Visual deprivation and distance estimation in the praying mantis larva

Abstract. Young larvae of the praying mantis, Tenodera sinensis Saussure, were placed on an off-centre island surrounded by a round arena with six black bars painted on a white inner wall. In this situation, it was shown that the horizontal peering movements of the head often seen in mantids are in fact used to measure distances; motion parallax may be involved in this process. Aimed jumps that followed peering were taken to be the distinct result of an absolute distance measurement. Specific visual deprivation such as painting over of certain parts of the eye with opaque black varnish or degeneration of the fovea with sulforhodamine showed that: absolute evaluation of distance is only possible with two fully intact eyes; the peering mechanism is under visual control; and visual experience has a long-term effect on distance measurement involving peering movements.     

Similarities and differences in the peering-jump behavior of three grasshopper species (Orthoptera: Caelifera)

The peering-jump behavior was studied for the common field grasshopper Chorthippus brunneus , the meadow grasshopper C. parallelus and the alpine grasshopper Miramella alpina (Orthoptera, Caelifera). It was found that immediately before jumping M. alpina executes primarily unilateral object-related peering movements, with approximately twice the amplitude and velocity of the predominantly bilateral object-related peering movements of the other two species. Whereas M. alpina almost always jumped toward the black stripes in the experimental arena, the other species jumped toward both the black stripes and the white spaces between them. All three species preferred the same pattern of black stripes, which permitted them to view one black stripe frontally, with an additional black stripe to the left and right, in the lateral visual field. The similarities and differences in the peering-jump behavior of the three grasshopper species is discussed with regard to visual perception (parallax cues) and environmental adaptation.     

Responses of the Toad Bufo bufo (L.) to Stationary Prey Stimuli1

In the present work toads (Bufo bufo) are shown to respond with prey catching to stationary dummies without previous or accompanying visual or olfactory stimulation. The subjects very rarely showed jerky head movements which, therefore, cannot be necessary for perception of stationary objects. Size preference with respect to stationary stimuli is about the same as in experiments with moving stimuli. However, differences exist between the effects of stationary and moving stimuli with respect to shape and orientation. If a square measuring 10 times 10 mm and a rectangle measuring 5 times 20 mm, oriented either horizontally or vertically, are presented within the frontal-vertical plane, the square is preferred to the rectangles, and among these the horizontal rectangle is to the vertical one. This latter preference is due to the negative effect of the vertical extension: If the vertical rectangle is reduced in length, it becomes more effective as compared to the horizontal rectangle. In the horizontal (X-Z) plane the square and the rectangle oriented parallel to the Z-axis are equally superior to the bar oriented parallel to the X-axis. At presentation of a pair of stimuli in both planes, the one in the frontal-vertical plane is always preferred to that in the horizontal plane. Correspondences and differences of these results to those from experiments with moving prey dummies are discussed.     

Motion parallax as a source of distance information in locusts and mantids

This review article is devoted to results on distance measurement in locusts (e.g., Wallace, 1959; Collett, 1978; Sobel, 1990) and mantids. Before locusts or mantids jump toward a stationary object, they perform characteristic pendulum movements with the head or body, called peering movements, in the direction of the object. The fact that the animals over- or underestimate the distance to the object when the object is moved with or against the peering movement, and so perform jumps that are too long or short, would seem to indicate that motion parallax is used in this distance measurement. The behavior of the peering parameters with different object distances also indicates that not only retinal image motion but also the animal’s own movement is used in calculating the distance.     

Motion detection in the presence and absence of background motion in anAnolis lizard

Anolis lizards respond to a moving object viewed in the periphery of their visual field by turning their eye to fixate the object with their central fovea. This paper describes the relative effectiveness of different patterns of motion of a small black lure in eliciting these eye movements and the way motion of a backdrop of vegetation affects the response. The stimulus was positioned 45 degrees from the animal's line of gaze and oscillated in the vertical axis at different frequencies between 0.5 and 10 Hz. At each frequency, the amplitude of the oscillation was increased until the lizard flicked its eye towards the stimulus. The minimum amplitude needed for response (0.22 degrees of visual angle) was independent of frequency and waveform. The probability of any response occurring was, however, lower at higher frequencies (7 and 10 Hz) and a 1.5 Hz square wave evoked the greatest proportion of responses. Sinusoidal oscillation of a background of vegetation at 1.6 Hz during or before motion of the stimulus lure reduced the probability of an eye flick but did not raise the minimum amplitude needed for a response. The suppressive effect was greatest when the lure was oscillated at frequencies close to that of the background. It is concluded that Anolis, which rely upon motion to detect objects in the periphery of the visual field, filter out irrelevant motion such as that of windblown vegetation by responding preferentially to particular patterns of motion and short term habituation to commonly present patterns of motion.     

