Horizontal-vertical filters in early vision predict anomalous line-orientation identification frequencies.

A characteristic of early visual processing is a reduction in the effective number of filter mechanisms acting in parallel over the visual field. In the detection of a line target differing in orientation from a background of lines, performance with brief displays appears to be determined by just two classes of orientation-sensitive filter, with preferred orientations close to the vertical and horizontal. An orientation signal represented as a linear combination of responses from such filters is shown to provide a quantitative prediction of the probability density function for identifying the perceived orientation of a target line. This prediction was confirmed in an orientation-matching experiment, which showed that the precision of orientation estimates was worst near the vertical and horizontal and best at about 30 degrees each side of the vertical, a result that contrasts with the classical oblique effect in vision, when scrutiny of the image is allowed. A comparison of predicted and observed frequency distributions showed that the hypothesized orientation signal was formed as an opponent combination and horizontal and vertical filter responses.     

Matched filters in human vision

The technique of subthreshold addition of sinusoidal gratings was used to analyse the visual system of man during the perception of edges, lines, and bars. The experimentally obtained sensitivity function varies in close relationship to the test pattern, and can be factorized into the conjugate complex spectrum of the test pattern at threshold and a pattern-invariant function of spatial frequency. Interpreting the sensitivity function as transfer function, which is possible under certain conditions, we can describe the visual system as a matched filter which extracts an input signal contaminated with noise of specified spectral energy density. Questions discussed refer to the spatial operations occurring in matched filters, the relationship between the modulation transfer function for sine-wave gratings and the pattern-invariant transfer component, the exact determination of elements within the theoretical concept, and the realization of matched filters by the nervous system.These investigations were supported as a project of the Sonderforschungsbereich 50 Kybernetik, München.     

Two spatio-temporal filters in human vision

We have used the psychophysical methods described in the first paper of this series (Holliday and Ruddock, 1983) to determine selected spatial and temporal response characteristics of the ST1 and ST2 filters for subjects suffering visual defects. Data are given for 19 amblyopes, an albino and a hemianope, and comparison data are also given for a number of subjects with normal vision. The ST1 spatial responses for both the "normal" and "amblyopic" eyes of 12 convergent strabismic amblyopes are displaced to low spatial frequencies compared to the normal curve, which implies that there is a loss of fine spatial tuning. In all but one subject, the curve for the "amblyopic" eye peaks at a spatial frequency lower than that for the "normal" eye, thus the former deviates further from the normal pattern than the latter. The ST1 spatial responses of 6 refractive amblyopes are also displaced to the low frequency side of the normal curve, although on average the shift is smaller than in the case of the strabismic amblyopes. For each subject, the response curve of the "amblyopic" eye peaks at a lower spatial frequency than does that for the "normal" eye. ST1 spatial responses were measured for targets located up to 30 degrees off-axis along the horizontal meridian and sample data are given for one strabismic and one refractive amblyope and for two normal subjects. It is concluded from these data that the changes in the spatial responses associated with amblyopia do not simply reflect eccentric fixation of the target. The ST2 spatial response was measured for the "normal" and "amblyopic" eyes of 9 amblyopes (7 strabismic and 2 refractive). There is no significant difference between the average amblyopic response and that of normal subjects, and only in one case does the response for an "amblyopic" eye peak at a frequency lower than the peak frequency for normal vision. The ST2 temporal response for 9 amblyopes shows no systematic deviations from the normal response. For the albino, both the ST1 and ST2 spatial responses peak at around 0.3 cycles deg-1, and both curves are displaced considerably to the low spatial frequency side of the normal ST2 spatial response. The albino's ST2 temporal response is essentially normal. Measurements for the hemianope's "blind" hemifield under conditions appropriate to the isolation of the ST1 and ST2 spatial responses reveal no tuning curves. The ST2 temporal response for the "blind" hemifield, however, is of large amplitude, with a peak at 2 Hz, well below the normal frequency response peak. It is argued that the loss of fine spatial tuning which occurs in the ST1, but not the ST2, spatial responses of the amblyopes is consistent with the sequential organisation of these two filter classes proposed by Holliday and Ruddock (1983). Further, for the only two subjects whose ST2 spatial response curves are displaced to abnormally low frequencies (the albino and a strabismic amblyope) the ST1 spatial response is shifted to low spatial frequencies compared to the normal ST2 curve...     

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.     

Human vision combines oriented filters to compute edges.

