The Cell Wall
The internal osmotic pressure of most bacteria ranges
the dye–iodine complex and become translucent, but
from 5 to 20 atm as a result of solute concentration via
they can then be counterstained with safranin (a red
active transport. In most environments, this pressure
dye). Thus, gram-positive bacteria look purple under the
would be sufficient to burst the cell were it not for the
microscope, and gram-negative bacteria look red. The
presence of a high-tensile strength cell wall (Figure 2-
distinction between these two groups turns out to reflect
14). The bacterial cell wall owes its strength to a layer
fundamental differences in their cell envelopes (Figure
composed of a substance variously referred to as murein,
2-15).
mucopeptide, or peptidoglycan (all are synonyms). The
structure of peptidoglycan is discussed below. Most Special Components of Gram-Positive Cell Walls
bacteria are classified as gram-positive or gram-negative
according to their response to the Gram-staining Most gram-positive cell walls contain considerable
procedure. This procedure was named for the histologist amounts of teichoic and teichuronic acids, which may
Hans Christian Gram, who developed this differential account for up to 50% of the dry weight of the wall and
staining procedure in an attempt to stain bacteria in 10% of the dry weight of the total cell. In addition, some
infected tissues. The Gram stain depends on the ability gram-positive walls may contain polysaccharide
of certain bacteria (the gram-positive bacteria) to retain molecules.
a complex of crystal violet (a purple dye) and iodine 1. Teichoic and teichuronic acids
after a brief wash with alcohol or acetone. Gram-
negative bacteria do not retain There are two types of teichoic acids: wall teichoic acid
(WTA), covalently linked to peptidoglycan, and
membrane teichoic acid, covalently linked to membrane
glycolipid. Because the latter are intimately associated
with lipids, they have been called lipoteichoic acids
(LTA). Together with peptidoglycan, WTA and LTA
make up a polyanionic network or matrix that provides
functions relating to the elasticity, porosity, tensile
strength, and electrostatic properties of the envelope.
Although not all gram-positive bacteria have
conventional LTA and WTA, those that lack these
polymers generally have functionally similar ones.
The teichoic acids constitute major surface antigens of
those gram-positive species that possess them, and their
accessibility to antibodies has been taken as evidence
that they lie on the outside surface of the peptidoglycan.
The teichuronic acids are synthesized in place of teichoic
acids when phosphate is limiting.
2. Polysaccharides Gram-positive cell wall consists of a plasma membrane
and thick peptidoglycan layer; the gram-negative cell
The hydrolysis of gram-positive walls has yielded, from wall consists of a plasma membrane, a thin
certain species, neutral sugars such as mannose, peptidoglycan layer, and an outer membrane containing
arabinose, rhamnose, and glucosamine and acidic sugars lipopolysaccharide (endotoxin). The space between the
such as glucuronic acid and mannuronic acid. plasma membrane and outer membrane is referred to as
the periplasmic space.
Special Components of Gram-Negative Cell Walls
Gram-negative cell walls contain three components that Most bacteria are classified as gram-positive or gram–
lie outside of the peptidoglycan layer: lipoprotein, outer negative according to their response to the Gram-
membrane, and lipopolysaccharide. staining procedure. The differences between these two
groups are reflected by fundamental differences in their
cell envelopes.
1. Outer membrane—The outer membrane is
chemically distinct from all other biological membranes.
It is a bilayered structure; its inner leaflet resembles in Cell Wall
composition that of the cell membrane, and its outer In addition to a plasma membrane, some eukaryotic cells
leaflet contains a distinctive component, a have a cell wall. Cells of fungi, algae, plants, and even
lipopolysaccharide (LPS). some protists have cell walls. Depending upon the type
of eukaryotic cell, cell walls can be made of a wide range
2. Lipopolysaccharide (LPS)—The LPS of gram- of materials, including cellulose (fungi and plants);
negative cell walls consists of a complex glycolipid, biogenic silica, calcium carbonate, agar, and
called lipid A, to which is attached a polysaccharide carrageenan (protists and algae); or chitin (fungi). In
made up of a core and a terminal series of repeat units. general, all cell walls provide structural stability for the
cell and protection from environmental stresses such as
3. Lipoprotein—Molecules of an unusual lipoprotein desiccation, changes in osmotic pressure, and traumatic
cross-link the outer membrane and peptidoglycan layers. injury
Lipoprotein is numerically the most abundant protein of
gram-negative cells (ca 700,000 molecules per cell). Its
Gram-Positive Bacteria
function (inferred from the behavior of mutants that lack
A gram-positive bacterium has a thick, multilayered cell
it) is to stabilize the outer membrane and anchor it to the
wall consisting mainly of peptidoglycan (150 to 500 Å)
peptidoglycan layer.
surrounding the cytoplasmic membrane (Figure 12-5).
