,
absorption and transport of carbohydrates, proteins and lipids. We shall begin with
digestion.

5.3 DIGESTION
Digestive enzymes break down food particles into smaller units. You will see that the
final breakdown products of protein digestion are single amino acids or small chains
of two or three amino acids. The final products of carbohydrate digestion are
monosaccharides. The final digestive products of triacylglycerol digestion are free
fatty acids and glycerol and monoacylglycerols. Vitamins, minerals, water and some
larger fat-like compounds such as cholesterol are not broken down before they are
absorbed.

Where does the digestion of food occur in our body? Obviously, it occurs in the
digestive system. You may recall reading about the anatomy of the digestive system
in the Applied Physiology Course, Unit 6. If you have not gone through this Unit, we
suggest you read the Unit now, as it will help in understanding the concepts
explained in this Unit. The human digestive system is a coiled, muscular tube (6-9
meters long when fully extended) extending from the mouth to the anus. Several
specialized compartments occur along this length- mouth, pharynx, orsophagus,
stomach, small intestine, large intestine and anus. Basically, the digestive system is
made up of two groups of organs:
• Alimentary tract, which includes mouth, pharynx, oesophagus, stomach, small
intestine, large intestine, appendix, rectum, anal canal, and
• Accessory organs, which include tongue, teeth, salivary glands, liver, gall
bladder and pancreas.

The major function of the digestive system is to ingest the food materials, digest it to
absorbable end products, absorb these products and eliminate the unusable material.
That sounds simple. But actually the process involved is not all that simple. Starting
from the mouth, the process of digestion through the different compartments of the
digestive system is discussed next.

5.3.1 Digestion in the Mouth

The mouth receives food. The tongue serves in swallowing, manipulating the food
for chewing and in perceiving taste. The teeth mechanically subdivide the food for
easier digestion. Then, there are the salivary glands. There are three pairs of salivary
glands namely parotid, submaxillary and sublingual which secrete the colourless
viscous fluid called ‘saliva’. Saliva helps in swallowing the food by lubricating the
food and the neutral pH of saliva prevents the decalcification of teeth and keeps the
124
mouth clean. In the mouth, is also present the salivary amaylase, originally called Digestion, Absorption
ptyalin. The presence of salivary amylase starts the starch digestion by breaking it and Transport
of Carbohydrates,
into dextrins. Dextrins are polysaccharides containing fewer simple sugar units than Proteins and Lipids
starches. The salivary secretion is controlled by nervous reflexes. From the mouth,
the partly digested food enters the stomach. What happens there? Let’s find out.

5.3.2 Digestion in the Stomach

To understand the digestion mechanism in the stomach, it is important to know about

the anatomy of the stomach. Look up Unit 6, in the Applied Physiology Course. You
would notice that the stomach is a ‘J’ shaped organ and secretes gastric juice. The
major components of gastric juice are hydrochloric acid and the enzyme pepsin,
which is secreted in an inactive form called pepsinogen. Small amount of lipase is
also produced but remains inactive because of the acidic nature of gastric juice.
Rennin is present only in infants and helps in the digestion of milk protein ‘casein’.
Hydrochloric acid (pH 1.2 – 1.5) activates the inactive pepsinogen to the active form
‘pepsin’ as indicated in the Figure 5.1. Hydrochloric acid denatures the food proteins
for easy digestion. The germicidal effect of hydrochloric acid also prevents the
growth of microorganisms in the stomach.

Digestion of proteins is initiated in the stomach by pepsin. Pepsin acts upon food
proteins and converts them to proteoses and peptones which are large polypeptide
derivatives as can be seen in Figure 5.1. Pepsin is an endopeptidase since it
hydrolyses peptide bonds within the main polypeptide structure. Pepsin specifically
acts upon the peptide bonds formed by amino acids with an aromatic ring
(e.g. tyrosine) or dicarboxylic amino acid i.e. amino acids with 2 COOH groups
(e.g. glutamate). You may recall reading about the structure of amino acid earlier in
Unit 2 under the protein section.

In infants, the presence of rennin helps in digestion of casein to paracasein which is

then acted upon by pepsin.
HCl
Pepsinogen Pepsin + Peptides

The pepsin formed as an active proteolytic enzyme activates the remainder of the
pepsinogen molecules by proteolytically converting them to pepsin.
Pepsin
Pepsinogen Pepsin + Peptides
–2
pH

Food proteins Proteoses and peptones

(large polypeptides)

Figure 5.1: Role of pepsin

A word about the control of gastric secretion. Gastric secretion is initiated by nervous
reflexes and the continued secretion is controlled by the hormone ‘gastrin’ produced
by the stomach. Histamine, the decarboxylated amino acid ‘histidine’ also stimulates
gastric juice production.

Next, from the stomach the partly digested food passes into the intestine via the
pancreas. We shall look at the role of pancreas in digestion now.

