Retinal scanning and visual inputs in freely walking crickets

ABSTRACT. Freely walking crickets were filmed from above during their visual orientation towards a black stripe. A frame-by-frame analysis enabled head and body movements to be recorded. The animals walk in 200ms bouts (runs) separated by pauses of similar duration. During each run, rotations of the body axis are observed and some corrections of the course direction occur between successive runs. Generally, the crickets do not walk straight ahead but slightly sideways. Because no lateral head movements were observed during visually orientated locomotion, retinal scanning results from both rotations of the body axis and translation of the head. While walking, one of the target edges is maintained by the cricket on a relatively limited area of the retina, generally between 10 and 25 laterally. Thus, the cricket often records three pieces of information about each edge: one in the monocular visual field, and two in the binocular visual field. Nevertheless, between two pauses, the images of each edge shift asymmetrically on the retinae. Such movement could prevent receptor adaptation by modulation of the ommatidial excitation, or by stimulation of the neighbouring ommatidia. It is also suggested that antennal movements are influenced by the positions of the visually fixated target edges.     

Quality of Visual Cue Affects Visual Reweighting in Quiet Standing

Sensory reweighting is a characteristic of postural control functioning adopted to accommodate environmental changes. The use of mono or binocular cues induces visual reduction/increment of moving room influences on postural sway, suggesting a visual reweighting due to the quality of available sensory cues. Because in our previous study visual conditions were set before each trial, participants could adjust the weight of the different sensory systems in an anticipatory manner based upon the reduction in quality of the visual information. Nevertheless, in daily situations this adjustment is a dynamical process and occurs during ongoing movement. The purpose of this study was to examine the effect of visual transitions in the coupling between visual information and body sway in two different distances from the front wall of a moving room. Eleven young adults stood upright inside of a moving room in two distances (75 and 150 cm) wearing a liquid crystal lenses goggles, which allow individual lenses transition from opaque to transparent and vice-versa. Participants stood still during five minutes for each trial and the lenses status changed every one minute (no vision to binocular vision, no vision to monocular vision, binocular vision to monocular vision, and vice-versa). Results showed that farther distance and monocular vision reduced the effect of visual manipulation on postural sway. The effect of visual transition was condition dependent, with a stronger effect when transitions involved binocular vision than monocular vision. Based upon these results, we conclude that the increased distance from the front wall of the room reduced the effect of visual manipulation on postural sway and that sensory reweighting is stimulus quality dependent, with binocular vision producing a much stronger down/up-weighting than monocular vision.     

Fix your eyes in the space you could reach: neurons in the macaque medial parietal cortex prefer gaze positions in peripersonal space

Interacting in the peripersonal space requires coordinated arm and eye movements to visual targets in depth. In primates, the medial posterior parietal cortex (PPC) represents a crucial node in the process of visual-to-motor signal transformations. The medial PPC area V6A is a key region engaged in the control of these processes because it jointly processes visual information, eye position and arm movement related signals. However, to date, there is no evidence in the medial PPC of spatial encoding in three dimensions. Here, using single neuron recordings in behaving macaques, we studied the neural signals related to binocular eye position in a task that required the monkeys to perform saccades and fixate targets at different locations in peripersonal and extrapersonal space. A significant proportion of neurons were modulated by both gaze direction and depth, i.e., by the location of the foveated target in 3D space. The population activity of these neurons displayed a strong preference for peripersonal space in a time interval around the saccade that preceded fixation and during fixation as well. This preference for targets within reaching distance during both target capturing and fixation suggests that binocular eye position signals are implemented functionally in V6A to support its role in reaching and grasping.     

