Chapt 26

Biotechnology
Chapter Concepts
26.1 Cloning of a Gene
Using recombinant DNA technology, bacteria
and viruses can be genetically altered to clone a
gene. 530
A genomic library contains bacteria or viruses
that carry fragments of the DNA of a particular
organism. 532
The polymerase chain reaction (PCR) makes
multiple copies of DNA segments. Analysis of
the DNA usually follows. 532
26.2 Biotechnology Products
Bacteria, agricultural plants, and farm animals
have been genetically engineered to produce
commercially available products. 534
Agricultural plants and farm animals have
been genetically engineered to improve their
yield. 535
Farm animals have been genetically engineered
to serve as a source of organs for human
transplant patients. 536
It is now possible to clone animals, and
cloning is used to produce multiple copies of
farm animals that have been genetically
engineered. 536
26.3 The Human Genome Project
The Human Genome Project has two goals: to
genetically map each chromosome and to
sequence the DNA bases of each chromosome.
538
26.4 Gene Therapy
Gene therapy is now being used to replace
defective genes with healthy genes and to help
cure various human ills. 539
DNAcan be extracted from a cell and manipulated using standard
laboratory techniques. This discovery has led to the commercial
availability of gene products and gene therapy in humans.
529
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26.1 Cloning of a Gene M
The cloning of a gene produces many identical copies. Re-
combinant DNAtechnology is used when a very large quan-
tity of the gene is required. The use of the polymerase chain
reaction (PCR) creates a lesser number of copies within a
laboratory test tube.
Recombinant DNA Technology
Recombinant DNA(rDNA) contains DNAfrom two differ-
ent sources. To make rDNA, a technician often begins by se-
lecting a vector, the means by which recombinant DNA is
introduced into a host cell. One common type of vector is a
plasmid. Plasmids are small accessory rings of DNA. The
ring is not part of the bacterial chromosome and can be repli-
cated independently. Plasmids were discovered by investi-
gators studying the sex life of the intestinal bacterium
Escherichia coli.
Two enzymes are needed to introduce foreign DNAinto
vector DNA(Fig. 26.1). The rst enzyme, called a restriction
enzyme, cleaves plasmid DNA, and the second, called DNA
ligase, seals foreign DNAinto the opening created by the re-
striction enzyme.
Restriction enzymes occur naturally in bacteria, where
they stop viral reproduction by cutting up viral DNA. They
are called restriction enzymes because they restrict the
growth of viruses. Hundreds of different restriction
530 Part 5 Continuance of the Species 26-2
T
he experiment was Freds last hope. The antiviral
drugs had done little to stop the AIDS virus from rav-
aging his immune system, so Freds physicians de-
cided to try to build him HIV-proof immune cells. After
removing some of Freds bone marrow, the source of the
bodys immune cells, they grew these cells in the laboratory
and added to them several genes they hoped would make
the cells less susceptible to infection by the AIDS virus. They
then transplanted the genetically altered cells back into
Fred, hoping that they would multiply and reconstitute his
weakened immune system with one that HIV could no longer
defeat.
Since birth Mary had been plagued with cystic brosis,
which causes excess mucus in the lungs and digestive tract.
Just a few years before, scientists had isolated the gene
whose mutant form is responsible for the torturous effects of
the disease. A modied virus that normally causes a cold had
been genetically engineered to carry the normal gene deep
into the cells that line the lungs. A single therapeutic dose
would place the modied virus in the lungs of the little girl now
racked with cough. Maybe someday Mary would be able to
run and play just like all the other children her age. These are
just two examples of gene therapy in humans, a eld that is
burgeoning with new ideas and treatments every day.
Genetic engineering is the use of technology to alter
the genome of viruses, bacteria, and other cells for medical
or industrial purposes. Biotechnology includes genetic engi-
neering and other techniques that make use of natural bio-
logical systems to produce a product or to achieve an end
desired by human beings. Genetic engineering has the ca-
pability of altering the genotype of unicellular organisms and
the genotype of plants and animals, including ourselves.
bacterium
restriction
enzyme
cleaves DNA
DNA ligase seals human gene and plasmid
recombinant DNA
cloning
cloned insulin gene insulin
host cell takes up recombined plasmid
human cell
insulin gene
plasmid
human
DNA
Figure 26.1 Cloning of a gene.
Human DNA and plasmid DNA are cleaved by the same type of
restriction enzyme and spliced together by the enzyme DNA ligase.
Gene cloning is achieved when a host cell takes up the recombined
plasmid and the plasmid reproduces. Multiple copies of the gene are
now available to an investigator. If the insulin gene functions normally
as expected, the product (insulin) may also be retrieved.
enzymes have been isolated and puried. Each one cuts
DNAat a specic cleavage site. For example, the restriction
enzyme called EcoRI always cuts double-stranded DNA
when it has this sequence of bases at the cleavage site:
Notice there is now a gap into which a piece of foreign DNA
can be placed if it ends in bases complementary to those ex-
posed by the restriction enzyme. To assure this, it is only
necessary to cleave the foreign DNA with the same type of
restriction enzyme. The single-stranded but complemen-
tary ends of the two DNA molecules are called sticky
A
C T
A A
G
T T
A
T
A
C
G
restriction
enzyme
A
C T
A A
G
T T
A
T
A
C
G
T
C
G
C
G
T
C
G
C
G
sticky ends
DNA
duplex
ends because they can bind by complementary base pair-
ing. They therefore facilitate the insertion of foreign DNA
into vector DNA.
The second enzyme needed for preparation of rDNA,
DNA ligase, is a cellular enzyme that seals any breaks in a
DNA molecule. Genetic engineers use this enzyme to seal
the foreign piece of DNA into the vector. DNA splicing is
now complete; an rDNAmolecule has been prepared.
Plasmid Vector Compared to Viral Vector
A clone can be a large number of molecules (i.e., cloned
genes) or cells (i.e., cloned bacteria) or organisms that are
identical to an original specimen. Figure 26.2 compares the
use of a plasmid and a virus to clone a gene.
