Egocentric and allocentric spatial reference frames in aging: A systematic review

Aging affects many aspects of everyday living, such as autonomy, security and quality of life. Among all, spatial memory and spatial navigation show a gradual but noticeable decline, as a result of both neurobiological changes and the general slowing down of cognitive functioning. We conducted a systematic review using the Preferred Reporting Items for Systematic Reviews and Meta-Analysis (PRISMA) guidelines to identify studies that specifically investigated the role of allocentric and egocentric frames in healthy aging. Concerning spatial navigation, our results showed a preservation of egocentric strategies, along with specific impairments in the use of allocentric and switching abilities. Regarding spatial memory, instead, outcomes were more divergent and not frame-specific. With this perspective, spatial impairments were discussed considering the cognitive profile of mild cognitive impairment (MCI) and Alzheimer’s Disease (AD).     

Electroencephalographic analysis of cortico-muscular coherence: reference effect, volume conduction and generator mechanism.

OBJECTIVE: To measure the synchrony between cortical and muscle oscillatory activities, the coherence estimate between EEG and EMG was computed. METHODS: The multichannel electroencephalogram (EEG) and electromyogram (EMG) of the right abductor pollicis brevis muscle were recorded in 5 normal volunteers. Various types of EEG derivation methods were systematically compared to establish a standard method to study cortico-muscular coupling. RESULTS: The use of a reference-free EEG derivation (current source density) greatly improved cortico-muscular coherence. In all subjects, EEGs over the left sensorimotor cortex were coherent with EMG (mean peak frequency: 18.7 Hz, mean highest coherence: 0.124). The time lag from cortex to muscle in 14-50 Hz was 14.3 ms. EEG source derivation revealed that both radial and tangential generators in the precentral cortex might contribute to this phenomenon. In the EEG signals using common average reference, an artifactual coherence peak over the medial frontal area was observed, which might largely be explained by volume conduction from the primary sensorimotor cortex. CONCLUSIONS: We conclude that the current source density or its approximation is preferable to estimate the cortico-muscular coherence and that the interpretation of such coherence using referenced EEGs should be taken with care.     

Material-specific long-term memory representations of faces and spatial positions: evidence from slow event-related brain potentials

Motivated by models that propose material-specific cortical long-term memory representations we expected different topographies of event-related slow waves of the EEG during cued retrieval of two distinct types of information (faces and spatial positions), which are assumed to be processed and stored in topographically distinct cortical areas, i.e., in either the ventral or the dorsal visual pathway. Seventeen participants learned associations either between words and spatial positions or between words and faces. Each word was associated with either one or two positions or faces. In a cued recall test, one day later, participants saw two words and had to decide whether these were linked to each other via an associated spatial position or a face. Slow event-related potentials (ERPs) of the EEG were recorded from 61 scalp electrodes during both acquisition and recall. Response times increased monotonically with the number of faces and positions to be reactivated. Negative slow ERPs showed a comparable topography during anticipation learning and cued recall, but dissociated topographically for positions and faces. The maximum of the negativity increased when items were presented repetitively (compared to the first presentation) during learning, and also with the number of the to-be-reactivated associations during retrieval. These results are consistent with an information-processing model that assumes material-specific cortical representations of episodic memory contents, which are established as localized cortical cell assemblies during encoding, and which are being reactivated during recall.     

The effect of contextual auditory stimuli on virtual spatial navigation in patients with focal hemispheric lesions

Topographical disorientation is a frequent deficit among patients suffering from brain injury. Spatial navigation can be explored in this population using virtual reality environments, even in the presence of motor or sensory disorders. Furthermore, the positive or negative impact of specific stimuli can be investigated. We studied how auditory stimuli influence the performance of brain-injured patients in a navigational task, using the Virtual Action Planning–Supermarket (VAP-S) with the addition of contextual (“sonar effect” and “name of product”) and non-contextual (“periodic randomised noises”) auditory stimuli. The study included 22 patients with a first unilateral hemispheric brain lesion and 17 healthy age-matched control subjects. After a software familiarisation, all subjects were tested without auditory stimuli, with a sonar effect or periodic random sounds in a random order, and with the stimulus “name of product”. Contextual auditory stimuli improved patient performance more than control group performance. Contextual stimuli benefited most patients with severe executive dysfunction or with severe unilateral neglect. These results indicate that contextual auditory stimuli are useful in the assessment of navigational abilities in brain-damaged patients and that they should be used in rehabilitation paradigms.     

