Behavioral analysis of two distinct visual responses in the larva of the tiger beetle (Cicindela chinensis)

The tiger beetle larva shows two distinct visual responses, a predatory jump and an evasive withdrawal into the burrow (escape). In the present study the visual stimuli controlling these two responses have been behaviorally analyzed in the larva of Cicindela chinensis. The threshold size needed for a target to elicit both responses is a visual angle of 5–7°. The velocities of moving targets needed to elicit the responses are 0.4–33° s⁻¹ for the jump and 0.76–90° s⁻¹ for the escape. Choice between the two responses appears to be controlled by the actual target size rather than by the angular size. It also appears to be controlled by the target height. As the height of the target increases, the probability for the jump decreases, whereas the probability for the escape increases. Response properties of the larva with only a single functional stemma, the other stemmata being occluded, are different from those of the intact larva, which suggests cooperation of at least two stemmata for the release of different visual responses. Visual responses of the one-stemma larva still vary, however, with target size and target height, which suggests the visual responses are partially controlled even by a single stemma. Although our data do not resolve these conflicting results, more than one stemma is necessary for a firm choice between the two responses. Accepted: 13 May 1997     

Temporal modulation of luminance adapts time constant of fly movement detectors

The time constant of movement detectors in the fly visual system has been proposed to adapt in response to moving stimuli (de Ruyter van Steveninck et al. 1986). The objective of the present study is to analyse, whether this adaptation can be induced as well, if the luminance of a stationary uniform field is modulated in time. The experiments were done on motion-sensitive wide-field neurones of the lobula plate, the posterior part of the third visual ganglion of the blowfly, calliphora erythrocephala. These cells are assumed to receive input from large retinotopic arrays of movement detectors. In order to demonstrate that our results concern the properties of the movement detectors rather than those of a particular wide-field cell we recorded from two different types of them, the H1- and the HSE-cell. Both cell types respond to a brief movement stimulus in their preferred direction with a transient excitation. This response decays about exponentially. The time constant of this decay reflects, in a first approximation, the time constant of the presynaptic movement detectors. It was determined after prestimulation of the cell by the following stimuli: (a) periodic stationary grating; (b) uniform field, the intensity of which was modulated sinusoidally in time (flicker stimulation); (c) periodic grating moving front-to-back; (d) periodic grating moving back-to-front. The decay of the response is significantly faster not only after movement but also after flicker stimulation as compared with pre-stimulation with a stationary stimulus. This is interpreted as an adaptation of the movement detector's time constant. The finding that flicker stimulation also leads to an adaptation shows that movement is not necessary for this process. Instead the adaptation of the time constant appears to be governed mainly by the temporal modulation (i.e., contrast frequency) of the signal in each visual channel.     

The angular orientation of the movement detectors acting on the flight lift response in flies

The lift response of houseflies Musca domestica in fixed flight to periodic gratings movins in 12 different orientations has been measured. Two projectors were arranged symmetrically to the flies stimulating successively 18 circular patches of 50° (25°) diameter (9 for each eye) in their visual field. The shapes of the lift responses measured as a function of the orientation of the moving grating varied when different patches in the visual field were stimulated. A qualitative comparison of these response curves leads to the conclusion that the orientation of the movement detecting substrate acting on the flight lift response varies as a function of the stimulated area in the visual field. A straightforward correlation between the geometry of the ommatidial pattern and the orientation of the movement detecting substrate valid for all stimulated areas of the compound eyes does not seem very likely.     

Influence of Retinal Image Shifts and Extra-Retinal Eye Movement Signals on Binocular Rivalry Alternations

Previous studies have indicated that saccadic eye movements correlate positively with perceptual alternations in binocular rivalry, presumably because the foveal image changes resulting from saccades, rather than the eye movement themselves, cause switches in awareness. Recently, however, we found evidence that retinal image shifts elicit so-called onset rivalry and not percept switches as such. These findings raise the interesting question whether onset rivalry may account for correlations between saccades and percept switches.We therefore studied binocular rivalry when subjects made eye movements across a visual stimulus and compared it with the rivalry in a ‘replay’ condition in which subjects maintained fixation while the same retinal displacements were reproduced by stimulus displacements on the screen. We used dichoptic random-dot motion stimuli viewed through a stereoscope, and measured eye and eyelid movements with scleral search-coils.Positive correlations between retinal image shifts and perceptual switches were observed for both saccades and stimulus jumps, but only for switches towards the subjects'' preferred eye at stimulus onset. A similar asymmetry was observed for blink-induced stimulus interruptions. Moreover, for saccades, amplitude appeared crucial as the positive correlation persisted for small stimulus jumps, but not for small saccades (amplitudes < 1°). These findings corroborate our tenet that saccades elicit a form of onset rivalry, and that rivalry is modulated by extra-retinal eye movement signals.     

