A number of cortical structures are reported to have elevated single unit firing rates sustained throughout the memory period of a working memory task. How the nervous system forms and maintains these memories is unknown but reverberating neuronal network activity is thought to be important. We studied the temporal structure of single unit (SU) activity and simultaneously recorded local field potential (LFP) activity from area LIP in the inferior parietal lobe of two awake macaques during a memory-saccade task. Using multitaper techniques for spectral analysis, which play an important role in obtaining the present results, we find elevations in spectral power in a 50-90Hz (gamma) frequency band during the memory period in both SU and LFP activity. The activity is tuned to the direction of the saccade providing evidence for temporal structure that codes for movement plans during working memory. We also find SU and LFP activity are coherent during the memory period in the 50-90Hz gamma band and no consistent relation is present during simple fixation. Finally, we find organized LFP activity in a 15-25Hz frequency band that may be related to movement execution and preparatory aspects of the task. Neuronal activity could be used to control a neural prosthesis but SU activity can be hard to isolate with cortical implants. As the LFP is easier to acquire than SU activity, our finding of rich temporal structure in LFP activity related to movement planning and execution may accelerate the development of this medical application.Keywords: parietal, prosthesis, local field potential, gamma band, coherence, temporal structure.Pesaran et. al. 3Working memory is a brain system requiring the temporary storage and manipulation of information necessary for the performance of complex cognitive tasks (Baddeley, 1992). The neurophysiological basis of working memory is studied in non-human primates by recording neural activity during delayed-response tasks (Fuster, 1995). Cue-selective elevated single unit firing rates have been recorded during the delay period in many brain areas during different versions of the task (Fuster and Jervey, 1982;Bruce and Goldberg, 1985;Gnadt and Andersen, 1988;Miyashita and Chang, 1988;Funahashi et al., 1989;Koch and Fuster, 1989;Miller et al., 1996;Zhou and Fuster, 1996). How this neural activity is sustained is unknown but may be important to understanding the neural basis of working memory (Goldman-Rakic, 1995). Converging evidence points to the importance of a distributed recurrent neuronal network (Goldman-Rakic, 1988) and reverberating network activity has long been suggested as a possible mechanism for short-term memory (Lorente de No, 1938;Hebb, 1949;Amit, 1995;Seung, 1996;Wang, 1999).Measures with the potential to capture correlated neural activity on a millisecond time scale may be needed to resolve reverberating memory activity. The dynamical structure of neuronal activity has been the source of much interest as a temporal code (for a review see Singer and Gray (1995) ) however ...

To look at or reach for what we see, spatial information from the visual system must be transformed into a motor plan. The posterior parietal cortex (PPC) is well placed to perform this function, because it lies between visual areas, which encode spatial information, and motor cortical areas. The PPC contains several subdivisions, which are generally conceived as high-order sensory areas. Neurons in area 7a and the lateral intraparietal area fire before and during visually guided saccades. Other neurons in areas 7a and 5 are active before and during visually guided arm movements. These areas are also active during memory tasks in which the animal remembers the location of a target for hundreds of milliseconds before making an eye or arm movement. Such activity could reflect either visual attention or the intention to make movements. This question is difficult to resolve, because even if the animal maintains fixation while directing attention to a peripheral location, the observed neuronal activity could reflect movements that are planned but not executed. To address this, we recorded from the PPC while monkeys planned either reaches or saccades to a single remembered location. We now report that, for most neurons, activity before the movement depended on the type of movement being planned. We conclude that PPC contains signals related to what the animal intends to do.

The posterior parietal cortex (PPC), historically believed to be a sensory structure, is now viewed as an area important for sensory-motor integration. Among its functions is the forming of intentions, that is, high-level cognitive plans for movement. There is a map of intentions within the PPC, with different subregions dedicated to the planning of eye movements, reaching movements, and grasping movements. These areas appear to be specialized for the multisensory integration and coordinate transformations required to convert sensory input to motor output. In several subregions of the PPC, these operations are facilitated by the use of a common distributed space representation that is independent of both sensory input and motor output. Attention and learning effects are also evident in the PPC. However, these effects may be general to cortex and operate in the PPC in the context of sensory-motor transformations.

