Grouping and Crowding Affect Target Appearance over Different Spatial Scales

Crowding is the impairment of peripheral target perception by nearby flankers. A number of recent studies have shown that crowding shares many features with grouping. Here, we investigate whether effects of crowding and grouping on target perception are related by asking whether they operate over the same spatial scale. A target letter T had two sets of flanking Ts of varying orientations. The first set was presented close to the target, yielding strong crowding. The second set was either close enough to cause crowding on their own or too far to cause crowding on their own. The Ts of the second set had the same orientation that either matched the target’s orientation (Grouped condition) or not (Ungrouped condition). In Experiment 1, the Grouped flankers reduced crowding independently of their distance from the target, suggesting that grouping operated over larger distances than crowding. In Experiments 2 and 3 we found that grouping did not affect sensitivity but produced a strong bias to report that the grouped orientation was present at the target location whether or not it was. Finally, we investigated whether this bias was a response or perceptual bias, rejecting the former in favor of a perceptual grouping explanation. We suggest that the effect of grouping is to assimilate the target to the identity of surrounding flankers when they are all the same, and that this shape assimilation effect differs in its spatial scale from the integration effect of crowding.     

Crowding by invisible flankers

Background

Human object recognition degrades sharply as the target object moves from central vision into peripheral vision. In particular, one''s ability to recognize a peripheral target is severely impaired by the presence of flanking objects, a phenomenon known as visual crowding. Recent studies on how visual awareness of flanker existence influences crowding had shown mixed results. More importantly, it is not known whether conscious awareness of the existence of both the target and flankers are necessary for crowding to occur.

Methodology/Principal Findings

Here we show that crowding persists even when people are completely unaware of the flankers, which are rendered invisible through the continuous flash suppression technique. Contrast threshold for identifying the orientation of a grating pattern was elevated in the flanked condition, even when the subjects reported that they were unaware of the perceptually suppressed flankers. Moreover, we find that orientation-specific adaptation is attenuated by flankers even when both the target and flankers are invisible.

Conclusions

These findings complement the suggested correlation between crowding and visual awareness. What''s more, our results demonstrate that conscious awareness and attention are not prerequisite for crowding.     

目标和干扰子之间的拓扑性质差异可以削弱拥挤效应

"拥挤效应"被认为是外周视觉物体辨认过程中的一个重要瓶颈.它是指当目标被干扰子包围,在外周视野呈现时,观察者辨认目标的能力被大大削弱,尤其是当目标和干扰子之间存在某种相似性时.许多研究分别试图在不同层次上提出解释这一现象的机制.本文通过三个实验,使用了不同的视觉刺激图形的辨认任务(例如,三角形和箭头的朝向判断、数字和字母的辨认以及S形图形的朝向辨认),测量了目标和干扰子之间中心距离的阈值,结果一致地发现,当目标和干扰子之间存在拓扑性质差异(洞的个数差异)时,拥挤效应会显著降低,并且排除了目标和干扰子之间的主观相似性、形状和面积差异等可能的因素.从知觉组织的角度验证了当目标和干扰子之间存在拓扑性质差异时,拥挤效应会显著降低,这是首次发现的一个影响拥挤效应的新的维度.本文结果不仅为拥挤效应的机制提供了一个新的解释,也为大范围首先拓扑知觉在知觉物体形成中的作用提供了支持性证据.     

A Neurophysiologically Plausible Population Code Model for Feature Integration Explains Visual Crowding

An object in the peripheral visual field is more difficult to recognize when surrounded by other objects. This phenomenon is called “crowding”. Crowding places a fundamental constraint on human vision that limits performance on numerous tasks. It has been suggested that crowding results from spatial feature integration necessary for object recognition. However, in the absence of convincing models, this theory has remained controversial. Here, we present a quantitative and physiologically plausible model for spatial integration of orientation signals, based on the principles of population coding. Using simulations, we demonstrate that this model coherently accounts for fundamental properties of crowding, including critical spacing, “compulsory averaging”, and a foveal-peripheral anisotropy. Moreover, we show that the model predicts increased responses to correlated visual stimuli. Altogether, these results suggest that crowding has little immediate bearing on object recognition but is a by-product of a general, elementary integration mechanism in early vision aimed at improving signal quality.     

