Colour constancy under simultaneous changes in surface position and illuminant

Two kinds of constancy underlie the everyday perception of surface colour: constancy under changes in illuminant and constancy under changes in surface position. Classically, these two constancies seem to place conflicting demands on the visual system: to both take into account the region surrounding a surface and also discount it. It is shown here, however, that the ability of observers to make surface-colour matches across simultaneous changes in test-surface position and illuminant in computer-generated 'Mondrian' patterns is almost as good as across changes in illuminant alone. Performance was no poorer when the surfaces surrounding the test surface were permuted, or when information from a potential comparison surface, the one with the highest luminance, was suppressed. Computer simulations of cone-photoreceptor activity showed that a reliable cue for making surface-colour matches in all experimental conditions was provided by the ratios of cone excitations between the test surfaces and a spatial average over the whole pattern.     

Cone contrast computations: physical versus perceived background and colour constancy

The light reflected from an object depends both on the object's surface and on the illuminant. Visual systems attempt to resolve this intrinsic ambiguity by comparing the light reflected from the object with respect to the background by computing the difference between the object-background light sampled by three cones. The cone-contrasts for the sample-background stimulus under the test illumination should correspond to the cone-contrasts for samples matched in appearance under the standard background (C). The validity of this cone-contrast rule (Whittle, 2003) and its possible link with stability of perceived colour was studied here using six test illuminants. A successive asymmetric colour-matching task with 40 simulated Munsell samples (value 7, chroma 4) on a neutral background (N7) was used. The subject adjusted the sample under standard illuminant C to match the colour appearance of the test sample under one of the test illuminants. Brunswik Ratio (BR) was used as an index of stability of colour appearance. When computed with respect to the reference illuminant C, the cone contrast rule was violated (particularly for S-cone-contrast). However, if a new reference point based on the perceived colour of the neutral background under the test illumination was used, the cone contrast rule applied. That is, when cone contrasts of the matching samples are computed with respect to this perceived background, they correspond to cone contrasts of the test stimuli. This represents a form of discounting the illuminant for the purpose of determining an object's cone-contrast against the background, which does not vary with background illumination. These cone contrasts, however, do not provide any information about the colour appearance of objects under particular viewing conditions, unless calibrated against a standard by allowing subjects to learn particular colours.     

Estimating photoreceptor excitations from spectral outputs of a personal light exposure measurement device

The intrinsic circadian clock requires photoentrainment to synchronize the 24-hour solar day. Therefore, light stimulation is an important component of chronobiological research. Currently, the chronobiological research field overwhelmingly uses photopic illuminance that is based on the luminous efficiency function, V(λ), to quantify light levels. However, recent discovery of intrinsically photosensitive retinal ganglion cells (ipRGCs), which are activated by self-contained melanopsin photopigment and also by inputs from rods and cones, makes light specification using a one-dimensional unit inadequate. Since the current understanding of how different photoreceptor inputs contribute to the circadian system through ipRGCs is limited, it is recommended to specify light in terms of the excitations of five photoreceptors (S-, M-, L-cones, rods and ipRGCs; Lucas et?al., 2014 Lucas RJ, Peirson SN, Berson DM, et al. (2014). Measuring and using light in the melanopsin age. Trends Neurosci. 37:1–9[Crossref], [PubMed], [Web of Science ®] , [Google Scholar]). In the current study, we assessed whether the spectral outputs from a commercially available spectral watch (i.e. Actiwatch Spectrum) could be used to estimate photoreceptor excitations. Based on the color sensor spectral sensitivity functions from a previously published work, as well as from our measurements, we computed spectral outputs in the long-wavelength range (R), middle-wavelength range (G), short-wavelength range (B) and broadband range (W) under 52 CIE illuminants (25 daylight illuminants, 27 fluorescent lights). We also computed the photoreceptor excitations for each illuminant using human photoreceptor spectral sensitivity functions. Linear regression analyses indicated that the Actiwatch spectral outputs could predict photoreceptor excitations reliably, under the assumption of linear responses of the Actiwatch color sensors. In addition, R, G, B outputs could classify illuminant types (fluorescent versus daylight illuminants) satisfactorily. However, the assessment of actual Actiwatch recording under several testing light sources showed that the spectral outputs were subject to great non-linearity, leading to less accurate estimation of photoreceptor excitations. Based on our analyses, we recommend that each spectral watch should be calibrated to measure spectral sensitivity functions and linearization characteristics for each sensor to have an accurate estimation of photoreceptor excitations. The method we provided to estimate photoreceptor excitations from the outputs of spectral watches could be used for chronobiological studies that can tolerate an error in the range of 0.2–0.5 log units. Our method can be easily expanded to incorporate linearization functions to have more accurate estimations.     

