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Navigation is a ubiquitous challenge to mobile animals as it is essential for finding mates, food and shelter. It can rely on self-generated (idiothetic) as well as external (allothetic) information. The contribution of either source of information depends on multiple factors, including the sensory modality. Far-range modalities such as vision have frequently been studied in the context of navigation, but the extent to which near-range sensory systems provide information for navigation is much less understood. Here we focused on spatial learning in the weakly electric fish Gnathonemus petersii. During their nocturnal excursions these fish typically rely on their short-range active electric sense to explore their environment. We addressed how these fish navigate and how electrosensory information is integrated in navigation. All fish learned to localize a target in a Barnes-like maze. In a series of transfer tests, we found that fish followed an idiothetic navigation strategy. When this strategy failed, fish were able to integrate electrosensory information to complete the task. Our results indicate that the active electric sense contributes to navigation in a resource-efficient and context-dependent manner. Together they show that weakly electric fish can incorporate highly localized sensory input in egocentric navigation. Extending these results will be important to reveal the sensory mechanisms of egocentric navigation in fish as well as to research whether and how spatially confined near-range sensory information might be used to form global representations of space. (C) 2019 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

In humans and other vertebrates, sensory information is sometimes lateralized towards one brain hemisphere that dominates the control of a task. Although sensory lateralization may depend on the stimuli being processed, the degree or direction of lateralization can differ according to behavioural phenotype. Accordingly, personality may play an important role in lateralization, yet there is a lack of evidence regarding how lateralizations are utilized to process information and promote a personality-based response to a particular situation. Here we show that simultaneous stimulus processing and organization of personality-based responses can be accomplished via differences in laterality between senses. We demonstrate this by examining novel object inspection in the weakly electric fish Gnathonemus petersii. We found that electrosensing is lateralized in this species, but differently between personality phenotypes: bold fish were lateralized towards the right hemisphere and timid fish the left. By contrast, visual laterality did not vary with personality; rather the left hemisphere was dominant across the population, as is common for fish when visually analysing unfamiliar objects. This evidence reveals differences in functional laterality between sensory systems and the role of personality in eliciting these differences. The species has a stronger input of electrical signals than visual signals in its brain; therefore, sensory representation in the brain might drive the laterality differences.

Most animals possess multiple sensory systems, which can be used during navigation. Different senses obtain environmental information on different spatial scales and thus provide a different basis for efficient navigation. Here we used the weakly electric fish Gnathonemus petersii to investigate how different sensory inputs influence the navigational strategy and whether landmark information can be transferred flexibly between two sensory systems. Fish were trained to swim through a maze using a particular route indicated by either visual landmarks, electrical landmarks or without any landmarks. In subsequent tests, egocentric (internal cues, such as motion patterns) and allocentric cues (external cues like landmarks) were put in conflict by relocating the local landmarks. We found that all fish, independent of the available sensory input, chose the egocentric over the allocentric route. However, visual landmarks significantly improved the training duration compared to the other groups, suggesting an involvement of allocentric visual cues during route acquisition. In a second experimental series, fish were trained to use either visual or electrical landmarks for navigation and were subsequently tested in sensory transfer tests. Fish trained with visual landmarks were able to learn this allocentric navigation task and were capable of cross-modal landmark recognition, although navigation based on electrical landmarks was less efficient. The fish trained with electrical landmarks did not learn the task at all, suggesting that the short perceptual range of the electric sense prevented learning of allocentric navigation. Together our results show that the type of sensory input influences the efficiency of allocentric navigation in G. petersii and that these fish are able to use egocentric and allocentric strategies flexibly to navigate successfully under varying environmental conditions.

