Extension of perceived arm length following tool-use: clues to plasticity of body metrics

Humans hold a very accurate representation of the metrics of their body parts. Recent evidence shows that the spatial estimation of body parts length, as assessed through a bisection task, is even more accurate than that of non-corporeal extrapersonal objects (Sposito, Bolognini, Vallar, Posteraro, & Maravita (2009)). In the present paper we show that human participants estimate the mid-point of their forearm, which was kept in a radial posture, to be more distal following a 15-min training with a 60 cm-long tool as compared to pre tool-use. This outcome is compatible with an increased representation of the participants' forearm length. Control experiments show that this result was not due to a mere distal proprioceptive shift induced by tool-use, and was not replicated following the use of a 20 cm-long, functionally irrelevant tool. These results strongly support the view that, although the inner knowledge of one's own body metrics appears to be one of the more stable features of body representation, body-space interactions requiring the use of tools that extend the natural range of action, entail measurable dynamic changes in the representation of body metrics.     

Processing of peripersonal and extrapersonal space using tools: evidence from visual line bisection in real and virtual environments

The dissociation between peripersonal space (within reaching) and extrapersonal space (beyond reaching) has been reported in studies using the line bisection task in left neglect patients and in healthy participants. Furthermore, this dissociation can be modulated by tool use. We conducted two experiments to compare line bisection in peripersonal (i.e., 30, 60 cm) and extrapersonal space (i.e., 90, 120 cm). Healthy participants bisected visual lines using sticks and a laser pointer, according to the experimental paradigm of Longo and Lourenco [On the nature of near space: Effects of tool use and the transition to far space. Neuropsychologia, 44, 977–981, 2006]. In Experiment 1 participants performed line bisection in a real environment, whereas in Experiment 2 participants performed line bisection in a virtual environment. Results from both experiments revealed an abrupt midpoint shift from the peripersonal to the extrapersonal space but only when a laser pointer was used. In addition, we confirmed that peripersonal space can be extended to extrapersonal space when participants used a stick. Notably, virtual reality can be a useful technique for studying the dissociation between peripersonal and extrapersonal space and their interaction by means of tool use.     

Differences between left- and right-sided neglect revisited: A large cohort study across multiple domains

Unilateral spatial neglect (USN) is a syndrome that can occur after right- and left-hemisphere damage. It is generally accepted that left-sided USN is more severe than right-sided USN. Evidence for such a difference in other domains is lacking. Primary aims were to compare frequency, severity, region specificity, cognition, physical functioning, and physical independence between left and right USN. Secondary aims were to compare lesion characteristics. A total of 335 stroke patients admitted for inpatient rehabilitation were included. The severity of the lateralized attentional deficit was measured with a shape cancellation and line bisection test (in peripersonal and extrapersonal space) and the Catherine Bergego scale. The Mini-Mental State Examination, Stichting Afasie Nederland score, search organization (i.e., best R and intersections rate), Motricity Index, balance, mobility, and self-care were assessed. Measures were statistically compared between left, right, and no USN patients. Lesion overlay plots were compared with lesion subtraction analyses. Results: Left USN (15.82%) was more frequent than right USN (9.25%). Demographic and stroke characteristics were comparable between groups. The lateralized attentional deficit was most severe in left USN. USN in both peripersonal and extrapersonal space was more frequently left-sided in nature. Search efficiency was lower in left USN. Balance was poorer in right USN. No differences between left and right USN were found for cognitive ability, communication, motor strength, mobility, and self-care. Most patients with left USN had right-hemispheric lesions, whereas patients with right USN could have lesions in either the left or the right hemisphere. To conclude, left and right USN are both common after stroke. Although the lateralized attention deficit is worse in left than in right USN, consequences at the level of physical functioning and physical independence are largely comparable. From a clinical perspective, it is important to systematically screen for USN, both after right- and after left-hemisphere damage.     