Components of Visual Prey Recognition by the Mexican Aquatic Garter Snake Thamnophis melanogaster

Garter snakes (genus Thamnophis) rely mainly on chemical cues to recognize prey, but in some of the more aquatic species visual stimuli may suffice to elicit predatory attacks. However, the only visual components known to be involved in the visual release of attacks are movement and contrast with background. We explored other visual components by presenting visual models varying only in size, shape, or path of movement to an aquatically specialized species, Thamnophis melanogaster. Snakes responded preferentially to models consistent in size with natural prey, to models having non-elongate shapes regardless of type of contour (rounded or angular), and to those following paths with vertical, rather than only horizontal, components.     

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)     

Dynamic visual cues induce jaw opening and closing by tiger beetles during pursuit of prey

In dynamic locomotory contexts, visual cues often trigger adaptive behaviour by the viewer, yet studies investigating how animals determine impending collisions typically employ either stationary viewers or objects. Here, we describe a dynamic situation of visually guided prey pursuit in which both impending prey contact and escape elicit observable adaptive behaviours in the pursuer, a predatory beetle. We investigated which visual cues may independently control opening and closing of the beetle''s jaws during chases of prey dummies. Jaw opening and closing typically occur when prey is within the 60° binocular field, but not at specific distances, angular sizes or time-to-collision. We show that a sign change in the expansion rate of the target image precedes jaw opening (16 ms) and closing (35 ms), signalling to the beetle that it is gaining on the target or that the target is getting away. We discuss the ‘sloppiness'' of such variation in the lag of the behavioural response, especially jaw closing, as an adaptation to uncertainty about target position due to degradation of the target image by motion blur from the fast-running beetle.     

Factors affecting feeding selectivity of visual predators on the copepod Acartia tonsa: locomotion,visibility and escape responses

Visual predation by fish on copepods involves prey encounter, attack and capture; during any of these processes prey selection can occur. Developmental changes in copepods, including increases in swimming speed, size and image contrast increase the encounter rate and distance at which they can be detected by predators. Copepods compensate for this increase vulnerability with age through diel vertical migration and improved escape capabilities. This study quantifies the changes in swimming speed and movement pattern with developmental stage of the copepod Acartia tonsa, using a video-computer system for motion analysis. Changes in visible size and image contrast with developmental stage were quantified under simulated natural illumination conditions using a video based image analysis system. The escape responses of the naupliar stages of the copepod Acartia tonsa were quantified in response to a stationary pipette sucking in water at a constant speed. Accurate quantification of the parameters that affect feeding selectivity of planktivorous fish will provide the basis for evaluation of their relative importance in future studies.     

No impact of a presumed manipulative parasite on the responses and susceptibility of fish to simulated predation

Parasites manipulating their host to facilitate trophic transmission is a widespread and diverse phenomenon. Trematode eye‐flukes in the family Diplostomidae infect a variety of fish species as metacercariae, many residing in the eyes. A recently described diplostomid, Tylodelphys darbyi, from the South Island of New Zealand has been found to infect common bully Gobiomorphus cotidianus, an endemic freshwater fish. Within the fish, the metacercariae move about freely in the liquid parts of the eye and are quite large. We hypothesized that increasing intensity of T. darbyi infection will result in increasing visual impairment, thus reducing the ability of G. cotidianus to identify and react to a predatory threat. To test this hypothesis, we performed experiments to (a) examine the fish's reaction to a purely visual predator cue and (b) test their ability to avoid simulated predation under natural levels of infection. Among the 64 fish used in our experiments, T. darbyi had a prevalence of 98.7% with an average of 17.6 worms per fish. However, there was no relationship between T. darbyi intensity and either the fish's reaction to a visual predator stimulus or their ability to escape a simulated predator. Our findings indicate that despite being present in large numbers in the eyes of its fish host, the parasite appears incapable of improving its chances of trophic transmission to its avian definitive host. The results also suggest that the fish G. cotidianus could be using other senses (e.g., olfaction and lateral line) to compensate for visual impairment, and detect and respond to predators.     

Analyse der auslösenden Reizsituationen für die Anschwimm-, Eindring- und Fluchtreaktion junger Hemihaplochromis multicolor (Cichlidae)

To investigate the selective response to stimulation the stimuli effecting release of three different reactions in young Hemihaplochromis multicolor were analysed: approach, contact making, and escape. Responses to grey, coloured, and patterned dummies were compared in double-choice experiments. These experiments show that pattern shape has no meaning but that the size of the dark area of the pattern and the contrast in brightness between the dark area and that surrounding it are important sign stimuli for each of the three reactions. Therefore, selective responsiveness does not appear to result from heterogeneous summation but from the quantity of each stimulus parameter and the special combination of sign stimuli.     

