Saturday, 22 August 2026

Microbial Genetics

Microbial Genetics

Microbial Genetics Conjugation Transformation Transduction Gene Mapping Bacteriophage Mapping CSIR-NET GATE

1. Introduction to Microbial Genetics

Microbial genetics is the branch of genetics concerned with the study of heredity, genetic variation, gene organization, mutation, recombination, and gene transfer in microorganisms. Microorganisms, particularly bacteria and bacteriophages, have played a fundamental role in establishing many basic principles of molecular genetics.

Bacteria are especially useful for genetic studies because they have relatively small genomes, short generation times, and can be grown in very large populations. Their genetic systems also allow researchers to study recombination and gene transfer efficiently.

Unlike sexually reproducing eukaryotes, bacteria do not undergo meiosis and fertilization in the classical sense. Nevertheless, bacteria can acquire DNA from other cells through several mechanisms. The three classical mechanisms of horizontal gene transfer are:

  • Transformation – uptake of naked DNA from the environment.
  • Conjugation – DNA transfer through direct cell-to-cell contact.
  • Transduction – DNA transfer mediated by bacteriophages.

These processes can introduce new genes into bacterial cells and generate genetic recombination. They are therefore extremely important in microbial evolution and in the study of bacterial genetics.

Core concept:

Transformation = naked DNA
Conjugation = cell-to-cell contact
Transduction = bacteriophage-mediated transfer

2. Index / Table of Contents

3. Horizontal Gene Transfer

Horizontal gene transfer means the movement of genetic information between organisms other than by direct parent-to-offspring transmission. In bacteria, horizontal gene transfer is a major source of genetic variation.

Three classical mechanisms

Mechanism DNA Source Main Requirement Important Feature
Transformation Naked DNA Competent recipient DNA taken up from environment
Conjugation Donor bacterium Cell-to-cell contact Often involves F plasmid
Transduction Bacteriophage-associated DNA Phage infection Phage transfers bacterial DNA

4. Bacterial Conjugation

Conjugation is a process in which genetic material is transferred from one bacterial cell to another through direct physical contact. It is commonly associated with the F plasmid, also called the fertility factor, in Escherichia coli.

The classical F plasmid contains genes required for conjugative transfer. These genes allow the donor cell to establish contact with a recipient cell and transfer DNA.

Important terms

  • F+ cell: contains the F factor.
  • F cell: lacks the F factor.
  • F factor: fertility plasmid involved in conjugation.
  • F pilus: structure involved in initial donor-recipient contact.
  • Hfr cell: cell in which the F factor has integrated into the bacterial chromosome.
  • F′ cell: cell carrying an F plasmid that has acquired some bacterial chromosomal genes.

5. F+ × F Conjugation

The simplest conjugation experiment involves an F+ donor and an F recipient.

F+ donor
+
F recipient

Transfer of F factor

F+ recipient

During conjugation, the donor transfers a copy of the F plasmid to the recipient. At the end of the process, the recipient generally becomes F+.

The recipient does not normally become an Hfr cell simply because it receives an F plasmid. Hfr status requires integration of the F factor into the bacterial chromosome.

Exam point: F+ × F conjugation primarily transfers the F plasmid. It does not normally transfer large numbers of chromosomal genes.

6. Hfr Cells

An Hfr cell is a bacterial cell in which the F factor has become integrated into the bacterial chromosome.

Hfr stands for high frequency of recombination.

Because the F factor is integrated into the chromosome, conjugation from an Hfr donor can result in transfer of chromosomal genes into a recipient cell.

F plasmid

Integration into bacterial chromosome

Hfr cell

Chromosomal gene transfer during conjugation

Important characteristics of Hfr cells

  • The F factor is integrated into the bacterial chromosome.
  • Hfr cells can transfer bacterial chromosomal genes at high frequency.
  • Chromosomal genes are transferred in a particular order.
  • The gene closest to the origin of transfer enters first.
  • Genes farther away enter later.
  • The complete transfer of the entire chromosome is often interrupted before the F factor itself is completely transferred.
  • Therefore, the recipient usually remains F even after receiving chromosomal genes from an Hfr donor.

7. Origin and Direction of Transfer in Hfr Conjugation

The integrated F factor contains a site called oriT, or origin of transfer. DNA transfer begins at this region.