Bacterial cells take up recombined plasmids, especially
if they are treated with calcium chloride to make them more
permeable. Thereafter, as the host cell reproduces, a bacter-
ial clone forms and each new cell contains at least one plas-
mid. Therefore, each of the bacteria contains the gene of
interest which hopefully is expressing itself and producing a
product. The investigator can recover either the cloned gene,
or the protein product from this bacterial clone (see also
Fig. 26.1).
Chapter 26 Biotechnology 531 26-3
bacterium
plasmid DNA
foreign gene
host cell
recombined
plasmid
recombinant DNA
Removal of Vector Recombinant DNA Taken up by Host Cloning of Gene
a.
host cell
foreign gene
recombinant
DNA
recombinant DNA
b.
viral DNA
virus
Figure 26.2 Preparation of a genomic library.
Each bacterial or viral clone in a genomic library contains a segment of DNA from a foreign cell. a. A plasmid is removed from a bacterium and is
used to make recombinant DNA. After the recombined plasmid is taken up by a host cell, replication produces many copies. b. Viral DNA is
removed from a bacteriophage such as lambda and is used to make recombinant DNA. The virus containing the recombinant DNA infects a host
bacterium. Cloning is achieved when the virus reproduces and then leaves the host cell.
Viruses that infect bacteria are called bacteriophages. In
Figure 26.2b, the DNA of a bacteriophage called lambda is
being used as a vector. After lambda attaches to a host bac-
terium, recombined DNAis released from the virus and en-
ters the bacterium. Here, it will direct the reproduction of
many more viruses. Each virus in the bacteriophage clone
contains a copy of the gene being cloned.
Genomic Library
Agenome is the full set of genes of an individual. Agenomic
library is a collection of bacterial or bacteriophage clones;
each clone contains a particular segment of DNA from the
source cell. When you make a genomic library, an organ-
isms DNAis simply sliced up into pieces, and the pieces are
put into vectors (i.e., plasmids or viruses) that are taken up
by host bacteria. The entire collection of bacterial or bacte-
riophage clones that result contains all the genes of that
organism.
In order for human gene expression to occur in a bac-
terium, the gene has to be accompanied by the proper regu-
latory regions. Also, the gene should not contain introns
because bacterial cells do not have the necessary enzymes to
process primary messenger RNA (mRNA). Its possible to
make a human gene that lacks introns, however. The en-
zyme called reverse transcriptase can be used to make a
DNA copy of all the mature mRNA molecules from a cell.
This DNA molecule, called complementary DNA (cDNA),
does not contain introns. Notice that a genomic library made
from cDNAwill contain only those genes that are being ex-
pressed in the source cell.
You can use a particular probe to search a genomic library
for a certain gene. A probe is a single-stranded nucleotide
sequence that will hybridize (pair) with a certain piece of
DNA. Location of the probe is possible because the probe is
either radioactive or uorescent. After the probe hybridizes
with the gene of interest, the gene can be isolated from the
fragment. Now this particular fragment can be cloned fur-
ther or even analyzed for its particular DNAsequence.
The Polymerase Chain Reaction
The polymerase chain reaction (PCR) can create millions of
copies of a single gene or any specic piece of DNAin a test
tube. PCR is very specicthe targeted DNAsequence can
be less than one part in a million of the total DNA sample!
This means that a single gene, or smaller piece of DNA,
among all the human genes can be amplied (copied) using
PCR.
PCR takes its name from DNApolymerase, the enzyme
that carries out DNA replication in a cell. It is considered a
chain reaction because DNApolymerase will carry out repli-
cation over and over again, until there are millions of copies
of the desired DNA. PCR does not replace gene cloning,
which is still used whenever a large quantity of gene or pro-
tein product is needed.
Before carrying out PCR, primerssequences of about
20 bases that are complementary to the bases on either side
of the target DNAmust be available. The primers are
needed because DNApolymerase does not start the replica-
tion process; it only continues or extends the process. After
the primers bind by complementary base pairing to the
DNA strand, DNA polymerase copies the target DNA
(Fig. 26.3).
PCR has been in use for several years, and now almost
every laboratory has automated PCR machines to carry out the
procedure. Automation became possible after a temperature-
insensitive (thermostable) DNA polymerase was extracted
from the bacterium Thermus aquaticus, which lives in hot
springs. This enzyme can withstand the high temperature
used to separate double-stranded DNA; therefore, replica-
tion need not be interrupted by the need to add more
enzyme.
5 3
5 3
3 5
3 5
5 3
5
5
3
3
3 5
3 5
3 5
heat for 1 minute
to denature (separate)
cool for 2 minutes
and add primers
add DNA polymerase
and wait 1.5 minutes
primer
primer
repeat cycle
Figure 26.3 Polymerase chain reaction (PCR).
PCR is performed in laboratory test tubes. Primers (red), which are
DNA sequences complementary to the 3 end of the targeted DNA,
are necessary for DNA polymerase to make a copy of the DNA
strand.
Analyzing DNA
The entire genome of an individual can be subjected to DNA
ngerprinting, a process described in Figure 26.4. The
genome is treated with restriction enzymes, which results in
a unique collection of different-sized fragments. Therefore,
restriction fragment length polymorphisms (RFLPs) exist
between individuals. During a process called gel elec-
trophoresis, the fragments can be separated according to
their lengths, and the result is a number of bands that are so
close together they appear as a smear. However, the use of
probes for genetic markers produces a distinctive pattern
that can be recorded on X-ray lm.
The DNA from a single sperm is enough to identify a
suspected rapist. Since DNA is inherited, its ngerprint re-
sembles that of ones parents. DNA ngerprinting success-
fully identied the remains of a teenager who had been
murdered eight years before because the skeletal DNA was
similar to that of the parents DNA. DNAngerprinting has
also been helpful to evolutionists. For example, it was used
Figure 26.4 DNA ngerprinting.
