Schema cells in the macaque hippocampus.

Concept cells in the human hippocampus encode the meaning conveyed by stimuli over their perceptual aspects. Here we investigate whether analogous cells in the macaque can form conceptual schemas of spatial environments. Each day, monkeys were presented with a familiar and a novel virtual maze, sharing a common schema but differing by surface features (landmarks). In both environments, animals searched for a hidden reward goal only defined in relation to landmarks. With learning, many neurons developed a firing map integrating goal-centered and task-related information of the novel maze that matched that for the familiar maze. Thus, these hippocampal cells abstract the spatial concepts from the superficial details of the environment and encode space into a schema-like representation.

Leftward oculomotor prismatic training induces a rightward bias in normal subjects.

Wedge prisms shifting the visual field laterally create a mismatch between the straight ahead position signalled by vision and that encoded by extraretinal and head-on-trunk proprioceptive information. Short-term adaptation to left-deviating prisms in normal subjects results in a visuomotor attentional bias towards the right-hand side (aftereffect). Prismatic adaptation (PA) is usually induced through a training consisting in repeated ballistic movements of the dominant arm towards visual targets, while participants are wearing prismatic goggles. The present study demonstrates that an original oculomotor PA procedure with leftward deviating prisms-without pointing movements and only consisting in repeated gaze shifts towards visual targets-can induce a rightward bias in normal subjects as assessed by visual straight ahead and line bisection tasks (Experiments 1 and 2). We show that oculomotor PA induces a bias in line bisection similar to that reported after visuomotor PA (Experiment 2). We suggest that a conflict between retinal, extraretinal and proprioceptive information about the straight ahead location causes the observed effects. In follow-up experiments 3, 4, and 5, we demonstrate that neither eye deviation without prisms nor shift of the visual field without eye deviation induces PA biases. We propose that an optimal integration model of visual and proprioceptive inputs can best account for the observed results.

A real-time 3D video tracking system for monitoring primate groups.

To date, assessing the solitary and social behaviors of laboratory primates' colonies relies on time-consuming manual scoring methods. Here, we describe a real-time multi-camera 3D tracking system developed to measure the behavior of socially-housed primates. Their positions are identified using non-invasive color markers such as plastic collars, thus allowing to also track colored objects and to measure their usage. Compared to traditional manual ethological scoring, we show that this system can reliably evaluate solitary behaviors (foraging, solitary resting, toy usage, locomotion) as well as spatial proximity with peers, which is considered as a good proxy of their social motivation. Compared to existing video-based commercial systems currently available to measure animal activity, this system offers many possibilities (real-time data, large volume coverage, multiple animal tracking) at a lower hardware cost. Quantitative behavioral data of animal groups can now be obtained automatically over very long periods of time, thus opening new perspectives in particular for studying the neuroethology of social behavior in primates.

The role of the right parietal lobe in anorexia nervosa.

BACKGROUND: Patients with anorexia nervosa (AN) overestimate their size despite being severely underweight. Whether this misperception echoes an underlying emotional disturbance or also reflects a genuine body-representation deficit is debatable. Current measures inquire directly about subjective perception of body image, thus distinguishing poorly between top-down effects of emotions/attitudes towards the body and disturbances due to proprioceptive disorders/distorted body schema. Disorders of body representation also emerge following damage to the right parietal lobe. The possibility that parietal dysfunction might contribute to AN is suspected, based on the demonstrated association of spatial impairments, comparable to those found after parietal lesion, with this syndrome. METHOD: We used a behavioral task to compare body knowledge in severe anorexics (n=8), healthy volunteers (n=11) and stroke patients with focal damage to the left/right parietal lobe (n=4). We applied a psychophysical procedure based on the perception, in the dark, of an approaching visual stimulus that was turned off before reaching the observer. Participants had to predict whether the stimulus would have hit/missed their body, had it continued its linear motion. RESULTS: Healthy volunteers and left parietal patients estimated body boundaries very close to the real ones. Conversely, anorexics and right parietal patients underestimated eccentricity of their left body boundary. CONCLUSIONS: These findings are in line with the role the parietal cortex plays in developing and maintaining body representation, and support the possibility for a neuropsychological component in the pathogenesis of anorexia, offering alternative approaches to treatment of the disorder.

Parallel integral projection transform for straight electrode localization in 3-D ultrasound images.

