CHAPTER

3
Cell
s
About 200 different types of cells compose your body. Each cell is a
living structural and functional unit that is enclosed by a membrane.
All cells arise from existing cells by the process, in which one cell
divides into two new cells. In your body, different types of cells
fulfill unique roles that support homeostasis and contribute to the
many functional capabilities of the human organism. As you study
the vari­ ous parts of a cell and their relationships to each other,
you will learn that cell structure and function are intimately related.

Looking Back to Move Ahead...

, Levels of Organization and Body Systems (Section 1.1)
• Ions, Molecules, and Compounds (Section 2.1)
• Carbohydrates (Section 2.2)
• Lipids (Section 2.2)
• Proteins (Section 2.2)
• Deoxyribonucleic Acid (DNA) and Ribonucleic.Acid
(RNA) (Section 2.2)
Cancer cells dividing

Q Did you ever wonder why cancer is so difficult to

treat? You can find out under Common Disorders:
Cancer.

E] A Generalized ease of study, we can divide a cell into three main parts: the
plasma membrane, cytoplasm, and nucleus.

View 1. The plasma membrane forms a cell's flexible outer surface,

of the Cell separat­ ing the cell's internal environment (inside the cell) from
its external environment (outside the cell). It regulates the flow
OBJECTIVE
of materials into and out of a cell to maintain the appropriate
• Name and describe the three main parts of a environment for normal cellular activities. The plasma membrane
cell. also plays a key role in communication among cells and between
cells and their ex­ternal environment.
Cell biology is the study of cellular structure and function. Figure 3.1 2. The cytoplasm (Si-to-plasm; plasm = formed or molded) con­
is a generalized view of a cell that shows the main cellular compo­ sists of all the cellular contents between the plasma membrane
nents. Though some body cells lack some cellular structures shown and the nucleus. Cytoplasm can be divided into two compo­
in this diagram, many body cells include most of these components. nents: cytosol and organelles. Cytosol (Si-to-sol) is the liquid
For

40
 3.2 The Plasma Membrane 41

hj§,l;jJHI Generalized view of a body cell.

The cell is the basic, living, structural and functional unit of the
body.

; " " " ' " co,

m
Nucleus:
Chromatin
Cytoskeleton: Proteasome ribosome
Nuclear pore
s
Microtubule-- - - - - - - ; : : : - - ! S : ; l \ Nuclear
envelope
Microfilament /

.
Nucleolu
Intermediate filament 'itfit";" s
• l,J ;::E:;::: ---
M1crov,111 - - - - ­
Glycogen
Centrosome: • • :; ;i,:,
, ' granules
Pericentriolar Plasma
' • membrane
- • r-'
material •f
Centrioles- : : : : . . , , , . , , - .-._.,,,...- " , \o
L,, •
= . . i - - Rough
Secretory endoplasmic
reticulum
vesicle (ER)
Ribosome
Lysosome attached to
ER

Golgi complex
Peroxiso

Milochondrion ..
Microtubule

Microfilament

Sectional
view

Q What are the three principal parts of a

cell?

portion of cytoplasm that consists mostly of water plus

dissolved solutes and suspended particles. It is also called The Plasma Membrane
intracellular fluid. Within the cytosol are several different types
of organelles
(or-ga-NELZ = little organs), each of which has a characteristic
OBJECTIVE
structure and specific functions.
• Describe the structure and functions of the plasma
3. The nucleus (NOO-kle-us = nut kernel) is the largest organelle of membrane.
a cell. The nucleus acts as the control center for a cell because it
con­ tains the genes, which control cellular structure and most
cellular The plasma membrane is a flexible yet sturdy barrier that consists
activities. mostly of lipids and proteins. The basic framework of the plasma
Checkpoint membrane is the lipid bilayer, two tail-to-tail layers made up of
1. what are the general functions of the three main parts of a three types of lipid molecules: phosphalipids (lipids that contain
cell? phosphorus), cholesterol, and glycolipids (lipids attached to

Ca - ·Sc· nn
 42 CHAPTER 3 Cells

OfuiMJii Chemistry and structure of the plasma

membrane.
The plasma membrane consists mostly of phospholipids, arranged in a bilayer, and
proteins, most f which are glycoproteins.

3. Helps identify the cell to other

1. Acts as a barrier separating cells (e.g., immune cells).
inside and outside of the 4. Participatesin
cell. intercellular signaling.
2. Controls the flow of substances Channel
into and out of protein
the cell. Extracellular
fluid
Glycoproteln:
carbohydrate Lipid
Protein } bilayer

Peripheral protein

lipid:
ohydrate

Phosphollplds:
. :
. ; , -
Head - - - - - , ,
er,· ,-1
j'
I
Fatty •
acid 'j i';
tails
Peripheral protein

Q Name several functions carried out by membrane

proteins.

carbohydrates; glyco = carbohydrate) (Figure 3.2). The proteins move a substance from one side of the membrane to the other
in a membrane are of two types-integraland peripheral (Figure (see Figure 3.6). Large molecules such as proteins are unable to
3.2). Integral proteins extend into or through the lipid bilayer. pass through the plasma membrane except by transport within
Perlph• era/ proteins (pe-RIF-er-al) are loosely attached to the vesicles (discussed later in this chapter).
exterior or interior surface of the membrane. Some peripheral Most functions of the plasma membrane depend on the types
proteins, called gfycoprateins, are proteins attached to of proteins that are present. Integral proteins called receptors
carbohydrates. recognize and bind a specific molecule that governs some cellular
The plasma membrane allows some substances to move into function, for example, a hormone such as insulin. Some integral
and out of the cell but restricts the passage of other substances. This proteins act as enzymes, speeding up specific chemical reactions.
property of membranes is called selective permeability (per'-me-a­ Membrane glyco­ proteins and glycolipids often are cell identity
BIL-i-te). The lipid bilayer part of the membrane is permeable to wa­ markers. They enable a cell to recognize other cells of its own kind
ter and to nonpolar (lipid-soluble) molecules, such as fatty acids, during tissue formation, or to recognize and respond to potentially
fat-soluble vitamins, steroids, oxygen, and carbon dioxide. The lipid dangerous foreign cells.
bilayer is not permeable to ions and large, uncharged polar mole­
cules such as glucose and amino acids. These small and medium­ Checkpoint
sized water-soluble materials may cross the membrane with the 2. What molecules make up the plasma membrane and
assistance of integral proteins. Some integral proteins form ion what are theirfunctions7
channels through which specific ions, such as potassium ions (K+),
3. What is meant by selective permeability?
can move into and out of cells (see Figure 3.S). Other membrane
proteins act as carriers (transporters), which change shape as they

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 a I(,(.tr.4-S ti{\I e.i.t- ..., OA{•
L < (,
I• ,:{i,,l.l 3.3 Transport Across th Plasma Membrane 4]
L
llO f dl j ; j J f f l
Prlnclple of diffusion. A crystal of dye placed in a cy i der
Transport Across the of water dissolves (a}, and there is net diffusion lrom the region of higher dye
concentration to regions of lower dye concentration (b}. At equilibrium (c),
Plasma Membrane dye concentration is uniform throughout the solution.

At equilibrium, net diffusion slops but random movemenu continue.

• Desen'be the processes that transport substances across the

plasma membrane.

Movement of materials across its plasma membrane is essential to

the life of a cell. Certain substances must move into the cell to
support metabolicreactions. Other materials must be moved out
because they have been°produced by the cell for export or are
cellular waste products . Before discussing how ,.;;-aterials move into
and out of a cell, we need to understand what exactly IS being
moved as well as ffie form it needs to take to make-its journey.•
•Aboutlwo-thiffis of the fluid in your body is contained inside
body cells and is called intracellular fluid (/CF) (intra- = within). ICF,
c a t e d earlier, is actually the cytosol of a cell. Fluid o e body cells Beginnin Intermediate Equilibrium
as indi­
is called extracellular fluid tf.Ci:rrexfro- = outside). The ECF int he g (b) (c)
micro­ scopic spaces between the cells of tissues is interstitial fluid (a)
(in;·ter­ AndyWnhn
STISH-al; inter-'= betwe-en) The ECf: iii oiood vessels is called Q How does simple
ik
diffusion differ from f«jlitated diffusi ?
blood plasma (PLAZ-ma), and that in lym atIc vessels is called
lymph. The ECF within and around the brain and spinal cord of the substance iffuse from the region of hieh concentration to
iscalled cerebrospinal fluid (CSF) (se-re'-bro-SPl-nal). . the region of low concentration than in the opposite direction. The
Materials dissolved in body fluids includ gases, n u t r i Q diffusion of more molecules in one direction than the other is called
and other substances neede'atomaintain life.Ali net diffusion. Substances undergoing net diffusion move from a high
a t e r i ' a l - d i s s o l v a fluid is called a solute, and the fluid in which•it to a low concentration, or down their concentration gradient After
is dissolved is'the· solvent. Body fluids are dilute solutionsin which a some time, equilibrium (e-kwi-LIB-re-um) is reached: The substance
va'riety of solutes are dissolved in a·very familiar solvent, waler. The becomes evenly distributed throughout the solution and the
amount of a solute in a solution isits concentratiO!], A concentration concentration gradient disappears.
gradient isa differe ce i Placing a crystal of dye in a water-filled container provides an
co centration between two different areas, for' xampfe, the ICF example of diffusion (Figure 3.3). At the beginning, the color is most
and· _ intense just next to the crystal because the crystal is dissolving
-
them) Solutes
are said°to move downmoving fromconcentration
or witnthe a·high-concentration_Mea
gradient. and the dye concentration is greatest there. At increasing
(wherethere
Sol- are, more of
utes mov(ngfrom them) to a low-concentratio
a low-concentration areah-i g hare.a
- c (where' color is lighter
distances, the and lighter because the dye concentration is lower
there
a t i are
rarea o n fewer of d t moo v e u p or against the concen ra i n and lower. The dye molecules undergo net diffusion, down their
radient. u b s t a n c e s move a • s cellular membranes by concentration gradient, until they are evenly mixed in the water. At
passive proc'esses and active processes. sii e processes, 1ri equilibrium the solution has a uniform color. In the example of
which·a subs ance moves do-.yri its concel)trationgradient through dye diffusion, no membrane was involved. Substances may also
the membrane, using only its own energy o otlon diffuse across a membrane, if the membrane is permeable to them.
(kinetic energy), include simple diffusion and osmosis.1/\ottive Now that you have a basic understanding of the nature of diffu-
processes, cellular energy, usually in the form of ATP, i d to " p s sion, we will consider two types of diffusion: simple diffusion and
• the substance through the membran "uphill" again t its fa- cilitated diffusion.
concentration gradient. Ari example is active transport. Another way
that some substances may enter and leave cells is an active process SIMPLE DIFFUSION In simple diffusion, substances diffuse across
in wl'l!ch tiny membrane sacs r ferred to as vesicles are used. a membrane through the.lipid bilayer (Figure 3.4). Lipid-soluble s
b­ s t h t move across membranes by simple
diffusion through the Ii id bila er inclLJie.oxygen, ca!,_bon dioxide,
fatty
and nlacids;
o g e steroids;
n ases; and fat-soluble vitamins (A D, E and K . Polar
Diffusion: The Principle Diffusion (di-FU-zhun; diffus- = molecules su WJter and u a_also move th rough the lipid
spreading) is a passive process in which a substance moves due to Simple
bilayer. diffusion through the lipid b_ilayerisimportant in theexchange
its kinetic energy. If a particular substance is present in