Detection of unexpected events during spatial navigation in humans: bottom-up attentional system and neural mechanisms

Navigation is a complex cognitive ability requiring the processing and integration of several different types of information extracted from the environment. While navigating, however, an unexpected event may suddenly occur, which individuals are required to detect promptly in order to apply an appropriate behavioural response. The alerting mechanism that is integral to the detection of unexpected events is referred to as the bottom-up attentional system. Using event-related functional magnetic resonance imaging, we investigated the neural basis of bottom-up detection of unexpected events while individuals moved within a virtual environment. We identified activation within a right fronto-temporo-parietal network in response to unexpected events while navigating in this virtual environment. Furthermore, when an unexpected event requires an adjusted behavioural response, a region of the right ventrolateral pre-frontal cortex (areas 45 and 47/12) is selectively activated. Our data replicate earlier findings on the neural mechanisms underlying visual attention and extend these findings to the more complex real-life ability of spatial navigation, thereby suggesting that these neural mechanisms subserve the bottom-up attentional systems that are crucial for effective locomotion in real surroundings.     

The neural basis of aphasia rehabilitation: evidence from neuroimaging and neurostimulation

This article is a selective review of functional imaging investigations and brain stimulation studies addressing the neural mechanisms of recovery of stroke-associated aphasia. The imaging results show that aphasia recovery is associated with a complex pattern of brain reorganisation, involving both ipsilateral and contralateral brain regions, which is modulated by lesion size and site, time post-onset, type of training, and language task. The information provided by the imaging investigations needs to be integrated with the results of brain stimulation studies, in order to specify the most effective protocols in term of modality, locus and timing of stimulation. Further studies, using multiple imaging and neuromodulation approaches, are required to reach sound conclusions about the potential usefulness of brain stimulation approaches as an adjunct to aphasia rehabilitation.     

The spatial representation of numerical and non-numerical sequences: evidence from neglect

Psychophysical and neuropsychological studies have revealed that humans represent numbers along a continuous, left-to-right oriented mental line. However, it has been recently claimed that this format of representation is not special to numbers because non-numerical sequences would be spatially coded in the same way. To test this hypothesis, the present study investigated the effects of left neglect upon the bisection of numerical and non-numerical intervals. Eight patients with left neglect performed a visual line bisection task and three mental bisection tasks with number, letter, and month intervals. The error pattern in the number bisection task, indexed by the modulating effect of interval length, mirrored that of the visual task and confirmed the left-to-right spatial orientation of the mental number line. In contrast, the bisection of non-numerical intervals showed a very different pattern. The results suggest that the spatial layout characterizing numerical representations constitutes a specific property of numbers rather than a general characteristic of ordered sequences.     

The role of ventromedial prefrontal cortex in navigation: a case of impaired wayfinding and rehabilitation

A patient with ventromedial prefrontal damage, LG, is described who had a severe difficulty in wayfinding in his home town, despite intact knowledge for landmarks and routes in town. Although able to recall his spatial goals, LG often headed to familiar, "attractor" locations while navigating, losing his way in the process. Both a laboratory and an ecological study showed that spatial navigation improved when the patient was periodically reminded of, or asked to recall, the goal destination along his route. It is suggested that the ventromedial prefrontal cortex is necessary to maintain actively the goal destination in working memory, for use in navigation.     

The neural dynamics of timed motor tasks: evidence from a synchronization-continuation paradigm

Distinct processing that integrates an accurate time scale is necessary for optimal motor behaviour. In the present study, corticocortical interactions as determined by EEG coherence were assessed in a synchronization–continuation paradigm during which subjects initially performed tapping movements in synchrony with external cues, followed by internal pacing of the target interval when the metronome was switched off. Unimanual and bimanual tasks were executed, and continuation of tapping was conducted with the same or different effector(s). The data showed an increased degree of mesial–central connectivity in the unpaced as compared to paced performance that was independent of task complexity, pointing to a general intensified demand of temporal processing when external cues are unavailable. When switching temporal information between effectors, coherence increased across the motor network. This increase depended upon preceding task complexity, and was most prominent for interhemispheric connections when performing unimanual tasks following bimanual pacing. Overall the data illustrate that timing of skilled actions can easily be transferred between effectors, although increased neural resources are required to conform to the temporal and motor constraints.     

