Peripheral vision is mainly for looking rather than seeing.

Vision includes looking and seeing. Looking, mainly via gaze shifts, selects a fraction of visual input information for passage through the brain's information bottleneck. The selected input is placed within the attentional spotlight, typically in the central visual field. Seeing decodes, i.e., recognizes and discriminates, the selected inputs. Hence, peripheral vision should be mainly devoted to looking, in particular, deciding where to shift the gaze. Looking is often guided exogenously by a saliency map created by the primary visual cortex (V1), and can be effective with no seeing and limited awareness. In seeing, peripheral vision not only suffers from poor spatial resolution, but is also subject to crowding and is more vulnerable to illusions by misleading, ambiguous, and impoverished visual inputs. Central vision, mainly for seeing, enjoys the top-down feedback that aids seeing in light of the bottleneck which is hypothesized to starts from V1 to higher areas. This feedback queries for additional information from lower visual cortical areas such as V1 for ongoing recognition. Peripheral vision is deficient in this feedback according to the Central-peripheral Dichotomy (CPD) theory. The saccades engendered by peripheral vision allows looking to combine with seeing to give human observers the impression of seeing the whole scene clearly despite inattentional blindness.

Eye movement evidence for the V1 Saliency Hypothesis and the Central-peripheral Dichotomy theory in an anomalous visual search task.

Typically, searching for a target among uniformly tilted non-targets is easier when this target is perpendicular, rather than parallel, to the non-targets. The V1 Saliency Hypothesis (V1SH) - that V1 creates a saliency map to guide attention exogenously - predicts exactly the opposite in a special case: each target or non-target is a pair of equally-sized disks, a homo-pair of two disks of the same color, black or white, or a hetero-pair of two disks of the opposite color; the inter-disk displacement defines its orientation. This prediction - parallel advantage - was supported by the finding that parallel targets require shorter reaction times (RTs) to report targets' locations. Furthermore, it is stronger for targets further from the center of search images, as predicted by the Central-peripheral Dichotomy (CPD) theory entailing that saliency effects are stronger in peripheral than in central vision. However, the parallel advantage could arise from a shorter time required to recognize - rather than to shift attention to - the parallel target. By gaze tracking, the present study confirms that the parallel advantage is solely due to the RTs for the gaze to reach the target. Furthermore, when the gaze is sufficiently far from the target during search, saccade to a parallel, rather than perpendicular, target is more likely, demonstrating the Central-peripheral Dichotomy more directly. Parallel advantage is stronger among observers encouraged to let their search be guided by spontaneous gaze shifts, which are presumably guided by bottom-up saliency rather than top-down factors.

Peripheral and central sensation: multisensory orienting and recognition across species.

Attentional bottlenecks force animals to deeply process only a selected fraction of sensory inputs. This motivates a unifying central-peripheral dichotomy (CPD), which separates multisensory processing into functionally defined central and peripheral senses. Peripheral senses (e.g., human audition and peripheral vision) select a fraction of the sensory inputs by orienting animals' attention; central senses (e.g., human foveal vision) allow animals to recognize the selected inputs. Originally used to understand human vision, CPD can be applied to multisensory processes across species. I first describe key characteristics of central and peripheral senses, such as the degree of top-down feedback and density of sensory receptors, and then show CPD as a framework to link ecological, behavioral, neurophysiological, and anatomical data and produce falsifiable predictions.

Li and Atick's theory of efficient binocular coding: A tutorial and mini-review.

Li and Atick (Network: Computation in Neural Systems 5 (1994) 157-174) presented a theory of efficient binocular encoding that explains a number of experimental findings. A binocular neuron is conventionally described in terms of two channels: the left and right eyes. Li and Atick's theory instead describes the neuron in terms of two alternative channels: the binocular sum and difference. The advantage of the latter description is that, unlike the left and right eye channels, the summation and differencing channels are usually uncorrelated; this means that each channel can be optimised independently of the other. The theory shows how to derive optimal receptive fields for the binocular summation and differencing channels; from these, it is easy to derive the neuron's optimal left and right eye receptive fields. The functional reality of the summation and differencing channels is demonstrated by a series of adaptation studies that confirm some counterintuitive predictions of the theory. Here we provide an accessible account of the theory, and review the evidence supporting it.

The central-peripheral dichotomy and metacontrast masking.

