Prokaryotes

Typical organisms bacteria, archaea

Eukaryotes
protists, fungi, plants, animals

~ 110 m

~ 10100 m (sperm cells, apart from the tail, are

smaller)

nucleoid region; no
real nucleus

real nucleus with double membrane

circular (usually)

linear molecules (chromosomes) with histone

proteins

coupled in cytoplasm

RNA-synthesis inside the nucleus

protein synthesis in cytoplasm

50S+30S

60S+40S

Cytoplasmatic
structure

very few structures

highly structured by endomembranes and a

cytoskeleton

Cell movement

flagella made of
flagellin

flagella and cilia containing microtubules;

lamellipodia and filopodia containing actin

Mitochondria

none

one to several thousand (though some lack

mitochondria)

Chloroplasts

none

in algae and plants

Organization

usually single cells

single cells, colonies, higher multicellular organisms

with specialized cells

Cell division

Binary fission (simple

division)

Mitosis (fission or budding)

Meiosis

Typical size
Type of nucleus
DNA
RNA-/proteinsynthesis
Ribosomes

Cell Glosaries:
Cell

Prokaryote

Eukaryote
Flagella

the basic structural unit of all living organisms; it typically

comprises a small, usually microscopic, discrete mass of
organellecontaining
cytoplasm, bounded externally by a membrane. Each
cell is capable of interacting with other cells and
performing all the
fundamental functions of life. In plants the cell usually also
includes
the cell wall. Originally the term referred to the bounding
wall of a
plant cell that had lost its living content.
any organism in which the genomic
DNA is not enclosed by a nuclear membrane within the
cells;
i.e. any organism whose cells possess a prokaryon. The
prokaryotes
comprise the bacteria, cyanobacteria, and archaea and
are classified in their own kingdom, variously named
Monera
or Procaryotae.
any organism whose cells contain
a eukaryon (or eukarya) and undergo meiosis.
the specialized locomotory
appendage of bacteria of certain taxonomic groups,
consisting of a
noncontractile, filiform extension through the cell surface,
borne
singly, in groups, or as a covering layer. It is commonly 3
20 lm
long and 1225 nm in diameter, is built up of several

(often three)
longitudinally arranged chains of flagellin subunits (often
in a spiral),
and is anchored to the cell envelope by a basal body (def.
1),
which imparts rotatory motion to it. Bacterial flagella carry
the H
(or flagellar) antigens, which are useful in some instances,
e.g. in
Salmonella spp., for classification.

Flagellin

Cilia

any member of a group of similar monomeric soluble

globular
proteins that constitute the subunits of bacterial flagella.
They
also comprise the major antigens of several bacteria,
including Salmonella
spp., which alternate between the production of two
antigenic
forms of flagella, termed phase 1 and phase 2.
a specialized eukaryotic locomotor organelle that
consists of a filiform extrusion of the cell surface. Each
cilium is
bounded by an extrusion of the cytoplasmic membrane,
and contains
a regular longitudinal array of microtubules, anchored
basally
in a centriole.

Microvilli

any of the very small, finger-like projections

that occur on the exposed surfaces of epithelial cells,
especially
where cellular function requires a maximal surface area for
absorption or secretion, as in the proximal renal tubule
and the
small intestine, where they constitute the brush border.
The projections
are about 12 lm long and 100 nm in diameter. The core of
a
microvillus contains a bundle of about 40 actin filaments
of uniform
polarity with their barbed (+) ends anchored in a cap of
illdefined
amorphous material in the microvillar tip. At the base the
actin filaments extend into the perpendicular network, also
made
up largely of actin, known as the terminal web. In the core
the actin
filaments are bound together by the actin-binding proteins
fimbrin
and villin. The renal and intestinal microvilli contain a wide
range
of hydrolases that are integral membrane proteins; their
hydrophilic
domains, containing the catalytic sites, face outwards
from the cell.

