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Cytology
The Study of Cells
CHAPTER OUTLINE
2.1
2.4
2.2
Study Guide
2.3
DEEPER INSIGHTS
2.1 When Desmosomes Fail
2.2 The Origin of Mitochondria
2.3 Cancer
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2.1
Microscopy
Cytology would not exist without the microscope. Throughout this
book, you will find many photomicrographsphotos of tissues
and cells taken through the microscope. The microscopes used
1
to produce them fall into three basic categories: the light microscope, transmission electron microscope, and scanning electron
microscope.
The light microscope (LM) uses visible light to produce its
images. It is the least expensive type of microscope, the easiest to
use, and the most often used, but it is also the most limited in the
amount of useful magnification it can produce. Light microscopes
today magnify up to 1,200 times. There are several varieties of light
microscopes, including the fluorescence microscope used to produce figure 2.15b.
Most of the structure we study in this chapter is invisible to
the LM, not because the LM cannot magnify enough but because
it cannot reveal enough detail. The most important thing about a
good microscope is not magnification but resolutionthe ability
to reveal detail. Any image can be photographed and enlarged as
much as we wish, but if enlargement fails to reveal greater detail, it
is empty magnification. A large blurry image is not nearly as informative as one that is small and sharp. For reasons of physics beyond
the scope of this chapter, it is the wavelength of light that places a
limit on resolution. At the wavelengths of visible light (about 400 to
700 nanometers, or nm), the LM cannot distinguish between two
objects any closer together than 200 nm (0.2micrometers, or m).
Resolution improves when objects are viewed with radiation of shorter wavelengths. Electron microscopes achieve higher
resolution by using not visible light but beams of electrons with
very short wavelength (0.005 nm). The transmission electron
microscope (TEM), invented in the mid-twentieth century, is usually used to study specimens that have been sliced ultrathin with
diamond knives and stained with heavy metals such as osmium,
which absorbs electrons. The TEM resolves details as small as
0.5nm and attains useful magnifications of biological material up
to 600,000 times. This is good enough to see even things as small as
proteins, nucleic acids, and other large molecules. Such fine detail
is called cell ultrastructure. Even at the same magnifications as the
LM, the TEM reveals far more detail (fig. 2.1). It usually produces
two-dimensional black-and-white images, but electron photo
micrographs are often colorized for instructional purposes.
The scanning electron microscope (SEM) uses a specimen
coated with vaporized metal (usually gold). The electron beam
strikes the specimen and discharges secondary electrons from the
metal coating. These electrons then strike a fluorescent screen
and produce an image. The SEM yields less resolution than the
TEM and is used at lower magnification, but it produces dramatic
three-dimensional images that are sometimes more informative
than TEM images, and it does not require that the specimen be
cut into thin slices. The SEM can view only the surfaces of specimens; it does not see through an object as the LM or TEM does.
Cell interiors can be viewed, however, by a freeze-fracture method
in which a cell is frozen, cracked open, coated with gold vapor,
and then viewed by either TEM or SEM. Figure2.2 compares red
blood cells photographed with the LM, TEM, and SEM.
Apply What You Know
Beyond figure 2.2, list all of the photomicrographs in this chapter
that you believe were made with the LM, with the TEM, and with
the SEM.
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10.0 m
Blood vessel
10.0 m
2.0 m
10.0 m
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Squamous
Cuboidal
Columnar
Polygonal
Stellate
Spheroid
Discoid
Fusiform (spindle-shaped)
Fibrous
20 m
Growth
20 m
10 m
10 m
Large cell
Diameter
= 20 m
Surface area = 20 m 20 m 6 = 2,400 m2
Volume
= 20 m 20 m 20 m = 8,000 m3
Small cell
Diameter
= 10 m
Surface area = 10 m 10 m 6 = 600 m2
Volume
= 10 m 10 m 10 m = 1,000 m3
Effect of cell growth:
Diameter (D ) increased by a factor of 2
Surface area increased by a factor of 4 (= D 2)
Volume increased by a factor of 8 (= D 3)
Figure 2.4 The Relationship Between Cell Surface Area and Volume.
As a cell doubles in width, its volume increases eightfold, but its surface area
increases only fourfold. A cell that is too large may have too little plasma
membrane to support the metabolic needs of its volume of cytoplasm.