Is discrimination enhanced at the boundaries of perceptual categories? A negative case

The human visual system imposes discrete perceptual categories on the continuous input space that is represented by the ratios of excitations of the cones in the retina. Is discrimination enhanced at the boundaries between perceptual hues, in the way that discrimination may be enhanced at the boundaries between speech sounds in hearing? In the chromaticity diagram, the locus of unique green separates colours that appear yellowish from those that appear bluish. Using a two-alternative spatial forced choice and an adapting field equivalent to the Daylight Illuminant D65, we measured chromatic discrimination along lines orthogonal to the locus of unique green. In experimental runs interleaved with these performance measurements, we obtained estimates of the phenomenological boundary from the same observers. No enhancement of objectively measured discrimination was observed at the category boundary between yellowish and bluish hues. Instead, thresholds were minimal at chromaticities where the ratio of long-wave to middle-wave cone excitation was the same as that for the background adapting field.     

CONIFER OVULATE CONE MORPHOLOGY: IMPLICATIONS ON POLLEN IMPACTION PATTERNS

Empirically determined patterns of pollen impaction on the surfaces of pine ovulate cones are correlated with regions of nonlaminar flow created by the spatial arrangement and morphology (aspect ratios) of scale-bract complexes. Results from the serial discharge of pollen, upwind of ovulate cones, indicate that ovules on ovulate cones are preferentially impacted by pollen from their own species. Analyses indicate that while aerodynamic factors dominate the entrapment of pollen by ovulate cones, other factors such as pollen impaction-rebound and rebound-reentrainment are significant. Surface characteristics in addition to the settling velocities of pollen may play important roles in determining pollination efficiency. Wind tunnel analyses of the aerodynamic effects of scale-bract arrangement and aspect ratios indicate that each complex behaves as an aerofoil, deflecting air eddies toward the micropylar ends of ovules. The ovulate cone geometry, as a whole, deflects unidirectional wind into cyclonic vortices around the cone axis, each scale-bract deflecting nonimpacted pollen along orthostichies and parastichies. The morphology of the typical conifer ovulate cone is interpreted as a structure that optimizes anemophilous reproduction.     

Age related size invariance in perception of illusory and fragmented contours

We investigated invariant perception to sizes of images. Observers were schoolmates of 7-17 years and adults. Two types of stimuli were used: fragmented line drawing of common objects and discs with deleted sectors, which represented illusory Kanizsa contours when discs were in particular positions. In experiments with fragmented images, we found an improvement in image recognition with observers' age, increasing up to 13-14 years. The probability of recognition of fragmented line drawings increased significantly with decreasing stimulus size for children of 7-12 years, indicating that size invariance at recognition for fragmented line drawings was absent in these children. However, size invariance was found for observers of 13-17 years and for adults in this task. At Kanizsa illusion appearance, the ratio of the separation between discs and disc diameter was smaller when we used larger disc diameters. This ratio increased with increasing age of observers. Obtained results provide evidence for the absence of size invariance when perceiving the Kanizsa illusion in our experimental conditions.     