Protoplasm

the living contents of a cell; i.e. the matter contained

within (and including) the plasma membrane, usually
taken to exclude
large vacuoles and masses of secretory or ingested
material.
In eukaryotes it includes the nucleus and cytoplasm.
the rigid or semi-rigid envelope lying outside the cell

Cell Wall

membrane

of plant, fungal, and most prokaryotic cells maintaining

their
shape and protecting them from osmotic lysis. In
prokaryotes it lies
inside any capsule and slime layer, usually consists mainly
of peptidoglycan,
and can be removed by various techniques with retention
of its three-dimensional form. In fungi the cell wall is
composed
largely of polysaccharides, while in plants it is made up of
cellulose
and, often, lignin.

Cell

any membrane found in a cell. The term is sometimes

Membrane

used loosely to indicate the plasma membrane.

Cytoplasmic
Membrane

any of the membranous structures

within the cytoplasm of eukaryotes, such as the rough and
smooth
endoplasmic reticulum and the Golgi apparatus.
plasma membrane but including the other intracellular
organelles
and structures. Some authors currently also exclude
mitochondria,
chloroplasts, or other structures containing indepen-

Cytoplasm

163

cytochrome m cytoplasm

dently replicating DNA. The term is not synonymous with

cytoplasmic
fraction.

Cytosol

Cytoskeleto
n

the part of the cytoplasm that does not appear to contain

membranous or particulate subcellular components. The
term is
not synonymous with soluble fraction.
any of the various filamentous elements within the
cytoplasm of eukaryotic cells that remain after treatment
of the
cells with mild detergent to remove membrane
constituents and soluble
components of the cytoplasm. The term embraces
intermediate
filaments, microfilaments, microtubules, and the
microtrabecular lattice.

The various elements of the cytoskeleton not only serve in

the
maintenance of cellular shape but also have roles in other
cellular
functions, including cellular movement, cell division,
endocytosis,
and movement of organelles.

Microtubules

any of the long, generally straight, hollow tubes of internal

diameter 1215 nm and external diameter 24 nm found in
a
wide variety of eukaryotic cells. Each consists (usually) of
13
protofilaments of polymeric tubulin, staggered in such a
manner that
the tubulin monomers are arranged in a characteristic
helical pattern
on the surface, and with the a/b axes of the tubulin
parallel to
the long axis of the tubule. Microtubules exist in
equilibrium with a
pool of tubulin monomers in the cytoplasm and can be
rapidly assembled
and disassembled in response to various physiological
stimuli; there are also a number of microtubuleassociated proteins.
Most microtubules in a cell appear to be initiated at
microtubule-organizing
centres and some may also become attached to the
kinetochores.

Microtubules are involved in force generation in cilia and

flagella, where they exist in a characteristic 9 + 2 array (a
ring of
nine doublet microtubules with two single microtubules in
the centre),
and the integrity of the microtubule network appears to be
necessary
for certain aspects of mitotic spindle function and for the
saltatory movement of cell organelles. Taxol prevents
depolymerization
of microtubules, and is thus antimitotic.

Microfilamen
t

a type of actin filament that contributes to the

cytoskeleton
of eukaryotic cells. Microfilaments are elongated
structures
consisting of two strands of globular actin monomers
twisted
into a helix with 13.5 molecules per turn. Microfilaments

form the
major component of the cells contractile machinery and
are implicated
in cell division, cell movement, muscle contraction, nerve
outgrowth,
tubular gland formation, gastrulation, and neurulation.

Organelle

any discrete structure in a unicellular organism, or in an

individual cell of a multicellular organism, that is adapted
and/or
specialized for the performance of one or more vital
functions.

Nucleus

the most conspicuous organelle of

a eukaryotic cell; it contains the chromosomes and (except
for mitochondria
and chloroplasts) is the sole site of DNA replication and
of RNA synthesis in the cell. Usually a spheroidal body, it is
separated
from the cytoplasm by the nuclear envelope.

