How cognitive aging affects multisensory integration of navigational cues

Entorhinal grid cells and hippocampal place cells are key systems for mammalian navigation. By combining information from different sensory modalities, they provide abstract representations of space. Given that both structures are among the earliest to undergo age-related neurodegenerative changes, we asked whether age-related navigational impairments are related to deficient integration of navigational cues. Younger and older adults performed a homing task that required using visual landmarks, self-motion information, or a combination of both. Further, a conflict between cues assessed the influence of each sensory domain. Our findings revealed performance impairments in the older adults, suggestive of a higher noise in the underlying spatial representations. In addition, even though both groups integrated visual and self-motion information to become more accurate and precise, older adults did not place as much influence on visual information as would have been optimal. As these findings were unrelated to potential changes in balance or spatial working memory, this study provides the first evidence that increasing noise and a suboptimal weighting of navigational cues might contribute to the common problems with spatial representations experienced by many older adults. These findings are discussed in the context of the known age-related changes in the entorhinal-hippocampal network.     

Measurement of instantaneous perceived self-motion using continuous pointing

In order to optimally characterize full-body self-motion perception during passive translations, changes in perceived location, velocity, and acceleration must be quantified in real time and with high spatial resolution. Past methods have failed to effectively measure these critical variables. Here, we introduce continuous pointing as a novel method with several advantages over previous methods. Participants point continuously to the mentally updated location of a previously viewed target during passive, full-body movement. High-precision motion-capture data of arm angle provide a measure of a participant’s perceived location and, in turn, perceived velocity at every moment during a motion trajectory. In two experiments, linear movements were presented in the absence of vision by passively translating participants with a robotic wheelchair or an anthropomorphic robotic arm (MPI Motion Simulator). The movement profiles included constant-velocity trajectories, two successive movement intervals separated by a brief pause, and reversed-motion trajectories. Results indicate a steady decay in perceived velocity during constant-velocity travel and an attenuated response to mid-trial accelerations. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.

Circular trajectory formation during blind locomotion: a test for path integration and motor memory

 Eight healthy subjects were asked to walk blindfolded along circular paths of different radii after several practice trials with vision. Their task was to stop after completing two full revolutions. They always walked counter-clockwise (CCW) in (a) a control condition (CONTROL), including the instructions mentioned above, (b) with the further instruction to count backwards in twos (MENTAL), (c) with the instruction to count loudly (LOUD). The movement of two markers lying along the head naso-occipital axis was recorded by means of an ELITE system. Total walked distance (DISTANCE), total head turning angle (ANGLE) and average radius (RADIUS) of the trajectories performed were measured. All subjects were able to perform approximately circular trajectories. They consistently overshot the ideal radius independently of the condition and circle size, undershot the total angle and overshot total distance. The LOUD condition induced greater errors in the performance but only on total distance (P<0.05). A strong correlation was found between the errors in radius and total distance but not between distance and total angle. Principal components analysis suggested that radius and distance share a common source of errors while total angle produced independent errors. The results indicate that (a) circular trajectories can be generated starting from spatial and/or motor memory, without the aid of visual information; (b) the task needs some attentional control and does not involve simple automatic processing of afferent information; (c) different sensory information or different processing modes are probably involved in the estimation of the curvature and length of the walked path on the one hand, and of the total rotation angle on the other. Received: 1 June 1996 / Accepted: 18 January 1997     

Grammatical class in lexical production and morhpological processing: Evidence from a case of fluent aphasia

We present the case of a fluent aphasic patient who is impaired at producing nouns relative to verbs in picture naming, sentence completion, and sentence generation tasks, but is better at both producing and comprehending concrete nouns than abstract nouns. Moreover, he displays a selective difficulty in producing the plural forms of some nouns and pseudowords presented as nouns, but was able to produce the phonologically identical third-person singular forms of corresponding verb homonyms and of the same pseudowords presented as verbs. This pattern of performance casts doubt on the hypothesis that grammatical class effects are always epiphenomena of more general semantic impairments that affect the naming of actions or of concrete objects, and suggests that these effects may arise instead from damage to syntactic processes pertaining specifically to the grammatical properties of words. We also discuss the implications of such damage for models of morphological processing.     