Tuning Properties of MT and MSTd and Divisive Interactions for Eye-Movement Compensation

The primate brain intelligently processes visual information from the world as the eyes move constantly. The brain must take into account visual motion induced by eye movements, so that visual information about the outside world can be recovered. Certain neurons in the dorsal part of monkey medial superior temporal area (MSTd) play an important role in integrating information about eye movements and visual motion. When a monkey tracks a moving target with its eyes, these neurons respond to visual motion as well as to smooth pursuit eye movements. Furthermore, the responses of some MSTd neurons to the motion of objects in the world are very similar during pursuit and during fixation, even though the visual information on the retina is altered by the pursuit eye movement. We call these neurons compensatory pursuit neurons. In this study we develop a computational model of MSTd compensatory pursuit neurons based on physiological data from single unit studies. Our model MSTd neurons can simulate the velocity tuning of monkey MSTd neurons. The model MSTd neurons also show the pursuit compensation property. We find that pursuit compensation can be achieved by divisive interaction between signals coding eye movements and signals coding visual motion. The model generates two implications that can be tested in future experiments: (1) compensatory pursuit neurons in MSTd should have the same direction preference for pursuit and retinal visual motion; (2) there should be non-compensatory pursuit neurons that show opposite preferred directions of pursuit and retinal visual motion.     

The Visual Orientation Strategies of Mantis religiosa and Empusa fasciata Reflect Differences in the Structure of Their Visual Surroundings

In the present study, peering behaviour, which is used to measure distance by the image motion caused by head movement, is examined in two types of mantid. Mantis religiosa inhabits a region of dense grass consisting of uniform, generally uniformly aligned, and closely spaced elements and executes slow, simple peering movements. In contrast, Empusa fasciata climbs about in open regions of shrubs and bushes which consist of irregular, variably aligned and variably spaced elements and it executes comparatively quick, complex peering movements. Hence, it seems that in these two species of mantid, the same orientation mechanism has been adapted to the unique structures of their visual surroundings. Apparently M. religiosa uses motion parallax and E. fasciata uses a combination of motion parallax and forward and backward movements (image expansion/contraction over time) to detect object distances.     

Role of visual cues and interaction with host odour during the host-finding behaviour of the cabbage moth

The approach and landing responses of female Mamestra brassicae (L.) (Lepidoptera: Noctuidae) to visual cues from artificial plant leaves of different shapes and presence/absence of cabbage plant odour were investigated in a laboratory wind tunnel. The leaves were painted with cadmium yellow colour and observed under dim red light. Females showed oriented flight towards plant odours but landed significantly more often when the odour was presented with an artificial leaf. In three-choice tests, the shape of the leaf targets (circle, square or triangle) did not influence the female response. However, the size of the target did influence the insect response: the females preferred landing on square targets with sides of 5 or 10 cm rather than on the largest target, with sides of 15 cm. The orientation of the target influenced the insects' response: females landed significantly more often on the target positioned vertically than horizontally.     

