Alzheimer'S Disease and Aluminium
The possible role of aluminium in thc aetiopathogenesis of Alzheimer's
disease (AD) is one of the more controversial issues in clinical neuroscience.
Recent advances in understanding the molecular pathology of this disorder and
the finding that some familial forms of presenile AD are associated with a
mutation in the b-amyloid precursor protein (APP) gene have not
resolved the problem, since many elderly cases appear to be sporadic, and the
finding of discordance in monozygotic twin pairs argues strongly for the
importance of environmental factors. While over 20 risk factors for AD have
been claimed the evidence for most of these is weak except in the case of age
and familial history. Aluminium (Al ) is known to be encephalopathic in special
circumstances, such as renal failure or following massive occupational
exposure, but there is no firm evidence implicating this metal in the aetiology
or pathogenesis of AD.
Historical Perspective
The first suggestion that Al might be involved in the pathogenesis of AD
came with the finding of extensive neurofibrillary changes in the spinal cord
and various cortical regions of rabbits injected with Al salts and with the discovery of similar
changes and cognitive deficits in cats treated in this manner. Subsequently, it
was reported that the Al content of the neocortex was significantly elevated in
AD with some patients showing increases to levels known to induce
encephalopathy in susceptible animal species. Evidence that Al was responsible
for dialysis encephalopathy, or dialysis dementia as it was also called was
considered by some to lend support to the aluminium hypothesis' of AD. However
there are fundamental problems with such an interpretation of these early data.
The neurofibrillary changes induced by Al in animals consist of straight 10 mm filaments and involve neurofilament
proteins, unlike the paired helical filaments characteristic of the human
tangle, first described by Kidd and which contain the microtubule-associated
protein tau, as a major component.
Furthermore, neurofibrillary tangles (NFTs) are not found extensively in
the spinal cord in AD, and animals treated with Al do not develop b-amyloid containing neuritic plaques.
Although an age-related increase in the Al content of the brain in the elderly
was subsequently shown, a significant increase in AD compared with age-matched
controls was not confirmed. In
addition, although the clinical features of dialysis encephalopathy include
progressive dementia there are focal neurological symptoms which readily
differentiate this condition from AD, and the majority of neuropathological
studies in this condition have failed to find senile plaques and NETs, the
hallmark lesions of AD.
More recent suggestions
that Al may be implicated in the aetiopathogenesis of AD have been based on
other lines of evidence, notably: (i) claims that Al accumulates in those brain
regions which are selectively vulnerable in AD and that the transport of Al may
be defective in this disorder. (ii) the association of Al with senile plaques
and NETs. (iii) the finding of premature b-amyloid deposition in the brains of
chronic renal dialysis patients, (iv) epidemiological studies, and (v) preliminary
evidence that treatment with the chelating agent, desferrioxamine (DFO) can
retard the rate of behavioural decline in AD.
Uptake of Aluminium by the Brain
Although aluminium is the most abundant metal in the earth's crust, it
is largely present in highly insoluble forms such as aluminosilicates which are
not bioavailable. It has no known role as an essential trace element and its
natural abundance contrasts with the small amount (30-50 mg present in the
human body, reflecting: (i) the insolubility of most naturally occurring forms
of Al, (ii) the gastrointestinal barrier to the absorption of soluble species.
and (iii) the efficiency with which the kidneys excrete Al in healthy
individuals. Dietary sources of Al include food additives, naturally occurring
forms present in sources such as tea (which is rich in Al), drinking water
(especially where aluminium is used as a flocculation agent to remove organic
residues), and relatively large amounts associated with medications such as
antacids or buffered aspirin. Little is known about the mechanisms of Al
absorption from the gastrointestinal tract, or the chemical interactions with
other dietary constituents which may affect these processes. although it is
clear that the relative amounts of Al present in various dietary sources do not
necessarily reflect their bioavailability.
Plasma concentrations of Al in normal subjects are in the range of 1-10
pg/I, and most if not all plasma Al is bound to the iron-transpoiting protein.
transferrin. There is no convenient isotope of Al, but used gallium, which
also binds with high affinity to transferrin, as a marker of Al transport in
the rat. The uptake of 67Ca into brain was found to be
unidirectional and to correspond largely with the distribution of transferrin
receptors.
