GENETICS OF MOTOR NEURONE DISEASE
Motor neurone
disease (MND) or amyotrophic lateral sclerosis (ALS) is a neurodegenerative
disorder of unknown cause pathologically characterised by progressive
degeneration of motor neurones in the cortex, brain stem and spinal cord. It is
mainly sporadic, however 5-10% of MND/ALS cases are familial. In these cases,
the clinical and pathological features are indistinguishable from those in the
sporadic form of the disease. Furthermore, a number of inherited conditions,
clinically and pathologically related to classical MND/ALS, exist. Thus, much
effort has been dedicated to the identification of the genetic abnormalities
involved in motor neurone degeneration. Elucidation of the molecular basis of
motor neurone death would, in fact, provide targets for therapy. To date,
mutations leading to motor neurone degeneration have been found in:
1) the superoxide
dismutase (SOD 1) gene in chromosome 21 -linked /1 MND/ALS families
2) the
neurofilament heavy (NF-H) gene_In sporadic MND/ALS,
3) the_androgen
receptor gene in X-linked bulbospinal neuronopathy (SBMA or Kennedy's disease).
SODI gene mutations in familial MND/ALS
Familial MND/ALS,
clinically indistinguishable from sporadic MND/ALS, is expressed as an
age-dependent autosomal dominant trait. In a subset of MND/ALS families,
genetic linkage has recently been established with markers on chromosome 2
1q22. 1. In these families, SOD 1, the gene encoding the cytosolic antioxidant enzyme
Cu, Zn superoxide dismutase (SOD) has subsequently been studied as a candidate
gene because of its proximity to the familial MND/ALS locus.
Genomic DNA from
members of the chromosome 21-linked MND/ALS families and normal controls has
been amplified by the polymerase chain reaction (PCR) and examined for
single-stranded conformation polymorphism (SSCP), by changes in DNA migration
on nondenaturing polyacrylamide gels. In 30% of the chromosome 21 -linkcd
families, this approach has led to the identification of 14
point mutations
in exons 1,2,4 and 5 of the SODi gene. No changes in the sequence have been
found in exon 3 or In genomic DNA from normal individuals.
Although no
mutations have been found in the SOD 1 gene in sporadic MND/ALS, biochemical
studies have now provided evidence of increased oxidative damage to proteins
also in these cases.
Superoxide
dismutase is involved in the removal of free radicals normally produccd in
cellular metabolism. All the mutations described in familial MND/ALS code for
amino acid residues which disrupt the normal three dimensional structure of
the enzyme. Therefore, this discovery
suggests free radical toxicity as the primary pathogenic mechanism for motor
neurone death in familial MND/ALS. Studies are now being carried out to
establish the functional consequences of the gene mutations as well as the
involvement of sporadic MND/ALS
oxidative damage in the pathogenesis of familial a
A direct role of
neurofilament in the pathogeneis of
MND/ALS has been suggested by the abnormal accumulations of neurofilaments
which are one of the pathological hallmarks of MND/ALS. Furthermore, transgenic
mice models have shown that overexpression of neurofilament subunits leads to
motor neurone dysfunction. The NF-H protein includes a unique phosphorylation
domain of multiple lysine-serine-proline (KSP) repeats located in the
side-arms, appearing to modulate the spacing between neurofilaments. The liumm
NF-H sequence contains 43 KSP repeats. However, an NF-H allelic variant containing
44 KSP repeats has been identified. The distribution of the 43 and 44 NF-H
allelic variants has been examined in DNA samples from 148 control individuals
and 273 non related individuals with sporadic MND/ALS. The allelic distribution
between the two groups varies significantly. Moreover, in 3 MND/ALS patients,
mutations have been found in the phosphorviation domain of NF-H. One MND/ALS
patient has a 102 bp deletion, which includes 5 KSP repeats, while two other
MND/ALS patients have a mutant NF-H allele with Q3 bp deletion
including a lysine residue. These mutations may alter the cross-linking
properties of NF-H, therefore resulting in an impairment of neurofilament
transport.
X-_linked spinal
and bulbar muscular atro rophy or Kennedy's disease is a recessive,
adult-onset form
of lower motor neurone degeneration also associated with signs of androgen
insensitivity. The androgen receptor gene has been mapped to chromosome Xq 11 -
12, where linkage studies have localysed the SBMA gene defect. The first exon
of the gene contains a polymorphic CAG repeat, coding_for a polyglutamine
strech. The number of the CAG repeats normally varies In the population between
15 to 33. However, In Kennedy's disease patients, the number of the repeats
varies from 40 to 52. There is an absolute association of the larger
polyglutamine stretch with the disease phenotype and, furthermore, the number
of the repeats correlates with the severity of the disease. This is the only
known mutation of the androgen receptor gene associated with motor neurone
degeneration. Functional studies of the receptor carrying the expanded
polyglutamine stretch have now shown that the mutated protein exhibits reduced
transcriptional competence, The molecular mechanisms by which this may impair
motor neuron-specific survival remain to be clarified.
Conclusions
The specific
genetic lesions which, so far, have been identified in classical sporadic and
familial MND /ALS, as well as the related condition Kennedy's disease, point to
a wide range of causes underlying motor neurone degeneration. The functional
effects of these various genetic abnormalities need to be fully Investigated, in
order to identify putative targets for therapy.