Maria Concetta Morrone
Professor of Physiology, Faculty of Medicine, University of Pisa
Editor-in-Chief of Multisensory Research
Multisensory Research is an interdisciplinary archival journal covering all aspects of multisensory processing including the control of action, cognition and attention. We publish research using any approach to increase our understanding of multisensory perceptual, behavioural, neural and computational mechanisms.
Current research and interests
- Development and plasticity of human visual system
- fMRI of human visual cortex
- Multi-sensory perception
- Infant vision
- Motion perception
- Neural oscillations
- Visual stability
- Perception of time
- Numerosity perception
2016 (back to top)
Tamietto, M. & Morrone, M. C. (2016). Visual Plasticity: Blindsight Bridges Anatomy and Function in the Visual System, Curr Biol, 2 (26), R70-73. PDF
Some people who are blind due to damage to their primary visual cortex, V1, can discriminate stimuli presented within their blind visual field. This residual function has been recently linked to a pathway that bypasses V1, and connects the thalamic lateral geniculate nucleus directly with the extrastriate cortical area MT.
Lunghi, C., Morrone, M. C., Secci, J. & Caputo, R. (2016). Binocular Rivalry Measured 2 Hours After Occlusion Therapy Predicts the Recovery Rate of the Amblyopic Eye in Anisometropic Children, Invest Ophthalmol Vis Sci, 4 (57), 1537-1546. PDF
PURPOSE. Recent studies on adults have shown that short-term monocular deprivation boosts the deprived eye signal in binocular rivalry, reflecting homeostatic plasticity. Here we investigate whether homeostatic plasticity is present also during occlusion therapy for moderate amblyopia. METHODS. Binocular rivalry and visual acuity (using Snellen charts for children) were measured in 10 children (mean age 6.2 ± 1 years) with moderate anisometropic amblyopia before the beginning of treatment and at four intervals during occlusion therapy (2 hours, 1, 2, and 5 months). Visual stimuli were orthogonal gratings presented dichoptically through ferromagnetic goggles and children reported verbally visual rivalrous perception. Bangerter filters were applied on the spectacle lens over the best eye for occlusion therapy. RESULTS. Two hours of occlusion therapy increased the nonamblyopic eye predominance over the amblyopic eye compared with pretreatment measurements, consistent with the results in adults. The boost of the nonamblyopic eye was still present after 1 month of treatment, steadily decreasing afterward to reach pretreatment levels after 2 months of continuous occlusion. Across subjects, the increase in nonamblyopic eye predominance observed after 2 hours of occlusion correlated (rho = -0.65, P = 0.04) with the visual acuity improvement of the amblyopic eye measured after 2 months of treatment. CONCLUSIONS. Homeostatic plasticity operates during occlusion therapy for moderate amblyopia and the increase in nonamblyopic eye dominance observed at the beginning of treatment correlates with the amblyopic eye recovery rate. These results suggest that binocular rivalry might be used to monitor visual cortical plasticity during occlusion therapy, although further investigations on larger clinical populations are needed to validate the predictive power of the technique.
Benedetto, A., Spinelli, D. & Morrone, M. C. (2016). Rhythmic modulation of visual contrast discrimination triggered by action, Proceedings of the Royal Society of London B: Biological Sciences, 1831 (283), PDF
Recent evidence suggests that ongoing brain oscillations may be instrumental in binding and integrating multisensory signals. In this experiment, we investigated the temporal dynamics of visual–motor integration processes. We show that action modulates sensitivity to visual contrast discrimination in a rhythmic fashion at frequencies of about 5 Hz (in the theta range), for up to 1 s after execution of action. To understand the origin of the oscillations, we measured oscillations in contrast sensitivity at different levels of luminance, which is known to affect the endogenous brain rhythms, boosting the power of alpha-frequencies. We found that the frequency of oscillation in sensitivity increased at low luminance, probably reflecting the shift in mean endogenous brain rhythm towards higher frequencies. Importantly, both at high and at low luminance, contrast discrimination showed a rhythmic motor-induced suppression effect, with the suppression occurring earlier at low luminance. We suggest that oscillations play a key role in sensory–motor integration, and that the motor-induced suppression may reflect the first manifestation of a rhythmic oscillation.
2015 (back to top)
Tinelli, F., Anobile, G., Gori, M., Aagten-Murphy, D., Bartoli, M., Burr, D. C., et al. Time, number and attention in very low birth weight children,Neuropsychologia, 2015 PDF
Abstract Premature birth has been associated with damage in many regions of the cerebral cortex, although there is a particularly strong susceptibility for damage within the parieto-occipital lobes (Volpe, 2009). As these areas have been shown to be critical for both visual attention and magnitudes perception (time, space, and number), it is important to investigate the impact of prematurity on both the magnitude and attentional systems, particularly for children without overt white matter injuries, where the lack of obvious injury may cause their difficulties to remain unnoticed. In this study, we investigated the ability to judge time intervals (visual, audio and audio-visual temporal bisection), discriminate between numerical quantities (numerosity comparison), map numbers onto space (numberline task) and to maintain visuo-spatial attention (multiple-object-tracking) in school-age preterm children (N29). The results show that various parietal functions may be more or less robust to prematurity-related difficulties, with strong impairments found on time estimation and attentional task, while numerical discrimination or mapping tasks remained relatively unimpaired. Thus while our study generally supports the hypothesis of a dorsal stream vulnerability in children born preterm relative to other cortical locations, it further suggests that particular cognitive processes, as highlighted by performance on different tasks, are far more susceptible than others.
Tomassini, A., Spinelli, D., Jacono, M., Sandini, G. & Morrone, M. C. (2015). Rhythmic oscillations of visual contrast sensitivity synchronized with action,J Neurosci, 18 (35), 7019-7029. PDF
It is well known that the motor and the sensory systems structure sensory data collection and cooperate to achieve an efficient integration and exchange of information. Increasing evidence suggests that both motor and sensory functions are regulated by rhythmic processes reflecting alternating states of neuronal excitability, and these may be involved in mediating sensory-motor interactions. Here we show an oscillatory fluctuation in early visual processing time locked with the execution of voluntary action, and, crucially, even for visual stimuli irrelevant to the motor task. Human participants were asked to perform a reaching movement toward a display and judge the orientation of a Gabor patch, near contrast threshold, briefly presented at random times before and during the reaching movement. When the data are temporally aligned to the onset of movement, visual contrast sensitivity oscillates with periodicity within the theta band. Importantly, the oscillations emerge during the motor planning stage, approximately 500 ms before movement onset. We suggest that brain oscillatory dynamics may mediate an automatic coupling between early motor planning and early visual processing, possibly instrumental in linking and closing up the visual-motor control loop.
Lunghi, C., Emir, U. E., Morrone, M. C. & Bridge, H. (2015). Short-Term Monocular Deprivation Alters GABA in the Adult Human Visual Cortex,Curr Biol, 11 (25), 1496-1501. PDF
Neuroplasticity is a fundamental property of the nervous system that is maximal early in life, within the critical period [1-3]. Resting GABAergic inhibition is necessary to trigger ocular dominance plasticity and to modulate the onset and offset of the critical period [4, 5]. GABAergic inhibition also plays a crucial role in neuroplasticity of adult animals: the balance between excitation and inhibition in the primary visual cortex (V1), measured at rest, modulates the susceptibility of ocular dominance to deprivation [6-10]. In adult humans, short-term monocular deprivation strongly modifies ocular balance, unexpectedly boosting the deprived eye, reflecting homeostatic plasticity [11, 12]. There is no direct evidence, however, to support resting GABAergic inhibition in homeostatic plasticity induced by visual deprivation. Here, we tested the hypothesis that GABAergic inhibition, measured at rest, is reduced by deprivation, as demonstrated by animal studies. GABA concentration in V1 of adult humans was measured using ultra-high-field 7T magnetic resonance spectroscopy before and after short-term monocular deprivation. After monocular deprivation, resting GABA concentration decreased in V1 but was unaltered in a control parietal area. Importantly, across participants, the decrease in GABA strongly correlated with the deprived eye perceptual boost measured by binocular rivalry. Furthermore, after deprivation, GABA concentration measured during monocular stimulation correlated with the deprived eye dominance. We suggest that reduction in resting GABAergic inhibition triggers homeostatic plasticity in adult human V1 after a brief period of abnormal visual experience. These results are potentially useful for developing new therapeutic strategies that could exploit the intrinsic residual plasticity of the adult human visual cortex.
Greco, V., Frijia, F., Mikellidou, K., Montanaro, D., Farini, A., D'Uva, M., et al. (2015). A low-cost and versatile system for projecting wide-field visual stimuli within fMRI scanners,Behav Res Methods, PDF
We have constructed and tested a custom-made magnetic-imaging-compatible visual projection system designed to project on a very wide visual field (~80 degrees ). A standard projector was modified with a coupling lens, projecting images into the termination of an image fiber. The other termination of the fiber was placed in the 3-T scanner room with a projection lens, which projected the images relayed by the fiber onto a screen over the head coil, viewed by a participant wearing magnifying goggles. To validate the system, wide-field stimuli were presented in order to identify retinotopic visual areas. The results showed that this low-cost and versatile optical system may be a valuable tool to map visual areas in the brain that process peripheral receptive fields.
