Effects of mean luminance changes on human contrast perception: contrast dependence, time-course and spatial specificity

Background

When we are viewing natural scenes, every saccade abruptly changes both the mean luminance and the contrast structure falling on any given retinal location. Thus it would be useful if the two were independently encoded by the visual system, even when they change simultaneously. Recordings from single neurons in the cat visual system have suggested that contrast information may be quite independently represented in neural responses to simultaneous changes in contrast and luminance. Here we test to what extent this is true in human perception.

Methodology/Principal Findings

Small contrast stimuli were presented together with a 7-fold upward or downward step of mean luminance (between 185 and 1295 Td, corresponding to 14 and 98 cd/m²), either simultaneously or with various delays (50–800 ms). The perceived contrast of the target under the different conditions was measured with an adaptive staircase method. Over the contrast range 0.1–0.45, mainly subtractive attenuation was found. Perceived contrast decreased by 0.052±0.021 (N = 3) when target onset was simultaneous with the luminance increase. The attenuation subsided within 400 ms, and even faster after luminance decreases, where the effect was also smaller. The main results were robust against differences in target types and the size of the field over which luminance changed.

Conclusions/Significance

Perceived contrast is attenuated mainly by a subtractive term when coincident with a luminance change. The effect is of ecologically relevant magnitude and duration; in other words, strict contrast constancy must often fail during normal human visual behaviour. Still, the relative robustness of the contrast signal is remarkable in view of the limited dynamic response range of retinal cones. We propose a conceptual model for how early retinal signalling may allow this.     

The detection of spatial discontinuities: interactions between contrast and spatial contiguity

Thresholds were measured for the detection of spatial discontinuities (notches and bumps) along luminance boundaries. At high contrasts of the boundary, thresholds expressed in terms of the spatial notch/bump height fell well inside the hyperacuity range. Expressed as luminance increment thresholds between adjacent photoreceptors, the same thresholds were similar to those previously reported by Hartridge and by Hecht and Mintz for the detection of a single line. The ability of observers to detect differences in the height of a boundary on either side of a mean luminance gap was also investigated, and the effect of the gap was found to depend upon stimulus contrast. At high contrasts the introduction of a gap increased thresholds, but at the lowest contrasts, thresholds were unaffected by a gap. The role of different spatial frequency and orientational mechanisms in vernier acuity is discussed.     

The spatial position and replication timing of chromosomal domains are both established in early G1 phase

Mammalian chromosomal domains replicate at defined, developmentally regulated times during S phase. The positions of these domains in Chinese hamster nuclei were established within 1 hr after nuclear envelope formation and maintained thereafter. When G1 phase nuclei were incubated in Xenopus egg extracts, domains were replicated in the proper temporal order with nuclei isolated after spatial repositioning, but not with nuclei isolated prior to repositioning. Mcm2 was bound both to early- and late-replicating chromatin domains prior to this transition whereas specification of the dihydrofolate reductase replication origin took place several hours thereafter. These results identify an early G1 phase point at which replication timing is determined and demonstrate a provocative temporal coincidence between the establishment of nuclear position and replication timing.     

Costs and benefits of within-group spatial position: a feeding competition model

An animal's within-group spatial position has several important fitness consequences. Risk of predation, time spent engaging in antipredatory behavior and feeding competition can all vary with respect to spatial position. Previous research has found evidence that feeding rates are higher at the group edge in many species, but these studies have not represented the entire breadth of dietary diversity and ecological situations faced by many animals. In particular the presence of concentrated, defendable food patches can lead to increased feeding rates by dominants in the center of the group that are able to monopolize or defend these areas. To fully understand the tradeoffs of within-group spatial position in relation to a variety of factors, it is important to be able to predict where individuals should preferably position themselves in relation to feeding rates and food competition. A qualitative model is presented here to predict how food depletion time, abundance of food patches within a group, and the presence of prior knowledge of feeding sites affect the payoffs of different within-group spatial positions for dominant and subordinate animals. In general, when feeding on small abundant food items, individuals at the front edge of the group should have higher foraging success. When feeding on slowly depleted, rare food items, dominants will often have the highest feeding rates in the center of the group. Between these two extreme points of a continuum, an individual's optimal spatial position is predicted to be influenced by an additional combination of factors, such as group size, group spread, satiation rates, and the presence of producer-scrounger tactics.     