According to the central-peripheral dichotomy (CPD), feedback from higher to lower cortical areas along the visual pathway to aid recognition is weaker in the more peripheral visual field. Metacontrast masking is predominantly a reduced visibility of a brief target by a brief and spatially adjacent mask when the mask succeeds rather than precedes or coincides with the target. If this masking works mainly by interfering with the feedback mechanisms for target recognition, then, by the CPD, this masking should be weaker at more peripheral visual locations. We extended the metacontrast masking at fovea by Enns and Di Lollo to visual field eccentricities 1∘, 3∘, and 9∘. Relative to the target's onset, the mask appeared at a stimulus onset asynchrony (SOA) of -50, 0, 50, 92, or 142 milliseconds (ms). Enlarged stimuli were used for larger eccentricities to equalize target discrimination performance across eccentricities as best as possible for zero SOA and when SOA was too long for substantial masking. At each eccentricity, the masking was weakest at 0 or -50 ms SOA, strongest at 50 ms SOA, and weakened with larger (positive) SOAs. Consistent with the CPD, larger eccentricities presented weaker maskings at all nonzero, and particularly the positive, SOAs.

Parallel Advantage: Further Evidence for Bottom-up Saliency Computation by Human Primary Visual Cortex.

Finding a target among uniformly oriented non-targets is typically faster when this target is perpendicular, rather than parallel, to the non-targets. The V1 Saliency Hypothesis (V1SH), that neurons in the primary visual cortex (V1) signal saliency for exogenous attentional attraction, predicts exactly the opposite in a special case: each target or non-target comprises two equally sized disks displaced from each other by 1.2 disk diameters center-to-center along a line defining its orientation. A target has two white or two black disks. Each non-target has one white disk and one black disk, and thus, unlike the target, activates V1 neurons less when its orientation is parallel rather than perpendicular to the neurons' preferred orientations. When the target is parallel, rather than perpendicular, to the uniformly oriented non-targets, the target's evoked V1 response escapes V1's iso-orientation surround suppression, making the target more salient. I present behavioral observations confirming this prediction.

Contrast-reversed binocular dot-pairs in random-dot stereograms for depth perception in central visual field: Probing the dynamics of feedforward-feedback processes in visual inference.

In a random-dot stereogram (RDS), the spatial disparities between the interocularly corresponding black and white random dots determine the depths of object surfaces. If a black dot in one monocular image corresponds to a white dot in the other, disparity-tuned neurons in primary visual cortex (V1) respond as if their preferred disparities become non-preferred and vice versa, reversing the disparity sign reported to higher visual areas. Reversed depth is perceptible in the peripheral but not the central visual field. This study demonstrates that, in central vision, adding contrast-reversed dots to a noisy RDS (containing the normal contrast-matched dots) can augment or degrade depth perception. Augmentation occurs when the reversed depth signals are congruent with the normal depth signals to report the same disparity sign, and occurs regardless of the viewing duration. Degradation occurs when the reversed and normal depth signals are incongruent with each other and when the RDS is viewed briefly. These phenomena reflect the Feedforward-Feedback-Verify-and-reWeight (FFVW) process for visual inference in central vision, and are consistent with the central-peripheral dichotomy that central vision has a stronger top-down feedback from higher to lower brain areas to disambiguate noisy and ambiguous inputs from V1. When a RDS is viewed too briefly for feedback, augmentation and degradation work by adding the reversed depth signals from contrast-reversed dots to the feedforward, normal, depth signals. With a sufficiently long viewing duration, the feedback vetoes incongruent reversed depth signals and amends or completes the imperfect, but congruent, reversed depth signals by analysis-by-synthesis computation.

The Flip Tilt Illusion: Visible in Peripheral Vision as Predicted by the Central-Peripheral Dichotomy.

Consider a gray field comprising pairs of vertically aligned dots; in each pair, one dot is white the other black. When viewed in a peripheral visual field, these pairs appear horizontally aligned. By the Central-Peripheral Dichotomy, this flip tilt illusion arises because top-down feedback from higher to lower visual cortical areas is too weak or absent in the periphery to veto confounded feedforward signals from the primary visual cortex (V1). The white and black dots in each pair activate, respectively, on and off subfields of V1 neural receptive fields. However, the sub-fields' orientations, and the preferred orientations, of the most activated neurons are orthogonal to the dot alignment. Hence, V1 reports the flip tilt to higher visual areas. Top-down feedback vetoes such misleading reports, but only in the central visual field.