-
of oxygen and_£arbon dioxide betw en ,!,lood and body cells and
highconcentration in one area and in low concentration in another
area, more particles between- blood and air"within the lu ngs during breathing. It·also
- - -
is

Ca
 44 CHAPTER 3
Cells
OM'Mi&iSimple diffusion. Lipid- i4(31j;jjiJ flcllitated diffusion of potassium Ions (K' I through ii gated
solublemoleculesdiffusethrough t h e l i p i d b i l a y K' channel. A gatedchannel isoneinwhicha portionof
thechannelprotein acts as a gate to open or close thechannel'spore
In simple diffusion there is a net (greater) movement of tothepassageof ions.
substances from a region of their higher concentration to a region Ion channels are Integral membrane proteins that
of their lower concentration, allowspecificsmall, Inorganic ions to pass across
themembrane.
Extracellula Extracellular
r ftuid
fluid

Cytosol
··•••
_,:'./
Gate
open Gate
Cytoso closed
0 Lipid- l
solublemolecule Detailsof the W channel
Q What are some examples of substances that Q Is the concentration of K+higher in the cytosol of
diffuse through the lipid bilayer?
body cells or in the extracellular fluid?

the transport method for absorption of lipid-soluble nutrients and 8 As the carrier undergoes a change in shape, glucose
release of some wastes from body cells. passes
through the membrane.
FACILITATED DIFFUSION Some substances that cannot move
through the lipid bilayer by simple diffusion do cross the plasma 8 The carrier releases glucose on the other side of the
membrane by a passive process called facilitated diffusion. In 14iJlijlik)
membrane.
FaclUtated diffusion of glucose across ii plasma
this process, an integral membrane protein assists a specific membrane using ii carrier. The carrier protein binds to glucosein the
extracellular fluid and releases it into the cytosol.
substance
move across to the membrane. The membrane protein can either
from channels or function as carriers.
Facilitated diffusion across a membrane involving a carrier is
c i l i t a t e d diff;;;ion i-;;;olving ion channels, ions move
an important mechanism for transporting sugars such as glucose,
down
fructose, and galactoseinto cells.
their concentration gradients across the lipid bilayer. Most mem­
brane channels are ion channels, which allow a specific type of
ion to move across the membrane through the channel's pore. In Glucose Carrier Extracellular
typi• cal plasma membranes, the most common ion channels are
I protein fluid
(glucose
selec­ tive for K+ (potassium ions) or c1- (chloride ions); fewer • -._ tr11naport11r)
channels are available for Na+ (sodium ions) or ca2+(calcium
lucose
ions). Many ion channels are gated; that is, a portion of the
channel protein acts as a •gate," moving in one direction to open
the pore and in another direction to close it (Figure 3.5). When
the gates are open, ions diffuse into or out of cells, down their
concentration gradient. Gated channels are important for the
production of electrical sig­naIs by body cells.
In facilitated diffusion involving a carrier, the substance binds
to
a specific carrier on one side of the membrane and is released on
the other side after the carrier undergoes a change in shape.
Glucos
Substances that move across plasma membranes by
e
facilitated Cytoso
diffusion involving carriers include glucose, fructose, galactose, and l
0
someGlucose
vitamins.Glucose enterscarrier
binds to a glucose many bodyon cellsby
protein facilitated
the outside Q How does insulin alter glucose transport by
diffusion as follows (Figure3.6): facilitated diffusion? , •
surface
' ·
of the membrane.

Sc •
•dw 'th CamScanner
 3.J Transport Across the Plasma Membrane 45

The selective permeability of the plasma membrane is often the nuid rises in the tube, its water pressure forces some water
regulated to achieve homeostasis. For example, the hormone insulin molecules from the sac back into the beaker. At equilibrium, just
promotes the insertion of more glucose carriers into the plasma as many water molecules are moving into the beaker due to the
membranes of certain cells. 'tlius, t e effeJ:t of water pressure as are moving into the sac due to osmosis.
insu.lin..lLlq.J_nqease entry of g ose mtooo'd cells b eans
A solution containing solute particles that cannot pass through
otfac1hlated dlrtosron.
a membrane exerts a pressure on the membrane, called osmotic
Osm,osis (oz-MO-sis) is a passive process in pres­ sure. The osmotic pressure of a solution depends on the
which ere is a net movement of water through a selectively concentra­ tion of its solute particles-the higher the solute
permeable membrane. Wat r moves by osmosi's-from an area of concentration, the higher the solution's osmotic pressure. Because
higher water concentration to an area ofiowerwaterconcen"tration°(or the osmotic pressure of cytosol and interstitial fluid is the same, cell
from a area of1ower solute-concentration to an area of hi her solute volume remains con­ stant. Cells neither shrink due to water loss by
ioncent;atian). Wa! r mq!ecules pass t h ; g h plasma m:mbranes osmosis nor swell due to water gain by osmosis.
in iWo places: th ough th lieid bilayer and through integral Any solution in which cells maintain their normal shape and vol­
membrane proteins ume is called ao Isotonic solutjaa (i'-so-TON-ik; iso- = same;
that function as water channels. t o n ic = tension). This is a solution in which the concentrations of
The experiment ln Figure 3.7 demonstrates solutes are the same on both sides. For example, a 0.9% NaCl
osmosis. (sodium chloride, or table salt) solution, called a normal saline
solution, is isotonic for red blood cells. When red blood cells are
1. A sac mad; of cellophane, a selectively permeable membrane
bathed in 0.9% NaCl, water molecules enter and exit the cells at
that permits water but not sucrose (sugar) molecules to pass, is
the same rate, allowing the red blood cells to maintain their
filled with a"so.lution that is 20% sucrose and 80% water. the
normal shape and volume (Figure
upper part of the cellophane sac is wrapped tightly about a
3.Sa).
stopper through which a glass tulje is fitted.
If red blood cells are placed in aJ,ypotonlc solution !hi'-po-TON­
2. The sac is then placed into a-beaker containing pure (100%)
ik; hypo- = less than), a solution that has a lower concentration of
water (Figure 3.7a). Notice that the cellophane now separates two
solutes (higher concentration of water) than the cytosol inside the
fluids having different water concentrations.
red blood cells (Figure 3.Sb), water molecules enter the cells by
3. Water begins to move by osmosis from the region where its osmosis
con­ centration is higher (100% water in the beaker) through the faster than they leave. This situation causes the red blood cells
cello­ phane to where its concentration is lower (80% water to swell and eventually to burst. Rupture of red blood cells is
inside the sac). Because the cellophane is not permeable to called
sucrose, how­ ever, all the sucrose molecules remain inside the hemolysis (he-MOL-i-sis).
sac. Ahypertonic solutio.aJhi'-per-TON-ik; hyper-= greater than)
4. As water moves into the sac, the volume of the sucrose has a highe concentration of solutes (lower concentration of water)
bfai);ij&I Principle of osmosis.
solution increases and the fluid rises into the glass tube (Figure than does the cytosol inside red blood cells (Figure 3.Sc). When
3. 7b). As cells are placed in a hypertonic solution, water molecules move out
Osmosisis the net movement of water molecules through a of the cells by osmosis faster than they enter, causing the cells to
selectively permeable membrane. shrink. Such shrinkage of red blood cells is called crenation (kre-NA-
Clinical Connection
shun).
Grass
tube Medical Uses of Isotonic, Hypertonic, and
Rubber Hypotonic Solutions
stopper
RBCs and other body cells may be damaged or destroyed If exposed to
O 00
hypertonic or hypotonic solutions. For this reason, most Intravenous
0
Sucrose 0 o o (IV) solutions, liquids Infused into the blood of a vein, are Isotonic:
0
molecules o o solutions. Examples are Isotonic saline (0.9% NaCl) and DSW, which
o
ooo
o0 0
stands foi dextrose 5% in water. Sometimes infusion of hypertonlc:
Select,vel 0o solutions Is use­ful to treat patients who have cerebral edema, excess
O
permeable O o
membrane o o
Interstitial fluid ir the brain. Infusion of such a solution relieves fluid
o o,.., o overload by causini osmosis of water from interstitial fluid into the
"'o .,• o• C
Water ,'
blood. The kidneys the1 excrete the excess water from the blood into
molecules the urine. Hypotonlc solu tlons, given either orally or through an IV,
can be used to treat people wh1 are dehydrated. The water in the
(a) At start of experiment (b) At equilibrium hypotonic solution moves from the bloo1 into interstitial fluid and then
into body cells to rehydrate them. Water an most sports drinks that you
Q Will the fluid level in the tube continue to rise until the consume to "rehydrate" after a workout a, hypotonic relative to your
sucrose concentrations are the same in the beaker and in the body cells.
sac?

ned w'ilh Ca
46 CHAPTER 3 Cells

OfalMJH,I Prlndpleofosmoslnpplled to red blood cells (RBCs), The arrows Indicatethe

direction and degree of water movement Into and out of the cells.The scanning electron
mlcrographs have magnificationsof 15,000X.

An Isotonic solution Is one In which cells maintaintheir normal shape and volume.

Isotonic Hypotonlc Hypertonlc 1

solution aolutlon aolutlon j
f
i========-
l
6 ,) j
Gm1soox
(
(b) ABC undergoes (c) ABC undergoes
hemolysis crenallon

Question Isotonic Hypotonlc: Hypertonic:

Is the membrane Yes Yes Yes

per- meable to
water7
Where is the solute Equal on both sides of cell Inside the cell Outside the cell
concentration higher?

• Where is the solute Equal on both sides of cell Outside the cell Inside the cell
concentration lower?