DNA samples I and II are from the same individual. DNA sample III is from a different individual. Notice, therefore, that the restriction enzyme cuts
are different for sample III. Gel electrophoresis separates the DNA fragments according to their length because shorter fragments migrate farther
in an electrical eld than do longer fragments. The fragments are denatured (separated) and transferred to a membrane where a radioactive probe
can be applied. The resulting pattern (the DNA ngerprint) can then be detected by autoradiography. In a theoretical rape case, for example,
sample I could be from the suspects white blood cells, sample II could be from sperm in the victims vagina, and sample III could be from the
victims white blood cells.
to determine that the quagga, an extinct zebralike animal,
was a zebra rather than a horse. The only remains of the
quagga consisted of dried skin.
Following PCR, DNA segments, as opposed to the full
genome, can also be cut by restriction enzymes and sub-
jected to gel electrophoresis. Aprobe is not needed because
the restriction fragments will appear as distinctive bands.
PCR amplication and analysis can be used to diagnose vi-
ral infections, genetic disorders, and cancer. When the am-
plied DNA matches that of a virus, mutated gene, or
oncogene, then we know that a viral infection, genetic disor-
der, or cancer is present.
Sometimes DNAis sequenced following PCR. Sequenc-
ing mitochondrial DNA segments helped determine the
evolutionary history of human populations. It has even been
possible to sequence DNA taken from a 76,000-year-old
mummied human brain and from a 17- to 20-million-year-
old plant fossil following PCR amplication. Sequencing
can be done quickly with present-day DNAsequencers that
make use of computers.
Recombinant DNA technology and the polymerase
chain reaction are two ways to clone a gene.
Today, bacteria, plants, and animals are genetically engi-
neered to produce biotechnology products. Organisms that
have had a foreign gene inserted into them are called trans-
genic organisms.
From Bacteria
Recombinant DNA technology is used to produce bacteria
that reproduce in large vats called bioreactors. If the foreign
gene is replicated and actively expressed, a large amount of
protein product can be obtained. Biotechnology products
produced by bacteria, such as insulin, human growth hor-
mone, t-PA (tissue plasminogen activator), and hepatitis B
vaccine, are now on the market (Fig. 26.5).
Transgenic bacteria have been produced to promote the
health of plants. For example, bacteria that normally live on
plants and encourage the formation of ice crystals have been
changed from frost-plus to frost-minus bacteria. Also, a bac-
terium that normally colonizes the roots of corn plants has
now been endowed with genes (from another bacterium)
that code for an insect toxin. The toxin protects the roots
from insects.
Bacteria can be selected for their ability to degrade a
particular substance, and then this ability can be enhanced
by genetic engineering. For instance, naturally occurring
bacteria that eat oil can be genetically engineered to do an
even better job of cleaning up beaches after oil spills
(Fig. 26.6). Industry has found that bacteria can be used as
biolters to prevent airborne chemical pollutants from being
vented into the air. They can also remove sulfur from coal
before it is burned and help clean up toxic waste dumps.
One such strain was given genes that allowed it to clean up
levels of toxins that would have killed other strains. Further,
these bacteria were given suicide genes that caused them
to self-destruct when the job had been accomplished.
Organic chemicals are often synthesized by having cata-
lysts act on precursor molecules or by using bacteria to carry
out the synthesis. Today, it is possible to go one step further
and to manipulate the genes that code for these enzymes. For
instance, biochemists discovered a strain of bacteria that is es-
pecially good at producing phenylalanine, an organic chemi-
cal needed to make aspartame, the dipeptide sweetener better
known as NutraSweet. They isolated, altered, and formed a
vector for the appropriate genes so that various bacteria could
be genetically engineered to produce phenylalanine.
Many major mining companies already use bacteria to
obtain various metals. Genetic engineering may enhance the
ability of bacteria to extract copper, uranium, and gold from
low-grade sources. Some mining companies are testing ge-
netically engineered organisms that have improved bio-
leaching capabilities.
Bacteria are being genetically altered to perform all
sorts of tasks, not only in the factory, but also in
the environment.
Figure 26.5 Biotechnology products.
Products like clotting factor VIII, which is administered to
hemophiliacs, can be made by transgenic bacteria, plants, or
animals. After being processed and packaged, it is sold as a
commercial product.
Figure 26.6 Bioremediation.
Bacteria capable of decomposing oil have been engineered and
patented by the investigator, Dr. Chakrabarty. In the inset, the ask
toward the rear contains oil and no bacteria; the ask toward the
front contains the bacteria and is almost clear of oil.
In a hungry world you would expect herbicide-resistant crops
to be greeted with enthusiasmgenetically engineered wheat
and corn offer the possibility of a more bountiful harvest and
the feeding of many more people. There are some, however,
who see dangers lurking in the use of biotechnology to develop
new and different strains of plants and animals. And these
doomsayers are not just anybodythey are ecologists.
First, we have to consider that herbicide-resistant crops will
allow farmers to use more herbicide than usual in order to kill
off weeds. Then, too, suppose this new form of wheat is better
able to compete in the wild. Certainly we know of plants that
have become pests when transported to a new environment:
prickly pear cactus took over many acres of Australia; an orna-
mental tree, the melaleuca, has invaded and is drying up many
of the swamps in Florida. Such plants spread because they are
able to overrun the native plants of an area. Perhaps genetically
engineered plants will also spread beyond their intended areas
and be out of control. Or worse, suppose herbicide-resistant
wheat were to hybridize with a weed, making the weed also re-
sistant and able to take over other agricultural elds. As more
and different herbicides are used to kill off the weed, the envi-
ronment would be degraded. And similar concerns pertain to
any transgenic organism, whether a bacterium, animal, or plant.
In the past, humans have been quick to believe that a new ad-
vance was the answer to a particular problem. The pesticide
DDT was going to kill off mosquitoes, making malaria a disease
of the past. And this worked until mosquitoes became resistant.
Today, we know that DDT accumulates in the tissues of hu-
mans, possibly contributing to all manner of health problems,
from reduced immunity to reproductive infertility. When antibi-
otics were rst introduced, it was hoped a disease like tubercu-
losis would be licked forever. Resistant strains of tuberculosis
have now evolved to threaten us all.
More and more transgenic varieties have been developed
and are being tested in agricultural elds. Afew of these have a
weedy relative in the wild with which they could hybridize.
