Instruction Cue Research Articles

Differential roles of neuronal activity in the supplementary and presupplementary motor areas: from information retrieval to motor planning and execution.

We explored functional differences between the supplementary and presupplementary motor areas (SMA and pre-SMA, respectively) systematically with respect to multiple behavioral factors, ranging from the retrieval and processing of associative visual signals to the planning and execution of target-reaching movement. We analyzed neuronal activity while monkeys performed a behavioral task in which two visual instruction cues were given successively with a delay: one cue instructed the location of the reach target, and the other instructed arm use (right or left). After a second delay, the monkey received a motor-set cue to be prepared to make the reaching movement as instructed. Finally, after a GO signal, it reached for the instructed target with the instructed arm. We found the following apparent differences in activity: 1) neuronal activity preceding the appearance of visual cues was more frequent in the pre-SMA; 2) a majority of pre-SMA neurons, but many fewer SMA neurons, responded to the first or second cue, reflecting what was shown or instructed; 3) in addition, pre-SMA neurons often reflected information combining the instructions in the first and second cues; 4) during the motor-set period, pre-SMA neurons preferentially reflected the location of the target, while SMA neurons mainly reflected which arm to use; and 5) when executing the movement, a majority of SMA neurons increased their activity and were largely selective for the use of either the ipsilateral or contralateral arm. In contrast, the activity of pre-SMA neurons tended to be suppressed. These findings point to the functional specialization of the two areas, with respect to receiving associative cues, information processing, motor behavior planning, and movement execution.

Differential reward outcome learning in adult humans

Differential reward outcome learning provides a unique reward outcome for each condition of a conditional discrimination task, and this increases the rate at which these tasks can be learned. The present experiment aims to show that irrespective of any difference in learning rate, conditional discrimination tasks and differential reward outcome tasks are solved using different strategies. Two groups of 30 adult subjects were taught a series of conditional visual discriminations. One group received different reward outcomes for each condition, whilst the other group received the same reward outcome for each condition. Both groups learned the visual discriminations at the same rate. Subjects were then taught a new conditional cue with a subset of previously learnt discrimination problems and then required to transfer the newly learnt instruction cue to the remaining discrimination problems that had been previously learnt. Although both groups appeared to transfer the newly learnt cue at the same rate, the subjects performing the differential reward outcome task learnt to a criterion level of performance with fewer errors. The results are discussed in relation to evidence from monkeys indicating different neural mechanisms underlying the learning of both tasks.

Long- and short-range reward expectancy in the primate orbitofrontal cortex.

The orbitofrontal cortex (OFC) is important in motivation and emotion. We previously reported reward expectancy-related delay activities during a delayed reaction time task in primate OFC neurons. To further investigate the significance of the OFC in motivational operations, we examined pre-instruction, baseline activities of OFC neurons in relation to reward expectancy during the delayed reaction time task. In this task, an instruction cue indicated whether reward would be present or absent in the trial. Each set of four consecutive trials constituted one block within which three different kinds of rewards and one trial with no reward were given in a fixed order that differed from the monkey's reward preference. We identified two types of OFC neurons with reward expectancy-related pre-instruction activities: Step-type neurons showed stepwise changes (increase or decrease) in pre-instruction activity toward the trial with a particular outcome, which usually was the most or least attractive within a block; Pref-type neurons showed pre-instruction activity changes according to the monkey's preference for each trial's outcome. We propose that Step-type and Pref-type neurons are related to long-range and short-range reward expectancies of a particular outcome, respectively. The OFC is considered to play important roles in goal-directed behaviour by adjusting the motivational level toward a certain (current or future) outcome of a particular motivational significance based on the two kinds of reward expectancy processes. Impairments in goal-directed behaviour by OFC patients may be caused by a lack of long-range expectancy or by a deficit in compromising between short-range and long-range expectancies.

Neuronal activity in the primate substantia nigra pars reticulata during the performance of simple and memory-guided elbow movements.

