Responses of the goldfish trunk lateral line to moving objects

We recorded the responses of single afferent fibers in the posterior lateral-line nerve of the goldfish, Carassius auratus, to a small object moving in the water. Responses consisted of a dominant and reproducible pattern of discharge which was characterized by excitation followed by inhibition or vice versa. The pattern depended on the direction in which the object moved and was inverse when the direction was reversed. About half of the fibers continued to discharge bursts of spikes for a long time after the object had passed the fish. These spike bursts were not reproducible from one stimulus presentation to the next. In many fibers, the pattern of the response changed with speed and lateral distance of the moving object. Response strength increased with increasing object speed and decreasing lateral distance. Measurements of water motions revealed that the object generated complex water movements, aspects of which were reflected in the discharges of primary lateral-line afferents. The observed uniformity of the responses in the periphery suggests that many, but not all, of the response patterns of central lateral-line units to moving objects are due to additional information processing by the central nervous system and not to peripheral hydrodynamic effects. Accepted: 6 October 1997     

Structure-from-motion based on information at surface boundaries

Existing computational models of structurefrom-motion — the appearance of three-dimensional motion generated by moving two-dimensional patterns — are all based on variations of optical flow or feature point correspondence within the interior of single objects. Three separate phenomena provide strong evidence that in human vision, structure-from-motion is significantly affected by surface boundary cues. In the first, a rotating cylinder is seen, though no variation in optical flow exists across the apparent cylinder. In the second, the shape of the bounding contour of a moving pattern dominates the actual differential motion within the pattern. In the third, the appearance of independently moving objects changes significantly when the boundary between them becomes indistinct. We describe a simple computational model sufficient to account for these effects. The model is based on qualitative constraints relating possible object motions to patterns of flow, together with an understanding of the patterns of flow that can be discriminated in practice.     

Distance discrimination during active electrolocation in the weakly electric fish Gnathonemus petersii.

Weakly electric fish use active electrolocation for orientation at night. They emit electric signals (electric organ discharges) which generate an electrical field around their body. By sensing field distortions, fish can detect objects and analyze their properties. It is unclear, however, how accurately they can determine the distance of unknown objects. Four Gnathonemus petersii were trained in two-alternative forced-choice procedures to discriminate between two objects differing in their distances to a gate. The fish learned to pass through the gate behind which the corresponding object was farther away. Distance discrimination thresholds for different types of objects were determined. Locomotor and electromotor activity during distance measurement were monitored. Our results revealed that all individuals quickly learned to measure object distance irrespective of size, shape or electrical conductivity of the object material. However, the distances of hollow, water-filled cubes and spheres were consistently misjudged in comparison with solid or more angular objects, being perceived as farther away than they really were. As training continued, fish learned to compensate for these 'electrosensory illusions' and erroneous choices disappeared with time. Distance discrimination thresholds depended on object size and overall object distance. During distance measurement, the fish produced a fast regular rhythm of EOD discharges. A mechanisms for distance determination during active electrolocation is proposed.     

Distance discrimination during active electrolocation in the weakly electric fish Gnathonemus petersii

Weakly electric fish use active electrolocation for orientation at night. They emit electric signals (electric organ discharges) which generate an electrical field around their body. By sensing field distortions, fish can detect objects and analyze their properties. It is unclear, however, how accurately they can determine the distance of unknown objects. Four Gnathonemus petersii were trained in two-alternative forced-choice procedures to discriminate between two objects differing in their distances to a gate. The fish learned to pass through the gate behind which the corresponding object was farther away. Distance discrimination thresholds for different types of objects were determined. Locomotor and electromotor activity during distance measurement were monitored. Our results revealed that all individuals quickly learned to measure object distance irrespective of size, shape or electrical conductivity of the object material. However, the distances of hollow, water-filled cubes and spheres were consistently misjudged in comparison with solid or more angular objects, being perceived as farther away than they really were. As training continued, fish learned to compensate for these 'electrosensory illusions' and erroneous choices disappeared with time. Distance discrimination thresholds depended on object size and overall object distance. During distance measurement, the fish produced a fast regular rhythm of EOD discharges. A mechanisms for distance determination during active electrolocation is proposed.     