Perception action interaction: the oblique effect in the evolving trajectory of arm pointing movements

In previous studies, we provided evidence for a directional distortion of the endpoints of movements to memorized target locations. This distortion was similar to a perceptual distortion in direction discrimination known as the oblique effect so we named it the “motor oblique effect”. In this report we analyzed the directional errors during the evolution of the movement trajectory in memory guided and visually guided pointing movements and compared them with directional errors in a perceptual experiment of arrow pointing. We observed that the motor oblique effect was present in the evolving trajectory of both memory and visually guided reaching movements. In memory guided pointing the motor oblique effect did not disappear during trajectory evolution while in visually guided pointing the motor oblique effect disappeared with decreasing distance from the target and was smaller in magnitude compared to the perceptual oblique effect and the memory motor oblique effect early on after movement initiation. The motor oblique effect in visually guided pointing increased when reaction time was small and disappeared with larger reaction times. The results are best explained using the hypothesis that a low level oblique effect is present for visually guided pointing movements and this effect is corrected by a mechanism that does not depend on visual feedback from the trajectory evolution and might even be completed during movement planning. A second cognitive oblique effect is added in the perceptual estimation of direction and affects the memory guided pointing movements. It is finally argued that the motor oblique effect can be a useful probe for the study of perception–action interaction.     

Neural correlates of gesture processing across human development

Co-speech gesture facilitates learning to a greater degree in children than in adults, suggesting that the mechanisms underlying the processing of co-speech gesture differ as a function of development. We suggest that this may be partially due to children's lack of experience producing gesture, leading to differences in the recruitment of sensorimotor networks when comparing adults to children. Here, we investigated the neural substrates of gesture processing in a cross-sectional sample of 5-, 7.5-, and 10-year-old children and adults and focused on relative recruitment of a sensorimotor system that included the precentral gyrus (PCG) and the posterior middle temporal gyrus (pMTG). Children and adults were presented with videos in which communication occurred through different combinations of speech and gesture during a functional magnetic resonance imaging (fMRI) session. Results demonstrated that the PCG and pMTG were recruited to different extents in the two populations. We interpret these novel findings as supporting the idea that gesture perception (pMTG) is affected by a history of gesture production (PCG), revealing the importance of considering gesture processing as a sensorimotor process.     

An electrophysiological measure of access to representations in visual working memory

Previous research has demonstrated that the maintenance of visual information in working memory is associated with a sustained posterior contralateral negativity. Here we show that this component is also elicited during the spatially selective access to visual working memory. Participants memorized a bilateral visual search array that contained two potential targets on the left and right side. The task-relevant side was signalled by post-cues that were presented either 150 ms after array offset or after a longer interval (700–1000 ms). Enhanced negativities at posterior electrodes contralateral to the cued side of a target were elicited in response to both early and late post-cues, suggesting that they reflect not only memory maintenance, but also processes involved in the access to stored visual working memory representations. Results provide new electrophysiological evidence for the retinotopic organization of visual working memory.     

Estradiol treatment in preadolescent females enhances adolescent spatial memory and differentially modulates hippocampal region-specific phosphorylated ERK labeling

Estrogen levels in rats are positively correlated with enhanced memory function and hippocampal dendritic spine density. There is much less work on the long-term effects of estradiol manipulation in preadolescent rats. The present work examined how injections of estradiol during postnatal days 19–22 (p19–22; preadolescence) affected water maze performance and hippocampal phosphorylated ERK labeling. To investigate this, half of the estradiol- and vehicle-treated female rats were trained on a water maze task 24 h after the end of estradiol treatment (p23–27) while the other half was not trained. All female rats were tested on the water maze from p40 to p44 (adolescence) and hippocampal pERK1/2 labeling was assessed as a putative marker of neuronal plasticity. During adolescence, preadolescent-trained groups showed lower latencies than groups without preadolescent training. Retention data revealed lower latencies in both estradiol groups, whether preadolescent trained or not. Immunohistochemical detection of hippocampal pERK1/2 revealed elevations in granule cell labeling associated with the preadolescent trained groups and reductions in CA1 labeling associated with estradiol treatment. These results show a latent beneficial effect of preadolescent estradiol treatment on adolescent spatial performance and suggest an organizational effect of prepubescent exogenously applied estradiol.     

