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In order to execute movements to targets in the environment, we must first select a target in which to move, generally from an array of alternatives. Hick's Law states that reaction time (RT) increases as the number of response alternatives increases. Violations of this law, however, generally in the form of the absence of a relationship between response alternatives and RT have been reported in the literature. K. Kveraga, L. Boucher, and H. C. Hughes (2002), for example, found that saccades to visual targets violate Hick's Law. To examine this violation further, we measured saccade RTs in monkeys and humans and found that saccade RTs actually decreased as the number of potential target locations increased. We hypothesize that this arises because subjects must actively inhibit premature saccades, and that the required inhibition increases as the certainty of a movement to a particular location increases. With increased inhibition, saccade onset is delayed, resulting in an anti-Hick's effect.

Numerous human task-switching studies have shown that decision making that follows a task switch is slower and less accurate than that which follows a task repetition. Stoet and Snyder tested humans and rhesus monkeys on a task-switching paradigm, but found, surprisingly, no switch costs in the monkeys. We hypothesized that the exceptional monkey behavior may have been due to the more extensive practice the monkeys received in comparison with human subjects. In the present study, we tested the hypothesis that extensive practice can abolish switch costs in humans. Four human subjects each performed 23,000 trials in a task-switching paradigm. We found that this amount of practice does not abolish switch costs.

The brain's sensitivity to self-generated movements is critical for behavior, and relies on accurate internal representations of movements that have been made. In the present study, we stimulated neurons below saccade threshold in the frontal eye fields of monkeys performing an oculomotor delayed response task. Stimulation during, but not before, the memory period caused small but consistent displacements of memory-guided saccade endpoints. This displacement was in the opposite direction of the saccade that was evoked by stronger stimulation at the same site, suggesting that weak stimulation induced an internal saccade signal without evoking an actual movement. Consistent with this idea, the stimulation effect was nearly absent on a task where an animal was trained to ignore self-generated eye movements. These findings support a role for the frontal eye fields in accounting for self-generated movements, and indicate that corollary discharge signals can be manipulated independent of motor output.

The traditional approach to studying brain function is to measure physiological responses to controlled sensory, motor and cognitive paradigms. However, most of the brain's energy consumption is devoted to ongoing metabolic activity not clearly associated with any particular stimulus or behaviour. Functional magnetic resonance imaging studies in humans aimed at understanding this ongoing activity have shown that spontaneous fluctuations of the blood-oxygen-level-dependent signal occur continuously in the resting state. In humans, these fluctuations are temporally coherent within widely distributed cortical systems that recapitulate the functional architecture of responses evoked by experimentally administered tasks. Here, we show that the same phenomenon is present in anaesthetized monkeys even at anaesthetic levels known to induce profound loss of consciousness. We specifically demonstrate coherent spontaneous fluctuations within three well known systems (oculomotor, somatomotor and visual) and the 'default' system, a set of brain regions thought by some to support uniquely human capabilities. Our results indicate that coherent system fluctuations probably reflect an evolutionarily conserved aspect of brain functional organization that transcends levels of consciousness.

To form an accurate internal representation of visual space, the brain must accurately account for movements of the eyes, head or body. Updating of internal representations in response to these movements is especially important when remembering spatial information, such as the location of an object, since the brain must rely on non-visual extra-retinal signals to compensate for self-generated movements. We investigated the computations underlying spatial updating by constructing a recurrent neural network model to store and update a spatial location based on a gaze shift signal, and to do so flexibly based on a contextual cue. We observed a striking similarity between the patterns of behaviour produced by the model and monkeys trained to perform the same task, as well as between the hidden units of the model and neurons in the lateral intraparietal area (LIP). In this report, we describe the similarities between the model and single unit physiology to illustrate the usefulness of neural networks as a tool for understanding specific computations performed by the brain.

