Cell Capsule
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The bacterial capsule is a large structure common to many
bacteria Bacteria (; : bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of Prokaryote, prokaryotic microorganisms. Typically a few micr ...
. It is a
polysaccharide Polysaccharides (), or polycarbohydrates, are the most abundant carbohydrates found in food. They are long-chain polymeric carbohydrates composed of monosaccharide units bound together by glycosidic linkages. This carbohydrate can react with wat ...
layer that lies outside the
cell envelope The cell envelope comprises the inner cell membrane and the cell wall of a bacterium. In Gram-negative bacteria an bacterial outer membrane, outer membrane is also included. This envelope is not present in the Mollicutes where the cell wall is abse ...
, and is thus deemed part of the outer envelope of a bacterial cell. It is a well-organized layer, not easily washed off, and it can be the cause of various diseases. The capsule—which can be found in both
gram negative Gram-negative bacteria are bacteria that, unlike gram-positive bacteria, do not retain the crystal violet stain used in the Gram staining method of bacterial differentiation. Their defining characteristic is that their cell envelope consists of ...
and
gram-positive bacteria In bacteriology, gram-positive bacteria are bacteria that give a positive result in the Gram stain test, which is traditionally used to quickly classify bacteria into two broad categories according to their type of cell wall. The Gram stain ...
—is different from the second lipid membrane –
bacterial outer membrane The bacterial outer membrane is found in gram-negative bacteria. Gram-negative bacteria form two lipid bilayers in their cell envelopes - an inner membrane (IM) that encapsulates the cytoplasm, and an outer membrane (OM) that encapsulates the p ...
, which contains
lipopolysaccharide Lipopolysaccharide (LPS), now more commonly known as endotoxin, is a collective term for components of the outermost membrane of the cell envelope of gram-negative bacteria, such as '' E. coli'' and ''Salmonella'' with a common structural archit ...
s and
lipoprotein A lipoprotein is a biochemical assembly whose primary function is to transport hydrophobic lipid (also known as fat) molecules in water, as in blood plasma or other extracellular fluids. They consist of a triglyceride and cholesterol center, sur ...
s and is found only in gram-negative bacteria. When the amorphous viscid secretion (that makes up the capsule) diffuses into the surrounding medium and remains as a loose undemarcated secretion, it is known as a slime layer. Capsule and slime layer are sometimes summarized under the term
glycocalyx The glycocalyx (: glycocalyces or glycocalyxes), also known as the pericellular matrix and cell coat, is a layer of glycoproteins and glycolipids which surround the cell membranes of bacteria, epithelial cells, and other cells. Animal epithe ...
.


Composition

Most bacterial capsules are composed of
polysaccharide Polysaccharides (), or polycarbohydrates, are the most abundant carbohydrates found in food. They are long-chain polymeric carbohydrates composed of monosaccharide units bound together by glycosidic linkages. This carbohydrate can react with wat ...
, but some species use other materials, such as poly-D-
glutamic acid Glutamic acid (symbol Glu or E; known as glutamate in its anionic form) is an α- amino acid that is used by almost all living beings in the biosynthesis of proteins. It is a non-essential nutrient for humans, meaning that the human body can ...
in ''
Bacillus anthracis ''Bacillus anthracis'' is a gram-positive and rod-shaped bacterium that causes anthrax, a deadly disease to livestock and, occasionally, to humans. It is the only permanent (obligate) pathogen within the genus ''Bacillus''. Its infection is a ty ...
''. Because most capsules are so tightly packed, they are difficult to stain because most standard stains cannot penetrate the capsule. To visualize encapsulated bacteria using a microscope, a sample is treated with a dark stain, such as
India ink India ink (British English: Indian ink; also Chinese ink) is a simple black or coloured ink once widely used for writing and printing and now more commonly used for drawing and outlining, especially when inking comic books and comic strips. In ...
. The structure of the capsule prevents the stain from penetrating the cell. When viewed,JB bacterial capsules appear as a bright halo around the cell on a dark background.


Function

The bacterial capsule serves as a shield, giving protection from toxins, and from drying out. Capsules allow adhesion to surfaces and help enable the bacteria to evade the host immune system. The water content in the capsule gives the protection against drying out. The capsule is considered a
virulence factor Virulence factors (preferably known as pathogenicity factors or effectors in botany) are cellular structures, molecules and regulatory systems that enable microbial pathogens (bacteria, viruses, fungi, and protozoa) to achieve the following: * c ...
because it enhances the ability of bacteria to cause disease (e.g. prevents
phagocytosis Phagocytosis () is the process by which a cell (biology), cell uses its plasma membrane to engulf a large particle (≥ 0.5 μm), giving rise to an internal compartment called the phagosome. It is one type of endocytosis. A cell that performs ph ...
). The capsule can protect cells from engulfment by eukaryotic cells, such as macrophages. A capsule-specific antibody may be required for
phagocytosis Phagocytosis () is the process by which a cell (biology), cell uses its plasma membrane to engulf a large particle (≥ 0.5 μm), giving rise to an internal compartment called the phagosome. It is one type of endocytosis. A cell that performs ph ...
to occur. They also exclude bacterial
viruses A virus is a submicroscopic infectious agent that replicates only inside the living cells of an organism. Viruses infect all life forms, from animals and plants to microorganisms, including bacteria and archaea. Viruses are found in almo ...
and most
hydrophobic In chemistry, hydrophobicity is the chemical property of a molecule (called a hydrophobe) that is seemingly repelled from a mass of water. In contrast, hydrophiles are attracted to water. Hydrophobic molecules tend to be nonpolar and, thu ...
toxic materials such as detergents. Immunity to one capsule type does not result in immunity to the other types. Capsules also help cells adhere to surfaces. As a group where the capsule is present they are known as polysaccharide encapsulated bacteria or encapsulated bacteria.