Artificial and Natural Intelligence: From Invention to Discovery.

An international group of researchers met in November 2019 in Beijing to explore the intersection of neuroscience and AI. The aim was to offer a fertile ground for stimulating discussions and ideas, including issues such as policy making and the future of neuroscience and AI across the globe.

Face perception inherits low-level binocular adaptation.

In previous work (May & Zhaoping, 2016; May, Zhaoping, & Hibbard, 2012), we have provided evidence that the visual system efficiently encodes binocular information using separately adaptable binocular summation and differencing channels. In that work, binocular test stimuli delivered different grating patterns to the two binocular channels; selective adaptation of one of the binocular channels made participants more likely to see the other channel's grating pattern. In the current study, we extend this paradigm to face perception. Our test stimuli delivered different face images to the two binocular channels, and we found that selective adaptation of one binocular channel biased the observer to perceive the other channel's face image. We show that the perceived identity, gender, emotional expression, or direction of 3-D rotation of a facial test image can be influenced by pre-exposure to binocular random-noise patterns that contain no meaningful spatial structure. Our results provide compelling evidence that face-processing mechanisms can inherit adaptation from low-level sites. Our adaptation paradigm targets the low-level mechanisms in such a way that any response bias or inadvertent adaptation of high-level mechanisms selective for face categories would reduce, rather than produce, the measured effects of adaptation.

A new framework for understanding vision from the perspective of the primary visual cortex.

Visual attention selects only a tiny fraction of visual input information for further processing. Selection starts in the primary visual cortex (V1), which creates a bottom-up saliency map to guide the fovea to selected visual locations via gaze shifts. This motivates a new framework that views vision as consisting of encoding, selection, and decoding stages, placing selection on center stage. It suggests a massive loss of non-selected information from V1 downstream along the visual pathway. Hence, feedback from downstream visual cortical areas to V1 for better decoding (recognition), through analysis-by-synthesis, should query for additional information and be mainly directed at the foveal region. Accordingly, non-foveal vision is not only poorer in spatial resolution, but also more susceptible to many illusions.

Exploring the neural correlates of the reversed letter effect: Evidence from left and right parietal patients.

To investigate the hemispheric lateralization of attentional processes during visual search tasks depending on the stimulus material embedding the target, twelve patients with unilateral left (n = 7) or right (n = 5) parietal lesions and 20 age and education matched healthy controls (HC) were recruited. We used a visual search task for a uniquely tilted oblique bar embedded in an object shape 'N' or in its mirror reversal 'И'. The accuracy and the averaged reaction times (RTs) in each stimulus type ('N' or 'И') were analysed. HC presented significantly longer RTs when the target bar was embedded in 'N' among its mirror reversed 'И' (p < .05). This "reversed letter effect" was also found in the right parietal patients (p < .001), while no evidence of a reversed letter effect was found in the left parietal patients.

Bottom-up saliency and top-down learning in the primary visual cortex of monkeys.

Early sensory cortex is better known for representing sensory inputs but less for the effect of its responses on behavior. Here we explore the behavioral correlates of neuronal responses in primary visual cortex (V1) in a task to detect a uniquely oriented bar-the orientation singleton-in a background of uniformly oriented bars. This singleton is salient or inconspicuous when the orientation contrast between the singleton and background bars is sufficiently large or small, respectively. Using implanted microelectrodes, we measured V1 activities while monkeys were trained to quickly saccade to the singleton. A neuron's responses to the singleton within its receptive field had an early and a late component, both increased with the orientation contrast. The early component started from the outset of neuronal responses; it remained unchanged before and after training on the singleton detection. The late component started ∼40 ms after the early one; it emerged and evolved with practicing the detection task. Training increased the behavioral accuracy and speed of singleton detection and increased the amount of information in the late response component about a singleton's presence or absence. Furthermore, for a given singleton, faster detection performance was associated with higher V1 responses; training increased this behavioral-neural correlate in the early V1 responses but decreased it in the late V1 responses. Therefore, V1's early responses are directly linked with behavior and represent the bottom-up saliency signals. Learning strengthens this link, likely serving as the basis for making the detection task more reflexive and less top-down driven.

Reversed Depth in Anticorrelated Random-Dot Stereograms and the Central-Peripheral Difference in Visual Inference.

