Detection of sinusoidal gratings by pattern-specific detectors: further evidence for the correlation principle in human vision

 The detection of compound sinusoidal gratings of various spatial frequency separations and four different grating sizes has been studied using the summation-to-threshold paradigm. Contrast interrelation functions have been measured and spatial frequency tuning estimates, based on the slope of the contrast interrelation function at two definite points, were derived using the “negative gradient technique” proposed by Logvinenko [Logvinenko (1995) Biol Cybern 73: 547–552]. It is shown that compound grating detection can be modelled by assuming pattern-specific sensory mechanisms for each of the spatial frequency components, which adapt to the periodicity and the size of the stimulus but not to its envelope function. Further, it is shown that relative sensitivity for a given spatial frequency separation can be predicted with good accuracy by the correlation of the grating components used for superposition. It is suggested that the most plausible implementation of the pattern correlation principle in human grating detection is the “grating cell” model. Received: 10 March 2000 / Accepted in revised form: 4 December 2000     

Two spatio-temporal filters in human vision

1. We have studied visual detection of a circular target moving across a spatially and/or temporally modulated background. Illumination, I _t , for threshold detection of the target has been measured as a function of background modulation frequency and changes in I _t associated with background modulation provide a means of determining the frequency response characteristics of visual channels. 2. Temporal frequency responses obtained with temporally modulated, spatially uniform backgrounds have pass-band characteristics and the temporal frequency for peak response increases with increase in mean background illumination. These temporal frequency responses resemble those of the de Lange (1954) filter, but the latter incorporates the incremental thresholds for steady backgrounds. 3. The amplitude of this temporal response saturates at low (40%) background modulation, decreases to zero as the target velocity falls to zero, and is maximum for a circular target of diameter 2°. 4. The spatial characteristics of this temporal filter were measured with a background field consisting of alternate steady and flickering bars. The resulting spatial frequency curve peaks at 1 cycle deg^-1 for all background illuminations and is independent of the background grating orientation. This spatial response differs significantly from the IMG spatial functions observed with a background grating (Barbur and Ruddock, 1980). 5. The spatial and temporal responses reviewed above exhibit similar parametric variations and we therefore associate them with a single spatiotemporal filter, ST2. 6. A second temporal response, with low-pass frequency characteristics, was observed with a background field consisting of two matched gratings, presented in spatial and temporal antiphase. This response has parametric properties similar to those of the IMG spatial response described previously by Barbur and Ruddock (1980), thus we associated the two sets of data with a single spatio-temporal filter, ST1. 7. We show that the ST2 responses can be obtained by combining ST1 responses, and we present a network incorporating the two filters. 8. We review other psychophysical studies which imply the activity of two spatio-temporal filters with properties of the kind revealed in our studies. We argue that filter ST1 has properties equivalent to those of X-type and filter ST2 has properties equivalent to those of Y-type electrophysiological mechanisms.     

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.     

Interocular Transfer of Adaptation to Spatial Frequency during Retinal Ischaemia

PROLONGED inspection of a grating pattern of repetitive light and dark contours produces a transient reduction in the visibility of low-contrast gratings the spatial frequency and orientation of which are similar to that of the inspected grating. This after-effect, often referred to as adaptation, occurs even when one eye performs the inspection and its partner is tested. Certain properties of this after-effect, with other data, suggest that its locus is visual cortex^1–7, but the evidence is largely indirect because of the difficulty of identifying sites of action within the stages of the human visual system. One of the few techniques available for inferentially partitioning the visual system stages is retinal pressure blinding, a heroic but effective method for functionally uncoupling the retina from higher stages^8–11.     

Visual sensitivity of ground squirrels to spatial and temporal luminance variations

