PROKARYOTIC
CELL STRUCTURE
Living
organisms are differentiated from nonliving matter by their (1) ability to
reproduce (2) ability to ingest or assimilate food and metabolize them for
energy and growth (3) ability to excrete waste products (4) ability to react to
changes in their environment (irritability) and (5) Susceptibility to mutation.
The living organisms include a variety of micro and macro organisms of
differing size , shape morphology, and behaviour. They include tiny bacteria,
protozoans, worms, plants and animals like man,
whale
and elephants.
Carlous
Linnaeus (1707-1778), the Swedish botanist was the first to introduce
nomenclature for plants and animals. Until 18thcentury only plant
and animal kingdoms were recognized. However some organisms are predominately
plant like, some animal like and some do not fall in both the groups. Therefore
it was felt a third kingdom was necessary. Haeckel (1866), a German zoologist
suggested a third kingdom Protista to include those organisms that are not
typically plants and animals. Bacteria, algae, fungi and protozoa are cellular
organisms placed under protista. Viruses are not cellular organisms and
therefore not classified as protists. Bacteria were lower protists while algae,
fungi and protozoa were higher protists. A satisfactory criteria to
differentiate bacteria, fungi and algae could not be made until the development
of electron microscope, which depicted the internal structure of these
organisms. The absence of membrane bound internal structures in bacteria and
their presence in fungi, algae, protozoa, plants and animal cells was taken as
criterion to differentiate prokaryote and eukaryote.
Whittakar
(1969) proposed five kingdoms based on three levels of cellular organization
and three principal modes of nutrition, photosynthesis, absorption and
ingestion. The prokaryotes lacking ingestive mode of nutrition are included in
the kingdom. Monera. In the kingdom protista unicellular eukaryotic
microorganisms representing all the three modes of nutrition are included. The
multicellular green plants and higher algae were placed in the kingdom plantae
while multinucleate higher fungi in the kingdom fungi and the multicellular
animals in the kingdom Animalea.
Bacteria
and cyanobacteria (the blue green algae) of monera, microalgae and protozoa of
protists and yeasts molds and fungi represent the microorganisms. Most of them
are invisible to the naked eye and requires magnification. The oratically a
black dot of 4mμ in diameter on a white background can be perceived by retina
of human eye but in reality an object of above 30mμ in size only will be
visible to the eyes and objects lesser than that requires magnification.
Prokaryotes
are organisms with primitive type of nucleus lacking a well-defined membrane a
less complex nuclear division than mitosis. The nuclear material is a DNA
molecule in prokaryotes compared to chromosomes of higher organisms. Eukaryotes
are
organisms with cells having true nuclei enclosed in a nuclear membrane and are
structurally more complex them prokaryotes. A varying degree of localization of
cellular functions in distinct membrane bound intracellular organelles like
nuclei, mitochondria
chloroplasts
etc. The cells of living organisms are either prokaryotic or eukaryotic in
nature and there is not any intermediate condition. The size, shape, morphology
and the internal cellular organizations are different in these two groups.
The size of the microorganisms varies
from unicellular tiny bacteria to large brown algae and mushroom. Bacteria are
unicellular, small 0.5-1.0mm in diameter, which multiply by binary fission. The
algae are photosynthetic simple organism withunicellular primitive types to aggregates of similar cells
and to large brown algae with complex structure. Protozoa are unicellu-lar,
most of them living freely in soil and water while a few cause disease of man
and animals.
The
rigid cell wall of the bacterium confers shape. Bacteria vary in shape from
spherical (Coccus) rods (Bacillus) and heli-cally curved rods (Spirillum). Most
bacteria possess a constant shape but some exhibit polymorphism (variety of
shape).
Bacterial
cells are arranged in a characteristic manner in a particular species. In cocci
the arrangement is known as diplo-cocci when cells divide in one plane and
remain attached in pairs, streptococci when divide in one plane and remain
attached to form chains; tetrococci, when they divide in two planes and form
group of four cells (tetrads), staphylococci when they divide in three planes
and form bundles, sarcinae when they divide in three planes in a regular manner
and form a cutridal arrangements.