Leopard frog priorities in choosing between prey at different locations

Frogs are able to respond to a prey stimulus throughout their 360° ground-level visual field as well as in the superior visual field. We compared the likelihood of frogs choosing between a more nasally located, ground-level prey versus a more temporally located ground-level prey, when the prey at the nasal location is further away from the frog. Two crickets were presented simultaneously at 9 pairs of angles that included both crickets in the binocular visual field, both crickets in the monocular visual field, or one cricket in the binocular field and one in the monocular field. Frogs chose the more nasally located prey at least 71% of the time when the more temporal prey was in the monocular field; and 64% of the time when both prey were in the binocular field. Frogs tended to choose the more nasally located prey, even though it takes the frog longer to reach the prey. In addition, when given a choice between a prey located at ground level versus a prey located in the superior field, frogs tend to choose the prey at ground-level. These results suggest that there is a neural mechanism that biases frogs’ responses to prey stimuli.     

Comparison of player movement patterns between 1-day and test cricket

Physical demands of cricket presumably vary by both game format and performance level. Differences in player movement patterns between 2 game formats (1 day and multiday) and 2 levels of elite performance (state and international) were quantified with global positioning system technology. Five movement categories were established, and 15 movement pattern variables were reported. Data from state (n = 42, 200 files) and international (n = 12, 63 files) cricketers were scaled to hourly values to compare movement demands. Cricketers generally covered similar distances in both formats, except for state 1-day fielders who covered moderately greater distance (~0.7 km·h?1 more; 21 ± 8%; mean ± 90% confidence interval) than state multiday (first-class) fielders. State 1-day cricketers also covered small to moderately greater distances (running 41 ± 13%; striding 38 ± 16%; sprinting 39 ± 36%) in the faster movement patterns and consequently had moderately less recovery time (13-67%) between high-intensity efforts as first-class cricketers. Comparisons of movements between performance levels revealed similar total distances between state and international cricketers. However, Test fielders covered moderately greater (29-48%) distances at the higher-intensity movement patterns (running, striding, and sprinting) than first-class fielders. In summary, although movement patterns were broadly similar between formats and levels, it appears that one day cricket (compared with multiday games) and test matches (compared with state-level competition) require more higher-intensity running. Conditioning coaches should train state and international 1-day cricket players similarly, but should account for the higher physical demands of international multiday cricket.     

On the role of attention in binocular rivalry: electrophysiological evidence

During binocular rivalry visual consciousness fluctuates between two dissimilar monocular images. We investigated the role of attention in this phenomenon by comparing event-related potentials (ERPs) when binocular-rivalry stimuli were attended with when they were unattended. Stimuli were dichoptic, orthogonal gratings that yielded binocular rivalry and dioptic, identically oriented gratings that yielded binocular fusion. Events were all possible orthogonal changes in orientation of one or both gratings. We had two attention conditions: In the attend-to-grating condition, participants had to report changes in perceived orientation, focussing their attention on the gratings. In the attend-to-fixation condition participants had to report changes in a central fixation target, taking attention away from the gratings. We found, surprisingly, that attending to rival gratings yielded a smaller ERP component (the N1, from 160-210 ms) than attending to the fixation target. To explain this paradoxical effect of attention, we propose that rivalry occurs in the attend-to-fixation condition (we found an ERP signature of rivalry in the form of a sustained negativity from 210-300 ms) but that the mechanism processing the stimulus changes is more adapted in the attend-to-grating condition than in the attend-to-fixation condition. This is consistent with the theory that adaptation gives rise to changes of visual consciousness during binocular rivalry.     

Subitization and attentional engagement by transient stimuli

A series of experiments investigated the visual selection of moving and static items during enumeration. Small numbers of visual targets can be enumerated with little increase in reaction time and error with set size, a process referred to as 'subitization'. The number of items that can be subitized' is typically between one and four and known as the subitization range. This study looked for evidence of subitizing of subsets of items presented on a computer display. Fast and accurate enumeration was found for random configurations of moving targets even when presented among static distracters. This was not the case for static targets presented among moving or transient distracters. RTs to these targets were longer and showed a steady increase in RT with target number, even in the subitization range. However, when static targets and moving distracters were presented foveally, fast enumeration/subitization of the static targets was again possible. This was not due to reduced inter-item spacing, since linear effects of the number of targets still emerged when stimuli were presented peripherally but the size-spacing ratio was matched to the foveal presentations. There was indication that instead performance reflected perceived differences in movement speed for stimuli presented in parafoveal and more peripheral retinal regions. In support of this, subitization of static items improved as the movement speed of the distracters increased. The data suggest that the processes supporting subitization are highly sensitive to dynamic stimuli and depend on the ease of segmentation between static and moving arrays.     