Summary We have investigated the visual sensitivity of the California ground squirrel (Speromphilus beecheyi) to spatial and temporal luminance patterns. Spatial contrast sensitivity functions were determined in behavioral discrimination experiments in which the stimuli were sinusoidally-modulated luminance gratings. These squirrels were found to be maximally sensitive to spatial frequencies of about 0.7 cycles/ degree (c/d), and they are unable to discriminate gratings whose frequencies exceed 4 c/d. Similar results were obtained in electrophysiological experiments when the visually evoked cortical potential (VECP) was recorded from anesthetized squirrels. A third experiment involved tests of the ability of ground squirrels to discriminate square-wave gratings of much higher luminance (340 cd/m²). The finest gratings which were discriminable at this luminance level did not exceed 3.9–4.3 c/d and, thus, we conclude that the maximal spatial resolution of the California ground squirrel is about 4 c/d (corresponding to a bar separation of 7.5). In another behavioral experiment the abilities of ground squirrels to discriminate sinusoidally flickering lights (mean luminance = 3.4 cd/m²) was measured. The results show that ground squirrels are maximally sensitive to lights flickering at a rate of about 18 Hz, and that the highest rates that are still discriminable are slightly above 60 Hz.Abbreviations c/d cycles/degree - CFF critical flicker frequency - VECP visually evoked cortical potential This research was supported by Grant EY 00105 from the National Eye Institute. We thank David Birch who participated in some preliminary behavioral experiments and Kenneth Long who provided the histological material from which measurements of receptor spacing were made.     

Phase channels and the square-wave illusion

We have utilized a phase-sensitive effect, the square-wave illusion, to investigate the presence and tuning of phase-sensitive mechanisms. Subjects viewed a 1 c/deg triangular-wave grating and reported the presence or absence of the illusion, before and after adaptation to square-wave gratings of various spatial frequencies. Illusion strength declined with adaptation and was a function of spatial frequency. This function may reflect the spatial frequency tuning of a 'phase channel'. We have also measured by adaptation a contrast sensitivity function, maximally sensitive to a frequency of 1 c/deg. When normalized to the same maximal depression from baseline sensitivity, the phase channel and spatial-frequency channel shared the same low frequency tuning, but the phase channel was broader, with extended sensitivity to higher frequencies. These results can be understood if the phase channel is constructed from combinations of phase sensitive spatial-frequency channels.     

Effect of Pre-natal Protein Deprivation on Neonatal Cerebellar Development <Emphasis Type="Italic">in vitro</Emphasis>

MOST biochemical and morphological studies of the effects of nutritional deprivation on the mammalian nervous system have dealt with quantitative undernutrition during the first 21 postnatal days. Findings have included a reduction in brain cellularity^1–4; a delay in neuronal and glial differentiation³; and a decrease in the lipids^3,3–7, lamellae numbers⁸ and stainability^3,5,9 of the myelin sheath. Such studies, even of those regions of the nervous system developing maximally post-natally, may be criticized as enabling the anlage, however undifferentiated, to develop normally until birth. I have, therefore, studied in a tissue culture system the morphogenetic effects of pre-natal maternal protein deprivation on the growth and development of the cerebellum, a region of the brain developing maximally after birth. Cultures of neonatal mouse cerebellum from mothers deprived of protein were analysed in a nutritionally normal environment to determine the effect of pre-natal protein restriction on the post-natal developmental potential of the nervous system.     

Visual acuity of the midland banded water snake estimated from evoked telencephalic potentials

The visual acuity of seven midland banded water snakes was measured by recording evoked responses from telencephalon to temporally modulated square wave grating patterns. Using conventional electrophysiological techniques and signal averaging, high contrast square wave gratings of different spatial frequencies were presented. Acuity was estimated by extrapolating relative response amplitude/log₁₀ spatial frequency functions which yielded an average acuity of 4.25 cycles/degree. Refractive state was also estimated by recording evoked potentials to intermediate spatial frequencies with different lenses in front of the eye. Polynomial fits indicated that under the experimental conditions the snakes were around 6.4 diopters hyperopic suggesting a corrected acuity of 4.89 cycles/degree. Reduction of grating luminance resulted in a reduction in evoked potential acuity measurements. These results indicate that the spatial resolution of midland banded water snakes is the equal of cat; about 20/120 in human clinical terms.     

MRI pattern recognition in multiple sclerosis normal-appearing brain areas

Objective

Here, we use pattern-classification to investigate diagnostic information for multiple sclerosis (MS; relapsing­remitting type) in lesioned areas, areas of normal­appearing grey matter (NAGM), and normal-appearing white matter (NAWM) as measured by standard MR techniques.

Methods

A lesion mapping was carried out by an experienced neurologist for Turbo Inversion Recovery Magnitude (TIRM) images of individual subjects. Combining this mapping with templates from a neuroanatomic atlas, the TIRM images were segmented into three areas of homogenous tissue types (Lesions, NAGM, and NAWM) after spatial standardization. For each area, a linear Support Vector Machine algorithm was used in multiple local classification analyses to determine the diagnostic accuracy in separating MS patients from healthy controls based on voxel tissue intensity patterns extracted from small spherical subregions of these larger areas. To control for covariates, we also excluded group-specific biases in deformation fields as a potential source of information.

