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Alessandro Benedetto

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Alessandro Benedetto

Post-Doc in Department of Translational Research and New Technologies in Medicine and Surgery, University of Pisa


  • Email: alessandro.benedetto ( AT ) med.unipi.it

Research laboratories

  • Department of Translational Research on New Technologies in Medicine and Surgery, University of Pisa
  • CNR Institute of Neuroscience, Pisa



  • 2013-2016: PhD course in Neuroscience - Dottorato Toscano di Neuroscienze (XXIX ciclo).
  • 2011-2013: Master degree in Psychology - Cognitive Neuroscience. University Vita-Salute San Raffaele, Milano.
  • 2008-2011: Bachelor in Philosophy of Mind and Language. University Vita-Salute San Raffaele, Milano.


Current research and interests

  • Behavioral and neural oscillations
  • Action and perception
  • Time perception



  • "A Behavioral Study of Sensorimotor Integration in Metrical Coding". Master's Thesis, University Vita-Salute San Raffaele Milano (2013). Supervisor: G. Baud-Bovy.
  • "The temporal dynamics of vision for action and perception". Doctoral Thesis, University of Florence (03/04/2017). Supervisor: M.C. Morrone.


Benedetto, A., Morrone, C. & Tomassini, A. (2019). The Common Rhythm of Action and Perception., Journal of Cognitive Neuroscience, PDF

Research in the last decade has undermined the idea of perception as a continuous process, providing strong empirical support for its rhythmic modulation. More recently, it has been revealed that the ongoing motor processes influence the rhythmic sampling of sensory information. In this review, we will focus on a growing body of evidence suggesting that oscillation-based mechanisms may structure the dynamic interplay between the motor and sensory system and provide a unified temporal frame for their effective coordination. We will describe neurophysiological data, primarily collected in animals, showing phase-locking of neuronal oscillations to the onset of (eye) movements. These data are complemented by novel evidence in humans, which demonstrate the behavioral relevance of these oscillatory modulations and their domain-general nature. Finally, we will discuss the possible implications of these modulations for action-perception coupling mechanisms.

Benedetto, A. & Morrone, M. C. (2019). Visual sensitivity and bias oscillate phase-locked to saccadic eye movements, Journal of Vision, 14 (19), 15. PDF

Oscillations in perceptual performance have been observed before and after a voluntary action, like hand, finger, and eye movements. In particular, discrimination accuracy of suprathreshold contrast stimuli oscillates in the delta range (2–3 Hz) phase-locked to saccadic eye movements. Importantly, saccadic suppression is embedded in phase with these long-lasting perceptual oscillations. It is debated whether these rhythmic modulations affect only appearance of high-contrast stimuli or whether absolute detection threshold is also modulated rhythmically. Here we measured location discrimination of a brief Gabor patch presented randomly between 1 s before and after a voluntary saccade and demonstrated that absolute contrast thresholds oscillated at a similar frequency to suprathreshold contrast discrimination. Importantly, saccadic suppression is also embedded in phase with absolute threshold oscillations. Interestingly, response bias was also found to oscillate at the same frequency in both tasks. However, the frequency was in the alpha range for bias, while it was in the delta range for sensitivity. These results demonstrate the presence of perisaccadic delta oscillations in visual sensitivity phase-locked to saccadic onset, and independent from response bias alpha oscillations. Overall, the present findings reinforce the suggestion of a leading role of oscillations in the temporal binding between eye-movement and visual processing timing.


Benedetto, A., Burr, D. C. & Morrone, M. C. (2018). Perceptual Oscillation of Audiovisual Time Simultaneity, eNeuro, 3 (5), PDF

Action and perception are tightly coupled systems requiring coordination and synchronization over time. How the brain achieves synchronization is still a matter of debate, but recent experiments suggest that brain oscillations may play an important role in this process. Brain oscillations have been also proposed to be fundamental in determining time perception. Here, we had subjects perform an audiovisual temporal order judgment task to investigate the fine dynamics of temporal bias and sensitivity before and after the execution of voluntary hand movement (button-press). The reported order of the audiovisual sequence was rhythmically biased as a function of delay from hand action execution. Importantly, we found that it oscillated at a theta range frequency, starting approximately 500 ms before and persisting approximately 250 ms after the button-press, with consistent phase-locking across participants. Our results show that the perception of cross-sensory simultaneity oscillates rhythmically in synchrony with the programming phase of a voluntary action, demonstrating a link between action preparation and bias in temporal perceptual judgments.


Benedetto, A. & Morrone, M.C. (2017). Saccadic suppression is embedded within extended oscillatory modulation of sensitivity, J Neurosci, PDF

Action and perception are intimately coupled systems; one clear case is saccadic suppression, the reduced visibility around the time of saccades, important in mediating visual stability; another is the oscillatory modulation of visibility synchronized with hand action. To suppress effectively the spurious retinal motion generated by the eye movements, it is crucial that saccadic suppression and saccadic onset be temporally synchronous. However, the mechanisms that determine this temporal synchrony are unknown. We investigated the effect of saccades on contrast discrimination sensitivity over a long period stretching over more than 1 second before and after saccade execution. Human subjects made horizontal saccades at will to two stationary saccadic targets separated by 20 degrees degrees. At a random interval, a brief Gabor patch was displayed between the two fixations in either the upper or lower visual field, and the subject had to detect its location. Strong saccadic suppression was measured between -50 and 50 ms from saccadic onset. However, the suppression was systematically embedded in a trough of oscillations of contrast sensitivity that fluctuated rhythmically in the delta range (at about 3 Hz), commencing about one second before saccade execution and lasting for up to one second after the saccade. The results show that saccadic preparation and visual sensitivity oscillations are coupled, and the coupling might be instrumental in temporally aligning the initiation of the saccade with the visual suppression.Significant statementSaccades are known to produce a suppression of contrast sensitivity at saccadic onset and an enhancement after saccadic offset. Here we show that these dynamics are systematically embedded in visual oscillations of contrast sensitivity that fluctuate rhythmically in the delta range (at about 3 Hz), commencing about one second before saccade execution and lasting for up to one second after the saccade. The results show that saccadic preparation and visual sensitivity oscillations are coupled, and the coupling might be instrumental in aligning temporally the initiation of the saccade with the visual suppression.

