Bacteria

Morphology and fine structure of bacteria

Bacteria:

Bacteria are ubiquitous, mostly free-living organisms often consisting of one biological cell. They are
prokaryotes. The word bacteria is the plural of the New Latin bacterium, which is the latinisation of the
Ancient Greek (baktḗrion) bacterium meaning rod. They are unicellular prokaryotes (Greek word meaning
prenucleus). Humans and most other animals carry vast numbers (approximately 1013 to 1014) of bacteria. Most
are in the gut, and there are many on the skin as well.

Morphology: The study of size, shape, structure and arrangement.

 Size of Bacteria

Bacteria come in many sizes

1- Spherical Bacteria Example; Cocci has size 0.5 to 2 µm

2- Rod Shape Bacteria Example; Bacilli has size 1 – 10 µm in length and 0.2 - 2µm in width.

Largest bacteria Thiomargarita nambiensis is about 0.3mm in size and coccoid in shape

World smallest bacteria is Mycoplasma genitalium 200-300 nm in size and has flask shaped cells.

 Shape of bacteria:

Is due to presence of rigid cell wall. Few basic shapes of bacteria are.

1- Spherical bacteria called cocci meaning berries.

2- Rod Shape bacteria called bacilli meaning little staff
3- Helical curved rods (Spiral)

Helical curved bacteria may be of following basic shapes

a) Bacteria that look like curved rods (coma shape) are called vibrios
Vibrio cholerae
b) Spirilla, rigid spiral shaped cells (like cork screw)
Helicobacter pylori
c) Spirochetes flexible multiple spiral shaped bacteria (Treponema pallidum)

4) Coccobacilli. Some bacilli are oval and look so much like cocci that they are called coccobacilli
(Haemophilus influenza) Besides These basic shapes few bacteria are star shaped, rectangular, and triangular.
Most bacterial cells have constant single shape and hence are monomorphic. Some bacteria are pleomorphic
being variable in shape and lacking a single characteristic forms.

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  Arrangement of Bacteria:

Bacterial cells are usually arranged in manner characteristics of their particular genra. Arrangement of bacteria
depend upon the plane of cell division and behavior of cell after division

Cocci: They appear in several characteristic arrangement

1- Micrococcus: Cocci that remain single are called micrococcus (Micrococcus lylae)
2- Diplococci: Cocci that appear in pair form are called diplococci. (Neisseria gonorrhoeae)
3- Streptococci: Cocci that remains attached in chain like arrangement are called streptococci.
(Streptococcus pyogens)
4- Tetrad: Those cocci that divides into two planes and remains in group of four is known as tetrad
arrangement. (Aerococcus)
5- Staphylococci: Cocci that divides in multiple planes and form grape like structure or arrangement.
(Staphylococcus aureus)
6- Sarcinae: Division of cocci bacteria in three planes produced cuboidal shape.

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b) Bacilli have fewer grouping than cocci

1) Single bacillus: Most bacilli appear as single rods

2) Diplobacilli : Bacilli appear in pairs after division.

3) Sterptobacilli . Bacilli occur in chains

4) Trichomes are like Streptobacilli but have much larger area of contact between cells.

5) Palisade arrangement. Cells are lined side by side like matchsticks.

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 Structure of Bacteria:

Bacterial Cell Wall

Prokaryotic eubacteria (bacteria) have a cell wall made of a unique substance called peptidoglycan. The
rigid structure of peptidoglycan gives the bacterial cell shape, surrounds the plasma membrane and provides
prokaryotes with protection from their environment

Beyond the cell wall, some bacteria have an additional layer called the glycocalyx. The structural features
and chemical composition of glycocalyces differ depending on the species of bacteria, but in general this
additional layer can come in one of two forms:
Structure External to the cell wall:

a) Glycocalyx (Sugar coat)

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Many bacteria secrete a sticky, viscous material that forms an extracellular coating around the cell. The material
is usually a polysaccharide, however, in the case of pathogenic Bacillus anthracis, is poly-D-glutamic acid.
Negative charge on the capsule repel phagocytes.

If the glycocalyx is thick, covalently bound to the cell and has an organized structure, it is called a capsule. If
the material is loosely bound and thinner, it is called a slime layer. If this layer can be visualized by light
microscopy using special staining methods, it is termed a capsule if the layer is too thin to be seen by light
microscopy it is termed a microcapsule.