Nucleolus

a small, dense body one or more of which

are present in the nucleus of eukaryotic cells. It is rich in
RNA and
protein, is not bounded by a limiting membrane, and is not
seen
during mitosis. Its prime function is the transcription of the
nucleolar
DNA into 45S ribosomal-precursor RNA, the processing of
this
RNA into 5.8S, 18S, and 28S components of ribosomal
RNA, and
the association of these components with 5S RNA and
proteins
synthesized outside the nucleolus. This association results
in the
formation of ribonucleoprotein precursors; these pass into
the cytoplasm
and mature into the 40S and 60S subunits of the ribosome

Nuclear
Envelope

the membrane that surrounds

the nucleus of a eukaryotic cell. It is a double membrane
composed of two lipid bilayers separated by a gap of width
2040
nm (the perinuclear space). The outer nuclear membrane
is continuous
with the endoplasmic reticulum of the cell and is
sometimes
studded with ribosomes. At specialized regions ( nuclear
pores) the
outer nuclear membrane is connected to the inner nuclear
membrane.

Nucleoid

resembling a nucleus

Nucleoplasm

all the protoplasm contained within the nuclear envelope

of a living cell.

Chromosom
e

a structure composed of a very long molecule of DNA

(in humans 50250 Mb long) and associated proteins (e.g.
histones)
that carries hereditary information. Chromosomes are
especially
evident in plant or animal cells undergoing mitosis or
meiosis,
where each chromosome becomes condensed into a
compact, readily
visible thread. In nondividing cells chromosomes typically
assume
a more dispersed form called chromatin. The number of
chromosomes
is characteristic for the species concerned. In a bacterium
only one chromosome is evident as the cell is about to
divide. This is
a circular molecule of DNA, 4.6 Mb long in E. coli, with
attached
proteins. After DNA replication, the two new chromosomes
attach

to a specialized site on the bacterial plasma membrane for

segregation
to the two daughter cells. Mitochondria and chloroplasts
contain
several copies of a much smaller circular DNA molecule
with
some attached proteins.

Histone

any of a group of evolutionarily highly conserved basic

proteins,
molecular mass 1121 kDa, that constitute about half the
mass of the chromosomes of all eukaryotic cells, except
spermatozoa.
They comprise single polypeptide chains with a lysineplusarginine
content of about 25%, and are concerned in the packing of
DNA in chromatin.

DNA

deoxyribonucleic
acid; one of the two main types of nucleic acid, consisting

of
a long, unbranched macromolecule formed from one, or
more commonly,
two, strands of linked deoxyribonucleotides, the 3phosphate
group of each constituent deoxyribonucleotide being
joined
in 3,5-phosphodiester linkage to the 5-hydroxyl group of
the deoxyribose
moiety of the next one.

RNA

Mitochondri
on

ribonucleic
acid; one of the two main types of nucleic acid, consisting

of a long,
unbranched macromolecule formed from ribonucleotides,
the 3phosphate group of each constituent ribonucleotide
(except the
last) being joined in 3,5-phosphodiester linkage to the 5hydroxyl
group on each ribose moiety of the next one. The presence
of a free
2-hydroxyl group on each ribose moiety renders these
phosphodiester
bonds susceptible to hydrolytic attack by alkali, in contrast
to
those of DNA. The RNA chain has polarity, with one 5 end
and one
3 end. Two purines, adenine and guanine, and two
pyrimidines, cytosine
and uracil, are the major bases usually present. In
addition,
minor bases may occur; transfer RNA, however, contains
unusual
bases in relatively large amounts. The sequence of bases
carries information,
whereas the sugar and phosphate groups play a structural
role. RNA is fundamental to protein biosynthesis in all
living
cells.
a semiautonomous, self-replicating
organelle that occurs in varying numbers, shapes, and
sizes in
the cytoplasm of virtually all eukaryotic cells. It is the site
of tissue
respiration Conventionally,
mitochondria are represented as elongated cylinders with
a
diameter of 0.51.0 lm; however, in living cells they show
great mobility
and plasticity, in some cells forming long, moving chains
while in others being fixed in position near sites of high
ATP consumption;
e.g. in cardiac muscle they are packed between the
myofibrils
while in a sperm they are tightly wrapped around the
flagellum.
Mammalian hepatocytes each contain 10002000
mitochondria, roughly 20% of the cell volume. A
mitochondrion
has two functionally distinct membrane systems, the outer

