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Traditionally, the recognition of these broad categories of genetically determined diseases has rested on their pattern of occurrence in families, segregated according to mendelian inheritance into autosomal dominant, autosomal recessive, and sex-linked types Mutations of nuclear DNA account for the majority of heritable autosomal and sex-linked diseases described in this chapter, and they are remarkably diverse in nature Some are lethal and, as mentioned, are therefore not transmitted to successive generations; others are less harmful and may conform to one of the classic mendelian patterns The mutation may be large and result in duplication (or deletion) of a major part of a chromosome or even of the entire genome (diploidy), or it may create a third copy of the entire complement of genes (triploidy) Other mutations are very small, involving only a single base pair ( point mutation ) Between these two extremes are deletions or duplications that include a portion of a gene, an entire gene, or contiguous genes The factors conducive to mutations are poorly understood An increasing age of the parent is important in relation to some mutations; the size, structure, and placement of the gene on the chromosome are important in others A mutation of the DNA of a germ cell leaves unchanged the somatic phenotype of the individual in whom it occurs, but it may have a devastating effect on the descendants Conversely, a DNA mutation of a somatic cell affecting only part of the cell population may change the individual harboring it but is not passed on to the descendants Such an individual, with both normal cells and cells containing the mutant gene, is referred to as a mosaic Mutations of somatic cells appear to be most pertinent to cancer and aging In the monogenic inheritance of all three mendelian patterns, the mutation usually causes an abnormality of a single protein It may involve an enzyme, peptide hormone, immunoglobulin, collagen, or coagulation factor Such abnormalities of single genes have been isolated in several hundred diseases, but little is known of their protein products About one-quarter of these diseases are apparent soon after birth and more than 90 percent by puberty More than half of them affect more than one organ Of the 10 in every 1000 live births with genetic diseases, 7 are dominant, 25 are recessive, and the remainder are sex-linked Autosomal dominant mutations usually cause manifest disease in heterozygotes, but variations in the size of the gene abnor-.

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mality can produce any one of several phenotypes This poses a challenge to the current clinical and pathologic classi cations of disease Moreover, an identical clinical syndrome may be traced to a gene on two different chromosomes Even more surprising, an estimated 28 percent of all gene loci have polymorphic rather than monomorphic effects that is, the same mutation has several different phenotypic expressions Another problem is that of differentiating dominant from recessive inheritance In small families, in which only one descendant is af icted and the parent is seemingly normal, one may mistakenly conclude that the inheritance is recessive Variable degrees of penetrance and expressivity are characteristic features of dominant patterns of inheritance but not of recessive ones There is also a general tendency for dominantly inherited disease to rst appear long after birth Autosomal recessive forms of inherited metabolic diseases, in contrast to dominant ones, occur only in the homozygous state (both alleles abnormal) They are usually characterized by an onset soon after birth The basic abnormality in the recessively inherited diseases discussed in this chapter is more often an enzyme de ciency than an abnormality of some other protein In disorders of X-linked genes, in which the mutant gene affects mainly one sex, the female will suffer the same fate as the male if one X chromosome has been inactivated, as happens in most cells during embryonic development (the Lyon phenomenon) However, even if the abnormal X chromosome is not widely expressed, the female carrier may still exhibit minor abnormalities In the latter case, sex-linked inheritance becomes dif cult to distinguish from dominant inheritance Also, sex linkage is deceptive when a disease is lethal to one sex In contrast to autosomal recessive mutations, the biochemical abnormality has more often been one of a basic protein than an enzyme de ciency Multifactorial genetic diseases are also by nature familial They may present as constitutional disorders with gene abnormalities located on several chromosomes (polygenic, or complex genetics ) The precise number of such genes required to produce a given abnormality is not known, making the risk of inheritance dif cult to calculate Here also the relative contributions of genetic and environmental in uences are highly variable The methodology for identifying this category of genetic disease has usually required the comparison of large numbers of affected patients and controls or of progeny living with their biologic parents and those adopted by normal families at an early age Monozygotic and dizygotic twin comparisons provide essential information about risk genes The occurrence of many disorders that display high degrees of familial incidence, such as schizophrenia and Tourette syndrome, but do not strictly conform to classic genetic principles has been attributed to this type of complex genetics.

In the past few decades, an entirely new type of genetic transmission relating to the DNA that lies in the mitochondria has been elucidated Mitochondria contain their own extrachromosomal DNA, distinct from nuclear DNA Mitochondrial DNA ( the other human genome ) is a double-stranded, circular molecule that encodes the protein subunits required mainly for translation of the proteins located on the mitochondrial inner membrane Of the 37 mitochondrial genes, small in number by comparison with nuclear DNA, 13 partake in the cellular processes of oxidative phosphorylation and the production of adenosine triphosphate (ATP) A small number of genes in the cell s nucleus also code for a consid-

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