The escape behavior of marine copepods in response to a quantifiable fluid mechanical disturbance

The threshold shear values needed to elicit the escape reactionto a quantifiable fluid mechanical disturbance were comparedbetween five free-swimming oceanic copepod species. The resultsindicate a significant difference in the threshold for differentspecies of copepods and between different age groups withina single species. In general, animals captured from more energeticregimes required a higher threshold than those captured frommore pacific locations. Labidocera madurae required the highestshear values with 51.5 s^–1 for 50% of the animals testedto elicit an escape reaction (S₅₀). Acartia tonsa and Euchaetarimana, in contrast, were behaviorally the most sensitive requiringan S₅₀ of only 1.5 and 4.1 s^–1, respectively, to initiatean escape reaction. Pleuromamma xiphias and Oithona requiredintermediate shear values with an S₅₀ of 7.2 and 8.1 s^–1.When compared to literature values, the threshold needed toelicit an escape reaction was consistently higher than averageenvironmental shear values. Threshold shear values also variedsignificantly with developmental stage. Naupliar stages of A.tonsarequired greater than six times the S₅₀ value required by adultsof the same species. This suggests that the higher vulnerabilityto predation of naupliar stages of copepods may not only reflectinferior escape strength, but may also result from the higherthreshold needed to elicit an escape reaction. This study supportsthe hypothesis that selective feeding patterns exhibited bypredators of copepods may be the result of the differentialbehavioral sensitivities of different species and developmentalstages of copepods.     

Responses to computer-generated visual stimuli by the male praying mantis, Sphodromantis lineola (Burmeister)

We tested male mantis' responses to square, computer-generated visual stimuli of various sizes to determine the stimulus parameters that affect their striking behaviour. Males in a high-hunger group displayed a sharp preference for solid black 12×12-degree squares moving linearly against a white background, especially when the squares moved downward (versus horizontally). Males in a low-hunger group visually tracked but, with rare exception, did not strike at any linearly moving squares. In contrast, when solid black squares moved erratically around visual field centre, males struck regularly at them irrespective of hunger level, even when the squares were as large as 47×47 degrees. Males also recognized black-and-white Julesz-patterned square stimuli moving against a similarly patterned background, indicating that they recognized the synchronous movement of a group of stimulus elements as a single moving object. Finally, we compared allometric and life history data between male and females S. lineola to elucidate the possible reasons for the differences between male response patterns and previously published data on females. These comparisons suggest that males employ a different behavioural strategy than do females when faced with large, erratically moving visual stimuli.     

Are the large monopolar cells of the insect lamina on the optomotor pathway?

Evidence is presented here from experiments on the visual system of the fly that questions participation of the large monopolar cells (LMCs) in the optomotor response.

Nonconsumptive effects of hunting on a nontarget game bird

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Speed tuning in elementary motion detectors of the correlation type

A prominent model of visual motion detection is the so-called correlation or Reichardt detector. Whereas this model can account for many properties of motion vision, from humans to insects (review, Borst and Egelhaaf 1989), it has been commonly assumed that this scheme of motion detection is not well suited to the measurement of image velocity. This is because the commonly used version of the model, which incorporates two unidirectional motion detectors with opposite preferred directions, produces a response which varies not only with the velocity of the image, but also with its spatial structure and contrast. On the other hand, information on image velocity can be crucial in various contexts, and a number of recent behavioural experiments suggest that insects do extract velocity for navigational purposes (review, Srinivasan et al. 1996). Here we show that other versions of the correlation model, which consists of a single unidirectional motion detector or incorporates two oppositely directed detectors with unequal sensitivities, produce responses which vary with image speed and display tuning curves that are substantially independent of the spatial structure of the image. This surprising feature suggests simple strategies of reducing ambiguities in the estimation of speed by using components of neural hardware that are already known to exist in the visual system. Received: 30 April 1998 / Accepted in revised form: 18 September 1998     

Exploratory Activity of Rats in Three Different Environments

Adult male hooded (Long‐Evans) rats (Rattus norvergicus) were used to compare the efficiency of three types of tests (plain open field – OF, open field with a refuge – OD, and complex environment with a refuge – EX), for the evaluation of exploratory behaviour. The results confirmed that OF is a highly aversive situation with the animals showing elevated emotional response (defecation and urination) when compared to situations containing a refuge (OD and EX). Presence of a plain arena (OF and OD) does not elicit high exploratory performance, as shown by the delayed and reduced exit from the refuge in OD when compared to EX. Thus locomotor activity in OF (covered distance, number of rearings) probably reflects more of an escape reaction than a genuine exploratory behaviour. Conversely the presence of a complex environment seems to elicit a high exploratory performance expressed as a quick exit from the refuge, an increased time in the environment (more than 75% of the observation time) and intense locomotor patterns (large covered distances, great number of rearings) but with little emotional display (reduced defecation and urination). The results thus show the importance of considering ethological factors in the choice or development of laboratory tests.     