Once transfer begins, DNA enters the recipient in a linear sequence. Therefore, the order in which genes enter the recipient can be used to determine the relative order of genes on the chromosome.

oriT

Gene A

Gene B

Gene C

Gene D

If gene A enters before gene B, and gene B enters before gene C, then the relative order can be represented as:

A → B → C → D

8. Interrupted Mating and Gene Mapping

One of the most important methods for mapping bacterial genes is the interrupted mating experiment.

In this method, an Hfr donor is mixed with an F recipient. At different time points, mating pairs are disrupted mechanically, preventing further DNA transfer.

The genes that have already entered the recipient can then be identified.

Basic principle

More time required for entry
=
Gene is farther from origin of transfer

For example, suppose the following results are obtained:

Gene First detected at
lac 5 min
gal 10 min
bio 15 min

The gene order is therefore:

oriT → lac → gal → bio
Remember: The earliest entering gene is closest to the origin of transfer. The latest entering gene is farther away from the origin.

9. F′ Plasmids and Sexduction

Sometimes an integrated F factor can excise incorrectly from the bacterial chromosome. During abnormal excision, the F plasmid may carry adjacent bacterial chromosomal genes with it.

The resulting plasmid is called an F′ plasmid.

Integrated F factor

Incorrect excision

F′ plasmid carrying bacterial gene(s)

Transfer to F recipient

Partial diploid / merodiploid state

Transfer of an F′ plasmid to a recipient can create a cell containing two copies of certain genes: one chromosomal copy and one plasmid-borne copy. Such a bacterial cell is called a merodiploid or partial diploid.

This process is historically associated with the term sexduction.

10. Bacterial Transformation

Transformation is the uptake of free or naked DNA from the surrounding environment by a bacterial cell, followed by genetic change resulting from that DNA.

The recipient must possess the appropriate ability to take up DNA. Such a cell is called competent.

Donor cell

Cell lysis

DNA released into environment

Competent recipient

DNA uptake

Recombination / maintenance

New genotype

11. Competence in Transformation

Competence refers to the physiological ability of a bacterial cell to take up DNA from the environment.

Some bacteria naturally become competent under specific physiological conditions. In laboratory genetics, competence can also be artificially induced using appropriate methods.

Important points

  • Only competent cells can efficiently undergo natural transformation.
  • DNA present in the extracellular environment can be taken up.
  • The DNA may be degraded, maintained as an independent element, or integrated into the bacterial chromosome depending on the system.
  • Chromosomal transformation can generate recombinant bacteria.
  • Transformation is different from conjugation because direct cell-to-cell contact is not the defining feature.
  • Transformation is different from transduction because a bacteriophage is not required as the transfer vehicle.

12. Griffith's Transformation Experiment

One of the most famous experiments in microbial genetics was performed by Frederick Griffith using Streptococcus pneumoniae.

He studied two bacterial forms:

  • S strain: smooth colonies and associated with a capsule.
  • R strain: rough colonies and lacking the capsule characteristic of the S strain.

Griffith observed that heat-killed S bacteria could somehow cause living R bacteria to acquire characteristics associated with the S type.

This phenomenon was called transformation.

Exam memory: Griffith → Streptococcus pneumoniae → Transformation

13. Avery, MacLeod and McCarty

The molecular nature of the transforming principle was investigated by Avery, MacLeod, and McCarty.

Their experiments demonstrated that DNA was responsible for the transforming activity in the pneumococcus system.

This work provided important evidence that DNA is the genetic material.

S bacterial extract

Remove protein → transformation remains

Remove RNA → transformation remains

Destroy DNA → transformation is lost

DNA is transforming principle

14. Bacterial Transduction

Transduction is the transfer of bacterial genetic material from one bacterium to another through a bacteriophage.

Bacteriophages, or phages, are viruses that infect bacteria.

During certain phage infections, bacterial DNA can become associated with phage particles and subsequently be delivered to another bacterial cell.

Donor bacterium

Phage infection

Bacterial DNA becomes associated with phage particle

Phage infects recipient

Bacterial DNA enters recipient

Recombination

Transductant

15. Generalized Transduction

In generalized transduction, potentially any bacterial gene can be transferred by a suitable generalized transducing phage.

The process generally results from an error during phage particle assembly, where a fragment of bacterial DNA is packaged into a phage particle instead of phage DNA.

When this particle infects another bacterium, the bacterial DNA can enter the recipient cell.

Key features

  • Potentially any region of the bacterial chromosome can be transferred.
  • The phage particle carries bacterial DNA rather than a normal phage genome.
  • The transferred bacterial DNA may recombine with the recipient chromosome.
  • The transferred DNA can be used for bacterial gene mapping.
Memory: Generalized = Generally any bacterial gene.