5.3.3 Role of Pancreas in Digestion

Pancreatic juices secreted from the pancreas aid in digestion of the food. About
600-800 ml of fluid i.e. the pancreatic juice is secreted per day by pancreas.
Pancreatic juice is alkaline in nature (pH about 8.0) and contains both organic and
inorganic substances. The enzyme components are trypsin, chymotrypsin, elastase
125
Nutritional Biochemistry and carboxypeptidase, all of which are secreted as inactive precursors called
zymogens. The other enzymes are amylase, lipase, cholesteryl ester hydrolase,
phospholipase, ribonucleases, deoxyribonucleases and phosphodiesterases.
Sodium, chloride and bicarbonate are the major inorganic constituents. Before we
proceed further, let us understand what we mean by zymogens. Zymogen is actually
the enzyme molecule synthesized, but in an inactive form. Once the food comes into
the stomach/intestine, the inactive zymogen gets converted into the active enzyme.
The example given herewith explains the concept clearly. Pepsinogen, we learnt
earlier in the sub-section 5.3.2 is the inactive form, which actually is a zymogen. HCl
in the stomach activates the inactive pepsinogen to the active enzyme pepsin.

Pepsinogen Pepsin
(Zymogen) (Active Enzyme)

Now, let us learn about the different pancreatic enzymes.

A. Trypsin

Trypsin is secreted in the inactive form trypsinogen which is converted into the active
form trypsin by the enzyme enterokinase secreted by the duodenal mucosa. Trypsin
acts upon the native proteins, proteoses and peptones and converts them to
polypeptides and peptides as indicated in Figure 5.2. It attacks the peptide linkages
containing arginine or lysine residues.

Enterokinase
Trypsinogen Trypsin + Hexapeptide

Proteoses & Peptones Polypeptides + Peptides

Figure 5.2: Role of trypsin

B. Chymotrypsin

Chymotrypsin is secreted in an inactive form ‘chymotrypsinogen’, which is activated

by trypsin as shown in Figure 5.3. Chymotrypsin is specific for peptide bonds
containing uncharged amino acid residues such as aromatic amino acids.

C. Elastase

The inactive proelastase is activated by trypsin to the active form elastase as can be
seen in Figure 5.3. Elastase attacks peptide bonds next to the small amino acid
residues such as glycine, alanine and serine and has a broader specificity than the
other two enzymes.

All the three enzymes viz. trypsin, chymotrypsin and elastase are endopeptidases
(a subclass of peptide hydrolases that hydrolyse the more centrally situated peptide
bonds). You have already seen that pepsin is also an endopeptidase.

D. Carboxypeptidase

The inactive zymogen procarboxypeptidase is activated by trypsin as shown in

Figure 5.3. The further action on the polypeptides is carried out by carboxypeptidase
which attacks the carboxy terminal peptide bond, liberating single amino acids.
So you realize that carboxypeptidase hydrolyses peptide bonds from the end of the
peptide chain. Therefore, it is called as an exopeptidase (unlike trypsin, chymotrypsin
and elastase). Further, since this enzyme acts on the peptide bonds of the peptide
chain containing free COOH (carboxylic) group, it is specifically called
carboxypeptidase.
126
The action of each of the specific enzyme discussed above is summarized in Digestion, Absorption
Figure 5.3. and Transport
of Carbohydrates,
Enterokinase
Proteins and Lipids
Trypsinogen Trypsin + Hexapeptide

Chymotrypsinogen Chymotrypsin + Peptide

Trypsin
Proelastase Elastase

Trypsin
Procarboxypeptidase Carboxypeptidase

Figure 5.3: Activation of zymogens by trypsin

E. Amylase

The presence of pancreatic amylase brings about the breakdown of starch and
glycogen and its action is similar to salivary amylase. The hydrolysis of starch and
glycogen produces maltose, maltotriose and a mixture of branched oligosaccharides
(α- limit dextrin), non branched oligosaccharides and some glucose.

F. Lipase

Pancreatic lipase specifically hydrolyses the primary ester linkages i.e. at the position
1 and 3 of triacylglycerols. The presence of bile salt helps in emulsification of fat,
thereby, increasing fat digestion. 2-monoglyceride (2-monoglycerol) and fatty acids
are the major end products of fat digestion and one fourth of the dietary
triacylglycerol is completely broken down to glycerol and fatty acids as shown in
Figure 5.4.

Triacylglycerol Glycerol Fatty Acids

Figure 5.4: Breakdown of Triacylglycerol to glycerol and fatty acids

Cholestryl ester hydrolase acts upon cholesteryl esters to liberate free cholesterol and
fatty acids. Phospholipases break down phospholipids.

G. Ribonucleases, Deoxyribonucleases, Phosphodiesterases

These enzymes are responsible for the degradation of dietary nucleic acids.
Ribonucleases and deoxyribonucleases secreted by the pancreatic juice hydrolyze
RNA and DNA to oligonucleotides. These are further hydrolyzed by pancreatic
phosphodiesterases producing a mixture of 3' and 5' mononucleotides. Nucleotidases
remove the phosphate groups hydrolytically releasing nucleosides. These are
absorbed by the mucosal cells or may be further degraded to free bases before uptake.