Slime Layer
A glycocalyx is considered a slime layer when the glycoprotein molecules are loosely associated with the
cell wall. Bacteria that are covered with this loose shield are protected from dehydration and loss of
nutrients.
Capsule
The glycocalyx is considered a capsule when the polysaccharides are more firmly attached to the cell wall.
Capsules have a gummy, sticky consistency and provide protection as well as adhesion to solid surfaces and
to nutrients in the environment.

Bacteria that possess capsules are considered to be encapsulated, and generally have greater pathogenicity
(ability to cause disease) because capsules protect bacteria, even from phagocytic white blood cells of the
immune system. The adhesive power of capsules is also a major factor in the initiation of some bacterial
diseases.

Diagram of bacterial mucoid-like structures:

1. capsule, 2. slime layer, 3. biofilm

What Is a Biofilm?
Glycocalyces are instrumental in the formation of biofilms. A biofilm is a living ecosystem made of
millions of bacterial cells, their wastes and other extracellular products.
Oral Biofilm & Plaque
The slime layer of Streptococcus mutans allows this bacteria and others to accumulate on tooth enamel
(yuck mouth and one of the causes of cavities). Other bacteria in the mouth become trapped in the slime and
form a biofilm, eventually building up as plaque.

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Functions
Contributing to the organisms' pathogenicity

The capsule and glycocalyx allow cells to adhere to surfaces, protect bacteria from antibodies and phagocytosis,
act as diffusion barriers against some antibiotics thus contributing to the organisms' pathogenicity. In
Streptococcus pneumonia only capsulated strains are pathogenic.
Steptococcus mutans attach to dental surface and cause dental carries

Provide protection to organism

Against temporary drying by binding water molecules.
Block attachment of bacteriophage

b) Appendages

Many bacteria have hair like appendages that project from the cell wall. There are of two kinds of appendages:
 Flagella (singular, flagellum) and pili (singular, pilus).
Flagella. Prokaryotic flagella are long, semi-rigid, helical, hollow tubular structures composed of several
thousand molecules of the protein flagellin. Cells may have one or many flagella. Flagella are highly antigenic

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Size and Structure; Prokaryotic flagella is thin and 20nm in width and 10-20um in length and It is compose of
three parts:

Basal body; Flagella are attached to the cell wall, cell membrane, or both by a basal body, which is a complex
molecular machine that rotates the flagellum like the screw propeller of a ship. It is a ring like structure. Its
composition is unknown.
Hook; it attaches filament to the basal body.
Filament; it is several time as long as hook cell and is composed of protein sub units called flagellin.

Arrangement:

Arrangement of flagella in bacteria is unique to species observed.

i. Monotrichous:
A single polar flagellum

Example. Pseudomonas aeruginosa

ii. Lophotrichous:
A tuft/cluster of polar flagella.

Example. Pseudomonas fluorescens.

iii. Amphitrichous:
Either a single flagellum or tuft of flagella on both poles.

Aquaspirillum serpens

iv. Peritrichous:
Whole bacterial body is surrounded by lateral flagella.

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 Example. Salmonella typhi

Function. They enable bacteria to propel themselves, for example, in response to a chemotactic stimulus.
Bacteria having polar flagella swim in a back and forth fashion .They reverse the direction of the swimming by
reversing the direction of flagella rotation.

Bacteria having lateral flagella move in a more complicated manner. Their flagella operate in synchrony to form
bundle that extend behind the cell.

Bacterial movement:

Bacterial chemotaxis; most motile bacteria are capable of directed swimming toward or away from a various
chemical compounds, process is called chemotaxis. Stimulus is not the chemical but concentration of chemical
i.e. temporal gradient which is sensed by protein chemoreceptors

i- Positive Chemotaxis: Swimming towards a chemical called positive chemotaxis.

ii- Negative Chemotaxis: Swimming away from a chemical is called negative chemotaxis

iii- Phototaxis; Phototropic bacteria exhibit phototaxis i.e. movement towards light.

iv- Glinding motility; some bacterial species are motile only when they are in contact with solid surface.
Myxobacteria show gliding motility by its pili.

Periplasmic Flagella. Certain helical bacteria like spirochetes lack external flagella. They have gel like
structure located within the cell called periplasmic flagella or endoflagella or axial filament.