Somatosensory cortical representation of the body size

The knowledge of the size of our own body parts is essential for accurately moving in space and efficiently interact with objects. A distorted perceptual representation of the body size often represents a core diagnostic criterion for some psychopathological conditions. The metric representation of the body was shown to depend on somatosensory afferences: local deafferentation indeed causes a perceptual distortion of the size of the anesthetized body part. A specular effect can be induced by altering the cortical map of body parts in the primary somatosensory cortex. Indeed, the present study demonstrates, in healthy adult participants, that repetitive Transcranial Magnetic Stimulation to the somatosensory cortical map of the hand in both hemispheres causes a perceptual distortion (i.e., an overestimation) of the size of the participants' own hand (Experiments 1–3), which does not involve other body parts (i.e., the foot, Experiment 2). Instead, the stimulation of the inferior parietal lobule of both hemispheres does not affect the perception of the own body size (Experiment 4). These results highlight the role of the primary somatosensory cortex in the building up and updating of the metric of body parts: somatosensory cortical activity not only shapes our somatosensation, it also affects how we perceive the dimension of our body.     

Heterotopagnosia: When I point at parts of your body

Heterotopagnosia is the acquired inability of brain-lesioned patients to point at someone else's body parts when prompted. The cognitive basis of this disorder is unclear. It might result from a biological function deficit critical for communication in human beings; alternatively, it could result from the disruption of a body representation. Here, we report three patients with heterotopagnosia following a recent left parieto-occipital stroke and a previous insular lesion. The patients were tested on their ability to name, point out and grasp several targets including body parts (own, real others’ and figurative others’). Language, visuo-spatial deficits or any confounding neuropsychological disorders were controlled for. We found that the patients erroneously pointed to their own body parts when asked to point at someone else's. Strikingly, their ability to grasp someone else's body parts was largely unimpaired. The dissociation between their grasping and communicative pointing abilities supports the hypothesis that heterotopagnosia is a disorder of communicative function conveyed by pointing but not by grasping. In addition, pointing performance in our patients varied according to the target: the more similar the target was to a real person, the worse the patients’ pointing performance. We suggest that communicative pointing might require a specific representation of the addressee's body and point of view, a heterocentric representation. In the patients described here this phenomenon resulted from a combined insulo-parietal lesion, which may explain why, in contrast to other patients described previously, the heterotopagnosia was long-lasting.     

Multiple perceptual distortions and their modulation in leftsided visual neglect

There is an ongoing debate concerning the perception and neural representation of space in neglect. Four experiments are here reported designed to further investigate the nature of perceptual distortions and their modifiability in patients with neglect. In Experiment 1 it was found that neglect patients, in contrast to left- or right-hemisphere lesioned control patients and normal subjects, show similar distortions of perceived visual space when judging the extension of horizontal distances (space distortion) as compared to the horizontal size of objects (size distortion). Similar deficits were present in most neglect patients in a newly developed space bisection task. These results attest that neglect patients have perceptual distortions related to within-object (size) and between-object (distance) spatial processing in their horizontal plane. Objects were oversized by 33% and distances by 19% horizontally in neglect patients, whereas all control groups showed nearly veridical spatial coding (deviations<5%). In Experiment 2 the modifiability of these distortions was tested by the use of slow visual background motion. Leftward, coherent background motion transiently restored normal horizontal size and distance coding in neglect patients, whereas rightward motion aggravated the deficit significantly in the distance task, but not in the size task. None of the other subject groups showed any influence of background motion on spatial judgments. Experiment 3 evaluated possible effects of simultaneous vs successive stimulus presentation in perceptual distortions, thus modulating attentional factors. Neglect patients performed significantly better - although not normal - with a successive presentation of the spatial stimuli (2 s, 10 s delay) as compared to the simultaneous condition in the size judgment task, but not in the distance task. In contrast, this manipulation had no effects in any of the control groups. Experiment 4 reports more detailed results of subject J.S., a neglect patient with a right mediotemporal lesion, who showed a marked horizontal size distortion, but normal horizontal distance judgments. Despite some fluctuation in J.S.'s size judgments he showed a significant overestimation of horizontal object size by +22% to +40% across several test blocks and testing sessions. Thus, performance fluctuations due to attentional or other reasons cannot fully account for J.S.'s dissociation in size vs distance judgments. He thus shows that the visual coding of horizontal spatial extension within an object can be dissociated from that of the spatial extension between objects along the horizontal plane. Finally, performance in the three spatial tasks (used in Experiment 1) was found to correlate significantly with three typical tests of spatial neglect (line bisection, cancellation, copying, r=0.42-0.77), thereby indicating a significant relationship to the neglect syndrome.Together, the results of the four experiments are interpreted in support of multiple spatial-perceptual distortions in visual neglect, which are influenced by visual motion and attention. Perceptual distortions relating to objects and space between objects are present in most neglect patients, but may dissociate, as in case J.S. It is argued that these might reflect the existence of several, partially overlapping and diverging neural maps for the representation of different spatial attributes in the horizontal plane. Furthermore, it is hypothesized that these perceptual distortions constitute an important element of the spatial-perceptual deficits encountered in the syndrome, contribute to its severity, but are not the key deficit of the disorder.     