How Can Dolphins Recognize Fish According to Their Echoes? A Statistical Analysis of Fish Echoes

Echo-based object classification is a fundamental task of animals that use a biosonar system. Dolphins and porpoises should be able to rely on echoes to discriminate a predator from a prey or to select a desired prey from an undesired object. Many studies have shown that dolphins and porpoises can discriminate between objects according to their echoes. All of these studies however, used unnatural objects that can be easily characterized in human terminologies (e.g., metallic spheres, disks, cylinders). In this work, we collected real fish echoes from many angles of acquisition using a sonar system that mimics the emission properties of dolphins and porpoises. We then tested two alternative statistical approaches in classifying these echoes. Our results suggest that fish species can be classified according to echoes returning from porpoise- and dolphin-like signals. These results suggest how dolphins and porpoises can classify fish based on their echoes and provide some insight as to which features might enable the classification.     

Visual discrimination of objects differing in spatial depth by goldfish

Training experiments were performed to investigate the ability of goldfish to discriminate objects differing in spatial depth. Tests on size constancy should give insight into the mechanisms of distance estimation. Goldfish were successfully trained to discriminate between two black disk stimuli of equal size but different distance from the tank wall. Each stimulus was presented in a white tube so that the fish could see only one stimulus at a time. For each of eight training stimulus distances, the just noticeable difference in distance was determined at a threshold criterion of 70% choice frequency. The ratio of the retinal image sizes between training stimulus and comparison stimulus at threshold was about constant. However, in contrast to Douglas et al. (Behav Brain Res 30:37–42, 1988), goldfish did not show size constancy in tests with stimuli of the same visual angle. This indicates that they did not estimate distance, but simply compared the retinal images under our experimental conditions. We did not find any indication for the use of accommodation as a depth cue. A patterned background at the rear end of the tubes did not have any effect, which, however, does not exclude the possibility that motion parallax is used as a depth cue under natural conditions.     

Fish perform spatial pattern recognition and abstraction by exclusive use of active electrolocation

The field generated by the electric organ of weakly electric fish varies with the electrical properties of nearby objects. Correspondingly, current fluxes in this field differentially stimulate the electroreceptors in the fish's skin. Thus, resistors are to conductors and insulators as gray is to black and white in optics. Additionally, the capacitances of plants and insect larvae contrast with those of water or stones, giving effects comparable to "coloration". Receptors arrayed over a large area of the skin act like a retina upon which the discharge projects "electric images". By further central processing, the fish also discriminate between objects according to their composition, size, or distance, a procedure termed "electrolocation", analogous to echolocation in bats. Here we demonstrate that G. petersii and S. macrurus can also recognize 3D orientations and configurations and extract and generalize spatial features solely with their electrical sense. We presented fish with virtual electrical "objects" formed from electrodes set flush in the inner surface of a Y maze with various patterns of external connectivity. With reward and aversion training, the fish could recognize similar electrode configurations and extract a feature, e.g., a vertical connectivity, present in various novel configurations. Previously, shape recognition has only been shown in electrolocating fish when they are in full mechanical contact with solid objects.     

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.     

The role of background movement in goldfish vision

In experiments described in the literature objects presented to restrained goldfish failed to induce eye movements like fixation and/or tracking. We show here that eye movements can be induced only if the background (visual surround) is not stationary relative to the fish but moving. We investigated the influence of background motion on eye movements in the range of angular velocities of 5–20° s⁻¹. The response to presentation of an object is a transient shift in mean horizontal eye position which lasts for some 10 s. If an object is presented in front of the fish the eyes move in a direction such that it is seen more or less symmetrically by both eyes. If it is presented at ±70° from the fish's long axis the eye on the side of the object moves in the direction that the object falls more centrally on its retina. During these object induced eye responses the typical optokinetic nystagmus of amplitude of some 5° with alternating fast and slow phases is maintained, and the eye velocity during the slow phase is not modified by presentation of the object. Presenting an object in front of stationary or moving backgrounds leads to transient suppression of respiration which shows habituation to repeated object presentations. Accepted: 14 April 2000     