Age-related size invariance in perception of illusory and fragmented contours

We investigated invariant perception to sizes of images. Observers were secondary school students aged 7–17 years and adults. Two types of stimuli were used: fragmented line drawing of common objects and discs with deleted sectors which represented illusory Kanizsa contours, when discs were in particular positions. In experiments with fragmented images, we found an improvement in image recognition with observer’s age, increasing up to 13–14 years. The probability of recognition of fragmented line drawings increased significantly with decreasing stimulus size for children aged 7–12 years, indicating that size invariance at recognition for fragmented line drawings was absent in these children. However, size invariance was found for observers aged 7–12 years and for adults in this task. Upon the Kanizsa illusion appearance, the ratio of the separation between discs and disc diameter was smaller when we used larger disc diameters. This ratio increased with increasing age of observers. The obtained results provide evidence for the absence of size invariance when perceiving the Kanizsa illusion under our experimental conditions.     

Measurements of growth cone adhesion to culture surfaces by micromanipulation

Neurons were grown on plastic surfaces that were untreated, or treated with polylysine, laminin, or L1 and their growth cones were detached from their culture surface by applying known forces with calibrated glass needles. This detachment force was taken as a measure of the force of adhesion of the growth cone. We find that on all surfaces, lamellipodial growth cones require significantly greater detachment force than filopodial growth cones, but this differences is, in general, due to the greater area of lamellipodial growth cones compared to filopodial growth cones. That is, the stress (force/unit area) required for detachment was similar for growth cones of lamellipodial and filopodial morphology on all surfaces, with the exception of lamellipodial growth cones on L1-treated surfaces, which had a significantly lower stress of detachment than on other surfaces. Surprisingly, the forces required for detachment (760-3,340 mudynes) were three to 15 times greater than the typical resting axonal tension, the force exerted by advancing growth cones, or the forces of retraction previously measured by essentially the same method. Nor did we observe significant differences in detachment force among growth cones of similar morphology on different culture surfaces, with the exception of lamellipodial growth cones on L1-treated surfaces. These data argue against the differential adhesion mechanism for growth cone guidance preferences in culture. Our micromanipulations revealed that the most mechanically resistant regions of growth cone attachment were confined to quite small regions typically located at the ends of filopodia and lamellipodia. Detached growth cones remained connected to the substratum at these regions by highly elastic retraction fibers. The closeness of contact of growth cones to the substratum as revealed by interference reflection microscopy (IRM) did not correlate with our mechanical measurements of adhesion, suggesting that IRM cannot be used as a reliable estimator of growth cone adhesion.     

Modelling of the vertebrate visual system 3. topological analysis of the cone mosaic

Spatial organization of the cone mosaic of the generalized vertebrate retina consists of rows of red and green cones alternating with rows of blue and blank cones. Cone inputs to retinal elements are defined spatially by red and green unit hexagons. Topological analysis entails determining for each cone in the mosaic the number of each cone type present in the unit hexagon which the activated cone can influence via electrical coupling between cones and/or stray light. Only weighted inputs in one-half of a sextant of the unit hexagon need be designated, since all other weighted inputs can be determined by rules giving systematic transformations of all cone types from one sextant to another: these rules arise from symmetries of the cone mosaic. Four retinal types are possible depending on replacement of blank cones by specific cone types; three cone-dominant retinas, where all blank cones are replaced by a specific cone type, and two forms of a trichromatic retina, where blank cones are replaced by equal numbers of red and green cones. The weighted input is the sum of individual cone type contributions and depends on the number of each cone type in the unit hexagon which can influence the cone in question. Weighted inputs for cone-dominant retinas are readily found by replacing blank cones with the proper cone type, while weighted inputs for trichromatic retinas require use of a specified cone mosaic to determine extra red and green cones. Receptive field size of post-cone elements as well as overlap of the center and surround fields of annular organized receptive fields of retinal elements increased with increasing values for attenuation factors.     