Primate behavior is flexible: The response to a stimulus often depends on the task in which it occurs. Here we study how single neurons in the posterior parietal cortex (PPC) respond to stimuli which are associated with different responses in different tasks. Two rhesus monkeys performed a task-switching paradigm. Each trial started with a task cue instructing which of two tasks to perform, followed by a stimulus requiring a left or right button press. For half the stimuli, the associated responses were different in the two tasks, meaning that the task context was necessary to disambiguate the incongruent stimuli. The other half of stimuli required the same response irrespective of task context (congruent). Using this paradigm, we previously showed that behavioral responses to incongruent stimuli are significantly slower than to congruent stimuli. We now demonstrate a neural correlate in the PPC of the additional processing time required for incongruent stimuli. Furthermore, we previously found that 29% of parietal neurons encode the task being performed (task-selective cells). We now report differences in neuronal timing related to congruency in task-selective versus task nonselective cells. These differences in timing suggest that the activity in task nonselective cells reflects a motor command, whereas activity in task-selective cells reflects a decision process.

To acquire something that we see, visual spatial information must ultimately result in the activation of the appropriate set of muscles. This sensory to motor transformation requires an interaction between information coding target location and information coding which effector will be moved. Activity in the monkey parietal reach region (PRR) reflects both spatial information and the effector (arm or eye) that will be used in an upcoming reach or saccade task. To further elucidate the functional role of PRR in visually guided movement tasks and to obtain evidence that PRR signals are used to drive arm movements, we tested the hypothesis that increased neuronal activity during a preparatory delay period would lead to faster reach reaction times but would not be correlated with saccade reaction times. This proved to be the case only when the type of movement and not the spatial goal of that movement was known in advance. The correlation was strongest in cells that showed significantly more activity on arm reach compared with saccade trials. No significant correlations were found during delay periods in which spatial information was provided in advance. These data support the idea that PRR constitutes a bottleneck in the processing of spatial information for an upcoming arm reach. The lack of a correlation with saccadic reaction time also supports the idea that PRR processing is effector specific, that is, it is involved in specifying targets for arm movements but not targets for eye movements.

We previously demonstrated that the activities of neurons in the lateral intraparietal area (LIP) and the parietal reach region (PRR) of the posterior parietal cortex (PPC) are modulated by nonspatial effector-specific information. We now report similar modulation in FEF, an area of frontal cortex that is reciprocally connected with LIP. Although it is possible that these effector-specific signals originate in LIP and are conveyed to FEF, it is also possible that these signals originate in FEF and are "fed back" to LIP. We found that signal magnitude was no larger, and onset time no earlier, in FEF compared with LIP. Moreover, effector-specific activity in FEF, but not in LIP, was largely driven by spatial prediction. These results suggest that the saccade-related effector-specific signals found in LIP do not originate in FEF. Conversely, LIP may contribute to the effector-specific signals found in FEF, but does not wholly account for them.

In humans, the effects of subanesthetic doses of ketamine, an N-methyl-D-aspartate (NMDA) receptor antagonist, substantially impair executive control functions. Here, we consider whether ketamine exposure can provide an animal model for the effects of ketamine on executive control. Two monkeys (Macaca mulatta) performed a cued task-switching paradigm. We studied their behavior before and after a range of ketamine doses. We found that ketamine slowed overall performance and decreased overall accuracy, strongly impaired the capacity to ignore task-irrelevant information and, to a lesser degree, decreased accuracy when a task switch was required. This pattern of results is very similar to that found in studies of schizophrenic patients performing task-switching paradigms or the Stroop task. We conclude that ketamine in monkeys provides a good animal model for exploring the relationship between the glutamate system, executive control, and the symptoms of schizophrenia.

To examine the distribution of visual and oculomotor activity across the macaque brain, we performed functional magnetic resonance imaging (fMRI) on awake, behaving monkeys trained to perform visually guided saccades. Two subjects alternated between periods of making saccades and central fixations while blood oxygen level dependent (BOLD) images were collected [3 T, (1.5 mm)3 spatial resolution]. BOLD activations from each of four cerebral hemispheres were projected onto the subjects' cortical surfaces and aligned to a surface-based atlas for comparison across hemispheres and subjects. This surface-based analysis revealed patterns of visuo-oculomotor activity across much of the cerebral cortex, including activations in the posterior parietal cortex, superior temporal cortex and frontal lobe. For each cortical domain, we show the anatomical position and extent of visuo-oculomotor activity, including evidence that the dorsolateral frontal activation, which includes the frontal eye field (on the anterior bank of the arcuate sulcus), extends anteriorly into posterior principal sulcus (area 46) and posteriorly into part of dorsal premotor cortex (area 6). Our results also suggest that subcortical BOLD activity in the pulvinar thalamus may be lateralized during voluntary eye movements. These findings provide new neuroanatomical information as to the complex neural substrates that underlie even simple goal-directed behaviors.