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Microvillus
Terminal web
Desmosome
Secretory vesicle
undergoing
exocytosis
Fat droplet
Secretory vesicle
Intercellular
space
Golgi vesicles
Centrosome
Centrioles
Golgi complex
Free ribosomes
Nucleus
Lysosome
Nucleolus
Microtubule
Nuclear
envelope
Rough endoplasmic
reticulum
Smooth endoplasmic
reticulum
Mitochondrion
Plasma membranes
Hemidesmosome
Basal cell surface
Basement
membrane
Figure 2.5 Structure of a Generalized Cell. The cytoplasm is usually more crowded with organelles than shown here. The organelles are not all drawn
to the same scale.
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Table 2.1
Object
Size
100 m
1015 m
Cilia, length
710 m
0.2 4 m
13 m
Microvilli, length
12 m
Before You Go On
Answer the following questions to test your understanding of the
preceding section:
1.
2.
3.
4.
2.2
30100 nm
Ribosomes, diameter
15 nm
510 nm
7.5 nm
2.0 nm
0.8 nm
were little aware of this detail simply because most of these structures are too small to be resolved by the light microscope (table2.1).
We now regard cells as having the following major components:
Plasma membrane
Cytoplasm
Cytoskeleton
Organelles (including the nucleus)
Inclusions
Cytosol
The plasma membrane (cell membrane) forms the cells surface boundary. The material enclosed by the plasma membrane is
the cytoplasm,7 and the material within the nucleus (usually the
cells largest organelle) is the nucleoplasm. The cytoplasm contains
the cytoskeleton, a supportive framework of protein filaments and
tubules; an abundance of organelles, diverse structures that perform various metabolic tasks for the cell; and inclusions, which are
foreign matter or stored cell products. The cytoskeleton, organelles,
and inclusions are embedded in a clear gel called the cytosol.
The cytosol is also called the intracellular fluid (ICF). All
body fluids not contained in the cells are collectively called the
extracellular fluid (ECF). The ECF located between the cells is
also called tissue (interstitial) fluid. Some other extracellular
fluids include blood plasma, lymph, and cerebrospinal fluid.
7
Membrane Lipids
The plasma membrane is an oily, two-layered lipid film with proteins embedded in it (fig. 2.6b). By weight, it is about half lipid
and half protein. Since the lipid molecules are smaller and lighter,
however, they constitute about 90% to 99% of the molecules in the
membrane.
About 75% of the membrane lipid molecules are phospholipids.
A phospholipid (fig. 2.7) consists of a three-carbon backbone called
glycerol, with fatty acid tails attached to two of the carbons and a
phosphate-containing head attached to the third. The two fatty acid
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Plasma membrane
of upper cell
Intercellular space
Plasma membrane
of lower cell
Nuclear envelope
Nucleus
(a)
100 nm
Extracellular fluid
Peripheral
protein
Glycolipid
Glycoprotein
Carbohydrate
chains
Extracellular
face of
membrane
Phospholipid
bilayer
Channel
Peripheral
protein
Cholesterol
Transmembrane
protein
Intracellular fluid
Intracellular
face of
membrane
Proteins of
cytoskeleton
(b)
Figure 2.6 The Plasma Membrane. (a) Plasma membranes of two adjacent cells (TEM). Note also that the nuclear envelope is composed
of a double membrane, each layer of which is similar to a plasma membrane. (b)Molecular organization of the plasma membrane.
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Nitrogencontaining
group
(choline)
Hydrophilic
head
Phosphate
group
Glycerol
Fatty acid
tails
Hydrophobic
fatty acid
tails
(b)
Key
Carbon
Hydrogen
Nitrogen
Oxygen
Phosphorus
(a)
Membrane Proteins
Carbohydrate
Transmembrane protein:
Hydrophilic region
Hydrophobic region
Phospholipid
bilayer
Cytoskeletal
protein
Anchoring peripheral
protein
Figure 2.8 Transmembrane Proteins. A transmembrane protein has hydrophobic regions embedded in the phospholipid bilayer and hydrophilic
regions projecting into the extracellular and intracellular fluids. The protein may cross the membrane once (left) or multiple times (right). The intracellular
domain of the protein is often anchored to the cytoskeleton by peripheral proteins.
What other regions of the protein on the right would be hydrophilic in addition to the one labeled?
