Marit F. L. Ruitenberg

Control of automated behavior: insights from the discrete sequence production task

Work with the discrete sequence production (DSP) task has provided a substantial literature on discrete sequencing skill over the last decades. The purpose of the current article is to provide a comprehensive overview of this literature and of the theoretical progress that it has prompted. We start with a description of the DSP task and the phenomena that are typically observed with it. Then we propose a cognitive model, the dual processor model (DPM), which explains performance of (skilled) discrete key-press sequences. Key features of this model are the distinction between a cognitive processor and a motor system (i.e., motor buffer and motor processor), the interplay between these two processing systems, and the possibility to execute familiar sequences in two different execution modes. We further discuss how this model relates to several related sequence skill research paradigms and models, and we outline outstanding questions for future research throughout the paper. We conclude by sketching a tentative neural implementation of the DPM.

Motor skill learning in the middle-aged: limited development of motor chunks and explicit sequence knowledge

The present study examined whether middle-aged participants, like young adults, learn movement patterns by preparing and executing integrated sequence representations (i.e., motor chunks) that eliminate the need for external guidance of individual movements. Twenty-four middle-aged participants (aged 55–62) practiced two fixed key press sequences, one including three and one including six key presses in the discrete sequence production task. Their performance was compared with that of 24 young adults (aged 18–28). In the middle-aged participants motor chunks as well as explicit sequence knowledge appeared to be less developed than in the young adults. This held especially with respect to the unstructured 6-key sequences in which most middle-aged did not develop independence of the key-specific stimuli and learning seems to have been based on associative learning. These results are in line with the notion that sequence learning involves several mechanisms and that aging affects the relative contribution of these mechanisms.

Context-dependent motor skill and the role of practice.

Research has shown that retrieval of learned information is better when the original learning context is reinstated during testing than when this context is changed. Recently, such contextual dependencies have also been found for perceptual-motor behavior. The current study investigated the nature of context-dependent learning in the discrete sequence production task, and in addition examined whether the amount of practice affects the extent to which sequences are sensitive to contextual alterations. It was found that changing contextual cues—but not the removal of such cues—had a detrimental effect on performance. Moreover, this effect was observed only after limited practice, but not after extensive practice. Our findings support the notion of a novel type of context-dependent learning during initial motor skill acquisition and demonstrate that this context-dependence reduces with practice. It is proposed that a gradual development with practice from stimulus-driven to representation-driven sequence execution underlies this practice effect.

Context-dependent motor skill: perceptual processing in memory-based sequence production

Previous studies have shown that motor sequencing skill can benefit from the reinstatement of the learning context—even with respect to features that are formally not required for appropriate task performance. The present study explored whether such context-dependence develops when sequence execution is fully memory-based—and thus no longer assisted by stimulus–response translations. Specifically, we aimed to distinguish between preparation and execution processes. Participants performed two keying sequences in a go/no-go version of the discrete sequence production task in which the context consisted of the color in which the target keys of a particular sequence were displayed. In a subsequent test phase, these colors either were the same as during practice, were reversed for the two sequences or were novel. Results showed that, irrespective of the amount of practice, performance across all key presses in the reversed context condition was impaired relative to performance in the same and novel contexts. This suggests that the online preparation and/or execution of single key presses of the sequence is context-dependent. We propose that a cognitive processor is responsible both for these online processes and for advance sequence preparation and that combined findings from the current and previous studies build toward the notion that the cognitive processor is highly sensitive to changes in context across the various roles that it performs.

Sequential movement skill in Parkinson's disease: a state-of-the-art.

The present work reviews research on the learning and skilled performance of movement sequences in Parkinsons disease (PD). We focus specifically on the serial reaction time (SRT) task, and start by outlining behavioral studies on PD patients and healthy control participants. The literature is not unequivocal: Whereas the majority of studies indicate impaired sequencing skill in PD, still a considerable set of studies opposes this conclusion. We identify and discuss various determinants of sequence skill in PD that may contribute to the inconclusiveness of the literature. One major determinant is the role of dopaminergic medication. It has been hypothesized that while such medication restores dopamine levels in depleted parts of the brain, it may also overdose brain regions in which dopamine depletion is less pronounced. As sequence learning involves the contribution of both affected and unaffected brain areas, dopaminergic medication may enhance particular (motor-related) processes involved in sequence learning, but hinder other (cognition-related) processes that are still intact in PD. We discuss studies supporting this notion and finish with some recommendations for future research: systematically consider the impact of medication, build on models of sequence learning that include both cognitive and motor components, and include more elaborated motor skill to be able to better dissociate cognitive and motor-based problems and explore their interactions.