We decoded on a trial-by-trial basis the location of visual targets, as a marker of the locus of attention, and intentions to reach and to saccade in different directions using the activity of neurons in the posterior parietal cortex of two monkeys. Predictions of target locations were significantly worse than predictions of movement plans for the same target locations. Moreover, neural signals in the parietal reach region (PRR) gave better predictions of reaches than saccades, whereas signals in the lateral intraparietal area (LIP) gave better predictions of saccades than reaches. Taking together the activity of both areas, the prediction of either movement in all directions became nearly perfect. These results cannot be explained in terms of an attention effect and support the idea of two segregated populations in the posterior parietal cortex, PRR and LIP, that are involved in different movement plans.

In the present study, we examined the role of frontal eye field neurons in the maintenance of spatial information in a delayed-saccade paradigm. We found that visual, visuomovement, and movement neurons conveyed roughly equal amounts of spatial information during the delay period. Although there was significant delay-period activity in individual movement neurons, there was no significant delay-period activity in the averaged population of movement neurons. These contradictory results were reconciled by the finding that the population of movement neurons with memory activity consisted of two subclasses of neurons, the combination of which resulted in the cancellation of delay-period activity in the population of movement neurons. One subclass consisted of neurons with significantly greater delay activity in the preferred than in the null direction ("canonical"), whereas the other subclass consisted of neurons with significantly greater delay activity in the null direction than in the preferred direction ("paradoxical"). Preferred direction was defined by the saccade direction that evoked the greatest movement-related activity. Interestingly, the peak saccade-related activity of canonical neurons occurred before the onset of the saccade, whereas the peak saccade-related activity of paradoxical neurons occurred after the onset of the saccade. This suggests that the former, but not the latter, are directly involved in triggering saccades. We speculate that paradoxical neurons provide a mechanism by which spatial information can be maintained in a saccade-generating circuit without prematurely triggering a saccade.

Response inhibition, or impulse control, is critical for normal cognitive function. In this issue of Neuron, Hasegawa and colleagues use a spatial nonmatch-to-sample task to reveal neurons in and around the frontal eye fields that encode where an animal should not look., (Copyright 2004 Cell Press)

The primate posterior parietal cortex (PPC), part of the dorsal visual pathway, is best known for its role in encoding salient spatial information. Yet there are indications that neural activity in the PPC can also be modulated by nonspatial task-related information. In this study, we tested whether neurons in the PPC encode signals related to cognitive set, that is, the preparation to perform a particular task. Cognitive set has previously been associated with the frontal cortex but not the PPC. In this study, monkeys performed a cognitive set shifting paradigm in which they were cued in advance to apply one of two different task rules to the subsequent stimulus on every trial. Here we show that a subset of neurons in the PPC, concentrated in the lateral bank of the intraparietal sulcus and on the angular gyrus, responds selectively to cues for different task rules., (Copyright 2004 Cell Press)

Neurons in many cortical areas involved in visuospatial processing represent remembered spatial information in retinotopic coordinates. During a gaze shift, the retinotopic representation of a target location that is fixed in the world (world-fixed reference frame) must be updated, whereas the representation of a target fixed relative to the center of gaze (gaze-fixed) must remain constant. To investigate how such computations might be performed, we trained a 3-layer recurrent neural network to store and update a spatial location based on a gaze perturbation signal, and to do so flexibly based on a contextual cue. The network produced an accurate readout of target position when cued to either reference frame, but was less precise when updating was performed. This output mimics the pattern of behavior seen in animals performing a similar task. We tested whether updating would preferentially use gaze position or gaze velocity signals, and found that the network strongly preferred velocity for updating world-fixed targets. Furthermore, we found that gaze position gain fields were not present when velocity signals were available for updating. These results have implications for how updating is performed in the brain.

