Paré M, Wurtz R H
Laboratory of Sensorimotor Research, National Eye Institute, National Institutes of Health, Bethesda, Maryland 20892,
J Neurophysiol. 2001 Jun;85(6):2545-62. doi: 10.1152/jn.2001.85.6.2545.
Neurons in both the lateral intraparietal area (LIP) of the monkey parietal cortex and the intermediate layers of the superior colliculus (SC) are activated well in advance of the initiation of saccadic eye movements. To determine whether there is a progression in the covert processing for saccades from area LIP to SC, we systematically compared the discharge properties of LIP output neurons identified by antidromic activation with those of SC neurons collected from the same monkeys. First, we compared activity patterns during a delayed saccade task and found that LIP and SC neurons showed an extensive overlap in their responses to visual stimuli and in their sustained activity during the delay period. The saccade activity of LIP neurons was, however, remarkably weaker than that of SC neurons and never occurred without any preceding delay activity. Second, we assessed the dependence of LIP and SC activity on the presence of a visual stimulus by contrasting their activity in delayed saccade trials in which the presentation of the visual stimulus was either sustained (visual trials) or brief (memory trials). Both the delay and the presaccadic activity levels of the LIP neuronal sample significantly depended on the sustained presence of the visual stimulus, whereas those of the SC neuronal sample did not. Third, we examined how the LIP and SC delay activity relates to the future production of a saccade using a delayed GO/NOGO saccade task, in which a change in color of the fixation stimulus instructed the monkey either to make a saccade to a peripheral visual stimulus or to withhold its response and maintain fixation. The average delay activity of both LIP and SC neuronal samples significantly increased by the advance instruction to make a saccade, but LIP neurons were significantly less dependent on the response instruction than SC neurons, and only a minority of LIP neurons was significantly modulated. Thus despite some overlap in their discharge properties, the neurons in the SC intermediate layers showed a greater independence from sustained visual stimulation and a tighter relationship to the production of an impending saccade than the LIP neurons supplying inputs to the SC. Rather than representing the transmission of one processing stage in parietal cortex area LIP to a subsequent processing stage in SC, the differences in neuronal activity that we observed suggest instead a progressive evolution in the neuronal processing for saccades.
猴子顶叶皮层的外侧顶内区(LIP)和上丘(SC)中间层的神经元,在扫视眼动开始之前就被很好地激活了。为了确定从LIP区到SC区是否存在对扫视的隐蔽加工过程,我们系统地比较了通过逆向激活鉴定的LIP输出神经元与从同一猴子收集的SC神经元的放电特性。首先,我们比较了延迟扫视任务期间的活动模式,发现LIP和SC神经元在对视觉刺激的反应以及延迟期间的持续活动方面表现出广泛的重叠。然而,LIP神经元的扫视活动明显弱于SC神经元,并且在没有任何先前延迟活动的情况下从未发生过。其次,我们通过对比它们在延迟扫视试验中的活动来评估LIP和SC活动对视觉刺激存在的依赖性,在这些试验中,视觉刺激的呈现要么是持续的(视觉试验),要么是短暂的(记忆试验)。LIP神经元样本的延迟和扫视前活动水平都显著依赖于视觉刺激的持续存在,而SC神经元样本则不然。第三,我们使用延迟的GO/NOGO扫视任务研究了LIP和SC延迟活动与未来扫视产生的关系,在该任务中,注视刺激颜色的变化指示猴子要么向周边视觉刺激进行扫视,要么抑制其反应并保持注视。对LIP和SC神经元样本的平均延迟活动而言,提前做出扫视的指令使其显著增加,但LIP神经元比SC神经元对反应指令的依赖性显著更低,并且只有少数LIP神经元受到显著调制。因此,尽管它们的放电特性存在一些重叠,但SC中间层的神经元比向SC提供输入的LIP神经元表现出对持续视觉刺激更大的独立性,以及与即将发生的扫视产生更紧密的关系。我们观察到的神经元活动差异并非代表顶叶皮层LIP区的一个加工阶段向SC区的后续加工阶段的传递,而是表明扫视的神经元加工过程在逐步演变。