The consistency of element transformations affects the visibility but not the direction of illusory motion

Two experiments tested whether the consistency of element transformations affected perceptions of long-range apparent motion. Vertical lines were used to generate apparent motion against one of two different backgrounds, control and depth. Consistency was manipulated by changing the size of the vertical lines. In some displays, the size of the vertical lines remained constant during movement. In other displays, the size of the vertical lines changed during movement. Consistent movement occurred when the size manipulation was in agreement with the type of background used. In Experiment I, points of subject equality for the quality of motion relative to a standard display were measured. These PSEs indicated that consistent movement (e.g., line sizes held constant for control background displays) was more visible than inconsistent movement (e.g., line sizes constant for depth background displays). In Experiment II, a motion competition display was used to measure thresholds for perceived direction of motion. The depth background was used to make motion in one direction more consistent than motion in the opposite direction. However, no significant differences were noted between thresholds obtained in this condition and those obtained in a control condition. Thus the consistency of element transformations affected the quality of motion, but did not affect the perceived direction of motion. These results are consistent with Ullman's (The Interpretation of Visual Motion, MIT Press, 1979) two-component theory of apparent motion.     

The Effect of Visual Apparent Motion on Audiovisual Simultaneity

Visual motion information from dynamic environments is important in multisensory temporal perception. However, it is unclear how visual motion information influences the integration of multisensory temporal perceptions. We investigated whether visual apparent motion affects audiovisual temporal perception. Visual apparent motion is a phenomenon in which two flashes presented in sequence in different positions are perceived as continuous motion. Across three experiments, participants performed temporal order judgment (TOJ) tasks. Experiment 1 was a TOJ task conducted in order to assess audiovisual simultaneity during perception of apparent motion. The results showed that the point of subjective simultaneity (PSS) was shifted toward a sound-lead stimulus, and the just noticeable difference (JND) was reduced compared with a normal TOJ task with a single flash. This indicates that visual apparent motion affects audiovisual simultaneity and improves temporal discrimination in audiovisual processing. Experiment 2 was a TOJ task conducted in order to remove the influence of the amount of flash stimulation from Experiment 1. The PSS and JND during perception of apparent motion were almost identical to those in Experiment 1, but differed from those for successive perception when long temporal intervals were included between two flashes without motion. This showed that the result obtained under the apparent motion condition was unaffected by the amount of flash stimulation. Because apparent motion was produced by a constant interval between two flashes, the results may be accounted for by specific prediction. In Experiment 3, we eliminated the influence of prediction by randomizing the intervals between the two flashes. However, the PSS and JND did not differ from those in Experiment 1. It became clear that the results obtained for the perception of visual apparent motion were not attributable to prediction. Our findings suggest that visual apparent motion changes temporal simultaneity perception and improves temporal discrimination in audiovisual processing.     

Interaction of perceptual grouping and crossmodal temporal capture in tactile apparent-motion

Previous studies have shown that in tasks requiring participants to report the direction of apparent motion, task-irrelevant mono-beeps can "capture" visual motion perception when the beeps occur temporally close to the visual stimuli. However, the contributions of the relative timing of multimodal events and the event structure, modulating uni- and/or crossmodal perceptual grouping, remain unclear. To examine this question and extend the investigation to the tactile modality, the current experiments presented tactile two-tap apparent-motion streams, with an SOA of 400 ms between successive, left-/right-hand middle-finger taps, accompanied by task-irrelevant, non-spatial auditory stimuli. The streams were shown for 90 seconds, and participants' task was to continuously report the perceived (left- or rightward) direction of tactile motion. In Experiment 1, each tactile stimulus was paired with an auditory beep, though odd-numbered taps were paired with an asynchronous beep, with audiotactile SOAs ranging from -75 ms to 75 ms. Perceived direction of tactile motion varied systematically with audiotactile SOA, indicative of a temporal-capture effect. In Experiment 2, two audiotactile SOAs--one short (75 ms), one long (325 ms)--were compared. The long-SOA condition preserved the crossmodal event structure (so the temporal-capture dynamics should have been similar to that in Experiment 1), but both beeps now occurred temporally close to the taps on one side (even-numbered taps). The two SOAs were found to produce opposite modulations of apparent motion, indicative of an influence of crossmodal grouping. In Experiment 3, only odd-numbered, but not even-numbered, taps were paired with auditory beeps. This abolished the temporal-capture effect and, instead, a dominant percept of apparent motion from the audiotactile side to the tactile-only side was observed independently of the SOA variation. These findings suggest that asymmetric crossmodal grouping leads to an attentional modulation of apparent motion, which inhibits crossmodal temporal-capture effects.     

