Chapter 9: Genetics of Bacteria

Student Learning Objectives

Upon completion of this chapter you should be able to:
1.
2.
3.
4.

Know the three mechanisms by which gene transfer can occur in bacteria.
Understand the key experiment that initiated gene mapping in bacteria.
Understand how gene transfer in bacteria can be used to map bacterial genes.
Recognize the significance of horizontal gene transfer.

9.1 Overview of Genetic Transfer in Bacteria

Overview
The previous chapters in this course have primarily examined the patterns of inheritance
between parents and offspring, or what is called vertical gene transfer. This chapter explores
various types of gene transfer in bacteria, all of which are examples of horizontal gene transfer.
Bacteria may transfer genetic information primarily by one of three methods: conjugation,
transduction, and transformation. Study Table 9.1 carefully as it provides a very good
introduction to the material in the rest of the chapter.

Outline of Key Terms

Genetic transfer
Conjugation
Transduction
Transformation

Focal Points

Three mechanisms of genetic transfer found in bacteria (Table 9.1)

Exercises and Problems

For questions 1 to 3, match the method of genetic transfer to its correct description.
a. transformation
b. transduction
c. conjugation
_____ 1. Donor cell physically contacts recipient cell and transfers its DNA
_____ 2. Donor cell dies releasing its DNA in the environment; recipient cell then picks it up
_____ 3. Virus infects donor cells, picks up DNA, then transfers it to recipient cell

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9.2 Bacterial Conjugation

Overview
Conjugation is the transfer of genetic information between two bacteria that are in contact
with one another. The process was discovered by Joshua Lederberg and Edward Tatum, who
analyzed auxotrophic strains of E. coli (Refer to Figure 9.1). The requirement for cell-to-cell
contact was established by Bernard Davis, in a very clever experiment employing a U-tube
apparatus (Figure 9.2). After this historical introduction, the section examines the steps of
conjugation at the molecular and cellular levels. This transfer of genetic information usually
involves the movement of small circular pieces of DNA, called plasmids (e.g. the F factor). While
bacteria do not have sexes, they are classified as donor (F+) and recipient (F-) strains, which differ
slightly in their genetic composition. The two cells make contact with each other via the sex pilus
of the donor cell. A conjugation bridge then forms, which provides a passageway for the transfer
of DNA (Refer to Figure 9.4).

Outline of Key Terms

F+ cell
F cell
Sex pilus
Conjugation bridge
Origin of transfer
Relaxosome

Minimal medium
Auxotroph
Prototroph
Conjugation
Plasmids
F factor
Episome

Focal Points

Discovery of conjugation by Lederberg and Tatum (Figure 9.1)

Genes on the F factor that play a role in conjugation (Figure 9.3)
The transfer of an F factor during bacterial conjugation (Figure 9.4)

Exercises and Problems

Complete the following sentences with the most appropriate term(s):
1. A(n) ____ is a bacterial strain that cannot make a particular nutrient which must be given to it.
2. The sex pilus of E. coli is made up of the protein ________.
3. The movement of DNA by conjugation begins at a site on the plasmid called the ________.
4. The F factor encodes a protein complex called the _________, which cuts the DNA molecule,
and catalyzes the separation of the strands, so that only the cut DNA strand is transferred.
5. Plasmids such as the F factor are ________, and so can integrate into bacterial chromosomes.
6. The result of conjugation is that the recipient cell has acquired an F factor, converting it from
a(n) ______ cell to a(n) ______.

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9.3 Conjugation and Mapping via Hfr Strains

Overview
In addition to F+ cells, there exists two other types of donor cells in E. coli: 1) Hfr strains
(High frequency of recombination), which are produced when the F factor integrates into the
bacterial chromosome; 2) F factors, which are created via an imprecise excision of the F factor
from the chromosome, thereby producing a plasmid that carries a small part of the bacterial
chromosome (Figure 9.5). Hfr strains can actually transfer portions of the bacterial chromosome
during conjugation (Figure 9.6). Indeed, this process is the basis of the interrupted mating
technique, which was developed by Elie Wollman and Franois Jacob as a way to map genes
along the E. coli chromosome (Figure 9.7). You should familiarize yourself with this classic
experiment in the history of genetics.