Agricultural ofcials point out, however, that genetically engi-
neered plants have been growing in elds since 1994, and al-
though hybridization with weedy relatives may have occurred,
a superweed has not emerged. Still, say botanists, it could
happen in the future. Laboratory studies in Denmark showed
that a hybrid of transgenic oilseed rape and eld mustard did
resist herbicides and was able to produce highly fertile pollen.
Ecologists maintain it may be only a matter of time before a su-
perweed does appear in the wild.
Biotechnology: Friend or Foe?
From Plants
Techniques have been developed to introduce foreign genes
into immature plant embryos, or into plant cells that have
had the cell wall removed and are called protoplasts. It is
possible to treat protoplasts with an electric current while
they are suspended in a liquid containing foreign DNA. The
electric current makes tiny, self-sealing holes in the plasma
membrane through which genetic material can enter. Then a
protoplast will develop into a complete plant.
Foreign genes transferred to cotton, corn, and potato
strains have made these plants resistant to pests because
their cells now produce an insect toxin. Similarly, soybeans
have been made resistant to a common herbicide. Some corn
and cotton plants are both pest and herbicide resistant.
These and other genetically engineered crops are now reach-
ing the market place.
Plants are also being engineered to produce human pro-
teins, such as hormones, clotting factors, and antibodies, in
their seeds. One type of antibody made by corn can deliver
radioisotopes to tumor cells, and another made by soybeans
can be used as treatment for genital herpes. A weed called
mouse-eared cress has been engineered to produce a
biodegradable plastic (polyhydroxybutyrate, or PHB) in cell
granules.
Genetically engineered crops are now reaching the
market, and medicines made by genetically
engineered plants will soon be used to treat cancer
and other types of diseases.
From Animals
Techniques have been developed to insert genes into the
eggs of animals. It is possible to microinject foreign genes
into eggs by hand, but another method uses vortex mixing.
The eggs are placed in an agitator with DNA and silicon-
carbide needles, and the needles make tiny holes through
which the DNA can enter. When these eggs are fertilized,
the resulting offspring is a transgenic animal. Using this
technique, many types of animal eggs have taken up the
gene for bovine growth hormone (bGH). The procedure
has been used to produce larger shes, cows, pigs, rabbits,
and sheep. Genetically engineered fishes are now being
kept in ponds that offer no escape to the wild because there
is much concern that they will upset or destroy natural
ecosystems.
Gene pharming, the use of transgenic farm animals to
produce pharmaceuticals, is being pursued by a number of
rms. Genes that code for therapeutic and diagnostic pro-
teins are incorporated into the animals DNA, and the pro-
teins appear in the animals milk. There are plans to produce
drugs for the treatment of cystic brosis, cancer, blood dis-
eases, and other disorders. Antithrombin III, for preventing
blood clots during surgery, is currently being produced by a
herd of goats, and clinical trials have begun. Figure 26.7b, c
outlines the procedure for producing transgenic mammals:
DNA containing the gene of interest is injected into donor
eggs. Following in vitro fertilization, the zygotes are placed
in host females where they develop. After female offspring
mature, the product is secreted in their milk.
USDA scientists have been able to genetically engineer
mice to produce human growth hormone in their urine in-
stead of in milk. They expect to be able to use the same tech-
nique on larger animals. Urine is a preferable vehicle for a
biotechnology product rather than milk because all animals
in a herd urinateonly females produce milk; animals start
to urinate at birthfemales dont produce milk until matu-
rity; and its easier to extract proteins from urine than from
milk.
Xenotransplantation
Scientists have begun the process of genetically engineering
animals to serve as organ donors for humans who need a
transplant. Xenotransplantation is the use of animal organs
instead of human organs in transplant patients. We now
have the ability to transplant kidneys, heart, liver, pancreas,
lung, and other organs for two reasons. First, solutions have
been developed that preserve donor organs for several
hours, and second, rejection of transplanted organs can be
prevented by immunosuppressive drugs. Unfortunately,
however, there are not enough human donors to go around.
Fifty thousand Americans needed transplants in 1996, but
only 20,000 patients got them. As many as 4,000 died that
year while waiting for an organ.
Its no wonder, then, that scientists are suggesting that
we should get organs from a source other than another hu-
man. You might think that apes, such as the chimpanzee or
the baboon might be a scientically suitable species for this
purpose. But apes are slow breeders and probably cannot be
counted on to supply all the organs needed. Anyway, many
people might object to using apes for this purpose. In con-
trast, animal husbandry has long included the raising of
pigs as a meat source, and pigs are prolic. Afemale pig can
become pregnant at six months and can have two litters a
year, each averaging about ten offspring.
Ordinarily, humans would violently reject transplanted
pig organs. Genetic engineering, however, can make these
organs less antigenic. Scientists have produced a strain of
pigs whose organs would most likely, even today, survive
for a few months in humans. They could be used to keep a
patient alive until a human organ was available. The ulti-
mate goal is to make pig organs as widely accepted by hu-
mans as type O blood. Aperson with type O blood is called
a universal donor because the red blood cells carry no AB
nor B antigens.
As xenotransplantation draws near, other concerns
have been raised. Some experts fear that animals might be
infected with viruses, akin to Ebola virus or the virus that
causes mad cow disease. After infecting a transplant pa-
tient, these viruses might spread into the general populace
and begin an epidemic. Scientists believe that HIV was
spread to humans from monkeys when humans ate mon-
key meat. Those in favor of using pigs for xenotransplan-
tation point out that pigs have been around humans for
centuries without infecting them with any serious
diseases.
Cloning of Animals
Imagine that an animal has been genetically altered to pro-
duce a biotechnology product or to serve as an organ
donor. What would be the best possible way to get identi-
cal copies of this animal? If cloning of the animal was pos-
sible, you could get many exact copies of this animal.