Diversity

The capsule is found most commonly among gram-negative bacteria: * ''
Escherichia coli ''Escherichia coli'' ( )Wells, J. C. (2000) Longman Pronunciation Dictionary. Harlow ngland Pearson Education Ltd. is a gram-negative, facultative anaerobic, rod-shaped, coliform bacterium of the genus '' Escherichia'' that is commonly fo ...
'' (in some strains) * ''
Neisseria meningitidis ''Neisseria meningitidis'', often referred to as the meningococcus, is a Gram-negative bacterium that can cause meningitis and other forms of meningococcal disease such as meningococcemia, a life-threatening sepsis. The bacterium is referred to a ...
'' * ''
Klebsiella pneumoniae ''Klebsiella pneumoniae'' is a Gram-negative, non-motile, encapsulated, lactose- fermenting, facultative anaerobic, rod-shaped bacterium. It appears as a mucoid lactose fermenter on MacConkey agar. Although found in the normal flora of the mo ...
'' * ''
Haemophilus influenzae ''Haemophilus influenzae'' (formerly called Pfeiffer's bacillus or ''Bacillus influenzae'') is a Gram-negative, Motility, non-motile, Coccobacillus, coccobacillary, facultative anaerobic organism, facultatively anaerobic, Capnophile, capnophili ...
'' * ''
Pseudomonas aeruginosa ''Pseudomonas aeruginosa'' is a common Bacterial capsule, encapsulated, Gram-negative bacteria, Gram-negative, Aerobic organism, aerobic–facultative anaerobe, facultatively anaerobic, Bacillus (shape), rod-shaped bacteria, bacterium that can c ...
'' * ''
Salmonella ''Salmonella'' is a genus of bacillus (shape), rod-shaped, (bacillus) Gram-negative bacteria of the family Enterobacteriaceae. The two known species of ''Salmonella'' are ''Salmonella enterica'' and ''Salmonella bongori''. ''S. enterica'' ...
'' * ''
Acinetobacter baumannii ''Acinetobacter baumannii'' is a typically short, almost round, rod-shaped (coccobacillus) Gram-negative bacterium. It is named after the bacteriologist Paul Baumann. It can be an opportunistic pathogen in humans, affecting people with compromis ...
'' However, some gram-positive bacteria may also have a capsule: * ''
Bacillus megaterium ''Priestia megaterium'' (''Bacillus megaterium'' prior to 2020) is a rod-like, Gram staining, Gram-positive, mainly aerobic, endospore, spore forming bacterium found in widely diverse habitats.De Vos, P. ''et al.'' Bergey's Manual of Systematic B ...
'' for example, synthesizes a capsule composed of polypeptide and polysaccharides. * ''
Bacillus anthracis ''Bacillus anthracis'' is a gram-positive and rod-shaped bacterium that causes anthrax, a deadly disease to livestock and, occasionally, to humans. It is the only permanent (obligate) pathogen within the genus ''Bacillus''. Its infection is a ty ...
'' * ''
Streptococcus pyogenes ''Streptococcus pyogenes'' is a species of Gram-positive, aerotolerant bacteria in the genus '' Streptococcus''. These bacteria are extracellular, and made up of non-motile and non-sporing cocci (round cells) that tend to link in chains. They ...
'' synthesizes a
hyaluronic acid Hyaluronic acid (; abbreviated HA; conjugate base hyaluronate), also called hyaluronan, is an anionic, nonsulfated glycosaminoglycan distributed widely throughout connective, epithelial, and neural tissues. It is unique among glycosaminog ...
capsule. * ''
Streptococcus pneumoniae ''Streptococcus pneumoniae'', or pneumococcus, is a Gram-positive, spherical bacteria, hemolysis (microbiology), alpha-hemolytic member of the genus ''Streptococcus''. ''S. pneumoniae'' cells are usually found in pairs (diplococci) and do not f ...
'' has at least 91 different capsular serotypes. These serotypes are the basis for the
pneumococcal vaccine Pneumococcal vaccines are vaccines against the bacterium ''Streptococcus pneumoniae''. Their use can prevent some cases of pneumonia, meningitis, and sepsis. There are two types of pneumococcal vaccines: conjugate vaccines and polysaccharide ...
s. * ''
Streptococcus agalactiae ''Streptococcus agalactiae'' (also known as group B streptococcus or GBS) is a gram-positive coccus (round bacterium) with a tendency to form chains (as reflected by the genus name ''Streptococcus''). It is a beta-hemolytic, catalase-negative, an ...
'' produces a polysaccharide capsule of nine antigenic types that all contain sialic acid (Ia, Ib, II, III, IV, V, VI, VII, VIII). * ''
Staphylococcus epidermidis ''Staphylococcus epidermidis'' is a Gram-positive bacterium, and one of over 40 species belonging to the genus ''Staphylococcus''. It is part of the human flora, normal human microbiota, typically the skin flora, skin microbiota, and less commonl ...
'' * ''
Staphylococcus aureus ''Staphylococcus aureus'' is a Gram-positive spherically shaped bacterium, a member of the Bacillota, and is a usual member of the microbiota of the body, frequently found in the upper respiratory tract and on the skin. It is often posi ...
'' * ''
Lactococcus garvieae ''Lactococcus garvieae'' is a known fish pathogen affecting saltwater fish in the Far East, specifically in rainbow trout, Japanese yellowtail, Cobia (''Rachycentron canadum'') and grey mullet (''Mugil cephalus''). This bacteria causes lesions ...
'' synthesizes capsular gene clusters and some time synthesizes a
hyaluronic acid Hyaluronic acid (; abbreviated HA; conjugate base hyaluronate), also called hyaluronan, is an anionic, nonsulfated glycosaminoglycan distributed widely throughout connective, epithelial, and neural tissues. It is unique among glycosaminog ...
capsule. The
yeast Yeasts are eukaryotic, single-celled microorganisms classified as members of the fungus kingdom (biology), kingdom. The first yeast originated hundreds of millions of years ago, and at least 1,500 species are currently recognized. They are est ...
''
Cryptococcus neoformans ''Cryptococcus neoformans'' is an encapsulated basidiomycetous yeast belonging to the class Tremellomycetes and an obligate aerobe that can live in both plants and animals. Its teleomorph is a filamentous fungus, formerly referred to ''Filob ...
'', though not a bacterium, has a similar capsule. Capsules too small to be seen with an ordinary microscope, such as the M protein of ''
Streptococcus pyogenes ''Streptococcus pyogenes'' is a species of Gram-positive, aerotolerant bacteria in the genus '' Streptococcus''. These bacteria are extracellular, and made up of non-motile and non-sporing cocci (round cells) that tend to link in chains. They ...
'', are called microcapsules.