Visual acuity and spatial contrast sensitivity in tree squirrels

The visual sensitivity of tree squirrels from three different species (two western gray squirrels, Sciurus griseus; three fox squirrels, Sciurus niger; and an eastern gray squirrel, Sciurus carolinensis) was measured for spatial patterns defined by luminance differences. Spatial contrast sensitivity functions were determined in behavioral discrimination experiments in which the stimuli were sinusoidally-modulated gratings. At an average luminance level of 3.4 cd/m² these squirrels were maximally sensitive to spatial frequencies of about 0.5 cycles/degree (c/d). These experiments, in conjunction with additional measurements involving discrimination of square-wave grating patterns, also indicate that the finest gratings these squirrels can discriminate average 2.2 c/d (SD = 0.42 c/d). There are no obvious differences in resolution acuity among the three species. Grating acuity also was measured at a much higher luminance level (340 cd/m²). The results of this experiment indicate that the spatial acuity of the tree squirrel increases with luminance level, reaching a maximum of 3.9 c/d at 340 cd/m².     

Discrimination of the dynamic properties of sound source spatial location in humans: Electrophysiology and psychophysics

The spatial resolution of the human auditory system was studied under conditions, where the location of the sound source was changed according to different temporal patterns of interaural time delay. Two experimental procedures were run in the same group of subjects: a psychophysical procedure (the transformed staircase method) and an electrophysiological one (which requires recording of mismatch negativity, the auditory evoked response component). It was established that (1) the value of the mismatch negativity reflected the degree of spatial deviation of the sound source; (2) the mismatch negativity was elicited even at minimum (20μs) interaural time delays under both temporal patterns (abrupt azimuth change and gradual sound movement at different velocities); (3) an abrupt change of the sound source azimuth resulted in a greater mismatch negativity than gradual sound movement did if the interaural time delay exceeded 40 μs; (4) the discrimination threshold values of the interaural delay obtained in the psychophysical procedure were greater than the minimum interaural delays that elicited mismatch negativity, with the exception of the expert listeners, who exhibited no significant difference.     

Phase contrast electron microscopy: development of thin-film phase plates and biological applications

Phase contrast transmission electron microscopy (TEM) based on thin-film phase plates has been developed and applied to biological systems. Currently, development is focused on two techniques that employ two different types of phase plates. The first technique uses a Zernike phase plate, which is made of a uniform amorphous carbon film that completely covers the aperture of an objective lens and can retard the phase of electron waves by pi/2, except at the centre where a tiny hole is drilled. The other technique uses a Hilbert phase plate, which is made of an amorphous carbon film that is twice as thick as the Zernike phase plate, covers only half of the aperture and retards the electron wave phase by pi. By combining the power of efficient phase contrast detection with the accurate preservation achieved by a cryotechnique such as vitrification, macromolecular complexes and supermolecular structures inside intact bacterial or eukaryotic cells may be visualized without staining. Phase contrast cryo-TEM has the potential to bridge the gap between cellular and molecular biology in terms of high-resolution visualization. Examples using proteins, viruses, cyanobacteria and somatic cells are provided.     

Regrouping of peptide chains in molecules and molecular complexes: recognition and spatial conjugation codes of amino acids

A two-step model for reactions between peptide and protein molecules in aqueous medium is considered. The first stage of the reaction involves specific recognition and primary complex formation. This process is governed by the amino acid interaction code a-a as a part of genetic code (algorithm a-n-n-a, amino acid-codon-anticodon-anti-amino acid). According to the a-a code, the primary complex formation is determined by amino acid pairs of opposite polarity. During the second stage of the reaction, when the contacting ligand and receptor surfaces undergo dehydration, the primary complex becomes rearranged. The new structure is mainly determined by pairwise contacts of amino acids having similar polarity and belonging to the same amino acid family.     

Aesthetic issues in spatial composition: effects of position and direction on framing single objects

Artists who work in two-dimensional visual media regularly face the problem of how to compose their subjects in aesthetically pleasing ways within a surrounding rectangular frame. We performed psychophysical investigations of viewers' aesthetic preferences for the position and facing direction of single, directed objects (e.g. people, cars, teapots and flowers) within such rectangular frames. Preferences were measured using two-alternative forced-choice preference judgments, the method of adjustment, and free choice in taking photographs. In front-facing conditions, preference was greatest for pictures whose subject was located at or near the center of the frame and decreased monotonically and symmetrically with distance from the center (the center bias). In the left- or right-facing conditions, there was an additional preference for objects to face into rather than out of the frame (the inward bias). Similar biases were evident using a method of adjustment, in which participants positioned objects along a horizontal axis, and in free choice photographs, in which participants were asked to take 'the most aesthetically pleasing picture' they could of everyday objects. The results are discussed as affirming the power of the center and facing direction in the aesthetic biases viewers bring to their appreciation of framed works of visual art (e.g. Alexander, 2002; Arnheim, 1988).