Asexual reproduction through cloning would be the pre-
ferred procedure to use. Cloning is a form of asexual repro-
duction because it requires only the genes of that one
animal. For many years it was believed that adult verte-
brate animals could not be cloned. Although each cell con-
tains a copy of all the genes, certain genes are turned off in
mature specialized cells. Different genes are expressed in
muscle cells, which contract, compared to nerve cells
which conduct nerve impulses, and to glandular cells,
which secrete. Cloning of an adult vertebrate would re-
quire that all genes of an adult cell be turned on again if de-
velopment is to proceed normally. It has long been thought
this would be impossible.
But in 1997 scientists at the Raslin Institute in Scotland
announced that they achieved this feat and had produced
a cloned sheep called Dolly. In 1998, genetically altered
calves were cloned in the United States using the same
26-8 536 Part 5 Continuance of the Species
method. An alternate method, used at the Univer-
sity of Hawaii for the cloning of mice, was so suc-
cessful that clones of clones were produced. Figure
26.7c suggests that after enucleated eggs have been
injected with 2n nuclei of adult cells, they can be
coaxed to begin development. The offspring have
the genotype and phenotype of the adult that do-
nated the nuclei; therefore, the adult has been
cloned. In the procedure that produced cloned
mice, the 2n nuclei were taken from cumulus cells.
Cumulus cells are those that cling to an egg after
ovulation occurs. A specially prepared chemical
bath was used to stimulate the eggs to divide and
begin development. Now that scientists have a
method to clone mammals, this procedure will un-
doubtedly be used routinely. Some are even begin-
ning to think about the cloning of humans despite
the objections of many and a presidential order that
the procedure is not to be developed in the United
States.
Transgenic animals, which secrete a
biotechnology product in their milk, and
pigs, whose organs can be used for
xenotransplantation, have been produced.
Procedures have been developed to allow
the cloning of these animals.
human gene
egg donor
microinjection of human gene
development within a host goat
transgenic goat
milk from transgenic goat
b. Making a transgenic animal
2n nuclei
adult cells from
transgenic goat
microinjection of 2n nuclei
egg donor
enucleated eggs
development within host goats
cloned transgenic goats
milk from cloned goats
c. Making clones of a transgenic animal
egg
a.
Figure 26.7 Genetically engineered animals.
a. This goat is genetically engineered to produce
antithrombin III, which is secreted in her milk. This
researcher and many others are involved in the project.
b. The procedure to produce a transgenic animal.
c. The procedure to clone a transgenic animal.
The Human Genome Project is a massive effort originally
funded by the U.S. government and now increasingly by
U.S. pharmaceutical companies to map the human chromo-
somes. Many nonprot and for prot biochemical laborato-
ries about the world are now involved in the project which
has two primary goals.
The rst goal is to construct a genetic map of the human
genome. The aim is to show the sequence of genes along the
length of each type chromosome, such as depicted for the X
chromosome in Figure 26.8. If the estimate of 1,000+ human
genes is correct, each chromosome on average would con-
tain about 50 alleles.
The map for each chromosome is presently incomplete,
and in many instances scientists rely on the placement of
RFLPs (see page 533). These sites eventually allow scientists
to pinpoint disease-causing genes because a particular RFLP
and a defective gene are often inherited together. For
example, it is known that persons with Huntington disease
have a unique site where a restriction enzyme cuts DNA.
The test for Huntington disease relies on this difference from
the normal.
The genetic map of a chromosome can be used not only
to detect defective genes, but possibly also to tailor treat-
ments to the individual. Only certain hypertension patients
benet from a low-salt diet, and it would be useful to know
which patients these are. Myriad Genetics, a genome com-
pany, has developed a test for a mutant angiotensinogen
gene because they want to see if patients with this mutation
are the ones that benet from a low-salt diet. Several other
mutant genes have also been correlated with specic drug
treatments (Table 26.1). One day the medicine you take
might carry a label that it is effective only in persons with
genotype #101!
The second goal is to construct a base sequence map.
There are three billion base pairs in the human genome, and
its estimated it would take an encyclopedia of 200 volumes,
each with 1,000 pages, to list all of these. Yet, this goal of the
Human Genome Project is expected to be reached by the
year 2004, if not earlier.
The methodology, thus far, has been to rst chop up the
genome into small pieces, each just 1,000 to 2,000 base pairs
long. PCR instruments copy the pieces many times, and then
an automatic DNA sequencer determines the order of the
base pairs. You need many DNAcopies because of the way
the sequencer works. A computer program later strings the
sequenced pieces together in the correct order by looking for
base sequence overlaps between them. Instrumentation has
gradually improved, and recently one scientist, J. Craig Ven-
ter, has founded a company which he says will sequence the
entire genome in three years.
Venter plans on using what is called a whole-genome
shotgun sequencing method. He plans on working with the
entire human genome at once. Each overlapping fragment
will be about 5,000 bases long, and he will sequence the ends
of each fragment using only powerful new sequencing ma-
chines. Again, a computer program will string the fragments
together by looking for overlapping regions. Why is Venter
going off on his own, instead of participating in the world-
wide effort by many laboratories and scientists to sequence
the human genome? His backers expect to market the
whole-genome database to subscribers, and to patent rare
but pharmacologically interesting genes. Private enterprise
is giving new impetus to the eld now called genomics.
Knowing the base sequence of normal genes may make
it possible one day to treat certain human ills by administer-
ing normal genes and/or their protein products to those
who suffer from a genetic disease.
ichthyosis, X linked
hypophosphatemia
ocular albinism
Duchenne muscular dystrophy
retinitis pigmentosa
Lesch-Nyhan syndrome
hemophilia B
fragile X syndrome
hemophilia A
color blindness (several forms)
spastic paraplegia, X linked
Mutant Gene for Disease Treatment
Apolipoprotein E Alzheimer Experimental Glaxo
Wellcome drug
Cytochrome P-450 Cancer Amonade
Chloride gate Cystic brosis Pulmozyme
Dopamine receptor D4 Schizophrenia Clozapine
Angiotensinogen Hypertension Low-salt diet
Table 26.1 Customizing Drug Treatments
Figure 26.8 Genetic map of X chromosome.
The human X chromosome has been partially mapped, and this is the
order of some of the genes now known to be on this chromosome.