The Brentano Illusion Test (BRIT): An implicit task of perceptual processing for the assessment of visual field defects in neglect patients

ABSTRACT

In brain damaged patients with unilateral spatial neglect (USN), the differential diagnosis between the presence and absence of a unilateral visual half-field deficit (VHFD) is hampered by the similarity of their phenomenology. The absence of stimuli detection in the contralateral visual field, indeed, can be due to the co-occurrence of USN and VHFD or the sole presence of the USN. The disentangling of the two conditions is required to devise more specific rehabilitation programmes. Daini et al. [2002. Exploring the syndrome of spatial unilateral neglect through an illusion of length. Experimental Brain Research, 144(2), 224–237.] reported a difference in performance for the two conditions when the tasks required the bisection of Brentano illusory stimuli. Only when USN and VHFD co-occurred, the leftward illusory effect was disrupted. Based on previous findings, in this cross-sectional study, we developed the Brentano Illusion Test (BRIT), a clinical tool that helps the identification of VHFD in USN patients. The BRIT is a simple behavioural test of line bisection aimed at verifying the presence or absence of implicit processing in USN and thus helping the diagnosis of VHFD in USN patients; it also provides normative data for the line bisection task and the length effect.     

Listening to numbers affects visual and haptic bisection in healthy individuals and neglect patients

There is evidence that humans represent numbers in the form of a mental number line (MNL). Here we show that the MNL modulates the representation of visual and haptic space both in healthy individuals and right-brain-damaged patients, both with and without left unilateral spatial neglect (USN). Participants were asked to estimate the midpoint of visually or haptically explored rods while listening to task-irrelevant stimuli: a small digit ("2"), a large digit ("8"), or a non-numerical auditory stimulus ("blah"). In a control silent condition, the bisection error of USN patients was biased rightwards (namely, the marker of USN) only in the visual modality. Regardless of the direction of the bisection error committed in silent trials, listening to the small digit shifted the perceived midline leftwards, and listening to the large digit shifted the perceived midline rightwards, compared to a control condition in which a neutral syllable ("blah") was presented. The shift induced by listening to numbers occurred independently of the modality of response (i.e., both in vision and haptics), and in every group of participants. Interestingly, the effect of auditory numbers processing on space estimation was overall larger for haptically than for visually explored space in all participants. In conclusion, the present data show that listening to irrelevant numbers affects space perception also in patients with left USN, indicating that the spatial representation and attention processes disrupted by USN are not involved in these numerical magnitude-spatial effects.     

