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The vehicle most commonly used in higher animals is the DNA tumor virus SV40. (SV, or simian vacuolating virus, was rst isolated in monkeys; however, it can transform normal mouse, rabbit, and hamster cells. Unlike the use of the word transformation in bacteria, transformation in eukaryotes refers to the changing of a normal cell into a rapidly growing, cancerous one.) SV40 is an icosahedral particle with a small (5,224 base pairs) chromosome, which is a circular, double-stranded DNA molecule. Like vectors, SV40 virions allow foreign DNA to replace part of their DNA. The viruses can then be used in recombinant DNA studies in one of two ways ( g. 13.17). They can replicate and complete their life cycle with the help of nonrecombinant viruses, or they can replicate in the host without making active virus particles by existing as circular plasmids in the cytoplasm or by integrating into the host s chromosomes. SV40 has become a valuable tool in mammalian genomic studies. For example,
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the rabbit -globin gene was cloned in SV40, and enhancer sequences (see chapter 10) were discovered in SV40. DNA tumor viruses have also deepened our understanding of transformation in eukaryotes (oncogenesis).
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The best-studied system for introducing foreign genes into plants is the naturally occurring crown gall tumor system. The soil bacterium Agrobacterium tumefaciens causes tumors, known as crown galls, in many dicotyledonous plants ( g. 13.18). In essence, the crown gall is made of transformed plant cells.These cells have been transformed by a plasmid within the bacterium called the tumorinducing, or Ti, plasmid. Transformation occurs when a piece of the plasmid called T-DNA (for transferred DNA) is integrated into the chromosome of the plant host. Crown gall cells produce amino acid derivatives, termed opines, that the A. tumefaciens cells use. By manipulating this
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Tamarin: Principles of Genetics, Seventh Edition
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III. Molecular Genetics
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13. Genomics, Biotechnology, and Recombinant DNA
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Genomics, Biotechnology, and Recombinant DNA
Antibiotic resistance (e.g., ampr) Bacterial origin of replication (ori)
SV40 Yeast origin of DNA replication (ARS) Centromeric region (CEN) Early genes
Protein coat DNA Remove DNA from protein coat
Origin of DNA replication Late genes SV40 DNA Replace part of SV with foreign DNA
Make linear cut (with endonuclease) and add telomeric ends
or Infect cell
Telomere
ampr ori CEN ARS
Telomere YAC
Figure 13.16 Escherichia coli plasmid pBR322 modi ed for
After infection of host, three choices for continued life cycle Add no helper SV40 Add normal helper SV40 virus 1 SV40 replicates with aid of helper virus and lyses cell or 2 Replicates as a plasmid 3 Integrates into host chromosome
use in yeast. This plasmid survives and replicates in both yeast and E. coli because it contains the origin of replication for both, as well as a yeast centromeric region (CEN). When it is made linear and telomeres are added, the yeast arti cial chromosome (YAC) becomes suitable for cloning large pieces of DNA.
system, geneticists have begun to understand the transformation process in plants as well as to develop a manipulatable system for introducing foreign genes into plants. The study of genetics in plants has been boosted a great deal by the availability of model organisms similar to E. coli, yeast, and fruit ies. Recently, much attention has focused on the meadow weed, Arabidopsis thaliana ( g. 13.19). This small plant is ideal for studying plant genetics because its genome is small, approximately 100 million base pairs located in only ve chromosomes (2n 10). This is only about ve times the genome of yeast or twenty times the genome of E. coli. Thus, in terms of genome size, it is quite manageable. A. thaliana has joined the ranks of organisms whose genomes have been sequenced. The plants are easy to grow in very large numbers, and each plant produces as many as ten thousand seeds. Hence, this organism compares very favorably with fruit ies and yeast for studying questions of gene control in a eukaryote, in this case a plant.
SV40 vehicles
Normal SV40
Figure 13.17 SV40 virus can be used as a gene cloning
vehicle. Although part of the virus is replaced by inserted DNA during cloning, it can still replicate with the aid of normal helper viruses (nonrecombinant SV40). Without the aid of helper viruses, it can either replicate as a plasmid or integrate into the host chromosome.
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