Solution Manual for Microbiology: An Introduction, 13th Edition, Gerard J.

Tortora, Berdell

Solution Manual for Microbiology: An Introduction,

13th Edition, Gerard J. Tortora, Berdell R. Funke,
Christine L. Case, Derek Weber, Warner Bair

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 CHAPTER
Microbial Genetics
8 Global Edition

Learning Objectives Check Your Understanding

8-1 Define genetics, genome, Give a clinical application of genomics.
chromosome, gene, genetic code,
genotype, phenotype, and genomics.
8-2 Describe how DNA serves as genetic Why is the base pairing in DNA important?
information.
8-3 Describe the process of DNA Describe DNA replication, including the
replication. functions of DNA gyrase, DNA ligase, and DNA
polymerase.
8-4 Describe protein synthesis, including What is the role of the promoter, terminator, and
transcription, RNA processing, and mRNA in transcription?
translation.
8-5 Compare protein synthesis in prokary- How does mRNA production in eukaryotes
otes and eukaryotes. differ from the process in prokaryotes?
8-6 Define operon. Use the following metabolic pathway to answer
the questions that follow it.

Substrate A ⎯⎯⎯⎯
→ Intermediate B
enzyme a

8-7 Explain pre-transcriptional regulation
of gene expression in bacteria.
8-8 Explain post-transcriptional regulation
of gene expression.
8-9 Classify mutations by type.
8-10 Describe two ways mutations can be
Copyright © 2021 Pearson Education Ltd.
⎯⎯⎯⎯
enzyme b
→ End-product C
a. If enzyme a is inducible and is not being
synthesized at present, a (1) _____ protein must
be bound tightly to the (2) _____ site. When the
inducer is present, it will bind to the (3) _____ so
that (4) _____ can occur.
b. If enzyme a is repressible, end-product C,
called a (1) _____, causes the (2) _____ to bind
to the (3) _____. What causes derepression?
What is the role of cAMP in regulating gene
expression?
How does miRNA stop protein synthesis?
How can a mutation be beneficial?
How can mutations be repaired?
79
 repaired.
8-11 Describe the effect of mutagens on the
mutation rate.
8-12 Outline the methods of direct and
indirect selection of mutants.
8-13 Identify the purpose of and outline the
procedure for the Ames test.
8-14 Describe the functions of plasmids and
transposons.
8-15 Differentiate horizontal and vertical
gene transfer.
8-16 Compare the mechanisms of genetic
recombination in bacteria.
8-17 Discuss how genetic mutation and
recombination provide material for
natural selection to act upon.
How do mutagens affect the mutation rate?
How would you isolate an antibiotic-resistant
bacterium? An antibiotic-sensitive bacterium?
What is the principle behind the Ames test?
What types of genes do plasmids carry?
Differentiate horizontal and vertical gene
transfer.
Compare conjugation between the following
pairs:
F+  F–, Hfr  F–.
Natural selection means that the environment
favors survival of some genotypes. From where
does diversity in genotypes come?

New in This Edition

• The discussion of operons, induction, and repression has been revised.
• Riboswitches are defined.
• A new box about tracking Zika virus is included.

Chapter Summary
Structure and Function of the Genetic Material (pp. 231–241)
ASM 4.2: Although the central dogma is universal in all cells, the
processes of replication, transcription, and translation differ in
Bacteria, Archaea, and Eukaryotes.
1. Genetics is the study of what genes are, how they carry information, how their
information is expressed, and how they are replicated and passed to subsequent
generations or other organisms.
2. DNA in cells exists as a double-stranded helix; the two strands are held together by
hydrogen bonds between specific nitrogenous base pairs: AT and CG.
3. A gene is a sequence of nucleotides, that encodes a functional product, usually a protein.
4. The DNA in a cell is duplicated before the cell divides, so each offspring cell receives the
same genetic information.

Genotype and Phenotype (pp. 231–234)

5. Genotype is the genetic composition of an organism, its entire complement of DNA.

80 INSTRUCTOR'S GUIDE FOR MICROBIOLOGY: AN INTRODUCTION, GE, 13e Copyright © 2021 Pearson Education Ltd.
 6. Phenotype is the expression of the genes: the proteins of the cell and the properties they
confer on the organism.