Atrichous: Bacteria without flagella.

Pili:

Pilus (hair)

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“These are hollow, non-helical, filamentous structures that are thinner shorter and more numerous than flagella.
They have no role in bacterial motility. They are mostly fond on Gram negative bacteria”

Functions

 Pili is involved in conjugation. One type known as F-Pilus or Sex-Pilus serves as a port of entry of
genetic material during bacterial mating.

 Pili play role in bacterial virulence. They help in attachment of pathogenic bacteria to attach to epithelial
cell lining of respiratory of respiratory, intestinal or genito-urinary tract and prevent the washing of
bacteria hence help in the establishment of disease or infection in human. E.coli causing severe watery
diarrhea, attack to lining of small intestine by Pili. Neisseria gonorrhoeae causing genitourinary tract
infections attach to the lining with Pili.

Some of the differences between fimbriae and Pili are as follows:

Characteristics Fimbriae Pili

Pili are hair like

Fimbriae are tiny bristle-like
microfibers that are thick
1 Definition fibers arising from the surface
tubular structure made up
of bacterial cells.
of pilin.

2 Length Shorter than pili Longer than fimbriae.

3 Diameter Thin Thicker than fimbriae.

No. of fimbriae are 200-400 No of pili are less 1-10 per

4 Number
per cell. cell.

5 Made up of Fimbrillin protein. Pilin protein.

Responsible for bacterial

Responsible for cell to surface conjugation.
6 Function attachment. Specialized for
attachment i.e. enable the cell
to adhere the surfaces of other Two basic function of pili.
They are gene transfer and

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 bacteria. attachment.

Do not function in active Type IV pili shows

7 Motility
motility. twitching type of motility.

Escherichia coli, Neisseria

gonorrhoeae.
Salmonella typhimurium, Neisseria gonorrhoeae, the
Shigella dysenteriae. cause of gonorrhea, uses
Shigella dysenteriae uses its pili to attach to the
8 Examples
fimbriae to attach to the urogenital and cervical
intestine and then produces a epithelium when it causes
toxin that causes diarrhea. disease.

Sheath

Some bacterial species those from fresh or marine environment from chain or trichomes enclosed by a hollow,
tube like sheath which play a role in bacterial survival

Prosthecae (Prosthecate Bacteria)

Prosthecae are semi-rigid extension of cell wall and membrane that increase the surface area of cell for nutrient
absorption.

Stalks

Stalks are certain non-living ribbon like structures secreted by cells which help in attachment of cell to various
surfaces.

Cell Wall:

The cell wall of bacterial cell is a complex semi rigid structure responsible for the shape of the cell.
Thickness ranges from 10-25 nm. Its weight ranges from 20-25% of dry weight of cell.

Function of cell wall:

i- It maintains the shape of bacteria.

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 ii- It protects the cell from expending a bursting because of uptake of water because most of the
bacterium live in hypotonic environment.
iii- It is the site of attachment of flagella.
iv- It is essential for bacterial growth and division. Cells whose walls have been removed are
incapable of normal growth and division.
v- It contains various virulence factors like Endotoxins which contribute in bacterial pathogenicity.
vi- It is the site of action of several antibiotics.

Structure and chemical composition

The bacterial cell wall is composed of a polymer called peplidoglycan (also known as murein). It is composed
of Sugar and amino acid. The disaccharide portion is made up of monosaccharides called N-acetylglucosamine
(NAG) and N-acetylmuramic acid (NAM). Alternating NAM and NAG molecules are linked in rows of 10 to
65 sugars to form a carbohydrate "backbone" (the glycan portion of peptidoglycan). Adjacent rows are linked
by polypepides (the peptide portion of peptidoglycan). The amino acids of Gram Positive bacteria are L
Alanine, D Glutamine, L Lysine and D Alanine while The major variation is observed at the third amino acid of
the peptide interbridge. Most Gram-negative bacteria and Gram-positive bacilli have meso-diaminopimelic
acid (meso-Dap), whereas other Gram-positive bacteria use l-lysine (l-Lys) at this position.

Gram-Positive Cell Walls

In most gram-positive bacteria, the cell wall consists of many layers of peptidoglycan, forming a thick, rigid
structure. In addition, the cell walls of gram-positive bacteria contain teichoic acids and Lipoteichoic acids,
which consist primarily of an alcohol (such as glycerol or ribitol) and phosphate (this carries strong negative)
which regulate movement of cations into and out of the cell.