Detecting binocular 3D motion in static 3D noise: no effect of viewing distance

Relative binocular disparity cannot tell us the absolute 3D shape of an object, nor the 3D trajectory of its motion, unless the visual system has independent access to how far away the object is at any moment. Indeed, as the viewing distance is changed, the same disparate retinal motions will correspond to very different real 3D trajectories. In this paper we were interested in whether binocular 3D motion detection is affected by viewing distance. A visual search task was used, in which the observer is asked to detect a target dot, moving in 3D, amidst 3D stationary distractor dots. We found that distance does not affect detection performance. Motion-in-depth is consistently harder to detect than the equivalent lateral motion, for all viewing distances. For a constant retinal motion with both lateral and motion-in-depth components, detection performance is constant despite variations in viewing distance that produce large changes in the direction of the 3D trajectory. We conclude that binocular 3D motion detection relies on retinal, not absolute, visual signals.     

On the mechanism of prometaphase congression: Chromosome velocity as a function of position on the spindle

Rates of movement of univalents at prometaphase and of half-bivalents at anaphase in living cricket and grasshopper spermatocytes were determined as a function of the distance from the pole toward which the movement was directed. In the artificially produced univalents of cricket cells, correlation coefficients for rate versus distance form the pole were widely disparate from movement to movement and there was no consistent relationship between velocity and distance from the pole. However, in the naturally occurring univalents of grasshopper cells, there was a significant positive correlation between velocity and distance from the pole. In both cricket and grasshopper cells, there was no consistent correlation between rate of movement and distance from the pole for half-bivalents at anaphase. The prometaphase data from grasshopper cells support the simple hypothesis of Östergren (1950) that congression results from the application to chromosomes of forces which increase with increasing distance from the pole. Furthermore, these data are consistent with models of force production which suppose that the relationship between force (reflected as velocity) and distance from the pole is a linear one.     

Linking Perception,Cognition, and Action: Psychophysical Observations and Neural Network Modelling

It has been argued that perception, decision making, and movement planning are in reality tightly interwoven brain processes. However, how they are implemented in neural circuits is still a matter of debate. We tested human subjects in a temporal categorization task in which intervals had to be categorized as short or long. Subjects communicated their decision by moving a cursor into one of two possible targets, which appeared separated by different angles from trial to trial. Even though there was a 1 second-long delay between interval presentation and decision communication, categorization difficulty affected subjects’ performance, reaction (RT) and movement time (MT). In addition, reaction and movement times were also influenced by the distance between the targets. This implies that not only perceptual, but also movement-related considerations were incorporated into the decision process. Therefore, we searched for a model that could use categorization difficulty and target separation to describe subjects’ performance, RT, and MT. We developed a network consisting of two mutually inhibiting neural populations, each tuned to one of the possible categories and composed of an accumulation and a memory node. This network sequentially acquired interval information, maintained it in working memory and was then attracted to one of two possible states, corresponding to a categorical decision. It faithfully replicated subjects’ RT and MT as a function of categorization difficulty and target distance; it also replicated performance as a function of categorization difficulty. Furthermore, this model was used to make new predictions about the effect of untested durations, target distances and delay durations. To our knowledge, this is the first biologically plausible model that has been proposed to account for decision making and communication by integrating both sensory and motor planning information.     

Hawk Eyes I: Diurnal Raptors Differ in Visual Fields and Degree of Eye Movement

Background

Different strategies to search and detect prey may place specific demands on sensory modalities. We studied visual field configuration, degree of eye movement, and orbit orientation in three diurnal raptors belonging to the Accipitridae and Falconidae families.