DNA and Chromosomes (p. 234)

7. The DNA in a chromosome exists as one long double helix associated with various
proteins that regulate genetic activity.
8. Genomics is the molecular characterization of genomes.

The Flow of Genetic Information (p. 234)

9. Following cell division, each offspring cell receives a chromosome that is virtually
identical to the parent’s.
10. Information contained in the DNA is transcribed into RNA and translated into proteins.

DNA Replication (pp. 234–238)

11. During DNA replication, the two strands of the double helix separate at the replication
fork, and each strand is used as a template by DNA polymerases to synthesize two new
strands of DNA according to the rules of complementary base pairing.
12. The result of DNA replication is two new strands of DNA, each having a base sequence
complementary to one of the original strands.
13. Because each double-stranded DNA molecule contains one original and one new strand,
the replication process is called semiconservative.
14. DNA is synthesized in one direction designated 5 → 3. At the replication fork, the
leading strand is synthesized continuously and the lagging strand discontinuously.
15. DNA polymerase proofreads new molecules of DNA and removes mismatched bases
before continuing DNA synthesis.

RNA and Protein Synthesis (pp. 238–241)

16. During transcription, the enzyme RNA polymerase synthesizes a strand of RNA from
one strand of double-stranded DNA, which serves as a template.
17. RNA is synthesized from nucleotides containing the bases A, C, G, and U, which pair
with the bases of the DNA strand being transcribed.
18. RNA polymerase binds the promoter; transcription begins at AUG; the region of DNA
that is the end point of transcription is the terminator; RNA is synthesized in the 5 → 3
direction.
19. Translation is the process in which the information in the nucleotide base sequence of
mRNA is used to dictate the amino acid sequence of a protein.
20. The mRNA associates with ribosomes, which consist of rRNA and protein.
21. Three-base codons of mRNA specify amino acids.
22. The genetic code refers to the relationship among the nucleotide base sequence of DNA,
the corresponding codons of mRNA, and the amino acids for which the codons code.
23. Specific amino acids are attached to molecules of tRNA. Another portion of the tRNA
has a base triplet called an anticodon.
24. The base pairing of codon and anticodon at the ribosome results in specific amino acids
being brought to the site of protein synthesis.
Copyright © 2021 Pearson Education Ltd. CHAPTER 8 Microbial Genetics 81
 25. The ribosome moves along the mRNA strand as amino acids are joined to form a grow-
ing polypeptide; mRNA is read in the 5 → 3 direction.
26. Translation ends when the ribosome reaches a stop codon on the mRNA.

The Regulation of Bacterial Gene Expression (pp. 241–247)

ASM 4.3: The regulation of gene expression is influenced by external

and internal molecular cues and/or signals.
1. Regulating protein synthesis at the gene level is energy-efficient because proteins are
synthesized only as they are needed.
2. Constitutive genes are expressed at a fixed rate. Examples are genes for the enzymes in
glycolysis.

Pre-transcriptional Control (pp. 241–246)

3. In bacteria, a group of coordinately regulated structural genes with related metabolic
functions, plus the promoter and operator sites that control their transcription, is called an
operon.
4. In the operon model for an inducible system, a regulatory gene codes for the repressor
protein.
5. When the inducer is absent, the repressor binds to the operator, and no mRNA is
synthesized.
6. When the inducer is present, it binds to the repressor so that it cannot bind to the
operator; thus, mRNA is made, and enzyme synthesis is induced.
7. In repressible systems, the repressor requires a corepressor in order to bind to the
operator site; thus, the corepressor controls enzyme synthesis.
8. Transcription of structural genes for catabolic enzymes (such as β-galactosidase) is
induced by the absence of glucose. Cyclic AMP and CRP must bind to a promoter in the
presence of an alternative carbohydrate.
9. Methylated nucleotides are not transcribed in epigenetic control.

Post-transcriptional Control (pp. 246–247)

10. mRNA as a riboswitch regulates translation.
11. MicroRNAs combine with mRNA; the resulting double-stranded RNA is destroyed.

Changes in Genetic Material (pp. 247–254)

ASM 4.1: Genetic variations can impact microbial functions

(e.g., in biofilm formation, pathogenicity, and drug resistance).

ASM 1.2: Mutations and horizontal gene transfer, and the immense
variety of microenvironments, have selected for a huge diversity of
microorganisms.
1. Mutations and horizontal gene transfer can change a bacterium’s genotype.

