There Are Two Types of Cells: Prokaryotes and Eukaryotes

All modern organisms are based on the same morphological unit, the cell.
There are two major classifications of cells: the eukaryotes (Greek: eu, good or true
+ karyon, kernel or nut), which have a membrane-enclosed nucleus encapsulating
their DNA; and the prokaryotes (Greek: pro, before), which lack a nucleus.
Prokaryotes, comprising the various types of bacteria, have relatively simple
structures and are almost all unicellular (although they may form filaments or
colonies of independent cells). Eukaryotes, which are multicellular as well as
unicellular, are vastly more complex than prokaryotes. (Viruses are much simpler
entities than cells and are not classified as living because they lack the metabolic
apparatus to reproduce outside their host cells.) Prokaryotes are the most numerous
and widespread organisms on the earth. This is because their varied and often highly
adaptable metabolisms suit them to an enormous variety of habitats. Prokaryotes
range in size from 1 to 10 m and have one of three basic shapes (Figure 1):
spheroidal (cocci), rodlike (bacilli), and helically coiled (spirilla). Except for an
outer cell membrane, which in most cases is surrounded by a protective cell wall,
nearly all prokaryotes lack cellular membranes. However, the prokaryotic
cytoplasm (cell contents) is by no means a homogeneous soup.
Prokaryotes comprise two separate but related groups: the bacteria (or
eubacteria) and the archaea (or archaebacteria). These two distinct groups of
prokaryotes diverged early in the history of life on Earth. The bacteria are the
commonly encountered prokaryotes in soil, water and living in or on larger
organisms, and include Escherichia coli and the Bacillus species, as well as the
cyanobacteria (photosynthetic blue-green algae). The archaea mainly inhabit
unusual environments such as salt brines, hot acid springs, bogs and the ocean
depths, and include the sulfur bacteria and the methanogens, although some are
found in less hostile environments. The best characterized prokaryote is Escherichia
coli, a 2 m by 1 m rodlike bacterium that inhabits the mammalian colon.

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 Figure 1a. Scale drawings of some prokaryotic cells.

Figure 1b. Prokaryote cell structure.

Eukaryotic cells are generally 10 to 100 m in diameter and thus have a

thousand to a million times the volume of typical prokaryotes. It is not size, however,
but a profusion of membrane-enclosed organelles that best characterizes eukaryotic
cells (Figure 2). In addition to a nucleus, eukaryotes have an endoplasmic
reticulum, the site of synthesis of many cellular components, some of which are
subsequently modified in the Golgi apparatus. The bulk of aerobic metabolism
takes place in mitochondria in almost all eukaryotes, and photosynthetic cells
contain chloroplasts. Other organelles, such as lysosomes and peroxisomes,
perform specialized functions. Vacuoles, which are more prominent in plant cells,
usually function as storage depots. The cytosol (the cytoplasm minus its membrane

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bounded organelles) is organized by the cytoskeleton, an extensive array of
filaments that also gives the cell its shape and the ability to move. The various
organelles that compartmentalize eukaryotic cells represent a level of complexity
that is largely lacking in prokaryotic cells. Nevertheless, prokaryotes are more
efficient than eukaryotes in many respects. Prokaryotes have exploited the
advantages of simplicity and miniaturization. Their rapid growth rates permit them
to occupy ecological niches in which there may be drastic fluctuations of the
available nutrients. In contrast, the complexity of eukaryotes, which renders them
larger and more slowly growing than prokaryotes, gives them the competitive
advantage in stable environments with limited resources. It is therefore erroneous to
consider prokaryotes as evolutionarily primitive compared to eukaryotes. Both types
of organisms are well adapted to their respective lifestyles.

Figure 2a. The structure of the pant cell.

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 Figure 2b. Diagram of a typical animal cell accompanied by electron
micrographs of its organelles.