Accumulation of 67Ca
was highest in regions such as the cortex, hippocampus and amygdala, areas
containing the highest densities of transferrin receptors and which are
selectively vulnerable in AD. The uptake of Al by the human brain appears to be
similar, as evidenced by a recent study in patients with chronic renal failure
who are unable to excrete Al and are regularly treated with large doses of
Al-containing compounds to prevent hyperphosphataemia. By using imaging secondary
ion mass spectrometry, the presence of numerous, focal concentrations of Al has
been shown in the brains of such patients.
This uptake appeared to reflect the distribution of pyramidal cells and
other neurones with a high requirement for iron in the synthesis of respiratory
chain enzymes, and which have high densities of transferrin receptors on the
cell surface
It appears that transferrin-mediated uptake into brain is the major
route of entry of Al, even when the gastrointestinal barrier is bypassed, since
iron-deficient rats receiving Al by intraperitoneal injection showed greater
cerebral accumulation than control animals given Al by the same route,
presumably, due to increased synthesis of brain transferrin receptors. It has
been suggested recently that the plasma binding of 67Ca (and by
implication Al) to transferrin is defective in AD and Down's syndrome, and that
this might lead to the abnormal accumulation of Al in AD. However, this study was carried out under
conditions which were markedly suboptimal for metal ion-binding to transferrin,
and it has also been criticized on other grounds. These findings need to be
replicated with a more rigorous investigation.
The observation that Al salts applied to the olfactory mucosa in
experimental animals can result in the translocation of Al into the brain
[24], and the predominant occurrence of Alzheimer-type neuropathological
changes in regions connected with the olfactory system, have led to the suggestion that the olfactory route could be
important for the entry of this agent. However, the recent report of NETs in
brain regions normally connected to the olfactory bulb in a patient with
congenital olfactory dysgenesis, who had never developed an olfactory bulb or
olfactory tracts, makes this hypothesis difficult to sustain.
Plaques
Several groups have reported the association of Al with senile plaques
and NFTs, the two diagnostic, neuropathological lesions of AD. However,
negative findings have also been reported and there is neither general
agreement that this association is consistent nor specific. Thus, Perl and
Brody have claimed that the Al content of NFT-bearing neurones in the
hippocampus in AD is significantly increased compared with that of adjacent
neurones without tangles, and Candy et al
have suggested that a focal deposit of Al in the form of aluminosilicates
is a consistent feature of the central region of the core of senile plaques.
Some groups have failed to detect Al associated with these lesions and,
although negative evidence is more difficult to interpret - especially when
standards suitable for imaging microanalysis have not been used, it can be
argued that the presence of Al does not prove the involvement of this element
in the pathogenesis of plaques and tangles. A more important issue is, rather,
whether such lesions appear prematurely in patients known to be exposed
chronically to high levels of aluminium. This question was addressed recently
in a postmortem study of the brains of patients with chronic renal failure who
did not have dialysis encephalopathy.
Increased immunostaining with an antibody to APP was found in cortical
pyramidal neurones in the majority of patients, and such changes were not
evident in age-matched control subjects. Furthermore, one third of the dialysis
patients had high densities of amorphous b-amyloid deposits in the cerebral cortex, including
patients below the age at which such changes normally occur, except in
presenile AD or Down's syndrome. Although APP is a cell stress' protein and
dialysis patients suffer from many metabolic abnormalities, similar changes
have not been found in patients with hepatic encephalopathy and it is
remarkable that intracellular staining for APP occurs in a neuronal population
which selectively accumulates Al. The data suggest that chronic accumulation
of Al in neurones may lead to increased synthesis or abnormal processing of
APP, and, in some individuals, the extracellular deposition of b-amyloid fibrils. Cortical NFTs were not
observed in any of the dialysis patients studied, suggesting that it is
unlikely that Al plays any direct role in NFT formation.