Results

Among regions containing lesions a posterior parietal WM area was maximally informative about the clinical status (96% accuracy, p<10⁻¹³). Cerebellar regions were maximally informative among NAGM areas (84% accuracy, p<10⁻⁷). A posterior brain region was maximally informative among NAWM areas (91% accuracy, p<10⁻¹⁰).

Interpretation

We identified regions indicating MS in lesioned, but also NAGM, and NAWM areas. This complements the current perception that standard MR techniques mainly capture macroscopic tissue variations due to focal lesion processes. Compared to current diagnostic guidelines for MS that define areas of diagnostic information with moderate spatial specificity, we identified hotspots of MS associated tissue alterations with high specificity defined on a millimeter scale.     

Tumor eradication after cyclophosphamide depends on concurrent depletion of regulatory T cells: a role for cycling TNFR2-expressing effector-suppressor T cells in limiting effective chemotherapy

Tumor cell death potentially engages with the immune system. However, the efficacy of anti-tumor chemotherapy may be limited by tumor-driven immunosuppression, e.g., through CD25⁺ regulatory T cells. We addressed this question in a mouse model of mesothelioma by depleting or reconstituting CD25⁺ regulatory T cells in combination with two different chemotherapeutic drugs. We found that the efficacy of cyclophosphamide to eradicate established tumors, which has been linked to regulatory T cell depletion, was negated by adoptive transfer of CD25⁺ regulatory T cells. Analysis of post-chemotherapy regulatory T cell populations revealed that cyclophosphamide depleted cycling (Ki-67^hi) T cells, including foxp3⁺ regulatory CD4⁺ T cells. Ki-67^hi CD4⁺ T cells expressed increased levels of two markers, TNFR2 and ICOS, that have been associated with a maximally suppressive phenotype according to recently published studies. This suggest that cyclophosphamide depletes a population of maximally suppressive regulatory T cells, which may explain its superior anti-tumor efficacy in our model. Our data suggest that regulatory T cell depletion could be used to improve the efficacy of anti-cancer chemotherapy regimens. Indeed, we observed that the drug gemcitabine, which does not deplete cycling regulatory T cells, eradicates established tumors in mice only when CD25⁺ CD4⁺ T cells are concurrently depleted. Cyclophosphamide could be used to achieve regulatory T cell depletion in combination with chemotherapy. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The latency of the light response is modulated by the phosphorylation state of Drosophila TRP at a specific site

Drosophila photoreceptors respond to oscillating light of high frequency (～100 Hz), while increasing the oscillating light intensity raises the maximally detected frequency. Recently, we reported that dephosphorylation of the light-activated TRP ion channel at S936 is a fast, graded, light-, and Ca²⁺-dependent process. We further found that this process affects the detection limit of high frequency oscillating light. Accordingly, transgenic Drosophila, which do not undergo phosphorylation at the S936-TRP site (trp^S936A), revealed a short time-interval before following the high stimulus frequency (oscillation-lock response) in both dark- and light-adapted flies. In contrast, the trp^S936D transgenic flies, which mimic constant phosphorylation, showed a long-time interval to oscillation-lock response in both dark- and light-adapted flies. Here we extend these findings by showing that dark-adapted trp^S936A flies reveal light-induced current (LIC) with short latency relative to trp^WT or trp^S936D flies, indicating that the channels are a limiting factor of response kinetics. The results indicate that properties of the light-activated channels together with the dynamic light-dependent process of TRP phosphorylation at the S936 site determine response kinetics.     

An automated paradigm for Drosophila visual psychophysics

Background

Mutations that cause learning and memory defects in Drosophila melanogaster have been found to also compromise visual responsiveness and attention. A better understanding of attention-like defects in such Drosophila mutants therefore requires a more detailed characterization of visual responsiveness across a range of visual parameters.

