Adaptation to numerosity affects the pupillary light response.

We recently showed that the gain of the pupillary light response depends on numerosity, with weaker responses to fewer items. Here we show that this effect holds when the stimuli are physically identical but are perceived as less numerous due to numerosity adaptation. Twenty-eight participants adapted to low (10 dots) or high (160 dots) numerosities and subsequently watched arrays of 10-40 dots, with variable or homogeneous dot size. Luminance was constant across all stimuli. Pupil size was measured with passive viewing, and the effects of adaptation were checked in a separate psychophysical session. We found that perceived numerosity was systematically lower, and pupillary light responses correspondingly smaller, following adaptation to high rather than low numerosities. This is consistent with numerosity being a primary visual feature, spontaneously encoded even when task irrelevant, and affecting automatic and unconscious behaviours like the pupillary light response.

Investigating cross-orientation inhibition with continuous tracking.

We investigated cross-orientation inhibition with the recently developed continuous tracking technique. We designed an experiment where participants tracked the horizontal motion of a narrow vertical grating. The target was superimposed on one of three different backgrounds, in separate sessions: a uniform gray background or a sinusoidal grating oriented either parallel or orthogonal to the target. Both mask and target where phase reversed. We cross-correlated target and mouse movements and compared the peaks and lags of response with the different masks. Our results are in agreement with previous findings on cross-orientation inhibition: The orthogonal mask had a weak effect on the peaks and lags of correlation as a function of target contrast, consistently with a divisive effect of the mask, while the parallel mask acted subtractively on the response. Interestingly, lags of correlation decreased approximately linearly with contrast, with decrements of the order of 100 ms, even at 10 times the detection threshold, confirming that it is possible to investigate behavioral differences above threshold using the continuous tracking paradigm.

Serial Dependence in Perception.

Much evidence has shown that perception is biased towards previously presented similar stimuli, an effect recently termed serial dependence. Serial dependence affects nearly every aspect of perception, often causing gross perceptual distortions, especially for weak and ambiguous stimuli. Despite unwanted side-effects, empirical evidence and Bayesian modeling show that serial dependence acts to improve efficiency and is generally beneficial to the system. Consistent with models of predictive coding, the Bayesian priors of serial dependence are generated at high levels of cortical analysis, incorporating much perceptual experience, but feed back to lower sensory areas. These feedback loops may drive oscillations in the alpha range, linked strongly with serial dependence. The discovery of top-down predictive perceptual processes is not new, but the new, more quantitative approach characterizing serial dependence promises to lead to a deeper understanding of predictive perceptual processes and their underlying neural mechanisms.

Crowding results from optimal integration of visual targets with contextual information.

Crowding is the inability to recognize an object in clutter, usually considered a fundamental low-level bottleneck to object recognition. Here we advance and test an alternative idea, that crowding, like predictive phenomena such as serial dependence, results from optimizing strategies that exploit redundancies in natural scenes. This notion leads to several testable predictions: crowding should be greatest for unreliable targets and reliable flankers; crowding-induced biases should be maximal when target and flankers have similar orientations, falling off for differences around 20°; flanker interference should be associated with higher precision in orientation judgements, leading to lower overall error rate; effects should be maximal when the orientation of the target is near that of the average of the flankers, rather than to that of individual flankers. Each of these predictions were supported, and could be simulated with ideal-observer models that maximize performance. The results suggest that while crowding can affect object recognition, it may be better understood not as a processing bottleneck, but as a consequence of efficient exploitation of the spatial redundancies of the natural world.

Numerosity perception is tuned to salient environmental features.

Numerosity perception is a key ability to guide behavior. However, current models propose that number units encode an abstract representation of numerosity regardless of the non-numerical attributes of the stimuli, suggesting rather coarse environmental tuning. Here we investigated whether numerosity systems spontaneously adapt to all visible items, or to subsets segregated by salient attributes such as color or pitch. We measured perceived numerosity after participants adapted to highly numerous stimuli with color either matched to or different from the test. Matched colors caused a 25% underestimation of numerosity, while different colors had virtually no effect. This was true both for physically different colors, and for the same colors perceived as different, via a color-assimilation illusion. A similar result occurred in the acoustic domain, where adaptation magnitude was halved when the adaptor and test differed in pitch. Taken together, our results support the idea that numerosity perception is selectively tuned to salient environmental attributes.

Ideal observer analysis for continuous tracking experiments.

