Visual attention: The past 25 years

What determines what we see? In contrast to the traditional "modular" understanding of perception, according to which visual processing is encapsulated from higher-level cognition, a tidal wave of recent research alleges that states such as beliefs, desires, emotions, motivations, intentions, and linguistic representations exert direct top-down influences on what we see. There is a growing consensus that such effects are ubiquitous, and that the distinction between perception and cognition may itself be unsustainable. We argue otherwise: none of these hundreds of studies - either individually or collectively - provide compelling evidence for true top-down effects on perception, or "cognitive penetrability". In particular, and despite their variety, we suggest that these studies all fall prey to only a handful of pitfalls. And whereas abstract theoretical challenges have failed to resolve this debate in the past, our presentation of these pitfalls is empirically anchored: in each case, we show not only how certain studies could be susceptible to the pitfall (in principle), but how several alleged top-down effects actually are explained by the pitfall (in practice). Moreover, these pitfalls are perfectly general, with each applying to dozens of other top-down effects. We conclude by extracting the lessons provided by these pitfalls into a checklist that future work could use to convincingly demonstrate top-down effects on visual perception. The discovery of substantive top-down effects of cognition on perception would revolutionize our understanding of how the mind is organized; but without addressing these pitfalls, no such empirical report will license such exciting conclusions. Language: en

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Cognition does not affect perception: Evaluating the evidence for "top-down" effects.

Active inference provides a simple and neurobiologically plausible account of how action and perception are coupled in producing (Bayes) optimal behaviour. This can be seen most easily as minimising prediction error: we can either change our predictions to explain sensory input through perception. Alternatively, we can actively change sensory input to fulfil our predictions. In active inference, this action is mediated by classical reflex arcs that minimise proprioceptive prediction error created by descending proprioceptive predictions. However, this creates a conflict between action and perception; in that, self-generated movements require predictions to override the sensory evidence that one is not actually moving. However, ignoring sensory evidence means that externally generated sensations will not be perceived. Conversely, attending to (proprioceptive and somatosensory) sensations enables the detection of externally generated events but precludes generation of actions. This conflict can be resolved by attenuating the precision of sensory evidence during movement or, equivalently, attending away from the consequences of self-made acts. We propose that this Bayes optimal withdrawal of precise sensory evidence during movement is the cause of psychophysical sensory attenuation. Furthermore, it explains the force-matching illusion and reproduces empirical results almost exactly. Finally, if attenuation is removed, the force-matching illusion disappears and false (delusional) inferences about agency emerge. This is important, given the negative correlation between sensory attenuation and delusional beliefs in normal subjects--and the reduction in the magnitude of the illusion in schizophrenia. Active inference therefore links the neuromodulatory optimisation of precision to sensory attenuation and illusory phenomena during the attribution of agency in normal subjects. It also provides a functional account of deficits in syndromes characterised by false inference and impaired movement--like schizophrenia and Parkinsonism--syndromes that implicate abnormal modulatory neurotransmission.

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Active inference, sensory attenuation and illusions

The more we anticipate a response to a predictable stimulus, the faster we react. This empirical observation has been confirmed and quantified by many investigators suggesting that the processing of behaviorally relevant stimuli is facilitated by probability-based confidence of anticipation. However, the exact neural mechanisms underlying this phenomenon are largely unknown. Here we show that performance changes related to different levels of expectancy originate in dynamic modulation of delta oscillation phase. Our results obtained in rhythmic auditory target detection tasks indicated significant entrainment of the EEG delta rhythm to the onset of the target tones with increasing phase synchronization at higher levels of predictability. Reaction times correlated with the phase of the delta band oscillation at target onset. The fastest reactions occurred during the delta phase that most commonly coincided with the target event in the high expectancy conditions. These results suggest that low-frequency oscillations play a functional role in human anticipatory mechanisms, presumably by modulating synchronized rhythmic fluctuations in the excitability of large neuronal populations and by facilitating efficient task-related neuronal communication among brain areas responsible for sensory processing and response execution.

Phase entrainment of human delta oscillations can mediate the effects of expectation on reaction speed (Journal of Neuroscience (2010) (13578-13585))

The size of the pupil has a large effect on visual function, and pupil size depends mainly on the adapting luminance, modulated by other factors. Over the last century, a number of formulas have been proposed to describe this dependence. Here we review seven published formulas and develop a new unified formula that incorporates the effects of luminance, size of the adapting field, age of the observer, and whether one or both eyes are adapted. We provide interactive demonstrations and software implementations of the unified formula.