In surgical practice, small metallic instruments are frequently used to perform various tasks inside the human body. We address the problem of their accurate localization in the tissue. Recent experiments using medical ultrasound have shown that this modality is suitable for real-time visualization of anatomical structures as well as the position of surgical instruments. We propose an image-processing algorithm that permits automatic estimation of the position of a line-segment-shaped object. This method was applied to the localization of a thin metallic electrode in biological tissue. We show that the electrode axis can be found through maximizing the parallel integral projection transform that is a form of the Radon transform. To accelerate this step, hierarchical mesh-grid algorithm is implemented. Once the axis position is known, localization of the electrode tip is performed. The method was tested on simulated images, on ultrasound images of a tissue mimicking phantom containing a metallic electrode, and on real ultrasound images from breast biopsy. The results indicate that the algorithm is robust with respect to variations in electrode position and speckle noise. Localization accuracy is of the order of hundreds of micrometers and is comparable to the ultrasound system axial resolution.

The influence of hand posture on corticospinal excitability during motor imagery: a transcranial magnetic stimulation study.

In order to study the interaction between proprioceptive information and motor imagery, we herein investigate how compatible and incompatible postural signals influence corticospinal excitability during the mental simulation of hand movements. Subjects were asked to imagine themselves joining the tips of the thumb and the little finger while they maintained one of the two following hand postures: posture A (PA, compatible), little finger, index and thumb extended, the remaining fingers flexed; or posture B (PB, incompatible), index and thumb extended, other fingers flexed. All subjects rated the imagined finger opposition movements as easier to perform when the hand was kept in PA than in PB (P < 0.01) and the correlation between the duration of motor imagery and movement execution was also higher for PA than PB (P < 0.01). For each posture, motor evoked potentials (MEPs) elicited by focal transcranial magnetic stimulation (TMS) of the left motor cortex were recorded from the right opponens pollicis muscle during both motor imagery (MI) and rest (R) conditions. MEP area varied according to the hand posture: PA induced a higher increase in corticospinal excitability, when compared with PB. These results indicate that the actual limb posture affects the process of motor imagery. The source of this postural modulation effect is discussed.

Ocular fixation and visual activity in the monkey lateral intraparietal area.

The macaque lateral intraparietal area (LIP) has been implicated in visuospatial attention and saccade planning. Since area LIP also contains a representation of the central visual field, we investigated its possible role in fixation and foveal attention in a visual fixation task with gap (momentary disappearance of fixation point). In addition to the expected visual neurons ( n=119), two main categories were identified: (1) cells responding tonically both during the presence and momentary absence of the fixation stimulus( n=47); a subset of these neurons studied in a saccade task showed perisaccadic inhibition in half of the cases (14/27). The timing of this inhibition, however, is only loosely related to saccade timing; (2) cells responding mainly to the absence of the fixation stimulus, with either abrupt or gradual onset of activity during the gap ( n=62). During saccades, these neurons showed presaccadic buildup and/or postsaccadic activity, which was spatially tuned in about half of the tested cells (28/53). Ninety-one percent of the cells in the first category and 59% of the cells in the second category were located in the dorsal portion of area LIP (LIPd). These results are consistent with the hypothesis of an oculomotor-attentional network contributing to fixation engagement and disengagement in a subregion of LIP.

Visual receptive field modulation in the lateral intraparietal area during attentive fixation and free gaze.

The receptive field (RF) of neurons recorded from the lateral intraparietal area (LIP) was quantified using a rapid, computer-driven mapping procedure. For each neuron, the RF was mapped: (1) during attentive fixation and (2) during free visual exploration. RF location, size and internal structure were modulated by the mapping context in over two-thirds of the recorded neurons. The major trend was a proportionally larger amount of neuronal visual resources allocated to central space during fixation, and an attenuated center-to-periphery gradient in the visual field representation during free gaze. A population approach shows that these spatial modulations are accompanied by changes in the signal-to-noise ratio of the information carried in the RF substructure. We related these neurophysiological observations to behavior, by comparing the characteristics of saccades elicited during fixation and free gaze. Together, the results suggest that the dynamics of LIP visual RFs may characterize both the state of engagement of attention and the power of resolution of visual analysis: during fixation, the neural population is locked in a filter state concentrating the processing resources at the fovea, while during free gaze, the population shifts to a detector state spreading the resources more evenly across the visual field.

Motor and visual imagery as two complementary but neurally dissociable mental processes.