Where is the water Equal on both sides of cell Outside the cell Inside the cell
concentration higher?

Where is the water Equal on both sides of cell Inside the cell Outside the cell
concentration lower?

Which way will there be None Outside to inside Inside to outside

net movement of water?

What will happen to the Stay the same Swell (cell may burst Shrink
cell size? open)

Q Will a 2% solution of NaCl cause hemolysis or crenation of

RBCs?

Active.,ELPresses against the Na+ concentration gradient. At the same time, the
pump moves potassium ions into cells against the K+ concentration
Active Transport Active transport is an active process in gradient. Because K+ and Na+ slowly leak back across the plasma
which cellular energy is used to transport substances across membrane down their gradients, the sodium-potassium pumps
the membrane against a concentration gradient (from an area of must operate continually to maintain a low concentration of Na+
low concentration to an area of high concentration). and a high concentration of K+ in the cytosol. These differing con­
Energy derived from splitting ATP changes the shape of a centrations are crucial for osmotic balance of the two fluids and
carrier protein, called a pump, which moves a substance across a also for the ability of some cells to generate electrical signals such
cellular membrane against its concentration gradient. A typical as action potentials.
body cell ex­pends about 40% of its ATP on active transport. Drugs Figure 3.9 shows how the sodium-potassium pump operates.
that turn off ATP production, such as the poison cyanide, are lethal
0 Three sodium ions (Na+) in the cytosol bind to the pump protein.
because they shut down active transport in cells throughout the
body. Substances 8 Na+ binding triggers the splitting of ATP into ADP plus a
transported across the plasma membrane by active transport phosphate
are mainly ions, primarily Na+, K+, H+, Cai+, 1·, and ct·. group (®), which also becomes attached to the pump protein.
The most important active transport pump expels sodium ions This chemical reaction changes the shape of the pump protein,
(Na+) from cells and brings in potassium ions (K+}. The pump pro­ expelling the three Na+ into the extracellular fluid. The
tein also acts as an enzyme to split ATP. Because of the ions it changed shape of the pump protein then favors binding of two
moves, this p u mp is called the sodium.:potassium (Na+-K+) pump. potassium ions (K+) in the extracellular fluid to the pump
All cells have thousands of sodium-potassiumpumps in their plasma protein.
membranes. These pumps maintain a low concentration of sodium
E) The binding of K+ causes the pump protein to release the
ions in the cytosol by pumping Na+ into the extracellular fluid
phosphate group, which causes the pump protein to return to its
original shape.

-ned w C ·Sc
 3.3 Transport Across the PlasmaMembrane 47

I J t S , M i U Operation of the sodlum-potanlumpump, Sodium ions (Na') are expelled from the
cell, and potassium ions (K ' ) are Imported Into the cell. The pump does not work unless Na· and ATP
are present in the cytosol and K• Is present in the extracellular fluid.

The sodium-potassium pump maintainsa low intracellular concentrationof Na•.

Ex1racellular fluid

3 sodium ions (Na•) from the Na" binding triggers ATP to 2 potassium ions (K") The release of the(J
cytosol bind to the inside surface bind to the pu and land on the outside causes the pump to
of the sodium-potassium pump. be split into ADP and! surface of the pump return to its original
JJ(phosphate). The energy and cause the shape, which moves
from ATP splitting causes the K' into the cell.
the protein to change shape, to be released.
which moves the Na• to the
outside.
Q What is the role of ATP in the operation of this
pump?

0 As the pump protein returns to its original shape, it releases the receptor, causing the cell to extend projections of its plasma mem­
two K+ into the cytosol. At this point, the pump is ready again to brane and cytoplasm, called pseudapods (SOO-do-pods;pseudo- ==
bind Na+, and the cycle repeats. false;-pods == feet). Two or more pseudopods surround the particle,
and portions of their membranes fuse to form a vesicle called a
Tr a n s p o r t in Vesicles A vesicle (VES-i-kul) is a small round phogosome that enters the cytoplasm. The phagosome fuses with
sac formed by budding off from an existing membrane. Vesicles one or more lysosomes, and lysosomal enzymes break down the
transport substances from one structure to another within cells, take in ingested material. In most cases, any undigested materials may
substances from extracellular fluid, and release substances into remain indefinitely in a vesicle called a residual body or be excreted
extracellular fluid. Movement of vesicles requires energy supplied by the cell.
by ATP and is therefore an active process. The two main types of Phagocytosis occurs only in phagocytes, cells that are spe­
transport in vesicles between a cell and the extracellular fluid that cialized to engulf and destroy bacteria and other foreign substances.
surrounds it are (1) endocytasis (en'-d6-si-T6-sis; endo- = within), in Phagocytes include certain types of white blood cells and mac­
which materials move into a cell in a vesicle formed from the plasma rophages, which are present in most body tissues. The process of
membrane, and (2) exacytosis (ek'-s6-si-T6-sis; exo- = out), in which phagocytosis is a vital defense mechanism that helps protect th
materials move out of a cell by the fusion of a vesicle formed inside a body from disease.
cell with the plasma membrane.
2. Bulk-phase endocytosis. In bulk-phase endocytosis (pinocytosis)
or "cell drinking," cells take up tiny droplets of extracellular fluid.
ENDOCYTOStS Substances brought into the cell by endocytosis are
The process occurs in most body cells and takes in any and all
surrounded by a piece of the plasma membrane, which buds off inside
solutes dissolved in the extracellular fluid. During bulk-phase en­
the cell to form a vesicle containing the ingested substances. The two
types of endocytosis we will consider are phagocytosis and bulk­ docytosis the plasma membrane folds inward and forms a vesicle

phase endocytosis. containing a droplet of extracellular fluid. The vesicle detaches or

"pinches off" from the plasma membrane and enters the cytosol.
1. Phagocytosis. In phagacytasis (fag'-6-si-T6-sis; phogo- == to eat) Within the cell, the vesicle·fuses with a lysosome, where enzymes
or "cell eating," large solid particles, such as whole bacteria or vi­ degrade the engulfed solutes. The resulting smaller molecules
ruses or aged or dead cells, are taken in by the cell (Figure 3.10). such as amino acids and fatty acids, leave the lysosome to be used
Phagocytosis begins as the particle binds to a plasma membrane elsewhere in the cell.

ned w'ilh
48 CHAPTER 3 Cells

■ U41j;jJHt,1Phagocytosls.

Phagocytosis is a vital defense mechanism that helps protect the body from disease.

Phagosome

Digestive
enzymes

Fusion of
lysosome and
phagosome

Digestion by
lysosomat
enzymes
B
Undigested a
materials c
t
Pseudopods
e
r
i
White blood
u
cell
m
Sebastian Kaulitzki/Shutterstock
(a) Diagram of the process (b) White blood cell engulfing
a bacterium
Q What triggers pseudopod
formation?
Exocnos1s In contrast with endocytosis, which brings materials
i nto a cell, exocytosis results in secretion, the liberation of materials
from a cell. All cells carry out e,c:ocytosis, but it is especially impor­
B Cytoplasm
tant in t w o types of cells: (1) secretory cells that liberate digestive
enzymes, hormones, mucus, or other secretions; (2) nerve cells that OBJECTIVE
release substances called neurotransmitters via exocytosis (see
Figure 9.7). During exocytosis, membrane-enclosed vesicles called • Describe the structure and functions of cytoplasm, cytosol, and
secretory vesicles form inside the cell, fuse with the plasma mem­ organelles.
brane, and release their contents into the extracellular fluid.
Segments ofthe plasma membrane lost through endocytosis are
Cytoplasm consists of all of the cellular contents between the
recovered or recycled by exocytosis. The balance between endocyto­
plasma membrane and the nucleus and includes both cytosol and
sis and exocytosis keeps the surface area of a cell's plasma membrane
organelles.
relatively constant.
Table 3.1 summarizes the processes by which materials move
i nto a n d o u t of cells.
Cytosol
Checkpoint
The cytoso/ (intracellular fluid') is the liquid portion of the cyto­
4. What is the key difference between passive and active processes? plasm that surrounds organelles and accounts for about 55% of the
5. How does simple diffusion compare to facilitated diffusion? total cell volume. Although cytosol varies in composition and
6. In what ways are endocytosis and exocytosis similar and different? consistency from one part of a cell to another, typically it is 75-90%
water plus various dissolved solutes and suspended particles.

• nedw I: Ca ·Scn
 I

l.◄ Cyt o p ln m 49

TABLE 3,1 Transport of M•terl •l s Into •nd out of Cells

Transport Process Description Substancn Transpor1 d

Passive Processes Movement of subst.ancesdown a conc.entratlongradient until
equilibrium Is reached; do not require cellular energy In the
form of ATP
Diffusion Movement of a substanceby kinetic energydown a
concentration gradient until equilibrium Is reached
Simple diffusion Passive movement of a substance through the IJpld bilayer of Lipid•solublemolecules: oxygen,carbon
the plasmamembrane dioxide, and nitrogen gases; fatty acids;
steroids; and lat-soluble vitamins (A, D, E,
K). Polarmolecules: water and urea
Facilitated diffusion Passivemovement of a subslance down Its concentration K•, c1-, Na', Ca'·, glucose, fructose,
gradient aided by Ionchannels and carriers galactose, and some vitamins
Osmosis Movement of water molecules across a selectively permeable Water
membrane from an area of higher water concentration to an area
of lower water concentration

Active Processes Movement of substances against a concentration gradient;

requires cellular energy in the form of ATP
Active Transport Transport in which cell expends energy to move a substance Na., K., Ca'., H+, 1-, c1-, and other
across the membrane against its concentration gradient aided by ions
membrane proteins that act as pumps; these integral membrane
proteins use energysupplied by ATP
Transport In Vesicles Movement of substances into or out of a cell in vesicles that bud
from the plasma membrane; requiresenergy supplied by ATP
Endocytosis Movement of substances into a cell in vesicles
Phogocytosis "Cell eating"; movement of a solid particle into a cell Bacteria, viruses, and aged or dead
after pseudopods engulf it cells
Bulk•phose endocytosis "Cell drinking"; movement of extracellular fluid into a cell by Solutes in extracellular fluid
lnfolding of plasma membrane
Exocytosls Movement of substances out of a cell in secretory vesicles that Neurotransmitters, hormones, and
fuse with the plasma membrane and release their contents into digestive enzymes
the extracellular fluid

