Cell wall is the outermost layer of most cells that protects the bacterial cell and gives it shape.

One exception is Mycoplasma which lacks cell wall. Bacterial cell walls are made of
peptidoglycan which is made from polysaccharide chains cross-linked by unusual peptides
containing D-amino acids. Bacterial cell walls are different from the cell walls of plants and
fungi which are made of cellulose and chitin, respectively. The cell wall of bacteria is also
distinct from that of Archaea, which do not contain peptidoglycan. The cell wall is essential
to the survival of many bacteria. The antibiotic penicillin is able to kill bacteria by preventing
the cross-linking of peptidoglycan and this causes the cell wall to weaken and lyse.
Lysozyme enzyme can also damage bacterial cell walls. There are broadly speaking two
different types of cell wall in bacteria, called Gram-positive and Gram-negative (Figure ).
The names originate from the reaction of cells to the Gram stain, a test long-employed for the
classification of bacterial species. Gram-positive bacteria possess a thick cell wall containing
many layers of peptidoglycan and teichoic acids. In contrast, Gram-negative bacteria have a
relatively thin cell wall consisting of a few layers of peptidoglycan surrounded by a second
lipid membrane containing lipopolysaccharides and lipoproteins. These differences in
structure can produce differences in property as antibiotic susceptibility. For example
vancomycin can kill only Gram-positive bacteria and is ineffective against Gram-negative
pathogens, such as Pseudomonas aeruginosa or Haemophilus influenzae.
A: Gram positive cell wall B: Gram negative cell wall

It is the extracellular structure surrounding plasma membrane. The cell wall is composed of
cellulose, hemicellulose, pectin and in many cases lignin, is secreted by the protoplast on the
outside of the cell membrane. This contrasts with the cell walls of fungi (which are made of
chitin), and of bacteria, which are made of peptidoglycan. An important function of the cell
wall is that it controls turgity. The cell wall is divided into the primary cell wall and the
secondary cell wall. The Primary cell wall: extremely elastic and the secondary cell wall forms
around primary cell wall after growth are complete.

The plant cell wall is a defining feature that distinguishes plant cells from animal cells, which
lack this rigid outer layer. It is a non-living structure synthesized by the living protoplast, and
a cell without its wall is referred to as a protoplast. Composed primarily of cellulose,
hemicellulose, pectin, and proteins, the plant cell wall provides mechanical support and
protection to the cell. In contrast, fungal cell walls are made of chitin, while bacterial cell walls
consist of protein-lipid-polysaccharide complexes. The cell wall not only safeguards the
plasma membrane but also determines the shape of plant cells, leading to the classification of
various cell types such as parenchymatous and collenchymatous cells based on wall structure
and function.
Chemically speaking, the plant cell wall is composed of a variety of polysaccharides
(carbohydrates), lipids, proteins and mineral deposits, all exhibiting distinct staining reactions
Table 5-5.

Chemical nature and staining reaction of various components of plant cell wall.
Eukaryotic Cell wall
The cell wall is complex in nature and is differentiated in the following layers :
(i) Primary cell wall; (ii) Secondary cell wall; (iii) Tertiary cell wall.
• Primary cell wall. The first formed cell wall is known as primary cell wall. It is the
outermost layer of the cell and in the immature meristematic and parenchymatous cells
it forms the only cell wall. The primary cell is comparatively thin and permeable.
Certain epidermal cells of the leaf and the stem also possess the cutin and cutin waxes
which make the primary cell wall impermeable. The primary cell wall of the yeast and
the fungi composed of the chitin.
• Secondary cell wall. The primary cell wall is followed by secondary cell wall. The
secondary cell wall is thick, permeable and lies near the plasma membrane of the
tertiary cell wall, if the latter occurs. It is composed of three concentric layers (S1, S2
and S3) which occur one after another. Chemically the secondary cell wall is composed
of compactly arranged macrofibrils of the cellulose, in between which sometimes
occurs lignin as an inter-fibrillar material.
• Tertiary cell wall. In certain plant cells, there occurs another cell wall beneath the
secondary cell wall which is known as tertiary cell wall. The tertiary cell wall differs
from the primary and secondary cell wall in its morphology, chemistry and staining
properties. Besides the cellulose, the tertiary cell wall consists of another chemical
substance known as the xylan.
Middle lamella. The cells of plant tissues generally remain cemented together by an inter-
cellular matrix known as the middle lamella. The middle lamella is mainly composed of the
pectin, lignin and some proteins.