Prokaryote Cell Structure

Cell structure: Prokaryotes generally range in size from 0.1 to 10 μm, and have one
of three basic shapes: spherical (cocci), rod-like (bacilli) or helically coiled (spirilla).
Like all cells, a prokaryotic cell is bounded by a plasma membrane that completely
encloses the cytosol and separates the cell from the external environment. The
plasma membrane, which is about 8 nm thick, consists of a lipid bilayer containing
proteins. Although prokaryotes lack the membranous subcellular organelles
characteristic of eukaryotes, their plasma membrane may be infolded to form
mesosomes. The mesosomes may be the sites of deoxyribonucleic acid (DNA)
replication and other specialized enzymatic reactions. In photosynthetic bacteria, the
mesosomes contain the proteins and pigments that trap light and generate adenosine
triphosphate (ATP). The aqueous cytosol contains the macromolecules [enzymes,
messenger ribonucleic acid (mRNA), transfer RNA (tRNA) and ribosomes], organic
compounds and ions needed for cellular metabolism. Also, within the cytosol is the
prokaryotic ‘chromosome’ consisting of a single circular molecule of DNA
which is condensed to form a body known as the nucleoid.

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Bacterial cell walls: To protect the cell from mechanical injury and osmotic
pressure, most prokaryotes are surrounded by a rigid 3–25 nm thick cell wall. The
cell wall is composed of peptidoglycan, a complex of oligosaccharides and proteins.
The oligosaccharide component consists of linear chains of alternating
N-acetylglucosamine (GlcNAc) and N-acetylmuramic acid (NAM) linked β (1–4).
Attached via an amide bond to the lactic acid group on NAM is a D-amino acid
containing tetrapeptide. Adjacent parallel peptidoglycan chains are covalently cross-
linked through the tetrapeptide side-chains by other short peptides. The extensive
cross-linking in the peptidoglycan cell wall gives it its strength and rigidity. The
presence of D-amino acids in the peptidoglycan renders the cell wall resistant to the
action of proteases which act on the more commonly occurring L-amino acids, but
provides a unique target for the action of certain antibiotics such as penicillin.
Penicillin acts by inhibiting the enzyme that forms the covalent cross-links in the
peptidoglycan, thereby weakening the cell wall. The β(1–4) glycosidic linkage
between NAM and GlcNAc is susceptible to hydrolysis by the enzyme lysozyme
which is present in tears, mucus and other body secretions. Bacteria can be classified
as either Gram-positive or Gram-negative depending on whether or not they take
up the Gram stain. Gram-positive bacteria (e.g. Bacillus polymyxa) have a thick (25
nm) cell wall surrounding their plasma membrane, whereas Gram-negative bacteria
(e.g. Escherichia coli) have a thinner (3 nm) cell wall and a second outer membrane
(Figure 3). In contrast with the plasma membrane, this outer membrane is very
permeable to the passage of relatively large molecules (molecular weight > 1000
Da) due to porin proteins which form pores in the lipid bilayer. Between the outer
membrane and the cell wall is the periplasm, a space occupied by proteins secreted
from the cell.

Figure 3. Cell wall structure of (a) Gram-positive and (b) Gram-negative bacteria.

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Bacterial flagella: Many bacterial cells have one or more tail-like appendages
known as flagella. By rotating their flagella, bacteria can move through the
extracellular medium towards attractants and away from repellents, so called
chemotaxis. Bacterial flagella are different from eukaryotic cilia and flagella in two
ways: (1) each bacterial flagellum is made of the protein flagellin (53 kDa subunit)
as opposed to tubulin; and (2) it rotates rather than bends. An E. coli bacterium has
about six flagella that emerge from random positions on the surface of the cell.
Flagella are thin helical filaments, 15 nm in diameter and 10 μm long. Electron
microscopy has revealed that the flagellar filament contains 11 subunits in two
helical turns which, when viewed end-on, has the appearance of an 11-bladed
propeller with a hollow central core. Flagella grow by the addition of new flagellin
subunits to the end away from the cell, with the new subunits diffusing through the
central core. Between the flagellar filament and the cell membrane is the flagellar
hook composed of subunits of the 42 kDa hook protein that forms a short, curved
structure. Situated in the plasma membrane is the basal body or flagellar motor, an
intricate assembly of proteins. The flexible hook is attached to a series of protein
rings which are embedded in the inner and outer membranes. The rotation of the
flagella is driven by a flow of protons through an outer ring of proteins, called the
stator.
Eukaryote Cell Structure