26.4 Gene Therapy
Gene therapy is the insertion of genetic material into human
cells for the treatment of a disorder. It includes procedures
that give a patient healthy genes to make up for faulty genes
and also includes the use of genes to treat various other hu-
man illnesses such as cancer and cardiovascular disease.
Currently there are approximately 1,000 patients enrolled in
nearly 200 approved gene therapy trials in the United States.
There are ex vivo (outside the body) and in vivo (inside the
body) methods of gene therapy.
Genetic Disorders
As an example of an ex vivo method of gene therapy, con-
sider Figure 26.9, which describes a methodology for the
treatment of children with severe combined immunode-
ciency syndrome (SCID). These children lack an enzyme
called adenosine deaminase (ADA) that is involved in the
maturation of T and B cells, and therefore they are subject to
life-threatening infections. Bone marrow stem cells are re-
moved from the blood and infected with a retrovirus (RNA
virus) that carries a normal gene for the enzyme. Then the
cells are returned to the patient. Bone marrow stem cells are
preferred for this procedure because they divide to produce
more cells with the same genes. With an ex vivo method of
genetic therapy, it is possible to test and make sure gene
transfer has occurred before the cells are returned to the pa-
tient. Patients who have undergone this procedure do have
a signicant improvement in their immune function that is
associated with a sustained rise in the level of ADAenzyme
activity in the blood.
Among the many gene therapy trials, one is for the treat-
ment of familial hypercholesterolemia, a condition that de-
velops when liver cells lack a receptor for removing
cholesterol from the blood. The high levels of blood choles-
terol make the patient subject to fatal heart attacks at a
young age. In a newly developed procedure, a small portion
of the liver is surgically excised and infected with a retro-
virus containing a normal gene for the receptor. Several pa-
tients have experienced a lowering of serum cholesterol
levels following this procedure.
Cystic brosis patients lack a gene that codes for trans-
membrane carrier of the chloride ion. Patients often die due
to numerous infections of the respiratory tract. An in vivo
method of treatment is being tried. Liposomesmicro-
scopic vesicles that spontaneously form when lipoproteins
are put into a solutionhave been coated with the gene
needed to cure cystic brosis. Then the solution is sprayed
into patients nostrils. Due to limited gene transfer this
methodology has not as yet been successful.
In a recent and surprising move, some researchers are
investigating the possibility of directly correcting the base
sequence of patients with a genetic disorder. The exact pro-
cedure has not yet been published.
Cancer
Chemotherapy in cancer patients often kills off healthy cells
as well as cancer cells. In clinical trials, researchers have
given genes to cancer patients that either make healthy cells
more tolerant of chemotherapy or make tumors more vul-
nerable to it. In one ex vivo clinical trial, bone marrow stem
cells from about 30 women with late-stage ovarian cancer
were infected with a virus carrying a gene to make them
defective
gene
1.
2.
2. Use retroviruses to
infect stem cells
with normal gene.
3. Recombinant
DNA carries
normal gene
into genome.
4. Return genetically
engineered cells
to patient.
retrovirus
normal
gene
recombinant
RNA
reverse
transcription
recombinant
DNA
4.
1. Remove stem cells.
3.
Figure 26.9 Ex vivo gene therapy in humans.
Bone marrow stem cells are withdrawn from the body, a virus is used to insert a normal gene into them, and they are returned to the body.
Other Illnesses
During coronary artery angioplasty, a balloon catheter is
sometimes used to open up a closed artery. Unfortunately,
the artery has a tendency to close up again. But investiga-
tors have come up with a new procedure. The balloon is
coated with a plasmid that contains a gene for VEGF (vas-
cular endothelial growth factor). The expression of the
gene, which promotes the proliferation of blood vessels to
bypass the obstructed area, has been observed in at least
one patient.
Perhaps it will be possible also to use in vivo therapy to
cure hemophilia, diabetes, Parkinson disease, or AIDS. To
treat hemophilia, patients could get regular doses of cells
that contain normal clotting-factor genes. Or such cells
could be placed in organoids, articial organs that can be im-
planted in the abdominal cavity. To cure Parkinson disease,
dopamine-producing cells could be grafted directly into the
brain.
The Human Genome Project produces information
useful to gene therapists. Researchers are
envisioning all sorts of ways to cure human genetic
disorders as well as many other types of illnesses.
more tolerant of chemotherapy. Once the bone marrow
stems cells were protected, it was possible to increase the
level of chemotherapy to kill the cancer cells
In a recent in vivo study, a retrovirus carrying a normal
p53 gene was injected directly into the tumor of lung cancer
patients. This gene, which helps regulate the cell cycle and
also brings about apoptosis in cells with damaged DNA, is
often mutated in tumor cells. No cures were reported but the
tumors shrank in three patients and stopped growing in
three others. Expression of p53 is only needed for a short
time and an elevated amount in normal cells seems to do
them no harm.
Some investigators prefer the use of adenoviruses rather
than retroviruses. Adenoviruses can be produced in much
larger quantities and they are active even when they are not
dividing. (Retroviruses integrate DNAinto the host chromo-
some where it is inactive unless replication occurs.) When
adenoviruses infect a cell, they normally produce a protein
that binds to p53 and inactivates it. In a cleverly designed
procedure, researchers genetically engineered adenoviruses
to lack the gene that produces this protein. Therefore, the
virus will kill tumor cells that lack p53 but not healthy cells
that have a p53 gene. Clinical trials are underway, and it is ex-
pected that the virus will spread through the cancer, killing
the cancer cells, and stopping when it reaches normal cells.
S
omatic gene therapy attempts to treat
or prevent human illnesses. Some day,
for example, it may be possible to give
children who have cystic brosis, Hunt-
ington disease, or any other genetic disor-
der a normal gene to make up for the
inheritance of a faulty gene. And gene
therapy is even more likely for treatment
of diseases like cancer, AIDS, and heart
disease. Germ line gene therapy is the
term that is now being used to mean the
use of gene therapy solely to improve the
traits of an individual. It would be the ge-
netic equivalent of procedures like body
building, liposuction, or hair transplants.