Prokaryotic cell wall

It is important to note that not all bacteria have a cell wall. Having said that though, it is also
important to note that most bacteria (about 90%) have a cell wall and they typically have one
of two types: a gram positive cell wall or a gram negative cell wall. The two different cell
wall types can be identified in the lab by a differential stain known as the Gram stain.
Developed in 1884, it’s been in use ever since. Originally, it was not known why the Gram
stain allowed for such reliable separation of bacterial into two groups. Once the electron
microscope was invented in the 1940s, it was found that the staining difference correlated with
differences in the cell walls.

A cell wall, not just of bacteria but for all organisms, is found outside of the cell membrane.
It’s an additional layer that typically provides some strength that the cell membrane lacks, by
having a semi-rigid structure.

Both gram positive and gram negative cell walls contain an ingredient known
as peptidoglycan (also known as murein). This particular substance hasn’t been found
anywhere else on Earth, other than the cell walls of bacteria. But both bacterial cell wall types
contain additional ingredients as well, making the bacterial cell wall a complex structure
overall, particularly when compared with the cell walls of eukaryotic microbes. The cell walls
of eukaryotic microbes are typically composed of a single ingredient, like the cellulose found
in algal cell walls or the chitin in fungal cell walls.

The bacterial cell wall performs several functions as well, in addition to providing overall
strength to the cell. It also helps maintain the cell shape, which is important for how the cell
will grow, reproduce, obtain nutrients, and move. It protects the cell from osmotic lysis, as the
cell moves from one environment to another or transports in nutrients from its surroundings.
Since water can freely move across both the cell membrane and the cell wall, the cell is at risk
for an osmotic imbalance, which could put pressure on the relatively weak plasma membrane.
Studies have actually shown that the internal pressure of a cell is similar to the pressure found
inside a fully inflated car tire. That is a lot of pressure for the plasma membrane to withstand!
The cell wall can keep out certain molecules, such as toxins, particularly for gram negative
bacteria. And lastly, the bacterial cell wall can contribute to the pathogenicity or disease –
causing ability of the cell for certain bacterial pathogens.

Structure of Peptidoglycan

Let us start with peptidoglycan, since it is an ingredient that both bacterial cell walls have in
common. Peptidoglycan is a polysaccharide made of two glucose derivatives, N-
acetylglucosamine (NAG) and N-acetylmuramic acid (NAM), alternated in long chains. The
chains are cross-linked to one another by a tetrapeptide that extends off the NAM sugar unit,
allowing a lattice-like structure to form. The four amino acids that compose the tetrapeptide
are: L-alanine, D-glutamine, L-lysine or meso-diaminopimelic acid (DPA), and D-alanine.
Typically only the L-isomeric form of amino acids are utilized by cells but the use of the mirror
image D-amino acids provides protection from proteases that might compromise the integrity
of the cell wall by attacking the peptidoglycan. The tetrapeptides can be directly cross-
linked to one another, with the D-alanine on one tetrapeptide binding to the L-lysine/ DPA on
another tetrapeptide. In many gram positive bacteria there is a cross-bridge of five amino acids
such as glycine (peptide interbridge) that serves to connect one tetrapeptide to another. In
either case the cross-linking serves to increase the strength of the overall structure, with more
strength derived from complete cross-linking, where every tetrapeptide is bound in some way
to a tetrapeptide on another NAG-NAM chain.
While much is still unknown about peptidoglycan, research in the past ten years suggests that
peptidoglycan is synthesized as a cylinder with a coiled substructure, where each coil is cross-
linked to the coil next to it, creating an even stronger structure overall.

Peptidoglycan Structure.

Functions of Bacterial Cell Walls

• Adds overall strength to the cell structure.
• Maintains cell shape, essential for growth, movement, reproduction, and nutrient
intake.
• Protects against osmotic lysis by managing internal pressure and water balance.
• Internal pressure can match that of a fully inflated car tire—the cell wall helps
withstand this!
• Filters out harmful molecules like toxins—especially effective in Gram-negative
bacteria.
• Enhances pathogenicity in some bacteria, contributing to their ability to cause disease.