Plasma membrane: The plasma membrane envelops the cell, separating it from the
external environment and maintaining the correct ionic composition and osmotic
pressure of the cytosol. The plasma membrane, like all membranes, is impermeable
to most substances but the presence of specific proteins in the membrane allows
certain molecules to pass through, therefore making it selectively permeable. The
plasma membrane is also involved in communicating with other cells, in particular
through the binding of ligands (small molecules such as hormones,
neurotransmitters, etc.) to receptor proteins on its surface. The plasma membrane is
also involved in the exocytosis (secretion) and endocytosis (internalization) of
proteins and other macromolecules.
In 1972 S. Jonathan Singer and Garth Nicholson proposed the fluid mosaic
model for the overall structure of biological membranes, in which the membranes
can be viewed as two-dimensional solutions of orientated lipids and globular
proteins (Figure 4). The integral membrane proteins can be considered as
‘iceberg’ floating in a two-dimensional lipid ‘sea’. They proposed that the
bilayer organization of the lipids would act both as a solvent for the amphipathic
integral membrane proteins and as a permeability barrier. They also proposed that
some lipids may interact with certain membrane proteins, that these interactions
would be essential for the normal functioning of the protein, and that membrane
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proteins would be free to diffuse laterally in the plane of the bilayer unless restricted
in some way, but would not be able to flip from one side of the bilayer to the other.
This model is now supported by a wide variety of experimental observations.

Figure 4a. The fluid mosaic model of membrane structure.

Figure 4b. Protein associations with membranes.

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 Figure 4c. Functions of membrane proteins.

Figure 4d. Arrangements of membrane phospholipids in a bilayer.

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Figure 4e. Structures of some phospholipids.

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Figure 4f. Structures of membrane glycerophospholipids. R1 and R2
represent hydrocarbon chains of fatty acids.

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 Figure 4f. Structures of (a) the sphingolipids sphingomyelin and
galactocerebroside; (b) cholesterol. R1 represents the hydrocarbon chain of
fatty acids.

Figure 4g. Asymmetry of Figure 4h. Structure Figure 4i.

membranes. of cholesterol. Cholesterol and
phospholipids in
membranes.

Nucleus: The nucleus is bounded by two membranes, the inner and outer nuclear
membranes. These two membranes fuse together at the nuclear pores through which
molecules [messenger ribonucleic acid (mRNA), proteins, ribosomes, etc.] can
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move between the nucleus and the cytosol. Other proteins, for example those
involved in regulating gene expression, can pass through the pores from the cytosol
to the nucleus. The outer nuclear membrane is often continuous with the rough
endoplasmic reticulum (RER). Within the nucleus the DNA is tightly coiled around
histone proteins and organized into complexes called chromosomes. Visible under
the light microscope is the nucleolus, a subregion of the nucleus which is the site of
ribosomal ribonucleic acid (rRNA) synthesis.
Endoplasmic reticulum: The endoplasmic reticulum (ER) is an interconnected
network of membrane vesicles. The rough endoplasmic reticulum (RER) is studded
on the cytosolic face with ribosomes, the sites of membrane and secretory protein
biosynthesis. Within the lumen of the RER are enzymes involved in the
posttranslational modification (glycosylation, proteolysis, etc.) of membrane and
secretory proteins. The smooth endoplasmic reticulum (SER), which is not studded
with ribosomes, is the site of phospholipid biosynthesis, and is where a number of
detoxification reactions take place.
Golgi apparatus: The Golgi apparatus, a system of flattened membrane-bound sacs,
is the sorting and processing center of the cell. Membrane vesicles from the RER,
containing membrane and secretory proteins, fuse with the Golgi apparatus and
release their contents into it. On transit through the Golgi apparatus, further
posttranslational modifications to these proteins take place and they are then sorted
and packaged into different vesicles. These vesicles bud off from the Golgi apparatus
and are transported through the cytosol, eventually fusing either with the plasma
membrane to release their contents into the extracellular space (a process known as
exocytosis) or with other internal organelles (e.g. lysosomes).
Mitochondria: A mitochondrion has an inner and an outer membrane between
which is the intermembrane space (Figure 5). The outer membrane contains porin
proteins which make it permeable to molecules of up to 10 kDa. The inner
membrane, which is considerably less permeable, has large infoldings called cristae,
which protrude into the central matrix. The inner membrane is the site of oxidative
phosphorylation and electron transport involved in ATP production. The central
matrix is the site of numerous metabolic reactions including the citric acid cycle and
fatty acid breakdown. Also, within the matrix is found the mitochondrial DNA which
encodes some of the mitochondrial proteins.

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 Figure 5. The structure of a mitochondrion.