The basal ganglia participate in motor functions and are implicated in the pathophysiology of movement disorders. It has been shown in primates that the activity of many neurons in one of the basal ganglia output nuclei, the internal segment of the globus pallidus, changes with active or passive movements. The involvement of the second major output nucleus, the substantia nigra pars reticulata (SNr), in movement is less well established. In this study, the electrophysiologic activity of SNr neurons was studied in two awake Rhesus monkeys while the animals were examined and while they performed elbow movements in two different motor tasks (n = 261 cells). Responses to examination were uncommon and subtle. Twenty-one percent of neurons responded to the target step in a step tracking task, mostly with anticipatory responses, although some cells showed directional, movement-related activity. In a delayed-response task, 17% of cells showed anticipatory activity to an instruction cue preceding the target jump, 11% responded directly to the cue, and 11% showed long-lasting postcue activity. Movement-related responses were seen in 21% in this task. Reward responses occurred in 10% of neurons. Responses to more than one event were common. The results demonstrate that few neurons in the SNr respond directly to passive or active movements, but a large proportion shows responses that may be related to memory, attention, or movement preparation. While internal pallidal segment neurons may be preferentially concerned with controlling elemental movement parameters, neurons in the SNr may be more involved in higher motor functions or nonmotor aspects of behavior.

Dorsoventral Patterning of the Mouse Coat by Tbx15

Many members of the animal kingdom display coat or skin color differences along their dorsoventral axis. To determine the mechanisms that control regional differences in pigmentation, we have studied how a classical mouse mutation, droopy ear (deH), affects dorsoventral skin characteristics, especially those under control of the Agouti gene. Mice carrying the Agouti allele black-and-tan (at) normally have a sharp boundary between dorsal black hair and yellow ventral hair; the deH mutation raises the pigmentation boundary, producing an apparent dorsal-to-ventral transformation. We identify a 216 kb deletion in deH that removes all but the first exon of the Tbx15 gene, whose embryonic expression in developing mesenchyme correlates with pigmentary and skeletal malformations observed in deH/deH animals. Construction of a targeted allele of Tbx15 confirmed that the deH phenotype was caused by Tbx15 loss of function. Early embryonic expression of Tbx15 in dorsal mesenchyme is complementary to Agouti expression in ventral mesenchyme; in the absence of Tbx15, expression of Agouti in both embryos and postnatal animals is displaced dorsally. Transplantation experiments demonstrate that positional identity of the skin with regard to dorsoventral pigmentation differences is acquired by E12.5, which is shortly after early embryonic expression of Tbx15. Fate-mapping studies show that the dorsoventral pigmentation boundary is not in register with a previously identified dermal cell lineage boundary, but rather with the limb dorsoventral boundary. Embryonic expression of Tbx15 in dorsolateral mesenchyme provides an instructional cue required to establish the future positional identity of dorsal dermis. These findings represent a novel role for T-box gene action in embryonic development, identify a previously unappreciated aspect of dorsoventral patterning that is widely represented in furred mammals, and provide insight into the mechanisms that underlie region-specific differences in body morphology.

Neural activity in primary motor and dorsal premotor cortex in reaching tasks with the contralateral versus ipsilateral arm.

To investigate the effector dependence of task-related neural activity in dorsal premotor (PMd) and primary motor cortex (M1), directional tuning functions were compared between instructed-delay reaching tasks performed separately with either the contralateral or the ipsilateral limb. During presentation of the instructional cue, the majority (55/90, 61%) of cells in PMd were tuned with both arms, and their dynamic range showed a trend for stronger discharge with the contralateral arm. Most strikingly, however, the preferred direction of most of these latter cells (41/55, 75%) was not significantly different between arms. During movement, many PMd cells continued to be tuned with both arms (53/90, 59%), with a trend for increasing directional differences between the arms over the course of the trial. In contrast, during presentation of the instructional cue only 5/74 (7%) cells in M1 were tuned with both arms. During movement, about half of M1 cells (41/74, 55%) were tuned with both arms but the preferred directions of their tuning functions were often very different and there was a strong bias toward greater discharge rates when the contralateral arm was used. Similar trends were observed for EMG activity. In conclusion, M1 is strongly activated during movements of the contralateral arm, but activity during ipsilateral arm movements is also common and usually different from that seen with the contralateral arm. In contrast, a major component of task-related activity in PMd represents movement in a more abstract or task-dependent and effector-independent manner, especially during the instructed-delay period.