Motion, not Shape, Facilitates Association of Predation Risk with Novel Objects by Fathead Minnows (Pimephales promelas)

Injury‐released chemical cues are reliable indicators of predation risk among many aquatic taxa. When a novel, neutral stimulus is presented in tandem with chemical cues from an injured conspecific, an association is formed between the novel stimulus and apparent risk. Learned recognition of predation risk is well documented for fathead minnows, Pimephales promelas. When minnows detect alarm cues in nature they are also potentially exposed to multiple environmental stimuli, few of which are likely to be relevant indicators of risk. How do minnows discern among candidate stimuli potentially associated with predation risk? Two possibilities are shape and motion. In this study, individual piscivore‐naïve minnows were presented simultaneously with conspecific chemical alarm cues and two stimulus objects. One object was a darkened tube with its long axis in the horizontal plane (fish‐like). The second object was a black disk. Following introduction of chemical alarm cues, one of the objects was raised and lowered repeatedly. After a single conditioning trial, minnows associated risk significantly more with the previously moving object than the previously stationary object, as indicated by reduced activity. Object shape had no significant effect on response intensity in test trials. Our data suggest that minnows have been selected to form aversive responses to moving objects at a site of recent predation because movement is a more predictable indicator of predator identity than shape.     

Mind the gap: the minimal detectable separation distance between two objects during active electrolocation

In a food‐rewarded two‐alternative forced‐choice procedure, it was determined how well the weakly electric elephantnose fish Gnathonemus petersii can sense gaps between two objects, some of which were placed in front of complex backgrounds. The results show that at close distances, G. petersii is able to detect gaps between two small metal cubes (2 cm × 2 cm × 2 cm) down to a width of c. 1·5 mm. When larger objects (3 cm × 3 cm × 3 cm) were used, gaps with a width of 2–3 mm could still be detected. Discrimination performance was better (c. 1 mm gap size) when the objects were placed in front of a moving background consisting of plastic stripes or plant leaves, indicating that movement in the environment plays an important role for object identification. In addition, the smallest gap size that could be detected at increasing distances was determined. A linear relationship between object distance and gap size existed. Minimal detectable gap sizes increased from c. 1·5 mm at a distance of 1 cm, to 20 mm at a distance of 7 cm. Measurements and simulations of the electric stimuli occurring during gap detection revealed that the electric images of two close objects influence each other and superimpose. A large gap of 20 mm between two objects induced two clearly separated peaks in the electric image, while a 2 mm gap caused just a slight indentation in the image. Therefore, the fusion of electric images limits spatial resolution during active electrolocation. Relative movements either between the fish and the objects or between object and background might improve spatial resolution by accentuating the fine details of the electric images.     

Distance and shape: perception of the 3-dimensional world by weakly electric fish.

Weakly electric fish orient at night in complete darkness by employing their active electrolocation system. They emit short electric signals and perceive the consequences of these emissions with epidermal electroreceptors. Objects are detected by analyzing the electric images which they project onto the animal's electroreceptive skin surface. This process corresponds to similar processes during vision, where visual images are cast onto the retinas of eyes. Behavioral experiments have shown that electric fish can measure the distance of objects during active electrolocation, thus possessing three-dimensional depth perception of their surroundings. The fundamental mechanism for distance determination differs from stereopsis used during vision by two-eyed animals, but resembles some supplementary mechanisms for distance deduction in humans. Weakly electric fish can also perceive the three-dimensional shape of objects. The fish can learn to identify certain objects and discriminate them from all other objects. In addition, they spontaneously categorize objects according to their shapes and not according to object size or material properties. There is good evidence that some fundamental types of perceptional invariances during visual object recognition in humans are also found in electric fish during active electrolocation. These include size invariance (maybe including size constancy), rotational invariance, and translational invariance. The mechanisms of shape detection during electrolocation are still unknown, and their discoveries require additional experiments.     