Chloroplasts: Chloroplasts, present exclusively in plant cells, also have inner and
outer membranes. In addition, there is an extensive internal membrane system made
up of thylakoid vesicles (interconnected vesicles flattened to form discs) stacked
upon each other to form grana (Figure 6). Within the thylakoid vesicles is the green
pigment chlorophyll, along with the enzymes that trap light energy and convert it
into chemical energy in the form of ATP. The stroma, the space surrounding the
thylakoid vesicles, is the site of carbon dioxide (CO2) fixation-the conversion of CO2
into organic compounds. Chloroplasts, like mitochondria, contain DNA which
encodes some of the chloroplast proteins.

Figure 6: The structure of a chloroplast.

Lysosomes: Lysosomes, which are found only in animal cells, have a single
boundary membrane. The internal pH of these organelles is mildly acidic (pH 4–5),
and is maintained by integral membrane proteins which pump H + ions into them.
The lysosomes contain a range of hydrolases that are optimally active at this acidic
pH (and hence are termed acid hydrolases), but which are inactive at the neutral pH

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of the cytosol and extracellular fluid. These enzymes are involved in the degradation
of host and foreign macromolecules into their monomeric subunits; proteases
degrade proteins, lipases degrade lipids, phosphatases remove phosphate groups
from nucleotides and phospholipids, and nucleases degrade DNA and RNA.
Lysosomes are involved in the degradation of extracellular macromolecules that
have been brought into the cell by endocytosis as well as in the degradation and
recycling of normal cellular components.
Peroxisomes: These organelles have a single boundary membrane and contain
enzymes that degrade fatty acids and amino acids. A byproduct of these reactions is
hydrogen peroxide, which is toxic to the cell. The presence of large amounts of the
enzyme catalase in the peroxisomes rapidly converts the toxic hydrogen peroxide
into harmless H2O and O2:
Catalase
2H2O2 2H2O + O2

Cytosol: The cytosol is that part of the cytoplasm not included within any of the
subcellular organelles, and is a major site of cellular metabolism, containing a large
number of different enzymes and other proteins. For example, glycolysis,
gluconeogenesis, the pentose phosphate pathway and fatty acid synthesis all take
place in the cytosol. The cytosol is not a homogeneous ‘soup’ but has within it
the cytoskeleton, a network of fibers criss-crossing through the cell that helps to
maintain the shape of the cell. The cytoskeletal fibers include microtubules (30 nm
in diameter), intermediate filaments (10 nm in diameter) and microfilaments (8 nm
in diameter). Also found within the cytosol of many cells are inclusion bodies
(granules of material that are not membrane-bounded) such as glycogen granules in
liver and muscle cells, and droplets of triacylglycerol in the fat cells of adipose
tissue.
Plant cell wall: Surrounding the plasma membrane of a plant cell is the cell wall,
which imparts strength and rigidity to the cell. This is built primarily of cellulose, a
rod-like polysaccharide of repeating glucose units linked β(1–4). These cellulose
molecules are aggregated together by hydrogen bonding into bundles of fibers, and
the fibers in turn are cross-linked together by other polysaccharides. In wood another
compound, lignin, imparts added strength and rigidity to the cell wall. Lignin is a
complex water-insoluble phenolic polymer.
Plant cell vacuole: Plant cells usually contain one or more membrane-bounded
vacuoles. These are used to store nutrients (e.g. sucrose), water, ions and waste
products (especially excess nitrogen-containing compounds). Like lysosomes in
animal cells, vacuoles have an acidic pH maintained by H+ pumps in the membrane
and contain a variety of degradative enzymes. Entry of water into the vacuole causes

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it to expand, creating hydrostatic pressure (turgor) inside the cell which is balanced
by the mechanical resistance of the cell wall.

Basic Properties of Cells

Just as plants and animals are alive, so too are cells. Life, in fact, is the most
basic property of cells, and cells are the smallest units to exhibit this property. Unlike
the parts of a cell, which simply deteriorate if isolated, whole cells can be removed
from a plant or animal and cultured in a laboratory where they will grow and
reproduce for extended periods of time. If mistreated, they may die. Death can also
be considered one of the most basic properties of life, because only a living entity
faces this prospect. Remarkably, cells within the body generally die “by their own
hand”- the victims of an internal program that causes cells that are no longer needed
or those that pose a risk of becoming cancerous to eliminate themselves. The first
culture of human cells was begun by George and Martha Gey of Johns Hopkins
University in 1951. The cells were obtained from a malignant tumor and named
HeLa cells after the donor, Henrietta Lacks. HeLa cells- descended by cell division
from this first cell sample- are still being grown in laboratories around the world
today (Figure 7). Because they are so much simpler to study than cells situated within
the body, cells grown in vitro (i.e., in culture, outside the body) have become an
essential tool of cell and molecular biologists. In fact, much of the information that
will be discussed in this book has been obtained using cells grown in laboratory
cultures.