When far becomes near: remapping of space by tool use

Far (extrapersonal) and near (peripersonal) spaces are behaviorally defined as the space outside the hand-reaching distance and the space within the hand-reaching distance. Animal and human studies have confirmed this distinction, showing that space is not homogeneously represented in the brain. In this paper we demonstrate that the coding of space as "far" and "near" is not only determined by the hand-reaching distance, but it is also dependent on how the brain represents the extension of the body space. We will show that when the cerebral representation of body space is extended to include objects or tools used by the subject, space previously mapped as far can be remapped as near. Patient P.P., after a right hemisphere stroke, showed a dissociation between near and far spaces in the manifestation of neglect. Indeed, in a line bisection task, neglect was apparent in near space, but not in far space when bisection in the far space was performed with a projection lightpen. However, when in the far space bisection was performed with a stick, used by the patient to reach the line, neglect appeared and was as severe as neglect in the near space. An artificial extension of the patient's body (the stick) caused a remapping of far space as near space.     

Selective impairment of self body-parts processing in right brain-damaged patients

To investigate whether the processing of the visual appearance of one's own body, that is the corporeal self is a unified or modular function we submitted eight right brain-damaged (RBD) patients and a group of fourteen age-matched neurologically healthy subjects, to a visual matching-to-sample task testing for corporeal self processing. If corporeal self processing is a unique function (i.e., body- and face-parts are processed by the same network), patients impaired in self body-parts (i.e., showing no self-advantage) should be impaired also in self face-parts; alternatively, if corporeal self processing is a modular function (i.e., body- and face-parts are processed by different networks), patients impaired in self body-parts should be unimpaired in self face-parts, unless the face-module is also damaged by the lesion. Results showed that healthy participants were more accurate in processing pictures representing their own as compared to other people's body- and face-parts, showing the so-called self-advantage. The patients' findings revealed a simple dissociation, in that patients who were impaired in the processing of self-related body-parts showed a preserved self-advantage when processing self-related face-parts, thus providing initial evidence of a modular representation of the corporeal self.     

Viewing another person's body as a target object: a behavioural and PET study of pointing

Humans usually point at objects to communicate with other persons, although they generally avoid pointing at the other's body. Moreover, patients with heterotopagnosia after left parietal damage cannot point at another person's body parts, although they can point at objects and at their own body parts and although they can grasp the others' body parts. Strikingly, their performance gradually improves for figurative human body targets. Altogether, this suggests that the body of another real person holds a specific status in communicative pointing. Here, we test in healthy individuals whether performance for communicative pointing is influenced by the communicative capacity of the target. In Experiment 1, pointing at another real person's body parts was compared to pointing at objects, and in Experiment 2, the person was replaced by a manikin. While reaction times for pointing at objects were shorter compared to pointing at other person's body parts, they were similar for objects and manikin body parts. By adapting Experiment 1 to PET-scan imaging (Experiment 3), we showed that, compared to pointing at objects, the brain network for pointing at other person's body parts involves the left posterior intraparietal sulcus, lesion of which could cause heterotopagnosia. Taken together, our results indicate that the specificity of pointing at another person's body goes beyond the visuo-spatial features of the human body and might rather rely on its communicative capacity.     

Unilateral spatial neglect: a dramatic but often neglected consequence of right hemisphere damage

INTRODUCTION: Unilateral Spatial Neglect (USN) is a common consequence of right brain damage. In the most severe cases, behavioral signs of USN can last several years and compromise patients' autonomy and social rehabilitation. These clinical facts stress the need for reliable procedures of diagnosis and rehabilitation. STATE OF THE ART: The last 3 decades have witnessed an explosion of studies on USN, which raises issues related to complex cognitive activities such as mental representation, spatial attention and consciousness. USN is probably a heterogeneous syndrome, but some of its underlying mechanisms might be understood as an association of disorders of spatial attention. A bias of automatic orienting towards right-sided objects seems typical of left USN. Afterwards, patients find it difficult to disengage their attention in order to explore the rest of the visual scene. Neglected objects are sometimes processed in an "implicit" way. PERSPECTIVES: The development of behavioural paradigms and of neuroimaging techniques and their application to the study of USN has advanced our understanding of the functional mechanisms of attention and spatial awareness, as well as of their neural bases. A number of new procedures for rehabilitation have recently been proposed. CONCLUSION: The present review describes the clinical presentation of USN, its anatomical basis and some of possible accounts of different aspects of neglect behavior. Results of computer simulations and of rehabilitation techniques are also presented with implications for the functioning of normal neurocognitive systems.     