Cicchini, G. M., Marino, C., Mascheretti, S., Perani, D. & Morrone, M. C. (2015). Strong Motion Deficits in Dyslexia Associated with DCDC2 Gene Alteration,J Neurosci, 21 (35), 8059-8064. PDF
Dyslexia is a specific impairment in reading that affects 1 in 10 people. Previous studies have failed to isolate a single cause of the disorder, but several candidate genes have been reported. We measured motion perception in two groups of dyslexics, with and without a deletion within the DCDC2 gene, a risk gene for dyslexia. We found impairment for motion particularly strong at high spatial frequencies in the population carrying the deletion. The data suggest that deficits in motion processing occur in a specific genotype, rather than the entire dyslexia population, contributing to the large variability in impairment of motion thresholds in dyslexia reported in the literature.
Biagi, L., Crespi, S. A., Tosetti, M. & Morrone, M. C. (2015). BOLD Response Selective to Flow-Motion in Very Young Infants, PLoS Biol, 9 (13), e1002260. PDF
In adults, motion perception is mediated by an extensive network of occipital, parietal, temporal, and insular cortical areas. Little is known about the neural substrate of visual motion in infants, although behavioural studies suggest that motion perception is rudimentary at birth and matures steadily over the first few years. Here, by measuring Blood Oxygenated Level Dependent (BOLD) responses to flow versus random-motion stimuli, we demonstrate that the major cortical areas serving motion processing in adults are operative by 7 wk of age. Resting-state correlations demonstrate adult-like functional connectivity between the motion-selective associative areas, but not between primary cortex and temporo-occipital and posterior-insular cortices. Taken together, the results suggest that the development of motion perception may be limited by slow maturation of the subcortical input and of the cortico-cortical connections. In addition they support the existence of independent input to primary (V1) and temporo-occipital (V5/MT+) cortices very early in life.
Lunghi, C., Berchicci, M., Morrone, M. C. & Di Russo, F. (2015). Short-term monocular deprivation alters early components of visual evoked potentials, J Physiol, 19 (593), 4361-4372. PDF
Very little is known about plasticity in the adult visual cortex. In recent years psychophysical studies have shown that short-term monocular deprivation alters visual perception in adult humans. Specifically, after 150 min of monocular deprivation the deprived eye strongly dominates the dynamics of binocular rivalry, reflecting homeostatic plasticity. Here we investigate the neural mechanisms underlying this form of short-term visual cortical plasticity by measuring visual evoked potentials (VEPs) on the scalp of adult humans during monocular stimulation before and after 150 min of monocular deprivation. We found that monocular deprivation had opposite effects on the amplitude of the earliest component of the VEP (C1) for the deprived and non-deprived eye stimulation. C1 amplitude increased (+66%) for the deprived eye, while it decreased (-29%) for the non-deprived eye. Source localization analysis confirmed that the C1 originates in the primary visual cortex. We further report that following monocular deprivation, the amplitude of the peak of the evoked alpha spectrum increased on average by 23% for the deprived eye and decreased on average by 10% for the non-deprived eye, indicating a change in cortical excitability. These results indicate that a brief period of monocular deprivation alters interocular balance in the primary visual cortex of adult humans by both boosting the activity of the deprived eye and reducing the activity of the non-deprived eye. This indicates a high level of residual homeostatic plasticity in the adult human primary visual cortex, probably mediated by a change in cortical excitability.
Zimmermann, E., Morrone, M. C. & Burr, D. (2015). Visual mislocalization during saccade sequences,Exp Brain Res, 2 (233), 577-585. PDF
Visual objects briefly presented around the time of saccadic eye movements are perceived compressed towards the saccade target. Here, we investigated perisaccadic mislocalization with a double-step saccade paradigm, measuring localization of small probe dots briefly flashed at various times around the sequence of the two saccades. At onset of the first saccade, probe dots were mislocalized towards the first and, to a lesser extent, also towards the second saccade target. However, there was very little mislocalization at the onset of the second saccade. When we increased the presentation duration of the saccade targets prior to onset of the saccade sequence, perisaccadic mislocalization did occur at the onset of the second saccade.
2014 (back to top)
Lunghi, C., Morrone, M. C. & Alais, D. (2014). Auditory and tactile signals combine to influence vision during binocular rivalry,J Neurosci, 3 (34), 784-792. PDF
Resolution of perceptual ambiguity is one function of cross-modal interactions. Here we investigate whether auditory and tactile stimuli can influence binocular rivalry generated by interocular temporal conflict in human subjects. Using dichoptic visual stimuli modulating at different temporal frequencies, we added modulating sounds or vibrations congruent with one or the other visual temporal frequency. Auditory and tactile stimulation both interacted with binocular rivalry by promoting dominance of the congruent visual stimulus. This effect depended on the cross-modal modulation strength and was absent when modulation depth declined to 33%. However, when auditory and tactile stimuli that were too weak on their own to bias binocular rivalry were combined, their influence over vision was very strong, suggesting the auditory and tactile temporal signals combined to influence vision. Similarly, interleaving discrete pulses of auditory and tactile stimuli also promoted dominance of the visual stimulus congruent with the supramodal frequency. When auditory and tactile stimuli were presented at maximum strength, but in antiphase, they had no influence over vision for low temporal frequencies, a null effect again suggesting audio-tactile combination. We also found that the cross-modal interaction was frequency-sensitive at low temporal frequencies, when information about temporal phase alignment can be perceptually tracked. These results show that auditory and tactile temporal processing is functionally linked, suggesting a common neural substrate for the two sensory modalities and that at low temporal frequencies visual activity can be synchronized by a congruent cross-modal signal in a frequency-selective way, suggesting the existence of a supramodal temporal binding mechanism.
Morrone, M. C. (2014). Interaction between Eye Movements and Vision: Perception during Saccades. In J. S. W. L. M. Chalupa (Ed.), The New Visual Neuroscience (2nd ed., pp. 947 -962): MIT Press. PDF
Tomassini, A., Gori, M., Baud-Bovy, G., Sandini, G. & Morrone, M. C. (2014). Motor commands induce time compression for tactile stimuli,J Neurosci, 27 (34), 9164-9172. PDF
Saccades cause compression of visual space around the saccadic target, and also a compression of time, both phenomena thought to be related to the problem of maintaining saccadic stability (Morrone et al., 2005; Burr and Morrone, 2011). Interestingly, similar phenomena occur at the time of hand movements, when tactile stimuli are systematically mislocalized in the direction of the movement (Dassonville, 1995; Watanabe et al., 2009). In this study, we measured whether hand movements also cause an alteration of the perceived timing of tactile signals. Human participants compared the temporal separation between two pairs of tactile taps while moving their right hand in response to an auditory cue. The first pair of tactile taps was presented at variable times with respect to movement with a fixed onset asynchrony of 150 ms. Two seconds after test presentation, when the hand was stationary, the second pair of taps was delivered with a variable temporal separation. Tactile stimuli could be delivered to either the right moving or left stationary hand. When the tactile stimuli were presented to the motor effector just before and during movement, their perceived temporal separation was reduced. The time compression was effector-specific, as perceived time was veridical for the left stationary hand. The results indicate that time intervals are compressed around the time of hand movements. As for vision, the mislocalizations of time and space for touch stimuli may be consequences of a mechanism attempting to achieve perceptual stability during tactile exploration of objects, suggesting common strategies within different sensorimotor systems.
Zimmermann, E., Morrone, M. C. & Burr, D. C. (2014). Buildup of spatial information over time and across eye-movements,Behavioural brain research, PDF
To interact rapidly and effectively with our environment, our brain needs access to a neural represen-tation of the spatial layout of the external world. However, the construction of such a map poses majorchallenges, as the images on our retinae depend on where the eyes are looking, and shift each time wemove our eyes, head and body to explore the world. Research from many laboratories including ourown suggests that the visual system does compute spatial maps that are anchored to real-world coordi-nates. However, the construction of these maps takes time (up to 500 ms) and also attentional resources.We discuss research investigating how retinotopic reference frames are transformed into spatiotopicreference-frames, and how this transformation takes time to complete. These results have implicationsfor theories about visual space coordinates and particularly for the current debate about the existence ofspatiotopic representations.
Zimmermann, E., Morrone, M. C. & Burr, D. C. (2014). The visual component to saccadic compression,J Vis, 12 (14), PDF
Visual objects presented around the time of saccadic eye movements are strongly mislocalized towards the saccadic target, a phenomenon known as "saccadic compression." Here we show that perisaccadic compression is modulated by the presence of a visual saccadic target. When subjects saccaded to the center of the screen with no visible target, perisaccadic localization was more veridical than when tested with a target. Presenting a saccadic target sometime before saccade initiation was sufficient to induce mislocalization. When we systematically varied the onset of the saccade target, we found that it had to be presented around 100 ms before saccade execution to cause strong mislocalization: saccadic targets presented after this time caused progressively less mislocalization. When subjects made a saccade to screen center with a reference object placed at various positions, mislocalization was focused towards the position of the reference object. The results suggest that saccadic compression is a signature of a mechanism attempting to match objects seen before the saccade with those seen after.