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Breakdown
products
Chemical
messenger
Ions
(a) Receptor
A receptor that
binds to chemical
messengers such
as hormones sent
by other cells
(b) Enzyme
An enzyme that
breaks down
a chemical
messenger and
terminates its
effect
CAM of
another cell
(f) Cell-adhesion
molecule (CAM)
A cell-adhesion
molecule (CAM)
that binds one
cell to another
Membrane Transport
One of the most important functions of the plasma membrane is
to control the passage of materials into and out of the cell. Figure 2.10 illustrates three methods of movement through plasma
Filtration
Simple Diffusion
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Lipid-soluble solutes
diffuse through
membrane
phospholipid.
High
concentration
Water-soluble solutes
diffuse through
channel proteins.
Water
Capillary wall
Solute moves
down its
concentration
gradient
Red blood
cell
Clefts hold back
blood cells and
other large particles.
Low
concentration
(a) Filtration
High
concentration
Solute binds to
receptor site on
transport protein.
Protein
changes
shape.
Protein releases
solute on other side
of membrane.
Protein releases
solute to other
side of membrane
and releases Pi.
Solute moves
up its
concentration
gradient.
Solute moves
down its
concentration
gradient.
Low
concentration
High
concentration
Solute binds
to receptor
site on
transport
protein.
Receptor site
Low
concentration
Osmosis
Osmosis10 (oz-MO-sis) is the net flow of water through a selectively permeable membrane from the more watery side (the one
with less dissolved matter) to the less watery side (with more dissolved matter). Water molecules tend to associate with particles of
dissolved matter and thus resist going back through the membrane
in the opposite directionhence the net accumulation of water on
the side with more solute. An exception to this is reverse osmosis,
in which a physical force drives water back to the more dilute side
of the membrane. This principle is used to desalinate seawater, converting it todrinkable freshwater, in arid regions and ships at sea.
In the human body, the force generated by the heartbeat causes
10
11
Facilitated Diffusion
facil = easy
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10
Pseudopod
Active Transport
Bacteria
Phagosome
Nucleus
(a) Phagocytosis
Pinocytotic vesicle
(b) Pinocytosis
Vesicular Transport
Solute
Receptor
Vesicle
(c) Receptor-mediated endocytosis
Cell product
Secretory
vesicle
(d) Exocytosis
Figure 2.11 Modes of Vesicular Transport. (a) Phagocytosis. A white blood cell engulfing bacteria with its
12
pseudopods. (b) Pinocytosis. A cell imbibing droplets of extracellular fluid. (c) Receptor-mediated endocytosis.
The Ys in the plasma membrane represent membrane receptors. The receptors bind a solute in the extracellular
fluid, then cluster together. The membrane caves in at that point until a vesicle pinches off into the cytoplasm
bearing the receptors and bound solute. (d) Exocytosis. A cell releasing a secretion or waste product.
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chains of sugars belonging to the membrane glycolipids and glycoproteins (fig. 2.12). The glycocalyx has multiple functions. It cushions
the plasma membrane and protects it from physical and chemical injury, somewhat like the Styrofoam peanuts in a shipping carton. It
functions in cell identity and thus in the bodys ability to distinguish
its own healthy cells from diseased cells, invading organisms, and
transplanted tissues. Human blood types and transfusion compatibility are determined by the glycocalyx. The glycocalyx also includes
the cell-adhesion molecules described earlier and, thus, helps to bind
tissues together and enables a sperm to bind to an egg and fertilize it.
Microvilli
All of our cells are essentially sugar-coated. They have a fuzzy surface
coat called the glycocalyx16 (GLY-co-CAY-licks), composed of short
Microvilli17 (MY-cro-VIL-eye; singular, microvillus) are extensions of the plasma membrane that serve primarily to increase its
surface area (fig. 2.12). They are best developed in cells specialized for absorption, such as the epithelial cells of the intestines and
kidney tubules. The small intestine has about 200 million microvilli per square millimeter, with about 3,000 on the surface of each
16
17
The Glycocalyx
Glycocalyx
Microvillus
Actin
microfilaments
Rootlet of
actin
microfilaments
(a)
1.0 m
(b)
0.1 m
Figure 2.12 Microvilli and the Glycocalyx. The microvilli are anchored by bundles of actin microfilaments, which occupy the core of each microvillus
and project into the cytoplasm. (a) Longitudinal sections, perpendicular to cell surface. (b) Cross sections.
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12
Cilia
Flagella
cilium = eyelash
axo = axis; neme = thread
20
dyn = power, energy; in = protein
21
flagellum = whip
19
an identical axoneme, but between the axoneme and plasma membrane it also has a complex sheath of coarse cytoskeletal filaments
that stiffen the tail and give it more propulsive power.