The experiments examined the perceived spatial structure of plaid patterns, composed of two or three sinusoidal gratings of the same spatial frequency, superimposed at different orientations. Perceived structure corresponded well with the pattern of zero crossings in the output of a circular spatial filter applied to the image. This lends some support to Marr & Hildreth's (Proc. R. Soc. Lond. B 207, 187 (1980)) theory of edge detection as a model for human vision, but with a very different implementation. The perceived structure of two-component plaids was distorted by prior exposure to a masking or adapting grating, in a way that was perceptually equivalent to reducing the contrast of one of the plaid components. This was confirmed by finding that the plaid distortion could be nulled by increasing the contrast of the masked or adapted component. A corresponding reduction of perceived contrast for single gratings was observed after adaptation and in some masking conditions. I propose the outlines of a model for edge finding in human vision. The plaid components are processed through cortical, orientation-selective filters that are subject to attenuation by forward masking and adaptation. The outputs of these oriented filters are then linearly summed to emulate circular filtering, and zero crossings (zcs) in the combined output are used to determine edge locations. Masking or adapting to a grating attenuates some oriented filters more than others, and although this changes only the effective contrast of the components, it results in a geometric distortion at the zc level after different filters have been combined. The orientation of zcs may not correspond at all with the orientation of Fourier components, but they are correctly predicted by this two-stage model. The oriented filters are not 'orientation detectors', but are precursors to a more subtle stage that locates and represents spatial features.     

Cytoplasmic membrane transplants in early Xenopus embryos indicate cell-cycle-specific effects.

Fragments of rough endoplasmic reticulum or Golgi complex isolated from normal adult rat liver homogenates were injected into one cell of cleaving two-cell Xenopus laevis embryos and the effects on development were monitored during early cleavage by morphological analysis. Scanning electron microscopy revealed the formation of large cells on the injected side of the embryos. Such large cells were not present in controls and thus were considered to have been formed as a consequence of delayed cleavage. Delay of cleavage was obtained with as little as 1 ng of membrane protein giving a ratio of membrane protein to embryo protein of 1:10(5). Cytological observations of microinjected embryos confirmed the occurrence of delayed cytokinesis and suggested that nuclear division became asynchronous. Since rough microsomes from proliferating tissues (i.e., livers with primary tumors and livers undergoing regeneration) showed little or no effect on cytokinesis after microinjection into early embryos, we conclude that cytoplasmic membranes may exhibit cell-cycle-specific properties important for normal development.     

Early vision is early in time

Roelfsema, Tolboom, and Khayat have found that neurons in primary visual cortex, V1, increase their spike firing rates to signal image segmentation and attention. V1 responses were in a temporal sequence: first to image motion, next to segmentation, last to attentional signals. The involvement of V1 with segmentation and attention suggests modifying the hierarchical view of visual perception.     

Cross- and auto-correlation in early vision

Neurons that respond selectively to the orientation of visual stimuli were discovered in V1 more than 50 years ago, but it is still not fully understood how or why this is brought about. We report experiments planned to show whether human observers use cross-correlation or auto-correlation to detect oriented streaks in arrays of randomly positioned dots, expecting that this would help us to understand what David Marr called the 'computational goal' of V1. The streaks were generated by two different methods: either by sinusoidal spatial modulation of the local mean dot density, or by introducing coherent pairs of dots to create moiré patterns, as Leon Glass did. A wide range of dot numbers was used in the randomly positioned arrays, because dot density affects cross- and auto-correlation differently, enabling us to infer which method was used. This difference stems from the fact that the cross-correlation task is limited by random fluctuations in the local mean density of individual dots in the noisy array, whereas the auto-correlation task is limited by fluctuations in the numbers of randomly occurring spurious pairs having the same separation and orientation as the deliberately introduced coherent pairs. After developing a new method using graded dot luminances, we were able to extend the range of dot densities that could be used by a large factor, and convincing results were obtained indicating that the streaks generated by amplitude modulation were discriminated by cross-correlation, while those generated as moiré patterns were discriminated by auto-correlation. Though our current results only apply to orientation selectivity, it is important to know that early vision can do more than simple filtering, for evaluating auto-correlations opens the way to more interesting possibilities, such as the detection of symmetries and suspicious coincidences.     

