Mental rotation meets the motion aftereffect: the role of hV5/MT+ in visual mental imagery

A growing number of studies show that visual mental imagery recruits the same brain areas as visual perception. Although the necessity of hV5/MT+ for motion perception has been revealed by means of TMS, its relevance for motion imagery remains unclear. We induced a direction-selective adaptation in hV5/MT+ by means of an MAE while subjects performed a mental rotation task that elicits imagined motion. We concurrently measured behavioral performance and neural activity with fMRI, enabling us to directly assess the effect of a perturbation of hV5/MT+ on other cortical areas involved in the mental rotation task. The activity in hV5/MT+ increased as more mental rotation was required, and the perturbation of hV5/MT+ affected behavioral performance as well as the neural activity in this area. Moreover, several regions in the posterior parietal cortex were also affected by this perturbation. Our results show that hV5/MT+ is required for imagined visual motion and engages in an interaction with parietal cortex during this cognitive process

Tell it to the hand: Attentional modulation in the identification of misoriented chiral objects

Research in the field of cognitive neuroscience and neuropsychology on spatial cognition and mental imagery has increased considerably over the last few decades. While at the beginning of the XX century studying imagery was considered an object of derision \u2013 a \u2015sheer bunk\u2016 (Watson, 1928) \u2013 at the present, imagery researchers have successfully developed models and improved behavioral and neurophysiological measures (e.g., Kosslyn et al., 2006). Mental rotation constituted a major advance in terms of behavioral measures sensitive to imaginative operations executed on visual representations (i.e., Shepard & Cooper, 1982). The linearity of modulation between response times and angular disparity of the images allowed a quantitative estimate of imagery processes. The experiments described in the present thesis were motivated by the intent to continue and extend the understanding of such fascinating mental phenomena. The evolution of the present work took initial steps from the adoption of a behavioral paradigm, the hand laterality judgment task, as privileged tool for studying motor imagery in healthy individuals and brain-damaged patients. The similarity with mental rotation tasks and the implicit nature of the task made it the best candidate to test hypotheses regarding the mental simulation of body movements. In this task, response times are linearly affected by the angular departures the hand pictures are shown in, as for mental rotation, and their distributions are asymmetric between left and right hands. Drawing from these task features a widely held view posits that laterality judgment of rotated hand pictures requires participants to imagine hand-arm movements, although they receive no instruction to do so (e.g., Parsons, 1987a; Parsons, 1994). In Chapter 1, I provided a review of the relevant literature on visual and motor imagery. Particular aspects of the mental rotation literature are also explored. In Chapter 2, I examined the hand laterality task and the vast literature of studies that employed this task as means to test motor imagery processes. An alternative view to the motor imagery account is also discussed (i.e., the disembodied account). In Chapter 3, I exploited the hand laterality task, and a visual laterality task (Tomasino et al., 2010) to test motor and visual imagery abilities in a group of healthy aged individuals. In Chapter 4, I described an alternative view that has been proposed by others to explain the pattern of RTs in the hand laterality task: The multisensory integration account (Grafton & Viswanathan, 2014). In this view, hand laterality is recognized by pairing information between the seen hand's visual features and the observer's felt own hand. In Chapter 5, I tested and found evidence for a new interpretation of the particular configuration of response times in the hand laterality task. I demonstrated a spatial compatibility effect for rotated pictures of hands given by the interaction between the direction of stimulus rotation (clockwise vs. counterclockwise) and the laterality of the motor response. These effects changed by following temporal dynamics that were attributed to shifts of spatial attention. In the same chapter, I conducted other psychophysics experiments that confirmed the role of spatial attention and that ruled out the view of multisensory integration as the key aspect in determining the asymmetries of the response times' distribution. In Chapter 6, I conducted a study with patients suffering from Unilateral Neglect in which they performed the hand laterality task and a visual laterality task. The findings indicated that patients failed to integrate visual information with spatially incompatible responses irrespective of the type of task, and depending on egocentric stimulus-response spatial codes. A general discussion is presented in Chapter 7