A visually evoked escape response of the housefly

Summary Flies (Musca domestica) avoid danger by initiating a rapid jump followed by flight. To identify the visual cues that trigger the escape response in the housefly, we measured the timing and probability of escapes when the fly was presented with a variety of visual stimuli created by moving targets toward it. Our results show that an escape response is triggered by an approaching dark disk, but not by a receding dark disk. On the other hand, a bright disk elicits escape only when it recedes. A disk with black and white rings is less effective at eliciting escape than is a dark solid disk of the same size. This indicates that the darkening contrast produced by an approaching stimulus is a more crucial parameter than expansion cues contained in the optical flow. Escape is also triggered by a horizontally moving dark edge, but not by a moving bright edge or by a grating. An examination of several visual parameters reveals that the darkening contrast, measured from the onset of stimulation to the start of escape is nearly constant for a variety of stimuli that trigger escape reliably. Thus darkening contrast, coupled with motion may be crucial in eliciting the visually evoked escape response. Other visual parameters such as time-to-contact or target angular velocity seem to be relatively unimportant to the timing of escapes.Abbreviations P _s Probability of successful escape - r _disk radius of disk target - r _arena radius of shielding arena - v _disk linear velocity of disk target - v _edge linear velocity of edge - d _disk angular velocity of disk target boundary - _edge angular velocity of edge - _escape target distance at escape - d _start target distance before onset of target movement - h _edge height of the edge above fly - x _start distance from corner of triangle to start position of edge (0 or 50 mm) - x _escape distance from corner of triangle to the position of the edge when the fly escapes - x _center distance from corner of triangle to point above the center of the pad - x _total distance from the corner of the triangle to the base (height of triangle = base of triangle)     

Segregation of object and background motion in visual area MT: effects of microstimulation on eye movements

To track a moving object, its motion must first be distinguished from that of the background. The center-surround properties of neurons in the middle temporal visual area (MT) may be important for signaling the relative motion between object and background. To test this, we microstimulated within MT and measured the effects on monkeys' eye movements to moving targets. We found that stimulation at "local motion" sites, where receptive fields possessed antagonistic surrounds, shifted pursuit in the preferred direction of the neurons, whereas stimulation at "wide-field motion" sites shifted pursuit in the opposite, or null, direction. We propose that activating wide-field sites simulated background motion, thus inducing a target motion signal in the opposite direction. Our results support the hypothesis that neuronal center-surround mechanisms contribute to the behavioral segregation of objects from the background.     

Scototaxis and target perception in the camel tickHyalomma dromedarii

The camel tick,Hyalomma dromedarii, exhibited positive scototaxis in an arena, e.g. it oriented towards a black or grey target in front of a white background. The degree of the scototactic response varied with the size and the elevation of the target, with its luminance contrast, with its shape and with the speed by which the target was moved: (1) the response to stationary and moving targets increased with increasing target size; (2) presentation of the targets at an elevation of 11^o–15^o induced the highest response; (3) the response decreased with decreasing luminance contrast of the target; (4) targets with the shape of a disk, a triangle standing on a vertex, a vertical bar or a silhouette of a dromedary caused high responses; a low response was observed when the target was a horizontal bar and there was no response to a striped pattern; (5) the smaller the size of a disk, the faster it had to be moved to elicit an optimum response.The smallest disk which elicited a significant response appeared under a visual angle of 4.8^o for a thick at the starting point. The smallest dromedary-shaped silhouette which elicited a significant response corresponded to the silhouette of a real dromedary at a distance of 18 m.     

Responses of the blister beetle Hycleus apicicornis to visual stimuli

Insect attraction to host plants may be partly mediated by visual stimuli. In the present study, the responses of adult Hycleus apicicornis (Guér.) (Coleoptera: Meloidae) to plant models of different colours, different combinations of two colours, or three hues of blue of different shapes are compared. Single‐colour models comprised the colours sky blue, bright green, yellow, red, white and black. Sky blue (reflecting light in the 440–500 nm region) is the most attractive, followed by white, which reflects light over a broader range (400–700 nm). On landing on sky blue targets, beetles exhibit feeding behaviour immediately. When different hues of blue (of different shapes) are compared, sky blue is preferred over turquoise, followed by dark blue, indicating that H. apicicornis is more attracted to lighter hues of blue than to darker ones. No significant differences are found between the three shapes (circle, square and triangle) tested, suggesting that reflectance associated with colour could be a more important visual cue than shape for host location by H. apicicornis. The preference of H. apicicornis for sky blue can be exploited in designing an attractive trap for its management.     

