Transduction is one of three basic mechanisms for genetic exchange in bacteria. Like transformation and conjugation, transduction allows the movement of genetic information from a donor cell to a recipient. Unlike the other mechanisms, however, transduction requires the participation of a type of virus called a bacteriophage in order to accomplish this movement. While transduction has been studied in the laboratory since the 1950s, more recently scientists have shown that the process also occurs in nature and probably plays an important role in the evolution of bacteria.
While transduction is common to many bacteria (but not all), the processes can be divided into two basic mechanisms. Generalized transduction tends to transfer all bacterial genes with similar frequencies, or number of cells genetically altered as a function of the total number of potential recipient cells. Specialized transduction tends to transfer only specific genes. It is the life cycle of the particular virus involved in transduction that determines which mechanism will occur, because the transduction of bacterial chromosomal genes is, in fact, the result of an error in the mechanism for viral replication. Thus, to understand the processes of transduction, one must understand the basic mechanisms of viral replication.
Generalized transduction is usually mediated by certain lytic viruses. A lytic virus is one that normally infects a host cell and redirects the resources of the host away from its own cellular replication toward viral replication and the eventual lysis, or breaking open, of the bacterial cell. Typical lytic viruses use the transcription and translation mechanisms of the host cell to express viral genes.
Viral genes are expressed in a specific order. First, the genes needed to hijack host cell metabolism are expressed. Next, the viral genetic information is copied many times. The third step involves transcription and translation of viral genes for protein components. The fourth step requires the insertion of viral DNA into the capsid, the protein shell of the virus. Finally, the virus expresses genes needed to break open the host cell, releasing hundreds of new viral particles that move on to infect new host cells.
Among the best studied of the generalized transducing viruses is P22, a virus of the bacterium Salmonella choleraesuis (subspp. typhimurium ). In the course of its normal life cycle, the virus binds to a specific receptor on the surface of the bacterial cell, injects the viral DNA into the host cell, and begins to express viral genes. The ends of the viral DNA are redundant; that is, they have small sections in which the genetic information is duplicated. The DNA repair enzymes of the host cell joins these ends so that the viral DNA forms a circle of double-stranded DNA. A DNA nuclease enzyme then makes a nick in one of the strands of DNA. The unbroken strand then serves as a template, allowing the broken strand to extend itself. This forms a long stretch of DNA that repeats the genome of the virus many times.
The long repeating sequence of DNA is known as a concatamer. The genes for viral capsid proteins (the proteins that make up the coat of the virus) are then expressed, and empty viral capsids are produced. Next, a viral DNA-cutting enzyme cuts the concatamer of DNA at a specific sequence. Once the concatamer is cut, the end of the DNA is pushed into an empty viral capsid until the viral head is full. A new empty capsid then positions itself on the end of the concatamer and the process is repeated until all the viral DNA is packaged.
Generalized transduction occurs when the enzyme that normally cuts the concatamer cuts the host chromosome instead. The viral capsids cannot distinguish viral DNA from host DNA; thus, once the initial cut is made, the empty capsids are filled with host chromosomal DNA instead of viral DNA. Since each empty capsid picks up where the last ended, it is possible for all the genes on the host chromosome to be packaged at similar rates: thus, this process is generalized. When the host cell breaks open, the viral capsids containing host chromosomal DNA, now called transducing particles, are able to bind to a new host cell. The DNA they carry is injected into the new host, but since there are no viral genes included on the injected DNA, the newly infected host cell is not killed. If the original host cell (the donor) was genetically different from the new host cell (the recipient), the DNA recombination enzymes of the recipient cell will insert the new genes in place of the old, thus altering its own genetic makeup. This genetic change is integrated into the recipient genome and is passed on to future progeny cells.