Selective Stimulation of Penumbral Cones Reveals Perception in the Shadow of Retinal Blood Vessels

In 1819, Johann Purkinje described how a moving light source that displaces the shadow of the retinal blood vessels to adjacent cones can produce the entopic percept of a branching tree. Here, we describe a novel method for producing a similar percept. We used a device that mixes 56 narrowband primaries under computer control, in conjunction with the method of silent substitution, to present observers with a spectral modulation that selectively targeted penumbral cones in the shadow of the retinal blood vessels. Such a modulation elicits a clear Purkinje-tree percept. We show that the percept is specific to penumbral L and M cone stimulation and is not produced by selective penumbral S cone stimulation. The Purkinje-tree percept was strongest at 16 Hz and fell off at lower (8 Hz) and higher (32 Hz) temporal frequencies. Selective stimulation of open-field cones that are not in shadow, with penumbral cones silenced, also produced the percept, but it was not seen when penumbral and open-field cones were modulated together. This indicates the need for spatial contrast between penumbral and open-field cones to create the Purkinje-tree percept. Our observation provides a new means for studying the response of retinally stabilized images and demonstrates that penumbral cones can support spatial vision. Further, the result illustrates a way in which silent substitution techniques can fail to be silent. We show that inadvertent penumbral cone stimulation can accompany melanopsin-directed modulations that are designed only to silence open-field cones. This in turn can result in visual responses that might be mistaken as melanopsin-driven.     

Effects of Specular Highlights on Perceived Surface Convexity

Shading is known to produce vivid perceptions of depth. However, the influence of specular highlights on perceived shape is unclear: some studies have shown that highlights improve quantitative shape perception while others have shown no effect. Here we ask how specular highlights combine with Lambertian shading cues to determine perceived surface curvature, and to what degree this is based upon a coherent model of the scene geometry. Observers viewed ambiguous convex/concave shaded surfaces, with or without highlights. We show that the presence/absence of specular highlights has an effect on qualitative shape, their presence biasing perception toward convex interpretations of ambiguous shaded objects. We also find that the alignment of a highlight with the Lambertian shading modulates its effect on perceived shape; misaligned highlights are less likely to be perceived as specularities, and thus have less effect on shape perception. Increasing the depth of the surface or the slant of the illuminant also modulated the effect of the highlight, increasing the bias toward convexity. The effect of highlights on perceived shape can be understood probabilistically in terms of scene geometry: for deeper objects and/or highly slanted illuminants, highlights will occur on convex but not concave surfaces, due to occlusion of the illuminant. Given uncertainty about the exact object depth and illuminant direction, the presence of a highlight increases the probability that the surface is convex.     

A cone formation theory of contamination in high resolution transmission electron microscopy

The formation theory of the contamination cone in high resolution transmission electron microscopy is discussed on the basis of a time dependent series of micrographs and their tilted images for evaluating the contamination process. For a high current density of an electron probe focused on the specimen, it was observed that two contaminated cones covering both upper and lower surfaces of a carbon substrate are formed as a function of the exposure time of the electron beam. From the measurement of the cross-section and height of contaminated cones, the contamination rate together with the volume rate, and time dependent shapes of various cones were analysed. The analysis was dependent on the diffusion cross-section and current density together with an average lifetime of adsorption molecules.     

Growth cone-growth cone interactions in cultures of rat sympathetic neurons

Growth cones of sympathetic neurons from the superior cervical ganglia of neonatal rats were studied using video-microscopy to determine events following contact between growth cones and other cell surfaces, including other growth cones and neurites. A variety of behaviors were observed to occur upon contact between growth cones. Most commonly, one growth cone would collapse and subsequently retract upon establishing filopodial contact with the growth cone of another sympathetic neuron. Contacts resulting in collapse and retraction were often accompanied by a rapid and transient burst of lamellipodial activity along the neurite 30-50 microns proximal to the retracting growth cone. In no instances did interactions between growth cones and either fibroblasts or red blood cells result in the growth cone collapsing, suggesting that a specific recognition event was involved. On several occasions, growth cones were seen to track other growth cones, although fasciculation was rare. In some cases, there was no obvious response between contacting growth cones. Growth cone-growth cone contact was almost four times more likely to result in collapse and retraction than was growth cone-neurite contact (45% vs 12%, respectively). These observations suggest that the superior cervical ganglion may be composed of neurons with different cell surface determinants and that these determinants are more concentrated on the surface of growth cones than on neurites. These results further suggest that contact-mediated inhibition of growth cone locomotion may play an important role in growth cone guidance.     