Effects of expectations for different reward magnitudes on neuronal activity in primate striatum.

In behavioral science, it is well known that humans and nonhuman animals are highly sensitive to differences in reward magnitude when choosing an outcome from a set of alternatives. We know that a realm of behavioral reactions is altered when animals begin to expect different levels of reward outcome. Our present aim was to investigate how the expectation for different magnitudes of reward influences behavior-related neurophysiology in the anterior striatum. In a spatial delayed response task, different instruction pictures are presented to the monkey. Each image represents a different magnitude of juice. By reaching to the spatial location where an instruction picture was presented, animals could receive the particular liquid amount designated by the stimulus. Reliable preferences in reward choice trials and differences in anticipatory licks, performance errors, and reaction times indicated that animals differentially expected the various reward amounts predicted by the instruction cues. A total of 374 of 2,000 neurons in the anterior parts of the caudate nucleus, putamen, and ventral striatum showed five forms of task-related activation during the preparation or execution of movement and activations preceding or following the liquid drop delivery. Approximately one-half of these striatal neurons showed differing response levels dependent on the magnitude of liquid to be received. Results of a linear regression analysis showed that reward magnitude and single cell discharge rate were related in a subset of neurons by a monotonic positive or negative relationship. Overall, these data support the idea that the striatum utilizes expectancies that contain precise information concerning the predicted, forthcoming level of reward in directing general behavioral reactions.

Disruption of information processing in the supplementary motor area of the MPTP-treated monkey: a clue to the pathophysiology of akinesia?

It has been suggested that the underactivity of mesial frontal structures induced by dopamine depletion could constitute one of the main substrates underlying akinesia in Parkinson's disease. Functional imaging and movement-related potential recordings indicate an implication of the frontal lobes in this pathological process, but the question has not yet been investigated at a cellular level using single unit recording. We therefore compared neuronal activity in both the presupplementary motor area (pre-SMA) and the supplementary motor area proper (SMAp) of the Macaca mulatta monkey during a delayed motor task, before and after MPTP treatment. In the pre-SMA, which receives strong inputs from the prefrontal cortex, the baseline firing frequency and the percentage of neurons responding to visual instruction cues decreased in lesioned monkeys. In the SMAp, which sends direct outputs to the primary motor cortex, not only was the response to visual cues impaired, but the percentage of SMAp neurons responding to intracortical microstimulation fell and the threshold of response rose. Neuronal activity after the Go signal diminished sharply in both structures in the symptomatic animal and the discharge pattern became more irregular; in the SMAp neuronal activity remained modified longer. Most of these changes could already be observed in the presymptomatic animal presenting no clinical signs of parkinsonism. These data would indicate that, at the moment when dopamine depletion has impaired the ability of cortical neurons to operate the focused selection of incoming information giving instructions for movement, pre-SMA and SMAp neurons are also in a state of severe hypoactivity. The conjunction of these phenomena could play a critical role in the genesis of akinesia.

Effects of a cross-modal manipulation of attention on somatosensory cortical neuronal responses to tactile stimuli in the monkey.