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Gram-Negative Cell Walls

The cell walls of gram-negative bacteria consist of one or a very few layers of peptidoglycan and an outer
membrane, a bilayer consisting of phospholipid, protein, and lipopolysaccharide .Lipids portion of
lipopolysaccharides is called lipid A and is referred as endotoxin and is a toxic in host blood stream or GIT. It
cause fever and shock.
The peptidoglycan is bonded to lipoproteins (lipids covalently linked to proteins) in the outer membrane. The
periplasm, a gel-like fluid between the outer membrane and the plasma membrane is also present in Gram
negative bacteria. The periplasm contains a high concentration of degradative enzymes and transport proteins.
Gram-negative cell walls do not contain teichoic acids. Because the cell walls of gram-negative bacteria contain
only a small amount of peptidoglycan, they are more susceptible to mechanical breakage.

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Difference between Gram positive and Gram negative cell wall.

Gram negative cell wall Gram positive cell wall

It contain less peptidoglycan almost 10% of It contains more peptidoglycan almost 50% of
cell wall cell wall.
Less thick almost 2-7 nm More thick almost 20-80 nm
Cell wall structure is more complex Simple structure of cell wall
Outer membrane is present i.e. composed of An outer membrane is absent so it contains less
phospholipids, proteins and lipids.
lipopolysaccharides. so it contain more lipids.
Periplasmic space is present. It is a fluid is A much smaller periplasmic space is also
filled space between outer membrane and present in gram-positive bacteria (more
plasma membrane. It contains high accurately "monoderm"), between cell wall and
concentration of enzymes and transport the plasma membrane.
proteins.
Lipoteichoic acid is absent Lipoteichoic acid is present

Function of Cell Wall in Gram Negative Bacteria

It carries strong negative charge which is an important factor in invading phagocytosis and action of
complement system. It also provides a barrier to anti-biotics.

Clinical Importance

Clinically the cell wall is important because it contributes to the ability of some species to cause disease and is
the site of action of some antibiotics. Chemicals that damage bacterial cell walls, or interfere with their
synthesis, often do not harm the cells of an animal host because the bacterial cell wall is made of chemicals
unlike those in eukaryotic cells. Thus, cell wall synthesis is the target for some antimicrobial drugs certain
antibiotics, such as penicillin, destroy bacteria by interfering with the formation of the peptide cross-bridges of
peptidoglycan, thus preventing the formation of a functional cell wall. Most gram-negative bacteria are not as
susceptible to penicillin as gram-positive bacteria are because the outer membrane of gram-negative bacteria
forms a barrier that inhibits the entry of this and other substances, and gram- negative bacteria have fewer
peptide cross-bridges. However, gram-negative bacteria are quite susceptible to some beta lactam antibiotics
that penetrate the outer membrane better than penicillin.

ATYPICAL CELL WALL

Mycoplasma

They have no cell wall. Their plasma membrane has lipids called sterols and enclosing the cytoplasm of cell
which protect them from lysis. They are small in size and pass through bacterial filters so are sometime
mistakenly considered as viruses.

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Mycoplasma are opportunistic pathogens. They cause pneumonia (Mycoplasma pneumoniae) in children.

Acid Fast Bacteria

Few bacterial genra like Mycobacterium and Nocardia have 60% of waxy material mycolic acid in their cell
wall. Because of mycolic acid they are resistant to staining. They are stained by special staining techniques so
are called Acid Fast Bacteria.

Archaeobacteria

Although most Archaeobacteria possess cell walls, these donot contain peptidoglycan. Their cell wall is usually
composed of protein, glycoprotein or polysaccharide. A few genra like Methanobacterium have cell wall
composed of pseudomurein.

Cytoplasmic Membrane

Cytoplasmic membrane lying inside the cell wall

Structure

Its thickness is 7.5nm. It is composed of phospholipid (20-30%) and proteins (60-70%).

Phospholipid form a bilayer in which most of proteins are strongly held (integral proteins) and other types of
protein that are loosely attached (peripheral proteins) and may be removed by a mild treatment of membrane
such as osmotic shock.