2013 (back to top)
Cicchini, G. M., Binda, P., Burr, D. C. & Morrone, M. C. (2013). Transient spatiotopic integration across saccadic eye movements mediates visual stability,J Neurophysiol, 4 (109), 1117-1125. PDF
Eye movements pose major problems to the visual system, because each new saccade changes the mapping of external objects on the retina. It is known that stimuli briefly presented around the time of saccades are systematically mislocalized, whereas continuously visible objects are perceived as spatially stable even when they undergo large transsaccadic displacements. In this study we investigated the relationship between these two phenomena and measured how human subjects perceive the position of pairs of bars briefly displayed around the time of large horizontal saccades. We show that they interact strongly, with the perisaccadic bar being drawn toward the other, dramatically altering the pattern of perisaccadic mislocalization. The interaction field extends over a wide range (200 ms and 20 degrees ) and is oriented along the retinotopic trajectory of the saccade-induced motion, suggesting a mechanism that integrates pre- and postsaccadic stimuli at different retinal locations but similar external positions. We show how transient changes in spatial integration mechanisms, which are consistent with the present psychophysical results and with the properties of "remapping cells" reported in the literature, can create transient craniotopy by merging the distinct retinal images of the pre- and postsaccadic fixations to signal a single stable object.
Knoll, J., Morrone, M. C. & Bremmer, F. (2013). Spatio-temporal topography of saccadic overestimation of time,Vision Res, (83C), 56-65. PDF
Rapid eye movements (saccades) induce visual misperceptions. A number of studies in recent years have investigated the spatio-temporal profiles of effects like saccadic suppression or perisaccadic mislocalization and revealed substantial functional similarities. Saccade induced chronostasis describes the subjective overestimation of stimulus duration when the stimulus onset falls within a saccade. In this study we aimed to functionally characterize saccade induced chronostasis in greater detail. Specifically we tested if chronostasis is influenced by or functionally related to saccadic suppression. In a first set of experiments, we measured the perceived duration of visual stimuli presented at different spatial positions as a function of presentation time relative to the saccade. We further compared perceived duration during saccades for isoluminant and luminant stimuli. Finally, we investigated whether or not saccade induced chronostasis is dependent on the execution of a saccade itself. We show that chronostasis occurs across the visual field with a clear spatio-temporal tuning. Furthermore, we report chronostasis during simulated saccades, indicating that spurious retinal motion induced by the saccade is a prime origin of the phenomenon.
Zimmermann, E., Morrone, M. C., Fink, G. R. & Burr, D. (2013). Spatiotopic neural representations develop slowly across saccades,Curr Biol, 5 (23), R193-194. PDF
One of the long-standing unsolved mysteries of visual neuroscience is how the world remains apparently stable in the face of continuous movements of eyes, head and body. Many factors seem to contribute to this stability, including rapid updating mechanisms that temporarily remap the visual input to compensate for the impending saccade . However, there is also a growing body of evidence pointing to more long-lasting spatiotopic neural representations, which remain solid in external rather than retinal coordinates [2-6]. In this study, we show that these spatiotopic representations take hundreds of milliseconds to build up robustly.
Lunghi, C., Burr, D. C. & Morrone, M. C. (2013). Long-term effects of monocular deprivation revealed with binocular rivalry gratings modulated in luminance and in color,J Vis, 6 (13), PDF
During development, within a specific temporal window called the critical period, the mammalian visual cortex is highly plastic and literally shaped by visual experience; to what extent this extraordinary plasticity is retained in the adult brain is still a debated issue. We tested the residual plastic potential of the adult visual cortex for both achromatic and chromatic vision by measuring binocular rivalry in adult humans following 150 minutes of monocular patching. Paradoxically, monocular deprivation resulted in lengthening of the mean phase duration of both luminance-modulated and equiluminant stimuli for the deprived eye and complementary shortening of nondeprived phase durations, suggesting an initial homeostatic compensation for the lack of information following monocular deprivation. When equiluminant gratings were tested, the effect was measurable for at least 180 minutes after reexposure to binocular vision, compared with 90 minutes for achromatic gratings. Our results suggest that chromatic vision shows a high degree of plasticity, retaining the effect for a duration (180 minutes) longer than that of the deprivation period (150 minutes) and twice as long as that found with achromatic gratings. The results are in line with evidence showing a higher vulnerability of the P pathway to the effects of visual deprivation during development and a slower development of chromatic vision in humans.
Orchard-Mills, E., Leung, J., Burr, D., Morrone, M. C., Wufong, E., Carlile, S., et al. (2013). A mechanism for detecting coincidence of auditory and visual spatial signals,Multisens Res, 4 (26), 333-345. PDF
Information about the world is captured by our separate senses, and must be integrated to yield a unified representation. This raises the issue of which signals should be integrated and which should remain separate, as inappropriate integration will lead to misrepresentation and distortions. One strong cue suggesting that separate signals arise from a single source is coincidence, in space and in time. We measured increment thresholds for discriminating spatial intervals defined by pairs of simultaneously presented targets, one flash and one auditory sound, for various separations. We report a 'dipper function', in which thresholds follow a 'U-shaped' curve, with thresholds initially decreasing with spatial interval, and then increasing for larger separations. The presence of a dip in the audiovisual increment-discrimination function is evidence that the auditory and visual signals both input to a common mechanism encoding spatial separation, and a simple filter model with a sigmoidal transduction function simulated the results well. The function of an audiovisual spatial filter may be to detect coincidence, a fundamental cue guiding whether to integrate or segregate.
Castaldi, E., Frijia, F., Montanaro, D., Tosetti, M. & Morrone, M. C. (2013). BOLD human responses to chromatic spatial features,Eur J Neurosci, 2 (38), 2290-2299. PDF
Animal physiological and human psychophysical studies suggest that an early step in visual processing involves the detection and identification of features such as lines and edges, by neural mechanisms with even- and odd-symmetric receptive fields. Functional imaging studies also demonstrate mechanisms with even- and odd-receptive fields in early visual areas, in response to luminance-modulated stimuli. In this study we measured fMRI BOLD responses to 2-D stimuli composed of only even or only odd symmetric features, and to an amplitude-matched random noise control, modulated in red-green equiluminant colour contrast. All these stimuli had identical power but different phase spectra, either highly congruent (even or odd symmetry stimuli) or random (noise). At equiluminance, V1 BOLD activity showed no preference between congruent- and random-phase stimuli, as well as no preference between even and odd symmetric stimuli. Areas higher in the visual hierarchy, both along the dorsal pathway (caudal part of the intraparietal sulcus, dorsal LO and V3A) and the ventral pathway (V4), responded preferentially to odd symmetry over even symmetry stimuli, and to congruent over random phase stimuli. Interestingly, V1 showed an equal increase in BOLD activity at each alternation between stimuli of different symmetry, suggesting the existence of specialised mechanisms for the detection of edges and lines such as even- and odd-chromatic receptive fields. Overall the results indicate a high selectivity of colour-selective neurons to spatial phase along both the dorsal and the ventral pathways in humans.
Burr, D., Rocca, E. D. & Morrone, M. C. (2013). Contextual effects in interval-duration judgements in vision, audition and touch,Exp Brain Res, PDF
We examined the effect of temporal context on discrimination of intervals marked by auditory, visual and tactile stimuli. Subjects were asked to compare the duration of the interval immediately preceded by an irrelevant "distractor" stimulus with an interval with no distractor. For short interval durations, the presence of the distractor affected greatly the apparent duration of the test stimulus: short distractors caused the test interval to appear shorter and vice versa. For very short reference durations (< o =100 ms), the contextual effects were large, changing perceived duration by up to a factor of two. The effect of distractors reduced steadily for longer reference durations, to zero effect for durations greater than 500 ms. We found similar results for intervals defined by visual flashes, auditory tones and brief finger vibrations, all falling to zero effect at 500 ms. Under appropriate conditions, there were strong cross-modal interactions, particularly from audition to vision. We also measured the Weber fractions for duration discrimination and showed that under the conditions of this experiment, Weber fractions decreased steadily with duration, following a square-root law, similarly for all three modalities. The magnitude of the effect of the distractors on apparent duration correlated well with Weber fraction, showing that when duration discrimination was relatively more precise, the context dependency was less. The results were well fit by a simple Bayesian model combining noisy estimates of duration with the action of a resonance-like mechanism that tended to regularize the sound sequence intervals.
Burr, D., Della Rocca, E. & Morrone, M. C. (2013). Erratum to: Contextual effects in interval-duration judgements in vision, audition and touch,Exp Brain Res, PDF
Lunghi, C. & Morrone, M. C. (2013). Early interaction between vision and touch during binocular rivalry,Multisens Res, 3 (26), 291-306. PDF
Multisensory integration is known to occur at high neural levels, but there is also growing evidence that cross-modal signals can be integrated at the first stages of sensory processing. We investigated whether touch specifically affected vision during binocular rivalry, a particular type of visual bistability that engages neural competition in early visual cortices. We found that tactile signals interact with visual signals outside of awareness, when the visual stimulus congruent with the tactile one is perceptually suppressed during binocular rivalry and when the interaction is strictly tuned for matched visuo-tactile spatial frequencies. We also found that voluntary action does not play a leading role in mediating the effect, since the interaction was observed also when tactile stimulation was passively delivered to the finger. However, simultaneous presentation of visual and tactile stimuli is necessary to elicit the interaction, and an asynchronous priming touch stimulus is not affecting the onset of rivalry. These results point to a very early cross-modal interaction site, probably V1. By showing that spatial proximity between visual and tactile stimuli is a necessary condition for the interaction, we also suggest that the two sensory spatial maps are aligned according to retinotopic coordinates, corroborating the hypothesis of a very early interaction between visual and tactile signals during binocular rivalry.