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 Figure 7. HeLa cells were the first human cells to be kept in culture for long periods of
time and are still in use today. Unlike normal cells, which have a finite lifetime in
culture, these cancerous HeLa cells can be cultured indefinitely as long as conditions
are favorable to support cell growth and division.

Cells Possess a Genetic Program and the Means to Use It

Organisms are built according to information encoded in a collection of genes,
which are constructed of DNA. If converted to words, the human genetic program would
contain enough information to fill millions of pages of text. Remarkably, this vast amount
of information is packaged into a set of chromosomes that occupies the space of a cell
nucleus—hundreds of times smaller than the dot on this i.
Genes are more than storage lockers for information: They constitute the recipes
for constructing cellular structures, the directions for running cellular activities, and the
program for making more of themselves. The molecular structure of genes allows for
changes in genetic information (mutations) that lead to variation among individuals,
which forms the basis of biological evolution. Discovering the mechanisms by which cells
use and transmit their genetic information has been one of the greatest achievements of
science during the last century.
Cells Are Capable of Producing More of Themselves
Just as individual organisms are generated by reproduction, so too are individual
cells. Cells reproduce by division, a process in which the contents of a mother cell are
distributed into two daughter cells. Prior to division, the genetic material is faithfully

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duplicated, and each daughter cell receives a complete and equal share of genetic
information. In most cases, the two daughter cells have approximately equal volume. In
some cases, however, as occurs when a human oocyte undergoes division, one of the cells
can retain nearly all of the cytoplasm, even though it receives only half of the genetic
material.
Cells Acquire and Utilize Energy
Every biological process requires the input of energy. Virtually all of the energy
utilized by life on the Earth’s surface arrives in the form of electromagnetic radiation from
the sun. The energy of light is trapped by light-absorbing pigments present in the
membranes of photosynthetic cells. Light energy is converted by photosynthesis into
chemical energy that is stored in energy-rich carbohydrates, such as sucrose or starch.
For most animal cells, energy arrives prepackaged, often in the form of the sugar glucose.
In humans, glucose is released by the liver into the blood, where it circulates through the
body delivering chemical energy to all the cells. Once in a cell, the glucose is disassembled
in such a way that its energy content can be stored in a readily available form (usually as
ATP) that is later put to use in running all of the cell’s myriad energy-requiring activities.
Cells expend an enormous amount of energy simply breaking down and rebuilding the
macromolecules and organelles of which they are made. This continual turnover
maintains the integrity of cell components in the face of inevitable wear and tear and
enables the cell to respond rapidly to changing conditions.
Cells Carry Out a Variety of Chemical Reactions
Cells function like miniaturized chemical plants. Even the simplest bacterial cell is
capable of hundreds of different chemical transformations, none of which occurs at any
significant rate in the inanimate world. Virtually all chemical changes that take place in
cells require enzymes-molecules that greatly increase the rate at which a chemical
reaction occurs. The sum total of the chemical reactions in a cell represents that cell’s
metabolism.
Cells Engage in Mechanical Activities
Cells are sites of bustling activity. Materials are transported from place to place,
structures are assembled and then rapidly disassembled, and, in many cases, the entire
cell moves itself from one site to another. These types of activities are based on dynamic,
mechanical changes within cells, many of which are initiated by changes in the shape of
motor proteins. Motor proteins are just one of many types of molecular “machines”
employed by cells to carry out mechanical activities.

Cells Are Able to Respond to Stimuli

Some cells respond to stimuli in obvious ways; a single-celled protist, for example,
moves away from an object in its path or moves toward a source of nutrients. Cells within
a multicellular plant or animal respond to stimuli less obviously. Most cells are covered
with receptors that interact with substances in the environment in highly specific ways.
Cells possess receptors to hormones, growth factors, and extracellular materials, as well
as to substances on the surfaces of other cells. Receptors provide pathways through which
external stimuli can evoke specific responses in target cells. Cells may respond to specific
stimuli by altering their metabolic activities, moving from one place to another, or even
committing suicide.

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