Spatial analysis in the human cerebral cortex : behavioural and functional magnetic resonance studies of spatial transformations in visual perception and imagery

Die vorliegende Dissertation berichtet über eine Serie von Verhaltens- und funktionellen Bildgebungsstudien zu experimentalpsychologischen Paradigmata, die eine räumliche Analyse und Koordinatentransformation von Material der visuellen Wahrnehmung oder Vorstellung beinhalten. Nach einer Einführung in die Prinzipien und Techniken der funktionellen Kernspintomographie, der hier benutzten Methode für die Messung von Gehirnaktivität, werden die Versuche einer Replikation des berühmten Stratton'schen Umkehrbrillen-Experiments dargestellt (Kapitel 1). Unsere vier Probanden zeigten zwar eine zügige Anpassung der visuomotorischen Funktionen an die neue visuelle Umwelt, berichteten aber, anders als Stratton, nicht, daß sie die Welt nach einigen Tagen mit der Umkehrbrille wieder normal sähen. Diese Persistenz des umgekehrten Bildes wurde durch eine psychphysische Testbatterie bestätigt. Des weiteren ergaben die funktionellen Kernspinmessungen, daß sich die kortikale retinotope Organisation im Verlaufe des Experiments nicht geändert hat. Da sich also Strattons Haupteffekt, das Aufrechtsehen durch die Umkehrbrille nach einwöchiger Adaptation, nicht replizieren ließ, werden andere Möglichkeiten der Interpretation der verschiedenen Umkehrexperimente der letzten hundert Jahre vorgeschlagen. Dieses Ergebnis einer funktionellen Anpassung ohne größere Änderungen der visuellen Wahrnehmung (und ohne Veränderungen der Repräsentation der Netzhautareale in der Sehrinde) führte zu der Hypothese, daß die erforderlichen Transformationen auf einer höheren Stufe der kortikalen Hierarchie der visuellen Verarbeitung erfolgen. Zur Testung dieser Hypothese wurde eine funktionelle Kernspinstudie des Umkehrlesens durchgeführt (Kapitel 2). Hierbei lasen die Probanden Wörter und Sätze in Spiegelschrift oder auf dem Kopf. Der neuronale Mechanismus der räumlichen Transformationen, die zur Bewältigung dieser Aufgabe nötig sind, konnte in bestimmten Regionen des Parietallappens, die zwischen den Probanden sehr konstant waren, lokalisiert werden. Weiterhin fand sich eine Koaktivierung okzipitootemporaler Objekterkennungs-Areale. Die Spezifizität der parietalen Aktivierungsfoci wurde durch ein Kontrollexperiment bestätigt, in welchem das kortikale System für räumliche Transformationen von den Netzwerken der allgemeinen visuellen Aufmerksamkeit und der Augenbewegungskontrolle unterschieden werden konnte. In einem weiteren Experiment wurden die räumlichen Funktionen des Parietallappens unter dem Vorzeichen der visuellen Vorstellung untersucht. Als Paradigma wurde der "mental clock" - Test verwendet, bei welchem die Probanden die Winkel der Zeiger zweier Uhren vergleichen müssen, deren Zeiten nur akustisch vorgegeben werden (Kapitel 3). Diese Aufgabe erfordert die Generierung eines entsprechenden Vorstellungsbildes und dessen räumliche Analyse, stellt also sowohl ein kontrolliertes Vorstellungs-Paradigma als auch einen Test räumlicher Funktionen dar, der nicht auf visuell präsentiertem Material beruht. Das parietale Aktivierungsmuster, das der Analyse der Winkel dieser vorgestellten Uhren zugeschrieben werden konnte, entsprach weitgehend demjenigen, das mit der räumlichen Transformation von Buchstaben verbunden war. Es handelt sich also wahrscheinlich um ein kortikales System für räumliche Analyse und Koordinatentransformationen, das nicht auf eine visuelle Stimulation angewiesen ist, sondern auch bei bloßer visueller Vorstellung aktiv werden kann. Die vorgelegten Resultate werden im Kontext neuerer neuropsychologischer Befunde zu Defiziten räumlicher Analyse und Vorstellung bei Läsionen des Parietallappens diskutiert (Kapitel 4). Auch die methodologischen Probleme der kognitiven Subtraktion, die in unseren Studien teilweise benutzt wurde, werden behandelt. Dabei wird erläutert, inwiefern diese für die Beurteilung der vorgelegten Studien nur von untergeordneter Bedeutung sind. Nichtsdestoweniger schlagen wir Modifikationen der experimentellen Paradigmata im Sinne des parametrischen Designs und des "event-related functional magnetic resonance imaging" vor, die bei zukünftigen Studien einen vollständigen Verzicht auf die kognitive Subtraktion ermöglichen dürften. Schließlich wird die Bedeutung der vorgelegten Ergebnisse für die Erforschung der Anpassungsfähigkeit des menschlichen Gehirns und des Verhältnisses von Vorstellung und visueller Wahrnehmung dargelegt

