Johnson M T, Coltz J D, Hagen M C, Ebner T J
Department of Neuroscience, University of Minnesota, Minneapolis, Minnesota 55455, USA.
J Neurophysiol. 1999 Feb;81(2):875-94. doi: 10.1152/jn.1999.81.2.875.
Premotor and primary motor cortical neuronal firing was studied in two monkeys during an instructed delay, pursuit tracking task. The task included a premovement "cue period," during which the target was presented at the periphery of the workspace and moved to the center of the workspace along one of eight directions at one of four constant speeds. The "track period" consisted of a visually guided, error-constrained arm movement during which the animal tracked the target as it moved from the central start box along a line to the opposite periphery of the workspace. Behaviorally, the animals tracked the required directions and speeds with highly constrained trajectories. The eye movements consisted of saccades to the target at the onset of the cue period, followed by smooth pursuit intermingled with saccades throughout the cue and track periods. Initially, an analysis of variance (ANOVA) was used to test for direction and period effects in the firing. Subsequently, a linear regression analysis was used to fit the average firing from the cue and track periods to a cosine model. Directional tuning as determined by a significant fit to the cosine model was a prominent feature of the discharge during both the cue and track periods. However, the directional tuning of the firing of a single cell was not always constant across the cue and track periods. Approximately one-half of the neurons had differences in their preferred directions (PDs) of >45 degrees between cue and track periods. The PD in the cue or track period was not dependent on the target speed. A second linear regression analysis based on calculation of the preferred direction in 20-ms bins (i.e., the PD trajectory) was used to examine on a finer time scale the temporal evolution of this change in directional tuning. The PD trajectories in the cue period were not straight but instead rotated over the workspace to align with the track period PD. Both clockwise and counterclockwise rotations occurred. The PD trajectories were relatively straight during most of the track period. The rotation and eventual convergence of the PD trajectories in the cue period to the preferred direction of the track period may reflect the transformation of visual information into motor commands. The widely dispersed PD trajectories in the cue period would allow targets to be detected over a wide spatial aperture. The convergence of the PD trajectories occurring at the cue-track transition may serve as a "Go" signal to move that was not explicitly supplied by the paradigm. Furthermore, the rotation and convergence of the PD trajectories may provide a mechanism for nonstandard mapping. Standard mapping refers to a sensorimotor transformation in which the stimulus is the object of the reach. Nonstandard mapping is the mapping of an arbitrary stimulus into an arbitrary movement. The shifts in the PD may allow relevant visual information from any direction to be transformed into an appropriate movement direction, providing a neural substrate for nonstandard stimulus-response mappings.
在一项指令延迟追踪任务中,对两只猴子的运动前区和初级运动皮层神经元放电进行了研究。该任务包括一个运动前的“提示期”,在此期间,目标出现在工作空间的周边,并以四种恒定速度之一沿八个方向之一移动到工作空间的中心。“追踪期”包括一个视觉引导、误差受限的手臂运动,在此期间,动物在目标从中央起始框沿一条线移动到工作空间的相对周边时追踪目标。从行为上来说,动物以高度受限的轨迹追踪所需的方向和速度。眼球运动包括在提示期开始时向目标的扫视,随后在整个提示期和追踪期,平稳追踪与扫视相互交织。最初,使用方差分析(ANOVA)来测试放电中的方向和时期效应。随后,使用线性回归分析将提示期和追踪期的平均放电拟合到余弦模型。通过与余弦模型的显著拟合确定的方向调谐是提示期和追踪期放电的一个突出特征。然而,单个细胞放电的方向调谐在提示期和追踪期并不总是恒定的。大约一半的神经元在提示期和追踪期之间的偏好方向(PDs)差异大于45度。提示期或追踪期的PD不依赖于目标速度。基于在20毫秒时间间隔内(即PD轨迹)计算偏好方向的第二次线性回归分析,用于在更精细的时间尺度上检查这种方向调谐变化的时间演变。提示期的PD轨迹不是直线,而是在工作空间上旋转以与追踪期的PD对齐。顺时针和逆时针旋转都有发生。追踪期的大部分时间里,PD轨迹相对较直。提示期PD轨迹向追踪期偏好方向的旋转和最终收敛可能反映了视觉信息向运动指令的转换。提示期广泛分散的PD轨迹将允许在宽空间孔径内检测目标。提示 - 追踪转换时PD轨迹的收敛可能作为一个未由该范式明确提供的“开始”运动信号。此外,PD轨迹的旋转和收敛可能为非标准映射提供一种机制。标准映射是指一种感觉运动转换,其中刺激是伸手可及的物体。非标准映射是将任意刺激映射到任意运动。PD的变化可能允许来自任何方向的相关视觉信息被转换为适当的运动方向,为非标准刺激 - 反应映射提供神经基础。