Zimmerman, E., Morrone,M.C. & Burr, DC.. (2013).Spatial position information accumulates steadily over time, J Neurosci 33(47):18396-18401. PDF
One of the more enduring mysteries of neuroscience is how the visual system constructs robust maps of the world that remain stable in the face of frequent eye-movements. Here we show that encoding the position of objects in external space is a relatively slow process, building up over hundreds of milliseconds. We display targets to which human subjects saccade after a variable preview duration. As they saccade, the target is displaced leftwards or rightwards, and subjects report the displacement direction. When subjects saccade to targets without delay, sensitivity is poor: but if the target is viewed for 300-500 ms before saccading, sensitivity is similar to that during fixation with a strong visual mask to dampen transients. These results suggest that the poor displacement thresholds usually observed in the “saccadic suppression of displacement” paradigm do not reflect the action of special mechanisms conferring saccadic stability, but the fact that the target has had insufficient time to be encoded in memory. Under more natural conditions, trans-saccadic displacement detection is as good as in fixation, when the displacement transients are masked.
Pooresmaeili, A., Cicchini, G.M., Morrone, M. C. & Burr, D.C. (2013). Spatiotemporal filtering and motion illusions, Journal of Vision, (13)10-21. PDF
Our group has long championed the idea that perceptual processing of information can be anchored in a dynamic coordinate system that need not correspond to the instantaneous retinal representation that need not correspond to the istantaneus retinal representation...
Pooresmaeili, A., Arrighi, R., Biagi, L. & Morrone, M. C. (2013). Blood oxygen level-dependent activation of the primary visual cortex predicts size adaptation illusion,J Neurosci, 40 (33), 15999-16008. PDF
In natural scenes, objects rarely occur in isolation but appear within a spatiotemporal context. Here, we show that the perceived size of a stimulus is significantly affected by the context of the scene: brief previous presentation of larger or smaller adapting stimuli at the same region of space changes the perceived size of a test stimulus, with larger adapting stimuli causing the test to appear smaller than veridical and vice versa. In a human fMRI study, we measured the blood oxygen level-dependent activation (BOLD) responses of the primary visual cortex (V1) to the contours of large-diameter stimuli and found that activation closely matched the perceptual rather than the retinal stimulus size: the activated area of V1 increased or decreased, depending on the size of the preceding stimulus. A model based on local inhibitory V1 mechanisms simulated the inward or outward shifts of the stimulus contours and hence the perceptual effects. Our findings suggest that area V1 is actively involved in reshaping our perception to match the short-term statistics of the visual scene.
Sani, I., Santandrea, E., Golzar, A., Morrone, M. C. & Chelazzi, L. (2013). Selective tuning for contrast in macaque area V4,J Neurosci, 47 (33), 18583-18596. PDF
Visually responsive neurons typically exhibit a monotonic-saturating increase of firing with luminance contrast of the stimulus and are able to adapt to the current spatiotemporal context by shifting their selectivity, therefore being perfectly suited for optimal contrast encoding and discrimination. Here we report the first evidence of the existence of neurons showing selective tuning for contrast in area V4d of the behaving macaque (Macaca mulatta), i.e., narrow bandpass filter neurons with peak activity encompassing the whole range of visible contrasts and pronounced attenuation at contrasts higher than the peak. Crucially, we found that contrast tuning emerges after a considerable delay from stimulus onset, likely reflecting the contribution of inhibitory mechanisms. Selective tuning for luminance contrast might support multiple functions, including contrast identification and the attentive selection of low contrast stimuli.
2012 (back to top)
Pooresmaeili, A., Cicchini, G. M., Morrone, M. C. & Burr, D. (2012). "Non-retinotopic processing" in Ternus motion displays modeled by spatiotemporal filters,J Vis, 1 (12), PDF
Recently, M. Boi, H. Ogmen, J. Krummenacher, T. U. Otto, & M. H. Herzog (2009) reported a fascinating visual effect, where the direction of apparent motion was disambiguated by cues along the path of apparent motion, the Ternus-Pikler group motion, even though no actual movement occurs in this stimulus. They referred to their study as a "litmus test" to distinguish "non-retinotopic" (motion-based) from "retinotopic" (retina-based) image processing. We adapted the test to one with simple grating stimuli that could be more readily modeled and replicated their psychophysical results quantitatively with this stimulus. We then modeled our experiments in 3D (x, y, t) Fourier space and demonstrated that the observed perceptual effects are readily accounted for by integration of information within a detector that is oriented in space and time, in a similar way to previous explanations of other motion illusions. This demonstration brings the study of Boi et al. into the more general context of perception of moving objects.
Zimmermann, E., Morrone, M. C. & Burr, D. (2012). Visual motion distorts visual and motor space, J Vis, 2 (12), PDF
Mapping of number onto space is fundamental to mathematics and measurement. Previous research suggests that while typical adults with mathematical schooling map numbers veridically onto a linear scale, pre-school children and adults without formal mathematics training, as well as individuals with dyscalculia, show strong compressive, logarithmic-like non-linearities when mapping both symbolic and non-symbolic numbers onto the numberline. Here we show that the use of the linear scale is dependent on attentional resources. We asked typical adults to position clouds of dots on a numberline of various lengths. In agreement with previous research, they did so veridically under normal conditions, but when asked to perform a concurrent attentionally-demanding conjunction task, the mapping followed a compressive, non-linear function. We model the non-linearity both by the commonly assumed logarithmic transform, and also with a Bayesian model of central tendency. These results suggest that veridical representation numerosity requires attentional mechanisms.
Binda, P., Morrone, M. C. & Bremmer, F. (2012). Saccadic compression of symbolic numerical magnitude,PLoS One, 11 (7), e49587. PDF
Stimuli flashed briefly around the time of saccadic eye movements are subject to complex distortions: compression of space and time; underestimate of numerosity. Here we show that saccadic distortions extend to abstract quantities, affecting the representation of symbolic numerical magnitude. Subjects consistently underestimated the results of rapidly computed mental additions and subtractions, when the operands were briefly displayed before a saccade. However, the recognition of the number symbols was unimpaired. These results are consistent with the hypothesis of a common, abstract metric encoding magnitude along multiple dimensions. They suggest that a surprising link exists between the preparation of action and the representation of abstract quantities.
Panichi, M., Burr, D., Morrone, M. C. & Baldassi, S. (2012). Spatiotemporal dynamics of perisaccadic remapping in humans revealed by classification images,J Vis, 4 (12), 11. PDF
We actively scan our environment with fast ballistic movements called saccades, which create large and rapid displacements of the image on the retina. At the time of saccades, vision becomes transiently distorted in many ways: Briefly flashed stimuli are displaced in space and in time, and spatial and temporal intervals appear compressed. Here we apply the psychophysical technique of classification images to study the spatiotemporal dynamics of visual mechanisms during saccades. We show that saccades cause gross distortions of the classification images. Before the onset of saccadic eye movements, the positive lobes of the images become enlarged in both space and in time and also shifted in a systematic manner toward the pre-saccadic fixation (in space) and anticipated in time by about 50 ms. The transient reorganization creates a spatiotemporal organization oriented in the direction of saccadic-induced motion at the time of saccades, providing a potential mechanism for integrating stimuli across saccades, facilitating stable and continuous vision in the face of constant eye movements.
Tinelli, T., Cicchini, G.M., Arrighi, R., Tosetti, M., Cioni, G., Morrone M. C. (2012). Blindsight in children with congenital and acquired cerebral lesions, Cortex (published online 10 August 2012) PDF
It has been shown that unconscious visual function can survive lesions to optical radiations and/or primary visual cortex (V1), a phenomenon termed “blindsight”. Studies on animal models (cat and monkey) show that the age when the lesion occurs determines the extent of residual visual capacities. Much less is known about the functional and underlying neuronal repercussions of early cortical damage in humans. We measured sensitivity to several visual tasks in four children with congenital unilateral brain lesions that severely affected optic radiations, and in another group of three children with similar lesions, acquired in childhood. In two of the congenital patients, we measured blood oxygenation level dependent (BOLD) activity in response to stimulation of each visual field quadrants. Results show clear evidence of residual unconscious processing of position, orientation and motion of visual stimuli displayed in the scotoma of congenitally lesioned children, but not in the children with acquired lesions. The calcarine cortical BOLD responses were abnormally elicited by stimulation of the ipsilateral visual field and in the scotoma region, demonstrating a profound neuronal reorganization. In conclusion, our data suggest that congenital lesions can trigger massive reorganization of the visual system to alleviate functional effects of early brain insults.
Tomassini, A., Gori, M., Burr, D., Sandini, G. & Morrone, M. C. (2012). Active movement restores veridical event-timing after tactile adaptation,J Neurophysiol, 8 (108), 2092-2100. PDF
Growing evidence suggests that time in the subsecond range is tightly linked to sensory processing. Event-time can be distorted by sensory adaptation, and many temporal illusions can accompany action execution. In this study, we show that adaptation to tactile motion causes a strong contraction of the apparent duration of tactile stimuli. However, when subjects make a voluntary motor act before judging the duration, it annuls the adaptation-induced temporal distortion, reestablishing veridical event-time. The movement needs to be performed actively by the subject: passive movement of similar magnitude and dynamics has no effect on adaptation, showing that it is the motor commands themselves, rather than reafferent signals from body movement, which reset the adaptation for tactile duration. No other concomitant perceptual changes were reported (such as apparent speed or enhanced temporal discrimination), ruling out a generalized effect of body movement on somatosensory processing. We suggest that active movement resets timing mechanisms in preparation for the new scenario that the movement will cause, eliminating inappropriate biases in perceived time. Our brain seems to utilize the intention-to-move signals to retune its perceptual machinery appropriately, to prepare to extract new temporal information.