Cell Junctions
Also at the cell surface are certain cell junctions that link cells
together and attach them to the extracellular material. Such attachments enable cells to grow and divide normally, resist stress, communicate with each other, and control the movement of substances
through the gaps between cells. Without them, cardiac muscle cells
would pull apart when they contracted, and every swallow of food
would scrape away the lining of the esophagus. We will examine
three types of junctionstight junctions, desmosomes, and gap
junctions (fig. 2.14). Each type serves a different purpose, and two
or more types often occur in a single cell.
Tight Junctions
Desmosomes
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Axoneme:
Peripheral microtubules
Central microtubules
Dynein arms
Cilia
Shaft of cilium
Basal body
(a)
Plasma membrane
10 m
(b)
Cilia
Microvilli
Axoneme
Dynein
arm
Central
microtubule
Peripheral
microtubules
(c)
0.15 m
(d)
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14
Proteins
Connexon
Pore
Basement membrane
(d) Hemidesmosome
Which of these junctions allows material to pass from one cell directly into the next?
DEEPER INSIGHT
2.1
Gap Junctions
23
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Before You Go On
Answer the following questions to test your understanding of the
preceding section:
5. Generally speaking, what sort of substances can enter a cell
by diffusing through its phospholipid membrane? What sort
of substances must enter primarily through the channel proteins?
6. Compare the structure and function of transmembrane proteins with peripheral proteins.
7. What membrane transport processes get all the necessary
energy from the spontaneous movement of molecules? What
ones require ATP as a source of energy? What membrane
transport processes are carrier mediated? What ones are not?
8. Identify several reasons why the glycocalyx is important to
human survival.
9. How do microvilli and cilia differ in structure and function?
10. What are the functional differences between tight junctions,
gap junctions, and desmosomes?
2.3
The Cytoskeleton
The cytoskeleton is a network of protein filaments and tubules
that structurally support a cell, determine its shape, organize its
contents, direct the movement of substances within the cell, and
contribute to movements of the cell as a whole. It can form a very
dense supportive web in the cytoplasm (fig. 2.15). It is connected
to transmembrane proteins of the plasma membrane and they, in
Organelles
The minute, metabolically active structures within a cell are called
organelles (literally, little organs) because they are to the cell what
organs are to the bodystructures that play individual physiological roles in the survival of the whole (see fig. 2.5). A cell may have
10 billion protein molecules, some of which are powerful enzymes
with the potential to destroy the cell if they are not contained and
isolated from other cellular components. You can imagine the
enormous problem of keeping track of all this material, directing molecules to the correct destinations, and maintaining order
against the incessant tug of disorder. Cells maintain order partly by
compartmentalizing their contents in organelles. Figure2.17 shows
some major organelles.
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16
Microvilli
Microfilaments
Terminal web
Secretory
vesicle in
transport
Lysosome
Desmosome
Kinesin
Microtubule
Intermediate
filaments
Intermediate
filaments
Microtubule
in the process
of assembly
Centrosome
Microtubule
undergoing
disassembly
Nucleus
Mitochondrion
(a)
Basement
membrane
Hemidesmosome
(b)
Figure 2.15 The Cytoskeleton. (a) Components of the cytoskeleton. Few organelles are
shown in order to emphasize the cytoskeleton. Note that all microtubules radiate from the
centrosome; they often serve as tracks for motor proteins transporting organelles. (b) Cells with
their cytoskeletons labeled with fluorescent antibodies, photographed through a fluorescence
microscope. The density of a typical cytoskeleton far exceeds even that shown in part (a).
What cytoskeletal structures shown here also play structural roles in microvilli, cilia, and flagella?
15 m
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(b)
(c)
Microtubule
Endoplasmic Reticulum
Protofilaments
Dynein arms
Tubulin
The Nucleus
The nucleus (fig. 2.17a) is the largest organelle and usually the only
one visible with the light microscope. It contains the cells chromosomes and is therefore the genetic control center of cellular activity. Granular organelles called ribosomes are produced here, and
the early steps in protein synthesis occur here under the direction
of the genes. Most cells have only one nucleus, but there are exceptions. Mature red blood cells have none; they are anuclear.24 A
few cell types are multinucleatehaving 2 to 50 nucleiincluding
some liver cells, skeletal muscle cells, and certain bone-dissolving
and platelet-producing cells.