Continuous tracking is a newly developed technique that allows fast and efficient data acquisition by asking participants to "track" a stimulus varying in some property (usually position in space). Tracking is a promising paradigm for the investigation of dynamic features of perception and could be particularly well suited for testing ecologically relevant situations difficult to study with classical psychophysical paradigms. The high rate of data collection may be useful in studies on clinical populations and children, who are unable to undergo long testing sessions. In this study, we designed tracking experiments with two novel stimulus features, numerosity and size, proving the feasibility of the technique outside standard object tracking. We went on to develop an ideal observer model that characterizes the results in terms of efficiency of conversion of stimulus strength into responses, and identification of early and late noise sources. Our ideal observer closely modeled results from human participants, providing a generalized framework for the interpretation of tracking data. The proposed model allows to use the tracking paradigm in various perceptual domains, and to study the divergence of human participants from ideal behavior.

Visual Duration but Not Numerosity Is Distorted While Running.

There is increasing evidence that action and perception interact in the processing of magnitudes such as duration and numerosity. Sustained physical exercise (such as running or cycling) increases the apparent duration of visual stimuli presented during the activity. However, the effect of exercise on numerosity perception has not yet been investigated. Here, we asked participants to make either a temporal or a numerical judgment by comparing the duration or numerosity of standard stimuli displayed at rest with those presented while running. The results support previous reports in showing that physical activity significantly expands perceived duration; however, it had no effect on perceived numerosity. Furthermore, the distortions of the perceived durations vanished soon after the running session, making it unlikely that physiological factors such as heart rate underlie the temporal distortion. Taken together, these results suggest a domain-selective influence of the motor system on the perception of time, rather than a general effect on magnitude.

Grouping-Induced Numerosity Biases Vary with Autistic-Like Personality Traits.

Individuals with autism spectrum disorder are thought to have a more local than global perceptual style. We used a novel paradigm to investigate how grouping-induced response biases in numerosity judgments depend on autistic-like personality traits in neurotypical adults. Participants judged the numerosity of clouds of dot-pairs connected by thin lines, known to cause underestimation of numerosity. The underestimation bias correlated strongly with autism-spectrum quotient (r = 0.72, Bayes factor > 100), being weaker for participants with high autistic traits. As connecting dots probably activates global grouping mechanisms, causing dot-pairs to be processed as an integrated whole rather than as individual dots, the results suggest that these grouping mechanisms may be weaker in individuals self-reporting high levels of autistic-like traits.

The dynamics of grouping-induced biases in apparent numerosity revealed by a continuous tracking technique.

Connecting pairs of items causes robust underestimation of the numerosity of an ensemble, presumably by invoking grouping mechanisms. Here we asked whether this underestimation in numerosity judgments could be revealed and further explored by continuous tracking, a newly developed technique that allows for fast and efficient data acquisition and monitors the dynamics of the responses. Participants continuously reproduced the perceived numerosity of a cloud of dots by moving a cursor along a number line, while the number of dots and the proportion connected by lines varied over time following two independent random walks. The technique was robust and efficient, and correlated well with results obtained with a standard psychophysics task. Connecting objects with lines caused an underestimation of approximately 15% during tracking, agreeing with previous studies. The response to the lines was slower than the response to the physical numerosity, with a delay of approximately 150 ms, suggesting that this extra time is necessary for processing the grouping effect.

Objective pupillometry shows that perceptual styles covary with autistic-like personality traits.

We measured the modulation of pupil size (in constant lighting) elicited by observing transparent surfaces of black and white moving dots, perceived as a cylinder rotating about its vertical axis. The direction of rotation was swapped periodically by flipping stereo-depth of the two surfaces. Pupil size modulated in synchrony with the changes in front-surface color (dilating when black). The magnitude of pupillary modulation was larger for human participants with higher Autism-Spectrum Quotient (AQ), consistent with a local perceptual style, with attention focused on the front surface. The modulation with surface color, and its correlation with AQ, was equally strong when participants passively viewed the stimulus. No other indicator, including involuntary pursuit eye movements, covaried with AQ. These results reinforce our previous report with a similar bistable stimulus (Turi, Burr, & Binda, 2018), and go on to show that bistable illusory motion is not necessary for the effect, or its dependence on AQ.

Location- and object-based attention enhance number estimation.

Humans and non-humans can extract an estimate of the number of items in a collection very rapidly, raising the question of whether attention is necessary for this process. Visual attention operates in various modes, showing selectivity both to spatial location and to objects. Here, we tested whether each form of attention can enhance number estimation, by measuring whether presenting a visual cue to increase attentional engagement will lead to a more accurate and precise representation of number, both when attention is directed to location and when it is directed to objects. Results revealed that enumeration of a collection of dots in the location previously cued led to faster, more precise, and more accurate judgments than enumeration in un-cued locations, and a similar benefit was seen when the cue and collection appeared on the same object. This work shows that like many other perceptual tasks, numerical estimation may be enhanced by the spread of active attention inside a pre-cued object.

Higher attentional costs for numerosity estimation at high densities.