Mutation (p. 248)

82 INSTRUCTOR'S GUIDE FOR MICROBIOLOGY: AN INTRODUCTION, GE, 13e Copyright © 2021 Pearson Education Ltd.
 2. A mutation is a change in the nitrogenous base sequence of DNA; that change causes a
change in the product coded for by the mutated gene.
3. Many mutations are neutral, some are disadvantageous, and others are beneficial.

Types of Mutations (pp. 248–249)

4. A base substitution occurs when one base pair in DNA is replaced with a different base
pair.
5. Alterations in DNA can result in missense mutations or nonsense mutations.
6. Spontaneous mutations occur without the presence of any mutagen.

Mutagens (pp. 249–252)

7. Mutagens are agents in the environment that cause permanent changes in DNA.
8. Ionizing radiation causes the formation of ions and free radicals that react with DNA; it
results in base substitutions or breakage of the sugarphosphate backbone.
9. Ultraviolet (UV) radiation is nonionizing; it causes bonding between adjacent thymines.

The Frequency of Mutation (p. 252)

10. Mutation rate is the probability that a gene will mutate when a cell divides; the rate is
expressed as 10 to a negative power.
11. A low rate of spontaneous mutations is beneficial in providing the genetic diversity
needed for evolution.

Identifying Mutants (pp. 252–253)

12. Mutants can be detected by selecting or testing for an altered phenotype.
13. Positive selection involves the selection of mutant cells and the rejection of nonmutated
cells.
14. Replica plating is used for negative selection—to detect, for example, auxotrophs that
have nutritional requirements not possessed by the parent (nonmutated) cell.

Identifying Chemical Carcinogens (pp. 253–254)

15. The Ames test is a relatively inexpensive and rapid test for identifying possible chemical
carcinogens.
16. The test assumes that a mutant cell can revert to a normal cell in the presence of a
mutagen and that many mutagens are carcinogens.

Genetic Transfer and Recombination (pp. 255–263)

ASM 1.4: The traditional concept of species is not readily applicable to

microbes due to asexual reproduction and the frequent occurrence of
horizontal gene transfer.
1. Genetic recombination, the rearrangement of genes from separate groups of genes,
usually involves DNA from different organisms; it contributes to genetic diversity.

Copyright © 2021 Pearson Education Ltd. CHAPTER 8 Microbial Genetics 83

 2. In crossing over, genes from two chromosomes are recombined into one chromosome
containing some genes from each original chromosome.
3. Vertical gene transfer occurs during reproduction when genes are passed from an
organism to its offspring.
4. Horizontal gene transfer in bacteria involves a portion of the cell’s DNA being
transferred from donor to recipient.
5. When some of the donor’s DNA has been integrated into the recipient’s DNA, the
resultant cell is called a recombinant.

Plasmids and Transposons (pp. 255–258)

6. Plasmids are self-replicating circular molecules of DNA carrying genes that are not
usually essential for the cell’s survival.
7. There are several types of plasmids, including conjugative plasmids, dissimilation
plasmids, plasmids carrying genes for toxins or bacteriocins, and resistance factors.
8. Transposons are small segments of DNA that can move from one region to another
region of the same chromosome or to a different chromosome or a plasmid.
9. Complex transposons can carry any type of gene, including antibiotic-resistance genes,
and are thus a natural mechanism for moving genes from one chromosome to another.

Transformation in Bacteria (pp. 258–260)

10. During this process, genes are transferred from one bacterium to another as “naked”
DNA in solution.

Conjugation in Bacteria (pp. 260–261)

11. This process requires contact between living cells.
12. One type of genetic donor cell is an F+; recipient cells are F–. F cells contain plasmids
called F factors; these are transferred to the F– cells during conjugation.

Transduction in Bacteria (pp. 261–263)

13. In this process, DNA is passed from one bacterium to another in a bacteriophage and is
then incorporated into the recipient’s DNA.
14. In generalized transduction, any bacterial genes can be transferred.

Genes and Evolution (p. 263)

ASM 1.2: Mutations and horizontal gene transfer, and the immense
variety of microenvironments have selected for a huge diversity of
microorganisms.
1. Diversity is the precondition for evolution.
2. Genetic mutation and recombination provide a diversity of organisms, and the process of
natural selection allows the growth of those best adapted to a given environment.