The nucleus is usually spheroidal to elliptical in shape and
averages about 5 m in diameter. It is surrounded by a nuclear
envelope consisting of two parallel membranes. The envelope is
perforated with nuclear pores, about 30 to 100 nm in diameter,
formed by a ring-shaped complex of proteins. These proteins
regulate molecular traffic into and out of the nucleus and bind
the two membranes together. The inside of the nuclear envelope
is lined by a web of intermediate filaments, like a cage enclosing
the DNA.
The material within the nucleus is called the nucleoplasm. In
most human cells, it contains 46 chromosomes25long strands of
DNA and protein. In nondividing cells, the chromosomes are in
the form of very fine filaments broadly dispersed throughout the
nucleus, visible only with the TEM. Collectively, this material is
called chromatin (CRO-muh-tin). When cells prepare to divide,
The term endoplasmic reticulum (ER) literally means little network within the cytoplasm. It is a system of interconnected channels called cisternae26 (sis-TUR-nee) enclosed by a membrane
(fig. 2.17b). In areas called rough endoplasmic reticulum, the
network consists of parallel, flattened cisternae covered with ribosomes, which give it its rough or granular appearance. The rough
ER is continuous with the outer membrane of the nuclear envelope,
and adjacent cisternae are connected by perpendicular bridges.
In areas called smooth endoplasmic reticulum, the membrane
lacks ribosomes, the cisternae are more tubular in shape, and they
branch more extensively. The cisternae of the smooth ER are continuous with those of the rough ER, so the two are different parts
of the same cytoplasmic network.
The endoplasmic reticulum synthesizes steroids and other lipids, detoxifies alcohol and other drugs, and manufactures all of the
membranes of the cell. Rough ER produces the phospholipids and
proteins of the plasma membrane. It also synthesizes proteins that
are either secreted from the cell or packaged in organelles called
lysosomes. Rough ER is most abundant in cells that synthesize large
amounts of protein, such as antibody-producing cells and cells of
the digestive glands.
Most cells have only a scanty smooth ER, but it is relatively
abundant in cells that engage extensively in detoxification, such as
liver and kidney cells. Long-term abuse of alcohol, barbiturates,
and other drugs leads to tolerance partly because the smooth ER
proliferates and detoxifies the drugs more quickly. Smooth ER is
also abundant in cells that synthesize steroid hormones, for example in the testes and ovaries. Skeletal and cardiac muscle contain
extensive networks of modified smooth ER called sarcoplasmic reticulum, which releases calcium to trigger muscle contraction and
stores the calcium between contractions.
Ribosomes
Golgi Complex
24
26
25
27
cistern = reservoir
Camillo Golgi (18431926), Italian histologist
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18
Nucleolus
Nuclear
envelope
Nuclear
pore
(a) Nucleus
Cisternae
Smooth endoplasmic
reticulum
(b) Endoplasmic reticulum
Cisterna
Golgi vesicle
Protein
crystal
(d) Lysosomes
Protein link
Longitudinal view
Crista
Cristae
Matrix
Microtubules
Cross section
(e) Mitochondrion
(f) Centrioles
Figure 2.17 Major Organelles. (a) Nucleus. (b) Endoplasmic reticulum, showing rough and smooth regions. (c) Golgi complex and Golgi vesicles.
(d)Lysosomes. (e) Mitochondrion. (f ) A pair of centrioles. Centrioles are typically found in pairs, perpendicular to each other so that an electron micrograph
shows one in cross section and one in longitudinal section.
With rare exceptions, only one of these organelles can normally be seen with a typical student light microscope. Which one?
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sorts these proteins, passes them along from one cisterna to the
next, cuts and splices some of them, adds carbohydrates to some
of them, and finally packages the proteins in membrane-bounded
Golgi vesicles. These vesicles bud off the rim of the cisterna farthest from the ER. They are seen in abundance in the neighborhood of the Golgi complex.
Some Golgi vesicles become lysosomes, discussed shortly;
some migrate to the plasma membrane and fuse with it, contributing fresh protein and phospholipid to the membrane; and some
become secretory vesicles that store a cell product, such as breast
milk protein, mucus, or digestive enzymes, for later release by
exocytosis.
1 Protein formed by
ribosomes on rough ER.
2 Protein packaged into transport
vesicle, which buds from ER.