Outline of Key Terms

F factors
Interrupted mating
Hfr strains
Minutes

Focal Points

Formation of Hfr cell and subsequent excision of F factor (Figure 9.5)

Transfer of bacterial genes from an Hfr strain to an F cell (Figure 9.6)
Conjugation experiment used to map bacterial genes (pages 198-201)

Exercises and Problems

Complete the following sentences with the most appropriate term(s):
The F factor can integrate into the bacterial chromosome, thus creating an (1) ________
strain. If the F factor is excised properly, the cell goes back to being (2) _______. However, an
imprecise excision creates an (3) _______ plasmid, which carries some bacterial genes on it.
(4) _________ is a technique that can be used to map genes along the bacterial
chromosome. This technique allows researchers to determine the relative (5) ________ of genes
based on the (6) ______ it takes for each gene to be transferred to the recipient cell. In this type of
conjugation experiment, the recipient cell remains (7) ________ after conjugation because not all
of the (8) ________ is transferred. Because the bacterial chromosome is (9) _________, a starting
point on the map must be arbitrarily assigned. Researchers scale genetic maps from such bacterial
conjugation studies in units of (10) _________.

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9.4 Bacterial Transduction

Overview
The second mechanism of gene transfer to be discussed is transduction. This involves the
transfer of genetic material from one bacterium to another using a bacterial virus (or
bacteriophage) as an intermediate. In order to understand how this is possible, it is important to
look a little ahead to Chapter 10, Figure 10.3, which describes the reproductive cycles of
bacteriophages.
There are two types of transduction: 1) Generalized transduction involves the transfer of
any piece of the bacterial chromosome from the donor to the recipient cell (Refer to Figure 9.10);
and 2) Specialized transduction, in which only certain bacterial genes can be transferred to
recipient cells (Refer to Solved Problem S5 at the end of the chapter). A rare form of
transduction, called cotransduction, can be used to map bacterial genes that are closely linked on
the chromosome (Figure 9.11).

Outline of Key Terms

Generalized transduction
Co-transduction

Focal Points

Transduction in bacteria (Figure 9.10)

The steps in a cotransduction experiment (Figure 9.11)

Exercises and Problems

Complete the following sentences with the most appropriate term(s):
Transduction is the transfer of DNA from one bacterium to another, via a bacterial virus
or (1) ________. These viruses can undergo either the (2) ________ or the (3) ________ cycles.
In the (4) _______ cycle, when the virus infects the bacterial cell, the host DNA is (5) ________.
When new viral particles are made, any piece of the bacterial chromosome can be incorporated
into the virus and transferred to a recipient cell. This process is called (6) ________ transduction.
This is in contrast to (7) ________ transduction, in which only certain bacterial genes can be
transferred to a recipient cell. In all cases, if two genes are close together along the chromosome,
a bacterial virus may package them both into a single piece and transfer them to a recipient cell.
This phenomenon, called (8) ________, can be used to map genes that are within (9) ________ of
each other on the bacterial chromosome.

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9.5 Bacterial Transformation

Overview
The third major form of gene transfer in bacteria is transformation. During
transformation, bacteria acquire DNA from their environment (Figure 9.12). Not all bacteria are
capable of transformation. Those that possess the ability are called competent cells.
Transformation is a natural event, but has been exploited by geneticists to introduce genes of
interest into bacteria. This process is called artificial transformation. Transformation may be used
to map two genes that are closely linked by a process called cotransformation.

Outline of Key Terms

Homologous recombination
Heteroduplex
Nonhomologous recombination
Illegitimate recombination
Cotransformation

Natural transformation
Artificial transformation
Competent cells
Competence factors
Competence-stimulating peptide (CSP)
DNA-uptake signal sequence

Focal Points

The steps of bacterial transformation (Figure 9.12)

Exercises and Problems

Complete the following sentences with the most appropriate term(s):
1. A technique known as ________ uses an electric current to allow bacteria to uptake the DNA.
2. Bacterial cells that are able to take up DNA from the environment are called ________ cells.
3. During homologous recombination, alignment of foreign and host DNA results in a region
called a ________ that contains one or more base sequence mismatches.
4. In Streptococcus pneumoniae, transformation is triggered by the secretion of a short peptide
called the ________.
5. Some bacterial species, such as Haemophilus influenzae, promote the uptake of DNA among
members of their own species via ________, which are 9-10 bp long.
6. If the DNA that enters the cell is not homologous to any host gene, the DNA strand may be
incorporated at a random site in the chromosome. This process is known as
nonhomologous, or ________, recombination.
7. If two genes are close together on the same chromosome, their ________ frequency is
expected to be high.

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9.6 Horizontal Gene Transfer

Overview
Organisms can undergo two types of gene transfer. In vertical gene transfer, the genetic
information is transferred from mother cell to daughter cell, or from parent to offspring. In
contrast, horizontal gene transfer is a process in which an organism incorporates genetic material
from another organism without being the offspring of that organism. Examples include all three
mechanisms of genetic transfer that have been already discussed in this chapter. Horizontal gene
transfer has great medical importance because it can lead to acquired antibiotic resistance.