If genetic interference occurred earlier
that is, on the eggs, sperm, or embryo, its
possible that it would indeed affect the
germ linethat is, all the future descen-
dants of the individual.
How might germ line gene therapy
become routine? Consider this scenario.
Presently, a gene for VEGF (vascular en-
dothelial growth factor), a protein pro-
duced by cells to grow new blood vessels,
is being used to treat atherosclerosis. Im-
proved arterial circulation in the legs of
some recent patients did away with the
threat of possible amputation. This same
gene is now being considered for the treat-
ment of blocked coronary arteries. There
may be instances, though, in which
healthy people want to grow new blood
vessels for enhancement purposes. Run-
ners might want to improve their circula-
tion in order to win races, and parents
might think that increased circulation to
the brain might increase the intelligence of
their children. Bioethicist Eric Juengst of
Case Western thinks that as a society, there
is nothing we can now do to prevent us
from crossing the line between the use of
gene therapy for therapeutic purposes and
its use for enhancement reasons.
Questions
1. Do you approve of gene therapy to treat
or prevent human illnesses? after they
occur? or possibly before they occur, if
genetic testing shows the likelihood of
their development? Why or why not?
2. Do you approve of gene therapy to
enhance the characteristics of an
individual? after a condition like baldness
occurs? or possibly before it occurs if
genetic testing shows the likelihood of it
occurring? Why or why not?
3. Do you approve of somatic gene therapy
of embryos? germ line gene therapy of
embryos? Explain.
Go To Student OLC
Summarizing the Concepts
26.1 Cloning of a Gene
Two methods are currently available for making copies of DNA. Re-
combinant DNA contains DNA from two different sources. A restric-
tion enzyme is used to cleave plasmid DNA and to cleave foreign
DNA. The sticky ends produced facilitate the insertion of foreign
DNAinto vector DNA. The foreign gene is sealed into the vector DNA
by DNAligase. Both bacterial plasmids and viruses can be used as vec-
tors to carry foreign genes into bacterial host cells.
Agenomic library can be used as a source of genes to be cloned. A
radioactive or uorescent probe is used to identify the location of a sin-
gle gene among the cloned fragments of an organisms DNA.
Restriction enzymes are used to fragment DNA. If the entire
genome is used for ngerprinting, the use of probes is necessary in or-
der to see a pattern following gel electrophoresis. If just a portion of the
genome is of interest or available, PCR uses the enzyme DNA poly-
merase to make multiple copies of target DNA. Analysis of DNA seg-
ments following PCR can involve gel electrophoresis to identify the
DNAas belonging to a particular organism or can involve determining
the base sequence of the DNAsegment.
Transgenic organisms are ones that have had a foreign gene inserted
into them. Genetically engineered bacteria, agricultural plants, and
farm animals now produce commercial products of interest to humans,
such as hormones and vaccines. Bacteria secrete the product. The seeds
of plants and the milk of animals contain the product.
Transgenic bacteria have been produced to promote the health of
plants, perform bioremediation, extract minerals, and produce chemi-
cals. Transgenic agricultural plants which have been engineered to re-
sist herbicides and pests are commercially available. Transgenic
animals have been given various genes, in particular the one for bovine
growth hormone (bGH). Pigs have been genetically altered to serve as
a source of organs for transplant patients. Cloning of animals is now
possible.
The Human Genome Project has two goals. The rst goal is to construct
a genetic map which will show the sequence of the genes on each chro-
mosome. Although the known location of genes is expanding, the ge-
netic map still contains many RFLPs (restriction fragment length
polymorphisms) as the rst step toward nding more genes. The sec-
ond goal of the project is to sequence the bases. To do this researchers
rely on PCR and automatic sequencing instruments. As the sequencers
have improved so has the speed with which DNA is sequenced. The
hope is that knowing the location and sequence of bases in a gene will
promote the possibility of gene therapy.
26.4 Gene Therapy
Gene therapy is used to correct the genotype of humans and to cure
various human ills. There are ex vivo and in vivo methods of gene
therapy. Gene therapy has apparently helped children with SCID to
lead a normal life; the treatment of cystic brosis has been less suc-
cessful. Anumber of imaginative therapies are being employed in the
war against cancer and other human illness such as cardiovascular
disease.
Studying the Concepts
1. What is the methodology for producing recombinant DNAto
be used in gene cloning? 530
2. Bacteria can be used to clone a gene or produce a product.
Explain. 531
3. What is a genomic library, and how do you locate a gene of
interest in the library? 532
4. What is the polymerase chain reaction (PCR), and how is
it carried out to produce multiple copies of a DNA
segment? 532
5. What is DNAngerprinting, a process that utilizes the entire
genome? 533
6. What are some practical applications of DNAsegments
analysis following PCR? 53233
7. For what purposes have bacteria, plants, and animals been
genetically altered? 53436
8. What is xenotransplantation, and what drawback is presently
preventing pigs from serving as source animals? 536
9. Explain how and why transgenic animals that secrete a prod-
uct are often cloned. 53637
10. Explain the two primary goals of the Human Genome Project.
What are the possible benets of the project? 538
11. Explain and give examples of ex vivo and in vivo gene thera-
pies in humans. 53940
Testing Yourself
Choose the best answer for each question.
1. Which of these is a true statement?
a. Both plasmids and viruses can serve as vectors.
b. Plasmids can carry recombinant DNAbut viruses cannot.
c. Vectors carry only the foreign gene into the host cell.
d. Only gene therapy uses vectors.
e. Both a and d are correct.
2. Which of these is a benet to having insulin produced by
biotechnology?
a. It is just as effective. d. It is less expensive.
b. It can be mass-produced. e. All of these are correct.
c. It is nonallergenic.
3. Restriction fragment length polymorphisms (RFLPs)
a. are achieved by using restriction enzymes.
b. identify individuals genetically.
c. are the basis for DNAngerprints.
d. can be subjected to gel electrophoresis.
e. All of these are correct.
