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16. Gene Expression: Control in Eukaryotes
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J 1 J 2 J 3
Transcription
V 50 J 4
C RNA
Splice out
seen in the undifferentiated (germ-line) genome. Mice that lack the gene for terminal deoxynucleotide transferase lack these N segments, implicating that gene in the process of N-segment formation. The enzyme adds nucleotides at the 3 ends of the DNA strands; these free ends are created during V(D)J joining. The enzyme is found in high levels in immature lymphocytes. There is a nal way in which variability is generated. Sequencing studies indicate that mutation occurs in variable regions after recombination has taken place. The mechanism of this speci c mutagenesis, called somatic hypermutation, is not known. Given the number of variable, constant, joining, and diversity genes, as well as the variation at the joining junctions, it is easy to see how 109 different immunoglobulin combinations could be generated (table 16.6).
V 50 J 4 C RNA Translation
Table 16.6 Three Hundred Immunoglobulin
Genes Can Generate 1.8 Billion Different Antibodies
Source Light Chains V genes
Protein
Factor
V J C Figure 16.39
40 5 10 200 5 9 100 2,000 900,000 900,000 1.8 billion 2,000
J genes V-J recombination* Heavy Chains V genes D genes J genes V-D, D-J recombination* Total
V-J joining in the human region. In this example, V 50 is joined to J 4 and then to C . First, V-J joining takes place using the heptamer-nonamer signals shown in gure 16.35. Then the region from the V 50 to the C genes is transcribed. Splicing the RNA removes the region containing the extra J gene, J 5. The nal RNA, containing V 50-J 4-C , is then translated into the light chain.
Source: Data modi ed from P. Leder, The genetics of antibody diversity, Scienti c American, 102 15, May 1982. * Junctional diversity, N-segment formation, and hypermutability.
Light V 1 V 80 J 1 J 2 J 3 J 4 J 5 C
V 1
V 2 J 1 J 2 J 3 J 4
Heavy VH1 VH200 D1 D5 JH1 JH9 C C C C C
Figure 16.40 Arrangement of the genes in the light and heavy chains of the human immunoglobulin complexes.
Tamarin: Principles of Genetics, Seventh Edition
III. Molecular Genetics
16. Gene Expression: Control in Eukaryotes
The McGraw Hill Companies, 2001
Sixteen
Gene Expression: Control in Eukaryotes
V1 DNA
V200
D1 D5
D-J joining V1 V200
Excise
D1 D3 J3
V-DJ joining V1 V50 D3 J3
Excise
V51 D2
Transcription V-D-J region V DJ RNA J9 C Exon 1 Hinge Exon 2 Exon 3 DNA
RNA processing
Hinge
S S S S
V DJ C
Heavy-chain protein
CH1 Figure 16.41
Formation of an immunoglobulin heavy chain (IgM). First, D-J joining takes place, followed by V-DJ joining. In each case, intervening DNA is spliced out by site-speci c recombination. Then, as in light-chain formation ( g. 16.39), the modi ed region is transcribed; RNA processing (splicing) then brings the nal regions together, which, when translated, form the V-D-J-C heavy chain. (Source: Data from F. W. Alt, et al., Development
VH Figure 16.42 The constant portion of heavy-chain genes is made of four domains, each transcribed from its own exon.
of the primary antibody repertoire, Science, 238:1079 87, 1987.)
T-Cell Receptors and MHC Proteins
As we mentioned earlier, genetic diversity also exists in the T-cell receptors and the major histocompatibility complex (MHC). From its function (recognizing both the antigen and the MHC self gene product), it seems evident that the T-cell receptor must show the same type of diversity that immunoglobulins have. In fact, the T-cell receptor genes are very similar to the immunoglobulin genes. T-cell receptors are composed of and subunits; there are V, J, and C components of the subunit and V, J, D, and C components of the subunits ( g. 16.43). In a sampling of T-cell receptors from one individual, approximately one million different chains and 25 different chains were found, yielding approximately 25 million (1 million 25) different T-cell receptors.
The major histocompatibility complex (MHC) region (also known as the human leukocyte antigen or HLA region in people) comprises a region of 3.6 million base pairs with 224 identi ed genetic loci. The genes are generally referred to as class I, II, and III genes. Class III genes code for proteins in the complement system, which is involved in the destruction of foreign cells. Class I and II genes code, in part, for proteins that present antigens to T cells. That is, class I and II proteins form structures with grooves on their surfaces that are shaped to hold small polypeptides. These polypeptides can be normal breakdown products of cellular metabolism in healthy cells ( self proteins) or parts of foreign invaders or their gene products in infected cells. Although similar, the two types of MHC proteins are found in different places and serve somewhat different functions. Class I MHC proteins consist of a membrane-bound chain and a second chain called 2 macroglobulin ( g. 16.44a). Class II MHC proteins consist of an and chain ( g. 16.44b). These two proteins present antigens somewhat differently. In the class I molecules, the
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