84 INSTRUCTOR'S GUIDE FOR MICROBIOLOGY: AN INTRODUCTION, GE, 13e Copyright © 2021 Pearson Education Ltd.
The Loop
Generalized transduction is covered in this chapter; specialized transduction is discussed in
Chapter 13. Genetic engineering techniques (Chapter 9) and industrial microbiology
(Chapter 28) can be covered with this chapter. Antibiotics that interfere with protein synthesis
can be included (see Figure 20.4).

Exploring the Microbiome

Horizontal Gene Transfer and the Unintended Consequences of

Antibiotic Usage
Horizontal gene transfer of naturally ocurring plasmids with antibiotic-resistance genes to
pathogenic bacteria is making it harder to treat bacterial infections. Antibiotic-resistant
bacteria accumulate in the human microbiome as a person ages.
Discussion questions:
• What bacterial structures are involved in horizontal gene transfer?
• If all treatments with antibiotics carry the risk of increasing the antibiotic-resistant
fraction of our microbiome, when is it appropriate to use antibiotics?
• Research has shown that the transfer of plasmids from one bacterium to another can
happen very quickly. The following reference documents that such occurrences
happen easily on stainless steel, but less so on copper surfaces. Would it make sense
to use copper-based beds or medicine poles in medical settings.
• Further reading: Warnes, S.L. et al. (2012) Horizontal Transfer of Antibiotic
Resistance Genes on Abiotic Touch Surfaces: Implications for Public Health. mBio
3: 6 e00489-12 doi: 10.1128/mBio.00489-12.

Answers
Figure Questions
Figure Question Answer
8.1 How many times longer than the 2 µm cell 1,000x longer
is the chromosome?
8.3 What is the advantage of semiconservative Each offspring cell gets one strand of
replication? DNA from the parent cell, which
survived to reproduce.
8.4 Why is one strand “upside down” relative Carbon 1 of each sugar must face each
to the other strand? Why can’t both strands other for base pairing causing the
“face” the same way? antiparallel arrangement. The nucleotide
bases would not face each other if the
strands were parallel.
8.5 Why is one strand of DNA synthesized DNA polymerase synthesizes in one
discontinuously? direction; consequently the lagging

Copyright © 2021 Pearson Education Ltd. CHAPTER 8 Microbial Genetics 85

 strand is read from the replication fork.
Okazaki fragments are made every time
the replication form moves down.
8.6 What is the origin of replication? Where DNA polymerase binds and starts
synthesizing new strands
8.7 When does transcription stop? When the terminator sequence is
transcribed
8.8 What is the advantage of the degeneracy of Mutations and wobble (misfit of tRNA)
the genetic code? may not change the amino acid placed in
the peptide.
8.9 When does translation stop? When a stop codon is in the ribosome
8.10 Why can translation begin before transcrip- In prokaryotes, mRNA is produced by
tion is complete in prokaryotes but not in transcription and is in the cytoplasm
eukaryotes? where it is available to ribosomes. In
eukaryotes, exons must be removed from
the RNA transcript before translation.
(Transcription is in the nucleus and
translation in the cytoplasm in
eukaryotes.)
8.11 Why can’t the RNA transcript be used for Introns must be removed from the RNA
translation? transcript before translation.
8.12 What causes transcription of an inducible The presence of an inducer, such as the
enzyme? substrate for the enzyme
8.13 What causes transcription of a repressible Repressible enzymes are transcribed
enzyme? unless the repressor is activated.
8.14 When both glucose and lactose are present, Glucose can be used immediately in
why will cells use glucose first? glycolysis.
8.15 Will transcription of the lac operon occur in Lactose + glucose: no
the presence of lactose and glucose? In the Lactose alone: yes
presence of lactose and the absence of
glucose? In the presence of glucose and the Glucose alone: no
absence of lactose?
8.16 In mammals, some miRNAs hybridize with The miRNA would not stop the virus.
viral RNA. What would happen if a
mutation occurred in the miRNA gene?
8.17 Does a base substitution always result in a No, because of the degeneracy of the
different amino acid? genetic code
8.18 What happens if base 9 in (a) is changed to No change; phenylalanine is put in the
a C? growing peptide.
8.19 What is a mutagen? An environmental factor (e.g., chemical,
radiation) that causes a mutation.
8.20 Why do these drugs kill cells? They cause many mutations, one or more
86 INSTRUCTOR'S GUIDE FOR MICROBIOLOGY: AN INTRODUCTION, GE, 13e Copyright © 2021 Pearson Education Ltd.