Methadone disrupts performance on the working memory version of the Morris water task

The aim of the study was to examine if administration of the mu-opiate agonist methadone hydrochloride resulted in deficits in performance on the Morris water tank task, a widely used test of spatial cognition. To this end, after initial training on the task, Long-Evans rats were administered saline or methadone at either 1.25, 2.5 or 5 mg/kg ip 15 min prior to testing. The performance of the highest-dose methadone group was inferior to that of the controls on the working memory version of the Morris task. There were also differences between the groups on the reference memory version of the task, but this result cannot be considered reliable. These data show that methadone has its most profound effect on cognition in rats when efficient performance on the task requires attention to and retention of new information, in this case, the relationship between platform location and the extramaze cues.     

Spatial orientation in humans: perception of angular whole-body displacements in two-dimensional trajectories

Vestibular perception of whole-body passive rotation in the horizontal plane was studied by applying two-dimensional (2D) motion to eight blindfolded healthy volunteers: pure rotations in place, corner-like trajectories and arcs of a circular trajectory were randomly applied by means of a remotely controlled robot. Angles embedded in the 2D trajectories were 45°, 90°, 135° and 180°. Stimulation of semicircular canals was the same for all trajectories but was accompanied by concurrent otolith stimulation during circular motion. Subjects participated in two successive experimental sessions. In the first session they were instructed to use a pointer to reproduce the total angular displacement after the motion (REPRODUCTION); in the second session they had to keep pointing towards a remote (15 m) memorised target during the motion (TRACKING). In REPRODUCTION subjects tended to overestimate their rotation angle by 28 ± 11% (mean ± SD). There was no systematic effect of the trajectory. Overestimation also occurred when subjects were required to rotate in darkness by 180° (by controlling a joystick). In TRACKING there was virtually no overestimation (6 ± 17%) and the movement of the pointer matched the dynamics of angular motion. We conclude that (a) the brain can separate and memorise the angular component of complex 2D motion; however, a large inter-individual variability in estimating its amplitude exists; (b) in the range of linear accelerations used in the study, no appreciable effect of otolith-canal perceptual interaction was shown; (c) angular displacements can be dynamically transformed into matched pointing movements; (d) overestimation seems to be typical of delayed judgements of angular displacement and of self-controlled rotations in place. This could be due to the characteristics of the physiological calibration of the vestibular input. Received: 30 October 1996 / Accepted: 18 June 1997     

Human movement coordination implicates relative direction as the information for relative phase

The current studies explore the informational basis of the coupling in human rhythmic movement coordination tasks. Movement stability in these tasks is an asymmetric U-shaped function of mean relative phase; 0 degrees is maximally stable, 90 degrees is maximally unstable and 180 degrees is intermediate. Bingham (2001, 2004a, 2004b) hypothesized that the information used to perform coordinated rhythmic movement is the relative direction of movement, the resolution of which is determined by relative speed. We used an experimental paradigm that entails using a circular movement to produce a linear motion of a dot on a screen, which must then be coordinated with a linearly moving computer controlled dot. This adds a component to the movement that is orthogonal to the display. Relative direction is not uniquely defined between orthogonal components of motion, but relative speed is; it was therefore predicted that the addition of the component would only introduce a symmetric noise component and not otherwise contribute to the U-shape structure of movement stability. Results for experiment 1 supported the hypothesis; movement that involved the additional component was overall less stable than movement that involved only the parallel component along which relative direction can be defined. Two additional studies ruled out alternative explanations for the pattern of data in experiment 1. Overall, the results strongly implicate relative direction as the information underlying performance in rhythmic movement coordination tasks.     