Microcircuitry of posterior cingulate cortex in vitro: electrophysiology and laminar analysis using the current source density method

We used current source density (CSD) analysis of a laminar profile of subicular stimulus-evoked field potentials recorded in cortical slices in vitro to characterize the interlaminar microcircuitry of posterior cingulate cortex. Neuroanatomic and electrophysiologic data indicate that subiculocingulate tract (SCT) afferents monosynaptically excite apical dendrites of deep laminae (V–VI) neurons, evoking pure EPSPs, while superficial laminae (II/III–IV) neurons are driven polysynaptically, evoking a mixture of longer latency EPSPs and IPSPs. Consistent with this model, CSD analysis of field potential laminar profiles supports the conclusion that activation of excitatory subicular afferent terminal fields in superfical laminae of cingulate cortex elicits primary monosynaptic activation of apical dendrites of deep lamina (V–VI) pyramids. Subsequent EPSP propagation to the somata of these pyramids generated synchronous action potential discharges which appeared to elicit delayed polysynaptic activation of superficial laminae pyramids and interneurons. Latency differences between SCT-stimulus-evoked EPSPs and action potentials in superficial and deep laminae were minimized by stimulus train frequencies of 5–8 Hz, indicating that the proposed microcircuitry can show functional tuning at frequencies characteristic of hippocampal neuronal activity (theta). Such tuning suggests that hippocampal output activity frequency and phase locked to theta rhythm will be preferentially gated through cingulate cortex.     

The conjoint importance of the hippocampus and anterior thalamic nuclei for allocentric spatial learning: evidence from a disconnection study in the rat

A disconnection procedure was used to test whether the hippocampus and anterior thalamic nuclei form functional components of the same spatial memory system. Unilateral excitotoxic lesions were placed in the anterior thalamic (AT) nuclei and hippocampus (HPC) in either the same (AT-HPC Ipsi group) or contralateral (AT-HPC Contra group) hemispheres of rats. The behavioral effects of these combined lesions were compared in several spatial memory tasks sensitive to bilateral hippocampal lesions. In all of the tasks tested, T-maze alternation, radial arm maze, and Morris water maze, those animals with lesions placed in the contralateral hemispheres were more impaired than those animals with lesions in the same hemisphere. These results provide direct support for the notion that the performance of tasks that require spatial memory rely on the operation of the anterior thalamus and hippocampus within an integrated neural network.     

The neural basis of reversible sentence comprehension: evidence from voxel-based lesion symptom mapping in aphasia

We explored the neural basis of reversible sentence comprehension in a large group of aphasic patients (n = 79). Voxel-based lesion symptom mapping revealed a significant association between damage in temporo-parietal cortex and impaired sentence comprehension. This association remained after we controlled for phonological working memory. We hypothesize that this region plays an important role in the thematic or what-where processing of sentences. In contrast, we detected weak or no association between reversible sentence comprehension and the ventrolateral pFC, which includes Broca's area, even for syntactically complex sentences. This casts doubt on theories that presuppose a critical role for this region in syntactic computations.     

The neural basis of perceptual bias and response bias in the Landmark task

Spatial neglect as a multifaceted syndrome may consist of perceptual/attentional as well as motor/intentional components. The present study investigated the lesion anatomy underlying perceptual and response bias using a manual response Landmark task (Bisiach, Ricci, Lualdi, & Colombo, 1998) in 68 patients with right-hemispheric stroke. The two differential aspects of the neglect syndrome were assessed by measuring response tendencies resulting from underestimations of the length of left line segments and from hypometric movement execution towards contralesional space, respectively. Perceptual and response bias were orthogonal components of task performance in the Landmark task. Perceptual as well as response bias both explained variance in the performance of standard paper-and-pencil neglect tests. While lesions within middle frontal, inferior parietal and parieto-occipital brain regions were related to perceptual bias, subcortical lesions within the caudate were related to response bias in the Landmark task. Our data suggest that perceptual/attentional and motor/intentional aspects of neglect are independent components of the syndrome with differential neural underpinnings in fronto-parietal and subcortical brain regions.     