Characteristics of the flux dimension cue in apparent motion correspondence.

It has been shown that element flux and size (but not luminance) serve as correspondence cues in the apparent motion visual system. Results are now presented of a study of the characteristics of the flux cue. It was found that flux rather than luminance is used by the system even when the size of the elements is greater than the size limit of Ricco's law. There were interactions between the apparent motion processing of the size and flux dimensions, beyond the obvious dependence of flux on size: positively correlated size and flux differences between elements have a greater effect on correspondence than do negatively correlated differences. Finally, when comparing the fluxes of different elements, the apparent motion system uses relative flux (above or below background) rather than absolute flux (relative to zero).     

Developmental features of protophloem sieve elements in roots of wheat (Triticum aestivum L.)

Summary Protophloem sieve elements (PSEs) in roots of wheat (Triticum aestivum L.) are arranged in single vertical files. The number of PSEs within the files increases by symmetrical divisions, which take place after the completion of asymmetrical (formative) divisions and before the initiation of differentiation. The divisions are preceded by well defined pre-prophase bands (PPB) of microtubules, which surround the nucleus in an equatorial position. In the cytoplasmic region between the nuclear surface and the PPB, perinuclear and endoplasmic microtubules were observed. The perinuclear microtubules are considered as part of the developing spindle, while the endoplasmic ones interlink the perinuclear microtubules with the PPB. Dividing cells do not show any signs of incipient differentiation. The first and most reliable indication of a commencing differentiation is provided by the sieve-element plastids that begin to accumulate dense crystalloid inclusions in the very young PSEs. In mature PSEs plastids contain two kinds of crystalloid inclusions, dense and thin, in a translucent stroma. Depending on the plastid-inclusions criterion it was shown that: (a) the PSEs of a given root do not initiate differentiation at exactly the same stage, (b) the developmental sequence extends to a span of 7–9 actively differentiating PSEs arranged in a single vertical file, and (c) each PSE needs about 16–21 h to pass through the whole developmental sequence. In the last two differentiating PSEs of a file, mitochondria were found to be enveloped by single cisternae of ER. The association is temporary as it is lost in the first PSEs with an autolysed lumen. During differentiation, Golgi bodies were abundant and active in producing vesicles involved in cell wall development. Golgi vesicles were also found among the microtubules of the PPB, but no local thickening was observed. Golgi bodies disorganize in the last stages of autolysis and disappear in mature sieve elements.Abbreviations ER endoplasmic reticulum - MSE metaphloem sieve element - PPB pre-prophase band - PSE protophloem sieve element Dedicated to Prof. Dr. Dr. h.c. Eberhard Schnepf on the occasion of his retirement     

High temporal frequency synchrony is insufficient for perceptual grouping

We used textures of randomly moving grating patches to assess the role of fine-grain temporal synchrony in texture segregation. In the target area, patches reversed direction simultaneously. In the surround, patches changed direction at random times. Thus, phase changes in the target area were precisely synchronous, whereas those in the surround were not. In agreement with work carried out by Lee and Blake, we found that the target area was frequently visible, and that observers could discriminate its shape (horizontal versus vertical) at frame rates of 100 Hz in brief exposures (200 ms). Further experiments suggested that the length of unidirectional motion sequences in the target area, rather than synchrony, determined its visibility. To eliminate completely contrast and motion cues, we made all the background elements identical to the target elements, but with a random starting phase. Despite the presence of synchrony in the target area but not the background, the target was generally very hard to see. Targets that remained visible contained low temporal frequency modulations of direction. We conclude that the human observer can detect synchrony, but only at modest temporal frequencies once motion and contrast artefacts have been eliminated.     