The role of attention in modulating tactile sensitivity in primary (SI) and secondary somatosensory cortex (SII) was addressed using a cross-modal manipulation of attention, somatosensory versus visual. Two adult monkeys (Macaca mulatta) were trained to perform two tasks: tactile discrimination of a change in the texture of a surface presented to digits 3 and 4 and visual discrimination of a change in the intensity of a light. In each trial, standard texture (2 mm spatial period, SP) and visual stimuli were presented. These were followed by an increase in SP and/or luminance. Each trial was preceded by an instruction cue (colored light) that directed the animal to attend and respond to the change in one modality while ignoring any change in the other modality. The two tasks were interleaved during the recording, on a trial-by-trial basis. Extracellular recordings were made from 178 neurons (SI, 102; SII, 76), all with a cutaneous receptive field on the stimulated digit tips. Discharge was quantified in both tasks during the instruction, the standard-stimuli, and the texture-change periods. The results showed that selective attention to tactile stimuli had qualitatively and quantitatively greater and earlier effects in SII than SI. Twenty-four of 102 SI cells showed a significant change in discharge with the direction of attention. For almost all cells (20/24), discharge was enhanced when attention was directed toward the tactile stimuli; the effects were most frequent in the analysis interval that encompassed the change in SP (16/24). A significantly higher proportion of SII cells were attention-sensitive (47/76). The effects were concentrated in the texture-change period (39/47) but also included earlier periods in the trial (instruction period, n = 15; standard-stimuli period, n = 32). Attention-related modulation that spanned all three intervals (n = 11) likely reflected baseline changes in discharge. For the texture-sensitive cells (43 in SI, 37 in SII), the mean change in discharge frequency (post texture change - pre-texture change) in each task was significantly increased in SII but not SI with selective attention. The results are consistent with a two-stage modulation of parietal cortical discharge, an initial stage (SI) in which there is some enhancement of sensory responses to the salient feature, the texture change, and a second stage (SII) in which baseline changes occur, along with further feature selection. These controls may be independently exerted on SI and SII, or they may reflect top-down controls from SII to SI.

Parietal Magnetic Stimulation Delays Visuomotor Mental Rotation at Increased Processing Demands

Visuomotor rotation (VMR) is a variant of the classic mental rotation paradigm. Subjects perform a center-out arm reaching movement, with the instruction to point clockwise or anticlockwise away from the direction of a reaction signal by a prespecified amount. Like classic mental rotation (MR) tasks, there is a linear relationship between reaction time (RT) and required angle of rotation (angular disparity). Although functional imaging studies have consistently demonstrated parietal activations centered around the intraparietal sulcus during MR tasks, the involvement of parietal cortex in VMR has not been investigated. The aim of the present experiments was to test in human subjects whether VMR also involves activity in parietal areas. We used short trains of transcranial magnetic stimulation (TMS) to produce a temporary “virtual lesion” of the posterior parietal cortex (PPC) around the intraparietal sulcus during the reaction period of a VMR task. Four pulses of 20-Hz rTMS were applied to the left PPC, right PPC, or vertex (control condition) 100 ms after the presentation of an instruction cue. Reaction times (RTs) were evenly prolonged by right or left parietal TMS compared with vertex stimulation, but only for large angles of rotation, and without affecting the spatial accuracy of the final response. A control experiment showed that parietal rTMS did not impair visual perception or the ability to judge the size of visual angles. The data thus provide evidence for bilateral involvement of the PPC in VMR that increases with processing demands.

Differential Involvement of Parietal and Precentral Regions in Movement Preparation and Motor Intention

Flexible goal-oriented behavior relies on spatial coordinate transformations and motor control mechanisms, but also on the capability to take advantage of contextual information for steering the sensorimotor machinery. Although accurate performance of a sensorimotor task requires parietal and frontal regions, their differential contribution and functional relationship with other associative regions remains obscure. We have used event-related functional magnetic resonance imaging to measure human cerebral activity associated with motor cognitive processes in the context of delayed performance of an associative visuomotor task. Movement instruction (specified by visual cues) and motor performance (specified by an auditory cue) were separated by a variable delay period. By manipulating the predictive value of the instruction cue, we distinguished delay-related activity influenced by response probabilities (motor preparation and motor inhibition) from delay-related activity unaffected by the likelihood of providing a motor response (motor intention). We found delay-related activity distributed across a cerebral network involving not only frontal circuitry, but also extrastriate and mediotemporal regions. Areas showing motor intentions and preparatory responses were spatially intermingled. Posterior parietal cortex deviated from this pattern, showing delay-related activity regardless of movement probability, but no specific preparatory responses. These results suggest that posterior parietal cortex and dorsal precentral cortex play different strategic roles in handling associative visuomotor problems. While parietal regions cover a range of potential responses defined by the task setting, precentral regions focus on a likely movement. Temporo-prefrontal regions might incorporate contextual information in the visuomotor process by defining potential and probable responses on the basis of the task contingencies.