During active electrolocation, the weakly electric fish Gnathonemus petersii judges the distance and impedance of nearby objects. Capacitive objects, which modulate local amplitude and waveform of the fish9s electric probing signals, cast amplitude and waveform images onto the fish9s electroreceptive skin. For an unambiguous estimation of the impedance and distance of an object, the animal has to deal with multiple dependencies of object and image parameters. Based on experimentally recorded amplitude and waveform images, we investigated possible strategies of the fish to unequivocally determine both the distance and the impedance of capacitive objects. We show that the relative slope in amplitude images, but not in waveform images, is independent of object impedance and is a measure of object distance. Distance-invariant impedance estimators were obtained by two different analytical strategies. The peak modulations of both image types were ‘calibrated’ with the relative slope of the amplitude image. Impedance estimators were obtained whenever these pairs of image features (peak and relative slope) were related dynamically over two consecutive distances. A static impedance estimator termed ‘electric colour’ is postulated to arise from the relationship of an amplitude and waveform image. Our results confirm that electric colour is indeed unaffected by object distance. For electric colour estimation we suggest a minimalistic approach of just relating the peak modulations of both image types to the basal amplitude and waveform condition. Our results are discussed with regard to the anatomical and physiological organization of the fish9s electrosensory neuronal pathways and behavioural strategies of electrolocating fish.

Gnathonemus petersii use electrolocation to navigate in unfamiliar environments. The goal of these experiments was to determine whether fish could learn the location of a fixed aperture after interference with selected sensory input. By manipulating environmental cues (aperture height and water depth) and comparing the fish's performance, the contributions of the electrosensory system, vision, and hydrostatic pressure were examined. The fish's task was to find a circular aperture in a wall dividing a 200-litre aquarium into two equal compartments. In experiment 1, the position of the aperture was raised by 10.1 cm after the fish had become familiar with its original location. In experiment 2, the water level was raised by 10 cm (leaving the aperture unchanged). When the aperture was raised, intact fish found the new aperture with no difficulty, whereas blind, electrically 'silent', and sham-operated fish were slow finding the new position. When the water level was raised, all fish increased the height at which they contacted the wall, increased their electric-organ discharge (EOD) rate, and located the aperture. This increase, in response to the rapid change in water depth, suggests that all fish used hydrostatic pressure cues to maintain depth orientation, and that those fish that learned the aperture height had used hydrostatic cues to locate its position. The data suggest that G. petersii develop an internal representation based on an electrosensory central expectation and hydrostatic cues. The fish develop a sensory 'image' of their immediate environment and associate a specific image with a specific depth. As the environment becomes more familiar, the fish apparently attend less to electrosensory information and navigate according to the internal representation, relying primarily on hydrostatic pressure cues.

We investigated the electrolocation performance of the weakly electric fish, Gnathonemus petersii, in novel and familiar environments. By selectively interfering with the fish's sensory input, we determined the sensory channels necessary for navigation and orientation. The fish's task was to locate a circular aperture (diameter: 64 mm) in a wall dividing a 200–1 aquarium into two equal compartments. To assess the fish's performance, we measured (1) the time it took the fish to locate the aperture, (2) the height at which it contacted the divider, (3) its electric organ discharge rate, and (4) the frequency of divider crossings. In the first experiment (novel environment), 50 naive G. petersii assigned to five groups of 10 fish each (intact, blind, electrically “silent,” blind and “silent,” and shamoperated animals) were tested with the aperture presented randomly in one of three positions (aperture center: 7.6, 17.7, 27.8 cm from the bottom). In a novel environment, G. petersii depend on active electrolocation. Despite the changing aperture position, over the 15 trials, fish with a functioning electric organ found the aperture, whereas those without one did not. The electric organ discharge rate was inversely correlated with the amount of time spent searching for the aperture. In a second experiment (familiar environment) 20 intact fish learned the position of a fixed aperture. When we subsequently denervated the electric organ in 10 of these animals, their performance did not differ significantly from that of their conspecifics. Thus, once the fish were familiar with the aperture's position, they no longer depended on active electrolocation. We interpret and discuss this behavior as evidence for a “central expectation” and discuss its possible role in electronavigation.

In four species of mormyrid weakly electric fish (Pollimyrus isidori, Gnathonemus petersii, Mormyrus rume, and M. caschive) the electric signalling and motor behaviour during the solution of a conditioned electrolocation task was investigated. All fish had to discriminate between two objects of different electric properties which were identical otherwise. The behaviour of the fish before, during, and after electrolocation of the objects was recorded and analysed. The same types of stereotyped motor behaviours (‘probing motor acts’, PMAs) were observed in all fish during probing of the objects. Independent of the occurrence of PMAs, ‘probing’ was accompanied by a regular pattern (regularization) of electric organ discharges (EODs). Interval durations between successive discharges were kept constant on an individual specific level which was maintained in successive tests under the same conditions. This level was negatively correlated with the spectral frequency of peak power of a single EOD but was also influenced by additional parameters such as motivation. It did not depend, however, on the type of electrolocation target. Before and after investigation of the objects, the discharge rhythm was variable with inter-pulse intervals spanning a wider range than during ‘probing’. In contrast to discharge rhythm, there was no change in waveform or spectral composition of single EODs of a fish, regardless of behavioural context. In conclusion I suggest that regularization of electric signalling is an important factor during electrolocation in all mormyrids but its level is not critical. No special behavioural strategies exist which might explain different electrolocation abilities of the four mormyrid species.