Spatial updating and the maintenance of visual constancy

Spatial updating is the means by which we keep track of the locations of objects in space even as we move. Four decades of research have shown that humans and non-human primates can take the amplitude and direction of intervening movements into account, including saccades (both head-fixed and head-free), pursuit, whole-body rotations and translations. At the neuronal level, spatial updating is thought to be maintained by receptive field locations that shift with changes in gaze, and evidence for such shifts has been shown in several cortical areas. These regions receive information about the intervening movement from several sources including motor efference copies when a voluntary movement is made and vestibular/somatosensory signals when the body is in motion. Many of these updating signals arise from brainstem regions that monitor our ongoing movements and subsequently transmit this information to the cortex via pathways that likely include the thalamus. Several issues of debate include (1) the relative contribution of extra-retinal sensory and efference copy signals to spatial updating, (2) the source of an updating signal for real life, three-dimensional motion that cannot arise from brain areas encoding only two-dimensional commands, and (3) the reference frames used by the brain to integrate updating signals from various sources. This review highlights the relevant spatial updating studies and provides a summary of the field today. We find that spatial constancy is maintained by a highly evolved neural mechanism that keeps track of our movements, transmits this information to relevant brain regions, and then uses this information to change the way in which single neurons respond. In this way, we are able to keep track of relevant objects in the outside world and interact with them in meaningful ways.     

Neural mechanisms of movement speed and <Emphasis Type="Italic">tau</Emphasis> as revealed by magnetoencephalography

A fundamental aspect of goal-directed behavior concerns the closure of motion-gaps in a timely fashion. In this context, the critical variable is the time-to-closure, called tau (Lee in Perception 5:437–459, 1976), and is defined as the ratio of the current distance-to-goal gap over the current instantaneous speed towards the goal. In this study, we investigated the neural mechanisms of speed and tau in pointing hand movements by recording MEG activity from the whole brain of 20 right-handed healthy human subjects operating a joystick with their right hand. The relations between neural signals and speed and tau were analyzed using an autoregressive multiple regression model, where the time-varying MEG signal was the dependent variable and the corresponding value of speed and tau were the independent variables. With respect to speed, we found that 81% of sensors showed significant relations over the left frontal-parietal, left parieto-temporal, and, less prominently, the right temporo-occipital sensor space. These results document the widespread involvement of brain areas with movement speed, especially in the left hemisphere (i.e., contralateral to the moving limb), in accord with previous studies. With respect to tau, 22% of sensors showed significant relations over the parietal (bilaterally), right parietal-temporal, and, less prominently, the left temporo-occipital sensor space. The tau effects often occurred concurrently with speed effects and spatially overlapped in the left fronto-parietal sensors. These findings document for the first time the time-varying, dynamic processing of information regarding tau in specific brain areas, including the right parietal cortex. This is of special interest, for that area has been found to be involved in processing information concerning the duration of time intervals in perceptual tasks (Harrington et al. in J Neurosci 18:1085–1095, 1998; Rao et al. in Nat Neurosci 4:317–323, 2001). Since tau is itself a time interval, we hypothesize that the right parietal focus of tau processing observed in this study reflects the ongoing processing of tau as an interval for a timely arrival of the hand to the target.     

Spatial learning and memory in African mole-rats: the role of sociality and sex

Spatial learning and memory is an important skill for the survival and fitness and may vary between the sexes depending on differences in space use. This is particularly true for animals that explore the subterranean niche as it is associated with high travelling costs. In subterranean rodents the complexity of burrow systems varies with differing degrees of sociality possibly posing stronger selective pressures regarding spatial abilities on species with more complex burrow structures. This could lead to superior abilities in spatial learning and memory in social compared to solitary subterranean species. We tested this hypothesis in two species of subterranean mole-rats, the eusocial Damaraland (Fukomys damarensis) and solitary Cape mole-rats (Georychus capensis) by comparing their ability to locate food in an artificial maze. Measurements of the time taken to the goal chamber, the number of wrong turns taken, and the average velocity at which animals travelled were used to compare performance between animals. We did not find marked sex-specific differences in either study species during the assessment of learning and memory retention. In accordance with our hypothesis significant differences between the species were apparent during both learning and memory trials with the social species exhibiting superior performances. However, in both species memory retention was generally high suggesting that the fossorial lifestyle poses a strong selective pressure on spatial abilities in subterranean mammals.     