8.21 How do excision repair enzymes “know”
which strand is incorrect?
8.22 What is an auxotroph?
8.23 Do all mutagens cause cancer?
8.24 What type of enzyme breaks the DNA?
8.25 Why are R factors important in the
treatment of infectious diseases?
8.26 Why are transposons sometimes referred to
as “jumping genes?”
8.27 Why did encapsulated bacteria kill the
mouse while nonencapsulated bacteria did
not? What killed the mouse in (d)?
8.28 What type of enzyme cuts the donor DNA?
8.29 What is an F+ cell?
8.30 Do bacteria reproduce during conjugation?
8.31 How many minutes of conjugation would
be needed to transfer genes for membrane
synthesis on this chromosome?
8.32 How would E. coli acquire the Shiga toxin
gene?
of which will be lethal.
The excision enzymes cut the non-
methylated (newly made) strand.
A microorganism having a nutritional
requirement that is absent in the parent
No.
Endonuclease
R factors carry antibiotic-resistance
genes.
They insert and excise from DNA
randomly.
Nonencapsulated bacteria were killed by
phagocytes (white blood cells). In (d),
transformed bacteria acquired the gene
for capsules.
Endonuclease and exonuclease
A cell with an F plasmid that can donate
DNA to a recipient (F –) cell.
No.
76 minutes
Transduction, transformation, and
conjugation are all possible.

Review
1. In the absence of DNA gyrase, the supercoiled DNA would fail to relax and there would
be no replication fork. In the absence of DNA ligase, a complete stretch of newly synthe-
sized daughter DNA would not form as the covalent bonds that join DNA strands, Oka-
zaki fragments, and new segments in excision repair would not exist. In the absence of
DNA polymerase, synthesis of new DNA strands will not occur. Proofreading of new
molecules of DNA and removal of mismatched bases before DNA synthesis would be
impaired, resulting in errors. In the absence of helicase, the unwinding of DNA would
fail.

Copyright © 2021 Pearson Education Ltd. CHAPTER 8 Microbial Genetics 87

 2.

3. a. 2 d. 1
b. 4 e. 5
c. 3
4. a. ATATTACTTTGCATGGACT f. Proline substituted for threonine
(missense mutation)
b. met-lys-arg-thr-(end). g. Frameshift mutation
c. TATAATGAAACGTACCTGA h. Adjacent thymines might polymerize
d. No change i. ACT
e. Cysteine substituted for arginine
5. An inducible operon has a repressor bound to its DNA and may be turned on by an in-
ducer, while a repressible operon does not have a repressor bound to its DNA and may be
turned off by the binding of a corepressor and repressor.
6. a. After translation
b. After transcription but before translation
c. Before transcription
d. Before transcription
7. Direct UV light on DNA results in the formation of pyrimidine dimers, leading to aber-
rant base pairing and errors in replication and transcription. Unless rectified, incorrect
proteins form and cause serious disease.
8. a. The F− cell gets converted into an F+ cell.
b. An Hfr cell is formed.
c. A recombinant F− cell is formed.
9. Transposons provide a natural mechanism for the movement of genes from one chromo-
some to another. They are capable of being carried between cells on plasmids or viruses,
and thus can spread from one organism or one species to another. Transposons are con-
sidered to be potential mediators of evolution in organisms.
10. Saccharomyces cerevisiae

Multiple Choice
1. a 6. b

88 INSTRUCTOR'S GUIDE FOR MICROBIOLOGY: AN INTRODUCTION, GE, 13e Copyright © 2021 Pearson Education Ltd.
 2. c 7. a
3. b 8. c
4. a 9. d
5. d 10. b

Analysis
1. Cancerous cells are growing faster than normal cells. Mutations have a greater effect
when a cell is growing because it is synthesizing DNA and enzymes. The probability of a
lethal mutation also is increased in rapidly growing cells.
2. If a DNA polymerase was defective and had poor proofreading ability, we would expect
many mismatches to occur within the DNA; the defective DNA polymerase would not be
able to excise the incorrect base and replace it with the correct one. Errors at the DNA
replication step would likely lead to further errors during protein synthesis, leading to
poor cell viability.
3. a. Mercuric ion
b. To detoxify it
c. Detoxifying mercuric ion will allow the cell to live where other organisms may not be
able to.