Humans can estimate numerosity over a large range, but the precision with which they do so varies considerably over that range. For very small sets, within the subitizing range of up to about four items, estimation is rapid and errorless. For intermediate numerosities, errors vary directly with the numerosity, following Weber's law, but for very high numerosities, with very dense patterns, thresholds continue to rise with the square root of numerosity. This suggests that three different mechanisms operate over the number range. In this study we provide further evidence for three distinct numerosity mechanisms, by studying their dependence on attentional resources. We measured discrimination thresholds over a wide range of numerosities, while manipulating attentional load with both visual and auditory dual tasks. The results show that attentional effects on thresholds vary over the number range. Both visual and auditory attentional loads strongly affect subitizing, much more than for larger numerosities. Attentional costs remain stable over the estimation range, then rise again for very dense patterns. These results reinforce the idea that numerosity is processed by three separates but probably overlapping systems.

Different reaction-times for subitizing, estimation, and texture.

Humans can estimate and encode numerosity over a large range, from very few items to several hundreds. Two distinct mechanisms have been proposed: subitizing, for numbers up to four and estimation for larger numerosities. We have recently extended this idea by suggesting that for very densely packed arrays, when items are less segregable, a third "texture" mechanism comes into play. In this study, we provide further evidence for the existence of a third regime for numerosity. Reaction times were very low in the subitizing range, rising rapidly for numerosities greater than four. However, for tightly packed displays of very high numerosities, reaction times became faster. These results reinforce the idea of three regimes in the processing of numerosity, subitizing, estimation, and texture.

Time dilation effect in an active observer and virtual environment requires apparent motion: No dilation for retinal- or world-motion alone.

It is known that moving visual stimuli are perceived to last longer than stationary stimuli with the same physical duration (Kanai, Paffen, Hogendoorn, & Verstraten, 2006), and that motor actions (Tomassini & Morrone, 2016) and eye movements (Morrone, Ross, & Burr, 2005) can alter perceived duration. In the present work, we investigated the contributions of stimulus motion and self-motion to perceived duration while observers stood or walked in a virtual reality environment. Using a visual temporal reproduction task, we independently manipulated both the participants' motion (stationary or walking) and the stimulus motion (retinal stationary, real-world stationary and negative double velocity). When the observers were standing still, drifting gratings were perceived as lasting longer than duration-matched static gratings. Interestingly, we did not see any time distortion when observers were walking, neither when the gratings were kept stationary relative to the observer's point of view (i.e., no retinal motion) nor when they were stationary in the external world (i.e., producing the same retinal velocity as the walking condition with stationary grating). Self-motion caused significant dilation in perceived duration only when the gratings were moving at double speed, opposite to the observers' walking direction. Consistent with previous work (Fornaciai, Arrighi, & Burr, 2016), this suggests that the system is able to suppress self-generated motion to enhance external motion, which would have ecological benefits, for example, for threat detection while navigating through the environment.

Perceptual Oscillation of Audiovisual Time Simultaneity.

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 ∼500 ms before and persisting ∼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.

Pupillometry reveals perceptual differences that are tightly linked to autistic traits in typical adults.

The pupil is primarily regulated by prevailing light levels but is also modulated by perceptual and attentional factors. We measured pupil-size in typical adult humans viewing a bistable-rotating cylinder, constructed so the luminance of the front surface changes with perceived direction of rotation. In some participants, pupil diameter oscillated in phase with the ambiguous perception, more dilated when the black surface was in front. Importantly, the magnitude of oscillation predicts autistic traits of participants, assessed by the Autism-Spectrum Quotient AQ. Further experiments suggest that these results are driven by differences in perceptual styles: high AQ participants focus on the front surface of the rotating cylinder, while those with low AQ distribute attention to both surfaces in a more global, holistic style. This is the first evidence that pupillometry reliably tracks inter-individual differences in perceptual styles; it does so quickly and objectively, without interfering with spontaneous perceptual strategies.

A shared numerical representation for action and perception.

Humans and other species have perceptual mechanisms dedicated to estimating approximate quantity: a sense of number. Here we show a clear interaction between self-produced actions and the perceived numerosity of subsequent visual stimuli. A short period of rapid finger-tapping (without sensory feedback) caused subjects to underestimate the number of visual stimuli presented near the tapping region; and a period of slow tapping caused overestimation. The distortions occurred both for stimuli presented sequentially (series of flashes) and simultaneously (clouds of dots); both for magnitude estimation and forced-choice comparison. The adaptation was spatially selective, primarily in external, real-world coordinates. Our results sit well with studies reporting links between perception and action, showing that vision and action share mechanisms that encode numbers: a generalized number sense, which estimates the number of self-generated as well as external events.