The effect of space location on neglect depends on the nature of the task

It has been often reported that in patients with visual neglect line bisection is more accurate in the right than in the left hemispace. However, no data are available on the effect of hemispace on reading errors associated with neglect. We examined a 62-year-old man who presented with severe left neglect following a large infarction in the right cerebral hemisphere. The patient was asked to read 180 words aloud and to bisect 90 lines. Stimuli were presented in three different spatial locations: across the centre, to the right or to the left of the body midline. Line bisection was significantly more accurate in the right hemispace compared with the centre, or the left hemispace. In contrast, reading was significantly more accurate with words presented on the left side than on the centre or right side. This is the first time that such dissociation has been reported. We hypothesize that the dissociation depends on the nature of the stimuli and on the different cognitive demands of the tasks.     

Object orientation agnosia: a failure to find the axis?

A dissociation between the ability to recognize misoriented objects and to determine their orientation has been reported in a small number of patients with vascular lesions. In this article, we describe a 57-year-old man with probable Alzheimer' s disease who shows the same dissociation. Neuroimaging findings indicated marked hypometabolism in the posterior cortical regions, particularly the postero-superior parietal lobes. Clinically, the patient had good object recognition accompanied by severely impaired spatial abilities. The experimental investigations comprised a variety of tasks in which he identified misoriented objects, evaluated the orientation of single objects, or discriminated the orientation of simultaneously presented items. Results revealed that his object recognition was independent of orientation and was largely mediated by salient features. With respect to orientation judgements, the patient displayed a profound inability to judge the orientation of nonupright objects, but remarkably intact (though largely implicit) knowledge of the upright orientation. Strikingly, his orientation judgements were also more accurate for upside-down objects than for other orientations (i.e., 90 degrees ). We interpret these results as evidence that judgements about object orientation are facilitated when the orientation of the principal axis of the object matches that of an internal representation. We propose that the inability to determine other orientations may be due to the failure of an "axis-finding" mechanism implemented in the posterior parietal lobes, that translates between object-centered and eye-centered coordinates appropriate for guiding visual scanning.     

"Mind the gap": the size-distance dissociation in visual neglect is a cueing effect

There is a growing body of evidence that the processes mediating the allocation of spatial attention within objects may be separable from those governing attentional distribution between objects. In the neglect literature, a related proposal has been made regarding the perception of (within-object) sizes and (between-object) distances. This proposal follows observations that, in size-matching and bisection tasks, neglect is more strongly expressed when patients are required to attend to the sizes of discrete objects than to the (unfilled) distances between objects. These findings are consistent with a partial dissociation between size and distance processing, but a simpler alternative must also be considered. Whilst a neglect patient may fail to explore the full extent of a solid stimulus, the estimation of an unfilled distance requires that both endpoints be inspected before the task can be attempted at all. The attentional cueing implicit in distance estimation tasks might thus account for their superior performance by neglect patients. We report two bisection studies that address this issue. The first confirmed, amongst patients with left visual neglect, a reliable reduction of rightward error for unfilled "gap" stimuli as compared with solid lines. The second study assessed the cause of this reduction, deconfounding the effects of stimulus type (lines vs. gaps) and attentional cueing, by applying an explicit cueing manipulation to line and gap bisection tasks. Under these matched cueing conditions, all patients performed similarly on line and gap bisection tasks, suggesting that the reduction of neglect typically observed for gap stimuli may be attributable entirely to cueing effects. We found no evidence that a spatial extent, once fully attended, is judged any differently according to whether it is filled or unfilled.     