Glycocalyx

The cell membrane of bacteria, epithelia is surrounded by a pericellular matrix, which is known
as Glycocalyx. It is a carbohydrate-enriched coating that covers both eukaryotic cells and
prokaryotic cells, particularly bacteria.
• It is made of glycoprotein and glycolipid.
• Martinez and Palomo first discovered a cell coat around the animal cells in 1970,
it is termed as the glycocalyx.
• The external surface of plasma membranes in most animal epithelial cells
contains a fuzz-like coating which is made of several carbohydrate moieties of
membrane glycolipids and glycoproteins. It assists as backbone molecules for
 maintenance. The carbohydrate portion of the glycolipids helps in cell–cell
recognition, communication, and intercellular adhesion.
• The body uses glycocalyx as an identifier to differentiate between its own normal
cells and transplanted tissues, diseased cells, or invading organisms.
• The glycocalyx performs a significant role in the regulation of endothelial
vascular tissue, including the modulation of red blood cell volume in capillaries.
• During the embryonic development, Glycocalyx permits the cells to adhere to
each other and control the movement of cells.
• The slime layer at the surface of a fish is a great example of glycocalyx.
• The term glycocalyx came from the Greek word glykys = sweet, kalyx = husk
which means “sugar coat”.
• It is a highly charged layer of membrane-bound biological macromolecules that
are connected to a cell membrane.
• It functions as a barrier within a cell and its surrounding.
• It helps in cell-cell interactions and protects the cell membrane from physical
forces and stresses. It also maintains the integrity of the membrane.
• They also help in the development and progression of many diseases.

Glycocalyx composition and structure

• It is made up of glycosaminoglycans, proteoglycans, and other glycoproteins
bearing acidic oligosaccharides and terminal sialic acids.
• It is thick approximately 500-2000nm in diameter hydrated gel-like coating on the
luminal exterior of the vascular endothelium.
• The term means “sweet husk”, which referrs to the high polysaccharide content.
• Most of the associated proteins of Glycocalyx are transmembrane and can be
connected to the cytoskeleton. This connection helps in the transduction of signals
from the external to the internal parts of a cell.
• It contains different types of polyanionic macromolecules such as Glycoproteins,
Polysaccharides, Proteoglycans, Glycosaminoglycans (For example heparan
sulfate, chondroitin sulfate, and hyaluronic acid). These are helps to repel
circulating platelets
• Based on the local microenvironment the exact composition is varied.
• At any granted time, it further comprises constituents of the routine molecular
traffic which moves within it or lodges within it, such as Plasma proteins, Enzymes
and enzyme inhibitors, Growth factors, Cytokines, Amino acids, Cations, Water.
• It is fragile but self-repairing. It can restore itself within less than 1 second by
adsorbing plasma constituents
Glycocalyx function
• Protection: It Cushions the plasma membrane and defends it from
chemical damage
• Immunity to infection: It facilitates the immune system to identify and
selectively attack the outside organisms.
• Inflammation regulation: In blood vessels, the rolling/binding of
leukocytes in healthy states are prevented by the Glycocalyx coating on
endothelial walls.
• Defense against cancer: In cancer cells, the changes in the glycocalyx
permit the immune system to identify and neutralize them.
• Fertilization: It allows the sperm to identify and adhere to eggs.
• Transplant compatibility: Forms the basis for compatibility of blood
transfusions, tissue grafts, and organ transplants.
 • Embryonic development: Glycocalyx guides embryonic cells to their
targets in the body
• Cell adhesion: It held cells together so that tissues do not fall apart.
Glycocalyx – Vascular Endothelial Tissue
• Glycocalyx is located - apical surface of vascular endothelial cells which line
the lumen.
• When vessels are stained with cationic dyes such as Alcian blue stain, transmission
electron microscopy shows a small, irregularly shaped layer extending approximately
50–100 nm into the lumen of a blood vessel.
• Another study used osmium tetroxide staining during freeze substitution, and showed
that the endothelial glycocalyx could be up to 11 μm thick.
• It is present throughout a diverse range of microvascular beds (capillaries) and
macrovessels (arteries and veins).
• Endothelial nitric oxide synthase (endothelial NOS)
• Extracellular superoxide dismutase (SOD3)
• Angiotensin converting enzyme
• Antithrombin-III
• Lipoprotein lipase
• Apolipoproteins
• Growth factors
• Chemokines
• The enzymes and proteins listed above serve to reinforce the glycocalyx barrier against
vascular and other diseases.
• Another main function of the glycocalyx within the vascular endothelium is that it
shields the vascular walls from direct exposure to blood flow, while serving as a
vascular permeability barrier.
• Its protective functions are universal throughout the vascular system, but its relative
importance varies depending on its exact location in the vasculature.
• In microvascular tissue, the glycocalyx serves as a vascular permeability barrier by
inhibiting coagulation and leukocyte adhesion.
• Leukocytes must not stick to the vascular wall because they are important components
of the immune system that must be able to travel to a specific region of the body when
needed.
• In arterial vascular tissue, the glycocalyx also inhibits coagulation and leukocyte
adhesion, but through mediation of shear stress-induced nitric oxide release.
• Another protective function throughout the cardiovascular system is its ability to affect
the filtration of interstitial fluid from capillaries into the interstitial space

