Special issue: ReviewThe human cerebellum contributes to motor, emotional and cognitive associative learning. A review
Introduction
The cerebellum is known to be involved in motor coordination and learning (Thach et al., 1992, Bastian, 2006). In addition, the cerebellum appears to contribute to certain non-motor functions, including cognition, emotion and behavior (Schmahmann, 2004, Timmann and Daum, 2007, Ito, 2008). Because of its simple and homogeneous micro-circuitry the cerebellum has long been thought to perform a common computation, which serves various functions (Ito, 2006). Functional heterogeneity is explained by the input and output structure of the cerebellum. One example in support of this view is cerebellar involvement in different forms of associative learning. Human and a vast number of animal studies provide good evidence that the cerebellum is involved in classical eyeblink conditioning, a form of associative motor learning (Attwell et al., 2002, Bracha, 2004, Thompson, 2005, Gerwig et al., 2007). Furthermore, in the animal literature there is growing evidence that the cerebellum is involved in associative learning of emotional responses (Sacchetti et al., 2005). Animal data suggest that the intermediate cerebellum is involved in conditioning of specific aversive reactions (for example, eyeblink), and the medial cerebellum in conditioning of unspecific aversive reactions such as fear-related slowing of heart rate (Supple and Leaton, 1990, Bracha et al., 1999) (Fig. 1). Finally, more cognitive forms of associative learning appear to depend on the integrity of the cerebellum (Drepper et al., 1999, Timmann et al., 2002). Given that the posterolateral cerebellum is thought to contribute to cognitive function, the lateral cerebellum may play a role in higher-order associative learning. In the present paper results of human lesion studies will be compared examining associative learning in the motor, emotional and cognitive domain.
Section snippets
Motor associative learning
Cerebellar patients present with disorders in motor coordination (ataxia). Disorders in motor learning likely contribute to impaired motor capacities in daily life. The best investigated form of associative motor learning is classical eyeblink conditioning. A behaviorally neutral conditioned stimulus (CS), such as a tone, is presented and followed by an unconditioned stimulus (US), such as an air-puff that reliably elicits an unconditioned eyeblink response (UR). Repeated paired presentations
Emotional associative learning
The amygdala are the key structure in fear learning (Maren, 2005, Kim and Jung, 2006, for reviews). However, fear learning-related plastic changes take place in a more distributed network including the hippocampus and prefrontal cortex (Sacchetti et al., 2005). A current idea is that the amygdala is essential for fear conditioning to discrete cues, and the hippocampus for fear conditioning to contextual cues. The prefrontal cortex is important in extinction of fear memories. The cerebellum is
Cognitive associative learning
An increasing number of human lesion and functional brain imaging studies appear to support the hypothesis that the cerebellum contributes to a wide range of cognitive functions, including memory, language and visuospatial functions (Timmann and Daum, 2007, Baillieux et al., 2008, Ito, 2008 for recent reviews). However, there are also negative findings (Schoch et al., 2004, Richter et al., 2005, Frank et al., 2007a, Haarmeier and Thier, 2007). Overall, clinical abnormalities appear to be mild
Conclusions
Human cerebellar lesion studies provide strong evidence that the cerebellum is involved in associative learning in the motor, emotional and cognitive domain. Motor and emotional learning were assessed using classical eyeblink and fear conditioning. Classical conditioning is a method to test a type of associative learning, in which subjects learn relationships among stimuli (Dudai, 2002). More specifically, subjects learn that one stimulus (the CS) predicts the other (the US). Cognitive learning
Acknowledgements
The authors like to thank Beate Brol for help in data analysis and preparation of the figures. The study was supported by grants of the Deutsche Forschungsgemeinschaft (Ti 239/7-1; Ti 239/5-2).
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