127
Nutritional Biochemistry We have so far studied about the different enzymes produced by the pancreas. How
are these pancreatic secretions controlled? Let’s find out. The pancreas releases its
digestive juice when stimulated by secretin, a hormone produced in the duodenum in
response to chyme (the complex contents of the stomach). Secretin causes the
secretion of an almost protein free electrolyte solution that has a high concentration
of bicarbonate. Pancreozymin or cholecystokinin-pancreozymin is a hormone
released from the mucosa of the duodenum, which causes the release of an enzyme–
rich juice from the pancreas.

Next, we shall look at the role of bile in digestion.

5.3.4 Role of Bile in Digestion

Bile is produced by liver and stored in gall bladder. During digestion, the gall bladder
contracts and supplies bile, rapidly to the duodenum through the common bile duct.

As compared to the bile produced by liver (hepatic bile), the bile stored in gall
bladder is more concentrated (since water is absorbed during the storage period in the
gall bladder) and high in bile salts, cholesterol and pigments. Bile acids are cholic
acid and deoxycholic acid and salt of these acids with glycine and taurine are known
as bile salts. Bile salts have a considerable ability to lower surface tension. By this
property, they prevent the coming together of the small fat droplets in the intestine
and thus allow more rapid digestion of the fat. This is called the emulsification action
of bile. Bile salts also combine with fatty acids and render them more easily
absorbed. This is called hydrotropic action of bile. The presence of bile in the
intestine is to accomplish digestion and absorption of fat and also of fat-soluble
vitamins.

Finally, the partly digested food enters the intestine. Let us study about the process of
digestion in the intestine.

5.3.5 Digestion in the Intestine

About 2 to 3 liters of alkaline fluid is secreted every day by the intestine and
completes the digestive process. Sodium bicarbonate and sodium chloride are the
main inorganic components. The organic components include enzymes such as
aminopeptidases, dipepetidases of various specificity, specific disaccharidases,
phospholipases etc. Let us get to know about these enzymes and about the control of
these (intestinal) secretions.

Enzymes of intestinal juice

Aminopepetidase is an exopeptidase and cleaves peptide bonds next to N-terminal

amino acids of polypeptides and oligopeptides (unlike carboxypeptidase which acts
on peptide bond at the C (carboxy) terminal). Dipeptidase completes the digestion of
dipeptides to free amino acids.

Next, let us see how the intestinal secretions are controlled.

Control of intestinal secretion

There are specific disaccharidases and oligosaccharidases i.e. α glucosidase (maltase)

which removes single glucose residues from α (1 4) linked oligosaccharides and
disaccharides, starting from the non-reducing ends. Isomaltase (α-dextrinase)
hydrolyses 1 6 bonds in α-limit dextrins. β-galactosidase (lactase) removes
galactose from lactose (i.e. hydrolyses lactose to galactose and glucose) and sucrase
hydrolyses sucrose to glucose and fructose.

128
The intestinal juice contains phospholipase that attacks phospholipids to produce Digestion, Absorption
glycerol, fatty acids, phosphoric acid and bases such as choline. and Transport
of Carbohydrates,
Proteins and Lipids
The intestinal secretion is stimulated by enterokinase produced by jejunum, of the
small intestine, which in turn is released by the entry of acid chyme from the stomach
to duodenum.

In this section we studied about the process of digestion in the mouth stomach
pancreas intestine. We have seen that the process of digestion is achieved
through the secretions (enzymes) secreted by the organs of the digestive system.
Next, we shall look at the digestion of food material. But, first let us try to answer the
questions given in check your progress 1. These will test your understanding on the
topic covered so far.

Check Your Progress Exercise 1

1) What do you understand by the term ‘digestion’? What is the major site of
absorption?
............................................................................................................................
............................................................................................................................
............................................................................................................................
2) List the various organs involved in digestion process.
............................................................................................................................
............................................................................................................................
............................................................................................................................
3) What is saliva? What is its role in digestion?
............................................................................................................................
............................................................................................................................
............................................................................................................................
4) Describe the digestive process occuring in the stomach.
............................................................................................................................
............................................................................................................................
............................................................................................................................
5) Describe the composition of pancreatic juice.
............................................................................................................................
............................................................................................................................
............................................................................................................................
6) Explain the emulsification action of bile.
............................................................................................................................
............................................................................................................................
............................................................................................................................
7) List the enzymes of intestinal juice.
............................................................................................................................
............................................................................................................................
............................................................................................................................

129
Nutritional Biochemistry
5.4 DIGESTION OF FOOD MATERIALS
Digestion, you know, involves the mixing of food, its movement through the digestive
tract and chemical breakdown of the large molecules of food into smaller molecules.
Digestion, you also learnt, begins in the mouth, when we chew and swallow, and is
completed in the small intestine. The chemical process varies somewhat for different
kinds of food/nutrients i.e. for carbohydrates, proteins, fats etc. In this section, we
shall learn about the chemical process involved during the digestion of these nutrients
in the body. We shall start with the digestion of carbohydrates.

5.4.1 Digestion of Carbohydrates

In the last section we saw that the digestion of food starts in the mouth itself by the
action of enzyme salivary amylase and the carbohydrates present in the food,
particularly the starch, are broken down by the salivary amylase to starch dextrins,
glucose and maltose. The presence of hydrochloric acid in the stomach stops further
action of salivary amylase when the food enters the stomach.