Methodology/Principal Findings

We used an ophthalmoscopic reflex technique and an integrated 3D digitizer system. We found inter-specific variation in visual field configuration and degree of eye movement, but not in orbit orientation. Red-tailed Hawks have relatively small binocular areas (∼33°) and wide blind areas (∼82°), but intermediate degree of eye movement (∼5°), which underscores the importance of lateral vision rather than binocular vision to scan for distant prey in open areas. Cooper''s Hawks'' have relatively wide binocular fields (∼36°), small blind areas (∼60°), and high degree of eye movement (∼8°), which may increase visual coverage and enhance prey detection in closed habitats. Additionally, we found that Cooper''s Hawks can visually inspect the items held in the tip of the bill, which may facilitate food handling. American Kestrels have intermediate-sized binocular and lateral areas that may be used in prey detection at different distances through stereopsis and motion parallax; whereas the low degree eye movement (∼1°) may help stabilize the image when hovering above prey before an attack.

Conclusions

We conclude that: (a) there are between-species differences in visual field configuration in these diurnal raptors; (b) these differences are consistent with prey searching strategies and degree of visual obstruction in the environment (e.g., open and closed habitats); (c) variations in the degree of eye movement between species appear associated with foraging strategies; and (d) the size of the binocular and blind areas in hawks can vary substantially due to eye movements. Inter-specific variation in visual fields and eye movements can influence behavioral strategies to visually search for and track prey while perching.     

Prey orienting in frogs: Accounting for variations in output with stimulus distance

Summary We have studied the responses of leopard frogs,Rana pipiens, to live mealworms presented at different distances on the mid-sagittal plane. The response of normal frogs to stimuli at nearer distances consists of a direct snap whose amplitude increases with stimulus distance. For greater distances, the response consists of a forward hop whose amplitude also varies with stimulus distance. Over an intermediate range of distances, responses may be either snaps or hops. Whichever response occurs is of appropriate amplitude. The distance at which frogs switch from predominantly snapping responses to predominantly hopping responses increases with body size.Like normal frogs, unilaterally blinded frogs respond to stimuli at nearer distances with snaps whose amplitude varies with stimulus distance, switch from snapping to hopping over an intermediate range of distances, and respond to stimuli at greater distances with hops whose amplitude also increases with stimulus distance. In many cases, unilateral blinding did however result in a decrease in the distance at which the frogs switched from snapping to hopping. Such changes were not accompanied by the changes in snap or hop amplitude which would be expected if unilateral blinding resulted in generalized changes in distance judgement. Normal variations in snap amplitude and switches from snapping to hopping were also observed in frogs subjected to unilateral eye removal prior to the metamorphic eye migration which creates the adult binocular visual field.These results imply that neither distance discrimination nor any of the kinds of variation in motor output which occur with increasing stimulus distance necessarily depend on binocular cues. The behaviors studied also appear to be largely independent of normal binocular experience. More generally, our results suggest that the movement triggered by a stimulus at a particular location is not determined entirely by the retinal and superficial tectal region activated but rather reflects a combination of a retinal local sign signal with other kinds of information. The latter probably include signals related to stimulus distance and body posture, and may include signals related to body size as well.     

Scototaxis and shape discrimination in the female cricket Acheta domesticus in an arena and on a compensatory treadmill

ABSTRACT. The scototactic responses of adult female Acheta domesticus L. were tested toward various shaped targets in an orientation arena and on a compensatory treadmill. In an arena, crickets oriented toward dark targets (positive scototaxis) if they had horizontal visual angles ≥30, and if the vertical dimensions of the target was ≤ its horizontal dimension. Unattractive targets did not result in negative scototaxis but caused crickets to orient randomly with no net directionality. When complex targets were composed of two or more simple rectangular targets which had been previously defined as attractive or unattractive, intermediate responses were obtained. Crickets oriented less toward complex targets than toward simple attractive targets, but oriented more than toward simple unattractive targets. The responses of a female cricket toward an attractive target can be modified by the presence of a chemical signal emitted by previously tested females. When the signal is present females are less likely to orient toward a target that would otherwise be very attractive. This effect was not as great when the females being tested were taken directly from densely populated colonies.
When running on a compensatory treadmill, female crickets exhibit scototactic tendencies similar to those displayed in the arena. When tested on the treadmill over long periods, the amount of time spent orienting toward an attractive target increased.     