Outline of Key Terms

Vertical gene transfer
Horizontal gene transfer
Acquired antibiotic resistance

Focal Point

Acquisition of antibiotic resistance via horizontal gene transfer (page 206)

Exercises and Problems

The following statements apply to the three types of bacterial horizontal gene transfer. Indicate if
the statements are true (T) or false (F). If false, change the statement to make it true.
_____ 1. Horizontal gene transfer is the movement of DNA from parent to offspring.
_____ 2. Genetic distance obtained from conjugation studies is measured in total nucleotides.
_____ 3. Bacteria that can uptake DNA from the environment are called competent cells.
_____ 4. Artificial conjugation may be used to introduce DNA into a bacterium from its
environment using laboratory procedures.
_____ 5. The movement of genetic material by conjugation begins at a site on the plasmid
called the start site.
_____ 6. Interrupted mating is a form of transformation study used to map bacterial genes.
_____ 7. Cotransformation and cotransduction experiments can be used to map closely linked
genes in bacteria.
-

_____ 8. Both Hfr and F+ cells can transfer bacterial genes to F cells during conjugation.
_____ 9. Plasmids that can integrate into the bacterial chromosome are called episomes.
_____ 10. The two types of bacteriophage life cycles are termed lytic and lysogenic.

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Chapter Quiz
1. Which of the following mechanisms of gene transfer in bacteria involves the use of a viral
intermediate?
a. transduction
b. transformation
c. conjugation
d. co-transformation
e. none of the above
2. Which of the following processes are sensitive to extracellular DNases?
a. transduction
b. transformation
c. conjugation
d. both transduction and conjugation
e. both conjugation and transformation
3. In which of the following do bacteria have to be in physical contact for gene transfer to occur?
a. transduction
b. transformation
c. conjugation
d. both transduction and conjugation
e. both conjugation and transformation
4. Which of the following bacteria are naturally competent?
a. Streptococcus penumoniae
b. Haemophilus influenzae
c. Both A and B
d. Neither A nor B
5. Which of the following procedures are used for genetic mapping in bacteria?
a. interrupted mating
b. cotransduction
c. cotransformation
d. two of the above
e. all of the above
6. The transfer of genetic material between two different species is called _______.
a. vertical gene transfer
b. horizontal gene transfer
c. conjugation
d. sexual reproduction
7. A bacterial strain that is a methionine _________ cannot grow on a minimal medium unless
methionine the medium contains methionine.
a. autotroph
b. auxotroph
c. prototroph
d. phototroph
e. paratroph

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8. After mating with an Hfr strain, what happens to an F cell?

a. It becomes F+ and picks up some chromosomal genes
b. It becomes Hfr and picks up some chromosomal genes
c. It becomes F+ but does not pick up any chromosomal genes
d. It stays F but picks up some chromosomal genes
e. It stays F and does not pick up any chromosomal genes
9. In which type of transduction does the transducing particle carry only specific portions of the
bacterial genome?
a. general transduction
b. abortive transduction
c. specialized transduction
d. natural transduction
e. lytic transduction
10. The _________ is a protein complex that cuts the DNA during conjugation to initiate its
transfer.
a. episome
b. transformasome
c. nucleosome
d. relaxosome
e. phagosome

Answer Key for Study Guide Questions

This answer key provides the answers to the exercises and chapter quiz for this chapter. Answers
in parentheses ( ) represent possible alternate answers to a problem, while answers marked with
an asterisk (*) indicate that the response to the question may vary.
9.1

1. c
2. a
3. b

9.2

1. auxotroph
2. pilin
3. origin of transfer
4. relaxosome
5. episomes
+
6. F to F

9.3

1. Hfr
+
2. F

3. F
4. Interrupted mating
5. order

6. time
7. F
8. F factor (F plasmid)
9. circular
10. minutes

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9.4

1. bacteriophages or phage
2. lytic (or lysogenic)
3. lysogenic (or lytic)
4. lytic
5. fragmented (digested, hydrolyzed, broken up, etc.)
6. generalized
7. specialized
8. cotransduction
9. 2 minutes

9.5

1. electroporation
2. competent
3. heteroduplex
4. competence-stimulating peptide
5. DNA-uptake signal sequences
6. illegitimate
7. cotransformation

9.6

1. F, vertical gene transfer

2. F, measured in minutes
3. T
4. F, artificial transformation
5. F, called the origin of transfer
6. F, conjugation study
7. T
8. F, only Hfr cells
9. T
10. T

Quiz
1. a
2. b
3. c
4. c
5. e
6. b
7. b
8. d
9. c
10. d

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