Body Context and Posture Affect Mental Imagery of Hands

Different visual stimuli have been shown to recruit different mental imagery strategies. However the role of specific visual stimuli properties related to body context and posture in mental imagery is still under debate. Aiming to dissociate the behavioural correlates of mental processing of visual stimuli characterized by different body context, in the present study we investigated whether the mental rotation of stimuli showing either hands as attached to a body (hands-on-body) or not (hands-only), would be based on different mechanisms. We further examined the effects of postural changes on the mental rotation of both stimuli. Thirty healthy volunteers verbally judged the laterality of rotated hands-only and hands-on-body stimuli presented from the dorsum- or the palm-view, while positioning their hands on their knees (front postural condition) or behind their back (back postural condition). Mental rotation of hands-only, but not of hands-on-body, was modulated by the stimulus view and orientation. Additionally, only the hands-only stimuli were mentally rotated at different speeds according to the postural conditions. This indicates that different stimulus-related mechanisms are recruited in mental rotation by changing the bodily context in which a particular body part is presented. The present data suggest that, with respect to hands-only, mental rotation of hands-on-body is less dependent on biomechanical constraints and proprioceptive input. We interpret our results as evidence for preferential processing of visual- rather than kinesthetic-based mechanisms during mental transformation of hands-on-body and hands-only, respectively

Effects of a Body-Oriented Response Measure on the Neural Substrate of Imagined Perspective Rotations

Previous behavioral studies suggest that response measures related to the body, such as pointing, serve to anchor participants to their physical body during mental rotation tasks in which their perspective must be shifted elsewhere. This study investigated whether such measures engage spatial and low-level cortical motor areas of the brain more readily than non-body-related measures. We directly compared activation found in two imagined perspective rotation tasks, using responses that varied in the degree to which they emphasized the human body. In the body minimize condition, participants imagined rotating themselves around an object and judged whether a prescribed part of the object would be visible from the imagined viewpoint. In the body maximize condition, participants imagined rotating around the object and then located the prescribed object part with respect to their bodies. A direct comparison of neural activation in both conditions revealed distinct yet overlapping neural regions. The body maximize condition yielded activation in low-level cortical motor areas such as premotor cortex and primary motor cortex, as well as bilateral spatial processing areas. The body minimize condition yielded activation in nonmotoric egocentric processing regions. However, both conditions showed activation in the parietal-occipital region that is thought to be involved in egocentric transformations. These findings are discussed in the context of recent hypotheses regarding the role of the body percept in imagined egocentric transformations. © 2009 Massachusetts Institute of Technology

Are all spatial reference frames egocentric? Reinterpreting evidence for allocentric, object-centered, or world-centered reference frames

The use and neural representation of egocentric spatial reference frames is well-documented. In contrast, whether the brain represents spatial relationships between objects in allocentric, object-centered, or world-centered coordinates is debated. Here, I review behavioral, neuropsychological, neurophysiological (neuronal recording), and neuroimaging evidence for and against allocentric, object-centered, or world-centered spatial reference frames. Based on theoretical considerations, simulations, and empirical findings from spatial navigation, spatial judgments, and goal-directed movements, I suggest that all spatial representations may in fact be dependent on egocentric reference frames