Visual crowding is correlated with awareness

Crowding by nearby features causes identification failures in the peripheral visual field. However, prominent visual features can sometimes fail to reach awareness, causing scenes to be incorrectly interpreted. Here we examine whether awareness of the flanking features is necessary for crowding to occur. Flankers that were physically present were rendered perceptually absent with adaptation-induced blindness. In a letter identification task, targets were presented unflanked or with up to four flanker letters. On each trial, observers reported both the number of letters they perceived and the identity of a target letter. This paradigm allowed trial-by-trial assessment of awareness and crowding and ensured that both targets and flankers were attended. Target-letter identification performance was correlated with the number of flanking letters that were perceived on a given trial, regardless of the number that were physically present. Our data demonstrate that crowding can be released when flanking elements at attended locations are suppressed from visual awareness.     

Zipf’s Law: Balancing Signal Usage Cost and Communication Efficiency

We propose a model that explains the reliable emergence of power laws (e.g., Zipf’s law) during the development of different human languages. The model incorporates the principle of least effort in communications, minimizing a combination of the information-theoretic communication inefficiency and direct signal cost. We prove a general relationship, for all optimal languages, between the signal cost distribution and the resulting distribution of signals. Zipf’s law then emerges for logarithmic signal cost distributions, which is the cost distribution expected for words constructed from letters or phonemes.     

视觉拥挤效应的神经机制

当出现在边缘视野的一个物体被周围其他物体包围时,视觉系统对它的识别会很困难,这种现象叫做视觉拥挤效应.研究拥挤效应既有利于理解人类进行客体识别的过程,也对治疗黄斑变性、弱视和阅读障碍等视觉病变有显著的临床意义.自拥挤效应被提出以来,对拥挤效应的特性、神经机制和影响因素等都做了深入地研究.本文将系统地综述拥挤效应的研究进展,包括其特性、现有的理论假设、计算模型、可能涉及的大脑区域以及近年来利用知觉学习消除拥挤效应的一些工作,最后对该领域的未来发展给出建议.尽管在这个领域已经获得了丰富的成果,但在许多问题上仍有争议,未来还需要更为巧妙的设计和精确的技术进一步解决这些问题.     

“Vision for Action” in Young Children Aligning Multi-Featured Objects: Development and Comparison with Nonhuman Primates

Effective vision for action and effective management of concurrent spatial relations underlie skillful manipulation of objects, including hand tools, in humans. Children’s performance in object insertion tasks (fitting tasks) provides one index of the striking changes in the development of vision for action in early life. Fitting tasks also tap children’s ability to work with more than one feature of an object concurrently. We examine young children’s performance on fitting tasks in two and three dimensions and compare their performance with the previously reported performance of adult individuals of two species of nonhuman primates on similar tasks. Two, three, and four year-old children routinely aligned a bar-shaped stick and a cross-shaped stick but had difficulty aligning a tomahawk-shaped stick to a matching cut-out. Two year-olds were especially challenged by the tomahawk. Three and four year-olds occasionally held the stick several inches above the surface, comparing the stick to the surface visually, while trying to align it. The findings suggest asynchronous development in the ability to use vision to achieve alignment and to work with two and three spatial features concurrently. Using vision to align objects precisely to other objects and managing more than one spatial relation between an object and a surface are already more elaborated in two year-old humans than in other primates. The human advantage in using hand tools derives in part from this fundamental difference in the relation between vision and action between humans and other primates.     

Sensory modulation of gait characteristics in human locomotion: A neuromusculoskeletal modeling study

The central nervous system of humans and other animals modulates spinal cord activity to achieve several locomotion behaviors. Previous neuromechanical models investigated the modulation of human gait changing selected parameters belonging to CPGs (Central Pattern Generators) feedforward oscillatory structures or to feedback reflex circuits. CPG-based models could replicate slow and fast walking by changing only the oscillation’s properties. On the other hand, reflex-based models could achieve different behaviors through optimizations of large dimensional parameter spaces. However, they could not effectively identify individual key reflex parameters responsible for gait characteristics’ modulation. This study investigates which reflex parameters modulate the gait characteristics through neuromechanical simulations. A recently developed reflex-based model is used to perform optimizations with different target behaviors on speed, step length, and step duration to analyze the correlation between reflex parameters and their influence on these gait characteristics. We identified nine key parameters that may affect the target speed ranging from slow to fast walking (0.48 and 1.71 m/s) as well as a large range of step lengths (0.43 and 0.88 m) and step duration (0.51, 0.98 s). The findings show that specific reflexes during stance significantly affect step length regulation, mainly given by positive force feedback of the ankle plantarflexors’ group. On the other hand, stretch reflexes active during swing of iliopsoas and gluteus maximus regulate all the gait characteristics under analysis. Additionally, the results show that the hamstrings’ group’s stretch reflex during the landing phase is responsible for modulating the step length and step duration. Additional validation studies in simulations demonstrated that the modulation of identified reflexes is sufficient to regulate the investigated gait characteristics. Thus, this study provides an overview of possible reflexes involved in modulating speed, step length, and step duration of human gaits.     