The peptidoglycan is a meshlike exoskeleton similar in
4. The periplasmic space—The space between the
function to the exoskeleton of an insect. Unlike the
inner and outer membranes, called the periplasmic
exoskeleton of the insect, however, the peptidoglycan of
space, contains the peptidoglycan layer and a gel-like
the cell is sufficiently porous to allow diffusion of
solution of proteins.
metabolites to the plasma membrane. A new model for
peptidoglycan suggests that the glycan extends out from
Cell Walls of the Archaea
the plasma membrane like bristles that are cross-linked
The Archaea do not have cell walls like the Bacteria.
with short peptide chains. The peptidoglycan is essential
Some have a simple S-layer often composed of
for structure, replication, and survival in the normally
glycoproteins. Some Archaea have a rigid cell wall
hostile conditions in which bacteria grow.
composed of polysaccharides or a peptidoglycan called
pseudomurein. The pseudomurein differs from the
The peptidoglycan can be degraded by lysozyme.
peptidoglycan of bacteria by having l-amino acids rather
Lysozyme is an enzyme in human tears and mucus but
than d-amino acids and disaccharide units with an a-1--
is also produced by bacteria and other organisms.
>3 rather than a beta-1-->4 linkage. Archaea that have a
Lysozyme cleaves the glycan backbone of the
pseudomurein cell wall are gram-positive.
peptidoglycan. Without the peptidoglycan, the bacteria
succumb to the large osmotic pressure differences across
SUMMARY the cytoplasmic membrane and lyse. Removal of the cell
wall produces a protoplast that lyses unless it is Comparison of gram-positive and gram-negative
osmotically stabilized. bacterial cell walls.
A, A gram-positive bacterium has a thick peptidoglycan
The gram-positive cell wall may also include other layer that contains teichoic and lipoteichoic acids. B, A
components such as proteins, teichoic and lipoteichoic gram-negative bacterium has a thin peptidoglycan layer
acids, and complex polysaccharides (usually called C and an outer membrane that contains
polysaccharides). The M protein of streptococci and lipopolysaccharide, phospholipids, and proteins. The
protein A of S. aureus are covalently bound to the periplasmic space between the cytoplasmic and outer
peptidoglycan. Teichoic acids are water-soluble anionic membranes contains transport, degradative, and cell
polymers of polyol phosphates that are covalently linked wall synthetic proteins. The outer membrane is joined to
to the peptidoglycan and essential to cell viability. the cytoplasmic membrane at adhesion points and is
Lipoteichoic acids have a fatty acid and are anchored in attached to the peptidoglycan by lipoprotein links.
the cytoplasmic membrane. These molecules are
common surface antigens that distinguish bacterial Bacteria
serotypes and promote attachment to other bacteria and Bacteria are unicellular organisms. The cells are
to specific receptors on mammalian cell surfaces described as prokaryotic because they lack a nucleus.
(adherence). Teichoic acids are important factors in They exist in four major shapes: bacillus (rod shape),
virulence. Lipoteichoic acids are shed into the media and coccus (spherical shape), spirilla (spiral shape), and
the host, and although weaker, they bind to pathogen vibrio (curved shape). Most bacteria have a
pattern receptors and initiate innate protective host peptidoglycan cell wall; they divide by binary fission;
responses similar to endotoxin. and they may possess flagella for motility. The
difference in their cell wall structure is a major feature
Gram-Negative Bacteria used in classifying these organisms.
Gram-negative cell walls are more complex than gram-
positive cell walls, both structurally and chemically (see According to the way their cell wall structure stains,
Figure 12-2). Structurally, a gram-negative cell wall bacteria can be classified as either Gram-positive or
contains two layers external to the cytoplasmic Gram-negative when using the Gram staining. Bacteria
membrane. Immediately external to the cytoplasmic can be further divided based on their response to gaseous
membrane is a thin peptidoglycan layer that accounts for oxygen into the following groups: aerobic (living in the
only 5% to 10% of the gram-negative cell wall by presence of oxygen), anaerobic
weight. There are no teichoic or lipoteichoic acids in the
gram-negative cell wall. External to the peptidoglycan
layer is the outer membrane, which is unique to gram-
negative bacteria.