16. Specialized Transduction

Specialized transduction differs from generalized transduction because only particular bacterial genes are transferred.

This occurs with certain temperate phages that integrate into specific sites of the bacterial chromosome as prophages.

If the prophage later excises incorrectly, it may carry neighboring bacterial genes with it.

Temperate phage

Integration into bacterial chromosome

Prophage

Incorrect excision

Phage carries adjacent bacterial genes

Specialized transduction
Memory: Generalized transduction → potentially any bacterial gene.

Specialized transduction → specific genes near the prophage integration site.

17. Generalized vs Specialized Transduction

Feature Generalized Specialized
Genes transferred Potentially any bacterial gene Specific bacterial genes
Basic cause Packaging error Incorrect prophage excision
Phage type Often lytic/generalized transducing phage Temperate phage
Location of transferred genes Can be from many chromosome regions Usually adjacent to prophage insertion site
Mapping use Cotransduction mapping Specific gene transfer studies

18. Transformation vs Conjugation vs Transduction

Feature Transformation Conjugation Transduction
DNA source Naked DNA Donor cell Bacteriophage-associated DNA
Cell contact Not required Required Not directly required
Phage required? No No Yes
Important element Competence F factor / conjugative plasmid Bacteriophage
Classic example Griffith F+ × F E. coli Phage-mediated bacterial gene transfer

19. Genetic Recombination in Bacteria

The DNA transferred through transformation, conjugation, or transduction can become incorporated into the recipient genome through homologous recombination or other genetic processes.

For homologous recombination, the incoming DNA must have sufficient sequence similarity with the recipient chromosome to allow strand pairing and exchange.

Donor DNA +
Recipient chromosome

Homologous pairing

DNA exchange

Recombinant chromosome

New phenotype

This process creates new combinations of alleles and therefore contributes to genetic variation.

20. Mapping of Genes in Bacteria

Gene mapping refers to determining the relative position and order of genes on a chromosome.

In bacterial genetics, several approaches can be used to establish gene order, including:

  • Interrupted mating.
  • Cotransduction frequency.
  • Transformation frequency.
  • Recombination frequency.

The principle is that genes that are physically close together are more likely to be transferred or recombined together than genes that are far apart.

21. Gene Mapping by Interrupted Mating

Interrupted mating is particularly useful with Hfr strains.

Suppose the following genes enter a recipient in this order:

A → B → C → D

The entry times can be used to determine gene order.

Gene Entry time
A 5 minutes
B 12 minutes
C 18 minutes
D 25 minutes

Therefore:

oriT → A → B → C → D

The differences in entry time provide an estimate of relative map distance.

22. Cotransduction and Gene Mapping

Cotransduction refers to the simultaneous transfer of two or more bacterial genes by the same transducing phage particle.

Cotransduction is useful for determining whether genes are physically close on the bacterial chromosome.

If two genes are close together, they are more likely to be packaged within the same DNA fragment and therefore transferred together.

High cotransduction frequency

Genes are relatively close

Low cotransduction frequency

Genes are relatively far apart

23. Cotransduction Frequency

The cotransduction frequency is the proportion of transductants carrying one marker that also carry another marker.

Cotransduction frequency = Number of double-marker transductants ÷ Total relevant transductants × 100

For example, if 30 out of 100 transductants carry both gene A and gene B:

Cotransduction frequency = 30 / 100 × 100 = 30%

A higher value generally indicates closer physical linkage, although the exact relationship between frequency and distance depends on the phage and mapping system.

24. Gene Mapping by Transformation

Transformation can also provide information about gene linkage.

If two bacterial genes are sufficiently close together, donor DNA fragments containing both genes can be taken up by a competent recipient.

If the two markers are frequently transformed together, they are likely to be closely linked.

Mapping principle: The closer two genes are, the more frequently they tend to be inherited together in processes involving limited DNA fragments.

25. Mapping Genes in Bacteriophages

Bacteriophages are viruses that infect bacteria. They have been extremely important in classical genetics because their genomes are relatively simple and their progeny can be studied in large numbers.

Phage genetic mapping uses recombination between different phage genomes to determine the relative position of genes.

A classical system involves infecting a bacterial cell with two genetically different phages. If both phages infect the same cell, their genomes can undergo recombination.

Phage A genome +
Phage B genome

Coinfection

Recombination

Parental + recombinant phage progeny

26. Recombination in Bacteriophages

When two genetically different phage genomes infect the same bacterial cell, genetic recombination can occur between homologous regions.

