Klam F, Petit J, Grantyn A, Berthoz A
LPPA, UMR 9950 CNRS-Collège de France, Paris.
Exp Brain Res. 2001 Jul;139(2):233-47. doi: 10.1007/s002210100759.
We presented a mechanical target moving at constant velocity to awake, nontrained, head-restrained cats, in order to study how naive animals pursue objects moving at a high speed with their gaze. Eye movements were recorded while the target was moving in different directions at a constant velocity (20-80 degrees/s) through the center of the visual field. We observed two oculomotor strategies: cats either made an interception saccade (IS) toward the target but opposite to its motion, or tracked it in the direction of motion. They used the interception strategy more frequently when the gaze position error at the onset of target motion was large, and the tracking strategy when it was small. Interception was always achieved by single saccades, which were faster than tracking saccades (TS). During tracking, cats generated sequences of two to six saccades separated by "smooth" eye movements. Tracking quality varied considerably from trial to trial. When the level of motivation was high, cats would track the target at 80 degrees/s over up to 75% of the oculomotor range, with relatively small position errors. We compared ISs and TSs with respect to their metric properties and timing. The amplitudes of ISs positively correlated with position error existing 100 ms before saccade onset, but saccade vectors were directed to a point ahead of the target along the target's track. We conclude that, in programming the ISs, target motion is used to predict the future target position so as to assure a spatial lead of the gaze at the saccade end, instead of attempting a precise capture of the target. The amplitude of TSs did not depend on preceding position errors. TSs were usually small at the onset of the first saccade, as if cats would wait till the target arrived near the line of sight. A majority of primary TSs were initiated before the target arrived near the direction of gaze. Thus they had a direction, opposite to the position error sampled 100 ms before the saccade, but the same as the direction of target motion. Prediction of the future target position from its velocity vector should therefore contribute to the programming of TSs. In addition, we observed that TSs were faster when they were initiated with a spatial lag relative to the target and they were slower if there was a spatial lead or target velocity was reduced. Such a modulation appears to be analogous to the predictive correction of the saccade amplitude during smooth pursuit in primates. Considering strong visual motion sensitivity and motor properties of output neurons of the superior colliculus, it is likely that, in cats, the colliculus makes a major contribution to the integration of eye movement-related and target motion-related signals.
我们向清醒、未受过训练、头部固定的猫呈现一个匀速移动的机械目标,以研究未经训练的动物如何用目光追踪高速移动的物体。在目标以恒定速度(20 - 80度/秒)沿不同方向穿过视野中心移动时,记录其眼球运动。我们观察到两种动眼策略:猫要么朝着目标但与目标运动方向相反地进行拦截扫视(IS),要么沿运动方向追踪目标。当目标运动开始时的注视位置误差较大时,它们更频繁地使用拦截策略,而当误差较小时则使用追踪策略。拦截总是通过单个扫视完成,这些扫视比追踪扫视(TS)更快。在追踪过程中,猫会产生由“平滑”眼动分隔的两到六个扫视序列。每次试验的追踪质量差异很大。当动机水平较高时,猫能够以80度/秒的速度在高达75%的动眼范围内追踪目标,且位置误差相对较小。我们比较了IS和TS的度量特性和时间安排。IS的幅度与扫视开始前100毫秒时存在的位置误差呈正相关,但扫视向量指向目标轨迹上目标前方的一点。我们得出结论,在规划IS时,目标运动被用于预测未来目标位置,以便确保扫视结束时注视在空间上领先,而不是试图精确捕捉目标。TS的幅度不依赖于先前的位置误差。TS在第一次扫视开始时通常较小,就好像猫会等到目标靠近视线。大多数初级TS在目标到达注视方向附近之前就开始了。因此,它们的方向与扫视前100毫秒采样的位置误差方向相反,但与目标运动方向相同。因此,根据目标速度向量对未来目标位置的预测应该有助于TS的规划。此外,我们观察到,相对于目标有空间滞后时启动的TS更快,而有空间超前或目标速度降低时则较慢。这种调制似乎类似于灵长类动物平滑追踪过程中扫视幅度的预测性校正。考虑到上丘输出神经元强大的视觉运动敏感性和运动特性,很可能在猫中,上丘对与眼动相关和与目标运动相关的信号整合起主要作用。