one
completely surrounding the whole organelle and the inner
one
being infolded into cristae. These membrane systems
define two
compartments: the mitochondrial matrix and the
intermembrane
space. The matrix contains enzymes for the oxidation of
pyruvate
and fatty acids and for the tricarboxylic-acid cycle; it also
contains
the mitochondrial DNA and the enzymes and structures
necessary
for expression of the mitochondrial genes. The inner
membrane
contains the enzymes of the respiratory chain, ATP
synthetase, and
specific transport proteins. The intermembrane space
contains a
number of kinases. The outer membrane, which is
permeable to
molecules of up to 10 kDa, contains monoamine oxidase,
cytochrome
b5, a number of transferases, and a fatty-acid elongation
system. The mitochondrion was formerly known as a
chondriosome
and by numerous other names.

Crista

Chloroplast

Thylakoid

Granum
Ribosome

any of the inward folds of the inner mitochondrial

membrane. Their number, extent, and shape differ in
mitochondria
from different tissues and organisms. They appear to be
devices for increasing the surface area of the inner
mitochondrial
membrane, where the enzymes of electron transport and
oxidative
phosphorylation are found. Their shape can vary with the
respiratory
state of the mitochondria.
chlorophyll-containing
plastid, found in cells of algae and higher plants, that is
the site of
photosynthesis. Chloroplasts are characterized by a
system of membranes
embedded in a hydrophobic proteinaceous matrix, or
stroma. The basic unit of the membrane system is a
flattened single
vesicle called the thylakoid; thylakoids stack into grana
(sing.
granum). All the thylakoids of a granum are connected
with each
other, and the grana are connected by intergranal
lamellae. Chloroplasts
are rich in various proteases (e.g. serine, aspartic, ATPdependent,
light-activated, for oxidized or photodamaged proteins,
for trafficking of protein precursors into the plastids).
Some of
these may have a role in senescence.
a saclike vesicle that bears the photosynthetic pigments in
photosynthetic organisms. In prokaryotes the thylakoids
are of various
shapes and are attached to the plasma membrane. In
eukaryotes
they are flattened, membrane-bound disclike structures
located
in the chloroplasts; in higher plants they form dense
stacks called
grana. Isolated thylakoid preparations can carry out
photosynthetic
electron transport and the associated phosphorylation.
a stack of disclike thylakoids resembling a pile of
coins. Grana occur in chloroplasts, especially in higher
plants.
an intracellular organelle, about 200 in diameter,
consisting
of RNA and protein. It is the site of protein biosynthesis
resulting

from translation of messenger RNA (mRNA). It consists of

one large and one small subunit, each containing only
protein and
RNA. Ribosomes and their subunits are characterized by
their sedimentation
coefficients, expressed in Svedberg units (symbol: S). The
prokaryotic ribosome (70S) comprises a large (50S) and a
small
(30S) subunit, while the eukaryotic ribosome (80S)
comprises a
large (60S) subunit and a small (40S) subunit. Several
ribosomes
may bind to one mRNA to form a polysome. Two sites on
the large
subunit are involved in translation, namely the aminoacyl
site (A
site) and peptidyl site (P site). Ribosomes from
prokaryotes, eukaryotes,
mitochondria, and chloroplasts have distinct ribosomal
proteins.
Prokaryotic ribosomes contain 3545% protein, while
eukaryotic
ribosomes contain 50% protein. Ribosomal RNA (rRNA) is
the
RNA part of a ribosome. In prokaryotes, a 30S subunit
contains
one 16S RNA molecule with associated proteins, while a
50S subunit
contains one 5S and one 23S RNA molecule. In eukaryotes,
the 60S subunit contains one 28S, one 5.8S, and one 5S
RNA molecule,
while the 40S subunit contains one 18S RNA molecule. The
structures are known. In Escherichia coli, the rRNAs form
part of a
large transcript containing transfer RNA (tRNA) molecules,
which
is then cleaved during processing. In eukaryotes, the
genes for the
5.8S, 18S, and 28S molecules reside in the nucleolus, and
code for a
40S transcript, which is then cleaved during processing;
this includes
splicing that occurs in the absence of protein. The gene for
the 5S rRNA is a separate transcription unit outside the
nucleolus

