Polysaccharides are polymeric carbohydrate molecules composed of long chains of monosaccharide
units bound together by glycosidic linkages and on hydrolysis give the constituent monosaccharides
or oligosaccharides.
They range in structure from linear to highly branched.
Examples include storage polysaccharides such as starch and glycogen, and structural polysaccharides
such as cellulose and chitin.
Polysaccharides are often quite heterogeneous, containing slight modifications of the repeating
unit.
Depending on the structure, these macromolecules can have distinct properties from their monosaccharide
building blocks.
They may be amorphous or even insoluble in water.
When all the monosaccharides in a polysaccharide are the same type, the polysaccharide is called
a homopolysaccharide or homoglycan, but when more than one type of monosaccharide is present
they are called heteropolysaccharides or heteroglycans.
Natural saccharides are generally of simple carbohydrates called monosaccharides with
general formulan where n is three or more.
Examples of monosaccharides are glucose, fructose, and glyceraldehyde Polysaccharides, meanwhile,
have a general formula of Cx(H2O)y where x is usually a large number between 200 and
2500.
Considering that the repeating units in the polymer backbone are often six-carbon monosaccharides,
the general formula can also be represented asn where 40≤n≤3000.
Polysaccharides contain more than ten monosaccharide units.
Definitions of how large a carbohydrate must be to fall into the categories polysaccharides
or oligosaccharides vary according to personal opinion.
Polysaccharides are an important class of biological polymers.
Their function in living organisms is usually either structure- or storage-related.
Starch is used as a storage polysaccharide in plants, being found in the form of both
amylose and the branched amylopectin.
In animals, the structurally similar glucose polymer is the more densely branched glycogen,
sometimes called 'animal starch'.
Glycogen's properties allow it to be metabolized more quickly, which suits the active lives
of moving animals.
Cellulose and chitin are examples of structural polysaccharides.
Cellulose is used in the cell walls of plants and other organisms, and is said to be the
most abundant organic molecule on earth.
It has many uses such as a significant role in the paper and textile industries, and is
used as a feedstock for the production of rayon, cellulose acetate, celluloid, and nitrocellulose.
Chitin has a similar structure, but has nitrogen-containing side branches, increasing its strength.
It is found in arthropod exoskeletons and in the cell walls of some fungi.
It also has multiple uses, including surgical threads.
Polysaccharides also include callose or laminarin, chrysolaminarin, xylan, arabinoxylan, mannan,
fucoidan and galactomannan.
Function Structure
Nutrition polysaccharides are common sources of energy.
Many organisms can easily break down starches into glucose, however, most organisms cannot
metabolize cellulose or other polysaccharides like chitin and arabinoxylans.
These carbohydrates types can be metabolized by some bacteria and protists.
Ruminants and termites, for example, use microorganisms to process cellulose.
Even though these complex carbohydrates are not very digestible, they provide important
dietary elements for humans.
Called dietary fiber, these carbohydrates enhance digestion among other benefits.
The main action of dietary fiber is to change the nature of the contents of the gastrointestinal
tract, and to change how other nutrients and chemicals are absorbed.
Soluble fiber binds to bile acids in the small intestine, making them less likely to enter
the body; this in turn lowers cholesterol levels in the blood.
Soluble fiber also attenuates the absorption of sugar, reduces sugar response after eating,
normalizes blood lipid levels and, once fermented in the colon, produces short-chain fatty acids
as byproducts with wide-ranging physiological activities.
Although insoluble fiber is associated with reduced diabetes risk, the mechanism by which
this occurs is unknown.
Not yet formally proposed as an essential macronutrient, dietary fiber is nevertheless
regarded as important for the diet, with regulatory authorities in many developed countries recommending
increases in fiber intake.
Storage polysaccharides Starches
Starches are glucose polymers in which glucopyranose units are bonded by alpha-linkages.
It is made up of a mixture of amylose and amylopectin.
Amylose consists of a linear chain of several hundred glucose molecules and Amylopectin
is a branched molecule made of several thousand glucose units.
Starches are insoluble in water.
They can be digested by hydrolysis, catalyzed by enzymes called amylases, which can break
the alpha-linkages.
Both humans and animals have amylases, so they can digest starches.
Potato, rice, wheat, and maize are major sources of starch in the human diet.
The formations of starches are the ways that plants store glucose
Glycogen Glycogen serves as the secondary long-term
energy storage in animal and fungal cells, with the primary energy stores being held
in adipose tissue.
Glycogen is made primarily by the liver and the muscles, but can also be made by glycogenesis
within the brain and stomach.
Glycogen is the analogue of starch, a glucose polymer in plants, and is sometimes referred
to as animal starch, having a similar structure to amylopectin but more extensively branched
and compact than starch.
Glycogen is a polymer of α(1→4) glycosidic bonds linked, with α(1→6)-linked branches.
Glycogen is found in the form of granules in the cytosol/cytoplasm in many cell types,
and plays an important role in the glucose cycle.
Glycogen forms an energy reserve that can be quickly mobilized to meet a sudden need
for glucose, but one that is less compact and more immediately available as an energy
reserve than triglycerides.
In the liver hepatocytes, glycogen can compose up to eight percent of the fresh weight soon
after a meal.
Only the glycogen stored in the liver can be made accessible to other organs.
In the muscles, glycogen is found in a low concentration of one to two percent of the
muscle mass.
The amount of glycogen stored in the body—especially within the muscles, liver, and red blood cells—varies
with physical activity, basal metabolic rate, and eating habits such as intermittent fasting.
Small amounts of glycogen are found in the kidneys, and even smaller amounts in certain
glial cells in the brain and white blood cells.
The uterus also stores glycogen during pregnancy, to nourish the embryo.