Spatial characteristics of movement detection mechanisms in human vision

Visual detection of targets moving against structured background fields has been studied with near-monochromatic stimuli, selected so as to isolate the different increment threshold spectral response mechanisms. It is shown that for foveal vision, the red-and green-sensitive mechanisms (₅ and ₄ respectively) yield IMG functions (Barbur and Ruddock, 1980), similar to those found with white light. In contrast, the blue-sensitive (₁) mechanism yields a low-frequency IMG response quite unlike that found for the other mechanisms. There is also considerable variation between subjects in this case. Measurements taken 30° off-axis with low (1.4 log trolands) background illumination level, yield a low frequency response IMG function for both rod and cone spectral mechanisms, similar to those found with white light stimuli. At high illumination levels (>2.2 log trolands), the IMG function for the ₅-mechanism is shifted to higher spatial frequencies, as is also observed with white light stimuli. A wavelength-selective binocular interaction effect, manifested in the detection of moving targets, is also described, and it is suggested that this may be of value in the study of defective colour vision.     

Overdiagnosis in early detection programs

Overdiagnosis refers to the situation where a screening exam detects a disease that would have otherwise been undetected in a person's lifetime. The disease would have not have been diagnosed because the individual would have died of other causes prior to its clinical onset. Although the probability of overdiagnosis is an important quantity for understanding early detection programs it has not been rigorously studied. We analyze an idealized early detection program and derive the mathematical expression for the probability of overdiagnosis. The results are studied numerically for prostate cancer and applied to a variety of screening schedules. Our investigation indicates that the probability of overdiagnosis is remarkably high.     

Cellular 'elementary organisms' in vitro. The early vision of Gottlieb Haberlandt and its realization

The botanist Gottlieb Haberlandt was the first who in 1898 (published in 1902) tried the systematic culture in vitro of single cells. His purpose was to study the mutual influences of cells as the last 'living units' within the multicellular body. Haberlandt visualized the theoretical potential of the culture approach in experimental plant morphology and physiology and nearly half a century was to elapse before his far-reaching ideas were realized. The history of cell and tissue culture is traced from the first concepts and origins in experimental embryology (1858/59) to its final verification almost 100 years later. The paper reminds of essential steps, theoretical backgrounds, accidental and planned discoveries and methodological approaches, and attempts to present all sources. The more or less known reviews of White (1931, 1936 and 1946), Fiedler (1938/39), Stapp (1947), Kandler (1948) and Street (1959) are summarized, and the mutual influence of plant and animal tissue culture on each other is discussed. Previous retrospective articles on the particular role of Haberlandt, namely those of Gautheret (1942) and Krikorian and Berquam (1969), are included in a general assessment of physiological anatomy in memory of the 50th anniversary of this great botanist's death.     

Area summation in human vision at and above detection threshold

The initial image-processing stages of visual cortex are well suited to a local (patchwise) analysis of the viewed scene. But the world's structures extend over space as textures and surfaces, suggesting the need for spatial integration. Most models of contrast vision fall shy of this process because (i) the weak area summation at detection threshold is attributed to probability summation (PS) and (ii) there is little or no advantage of area well above threshold. Both of these views are challenged here. First, it is shown that results at threshold are consistent with linear summation of contrast following retinal inhomogeneity, spatial filtering, nonlinear contrast transduction and multiple sources of additive Gaussian noise. We suggest that the suprathreshold loss of the area advantage in previous studies is due to a concomitant increase in suppression from the pedestal. To overcome this confound, a novel stimulus class is designed where: (i) the observer operates on a constant retinal area, (ii) the target area is controlled within this summation field, and (iii) the pedestal is fixed in size. Using this arrangement, substantial summation is found along the entire masking function, including the region of facilitation. Our analysis shows that PS and uncertainty cannot account for the results, and that suprathreshold summation of contrast extends over at least seven target cycles of grating.     

Feature detection in human vision: a phase-dependent energy model

This paper presents a simple and biologically plausible model of how mammalian visual systems could detect and identify features in an image. We suggest that the points in a waveform that have unique perceptual significance as 'lines' and 'edges' are the points where the Fourier components of the waveform come into phase with each other. At these points 'local energy' is maximal. Local energy is defined as the square root of the sum of the squared response of sets of matched filters, of identical amplitude spectrum but differing in phase spectrum by 90 degrees: one filter type has an even-symmetric line-spread function, the other an odd-symmetric line-spread function. For a line the main contribution to the local energy peak is in the output of the even-symmetric filters, whereas for edges it is in the output of the odd-symmetric filters. If both filter types respond at the peak of local energy, both edges and lines are seen, either simultaneously or alternating in time. The model was tested with a series of images, and shown to predict well the position of perceived features and the organization of the images.