Methodology/Principal Findings

We designed an automated behavioral paradigm for efficiently dissecting visual responsiveness in Drosophila. Populations of flies walk through multiplexed serial choice mazes while being exposed to moving visuals displayed on computer monitors, and infra-red fly counters at the end of each maze automatically score the responsiveness of a strain. To test our new design, we performed a detailed comparison between wild-type flies and a learning and memory mutant, dunce ¹. We first confirmed that the learning mutant dunce ¹ displays increased responsiveness to a black/green moving grating compared to wild type in this new design. We then extended this result to explore responses to a wide range of psychophysical parameters for moving gratings (e.g., luminosity, contrast, spatial frequency, velocity) as well as to a different stimulus, moving dots. Finally, we combined these visuals (gratings versus dots) in competition to investigate how dunce ¹ and wild-type flies respond to more complex and conflicting motion effects.

Conclusions/Significance

We found that dunce ¹ responds more strongly than wild type to high contrast and highly structured motion. This effect was found for simple gratings, dots, and combinations of both stimuli presented in competition.     

Perceptual spatial frequency — Orientation surface: Psychophysics and line element theory

We have investigated the discriminability of gratings which simultaneously vary in spatial frequency and orientation. Thirteen and nine reference gratings were used with two observers, and bivariate discrimination probability surfaces were determined around each grating. These data were then fitted to a general bivariate Gaussian function. The results clearly demonstrate local separability in this log frequency and orientation discrimination domain. Our results also show that the factor contributing most to the non-Euclidean nature of such frequency/orientation discrimination is orientation anisotropia, although we also find some evidence for smaller changes in the associated Riemannian line-element at different frequency ranges. These results cast doubt upon claims for a pseudo line-element for frequency discrimination based upon the nonlinear outputs of a fixed set of detectors.Study supported by Fraunhofer Gesellschaft Grant INSAN I 0784-V-6385 and Guest Professorship Mu 93/103-1 for TC by Deutsche Forschungsgemeinschaft     

A model for direction selectivity in threshold motion perception

Thresholds were measured for a moving line superimposed on moving sinusoidal gratings. When line and grating moved in the same direction significant subthreshold summation was observed over a range of spatial frequencies. For motion of the line and grating in opposite directions, summation was never observed. This supports the hypothesis that direction selective mechanisms are responsible for motion perception at threshold. Further analysis of the data produced estimates of the spatial frequency tuning of these mechanisms. A quantitative model is proposed to interpret the data, and it is suggested that flickering gratings are not decomposed into their moving components by the visual system.     

The molecular anatomy of spontaneous germline mutations in human testes

The frequency of the most common sporadic Apert syndrome mutation (C755G) in the human fibroblast growth factor receptor 2 gene (FGFR2) is 100–1,000 times higher than expected from average nucleotide substitution rates based on evolutionary studies and the incidence of human genetic diseases. To determine if this increased frequency was due to the nucleotide site having the properties of a mutation hot spot, or some other explanation, we developed a new experimental approach. We examined the spatial distribution of the frequency of the C755G mutation in the germline by dividing four testes from two normal individuals each into several hundred pieces, and, using a highly sensitive PCR assay, we measured the mutation frequency of each piece. We discovered that each testis was characterized by rare foci with mutation frequencies 10³ to >10⁴ times higher than the rest of the testis regions. Using a model based on what is known about human germline development forced us to reject (p < 10⁻⁶) the idea that the C755G mutation arises more frequently because this nucleotide simply has a higher than average mutation rate (hot spot model). This is true regardless of whether mutation is dependent or independent of cell division. An alternate model was examined where positive selection acts on adult self-renewing Ap spermatogonial cells (SrAp) carrying this mutation such that, instead of only replacing themselves, they occasionally produce two SrAp cells. This model could not be rejected given our observed data. Unlike the disease site, similar analysis of C-to-G mutations at a control nucleotide site in one testis pair failed to find any foci with high mutation frequencies. The rejection of the hot spot model and lack of rejection of a selection model for the C755G mutation, along with other data, provides strong support for the proposal that positive selection in the testis can act to increase the frequency of premeiotic germ cells carrying a mutation deleterious to an offspring, thereby unfavorably altering the mutational load in humans. Studying the anatomical distribution of germline mutations can provide new insights into genetic disease and evolutionary change.     