Recent studies indicate that covert mental activities, such as simulating a motor action and imagining the shape of an object, involve shared neural representations with actual motor performance and with visual perception, respectively. Here we investigate the performance, by normal individual and subjects with a selective impairment in either motor or visual imagery, of an imagery task involving a mental rotation. The task involved imagining a hand in a particular orientation in space and making a subsequent laterality judgement. A simple change in the phrasing of the imagery instructions (first-person or third-person imagery) and in actual hand posture (holding the hands on the lap or in the back) had a strong impact on response time (RT) in normal subjects, and on response accuracy in brain-damaged subjects. The pattern of results indicates that the activation of covert motor and visual processes during mental imagery depends on both top-down and bottom-up factors, and highlights the distinct but complementary contribution of covert motor and visual processes during mental rotation.

Representation of the visual field in the lateral intraparietal area of macaque monkeys: a quantitative receptive field analysis.

The representation of the visual field in the primate lateral intraparietal area (LIP) was examined, using a rapid, computer-driven receptive field (RF) mapping procedure. RF characteristics of single LIP neurons could thus be measured repeatedly under different behavioral conditions. Here we report data obtained using a standard ocular fixation task during which the animals were required to monitor small changes in color of the fixated target. In a first step, statistical analyses were conducted in order to establish the experimental limits of the mapping procedure on 171 LIP neurons recorded from three hemispheres of two macaque monkeys. The characteristics of the receptive fields of LIP neurons were analyzed at the single cell and at the population level. Although for many neurons the assumption of a simple two-dimensional gaussian profile with a central area of maximal excitability at the center and progressively decreasing response strength at the periphery can represent relatively accurately the spatial structure of the RF, about 19% of the cells had a markedly asymmetrical shape. At the population level, we observed, in agreement with prior studies, a systematic relation between RF size and eccentricity. However, we also found a more accentuated overrepresentation of the central visual field than had been previously reported and no marked differences between the upper and lower visual representation of space. This observation correlates with an extension of the definition of LIP from the posterior third of the lateral intraparietal sulcus to most of the middle and posterior thirds. Detailed histological analyses of the recorded hemispheres suggest that there exists, in this newly defined unitary functional cortical area, a coarse but systematic topographical organization in area LIP that supports the distinction between its dorsal and ventral regions, LIPd and LIPv, respectively. Paralleling the physiological data, the central visual field is mostly represented in the middle dorsal region and the visual periphery more ventral and posterior. An anteroposterior gradient from the lower to the upper visual field representations can also be identified. In conclusion, this study provides the basis for a reliable mapping method in awake monkeys and a reference for the organization of the properties of the visual space representation in an area LIP extended with respect to the previously described LIP and showing a relative emphasis of central visual field.

Space coding in primate posterior parietal cortex.

Neuropsychological studies of patients with lesions of right frontal (premotor) or posterior parietal cortex often show severe impairments of attentive sensorimotor behavior. Such patients frequently manifest symptoms like hemispatial neglect or extinction. Interestingly, these behavioral deficits occur across different sensory modalities and are often organized in head- or body-centered coordinates. These neuropsychological data provide evidence for the existence of a network of polymodal areas in (primate) premotor and inferior parietal cortex representing visual spatial information in a nonretinocentric frame of reference. In the monkey, a highly modular structural and functional specialization has been demonstrated especially within posterior parietal cortex. One such functionally specialized area is the ventral intraparietal area (VIP). This area is located in the fundus of the intraparietal sulcus and contains many neurons that show polymodal directionally selective discharges, i.e., these neurons respond to moving visual, tactile, vestibular, or auditory stimuli. Many of these neurons also encode sensory information from different modalities in a common, probably head-centered, frame of reference. Functional imaging data on humans reveal a network of cortical areas that respond to polymodal stimuli conveying motion information. One of these regions of activation is located in the depth of human intraparietal sulcus. Accordingly, it is suggested that this area constitutes the human equivalent of monkey area VIP. The functional role of area VIP for polymodal spatial perception in normals as well as the functional implications of lesions of area VIP in parietal patients needs to be established in further experiments.

Eye position encoding in the macaque ventral intraparietal area (VIP).