Specialized transduction results in the movement of only specific genes. The viruses that carry out specialized transduction are called lysogenic viruses. They have a mechanism for replication that is different from that of generalized transduction, for they integrate their DNA directly into the chromosome of its host's genome. Each time the host chromosome is duplicated, so is the integrated viral DNA. In many cases these viruses express genes that keep the viral DNA dormant; that is, the virus does not immediately replicate.
The best characterized model for a specialized transducing virus is phage Lambda of Escherichia coli. The Lambda virus begins its life cycle in much the same way as P22. A tail fiber on the end of the virus specifically binds to a receptor, called the maltose binding protein, on the surface of the E. coli cell. The viral DNA is then injected into the host cell. On the chromosome of the virus is a section of DNA that is almost identical to a DNA sequence found on the bacterial chromosome. The recombination enzymes of the E. coli host break and rejoin the viral DNA and host DNA together at this site, thus integrating the virus genome into the chromosome of the host bacterium.
A viral gene, the repressor gene, is then expressed and keeps the virus from activating its own replication. This integrated virus, called a prophage, can be maintained stably as a part of the host chromosome as long as the host cell remains healthy. If the host cell becomes damaged, enzymes are activated that destroy the Lambda repressor protein. Without this protein, the viral DNA will break out of the host chromosome and begin to replicate itself, much as a lytic virus does. Ultimately, the virus particles are packaged, released, and move on to infect a new host cell.
Specialized transduction occurs when the enzyme that cuts the viral DNA out of the host chromosome makes a mistake and cuts in the wrong place, removing some, but not all, of the viral genes. Since Lambda capsids fill by a "headful" mechanism, small bits of the host chromosome are packaged along with part of the viral genes. These viral particles are called defective particles because some of the viral genes are missing in the package and thus, when the virus infects a new host cell, not all the genes needed for viral replication are present. Lacking the ability to replicate, the virus cannot kill its new host. Because of this mechanism of viral packaging, specialized transducing viruses can pick up genes only on either side of the site where the virus integrates into the bacterial chromosome. Thus, while generalized transducing viruses can move any genes, specialized tranducing viruses move only specific genes.
Uses in Research
Generalized transducing viruses are the most useful in mapping bacterial chromosomal genes. Since the amount of DNA that is packaged by the virus is determined by the size of the head of the virus, each viral particle holds the same amount of DNA. The initial cutting of the host chromosome is a random event, giving all genes approximately the same probability of being packaged and transferred. Each piece of DNA that is packaged will be the same length, meaning that the closer together two genes are, the higher the probability that the two genes will be present on the same fragment of packaged DNA. In other words, the closer together the genetic markers are, the higher the frequency of cotransduction. Therefore the distance between closely linked chromosomal genes can be calculated by measuring the frequency that two genes or genetic markers are cotransduced.
When the distance between two genes is greater than the size of the viral genome, it is physically impossible for the two genes to be packaged in the same viral capsid. Thus, these genes are said to be "unlinked" with regard to viral mapping. Since most transducing viral capsids can hold only from about fifteen to fifty genes, transductional mapping of bacterial chromosomal genes is most effective for genes that are relatively close to one another.
Historically, viruses, including transducing viruses, have played an important role in defining the basic principles of molecular biology. Perhaps the most important contribution to the study of transduction was that made by Alfred Hershey and Martha Chase. During the 1940s and 1950s there was still a great deal of controversy over whether DNA or protein was the genetically inheritable material. Hershey and Chase recognized that the simplicity of the virus, consisting of DNA wrapped in a protein coat, was the ideal model to directly address the question of the basis for inheritance.
They began their experiments by growing viruses on host bacteria in media containing radioactive forms of sulfur and phosphorus. The radioactive sulfur labeled the protein components of the virus, while the radioactive phosphorus labeled the DNA portions. This allowed them to independently track the protein and DNA. After separating the radioactively labeled viruses from their host cells, they used the viruses to infect host bacteria that were not radioactively labeled. After infection, they separated the bacterial cells from the growth media. Radioactive phosphorus (viral DNA) was found inside the host cells, while the radioactive sulfur (viral proteins) was found outside the cell. This indicated that only the DNA of the virus enters the host cell, while the protein was left on the outside.