Among these are various ions, glucose, amino acids, fatty acids, line the small intestine. Some microfilaments extend beyond the
proteins, lipids, ATP, and waste products. Some cells also contain plasma membrane and help cells attach to one another or to extra­
lipid droplets that contain triglycerides and glycogen granules, clus­ cellular materials.
ters of glycogen molecules (see Figure 3.1). The cytosol is the site With respect to movement, microfilaments are involved in
of many of the chemical reactions that maintain cell structures and muscle contraction,cell division, and cell locomotion.Microfilament­
allow cellular growth. assisted movements include the migration of embryonic cells dur­
Extending throughout the cytosol, the cytoskeleton is a network ing development, the invasion of tissues by white blood cells
of three different types of protein filaments: microfilaments, interme• to fight infection, and the migration of skin cells during wound
diate filaments, and microtubules. healing.
The thinnest elements of the cytoskeleton are the microfila­ As their name suggests, intermediate filaments are thicker
ments (mi-kro-FIL-a•ments), which are concentrated at the pe• than microfilaments but thinner than microtubules (Figure J. l l b ) .
riphery of a cell and contribute to the cell's strength and shape They are found in parts of cells subject to tension (such as stretch­
{Figure 3. l l a). Microfilaments have two general functions: pro­ ing). help hold organelles such as the nucleus in place, and help
viding mechanical support and helping generate movements. attach cells to one another.
They also anchor the cytoskeleton to integral proteins in the plasma The largest of the cytoskeletal components, microtubules (mi-kr6-
membrane and provide support for microscopic, fingerlike projec­ TOO-buls) are long, hollow tubes (Figure l . l l c ) . Microtubulesr
tions of the plasma membrane called microvilli {mi-kr6-VIL-i; help determine cell shape and function in both the movement of
micro- = small; -villi= tufts of hair; singular is microvillus). organelles, such as secretory vesicles, within a cell and the migration
Because of chromosomes during cell division. They also are responsible for
they greatly increase the surface area of the cell, microvilli are movements of cilia and flagella.
abundant on cells involved in absorption, such as the cells that

ned w'ilh
 50 CHAPTER 3 Cells

nfoil;\Jlfl (Figure 3.12). The two centrioles are cylindrical structures, each
Cytoskeleton of which is composed of nine clusters of three microtubules(a
triplet) arranged in a circular pattern. Surrounding the centrioles
Extending throughout thecytosol,thecytoskeletonis a networkof
three kinds of protein filaments:microfilaments, is the pericentrio/ar matrix (per'-e-sen'-tre-6-lar), containing
intermediatefilaments, and microtubules. hundreds of ring-shaped proteins called tubulins. The tubulins
are the organizing centers for growth of the mitotic spindle,
Functions of the Cytoskeleton which plays a critical role in cell division, and for microtubule
formation in nondividing cells.
1. Serves as a scaffoldthat helpsto determinea cell'sshape and to
organize the cellular contents.
2. Aidsmovement of organelleswithinthe cell, of chromosomesduring Cilia and Flagella Microtubules are the main structural
cell division, and of wholecellssuch as and functional components of cilia and flagella, both of which
phagocytes. are motile projections of the cell surface. Cilia (SIL-e-a;
singular
is cilium = eyelash) are numerous, short, hairlike projections that
Microvillu extend from the surface of the cell (see Figure 3.1). In the human
s
body, cilia propel fluids across the surfaces of cells that are firmly

aem zr• anchored in place. The coordinated movement of many cilia on

the surface of a cell causes a steady movement of fluid along
Mlcrofllaments (a) Microfilament the cell's surface. Many cells of the respiratory tract, for
example, have hundreds of cilia that help sweep foreign
particles trapped in mucus away from the lungs. Their
movement is paralyzed by
nicotine in cigarette smoke. For this reason, smokers cough often

Uidli;lilfj
<'.entrosome.
Intermediate The pericentriolar material of a centrosome organizes the
filaments
mitotic spindle during cell division.
(b) Intermediate filament
Functions of Centrosomes
The pericentriolarmatrixof the centrosomecontains tubulins that
build microtubulesin nondividingcells and form the
mitoticspindleduring cell division.

(c) Microtubule

Q Which cytoskeletal components help form the structure of

centrioles, cilia, and flagella?

Microtubules
Organelles (triplets)

Organelles are specialized structures inside cells that have

character­istic shapes and specific functions. Each type of organelle
is a func­ tional compartment where specific processes take place,
and each has its own unique set of enzymes.

Centrosome The centrosome, located near the nucleus,

has two components-a pair of centrioles and pericentriolar Q What are the components of the
matrix centrosome?

Sc..nner
 l.-4 Cytoplnm 51

to remove foreign particles from their airways. Cells that line the li[%1l;IIHEI AlbosOfflH.
u t e r i n e (fallopian) tubes also have cilia that sweep oocytes
(egg cells) tow ard the uterus. Ribosomes, ttie sitesof proteinsynthesis, consist of a largesubunit and
Flagella (fla-JEL-a; singular is flagellum = whip) are similar in a small subunlL
structure to cilia but are much longer (see Figure 3.1). Flagella usu­
ally move an entire cell. The only example of a flagellum in the human Functions of Ribosomes
body is a sperm cell"s tail, which propels the sperm toward its possible 1. Ribosomes associated wilh endoplasmic reticulum synthesize
u n i o n with an oocyte. proteins destined for insertion In the plasma membrane
or secretion from the cell.
Ribosomes Ribosomes (Ri-b6-s6ms; -somes = bodies) are 2. Freerib omessynthesizeproteinsused In the c y t ol.
the sites of protein synthesis. Ribosomes are named for their high
content of ribo-nucleic acid (RNA). Besides ribosomal RNA (rRNA),
these tiny organelles contain ribosomal proteins. Structurally,
a ribosome consists of two subunits, large and small, one about
half the size of the other (Figure 3.13). The large and small subunits
are made in the nucleolus of the nucleus. Later, they exit the
nucleus and are assembled in the cytoplasm, where they form a
functional ribosome.
Some ribosomes are attached to the outer surface of the
nuclear membrane and to an extensively folded membrane called
the endoplasmic reticulum. These ribosomes synthesize proteins
destined for specific organelles, for insertion in the plasma mem­
brane, or for export from the cell. Other ribosomes are called free
ribosomes because they are not attached to other cytoplasmic
structures. Free ribosomes synthesize proteins used in the cytosol.
Ribosomes are also located within mitochondria, where they syn­
th_esize mitochondrial proteins. Large Small Complete
subunit subunit functional
ribosome
Endoplasmic Reticulum
Details of ribosomal subunits
Theendoplasmicreticulum(ER) (en'-do-PLAS-mik re-TIK-u-lum; -
plasmic = cytoplasm; reticulum = network) is a network of Q Where are ribosomal subunits synthesized and
folded membranes in the form of flattened sacs or tubules (Figure assembled?
3.14). The ER extends throughout the cytoplasm and is so
extensive that it constitutes more than half of the membranous
Clinical Connection
surfaces within the cytoplasm of most cells.
Cells contain two distinct forms of ER that differ in structure
Smooth ER and Increased Drug Tolerance
and function. Rough ER extends from the nuclear envelope
(membrane around the nucleus) and appears "rough" because its One of the functions of smooth ER, as noted earlier, is to detoxify
outer surface is studded with ribosomes (Figure 3.14). Proteins certain drugs. Individuals who repeatedly take such drugs, such as the
synthesized by ribosomes attached to rough ER enter the spaces sedatlve phenobarbital, develop changes in the smooth ER in their liver
within the ER for processing and sorting. These molecules cells. Pro­ longed administration of phenobarbital results in increased
(glycoproteins and phos­ pholipids) may be incorporated into tolerance to the drug; the same dose no longer produces the same
degree of sedation. With repeated exposure to the drug, the amount of
organelle membranes or the plasma membrane. Thus, rough ER
smooth ER and its en­ zymes increases to protect the cell from its toxic
is a factory for synthesizing
effects. As the amount of smooth ER increases, higher and higher
secretory proteins and membrane molecules.
dosages of the drug are needed to achieve the original effect. This
Smooth ER extends from the rough ER to form a network could result in an increased possibility of overdose and increased
of membranous tubules (Figure 3.14). As you may already drug dependence.
have
guessed, smooth ER appears "smooth" because it lacks ribosomes.
Smooth ER is where fatty acids and steroids such as estrogens and Golgi C o m p l e x After proteins are synthesized on a ribosome
testosterone are synthesized. In liver cells, enzymes of the smooth attached to rough ER, most are usually transported to anotherregion
ER also help release glucose into the bloodstream and inactivate of the cell. The first step in the transport pathway is through an
or detoxify a variety of drugs and potentially harmful substances, organelle called the Golgi complex (GOL-je). It consists of 3 to 20
including alcohol, pesticides, and carcinogens (cancer-causing cisterns (SIS­
agents). In muscle cells, calcium ions needed for muscle contrac­ terns = cavities), flattened membranous sacs with bulging edges,
t io n are stored and released from a form of smooth ER called sar­
coplasmic reticulum.

Sc· ned w'i'h C Scnn·

 52 CHAPTER 3 Cells

UB'MIHOEndoplasmic reticulum HB'Mfltj Golgl

(ER). complex.
The ER is a network of folded membrane.s that extend throughout Most proteins synthesized by ribosomes attached to rough ER
the cytoplasm and connect to the nuclear envelope. pass through the Golgi complex for processing.

Functions of Endoplasmic Reticulum Functions of Golgl Complex

1. Rough ER synthesizes glycoproteins and phospholipidsthat are 1. Modifies, sorts, packages, and transports proteinsreceived from
transferred into cellular organelles, inserted Into the plasma the rough ER.
membrane, or secreted during exocytosis. 2. Forms secretory vesicles that discharge processed proteins via
2. Smooth ER synthesizes fatty acids and steroids,such as estrogens and eiocytosis Ir.to extracellular fluid; forms membrane vesicles that ferry
testosterone; inactivates or detoxifies drugs and other potentially new molecules to the plasma membrane; forms transport vesiclesthat
harmful substances; removes the phosphate group from glucose-6- carry molecules to other organelles, such as lysosomes.
phosphate;and stores arid releases calcium ions that trigger
coritractiori in muscle cells.