We present evidence that neurons in the lateral intraparietal area (LIP) of monkey posterior parietal cortex (PPC) are activated by the instruction to make an eye movement, even in the complete absence of a spatial target. This study employed a visually guided motor task that dissociated the type of movement to make (saccade or reach) from the location where the movement was to be made. Using this task, animals were instructed to prepare a specific type of movement prior to knowing the spatial location of the movement target. We found that 25% of the LIP neurons recorded in two animals were activated significantly more by the instruction to prepare a saccade than by the instruction to prepare a reach. This finding indicates that LIP is involved in more than merely spatial attention and provides further evidence for nonspatial effector-specific signal processing in the dorsal stream.

Sensory-motor transformations are often studied using memory-guided movements to small numbers of targets. Whether target locations are directly converted into motor plans on every trial, or subjects use targets to select one of a small number of previously memorized trajectories is unknown. Well-trained monkeys made memory-guided saccades to familiar or nearby novel targets. Performance was superficially similar under the two conditions. However, saccades to novel targets close to the vertical meridian were repulsed away from the nearest familiar target. These findings suggest that sensory-to-motor transformations are performed on every trial, but that previous experience may bias the transformation.

A vertical asymmetry in memory-guided saccadic eye movements has been previously demonstrated in humans and in rhesus monkeys. In the upright orientation, saccades generally land several degrees above the target. The origin of this asymmetry has remained unknown. In this study, we investigated whether the asymmetry in memory saccades is dependent on body orientation in space. Thus animals performed memory saccades in four different body orientations: upright, left-side-down (LSD), right-side-down (RSD), and supine. Data in all three rhesus monkeys confirm previous observations regarding a significant upward vertical asymmetry. Saccade errors made from LSD and RSD postures were partitioned into components made along the axis of gravity and along the vertical body axis. Up/down asymmetry persisted only in body coordinates but not in gravity coordinates. However, this asymmetry was generally reduced in tilted positions. Therefore the upward bias seen in memory saccades is egocentric although orientation in space might play a modulatory role.

We investigated whether macaque monkeys possess the ability to prepare abstract tasks in advance. We trained two monkeys to use different stimulus-response (S-R) mappings. On each trial, monkeys were first informed with a visual cue which of two S-R mapping to use. Following a delay, a visual target was presented to which they would respond with a left or right button-press. We manipulated delay time between cue and target and found that performance was faster and more accurate with longer delays, suggesting that monkeys used the delay time to prepare each task in advance.

During a shift of gaze, an object can move along with gaze or stay fixed in the world. To examine the effect of an object's reference frame on spatial working memory, we trained monkeys to memorize locations of visual stimuli as either fixed in the world or fixed to gaze. Each trial consisted of an initial reference frame instruction, followed by a peripheral visual flash, a memory-period gaze shift, and finally a memory-guided saccade to the location consistent with the instructed reference frame. The memory-period gaze shift was either rapid (a saccade) or slow (smooth pursuit or whole body rotation). This design allowed a comparison of memory-guided saccade performance under various conditions. Our data indicate that after a rotation or smooth-pursuit eye movement, saccades to memorized world-fixed targets are more variable than saccades to memorized gaze-fixed targets. In contrast, memory-guided saccades to world- and gaze-fixed targets are equally variable following a visually guided saccade. Across all conditions, accuracy, latency, and main sequence characteristics of memory-guided saccades are not influenced by the target's reference frame. Memory-guided saccades are, however, more accurate after fast compared with slow gaze shifts. These results are most consistent with an eye-centered representational system for storing the spatial locations of memorized objects but suggest that the visual system may engage different mechanisms to update the stored signal depending on how gaze is shifted.