Investigations on the diurnal rhythm of the motoric and electric activity were performed with the electric fish Gnathonemus petersii. A group of 11 conspecifics as well as single fishes showed an increased electric and motoric activity at night, especially at the beginning and end of the dark phase and also during feeding. These normal diurnal rhythms were changed for at least 3 days by the aggressive behaviour of two fishes who were set together for the first time. During this time an order of precedence was established.

The present study has explored the optomotor response in two species of mormyrid fish, Gnathonemus petersii and Brienomyrus niger. Both species tended to follow a black and white striped stimulus pattern under illumination levels of 6, 12, and 60 lx. The optomotor response ceased to occur under 540 lx. The behavioral data support earlier histological findings implicating the mormyrid retina in dim light vision.

Gnathonemus petersii and other African weakly electric fish (Mormyridae) produce electric signals in order to sense their environment, a process called active electrolocation. During active electrolocation electrical images of objects in the environment are cast onto a mosaic of electroreceptors, which are embedded in the animal's skin. In Gn. petersii and four other species of mormyrid fish, the densities of electroreceptor organs (mormyromasts and ampullary organs) were determined at various skin regions, including the chin, the nasal region and the back. In all species investigated, the highest mormyromast densities were found at the chin, followed by the nasal region. In addition to this comparative approach, we investigated the electroreceptive epidermis of Gn. petersii in more detail. At the moveable chin appendix, the Schnauzenorgan, scanning electron microscopy analysis helped to elucidate the mosaic-like distribution and the morphology of the electroreceptor organs and their embedding in the epidermis. Because the number of mormyromasts increased very little when fish got bigger, receptor organ densities decreased with standard length. Mormyromast density at the Schnauzenorgan, especially at its tip, highly exceeded that at all other skin regions. Along the Schnauzenorgan, receptor densities decreased exponentially from the tip towards the base. At the nasal region, mormyromast densities were more than three times higher compared to the rest of the fish's head and trunk. A similar distribution to that of the mormyromasts was found for the ampullary electroreceptor organs, which are used for passive electroreception. Our findings support the hypothesis that at least in Gn. petersii the Schnauzenorgan and the nasal region are electroreceptive foveae.

Mormyrid fish use active electrolocation to detect and analyze objects. The electrosensory lateral line lobe in the brain receives input from electroreceptors and an efference copy of the command to discharge the electric organ. In curarized fish, we recorded extracellularly from neurons of the electrosensory lateral line lobe while stimulating in the periphery with either a local point stimulus or with a more natural whole-body stimulus. Two classes of neurons were found: (1) three types of E-cells, which were excited by a point stimulus; and (2) two types of I-cells, which were inhibited by point stimulus and responded with excitation to the electric organ corollary discharge. While all neurons responded to a point stimulus, only one out of two types of I-units and two of the three types of E-units changed their firing behavior to a whole-body stimulus or when an object was present. In most units, the responses to whole-body stimuli and to point stimuli differed substantially. Many electrosensory lateral line lobe units showed neural plasticity after prolonged sensory stimulation. However, plastic effects during whole body stimulation were often unlike those occurring during point stimuli, suggesting that under natural conditions electrosensory lateral line lobe network effects play an important role in shaping neural plasticity.

