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pp. 351–371. Amsterdam: Elsevier Science Publishers.
CARBOHYDRATES
Contents
Chemistry and Classification
Regulation of Metabolism
Requirements and Dietary Importance
Resistant Starch and Oligosaccharides
Chemistry and Classification
C L Stylianopoulos, Johns Hopkins University,
Baltimore, MD, USA
ª 2005 Elsevier Ltd. All rights reserved.
Introduction
Carbohydrates are the most abundant constituents
of cereals, fruits, vegetables, and legumes. They are
the major energy source in human nutrition and
contribute to the texture and flavor of processed
foods. They comprise a group of substances with
different structures and varying physical, chemical,
and physiological properties. Carbohydrates are
polyhydroxy aldehyde or ketone molecules and
their derivatives with the general formula (CH2O)n.
Dietary carbohydrates are important in maintaining
glycaemic homeostasis and gastrointestinal health.
Furthermore, they contain necessary micronutrients,
phytochemicals, and antioxidants.
Classification and Chemical Structure
Carbohydrates are classified into four categories
according to their chemical structure and degree of
polymerization: monosaccharides, disaccharides, oli-
gosaccharides, and polysaccharides.
Monosaccharides
Monosaccharides are the simplest form of carbohy-
drate and cannot be further hydrolyzed into smaller
subunits. According to their chain length, the
Williams GM and Weisburger JH (1991) Chemical carcinogen-
esis. In: Amdur MO, Doull J, and Klassen CD (eds.) Casarett
and Doull’s Toxicology. The Basic Science of Poisons, 4th
edn., pp. 127–200. New York: Pergamon Press.
World Health Organisation (1999) Principles for the assessment
of risks to human health of exposure to chemicals. Environ-
mental Health Perspectives, No. 210.
monosaccharides can be divided into several cate-
gories, the more nutritionally important being the
pentoses, with skeletons containing five carbon
atoms (e.g., ribose), and the hexoses, with skeletons
containing six carbon atoms (e.g., glucose).
The presence of asymmetrical carbon atoms in
monosaccharides with different functional groups
attached gives rise to optical activity, meaning that if
polarized light is passed through a solution of these
compounds, the plane of light will be rotated to the left
(levorotatory or L form) or to the right (dextrorotatory
or D form). As a result, mirror-imaged structures of the
same compound exist and are called stereoisomers.
Monosaccharides of the D form are more nutritionally
important because the metabolic and digestive
enzymes are specific for the D stereoisomers.
Monosaccharides demonstrate another type of
stereoisomerism due to their formation of cyclic struc-
tures. The pentoses form furanose (five-membered
rings), and the hexoses form pyranose (six-membered
rings). Cyclization can produce two stereoisomers (the
 and
configurations), and generally monosacchar-
ide solutions contain an equilibrium mixture of these
two forms. Figure 1 illustrates D-glucose in its pyranose
form in the  and
configurations. The isomerization
produces compounds with different properties and has
major metabolic importance, because of enzyme spe-
cificity for particular stereoisomers.
Glucose is the most abundant monosaccharide,
and is a major cell fuel in the human body, and
can be found unbound in body tissues and fluids.
Glucose is the building block of several polysacchar-
ides. Galactose and fructose are also used as cell
304 CARBOHYDRATES/Chemistry and Classification

H O
1 C
2
H OH
3 6C H2OH 6C H2OH
H H
5 O H 5 O OH
4 4 OH
H OH 1 + 4 1
OH
5
HO 3 2 OH HO 3 2 H
H OH
6
OH OH
CH2OH
D-glucose α-D-glucose β-D-glucose
(A) (B)
Figure 1 D-glucose molecule shown as (A) open chain and (B) a cyclic pyranose ring in the  and
configurations.

fuel. The most important monosaccharides and their Disaccharides

significance are outlined in Table 1.
Disaccharides consist of two monosaccharide units,
Several monosaccharide derivatives are constitu-
linked by glycosidic bonds in the  or
orientation.
ents of polysaccharides, as well as food ingredients.
The most important disaccharides are sucrose, lactose,
Some nutritionally important monosaccharide deriva-
and maltose. Sucrose consists of a molecule of -glu-
tives and their significance are outlined in Table 1.
cose and a molecule of
-fructose linked together
(Figure 2A). Lactose is found in milk and dairy pro-
ducts and consists of a molecule of galactose linked to
Table 1 Some nutritionally important monosaccharides a glucose molecule by a
-1,4glycosidic bond
Class Species Significance