Salivary amylase
Starch dextrins + glucose + maltose

When the chyme enters the duodenum, the alkaline pH of the pancreatic juice helps
in the digestion of starch / dextrin and glycogen by pancreatic amylase. The end
products are maltose, isomaltose etc.

Pancreatic amylase
Starch / dextrins maltose + isomaltose + α- limit dextrin

The enzymes from brush border membrane of small intestine complete the digestion
of various disaccharides of the diet and the product of pancreatic amylase action such
as maltose, isomaltose, sucrose, lactose. The final products of carbohydrate digestion
are monosaccharides i.e. glucose, fructose and galactose as shown herewith.

Maltase
Maltose glucose + glucose

Isomaltase
Isomaltose glucose + glucose

Sucrase
Sucrose fructose + glucose

Lactase
Lactose galactose + glucose

Remember, cellulose is not digested in the human GI tract due to the absence of the
enzyme cellulase.

Next, we shall study about the digestion of proteins in our body.

5.4.2 Digestion of Proteins

The proteolytic enzymes secreted by gastric juice, pancreatic juice and intestinal
juice cause the hydrolysis of proteins in the gastrointestinal tract. With the exception
of the intestinal peptidases, all proteolytic enzymes are activated by the conversion of
inactive large precursors called zymogens to functional enzymes. Recall reading
about this in the last section. The enzyme pepsin acts on food proteins and converts
them into proteoses and peptones.

Pepsin
Food proteins Proteoses and peptones
(large polypeptides)
130
The polypeptides formed in the stomach are digested in the intestine by trypsin, Digestion, Absorption
chymotrypsin, elastase and carboxypeptidases secreted in the pancreatic juice. and Transport
of Carbohydrates,
Proteins and Lipids
Trypsin, chymotrypsin
Polypeptides Peptides + amino acids

Carboxypeptidase
Peptides Amino acids

The products of these enzymes are free amino acids, dipeptides and small peptides.
The residual peptides are hydrolysed in the intestinal mucosal cells by
aminopeptidases and dipeptidases. Figure 5.5 graphically presents the process of
protein digestion. The final breakdown products of protein digestion are single amino
acids or small chains of two or three amino acids.

Figure 5.5: Process of protein digestion

The proteolytic enzymes have some remarkable specificity for clearing protein chain
at certain amino acid residues. The specificities are summarized in Table 5.1.

Table 5.1: Specificity of some proteolytic enzymes

pH Major site
Enzyme Occurrence
Optimum of action
Trypsin Intestine 7.5 to 8.5 Arginyl, lysyl bonds
Chymotrypsin Intestine 7.5 to 8.5 Aromatic amino acid
bonds (Phe, Trp, Tyr)
Pepsin Stomach 1.5 to 2.5 Wide range of specificity
Carboxypeptidase Intestine 7.5 to 8.5 C-terminal amino acid
Aminopeptidase Intestinal mucosa N-terminal amino acid

After proteins, it is the turn of fats. Let us get to know about the digestion of fats in
our body.

5.4.3 Digestion of Lipids

Even though lipase is present in the stomach, it remains inactive due to the acidic
environment of the stomach contents. Therefore, the ingested fat is primarily digested
in the small intestine.

131
Nutritional Biochemistry Fat present in the food is mixed with bile and pancreatic secretion while reaching the
duodenum. The bile salts emulsify the lipids before the action of pancreatic lipase.

Pancreatic lipase acts on the dietary triacylglycerols after it has been incorporated
into the mixed micelle in the intestinal lumen. The lipase acts at interface between
water and the triacylglycerol molecules, and its catalytic action is facilitated by the
presence of colipase (a small protein cofactor needed by pancreatic lipase for
efficient dietary lipid hydrolysis), which is also produced by the pancreas. Pancreatic
lipase is specific for the fatty acid residues at positions 1 and 3 of the glyceryl moiety
thus releasing 2-monoacylglycerol and two fatty acids from a triacylglycerol
molecule. The final digestive products of triacylglycerol digestion are free fatty acids,
glycerol and monoacylglycerols.

Triacylglycerol + 2H2O 2-Monoacylglycerol + 2 Fatty acids

After hydrolysis, the products diffuse from the micelle to the intestinal mucosal cell
membrane. The phosphoglycerides present in the diet are digested by pancreatic
phospholipase and cholesteryl esters are hydrolyzed to cholesterol and fatty acids
through the action of cholesteryl esterase.

Cholesteryl esterase
Cholesterol ester Cholesterol + FFA

The action of various phospholipases is presented in Figure 5.5. Phospholipase A1

hydrolyzes fatty acids attached to carbon 1 of the phospholipids, while phospholipase
A2 removes the fatty acid attached to carbon 2. It is found in pancreatic juice and
snake venom. In each case, the product is called lysophospholipid. Phospholipase B
can hydrolyse both the fatty acids (at carbons 1 and 2). Phospholipase C attacks the
ester bond in position 3 and removes phosphate + base, forming 1,2-diacylglycerol.
This enzyme is present in toxins secreted by bacteria. Phospholipase D, found
normally in plants, hydrolyses the nitrogenous base from phospholipids, forming
phosphatidic acid.