Nuclear
envelope

Ribosomes

Q What types of body cells are likely to have extensive

Golgi complexes?
Rough
ER
Q How do rough ER and smooth ER differ structurally contain as many as 60 different digestive enzymes; these
and functionally? enzymes can break down a wide variety of molecules once the
lysosome fuses with vesicles formed during endocytosis. The
piled on each other like a stack of pita bread (Figure 3.15). Most lysosomal membrane contains carrier proteins that allow the final
cells have several Golgi complexes. The Golgi complex is more products of digestion, such as monosaccharides, fatty acids,
extensive in cells that secrete proteins. and amino acids, to be transported into the cytosol.
The main function of the Golgi complex is to modify and pack­ Lysosomal enzymes also help recycle worn-out structures.
age proteins. Proteins synthesized by ribosomes on rough ER A lysosome can engulf another organelle, digest it, and return
enter the Golgi complex and are modified to form glycoproteins the digested components to the cytosol for reuse. In this way, old
and lipoproteins. Then, they are sorted and packaged into orga­ nelles are continually replaced. The process by which
vesicles. Some of the processed proteins are discharged from worn-out organelles are digested is called autophogy (aw-TOF-a-
the cell by exocytosis. Certain cells of the pancreas release the je; aut o-=
hormone insu­ lin this way. Other processed proteins become part self; -phagy = eating). During autophagy, the organelle
of the plasma membrane as existing parts of the membrane are to be digested is enclosed by a membrane derived from the ER to
lost. Still other processed proteins become incorporated into create
organelles called lysosomes. a vesicle that then fuses with a lysosome. In this way, a human liv­
er cell, for example, recycles about half its contents every week.
Lysosomes Lysosomes (Li-so-soms; lyso- == dissolving; - Lysosomal enzymes may also destroy the entire cell, a process
sames=
bodies) are membrane-enclosed vesicles (see Figure 3.1) that may known as autolysis (aw-TOL-i-sis). Autolysis occurs in some patho­
logical conditions and also is responsible for the tissue deteriora­
tion that occurs just after death.
 l.A Cyu,pusm 53

Clinical Connection Mitochondria Because they are the site of most ATP produc­
tion, the "powerhous.es- of a cell are Its mitochondria (mi-to-KON­
Tay-Sachs Disease dre-a; miro- = thread;-chondrio = granules; singular is
mitochondrion).
.Some
st
disorde
rs are caused by faulty or absent lysosomal enzymes. For A cell may have as few as one hundred or as many as several
in ance, Tay-Sachs disease (TA SAKS) which most often affects
thousand mitochondria, depending on how active the c.ell ls.
children OfAshkenaz1"(eastern European Jewish)' descent Is an Inherited Foreumple, active cells such as those found in muscles, the liver,
condition characterized by the absence of a single lysoso al enzyme. and kldneys use ATP at
ormally breaks down a membrane glycolipld called ganglioslde G.,,
This enzyme
that
15 a high rate and have largenumbers of mitochondria. A
especially prevalent in nerve cells. As the excess gangliosldeG.,,
mitochondrion consists of two membranes, each of which is similar
accumu­ lat s, because it is not broken down, the nerve cellsfunction
in structure to the
lessefficiently. Children with Tay-Sachs disease typically experience
seizures and muscle rigidity. They gradually become blind, demented, plasma membrane (Figure 3.16). The out•rmltochondrlol m•mbron•
and uncoordinated and usually die before the age of five. Tests can is smooth, but the lnn•r mitochondrial m•mbron• is arranged in
now reveal whether an adult is a carrier of the defective gene. a series of folds called mitochondrial crlsta• (KRIS-te; singular is
crlsta = ridge). The large central fluid-filled cavity of a
mitochondrion, enclosed by the Inner membrane and cristae, is
Peroxisomes Another group of organelles similar in structure the mitochondrial matrix. The elaborate folds of the cristae provide
to lysosomes, but smaller, are called peroxisomes (pe-ROKS-i-s6ms; an enormous surface
perox i- = peroxide; see Figure 3.1). Peroxisomes contain several area for a series of chemical reactions that provide most of a cell's
ATP. Enzymes that catalyze these reactions are located in the matrix
oxidases, which are enzymes that can oxidize (remove hydrogen atoms
and on
from) various organic substances. For example, amino acids and fatty
acids are oxidized in peroxisomes as part of normal metabolism. In U t acristae.
the ' M i & i JMitochondria also contain a small number of genes and a
Mltochondrlon.
few ribosomes, enabling them to synthesize some proteins.
addition, enzymes in peroxisomes oxidize toxic substances. Withinmitochondria, chemical reactions
Thus, peroxisomes are very abundant in the liver, where generate most of a cell's ATP.
detoxification of
alcohol and other damaging substances takes place. A byproduct Function of Mitochondria
of the oxidation reactions is hydrogen peroxide (H2O2), a potentially Generate ATP through reactions of aerobic cellular
toxic compound, and associated free radicals, such as superoxide. respiration.
However, peroxisomes also contain an enzyme called catalase that
decomposes the H2O2. Because the generation and degradation of
H 2 0 2 occurs within the same organelle, peroxisomes protect other
parts of the cell from the toxic effects of H2O2. Peroxisomes also
have
enzymes that destroy superoxide.
Proteasomes Although lysosomes degrade proteins delivered
to them in vesicles, proteins in the cytosol also require disposal at
certain times in the life of a cell.Continuous destruction of unneeded,
damaged, or faulty proteins is the function of tiny barrel-shaped
structures called proteasomes (PR6-te-a-s6ms = protein bodies) see
Figure 3.1. A typical body cell contains many thousands of Mitochondrial
matrix
proteasomes, in both the cytosol and the nucleus. Proteasomes were
so named because they contain myriad proteases, enzymes that cut
proteins into small peptides. Once the enzymes of a proteasome have
chopped up a protein into smaller chunks, other enzymes then break
down the peptides into amino acids, which can be recycled into new
proteins.

Clinical Connection

Proteasomes and Disease

Some diseases could result from failure of proteasomes to degrade ab­
normal proteins. For example, clumps of misfolded proteins accumulate Enzymes
in brain cells of people with Parkinson's disease and Alzheimer's
disease. Discovering why the proteasomes fail to clear these abnormal
proteins is a goal of ongoing research. Q How do the cristae of a mitochondrion contribute to
its ATP-producing function?

·ned wh Ca
54 CHAPTER 3 Cells

Checkpoint
ht3'j;jj9ij Nucleus,

The nucleus contains most of a cell's genes, which are located on

7. What does cytoplasm have that cytoso\ does not?
chromosomes.
8. What is an organelle?
9, Describe the structure and function of ribosomes, the Golgl Functions of the Nucleus
complex, and mitochondria. 1, Controls cellular structure.
2. DirectScellular activities.

0 Nucleus
3. Produces ribosomes in nucleoli.

OBJECTIVE

, Describe the structure and functions of the nucleus.

The nucleus is a spherical or oval structure that usually is the most

prominent feature of a cell (Figure 3.17). Most body cells have a single
nucleus, although some, such as mature red blood cells, have none. In
contrast, skeletal muscle cells and a few other types of cells have sev­
eral nuclei. A double membrane called the nuclear envelope separates
the nucleus from the cytoplasm. Both layers of the nuclear envelope
are lipid bilayers similar to the plasma membrane. The outer mem­
brane of the nudear envelope is continuous with the rough endoplas­
mic reticulum and resembles it in structure. Many openings called !5!!!!"!!!!!- -::f--Rough
endoplasmic
nuclear pores pierce the nudear envelope. Nuclear pores control the
reticulum
movement of substances between the nucleus and the cytoplasm.
Inside the nucleus are one or more spherical bodies called nucleoli
Details or the nucleus
(noo'-KLE-6-li; singular is nucleolus). These clusters of protein, DNA,
and RNA are the sites of assembly of ribosomes, which exit the nucleus Q What are the functions of nuclear
genes?

j Clinical Connection
Stem Cell Research themselves, which distinguishes them from non-stem ce\ls.
Examples include stem keratinocytes in the skin or satellite cells
Stem cells are unspecialized cells that have in the muscles. Pluripotent stem cells currently used in research
the ability to divide for indefinite periods are derived from (1) the five-day old embryos that were des­
and give rise to specialized cells. A stem cell tined to be used for infertility treatments but were not needed
with the potential to form an entireorganism and from (2) nonliving fetuses terminated during the first three
is known as a totipotent stem cell (t6-TIP-o­ months of pregnancy.
tent; totus- = whole; -potentia = power). The Scientists are also investigating the potential clinica I
cells of a zygote (fertilized ovum) and a four­ applications of odultoligopotentond unipotent stem cells-stem
day old embryo are totipotent stem cells. cells that remain in the body throughout adulthood. Recent
Five-day old embryo cells, by contrast, can experiments suggest that the ovaries of adult mice contain
give rise to many (but not all) different types stem cells that can develop into new ova (eggs). If these same
of cells. Such stem cells are called pluripotent types of stem cells are found in the ovaries of adult women,
stem cells (ploo-RIP-6-tent; p/ur- = several). scientists could potentially harvest some of them from a
Later, pluripotent stem cells can undergo fur­ woman about to undergo a sterilizing medical treatment (such
ther specialization into cells that have a spe­ aschemotherapy),store them, and then return the stem cells to
cific function and give rise to a closely related her ovaries after the medical treatment is completed in order to
family of cells. The primary role of these cells, restore fertility. Studies have also suggested that stem cells in
called multipatent stem cells (mul-TIP-6-tent), STEVE GSCHMEISSNER/SCIENCE human adult red bone marrow have the ability to differentiate
is to maintain or repair a tissue. 0/igopotent PHOTO LIBRARY/Getty Images into cells of the liver, kidney, heart, lung, skeletal muscle, skin,
stem cells (6-LIG-op-6-tent) give rise to a few Human embrvonic stem cell and organs of the gastrointestinal tract. In theory, adult stem
different cell types, such as myeloid and lym- cells from red bone marrow could be harvested from a patient
phoi d stem cells that develop into the different types of blood cells. and then used to repair other tissues and organs in that patient's
Unipotent stem cells (u-NIP-6-tent) produce only one cell type, but can renew body without having to use stem cells from embryos.