Spatial Attention and Memory Versus Motor Preparation: Premotor Cortex Involvement as Revealed by fMRI

Recent studies in both monkeys and humans indicate that the dorsal premotor cortex participates in spatial attention and working memory, in addition to its well known role in movement planning and execution. One important question is whether these functions overlap or are segregated within this frontal area. Single-cell recordings in monkeys suggest a relative specialization of the rostral portion of dorsal premotor cortex for attention and/or memory and of the caudal region for motor preparation. To test whether this possibility also holds true in humans, we used functional magnetic resonance imaging (fMRI) to compare, in the same set of subjects, brain activation related to strong spatial attention and memory demands to that elicited by long motor preparatory periods. The behavioral protocol was based on a task that had proved effective for dissociating neuronal properties related to these two functions in the monkey brain. The principle of the monkey task was that a first cue guided the focus of spatial attention and memory, whereas a second one instructed an arm movement. Based on this principle, two tasks were developed. One maximized spatial attention and memory demands by presenting long series of stimuli (4, 8, or 12) before the motor instructional cue, whereas the other extended the motor preparation phase by imposing long and variable delays (1-5.5 s) between the onset of the instructional cue and movement execution. The two tasks and their respective control conditions were arranged in two blocked-design sequences. The results indicate that the brain networks underlying the two functional domains overlap in the caudate nucleus and presupplementary motor area, and possibly in lateral prefrontal cortex as well, but involve different dorsal premotor fields. Motor preparation primarily recruited a dorsal premotor area located caudally, within the precentral gyrus (together with the supplementary motor area), whereas spatial attention and memory preferentially activated a more rostral site, in and anterior to the precentral sulcus (in addition to the posterior parietal cortex). These findings strengthen the idea that the primate dorsal premotor cortex contributes to both motor and nonmotor processes. Moreover, they corroborate emerging evidence from monkey physiology suggesting a relative functional segregation within this cortex, with attention to short-term storage of visuospatial information engaging a more rostral region than motor preparation.

Multiple movement representations in the human brain: an event-related fMRI study.

Neurovascular correlates of response preparation have been investigated in human neuroimaging studies. However, conventional neuroimaging cannot distinguish, within the same trial, between areas involved in response selection and/or response execution and areas specifically involved in response preparation. The specific contribution of parietal and frontal areas to motor preparation has been explored in electrophysiological studies in monkey. However, the associative nature of sensorimotor tasks calls for the additional contributions of other cortical regions. In this article, we have investigated the functional anatomy of movement representations in the context of an associative visuomotor task with instructed delays. Neural correlates of movement representations have been assessed by isolating preparatory activity that is independent from the performance of an actual motor act, or from the presence of a response's target. Movement instruction (specified by visual cues) and motor performance (specified by an auditory cue) were separated by a variable delay period. We have used whole-brain event-related fMRI to measure human brain activity during the performance of such a task. We have focused our analysis on specific preparatory activity, defined as a sustained response over variable delay periods between a transient visual instruction cue and a brief motor response, temporally independent from the transient events. Behavioral and electrophysiological controls ensured that preparatory activity was not contaminated by overt motor responses or working memory processes. We report suggestive evidence for multiple movement representations in the human brain. Specific sustained activity in preparation for an action was found not only in parieto-frontal regions but also in extrastriate areas and in the posterior portion of the superior temporal sulcus. We suggest that goal-directed preparatory activity relies on both visuomotor and visuoperceptual areas. These findings point to a functional-anatomical basis for the integration of perceptual and executive processes.

Participation of the thalamic CM-Pf complex in attentional orienting.

