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mer type of neuro brillary changes simply represent an acceleration of the natural aging process in the brain We are more inclined to the idea that the plaques and neuro brillary changes represent an acquired age-linked disease, analogous in this respect to certain cerebrovascular diseases In support of this latter view are several observations First, Homo sapiens is the only animal species in which Alzheimer neuro brillary changes and neuritic plaques are regularly found in the aging brain A few plaque-like structures (but no neuro brillary changes) have been seen occasionally in old dogs and monkeys but not in mice or rats It seems unbiologic that human aging should differ from that of all other animal species Second, some of the most severe forms of Alzheimer disease occur in middle adult life, long before the senium Third, these histopathologic changes in variable proportion occur in a number of other diseases unrelated to aging, such as dementia pugilistica ( punch-drunk state), Down syndrome, postencephalitic Parkinson disease, and progressive supranuclear palsy Fourth, neuro brillary tangles can be reproduced in the experimental animal by such toxins as aluminum, vincristine, vinblastine, and colchicine Finally, a small proportion of Alzheimer cases are de nitely familial Alzheimer disease is therefore more appropriately considered with degenerative diseases (see Chap 39, where the entire subject is discussed further) Virtually every molecular structure within the cell is subject to age-related biochemical modi cations, such as the formation of carbonyl proteins, glycation of sugars, and oxidative changes in lipids Some of these subcellular phenomena contribute to the aging process (see Smith CD et al and Mrak et al for details), as do the cumulation of mitochondrial DNA mutations and shortened lengths of the telomeres Among the visible biochemical alterations is an increasing accumulation of lipofuscin granules in the cytoplasm of neurons, sometimes extreme in degree Also, there is an age-related neuronal accumulation of iron and other pigment bodies Granulovacuolar changes are a regular nding in aging hippocampi, regardless of the mental state of the individual The accumulation of amyloid-containing concretions (corpora amylaceae) around nerve roots and diffusely in the subpial space is yet another aging effect Cerebral atherosclerosis is, of course, a frequent nding in the elderly, but it does not parallel aging with any degree of precision, being severe in some 30- to 40-year-old individuals and practically absent in some octogenarians In the normotensive individual, it tends to occur in scattered, discrete plaques mostly in the aorta and cervical arteries (carotid bifurcation and higher segments), proximal middle cerebral arteries, and at the vertebrobasilar junction and basilar portions of the cerebral arterial system In the hypertensive and diabetic, it is more diffuse and extends into ner branches of the cerebral and cerebellar arteries One or more cerebral ischemic softenings (infarcts) are found in about 25 percent of all individuals more than 70 years of age who have been carefully examined postmortem In addition to atherosclerotic disease, the basilar arteries become somewhat larger and more tortuous and opaque in the elderly Cerebral blood ow has been extensively investigated in the elderly population Most studies have shown that ow declines with age and that the cerebral metabolic rate declines in parallel There is also an age-related increase in cerebrovascular resistance Declines in ow are somewhat greater in the cortex than in white matter and greater in prefrontal regions than in other parts of the hemispheres Obrist demonstrated a 28 percent reduction in cerebral ow by age 80 It is noteworthy, however, that every cohort.

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of elderly persons tested in this way contained a signi cant proportion in which cerebral blood ow was equivalent to that in young control subjects In fact, in a group of 72-year-old men rigorously selected on the basis of freedom from disease, Sokoloff demonstrated that cerebral blood ow and oxygen consumption did not differ from those of normal men 22 years of age Cerebral glucose metabolism was reduced in all the elderly subjects, however With advancing age there is a general tendency for the electroencephalogram (EEG) to show a slowing of the alpha rhythm, an increase in beta activity, a decline in the percentage of slowwave sleep, and an increasing intrusion of theta rhythms, particularly over the temporal lobes There are large individual differences, however With respect to the neurotransmitters, it is generally agreed that the concentrations of acetylcholine, norepinephrine, and dopamine decline in the course of normal aging Also, the concentration of gamma-aminobutyric acid (GABA) has been shown to decline with age, particularly in the frontal cortex (Spokes et al) Analyses of postmortem human and animal brains have failed to demonstrate a decline with age in the concentration of serotonin or its metabolites (McEntee and Crook) Accurate assessment of other neurotransmitters has been more dif cult because of their marked lability in postmortem material Data from experiments in rats suggest that the glutamate content of the brain and the number of Nmethyl-D-aspartate (NMDA) receptors diminish with age, but the functional signi cance of this nding is unclear Unlike the case in Alzheimer disease, normal aging is associated with only slight and inconsistent abnormalities of cholinergic innervation of the hippocampus and cortex This is true also of the acetylcholine content and the activity of choline acetyltransferase (the synthesizing enzyme of acetylcholine) in these regions and the number of cholinergic neurons in the nucleus basalis of Meynert (substantia innominata) and other nuclei of the basal forebrain (Decker) Again, the signi cance of these changes is dif cult to judge They probably re ect the depletion of cells that occurs with aging The topics of cholinergic and glutamatergic function in the aging brain have been critically reviewed by McEntee and Crook.

Summary

With advancing age, skeletal muscles lose cells ( bers) and undergo a gradual reduction in their weight more or less parallel to that of the brain Atrophy of muscles and diminution in peak power and endurance are clinical expressions of these changes Many processes contribute to this age-dependent loss of lean muscle mass, described as sarcopenia These include decreased physical activity; diminished appetite associated with loss of smell and elevated levels of cholcystokinin, a satiety hormone; other endocrine changes such as diminished levels of growth hormone and androgens; and (as in the brain) the accumulation of subcellular defects such as nuclear and mitochondrial DNA mutations alluded to earlier Moreover, with aging, the slow loss of motor neurons contributes to a component of denervation atrophy Our own observations of neuropathologic material indicate that the wasting involves several processes, some principally myopathic and others relating to disuse or denervation, from loss of motor neurons In this material, denervation atrophy of the gastrocnemius muscles was found in 80 percent of individuals over 70 years of age The lost muscle bers are gradually replaced by endomysial connective tissue and fat cells The surviving bers are generally thinner than normal (pos-

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sibly due to disuse atrophy), but some enlarge, resulting in a wider than normal range of ber size Groups of bers all at the same stage of atrophy undoubtedly relate to loss of motor innervation The reduction in conduction velocity and decrease in amplitude of motor nerve potentials and, to a greater extent, of sensory nerves in the aged may be taken as other indices of loss of motor and sensory axons All these changes are more marked in the legs than elsewhere However, when Roos and colleagues examined the contractile speed and ring rates of the quadriceps muscle in young men and compared them to those of men close to 80 years old, they found little difference despite a 50 percent reduction in the maximum voluntary contraction force developed by the muscle in the older men It has been repeatedly observed that age is an important prognostic factor in a large number of human diseases This effect is very evident, for example, in the markedly slower and less complete recovery from Guillain-Barre polyneuropathy in older age groups compared with younger ones One presumes that the structural changes of aging in peripheral nerves limit the degree of myelin regeneration and lower the threshold for failure of electrical transmission

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