Modulation of growth cone morphology by substrate-bound adhesion molecules.

The growth cone, a terminal structure on developing and regenerating axons, is specialized for motility and guidance functions. In vivo the growth cone responds to environmental cues to guide the axon to its appropriate target. These cues are thought to be responsible for position-specific morphological changes in the growth cone, but the molecules that control growth cone behavior are poorly characterized. We used scanning electron microscopy to analyze the morphology of retinal ganglion cell growth cones in vitro on different adhesion molecules that axons normally encounter in vivo. L1/8D9, N-cadherin, and laminin each induced distinctive morphological characteristics in growth cones. Growth cones elaborated lamellipodial structures in response to the cell adhesion molecules L1/8D9 and N-cadherin, whereas laminin supported filopodial growth cones with small veils. On L1/8D9, the growth cones were larger and produced more filopodia. Filopodial associations between adjacent growth cones and neurites were frequent on L1/8D9 but were uncommon on laminin or N-cadherin. These results demonstrate that different adhesion molecules have profoundly different effects on growth cone morphology. This is consistent with previous reports suggesting that changes in growth cone morphology in vivo occur in response to changes in substrate composition.     

Growth cone behavior and production of traction force

The growth cone must push its substrate rearward via some traction force in order to propel itself forward. To determine which growth cone behaviors produce traction force, we observed chick sensory growth cones under conditions in which force production was accommodated by movement of obstacles in the environment, namely, neurites of other sensory neurons or glass fibers. The movements of these obstacles occurred via three, different, stereotyped growth cone behaviors: (a) filopodial contractions, (b) smooth rearward movement on the dorsal surface of the growth cone, and (c) interactions with ruffling lamellipodia. More than 70% of the obstacle movements were caused by filopodial contractions in which the obstacle attached at the extreme distal end of a filopodium and moved only as the filopodium changed its extension. Filopodial contractions were characterized by frequent changes of obstacle velocity and direction. Contraction of a single filopodium is estimated to exert 50-90 microdyn of force, which can account for the pull exerted by chick sensory growth cones. Importantly, all five cases of growth cones growing over the top of obstacle neurites (i.e., geometry that mimics the usual growth cone/substrate interaction), were of the filopodial contraction type. Some 25% of obstacle movements occurred by a smooth backward movement along the top surface of growth cones. Both the appearance and rate of movements were similar to that reported for retrograde flow of cortical actin near the dorsal growth cone surface. Although these retrograde flow movements also exerted enough force to account for growth cone pulling, we did not observe such movements on ventral growth cone surfaces. Occasionally obstacles were moved by interaction with ruffling lamellipodia. However, we obtained no evidence for attachment of the obstacles to ruffling lamellipodia or for directed obstacle movements by this mechanism. These data suggest that chick sensory growth cones move forward by contractile activity of filopodia, i.e., isometric contraction on a rigid substrate. Our data argue against retrograde flow of actin producing traction force.     