Function

1. Most important function of plasma membrane to act as selective barrier through which material enter and
exists the cells. Larger molecules such as proteins cannot pass through membrane but smaller molecules such as
oxygen, carbon dioxide and simple sugars can pass through it.

2. It also contains various enzymes that are involved in metabolism they break down nutrients and produced
energy in form of ATP.

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3. It also contains enzymes that are involved in synthesis of capsular or cell wall components.

Destruction of the Plasma Membrane by Antimicrobial Agents

Because the plasma membrane is vital to the bacterial cell, it is not surprising that several antimicrobial agents
exert their effects at this site. In addition to the chemicals that damage the cell wall and thereby indirectly
expose the membrane to injury, many compounds specifically damage plasma membranes. These compounds
include certain alcohols and quaternary ammonium compounds, which are used as disinfectants. By disrupting
the membrane's phospholipids, a group of antibiotics known as the polymyxins cause leakage of intracellular
contents and subsequent cell death
Protoplast

A protoplast is the portion of bacterial cell consisting of cytoplasmic membrane and cell material
bounded by it.

The term protoplast refers to the spherical shape assumed by Gram-positive bacteria. Spheroplast refers to the
spherical shape assumed by Gram-negative bacteria. The difference is essentially the presence of a single
membrane, in the case of the protoplast, and the two membranes (inner and outer) of the Gram-negative
spheroplasts. It is also possible to generate a gram-negative protoplast by the removal of the outer membrane.
Thus, in essence, protoplast refers to a bacterial sphere that is bounded by a single membrane and spheroplast
refers to a sphere that is bounded by two membranes.

Protoplast can be prepared from Gram positive bacteria by treating the cell with lysozyme or culturing the
bacteria in the presence of penicillin.

Spheroplast.

Round, osmotically fragile form of Gram negative bacteria by procedure similar as described in protoplast. It
contains two membranes, the cytoplasmic membrane and the outer membrane.

CYTOPLASM

Cytoplasm is the cell material bounded by the cytoplasmic membrane and it is divided into

1. Cytoplasmic area is granular in appearance and rich in ribosomes by which are protein synthesized.

2. Chromatic area is rich in DNA.

3. Fluid portion with dissolved substances.

Ribosomes of prokaryotes composed of two subunits 50S and 30S total are 70S. In contrast to ribosomes of
eukaryotes that has sedimentation co-efficient of 80S composed of two subunits 60S and 40S. In contrast to
eukaryotes, bacterial cell contains neither a distinct membrane enclosed nucleus nor a mitotic apparatus, so the
structure has been designated as NUCLEOID means nucleus like. It also contains a single circular DNA
molecule called PLASMID. Within cytoplasm
of prokaryotic cells there are several kind of reserved deposits known as inclusions. These include lipid
inclusion, sulphur granules, meta-chromatic granules (inorganic phosphate),polysaccharide granules, gas
vacuole and magnetosomes (Fe3O4).
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Some inclusions in Bacterial Cells Inclusion Composition Glycogen poly-glucose Reserve carbon and energy
source Poly-betahydroxybutyric acid (PHB) lipid Reserve carbon and energy source Poly-phosphates polymers
of PO4 Reserve phosphate, possibly high-energy PO4. Sulfur globules elemental S Reserve energy and or
electrons. Magnetosomes magnetite (iron oxide) Provide orientation in magnetic field, Gas vesicles protein
shells inflated with gases Provide buoyancy in aquatic environments, Parasporal crystals protein Produced by
endospore forming Bacilli - toxic to insects Function.

CLINICAL IMPORTANCE

Several antibiotics work by inhibiting protein synthesis on prokaryotic ribosomes. Antibiotics such as
Streptomycin and Gentamycin attach to 30S subunit and interfere with protein synthesis.

Other antibiotics such as erythromycin and Chloremphenicol interfere with protein synthesis by attaching to
50S subunit. Because of difference in prokaryotic and eukaryotic ribosomes, the microbial cell can be killed by
the antibiotic while the eukaryotic host cell remains unaffected.

Bacterial DNA replication is inhibited by certain quinolones such as ciprofloxacin and levofloxacin. They
prevent chromosomal topology by targeting DNA gyrase and Topoisomerase.

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Spores

Spore is a metabolic dormant form which under appropriate condition undergo generation to form a vegetative
cell. Certain species of bacteria produce spores either within cell (endospore) or external to cell(exospores).

