Retinotopic coding of extraretinal pursuit signals in early visual cortex.

During smooth pursuit, the image of the target is stabilized on the fovea, implying that speed judgments made during pursuit must rely on an extraretinal signal providing precise eye speed information. To characterize the introduction of such extraretinal signal into the human visual system, we performed a factorial, functional magnetic resonance imaging experiment, in which we manipulated the factor eye movement, with "fixation" and "pursuit" as levels, and the factor task, with "speed" and "form" judgments as levels. We hypothesized that the extraretinal speed signal is reflected as an interaction between speed judgments and pursuit. Random effects analysis yielded an interaction only in dorsal early visual cortex. Retinotopic mapping localized this interaction on the horizontal meridian (HM) between dorsal areas visual 2 and 3 (V2/V3) at 1-2 degrees azimuth. This corresponded to the position the pursuit target would have reached, if moving retinotopically, at the time of the subject's speed judgment. Because the 2 V2/V3 HMs are redundant, both may be involved in speed judgments, the ventral one involving judgments based on retinal motion and the dorsal one judgments requiring an internal signal. These results indicate that an extraretinal speed signal is injected into early visual cortex during pursuit.

Different spatial memory systems are involved in small- and large-scale environments: evidence from patients with temporal lobe epilepsy.

Recent reports show that humans and animals do not acquire information about routes and object locations in the same way. In spatial memory, a specific sub-system is hypothesized to be involved in encoding, storing and recalling navigational information, and it is segregated from the sub-system devoted to small-scale environment. We assessed this hypothesis in a sample of patients treated surgically for intractable temporal lobe epilepsy. We found double dissociations between learning and recall of spatial positions in large space versus small space. These results strongly support the hypothesis that two segregate systems process navigational memory for large-scale environments and spatial memory in small-scale environments.

Spatial memory of body linear displacement: what is being stored?

The ability to evaluate traveled distance is common to most animal species. Head trajectory in space is measured on the basis of the converging signals of the visual, vestibular, and somatosensory systems, together with efferent copies of motor commands. Recent evidence from human studies has shown that head trajectory in space can be stored in spatial memory. A fundamental question, however, remains unanswered: How is movement stored? In this study, humans who were asked to reproduce passive linear whole-body displacement distances while blindfolded were also able to reproduce velocity profiles. This finding suggests that a spatiotemporal dynamic pattern of motion is stored and can be retrieved with the use of vestibular and somesthetic cues.

Effects of rectilinear acceleration and optokinetic and caloric stimulations in space.

During the flight of Spacelab 1 the crew performed a number of experiments to explore changes in vestibular function and visual-vestibular interactions on exposure to microgravity. Measurements were made on the threshold for detection of linear oscillation, vestibulo-ocular reflexes elicited by angular and linear movements, oculomotor and posture responses to optokinetic stimulations, and responses to caloric stimulation. Tests were also conducted on the ground, during the 4 months before and on days 1 to 6 after flight. The most significant result was that caloric mystagmus of the same direction as on the earth could also be evoked in the weightless environment.

Distinct visual perspective-taking strategies involve the left and right medial temporal lobe structures differently.

This study assesses the role of the human medial temporal lobe (MTL) structures in the coordination of spatial information across perspective change and, in particular, in visual perspective taking--namely the capacity to know what another individual is seeing on the visual scene. Fourteen patients with unilateral temporal lobe resection and 21 control subjects performed two tasks, called 'object location memory' and 'viewpoint recognition', respectively. In the object location memory task, subjects had to memorize the position of a target object in the environment from an initial viewpoint. They were then shown the same environment from a new viewpoint and had to indicate whether or not the target object had moved. In the viewpoint recognition task, subjects had to imagine the perspective of an avatar from the initial viewpoint and then decide whether or not the new viewpoint was that of the avatar. The results showed a double dissociation, with left MTL patients being impaired in the object location memory task but not in the viewpoint recognition task and right MTL patients being impaired in the viewpoint recognition task but not in the object location memory task. Furthermore, based on multiple regression analyses between performance and the volumes of the different MTL structures, we discuss the specific involvement of the left temporopolar cortex and of the right hippocampus in different kinds of visual perspective taking.

Where neuroscience and dynamic system theory meet autonomous robotics: a contracting basal ganglia model for action selection.