Many neurons in area VIP encode the location of visual stimuli in a non-retinocentric frame of reference. In this context the question needed to be addressed whether the underlying coordinate transformation of the incoming visual signals could be generated within area VIP or whether this information would have to arrive from other areas. We tested 74 neurons in area VIP of two awake monkeys for an influence of eye position while animals performed a fixation task. More than half of the neurons (40/74) revealed an eye position effect. At the population level, however, this effect was balanced out. We suggest that local connections within area VIP could be used to generate an encoding of visual information in a non-retinocentric frame of reference.

Ventral intraparietal area of the macaque: congruent visual and somatic response properties.

In a previous report, we described the visual response properties in the ventral intraparietal area (area VIP) of the awake macaque. Here we describe the somatosensory response properties in area VIP and the patterns of correspondence between the responses of single neurons to independently administered tactile and visual stimulation. VIP neurons responded to visual stimulation only or to visual and tactile stimulation. Of 218 neurons tested, 153 (70%) were bimodal in the sense that they responded to stimuli that were independently applied in either sensory modality. Unimodal visual and bimodal neurons were intermingled within the recording area and could not be distinguished on the basis of their visual response properties alone. Most of the cells with a tactile receptive field (RF) responded well to light touch or air puffs. The distribution of RF locations principally emphasized the head (85%), with approximately equivalent representations of the upper and lower face areas. The tactile and visual RFs were aligned in a congruent manner, with the intersection of the visual vertical and horizontal meridian having its tactile counterpart in the nose/mouth area. Small foveal visual RFs were paired with small tactile RFs on the muzzle, and peripheral visual RFs were associated with tactile RFs on the side of the head or body. Most cells showed a strong sensitivity to moving stimuli, and the preferred directions of visual and tactile motion coincided in 85% of bimodal cells. In some cases, bimodal responses patterns were complementary: cells responding to motion in depth toward the monkey had responses, whereas cells responding to motion in depth away form the monkey had responses. Other forms of bimodal response congruence included orientation selectivity, and , , and / response types. The large proportion of bimodal tactile and visual neurons with congruent response properties in area VIP indicates that there are important functional differences between area VIP and other dorsal stream areas involved in the analysis of motion. We suggest that VIP is involved in the construction of a multisensory, head-centered representation of near extrapersonal space.

Spatial invariance of visual receptive fields in parietal cortex neurons.

Spatial information is conveyed to the primary visual cortex in retinal coordinates. Movement trajectory programming, however, requires a transformation from this sensory frame of reference into a frame appropriate for the selected part of the body, such as the eye, head or arms. To achieve this transformation, visual information must be combined with information from other sources: for instance, the location of an object of interest can be defined with respect to the observer's head if the position of the eyes in the orbit is known and is added to the object's retinal coordinates. Here we show that in a subdivision of the monkey parietal lobe, the ventral intraparietal area (VIP), the activity of visual neurons is modulated by eye-position signals, as in many other areas of the cortical visual system. We find that individual receptive fields of a population of VIP neurons are organized along a continuum, from eye to head coordinates. In the latter case, neurons encode the azimuth and/or elevation of a visual stimulus, independently of the direction in which the eyes are looking, thus representing spatial locations explicitly in at least a head-centred frame of reference.

Spatial representations for action in parietal cortex.

Parietal cortex contains multiple representations of visual space. Single neurons in area LIP encode attended locations relative to the fovea, while some VIP neurons encode stimulus location relative to the head and some MIP neurons may encode location relative to the arm. These multiple representations are tailored to guide specific kinds of actions: eye movements, head movements and arm movements, respectively. The function of parietal cortex is to signal the location of attended objects relative to the observer. It does so in order to allow the organism to act on its environment. The many different kinds of actions that can be performed are likely to be supported by these very different kinds of spatial representations.

Visual, presaccadic, and cognitive activation of single neurons in monkey lateral intraparietal area.