Executive control involves concentrating on one task without losing the ability to switch to a second task at will. We studied this ability in monkeys (Macaca mulatta) performing arbitrary stimulus-response mappings in a task-switching paradigm. We found relatively low switch costs but high task interference costs. This is the reverse of the typical human pattern of relatively large switch costs and small interference costs. This difference in the behavior of the two species may reflect anatomical differences in the sizes of the prefrontal and parietal cortices. These results indicate that monkeys are an excellent model for some but not all aspects of human task-switching.

A localized cluster of neurons in macaque posterior parietal cortex, termed the parietal reach region (PRR), is activated when a reach is planned to a visible or remembered target. To explore the role of PRR in sensorimotor transformations, we tested whether cells would be activated when a reach is planned to an as-yet unspecified goal. Over one-third of PRR cells increased their firing after an instruction to prepare a reach, but not after an instruction to prepare a saccade, when the target of the movement remained unknown. A partially overlapping population (two-thirds of cells) was activated when the monkey was informed of the target location but not the type of movement to be made. Thus a subset of PRR neurons separately code spatial and effector-specific information, consistent with a role in specifying potential motor responses to particular targets.

When primates reach for an object, they very often direct an eye movement toward the object as well. This pattern of directing both eye and limb movements to the same object appears to be fundamental to eye-hand coordination. We investigated interactions between saccades and reaching movements in a rhesus monkey model system. The amplitude and peak velocity of isolated eye movements are positively correlated with one another. This relationship is called the main sequence. We now report that the main sequence relationship for saccades is changed during coordinated eye and arm movements. In particular, peak eye velocity is approximately 4% faster for the same size saccade when the saccade is accompanied by a coordinated arm movement. Saccade duration is reduced by an equivalent amount. The main sequence relationship is unperturbed when the arm moves simultaneously but in the opposite direction as the eyes, suggesting that eye and arm movements must be tightly coordinated to produce the effect. Candidate areas mediating this interaction include the posterior parietal cortex and the superior colliculus.

Recent work on the coding of spatial information in the brain has significantly advanced our knowledge of sensory to motor transformations on several fronts. The encoding of information referenced to the retina (eye-centered) but modulated by eye position, called a gain field representation, has proved to be very common throughout parietal and occipital cortex. The use of an eye-centered representation as a working memory of spatial location is problematic if the eyes move during the memory period. Details regarding the manner in which the brain solves this problem are beginning to emerge. Finally, the discovery of eye-centered representations of ongoing or intended arm movements has changed the way we think about the order of operations in the sensory to motor coordinate transformation.

Behaviors such as sensing an object and then moving your eyes or your hand toward it require that sensory information be used to help generate a motor command, a process known as a sensorimotor transformation. Here we review models of sensorimotor transformations that use a flexible intermediate representation that relies on basis functions. The use of basis functions as an intermediate is borrowed from the theory of nonlinear function approximation. We show that this approach provides a unifying insight into the neural basis of three crucial aspects of sensorimotor transformations, namely, computation, learning and short-term memory. This mathematical formalism is consistent with the responses of cortical neurons and provides a fresh perspective on the issue of frames of reference in spatial representations.

In previous experiments, we showed that cells in the parietal reach region (PRR) in monkey posterior parietal cortex code intended reaching movements in an eye-centered frame of reference. These cells are more active when an arm compared with an eye movement is being planned. Despite this clear preference for arm movements, we now report that PRR neurons also fire around the time of a saccade. Of 206 cells tested, 29% had perisaccadic activity in a delayed-saccade task. Two findings indicate that saccade-related activity does not reflect saccade planning or execution. First, activity is often peri- or postsaccadic but seldom presaccadic. Second, cells with saccade-related activity were no more likely to show strong saccadic delay period activity than cells without saccade-related activity. These findings indicate that PRR cells do not take part in saccade planning. Instead, the saccade-related activity in PRR may reflect cross-coupling between reach and saccade pathways that may be used to facilitate eye-hand coordination. Alternatively, saccade-related activity may reflect eye position information that could be used to maintain an eye-centered representation of intended reach targets across eye movements.