Action selection, the problem of choosing what to do next, is central to any autonomous agent architecture. We use here a multi-disciplinary approach at the convergence of neuroscience, dynamical system theory and autonomous robotics, in order to propose an efficient action selection mechanism based on a new model of the basal ganglia. We first describe new developments of contraction theory regarding locally projected dynamical systems. We exploit these results to design a stable computational model of the cortico-baso-thalamo-cortical loops. Based on recent anatomical data, we include usually neglected neural projections, which participate in performing accurate selection. Finally, the efficiency of this model as an autonomous robot action selection mechanism is assessed in a standard survival task. The model exhibits valuable dithering avoidance and energy-saving properties, when compared with a simple if-then-else decision rule.

The influence of age in women in visuo-spatial memory in reaching and navigation tasks with and without landmarks.

Spatial memory and navigation capabilities tend to decline in normal aging, but few studies have assessed the impact of landmarks on route learning in a large-scale environment. The objectives were to examine age-related effects on visuo-spatial working memory capabilities in various environments and to determine the impact of landmarks in navigation skills in normal aging. 42 young women (23.6 ±â€¯4.9 years) and 37 older women (70.7 ±â€¯4.7 years) with no cognitive impairment have performed three visuo-spatial working memory tests: one in reaching space (computerized Corsi-Block-Tapping test) and two in locomotor navigation space (a condition without landmarks: Virtual Walking Corsi Test and a condition with landmarks: Virtual Room Walking Test). A two-way mixed ANOVA test showed that the young subjects performed better in all conditions than older subjects. The performance in visuo-spatial working memory thus decreases with age. Visuo-spatial working memory performances were identical in reaching and navigation spaces for both groups. The integration of landmarks into a navigational task decreases performance in older women, while this performance is not altered in younger women.

From brainstem to cortex: computational models of saccade generation circuitry.

The brain circuitry of saccadic eye movements, from brainstem to cortex, has been extensively studied during the last 30 years. The wealth of data gathered allowed the conception of numerous computational models. These models proposed descriptions of the putative mechanisms generating this data, and, in turn, made predictions and helped to plan new experiments. In this article, we review the computational models of the five main brain regions involved in saccade generation: reticular formation saccadic burst generators, superior colliculus, cerebellum, basal ganglia and premotor cortical areas. We present the various topics these models are concerned with: location of the feedback loop, multimodal saccades, long-term adaptation, on the fly trajectory correction, strategy and metrics selection, short-term spatial memory, transformations between retinocentric and craniocentric reference frames, sequence learning, to name the principle ones. Our objective is to provide a global view of the whole system. Indeed, narrowing too much the modelled areas while trying to explain too much data is a recurrent problem that should be avoided. Moreover, beyond the multiple research topics remaining to be solved locally, questions regarding the operation of the whole structure can now be addressed by building on the existing models.

Multisensory integration in spatial orientation.

Recent psychophysical studies on normal subjects, as well as brain imaging studies, have revised the concepts concerning the mechanisms underlying spatial orientation during navigation tasks. The emphasis has been put on internal models that allow the prediction of a planned trajectory and are essential in the steering of locomotion. Cognitive factors such as strategies and emotional parameters are now starting to be included in the research on spatial orientation. It is obvious that important individual and gender differences exist in the brain operations underlying spatial orientation in humans, which may help to understand the construction of a coherent perception and the organic neural disorders related to the internal representation of space.

Effect of gravity on human spontaneous 10-Hz electroencephalographic oscillations during the arrest reaction.

Electroencephalographic oscillations at 10 Hz (alpha and mu rhythms) are the most prominent rhythms observed in awake, relaxed (eye-closed) subjects. These oscillations may be considered as a marker of cortical inactivity or an index of the active inhibition of the sensory information. Different cortical sources may participate in the 10-Hz oscillation and appear to be modulated by the sensory context and functional demands. In microgravity, the marked reduction in multimodal graviceptive inputs to cortical networks participating in the representation of space could be expected to affect the 10-Hz activity. The effect of microgravity on this basic oscillation has heretofore not been studied quantitatively. Because the alpha rhythm has a functional role in the regulation of network properties of the visual areas, we hypothesised that the absence of gravity would affect its strength. Here, we report the results of an experiment conducted over the course of 3 space flights, in which we quantified the power of the 10-Hz activity in relation to the arrest reaction (i.e., in 2 distinct physiological states: eyes open and eyes closed). We observed that the power of the spontaneous 10-Hz oscillation recorded in the eyes-closed state in the parieto-occipital (alpha rhythm) and sensorimotor areas (mu rhythm) increased in the absence of gravity. The suppression coefficient during the arrest reaction and the related spectral perturbations produced by eye-opening/closure state transition also increased in on orbit. These results are discussed in terms of current theories on the source and the importance of the alpha rhythm for cognitive function.