Spatial learning and memory following fimbria-fornix transection and grafting of fetal septal neurons to the hippocampus

Summary The ability of intrahippocampal grafts of fetal septal-diagonal band tissue, rich in developing cholinergic neurons, to ameliorate cognitive impairments induced by bilateral fimbria-fornix transections in rats was examined in three experiments using the Morris water-maze to test different aspects of spatial memory. Experiment 1. Rats with fimbria-fornix lesions received either septal cell suspension grafts or solid septal grafts; normal rats and rats with lesions alone were used as controls. Sixteen weeks after surgery, the rats' spatial learning and memory were tested in the water-maze using a place test, designed to investigate place navigation performance, in which rats learned to escape from the water by swimming to a platform hidden beneath the water's surface. After 5 days of training, the rats were given a spatial probe test in which the platform was removed from the tank to test spatial reference memory. Experiment 2. The same rats used in Exp. 1 were tested in a delayed-match-to-sample, working memory version of the water-maze task. The platform was located in one of two possible locations during each trial, which was composed of 2 swims. If the rat remembered the location of the platform on the 2nd swim of a trial, it should find the platform more quickly on that swim, and thereby demonstrate working memory. Experiment 3. Prior to receiving fimbria-fornix lesions, normal rats were trained in a modification of the water-maze task using alternating cue navigation and place navigation trials (i.e., with visible or non-visible escape platforms). The retention and reacquisition of the place task and the spatial probe test were examined in repeated tests up to 6 months after the lesion and intrahippocampal grafting of septal cell suspensions. The effects of central muscarinic cholinergic receptor blockade with atropine were also tested. Normal rats performed well in both the place and spatial probe tests. In contrast, rats with fimbria-fornix lesions only were unable to acquire or retain spatial information in any test. Instead, these rats adopted a random, nonspatial search strategy, whereby their latencies to find the platform decreased in the place navigation tasks. Sixty to 80% of the rats with septal suspension or solid grafts had recovered place navigation, i.e., the ability to locate the platform site in the tank, in Exp. 1 and 3, and they showed a significantly improved performance in the working memory test in Exp. 2. Atropine abolished the recovered place navigation in the grafted rats, whereas normal rats were impaired to a lesser extent. In contrast, atropine had no effect on the non-spatial strategy adopted by rats with fimbria-fornix lesions only. The results show that: (1) fimbria-fornix lesions disrupt spatial learning and memory in both naive and pretrained rats; (2) with extended training the fimbria-fornix lesioned rats develop an efficient non-spatial strategy, which enables them to reduce their escape latency to levels close to those of intact controls; (3) intrahippocampal septal grafts can restore the ability of the lesioned rats to use spatial cues in the localization of the platform site; and (4) the behavioural recovery produced by grafts is dependent upon an atropine sensitive mechanism.     

Spatial coding of eye movements relative to perceived earth and head orientations during static roll tilt