Bacilli
are not arranged in such complex form as in cocci. Most of them occur singly or
in pairs (diplobacilli), form chains (streptobacilli) form trichomes, similar
to chains but with a larger area of contact between cells and lined side by
side like match sticks (palisade arrangement) at angles to one another.
Some
others form long branched multinucleate filaments called hypha as in fungi.
Hyphae ramify and collectively form mycelium. The curved bacteria are vibrioid
with less than one twist or turn of helical with one or more complete turns.
Rigid helical shape is in Spirilla and is flexible in spirochete.
Cell
wall is a very rigid structure that confers shape to the cell. This prevents expansion
of cells and bursting due to uptake of water as most bacteria live in
environments of higher osmotic pressure than that exists in cells (hypotonic
environments). A cell wall is common to almost all bacteria except in
mycoplasma that lacks typical cell wall and L-forms of bacteria like Streptobacillus
that are having walls but loose them
when grown in media con-taining sub lethal levels of cell wall synthesis
inhibiting antibiot-ics like penicillin. Mycoplasma lack cell wall permanently
and hence pleomorphic while L-forms of bacteria can revert back to walled
forms. The isolated cell walls without cellular constituents retain the
original contour of cells from which they are derived indicating that cell wall
confers shape. This is further strength-ened as the protoplast derived from any
type of cell cocci or bacilli show a spherical shape. Both eubacteria and
archaebacteria are grouped as Gram positive and Gram negative based on the wall
thickness. As the chemical composition of both eubacteria and archaeobacteria
differ it is only the thickness rather than the chemi-cal composition is the
key factor for Gram reaction.
Cell
wall constitutes 10-40% of cell. It is essential for growth and division. Cells
without walls (protoplasts) cannot grow and divide.
The
cell wall in eubacteria consists largely of an insoluble porous, cross-linked
polymer of enormous strength and rigidity viz., peptidoglycan (also called
murein).
This
is a bag shaped macromolecule and surrounds the cyto-plasmic membrane and found
only in prokaryotes. Although it is tough but in a dynamic state. It is a
polymer of N-acetyl glu-cosamine, N acetyl muramic acid, L-alanine, D-alanine,
D-glutamatic and a diamino acid (LL or meso diaminopimelic acid, L-lysine,
L-orthinine or L-diaminobutryic acid).
The
cell wall composition of archaeobacteria is different from eubacteria. Their
walls are composed of proteins, glycoproteins or polysaccharides. But in some
genera as Methanobacterium
the cell walls composed of pseudosuriein that have some superfecial
re-semblance to peptidoglycan but differs in chemical composition.
The
peptidoglycan constitutes about more than 50% of the dry weight of cells in
gram-positive eubacteria but only 10% in gram-negative bacteria. In addition to
peptidiglycan other sub-stances like polysaccharides in Streptococcus
pyrogenes teichoic
acids in Staphylococcus aureus, lipids as mycolic acids in Coryne-bacterium and Mycobacterium. The acid fast cord factor, a
my-colic acid derivative is toxic and plays a role in diseases due to Corynebacterium
diphtheriae and M.
tuberculosis.
The
wall of Gram negative consists of a thin peptidoglycan layer surrounded by an
outer membrane rich in lipids. The lipids in the wall constitute 11-12% of the
dry weight of the cells, The outer membrane is an impermeable barrier
preventing the escape of important enzymes from the periplasmic space between
the cy-toplasmic membrane and outer membrane. The outer membrane also prevents
external chemicals and enzymes that can destroy cells. Lysozyme, which dissolves
selectively the peptidoglycan can dam-age gram positive bacteria.
The
outer membrane, a bilayered structure consisting many of phospholipids,
proteins, and polysaccharides is anchored to the peptidoglycan layer by means
of Braun’s lipoprotein. The li-popolysaccharide (LPS) layer has toxic
properties and known as endotoxin. This occurs only in outer membrane and is
composed of lipid A, core polysaccharide and O antigen. The outer mem-brane is
impermeable to large molecules like protein but allow smaller molecules like
monosaccharides peptides and amino ac-ids through channels called porins.