Reproduction of self-rotation duration.

The vestibular system detects the velocity of the head even in complete darkness, and thus contributes to spatial orientation. However, during vestibular estimation of linear passive self-motion distance in darkness, healthy human subjects mainly rely on time, and they replicate also stimulus duration when required to reproduce previous self-rotation. We then made the hypothesis that the perception of vestibular-sensed motion duration is embedded within encoding of motion kinetics. The ability to estimate time during passive self-motion in darkness was examined with a self-rotation reproduction paradigm. Subjects were required to replicate through self-driven transport the plateau velocity (30, 60 and 90 degrees /s) and duration (2, 3 and 4s) of the previously imposed whole-body rotation (trapezoid velocity profile) in complete darkness; the rotating chair position was recorded (500 Hz) during the whole trials. The results showed that the peak velocity, but not duration, of the plateau phase of the imposed rotation was accurately reproduced. Suspecting that the velocity instruction had impaired the duration reproduction, we added a control experiment requiring subjects to reproduce two successive identical rotations separated by a momentary motion interruption (MMI). The MMI was of identical duration to the previous plateau phase. MMI duration was fidelitously reproduced whereas that of the plateau phase was hypometric (i.e. lesser reproduced duration than plateau) suggesting that subjective time is shorter during vestibular stimulation. Furthermore, the accurate reproduction of the whole motion duration, that was not required, indicates an automatic process and confirms that vestibular duration perception is embedded within motion kinetics.

Biologically based artificial navigation systems: review and prospects.

Diverse theories of animal navigation aim at explaining how to determine and maintain a course from one place to another in the environment, although each presents a particular perspective with its own terminologies. These vocabularies sometimes overlap, but unfortunately with different meanings. This paper attempts to define precisely the existing concepts and terminologies, so as to describe comprehensively the different theories and models within the same unifying framework. We present navigation strategies within a four-level hierarchical framework based upon levels of complexity of required processing (Guidance, Place recognition-triggered Response, Topological navigation, Metric navigation). This classification is based upon what information is perceived, represented and processed. It contrasts with common distinctions based upon the availability of certain sensors or cues and rather stresses the information structure and content of central processors. We then review computational models of animal navigation, i.e. of animats. These are introduced along with the underlying conceptual basis in biological data drawn from behavioral and physiological experiments, with emphasis on theories of "spatial cognitive maps". The goal is to aid in deriving algorithms based upon insights into these processes, algorithms that can be useful both for psychobiologists and roboticists. The main observation is, however, that despite the fact that all reviewed models claim to have biological inspiration and that some of them explicitly use "Cognitive Map"-like mechanisms, they correspond to different levels of our proposed hierarchy and that none of them exhibits the main capabilities of real "Cognitive Maps"--in Tolman's sense--that is, a robust capacity for detour and shortcut behaviors.

Assessing morphology and function of the semicircular duct system: introducing new in-situ visualization and software toolbox.

The semicircular duct system is part of the sensory organ of balance and essential for navigation and spatial awareness in vertebrates. Its function in detecting head rotations has been modelled with increasing sophistication, but the biomechanics of actual semicircular duct systems has rarely been analyzed, foremost because the fragile membranous structures in the inner ear are hard to visualize undistorted and in full. Here we present a new, easy-to-apply and non-invasive method for three-dimensional in-situ visualization and quantification of the semicircular duct system, using X-ray micro tomography and tissue staining with phosphotungstic acid. Moreover, we introduce Ariadne, a software toolbox which provides comprehensive and improved morphological and functional analysis of any visualized duct system. We demonstrate the potential of these methods by presenting results for the duct system of humans, the squirrel monkey and the rhesus macaque, making comparisons with past results from neurophysiological, oculometric and biomechanical studies. Ariadne is freely available at http://www.earbank.org.

Background, but not foreground, spatial cues are taken as references for head direction responses by rat anterodorsal thalamus neurons.