Dynamic Visuomotor Transformation Involved with Remote Flying of a Plane Utilizes the ‘Mirror Neuron’ System

Brain regions involved with processing dynamic visuomotor representational transformation are investigated using fMRI. The perceptual-motor task involved flying (or observing) a plane through a simulated Red Bull Air Race course in first person and third person chase perspective. The third person perspective is akin to remote operation of a vehicle. The ability for humans to remotely operate vehicles likely has its roots in neural processes related to imitation in which visuomotor transformation is necessary to interpret the action goals in an egocentric manner suitable for execution. In this experiment for 3rd person perspective the visuomotor transformation is dynamically changing in accordance to the orientation of the plane. It was predicted that 3rd person remote flying, over 1st, would utilize brain regions composing the ‘Mirror Neuron’ system that is thought to be intimately involved with imitation for both execution and observation tasks. Consistent with this prediction differential brain activity was present for 3rd person over 1st person perspectives for both execution and observation tasks in left ventral premotor cortex, right dorsal premotor cortex, and inferior parietal lobule bilaterally (Mirror Neuron System) (Behaviorally: 1st>3rd). These regions additionally showed greater activity for flying (execution) over watching (observation) conditions. Even though visual and motor aspects of the tasks were controlled for, differential activity was also found in brain regions involved with tool use, motion perception, and body perspective including left cerebellum, temporo-occipital regions, lateral occipital cortex, medial temporal region, and extrastriate body area. This experiment successfully demonstrates that a complex perceptual motor real-world task can be utilized to investigate visuomotor processing. This approach (Aviation Cerebral Experimental Sciences ACES) focusing on direct application to lab and field is in contrast to standard methodology in which tasks and conditions are reduced to their simplest forms that are remote from daily life experience

Beyond variability: Subjective timing and the neurophysiology of motor cognition.

Movement simulation helps increasing the chances to reach goals. A cognitive task used to study the neuro-behavioral aspects of movement simulation is mental rotation: people mentally re-orient rotated pictures of hands. However, the involvement of the primary motor cortex (M1) in mental rotation is largely controversial. Such inconsistency could arise from potential methodological flaws in experimental procedures and data analysis. In particular, until now, the timing of M1 activity has been computed in absolute terms: from the onset of mental rotation (onset-locked), neglecting intra- and inter-subject variability. A novel phase-locked approach is introduced to synchronize the same phases of cognitive processing among different subjects and sessions. This approach was validated in the particular case of corticospinal excitability of the motor cortex during mental rotation. We identified the relative time-windows during which the excitability of M1 is effector-specifically modulated by different features of mental rotation. These time windows correspond to the 55%-85% of the subjective timing. In sum, (i) we introduce a new method to study the neurophysiology of motor cognition, and (ii) validating this method, we shed new light on the involvement of M1 in movement simulation

Differential neural encoding of sensorimotor and visual body representations.

Sensorimotor processing specifically impacts mental body representations. In particular, deteriorated somatosensory input (as after complete spinal cord injury) increases the relative weight of visual aspects of body parts' representations, leading to aberrancies in how images of body parts are mentally manipulated (e.g. mental rotation). This suggests that a sensorimotor or visual reference frame, respectively, can be relatively dominant in local (hands) versus global (full-body) bodily representations. On this basis, we hypothesized that the recruitment of a specific reference frame could be reflected in the activation of sensorimotor versus visual brain networks. To this aim, we directly compared the brain activity associated with mental rotation of hands versus full-bodies. Mental rotation of hands recruited more strongly the supplementary motor area, premotor cortex, and secondary somatosensory cortex. Conversely, mental rotation of full-bodies determined stronger activity in temporo-occipital regions, including the functionally-localized extrastriate body area. These results support that (1) sensorimotor and visual frames of reference are used to represent the body, (2) two distinct brain networks encode local or global bodily representations, and (3) the extrastriate body area is a multimodal region involved in body processing both at the perceptual and representational level