Clinical Applications and Evaluation

1. a. Ciprofloxacin interferes with DNA replication; erythromycin interferes with
translation; acyclovir interferes with DNA replication.
b. Erythromycin is specific for bacterial ribosomes.
c. Acyclovir will have the most effects on the host because it affects eukaryotic DNA.
The effects of erythromycin on mitochondrial ribosomes are small for short-term use.
d. Acyclovir is used against Herpes virus infections. Erythromycin affects bacterial (70S)
ribosomes, not viruses or eukaryotes.
2. Sequence B is the most dissimilar and, therefore, probably not closely related to the
others. The amino acid sequence reflects the RNA (genome) of the virus.
3. About 28% of the nucleotides are different; however, they differ in only one of the seven
amino acids. Mutations account for the difference. HHV-8 causes Kaposi’s sarcoma.

Case Study: Cystic Fibrosis and the Microbiome

Background
One of the most common genetic (inherited) diseases in America is Cystic Fibrosis (CF).
CF is a chronic disease that affects the lungs and digestive systems, for which there currently
is no cure. Every person has two copies of the CFTR (Cystic Fibrosis Transmembrane-
conductance Regulator) gene, one from each parent. For a person to have CF, both parents
must be carriers of mutated CFTR.
The CFTR gene contains the instructions for making the CFTR protein, which is produced in
many organs, among them the lungs and the pancreas. The CFTR protein creates channels in

Copyright © 2021 Pearson Education Ltd. CHAPTER 8 Microbial Genetics 89

 the cell membrane to allow the movement of chloride ions in and out of the cell. When the
CFTR protein functions properly, the balance of chloride and fluid at the cell surface remains
normal. The mutated version of the CFTR gene causes the CFTR protein to malfunction,
leading to a buildup of thick mucus, especially in the lungs. This leads to lung infections and,
eventually, respiratory failure caused by a microbial biofilm. Pseudomonas aeruginosa is
especially problematic in CF patients. Scientists have found more than 1,700 different
mutations in the CFTR gene that can cause CF, which accounts for the fact that this genetic
disease is so common.

The FDA recently approved the drug Ivacaftor for treatment of splice mutations in CFTR, but
people with these types of mutations make a small amount of normal CFTR. Ivacaftor forces
the gate on the normal CFTR protein to stay open for longer, so that the channels can
function with a reduced amount of CFTR protein in the membrane.

Questions
1. How do genes direct the production of proteins like CFTR? (Hint: central dogma)
2. When both parents carry a CF gene, will all their children have CF?
3. Explain how a nonsense mutation in CFTR gene can cause CF.
4. What would be the effect of a mutation in the intron of the CFTR gene?
90 INSTRUCTOR'S GUIDE FOR MICROBIOLOGY: AN INTRODUCTION, GE, 13e Copyright © 2021 Pearson Education Ltd.
Solution Manual for Microbiology: An Introduction, 13th Edition, Gerard J. Tortora, Berdell

5. Why does the establishment of biofilms in the lungs of CF patients complicate the CF
treatment regimen?
6. Speculate on the role of Ivacaftor in the treatment of CF.

Answers
1. The CFTR gene is transcribed into mRNA, the mRNA is translated into protein.
2. No, not necessarily. Patients have a 50% chance of receiving the mutated gene from
either parent.
3. A nonsense mutation changes an amino-acid encoding codon to a stop codon, causing
premature termination of the amino acid chain, and shorter proteins. This might result in
a nonfunctional CFTR protein.
4. A mutation in an intron might have no effect on the final protein at all if it is spliced out.
Alternatively, if it is in a border region, it might change the way the mRNA is spliced,
resulting in defective proteins.
5. Biofilms are very hard to treat with antibiotics and can cause a decrease in lung function
in already compromised CF patients.
6. Ivacaftor makes CFTR gates more efficient, so that patients get more transport out of a
smaller number of proteins.

Copyright © 2021 Pearson Education Ltd. CHAPTER 8 Microbial Genetics 91

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