Neurokernel: An Open Source Platform for Emulating the Fruit Fly Brain

We have developed an open software platform called Neurokernel for collaborative development of comprehensive models of the brain of the fruit fly Drosophila melanogaster and their execution and testing on multiple Graphics Processing Units (GPUs). Neurokernel provides a programming model that capitalizes upon the structural organization of the fly brain into a fixed number of functional modules to distinguish between these modules’ local information processing capabilities and the connectivity patterns that link them. By defining mandatory communication interfaces that specify how data is transmitted between models of each of these modules regardless of their internal design, Neurokernel explicitly enables multiple researchers to collaboratively model the fruit fly’s entire brain by integration of their independently developed models of its constituent processing units. We demonstrate the power of Neurokernel’s model integration by combining independently developed models of the retina and lamina neuropils in the fly’s visual system and by demonstrating their neuroinformation processing capability. We also illustrate Neurokernel’s ability to take advantage of direct GPU-to-GPU data transfers with benchmarks that demonstrate scaling of Neurokernel’s communication performance both over the number of interface ports exposed by an emulation’s constituent modules and the total number of modules comprised by an emulation.     

The neural correlates of crowding-induced changes in appearance

Object recognition in the peripheral visual field is limited by crowding: the disruptive influence of nearby clutter. Despite its severity, little is known about the cortical locus of crowding. Here, we examined the neural correlates of crowding by combining event-related fMRI adaptation with a change-detection paradigm. Crowding can change the appearance of objects, such that items become perceptually matched to surrounding objects; we used this change in appearance as a signature of crowding and measured brain activity that correlated with the crowded percept. Observers adapted to a peripheral patch of noise surrounded by four Gabor flankers. When crowded, the noise appears oriented and perceptually indistinguishable from the flankers. Consequently, substitution of the noise for a Gabor identical to the flankers ("change-same") is rarely detected, whereas substitution for an orthogonal Gabor ("change-different") is rarely missed. We predicted that brain areas representing the crowded percept would show repetition suppression in change-same trials but release from adaptation in change-different trials. This predicted pattern was observed throughout cortical visual areas V1-V4, increasing in strength from early to late visual areas. These results depict crowding as a multistage process, involving even the earliest cortical visual areas, with perceptual consequences that are increasingly influenced by later visual areas.     

Reducing crowding by weakening inhibitory lateral interactions in the periphery with perceptual learning

We investigated whether lateral masking in the near-periphery, due to inhibitory lateral interactions at an early level of central visual processing, could be weakened by perceptual learning and whether learning transferred to an untrained, higher-level lateral masking known as crowding. The trained task was contrast detection of a Gabor target presented in the near periphery (4°) in the presence of co-oriented and co-aligned high contrast Gabor flankers, which featured different target-to-flankers separations along the vertical axis that varied from 2λ to 8λ. We found both suppressive and facilitatory lateral interactions at target-to-flankers distances (2λ - 4λ and 8λ, respectively) that were larger than those found in the fovea. Training reduces suppression but does not increase facilitation. Most importantly, we found that learning reduces crowding and improves contrast sensitivity, but has no effect on visual acuity (VA). These results suggest a different pattern of connectivity in the periphery with respect to the fovea as well as a different modulation of this connectivity via perceptual learning that not only reduces low-level lateral masking but also reduces crowding. These results have important implications for the rehabilitation of low-vision patients who must use peripheral vision to perform tasks, such as reading and refined figure-ground segmentation, which normal sighted subjects perform in the fovea.     

Top-down modulation of lateral interactions in early vision: does attention affect integration of the whole or just perception of the parts?

Attention can modulate sensitivity to local stimuli in early vision. But, can attention also modulate integration of local stimuli into global visual patterns? We recently measured effects of attention on the phenomenon of lateral interactions between collinear elements, commonly thought to reflect long-range mechanisms in early visual cortex underlying contour integration. We showed improved detection of low-contrast central Gabor targets in the context of collinear flankers, but only when the collinear flankers were attended for a secondary task rather than ignored in favor of an orthogonal flanker pair. Here, we contrast two hypotheses for how attention might modulate flanker influences on the target: by changing just local sensitivity to the flankers themselves (flanker-modulation-only hypothesis), or by weighting integrative connections between flanker and target (connection-weighting hypothesis). Modeled on the known nonlinear dependence of target visibility on collinear flanker contrast, the first hypothesis predicts that an increase in physical flanker contrast should readily offset any reduction in their effective contrast when ignored, thus eliminating attentional modulation. Conversely, the second hypothesis predicts that attentional modulation should persist even for the highest flanker contrasts. Our results showed the latter outcome and indicated that attention modulates flanker-target integration, rather than just processing of local flanker elements.     

