Department of Physiology and Biophysics and Washington National Primate Research Center, University of Washington, Seattle, Washington 98195-7330, USA.
J Neurosci. 2010 Mar 10;30(10):3715-27. doi: 10.1523/JNEUROSCI.4953-09.2010.
Adaptation of saccadic eye movements provides an excellent motor learning model to study theories of neuronal plasticity. When primates make saccades to a jumping target, a backward step of the target during the saccade can make it appear to overshoot. If this deception continues for many trials, saccades gradually decrease in amplitude to go directly to the back-stepped target location. We used this adaptation paradigm to evaluate the Marr-Albus hypothesis that such motor learning occurs at the Purkinje (P)-cell of the cerebellum. We recorded the activity of identified P-cells in the oculomotor vermis, lobules VIc and VII. After documenting the on and off error directions of the complex spike activity of a P-cell, we determined whether its saccade-related simple spike (SS) activity changed during saccade adaptation in those two directions. Before adaptation, 57 of 61 P-cells exhibited a clear burst, pause, or a combination of both for saccades in one or both directions. Sixty-two percent of all cells, including two of the four initially unresponsive ones, behaved differently for saccades whose size changed because of adaptation than for saccades of similar sizes gathered before adaptation. In at least 42% of these, the changes were appropriate to decrease saccade amplitude based on our current knowledge of cerebellum and brainstem saccade circuitry. Changes in activity during adaptation were not compensating for the potential fatigue associated with performing many saccades. Therefore, many P-cells in the oculomotor vermis exhibit changes in SS activity specific to adapted saccades and therefore appropriate to induce adaptation.
眼跳运动的适应为研究神经元可塑性理论提供了一个极好的运动学习模型。当灵长类动物向跳跃目标进行眼跳时,目标在眼跳过程中的向后移动会使其看起来超过目标位置。如果这种欺骗持续很多次试验,眼跳的幅度会逐渐减小,直接到达向后移动的目标位置。我们使用这种适应范式来评估 Marr-Albus 假说,即这种运动学习发生在小脑的浦肯野(P)细胞。我们记录了中脑导水管周围灰质、VIc 和 VII 叶的已识别 P 细胞的活动。在记录了一个 P 细胞的复杂峰活动的开和关误差方向之后,我们确定了其与眼跳相关的简单峰(SS)活动在这两个方向的眼跳适应过程中是否发生了变化。在适应之前,61 个 P 细胞中有 57 个在一个或两个方向的眼跳中表现出明显的爆发、停顿或两者的组合。在所有细胞中,包括最初无反应的四个中的两个,62%的细胞对大小因适应而改变的眼跳的行为与适应前收集的大小相似的眼跳不同。在至少 42%的细胞中,根据我们对小脑和脑干眼跳电路的当前知识,这些变化适当减小了眼跳幅度。适应过程中活动的变化不是为了补偿执行多次眼跳带来的潜在疲劳。因此,中脑导水管周围灰质中的许多 P 细胞表现出特定于适应眼跳的 SS 活动变化,因此适当诱导适应。