In a random-dot stereogram, the percept of object surfaces in a three-dimensional scene is generated by images presented to left and right eyes that comprise interocularly corresponding random black and white dots. The spatial disparities between the corresponding dots determine the depths of object surfaces. If the dots are anticorrelated, such that a black dot in one monocular image corresponds to a white dot in the other, disparity-tuned neurons in the primary visual cortex (V1) respond as if their preferred disparities become nonpreferred and vice versa, thereby reversing the disparity signs reported to higher visual areas. Typically, when viewing anticorrelated random-dot stereograms presented in the central visual field, humans have great difficulty perceiving the reversed depth or indeed any coherent depth at all. We report that the reversed depth is more easily perceived in the peripheral visual field, supporting a recently proposed central-peripheral dichotomy in the way that feedback from higher to lower visual cortical areas implements visual inference.

Ocularity Feature Contrast Attracts Attention Exogenously.

An eye-of-origin singleton, e.g., a bar shown to the left eye among many other bars shown to the right eye, can capture attention and gaze exogenously or reflexively, even when it appears identical to other visual input items in the scene and when the eye-of-origin feature is irrelevant to the observer's task. Defining saliency as the strength of exogenous attraction to attention, we say that this eye-of-origin singleton, or its visual location, is salient. Defining the ocularity of a visual input item as the relative difference between its left-eye input and its right-eye input, this paper shows the general case that an ocularity singleton is also salient. For example, a binocular input item among monocular input items is salient, so is a left-eye-dominant input item (e.g., a bar with a higher input contrast to the left eye than to the right eye) among right-eye-dominant items. Saliency by unique input ocularity is analogous to saliency by unique input colour (e.g., a red item among green ones), as colour is determined by the relative difference(s) between visual inputs to different photoreceptor cones. Just as a smaller colour difference between a colour singleton and background items makes this singleton less salient, so does a smaller ocularity difference between an ocularity singleton and background items. While a salient colour difference is highly visible, a salient ocularity difference is often perceptually invisible in some cases and discouraging gaze shifts towards it in other cases, making its behavioural manifestation not as apparent. Saliency by ocularity contrast provides another support to the idea that the primary visual cortex creates a bottom-up saliency map to guide attention exogenously.

Feedback from higher to lower visual areas for visual recognition may be weaker in the periphery: Glimpses from the perception of brief dichoptic stimuli.

Eye movements bring attended visual inputs to the center of vision for further processing. Thus, central and peripheral vision should have different functional roles. Here, we use observations of visual perception under dichoptic stimuli to infer that there is a difference in the top-down feedback from higher brain centers to primary visual cortex. Visual stimuli to the two eyes were designed such that the sum and difference of the binocular input from the two eyes have the form of two different gratings. These gratings differed in their motion direction, tilt direction, or color, and duly evoked ambiguous percepts for the corresponding feature. Observers were more likely to perceive the feature in the binocular summation rather than the difference channel. However, this perceptual bias towards the binocular summation signal was weaker or absent in peripheral vision, even when central and peripheral vision showed no difference in contrast sensitivity to the binocular summation signal relative to that to the binocular difference signal. We propose that this bias can arise from top-down feedback as part of an analysis-by-synthesis computation. The feedback is of the input predicted using prior information by the upper level perceptual hypothesis about the visual scene; the hypothesis is verified by comparing the feedback with the actual visual input. We illustrate this process using a conceptual circuit model. In this framework, a bias towards binocular summation can arise from the prior knowledge that inputs are usually correlated between the two eyes. Accordingly, a weaker bias in the periphery implies that the top-down feedback is weaker there. Testable experimental predictions are presented and discussed.

The evaluation of structural remodeling of the left atrium in elderly patients with non-valvular atrial fibrillation / 中华老年医学杂志

