Keisetsu Shima

Numerical representation for action in the parietal cortex of the monkey

The anterior part of the parietal association area in the cerebral cortex of primates has been implicated in the integration of somatosensory signals, which generate neural images of body parts and apposed objects and provide signals for sensorial guidance of movements. Here we show that this area is active in primates performing numerically based behavioural tasks. We required monkeys to select and perform movement A five times, switch to movement B for five repetitions, and return to movement A, in a cyclical fashion. Cellular activity in the superior parietal lobule reflected the number of self-movement executions. For the most part, the number-selective activity was also specific for the type of movement. This type of numerical representation of self-action was seen less often in the inferior parietal lobule, and rarely in the primary somatosensory cortex. Such activity in the superior parietal lobule is useful for processing numerical information, which is necessary to provide a foundation for the forthcoming motor selection.

Interval time coding by neurons in the presupplementary and supplementary motor areas

Interval timing is an essential guiding force of behavior. Previous reports have implicated the prefrontal and parietal cortex as being involved in time perception and in temporal decision making. We found that neurons in the medial motor areas, in particular the presupplementary motor area, participate in interval timing in the range of seconds. Monkeys were trained to perform an interval-generation task that required them to determine waiting periods of three different durations. Neuronal activity contributed to the process of retrieving time instructions from visual cues, signaled the initiation of action in a time-selective manner, and developed activity to represent the passage of time. These results specify how medial motor areas take part in initiating actions on the basis of self-generated time estimates.

Reorganization of activity in the supplementary motor area associated with motor learning and functional recovery

SummaryThe supplementary motor area (SMA) of primates has been implicated in the initiation and execution of limb movements. However, when a motor task was extensively overlearned, few SMA neurons, if any, were active before the movement onset. Subsequent lesions of the primary motor cortex gave rise to the appearance of premovement activity changes, indicating usedependent reorganization of the neuronal activity in SMA.

Categorization of behavioural sequences in the prefrontal cortex.

Although it has long been thought that the prefrontal cortex of primates is involved in the integrative regulation of behaviours, the neural architecture underlying specific aspects of cognitive behavioural planning has yet to be clarified. If subjects are required to remember a large number of complex motor sequences and plan to execute each of them individually, categorization of the sequences according to the specific temporal structure inherent in each subset of sequences serves to facilitate higher-order planning based on memory. Here we show, using these requirements, that cells in the lateral prefrontal cortex selectively exhibit activity for a specific category of behavioural sequences, and that categories of behaviours, embodied by different types of movement sequences, are represented in prefrontal cells during the process of planning. This cellular activity implies the generation of neural representations capable of storing structured event complexes at an abstract level, exemplifying the development of macro-structured action knowledge in the lateral prefrontal cortex.

Differences in spiking patterns among cortical neurons

Spike sequences recorded from four cortical areas of an awake behaving monkey were examined to explore characteristics that vary among neurons. We found that a measure of the local variation of interspike intervals, LV, is nearly the same for every spike sequence for any given neuron, while it varies significantly among neurons. The distributions of LV values for neuron ensembles in three of the four areas were found to be distinctly bimodal. Two groups of neurons classified according to the spiking irregularity exhibit different responses to the same stimulus. This suggests that neurons in each area can be classified into different groups possessing unique spiking statistics and corresponding functional properties.

Spatial distribution of cingulate cells projecting to the primary, supplementary, and pre-supplementary motor areas: a retrograde multiple labeling study in the macaque monkey

We examined the location and spatial distribution of cingulate cortical cells projecting to the forelimb areas of the primary motor cortex (MI), supplementary motor area (SMA), and pre-supplementary motor area (pre-SMA) using a multiple retrograde labeling technique in the monkeys (Macaca fuscata). The forelimb areas of the MI, SMA and pre-SMA were physiologically identified, based on the findings of intracortical microstimulation (ICMS) and single cell recording. Three different tracers, diamidino yellow (DY), fast blue (FB), and wheat germ agglutinin conjugated to horseradish peroxidase (WGA-HRP), were injected into each of the three motor areas in the same monkey. Retrogradely labeled cells in the cingulate cortex were plotted with an automated plotting system. Cells projecting to the forelimb area of the MI were distributed in the two separate regions situated rostrocaudally in the dorsal and ventral banks of the cingulate sulcus, namely the rostral cingulate motor area (CMAr) and caudal cingulate motor area (CMAc). These two regions corresponded to the forelimb areas identified by the ICMS in the same animal. The distribution of projection cells to the SMA overlapped extensively with that of projection cells to the MI. Although the MI received relatively sparse inputs from the CMAr than from the CMAc, the SMA received inputs from the CMAr and its adjacent areas as much as from the CMAc. The projection cells to the pre-SMA were distributed in the anterior portion of the cingulate cortex, including the anterior part of the CMAr and in a small part of the cingulate gyrus. These findings indicate that the MI and SMA share a considerable common information from the cingulate cortex, including the CMAr and CMAc, whereas the pre-SMA receives a different set of information from the anterior part of the cingulate cortex.