Sc· n
 J.S Nucl u• 55

th
rough nuclear pores and participate in protein synthesis in the cyto- chromutln (KRO-ma-tin) (Figure 3.17). The total genetic information
1
andasm.P . Cells th at synthesize• large amounts of carried in a cell or organism iscalled its genome (JE-n6m).
protein,• such as muscle The main parts of a cell and their functions are summarized in
liver cells, have prominent nucleoli.
Table 3.2.
ge Also wit• h'm the nucleus are most of the cell's hereditary units' called
t' ne s , Which control cellular structure and direct most cellular activi-
ies. The nuclear genes are arranged along chromosomes (chromo- = Checkpoint
colored) (see F1' gure 3.21). Human somati.c (body) cells have 46 chromo-
somes, 23 inherited from each parent. In a cell that is not dividing, the 10. Why is the nucleus so important In the life of a
46 "111
chromosomes appear as a diffuse, granular mass, which is called

TABLE 3,2 Cell Parts and Their

Functions
Part Description Functlon(s)
PLASMA MEMBRANE Composed of a lipid bilayer consisting of phospholipids, Protects cellular contents; makes contact with other cells;
cholesterol, and glycolfplds with various proteins inserted; contains channels, transporters, receptors, enzymes, and cell
surrounds cytoplasm identity markers; mediates the entry and e it of substances
CYTOPLASM Cellular contents between the plasma membrane and Site of all intracellular activities except those occurring in the
nucleus, including cytosol and organelles nucleus
Cytosol Composed of water, solutes, suspended particles, lipid Liquid in which many of the cell's chemical reactions occur
droplets, and glycogen granules
The cytoskeleton is a network in the cytoplasm Maintains shape and general organization of cellular contents;
composed responsible for cell movements
of three protein filaments: microfilaments, intermediate
Organelles filaments, and microtubules Each organelle has one or more specific functions
Specialized cellular structures with characteristic shapes
Centrosome and specific functions Pericentriolar matrix is organizing center for microtubules
Paired centrioles plus pericentriolar matrix and mitotic spindle
Cilia and flagella Motile cell surface projections with inner core of Cilia move fluids over a cell's surface; a flagellum moves an
microtubules entire cell
Ribosome Composed of two subunits containing ribosomal Protein synthesis
RNA
Endoplasmic and proteins; may be free in cytosol or attached Rough ER is the site of synthesis of glycoproteins and
reticulum (ER) to rough ER phospholipids; smooth ER is the site of fatty acid and steroid
Membranous network of folded membranes; rough ER is synthesis; smooth ER also releases glucose into the bloodstream,
studded with ribosomes and is attached to the nuclear inactivates or detoxifies drugs and potentially harmful substances,
membrane; smooth ER lacks ribosomes and stores and releases calcium ions for muscle contraction
Golgi complex A stack of 3-20 flattened membranous sacs called cisterns Accepts proteins from rough ER; forms glycoproteins and
lipoproteins; stores, packages, and exports proteins
Lysosome Vesicle formed from Golgi complex; contains digestive Fuses with and digests contents of vesicles; digests worn-out
enzymes organelles (autophagy), entire cells (autolysis), and extracellular
materials
Peroxisome Vesicle containing oxidative enzymes Detoxifies harmful substances, such as hydrogen peroxide and
associated free radicals
Proteasome Tiny barrel-shaped structure that contains proteases, Degrades unneeded, damaged, or faulty proteins by cutting
enzymes that cut proteins them into small peptides
Mitochondrion Consists of outer and inner membranes, cristae, and Site of reactions that produce most of a cell's ATP
matrix
NUCLEUS Consists of nuclear envelope with pores, nucleoli, and Contains genes, which control cellular structure and direct most
chromatin (or chromosomes) cellular activities

Flagellum --- / Cilium

Proteasome .
Intermediate - -'·
filament --- Cytoplasm
Centrosome - - L- Plasma
:;:;t::t:;::-:
1 membrane
Lysosome -~--......;t.:.....J
Ribosome on rough
Golgi complex
Smooth EA
Peroxisome ----= e'9 ;..,_: ER
Mitochondrion

Microfilament

·ned wh Ca
 56 CHAPTER 3
Cells
Information is stored in DNA in four types of nucleotides, the
Gene Action: repeating units of nucleic acids (see Figure 2.15). Each sequence
of three DNA nucleotides is transcribed as a complementary
Protein Synthesis (corresponding) sequence of three RNA nucleotides. Such a se­
quence of three successive DNA nucleotides is called a base
triplet. The three successive RNA nucleotides are called a
OBJECTIVE
codon. When translated, a given codon specifies a particular
amino acid.
• Outline the sequence of events involved in
protein synthesis.

Transcription
Although cells synthesize many chemicals to maintain homeostasis,
much of the cellular machinery is devoted to protein production. During transcription, which occurs in the nucleus, the genetic
Cells constantly synthesize large numbers of diverse proteins. The information in DNA base triplets is copied into a complementary
proteins in turn determine the physical and chemical sequence of codons in a strand of RNA. Transcription of DNA is
characteristics of cells and, on a larger scale, of organisms. catalyzed by the enzyme RNA polymerase, which must be
instructed where to start the transcription process and where to
The DNA contained in genes provides the instructions for
end it. The segment of DNA where RNA polymerase attaches to it
mak­ ing proteins. To synthesize a protein, the information
contained in a specific region of DNA is first transcribed (copied) is a special sequence of nucleotides called a promoter, located
to produce a specific molecule of RNA {ribonucleic acid). The RNA near the beginning of a gene {Figure 3.19a). Three kinds of RNA
then attaches to a ribosome, where the information contained are made from DNA:
in the RNA is translated into a corresponding specific sequence
of amino acids to form a new protein molecule (Figure 3.18). • Messenger RNA (mRNA) directs synthesis of a protein.
• Ribosomal RNA (rRNA) joins with ribosomal proteins to make ribo­
somes.
• Transfer RNA (tRNA) binds to an amino acid and holds it in
OB'Miih Overview of transcription and place on a ribosome until it is incorporated into a protein during
translation. translation. Each of the more than 20 different types of tRNA
Transcription occurs in the nucleus; translation takes place in binds to only one of the 20 different amino acids.
the cytoplasm.
During transcription, nucleotides pair in a complementary man­
ner: The nitrogenous base cytosine (C) in DNA dictates the comple­
mentary nitrogenous base guanine (G) in the new RNA strand, a G
- - - < " " - - - - - - Nucleus in DNA dictates a C in RNA, a thymine (T) in DNA dictates an
adenine {A) in RNA, and an A in DNA dictates a uracil (U) in RNA
------!--1-1...-DNA {Figure 3.19b). As an example, if a segment of DNA had the base
Nuclear pore sequence ATGCAT, the newly transcribed RNA strand would have the
complementary base sequence UACGUA.
RNA
Transcription of DNA ends at another special nucleotide se­
lLPlasma quence on DNA called a terminator, which specifies the end of the
membrane gene (see Figure 3.19a). Upon reaching the terminator, RNA poly­
merase detaches from the transcribed RNA molecule and the DNA
Cytoplasm strand. Once synthesized, mRNA, rRNA {in ribosomes), and tRNA
leave the nucleus of the cell by passing through a nuclear pore.
In the cytoplasm, they participate in the next step in protein
synthe­ sis, translation.
RNA
Ribosom
e
Protein Translation
In the process of translation, the nucleotide sequence in an
Why are proteins important in the life of a mRNA molecule specifies the amino acid sequence of a protein.
cell? Ribosomes
 1_, GeM Action: Prot ln SynthHIS 57

m,,..Sfti l
t Transcription In the nucleus. in the cytoplasm carry out translation. The small subunit of a ribo­
some has a binding site for mRNA; the larger subunit has three
. Duringtranscription,the geneticinformationIn DNA Iscopied to
bind· ing sites for tRNA molecules: a P site, A site, and E site
(Figure 3.20). The P (peptldy() site binds the tRNAcarrying the
RNA.
growingpolypeptide chain. The A (amlnoacyl) site binds the tRNA
Gene
DNA carrying the next amino acid to be added to the growing
polypeptide. The E (exit) site binds tRNA just before it isreleased
from the ribosome. Translation occurs in the following way (Figure
(a) 03.20):An mRNA molecule binds to the small ribosomal subunit at the
Overview mRNA binding site. A special tRNA, called initiator tRNA, binds to
the start codon (AUG) on mRNA, where translation begins.
8 Next, the large ribosomal subunit attaches to the small

ribosomal
subunit-mRNA complex, creating a functional ribosome. The
initiator tRNA fits into position on the ribosome. One end of a
DNA strand tRNA carries a specific amino acid, and the opposite and
being consists of a triplet of nucleotidescalled an ant/codon. By
transcribed pairing between complementary nitrogenous bases, the tRNA
anticodon attaches to the mRNA codon. For example, if the
mRNA codon is AUG, then a tRNA with the anticodon UAC
would attach to it.
8 The anticodon of another tRNA with its attached amino acid
DNA to mRNA ba■e pairing
during transcription pairs

Newly synthesized
pre-mRNA
,.
D N A
0C)
with the second mRNA codon at the A site of the ribosome.
Following the formation of the peptide bond, the resulting two­
A component of the large ribosomal subunit catalyzes the
peptide protein becomes attached to the tRNA at the A site.
T
G formation
() After peptide bond formation, the ribosome shifts the mRNA
C of a peptide
strand by onebond between
codon. the amino
The tRNA in the acid carried
P site entersbythe
theEinitiator
site and
C tRNA and the amino acid carried by the tRNA at the A site.
is subsequently released from the ribosome. The tRNA in the A
G site bearing the two-peptide protein shifts into the P site,
allowing another tRNA with its amino acid to bind to a newly
exposed
codon at the A site. Steps 8 through () occur repeatedly, and
the protein lengthens progressively.
O Protein synthesis ends when the ribosome reaches a stop codon
at the A site, which causes the completed protein to detach
from the final tRNA. In addition, tRNA vacates the P site and the
ribosome splits into its large and small subunits.

Protein synthesis progresses at a rate of about 15 amino acids

per second. As the ribosome moves along the mRNA and before it
com­ pletes synthesis of the whole protein, another ribosome may
attach behind it and begin translation of the same mRNA strand. In
Key: this way, several ribosomes may be attached to the same mRNA.
= Such a group of ribosomes is called a polyribosome. The
Adenine simultaneous movement of several ribosomes along the same
= mRNA strand permits a large amount of protein to be produced from
Guanine each mRNA.
=Thymine Checkpoint
=
Cytosine (b) Details 11. Define proteinsynthesis.
=Uracil 12, Distinguish between transcription and
Q What enzyme catalyzes transcription of translation.
DNA?

Sea C Scanner
58 CHAPTER 3 Cells

Hid'MJff,j Protein elongation and termination of protein synthesis during translatlon In

the cytoplasm,

During proteinsynthesisthe ribosomal subunits join,but they separate when the process
iscomplete.
P Sile
Aminoacid

P - - - - Large
subunit tRNA-
lil! !,,,,...,.C -
lnilialortRNA
y

( 0Large and small ribosomal

subunils join to form a
functional ribosome and
Amino acid ._. lnilialor tRNA
. fits Into P site. E) with
Anticodon of incoming
next mRNA codon tRNA pairs
at A sile. \
- - - - lniliator !RNA
(methionine)
J,

Start codon
0 Initiator tRNA attaches to
a start codon.