Hexoses D-glucose Major cell fuel, unbound CH2OH

in body fluids and O
tissues, building block HO CH2OH
O
of several HO
polysaccharides HO OH
D-fructose Cell fuel, constituent of O
sucrose CH2OH
D-galactose Cell fuel, constituent of OH
lactose
D-mannose Constituent of plant cell (A)
wall polysaccharides
and gums OH
Pentoses L-arabinose, Constituent of plant cell CH2OH
CH2OH
D-xylose wall polysaccharides O O O
D-ribulose, Metabolite in pentose
D-xylulose pathway HO
HO
D-ribose RNA constituent HO
HO
Uronic acids D-glucuronic, Constituent of plant cell OH
D-galacturonic wall polysaccharides (B)
D-mannuronic, Constituent of algal
D-guluronic polysaccharides CH2OH
Sugar D-glucitol, D-xylitol Food ingredient O
alcohols D-galactitol Metabolite of galactose HO
Deoxysugars D-deoxyribose DNA constituent HO
D-deoxygalactose Constituent of algal CH2OH
HO
polysaccharides O O
L-fucose Constituent of bacterial
HO
polysaccharides
L-rhamnose Constituent of pectic plant HO
OH
polysaccharides
Aminosugars D-glucosamine, Constituent of (C)
D-galactosamine aminosaminoglycans,
cartilage Figure 2 The molecular structures of (A) sucrose, (B) lactose,
and (C) maltose.
 CARBOHYDRATES/Chemistry and Classification 305

Table 2 Some nutritionally important disaccharides compounds consist of several hundred or even thou-
sands of monosaccharide units. The properties of
Class Species Significance
polysaccharides are determined by the species of
Disaccharide Sucrose Constituent of fruits, monosaccharides in the polymer backbone, the
vegetables, and sweetener types of linkages between residues, and the extent
Lactose Constituent of milk and dairy and type of chain branching.
products
Maltose, Constituent of starch
Glucans are polymers of glucose and the major
Isomaltose polysaccharides in the diet. The most important
Trehalose Food additive, constituent of glucans are starch, glycogen, and cellulose. Glyco-
mushrooms gen is the short-term storage form of glucose in
Lactulose Lactose derivative, laxative animal tissues. Starch is the most common digesti-
Disaccharide Maltitol Constituent of starch,
alcohols Lactitol sweetner
ble storage polysaccharide in plants, and cellulose
Constituent of lactose, is a major structural component of plant cell walls
sweetener (Figure 3). Some nutritionally important polysac-
charides and their significance are outlined in
Table 4.
(Figure 2B). Maltose is mainly produced by partial Polysaccharides with  linkages have a helical shape
hydrolysis of starch and consists of two glucose units (e.g., the amylose starch molecule), while those with