The centre médian-parafascicular (CM-Pf) complex is located at the posterior intralaminar nuclei of the thalamus and forms part of the nonspecific thalamocortical projection system and the internal circuit of the basal ganglia. However, the functional roles of this complex remain to be fully elucidated. Here we have examined whether the CM-Pf complex is involved in the process of covert attention. We trained two macaque monkeys to perform a task in which a visual target stimulus for button release appeared at either the same location as the preceding visual instruction cue (a "validly cued target") or a location on the opposite side (an "invalidly cued target"). Reaction times (RTs) to a validly cued target were significantly shorter than those to an invalidly cued target, leading to a "validity effect" of about 20 ms. We recorded the activity of 97 neurons in the CM-Pf while the monkeys performed the attention task with the hand that was contralateral to the neuronal recording. Seventy CM-Pf neurons showed task-related activity after the appearance of either the instruction cue or the target stimulus: 33 neurons responded with a prominent short-latency facilitation (SLF), whereas 37 responded with a short-latency suppression followed by a long-latency facilitation (LLF). Most of the SLF neurons responded preferentially to a cue appearing on the contralateral side (76%) and to an invalidly cued target appearing on the contralateral side (61%). In contrast, LLF neurons showed a short-latency suppression after the cue stimulus, regardless of whether the cue appeared on the contra- or ipsilateral side (84%). Inactivating the CM-Pf complex by local injection (1 microl) of the GABA(A) receptor agonist muscimol (1-5 microg/microl) resulted in a significant increase in the RT to a validly cued target presented on the contra- but not the ipsilateral side. In contrast, inactivating the CM-Pf complex did not affect RTs to invalidly cued targets on either the contra- or the ipsilateral side. Thus the validity effect was abolished only on the contralateral side. We conclude that the CM-Pf complex plays a specific and essential role in the process of attentional orienting to external events occurring on the contralateral side, probably through the projection of primary outputs to the striatum, which is involved in the action-selection mechanisms of the basal ganglia.

Coding and Monitoring of Motivational Context in the Primate Prefrontal Cortex

The prefrontal cortex is involved in acquiring and maintaining information about context, including the set of task instructions and/or the outcome of previous stimulus-response sequences. Most studies on context-dependent processing in the prefrontal cortex have been concerned with such executive functions, but the prefrontal cortex is also involved in motivational operations. We thus wished to determine whether primate prefrontal neurons show evidence of representing the motivational context learned by the monkey. We trained monkeys in a delayed reaction task in which an instruction cue indicated the presence or absence of reward. In random alternation with no reward, the same one of several different kinds of food and liquid rewards was delivered repeatedly in a block of approximately 50 trials, so that reward information would define the motivational context. In response to an instruction cue indicating absence of reward, we found that neurons in the lateral prefrontal cortex not only predicted the absence of reward but also represented more specifically which kind of reward would be omitted in a given trial. These neurons seem to code contextual information concerning which kind of reward may be delivered in following trials. We also found prefrontal neurons that showed tonic baseline activity that may be related to monitoring such motivational context. The different types of neurons were distributed differently along the dorsoventral extent of the lateral prefrontal cortex. Such operations in the prefrontal cortex may be important for the monkey to maximize reward or to modify behavioral strategies and thus may contribute to executive control.

Contrasting neuronal activity in the dorsal and ventral premotor areas during preparation to reach.

We compared neuronal activity in the dorsal and ventral premotor areas (PMd and PMv, respectively) when monkeys were preparing to perform arm-reaching movements in a motor-set period before their actual execution. They were required to select one of four possible movements (reaching to a target on the left or right, using either the left or right arm) in accordance with two sets of instruction cues, followed by a delay period, and a subsequent motor-set period. During the motor-set period, the monkeys were required to get ready for a movement-trigger signal to start the arm-reach promptly. We analyzed the activity of 211 PMd and 109 PMv neurons that showed selectivity for the combination of the two instruction cues during the motor-set period. A majority (53%) of PMd neurons exhibited activity significantly tuned to both target location and arm use, and an approximately equal number of PMd neurons showed selectivity to either forthcoming arm use or target location. In contrast, 60% of PMv neurons showed selectivity for target location only and not for arm use. These findings point to preference in the use of neuronal activity in the two areas: preparation for action in the PMd and preparation for target acquisition in the PMv.

Functional properties of primate spinal interneurones during voluntary hand movements.