Two populations of limbic neurons are likely neurophysiological substrates for cognitive operations required for spatial orientation and navigation: hippocampal pyramidal cells discharge selectively when the animal is in a certain place (the "firing field") in the environment, whereas head direction cells discharge when the animal orients its head in a specific, "preferred" direction. Cressant et al. (1997) showed that the firing fields of hippocampal place cells reorient relative to a group of three-dimensional objects only if these are at the periphery, but not the center of an enclosed platform. To test for corresponding responses in head direction cells, three objects were equally spaced along the periphery of a circular platform. Preferred directions were measured before and after the group of objects was rotated. (The rat was disoriented in total darkness between sessions). This was repeated in the presence or absence of a cylinder enclosing the platform. When the enclosure was present, the preferred directions of all 30 cells recorded shifted by the same angle as the objects. In the absence of the enclosure, the preferred directions did not follow the objects, remaining fixed relative to the room. These results provide a possible neurophysiological basis for observations from psychophysical experiments in humans that background, rather than foreground, cues are preferentially used for spatial orientation.

The "Stroop Walking Task": An innovative dual-task for the early detection of executive function impairment.

AIMS OF THE STUDY: To evaluate a dual-task named the "Stroop Walking Task", which is similar to the task of making a decision of whether to cross a street based on a pedestrian traffic light. PATIENTS AND METHODS: Fifty-one subjects (15 young adults, 21 subjectively healthy old subjects and 15 old subjects with mild cognitive impairment) had to respond to a visual signal (pictogram) with an appropriate motor response (walk or stop). We used an electronic walkway system to record the gait parameters and performed a cluster analysis on the obtained data. RESULTS: This dual-task enables the early detection of executive function impairment with 89% sensitivity and 87% specificity. CONCLUSION: The use of a dual-task that is inspired by an everyday event as an evaluation tool seems to facilitate the detection of ageing subjects' cognitive impairment, which is not detectable with traditional psychometric tests.

Development of anticipatory orienting strategies during locomotor tasks in children.

Some basic problems related to the development of goal-directed locomotion in humans are reviewed here. A preliminary study is presented which was aimed at investigating the emergence of anticipatory head orienting strategies during goal-directed locomotion in children. Eight children ranging from 3.5 to 8 years had to walk along a 90 degrees right corner trajectory to reach a goal, both in light and in darkness. The instantaneous orientation in space of the head, trunk, hips and left foot antero/posterior axes was computed by means of an ELITE four-TV camera, 100 Hz system. The results showed that predictive head orienting movements can occur also in the youngest children. The head starts to rotate toward the goal before the corner point of the trajectory is reached. In children, the head peak rotation coincides with the trajectory corner while in adults the peak is attained before. In children, the walking speed is largely decreased in darkness. The results suggest that feedforward control of goal-directed locomotion appears very early in gait development and becomes increasingly important afterwards.

Eye deviation during rotation in darkness in trait anxiety: an early expression of perceptual avoidance?

BACKGROUND: Patients with dizziness and patients with panic disorder and agoraphobia share a common symptomatology. Numerous studies have investigated a potential link between anxiety and the vestibular system, but few of them have addressed the specific topic of spatial representation. METHODS: Passive whole-body rotations in the horizontal plane were imposed on two groups of subjects who differed in their level of trait anxiety. Subjects were seated on a mobile robot in darkness. After each passive rotation, subjects were asked to reproduce the stimulus by driving the robot with a joystick and to perform a rotation of the same magnitude. Eye movements were recorded and analyzed. RESULTS: No difference in either perception (accuracy in the reproduction task) or in VOR gain was found between the two groups of subjects. Mean eye deviation, caused by fast phases of the nystagmus, differed in the two groups. It was typically in the anticompensatory direction in the non-anxious group, and in the compensatory direction the anxious group. Such compensatory movement may be explained by an egocentric orientation strategy, that may in turn indicate a lack of interest toward the visual surroundings. CONCLUSIONS: An egocentric strategy for self-orientation exhibited at a level below the threshold of awareness could reveal the existence of a physiological mode of processing leading to agoraphobic avoidance.

Lack of anticipatory gaze-orienting responses in patients with right brain damage.