0 Amino acid on tRNA al P site

forms a peptide bond with
amino acid at A site.

Stop codon
GProtein synthesis stops when
the ribosome reaches slop
codon on mANA.

ORibosome shifts by onecodon: tRNA .- C, The two-peplideprotein

previously at P site enlersE siteand creeled from the formation
is released from ribosome; tRNA "l"ifF--- thof e peptide bondbecomes
previously at A sile Is now at P site. attached lo !RNA at A site.

Key:
= Adenine m p l e t e protein

--= G u a n i n e t\ tRNA
= Cytosine

PL =Uracil
Summary of movement of ribosome along mRNA
Q What is the function of a stop
codon?

·
-ned w I:Ca
 J.7 Sorn,tic C.tl Division 59

B Nuclear Division: Mitosis

Somatic Cell Division mitos thread), the duplicated chromosomes become exa tly seg
During mittuls (mi-TO-sis;

egat_ed, one set into each of tv-10 separate nuclei. For

OBJECTIVE convenience, b1olog1sts divide the process into four stages:
prophase, metaphase, anaphase, and telophase. However, mitosis
is a continuous process, with one stage merging imperceptibly
• Discuss the stages, events, and significance of somatic cell
division. into the next.
PAOPHASE During early prophase (PRO-faz), the chromatin fibers cond
visible under the light microscope (Figure 3.21b). The condensation proc
strands as they move during mitosis. Recall that DNA replication took p
As body cells become damaged, diseased, or worn out, they are chromosome consists of a pair of identical, double-stranded chromatids. A
replaced by cell division, the process whereby cells reproduce
a cen­tromere (SEN-tro-mer),holds the chromatid pair together.
themselves. The two types of cell division are reproductive cell divi­
Later in prophase, the pericentriolar material of the two cen­
sion and somatic cell division. Reproductive cell division or meio­
microtubules called the mitotic spindle (Figure 3.21b). Lengthening of
sis is the process that produces gametes-sperm and oocytes-the between centrosomes pushes the centrosomes to opposite poles (end
cells needed to form the next generation of sexually reproducing
pole to pole. Then the nucleolus and nuclear envelope break down.
organisms. Meiosis is described in Chapter 23; here we will focus
on somatic cell division.
METAPHASE During metaphase (MET-a-faz), the centromeres of th
All body cells, except the gametes, are called somatic cells (so­
microtubules of the mitotic spindle at the exact center of the mi
MAT-ik; soma= body). In somatic cell division, a cell divides into alignment of the centromeres is called the metaphase plate.
two identical cells. An important part of somatic cell division is
replication (duplication) of the DNA sequences that make up genes ANAPHASE During anaphase (AN-a-faz) the centromeres split, sep­ arati
and chromo­ which move to opposite poles of the cell (Figure 3.21d). Once separated
somes so that the same genetic material can be passed on to the the chromosomes are pulled by the microtubules of the mitotic spin
newly formed cells. After somatic cell division, each newly formed because the centromeres lead the way and seem to drag the trailing arm
cell has the same number of chromosomes as the original cell.
Somatic cell division replaces dead or injured cells and adds new TELOPHASE The final stage of mitosis,telophase (TEL-6-fa-z), begins
ones for tis­ sue growth. For example, skin cells are continually 3.21e). The identical sets of chromosomes, now at opposite poles of
replaced by somatic cell divisions. chromatin form. A new nuclear envelope forms around each chroma
The cell cycle is the name for the sequence of changes that a mitotic spindle breaks up.
cell undergoes from the time it forms until it duplicates its contents
and divides into two cells. In somatic cells, the cell cycle consists of
two major periods: interphase, when a cell is not dividing, and the Cytoplasmic Division: Cytokinesis Division of a
mitotic phase, when a cell is dividing. cell's cytoplasm and organelles is called cytokinesis (si'-to-ki­
usually begins late in anaphase with formation of a cleavage furrow, a
of the plasma membrane, that extends around the center of the cell
lnterphase (Figure 3.21d,e). Microfilaments in the cleavage furrow pull the
plasma membrane progressively inward, constricting the center
During interphase the cell replicates its DNA. It also manufactures
ultimately pinching it in two. After cytokinesis there are two new an
additional organelles and cytosolic components, such as cen­
cytoplasm and organelles and identical sets of chromosomes. Whe
trosomes, in anticipation of cell division. lnterphase is a state of
(Figure 3.21f).
high metabolic activity, and during this time the cell does most of its
growing.
A microscopic view of a cell during interphase shows a clearly
de­ fined nuclear envelope, a nucleolus, and a tangled mass of
chromatin (Figure 3.21a). Once a cell completes its replication of
DNA and other activities of interphase, the mitotic phase begins.

Mitotic Phase
The mitotic phase (mi-TOT-ik) of the cell cycle consists of mitosis, divi­
sion of the nucleus, followed by cytokinesis, division of the cytoplasm
into two cells. The events that take place during mitosis and cytokine­
sis are plainly visible under a microscope because chromatin con­
denses into chromosomes.

Sc..nner
 60 CHAPTER 3 Cells

U@lj;jjijj 1.elldivlslon: mitosis and cytoklnesls. Begin the sequence at (a) at thetop of
thefigure and read clockwise until you completethe process.

In somatic cell division,a single cell divides to produce two identical cells.

Centrosome:
0
: : : 3 - - Cenlrioles
"' ::-;,;:x-,,-..,,.
.,,j] .. Perlcenlriolar material
." 0
. Nucleolus
.
I
-' 1
L Nuclear
Chromatin
// envelope
Plasma
l,l;.:..:..-"""'L .
l""/"l'-:-:"t"""::--:0::==r:-::----,!li:;.;.;;:...:.;_"-
membrane
Cytosol
(a)
INTERPHASE

\
Centromere - - 4 ; , ! , ¥ - : l - ­
fJ,"7ur;;;;::!il\ Mitotic
Chromosome -"\';;;.:-t spindle
(microtubules
(oj twni oedchartom
(I) IDENTICALCELLS. IN , r - : : : : b - , )Fragments of
INTERPHASE atids nuclear
0 centromere) • •- envelope

"-
(b) Late
PROPHASE
Metaphaseplate -.:••t'lillli, •

'
\ ,-

PHASE
0

Centromere Chromalids Cleavage furrow

- -

Chromosome I I - \ '
; ;
Early
9 6 0 5 0 x
Chromosome (d)
ANAPHASE
puring which phase of mitosis does cytokinesis begin?

c a-·Sc·
 .
,
Clinical Connection Oid'Hf ffFI DlvtrM • h • • ■nd 1lu1ofhuman cells.
Therelatr,e differenceInsizebetwttn thesmallest and larg t
ctflsIsactualty much greater thanshownhere.
Chemotherapy
One of the distinguishing featuresof cancer cellsIsuncontrolled Th@nearly 100uillion cellsIn an average adult can bt c.lasslf,td
division. The mass of cells resulting from thisdivisioniscalled a Into about 200different ctll types.
neoplasmor tumor. One of the ways to treat cancer Isby
chemotherapy, the use of anticancer drugs.Some of thesedrugsstop
cell divisionby Inhibitingthe formationof the
mitoticspindle.Unfortunately,thesetypesof anticancerdrugs alsokill all Smooth muscle
types of rapidlydividingcells inthe body,causingsideeffects,as will be cell
described inthe Common Disorders section.

Checkpoint
13. Distinguishbetween somatic and reproductive cell
division.Why is each important?
14. What are the major events of each stage of the mitotic
phase?

Cellular Diversity
Aed blOOd cell Epithelialcel
l
OBJECTIVE Q Why are sperm the only body cells that need to have
a flagellum?
• Describe how cells differ in size and
shape.
chapters, cellular diversity also permits organization of cells into
The body of an average human adult is composed of nearly 100 tril­ more complex tissues and organs.
lion cells. Cells vary considerably in size.
The sizes of cells are measured in units called micrometers. Checkpoint
One micrometer (µm) is equal to l one-millionth of a meter, or
10-6 rn 15, How iscell shape related to function? Give
(1/25,000 of an inch).High-powered microscopes are needed to see examples.
the smallest cells of the body. The largest cell, a single oocyte, has a
diam­ eter of about 140 µm and is barely visible to the unaided eye.
A red blood cell has a diameter of 8 µm. To better visualize this, an
average hair from the top of your head is approximately 100µmin
Aging and Cells
diameter.
The shapes of cells also vary considerably (Figure 3.22). OBJECTIVE
They may be round, oval, flat, cube-shaped, column-shaped,
elongated, • Describe the cellular changes that occur with aging.
star-shaped, cylindrical, or disc-shaped. A cell's shape is related to
its function in the body. For example, a sperm cell has a long
whip­ like tail (flagellum) that it uses for locomotion. Sperm cells Aging is a normal process accompanied by a progressive alteration of
are the only male cells required to move considerable distances. the body's homeostatic adaptive responses. It produces observable
The disc shape of a red blood cell gives it a large surface area that changes in structure and function and increases vulnerability to envi­
enhances its ability to pass oxygen to other cells. The long, spindle ronmental stress and disease. The specialized branch of medicine
shape of a relaxed smooth muscle cell shortens as it contracts. that deals with the medical problems and care of elderly persons is
This change in shape allows groups of smooth muscle cells to geriatrics (jer'-e-AT-riks; ger- = old age;-iotrics = medicine). Geron•
narrow or widen the passage for blood flowing through blood tology (jer'-on-TOL-6-je) is the scientific study of the process and
vessels. In this way, they regulate blood flow through various problems associated with aging.
tissues. Recall that some cells contain microvilli, which greatly Although many millions of new cells normally are produced each
increase their surface area. Micro­ villi are common in the epithelial minute, several kinds of cells in the body-including skeletal muscle
cells that line the small intestine, where the large surface area
speeds the absorption of digested food. Nerve cells have long
extensions that permit them to conduct nerve impulses over great
distances. As you will see in the following