This purpose of this study was to examine the spatial coding of eye movements during static roll tilt (up to ±45°) relative to perceived earth and head orientations. Binocular videographic recordings obtained in darkness from eight subjects allowed us to quantify the mean deviations in gaze trajectories along both horizontal and vertical coordinates relative to the true earth and head orientations. We found that both variability and curvature of gaze trajectories increased with roll tilt. The trajectories of eye movements made along the perceived earth-horizontal (PEH) were more accurate than movements along the perceived head-horizontal (PHH). The trajectories of both PEH and PHH saccades tended to deviate in the same direction as the head tilt. The deviations in gaze trajectories along the perceived earth-vertical (PEV) and perceived head-vertical (PHV) were both similar to the PHH orientation, except that saccades along the PEV deviated in the opposite direction relative to the head tilt. The magnitude of deviations along the PEV, PHH, and PHV corresponded to perceptual overestimations of roll tilt obtained from verbal reports. Both PEV gaze trajectories and perceptual estimates of tilt orientation were different following clockwise rather than counterclockwise tilt rotation; however, the PEH gaze trajectories were less affected by the direction of tilt rotation. Our results suggest that errors in gaze trajectories along PEV and perceived head orientations increase during roll tilt in a similar way to perceptual errors of tilt orientation. Although PEH and PEV gaze trajectories became nonorthogonal during roll tilt, we conclude that the spatial coding of eye movements during roll tilt is overall more accurate for the perceived earth reference frame than for the perceived head reference frame. Received: 22 April 1997 / Accepted: 18 December 1997     

Pitch body orientation influences the perception of self-motion direction induced by optic flow

We studied the effect of static pitch body tilts on the perception of self-motion direction induced by a visual stimulus. Subjects were seated in front of a screen on which was projected a 3D cluster of moving dots visually simulating a forward motion of the observer with upward or downward directional biases (relative to a true earth horizontal direction). The subjects were tilted at various angles relative to gravity and were asked to estimate the direction of the perceived motion (nose-up, as during take-off or nose-down, as during landing). The data showed that body orientation proportionally affected the amount of error in the reported perceived direction (by 40% of body tilt magnitude in a range of ±20°) and these errors were systematically recorded in the direction of body tilt. As a consequence, a same visual stimulus was differently interpreted depending on body orientation. While the subjects were required to perform the task in a geocentric reference frame (i.e., relative to a gravity-related direction), they were obviously influenced by egocentric references. These results suggest that the perception of self-motion is not elaborated within an exclusive reference frame (either egocentric or geocentric) but rather results from the combined influence of both.     

视觉工作记忆多通道脑电的不同频段能量空间分布

目的 工作记忆是重要的认知功能之一.研究工作记忆任务中多通道脑电(multi-channel,EEGs)在各个频率范围的能量分布,可为研究工作记忆的特征频段和关键脑区提供支持.方法 记录健康受试者在视觉工作记忆任务中的32通道EEGs,共16例受试者240次实验数据;对原始EEGs进行预处理;应用傅里叶变换计算预处理后的EEGs在δ、θ、α、β、γ频段能量的空间分布和各个频段能量占总能量的百分比.结果 δ、θ、α、β、γ频段能量占总能量的百分比分别为(22.44±0.86)％、(31.88±0.55)％、(24.66±1.43)％、(13.54±0.64)％、(7.47±0.48)％,θ频段的能量显著高于其他频段的能量,差异具有统计学意义(P＜0.001);在θ频段,能量主要集中于额中线区.结论 θ频段是工作记忆的特征频段,额区是工作记忆的关键脑区.     

A multisensory approach to spatial updating: the case of mental rotations

Mental rotation is the capacity to predict the outcome of spatial relationships after a change in viewpoint. These changes arise either from the rotation of the test object array or from the rotation of the observer. Previous studies showed that the cognitive cost of mental rotations is reduced when viewpoint changes result from the observer’s motion, which was explained by the spatial updating mechanism involved during self-motion. However, little is known about how various sensory cues available might contribute to the updating performance. We used a Virtual Reality setup in a series of experiments to investigate table-top mental rotations under different combinations of modalities among vision, body and audition. We found that mental rotation performance gradually improved when adding sensory cues to the moving observer (from None to Body or Vision and then to Body & Audition or Body & Vision), but that the processing time drops to the same level for any of the sensory contexts. These results are discussed in terms of an additive contribution when sensory modalities are co-activated to the spatial updating mechanism involved during self-motion. Interestingly, this multisensory approach can account for different findings reported in the literature.