Fredrik Ullén

We used functional magnetic resonance imaging to investigate if different brain regions are controlling the temporal and ordinal structure of movement sequences during performance. Human subjects performed overlearned spatiotemporal sequences of key‐presses using the right index finger. Under different conditions, the temporal and the ordinal structure of the sequences were varied systematically in relation to each other, using a factorial design: COMBINED had a rhythm of eight temporal intervals and a serial order of eight keys; TEMPORAL had an eight‐interval rhythm produced on one key; ORDINAL had an isochronous rhythm and an eight‐key serial order; two control conditions had an isochronous pulse performed on one or two keys, respectively. Brain regions involved in rhythmic and ordinal control of the sequences were revealed by analysing main effect contrasts for the corresponding factors. TEMPORAL and ORDINAL were also compared directly to test for significant differences. A dissociation was found between largely the presupplementary motor area, the right inferior frontal gyrus and precentral sulcus, and the bilateral superior temporal gyri, involved in temporal control, and lateral fronto‐parietal areas, the basal ganglia and the cerebellum, which were implicated in ordinal control. The vermis and the superior colliculus were the only regions with an activity increase specifically related to combining long temporal and ordinal sequences. We conclude that humans use different brain networks for temporal and ordinal sequence control, and that the performance of combined sequences activates both networks, the medial cerebellum, and the superior colliculus.

We investigated if the temporal and ordinal structures of sequences can be represented and learned independently. In Experiment 1, subjects learned three rhythmic sequences of key presses with the right index finger: Combined consisted of nine key presses with a corresponding temporal structure of eight intervals; Temporal had the temporal structure of Combined but was performed on one key; Ordinal had the ordinal structure of Combined but an isochronous rhythm. Subjects were divided into two groups. Group 1 first learned Combined, then Temporal and Ordinal; Group 2 first learned Temporal and Ordinal, then Combined. Strong transfer effects were seen in both groups. In Group 1, having learned combined facilitated the learning of the temporal ( Temporal) or ordinal ( Ordinal) sequence alone; in Group 2, having learned Temporal and Ordinal facilitated the learning of Combined, where the two are combined. This supports that subjects had formed independent temporal and ordinal representations. In Experiment 2, we investigated if these can be learned independently. Subjects repeatedly reproduced sequences with fixed temporal and random ordinal structure; random temporal and fixed ordinal structure; and random temporal and ordinal structures. Temporal and ordinal learning was seen only in the first and second sequences, respectively. In summary, we provide evidence for the existence of independent systems for learning and representation of ordinal and temporal sequences and for implicit learning of temporal sequences. This may be important for fast learning and flexibility in motor control.

Without practice, bimanual movements can typically be performed either in phase or in antiphase. Complex temporal coordination, e.g., during movements at different frequencies with a noninteger ratio (polyrhythms), requires training. Here, we investigate the organization of the neural control systems for in-phase, antiphase, and polyrhythmic coordination using functional magnetic resonance imaging (fMRI). Brisk rhythmic tapping with the index fingers was used as a model behavior. We demonstrate different patterns of brain activity during in-phase and antiphase coordination. In-phase coordination was characterized by activation of the right anterior cerebellum and cingulate motor area (CMA). Antiphase coordination was accompanied by extensive fronto-parieto-temporal activations, including the supplementary motor area (SMA), the preSMA, and the bilateral inferior parietal gyri, premotor cortex, and superior temporal gyri. When contrasting polyrhythmic tapping with in-phase tapping, activity was seen in the same set of brain regions, and in the posterior cerebellum and the CMA. Antiphase and polyrhythmic coordination may thus to a large extent use common neural control circuitry. In a separate experiment, we analyzed the neural control of the rhythmic structure and the serial order of finger movements during polyrhythmic tapping. Polyrhythmic tapping was compared with an isochronous coordination pattern that retained the same serial order of finger movements as the polyrhythm. This experiment showed that the preSMA and the bilateral superior temporal gyri may be crucial for the rhythmic control of polyrhythmic tapping, while the cerebellum, the CMA, and the premotor cortices presumably are more involved in the ordinal control of the sequence of finger movements.