Objective To assess the difference in left atrial properties between elderly and younger control subjects and the role of left atrium remodeling in patients of different ages with atrial fibrillation.Methods A total of 194 patients with non-valvular atrial fibrillation were enrolled from September 2014 to June 2016.Based on age,patients were divided into an elderly group (≥60 years,n=129) and a younger group (＜60 years,n=65).We evaluated remodeling parameters for the left atrium using an Ultrasound Cardiography (UCG) system in 125 elder subjects,together with 64 control subjects.All remodeling parameters were recorded,including left atrial diameter (LAD),left atrial square (LAS),left atrial pressure (LAP) and left ventricular end-diastolic dimension (LVEDD).Results The elderly group had more female patients and more patients with persistent atrial fibrillation.Meanwhile,scores of CHA2DS2Vsc and levels of N-terminal pro-brain natriuretic peptide (NT-ProBNP) were significantly increased in elderly patients (both P＜0.05).Moreover,the elderly group was associated with increased values of LAD and LAS[(39.1±4.4)mm vs.(37.1±5.3)mm,P＜0.01;and(23.3±4.5)cm2 vs.(21.4±4.8)cm2,P＜0.01;respectively],compared with those in the control group.Spearman's correlation analysis showed that LAD,LAS and LAP were all markedly related to age (r=0.213,P＜0.05;r=-0.175,P＜0.05;r=0.170,P＜0.05;respectively),persistent onset of atrial fibrillation (r=0.401,P＜0.05;r=0.446,P＜0.05;r=0.160,P＜0.05;respectively),and impaired heart function,measured by left ventricular ejection fraction (r=-0.4371,P＜0.05;r=-0.403,P＜0.05;r=-0.364,P＜0.05;respectively) and NT-ProBNP (r=0.485,P＜0.01;r=0.483,P＜ 0.01;r =0.293,P＜ 0.01;respectively).Conclusions Left atrial remodeling properties measured by the UCG system in the elderly with non-valvular atrial fibrillation are more serious than those in mid-aged and young subjects.As a convenient and accurate assessment of remodeling parameters,the UCG system is an excellent option for measuring left atrial remodeling in the elderly population.

From the optic tectum to the primary visual cortex: migration through evolution of the saliency map for exogenous attentional guidance.

Recent data have supported the hypothesis that, in primates, the primary visual cortex (V1) creates a saliency map from visual input. The exogenous guidance of attention is then realized by means of monosynaptic projections to the superior colliculus, which can select the most salient location as the target of a gaze shift. V1 is less prominent, or is even absent in lower vertebrates such as fish; whereas the superior colliculus, called optic tectum in lower vertebrates, also receives retinal input. I review the literature and propose that the saliency map has migrated from the tectum to V1 over evolution. In addition, attentional benefits manifested as cueing effects in humans should also be present in lower vertebrates.

Efficient Coding Theory Predicts a Tilt Aftereffect from Viewing Untilted Patterns.

The brain is bombarded with a continuous stream of sensory information, but biological limitations on the data-transmission rate require this information to be encoded very efficiently [1]. Li and Atick [2] proposed that the two eyes' signals are coded efficiently in the brain using mutually decorrelated binocular summation and differencing channels; when a channel is strongly stimulated by the visual input, such that sensory noise is negligible, the channel should undergo temporary desensitization (known as adaptation). To date, the evidence for this theory has been limited [3, 4], and the binocular differencing channel is missing from many models of binocular integration [5-10]. Li and Atick's theory makes the remarkable prediction that perceived direction of tilt (clockwise or counterclockwise) of a test pattern can be controlled by pre-exposing observers to visual adaptation patterns that are untilted or even have no orientation signal. Here, we confirm this prediction. Each test pattern consisted of different images presented to the two eyes such that the binocular summation and difference signals were tilted in opposite directions, to give ambiguous information about tilt; by selectively desensitizing one or other of the binocular channels using untilted or non-oriented binocular adaptation patterns, we controlled the perceived tilt of the test pattern. Our results provide compelling evidence that the brain contains binocular summation and differencing channels that adapt to the prevailing binocular statistics.

Brains studying brains: look before you think in vision.

Using our own brains to study our brains is extraordinary. For example, in vision this makes us naturally blind to our own blindness, since our impression of seeing our world clearly is consistent with our ignorance of what we do not see. Our brain employs its 'conscious' part to reason and make logical deductions using familiar rules and past experience. However, human vision employs many 'subconscious' brain parts that follow rules alien to our intuition. Our blindness to our unknown unknowns and our presumptive intuitions easily lead us astray in asking and formulating theoretical questions, as witnessed in many unexpected and counter-intuitive difficulties and failures encountered by generations of scientists. We should therefore pay a more than usual amount of attention and respect to experimental data when studying our brain. I show that this can be productive by reviewing two vision theories that have provided testable predictions and surprising insights.