The cortex of the inferior parietal lobule in primates is important for spatial perception and spatially oriented behavior. Recordings of single neurons in this area in behaving monkeys showed that the visual sensitivity of the retinotopic receptive fields changes systematically with the angle of gaze. The activity of many of the neurons can be largely described by the product of a gain factor that is a function of the eye position and the response profile of the visual receptive field. This operation produces an eye position-dependent tuning for locations in head-centered coordinate space.

Neurons in area 7a of the posterior parietal cortex of monkeys respond to both the retinal location of a visual stimulus and the position of the eyes and by combining these signals represent the spatial location of external objects. A neural network model, programmed using back-propagation learning, can decode this spatial information from area 7a neurons and accounts for their observed response properties.

Recent experiments are reviewed that indicate that sensory signals from many modalities, as well as efference copy signals from motor structures, converge in the posterior parietal cortex in order to code the spatial locations of goals for movement. These signals are combined using a specific gain mechanism that enables the different coordinate frames of the various input signals to be combined into common, distributed spatial representations. These distributed representations can be used to convert the sensory locations of stimuli into the appropriate motor coordinates required for making directed movements. Within these spatial representations of the posterior parietal cortex are neural activities related to higher cognitive functions, including attention. We review recent studies showing that the encoding of intentions to make movements is also among the cognitive functions of this area.

Cells in the dorsal division of the medial superior temporal area (MSTd) have large receptive fields and respond to expansion/contraction, rotation, and translation motions. These same motions are generated as we move through the environment, leading investigators to suggest that area MSTd analyzes the optical flow. One influential idea suggests that navigation is achieved by decomposing the optical flow into the separate and discrete channels mentioned above, that is, expansion/contraction, rotation, and translation. We directly tested whether MSTd neurons perform such a decomposition by examining whether there are cells that are preferentially tuned to intermediate spiral motions, which combine both expansion/contraction and rotation components. The finding that many cells in MSTd are preferentially selective for spiral motions indicates that this simple three-channel decomposition hypothesis for MSTd does not appear to be correct. Instead, there is a continuum of patterns to which MSTd cells are selective. In addition, we find that MSTd cells maintain their selectivity when stimuli are moved to different locations in their large receptive fields. This position invariance indicates that MSTd cells selective for expansion cannot give precise information about the retinal location of the focus of expansion. Thus, individual MSTd neurons cannot code, in a precise fashion, the direction of heading by using the location of the focus of expansion. The only way this navigational information could be accurately derived from MSTd is through the use of a coarse, population encoding. Positional invariance and selectivity for a wide array of stimuli suggest that MSTd neurons encode patterns of motion per se, regardless of whether these motions are generated by moving objects or by motion induced by observer locomotion.

Single cell and multiunit signals were recorded by a multichannel recording system (Plexon Inc, Texas) from 96 paralyne coated tungsten or platinum/iridium electrodes (impedance ≈ 300 kΩ) (Microprobe Inc. Maryland) implanted in the medial intraparietal area (MIP), a subdivision of the parietal reach region (PRR), and area 5 (1) of three rhesus monkeys trained to perform a memory reach task. One monkey (monkey S) also had 64 electrodes implanted in the dorsal premotor area (PMd) in a separate surgery. Each session consisted of a reach segment and a brain control segment. Trials in both segments were initiated in the same way: after the monkeys acquired a central red fixation point with the eyes and touched a central green target, a peripheral cue was flashed indicating the location of one out of four, five, six, or eight reach targets ( Figure 1a) (cue epoch). Reach targets were uniformly distributed around the central fixation point. As soon as the fixation point and central green target were acquired, hand and eye movements were restricted by a real time behavioural controller (LabVIEW, National Instruments). Eye position was monitored using a scleral search coil (CNC Engineering, monkeys S and O), or an infrared reflection system (ISCAN, monkey C) while hand position was monitored using an acoustic touch screen (ELO Touch). In order to successfully complete a trial, the monkeys were not allowed to move their eyes. In addition, the reaching hand had to be in contact with the centrally located green target at all times except after the GO signal which appeared during the reach segment of the session. After the offset of the cue, a delay of 1.5 ± 0.3 seconds ensued. During the reach segment, the green central target was extinguished after the memory period