Glycogen is composed of a branched chain of glucose residues.
It is stored in liver and muscles.
It is an energy reserve for animals.
It is the chief form of carbohydrate stored in animal body.
It is insoluble in water.
It turns red when mixed with iodine.
It also yields glucose on hydrolysis.
Structural polysaccharides Arabinoxylans
Arabinoxylans are found in both the primary and secondary cell walls of plants and are
the copolymers of two pentose sugars: arabinose and xylose.
Cellulose The structural component of plants are formed
primarily from cellulose.
Wood is largely cellulose and lignin, while paper and cotton are nearly pure cellulose.
Cellulose is a polymer made with repeated glucose units bonded together by beta-linkages.
Humans and many animals lack an enzyme to break the beta-linkages, so they do not digest
cellulose.
Certain animals such as termites can digest cellulose, because bacteria possessing the
enzyme are present in their gut.
Cellulose is insoluble in water.
It does not change color when mixed with iodine.
On hydrolysis, it yields glucose.
It is the most abundant carbohydrate in nature.
Chitin Chitin is one of many naturally occurring
polymers.
It forms a structural component of many animals, such as exoskeletons.
Over time it is bio-degradable in the natural environment.
Its breakdown may be catalyzed by enzymes called chitinases, secreted by microorganisms
such as bacteria and fungi, and produced by some plants.
Some of these microorganisms have receptors to simple sugars from the decomposition of
chitin.
If chitin is detected, they then produce enzymes to digest it by cleaving the glycosidic bonds
in order to convert it to simple sugars and ammonia.
Chemically, chitin is closely related to chitosan.
It is also closely related to cellulose in that it is a long unbranched chain of glucose
derivatives.
Both materials contribute structure and strength, protecting the organism.
Pectins Pectins are a family of complex polysaccharides
that contain 1,4-linked α-D-galactosyluronic acid residues.
They are present in most primary cell walls and in the non-woody parts of terrestrial
plants.
Acidic polysaccharides Acidic polysaccharides are polysaccharides
that contain carboxyl groups, phosphate groups and/or sulfuric ester groups.
Bacterial capsular polysaccharides Pathogenic bacteria commonly produce a thick,
mucous-like, layer of polysaccharide.
This "capsule" cloaks antigenic proteins on the bacterial surface that would otherwise
provoke an immune response and thereby lead to the destruction of the bacteria.
Capsular polysaccharides are water soluble, commonly acidic, and have molecular weights
on the order of 100-2000 kDa.
They are linear and consist of regularly repeating subunits of one to six monosaccharides.
There is enormous structural diversity; nearly two hundred different polysaccharides are
produced by E. coli alone.
Mixtures of capsular polysaccharides, either conjugated or native are used as vaccines.
Bacteria and many other microbes, including fungi and algae, often secrete polysaccharides
to help them adhere to surfaces and to prevent them from drying out.
Humans have developed some of these polysaccharides into useful products, including xanthan gum,
dextran, welan gum, gellan gum, diutan gum and pullulan.
Most of these polysaccharides exhibit useful visco-elastic properties when dissolved in
water at very low levels.
This makes various liquids used in everyday life, such as some foods, lotions, cleaners,
and paints, viscous when stationary, but much more free-flowing when even slight shear is
applied by stirring or shaking, pouring, wiping, or brushing.
This property is named pseudoplasticity or shear thinning; the study of such matters
is called rheology.
Aqueous solutions of the polysaccharide alone have a curious behavior when stirred: after
stirring ceases, the solution initially continues to swirl due to momentum, then slows to a
standstill due to viscosity and reverses direction briefly before stopping.
This recoil is due to the elastic effect of the polysaccharide chains, previously stretched
in solution, returning to their relaxed state.
Cell-surface polysaccharides play diverse roles in bacterial ecology and physiology.
They serve as a barrier between the cell wall and the environment, mediate host-pathogen
interactions, and form structural components of biofilms.
These polysaccharides are synthesized from nucleotide-activated precursors and, in most
cases, all the enzymes necessary for biosynthesis, assembly and transport of the completed polymer
are encoded by genes organized in dedicated clusters within the genome of the organism.
Lipopolysaccharide is one of the most important cell-surface polysaccharides, as it plays
a key structural role in outer membrane integrity, as well as being an important mediator of
host-pathogen interactions.
The enzymes that make the A-band and B-band O-antigens have been identified and the metabolic
pathways defined.
The exopolysaccharide alginate is a linear copolymer of β-1,4-linked D-mannuronic acid
and L-guluronic acid residues, and is responsible for the mucoid phenotype of late-stage cystic
fibrosis disease.
The pel and psl loci are two recently discovered gene clusters that also encode exopolysaccharides
found to be important for biofilm formation.
Rhamnolipid is a biosurfactant whose production is tightly regulated at the transcriptional
level, but the precise role that it plays in disease is not well understood at present.
Protein glycosylation, particularly of pilin and flagellin, became a focus of research
by several groups from about 2007, and has been shown to be important for adhesion and
invasion during bacterial infection.
Chemical identification tests for polysaccharides Periodic acid-Schiff stain
Polysaccharides with unprotected vicinal diols or amino sugars give a positive Periodic acid-Schiff
stain.
The list of polysaccharides that stain with PAS is long.
Although mucins of epithelial origins stain with PAS, mucins of connective tissue origin
have so many acidic substitutions that they do not have enough glycol or amino-alcohol
groups left to react with PAS.
See also Glycan
Oligosaccharide nomenclature Polysaccharide encapsulated bacteria
References
External links Polysaccharide Structure
Applications and commercial sources of polysaccharides European Polysaccharide Network of Excellence