Functions of delay-period activity in the prefrontal cortex and mnemonic scotomas revisited

Abstract

Working memory (WM) is one of key concepts to understand functions of the prefrontal cortex. Delay-period activity is an important neural correlate to understand the role of WM in prefrontal functions. The importance of delay-period activity is that this activity can encode not only visuospatial information but also a variety of information including non-spatial visual features, auditory and tactile stimuli, task rules, expected reward, and numerical quantity. This activity also participates in a variety of information processing including sensory-to-motor information transformation. These mnemonic features of delay-period activity enable to perform various important operations that the prefrontal cortex participates in, such as executive controls, and therefore, support the notion that WM is an important function to understand prefrontal functions. On the other hand, although experiments using manual versions of the delayed-response task had revealed many important findings, an oculomotor version of this task enabled us to use multiple cue positions, exclude postural orientation during the delay period, and further prove the importance of mnemonic functions of the prefrontal cortex. In addition, monkeys with unilateral lesions exhibited specific impairment only in the performance of memory-guided saccades directed toward visual cues in the visual field contralateral to the lesioned hemisphere. This result indicates that memories for visuospatial coordinates in each hemifield are processed primarily in the contralateral prefrontal cortex. This result further strengthened the idea of mnemonic functions of the prefrontal cortex. Thus, the mnemonic functions of the prefrontal cortex and delay-period activity may not need to be reconsidered, but should be emphasized.

Keywords: delay-period activity; delayed-response task; mnemonic scotoma; prefrontal cortex; spatial-information processing; working memory.

Figure 1

(A) Schematic drawings of the events in the oculomotor delayed-response (ODR) task. (B) Drawing of the Wisconsin general test apparatus (WGTA). (C) Schematic drawings of a monkey performing the delayed-response task in the WGTA. Reproduced from Goldman-Rakic (1987).

Figure 2

An example of directional delay-period activity observed in the prefrontal cortex. The visual cue was presented randomly at one of predetermined 8 peripheral positions. The position of each figure corresponds to the position of the visual cue. Significant activation was observed during the delay period only when the visual cue was presented at the 270° position. C: cue period (0.5 s). D: delay period (3.0 s). R: response period. Reproduced from Funahashi et al. (1989).

Figure 3

Effects of focal lesions of the dorsolateral prefrontal cortex on performance in the ODR task. In this monkey (Monkey JN), the first lesion was located in the left principal sulcal region and, several months later, a second lesion was applied in the right principal sulcal region. The locations of the ellipses indicate the locations where the visual cues were presented. The size of the ellipse indicates the magnitude of the behavioral deficit. Three delay lengths (1.5 s, 3.0 s, and 6.0 s) were randomly applied. Note that larger deficits were observed when the visual cues were presented in the contralateral visual field with respect to the lesioned hemisphere. Reproduced from Funahashi et al. (1993a).