The progeny can therefore contain parental genotypes as well as recombinant genotypes.

The frequency of recombinant progeny can be used to estimate genetic distance.

Recombination frequency = Recombinant progeny ÷ Total progeny × 100

For small genetic distances, recombination frequency is often used as an approximation of map distance.

27. Two-Point Mapping in Bacteriophages

Two-point mapping determines the distance between two genes.

Suppose two phage genes are called A and B. A cross between different genotypes produces parental and recombinant progeny.

If:

Recombinants = 80
Total progeny = 1000

Then:

Recombination frequency = 80 / 1000 × 100 = 8%

The approximate genetic distance would therefore be about 8 map units for this small interval.

28. Three-Point Mapping in Bacteriophages

Three-point mapping involves three genes and allows determination of gene order as well as distances between adjacent genes.

Suppose the genes are A, B, and C. A three-point cross produces different classes of progeny.

The general procedure is:

  1. Identify the most frequent parental classes.
  2. Identify the least frequent double-crossover classes.
  3. Compare parental and double-crossover classes.
  4. Determine which gene is in the middle.
  5. Calculate recombination frequencies for each interval.
Three-point mapping rule: The gene that changes relative to the parental arrangement in the double-crossover classes is the middle gene.

29. Fine-Structure Mapping

Classical bacteriophage genetics provided evidence that a gene is not necessarily an indivisible unit of recombination.

The work of Seymour Benzer using bacteriophage T4 helped establish a detailed picture of gene structure.

Benzer's studies introduced important concepts such as:

  • The gene can contain many mutable sites.
  • Recombination can occur between very closely spaced sites.
  • Mutations can occur at different positions within the same functional gene.
  • A gene can therefore be genetically divisible at the level of recombination.

Benzer used the terms rII and related mutations in T4 bacteriophage studies to investigate fine genetic structure.

30. Cistron, Recon and Muton

Classical fine-structure genetics introduced several functional concepts.

Term Meaning
Cistron Functional unit defined by the cis-trans complementation test; broadly related to a gene or transcriptional unit in classical genetics.
Recon Classical term for the smallest unit of genetic recombination.
Muton Classical term for the smallest unit capable of mutation.

These terms are particularly important in classical genetics and bacteriophage fine-structure mapping questions.

31. Benzer's Fine-Structure Studies

Seymour Benzer's work with bacteriophage T4 provided a detailed genetic analysis of the rII region.

The rII system was especially useful because different mutations produced different plaque phenotypes on suitable bacterial hosts.

By performing recombination and complementation experiments, Benzer was able to demonstrate that genetic regions contain many distinguishable sites.

Exam clue: If a question mentions Benzer + T4 + rII, think:

Fine-structure mapping of a gene.

32. Basic Principle of Bacteriophage Mapping

The fundamental principle of phage mapping is that recombination frequency reflects genetic distance.

Two genes that are very close together have fewer opportunities for recombination between them. Genes that are farther apart have a greater probability of recombination.

Small distance → Low recombination frequency

Large distance → Higher recombination frequency

For sufficiently small intervals, 1% recombination is conventionally approximated as 1 map unit or 1 centimorgan in many classical genetic mapping contexts.

33. Important Gene-Mapping Formulas

Formula 1: Recombination frequency

RF (%) = (Number of recombinant progeny ÷ Total progeny) × 100

Formula 2: Map distance

For small intervals: Map distance ≈ Recombination frequency

Formula 3: Cotransduction frequency

Cotransduction frequency (%) = Double-marker transductants ÷ Total relevant transductants × 100

Formula 4: Relationship of linkage and distance

Closer genes → greater cotransfer
Farther genes → lower cotransfer

34. Simple Numerical Example of Phage Mapping

Suppose a phage cross produces:

  • Total progeny = 5000
  • Recombinant progeny = 250

Then:

RF = 250 / 5000 × 100

RF = 5%

Therefore, for this small interval, the approximate map distance is 5 map units.

Important: Always use the total number of progeny in the denominator when calculating simple recombination frequency.

35. Coinfection in Bacteriophage Genetics

For phage recombination studies, a bacterial cell generally needs to be infected by two genetically different phage genomes.

This is called coinfection.

The two genomes are present in the same host cell and can interact through genetic recombination.