(living without oxygen), and facultative anaerobes (can
live in both environments).
According to the way they obtain energy, bacteria are
classified as heterotrophs or autotrophs. Autotrophs
make their own food by using the energy of sunlight or receptors that viruses use are molecules that are
chemical reactions, in which case they are called normally found on cell surfaces and have their own
chemoautotrophs. Heterotrophs obtain their energy by physiological functions. Viruses have simply evolved to
consuming other organisms. Bacteria that use decaying make use of these molecules for their own replication.
life forms as a source of energy are called saprophytes. Overall, the shape of the virion and the presence or
absence of an envelope tell us little about what disease
Viruses the virus may cause or what species it might infect, but
they are still useful means to begin viral classification
Unlike bacteria and fungi, viruses exist in a gray area
between “living” and “non-living” because of the Viruses are noncellular entities that consist of a nucleic
way they are structured and how they reproduce. acid core (DNA or RNA) surrounded by a protein coat.
Prokaryotes and eukaryotes can, for the most part, Although viruses are classified as microorganisms, they
reproduce on their own. Viruses, on the other hand, are not considered living organisms. Viruses cannot
depend on a host cell for nearly all of their functions, reproduce outside a host cell and cannot metabolize on
including reproduction. This is because viruses are their own. Viruses often infest prokaryotic and
essentially packets of genetic material and proteins— eukaryotic cells causing diseases.
they lack the cellular “machinery” required to
survive and reproduce on their own. \
Viruses are acellular, meaning they are biological
entities that do not have a cellular structure.
Therefore, they lack most of the components of cells,
such as organelles, ribosomes, and the plasma
membrane. They have A virion which is the virus
itself; it consists of a nucleic acid core, an outer
protein coating or capsid, and sometimes an
outer envelope made of protein and phospholipid
membranes derived from the host cell. The capsid is
made up of protein subunits called capsomeres. Viruses
may also contain additional proteins, such as enzymes
(depending on the type of virus) . The most obvious
difference between members of viral families is their
morphology, which is quite diverse. An interesting
feature of viral complexity is that host and virion
complexity are uncorrelated. Some of the most intricate
virion structures are observed in bacteriophages, viruses
that infect the simplest living organisms: bacteria.
Viruses don’t make cell membranes of their own, but Algae
some of them are coated with cell membranes from varies in chemical composition; substances commonly
their host. Just like the AIDS virus, HIV. In a single
found include cellulose, pectin, mannans, silicon
HIV, the “envelope” of glycoprotein and
phospholipid is the cell membrane that the virus dioxide, and calcium carbonate.
acquired when emerging from the infected host cell. Protozoa
Many viruses use some sort of glycoprotein to attach to The composition of the cell wall differs with each
their host cells via molecules on the cell called viral organism. In contrast, protozoa have no cell wall and
receptors. For these viruses, attachment is a requirement instead have a pellicle, which is a flexible, proteinaceous
for later penetration of the cell membrane, allowing covering. Protozoa are unicellular aerobic eukaryotes.
them to complete their replication inside the cell. The They have a nucleus, complex organ elles, and obtain
nourishment by absorption or ingestion through
specialized structures. They make up the largest group
of organisms in the world in terms of numbers, biomass,
and diversity. Their cell walls are made up of cellulose.
Protozoa have been traditionally divided based on their
mode of locomotion: flagellates produce their own food
and use their whip-like structure to propel forward,
ciliates have tiny hair that beat to produce movement,
amoeboids have false feet or pseudopodia used for
feeding and locomotion, and sporozoans are non-motile.
They also have different means of nutrition, which
groups them as autotrophs or heterotrophs.
Fungi
Most fungal species are multicellular organisms that
live on land rather than in water. Yeast and molds are
examples of fungi. Like algae, fungi possess cell walls.
Unlike algae cell walls, fungal cell walls contain chitin
rather than cellulose. Chitin is a tough, semitransparent
and complex molecule made up of repeating units of a
sugar called acetylglucosamine. It is better known as the
substance that makes up the hard outer coating of
crayfish, crabs, lobsters and some insects. Fungi
(mushroom, molds, and yeasts) are eukaryotic
cells (with a true nucleus). Most fungi are
multicellular and their cell wall is composed of
chitin. They obtain nutrients by absorbing
organic material from their environment
(decomposers), through symbiotic relationships
with plants (symbionts), or harmful
relationships with a host (parasites). They form
characteristic filamentous tubes called hyphae
that help absorb material. The collection of
hyphae is called mycelium. Fungi reproduce by
releasing spores.