1. Posterior parietal cortex contains neurons that are visually responsive and active in relation to saccadic eye movements. We recorded from single neurons in a subregion of parietal cortex, the lateral intraparietal area (LIP), in alert rhesus monkeys. To characterize more completely the circumstances under which LIP neurons are responsive, we used five tasks designed to test the impact of sensory, motor, and cognitive factors. We obtained quantitative data in multiple tasks in 91 neurons. We measured neural activity during central fixation and in relation to stimulus onset and saccade onset. 2. LIP neurons have visual responses to the onset of a stationary stimulus in the receptive field. These visual responses occurred both in tasks that require a subsequent eye movement toward the stimulus and in tasks in which eye movements are not permitted, indicating that this activity is sensory rather than presaccadic. 3. Visual responses were enhanced when the monkey had to use information provided by the stimulus to guide its behavior. The amplitude of the sensory response to a given stimulus was increased in a task in which the monkey would subsequently make a saccade to the location signaled by the stimulus, as compared with the amplitude of the visual response in a simple fixation task. 4. The visual response was also enhanced when the monkey attended to the stimulus without looking at it. This result shows that enhancement does not reflect saccade preparation because the response is enhanced even when the monkey is not permitted to make a saccade. Instead, enhancement reflects the allocation of attention to the spatial locus of the receptive field. 5. Many LIP neurons had saccade-related activity in addition to their visual responses. The visual response for most neurons was stronger than the saccade-related activation. 6. Saccade-related activity was independent of visual activity. Similar presaccadic activity was observed in trials that included a recent visual stimulus (memory-guided saccade task) and in trials with no visual stimulus (learned saccade task). 7. We observed increases in activity during fixation in tasks in which the monkey could anticipate the onset of a behaviorally significant stimulus. LIP neurons usually showed low levels of background firing in the fixation task during the period before stimulus onset. This background activity was increased in the peripheral attention and memory-guided saccade tasks during the period when the monkey was waiting for a behaviorally relevant stimulus to appear. 8. The results from these several tasks indicate that LIP neurons are activated in a variety of circumstances and are not involved exclusively in sensory processing or motor planning. The modulation of sensory responses by attention and anticipation suggests that cognitive factors play a major role in parietal function.

The mental representation of hand movements after parietal cortex damage.

Recent neuroimagery findings showed that the patterns of cerebral activation during the mental rehearsal of a motor act are similar to those produced by its actual execution. This concurs with the notion that part of the distributed neural activity taking place during movement involves internal simulations, but it is not yet clear what specific contribution the different brain areas involved bring to this process. Here, patients with lesions restricted to the parietal cortex were found to be impaired selectively at predicting, through mental imagery, the time necessary to perform differentiated finger movements and visually guided pointing gestures, in comparison to normal individuals and to a patient with damage to the primary motor area. These results suggest that the parietal cortex is important for the ability to generate mental movement representations.

Oculocentric spatial representation in parietal cortex.

Parietal cortex comprises several distinct areas. Neurons in each area are selective for particular stimulus dimensions and particular regions of space. The representation of space in a given area reflects a particular motor output by which a stimulus can be acquired. Neurons in the lateral intraparietal area (LIP) are active in relation to both visual and motor events. LIP neurons do not transmit an unambiguous saccadic command. Rather, they signal the location at which an event has occurred. These spatial locations are encoded in oculocentric coordinates, that is, with respect to the current or anticipated position of the center of gaze. When an eye movement brings the spatial location of a recently flashed stimulus into the receptive field of an LIP neuron, the neuron responds to the memory trace of that stimulus. This result indicates that, for nearly all LIP neurons, stored visual information is remapped in conjunction with saccades. Remapping of the memory trace maintains the alignment between the current image on the retina and the stored representation in cortex. Further, when an eye movement is about to occur, more than a third of LIP neurons transiently shift the location of their receptive fields. This anticipatory remapping allows the neuron to begin to respond to a visual stimulus even before the saccade is initiated that will bring the stimulus into the fixation-defined receptive field. Both kinds of remapping serve to create a constantly updated representation of stimulus location that is always in terms of distance and direction from the fovea. This oculocentric representation has the advantage that it already matches that known to exist in the frontal eye fields and the superior colliculus, the output targets of LIP, and it does not require further coordinate transformation in order to contribute to spatially accurate behavior. These results indicate that LIP can analyze visual space without ever forming a representation of absolute target position.

Congruent unilateral impairments for real and imagined hand movements.

The chronometry of imagined and actual movements was investigated in a patient with a unilateral lesion of the motor cortex. Motor imagery generated highly accurate estimates of motor performance in a variety of situations, reflecting the hypokinesia of the contralesional hand. There were parallel increases in mental and actual movement times from proximal to distal limb segments. Bimanual movements adopted the slower speed of the impaired hand in both conditions. Imagined motor sequences to the beat of a metronome predicted the maximum speed reached in actual performance. Finally, visually guided pointing showed the same target-size effects in the imagery and movement conditions. The results are in agreement with the hypothesis that common cerebral motor representations are activated when imaging and planning voluntary movements.