Retinal cytoarchitecture in Some Mountain-stream Teleosts of India

In this study, the cytoarchitecture of retina in 10 mountain-stream teleosts (seven cyprinid and three loach species) of India was examined by conventional light microscopy. The mountain-streams are shallow, cold water bodies having high turbidity in the Monsoon and clear water in the colder months. The incumbent fishes are, thus, periodically exposed to low and bright light of the streams, respectively. It was of particular interest to see whether their retina would reflect adaptations to the changing photic environments of the mountain streams. The retina of the cyprinids possess multiple cone types, a moderate cone density, high convergence ratios and prominent retinomotor responses. Triple cones occur only in the danio, Danio aequipinnatus; other cyprinids possess miniature, short and long single cones, and double or twin cones. In general, the cones are bulky in appearance. Cone droplets (ellipsosomes) are found in Garra lamta and G. gotyla gotyla. All cone types, including the miniature single cones, undergo elongation during dark adaptation. The common retinal features in the loaches are the bulky cones, ellipsosomes, low rod density and low convergence ratios. There is no area-like specialization of the retina in any of the species examined. No relationship is obvious between cone pattern and ambient light or food habit in any of those teleosts studied; row pattern is present throughout the retina of the surface-dwelling, total insectivores (cyprinids: Danio aequipinnatus and Barilius spp.), whereas square mosaic pattern is present in the bottom-dwelling total herbiovores (Garra spp.) as well as in the insectivores (loaches: Nemacheilus spp.). The retinal features have suggested that the cyprinids are well-adapted to both clear and turbid water (ellipsosomes, multiple cone types, bulky cones, high rod count and high convergence ratios); the loaches, on the other hand, are better adapted to clear water of the mountain streams.     

Effect of defoliation on grass growth. A quantitative review

This study describes the seasonal patterns of seed and cone abortion in natural Pinus halepensis trees and assesses the effects of pollen quantity, tree size and cone location on seed and cone abortion. The cone-set ratio was similar to the expected value, based on literature on woody perennials. The seed-ovule ratio, however, was almost twice as high as its expected value. The investment in protective cone elements was high and did not vary with the seed-ovule ratio of a cone. Pollination factors influenced cone abortion, as indicated by the high abortion rate at the end of the pollination period. Furthermore, abortive cones had lower pollination rates and lower pollen loads than well developed cones. Effects of resource availability were assessed as effects of tree size and cone position on twigs. Small trees aborted more cones than big trees and cone abortion was higher in apical cones than in basal cones. Abortion in P. halepensis is selective, mediated by both pollen quantity and resource availability. The relative importance of pollen and resources is suggested to be flexible, probably varying between trees and years. The high seed-ovule ratio of P. halepensis is the result of high pollination rate and selective abortion. The selective abortion might be due to the high allocation to protective cone elements relative to the allocation to seeds.     

Interactions among moths, crossbills, squirrels, and lodgepole pine in a geographic selection mosaic

Repeated patterns among biological communities suggest similar evolutionary and ecological forces are acting on the communities. Conversely, the lack of such patterns suggests that similar forces are absent or additional ones are present. Coevolution between a seed predator, the red crossbill (Loxia curvirostra complex), and lodgepole pine (Pinus contorta var. latifolia) exemplifies the ecological and evolutionary predictions for coevolving systems. In the absence of another seed predator and preemptive competitor (pine squirrels Tamiasciurus hudsonicus), natural selection by crossbills results in the evolution of larger cones with thicker distal scales, while relaxation of selection by squirrels results in the evolution of cones with more seeds and a greater ratio of seed mass to cone mass. However, in one range, the Little Rocky Mountains, distal scale thickness has diverged as expected but cone size has not. In these mountains seed predation by lodgepole pine cone borer moths (Eucosma recissoriana) was about 10 times greater than in other ranges lacking squirrels. We quantified moth predation and cone traits and found that moths select for smaller cones with fewer seeds. Thus, selection by moths in the Little Rocky Mountains counters both selection by crossbills for large cone size and relaxation of selection by squirrels favoring more seeds per cone and accounts for the relatively small and few-seeded cones in these mountains. It is also apparent that selection by crossbills changes seed defenses in a manner that favors seed predation by moths, whereas selection by squirrels likely reduces such predation. These results demonstrate the importance of considering the evolutionary consequences of community context in locally evolved (coevolved) traits and interactions.