Sequential motor skill in preadolescent children: The development of automaticity

This study investigated to what extent preadolescent children, like young adults, learn to perform sequential movements in an automatic fashion. A sample of 24 children (mean age = 11.3 years) practiced fixed 3-key and 6-key sequences in the discrete sequence production task by responding to key-specific stimuli via spatially compatible key presses. We compared their performance with that of 24 young adults (mean age = 22.0 years). Results showed that performance improved with practice for both age groups, although children were generally slower. Compared with young adults, children had less explicit knowledge but relied more on the available explicit knowledge when executing familiar 6-key sequences. Furthermore, they completed fewer of these sequences on the basis of just the first stimulus and showed a slower transition between successive segments within the sequences. Together, these findings provide insight into the degree to which preadolescent children develop automaticity in sequential motor skill, suggesting that preadolescent children automatize the processes underlying longer movement sequences slower and/or to a lesser extent than is the case with young adults. The current study is in line with the idea that there are several mechanisms that underlie sequencing skill and suggests that the use of these mechanisms may be dependent on age.

Cognitive and neural foundations of discrete sequence skill: a TMS study.

Executing discrete movement sequences typically involves a shift with practice from a relatively slow, stimulus-based mode to a fast mode in which performance is based on retrieving and executing entire motor chunks. The dual processor model explains the performance of (skilled) discrete key-press sequences in terms of an interplay between a cognitive processor and a motor system. In the present study, we tested and confirmed the core assumptions of this model at the behavioral level. In addition, we explored the involvement of the pre-supplementary motor area (pre-SMA) in discrete sequence skill by applying inhibitory 20 min 1-Hz off-line repetitive transcranial magnetic stimulation (rTMS). Based on previous work, we predicted pre-SMA involvement in the selection/initiation of motor chunks, and this was confirmed by our results. The pre-SMA was further observed to be more involved in more complex than in simpler sequences, while no evidence was found for pre-SMA involvement in direct stimulus-response translations or associative learning processes. In conclusion, support is provided for the dual processor model, and for pre-SMA involvement in the initiation of motor chunks.

Post-error slowing in sequential action: an aging study

Previous studies demonstrated significant differences in the learning and performance of discrete movement sequences across the lifespan: Young adults (18–28 years) showed more indications for the development of (implicit) motor chunks and explicit sequence knowledge than middle-aged (55–62 years; Verwey et al., 2011) and elderly participants (75–88 years; Verwey, 2010). Still, even in the absence of indications for motor chunks, the middle-aged and elderly participants showed some performance improvement too. This was attributed to a sequence learning mechanism in which individual reactions are primed by implicit sequential knowledge. The present work further examined sequential movement skill across these age groups. We explored the consequences of making an error on the execution of a subsequent sequence, and investigated whether this is modulated by aging. To that end, we re-analyzed the data from our previous studies. Results demonstrate that sequencing performance is slowed after an error has been made in the previous sequence. Importantly, for young adults and middle-aged participants the observed slowing was also accompanied by increased accuracy after an error. We suggest that slowing in these age groups involves both functional and non-functional components, while slowing in elderly participants is non-functional. Moreover, using action sequences (instead of single key-presses) may allow to better track the effects on performance of making an error.

Evidence for graded central processing resources in a sequential movement task

In the present experiment, we examined slowing of the individual key presses of a familiar keying sequence by four different versions of a concurrent tone counting task. This was done to determine whether the same cognitive processor that has previously been assumed by the dual processor model (DPM) to initiate familiar keying sequences and assist in their execution, is involved also in the central processes of a very different task (viz. identifying tones and counting target tones). The present results confirm this hypothesis. They also suggest that in this particular situation the central processing resources underlying the cognitive processor can be distributed across the central processes of different tasks in a graded manner, rather than that they continue to behave like a single, central processor that serially switches between the central processes of the concurrently performed tasks. We argue that the production of highly practiced movement sequences can be considered automatic in the sense that execution of familiar movement sequences can continue without cognitive control once they have been initiated.

Sequence learning in Parkinson's disease : focusing on action dynamics and the role of dopaminergic medication

Previous studies on movement sequence learning in Parkinsons disease (PD) have produced mixed results. A possible explanation for the inconsistent findings is that some studies have taken dopaminergic medication into account while others have not. Additionally, in previous studies the response modalities did not allow for an investigation of the action dynamics of sequential movements as they unfold over time. In the current study we investigated sequence learning in PD by specifically considering the role of medication status in a sequence learning task where mouse movements were performed. The focus on mouse movements allowed us to examine the action dynamics of sequential movement in terms of initiation time, movement time, movement accuracy, and velocity. PD patients performed the sequence learning task once on their regular medication, and once after overnight withdrawal from their medication. Results showed that sequence learning as reflected in initiation times was impaired when PD patients performed the task ON medication compared to OFF medication. In contrast, sequence learning as reflected in the accuracy of movement trajectories was enhanced when performing the task ON compared to OFF medication. Our findings suggest that while medication enhances execution processes of movement sequence learning, it may at the same time impair planning processes that precede actual execution. Overall, the current study extends earlier findings on movement sequence learning in PD by differentiating between various components of performance, and further refines previous dopamine overdose effects in sequence learning.