Porins span the membrane and are specific for different kinds of small
molecules.
There
are several structures external to cell wall in bacteria, which vary in structure
and composition depending upon the type of bacteria. They are flagella, pili or
fimbriae, capsules, sheath, prosthecae and stalk. Flagella are locomotory
organs in bacteria, which vary in number and arrangement. Some bacteria do not
have flagella.
Flagella
are hair like helical appendages 0.01 – 0.02 nm in diameter the flagellar
arrangements vary with the organisms. It may be polar if the flagella are at
one or both the ends or lateral if they are arranged on sides. They protrude
through the cell wall. A flagellum is composed of a basal body a short hook and
a helical filament longer than the cell. The basal body is associated with
cytoplasmic membrane and cell wall.
Bacteria
swim by rotating their helical flagella similar to cork screw. Bacteria with
polar flagella swim in a back and forth fash-ion. Those with lateral flagella
swim in a more complicated man-ner. Removal of flagella from a flagellate
bacterium will not result in death of bacterium and only motility will be
affected Spiro-chetes, the helical bacteria, swim even in viscous media,
without any external flagella. They have flagella like structure within the
cell located just beneath the cell envelope. They are known as periplasmic
flagella (also called endoflagella, axial filament). Spiroplasmas are also
helical in shape and swim in viscous media without even periplasmic flagella.
Some
bacteria like Cytophaga exhibit a gliding motility, which is a slow sinuous
flexing motion. This occurs when the cells come in contact with solid surface.
Pili
are short, hollow, non helical and filamentous append-ages. They are thinner
than flagella but more in number than fla-gella. They are found in both motile
and non motile bacteria and hence not involved in motility.
F
pilus (sex pilus), a type of pilus serves as port of entry for genetic material
during bacterial mating. Some pili in pathogenic bacteria serve as attachment
with host cells in human beings fa-cilitating infection without being washed
off easily by mucous.
Capsules
A viscous substance forming a
covering layer around the cell is found in some bacteria and is known as
capsule. If it is too thin it is called as microcapsule. If it is loose and
many cells are embedded in a matrix it is known as slime. The capsular material
is
not water soluble in many bacteria but in some it is highly water soluble, thus
making the medium in which they grow more viscous. Capsular material is
primarily polysaccharide in most bacteria. It may be a homopolysaccharide, made
up of a single kind of sugar, synthesized outside the cell from disaccharides.
The capsule of S.mutons is a glucan (a glucose polymer)
synthesized from sucrose. Capsules composing of several kinds of sugars are
termed
heteropolysaccharides.
These are synthesized from sugars within the cell, transported and polymerized
outside the cell. The capsule of Klebsiella pneumonia is a
heteropolysaccharide. The capsule of some bacteria is polypeptides. The capsule
of anthrax organism Bacillus anthracis is a polymer of D-glutamic acid.
Sheath
is a hollow tube that encloses cells in the form of chains or trichomes. This
is present in some bacteria living in fresh water and marine environment. The
cells some times move out of sheath. In a few cases the sheath is strengthened
by deposition of ferric and manganese hydroxides.
Aerobic
bacteria in fresh water and marine environment possess prosthecae, which
increases the surface area of cells for nutrient absorption from the dilute
aquatic environment. They are semirigid extension of cell wall and cytoplasm
membrane and smaller than the cell. Some bacteria have single prostheca (Caulobacter) and others have more than one (Stellar and Ancalomicrobium).
Stalks
are also found in some bacteria like Gallionella or Planctomyces. They are non-living ribbon like or tubular append-ages
that are excreted by cell. These stalks aid in attachments of cells to surface.
The
structures internal to cell are cytoplasmic membrane, protoplast, intracellular
membranes, the cytoplasm, cytoplasmic inclusions and nuclear material, the DNA.
The
cytoplasmic membrane is immediately beneath the cell wall and is about 7.5nm
thick. It is made up of phospholipids (about20-30 percent) forming a bilayer to
which both integral pro-teins and peripheral proteins are held. The membrane
has fluidity owing due to its lipid matrix and this allows components to move
laterally.