Endospore

The process of endospore formation within a vegetative cell takes several hours and is known as
sporogenesis/sporulation. These endospore are thick walled, highly durable bodies that are produce by a few
genera of Gram- positive bacteria when essentials nutrients are depleted.

Example of genera produce endospore are Bacillus, Clostridium, Sporosarcinae and few other genera.

SHAPE AND LOCATION OF ENDOSPORE

There are three types of shapes and location

Spore are elliptical and centrally located e.g. Bacillus cerus Spore are spherical and terminally located e.g.
Clostridium tetani Spore are ovoid (oval- like) and sub-terminally located. Clostridium sub-terminate

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Endospores are produced as intracellular structures within the cytoplasm of certain bacteria, most notably
Bacillus and Clostridium species. Endospore forming bacteria left to right: Clostridium botulinum,
Bacillusbrevis, Bacillus thuringiensis.

Functions

Endospore are resistant to dessication (drying) staining, radiation and disinfecting chemicals.
Composition All endospore contain large amount of DPA (dipilcolonic acid).
It is about 10% to 15% of spore weight and it occurs in combination with calcium.
Endospore remain dormant for thousands of years and returns to its vegetative state by a process called
germination.

Endospore formation is NOT a mechanism of reproduction. Rather it is a mechanism for survival in deleterious
environments. During the process of spore formation, one vegetative cell develops into one endospore. The
sequential steps of endospore formation in a Bacillus species. The process of endospore formation takes about
six hours. Eventually the mature endospore is released from its “mother cell” as a free spore Free endospore
Endospore within mother cell Vegetative cell

Medically-important Endospore-forming Bacteria Bacillus anthracis causes anthrax Bacillus cereus causes food
poisoning Clostridium tetani causes tetanus Clostridium botulinum causes botulism Clostridium perfringens
causes food poisoning and gas gangrene. Clostridium difficile causes antibiotic-induced diarrhea and
pseudomembranous colitis

Medical significance of sporulation

Some of the most notorious pathogens are spore-formers, including B. anthracis (anthrax,), Bacillus cereus
(gastroenteritis,), Clostridium tetani (tetanus,), Clostridium botulinum (botulism,), and Clostridium perfringens
(gas gangrene,). Spores of these organisms can remain viable for many years, and are generally not killed by
boiling, but can be killed by autoclaving

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 EXOSPORE: Exospore are spore external to the vegetative cell. EXAMPLE Cell of Methylosinus species
form exospores by budding at the one end of cell.

Function. They provide desiccation and heat resistant to cell.

EXOTOXIN: Heat sensitive protein molecules manufactured during metabolism by Gram Positive and Gram
Negative cells are released into host environment or in host cells categorized as

Cytotoxins: Kills Cells

Hemolysin: It Lyses RBC

Neurotoxin: Interfere with Nerve transmission

Enterotoxin: Effect intestinal Tract

Examples include the toxins responsible for cholera, diphtheria and botulism.

An exotoxin is a toxin secreted by bacteria. An exotoxin can cause damage to the host by destroying cells or
disrupting normal cellular metabolism. They are highly potent and can cause major damage to the host.
Exotoxins may be secreted, or, similar to endotoxins, may be released during lysis of the cell. Gram negative
pathogens may secrete outer membrane vesicles containing lipopolysaccharide endotoxin and some virulence
proteins in the bounding membrane along with some other toxins as intra-vesicular contents, thus adding a
previously unforeseen dimension to the well-known eukaryote process of membrane vesicle trafficking, which
is quite active at the host-pathogen interface.
They may exert their effect locally or produce systemic effects. Well-known exotoxins include: botulinum toxin
produced by Clostridium botulinum; Corynebacterium diphtheriae toxin, produced during life-threatening
symptoms of diphtheria; tetanospasmin produced by Clostridium tetani. The toxic properties of most exotoxins
can be inactivated by heat or chemical treatment to produce a toxoid. These retain their antigenic specificity and
can be used to produce antitoxins and, in the case of diphtheria and tetanus toxoids, are used as vaccines.
Exotoxins are susceptible to antibodies produced by the immune system, but some exotoxins are so toxic that
they may be fatal to the host before the immune system has a chance to mount defenses against them. In such
cases, antitoxin, anti-serum containing antibodies, can sometimes be injected to provide passive immunity.

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