The activity of cervical spinal interneurones (INs) was recorded in monkeys performing alternating hand movements. The contribution of INs to voluntary movement was determined by their response patterns during ramp-and-hold wrist movements and their postspike effects on forelimb muscle activity. Most INs were active during both flexion and extension, in contrast to the unidirectional activity of muscles and corticomotoneuronal cells. When recorded during performance of an instructed delay task, the activity of many INs was modulated during the delay period between the instruction cue and the subsequent go signal. Thus, spinal INs, like cortical neurones, participate in earliest stages of preparation for movement. The modulation of peripheral input to spinal INs was tested during an instructed delay task. The monosynaptic responses to electrical stimulation of a cutaneous nerve decreased during active movement, probably due to presynaptic inhibition. These results provide new insights into the role of spinal INs in preparation and execution of voluntary movement.

Reward-dependent spatial selectivity of anticipatory activity in monkey caudate neurons.

Many neurons show anticipatory activity in learned tasks. This phenomenon appears to reflect the brain's ability to predict future events. However, what actually is predicted is unknown. Using a memory-guided saccade task, in which only one out of four directions was rewarded in each block of trials, we found that a group of neurons in the monkey caudate nucleus (CD) showed activity before presentation of an instruction cue stimulus. Among 329 CD neurons that were related to memory-guided saccade tasks, 156 showed the precue activity and 91 of them were examined fully. Remarkably, the magnitude of the precue activity varied across the four blocks of the one-direction-rewarded (1DR) condition, depending on which direction was rewarded. A majority of neurons with precue activity (83/91, 91%) showed significant directional preference. The best and worst directions were usually in the contralateral and ipsilateral directions, respectively. Within a block, the precue activity increased rapidly for the best direction in 1DR and decreased gradually for the worst direction in 1DR and all-directions-rewarded (ADR) condition. The precue activity was weak in ADR. The precue activity did not reflect the likelihood of a particular cue stimulus, because the probability of the cue appearing in each direction was the same regardless of the rewarded direction. These results suggest that each CD neuron indicates a particular position-reward association prospectively, usually with contralateral preference. Assuming that the CD neurons have access to saccadic motor outputs, the precue activity would create a motivational bias toward the contralateral space, even before an instruction is given by the cue stimulus.

Selective activation of the ventrolateral prefrontal cortex in the human brain during active retrieval processing.

The present study examined the role of the prefrontal cortex in retrieval processing using functional magnetic resonance imaging in human subjects. Ten healthy subjects were scanned while they performed a task that required retrieval of specific aspects of visual information. In order to examine brain activity specifically associated with retrieval, we designed a task that had retrieval and control conditions that were perfectly matched in terms of depth of encoding, decision making and postretrieval monitoring and differed only in terms of whether retrieval was required. In the retrieval condition, based on an instructional cue, the subjects had to retrieve either the particular stimulus that was previously presented or its location. In the control condition, the cue did not instruct retrieval but shared with the instructional cues the function of alerting the subjects of the impending test phase. The comparison of activity between the retrieval and control conditions demonstrated a significant and selective increase in activity related to retrieval processes within the ventrolateral prefrontal cortical region, more specifically within area 47/12. These activity increases were bilateral but stronger in the right hemisphere. The present study by strictly controlling the level of encoding, postretrieval monitoring, and decision making has demonstrated a specific increase in the ventrolateral prefrontal region that could be clearly related to active retrieval processing, i.e. the active selection of particular stored visual representations.

Self-management of instruction cues for occupation: review of studies with people with severe and profound developmental disabilities

Helping people with severe and profound developmental disabilities acquire and maintain constructive occupation is an objective of great practical importance. During the last 15–20 years, studies directed at this goal have largely relied on five strategies of self-management of instruction cues. Those strategies consist of the use of (1) picture cues presented on sets of cards, (2) picture cues stored in computer-aided systems, (3) object cues attached to cards, (4) verbal cues stored in audio recording devices, and (5) self-verbalizations. This paper reviews the aforementioned strategies and discusses their overall effectiveness and their suitability (practicality). The paper also points out some relevant issues for future research.