linked through an -1,4glycosidic bond (Figure 2C). linkages generally have a linear or flat ribbon-like
Some nutritionally important disaccharides and their molecule (e.g., the cellulose molecule) (Figure 3).
significance are outlined in Table 2. Polysaccharide molecules can be linear or
branched. Branches can be formed through any
Oligosaccharides unlinked hydroxyl group and vary from alternating
Oligosaccharides consist of a chain of between three and consecutive single-unit branches to multiple-
and nine monosaccharide units covalently linked to unit branches (ramified structure). Polysaccharides
form large units and are named trioses, tetroses, etc, that are highly branched tend to be soluble in
depending on the number of carbon atoms in their water, because the chain structure prevents hydro-
molecules. Oligosaccharides are distributed widely gen bonding. Linear polysaccharides tend to be
in plants and when digested yield their constituent insoluble in water, unless they possess structural
monosaccharides. The major oligosaccharides are irregularities.
the raffinose series, formed by the linkage of galac-
tose, sucrose, and glucose units, and the maltose
series, formed by the linkage of glucose units. Nutritional Importance
Some nutritionally important oligosaccharides and Some carbohydrate types of specific importance in
their significance are outlined in Table 3. human nutrition are sugars and sugar alcohols,
starch, and dietary fiber.
Polysaccharides
Polysaccharides consist of long chains of monosac- Sugars
charide residues (more than nine) linked with glyco-
The Food and Agriculture Organization (FAO) and
sidic bonds. Polysaccharides found in nature
the World Health Organization (WHO) expert con-
usually have high molecular weights and are
sultation on carbohydrates use the term ‘sugar’ to
named after the component monosaccharide. These
describe monosaccharides and disaccharides. Sugars
can be separated analytically from the food matrix
Table 3 Some nutritionally important oligosaccharides
by gas–liquid chromatography (GLC), high perfor-
Class Species Significance mance liquid chromatography, and enzymatic
methods. Sugars are widely used in the food industry as
Maltoses Maltotriose, Constituent of starch
sweeteners and preservatives. They improve the tex-
Maltotetraose
Raffinoses Raffinose, Constituent of
ture, body, palatability, and viscosity of foods and
Stachyose, vegetables and beverages.
Verbascose legumes The UK Department of Health distinguishes
Fructoses Fructotriose Constituent of between ‘intrinsic’ and ‘extrinsic’ sugars. Intrinsic
cereals and tubers sugars are defined as those that occur naturally
Lactoses Fucosyl lactoses Constituent of
human breast milk as part of the plant cell walls. Extrinsic sugars were
defined as added sugars or those present when the
306 CARBOHYDRATES/Chemistry and Classification

CH2OH
HO O

HO
HO CH2OH
O O

HO
HO CH2OH
O O

HO
HO CH2OH
O O

HO
HO CH2OH
O O

HO
HO
O

(A)

CH2OH CH2OH CH2OH CH2OH

HO O O O O
O O O O
HO HO HO HO
HO HO HO HO

(B)
Figure 3 (A) Five units of an -1,4-D-glucopyranose chain from a starch molecule (amylose). (B) Four units of a
-1,4-D-glucopyranose
chain from a cellulose molecule.

food matrix has been disrupted. An additional term, can be found in cereals, potatoes, legumes, and
‘non-milk’ extrinsic sugar, is used to distinguish other vegetables, with amylopectin comprising
between milk and other extrinsic sugars. However, 80–85% and amylose 15–20% of total starch.
these terms have not been widely accepted.

Sugar Alcohols
Sugar alcohols are monosaccharide and disaccharide
derivatives, such as sorbitol and xylitol, which are
extensively used as sweeteners in the food industry.
They have received increased attention because of
their desirable properties of relative sweetness and
limited digestion and absorption.
Starch
The most important, abundant, and digestible poly-
saccharide in human nutrition is starch. Starch com-
prises large chains of -linked glucose residues, in
the form of amylose or amylopectin. Amylose is a
linear unbranched form of starch, which consists of
-1,4-linked glucose units (Figure 3A). Amylopectin
is a branched-chain polymer, which consists of
-1,6-linked glucose units. Both forms of starch
Dietary Fiber
There are several definitions of fiber, and no
consensus exists among international organizations.
The Association of Official Analytical Chemists
International defines fiber as nondigestible animal
and plant carbohydrates, based on the analytical
methods for fiber separation using an enzymatic–
gravimetric method (Table 5).
According to the new definition of the expert
panel on macronutrients appointed by the Insti-
tute of Medicine of the National Academies of
Science (Table 5), dietary fiber comprises intact
nondigestible carbohydrates and lignin derived
from plant sources. Functional fiber consists of
nondigestible carbohydrates, derived from either
plant or animal sources, that have shown favor-
able health outcomes for humans. Total fiber con-
sists of both dietary and functional fiber. With
this new definition of dietary and functional
 CARBOHYDRATES/Chemistry and Classification 307

