ReviewCompensatory mechanisms in the aging motor system
Introduction
Motor function declines with increasing age (Jebsen et al., 1969, Potvin et al., 1980, Hackel et al., 1992, Kolb et al., 1998). The factors leading to this decline are complex and multifactorial. They include changes in musculoskeletal architecture (Lexell, 1995, Lexell, 1997, Dutta et al., 1997) peripheral and central nerve conduction (Dorfman and Bosley, 1979), proprioception (Kaplan et al., 1985), neuromuscular coupling (Lexell, 1997, Delbono, 2003). In addition, although it is often widely assumed that there is a progressive loss of cortical and striatal neurons in normal aging, the evidence is against this (Winblad et al., 1985, Sawle et al., 1990, Haug and Eggers, 1991). In particular, there is no significant age-related neural loss in primary motor cortex (Haug and Eggers, 1991). There is evidence, however, of disruption of white matter integrity with age which will affect cortico-cortical and corticospinal connectivity (Madden et al., 2004). The effects of these changes are widespread from simple motor tasks such as repetitive finger tapping (Shimoyama et al., 1990), to more demanding timed tasks and visually guided hand movements (Houx and Jolles, 1993, Kauranen and Vanharanta, 1996, Smith et al., 1999), and the somatosensory and visual control of dexterity (Cole, 1991, Cole et al., 1999).
However, aging is not just the steady loss of these component parts. With increasing age comes an increase in experience which appears to have a marked effect on the central nervous system morphology, at least in animal models, many of which are compensatory (Kolb et al., 1998). Consequently, there is increasing interest in determining whether compensatory changes can be seen in the aging human nervous system.
Section snippets
Studying the working human brain
The tools available for studying the motor system in the working human brain, in either health or disease, are different to those used in animal models. In human subjects, experiments are performed at the level of neuronal systems rather than single cells or molecules. Both approaches have something to learn from the other and it is likely that for a complete understanding of the way the brain responds to injury, both will be required.
Functional brain imaging allows organisation of the human
Studying age-related cerebral reorganisation using haemodynamic responses
D’Esposito et al. (1999) studied over 50 subjects using fMRI whilst performing a button press task in response to a visual cue. These events were temporally sparse as the authors were primarily concerned with examining the effects of age on the haemodynamic response in primary motor cortex. The first thing to note was that task-related activation was detected in primary motor cortex in only 75% of the older subjects but 100% of the younger subjects (D’Esposito et al., 1999). Furthermore, for
Are age-related changes in cerebral organisation compensatory?
None of the studies described so far have addressed the issue of whether differences in motor system organisation are compensating for some impairment elsewhere in the motor system, whether peripheral or central. An important aspect of experimental design is to record some level of performance of the task under investigation. Heuninckx et al. (2005) were more explicit in modulating task complexity in their experimental design. Subjects were asked to perform hand flexion/extension, foot
What is the relationship between age and cerebral reorganisation?
The studies described so far have looked for categorical differences between a young and an old group of subjects. Ward and Frackowiak (2003) performed a study in which they examined the nature of the relationship between task-related signal change and age. Subjects were asked to perform a repetitive isometric hand grip task with continuous visual feed back of the force produced (Ward and Frackowiak, 2003). Behavioural data suggest that linear decreases in motor performance are seen as a
Compensatory reorganisation: lessons from stroke recovery studies
There is indirect evidence to suggest that the age-related changes in motor activation patterns are functionally useful and help to maintain performance. However, imaging studies in isolation are unable to provide direct evidence of the functional relevance of such changes.
Comparison can be made with the investigation of compensatory reorganisation of the motor system after stroke. Using fMRI, increases in task-related brain activation have been demonstrated in a number of motor-related brain
Conclusions
Concerns have been expressed over the validity of using a haemodynamic measure of neural activity in an older population with increasing risks for vascular disease. Despite these concerns, experiments using fMRI have produced a convincing body of evidence to suggest that older subjects recruit a wider network of brain regions during the performance of a variety of motor tasks in an attempt to maintain parity of performance levels with their younger counterparts. The idea of task-related
Acknowledgement
The author is supported by the Wellcome Trust.
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