Responses of the goldfish head lateral line to moving objects

We investigated how fibers in the anterior lateral line nerve of goldfish, Carassius auratus, respond to water motions generated by an object that was moved alongside the fish. Motion direction was from anterior to posterior or opposite, object diameter was between 0.1 and 4 cm and the distance between object and fish varied between 1 and 6 cm. Fibers exhibited monophasic responses characterized by a transient increase in discharge rate, biphasic responses consisting of an increase followed by a decrease in discharge rate or vice versa, or triphasic responses characterized by a rate increase followed by a decrease and again an increase or by the inverse pattern. In two-thirds of the fibers response patterns depended on object motion direction. Of these, about 60% responded to a reversal of motion direction with an inversion of the response pattern. Our results differ from previous data obtained from posterior lateral line nerve fibers in the relative proportions of the observed response patterns, and by a much smaller proportion of fibers that exhibited a direction-dependent response. These differences can be explained by the fact that the spatial orientations of the neuromasts on the head are more heterogenuous than on the trunk.     

Lateral line reception in still- and running water

The lateral line of fish is composed of neuromasts used to detect water motions. Neuromasts occur as superficial neuromasts on the skin and as canal neuromasts in subepidermal canals. Fibres of the lateral line nerves innervate both. There have been extensive studies on the responses of lateral line nerve fibres to dipole stimuli applied in still water. However, despite the fact that many fish live in rivers and/or swim constantly, responses of lateral line nerve fibres to dipole stimuli presented in running water have never been recorded. We investigated how the peripheral lateral line of still water fish ( Carassius auratus) and riverine fish ( Oncorhynchus mykiss) responds to minute sinusoidal water motions while exposed to unidirectional water flow. Both goldfish and trout have two types of posterior lateral line nerve fibres: Type I fibres, which most likely innervate superficial neuromasts, were stimulated by running water (10 cm s(-1)). The responses of type I fibres to water motions generated by a vibrating sphere were masked if the fish was exposed to running water. Type II fibres, which most likely innervate canal neuromasts, were not stimulated by running water. Consequently, responses of type II fibres to a vibrating sphere were not masked under flow conditions.     

Dynamic shape integration in extrastriate cortex

BACKGROUND: In anorthoscopic viewing conditions, observers can perceive a moving object through a narrow slit even when only portions of its contour are visible at any time. We used fMRI to examine the contribution of early and later visual cortical areas to dynamic shape integration. Observers' success at integrating the shape of the slit-viewed object was manipulated by varying the degree to which the stimulus was dynamically distorted. Line drawings of common objects were either moderately distorted, strongly distorted, or shown undistorted. Phenomenologically, increasing the stimulus distortion made both object shape and motion more difficult to perceive.RESULTS: We found that bilateral cortical activity in portions of the ventral occipital cortex, corresponding to known object areas within the lateral occipital complex (LOC), was inversely correlated with the degree of stimulus distortion. We found that activity in left MT+, the human cortical area specialized for motion, showed a similar pattern as the ventral occipital region. The LOC also showed greater activity to a fully visible moving object than to the undistorted slit-viewed object. Area MT+, however, showed more equivalent activity to both the slit-viewed and fully visible moving objects.CONCLUSIONS: In early retinotopic cortex, the distorted and undistorted stimuli elicited the same amount of activity. Higher visual areas, however, were correlated with the percept of the coherent object, and this correlation suggests that the shape integration is mediated by later visual cortical areas. Motion information from the dorsal stream may project to the LOC to produce the shape percept.     