Morrone, M. C. M. (2012). Plasticità ed adattabilità della visione,Giornale Italiano di Psicologia, (3), 517-522. PDF
Burr, D. C. & Morrone, M. C. (2012). Constructing stable spatial maps of the world,Perception, 11 (41), 1355-1372. PDF
To interact rapidly and effectively with our environment, our brain needs access to a neural representation—or map—of the spatial layout of the external world. However, the construction of such a map poses major challenges to the visual system, given that the images on our retinae depend on where the eyes are looking, and shift each time we move our eyes, head, and body to explore the world. Much research has been devoted to how the stability is achieved, with the debate often polarized between the utility of spatiotopic maps (that remain solid in external coordinates), as opposed to transiently updated retinotopic maps. Our research suggests that the visual system uses both strategies to maintain stability. f MRI, motion-adaptation, and saccade-adaptation studies demonstrate and characterize spatiotopic neural maps within the dorsal visual stream that remain solid in external rather than retinal coordinates. However, the construction of these maps takes time (up to 500 ms) and attentional resources. To solve the immediate problems created by individual saccades, we postulate the existence of a separate system to bridge each saccade with neural units that are ‘transiently craniotopic’. These units prepare for the effects of saccades with a shift of their receptive fields before the saccade starts, then relaxing back into their standard position during the saccade, compensating for its action. Psychophysical studies investigating localization of stimuli flashed briefly around the time of saccades provide strong support for these neural mechanisms, and show quantitatively how they integrate information across saccades. This transient system cooperates with the spatiotopic mechanism to provide a useful map to guide interactions with our environment: one rapid and transitory, bringing into play the high-resolution visual areas; the other slow, long-lasting, and low-resolution, useful for interacting with the world.
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2011 (back to top)
Burr, D. C., Cicchini, G. M., Arrighi, R. & Morrone, M. C. (2011). Spatiotopic selectivity of adaptation-based compression of event duration, J Vis, 2 (11), 21; author reply 21a. PDF
A. Bruno, I. Ayhan, and A. Johnston (2010) have recently challenged our report of spatiotopic selectivity for adaptation of event time (D. Burr, A. Tozzi, & M. C. Morrone, 2007) and also our claim that retinotopic adaptation of event time depends on perceived speed. To assist the reader judge this issue, we present here a mass of data accumulated in our laboratories over the last few years, all confirming our original conclusions. We also point out that where Bruno et al. made experimental measurements (rather than relying on theoretical reasoning), they too find clearly significant spatiotopically tuned adaptation-based compression of event time but of lower magnitude to ours. We speculate on the reasons for the differences in magnitude.
Lunghi C, Burr DC, Morrone C. (2011). Brief periods of monocular deprivation disrupt ocular balance in human adult visual cortex, Curr Biol. 2011 Jul 26;21(14):R538-9. PDF
Neuroplasticity is a fundamental property of the developing mammalian visual system, with residual potential in adult human cortex . A short period of abnormal visual experience (such as occlusion of one eye) before closure of the critical period has dramatic and permanent neural consequences, reshaping visual cortical organization in favour of the non-deprived eye [2,3]. We used binocular rivalry  - a sensitive probe of neural competition - to demonstrate that adult human visual cortex retains a surprisingly high degree of neural plasticity, with important perceptual consequences. We report that 150 minutes of monocular deprivation strongly affects the dynamics of binocular rivalry, unexpectedly causing the deprived eye to prevail in conscious perception twice as much as the non-deprived eye, with significant effects for up to 90 minutes. Apparent contrast of stimuli presented to the deprived eye was also increased, suggesting that the deprivation acts by up-regulation of cortical gain-control mechanisms of the deprived eye. The results suggest that adult visual cortex retains a good deal of plasticity that could be important in reaction to sensory loss.
Burr, D. C. & Morrone, M. C. (2011). Spatiotopic coding and remapping in humans,Philos Trans R Soc Lond B Biol Sci, 1564 (366), 504-515. PDF
How our perceptual experience of the world remains stable and continuous in the face of continuous rapid eye movements still remains a mystery. This review discusses some recent progress towards understanding the neural and psychophysical processes that accompany these eye movements. We firstly report recent evidence from imaging studies in humans showing that many brain regions are tuned in spatiotopic coordinates, but only for items that are actively attended. We then describe a series of experiments measuring the spatial and temporal phenomena that occur around the time of saccades, and discuss how these could be related to visual stability. Finally, we introduce the concept of the spatio-temporal receptive field to describe the local spatiotopicity exhibited by many neurons when the eyes move.
Crespi, S., Biagi, L., d'Avossa, G., Burr, D. C., Tosetti, M. & Morrone, M. C. (2011). Spatiotopic Coding of BOLD Signal in Human Visual Cortex Depends on Spatial Attention,PLoS One, 7 (6), e21661. PDF
The neural substrate of the phenomenological experience of a stable visual world remains obscure. One possible mechanism would be to construct spatiotopic neural maps where the response is selective to the position of the stimulus in external space, rather than to retinal eccentricities, but evidence for these maps has been inconsistent. Here we show, with fMRI, that when human subjects perform concomitantly a demanding attentive task on stimuli displayed at the fovea, BOLD responses evoked by moving stimuli irrelevant to the task were mostly tuned in retinotopic coordinates. However, under more unconstrained conditions, where subjects could attend easily to the motion stimuli, BOLD responses were tuned not in retinal but in external coordinates (spatiotopic selectivity) in many visual areas, including MT, MST, LO and V6, agreeing with our previous fMRI study. These results indicate that spatial attention may play an important role in mediating spatiotopic selectivity.
Binda, P., Morrone, M. C., Ross, J. & Burr, D. C. (2011). Underestimation of perceived number at the time of saccades,Vision Res, 1 (51), 34-42. PDF
Saccadic eye movements produce transient distortions in both space and time. Mounting evidence suggests that space and time perception are linked, and associated with the perception of another important perceptual attribute, numerosity. Here we investigate the effect of saccades on the perceived numerosity of briefly presented arrays of visual elements. We report a systematic underestimation of numerosity for stimuli flashed just before or during saccades, of about 35% of the reference numerosity. The bias is observed only for relatively large arrays of visual elements, in line with the notion that a distinct perceptual mechanism is involved with enumeration of small numerosities in the 'subitizing' range. This study provides further evidence for the notion that space, time and number share common neural representations, all affected by saccades.
Knoll, J., Binda, P., Morrone, M. C. & Bremmer, F. (2011). Spatiotemporal profile of peri-saccadic contrast sensitivity,J Vis, 14 (11), PDF
Sensitivity to luminance contrast is reduced just before and during saccades (saccadic suppression), whereas sensitivity to color contrast is unimpaired peri-saccadically and enhanced post-saccadically. The exact spatiotemporal map of these perceptual effects is as yet unknown. Here, we measured detection thresholds for briefly flashed Gaussian blobs modulated in either luminance or chromatic contrast, displayed at a range of eccentricities. Sensitivity to luminance contrast was reduced peri-saccadically by a scaling factor, which was almost constant across retinal space. Saccadic suppression followed a similar time course across all tested eccentricities and was maximal shortly after the saccade onset. Sensitivity to chromatic contrast was enhanced post-saccadically at all tested locations. The enhancement was not specifically linked to the execution of saccades, as it was also observed following a displacement of retinal images comparable to that caused by a saccade. We conclude that luminance and chromatic contrast sensitivities are subject to distinct modulations at the time of saccades, resulting from independent neural processes.
Tomassini A, Gori M, Burr D, Sandini G and Morrone MC (2011) Perceived duration of visual and tactile stimuli depends on perceived speed. Front. Integr. Neurosci. 5:51 PDF
It is known that the perceived duration of visual stimuli is strongly influenced by speed: faster moving stimuli appear to last longer. To test whether this is a general property of sensory systems we asked participants to reproduce the duration of visual and tactile gratings, and visuo-tactile gratings moving at a variable speed (3.5–15 cm/s) for three different durations (400, 600, and 800 ms). For both modalities, the apparent duration of the stimulus increased strongly with stimulus speed, more so for tactile than for visual stimuli. In addition, visual stimuli were perceived to last approximately 200 ms longer than tactile stimuli. The apparent duration of visuo-tactile stimuli lay between the unimodal estimates, as the Bayesian account predicts, but the bimodal precision of the reproduction did not show the theoretical improvement. A cross-modal speed-matching task revealed that visual stimuli were perceived to move faster than tactile stimuli. To test whether the large difference in the perceived duration of visual and tactile stimuli resulted from the difference in their perceived speed, we repeated the time reproduction task with visual and tactile stimuli matched in apparent speed. This reduced, but did not completely eliminate the difference in apparent duration. These results show that for both vision and touch, perceived duration depends on speed, pointing to common strategies of time perception.