The
phospholipids of eubacteria and archaeobacteria differ in composition. The
phospholipids of eubacteria are phosphoglycerides. In this straight chain fatty
acids are linked to glycerol by ester linkage. In archaeobacteria, the lipids
are polyisoprenoid branched chain lipids. In this phytanols, (long chain
branched alcohols) are ether linked glycerols.
The
cytoplasmic membrane is a barrier for penetration of most of water soluble
molecules. But the small molecules like nutrients and waste products are
transported across the membrane by specific proteins. The membrane also
contains various enzymes of respiratory metabolism and synthesis of cell wall
components and capsule. It is also the site of generation of proton motive
force that drives ATP synthesis, nutrition, transport system and flagellar
motility. The damage to membrane by physical or chemical agent lead to death of
cells.
The
cytoplasmic membrane and the cell material bounded by it are called protoplast.
The bacterial cell minus the cell wall is the protoplast. Protoplasts of gram
positive bacteria can be pre-pared by dissolving the cell wall by lysozyme or
by growing the bacteria in penicillin containing media. Penicillin prevents the
syn-thesis and formation of cell wall. Protoplasts thus prepared have to be
suspended in an isotonic medium, other wise the bacteria living in hypotonic
environments tend to absorb water and burst.
In Gram-negative bacteria lysozyme
treatment may destroy the cell wall. The outer membrane remains with the
cytoplasmic membrane enclosing the cell content. Such type of protoplasts with
the outer membrane is called as spheroplast.
The
bacteria that lack cell wall like mycoplasma are similar to protoplasts but
they are parasites of animals, plants or arthropods and hence live in
osmotically favourable or isotonic environments.
Bacteria
are prokaryotes that do not contain any membrane bound organelles inside the
cells. But bacteria have specialized invagination of cytoplasmic membrane that
increase the surface area for certain functions. Mesosomes are convoluted
tubules or vesicles formed by membranous invagination in bacteria. Central
mesosomes are located near the middle of the cell and penetrates deep into the
cytoplasm. It seem to be attached to the nuclear ma-terial. Peripheral
mesosomes shallowly penerate into the cytoplasm and seen to be invalid in
export of exocellular enzymes.
The
intracellular membrane is extensive in all phototrophic bacteria,
chemoautotrophs and in methane oxidizing bacteria. In phototrophic bacteria
they are the sites of photosynthesis as the increased surface area increase the
light absorbing pigments.
Thylakoids
are special intracellular membranes that occur in cyanobacteria but they are
separate from cytoplasmic membrane.
The
cytoplasmic membrane bound the cytoplasm. the cyto-plasm consists of a
cytoplasmic area, a chromatinic area and con-sists of Ribosome. Ribosomes are
macromolecular RNA protein bodies and are the sites of protein synthesis. The
chromatinic area is rich in DNA. The fluid proteins contain the dissolved
substances.
Ribosomes
of prokaryotes have a sedimentation coef of 70 Svederberg units (70S) and are
composed of two subunits of 50S and 30S. On the other hand ribosomes of
eukaryotes have a sedimentation coefficient of 80S and are composed of 60S and
40S subunits.
Cytoplasmic
inclusions are concentrated deposits of certain substances. Volutin granules or
metachromatic granules are polyphosphates deposits. It is a reserve of
phosphate. Poly-B-hy- droxy butyrate is a chloroform soluble lipid like
material and serve as carbonTheyandareenergyfoundsourceina.robic bacteria.
Polysaccharide
granules viz glycogen is present as inclusion. El-emental sulfur accumulates in
certain bacteria growing in envi-ronments rich in hydrogen sulphide.
Bacteria in aquatic habitats have
gas vacuoles to provide buoyancy. Gas vacuoles have water impermeable boundary
but permeable to dissolved gases which fill the cavity. Bacteria do not have a
nucleus with a nuclear membrane. The nuclear material is only a single circular
DNA molecule. This is called as nucleoid, the chromatin body, the nuclear
equivalent or functional chromosome.
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