Relating Neuronal Firing Patterns to Functional Differentiation of Cerebral Cortex

It has been empirically established that the cerebral cortical areas defined by Brodmann one hundred years ago solely on the basis of cellular organization are closely correlated to their function, such as sensation, association, and motion. Cytoarchitectonically distinct cortical areas have different densities and types of neurons. Thus, signaling patterns may also vary among cytoarchitectonically unique cortical areas. To examine how neuronal signaling patterns are related to innate cortical functions, we detected intrinsic features of cortical firing by devising a metric that efficiently isolates non-Poisson irregular characteristics, independent of spike rate fluctuations that are caused extrinsically by ever-changing behavioral conditions. Using the new metric, we analyzed spike trains from over 1,000 neurons in 15 cortical areas sampled by eight independent neurophysiological laboratories. Analysis of firing-pattern dissimilarities across cortical areas revealed a gradient of firing regularity that corresponded closely to the functional category of the cortical area; neuronal spiking patterns are regular in motor areas, random in the visual areas, and bursty in the prefrontal area. Thus, signaling patterns may play an important role in function-specific cerebral cortical computation.

Concept-based behavioral planning and the lateral prefrontal cortex

Many lines of evidence implicate the lateral prefrontal cortex (LPFC) in the executive control of behavior. In early studies, neuronal activity in this area was thought to retain information about forthcoming movements for a short period until they were executed. However, later studies have stressed its role in the cognitive aspects of behavioral planning, such as behavioral significance, behavioral rules and behavioral goals. The consequence of the intended action (i.e. a change in the state of the target object), rather than the intended movement, is primarily represented in the LPFC during planning. Recent studies show that the LPFC is involved in more abstract aspects of conceptual processes, such as in representing categories of multiple actions at the stage of behavioral planning.

Firing properties of two types of nucleus raphe dorsalia neurons during the sleep-walking cycle and their responses to sensory stimuli

Spontaneous activity of the nucleus raphe dorsalis (NRD) neurons during the sleep-waking cycle and effects of sensory stimuli upon NRD neurons were studied in cats. Seventy-one neurons recorded within the NRD were classified into two groups with the use of the coefficient of variation of firing intervals during waking (W): 41 regularly firing (clock-like) and 30 irregularly firing (non-clock-like) neurons. The majority of clock-like and one-third of non-clock-like neurons showed a decrease in their firing rate during slow-wave sleep (SWS) compared with W. All neurons of both types displayed their lowest level of activity during paradoxical sleep. During the late phase of SWS, many clock-like neurons reduced their firing prior to the occurrence of pontogeniculo-occipital waves, whereas non-clock-like neurons did not show such a specific property. Clock-like neurons were totally unresponsive to nociceptive and non-nociceptive somesthetic stimuli, while about half of the non-clock-like neurons were driven by these stimuli. Half of the clock-like and one-third of the non-clock-like neurons were driven by click stimulation, and the majority of them showed an excitatory response. Some of the clock-like and non-clock-like neurons exhibited inhibitory and excitatory response to flash stimulation, respectively. The results of this experiment show that two types of neurons do exist in the NRD and suggest that they play a functionally different role in the brain.

Prefrontal cortical cells projecting to the supplementary eye field and presupplementary motor area in the monkey

We examined the location and spatial distribution of prefrontal cortical (PF) cells projecting to the supplementary eye field (SEF) and presupplementary motor area (pre-SMA) using a double retrograde-labeling technique in monkeys (Macaca fuscata). The SEF and pre-SMA were physiologically identified based on the findings of intracortical microstimulation and single cell recordings. Two fluorescent tracers, diamidino yellow and fast blue, were injected into the SEF and pre-SMA of each monkey. Retrogradely labeled cells in the PF were plotted with an automated plotting system. The cells projecting to the SEF and pre-SMA were mainly distributed in the upper and lower banks of the principal sulcus (area 46), with little overlap. Cells projecting to the SEF, but not to the pre-SMA, were observed in areas 8a, 8b, 9, 12, and 45. These findings suggest that the SEF and pre-SMA receive different sets of information from the PF cells.