Figure 5.6: Action of various phospholipases

Finally, we shall look at the digestion of nucleic acids in our body.

5.4.4 Digestion of Nucleic Acids

Several nucleases present in the pancreatic juice like RNAase and DNAase digest
various nucleic acids. These enzymes are endonucleases which hydrolyze internal
phosphodiester bonds which are located more centrally in the nucleic acid. They form
shorter length chains with 3’-hydroxyl and 5’-phosphoryl or 5’-hydroxyl and 3’-
phosphoryl terminus. Some act on both the strands, whereas, others can hydrolyze
only single strand of nucleic acids. Phosphatases present in the intestinal juice
remove phosphate from hexophosphates, glycerophosphates and nucleotides derived
from the food. Further, the nucleotidases and nucleosidases present in the intestinal
juice hydrolyze it to the level of respective bases as shown in Figure 5.7.

132
 Digestion, Absorption
and Transport
of Carbohydrates,
Proteins and Lipids

Figure 5.7: Digestion of nucleic acids

With this, we end our study on digestion. We move to the absorption process next.
But before that, let us recapitulate what we have learnt till now.

Check Your Progress Exercise 2

1) Name the various enzymes involved in the digestion of carbohydrates.

Enumerate the reactions catalyzed by these.
............................................................................................................................
............................................................................................................................
............................................................................................................................
2) What are the final breakdown products of proteins? Explain giving suitable
reactions involved.
............................................................................................................................
............................................................................................................................
............................................................................................................................
3) Explain the role of lipase in lipid digestion.
............................................................................................................................
............................................................................................................................
............................................................................................................................
4) What are ‘endonucleases’? What is their site of action and end products?
............................................................................................................................
............................................................................................................................
............................................................................................................................

5.5 ABSORPTION AND TRANSPORT

We already know that the movement of substances into or across tissues, in
particular, the passage of nutrients and other substances into the walls of the
gastrointestinal tract and then into the bloodstream is referred to as absorption.

The small intestine is the main digestive and absorptive organ. Most absorption
occurs in the duodenum and jejunum (second third of the small intestine). You may
have already studied the anatomy of small intestine in the Applied Physiology
Course. If not, we suggest you look up Unit 6 of the Applied Physiology Course now.

Transport across the intestine may be active or passive. Active transport requires
energy, whereas passive transport does not. Also, active transport may involve
movement of a substance against a concentration gradient (that is, from a region of
lower to higher concentration), while substances that are passively transported always
move with the concentration gradient. Facilitated diffusion is a type of passive
transport which, unlike simple diffusion, uses a carrier. It is therefore more rapid
than simple diffusion. Active transport mechanisms have been identified for intestinal
absorption of many substances including glucose, galactose, amino acids, calcium,
133
Nutritional Biochemistry iron, folic acid, ascorbic acid, thiamin and bile acids. Fructose, riboflavin and vitamin
B12 (in combination with intrinsic factor) are among the substances absorbed by
facilitated diffusion. Look up Unit 8 in the Applied Physiology Course to know about
active and passive transport across cell-membrane. This information will help you
understand the absorption of different nutrients discussed here in this unit.

We shall study the absorption of different nutrients, namely carbohydrates, proteins

and fats in this section. We start with the absorption of carbohydrates.

5.5.1 Absorption of Carbohydrates

Absorption of carbohydrates is via both passive and active transport. Let us learn
how.

• Passive transport

The end products of carbohydrate digestion are monosaccharides. They are absorbed
from the jejunum into the portal blood system, through which they are transported
first to the liver and then to the rest of the body. Pentoses and fructose pass across the
intestinal barrier by simple passive diffusion and also by facilitated diffusion
involving carrier proteins.

• Active transport

D-glucose and D-galactose are absorbed by an active transport mechanism. The

active transport of glucose is illustrated in Figure 5.8. The transport of glucose is
facilitated by a carrier protein, which has separate binding sites for glucose and
sodium. Both are transported through the plasma membrane and after releasing
sodium and glucose into the cytosol (liquid medium of the cytoplasm), the carrier
protein return to take up fresh load. The sodium is transported down its concentration
gradient and at the same time causes the carrier to transport glucose against its
concentration gradient. The free energy required for this active transport is obtained
from the hydrolysis of ATP linked to sodium pump which expels Na+ from the cell.

The active transport of glucose is inhibited by Ouabain (a cardiac glycoside), an

inhibitor of the sodium pump and by phlorihizin, a known inhibitor of glucose
re-absorption in the kidney tubules.

Figure 5.8: Active transport of glucose

There is also a sodium independent carrier of glucose.

134
5.5.2 Absorption of Proteins Digestion, Absorption
and Transport
of Carbohydrates,
In sub-section 5.3.2, we have seen how the proteolytic enzymes break the dietary Proteins and Lipids
proteins into amino acids or di- and tripeptides. In fact, it is in this form that the
dietary proteins are absorbed. Dietary proteins are, with very few exceptions, not
absorbed. Digestion of proteins and absorption of amino acids goes on throughout
the small intestine. We shall look at the mechanism of absorption of amino acids and
peptides next.