Zimmerman, E., Burr D.C., and Morrone, M.C. (2011) Spatiotopic Visual Maps Revealed by Saccadic Adaptation in Humans, Curr Biol. 2011 Aug 23;21(16):1380-4 PDF
Saccadic adaptation is a powerful experimental paradigm to probe the mechanisms of eye movement control and spatial vision, in which saccadic amplitudes change in response to false visual feedback. The adaptation occurs primarily in the motor system, but there is also evidence for visual adaptation, depending on the size and the permanence of the postsaccadic error. Here we confirm that adaptation has a strong visual component and show that the visual component of the adaptation is spatially selective in external, not retinal coordinates. Subjects performed a memory-guided, double-saccade, outward-adaptation task designed to maximize visual adaptation and to dissociate the visual and motor corrections. When the memorized saccadic target was in the same position (in external space) as that used in the adaptation training, saccade targeting was strongly influenced by adaptation (even if not matched in retinal or cranial position), but when in the same retinal or cranial but different external spatial position, targeting was unaffected by adaptation, demonstrating unequivocal spatiotopic selectivity. These results point to the existence of a spatiotopic neural representation for eye movement control that adapts in response to saccade error signals.
2010 (back to top)
Morrone, M. C., Cicchini, M. & Burr, D. C. (2010). Spatial maps for time and motion,Exp Brain Res, 2 (206), 121-128. PDF
In this article, we review recent research studying the mechanisms for transforming coordinate systems to encode space, time and motion. A range of studies using functional imaging and psychophysical techniques reveals mechanisms in the human brain for encoding information in external rather than retinal coordinates. This reinforces the idea of a tight relationship between space and time, in the parietal cortex of primates.
Morrone, M. C. (2010). Brain development: critical periods for cross-sensory plasticity,Curr Biol, 21 (20), R934-936. PDF
Recent work has shown that visual deprivation of humans during a critical period leads to motion area MT+ responding to auditory motion. This cross-sensory plasticity, an important form of brain reorganization, may be mediated by top-down brain circuits from pre-frontal cortex.
Schutz, A. C. & Morrone, M. C. (2010). Compression of time during smooth pursuit eye movements,Vision Res, 24 (50), 2702-2713. PDF
Humans have a clear sense for the passage of time, but while implicit motor timing is quite accurate, explicit timing is prone to distortions particularly during action (Wenke & Haggard, 2009) and saccadic eye movements (Morrone, Ross, & Burr, 2005). Here, we investigated whether perceived duration is also affected by the execution of smooth pursuit eye movements, showing a compression of apparent duration similar to that observed during saccades. To this end, we presented two brief bars that marked intervals between 100 and 300 ms and asked subjects to judge their duration during fixation and pursuit. We found a compression of perceived duration for bars modulated in luminance contrast of about 32% and for bars modulated in chromatic contrast of 14% during pursuit compared to fixation. Interestingly, Weber ratios were similar for fixation and pursuit, if they are expressed as ratio between JND and perceived duration. This compression was constant for pursuit speeds from 7 to 14 deg/s and did not occur for intervals marked by auditory events. These results argue for a modality-specific component in the processing of temporal information.
Campanella, F., Sandini, G. & Morrone, M. C. (2010). Visual information gleaned by observing grasping movement in allocentric and egocentric perspectives,Proc Biol Sci, 1715 (278), 2142-2149. PDF
One of the major functions of vision is to allow for an efficient and active interaction with the environment. In this study, we investigate the capacity of human observers to extract visual information from observation of their own actions, and those of others, from different viewpoints. Subjects discriminated the size of objects by observing a point-light movie of a hand reaching for an invisible object. We recorded real reach-and-grasp actions in three-dimensional space towards objects of different shape and size, to produce two-dimensional 'point-light display' movies, which were used to measure size discrimination for reach-and-grasp motion sequences, release-and-withdraw sequences and still frames, all in egocentric and allocentric perspectives. Visual size discrimination from action was significantly better in egocentric than in allocentric view, but only for reach-and-grasp motion sequences: release-and-withdraw sequences or still frames derived no advantage from egocentric viewing. The results suggest that the system may have access to an internal model of action that contributes to calibrate visual sense of size for an accurate grasp.
Lunghi, C., Binda, P. & Morrone, M. C. (2010). Touch disambiguates rivalrous perception at early stages of visual analysis,Curr Biol, 4 (20), R143-144. PDF
Binocular rivalry is a powerful tool to study human consciousness: two equally salient stimuli are imaged on the retinae, but at any given instant only one is consciously perceived, the other suppressed.The suppression takes place early, probably in V1. However, a trace of the suppressed signal has been detected along the dorsal visual pathway (BOLD responses) and demonstrated with psychophysical experiments. The suppressed image of a rotating sphere during rivalry is restored to consciousness when the observer actively controls the rotation and a similar effect on the suppressed signal has been shown for motion perception and reflexive eye movements (see Supplemental References). Here, we asked whether cross-modal sensory signals could selectively interact with rivalrous visual signals that are analyzed at a very early stage, probably V1. An auditory stimulus, when attended, can influence binocular rivalry, extending dominance times for a congruent visual stimulus. Tactile information can also disambiguate unstable visual motion and can fuse with vision to improve discrimination (e.g. slant). Our results indicate that a haptic oriented stimulus can disambiguate visual perception during binocular rivalry of gratings of orthogonal orientation, not only by prolonging dominance but also by curtailing suppression of the visual stimulus of matched orientation. The effect is selective for the spatial frequency of the stimuli, suggesting that haptic signals interact with early visual representations to enhance access to conscious perception.
Burr, D. C., Ross, J., Binda, P. & Morrone, M. C. (2010). Saccades compress space, time and number,Trends Cogn Sci, 12 (14), 528-533. PDF
It has been suggested that space, time and number are represented on a common subjective scale. Saccadic eye movements provide a fascinating test. Saccades compress the perceived magnitude of spatial separations and temporal intervals to approximately half of their true value. The question arises as to whether saccades also compress number. They do, and compression follows a very similar time course for all three attributes: it is maximal at saccadic onset and decreases to veridicality within a window of approximately 50ms. These results reinforce the suggestion of a common perceptual metric, which is probably mediated by the intraparietal cortex; they further suggest that before each saccade the common metric for all three is reset, possibly to pave the way for a fresh analysis of the post-saccadic situation.
Burr, D. C. & Morrone, M. C. (2010). Vision: keeping the world still when the eyes move,Curr Biol, 10 (20), R442-444. PDF
A long-standing problem for visual science is how the world remains so apparently stable in the face of continual rapid eye movements. New experimental evidence, and computational models are helping to solve this mystery.
Binda, P., Morrone, M. C. & Burr, D. C. (2010). Temporal auditory capture does not affect the time course of saccadic mislocalization of visual stimuli,J Vis, 2 (10), 7 1-13. PDF
Irrelevant sounds can "capture" visual stimuli to change their apparent timing, a phenomenon sometimes termed "temporal ventriloquism". Here we ask whether this auditory capture can alter the time course of spatial mislocalization of visual stimuli during saccades. We first show that during saccades, sounds affect the apparent timing of visual flashes, even more strongly than during fixation. However, this capture does not affect the dynamics of perisaccadic visual distortions. Sounds presented 50 ms before or after a visual bar (that change perceived timing of the bars by more than 40 ms) had no measurable effect on the time courses of spatial mislocalization of the bars, in four subjects. Control studies showed that with barely visible, low-contrast stimuli, leading, but not trailing, sounds can have a small effect on mislocalization, most likely attributable to attentional effects rather than auditory capture. These findings support previous studies showing that integration of multisensory information occurs at a relatively late stage of sensory processing, after visual representations have undergone the distortions induced by saccades.
2009 (back to top)
Cicchini, G. M. & Morrone, M. C. (2009). Shifts in spatial attention affect the perceived duration of events,J Vis, 1 (9), 9 1-13. PDF
We investigated the relationship between attention and perceived duration of visual events with a double-task paradigm. The primary task was to discriminate the size change of a 2 degree circle presented 10 degrees left, right, above, or below fixation; the secondary task was to judge the temporal separation (from 133 ms to 633 ms) of two equiluminant horizontal bars (10 deg x 2 deg) briefly flashed 12 degrees above or below fixation. The stimulus onset asynchrony (SOA) between primary and secondary task ranged from -1300 ms to +1000 ms. Temporal intervals in proximity of the onset of the primary task stimuli were perceived strongly compressed by up to 40%. The effect was proportional to the size of the interval with a maximum effect at 100 ms SOA. Control experiments show that neither primary-task difficulty, nor the type of primary task discrimination (form or motion, or equiluminant or luminance contrast) nor spatial congruence between primary and secondary task alter the effect. Interestingly, the compression occurred only when the intervals are marked by bars presented in separated spatial locations: when the interval is marked by two bars flashed in the same spatial position no temporal distortion was found. These data indicate that attention can alter perceived duration when the brain has to compare the passage of time at two different spatial positions, corroborating earlier findings that mechanisms of time perception may monitor separately the various spatial locations possibly at high level of analysis.
Burr, D., Silva, O., Cicchini, G. M., Banks, M. S. & Morrone, M. C. (2009). Temporal mechanisms of multimodal binding,Proc Biol Sci, 1663 (276), 1761-1769. PDF
The simultaneity of signals from different senses-such as vision and audition-is a useful cue for determining whether those signals arose from one environmental source or from more than one. To understand better the sensory mechanisms for assessing simultaneity, we measured the discrimination thresholds for time intervals marked by auditory, visual or auditory-visual stimuli, as a function of the base interval. For all conditions, both unimodal and cross-modal, the thresholds followed a characteristic 'dipper function' in which the lowest thresholds occurred when discriminating against a non-zero interval. The base interval yielding the lowest threshold was roughly equal to the threshold for discriminating asynchronous from synchronous presentations. Those lowest thresholds occurred at approximately 5, 15 and 75 ms for auditory, visual and auditory-visual stimuli, respectively. Thus, the mechanisms mediating performance with cross-modal stimuli are considerably slower than the mechanisms mediating performance within a particular sense. We developed a simple model with temporal filters of different time constants and showed that the model produces discrimination functions similar to the ones we observed in humans. Both for processing within a single sense, and for processing across senses, temporal perception is affected by the properties of temporal filters, the outputs of which are used to estimate time offsets, correlations between signals, and more.