·ned wh Ca
 62 CHAPTER 3 Cells

cells and nerve cells-do not divide. Experiments have shown that Free radicals produce oxidative damage in lipids, proteins, or nu­
many other cell types have only a limited capability to divide. cleic acids. Some effects are wrinkled skin, stiff joints, and hardened
Normal cells grown outside the body divide only a certain number of arteries. Naturally occurring enzymes in peroxisomes and in the cyto­
times and then stop. These observations suggest that cessation of sol normally dispose of free radicals. Certain dietary substances,
mitosis is a normal, genetically programmed event. According to this such as vitamin E, vitamin C, beta carotene, zinc, and selenium, are
view, "aging genes" are part of the genetic blueprint at birth. These anti­oxidants that inhibit free radical formation.
genes have an important function in normal cells, but their activities Some theories of aging explain the process at the cellular
slow over time. They bring about aging by slowing down or halting level,
processes vital to life. while others concentrate on regulatory mechanisms operating within
Another aspect of aging involves telomeres (TE-16-merz), the entire organism. For example, the immune system may start to
specific attack the body's own cells. This autoimmune response might
DNA sequences found only at the tips of each chromosome. be caused by changes in certain plasma membrane glycoproteins
These pieces of DNA protect the tips of chromosomes from erosion and glycolipids (cell identity markers) that cause antibodies to
and from sticking to one another. However, in most normal body attach to and mark the cell for destruction. As changes in the
cells each cycle of cell division shortens the telomeres. Eventually, proteins on the plasma membrane of cells increase, the autoimmune
after many cycles of cell division, the telomeres can be response inten­sifies, producing the well-known signs of aging.
completely gone, and even some of the functional chromosomal
material may be lost. These observations suggest that erosion of Checkpoint
DNA from the tips of our chromosomes contributes greatly to the
16. Briefly outline the cellular changesinvolved
aging and death of cells. Individuals who experience high levels inaging.
of stress have significantly shorter telomere length.
Glucose, the most abundant sugar in the body, plays a role in
the aging process. It is haphazardly added to proteins inside and
...
Next, in Chapter 4, we will explore how cells associate to form
outside cells, forming irreversible cross-links between adjacent the tissues and organs that we will discuss later in the text.
protein mole­ cules. With advancing age, more cross-links form,
which contributes to the stiffening and loss of elasticity that occur
in aging tissues.

Common grows, it begins to compete with normal tissues for space and
nutrients. Eventually, the normal tissue decreases in size and dies.

Disorders Some malignant cells may detach from the initial (primary)
tumor . and invade a body cavity or enter the blood or lymph, then
Cancer circulate to and invade other body tissues, establishing secondary
Cancer is a group of diseases characterized byuncontrolled or tumors. Malignant cells resist the antitumor defenses of the body.
abnormal cell proliferation.When cells in a part of the body divide The pain associated with cancer develops when the tumor presses
without control, the excess tissue that develops is called a tumor or on nerves or blocks a passageway in an organ so that
neoplasm (NE-o­ secretions build up pressure, or as a result of dying tissue or
plazm; neo- = new). The study of tumors is called oncology (on- organs.
KOL­ o-je; onco- = swelling or mass). Tumors may be cancerous Cau ses of C a n c e r Several factors may trigger a normal
and often cell to lose control and become cancerous. One cause is environ­
fatal, or they may be harmless. A cancerous neoplasm is called a mental agents: substances in the air we breathe, the water we
malig­ nant tumor or malignancy. One property of most malignant drink, and the food we eat. A chemical agent or radiation that
tumors is their ability to undergo metastasis (me-TAS-ta-sis),the pro­ duces cancer is called a carcinogen (kar-SIN-6-jen).
spread of cancer­ Carcinogens induce mutations (mu-TA-shuns), permanent
ous cells to other parts of the body. A benign tumor is a neoplasm changes in the DNA base sequence of a gene. The World
that does not metastasize. An example is a wart. Most benign tumors Health Organization esti­ mates that carcinogens are associated
may
G row bet hsurgically
a n d Sremoved
p r e a d ifo fthis interferes with normalCells
Cancer body with 60-90% of all human cancers. Examples of carcinogens are
offunction or they
malignant tumors duplicate rapidly and continuously. Cells of hydrocarbons found in ciga­ rette tar, radon gas from the earth,
become
the body disfiguring.
that Some can be inoperable and perhaps fatal. and ultraviolet (UV) radiation in sunlight.
I hav e a high rate of cell division are more at risk for developing Intensive research efforts are now directed toward studying
cancer. cancer­ causing genes, or oncogenes (ON-ko-jenz). When
As malignant cells invade surrounding tissues, they often inappropriately activated, these genes have the ability to transform
trigger a normal cell into a cancerous cell. Most oncogenes derive from
angiogenesis (an'-je-6-JEN-e-sis), the growth of new networks of normal genes called proto-oncogenes that regulate growth and
Jblood vessels. Proteins that stimulate angiogenesis in tumors development. The proto­ oncogene undergoes some change that
are either causes it to be
• :aired tumor angiogenesis factors (TAFs). The formation of new
i l o o d vessels can occur either by overproduction of TAFs or by the
1ck of n a t u r a lly occurring angiogenesis inhibitors. As the
cancer
 Utdlcal Tumlnolo&Y and Conditions 13

expressed inappropriately or make its products in excessive cancer, the tumor begins as an area of inueased cell proliferation
amounts or at the wrong time. Some oncogenes cause excessive that results from one mutation. This growth then progresses to
production of growth factors, chemicals that stimulate cell growth. abnormal. but noncancerous, growths called adenomas. A er
Others may trig­ ger changes in a cell-surface receptor, causing it to several more mutations, a carcinoma develops. The fact that so
send signals as though it were being activated by a growth factor. many mutations are needed for a cancer to develop indicates
As a result, the growth pattern of the cell becomes abnormal. that cell grovllh is normally controlled with many sets of checks
Some cancers have a viral origin. Viruses are tiny packages of and balances.
nucleic acids, either RNA or DNA, that can reproduce only while inside
Treatment of Cancer Many cancers are removed
the cells they infect. Some viruses, termed oncogenlc viruses, cause surgically. However, when cancer is widely distributed throughout
cancer by stimulating abnormal proliferation of cells; for instance, the body or eKists in organs such as the brain whose functioning
the human popillomavirus(HPV) causes virtually all cervical cancers in would be greatly harmed by surgery, chemotherapy and radiation
women. The therapy may be used instead. Sometimes surgery, chemotherapy,
virus produces a protein that causes proteasomes to destroy a and radiation therapy are used in combination. Chemotherapy
protein that normally suppresses unregulated cell division. In the involves administering
absence of this suppressor protein, cells proliferate without control. drugs that cause the death of cancerous cells. Radiation therapy
Recent studies suggest that certain cancers may be linked to breaks chromosomes, thus blocking cell division. Because cancerous
a cell having abnormal numbers of chromosomes. As a result, the cells divide rapidly, they are more vulnerable to the destructive
cell could potentially have extra copies of oncogenes or too few effects of chemotherapy and radiation therapy than are normal cells.
copies of tumor-suppressor genes, which in either case could lead Unfortunately for the patients, hair follicle cells, red bone m,mow
to un­ controlled cell proliferation. There is also some evidence cells, and cells lining the gastrointestinal tract also are rapidly
suggest­ ing that cancer may be caused by normal stem cells that dividing. Hence, the side effects of chemotherapy and radiation
develop therapy include hair loss due to death of hair follicle cells, vomiting
into cancerous stem cells capable of forming malignant and nausea due to death of cells lining the stomach and intestines,
tumors.
and susceptibility to infection due to slowed production of white
Later in the book, we will discuss the process of inflammation,
blood cells in red bone marrow.
which is a defensive response to tissue damage. It appears that inflam­
Treating cancer is difficult because it is not a single disease and
mation contributes to various steps in the development of cancer.
because all the cells in a single tumor population rarely behave in the
Some evidence suggests that chronic inflammation stimulates the
same way. Although most cancers are thought to derive from a single
proliferation of mutated cells and enhances their survival, promotes
abnormal cell, by the time a tumor reaches a clinically detectable
angiogenesis, and contributes to invasion and metastasis of cancer
size, it may contain a diverse population of abnormal cells. For
cells. There is a clear relationship between certain chronic inflam­
example, some cancerous cells metastasize readily, and others do
matory conditions and the transformation of inflamed tissue into a
not. Some are sensitive to chemotherapy drugs and some are drug-
malignant tissue. For example, chronic gastritis (inflammation of
resistant. Because of differences in drug resistance, a single
the stomach lining) and peptic ulcers may be a causative factor in
chemotherapeutic agent may destroy susceptible cells but permit
60-90% of stomach cancers. Chronic hepatitis (inflammation of the
resistant cells to proliferate.
liver) and cirrhosis of the liver are believed to be responsible for about Another potential treatment for cancer that is currently under de­
80% of liver cancers. Colorectal cancer is 10 times more likely to
velopment is virotherapy, the use of viruses to kill cancer cells. The
occur in patients with chronic inflammatory diseases of the colon,
viruses employed in this strategy are designed so that they specifically
such as ulcerative colitis and Crohn's disease. And the relationship
target cancer cells without affecting the healthy cells of the body.
between asbestosis and silicosis, two chronic lung inflammatory For
conditions,
example, proteins (such as antibodies) that specifically bind to recep­
and lung cancer has long been recognized. Chronic inflammation
Carcinogenesis: A Multistep Process
is also an underlying contributor to rheumatoid arthritis, Alzheimer's
tors found only in cancer cells are attached to viruses. Once inside the
body, the viruses bind to cancer cells and then infect them. The cancer
carcinogenesis (kar'-si-no-JEN-e-sis),
dis­ ease, depression, the process by which
schizophrenia,cardiovascular cancer
disease, and
cells are eventually killed once the viruses cause cellular lysis.
develops, is a multistep process in which as many as 10 distinct
diabetes. Researchers are also investigating the role of metastasis regula­
mutations may have to accumulate in a cell before it becomes
tory genes that control the ability of cancer cells to undergo
cancerous. In colon
metastasis. Scientists hope to develop therapeutic drugs that can
manipulate
these genes and, therefore, block metastasis of cancer cells.
Medical Terminology and Conditions

Anaplasla (an'-a-PLA-ze-a; an- = not; -p/asia = to shape) The genes become activated. Enzymes produced by these genes disrupt
loss of tissue differentiation and function that is characteristic of the cytoskeleton and nucleus; the cell shrinks and pulls away from
most malignancies. neigh­ boring cells; the DNA within the nucleus fragments; and the
Apoptosls (ap-6-TO-sis; a falling off, like dead leaves from a tree) cytoplasm shrinks, although the plasma membrane remains intact.
An orderly, genetically programmed cell death in which "cell- Phagocytes in the vicinity then ingest the dying cell. Apoptosis
suicide" removes unneeded

ned w'ilh