Absorption of amino acids and peptides

Generally, the dietary proteins are almost completely digested to their constituent
amino acids and these are rapidly absorbed from the intestine into the portal blood.
Some of the dipeptides are hydrolyzed by peptidases located in the absorptive cells
so that only amino acids are released into the portal blood. D-amino acids are
absorbed by simple diffusion but the L-amino acids (occurring in foods) require a
carrier system in the absorption.

The mechanism by which amino acids are absorbed is conceptually identical to that
of monosaccharides. The luminal plasma membrane of the absorptive cell bears at
least four sodium-dependent amino acid transporters (one each for acidic, basic,
neutral and aromatic amino acids), as highlighted in Table 5.2. Each system
transports amino acids that are structurally similar. These transporters bind amino
acids only after binding sodium. The fully loaded transporter then undergoes a
conformational change that dumps sodium and the amino acid into the cytoplasm,
followed by its reorientation back to the original form. Thus, absorption of amino
acids is also absolutely dependent on the electrochemical gradient of sodium across
the epithelium. The energy dependent carrier system also involves vitamin B6
(pyridoxal phosphate) during the transport of amino acids.

Apart from amino acids, a substantial amount of small peptides are also absorbed by
stereospecific (determined by structure of amino acid) transport systems. These small
peptides are absorbed without dependence on sodium, probably by a single transport
molecule. The uptake mechanisms for peptides are separate from those for amino
acids.
Table 5.2: Amino acid transport systems

Amino acid specificity Examples of amino acid transported

1 Small neutral amino acids Alanine, serine and threonine
2 Large neutral and aromatic Isoleucine, leucine, valine, tyrosine,
amino acids tryptophan, phenylalanine
3 Basic amino acids Arginine, lysine, ornithine
4 Acidic amino acids Glutamic acid, aspartic acid

We have seen that dietary proteins as such cannot be absorbed. But, absorption of
intact proteins does occur only in a few circumstances. What are these
circumstances? Let’s find out.
Absorption of proteins in infants
In infants, permeability of the intestine appears to be greater than in later life, and
some large protein molecules such as antibodies in the mother’s milk and protein
allergens pass by diffusion through the wall. Pinocytosis appears to account for the
absorption of these large molecules in children and adults. Pinocytosis, as you may
recall reading in the Applied Physiology Course, is a process by which certain cells
can engulf and incorporate droplets of fluid. Look up Unit 2, sub-section 2.3.1 in the
Applied Physiology Course for understanding this process.

Next, let us look at the absorption of lipids.

135
Nutritional Biochemistry 5.5.3 Absorption of Lipids

Lipids are absorbed by a mechanism distinctly different from what we have seen for
monosaccharides and amino acids. In fact absorption of dietary triacylglycerols and
medium chain triacylglycerols in the food also varies. Let us get to know this process
in greater detail.

Absorption of long-chain triacylglycerols

The triacylglycerols present in the ordinary food that we eat contains long chain fatty
acids (mostly 16 and 18 carbon-atom saturated and unsaturated fatty acids).
Pancreatic lipase specifically acts upon the fatty acid residues at positions 1 and 3 of
the glyceryl moiety thereby producing 2-monoacylglycerols and the released fatty
acids can pass through cell membrane, and they are absorbed by diffusion into the
mucosal cells of the jejunum and ileum. There is resynthesis of triacylglycerol moiety
inside the mucosal cells and this reformed triacylglycerol is coated with protein,
cholesterol or phospholipid to form tiny globules called chylomicrons as shown in
Figure 5.9. Instead of being absorbed directly into capillary blood, these
chylomicrons are transported first into the lymphatic vessel that penetrates into each
villus. Chylomicron-rich lymph then drains into the system lymphatic system, which
rapidly flows into blood. Blood-borne chylomicrons are rapidly disassembled and
their constitutent lipids utilized throughout the body.

Absorption of medium-chain triacylglycerols

Medium chain triacylglycerols with fatty acids containing less than 10 to 12 carbons
are absorbed intact either onto the mucosal cell villi or into the cells and are
hydrolyzed to free fatty acids and glycerol by the lipase present inside the cell. The
released fatty acids pass directly into the portal vein. Plasma albumin acts as a carrier
for these fatty acids and delivers them to the liver as fatty acids by portal circulation.

The difference in digestion and absorption of dietary triacylglycerols and medium

chain triacylglycerols is illustrated in Figure 5.9.

Figure 5.9: Difference in digestion and absorption of dietary triacylglycerols and

medium chain triacylglycerols

136
Another lipid of importance that is absorbed in the small intestine is cholesterol. Digestion, Absorption
A specific transport protein has been identified that ferries cholesterol from the and Transport
of Carbohydrates,
intestinal lumen into the enterocyte. From there, cholesterol is incorporated into Proteins and Lipids
chylomicrons and shuttled into blood by the mechanisms described above.

So we have looked at how the fats are absorbed in the body. Now the fats (lipids)
absorbed from the diet must be transported in the blood. Blood plasma is a watery
environment. We also know that lipids are insoluble in water. So how are they
transported in blood? Well, fats are insoluble in water and so lipid compounds such
as cholesterol, fatty acids, oil soluble vitamins and triglycerides need to be associated
with proteins, forming water soluble lipoproteins, in order to be transported around
the body.

