Kornylo Krista, Dill Natalie, Saenz Melissa, Krauzlis Richard J
Systems Neurobiology Laboratory, Salk Institute for Biological Studies, La Jolla, California 92037, USA.
J Neurophysiol. 2003 Jun;89(6):2984-99. doi: 10.1152/jn.00859.2002.
The countermanding paradigm provides a useful tool for examining the mechanisms responsible for cancelling eye movements. The key feature of this paradigm is that, on a minority of trials, a stop signal is introduced some time after the appearance of the target, indicating that the subject should cancel the incipient eye movement. If the delay in giving the stop signal is too long, subjects fail to cancel the eye movement to the target stimulus. By modeling this performance as a race between a go process triggered by the appearance of the target and a stop process triggered by the appearance of the stop signal, it is possible to estimate the processing interval associated with cancelling the movement. We have now used this paradigm to analyze the cancelling of pursuit and saccades. For pursuit, we obtained consistent estimates of the stop process regardless of our technique or assumptions--it took 50-60 ms to cancel pursuit in both humans and monkeys. For saccades, we found different values depending on our assumptions. When we assumed that saccade preparation was under inhibitory control up until movement onset, we found that saccades took longer to cancel (humans: approximately 110, monkeys: approximately 80 ms) than pursuit. However, when we assumed that saccade preparation includes a final "ballistic" interval not under inhibitory control, we found that the same rapid stop process that accounted for our pursuit results could also account for the cancelling of saccades. We favor this second interpretation because cancelling pursuit or saccades amounts to maintaining a state of fixation, and it is more parsimonious to assume that this involves a single inhibitory process associated with the fixation system, rather than two separate inhibitory processes depending on which type of eye movement will not be made. From our behavioral data, we estimate that this ballistic interval has a duration of 9-25 ms in monkeys, consistent with the known physiology of the final motor pathways for saccades, although we obtained longer values in humans (28-60 ms). Finally, we examined the effect of trial sequence during the countermanding task and found that pursuit and saccade latencies tended to be longer if the previous trial contained a stop signal than if it did not; these increases occurred regardless of whether the preceding trial was associated with the same or different type of eye movement. Together, these results suggest that a common inhibitory mechanism regulates the initiation of pursuit and saccades.
反向指令范式为研究负责取消眼动的机制提供了一个有用的工具。该范式的关键特征是,在少数试验中,目标出现一段时间后会引入一个停止信号,指示受试者应取消初始眼动。如果给出停止信号的延迟过长,受试者就无法取消对目标刺激的眼动。通过将这种表现建模为目标出现触发的执行过程与停止信号出现触发的停止过程之间的竞赛,就有可能估计与取消眼动相关的处理间隔。我们现在已经使用这个范式来分析追踪眼动和扫视眼动的取消情况。对于追踪眼动,无论我们采用何种技术或假设,我们都得到了关于停止过程的一致估计——人类和猴子取消追踪眼动都需要50 - 60毫秒。对于扫视眼动,根据我们的假设我们得到了不同的值。当我们假设扫视眼动准备在运动开始前一直处于抑制控制之下时,我们发现扫视眼动取消所需的时间比追踪眼动更长(人类:约110毫秒,猴子:约80毫秒)。然而,当我们假设扫视眼动准备包括一个不受抑制控制的最终“弹道”间隔时,我们发现解释我们追踪眼动结果的相同快速停止过程也可以解释扫视眼动的取消。我们支持第二种解释,因为取消追踪眼动或扫视眼动相当于维持注视状态,并且假设这涉及与注视系统相关的单个抑制过程,而不是取决于不进行哪种类型眼动的两个单独抑制过程,这样更简洁。根据我们的行为数据,我们估计猴子的这个弹道间隔持续时间为9 - 25毫秒,这与扫视眼动最终运动通路的已知生理学一致,尽管我们在人类中得到了更长的值(28 - 60毫秒)。最后,我们研究了反向指令任务期间试验序列的影响,发现如果前一个试验包含停止信号,追踪眼动和扫视眼动潜伏期往往比不包含时更长;无论前一个试验与相同还是不同类型的眼动相关,都会出现这些增加。总之,这些结果表明一种共同的抑制机制调节追踪眼动和扫视眼动的启动。