Endoplasmic
Reticulum

ER; the irregular network of unit

membranes, visible only by electron microscopy, that
occurs in the
cytoplasm of many eukaryotic cells. The membranes form
a complex
meshwork of tubular channels, which are often expanded
into
slitlike cavities called cisternae. The ER may be rough (or
granular),
with ribosomes adhering to the outer surface, or smooth
(with no
ribosomes attached). When cells are disrupted by
homogenization,
the cisternae break up into small closed vesicles called
rough microsomes
or smooth microsomes. The ribosomes attached to the
rough
ER are the site of translation of the mRNA for those
proteins
which are to be retained within the cisternae (ER-resident
proteins),
targeted to the lysosomes, or destined for export from the
cell. Glycoproteins
undergo their initial glycosylation within the cisternae.
The smooth ER is the recipient of the proteins synthesized
in the
rough ER. Those proteins to be exported are passed to the
Golgi
complex (see Golgi apparatus), the resident proteins are
retained in
the rough ER, and the lysosomal proteins after
phosphorylation of
their mannose residues are passed to the lysosomes.

Glycosylation of
the glycoproteins also continues. The smooth ER is the site
of synthesis
of lipids, including the phospholipids. The major
membrane
phospholipids are phosphatidylcholines,
phosphatidylethanolamines,
phosphatidylserines, and sphingomyelins. The ER also
produces cholesterol, triacylglycerols, eicosanoids, and
ceramides,
and contains enzymes that catalyse reactions to detoxify
lipid-soluble
drugs and harmful products of metabolism. Large
quantities of
certain compounds such as phenobarbital cause an
increase in the
amount of the smooth ER

Microsome

Golgi
Apparatus

any of the small, heterogeneous, artefactual, vesicular

particles, 50150 nm in diameter, that are formed when
some eukaryotic
cells are homogenized and that sediment on centrifugation
at 105 g. Microsomes are formed mostly from disrupted
endoplasmic
reticulum membranes but some arise from the plasma
membrane.
Those formed from rough endoplasmic reticulum are
studded
with ribosomes on the outside (rough microsomes);
those
formed from smooth endoplasmic reticulum and from
plasma
membrane have no adhering ribosomes (smooth
microsomes).
a compound
membranous cytoplasmic organelle of eukaryotic cells,
consisting
of flattened, ribosome-free vesicles arranged in a more or
less regular stack. It differs from the endoplasmic
reticulum in often
having slightly thicker membranes, appearing in sections
as a characteristic
shallow semicircle so that the convex side (cis or entry
face) abuts the endoplasmic reticulum, secretory granules
emerging
from the concave side (trans or exit face). In vertebrate
cells there is
usually one such organelle, while in invertebrates and
plants, where
they are known usually as dictyosomes, there may be
several scattered
in the cytoplasm. The Golgi apparatus processes proteins
produced
on the ribosomes of the rough endoplasmic reticulum;
such
processing includes modification of the core
oligosaccharides of
glycoproteins, and the sorting and packaging of proteins
for transport
to a variety of cellular locations, e.g. to storage granules,
lysosomes,
or secretory vesicles. The Golgi is a major site of synthesis
of
polysaccharides, including the pectin and hemicellulose of
the plant
cell wall and most of the glycosaminoglycans of the
extracellular
matrix in animals. Three different regions are recognized
in terms
of structure and function: cis, in the vicinity of the cis face,
trans, in
the vicinity of the trans face, and medial, lying between
the cis and
trans regions. While the luminal proteins of the
endoplasmic reticulum
leave it together with the proteins to be secreted, they are
retrieved
from the cis network and returned to the endoplasmic
reticulum.