Phage genotype 1 +
Phage genotype 2

Same bacterial cell

Recombination

Recombinant phage

36. Bacterial Gene Mapping vs Bacteriophage Gene Mapping

Feature Bacterial Mapping Phage Mapping
Organism/system Bacteria Bacteriophage
Important methods Interrupted mating, transformation, transduction Phage crosses and recombination
Mapping basis Entry time / cotransfer / recombination Recombination frequency
Special example Hfr mapping T4 rII mapping
Important researcher Classical bacterial geneticists Seymour Benzer

37. CSIR-NET / GATE High-Yield Points

  • F+ cells contain the F factor.
  • F cells lack the F factor.
  • Hfr cells contain an F factor integrated into the chromosome.
  • Hfr means high frequency of recombination.
  • In Hfr conjugation, the gene closest to oriT enters first.
  • Interrupted mating is useful for determining bacterial gene order.
  • F′ plasmids can carry bacterial chromosomal genes.
  • F′ transfer can produce a merodiploid recipient.
  • Transformation involves uptake of naked DNA.
  • Competence is required for efficient natural transformation.
  • Griffith's experiment demonstrated transformation in pneumococcus.
  • Avery, MacLeod and McCarty identified DNA as the transforming principle in their experimental system.
  • Transduction is bacteriophage-mediated DNA transfer.
  • Generalized transduction can transfer potentially any bacterial gene.
  • Specialized transduction transfers specific genes associated with the prophage integration region.
  • Cotransduction frequency can be used to estimate linkage between bacterial genes.
  • Higher cotransduction frequency generally indicates closer genes.
  • Phage gene mapping is based heavily on recombination frequency.
  • Benzer's T4 rII experiments contributed greatly to fine-structure mapping.
  • Cistron, recon, and muton are classical fine-structure genetic concepts.

38. Common Mistakes Students Make

Mistake 1: Transformation requires a pilus

Incorrect. Transformation involves uptake of extracellular DNA and does not require the F pilus used in conjugation.

Mistake 2: Transduction means direct DNA transfer between two bacteria

Incorrect. A bacteriophage acts as the transfer vehicle.

Mistake 3: All transduction is generalized

Incorrect. Transduction can be generalized or specialized.

Mistake 4: Specialized transduction can transfer any bacterial gene

Incorrect. Specialized transduction is restricted to specific bacterial genes near the phage integration site.

Mistake 5: F+ means F factor integrated into chromosome

Not necessarily. F+ usually refers to a cell carrying the F plasmid. An Hfr cell has the F factor integrated into the chromosome.

Mistake 6: Hfr recipient automatically becomes F+

Usually not. In Hfr conjugation, transfer often stops before the entire integrated F factor is transferred.

Mistake 7: Low cotransduction means genes are close

Generally the opposite: higher cotransduction frequency indicates closer physical linkage.

Mistake 8: Recombination frequency can always be converted directly to exact physical distance

Not necessarily. Recombination frequency is a genetic distance measure, not a direct measurement of physical DNA length. Multiple crossover events can complicate interpretation over larger intervals.

39. Easy Memory Tricks

Transformation

"Transformation = Take DNA"

Bacterium takes naked DNA from the environment.

Conjugation

"Conjugation = Contact"

Two bacterial cells come into direct contact.

Transduction

"Transduction = Transport by Phage"

A bacteriophage transports bacterial DNA.

Generalized Transduction

"General = Generally any gene"

Specialized Transduction

"Special = Specific genes"

Hfr

"Hfr = High frequency of recombination"

F′

"F prime = F carrying bacterial genes"

Benzer

"Benzer → T4 → rII → Fine structure"

40. One-Line Revision

Transformation → Naked DNA

Conjugation → Cell contact + F factor

Hfr → Integrated F factor

F′ → F factor carrying bacterial genes

Transduction → Phage-mediated transfer

Generalized → Any bacterial gene potentially

Specialized → Specific genes near prophage site

Interrupted mating → Gene order by entry time

Cotransduction → Linkage by simultaneous transfer

Phage mapping → Recombination frequency

Benzer → T4 rII fine-structure mapping

41. Practice MCQs – 10 Important Questions

Attempt all questions before clicking the answer buttons. These questions are designed around important CSIR-NET, GATE and postgraduate genetics concepts.

