Bapi R S, Doya K, Harner A M
Computational Neurobiology Group, Kawato Dynamic Brain Project, ERATO, JST, Kyoto, Japan.
Exp Brain Res. 2000 May;132(2):149-62. doi: 10.1007/s002219900332.
To investigate the representation of motor sequence, we tested transfer effects in a motor sequence learning paradigm. We hypothesize that there are two sequence representations, effector independent and dependent. Further, we postulate that the effector independent representation is in visual/spatial coordinates, that the effector dependent representation is in motor coordinates, and that their time courses of acquisition during learning are different. Twelve subjects were tested in a modified 2x10 task. Subjects learned to press two keys (called a set) successively on a keypad in response to two lighted squares on a 3x3 display. The complete sequence to be learned was composed of ten such sets, called a hyperset. Training was given in the normal condition and sequence recall was assessed in the early, intermediate, and late stages in three conditions, normal, visual, and motor. In the visual condition, finger-keypad mapping was rotated 90 degrees while the keypad-display mapping was kept identical to normal. In the motor condition, the keypad-display mapping was also rotated 90 degrees, resulting in an identical finger-display mapping as in normal. Subjects formed two groups with each group using a different normal condition. One group learned the sequence in a standard keypad-hand setting and subsequently recalled the sequence using a rotated keypad-hand setting in the test conditions. The second group learned the sequence with a rotated keypad-hand setting and subsequently recalled the sequence with a standard keypad-hand setting in the test conditions. Response time (RT) and sequencing errors during recall were recorded. Although subjects committed more sequencing errors in both testing conditions, visual and motor, as compared to the normal condition, the errors were below chance level. Sequencing errors did not differ significantly between visual and motor conditions. Further, the sequence recall accuracy was over 70% even by the early stage when the subjects performed the sequence for the first time with the altered conditions, visual and motor. There were parallel improvements thereafter in all the conditions. These results of positive transfer of sequence knowledge across conditions that use dissimilar finger movements point to an effector independent sequence representation, possibly in visual/spatial coordinates. Initially the RTs were similar in the visual and the motor conditions, but with training RTs in the motor condition became significantly shorter than in the visual condition, as revealed by significant interaction for the testing stage and condition term in the repeated measures ANOVA. Moreover, using RTs for single key pressing in the three conditions as baseline indices, it was again observed that RTs in the visual and motor conditions were not significantly different in the early stage, but motor RTs became significantly shorter by the late testing stage. These results support the hypothesis that the motor condition benefits more than the visual because it uses identical effector movements to the normal condition. Further, these results argue for the existence of effector dependent sequence representation, in motor coordinates, which is acquired relatively slowly. The difference in the time course of learning of these two representations may account for the differential involvement of brain areas in early and late learning phases found in lesion and imaging studies.
为了研究运动序列的表征,我们在运动序列学习范式中测试了迁移效应。我们假设存在两种序列表征,即效应器独立表征和效应器依赖表征。此外,我们假定效应器独立表征处于视觉/空间坐标中,效应器依赖表征处于运动坐标中,并且它们在学习过程中的习得时间进程不同。12名受试者参与了一项改良的2x10任务测试。受试者学习在键盘上依次按下两个按键(称为一组),以响应3x3显示屏上的两个亮起的方块。待学习的完整序列由十个这样的数据组组成,称为超集。在正常条件下进行训练,并在正常、视觉和运动三种条件下的早期、中期和后期评估序列回忆情况。在视觉条件下,手指与键盘的映射旋转了90度,而键盘与显示屏的映射保持与正常情况相同。在运动条件下,键盘与显示屏的映射也旋转了90度,从而导致手指与显示屏的映射与正常情况相同。受试者分为两组,每组使用不同的正常条件。一组在标准键盘-手部设置下学习序列,随后在测试条件下使用旋转的键盘-手部设置回忆序列。第二组在旋转的键盘-手部设置下学习序列,随后在测试条件下使用标准键盘-手部设置回忆序列。记录回忆过程中的反应时间(RT)和序列错误。尽管与正常条件相比,受试者在视觉和运动这两种测试条件下都出现了更多的序列错误,但这些错误低于随机水平。视觉和运动条件下的序列错误没有显著差异。此外,即使在早期阶段,当受试者在视觉和运动这些改变的条件下首次执行序列时,序列回忆准确率也超过了70%。此后在所有条件下都有平行的提高。这些跨不同手指运动条件的序列知识正向迁移的结果表明存在一种效应器独立的序列表征,可能处于视觉/空间坐标中。最初,视觉和运动条件下的反应时间相似,但通过重复测量方差分析中测试阶段和条件项的显著交互作用表明,经过训练后,运动条件下的反应时间明显短于视觉条件下的反应时间。此外,将三种条件下单键按下的反应时间作为基线指标,再次观察到在早期阶段视觉和运动条件下的反应时间没有显著差异,但在测试后期运动反应时间明显缩短。这些结果支持了这样的假设,即运动条件比视觉条件受益更多,因为它使用了与正常条件相同的效应器运动。此外,这些结果支持存在一种效应器依赖的序列表征,处于运动坐标中,其习得相对较慢。这两种表征在学习时间进程上的差异可能解释了在损伤和成像研究中发现的大脑区域在早期和晚期学习阶段的不同参与情况。