Vacuole

Thus transport occurs in both directions. Brefeldin A

blocks
protein secretion through the Golgi but does not appear to
block
the return pathway. Galactosyltransferase is a marker
enzyme for the
trans membranes while N-acetylglucosaminyltransferase
III serves this
role for the medial membranes.
a closed structure, found only in eukaryotic cells, that is
completely surrounded by unit membrane and contains
liquid material.
Plant cells contain a conspicuous vacuole, which may
occupy
up to 95% of the cell volume. In these cells it has a role
similar to
that of lysosomes of animal cells, being rich in hydrolytic
enzymes
(especially various types of protease). In seed reserve
tissues, specialized
protein storage vacuoles contain abundant storage protein
deposited during seed maturation. Upon germination, a
specific
protease is imported to supply amino acids to the new
seedling.

Vesicle

a closed structure, found only in eukaryotic cells, that is

completely surrounded by a unit membrane but, unlike a
vacuole,
contains material that is not (or is not known to be) in the
liquid
state.

Microbody

a cytoplasmic organelle, spherical or oval in shape, and

0.11.5 nm across, that is bounded by a single membrane
and contains
oxidative enzymes, especially those utilizing H2O2.
Microbodies
include glyoxysomes and peroxisomes.

Peroxisome

a type of eukaryotic cell organelle that is rich in enzymes

that act on or generate hydrogen peroxide, especially the
enzymes
catalase, (S)-2-hydroxy-acid oxidase, and D-amino-acid
oxidase.
Peroxisomes occur in mammalian kidney and liver, and in
yeasts and certain protozoans. They contain over 50
matrix enzymes
involved in degradation of very-long-chain, branched, or
polyunsaturated fatty acids, in the conversion of
cholesterol to bile
acids, and in biosynthesis of other phospholipids and
isoprene compounds.
They are particularly numerous in leaf photosynthetic cells
of plants, photosynthesizing via the Calvin (C3) pathway; in
plants
they are closely associated with chloroplasts and
mitochondria,
which is consistent with their function in photorespiration

Glyoxysome

a cellular organelle, of density 1.25 gcm3, found in the

organs of fatty seedlings where fat is being converted to
carbohydrate.
It contains the enzymes of the glyoxylate cycle and most
of the
particulate beta-oxidation activity.

Lysosome

any of a group of related cytoplasmic, membrane-bound

organelles that are found in most animal cells and that
contain a variety
of hydrolases, most of which have their maximal activities
in
the pH range 56. The contained enzymes display latency
if properly
isolated. About 40 different lysosomal hydrolases are
known
and lysosomes have a great variety of morphologies and
functions.

Centrosome

a structure occurring close to the nucleus during

interphase

in many eukaryotic cells. It comprises a pair of centrioles,

satellite bodies, and a cytoplasmic zone from which the
mitotic microtubule
spindle is organized. The structure of the centrosome in
animal cells changes continually during the cell-division
cycle. The
protein PCM-1 recruits centrosomal proteins such as
centrin, pericentrin,
ninein, and dynactin. On phosphorylation by Cdk2, the
centrosomal nucleophosmin is released to initiate
centrosomal duplication.
In some cells, lack of p53 leads to multiple copies of the
centrosome being generated during a single cell cycle.

Centiole

a cellular organelle, found close to the nucleus in many

eukaryotic
cells, consisting of a small cylinder with microtubular
walls, 300500 nm long and 120150 nm in diameter. It
contains
nine short, parallel, peripheral microtubular fibrils, each
fibril consisting
of one complete microtubule fused to two incomplete
microtubules.
Cells usually have two centrioles, lying at right angles to
each other. At division, each pair of centrioles generates
another
pair and the twin pairs form the poles of the mitotic
spindle.