Temporal and spatial neural dynamics in the perception of basic emotions from complex scenes

The different temporal dynamics of emotions are critical to understand their evolutionary role in the regulation of interactions with the surrounding environment. Here, we investigated the temporal dynamics underlying the perception of four basic emotions from complex scenes varying in valence and arousal (fear, disgust, happiness and sadness) with the millisecond time resolution of Electroencephalography (EEG). Event-related potentials were computed and each emotion showed a specific temporal profile, as revealed by distinct time segments of significant differences from the neutral scenes. Fear perception elicited significant activity at the earliest time segments, followed by disgust, happiness and sadness. Moreover, fear, disgust and happiness were characterized by two time segments of significant activity, whereas sadness showed only one long-latency time segment of activity. Multidimensional scaling was used to assess the correspondence between neural temporal dynamics and the subjective experience elicited by the four emotions in a subsequent behavioral task. We found a high coherence between these two classes of data, indicating that psychological categories defining emotions have a close correspondence at the brain level in terms of neural temporal dynamics. Finally, we localized the brain regions of time-dependent activity for each emotion and time segment with the low-resolution brain electromagnetic tomography. Fear and disgust showed widely distributed activations, predominantly in the right hemisphere. Happiness activated a number of areas mostly in the left hemisphere, whereas sadness showed a limited number of active areas at late latency. The present findings indicate that the neural signature of basic emotions can emerge as the byproduct of dynamic spatiotemporal brain networks as investigated with millisecond-range resolution, rather than in time-independent areas involved uniquely in the processing one specific emotion.     

Generator localization by current source density (CSD): Implications of volume conduction and field closure at intracranial and scalp resolutions

The topographic ambiguity and reference-dependency that has plagued EEG/ERP research throughout its history are largely attributable to volume conduction, which may be concisely described by a vector form of Ohm’s Law. This biophysical relationship is common to popular algorithms that infer neuronal generators via inverse solutions. It may be further simplified as Poisson’s source equation, which identifies underlying current generators from estimates of the second spatial derivative of the field potential (Laplacian transformation). Intracranial current source density (CSD) studies have dissected the “cortical dipole” into intracortical sources and sinks, corresponding to physiologically-meaningful patterns of neuronal activity at a sublaminar resolution, much of which is locally cancelled (i.e., closed field). By virtue of the macroscopic scale of the scalp-recorded EEG, a surface Laplacian reflects the radial projections of these underlying currents, representing a unique, unambiguous measure of neuronal activity at scalp. Although the surface Laplacian requires minimal assumptions compared to complex, model-sensitive inverses, the resulting waveform topographies faithfully summarize and simplify essential constraints that must be placed on putative generators of a scalp potential topography, even if they arise from deep or partially-closed fields. CSD methods thereby provide a global empirical and biophysical context for generator localization, spanning scales from intracortical to scalp recordings.     

The neural basis of risk ratings: evidence from a functional magnetic resonance imaging (fMRI) study

Research investigating risk perception suggests that not only the quantitative parameters used in technical risk assessment (i.e., frequency and severity of harm) but also ‘qualitative’ aspects such as the dread a hazard provokes or its controllability influence risk judgments. It remains to be elucidated, however, which neural mechanism underlie risk ratings in healthy subjects. Using fMRI to detect changes in neural activity we compared the neural activations elicited by risk ratings with those elicited by a letter detection task performed on the same stimuli. The latter task served to control for basic stimulus processing, response selection and button-pressing during task performance. Risk ratings differentially activated the medial prefrontal cortex, the inferior frontal gyrus, the cerebellum (P < 0.05, FWE corrected, whole brain approach), and in an additional ROI analysis the amygdala (P < 0.05, FWE corrected). Of these structures, particularly the amygdala and the prefrontal cortex have been previously associated with decisions about affective interference. Furthermore our data suggest both, similarities and differences between the neural correlates of risk ratings and risk taking as involved, for e.g., in gambling tasks.     