OBJECTIVE: To study eye movements during cervical proprioceptive stimulation by passive body rotation in darkness, with the head held stationary, in patients with right brain damage and hemineglect. BACKGROUND: At very low frequency, this stimulation is reported to produce an illusion of head turning in space and eye deviations directed opposite to trunk rotation (in the direction of the illusory head rotation). METHODS: Ten normal subjects and seven patients with unilateral cerebral lesions (five right brain-damaged patients with mild to moderate visuospatial neglect, two left brain-damaged patients without neglect) were included in the study. Subjects were seated on a rotating chair. Stimuli consisted of slow sinusoidal passive trunk rotations (+/-30 degrees, 0.01 Hz) while the head was fixed in space. RESULTS: Eye movements directed opposite to trunk rotation were typical for normal subjects and for left brain-damaged patients. In contrast, all right brain-damaged patients showed either eye movements in the direction of trunk rotation or no eye deviations at all. CONCLUSION: This result could characterize a lack of anticipatory coordinating gaze behavior in patients with right brain damage.

A neurophysiological study of prepositus hypoglossi neurons projecting to oculomotor and preoculomotor nuclei in the alert cat.

The activity of 62 antidromically identified prepositus hypoglossi neurons was recorded in 10 alert cats during spontaneous, vestibular or visually induced eye movements. Neurons were antidromically activated from stimulating electrodes implanted in the ipsilateral medial longitudinal fasciculus (n = 24), the ipsilateral interstitial nucleus of Cajal (n = 6), the ipsilateral parabigeminal nucleus (n = 2), the contralateral superior colliculus (n = 6) and the contralateral cerebellar posterior peduncle (n = 24). Neurons were identified as eye-movement-related when their rate-position and/or rate-velocity plots showed correlation coefficients greater than or equal to 0.6. They were further classified as "position", "position-velocity" and "velocity-position" according to their relative eye position and velocity coefficients. However, they seemed to be distributed as a continuum in which a progressive decrease of eye velocity sensitivity was accompanied by a proportional increase in eye position sensitivity. "Position-velocity" neurons (n = 9) were mainly horizontal type II neurons projecting to the vicinity of the oculomotor complex; two of these neurons with vertical sensitivity were also activated from the interstitial nucleus of Cajal. Mean position and velocity sensitivity of these neurons were 5.2 spikes/s per degree and 0.62 spikes/s per degree per second, respectively. Pure "position" neurons (n = 7) also showed activation during ipsilateral eye fixations; their mean position gain was 7.3 spikes/s per degree and they projected to the ipsilateral oculomotor and Cajal nuclei, and to the contralateral superior colliculus. "Velocity-position" neurons (n = 18) were type I or II neurons with rather irregular tonic firing rates and a mean velocity gain of 0.75 spikes/s per degree per second. Type II "velocity-position" neurons projected mainly to the oculomotor area, while type I neurons projected preferentially to the cerebellum. A special type of "pause" neuron (n = 5), with very low firing rate and pausing mainly for contralateral saccades, was activated exclusively from the contralateral posterior peduncle. Many neurons with weak eye movement sensitivity (n = 22) were activated mainly (73%) from the cerebellum. It can be concluded that the prepositus hyperglossi nucleus distributes specific eye movement related signals to motor and premotor brainstem and cerebellar structures. The variability of interspike intervals of representative prepositus hypoglossi neurons of each class was compared to the discharge variability of identified abducens motoneurons.(ABSTRACT TRUNCATED AT 400 WORDS)

Motor processes in mental rotation.

Much indirect evidence supports the hypothesis that transformations of mental images are at least in part guided by motor processes, even in the case of images of abstract objects rather than of body parts. For example, rotation may be guided by processes that also prime one to see results of a specific motor action. We directly test the hypothesis by means of a dual-task paradigm in which subjects perform the Cooper-Shepard mental rotation task while executing an unseen motor rotation in a given direction and at a previously-learned speed. Four results support the inference that mental rotation relies on motor processes. First, motor rotation that is compatible with mental rotation results in faster times and fewer errors in the imagery task than when the two rotations are incompatible. Second, the angle through which subjects rotate their mental images, and the angle through which they rotate a joystick handle are correlated, but only if the directions of the two rotations are compatible. Third, motor rotation modifies the classical inverted V-shaped mental rotation response time function, favoring the direction of the motor rotation; indeed, in some cases motor rotation even shifts the location of the minimum of this curve in the direction of the motor rotation. Fourth, the preceding effect is sensitive not only to the direction of the motor rotation, but also to the motor speed. A change in the speed of motor rotation can correspondingly slow down or speed up the mental rotation.