Assessing Conformance with Benford’s Law: Goodness-Of-Fit Tests and Simultaneous Confidence Intervals

Benford’s Law is a probability distribution for the first significant digits of numbers, for example, the first significant digits of the numbers 871 and 0.22 are 8 and 2 respectively. The law is particularly remarkable because many types of data are considered to be consistent with Benford’s Law and scientists and investigators have applied it in diverse areas, for example, diagnostic tests for mathematical models in Biology, Genomics, Neuroscience, image analysis and fraud detection. In this article we present and compare statistically sound methods for assessing conformance of data with Benford’s Law, including discrete versions of Cramér-von Mises (CvM) statistical tests and simultaneous confidence intervals. We demonstrate that the common use of many binomial confidence intervals leads to rejection of Benford too often for truly Benford data. Based on our investigation, we recommend that the CvM statistic Ud2, Pearson’s chi-square statistic and 100(1 − α)% Goodman’s simultaneous confidence intervals be computed when assessing conformance with Benford’s Law. Visual inspection of the data with simultaneous confidence intervals is useful for understanding departures from Benford and the influence of sample size.     

Systematic Analysis of Compositional Order of Proteins Reveals New Characteristics of Biological Functions and a Universal Correlate of Macroevolution

We present a novel analysis of compositional order (CO) based on the occurrence of Frequent amino-acid Triplets (FTs) that appear much more than random in protein sequences. The method captures all types of proteomic compositional order including single amino-acid runs, tandem repeats, periodic structure of motifs and otherwise low complexity amino-acid regions. We introduce new order measures, distinguishing between ‘regularity’, ‘periodicity’ and ‘vocabulary’, to quantify these phenomena and to facilitate the identification of evolutionary effects. Detailed analysis of representative species across the tree-of-life demonstrates that CO proteins exhibit numerous functional enrichments, including a wide repertoire of particular patterns of dependencies on regularity and periodicity. Comparison between human and mouse proteomes further reveals the interplay of CO with evolutionary trends, such as faster substitution rate in mouse leading to decrease of periodicity, while innovation along the human lineage leads to larger regularity. Large-scale analysis of 94 proteomes leads to systematic ordering of all major taxonomic groups according to FT-vocabulary size. This is measured by the count of Different Frequent Triplets (DFT) in proteomes. The latter provides a clear hierarchical delineation of vertebrates, invertebrates, plants, fungi and prokaryotes, with thermophiles showing the lowest level of FT-vocabulary. Among eukaryotes, this ordering correlates with phylogenetic proximity. Interestingly, in all kingdoms CO accumulation in the proteome has universal characteristics. We suggest that CO is a genomic-information correlate of both macroevolution and various protein functions. The results indicate a mechanism of genomic ‘innovation’ at the peptide level, involved in protein elongation, shaped in a universal manner by mutational and selective forces.     

Fluctuation Scaling,Taylor’s Law,and Crime

Fluctuation scaling relationships have been observed in a wide range of processes ranging from internet router traffic to measles cases. Taylor’s law is one such scaling relationship and has been widely applied in ecology to understand communities including trees, birds, human populations, and insects. We show that monthly crime reports in the UK show complex fluctuation scaling which can be approximated by Taylor’s law relationships corresponding to local policing neighborhoods and larger regional and countrywide scales. Regression models applied to local scale data from Derbyshire and Nottinghamshire found that different categories of crime exhibited different scaling exponents with no significant difference between the two regions. On this scale, violence reports were close to a Poisson distribution (α = 1.057±0.026) while burglary exhibited a greater exponent (α = 1.292±0.029) indicative of temporal clustering. These two regions exhibited significantly different pre-exponential factors for the categories of anti-social behavior and burglary indicating that local variations in crime reports can be assessed using fluctuation scaling methods. At regional and countrywide scales, all categories exhibited scaling behavior indicative of temporal clustering evidenced by Taylor’s law exponents from 1.43±0.12 (Drugs) to 2.094±0081 (Other Crimes). Investigating crime behavior via fluctuation scaling gives insight beyond that of raw numbers and is unique in reporting on all processes contributing to the observed variance and is either robust to or exhibits signs of many types of data manipulation.     