Epidemiological Studies
Normal dietary intake of Al is small (10-20 mg/day) compared with
amounts ingested as antacid medication (50-1000
mg/day) but, surprisingly, there have been few epidemiological studies which
have investigated antacid use as a risk factor for AD. Excluding studies which
were small, with low statistical power to detect such an association, one
investigation has reported negative results, and two have reported evidence of
a weak positive relationship; overall, however, it does not appear that antacid
usage is a significant risk-factor for AD. In contrast to these data, studies
from four different countries have suggested a positive association between
exposure to Al in water and AD, while one has reported negative findings. Aluminium
in drinking water accounts for far less than 5% of dietary intake of Al, even in heavily treated supplies, and
the data indicate either that Al in water is particularly bioavailable, or that
some other factor in water is responsible for this association. While there is
no strong evidence to support the view that Al in water is more bioavailable
than Al from many other sources, Birehall and Chappell have noted that a broad
inverse correlation exists between dissolved silicon (in the form of silicic
acid) and Al in water, and that a possible explanation for these epidemiological
data is that high levels of silicic acid may prevent the absorption of Al not
just from water but also from other dietary sources. Heavy industrial exposure
to Al powder may also be significant. A recent Canadian study reported that
miners exposed to finely milled Al powder to prevent silicotic lung disease
showed evidence of a dose-related cognitive decline.
Effects of Chelation Therapy
A crucial test of the “aluminium hypothesis” would be to determine
whether the removal of this metal by chelation therapy has any effect on the
progression of AD. Crapper MeLachlan have recently reported the results of such
a study, in which intramuscular injections of the trivalent ion-specific
chelator, DFO, were used to treat AD patients over a period of 24 months.
Treatment with DFO led to a significant reduction of approximately 50% in the rate of decline of daily
living skills. Although the authors point out that the therapeutic effects of
DFO could be due to mechanisms other than Al chelation, such as a reduction in
iron-mediated free radical formation, these preliminary data clearly justify
further clinical trials with DEG and other chelating agents in the treatment of
AD.
Discussion and Conclusions
Alzheimer's disease is a heterogeneous disorder and there is strong
evidence for the involvement of both genetic and environmental risk factors in
its development. While it is clear that aluminium is neurotoxic in certain
conditions, such as dialysis encephalopathy, there is no conclusive evidence
that Al is a significant risk factor in the aetiopathogenesis of AD.
Aluminium is
abundant in the environment, occurring in many different chemical forms, and
relatively little is known about the factors which determine its
bioavailability. Aluminium absorbed from the gastrointestinal tract is
largely,if not totally, bound to transferrin and slow accumulation in the
nervous system occurs via transferrin-mediated transport. Accumulation is most
marked in regions containing neurones with high densities of transferrin
receptors, including areas that are selectively vulnerable in AD.
The study in
patients exposed to high blood levels of aluminium because of chronic renal
failure suggests that intracellular accumulation of Al may lead to altered
neuronal synthesis or processing of the b-APP and, in some patients, diffuse
deposits of b-amyloid in the cerebral cortex. A mutation in the
APP gene is responsible for some forms of familial AD, suggesting that the
pathogenic sequence in AD is abnormal synthesis/metabolism of APP which leads
to b-amyloid deposition and, eventually, tangle
formation, neuronal dysfunction and cell death. Such a sequence is supported by
findings that the direct intracerebral injection of b-amyloid in the rat causes
neurodegenerative changes, including neuronal loss, degenerating neuritic
processes and the appearance of Alz-50 immunoreactive proteins which
characterise the NFT-bearmg neurone in AD. It is possible, therefore, that Al
may contribute to or accelerate this cascade in AD, but the absence of
neurofibrillary changes in renal dialysis patients indicates that Al does not
directly lead to NFT formation. A further possibility is that chronic accumulation
of Al by the brain may be partly responsible, in a less specific way, for the
attrition of neuronal populations. This could contribute to age-related memory
dysfunction and reduce the threshold for the manifestation of clinical deficits
produced by more specific disease-related processes, such as b-amyloid deposition and NET formation.
Although several
studies have shown a geographical correlation between the prevalence of AD and
the concentration of Al in water supplies, it is possible, even likely, that
this association reflects exposure to other environmental factors, such as
dissolved silicon, which may limit the bioavailability of Al from all dietary
sources.
A single study
with the chelating agent DFO in AD suggests that such treatment may help to
slow the deterioration in living skills; further therapeutic trials with
chelating agents are required.
A cautious
interpretation of the above conclusions would suggest that it maybe appropriate
to limit Al intake in foods, fluids and medicines, and also where heavy
occupational exposure is involved. However, there is as yet no direct evidence
that such steps would reduce the risk of AD or slow progression of the
disorder. Further research is required before any consensus on the value of such
measures is likely to emerge.