Peculiar body representation alterations in hemineglect: a case report

We report the case of FP affected by personal and extrapersonal neglect and a body representation deficit characterized by delusional ideas. When FP performed the human figure, he placed body parts to the left, despite his extrapersonal neglect. Differently, when he performed the car figure, he placed all parts to the right, in line with his deficit. Comparing FP with a small patient group with the same clinical features without delusional ideas about body emerged that he was the only one to suffer from a specific body representation deficit characterized by a lack of body ownership sense.     

When action is not enough: tool-use reveals tactile-dependent access to Body Schema

Proper motor control of our own body implies a reliable representation of body parts. This information is supposed to be stored in the Body Schema (BS), a body representation that appears separate from a more perceptual body representation, the Body Image (BI). The dissociation between BS for action and BI for perception, originally based on neuropsychological evidence, has recently become the focus of behavioural studies in physiological conditions. By inducing the rubber hand illusion in healthy participants, Kammers et al. (2009) showed perceptual changes attributable to the BI to which the BS, as indexed via motor tasks, was immune. To more definitively support the existence of dissociable body representations in physiological conditions, here we tested for the opposite dissociation, namely, whether a tool-use paradigm would induce a functional update of the BS (via a motor localization task) without affecting the BI (via a perceptual localization task). Healthy subjects were required to localize three anatomical landmarks on their right arm, before and after using the same arm to control a tool. In addition to this classical task-dependency approach, we assessed whether preferential access to the BS could also depend upon the way positional information about forearm targets is provided, to subsequently execute the same task. To this aim, participants performed either verbally or tactually driven versions of the motor and perceptual localization tasks. Results showed that both the motor and perceptual tasks were sensitive to the update of the forearm representation, but only when the localization task (perceptual or motor) was driven by a tactile input. This pattern reveals that the motor output is not sufficient per se, but has to be coupled with tactually mediated information to guarantee access to the BS. These findings shade a new light on the action-perception models of body representations and underlie how functional plasticity may be a useful tool to clarify their operational definition.     

Normative data for distal line bisection and baking tray task

Line bisection is one of the tests used to diagnose unilateral spatial neglect (USN). Despite its wide application, no procedure or norms were available for the distal variant when the task was performed at distance with a laser pointer. Furthermore, the baking tray task was an ecological test aimed at diagnosing USN in a more natural context. The aim of this study was to collect normative values for these two tests in an Italian population. We recruited a sample of 191 healthy subjects with ages ranging from 20 to 89 years. They performed line bisection with a laser pointer on three different line lengths (1, 1.5, and 2 m) at a distance of 3 m. After this task, the subjects performed the baking tray task and a second repetition of line bisection to test the reliability of measurement. Multiple regression analysis revealed no significant effects of demographic variables on the performance of both tests. Normative cut-off values for the two tests were developed using non-parametric tolerance intervals. The results formed the basis for clinical use of these two tools for assessing lateralized performance of patients with brain injury and for diagnosing USN.     

Distance-mediated spatial neglect

Unilateral spatial neglect (USN) is usually assessed by means of individual stimuli or single arrays of stimuli. Seldom are stimuli presented as multiple objects or in spatially separated blocks, except in some tests for object-based neglect. The distance between individual objects or blocks of stimuli in such stimuli is implicitly considered irrelevant. We report on the case of a patient, EC, who showed severe USN in his everyday behavior, yet performed normally on standard tests for USN. Presented with stimuli in separate blocks, he performed flawlessly with 4 cm gaps between blocks, yet ignored all leftward blocks of stimuli when the gap was larger than this. EC’s dissociation between good performance on standard tasks and severe neglect with separate groups of stimuli, and the distance-mediated nature of his USN are novel observations with relevant theoretical and clinical implications.     

So near yet so far: neglect in far or near space depends on tool use.

The study of unilateral spatial neglect has shown that space can be dissociated on a peripersonal versus extrapersonal basis. We report a novel type of dissociation based on tool use in a patient suffering from left neglect. Line bisection was carried out in near and far space, using a stick and a laser pointer. A rightward bias was always found for the former, but not for the latter. Neglect thus appears to be contingent not only on distance, but also on the motor action required by the task.