Neuropsychologia 46 (2008) 521–539
Better or worse than expected? Aging, learning, and the ERN
Ben Eppinger
∗
, Jutta Kray, Barbara Mock, Axel Mecklinger
Department of Psychology, Saarland University, Campus Building A1 3, D-66041 Saarbr¨ ucken, Germany
Received 20 February 2007; received in revised form 23 August 2007; accepted 4 September 2007
Available online 7 September 2007
Abstract
This study examined age differences in error processing and reinforcement learning. We were interested in whether the electrophysiological
correlates of error processing, the error-related negativity (ERN) and the feedback-related negativity (FRN), reflect learning-related changes in
younger and older adults. To do so, we applied a probabilistic learning task in which we manipulated the validity of feedback. The results of our
study showed that learning-related changes were much more pronounced (a) in a response-locked positivity for correct trials compared to the
ERN and (b) in a feedback-locked positivity for positive feedback compared to the FRN. These findings provide an important extension to recent
theoretical accounts [Holroyd, C. B., & Coles, M. G. H. (2002). The neural basis of human error processing: Reinforcement learning, dopamine,
and the error-related negativity. Psychological Review, 109, 679–709; Nieuwenhuis, S., Ridderinkhof, K. R., Talsma, D., Coles, M. G. H., Holroyd,
C. B., Kok, A., et al. (2002). A computational account of altered error processing in older age: Dopamine and the error-related negativity. Cognitive,
Affective and Behavioral Neuroscience, 2, 19–36] since they suggest that positive learning signals on correct trials contribute to the reward-related
variance in the response- and feedback-locked ERPs. This effect has been overlooked in previous studies that have focused on the role of errors and
negative feedback for learning. Importantly, we did not find evidence for an age-related reduction of the ERN, when controlling for performance
differences between age groups, which questions the view that older adults are generally impaired in error processing. Finally, we observed a
substantial reduction of the FRN in the elderly, which indicates that older adults are less affected by negative feedback and rely more on positive
feedback during learning. This finding points to an age-related asymmetry in the processing of feedback valence.
© 2007 Elsevier Ltd. All rights reserved.
Keywords: Age differences; ACC; Dopamine; ERP; Error processing; Reinforcement learning
1. Introduction
The aim of this study is to examine age-related modulations in
reinforcement learning and their potential impact on the ability
to adaptively acquire and maintain new behavior. We focus on
the role that the processing of error information plays for rein-
forcement learning by investigating age differences in the behav-
ioral and electrophysiological correlates of error processing.
The basic principle of reinforcement learning has already
been formulated in the early 1920s by Thorndike. It suggests
that actions that are followed by feelings of satisfaction are more
likely to be generated again in the future, whereas actions that
are followed by negative outcomes are less likely to reoccur
(Thorndike, 1911). In recent years several researchers became
interested in the neurophysiological basis of reinforcement
∗
Corresponding author. Tel.: +49 681 3023698; fax: +49 681 3023871.
E-mail address: eppinger@mx.uni-saarland.de (B. Eppinger).
learning and especially in the role of dopamine for learn-
ing. For instance, electrophysiological studies in primates have
shown that learning based on rewarding outcomes depends
on the activity of dopaminergic neurons in the ventro-anterior
midbrain (substantia nigra and ventral tegmental area) (for
reviews, see Montague, Hyman, & Cohen, 2004; Schultz, 2002).
Schultz, Dayan, and Montague (1997) integrated these electro-
physiological findings in primates with learning models from
artificial intelligence. They showed that changes in the activity
of dopaminergic neurons during learning could be formalized
using a temporal difference learning model (see Barto & Sutton,
1997). Dopaminergic neurons from the ventral tegmental area
seem to signal the extent to which a rewarding outcome deviates
from a prediction during learning. That is, they code prediction
errors that reflect changes in the value of ongoing events, when
events are suddenly better or worse than expected (Schultz et
al., 1997). According to this model, learning is induced when a
reinforcer occurs that is better than predicted (positive prediction
error). In contrast, a reinforcer that is worse than predicted or
0028-3932/$ – see front matter © 2007 Elsevier Ltd. All rights reserved.
doi:10.1016/j.neuropsychologia.2007.09.001