Over the last few years it is becoming increasingly apparent that an important role of the posterior parietal cortex is to process sensory information for the purpose of planning actions. We review studies showing that a large component of neural activity in area LIP is related to planning saccades and activity in a nearby parietal reach region (PRR) to reaches. This intention related activity dominates the delay period in delayed movement tasks, and also comprises a substantial component of the transient response. These findings, along with additional anatomical and physiological evidence, lends support to the idea that different cortical areas within the PPC represent plans for different actions. We also found strong modulation of activity when movement plans were changed without changes in the locus of attention. This result suggests that PPC, which has been postulated to play a role in shifting attention, may also play a role in changing movement intentions. Sensory related activity was also present in these tasks and may be related to the stimulus or to attention. These experiments show that there are intention and sensory related activities in the PPC consistent with its proposed role in sensory-motor transformations. These studies also show that care must be taken to measure intention-related signals and not assume that all task dependent modulation in the PPC reflects attention.

The neural events associated with visually guided reaching begin with an image on the retina and end with impulses to the muscles. In between, a reaching plan is formed. This plan could be in the coordinates of the arm, specifying the direction and amplitude of the movement, or it could be in the coordinates of the eye because visual information is initially gathered in this reference frame. In a reach-planning area of the posterior parietal cortex, neural activity was found to be more consistent with an eye-centered than an arm-centered coding of reach targets. Coding of arm movements in an eye-centered reference frame is advantageous because obstacles that affect planning as well as errors in reaching are registered in this reference frame. Also, eye movements are planned in eye coordinates, and the use of similar coordinates for reaching may facilitate hand-eye coordination.

Vestibular signals play an important role in spatial orientation, perception of object location, and control of self-motion. Prior physiological research on vestibular information processing has focused on brainstem mechanisms; relatively little is known about the processing of vestibular information at the level of the cerebral cortex. Recent electrophysiological experiments examining the use of vestibular canal signals in two different perceptual tasks are described: computation of self motion and localization of visual stimuli in a world-centered reference frame. These two perceptual functions are mediated by different parts of the posterior parietal cortex, the former in the dorsal aspect of the medial superior temporal area (MSTd) and the latter in area 7a.

In order to direct a movement towards a visual stimulus, visual spatial information must be combined with postural information. For example, directing gaze (eye plus head) towards a visible target requires the combination of retinal image location with eye and head position to determine the location of the target relative to the body. Similarly, world-referenced postural information is required to determine where something lies in the world. Posterior parietal neurons recorded in monkeys combine visual information with eye and head position. A population of such cells could make up a distributed representation of target location in an extraretinal frame of reference. However, previous studies have not distinguished between world-referenced and body-referenced signals. Here we report that modulations of visual signals (gain fields) in two adjacent cortical fields, LIP and 7a, are referenced to the body and to the world, respectively. This segregation of spatial information is consistent with a streaming of information, with one path carrying body-referenced information for the control of gaze, and the other carrying world-referenced information for navigation and other tasks that require an absolute frame of reference.

The lateral intraparietal area (LIP) of macaque monkey, and a parietal reach region (PRR) medial and posterior to LIP, code the intention to make visually guided eye and arm movements, respectively. We studied the effect of changing the motor plan, without changing the locus of attention, on single neurons in these two areas. A central target was fixated while one or two sequential flashes occurred in the periphery. The first appeared either within the response field of the neuron being recorded or else on the opposite side of the fixation point. Animals planned a saccade (red flash) or reach (green flash) to the flash location. In some trials, a second flash 750 ms later could change the motor plan but never shifted attention: second flashes always occurred at the same location as the preceding first flash. Responses in LIP were larger when a saccade was instructed (n = 20 cells), whereas responses in PRR were larger when a reach was instructed (n = 17). This motor preference was observed for both first flashes and second flashes. In addition, the response to a second flash depended on whether it affirmed or countermanded the first flash; second flash responses were diminished only in the former case. Control experiments indicated that this differential effect was not due to stimulus novelty. These findings support a role for posterior parietal cortex in coding specific motor intention and are consistent with a possible role in the nonspatial shifting of motor intention.