Cell – Cell Interactions

Cell–cell interaction refers to the direct interactions between cell surfaces that play a crucial
role in the development and function of multicellular organisms. These interactions allow cells
to communicate with each other in response to changes in their microenvironment. This ability
to send and receive signals is essential for the survival of the cell. Interactions between cells
can be stable such as those made through cell junctions. These junctions are involved in the
communication and organization of cells within a particular tissue. Others are transient or
temporary such as those between cells of the immune system or the interactions involved in
tissue inflammation. These types of intercellular interactions are distinguished from other types
such as those between cells and the extracellular matrix. The loss of communication between
cells can result in uncontrollable cell growth and cancer.
Exocytosis - exiting the cell
Exocytosis is a process used by the cell to take out its trash and to incorporate proteins into the
cell membrane. During exocytosis, the phospholipid bilayer of the cell membrane surrounds
the waste proteins, creating a bubble-like structure called a vesicle. Vesicles are frequently
used in the cell for transportation of molecules across the cell membrane.

Waste proteins
A slightly different process occurs for waste products being ejected out of the cell, instead of
proteins being incorporated into the cell membrane. Once the vesicle has enclosed the waste
proteins on the inside of the cell, it moves towards the cell membrane. The vesicle merges with
the cell membrane, opening the bubble-like structure and ejecting the contents in the
environment surrounding the cell.

Proteins destined for the cell membrane

Exocytosis is also used to integrate new proteins into the cell membrane. In this process, the
new protein is formed inside the cell, and migrates to phospholipid bilayer of the vesicle. The
vesicle, containing the new protein as a part of the phospholipid bilayer, fuses with the cell
membrane. This allows the protein to be directly integrated into the cell membrane when the
vesicle, in the same way as with waste proteins, fuses and opens with the cell membrane.

Endocytosis - bringing in the goods

Endocytosis is the opposite process of exocytosis. Endocytosis brings molecules into the cell.
These molecules are important for the survival of the cell, such as glucose. There are three
different styles of endocytosis: 1) phagocytosis, 2) pinocytosis, and 3) receptor-mediated
endocytosis.
Phagocytosis
Phagocytosis is the process similar to eating, where the cell engulfs a molecule in order to move
it to the interior of the cell. The process starts by the molecule binding to specific receptors on
the surface of the cell membrane, triggering the cell membrane to reshape, surrounding the
molecule. The receptors allow this process to be specific, controlling what can enter the cell.
Then, the two ends of the cell fuse, creating a vesicle that surrounds the molecule. Eventually
the membrane around the molecule will be digested and its contents will be used! For example,
white blood cells recognize pathogens, such as viruses or bacterial cells, outside of the cell and
will use phagocytosis to bring it in to destroy it!

Pinocytosis
If phagocytosis is how the cell eats, then pinocytosis is how the cell drinks. Pinocytosis engulfs
dissolved ions and other solutes in the liquid medium surrounding the cell. This is different
than phagocytosis, which brings full, undissolved or insoluble molecules into the cell. The
distortion of the cell membrane to engulf the dissolved solutes is similar to that of phagocytosis.
Another important distinction is that pinocytosis is not specific to what is carried into the cell,
whereas phagocytosis can be highly specific. The liquid medium outside the cell is always
filled with dissolved particles and solutes that are handy for the cell, so the cell doesn’t need
this process to be specific.

Receptor-mediated endocytosis
Receptor-mediated endocytosis is very specific with respect to what is imported into the cell.
It’s actually a bit like a lock-and-key system. There are receptors embedded in the cell
membrane that, when bound by molecules with an exact match in shape, size, or other physical
attribute, will allow the molecule to enter into the cell through the same engulfment process as
phagocytosis or pinocytosis.

Cell junctions
There are many different ways that cells can connect to each other. The three main ways for
cells to connect with each other are: gap junctions, tight junctions, and desmosomes. These
types of junctions have different purposes, and are found in different places.
Intercellular Junctions

The extracellular matrix allows cellular communication within tissues through conformational
changes that induce chemical signals, which ultimately transform activities within the cell.
However, cells are also capable of communicating with each other via direct contact through
intercellular junctions.

