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on a given chromosome, the more likely it is that genetic exchanges will take place.

For example, recombination between HLA-A and HLA-DP occurs commonly, whereas recombination between HLA-DQ and HLA-DR is a rare event. Crossing over has the effect of rearranging the genes on the chromosome to produce new haplotypes in the general population.

Linkage Disequilibrium
An important characteristic of HLA antigens is the existence of linkage disequilibrium between the alleles of the loci. Linkage disequilibrium is the occurrence of HLA genes more frequently in the same haplotype than would be expected by chance alone. In the Rh system, the blood groups C, D, and e are found together more often than would be expected based on their individual gene frequencies. This is commonly found within the HLA system. In a randomly mating population at Hardy-Weinberg equilibrium,25 the occurrence of two alleles from closely linked genes will be the product of their individual gene frequencies. If the observed value of the joint frequency is significantly different from the expected frequency (the product of the individual allele frequencies), the alleles are said to be in linkage disequilibrium. For example, if HLAA1 and HLA-B8 gene frequencies are 0.16 and 0.1, respectively, in a population, the expected occurrence of an HLA haplotype bearing both A1 and B8 should be 0.16 _ 0.1 _ 100, or 1.6 percent. In certain white populations, however, the actual occurrence of this haplotype is as

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