Further details of their experiment make the case for DNA even more strongly. When viruses are grown on bacteria in a thin layer on the surface of agar plates, each virus will create a clear spot called a plaque, indicating infection. Hershey and Chase had two different mutants of their virus that resulted in plaques that looked different from the normal viral infection.
When the infected bacteria were replated, normal plaques were seen, indicating that the two different new mutants had both infected the same host cell and that recombination between the virus DNA occurred within, making a virus that had repaired both mutations. Consequently, since only DNA had entered the host cells and genetic change had occurred in the viruses, DNA had to be the inheritable material. Proteins could not be the source of inheritance because the viral proteins never entered the host cells.
see also Conjugation; DNA Repair; Escherichia Coli (E. coli bacterium); Eubacteria; Mapping; Nature of the Gene, History; Recombinant DNA; Transformation; Virus.
Curtis, Helen, and N. Susan Barnes. Invitation to Biology, 5th ed. New York: WorthPublishers, 1994.
Ingraham, John, and Catherine Ingraham. Introduction to Microbiology, 2nd ed. PacificGrove, CA: Brooks/Cole Publishing, 1999.
Madigan, Michael T., John Martinko, and Jack Parker. Brock Biology of Microorganisms, 10th ed. Upper Saddle River, NJ: Prentice Hall, 2000.
Streips, Uldis N., and Ronald E. Yasbin. Modern Microbial Genetics, 2nd ed. Hoboken, NJ: John Wiley & Sons, 2002.
Transduction is defined as the transfer of genetic information between cells using a type of virus particle called a bacteriophage . The virus contains genetic material from one cell, which is introduced into the other cell upon virus infection of the second cell. Transduction does not, therefore, require cell to cell contact and is resistant to enzymes that can degrade DNA .
Bacteriophage can infect the recipient cell and commandeer the host's replication machinery to produce more copies of itself. This is referred to as the lytic cycle. Alternatively, the phage genetic material can integrate into the host DNA where it can replicate undetected along with the host until such time as an activation signal stimulates the production of new virus particles. This is referred to as the lysogenic cycle. Transduction relies on the establishment of the lysogenic cycle, with the bacterial DNA becoming incorporated into the recipient cell chromosome along with the phage DNA. This means of transferring bacteria DNA has been exploited for genetic research with bacteria like Escherichia coli, Salmonella typhimurium, and Bacillus subtilis, which are specifically targeted by certain types of bacteriophage.
There are two types of transduction: generalized transduction and specialized transduction. In generalized transduction, the packaging of bacterial DNA inside the phage particle that subsequently infects another bacterial cell occurs due to error. The error rate is about one phage particle in 1,000. Experimental mutants of phage have been engineered where the error rate is higher. Once the bacterial DNA has been injected inside the second bacterium, there is approximately a 10percent change that the DNA will be stably incorporated into the chromosome of the recipient. A successful integration changes the genotype and phenotype of the recipient, which is called a transductant. A transductant will arise for about every 106 phage particles that contain bacterial DNA.
Specialized transduction utilizes specialized phage, in which some of the phage genetic material has been replaced by other genetic material, typically the bacterial chromosome. All of the phage particles carry the same portion of the bacterial chromosome. The phage can introduce their DNA into the recipient bacterium as above or via recombination between the chromosomal DNA carried by the phage and the chromosome itself.
Transduction has proved to be a useful means of transferring genetic traits from one bacterial cell to another.
See also Bacterial ultrastructure; Bacteriophage and bacteriophage typing; Molecular biology and molecular genetics; Viral genetics; Viral vectors in gene therapy; Virus replication; Viruses and responses to viral infection
transduction, in genetics: see recombination.