Prey Escape Direction is Influenced by the Pivoting Displays of Flush-Pursuing Birds

Painted redstart, Myioborus pictus, and its congeners in Central and South America, belong to a small fraction of insectivorous flush‐pursuing birds. Unlike most of the small insectivorous birds, which glean prey from substrates, the flush pursuers spread and pivot their conspicuously patterned tails and wings. This display triggers prey escape flights which are hypothesized to occur through visual stimulation of prey escape circuits [giant descending neuron cluster (GDNC) in Diptera] sensitive to the looming motion of an approaching bird, translational motion of a pivoting body with widely spread tail and contrast of the white‐black plumage pattern. In this paper, data from field observations of redstarts and experiments with bird models show an increase in the frequency of prey escapes away from the strong visual stimulation of an open tail, and in the direction opposite to that of the horizontal translational motion present in the pivots. We discuss how the effect on prey escape direction may enhance prey interception capabilities of redstarts during aerial pursuits. Combined with an earlier study the results show that, unlike the movements of typical gleaner–foragers, the flush displays by redstarts affect prey escape direction in a manner that may facilitate prey tracking and capture by birds. Because the GDNs, which mediate escape initiation, are not sensitive to motion direction, we hypothesize that other neurons, in addition to the GDNs, are involved in influencing the direction of escape responses.     

Visual feedback in the control of pheromone-mediated flight ofHeliothis virescens males (Lepidoptera: Noctuidae)

MaleHeliothis virescens (F.) (Lepidoptera: Noctuidae) were made to fly into a uniformly white and translucent tube within a large wind tunnel while responding to sex pheromone. Different visual patterns placed within the tube greatly affected the ability of the male moths to maintain upwind progress or remain oriented to the wind while in contact with the plume. Over 89% of males attempting to fly through a blank tube, lacking visual patterns, became disoriented, the males gaining or losing altitude and repeatedly hitting the sides of the tube. Patterns of 20–40 dots placed on the sides of the tube at or slightly above plume level resulted in high levels of sustained upwind flight (47–74%) relative to patterns placed directly below (30–40%), directly above (35%), or slightly below the level of the flight path (26–44%). Optimal upwind progression in pheromone-responding males occurred when image motion could be resolved both transversely (T), orthogonally to the longitudinal axis of the body relative to the horizontal plane of the environment, and longitudinally (L), along the body axis. Even very sparse patterns (single rows of dots) could elicit high levels of sustained upwind flight (53–63%) when positioned within the tube such that the males' movements would create both L and T image motion. However, successful negotiation of the tube was also unexpectedly facilitated by patterns apparently providing no horizontal transverse component for flying males but providing longitudinal flow while centering the moth in the plume through a symmetrical left-right input (4–40%).     

Effects of olfactory stimuli on arm-reaching duration

The aim of the present study was to investigate the effects of olfactory stimuli on visually guided reaching. In Experiment 1, participants reached toward and grasped either a small (almond/strawberry) or a large (apple/orange) visual target. Any 1 of 4 odors corresponding to the visual stimuli or odorless air was administered before movement initiation. Within the same block of trials, participants smelled 1) an odor associated with an object of a different size than the target, 2) an odor associated with an object of a size equal to that of the target, or 3) odorless air. Results indicated that reaching duration was longer for trials in which the odor "size" and the visual target did not match than when they matched. In Experiment 2, the same procedures were applied but the "no-odor" trials were administered in a separate block to the "odor" trials. Similar results as for Experiment 1 were found. However, in contrast to Experiment 1, the presence of an odor increased the level of alertness resulting in a shortening of reaching duration. We contend that olfactory stimuli have the capacity to elicit motor plans interfering with those programmed for a movement toward a visual stimulus.     

A model of the cerebellum in adaptive control of saccadic gain

 We review data showing that the cerebellum is required for adaptation of saccadic gain to repeated presentations of dual-step visual targets and thus, presumably, for providing adaptive corrections for the brainstem saccade generator in response to any error created by the open-loop saccadic system. We model the adaptability of the system in terms of plasticity of synapses from parallel fibers to Purkinje cells in cerebellar cortex, stressing the integration of cerebellar cortex and nuclei in microzones as the units for correction of motor pattern generators. We propose a model of the inferior olive as an error detector, and use a ‘window of eligibility’ to insure that error signals that elicit a corrective movement are used to adjust the original movement, not the secondary movement. In a companion paper we simulate this large, realistic network of neural-like units to study the complex spatiotemporal behavior of neuronal subpopulations implicated in the control and adaptation of saccades. Received: 25 November 1994/Accepted in revised form: 6 February 1996