Nucleus
Ribosomes
Golgi
complex
Rough ER
Lysosome
Figure 2.18 Organelle Collaboration in Protein Production. The steps in protein synthesis and secretion are numbered 1 through 6. (1) Acting on
instructions from the nucleus, each ribosome assembles amino acids in the correct sequence to make a particular protein. The protein is threaded into
the cisterna of the rough ER as it is synthesized. The rough ER cuts and splices proteins and may make other modifications. (2) The rough ER packages the
modified protein into transport vesicles that carry it to the Golgi complex. (3) The transport vesicle fuses with a cisterna of the Golgi complex and unloads its
protein. (4) The Golgi complex may further modify the protein. (5) The Golgi complex buds off Golgi vesicles containing the finished protein. (6) Some Golgi
vesicles become secretory vesicles, which travel to the plasma membrane and release the product by exocytosis.
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20
Proteasomes
DEEPER INSIGHT
Lysosomes
Peroxisomes
Peroxisomes (not illustrated) resemble lysosomes but contain different enzymes and are not produced by the Golgi complex. Their
general function is to use molecular oxygen (O2) to oxidize organic
molecules, especially to break down fatty acids into two-carbon
molecules that can be used as an energy source for ATP synthesis. Such reactions produce hydrogen peroxide (H2O2), hence the
name of the organelle. H2O2 is then used to oxidize other molecules, and the excess is broken down to water and oxygen by an enzyme called catalase. Peroxisomes also neutralize free radicals and
detoxify alcohol, other drugs, and a variety of blood-borne toxins.
Peroxisomes occur in nearly all cells but are especially abundant in
the liver and kidneys.
Mitochondria
Mitochondria30 (MY-toe-CON-dree-uh) (singular, mitochondrion) are organelles specialized for a process called aerobic respiration, which synthesizes most of the bodys ATP. They have a
variety of shapes: spheroidal, rod-shaped, bean-shaped, or threadlike (fig. 2.17e), and they constantly move, squirm, and change
shape. Like the nucleus, a mitochondrion is surrounded by a
double membrane. The inner membrane usually has folds called
2.2
Centrioles
Inclusions
Inclusions are of two kinds: accumulated cell products such as
pigments, fat droplets, and granules of glycogen (a starchlike carbohydrate); and internalized foreign matter such as dust, viruses, and
bacteria. Inclusions are never enclosed in a membrane, and unlike
the organelles and cytoskeleton, they are not essential to cell survival.
28
31
crista = crest
centro = central; some = body
32
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2.4
Anaphase
ase
aph
Met
se
ha
op
Pr
M Mitotic phase
G2
Second gap phase
Growth and preparation
for mitosis
is
es
kin
12. Briefly state how each of the following cell components can
be recognized in electron micrographs: the nucleus, a mitochondrion, a lysosome, and a centriole. What is the primary
function of each?
hase
Cy
to
Telo
p
Before You Go On
G1
First gap phase
Growth and normal
metabolic roles
S
Synthesis phase
DNA replication
Interphase
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22
DEEPER INSIGHT
2.3
Cancer
A tumor (neoplasm33) is a mass of tissue produced when the rate of cell
division exceeds the rate of cell death. A malignant34 tumor, or cancer
(fig. 2.20), is especially fast-growing, lacks a confining fibrous capsule, and has cells that are capable of breaking free and spreading to
other organs (metastasizing35). Cancer was named by Hippocrates,
who compared the distended veins in some breast tumors to the
outstretched legs of a crab.36
All cancer is caused by mutations (changes in DNA or chromosome structure), which can be induced by chemicals, viruses, or
radiation or simply occur through errors in DNA replication in the cell
cycle. Agents that cause mutation are called mutagens,37 and those
that induce cancer are also called carcinogens.38 Many forms of cancer stem from mutations in two gene families, the oncogenes and
tumor-suppressor genes. Oncogenes39 are mutated genes that promote the synthesis of excessive amounts of growth factors (chemicals that stimulate cell division) or excessive sensitivity of target cells
to growth factors. Tumor-suppressor (TS) genes inhibit the development of cancer by opposing oncogenes, promoting DNA repair, and
other means. Cancer occurs when TS genes are unable to perform
this function. Oncogenes are like an accelerator to the cell cycle, and
TS genes are like a brake.
Untreated cancer is almost always fatal. Tumors destroy healthy
tissue; they can grow to block major blood vessels or respiratory airways; they can damage blood vessels and cause hemorrhaging; they
can compress and kill brain tissue; and they tend to drain the body of
nutrients and energy as they hungrily consume a disproportionate
share of the bodys oxygen and nutrients.