The effect of contour closure on shape perception

We studied psychophysically whether 'contour closure' enhances the accuracy of shape perception. Stimulus configurations (presented on a blank background) always consisted of identical pattern elements, but the positions of the local elements were varied: the global contour shape either contained closure or not. In the first two stimulus conditions (Closure), the oriented pattern elements (Gabor patches) formed a 'closed' rectangular shape composed of either four long lines or four corners. In the third condition (No closure), the global shape was composed of the four corners, but they were outward oriented, and hence they did not form the outline of a closed contour. We measured the precision of shape perception using a discrimination task in which observers judged the aspect ratio of the outline shape i.e. whether the rectangular shape was tall or wide. We found that: (i) shape discrimination was better (more precise) for Closed contours than for Non-closed contours, i.e. the aspect ratio discrimination thresholds were lower for the Closed than Non-closed configurations. The improved performance could not be explained by differences in visibility of the local elements in the two conditions. (ii) For closed contours, shape discrimination was more precise when the local elements were aligned with the global shape, than when the local elements were orthogonal to it.     

Unconscious Local Motion Alters Global Image Speed

Accurate motion perception of self and object speed is crucial for successful interaction in the world. The context in which we make such speed judgments has a profound effect on their accuracy. Misperceptions of motion speed caused by the context can have drastic consequences in real world situations, but they also reveal much about the underlying mechanisms of motion perception. Here we show that motion signals suppressed from awareness can warp simultaneous conscious speed perception. In Experiment 1, we measured global speed discrimination thresholds using an annulus of 8 local Gabor elements. We show that physically removing local elements from the array attenuated global speed discrimination. However, removing awareness of the local elements only had a small effect on speed discrimination. That is, unconscious local motion elements contributed to global conscious speed perception. In Experiment 2 we measured the global speed of the moving Gabor patterns, when half the elements moved at different speeds. We show that global speed averaging occurred regardless of whether local elements were removed from awareness, such that the speed of invisible elements continued to be averaged together with the visible elements to determine the global speed. These data suggest that contextual motion signals outside of awareness can both boost and affect our experience of motion speed, and suggest that such pooling of motion signals occurs before the conscious extraction of the surround motion speed.     

The “lunching” effect: Pigeons track motion towards food more than motion away from it

Four experiments measured pigeons’ pecking at a small touch-screen image (the CS) that moved towards or away from a source of food (the US). The image's effectiveness as a CS was dependent on its motion, direction, and distance relative to the US. Pecking to the CS increased with proximity to the US when the CS moved towards the US (Experiment 1). This held true even when a departing CS signalled a US of greater magnitude (Experiment 2). Response rates to stationary stimuli were greater the closer they were to the hopper; but rate was less than when the same spot was part of a motion towards food, and greater than when it was part of a motion away (Experiment 3). The rate of responding in all three cases (motion towards, stationary, motion away) decreased exponentially with distance from the hopper. The distance and motion effects observed under these Pavlovian contingencies were different when pecking to the spot was required for reinforcement (Experiment 4).     

The influence of substrate on the functional response of an avian granivore and its implications for farmland bird conservation

Few studies to date have considered the effect of substrate on the functional response of an organism feeding on prey of varying visibility. Intake rates of lone captive canaries, Serinus canarius L., were measured at varying seed densities on patches of either earth or short grass (<1 cm). Experiment 1, using pale seeds, found intake rates were significantly higher and search times significantly lower on earth than on grass. Two measures of crypticity (contrast in light reflectance as measured using a spectrophotometer and an experiment with humans) found pale seeds to be more visible on earth. The results from experiment 1 could be explained by this difference in crypsis. Experiment 2 used identical seeds to those in experiment 1 except they were dyed to match their backgrounds. The two measures of crypticity both found that black seeds were less visible on earth than green seeds were on grass. However, intake rates were still significantly higher on earth than grass. Seed colour preference, vegetation impeding movement, and differences in vigilance rates or seed accessibility could not explain this result. We discuss three other potentially explanatory mechanisms, the most likely of which was the greater surface area needed for scanning created by the structure of grass. Crucially, regardless of the mechanism(s) involved, many vegetated substrates share similar properties with grass (structural complexity and shiny surfaces which reflect light) and so the outcome of our findings are likely to extend to many natural situations. Conservationists wishing to encourage granivorous birds should consider enhancing food accessibility by providing uniform substrates, such as bare earth, for them to forage on. In addition, behaviour-based models should incorporate the effects of habitat into their equations of the functional response.     

Do variables that affect similar bistable apparent-movement displays result in similar changes in perception?