Most lipids are transported in the blood as triacylglycerols within lipoprotein

(particles containing a core of hydrophobic lipids surrounded by a shell of proteins,
phospholipids and cholesterol). Four major groups of lipoproteins help in transport of
lipids. These include:

1. Chylomicrons-function is to carry dietary triacylglycerols

2. Very low density lipoprotein (VLDL)-function is to carry endogenously made

triacylglycerols

3. Low density lipoprotein (LDL)-function is to carry cholesterol and cholesterol

esters, and

4. High density lipoprotein (HDL)-functions in reverse cholesterol transport and

exchange of apoproteins.

We shall learn more about lipoproteins later in Unit 7 under lipoprotein metabolism.
With this, we come to an end of our study of digestion, absorption and transport of
carbohydrates, lipid and proteins.

Check Your Progress Exercise 3

1) List the various transport mechanisms involved in the absorption of various

nutrients in the intestine, giving suitable examples.
............................................................................................................................
............................................................................................................................
............................................................................................................................
2) How are carbohydrates absorbed in our body?
............................................................................................................................
............................................................................................................................
............................................................................................................................
3) Explain the active transport of glucose.
............................................................................................................................
............................................................................................................................
............................................................................................................................
4) Explain the process of absorption of amino acids and peptides.
............................................................................................................................
............................................................................................................................
............................................................................................................................

137
Nutritional Biochemistry 5) Bring out the major points of difference in the digestion and absorption of
dietary triacylglycerols and medium chain triacylglycerols.
............................................................................................................................
............................................................................................................................
............................................................................................................................
6) How are lipids transported in blood? List the four major groups of lipoproteins
involved in lipid transport.
............................................................................................................................
............................................................................................................................
............................................................................................................................

5.6 LET US SUM UP

In this unit we studied about the digestion, absorption and transport of carbohydrates,
fats and proteins in our body. The salient features discussed include:

1. The digestive system takes in food, digests it to the end products, which are
absorbable into the system. The non-absorbable products are eliminated.
2. The ingested food is broken down into end products through mechanical,
chemical and enzymatic digestion.
3. Digestion involves the participation of mouth, esophagus, stomach, intestine,
liver, pancreas and gall bladder.
4. Digestion also involves the participation of a number of enzymes viz.,
amylase, pepsin, trypsin, chymotrypsin, endopeptidases, aminopeptidases,
carboxypeptidases, nucleases and lipases, which break down the dietary
carbohydrates, proteins and lipids into simple units.
5. Control of pancreatic, liver and intestinal secretion is by hormones.
6. Most of the above enzymes are secreted in the form of pro-enzymes and get
converted into active form just before their action.
7. The end products are absorbed into the system through various transport
mechanisms such as active, passive mechanism etc.

5.7 GLOSSARY

Absorption : taking in the molecules from the GI tract.

Active transport : using carriers, energy (ATP) and enzymes of cell to cause a
substance to cross a membrane.
Bile : a clear yellow or orange fluid produced by the liver.
Brush border : a specialization of the free surface of a cell, consisting of
minute cylindrical processes (microvilli) that greatly
increase the surface area.
Chyme : the thick semi-fluid mixture of partially digested foods and
digestive juices found in the stomach and small intestine.
Dextrins : polysaccharides containing fewer simple sugar units than
starches.
Digestion : breaking of foods using chemical or mechanical means to
convert foods into chemical compounds, which could be
absorbed.
Facilitated : a type of passive transport that uses a carrier.
diffusion
Micelle : a submicroscopic aggregation of molecules, as a droplet in
138
 a colloidal system. Micelles are essentially small aggregates Digestion, Absorption
of mixed lipids and bile salts suspended within the ingesta. and Transport
of Carbohydrates,
Pinocytosis : the ingestion of dissolved materials by endocytosis. The Proteins and Lipids
cytoplasmic membrane invaginates and pinches off placing
small droplets of fluid in a pinocytic vesicle. The liquid
contents of the vesicle are then slowly transferred to the
cytosol.
Serum : spherical or ellipsoidal particles containing proteins,
lipoproteins cholesterol esters and triacylglycerols, encased within a
monolayer of phospholipids and cholesterol.
Zymogen : an inactive form of an enzyme; becomes active prior to its
action.

5.8 ANSWERS TO CHECK YOUR PROGRESS

EXERCISES

Check Your Progress Exercise 1

1). The process through which the major nutrients namely carbohydrates, proteins
and lipids are converted to simple sugars, amino acids, fatty acids and glycerol
respectively, while passing from mouth to small intestine, is referred to as
digestion. Microvilli of the small intestine is the major site of absorption.

2) The various organs involved in the digestion process are:

• Alimentary tract which includes mouth, pharynx, esophagus stomach, small
intestine, large intestine, appendix, rectum and anal canal, and
• Accessory organs which include tongue, teeth, salivary glands, liver, gall
bladder and pancreas.

3) Saliva is the colourless viscous fluid. It helps in swallowing the food by

lubricating the food and the neutral pH of saliva prevents the decalcification of
teeth and keeps the mouth clean. In the mouth, is also present the salivary
amylase. The presence of salivary amylase starts the starch digestion.