Benedetto, A., Lozano-Soldevilla, D. & VanRullen, R. (2017). Different Responses of Spontaneous and Stimulus-Related Alpha Activity to Ambient Luminance Changes, Eur J Neurosci, PDF

Alpha oscillations are particularly important in determining our percepts and have been implicated in fundamental brain functions. Oscillatory activity can be spontaneous or stimulus-related. Furthermore, stimulus-related responses can be phase- or non-phase-locked to the stimulus. Non-phase-locked (induced) activity can be identified as the average amplitude changes in response to a stimulation, while phase-locked activity can be measured via reverse correlation techniques (echo function). However, the mechanisms and the functional roles of these oscillations are far from clear. Here, we investigated the effect of ambient luminance changes, known to dramatically modulate neural oscillations, on spontaneous and stimulus-related alpha. We investigated the effect of ambient luminance on EEG alpha during spontaneous human brain activity at rest (experiment 1) and during visual stimulation (experiment 2). Results show that spontaneous alpha amplitude increased by decreasing ambient-luminance, while alpha frequency remained unaffected. In the second experiment, we found that under low-luminance viewing the stimulus-related alpha amplitude was lower, and its frequency was slightly faster. These effects were evident in the phase-locked part of the alpha response (echo function), but weaker or absent in the induced (non-phase-locked) alpha responses. Finally, we explored the possible behavioral correlates of these modulations in a monocular critical flicker frequency task (experiment 3), finding that dark adaptation in the left eye decreased the temporal threshold of the right eye. Overall, we found that ambient luminance changes impact differently on spontaneous and stimulus-related alpha expression. We suggest that stimulus-related alpha activity is crucial in determining human temporal segmentation abilities.


Benedetto, A. & Binda, P. (2016). Dissociable saccadic suppression of pupillary and perceptual responses to light, J Neurophysiol, 3 (115), 1243-1251. PDF

We measured pupillary constrictions in response to full-screen flashes of variable luminance, occurring either at the onset of a saccadic eye movement or well before/after it. A large fraction of perisaccadic flashes were undetectable to the subjects, consistent with saccadic suppression of visual sensitivity. Likewise, pupillary responses to perisaccadic flashes were strongly suppressed. However, the two phenomena appear to be dissociable. Across subjects and luminance levels of the flash stimulus, there were cases in which conscious perception of the flash was completely depleted yet the pupillary response was clearly present, as well as cases in which the opposite occurred. On one hand, the fact that pupillary light responses are subject to saccadic suppression reinforces evidence that this is not a simple reflex but depends on the integration of retinal illumination with complex "extraretinal" cues. On the other hand, the relative independence of pupillary and perceptual responses suggests that suppression acts separately on these systems-consistent with the idea of multiple visual pathways that are differentially affected by saccades.

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.


  • P.Binda, A.Benedetto. Cortical control of the pupillary light response. 9th FENS (Poster). Milan, Italy, 2014.
  • A.Benedetto, G.M.Cicchini, D.Spinelli, D.C.Burr, M.C.Morrone. Luminance level rhythmically modulates early sensory function. New Perspectives in Neuroscience: Research Results of Young Italian Neuroscientists (Poster). Naples, Italy, 26/02/2015.
  • A.Benedetto, M.C.Morrone, D.Burr. Audio-Visual Temporal-Order Judgment Reveals Rhythmic Oscillations in Temporal Bias. International Multisensory Research Forum 2015 (Poster). Pisa, Italy, 13-16/06/2015.
  • A.Benedetto, D.Burr, D.Spinelli, M.C.Morrone. Rhythmic modulation of human visual sensitivity depends on luminance. European Conference on Visual Perception 2015 (Poster). Liverpool, UK, 23-27/08/2015.
  • A.Benedetto, D.Spinelli, M.C. Morrone. Rhythmic modulation of visual contrast discrimination triggered by action. Oscillatory processes in perception and cognition (Talk). Toulouse, France, 9/12/2015.
  • A.Benedetto, M.C.Morrone. Saccadic preparation triggers visual oscillations in contrast sensitivity. International Multisensory Research Forum 2016 (Talk). Suzhou, Cina, 15-18/06/2016.
  • A.Benedetto,Lozano-Soldevilla D., Vanrullen R. Ambient luminance changes modulate oscillatory properties of the visual system. 17th Vision Science Society Meeting (Poster). St. Pete Beach, Florida, USA, May 19-24, 2017
  • Morrone MC., Benedetto A. Rhythmic modulation of human visual sensitivity synchronized with planning of saccades. 17th Vision Science Society Meeting (Talk). St. Pete Beach, Florida, USA, May 19-24, 2017


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