Table 4 Some nutritionally important polysaccharides Chemistry

Class Species Significance Monosaccharides share the same functional groups,
but their isomeric forms often exhibit differences in
Glucans Starch Storage polysaccharide in
plants
chemical reactions. Disaccharides exhibit a similar
Glycogen Short-term storage form of range of reactions to monosaccharides owing to
glucose in animal tissues the presence of similar functional groups. Oligosac-
Cellulose Major structural component charides generally exhibit properties similar to
of plant cell walls those of monosaccharides and disaccharides with
Galactans Major constituents of
noncellulosic matrix of
similar functional groups, but some oligosacchar-
plant cell wall ides with nine monosaccharide units may exhibit
Xylans Constituents of mature plant similar properties to polysaccharides. In general,
tissues polysaccharides show slower reaction rates because
Mannans Storage forms in several of steric effects.
plants
Uronans Galacturonans Major components of water-
soluble pectic fraction of Solubility
plants
Mannuronans Components of algal Monosaccharides, disaccharides, and oligosaccharides
polysaccharides have similar solubilities. Overall, they are very soluble
Guluronans Components of algal in water. Sucrose is extremely soluble in water, while
polysaccharides lactose is soluble to a lesser extent. Furthermore, they
Starch Amylose, Most common digestible
amylopectin plant polysaccharides
are insoluble in nonpolar organic solvents. They exhi-
Nonstarch Cellulose Major component of plant bit limited solubility in pure alcohols but are very
cell wall soluble in aqueous alcohol solutions (70–80% v/v),
Pectin Constituent of plant cell wall, and therefore these solutions are widely used for
food additive extraction and analysis. Oligosaccharides are less
Hemicellulose Constituent of plant cell wall
Gums, mucilages Plant hydrocolloids, food
soluble than monosaccharides in aqueous alcohol
additives solutions, and their solubility decreases as the number
Algal Constituents of algae and of monosaccharide units increases.
polysaccharides seaweed, food additives In general, polysaccharides form colloidal solu-
tions in water, while some other polymers are extre-
mely insoluble in water and require prior treatment
fiber, new analytical methods should be developed with acid, alkali, or organic solvents to get them to
and implemented to quantify accurately the total dissolve. For example,
-1,4-mannans and glucans
fiber component of foods. (e.g., cellulose) are very insoluble owing to hydrogen
Dietary and functional fiber cannot be digested bonding between parallel chains. On the other hand,
by mammalian enzymes, and therefore they pass arabinoxylans are readily soluble in water, because
almost intact through the small intestine. Fiber the arabinosyl chains inhibit hydrogen bonding.
consumption has potential health benefits, includ- Galactomannans are also readily soluble in water,
ing the promotion of general gastrointestinal health producing viscous solutions, and are used as food
and the prevention of several noncommunicable additive gums. The -linked glucans (e.g., amylose
diseases. and amylopectin) have completely different solu-
bilities. The glucan -1,4-amylose is very soluble
in warm water and forms colloidal solutions.
Table 5 Current definitions of dietary fiber When the amylose chains cool down, they form
an amylose gel, which subsequently forms an inso-
Source Definitions luble crystalline material. Amylopectins are also
AOACa Fiber: nondigestible animal and plant carbohydrates very soluble in hot water but do not form an
IOMb Dietary fiber: intact nondigestible carbohydrates and insoluble crystalline material to the same degree
lignin, derived from plant sources as amylose.
Functional fiber: nondigestible carbohydrates derived
from either plant or animal sources that have
shown favourable health outcomes for humans Reducing Properties
Total fiber: dietary and functional fiber
Monosaccharides are powerful reducing agents to a
a
AOAC: Association of Official Analytical Chemists range of metals in alkaline solution, owing to the
b
IOM: Institute of Medicine presence of aldo and keto groups. The extent of
308 CARBOHYDRATES/Chemistry and Classification
reduction varies among different monosaccharides.
Disaccharides and oligosaccharides have the same
reducing properties, except for sucrose, in which
both hemi-acetal groups are combined. Polysacchar-