The ups and downs (and lefts and rights) of synaesthetic number forms: Validation from spatial cueing and SNARC-type tasks

Typically, numbers are spatially represented using a mental ‘number line’ running from left to right. Individuals with number-form synaesthesia experience numbers as occupying specific spatial coordinates that are much more complex than a typical number line. Two synaesthetes (L and B) describe experiencing the numbers 1 through 10 running vertically from bottom to top, 10-20 horizontally from left to right, 21-40 from right to left, etc. We investigated whether their number forms could bias their spatial attention using a cueing paradigm and a SNARC-type task. In both experiments, the synaesthetes' responses confirmed their synaesthetic number forms. When making odd-even judgments for the numbers 1, 2, 8, and 9, they showed SNARC-compatibility effects for up-down movements (aligned with their number form), but not left-right (misaligned) movements. We conceptually replicated these biases using a spatial cueing paradigm. Both synaesthetes showed significantly faster response times to detect targets on the bottom of the display if preceded by a low number (1, 2), and the top of the display if preceded by a high number (8, 9), whereas they showed no cueing effects when targets appeared on the left or right (misaligned with their number forms). They were however reliably faster to detect left targets following the presentation of numbers 10 and 11, and right targets following numbers 19 and 20 (since 10-20 runs from left to right). In sum, cueing and SNARC tasks can be used to empirically verify synaesthetic number forms, and show that numbers can direct spatial attention to these idiosyncratic locations.     

Unavoidable errors: a spatio-temporal analysis of time-course and neural sources of evoked potentials associated with error processing in a speeded task

The detection of errors is known to be associated with two successive neurophysiological components in EEG, with an early time-course following motor execution: the error-related negativity (ERN/Ne) and late positivity (Pe). The exact cognitive and physiological processes contributing to these two EEG components, as well as their functional independence, are still partly unclear. Furthermore, these components are typically obtained in conditions where errors are rare events relative to correct trials, and thus presumably implicate other cognitive and motivational processes besides error monitoring. Here, we investigated error processing using high-density scalp ERPs and advanced topographical analyses in healthy participants, during a new Go/noGo task that led to many errors within a relatively short period of time, yet without generating frustration or insufficient motivation. ERP results showed the presence of two distinct electrophysiological markers of error monitoring (ERN/Ne and Pe) during this task, even though errors were practically as frequent as correct responses. Topographic mapping analyses showed for the first time that both the ERN/Ne and Pe elicited a specific distribution of electrical activity relative to correct responses (not just a change in the amplitude of electric signals), suggesting the activation of a distinct configuration of intracranial generators during error detection. This was confirmed by additional analyses using dipole source localization, showing generators in anterior cingulate cortex contributing to the ERN/Ne, but in more posterior cingulate regions for Pe. Moreover, we found that, across all participants, the magnitude of the ERN/Ne correlated with the level of state anxiety, even in the subclinical range, whereas the Pe was correlated negatively with the total number of errors and positively with the improvement of response speed on correct trials. By contrast, no significant relation was found between error monitoring ERPs and individual measures of impulsivity. Taken together, these data suggest that these two successive EEG components associated with errors reflect different monitoring processes, with distinct neural substrates in cingulate cortex. While ERN/Ne processes in anterior cingulate might primarily mediate error detection, Pe processes in posterior cingulate might be more directly related to behavioral adjustment based on the outcome of current actions.     

Neural coding of action in three dimensions: Task- and time-invariant reference frames for visuospatial and motor-related activity in parietal area V6A

Goal-directed movements involve a series of neural computations that compare the sensory representations of goal location and effector position, and transform these into motor commands. Neurons in posterior parietal cortex (PPC) control several effectors (e.g., eye, hand, foot) and encode goal location in a variety of spatial coordinate systems, including those anchored to gaze direction, and to the positions of the head, shoulder, or hand. However, there is little evidence on whether reference frames depend also on the effector and/or type of motor response. We addressed this issue in macaque PPC area V6A, where previous reports using a fixate-to-reach in depth task, from different starting arm positions, indicated that most units use mixed body/hand-centered coordinates. Here, we applied singular value decomposition and gradient analyses to characterize the reference frames in V6A while the animals, instead of arm reaching, performed a nonspatial motor response (hand lift). We found that most neurons used mixed body/hand coordinates, instead of “pure” body-, or hand-centered coordinates. During the task progress the effect of hand position on activity became stronger compared to target location. Activity consistent with body-centered coding was present only in a subset of neurons active early in the task. Applying the same analyses to a population of V6A neurons recorded during the fixate-to-reach task yielded similar results. These findings suggest that V6A neurons use consistent reference frames between spatial and nonspatial motor responses, a functional property that may allow the integration of spatial awareness and movement control.