The time course and frequency content of hydrodynamic events caused by moving fish,frogs, and crustaceans

Summary In the present study the time course and spectral-amplitude distribution of hydrodynamic flow fields caused by moving fish, frogs, and crustaceans were investigated with the aid of laser-Doppler-anemometry. In the vicinity of a hovering fish sinusoidal water movements can be recorded whose velocity spectra peak below 10 Hz (Fig. 2). Single strokes during startle responses or during steady swimming of fish, frogs, and crustaceans cause short-lasting, low-frequency (<10 Hz), transient water movements (Fig. 3). Low-frequency transients also occur if a frog approaches and passes a velocity-sensitive hydrodynamic sensor. In contrast, transient water movements caused by a rapidly struggling or startled fish or water motions measured in the wake of a slowly swimming (47 cm/s) trout can be broadbanded, i.e., these water movements can contain frequency components up to at least 100 Hz (Figs. 4, 5A, 6). High-frequency hydrodynamic events can also be measured behind obstacles submerged in running water (Fig. 5C). The possible biological advantage of the ability to detect high-frequency hydroynamic events is discussed with respect to the natural occurrence of high frequencies and its potential role in orientation and predator-prey interactions of aquatic animals.Abbreviations LDA laser Doppler anemometer     

Consequences of forced convection for the constraints on size and shape in embryos

Previously, predictions of the maximum size of biological objects based on oxygen availability have been made for both zero and infinite water velocity around the object. In reality, however, water velocity is always intermediate between zero and infinity. We predicted maximum size and optimal shape of biological objects, pending the velocity of water around them. We assumed oxygen inside the object to be transported by diffusion and outside the object by diffusion and convection. Fick's first law of diffusion describes the inner transport. For the outer transport, we relied on semi-empirical relations between mass transport and flow conditions (Friedlander's equations). To keep mathematical complexity acceptable, we restricted ourselves to the analysis of a sphere and a cylinder in cross flow. If water velocity is low, a spherical shape is most favourable for gas exchange. If water velocity is high, an elongated and flattened shape is more favourable. A size-dependent intermediate velocity exists where shape does not matter (10(-4) m s(-1)for teleost embryos). Teleost embryos are typically exposed to flow velocities equal to or larger than 10(-4) m s(-1), making an elongated shape more favourable than a spherical one. Although teleost eggs are typically spherical, the oxygen-consuming embryos inside are indeed elongated.     

3-D-orientation with the octavolateralis system.

Fish detect and localize a sound source with inner ear receptors and with the mechanosensory lateral line. The inner ear of fish is sensitive to the water displacements caused by sound waves through a direct, inertial response by hair cell epithelia of the ear. Hearing specialists, such as goldfish and herring, have accessory peripheral structures that provide additional sensitivity to the pressure component of a sound wave. While the inner ear of fish responds to the whole body motions caused by sound waves and--in case of hearing specialists--to sound pressure, the lateral line is only sensitive to water motions relative to the surface of the fish and to local pressure gradients. Using lateral line and/or acoustic input, some fish can determine the direction and the distance to a sound source. Most likely they do so by exploiting some of the mechanisms described in this paper. Piscivorous fish may use lateral line input to detect the wakes caused by swimming fish. Even in the absence of light catfish, for instance, can follow a 10 s old, three-dimensional wake left by a prey fish over distances up to 55 prey-body length.     

The Visual Priming of Motion-Defined 3D Objects

The perception of a stimulus can be influenced by previous perceptual experience, a phenomenon known as perceptual priming. However, there has been limited investigation on perceptual priming of shape perception of three-dimensional object structures defined by moving dots. Here we examined the perceptual priming of a 3D object shape defined purely by motion-in-depth cues (i.e., Shape-From-Motion, SFM) using a classic prime-target paradigm. The results from the first two experiments revealed a significant increase in accuracy when a “cloudy” SFM stimulus (whose object structure was difficult to recognize due to the presence of strong noise) was preceded by an unambiguous SFM that clearly defined the same transparent 3D shape. In contrast, results from Experiment 3 revealed no change in accuracy when a “cloudy” SFM stimulus was preceded by a static shape or a semantic word that defined the same object shape. Instead, there was a significant decrease in accuracy when preceded by a static shape or a semantic word that defined a different object shape. These results suggested that the perception of a noisy SFM stimulus can be facilitated by a preceding unambiguous SFM stimulus—but not a static image or a semantic stimulus—that defined the same shape. The potential neural and computational mechanisms underlying the difference in priming are discussed.