ides usually contain one reducing group at the term-
inal end of the polymer chain and, as a result, have
lower reducing properties.
Reactions in Acidic Solutions
When heated in strong acidic solutions, monosac-
charides dehydrate and condense into a range of
furans. The resulting furans condense with several
reagents to generate colored products; hence the
presence of monosaccharides and their derivatives
can be verified. Under weaker acidic conditions,
fructose is labile.
Reactions in Alkaline Solutions
In weak alkaline solutions, monosaccharides undergo
isomerization of the aldose-keto group (enolization).
In stronger alkaline solutions, they produce a series of
degradation compounds, namely saccharinic acids. In
the presence of ammonia, amino acids, and proteins,
they condense repeatedly to generate a series of highly
colored products (Maillard reaction); this reaction is
used in the food industry to produce caramel colors.
Hydrolysis
Acid Disaccharides and oligosaccharides in mild
acidic conditions are hydrolyzed to their constituent
monosaccharides. The fructofuranosyl linkages of the
fructooligosaccharides are quite susceptible to acid
hydrolysis. Polysaccharides are also hydrolyzed to
their constituent monosaccharides by acid hydrolysis,
but the conditions necessary for complete hydrolysis
depend on the solubilities of the polymers. The major-
ity of polysaccharides (e.g., starch) are completely
hydrolyzed under weak acidic conditions. However,
cellulose requires treatment with strong acid for sev-
eral hours prior to hydrolysis and subsequent heating
under weak acidic conditions for the completion of
the reaction. The uronans are very resistant to com-
plete acid hydrolysis, and disaccharides of aldobiuro-
nic acids are generally produced. Acid hydrolysis of
polysaccharides results in extensive losses of their
monosaccharide constituents.
Enzymatic Disaccharides are hydrolyzed in specific
enzymatic solutions, and, therefore, this is a useful
method for the analysis of sugar mixtures. Oligosac-
charides are also susceptible to enzymatic hydroly-
sis. The maltooligosaccharides can be rapidly
hydrolyzed by glucosidase enzymes.
Polysaccharides are more efficiently hydrolyzed to
their monosaccharide constituents using specific
enzymes. Fungal enzymes act specifically to hydro-
lyze different polysaccharides. The -1,4 glycosidic
linkages in starch can be hydrolyzed by various 
amylases (e.g., salivary and pancreatic), producing
maltose and isomaltose. The
-1,6 glycosidic lin-
kages in amylopectin are not as easily hydrolyzed
and require the presence of pullulanase – a fungal
enzyme – to complete the hydrolysis.
Ester Formation
Monosaccharides contain hydroxyl groups and react
with acids to form a variety of esters. The phosphate
esters play a main role in carbohydrate metabolism.
For example, the first step of glycolysis involves the
production of the glucose-6-phosphate ester in a
reaction catalyzed by the enzyme glucokinase in
the presence of adenosine triphosphate. The uronic
acids react with alcohols to form esters. The methyl
esters of uronic acids are the most important in
determining the physical properties of the uronans.
The presence of additional hydroxyl groups in dis-
accharides and oligosaccharides increases the number
of sites for esterification reactions. Sucrose reacts
with fatty acids to produce nondigestible esters,
which have similar properties to the triacylglycerols.
The polysaccharide galacturonans, which are
composed of an -1,4 galacturonic acid chain with
integrated rhamnose units, form salts with cations
and may be esterified with methoxyl groups.
Substitution
Monosaccharides undergo substitution reactions with
methyl iodide to produce methyl ether derivatives.
These compounds have been used to identify the struc-
ture of polymers, because the sites of nonmethyl sub-
stituted groups are indicative of the branch points after
hydrolysis. Monosaccharides undergo acetylation,
which occurs on the free or the reduced molecule to
produce acetylated alditols. These volatile compounds
have been used to identify sugar mixtures by GLC. The
presence of additional hydroxyl groups in disacchar-
ides and oligosaccharides increases the number of sites
for substitution reactions.
Abbreviations
D dextrorotatory
FAO Food and Agriculture Organization
GLC gas–liquid chromatography
L levorotatory
v/v volume/volume
WHO World Health Organization