Q1. Which mechanism involves uptake of naked DNA from the environment by a bacterial cell?
Correct Answer: B. Transformation

Transformation is the uptake of extracellular or naked DNA by a competent bacterial cell.
Q2. Which factor is classically associated with bacterial conjugation in E. coli?
Correct Answer: A. F factor

The F factor is a conjugative plasmid that contains genes involved in fertility and DNA transfer.
Q3. An Hfr cell is characterized by:
Correct Answer: B. F factor integrated into the bacterial chromosome

An Hfr cell is formed when the F factor integrates into the bacterial chromosome. Hfr stands for high frequency of recombination.
Q4. In an Hfr interrupted-mating experiment, which gene usually enters the recipient first?
Correct Answer: B. The gene closest to oriT

DNA transfer begins at the origin of transfer, oriT. Therefore, genes closer to oriT enter earlier.
Q5. Generalized transduction differs from specialized transduction because generalized transduction:
Correct Answer: B. Can potentially transfer any bacterial gene

Generalized transduction results from packaging errors and can potentially transfer DNA from many different regions of the bacterial chromosome.
Q6. Specialized transduction commonly results from:
Correct Answer: A. Incorrect excision of a prophage

Specialized transduction occurs when a temperate phage excises incorrectly from its integration site and carries adjacent bacterial genes with it.
Q7. Which experimental method is particularly useful for determining bacterial gene order based on the time of entry of genes?
Correct Answer: A. Interrupted mating

Interrupted mating with Hfr cells allows researchers to determine the order in which genes enter the recipient.
Q8. A high cotransduction frequency between two genes generally indicates that the genes:
Correct Answer: B. Are relatively close together

Genes located close together are more likely to be packaged in the same transducing DNA fragment and therefore show higher cotransduction frequency.
Q9. The classical fine-structure mapping studies of the rII region were performed by:
Correct Answer: B. Seymour Benzer

Seymour Benzer used bacteriophage T4 rII mutants to investigate the fine structure of genes and recombination at very small genetic intervals.
Q10. If 100 recombinant phage progeny are obtained from a total of 2000 progeny, the recombination frequency is:
Correct Answer: B. 5%

Recombination frequency = recombinant progeny / total progeny × 100.

= 100 / 2000 × 100

= 5%

42. Quick Revision Chart

Concept Remember
Transformation Naked DNA uptake
Competence Ability to take up extracellular DNA
Conjugation Direct cell-to-cell DNA transfer
F+ Contains F factor
F Lacks F factor
Hfr F factor integrated into chromosome
F′ F factor carrying bacterial genes
Transduction Phage-mediated gene transfer
Generalized transduction Potentially any bacterial gene
Specialized transduction Specific genes near prophage integration site
Interrupted mating Gene order based on entry time
Cotransduction Linked genes transferred together
Phage mapping Recombination frequency
Benzer T4 rII fine-structure mapping
TRANSFORMATION → NAKED DNA

CONJUGATION → CONTACT

TRANSDUCTION → PHAGE

GENERALIZED → ANY GENE

SPECIALIZED → SPECIFIC GENES

HFR → HIGH FREQUENCY RECOMBINATION

COTRANSDUCTION → LINKAGE

BENZER → T4 rII → FINE STRUCTURE

43. Final Exam Strategy

For CSIR-NET, GATE, DBT, ICMR and other Life Science examinations, microbial genetics questions are frequently based on distinguishing mechanisms of DNA transfer and interpreting mapping data.

  1. If the question says naked DNA, select transformation.
  2. If the question says direct cell-to-cell contact, think conjugation.
  3. If the question mentions F factor, think conjugation.
  4. If F is integrated into the bacterial chromosome, think Hfr.
  5. If F carries bacterial chromosomal genes, think F′ plasmid.
  6. If a bacteriophage transfers bacterial DNA, think transduction.
  7. If potentially any bacterial gene can be transferred, think generalized transduction.
  8. If only specific genes near a phage integration site are transferred, think specialized transduction.
  9. If genes enter at different times during Hfr mating, use the entry times to determine gene order.
  10. If two genes show high cotransduction frequency, they are generally close together.
  11. If recombinant and total phage progeny are given, calculate recombination frequency.
  12. If T4 rII and fine-structure mapping are mentioned, remember Seymour Benzer.

ULTIMATE MICROBIAl GENETICS MEMORY

Transformation → Naked DNA

Conjugation → Cell contact + F factor

Hfr → Integrated F factor + chromosomal transfer

F′ → F plasmid carrying bacterial genes

Transduction → Bacteriophage-mediated transfer

Generalized → Potentially any bacterial gene

Specialized → Specific genes near prophage site

Interrupted mating → Entry time → gene order

Cotransduction → Higher frequency = closer genes

Phage mapping → Recombination frequency

Benzer → T4 rII → fine-structure mapping

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