The posterior parietal cortex (PPC) has long been considered a sensory area specialized for spatial awareness and the directing of attention. However, a new, far reaching concept is now emerging that this area is involved in integrating sensory information for the purpose of planning action. Moreover, experiments by our group and others over the last two decades indicate that PPC is in fact anatomically organized with respect to action. PPC also is an 'association' cortex which must combine different sensory modalities which are coded in different coordinate frames. We have found, at least for two different cortical areas within PPC, that different sensory signals are brought into a common coordinate frame. This coordinate frame codes locations with respect to the eye, but also gain modulates the activity by eye and body position signals. An interesting feature of this coordinate representation at the population level is that it codes concurrently target locations in multiple coordinate frames (eye, head, body and world). Depending on how this population of neurons is sampled, different coordinate transformations can be accomplished by the same population of neurons.

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.

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.

The vestibulo-ocular reflex (VOR) has historically been considered a computationally simple reflex: to stabilize images on the retina against imposed head rotation, the eyes must be counterrotated by an equal amount in the opposite direction. During almost any head rotation, however, the eyes are also translated. We show that the VOR compensates for 90% of this translation, and suggest a computational scheme by which this is done, based on a temporal dissection of the VOR response to sudden head rotation. An initial response that corrects only for imposed rotation is refined by a series of three temporally delayed corrections of increasing complexity. The first correction takes only head rotation and viewing distance into account; the second, head rotation, viewing distance, and otolith translation; and the third, head rotation, viewing distance, otolith translation, and translation of the eyes relative to the otoliths. Responses of type I gaze velocity Purkinje (GVP) cells in the cerebellar flocculus and ventral paraflocculus of rhesus monkeys were recorded during sudden head rotation. We show that cell discharge was modulated both by axis location and by viewing distance, suggesting that GVP cells play a role in the VOR response to rotation-induced eye translation.

The mechanism for object location in the environment, and the perception of the external world as stable when eyes, head and body are moved, have long been thought to be centred on the posterior parietal cortex. However, head position signals, and their integration with visual and eye position signals to form a representation of space referenced to the body, have never been examined in any area of the cortex. Here we show that the visual and saccadic activities of parietal neurons are strongly affected by head position. The eye and head position effects are equivalent for individual neurons, indicating that the modulation is a function of gaze direction, regardless of whether the eyes or head are used to direct gaze. These data are consistent with the idea that the posterior parietal cortex contains a distributed representation of space in body-centred coordinates.

Coordinate transformations are an essential aspect of behavior. They are required because sensory information is coded in the coordinates of the sensory epithelia (e.g. retina, skin) and must be transformed to the coordinates of muscles for movement. In this review we will concentrate on recent studies of visual-motor transformations. The studies show that representations of space are distributed, being specified in the activity of many cells rather than in the activity of individual cells. Furthermore, these distributed representations appear to be derived by a specific operation, which systematically combines visual signals with eye and head position signals.

1. The vestibuloocular reflex (VOR) stabilizes images on the retina against movements of the head in space. Viewing distance, target eccentricity, and location of the axis of rotation may influence VOR responses because rotation of the head about most axes in space rotates and translates the eyes relative to visual targets. To study the VOR response to combined rotation and translation, monkeys were placed on a rate table and rotated briefly in the dark about a vertical axis that was located in front of or behind the eyes. The monkeys fixated a near or far visual target that was extinguished before the rotation. Eye movements were recorded from both eyes by the use of the search coil technique. 2. Peak eye velocity evoked by the VOR was linearly related to vergence angle for any axis of rotation. The percent change in the VOR with near target viewing relative to far target viewing at a vergence angle of 20 degrees was linearly related to the location of the axis of rotation. Axes located behind the eyes produced positive changes in VOR amplitude, and axes located in front of the eyes produced negative changes in VOR amplitude. An axis of rotation located in the coronal plane containing the centers of rotation of the eyes produced no modification of VOR amplitude. For any axis, the VOR compensated for approximately 90% of the translation of the eye relative to near targets. 3. The initial VOR response was not correct in magnitude but was refined by a series of three temporally delayed corrections of increasing complexity. The earliest VOR-evoked eye movement (10-20 ms after rotation onset) was independent of viewing distance and rotational axis location. In the next 100 ms, eye speed appeared to be sequentially modified three times: within 20 ms by viewing distance; within 30 ms by otolith translation; and within 100 ms by eye translation relative to the visual target. 4. These data suggest a formal model of the VOR consisting of four channels. Channel 1 conveys an unmodified head rotation signal with a pure delay of 10 ms. Channel 2 conveys an angular head velocity signal, modified by viewing distance with a pure delay of 20 ms, but invariant with respect to the location of the axis of rotation. Channel 3 conveys a linear head velocity signal, dependent on the location of the axis of rotation, that is modified by viewing distance with a pure delay of 30 ms.(ABSTRACT TRUNCATED AT 400 WORDS)