See also: Alcohol: Absorption, Metabolism and
Physiological Effects. Carbohydrates: Regulation of
Metabolism; Requirements and Dietary Importance;
Resistant Starch and Oligosaccharides. Colon: Structure
and Function. Dietary Fiber: Physiological Effects and
Effects on Absorption. Fructose. Galactose. Glucose:
Chemistry and Dietary Sources. Glycemic Index. Small
Intestine: Structure and Function. Stomach: Structure
and Function. Sucrose: Nutritional Role, Absorption and
Metabolism.
Further Reading
Brody T (1999) Nutritional Biochemistry, 2nd edn, pp. 1–56.
San Diego: Academic Press.
Eastwood M (2003) Principles of Human Nutrition, 2nd edn,
pp. 195–122, 418–426, 486–509. Oxford: Blackwell.
FAO/WHO (1998) Carbohydrates in human nutrition. Report of
a Joint FAO/WHO Expert Consultation. FAO. Food and
Nutrition Paper 66: 1–140.
Gray GM (2000) Digestion and absorption of carbohydrate. In:
In: Biochemical and Physiological Aspects of Human Nutrition,
1st edn, pp. 91–106. Philadelphia: WB Saunders Company.
Groff JL and Gropper SS (2000) Advanced Nutrition and Human
Metabolism, 3rd edn, pp. 70–105. Belmont: Wadsworth.
Institute of Medicine of the National Academies (2002) Dietary
Reference Intakes for Energy, Carbohydrate, Fiber, Fat, Fatty
Acids, Cholesterol, Protein, Amino Acids. Washington, DC:
The National Academies Press.
Lewis BA (2000) Structure and properties of carbohydrates. In:
In: Biochemical and Physiological Aspects of Human
Nutrition, 1st edn, pp. 3–22. Philadelphia: WB Saunders
Company.
Nantel G (2003) Glycemic carbohydrate: an international
approach. Nutrition Reviews 61: S34–S39.
Sanchez-Castillo CP, Hudson GJ, Englyst HN, Dewey P, and
James WPT (2002) The importance of dietary carbohydrates.
Archivos Latinoamericanos de Nutricion 52: 321–335.
Sullivan DM and Carpenter DM (eds.) (1993) Methods of Anal-
ysis for Nutritional Labeling. Arlington: Association of Offi-
cial Analytical Chemists.
Williams SR and Schlenker ED (2003) Essentials of Nutrition and
Diet Therapy, 8th edn, pp. 47–65. St Louis: Mosby.
Regulation of Metabolism
C L Stylianopoulos, Johns Hopkins University,
Baltimore, MD, USA
ª 2005 Elsevier Ltd. All rights reserved.
Introduction
The three basic monosaccharides important in human
nutrition are glucose, fructose, and galactose. Glucose
is the product of the digestion of starch. In human
metabolism, all simple sugars are converted into
CARBOHYDRATES/Regulation of Metabolism 309
glucose. Glucose is the circulating form of carbohy-
drate in the bloodstream. Fructose is the sweetest of
the simple sugars, and it is found in fruits and naturally
occurring substances such as honey. Fructose con-
sumption has increased greatly in the USA since the
1970s, when high-fructose corn syrup started to be
widely used in food processing. High-fructose corn
syrup is the major sweetening agent used by the food
industry. Galactose is produced by the digestion of
lactose, the major carbohydrate in milk.
Digestion
Most carbohydrates have to be converted to glucose in
order to be used for energy production. The digestion
of carbohydrates starts in the mouth, with mastication
and the enzymatic action of salivary amylase, which
converts starch to dextrins and maltose. Successive
contractions of the stomach (peristalsis) move the
food to the lower part of the stomach, while 20–30%
of the carbohydrate is already converted to maltose.
Peristalsis facilitates digestion in the small intestine,
while the chemical digestion of carbohydrates is com-
pleted by pancreatic amylase (which continues the
breakdown of starch to maltose) and intestinal disac-
charidases (sucrase, lactase, and maltase for the break-
down of fructose, lactose, and maltose, respectively).
The monosaccharide products of carbohydrate diges-
tion are then absorbed into the portal circulation.
Absorption
Glucose accounts for the largest quantity of
absorbed carbohydrate (80%), and galactose and
fructose account for only a small amount (20%).
The body quickly absorbs and transports the simple
sugars, which enter the portal circulation via the
capillaries of the intestinal villi and are transported
to the liver. In the liver, fructose and galactose are
converted to glucose, which is either used immedi-
ately for energy or stored in the form of glycogen.
The liver can store approximately 5% of its mass in
the form of glycogen, which can be readily con-
verted to glucose for the production of energy.
Transport
Monosaccharides traverse the epithelial lining of the
intestine by simple or facilitated diffusion or by active
transport. The transport system for the passage of
glucose and galactose through the apical membrane
of the intestinal villi is called the Naþ -dependent
glucose transporter (Naþ -dependent GLUT). Fructose
uses a different transporter, called GLUT5, for the
same passage. All monosaccharides are then trans-
ported from the enterocyte to the bloodstream by
another sugar transporter known as GLUT2. The