A new perceptual bias reveals suboptimal population decoding of sensory responses

Several studies have reported optimal population decoding of sensory responses in two-alternative visual discrimination tasks. Such decoding involves integrating noisy neural responses into a more reliable representation of the likelihood that the stimuli under consideration evoked the observed responses. Importantly, an ideal observer must be able to evaluate likelihood with high precision and only consider the likelihood of the two relevant stimuli involved in the discrimination task. We report a new perceptual bias suggesting that observers read out the likelihood representation with remarkably low precision when discriminating grating spatial frequencies. Using spectrally filtered noise, we induced an asymmetry in the likelihood function of spatial frequency. This manipulation mainly affects the likelihood of spatial frequencies that are irrelevant to the task at hand. Nevertheless, we find a significant shift in perceived grating frequency, indicating that observers evaluate likelihoods of a broad range of irrelevant frequencies and discard prior knowledge of stimulus alternatives when performing two-alternative discrimination.     

Moms are better nurses than dads: gender biased self‐facilitation in a dioecious Juniperus tree

Questions: Can gender of nurse plants affect regeneration patterns and spatial population structure? Is there a seed‐seedling conflict in the regeneration process? What factors are responsible for the clumped spatial population structure observed for adult trees? Location: Mediterranean cold semi‐arid high mountains in Spain. Methods: The spatial pattern of adult Juniperus thurifera trees was studied by means of Ripley's K‐analysis. χ² analyses were used to test for natural seedling frequency in each of three main microhabitats: (1) under female and (2) male tree canopies and (3) in open interspaces. The observed pattern was explained experimentally by studying seed and seedling survival for two years. Survival probabilities were calculated across life stages for each of three main microhabitats. Results: Adult J. thurifera trees were aggregated in space. Most seedlings were found underneath female J. thurifera trees. Experimental studies demonstrated that from seed dispersal to seedling survival all life stages showed the same positive or negative trend within a given microhabitat, indicating stage coupling and no seed‐seedling conflicts. Attraction of frugivo‐rous birds by reproductive female junipers and improvement of environmental conditions beneath tree canopies were the main factors responsible for the variation in seedling density among microhabitats; highest underneath female trees and lowest in open interspaces. Conclusions: In dioecious species, the gender of nurse plants can significantly determine the spatial population structure. In J. thurifera forests, facilitation beneath female trees occurs among all life stages without any sign of seed‐seedling conflict. The most critical factors shaping the spatial population structure were directed seed dispersal and environmental amelioration beneath female conspecific trees.     

Resource heterogeneity does not explain the diversity–productivity relationship across a boreal island fertility gradient

Many studies at the regional scale have found either negative or hump‐shaped relationships between productivity and diversity, and some theories propose that these occur because soil resource heterogeneity is either lower or less important in more productive environments. However, there have been few explicit tests of these theories in natural ecosystems. We evaluated the relationship between soil resource heterogeneity and plant richness within a well characterized system of 30 islands in northern Sweden across which soil fertility and productivity declines, and species richness increases, as a consequence of ecosystem retrogression. On each island we created a spatially explicit grid consisting of 49 sampling points in a 9.5 m quadrat, which we used to quantify spatial heterogeneity of five soil variables (NH₄⁺‐N, amino N, PO₄^?‐P, microbial biomass, and decomposition), and plant community composition. Using a hierarchical Bayesian approach, we estimated mean semivariograms of each variable for each island size class to compare three components of spatial heterogeneity: total variability, spatial grain, and patchiness. This analysis showed that variability within islands was usually lowest on small islands, where species richness was highest and productivity lowest; however, NH₄⁺‐N and amino N had greater patchiness and spatial grain on small islands. We did not detect any significant across‐island correlations between whole‐plot plant species richness and either whole‐plot standard deviation or coefficient of variation of any soil variable. Using partial Mantel tests, we found that mean correlation coefficients between within‐plot plant community composition and the soil variables were never significant for any island size class, and did not differ between island size classes. Our findings do not provide any evidence that soil resource heterogeneity controls the productivity–diversity relationship in this system, and suggests other mechanisms are primarily responsible.     

Evidence for different nonlinear summation schemes for lines and gratings at threshold

Within a wide class of multichannel models of the visual system it is suggested that spatial distributions of luminance are processed by the independent activation of grating detectors, or spatial frequency channels. Probability summation is often described in terms of Quick's nonlinear pooling model [Quick RF (1974) Kybernetik 16:65–67]. Using this model, we find evidence for the existence of different kinds of nonlinear summation at threshold; for compound gratings with well-separated spatial frequency components, the threshold functions indicate nonlinear summation which is not compatible with probability summation, while for line patterns well separated in the spatial domain the probability summation rule proves compatible with the data. Received: 24 June 1998 / Accepted in revised form: 16 March 1999