There are some differences in the ways that plant and animal cells communicate directly.
Plasmodesmata are junctions between plant cells, whereas animal cell contacts are carried out
through tight junctions, gap junctions, and desmosomes.

Gap Junctions
Gap junctions are essentially tubes that join two cells together. These tubes create a connection
that allows for the transport of water and ions to and from the connecting cells. The tubes also
help to spread electrochemical signals that are produced by action potentials that occur in the
nervous system (neurons) and in cardiac cells that make your heart beat. Gap junctions have a
pretty important job!

Lastly, similar to plasmodesmata in plant cells, gap junctions are the third type of direct
junction found within animal cells. These junctions are channels between adjacent cells that
allow for the transport of ions, nutrients, and other substances that enable cells to communicate.
Structurally, however, gap junctions and plasmodesmata differ. Gap junctions develop when a
set of six proteins (called connexins) in the plasma membrane arrange themselves in an
elongated doughnut-like configuration called a connexon. When the pores (“doughnut holes”)
of connexons in adjacent animal cells align, a channel between the two cells forms. Gap
junctions are particularly important in cardiac muscle. The electrical signal for the muscle to
contract is passed efficiently through gap junctions, which allows the heart muscle cells to
contract in tandem.

Junctions in Plant Cells

In general, long stretches of the plasma membranes of neighboring plant cells cannot touch one
another because they are separated by the cell wall that surrounds each cell. How then can a
plant transfer water and other soil nutrients from its roots, through its stems, and to its leaves?
This transport primarily uses the vascular tissues (xylem and phloem); however, there are also
structural modifications called plasmodesmata (singular: plasmodesma) that facilitate direct
communication in plant cells. Plasmodesmata are numerous channels that pass between cell
walls of adjacent plant cells and connect their cytoplasm; thereby, enabling materials to be
transported from cell to cell, and thus throughout the plant.

Plasmodesmata: A plasmodesma is a channel between the cell walls of two adjacent plant
cells. Plasmodesmata allow materials to pass from the cytoplasm of one plant cell to the
cytoplasm of an adjacent cell.

Tight Junctions
Tight junctions are different from gap junctions because they are the connections that form
when cells are squished up against one another. In this case, the cell membranes are connected,
but the contents of each cell are not connected in any way. There are no tubes here, but there
is an impermeable layer in between the cells. These types of cell connections are useful in
places that need to contain certain fluids, like in the bladder, the intestines or the kidneys.
Imagine if you didn’t have a watertight seal in those connections! Fluids like your urine would
be circulating through your body!

Junctions in Animal Cells

Communication between animal cells can be carried out through three types of junctions. The
first, a tight junction, is a watertight seal between two adjacent animal cells. The cells are held
tightly against each other by proteins (predominantly two proteins called claudins and
occludins). This tight adherence prevents materials from leaking between the cells. These
junctions are typically found in epithelial tissues that line internal organs and cavities and
comprise most of the skin. For example, the tight junctions of the epithelial cells lining your
urinary bladder prevent urine from leaking out into the extracellular space.
Tight Junctions: Tight junctions form watertight connections between adjacent animal cells.
Proteins create tight junction adherence.
Desmosomes
Finally, desmosomes are quite different from gap junctions and tight junctions. With
desmosomes, cell membranes are connected by thread like substances that connect the cells
across the space in between cells. Much like tight junctions, desmosomes physically hold the
cells together, but do not allow fluids or materials to pass from the inside of one cell to the
next. These connections are also attached to the scaffolding of the cell, called the cytoskeleton,
to help with structural support. The space in between the cells allows for water and solutes to
flow freely between each cell without compromising the connection. This is convenient for
areas of our body that experience high stress like in our skin or our intestines because the space
in between the cells offer flexibility that the other junctions can’t.

Also found only in animal cells are desmosomes, the second type of intercellular junctions in
these cell types. Desmosomes act like spot welds between adjacent epithelial cells, connecting
them. Short proteins called cadherins in the plasma membrane connect to intermediate
filaments to create desmosomes. The cadherins join two adjacent cells together and maintain
the cells in a sheet-like formation in organs and tissues that stretch, such as the skin, heart, and
muscles.
Desmosomes: A desmosome forms a very strong spot weld between cells. It is created by the
linkage of cadherins and intermediate filaments.