Two bistable apparent-movement displays (i.e. ones that generate two qualitatively different kinds of movement percepts under different conditions) were compared. They were designed to be as similar as possible spatially, and were studied with identical stimulus manipulations to see whether changes in balance between their bistable percepts would be similar. Results show that the two displays had different response characteristics to the same stimulus manipulations. Two models of motion perception that have previously predicted at least one kind of bistable apparent motion were considered in terms of how well they address the current data. As yet, neither model has been shown to predict the motion states and bistable behavior of the two displays studied here. It is concluded that results of the type described here (specifically, differences in the psychophysical functions yielded by two structurally similar but qualitatively different bistable displays) present a challenge for theories of motion perception.     

Flight Durations in Bumblebees Under Manipulation of Feeding Choices

Foraging bees spend less time flying between flowers of the same species than between individuals of different species. This time saving has been suggested as a possible advantage of flower-constant foraging. We hypothesized that the time required to switch flower type increases if (a) such switches are infrequent and (b) the bees need to decide whether to switch or not. Bumblebees were taught to forage on artificial feeders that were identical in morphology and reward schedule but differed in the color of their landing surface. In the first two experiments bees foraged alternatively between two feeders. In Experiment 1 the colors of the landing surfaces were switched every two or three visits, while in Experiment 2 they were switched every six or seven visits. In the third experiment, the bees were required to decide whether to make a color-constant or a color-shift flight. Intervisit time was defined as time elapsed between consecutive visits to feeders. When feeder colors were changed frequently (Experiment 1), we detected no difference between color-constant and color-shift intervisit times. When bees were repeatedly exposed to one color (Experiment 2), color shifts required a significantly longer time. When allowed to choose (Experiment 3), bees performed more color-constant flights than color-shift flights. Intervisit times were similar for color-constant and color-shift flights in this experiment. Intervisit times in Experiment 3 were significantly longer than in Experiment 2 and slightly but nonsignificantly longer than in Experiment 1. The results suggest that bees indeed save time though flower-constant foraging but that this time savings is a small (1 s/flower visit) under laboratory conditions, and appears only when switches between flower types are infrequent. The time saved may be more significant over long foraging trips, and when morphological differences between flower species are large, as often happens under natural conditions, providing a selective advantage to flower-constant foraging.     

A Comparison of Different Informative Vibrotactile Forward Collision Warnings: Does the Warning Need to Be Linked to the Collision Event?

Recent research demonstrates that auditory and vibrotactile forward collision warnings presenting a motion signal (e.g., looming or apparent motion across the body surface) can facilitate speeded braking reaction times (BRTs). The purpose of the present study was to expand on this work by directly comparing warning signals in which the motion conveyed was constant across all collision events with signals in which the speed of motion was dependent on the closing velocity (CV). Two experiments were conducted using a simulated car-following task and BRTs were measured. In Experiment 1, increasing intensity (looming) vibrotactile signals were presented from a single tactor attached to the driver''s waist. When the increase in intensity was CV-linked, BRTs were significantly faster as compared to a no-warning condition, however, they were not significantly different from constant intensity and CV-independent looming warnings. In Experiment 2, a vertical array of three tactors was used to create motion either towards (upwards) or away (downwards) from the driver''s head. When the warning signal presented upwards motion that was CV-linked, BRTs were significantly faster than all other warning types. Downwards warnings led to a significantly higher number of brake activations in false alarm situations as compared to upwards moving warnings. The effectiveness of dynamic tactile collision warnings would therefore appear to depend on both the link between the warning and collision event and on the directionality of the warning signal.     

The perception of apparent movement

When two similar pictures, overlapping but slightly displaced, were projected on a screen in alternation, apparent movement could be seen. How similar must successive pictures be to give apparent movement? This is the 'correspondence problem'. Manipulations of the local and global correspondences between pictures included motion phenomena such as reversed apparent movement; a four-stroke oscillatory cycle which gave an illusion of continuous motion in one direction; edges defined by texture, stereoscopic depth, or flicker, kinetic edges; and wave motion. It was concluded that human motion perception may comprise two separate mechanisms. Local point-by-point correlations between pictures are detected by a relatively peripheral system, probably based on directonally selective neural units. More subtle global correspondences are analysed by a more cognitive system which extracts edges before it process motion.     

