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V DNA V V J
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The point of crossover at the V-J junction is itself variable, generating junctional diversity. Not only are any two V and J genes capable of coming together, but also the sequence at the junction of the two genes can vary. For example, we see in gure 16.38 that the junction in the protein at amino acids 95 and 96 can be Pro-Trp, ProArg, or Pro-Pro, depending on exactly where the crossover occurred. In gure 16.39, we see the DNA after V 50 and J 4 join. This gene is now transcribed. The region between J 4 and C (the constant gene) is then removed by RNA splicing, leaving the nal messenger RNA product, which is then translated into a light chain. In this cell, the homologous region is repressed as well as both regions, a phenomenon known as allelic exclusion. Thus, this cell produces only one light chain, the V 50-J 4-C protein. Similar types of events take place in the heavy-chain gene and the light-chain gene if it has been activated. There are some differences, however ( g. 16.40). The complex in human beings has only two variable genes, with four J genes and one C gene. The heavy-chain
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Recombination V
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RAG1 RAG 2 HMG1 or HMG2 Figure 16.36 V-J joining. The variable genes are shown as blue boxes with blue arrowheads as the recombination signal sequences (the 12 spacer signal); the joining genes with their 23 spacer signals are in red. With the RAG1-RAG2 recombinase and either HMG1 or HMG2, a variable and a joining gene are brought together. Recombination results in double-strand breaks that are then repaired so that a variable and a joining gene are spliced together and the intermediate material is released. (Source: Diagram
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modeled from S. D. Fugmann, et al., The RAG proteins and V(D)J recombination: Complexes, ends, and transposition, Annual Review of Immunology, 18:495 527, 2000.)
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16. Gene Expression: Control in Eukaryotes
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Gene Expression: Control in Eukaryotes
Recombination signal sequence V DNA
J 1 2 34 5
1 2 34 5
J RAG1 RAG2 HMG1 or HMG2 3 OH
... T CT AG A
Ser 94
CCT CCC ACA GGA GGG TGT Pro 95
... ... ... T G G ACC
95 96 TGG TGG CGG CCG CCC Trp Trp Arg Pro Pro
T G G ACG ACC T G C Trp 96 Thr 97
3 OH
1. 2. 3. 4. 5. Figure 16.38
Pro Pro Pro Pro Pro
CCG CCT CCT CCT CCT
Transesterification
Variability in crossing over during V-J joining generates junctional diversity. In this case, V 41 and J 5 are shown. Amino acid codons 94 and 97 are always the same, TCT and ACG, respectively, as are the rst two bases of codon 95, CC. Depending on the exact point of crossover, ve different codon pairs can be generated. Codons for proline (Pro) are always the rst in each pair (95), but codons for tryptophan (Trp), arginine (Arg), or Pro are all possible second codons (96). Matching numbered arrows indicate crossover points for the ve possibilities. (Source: Data from E. E. Max, et al.,
Sequences of ve potential recombination sites encoded close to an immunoglobulin k constant region gene, Proceedings of the National Academy of Sciences, 76:3450 54, 1979.)
Processing
Repair V J
The mechanism of site-speci c recombination between a variable and joining gene. The RAG recombinase, with the help of HMG1 or HMG2, recognizes the recombination signal sequences (red and blue arrowheads). A nick at each signal is made, producing 3 OH ends; transesteri cation then forms hairpin loops and recombinational signals with doublestrand breaks. The hairpin loops are brought together, opened, and repaired, with some processing taking place. This creates junctional diversity, including crossover point variability and N segments (to be discussed later). The enzymes responsible for the processing are repair enzymes.
complex has about one hundred to three hundred V genes, nine J genes, and the ve C genes of the ve major types ( , , , , ). In addition, heavy-chain regions have another set of genes, called diversity (D) genes. At least ve such genes
are in the human heavy-chain complex, and they add still another variable region to the nal protein. In the heavy chain, D-J joining rst takes place, then V-DJ joining; lastly, splicing creates the nal heavy-chain product ( g. 16.41). As we pointed out earlier, the nal form of the heavychain protein in human beings has ve regions or domains CH3, CH2, hinge, CH1, and variable region (see g. 16.32). Each of the constant regions, as well as the hinge region, comes from its own exon ( g. 16.42). (The variable region, of course, comes from the extensive recombination just described: g. 16.41.) The heavy chain is thus another example of the relationship between exon structure and domain function, a topic we discussed in chapter 10. Heavy-chain structure would support the exon shuf ing view (introns early). V-J, D-J, and V-DJ joining, collectively called V(D)J joining, are the only known examples of site-speci c recombination in vertebrates.The genes responsible are active only in pre-B and pre-T cells. In addition to V(D)J joining, junctional diversity is also added during heavy-chain recombination by the addition of nucleotides in a template-free fashion. In other words, added nucleotides, called N segments, appear at the joining junctions; they are not speci ed in the DNA. For example, in one case, the sequence GTGGGGGCC (three codons long) was found at a D-J junction, but not
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