Evolution and Optimality of Similar Neural Mechanisms for Perception and Action during Search

A prevailing theory proposes that the brain''s two visual pathways, the ventral and dorsal, lead to differing visual processing and world representations for conscious perception than those for action. Others have claimed that perception and action share much of their visual processing. But which of these two neural architectures is favored by evolution? Successful visual search is life-critical and here we investigate the evolution and optimality of neural mechanisms mediating perception and eye movement actions for visual search in natural images. We implement an approximation to the ideal Bayesian searcher with two separate processing streams, one controlling the eye movements and the other stream determining the perceptual search decisions. We virtually evolved the neural mechanisms of the searchers'' two separate pathways built from linear combinations of primary visual cortex receptive fields (V1) by making the simulated individuals'' probability of survival depend on the perceptual accuracy finding targets in cluttered backgrounds. We find that for a variety of targets, backgrounds, and dependence of target detectability on retinal eccentricity, the mechanisms of the searchers'' two processing streams converge to similar representations showing that mismatches in the mechanisms for perception and eye movements lead to suboptimal search. Three exceptions which resulted in partial or no convergence were a case of an organism for which the targets are equally detectable across the retina, an organism with sufficient time to foveate all possible target locations, and a strict two-pathway model with no interconnections and differential pre-filtering based on parvocellular and magnocellular lateral geniculate cell properties. Thus, similar neural mechanisms for perception and eye movement actions during search are optimal and should be expected from the effects of natural selection on an organism with limited time to search for food that is not equi-detectable across its retina and interconnected perception and action neural pathways.     

Saccadic motor planning by integrating visual information and pre-information on neural dynamic fields

A functional model of target selection in the saccadic system is presented, incorporating elements of visual processing, motor planning, and motor control. We address the integration of visual information with pre-information. which is provided by manipulating the probability that a target appears at a certain location. This integration is achieved within a dynamic representation of planned eye movement which is modeled through distributions of activation on a topographic field. Visual input evokes activation, which is also constrained by lateral interaction within the field and by preshaping input representing pre-information. The model describes target selection observable in paradigms in which visual goals are presented at more than one location. Specifically, we model the transition from averaging, where endpoints of first saccades fall between two visual target locations, to decision making, where endpoints of first saccades fall accurately onto one of two simultaneously presented visual targets. We make predictions about how metrical biases of first saccades are induced by pre-information about target locations acquired by learning. When coupled to a motor control stage, activation dynamics on the planning level contribute to stabilizing gaze under fixation conditions. The neurophysiological relevance of our functional model is discussed.     

Visible binocular beats from invisible monocular stimuli during binocular rivalry

When two qualitatively different stimuli are presented at the same time, one to each eye, the stimuli can either integrate or compete with each other. When they compete, one of the two stimuli is alternately suppressed, a phenomenon called binocular rivalry [1,2]. When they integrate, observers see some form of the combined stimuli. Many different properties (for example, shape or color) of the two stimuli can induce binocular rivalry. Not all differences result in rivalry, however. Visual 'beats', for example, are the result of integration of high-frequency flicker between the two eyes [3,4], and are thus a binocular fusion phenomenon. It remains in dispute whether binocular fusion and rivalry can co-exist with one another [5-7]. Here, we report that rivalry and beats, two apparently opposing phenomena, can be perceived at the same time within the same spatial location. We hypothesized that the interocular difference in visual attributes that are predominantly processed in the Parvocellular pathway will lead to rivalry, and differences in visual attributes that are predominantly processed in the Magnocellular pathway tend to integrate. Further predictions based on this hypothesis were tested and confirmed.     