Both Physical Exercise and Progressive Muscle Relaxation Reduce the Facing-the-Viewer Bias in Biological Motion Perception

Biological motion stimuli, such as orthographically projected stick figure walkers, are ambiguous about their orientation in depth. The projection of a stick figure walker oriented towards the viewer, therefore, is the same as its projection when oriented away. Even though such figures are depth-ambiguous, however, observers tend to interpret them as facing towards them more often than facing away. Some have speculated that this facing-the-viewer bias may exist for sociobiological reasons: Mistaking another human as retreating when they are actually approaching could have more severe consequences than the opposite error. Implied in this hypothesis is that the facing-towards percept of biological motion stimuli is potentially more threatening. Measures of anxiety and the facing-the-viewer bias should therefore be related, as researchers have consistently found that anxious individuals display an attentional bias towards more threatening stimuli. The goal of this study was to assess whether physical exercise (Experiment 1) or an anxiety induction/reduction task (Experiment 2) would significantly affect facing-the-viewer biases. We hypothesized that both physical exercise and progressive muscle relaxation would decrease facing-the-viewer biases for full stick figure walkers, but not for bottom- or top-half-only human stimuli, as these carry less sociobiological relevance. On the other hand, we expected that the anxiety induction task (Experiment 2) would increase facing-the-viewer biases for full stick figure walkers only. In both experiments, participants completed anxiety questionnaires, exercised on a treadmill (Experiment 1) or performed an anxiety induction/reduction task (Experiment 2), and then immediately completed a perceptual task that allowed us to assess their facing-the-viewer bias. As hypothesized, we found that physical exercise and progressive muscle relaxation reduced facing-the-viewer biases for full stick figure walkers only. Our results provide further support that the facing-the-viewer bias for biological motion stimuli is related to the sociobiological relevance of such stimuli.     

Investigating the status of biological stimuli as objects of attention in multiple object tracking

Background

Humans are able to track multiple simultaneously moving objects. A number of factors have been identified that can influence the ease with which objects can be attended and tracked. Here, we explored the possibility that object tracking abilities may be specialized for tracking biological targets such as people.

Methodology/Principal Findings

We used the Multiple Object Tracking (MOT) paradigm to explore whether the high-level biological status of the targets affects the efficiency of attentional selection and tracking. In Experiment 1, we assessed the tracking of point-light biological motion figures. As controls, we used either the same stimuli or point-light letters, presented in upright, inverted or scrambled configurations. While scrambling significantly affected performance for both letters and point-light figures, there was an effect of inversion restricted to biological motion, inverted figures being harder to track. In Experiment 2, we found that tracking performance was equivalent for natural point-light walkers and ‘moon-walkers’, whose implied direction was incongruent with their actual direction of motion. In Experiment 3, we found higher tracking accuracy for inverted faces compared with upright faces. Thus, there was a double dissociation between inversion effects for biological motion and faces, with no inversion effect for our non-biological stimuli (letters, houses).

Conclusions/Significance

MOT is sensitive to some, but not all naturalistic aspects of biological stimuli. There does not appear to be a highly specialized role for tracking people. However, MOT appears constrained by principles of object segmentation and grouping, where effectively grouped, coherent objects, but not necessarily biological objects, are tracked most successfully.     

Load-displacement properties of lower cervical spine motion segments

The load-displacement behavior of 35 fresh adult cervical spine motion segments was measured in compression, shear, flexion, extension, lateral bending and axial torsion tests. Motion segments were tested both intact and with posterior elements removed. Applied forces ranged to 73.6 N in compression and to 39 N in shear, while applied moments ranged to 2.16 Nm. For each mode of loading, principal and coupled motions were measured and stiffnesses were calculated. The effect of disc degeneration on motion segment stiffnesses and the moments required for motion segment failure were also measured. In compression, the stiffnesses of the cervical motion segments were similar to those of thoracic and lumbar motion segments. In other modes of loading, cervical stiffnesses were considerably smaller than thoracic or lumbar stiffnesses. Removal of the posterior elements decreased cervical motion segment stiffnesses by as much as 50%. Degenerated cervical discs were less stiff in compression and stiffer in shear than less degenerated discs, but in bending or axial torsion, no statistically significant differences were evident. Bending moments causing failure were an order of magnitude lower than those for lumbar segments.     