Hedgehog signaling activation in the development of squamous cell carcinoma and adenocarcinoma of esophagus

Hedgehog (Hh) signaling is frequently activated in human cancer, including esophageal cancer. Most esophageal cancers are diagnosed in the advanced stages, therefore, identifying the very alterations that drive esophageal carcinogenesis may help designing novel strategies to diagnose and treat the disease. Analysis of Hh signaling in precancerous lesions is a critical first step in determining the significance of this pathway for carcinogenesis. Here we report our data on Hh target gene expression in 174 human esophageal specimens [28 esophageal adenocarcinomas (EAC), 19 Barrett’s esophagus, 103 cases of esophageal squamous cell carcinoma (ESCC), and 24 of squamous dysplastic lesions], and in two rat models of esophageal cancer. We found that 96% of human EAC express Hh target genes. We showed that PTCH1 expression is the most reliable biomarker. In contrast to EAC, only 38% of ESCC express Hh target genes. We found activation of Hh signaling in precancerous lesions of ESCCs and EACs in different degrees (21% and 58% respectively). Expression of Hh target genes is frequently detected in severe squamous dysplasia/ carcinoma in situ (p=0.04) and Barrett’s esophagus (p=0.01). Unlike EAC, sonic hedgehog (Shh) expression was rare in ESCCs. Consistent with the human specimen data, we found a high percentage of Hh signaling activation in precancerous lesions in rat models. These data indicate that Hh signaling activation is an early molecular event in the development of esophageal cancer, particularly EAC.     

Development of Eye Position Dependency of Slow Phase Velocity during Caloric Stimulation

The nystagmus in patients with vestibular disorders often has an eye position dependency, called Alexander’s law, where the slow phase velocity is higher with gaze in the fast phase direction compared with gaze in the slow phase direction. Alexander’s law has been hypothesized to arise either due to adaptive changes in the velocity-to-position neural integrator, or as a consequence of processing of the vestibular-ocular reflex. We tested whether Alexander’s law arises only as a consequence of non-physiologic vestibular stimulation. We measured the time course of the development of Alexander’s law in healthy humans with nystagmus caused by three types of caloric vestibular stimulation: cold (unilateral inhibition), warm (unilateral excitation), and simultaneous bilateral bithermal (one side cold, the other warm) stimulation, mimicking the normal push-pull pattern of vestibular stimulation. Alexander’s law, measured as a negative slope of the velocity versus position curve, was observed in all conditions. A reversed pattern of eye position dependency (positive slope) was found <10% of the time. The slope often changed with nystagmus velocity (cross-correlation of nystagmus speed and slope was significant in 50% of cases), and the average lag of the slope with the speed was not significantly different from zero. Our results do not support the hypothesis that Alexander’s law can only be observed with non-physiologic vestibular stimulation. Further, the rapid development of Alexander’s law, while possible for an adaptive mechanism, is nonetheless quite fast compared to most other ocular motor adaptations. These results suggest that Alexander’s law may not be a consequence of a true adaptive mechanism.     

Stimulus Roving and Flankers Affect Perceptual Learning of Contrast Discrimination in Macaca mulatta

‘Stimulus roving’ refers to a paradigm in which the properties of the stimuli to be discriminated vary from trial to trial, rather than being kept constant throughout a block of trials. Rhesus monkeys have previously been shown to improve their contrast discrimination performance on a non-roving task, in which they had to report the contrast of a test stimulus relative to that of a fixed-contrast sample stimulus. Human psychophysics studies indicate that roving stimuli yield little or no perceptual learning. Here, we investigate how stimulus roving influences perceptual learning in macaque monkeys and how the addition of flankers alters performance under roving conditions. Animals were initially trained on a contrast discrimination task under non-roving conditions until their performance levels stabilized. The introduction of roving contrast conditions resulted in a pronounced drop in performance, which suggested that subjects initially failed to heed the sample contrast and performed the task using an internal memory reference. With training, significant improvements occurred, demonstrating that learning is possible under roving conditions. To investigate the notion of flanker-induced perceptual learning, flanker stimuli (30% fixed-contrast iso-oriented collinear gratings) were presented jointly with central (roving) stimuli. Presentation of flanker stimuli yielded substantial performance improvements in one subject, but deteriorations in the other. Finally, after the removal of flankers, performance levels returned to their pre-flanker state in both subjects, indicating that the flanker-induced changes were contingent upon the continued presentation of flankers.