Binda, P., Cicchini, G. M., Burr, D. C. & Morrone, M. C. (2009). Spatiotemporal distortions of visual perception at the time of saccades,J Neurosci, 42 (29), 13147-13157. PDF
Both space and time are grossly distorted during saccades. Here we show that the two distortions are strongly linked, and that both could be a consequence of the transient remapping mechanisms that affect visual neurons perisaccadically. We measured perisaccadic spatial and temporal distortions simultaneously by asking subjects to report both the perceived spatial location of a perisaccadic vertical bar (relative to a remembered ruler), and its perceived timing (relative to two sounds straddling the bar). During fixation and well before or after saccades, bars were localized veridically in space and in time. In different epochs of the perisaccadic interval, temporal perception was subject to different biases. At about the time of the saccadic onset, bars were temporally mislocalized 50-100 ms later than their actual presentation and spatially mislocalized toward the saccadic target. Importantly, the magnitude of the temporal distortions co-varied with the spatial localization bias and the two phenomena had similar dynamics. Within a brief period about 50 ms before saccadic onset, stimuli were perceived with shorter latencies than at other delays relative to saccadic onset, suggesting that the perceived passage of time transiently inverted its direction. Based on this result we could predict the inversion of perceived temporal order for two briefly flashed visual stimuli. We developed a model that simulates the perisaccadic transient change of neuronal receptive fields predicting well the reported temporal distortions. The key aspects of the model are the dynamics of the "remapped" activity and the use of decoder operators that are optimal during fixation, but are not updated perisaccadically.
Burr, D. C., Baldassi, S., Morrone, M. C. & Verghese, P. (2009). Pooling and segmenting motion signals,Vision Res, 10 (49), 1065-1072. PDF
Humans are extremely sensitive to visual motion, largely because local motion signals can be integrated over a large spatial region. On the other hand, summation is often not advantageous, for example when segmenting a moving stimulus against a stationary or oppositely moving background. In this study we show that the spatial extent of motion integration is not compulsory, but is subject to voluntary attentional control. Measurements of motion coherence sensitivity with summation and search paradigms showed that human observers can combine motion signals from cued regions or patches in an optimal manner, even when the regions are quite distinct and remote from each other. Further measurements of contrast sensitivity reinforce previous studies showing that motion integration is preceded by a local analysis akin to contrast thresholding (or intrinsic uncertainty). The results were well modelled by two standard signal-detection-theory models.
Burr, D., Banks, M. S. & Morrone, M. C. (2009). Auditory dominance over vision in the perception of interval duration,Exp Brain Res, 1 (198), 49-57. PDF
The "ventriloquist effect" refers to the fact that vision usually dominates hearing in spatial localization, and this has been shown to be consistent with optimal integration of visual and auditory signals (Alais and Burr in Curr Biol 14(3):257-262, 2004). For temporal localization, however, auditory stimuli often "capture" visual stimuli, in what has become known as "temporal ventriloquism". We examined this quantitatively using a bisection task, confirming that sound does tend to dominate the perceived timing of audio-visual stimuli. The dominance was predicted qualitatively by considering the better temporal localization of audition, but the quantitative fit was less than perfect, with more weight being given to audition than predicted from thresholds. As predicted by optimal cue combination, the temporal localization of audio-visual stimuli was better than for either sense alone.
2008 (back to top)
Morrone, M. C., Guzzetta, A., Tinelli, F., Tosetti, M., Del Viva, M., Montanaro, D., et al. (2008). Inversion of perceived direction of motion caused by spatial undersampling in two children with periventricular leukomalacia,J Cogn Neurosci, 6 (20), 1094-1106. PDF
We report here two cases of two young diplegic patients with cystic periventricular leukomalacia who systematically, and with high sensitivity, perceive translational motion of a random-dot display in the opposite direction. The apparent inversion was specific for translation motion: Rotation and expansion motion were perceived correctly, with normal sensitivity. It was also specific for random-dot patterns, not occurring with gratings. For the one patient that we were able to test extensively, contrast sensitivity for static stimuli was normal, but was very low for direction discrimination at high spatial frequencies and all temporal frequencies. His optokinetic nystagmus movements were normal but he was unable to track a single translating target, indicating a perceptual origin of the tracking deficit. The severe deficit for motion perception was also evident in the seminatural situation of a driving simulation video game. The perceptual deficit for translational motion was reinforced by functional magnetic resonance imaging studies. Translational motion elicited no response in the MT complex, although it did produce a strong response in many visual areas when contrasted with blank stimuli. However, radial and rotational motion produced a normal pattern of activation in a subregion of the MT complex. These data reinforce the existent evidence for independent cortical processing for translational, and circular or radial flow motion, and further suggest that the two systems have different vulnerability and plasticity to prenatal damage. They also highlight the complexity of visual motion perception, and how the delicate balance of neural activity can lead to paradoxical effects such as consistent misperception of the direction of motion. We advance a possible explanation of a reduced spatial sampling of the motion stimuli and report a simple model that simulates well the experimental results.
Perna, A., Tosetti, M., Montanaro, D. & Morrone, M. C. (2008). BOLD response to spatial phase congruency in human brain,J Vis, 10 (8), 15 11-15. PDF
Human psychophysical observations, computational models, and the selectivity of neurons in primary visual cortex all suggest that an early step in visual processing is the detection of features such as lines and edges. However, previous fMRI experiments investigating the responses of early visual areas to phase coherence have led to apparently discordant results. We studied the human brain BOLD responses to structured periodic band-pass images of matched amplitude spectrum but of different phase spectra, arranged to create three distinct types of stimuli: pure edges; pure lines (matched global and local energy to the edges, but different phase); and random noise (random phase spectrum, hence no salient features, and a different spatial distribution of local energy from the lines and edges stimuli). Alternation of lines against edges did not activate primary visual cortex, but did activate two higher order visual areas. Alternation of these lines or edges against the random stimulus produced a strong activity in many visual areas, including primary visual cortex. Interestingly, the BOLD activity was higher for the edges and lines than for the random stimuli for a wide range of stimulus contrasts, indicating the presence of non-linear gain modulation in the cell response. These results show that phase congruency is coded at the level of primary visual cortex. We show that a stage of response gain modulation can explain our present and previous fMRI discordant results.
2007 (back to top)
Bruno, A. & Morrone, M. C. (2007). Influence of saccadic adaptation on spatial localization: comparison of verbal and pointing reports,J Vis, 5 (7), 16 11-13. PDF
Under conditions of short-term saccadic adaptation, stimuli presented long before saccadic onset are perceptually mislocalized in space. Here we study whether saccadic adaptation can also affect localization of objects by pointing. We measured localization performance during fixation and after normal saccades and adapted saccades, for a bar presented well before a saccadic eye movement, for both pointing and verbal localization, under open-loop conditions generated by a transient dark period about 300 ms after the presentation of the bar. During fixation and normal saccade, localization performance for verbal report was veridical, while for pointing there was an overestimation of the target eccentricity respect to gaze, in agreement with the idea of separate representations of space for action and perception. During saccadic adaptation, there was a significant shift of both pointing and verbal report localization in the direction of adaptation with similar spatial selectivity for both tasks. These results indicate that saccadic adaptation induces a similar re-calibration of the action map as well as of the perceptual map, suggesting a common site of operation in the transformation from eye-centered to gaze-centered coordinates.
Tozzi, A., Morrone, M. C. & Burr, D. C. (2007). The effect of optokinetic nystagmus on the perceived position of briefly flashed targets,Vision Res, 6 (47), 861-868. PDF
Stimuli flashed briefly around the time of an impending saccade are mislocalized in the direction of the saccade and also compressed towards the saccadic target. Similarly, targets flashed during pursuit eye movements are mislocalized in the direction of pursuit. Here, we investigate the effects of optokinetic nystagmus (OKN) on visual localization. Subjects passively viewed a wide-field drifting grating that elicited strong OKN, comprising the characteristic slow-phase tracking movement interspersed with corrected "saccade-like" fast-phase movements. Subjects reported the apparent position of salient bars flashed briefly at various positions on the screen. In general, bars were misperceived in the direction of the slow-phase tracking movement. Bars flashed around the onset of the fast-phase movements were subject to much less mislocalization, pointing to a competing shift in the direction of the fast-phase, as occurs with saccades. However, as distinct from saccades, there was no evidence for spatial compression around the time of the corrective fast-phase OKN. The results suggest that OKN cause perceptual mislocalizations similar to those of smooth pursuit and saccades, but there are some differences in the nature of the mislocalizations, pointing to different perceptual mechanisms associated with the different types of eye movements.