Seeing your error alters my pointing: observing systematic pointing errors induces sensori-motor after-effects

During the procedure of prism adaptation, subjects execute pointing movements to visual targets under a lateral optical displacement: as consequence of the discrepancy between visual and proprioceptive inputs, their visuo-motor activity is characterized by pointing errors. The perception of such final errors triggers error-correction processes that eventually result into sensori-motor compensation, opposite to the prismatic displacement (i.e., after-effects). Here we tested whether the mere observation of erroneous pointing movements, similar to those executed during prism adaptation, is sufficient to produce adaptation-like after-effects. Neurotypical participants observed, from a first-person perspective, the examiner's arm making incorrect pointing movements that systematically overshot visual targets location to the right, thus simulating a rightward optical deviation. Three classical after-effect measures (proprioceptive, visual and visual-proprioceptive shift) were recorded before and after first-person's perspective observation of pointing errors. Results showed that mere visual exposure to an arm that systematically points on the right-side of a target (i.e., without error correction) produces a leftward after-effect, which mostly affects the observer's proprioceptive estimation of her body midline. In addition, being exposed to such a constant visual error induced in the observer the illusion "to feel" the seen movement. These findings indicate that it is possible to elicit sensori-motor after-effects by mere observation of movement errors.     

Binocular information in the control of prehensile movements in multiple-object scenes

Recent evidence suggests that the visual control of prehension may be less dependent on binocular information than has previously been thought. Studies investigating this question, however, have generally only examined reaches to single objects presented in isolation, even though natural prehensile movements are typically directed at objects in cluttered scenes which contain many objects. The present study was designed, therefore, to assess the contribution of binocular information to the control of prehensile movements in multiple-object scenes. Subjects reached for and grasped objects presented either in isolation or in the presence of one, two or four additional 'flanking' objects, under binocular and monocular viewing conditions. So that the role of binocular information could be clearly determined, subjects made reaches both in the absence of a visible scene around the target objects (self-illuminated objects presented in the dark) and under normal ambient lighting conditions. Analysis of kinematic parameters indicated that the removal of binocular information did not significantly affect many of the major indices of the transport component, including peak wrist velocity. However, peak grip apertures increased and subjects spent more time in the final slow phase of movement, prior to grasping the object, during monocularly guided reaches. The dissociation between effects of binocular versus monocular viewing on transport and grasp parameters was observed irrespective of the presence of flanking objects. These results therefore further question the view that binocular vision is pre-eminent in the control of natural prehensile movements.     

Anteroposterior dynamic balance reactions induced by circular translation of the visual field

The anteroposterior sway of subjects under conditions of spontaneous dynamic balance on a wobbly platform was measured during visual stimulation by a visual target executing a circular trajectory in the frontal plane. The target was either a component of the whole moving visual scene or moving on a stationary background. With the former stimulation, obtained through the use of rotating prismatic glasses, every point of the visual field appeared to describe a circular trajectory around its real position so that the whole visual field apeared to be circularly translated, undistorted, inducing a binocular pursuit movement. Under these conditions, stereotyped anteroposterior dynamic balance reactions synchronous with the position of the stimulus were elicited. The latter stimulation consisted of pursuing a luminous target describing a trajectory similar to that of the fixation point seen through the rotating prisms on the same, this time stable, visual background. Although pursuit eye movements were comparable, as demonstrated by electro-oculographic recordings, no stereotyped equilibration reaction was induced. It is concluded that the translatory motion of the background image on the retina in the latter experiments contributed to the body's stability as well as to the perception of a stable environment.     

The role of binocular information in the 'on-line' control of prehension

Binocular visual information may be involved in the selection of appropriate motor programs before a reach is executed or it may be involved during the movement-execution phase in order to monitor and guide the hand to the target object. Here we introduced binocular information after 0%, 25%, 50% or 75% of the movement-execution phase and determined its effects on the kinematic indices of prehensile movements made to objects of different sizes placed at different distances. Kinematic indices linked to the transport component, such as peak velocity and time-to-peak velocity, were unaffected by the presence of binocular cues whereas later occurring indices, such as peak grip aperture and time in the slow phase, were significantly affected. Although the magnitude of the peak grip was affected by the presence of binocular cues, the time at which it occurred did not change. This pattern of results suggest that the visuo-motor control of prehensile movements utilises both feedforward and feedback strategies and that binocular cues are particularly important for the fine manual adjustments typical of the latter.