Orientation and Shape Discrimination in Juveniles and Adults of the Mangrove Crab Aratus pisonii (H. Milne Edwards, 1837): Effect of Predator and Chemical Cues

Behavioral responses of juveniles and adults of the mangrove crab Aratus pisonii (H. Milne Edwards, 1837) to black geometric shapes of equal surface area was measured. Crabs were tested either in presence or absence of chemicals generated from two common predator species, the portunid crab Callinectes ornatus Ordway, and the soapfish Haemulon aurolineatum Cuvier, 1830. The present study tested the hypothesis that A. pisonii (1) has the capacity to orient to visual cues; (2) it discriminates between different visual objects based on a combination of chemical and visual information and (3) this behavior changes with age. When presented with single black targets in background water, juveniles oriented toward all shapes. This behavioral response was interpreted as visual orientation toward potential shelter. Among shapes, juveniles showed preference for the vertical rectangle, probably due to the recognition of natural visual elements like mangrove roots. In predator conditioned water, juveniles exhibited a stronger response than in background water. Thus, juveniles were able to detect by odor the potential presence of predators. Change in responsiveness between adults and juveniles was also demonstrated.     

On the barn owl’s visual pre-attack behavior: I. Structure of head movements and motion patterns

Barn owls exhibit a rich repertoire of head movements before taking off for prey capture. These movements occur mainly at light levels that allow for the visual detection of prey. To investigate these movements and their functional relevance, we filmed the pre-attack behavior of barn owls. Off-line image analysis enabled reconstruction of all six degrees of freedom of head movements. Three categories of head movements were observed: fixations, head translations and head rotations. The observed rotations contained a translational component. Head rotations did not follow Listing’s law, but could be well described by a second-order surface, which indicated that they are in close agreement with Donder’s law. Head translations did not contain any significant rotational components. Translations were further segmented into straight-line and curved paths. Translations along an axis perpendicular to the line of sight were similar to peering movements observed in other animals. We suggest that these basic motion elements (fixations, head rotations, translations along a straight line, and translation along a curved trajectory) may be combined to form longer and more complex behavior. We speculate that these head movements mainly underlie estimation of distance during prey capture.     

Orientation and Shape Discrimination in Juveniles and Adults of the Mangrove Crab Aratus pisonii (H. Milne Edwards, 1837): Effect of Predator and Chemical Cues

Behavioral responses of juveniles and adults of the mangrove crab Aratus pisonii (H. Milne Edwards, 1837) to black geometric shapes of equal surface area was measured. Crabs were tested either in presence or absence of chemicals generated from two common predator species, the portunid crab Callinectes ornatus Ordway, and the soapfish Haemulon aurolineatum Cuvier, 1830. The present study tested the hypothesis that A. pisonii (1) has the capacity to orient to visual cues; (2) it discriminates between different visual objects based on a combination of chemical and visual information and (3) this behavior changes with age. When presented with single black targets in background water, juveniles oriented toward all shapes. This behavioral response was interpreted as visual orientation toward potential shelter. Among shapes, juveniles showed preference for the vertical rectangle, probably due to the recognition of natural visual elements like mangrove roots. In predator conditioned water, juveniles exhibited a stronger response than in background water. Thus, juveniles were able to detect by odor the potential presence of predators. Change in responsiveness between adults and juveniles was also demonstrated.     

Binocular onset rivalry at the time of saccades and stimulus jumps

Recent studies suggest that binocular rivalry at stimulus onset, so called onset rivalry, differs from rivalry during sustained viewing. These observations raise the interesting question whether there is a relation between onset rivalry and rivalry in the presence of eye movements. We therefore studied binocular rivalry when stimuli jumped from one visual hemifield to the other, either through a saccade or through a passive stimulus displacement, and we compared rivalry after such displacements with onset and sustained rivalry. We presented opponent motion, orthogonal gratings and face/house stimuli through a stereoscope. For all three stimulus types we found that subjects showed a strong preference for stimuli in one eye or one hemifield (Experiment 1), and that these subject-specific biases did not persist during sustained viewing (Experiment 2). These results confirm and extend previous findings obtained with gratings. The results from the main experiment (Experiment 3) showed that after a passive stimulus jump, switching probability was low when the preferred eye was dominant before a stimulus jump, but when the non-preferred eye was dominant beforehand, switching probability was comparatively high. The results thus showed that dominance after a stimulus jump was tightly related to eye dominance at stimulus onset. In the saccade condition, however, these subject-specific biases were systematically reduced, indicating that the influence of saccades can be understood from a systematic attenuation of the subjects' onset rivalry biases. Taken together, our findings demonstrate a relation between onset rivalry and rivalry after retinal shifts and involvement of extra-retinal signals in binocular rivalry.     