4. Which of these would you not expect to be a biotechnology
product?
a. vaccine d. protein hormone
b. modied enzyme e. steroid hormone
c. DNAprobe
5. What is the benet of using a retrovirus as a vector in gene
therapy?
a. It is not able to enter cells.
b. It incorporates the foreign gene into the host chromosome.
c. It eliminates a lot of unnecessary steps.
d. It prevents infection by other viruses.
e. Both b and c are correct.
6. Gel electrophoresis
a. cannot be used on nucleotides.
b. measures the size of plasmids.
c. tells whether viruses are infectious.
d. measures the charge and size of proteins and DNAfrag-
ments.
e. All of these are correct.
7. Using this key, put the phrases in the correct order to form a
plasmid-carrying recombinant DNA.
Key:
(1) use restriction enzymes
(2) use DNAligase
(3) remove plasmid from parent bacterium
(4) introduce plasmid into new host bacterium
a. 1,2,3,4
b. 4,3,2,1
c. 3,1,2,4
d. 2,3,1,4
8. Which of these is incorrectly matched?
a. xenotransplantationsource of organs
b. protoplastplant cell engineering
c. RFLPsDNAngerprinting
d. DNApolymerasePCR
e. DNAligasemapping human chromosomes
9. The restriction enzyme called EcoRI has cut double-stranded
DNAin this manner. The piece of foreign DNAto be inserted
has what bases from left to the right?
3. What experiment would you suggest to support the hypothe-
sis that plants engineered to contain genes for an insect toxin
against insects is safe for human consumption? Why might
your results not support the hypothesis?
4. What experiment would you suggest to support the hypothesis
that bacteria engineered to clean up a pollutant disappear when
the pollutant is gone? What would make them disappear?
C
A A
G
T T
A
T
A
T
C
G
a.
b.
c.
d.
e.
f.
g.
10. Label the following drawings, using these terms: retrovirus,
recombinant RNA(twice), defective gene, recombinant DNA,
reverse transcription, and human genome.
Thinking Scientically
sis that insulin produced by biotechnology will have fewer
side effects than insulin taken from the organs of livestock?
Why do you expect your results to support the hypothesis?
sis that meat from cattle bioengineered to contain extra
growth hormone genes is safe for human consumption? Why
do you expect your results to support the hypothesis? Your
answer should consider the fact that growth hormone is a
protein.
Understanding the Terms
bacteriophage 532
clone 531
complementary DNA
(cDNA) 532
DNAngerprinting 533
DNAligase 530
gene therapy 539
genetic engineering 530
genome 532
genomic library 532
plasmid 530
polymerase chain reaction
(PCR) 532
probe 532
recombinant DNA(rDNA)
530
restriction enzyme 530
restriction fragment length
polymorphism (RFLP) 533
transgenic organism 534
vector 530
xenotransplantation 536
Match the terms to these denitions:
a. Bacterial enzyme that stops viral reproduction by
cleaving viral DNA; used to cut DNAat specic points dur-
ing production of recombinant DNA.
b. Free-living organisms in the environment that
have had a foreign gene inserted into them.
c. Known sequences of DNAthat are used to nd
complementary DNAstrands; can be used diagnostically to
determine the presence of particular genes.
d. Production of identical copies; in genetic engi-
neering, the production of many identical copies of a gene.
e. Self-duplicating ring of accessory DNAin the
cytoplasm of bacteria.
Using Technology
Your study of biotechnology is supported by these available
technologies.
Essential Study Partner CD-ROM
Genetics Recombinant DNA
Visit the Mader web site for related ESP activities.
Exploring the Internet
The Mader Home Page provides resources and tools as
you study this chapter.
http://www.mhhe.com/biosci/genbio/mader
Life Science Animations 3D Video
24 Polymerase Chain Reaction
25 Gel Electrophoresis
26 Sucrose Gradient Centrifugation
27 Protein Purication/Column Chromatography
experiments in genetics and molecular biology that became the
foundation for todays research.
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279(4):60. Selective estrogen receptor modulators may protect
against breast and endometrial cancers, osteoporosis, and heart
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the initiation of human labor.
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Further Readings for Part 5
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focuses on the biology of AIDS.
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contraceptives in protecting women against STDs.
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American 278(62):4. New laser techniques allow manipulation of
chromosomes and other structures inside cells.
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American 274(2):104. Acid-loving pathogens are linked to
stomach ulcers and stomach cancer.
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Science & Medicine 4(6):52. The importance of supplemental
antioxidant vitamins depends on factors such as diet and life-
style.
Cox, F. D. 1996. The AIDS booklet. 4th ed. Dubuque, Iowa: Brown &
Benchmark Publishers. This easy to read, informative booklet
covers the transmission, prevention, and treatment of AIDS.
Crooks, R., and Baur, K. 1996. Our sexuality. 6th ed. Redwood City,
Calif.: Benjamin/Cummings Publishing. Introduction to the
biological, psychosocial, behavioral, and cultural aspects of
sexuality.
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malignancies. Science & Medicine 4(5):4. The eld of AIDS-
related gene therapies is advancing.
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disease. Scientic American 275(6):80. Failures in the processes of
cellular self-destruction may give rise to cancer, AIDS,
Alzheimer disease, and some genetic diseases.
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New York: Scientic American Library. This easy-to-read, well-
illustrated text describes how microbes respond to the physical
demands of their environment.
Galili, U. September/October 1998. Anti-Gal antibody prevents
xenotransplantation. Science & Medicine 5(5):28. Prevention of
interaction of the anti-Gal antibody with pig cells is necessary
to the progress of xenotransplantation.
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prostate cancer. Scientic American 279(6):74. Article details the
recent developments in diagnosis and treatment of prostate
cancer.
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Genes from genetically engineered plants can escape from crops
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cells.
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enzyme telomerase rebuilds the chromosomes of tumor cells;
this enzyme is being researched as a target for anticancer
treatments.
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being developed are a result of recent genetic analyses of the
human genome.
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The search is underway for the genes involved in cognitive
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Scientic American Special Issue. September 1996. What you need
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Scientic American 274(1):64. Consuming fewer calories may
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Chlamydial resistance to host defense. Science & Medicine
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suggest the same basic gene may be involved in the
development of certain features in vertebrates and
invertebrates.

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Biotechnology

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