Perna, A. & Morrone, M. C. (2007). The lowest spatial frequency channel determines brightness perception,Vision Res, 10 (47), 1282-1291. PDF
This study investigates the role played by individual spatial scales in determining the apparent brightness of greyscale patterns. We measured the perceived difference in brightness across an edge in the presence of notch filtering and high-pass filtering for two stimulus configurations, one that elicits the perception of transparency and one that appears opaque. For both stimulus configurations, the apparent brightness of the surfaces delimited by the border decreased monotonically with progressive (ideal) high-pass filtering, with a critical cut-off at 1 c/deg. Using two octave ideal notch filtering, the maximum detrimental effect on apparent brightness was observed at about 1c/deg. Critical frequencies for apparent brightness did not vary with contrast, viewing distance, or surface size, suggesting that apparent brightness is determined by the channel tuned at 1 c/deg. Modelling the data with the local energy model [Morrone, M. C., & Burr, D. C. (1988). Feature detection in human vision: a phase dependent energy model. Proceedings of the Royal Society (London), B235, 221-245] at 1c/deg confirmed the suggestion that this channel mediates apparent brightness for both opaque and transparent borders, with no need for pooling or integration across spatial channels.
Burr, D., Tozzi, A. & Morrone, M. C. (2007). Neural mechanisms for timing visual events are spatially selective in real-world coordinates,Nat Neurosci, 4 (10), 423-425. PDF
It is generally assumed that perceptual events are timed by a centralized supramodal clock. This study challenges this notion in humans by providing clear evidence that visual events of subsecond duration are timed by visual neural mechanisms with spatially circumscribed receptive fields, localized in real-world, rather than retinal, coordinates.
d'Avossa, G., Tosetti, M., Crespi, S., Biagi, L., Burr, D. C. & Morrone, M. C. (2007). Spatiotopic selectivity of BOLD responses to visual motion in human area MT,Nat Neurosci, 2 (10), 249-255. PDF
Many neurons in the monkey visual extrastriate cortex have receptive fields that are affected by gaze direction. In humans, psychophysical studies suggest that motion signals may be encoded in a spatiotopic fashion. Here we use functional magnetic resonance imaging to study spatial selectivity in the human middle temporal cortex (area MT or V5), an area that is clearly implicated in motion perception. The results show that the response of MT is modulated by gaze direction, generating a spatial selectivity based on screen rather than retinal coordinates. This area could be the neurophysiological substrate of the spatiotopic representation of motion signals.
Chirimuuta, M., Burr, D. & Morrone, M. C. (2007). The role of perceptual learning on modality-specific visual attentional effects,Vision Res, 1 (47), 60-70. PDF
Morrone et al. [Morrone, M. C., Denti, V., & Spinelli, D. (2002). Color and luminance contrasts attract independent attention. Current Biology, 12, 1134-1137] reported that the detrimental effect on contrast discrimination thresholds of performing a concomitant task is modality specific: performing a secondary luminance task has no effect on colour contrast thresholds, and vice versa. Here we confirm this result with a novel task involving learning of spatial position, and go on to show that it is not specific to the cardinal colour axes: secondary tasks with red-green stimuli impede performance on a blue-yellow task and vice versa. We further show that the attentional effect can be abolished with continued training over 2-4 training days (2-20 training sessions), and that the effect of learning is transferable to new target positions. Given the finding of transference, we discuss the possibility that V4 is a site of plasticity for both stimulus types, and that the separation is due to a luminance-colour separation within this cortical area.
Binda, P., Bruno, A., Burr, D. C. & Morrone, M. C. (2007). Fusion of visual and auditory stimuli during saccades: a Bayesian explanation for perisaccadic distortions,J Neurosci, 32 (27), 8525-8532. PDF
Brief stimuli presented near the onset of saccades are grossly mislocalized in space. In this study, we investigated whether the Bayesian hypothesis of optimal sensory fusion could account for the mislocalization. We required subjects to localize visual, auditory, and audiovisual stimuli at the time of saccades (compared with an earlier presented target). During fixation, vision dominates and spatially "captures" the auditory stimulus (the ventriloquist effect). But for perisaccadic presentations, auditory localization becomes more important, so the mislocalized visual stimulus is seen closer to its veridical position. The precision of the bimodal localization (as measured by localization thresholds or just-noticeable difference) was better than either the visual or acoustic stimulus presented in isolation. Both the perceived position of the bimodal stimuli and the improved precision were well predicted by assuming statistically optimal Bayesian-like combination of visual and auditory signals. Furthermore, the time course of localization was well predicted by the Bayesian approach. We present a detailed model that simulates the time-course data, assuming that perceived position is given by the sum of retinal position and a sluggish noisy eye-position signal, obtained by integrating optimally the output of two populations of neural activity: one centered at the current point of gaze, the other centered at the future point of gaze.
2006 (back to top)
Del Viva, M. M. & Morrone, M. C. (2006). A feature-tracking model simulates the motion direction bias induced by phase congruency,J Vis, 3 (6), 179-195. PDF
Here we report a new motion illusion where the prevailing motion direction is strongly influenced by the relative phase of the harmonic components of the stimulus. The basic stimulus is the sum of three sinusoidal contrast-reversing gratings: the first, the third, and the fifth harmonic of two square wave gratings that drift in opposite direction. The phase of one of the fifth components was kept constant at 180 deg, whereas the phase of the other fifth harmonic was varied over the range 0-150 deg. For each phase value of the fifth harmonic, the motion was strongly biased toward its direction, corresponding to the direction with stronger phase congruency between the three harmonics. The strength of the prevailing motion was assessed by measuring motion direction discrimination thresholds, by varying the contrast of the third and the fifth harmonics plaid pattern. Results show that the contrast of high harmonics had to be increased by more than a factor of 10, to achieve a balance of motion for phase differences greater than 60 deg between the 2 fifth harmonics. We also measured the dependence on the absolute phase of harmonic components and found that it is not an important parameter, excluding the possibility that local luminance cues could be mediating the effect. A feature-tracking model based on previous work is proposed to simulate the data. The model computes local energy function from a pair of space-time separable front stage filters and applies a battery of directional second stage mechanisms. It is able to simulate quantitatively the phase congruency dependence illusion and the insensitivity to overall phase. Other energy models based on directional filters fail to simulate the phase congruency dependency effect.
Bruno, A., Brambati, S. M., Perani, D. & Morrone, M. C. (2006). Development of saccadic suppression in children,J Neurophysiol, 3 (96), 1011-1017. PDF
We measured saccadic suppression in adolescent children and young adults using spatially curtailed low spatial frequency stimuli. For both groups, sensitivity for color-modulated stimuli was unchanged during saccades. Sensitivity for luminance-modulated stimuli was greatly reduced during saccades in both groups but far more for adolescents than for young adults. Adults' suppression was on average a factor of about 3, whereas that for the adolescent group was closer to a factor of 10. The specificity of the suppression to luminance-modulated stimuli excludes generic explanations such as task difficulty and attention. We suggest that the enhanced suppression in adolescents results from the immaturity of the ocular-motor system at that age.
Alais, D., Morrone, C. & Burr, D. (2006). Separate attentional resources for vision and audition,Proc Biol Sci, 1592 (273), 1339-1345. PDF
Current models of attention, typically claim that vision and audition are limited by a common attentional resource which means that visual performance should be adversely affected by a concurrent auditory task and vice versa. Here, we test this implication by measuring auditory (pitch) and visual (contrast) thresholds in conjunction with cross-modal secondary tasks and find that no such interference occurs. Visual contrast discrimination thresholds were unaffected by a concurrent chord or pitch discrimination, and pitch-discrimination thresholds were virtually unaffected by a concurrent visual search or contrast discrimination task. However, if the dual tasks were presented within the same modality, thresholds were raised by a factor of between two (for visual discrimination) and four (for auditory discrimination). These results suggest that at least for low-level tasks such as discriminations of pitch and contrast, each sensory modality is under separate attentional control, rather than being limited by a supramodal attentional resource. This has implications for current theories of attention as well as for the use of multi-sensory media for efficient informational transmission.
Burr, D. & Morrone, C. (2006). Perception: transient disruptions to neural space-time,Curr Biol, 19 (16), R847-849. PDF
How vision operates efficiently in the face of continuous shifts of gaze remains poorly understood. Recent studies show that saccades cause dramatic, but transient, changes in the spatial and also temporal tuning of cells in many visual areas, which may underly the perceptual compression of space and time, and serve to counteract the effects of the saccades and maintain visual stability.
Burr, D., McKee, S. & Morrone, C. M. (2006). Resolution for spatial segregation and spatial localization by motion signals,Vision Res, 6-7 (46), 932-939. PDF
We investigated two types of spatial resolution for perceiving motion-defined contours: grating acuity, the capacity to discriminate alternating stripes of opposed motion from transparent bi-directional motion; and alignment acuity, the capacity to localize the position of motion-defined edges with respect to stationary markers. For both tasks the stimuli were random noise patterns, low-pass filtered in the spatial dimension parallel to the motion. Both grating and alignment resolution varied systematically with spatial frequency cutoff and speed. Best performance for grating resolution was about 10 c/deg (for unfiltered patterns moving at 1-4 deg/s), corresponding to a stripe resolution of about 3'. Grating resolution corresponds well to estimates of smallest receptive field size of motion units under these conditions, suggesting that opposing signals from units with small receptive fields (probably located in V1) are contrasted efficiently to define edges. Alignment resolution was about 2' at best, under similar conditions. Whereas alignment judgment based on luminance-defined edges is typically 3-10 times better than resolution, alignment based on motion-defined edges is only 1.1-1.5 times better, suggesting motion contours are less effectively encoded than luminance contours.
Burr, D. & Morrone, C. (2006). Time perception: space-time in the brain,Curr Biol, 5 (16), R171-173. PDF