The axis of head rotation is usually different from the axis of eye rotation. Geometrical considerations show that the eyes translate with respect to visual targets located near the head. In order to accurately stabilize retinal images against this translation, the vestibulo-ocular reflex (VOR) must be modulated inversely with the target's distance. The neural correlate of viewing distance used to modulate the VOR is not known. Since vergence angle is related inversely to viewing distance, an efference copy of instantaneous vergence angle or proprioceptive afferent information from extraocular muscles could be used to adjust the VOR. To examine this hypothesis, we compared the time-course of changes in the VOR with the time-course of changes in vergence angle. The VOR was induced by briefly rotating monkeys about a vertical axis at different times during the execution of vergence eye movements. We found that the amplitude of the VOR changed systematically during the course of a vergence eye movement. On average, the changes in the amplitude of vestibular induced eye movements anticipated changes in vergence angle by 50 msec, but in some instances, up to 200 msec of anticipation was observed. These data suggest that a central command signal rather than an afferent or efferent copy of vergence eye position was used to modulate the VOR.

1. We studied eye velocity during the first 2 s of the vertical vestibuloocular reflex (VOR) elicited from cats placed on their sides (90 degrees roll position) and rotated about an earth vertical axis. Vestibular stimuli were presented in the dark and consisted of brief trapezoidal velocity profiles. Eye movements were recorded with a magnetic search coil, and eye velocity was analyzed with high temporal resolution. 2. The first 2 s of upward or downward eye velocity after the onset of head rotation was characterized and compared. Adaptive changes in VOR gain (eye/head velocity) were then induced, and upward and downward eye velocity responses were again compared. 3. The early time course of the vertical VOR was complex. After a latency of approximately 15 ms, eye velocity increased rapidly until it was equal in magnitude and opposite in direction to head velocity. The peak eye velocity decayed within less than 1 s to a plateau of slow-phase eye velocity (SPEV) equal to approximately -0.6 times the head velocity. Peak upward and downward eye velocity was symmetric. The transition from peak to plateau was more rapid for the downward VOR (slow phases downward) than for the upward VOR (slow phases upward). The plateau attained by upward SPEV was approximately 15% higher than the plateau attained by downward SPEV. 4. VOR gain adaptation was symmetric. The percentage change in adapted upward eye velocity equalled the percentage change in adapted downward eye velocity. Both peak and plateau SPEV adapted, but peak eye velocity adapted less than plateau eye velocity. VOR latency was unchanged by adaptation. 5. The trajectory of the VOR response to steps of head velocity could be divided into an invariant and a variant interval. The invariant interval consisted of the initial approximately 15 ms of the eye movement. Neither direction of head movement (upward vs. downward) nor adaptation of the VOR gain effected the eye movement trajectory during the invariant interval. The variant interval began approximately 30 ms after the onset of head movement and approximately 15 ms after the onset of eye movement. In unadapted animals, downward eye speed exceeded upward eye speed during the variant interval. In adapted animals, eye speed during the variant interval, but not during the invariant interval, diverged from eye speed in the unadapted state. We suggest that the initial invariant interval (approximately 15 ms) of the eye movement response trajectory may represent the direct response of the classically described three-neuron arc.(ABSTRACT TRUNCATED AT 400 WORDS)