4) The digestive process occurring in the stomach is discussed as:

The stomach secretes gastric juice. The major components of gastric juice are
hydrochloric acid and the enzyme pepsin which is secreted in an inactive form
called pepsinogen. Hydrochloric acid (pH 1.2 – 1.5) activates the inactive
pepsinogen to the active form pepsin. Hydrochloric acid denatures the food
proteins for easy digestion. The pepsin produced in an active proteolytic enzyme
that activates the remainder of the pepsinogen molecule by proteolytically
converting them to pepsin. Digestion of protein is initiated in the stomach by
pepsin. Pepsin acts upon food proteins and converts them to proteoses and
peptones.
In infants, the presence of another enzyme called rennin helps in digestion of
casein to paracasein which is then acted upon by pepsin.
5) Pancreatic juice is alkaline in nature (pH about 8.0) and contains both organic
and inorganic substances. The enzyme components are trypsin, chymotrypsin,
elastase and carboxypeptidase. The other enzymes are amylase, lipase,
cholesteryl ester hydrolase and phospholipases. Sodium, chloride and bicarbonate
are the major inorganic constituents.
6) Bile salts have a considerable ability to lower surface tension. By this property,
they prevent the coming together of the small fat droplets in the intestine and thus
allow more rapid digestion of the fat. This is called the emulsification action of
bile.

139
Nutritional Biochemistry 7). The enzymes of intestinal juice are aminopepetidase and dipeptidase.

Check Your Progress Exercise 2

1) Salivary amylase, pancreatic amylase, maltase, isomaltase, sucrase and lactase

are the enzymes involved in the digestion of carbohydrates. The reactions
catalyzed by these are:

Salivary amylase
Starch dextrins + glucose + maltose

Pancreatic amylase
Starch/dextrins maltose + isomaltose + α- limit dextrin

Maltase
Maltose glucose + glucose

Isomaltase
Isomaltose glucose + glucose

Sucrase
Sucrose fructose + glucose

Lactase
Lactose galactose + glucose

2) The final breakdown products of protein digestion are single amino acids or
small chains of two or three amino acids.

3) Pancreatic lipase acts on the dietary triacylglycerol after it has been incorporated
into the mixed micelle in the intestinal lumen. The lipase acts at interface
between water and the triacylglycerol molecules, and its catalytic action is
facilitated by the presence of colipase. Pancreatic lipase is specific for the fatty
acid residues at positions 1 and 3 of the glyceryl moiety thus releasing
2-monoacylglycerol and two fatty acids from the triacylglycerol molecule. The
final digestive products of triacylglycerol digestion are free fatty acids, glycerol
and monoacylglycerols.

4) Endonucleases are nucleases present in the pancreatic juice like RNAase and
DNAase. These hydrolyze internal phosphodiester bonds which are located more
centrally in the nucleic acid. Phosphatases present in the intestinal juice remove
phosphate from hexophosphates, glycerophosphates and nucleotides derived
from the food. Further, the nucleotidases and nucleosidases present in the
intestinal juice hydrolyze it.

140
Check Your Progress Exercise 3 Digestion, Absorption
and Transport
of Carbohydrates,
1) The various transport mechanisms involved in the absorption of various nutrients Proteins and Lipids
are active transport (glucose and galactose), passive transport (pentose and
fructose) and facilitated diffusion (fructose and riboflavin).

2) Carbohydrates are absorbed in our body via both active and passive transport.

3) The transport of glucose is facilitated by a carrier protein, which has separate

binding sites for glucose and sodium. Both are transported through the plasma
membrane and after releasing sodium and glucose into the cytosol (liquid
medium of the cytoplasm), the carrier protein returns to take up fresh load. The
sodium is transported down its concentration gradient and at the same time
causes the carrier to transport glucose against its concentration gradient. The free
energy required for this active transport is obtained from the hydrolysis of ATP
linked to sodium pump which expels Na+ from the cell.

4) The luminal plasma membrane of the absorptive cell bears at least four sodium-
dependent amino acid transporters (one each for acidic, basic, neutral and amino
acids). Each system transports amino acids that are structurally similar. These
transporters bind amino acids only after binding sodium. The fully loaded
transporter then undergoes a conformational change that dumps sodium and the
amino acid into the cytoplasm, followed by its reorientation back to the original
form. Thus, absorption of amino acids is also absolutely dependent on the
electrochemical gradient of sodium across the epithelium. The energy dependent
carrier system also involves vitamin B6 (pyridoxal phosphate) during the
transport of aminoacids.

Apart from amino acids, a substantial amount of small peptides are also absorbed
by stereospecific (determined by structure of amino acid) transport systems.
These small peptides are absorbed without dependence on sodium, probably by a
single transport molecule.

5) The major points of difference in the digestion and absorption of dietary

triacylglycerols and MCTs can be depicted as:

141
Nutritional Biochemistry 6) Most lipids are transported in the blood as triacylglycerols within lipoprotein.
Four major groups of lipoproteins help in transport of lipids. These include
chylomicrons, very low density lipoprotein, low density lipoprotein and high
density lipoprotein.

142