SUMMARY Afferent responses to the fish's own electric organ discharge were explored in the electrosensory lobe of the mormyrid fish Gnathonemus petersii. In order to understand the neural encoding of natural sensory images,responses were examined while objects of different conductivities were placed at different positions along the skin of the fish, i.e. at different points within, and also outside, peripheral receptive fields. The presence of an object in the fish's self-generated electric field produces local modulation of transcutaneous current density. Measurement of the local electric organ discharge shows that object images formed at the electroreceptive sensory surface have an opposing center-surround, `Mexican hat' profile. This is a pre-receptor phenomenon intrinsic to the physical nature of the sensory stimulus that takes place prior to neural lateral inhibition and is independent of such central inhibition. Stimulus intensity is encoded in the latency and number of action potentials in the response of primary afferent fibers. It is also reflected in changes in the amplitude and area of extracellular field potentials recorded in the deep granular layer of the electrosensory lobe. Since the object image consists of a redistribution of current density over the receptive surface,its presence is coded by change in the activity of receptors over an area much larger than the skin surface facing the object. We conclude that each receptor encodes information coming from the whole scene in a manner that may seem ambiguous when seen from a single point and that, in order to extract specific object features, the brain must process the electric image represented over the whole sensory surface.

The mormyrids are well known for their remarkable electrogenic and electrolocation capabilities and exceptionally large cerebellum that may account for much of their total oxygen consumption. Mormyrids living in oxygen-deficient waters may use oxygen efficiently, protecting the brain from hypoxia damage; and/or brain size may be reduced. This study compares the TFL, gill lamellar density, gill lamellar area, total gill surface area, and brain size of two species of mormyrids from extremely hypoxic waters in Uganda (Gnathonemus victoriae and Petrocephalus catostoma) to two open-water species (Mormyrus kannume and Gnathonemus longibarbis) from the same region. In addition, interdemic variation was considered by comparing swamp populations of G. victoriae and P. catostoma to open-water populations of the same species. Total gill surface area of all species fell within the upper range for freshwater fishes. However, there were both intraspecific and interspecific differences in gill characters. Interdemic comparisons showed larger gill size in swamp-dwelling populations. Brain size varied among species; larger brains were characteristic of species from well-oxygenated waters. Large gill surface area may permit survival of mormyrids in oxygen-stressed environments; however, mechanisms compensating for hypoxia seem to be inadequate to support a brain size as large as that seen in fish from well-oxygenated waters.

Weakly electric fish of the pulse type electrolocate objects in the dark by emitting discrete electric organ discharges (EODs) separated by intervals of silence. Two neighbouring pulse-type fish often reduce the risk of discharging simultaneously by means of an ‘echo response’: one fish will respond to a neighbour’s EOD with a discharge of its own following at a fixed short latency so that its EOD will occur long before the next EOD of its neighbour. Although working elegantly for two partners, this simple strategy should fail in larger groups because two fish could discharge in response to the same EOD of a third fish. Here, I show that the mormyrid fish Gnathonemus petersii could use a simple mechanism to reduce this problem. Individuals were stimulated with two closely spaced pulses, the second following so as to coincide with an echo given in response to the first. All the fish examined were able to respond more to the second pulse so that most of their echoes did not collide with the second pulse. An analysis was made of how echoing more to the second pulse depends on (i) the delay at which the stimulus followed the last spontaneous EOD, (ii) the spontaneous firing rate, (iii) the intensity of the stimulus, (iv) the number of stimulus pulses, (v) the interval between stimulus pulses, and (vi) the level of previous stimulation with double pulses. The results suggest that echoing more in response to the second pulse is probably because the first pulse causes an after-effect whose inferred properties would be compatible with the properties of the mormyromast afferences thought to be involved in the echo response.

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.

Saffan®, previously known as CT1341, is an injectable steroid anesthetic for use in cats and monkeys. It is also suited to induce anesthesia in fishes. It has the convenient property to induce anesthesia of the central nervous system, and to leave the sensory system of the integument operational. The active constituents are alphaxalone and alphadolone. A dose of 24 mg/kg i.m. is sufficient for 2 to 3 hours narcosis in fish of several hundreds of grams. For small specimens administration via bath-water in doses of 2 to 4.8 mg/l for induction and half this dose for maintenance are advisable. In catfish, Ictalurus nebulosus and I. melas, of 100 g or more the induction time after i.m. injection with 21 mg/kg is about 20 min and independent of weight or temperature. The recovery time varies with temperature from 213 min at 13°C to 19 min at 19°C. In 1000 g catfish, Clarias gariepinus, the onset of surgical anesthesia occurs after 26 min on average at doses of 24 mg/kg at 24°C. In 5 g catfish, Kryptopterus bicirrhis, 4.8 mg/l bath water at 25°C makes the opercular movements disappear after 5 min. For 15 cm TL Gnathonemus petersii a good induction dose is 2 mg/l bathwater. After 20 minutes at 23°C surgical narcosis is reached.