Cell Division
Cells divide by two mechanisms called mitosis and meiosis.
Meiosis, however, is restricted to one purpose, the production of
eggs and sperm, and is therefore treated in chapter 26 on reproduction. Mitosis serves all the other functions of cell division: the
development of an individual, composed of some 40 trillion cells,
from a one-celled fertilized egg; continued growth of all the organs
after birth; the replacement of cells that die; and the repair of damaged tissues. Four phases of mitosis are recognizableprophase,
metaphase, anaphase, and telophase (fig. 2.21).
In prophase,40 at the outset of mitosis, the chromosomes coil
into short, compact rods that are easier to distribute to daughter cells than the long, delicate chromatin of interphase. At this
stage, a chromosome consists of two genetically identical bodies
called chromatids, joined together at a pinched spot called the
centromere (fig. 2.22). There are 46 chromosomes, two chromatids
33
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1 Prophase
Chromosomes condense
and nuclear envelope
breaks down. Spindle
fibers grow from centrioles.
Centrioles migrate to
opposite poles of cell.
Aster
2
2 Metaphase
Chromosomes lie along midline
of cell. Some spindle fibers
attach to kinetochores.
Fibers of aster attach
to plasma membrane.
Spindle fibers
Centriole
3 Anaphase
Centromeres divide in two.
Spindle fibers pull sister
chromatids to opposite poles
of cell. Each pole (future
daughter cell) now has an
identical set of genes.
4 Telophase
Chromosomes gather
at each pole of cell.
Chromatin decondenses.
New nuclear envelope
appears at each pole.
New nucleoli appear
in each nucleus.
Mitotic spindle
vanishes.
Chromatids
Kinetochore
Cleavage furrow
Daughter
cells in
interphase
Nuclear envelope
re-forming
Chromatin
Nucleolus
chromosomes are relatively easy to observe. The drawings show a hypothetical cell
with only two chromosome pairs. In humans, there are 23 pairs.
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24
Kinetochore
Centromere
Sister
chromatids
(a)
(b)
700 nm
metaphase. From the end of the S phase of the cell cycle to the beginning
of anaphase in mitosis, a chromosome consists of two genetically identical
chromatids. (b) SEM of a metaphase chromosome.
Stem Cells
One of the most controversial scientific issues in the last several years
has been stem-cell research. Stem cells are immature cells with the
ability to develop into one or more types of mature, specialized cells.
Their ability to give rise to a diversity of mature cell types is called
developmental plasticity. Adult stem (AS) cells exist in most of the
bodys organs. Despite the name, they are not limited to adults, but
are found also in fetuses, infants, and children. They multiply and replace older cells that are lost to damage or normal cellular turnover.
Some AS cells are unipotent, able to develop into only one mature
45
cell type, such as the cells that develop into sperm or epidermal squamous cells. Some are multipotent, able to differentiate into multiple
mature cell types, such as certain bone marrow cells that give rise to
multiple types of white blood cells.
Embryonic stem (ES) cells comprise human embryos (technically, preembryos; see chapter 4) of up to 150 cells. They are
pluripotent, able to develop into any type of embryonic or adult
cell. ES cells are easily obtained from the excess embryos created
in fertility clinics when a couple attempts to conceive a child by in
vitro fertilization (IVF). In IVF, eggs are fertilized in glassware and
allowed to develop to about 8 to 16 cells. Some of these are then
transplanted into the mothers uterus. Excess embryos are created
to compensate for the low probability of success. Those that are not
transplanted to the uterus are usually destroyed, but they present a
potential source of stem cells for research and therapy.
Skin and bone marrow adult stem cells have been used in therapy for many years. There is hope that stem cells can be manipulated
to replace a broader range of tissues, such as injured spinal cords,
cardiac muscle damaged by a heart attack, brain tissue lost in Parkinson and Alzheimer diseases, or the insulin-secreting cells needed
by people with diabetes mellitus. Adult stem cells, however, seem to
have limited developmental potential and to be unable to produce
all cell types needed to treat a broad range of diseases. In addition,
they are present in very small numbers and are difficult to isolate
and culture in the quantities needed for therapy. Embryonic stem
cells are easier to obtain and culture and have more developmental flexibility, but their use remains embroiled in political, religious,
and ethical debate. Some would argue that if the excess embryos
from IVF are destined to be destroyed, it would seem sensible to use
them for beneficial purposes. Others argue, however, that potential
medical benefits cannot justify the destruction of a human embryo
or even a preembryo of scarcely more than 100 cells.