A Unified Formula For Light-adapted Pupil Size

Optical blur in the peripheral retina is known to be highly anisotropic due to nonrotationally symmetric wavefront aberrations such as astigmatism and coma. At the neural level, the visual system exhibits anisotropies in orientation sensitivity across the visual field. In the fovea, the visual system shows higher sensitivity for cardinal over diagonal orientations, which is referred to as the oblique effect. However, in the peripheral retina, the neural visual system becomes more sensitive to radially-oriented signals, a phenomenon known as the meridional effect. Here, we examined the relative contributions of optics and neural processing to the meridional effect in 10 participants at 0°, 10°, and 20° in the temporal retina. Optical anisotropy was quantified by measuring the eye's habitual wavefront aberrations. Alternatively, neural anisotropy was evaluated by measuring contrast sensitivity (at 2 and 4 cyc/deg) while correcting the eye's aberrations with an adaptive optics vision simulator, thus bypassing any optical factors. As eccentricity increased, optical and neural anisotropy increased in magnitude. The average ratio of horizontal to vertical optical MTF (at 2 and 4 cyc/deg) at 0°, 10°, and 20° was 0.96 ± 0.14, 1.41 ± 0.54 and 2.15 ± 1.38, respectively. Similarly, the average ratio of horizontal to vertical contrast sensitivity with full optical correction at 0°, 10°, and 20° was 0.99 ± 0.15, 1.28 ± 0.28 and 1.75 ± 0.80, respectively. These results indicate that the neural system's orientation sensitivity coincides with habitual blur orientation. These findings support the neural origin of the meridional effect and raise important questions regarding the role of peripheral anisotropic optical quality in developing the meridional effect and emmetropization.

Optical and neural anisotropy in peripheral vision.

Saccade Preparation Reshapes Sensory Tuning

It is well established that attention improves performance on many visual tasks. However, for more than 100 years, psychologists, philosophers, and neurophysiologists have debated its phenomenology-whether attention actually changes one's subjective experience. Here, we show that it is possible to objectively and quantitatively investigate the effects of attention on subjective experience. First, we review evidence showing that attention alters the appearance of many static and dynamic basic visual dimensions, which mediate changes in appearance of higher-level perceptual aspects. Then, we summarize current views on how attention alters appearance. These findings have implications for our understanding of perception and attention, illustrating that attention affects not only how we perform in visual tasks, but actually alters our experience of the visual world.

Spatial attention alters visual appearance.

Voluntary covert attention selects relevant sensory information for prioritized processing. The behavioral and neural consequences of such selection have been extensively documented, but its phenomenology has received little empirical investigation. Involuntary attention increases perceived spatial frequency (Gobell & Carrasco, 2005), but involuntary attention can differ from voluntary attention in its effects on performance in tasks mediated by spatial resolution (Yeshurun, Montagna, & Carrasco, 2008). Therefore, we ask whether voluntary attention affects the subjective appearance of spatial frequency—a fundamental dimension of visual perception underlying spatial resolution. We used a demanding rapid serial visual presentation task to direct voluntary attention and measured perceived spatial frequency at the attended and unattended locations. Attention increased the perceived spatial frequency of suprathreshold stimuli and also improved performance on a concurrent orientation discrimination task. In the control experiment, we ruled out response bias as an alternative account by using a lengthened interstimulus interval, which allows observers to disengage attention from the cued location. In contrast to the main experiment, the observers showed neither increased perceived spatial frequency nor improved orientation discrimination at the attended location. Thus, this study establishes that voluntary attention increases perceived spatial frequency. This phenomenological consequence links behavioral and neurophysiological studies on the effects of attention.

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Voluntary attention increases perceived spatial frequency

We summarize and discuss a series of psychophysical studies on the effects of spatial covert attention on spatial resolution, our ability to discriminate fine patterns. Heightened resolution is beneficial in most, but not all, visual tasks. We show how endogenous attention (voluntary, goal driven) and exogenous attention (involuntary, stimulus driven) affect performance on a variety of tasks mediated by spatial resolution, such as visual search, crowding, acuity, and texture segmentation. Exogenous attention is an automatic mechanism that increases resolution regardless of whether it helps or hinders performance. In contrast, endogenous attention flexibly adjusts resolution to optimize performance according to task demands. We illustrate how psychophysical studies can reveal the underlying mechanisms of these effects and allow us to draw linking hypotheses with known neurophysiological effects of attention.

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Antoine Barbot

Papers

Optical and neural anisotropy in peripheral vision.

Saccade Preparation Reshapes Sensory Tuning

Spatial attention alters visual appearance.

Voluntary attention increases perceived spatial frequency

How attention affects spatial resolution