Demonstration of capsule

# India ink staining: the capsule appears as a clear halo around the bacterium as the ink can't penetrate the capsule. # Maneval's capsule stain: the capsule appears as a clear halo between the pink-stained bacterium and the bluish-grey stained background. The background stain is the acidic stain Congo red (which changes color to bluish-grey due to the pH), and the pink stain is
fuchsine Fuchsine (sometimes spelled fuchsin) or rosaniline hydrochloride is a magenta dye with chemical formula C20H19N3·HCl.
. # Serological methods: Capsular material is antigenic and can be demonstrated by mixing it with a specific anticapsular serum. When examined under the microscope, the capsule appears 'swollen' due to an increase in its refractivity. This phenomenon is the basis of
quellung reaction The quellung reaction, also called the Neufeld reaction, is a biochemical reaction in which antibodies bind to the bacterial capsule of ''Streptococcus pneumoniae'', ''Klebsiella pneumoniae'', ''Neisseria meningitidis'', ''Bacillus anthracis'', ''H ...
.


Use in vaccination

Vaccination Vaccination is the administration of a vaccine to help the immune system develop immunity from a disease. Vaccines contain a microorganism or virus in a weakened, live or killed state, or proteins or toxins from the organism. In stimulating ...
using capsular material is effective against some organisms (e.g., '' H. influenzae'' type b, '' S. pneumoniae'', and '' N. meningitidis''). However, polysaccharides are not highly antigenic, especially in children, so many capsular vaccines contain polysaccharides conjugated with protein carriers, such as the
tetanus toxoid Tetanus vaccine, also known as tetanus toxoid (TT), is a toxoid vaccine used to prevent tetanus. During childhood, five doses are recommended, with a sixth given during adolescence. After three doses, almost everyone is initially immune, but ...
or diphtheria toxoid. This stimulates a much more robust immune response.


See also

*
Bacterial cell structure A bacterium, despite its simplicity, contains a well-developed cell structure which is responsible for some of its unique biological structures and pathogenicity. Many structural features are unique to bacteria, and are not found among archaea ...
*
Quellung reaction The quellung reaction, also called the Neufeld reaction, is a biochemical reaction in which antibodies bind to the bacterial capsule of ''Streptococcus pneumoniae'', ''Klebsiella pneumoniae'', ''Neisseria meningitidis'', ''Bacillus anthracis'', ''H ...
, a method to visualize capsule under a microscope


References

{{Bacteria Bacteriology Prokaryotic cell anatomy