Further scientific advances, however, are beginning to defuse
the controversy to some degree. Cell biologists have developed
methods to make adult stem cells reverse course on their developmental paths, going back to a pluripotent state that would enable them to go down new roads. In principle, an AS cell that was
originally destined to become smooth muscle, for example, might
be made to revert to a pluripotent state, then chemically guided
down a path leading to nervous tissue for Alzheimer patients or
persons paralyzed by spinal cord injury. Such modified adult cells,
now called induced pluripotent stem (iPS) cells, have begun to show
great promise for both medical benefit and reduced controversy.
Before You Go On
Answer the following questions to test your understanding of the
preceding section:
6. State what occurs in each of the four phases of the cell cycle.
1
17. State what occurs in each of the four phases of mitosis.
18. Explain how a cell ensures that each of its daughter cells gets
an identical set of genes.
19. Define unipotent, multipotent, and pluripotent stem cells. Give
an example of each.
20. Discuss the advantages and disadvantages of adult and embryonic stem cells for therapy.
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Study Guide
Assess Your Learning Outcomes
To test your knowledge, discuss the following
topics with a study partner or in writing, ideally
from memory.
2.1 The Study of Cells (p. 26)
1. The meaning of cytology and why it is important in medical anatomy and physiology
2. How the light microscope, transmission
electron microscope, and scanning electron
microscope differ, and the relative importance of magnification and resolution
3. The cell shapes denoted by the terms squamous, cuboidal, columnar, polygonal, stellate, spheroidal, ovoid, discoid, fusiform,
and fibrous
4. The distinction between the basal, apical,
and lateral surfaces of an epithelial cell, and
why it is important to differentiate them
5. The size of a micrometer and some common and extreme human cell sizes in terms
of micrometers
6. Reasons why cells cannot grow indefinitely
large; limitations on cell size
7. The meaning of plasma membrane, cytoplasm, nucleoplasm, cytoskeleton, organelles, inclusions, and cytosol
8. Terminology of the fluids on the inside and
outside of a cell
2.2 The Cell Surface (p. 30)
1. Why the cell surface is so important for an
understanding of human function
2. The molecular composition and organization of the plasma membrane
3. The general functions of membrane phospholipids, cholesterol, proteins, glycoproteins, and glycolipids
4. How integral proteins differ from the peripheral proteins of a plasma membrane
5. The diverse physiological roles of membrane proteins, and terms for the proteins
that play each role
6. The importance of filtration through biological membranes, and the principal place
in the body where this is relevant
7. The processes of simple diffusion, osmosis,
reverse osmosis, facilitated diffusion, and
active transport in relation to the plasma
membrane; the relationship of each process
to concentration gradients, the involvement of membrane proteins, and the necessity for ATP
8. The methods of vesicular transport
through the plasma membrane: endocytosis (by phagocytosis, pinocytosis, and
receptor-mediated endocytosis) and exocytosis; how each process is carried out; and
the purposes for which each process may
be used
9. The composition, location, and functions
of a cells glycocalyx
10. The structural and functional differences
between microvilli, cilia, and flagella
11. The cell junctions that bind cells to each
other and to extracellular materialtight
junctions, desmosomes, hemidesmosomes,
c. unipotent.
d. more developmentally limited than
adult stem cells.
e. more difficult to culture and harvest
than adult stem cells.
7. The amount of DNA in a cell doubles
during
a. prophase.
d. the S phase.
b. metaphase.
e. the G2 phase.
c. anaphase.
8. Fusion of a secretory vesicle with the
plasma membrane followed by release of
the vesicles contents is
a. exocytosis.
b. receptor-mediated endocytosis.
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c. active transport.
d. pinocytosis.
e. phagocytosis.
9. Most cellular membranes are made by
a. the nucleus.
b. the cytoskeleton.
c. enzymes in the peroxisomes.
d. the endoplasmic reticulum.
e. replication of existing membranes.
10. Matter can leave a cell by any of the
following means except
a. active transport.
d. pinocytosis.
b. simple diffusion. e. exocytosis.
c. facilitated diffusion.
True or False
Determine which five of the following statements
are false, and briefly explain why.
1. The shape of a cell is determined mainly
by its cytoskeleton.
2. The plasma membrane is composed
mostly of protein.
3. A plasma membrane is too thin to be seen
with the light microscope.
3.
4.
5.
6.
7.
-form
poly-philic
phagoendo-
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