Exposure to Organic Solvents Used in Dry Cleaning Reduces Low and High Level Visual Function

PurposeTo investigate whether exposure to occupational levels of organic solvents in the dry cleaning industry is associated with neurotoxic symptoms and visual deficits in the perception of basic visual features such as luminance contrast and colour, higher level processing of global motion and form (Experiment 1), and cognitive function as measured in a visual search task (Experiment 2).MethodsThe Q16 neurotoxic questionnaire, a commonly used measure of neurotoxicity (by the World Health Organization), was administered to assess the neurotoxic status of a group of 33 dry cleaners exposed to occupational levels of organic solvents (OS) and 35 age-matched non dry-cleaners who had never worked in the dry cleaning industry. In Experiment 1, to assess visual function, contrast sensitivity, colour/hue discrimination (Munsell Hue 100 test), global motion and form thresholds were assessed using computerised psychophysical tests. Sensitivity to global motion or form structure was quantified by varying the pattern coherence of global dot motion (GDM) and Glass pattern (oriented dot pairs) respectively (i.e., the percentage of dots/dot pairs that contribute to the perception of global structure). In Experiment 2, a letter visual-search task was used to measure reaction times (as a function of the number of elements: 4, 8, 16, 32, 64 and 100) in both parallel and serial search conditions.ResultsDry cleaners exposed to organic solvents had significantly higher scores on the Q16 compared to non dry-cleaners indicating that dry cleaners experienced more neurotoxic symptoms on average. The contrast sensitivity function for dry cleaners was significantly lower at all spatial frequencies relative to non dry-cleaners, which is consistent with previous studies. Poorer colour discrimination performance was also noted in dry cleaners than non dry-cleaners, particularly along the blue/yellow axis. In a new finding, we report that global form and motion thresholds for dry cleaners were also significantly higher and almost double than that obtained from non dry-cleaners. However, reaction time performance on both parallel and serial visual search was not different between dry cleaners and non dry-cleaners.ConclusionsExposure to occupational levels of organic solvents is associated with neurotoxicity which is in turn associated with both low level deficits (such as the perception of contrast and discrimination of colour) and high level visual deficits such as the perception of global form and motion, but not visual search performance. The latter finding indicates that the deficits in visual function are unlikely to be due to changes in general cognitive performance.     

Sensory response patterns and the evolution of visual signal design in anoline lizards

 The evolutionary relationship between visual system response and visual signal design was investigated in four species of anoline lizards which occupy distinctly different habitats. Anoles display with motion patterns of a colorful throat fan called the dewlap. We assessed signal visibility by recording evoked potentials from the optic tectum in response to a moving stimulus flag (a dewlap-like stimulus), and, in one species, by testing behavioral response. The motion pattern, intensity and spectral quality of the stimulus flag, and the background against which it was viewed, were independently manipulated. In all cases, high-velocity motion patterns with a high percentage of brightness contrast between stimulus and background produced the greatest response. Differences in spectral quality between stimulus and background (color contrast) had no effect on tectal responses, but did influence the behaviorally measured detection probability. Using habitat light data we estimated the visibility of the dewlap of each species in different natural habitats. Each species' dewlap was highly visible in its own habitat, but some were much less visible in the habitats of some other species. Habitat light conditions and visual system response properties appear to have constrained the evolution of dewlap design, in at least some of the species. Accepted: May 1998     

Identification of a DNA structural motif that includes the binding sites for Sp1, p53 and GA-binding protein.

We have analyzed predicted helical twist angles in the 21-bp repeat region of the SV40 genome, using a semi-empirical model previously shown to accurately predict backbone conformations. Unexpectedly, the pattern of twist angles characteristic of the six GC-boxes is repeated an additional five times at positions that are regularly interspersed with the six GC-box sequences. These patterns of helical twist angles are associated with a second, imperfectly-repeated sequence motif, the TR-box 5'-RRNTRGG. Unrelated DNA sequences that interact with trans-acting factors (p53 and GABP) exhibit similar twist angle patterns, due to elements of the general form 5'-RRRYRRR that occur as interspersed arrays with a spacing of 10-11 bp and an offset of 4-6 bp. Arrays of these elements, which we call pyrimidine sandwich elements (PSEs), may play an important role in the interaction of trans-acting factors with DNA control regions. In 13 human proto-oncogenes analyzed, we identified 31 PSE arrays, 11 of which were in the 5'-flanking regions of the genes. The most extensive array was found in the promoter region of the K-ras gene. Extending over 80 bp of DNA, it contained 16 PSEs that showed an average deviation from the SV40 criterion pattern of angles of only 1.2 degrees.