Visually targeted reaching in horse-head grasshoppers

Visually targeted reaching to a specific object is a demanding neuronal task requiring the translation of the location of the object from a two-dimensionsal set of retinotopic coordinates to a motor pattern that guides a limb to that point in three-dimensional space. This sensorimotor transformation has been intensively studied in mammals, but was not previously thought to occur in animals with smaller nervous systems such as insects. We studied horse-head grasshoppers (Orthoptera: Proscopididae) crossing gaps and found that visual inputs are sufficient for them to target their forelimbs to a foothold on the opposite side of the gap. High-speed video analysis showed that these reaches were targeted accurately and directly to footholds at different locations within the visual field through changes in forelimb trajectory and body position, and did not involve stereotyped searching movements. The proscopids estimated distant locations using peering to generate motion parallax, a monocular distance cue, but appeared to use binocular visual cues to estimate the distance of nearby footholds. Following occlusion of regions of binocular overlap, the proscopids resorted to peering to target reaches even to nearby locations. Monocular cues were sufficient for accurate targeting of the ipsilateral but not the contralateral forelimb. Thus, proscopids are capable not only of the sensorimotor transformations necessary for visually targeted reaching with their forelimbs but also of flexibly using different visual cues to target reaches.     

Effects of luminance, size, and angular velocity on the recognition of nonlocomotive prey models by the praying mantis

Adult females of the mantis Tenodera angustipennis were presented with the "nonlocomotive" prey model, a static rectangle with two lines oscillating regularly at its sides, generated on a computer display. The models were varied in rectangle luminance (black, gray, and light gray), rectangle height (0.72, 3.6, and 18 mm), rectangle width (0.72, 3.6, and 18 mm), and angular velocity of oscillating lines (65°, 260°, and 1040°/s) to examine their effects on prey recognition. Before striking the model, the mantis sometimes showed peering movements that involved swaying its body from side to side. The black model of medium size (both height and width) elicited higher rates of fixation, peering, and strike responses than the large, small, or gray model. The model of medium angular velocity elicited a higher strike rate than that of large or small angular velocity, but angular velocity had little effect on fixation and peering. We conclude that mantises respond to a rectangle in deciding whether to fixate, and to both rectangle and lines in deciding whether to strike after fixation. Received: September 2, 1999 / Accepted: March 21, 2000     

Visual memory of shapes in quail chicks: discrimination among 2-dimensional objects

Newly hatched chicks spontaneously peck at conspicuous objects, and soon learn to discriminate between edible food particles and inedible objects. To examine whether this discrimination is based on a chick's ability to memorize objects by shape cues, we analyzed the pecking behavior. One- to 3-day old quail chicks (Coturnix japonica) were presented with dry objects of different shapes (ball, disk, triangle and T-shape) of similar size (4 mm) and color (green). Habituation occurred after repeated presentation of any one of these objects (duration: 30 sec; interval: 4 min). When chicks showed significantly more pecks at a novel object (dishabituation), we assumed that chicks had memorized the habituated shapes and distinguished the novel object. Chicks did not show dishabituation between a ball and a disk. On the other hand, chicks discriminated a triangle or T-shape from the memorized image of disk, but did not memorize either triangle or T-shape by its shape. Similarly, chicks did not memorize the size of disks as a reference for subsequent pecking behavior. Chicks proved to have a limited ability to memorize shape and size cues for selective pecking behavior, in strong contrast to their accurate memorization of colors.     

Responses of female New World screwworm flies, Cochliomyia hominivorax, to coloured targets in the laboratory

The responses of unmated female New World screwworm flies, Cochliomyia hominivorax Coquerel, to visual targets were studied in a wind-tunnel. Both activity and frequency of contacts with targets increased greatly when the screwworm attractant mixture swormlure-4 was added to the airstream. Target-orientated responses depended on target colour, with red and black targets being preferred over blue, white and yellow ones; this preference was much greater in the presence of odour than in its absence. No preference was detected for different shapes and orientations of red targets, all of equivalent surface area. Omitting different components from swormlure-4 generally resulted in a large reduction in activation and target contacts. Attempts to substitute 1-octen-3-ol for the butanol fraction were unsuccessful, but skatole may substitute to some extent for indole; the two isomers of butanol normally present in swormlure-4 may substitute partly or completely for each other. This type of measurement forms a suitable bioassay in the development of attractive targets for monitoring and control of wild adult screwworm populations.