Both routine electron microscopy and immunocytochemistry with protein A–gold were used to identify the cell types within the islet organs of four species of teleosts ( Osteoglossum bicirrhosum, Pantodon buchholzi, Notopterus chitala, and Gnathonemus petersii ) within Osteoglossomorpha, a subdivision with an ancient lineage. Four primary endocrine cell types, A, B, D, and F, were identified within the islets of the four species examined. The B- and D-cells were located mainly in the central core of the islet in the four species. In general, the A-cells were located at the islet periphery in all of the four species but in P. buchholzi and N. chitala they were also differently distributed toward the islet core. F-cells were present only at the islet periphery. Granules of B-cells in three species had a relatively homogeneous shape of the matrix core, but in O. bicirrhosum, the shape varied greatly. Variation in matrix shape of B-cell granules may indicate a different conformation of insulin molecules among at least some species of osteoglossomorphs, and this observation may have some taxonomic significance. Two somatostatin-containing (SST) D-cell types (D1 and DX) with granules of different shape were observed in all four species of osteoglossomorphs. The granules of the two D-cells immunostained either with anti-SST-25 and anti-SST-14 (D1-cells) or with anti-SST-34 (DX-cells). Immunocytochemistry confirmed that A-cells, containing glucagon-family peptides, and F-cells, containing peptides of the pancreatic polypeptide family, had granules of different shape. The cells of the islet organs of these osteoglossomorphs are more similar to those in more derived teleosts than they are to those of nonteleost actinopterygians.

Weakly electric fish produce electric signals (electric organ discharges, EODs) with a specialised electric organ creating an electric field around their body. Objects within this field alter the EOD-induced current at epidermal electroreceptor organs, which are distributed over almost the entire body surface. The detection, localisation and analysis of objects performed by monitoring self-produced electric signals is called active electrolocation. Electric fish employ active electrolocation to detect objects that are less than 12 cm away and have electric properties that are different from those of the surrounding water. Within this range, the mormyrid Gnathonemus petersii can also perceive the distance of objects. Depth perception is independent of object parameters such as size, shape and material. The mechanism for distance determination through electrolocation involves calculating the ratio between two parameters of the electric image that the object projects onto the fish’s skin. Electric fish can not only locate objects but can also analyse their electrical properties. Fish are informed about object impedance by measuring local amplitude changes at their receptor organs evoked by an object. In addition, all electric fish studied so far can independently determine the capacitative and resistive components of objects that possess complex impedances. This ability allows the fish to discriminate between living and non-living matter, because capacitance is a property of living organisms. African mormyrids and South American gymnotiforms use different mechanisms for capacitance detection. Mormyrids detect capacitance-evoked EOD waveform distortions, whereas gymnotiforms perform time measurements. Gymnotiforms measure the temporal phase shift of their EODs induced at body parts close to the object relative to unaffected body parts further away.

Electroreceptive afferents from A- and B-electroreceptor cells of mormyromasts and Knollenorgans were tested for their sensitivity to different stimulus waveforms in the weakly electric fish Gnathonemus petersii. Both A- and B-mormyromast cells had their lowest sensitivity to a waveform similar to the self-generated electric organ discharge (EOD) (around 0° phase-shift). Highest sensitivities, i.e. lowest response thresholds, in both A- and B-cells were measured at phase shifts of +135°. Thus, both cell types were inversely waveform tuned. The sensitivity of B-cells increased sharply with increasing waveform distortions. Their tuning curves had a sharp minimum of sensitivity at +7° phase shift. A-cells had a much broader waveform tuning with a plateau level of low sensitivity from +24° to −15°. Across a 360° cycle of phase-shifts, the range of thresholds was 16 dB for individual B-cells and 4.5 dB for individual A-cells. Knollenorgan afferents were tuned to 0° phase-shifted EODs and had a dynamic range of 12 dB. Lowest sensitivities were measured at a phase shift of +165°. Experiments with computer-generated stimuli revealed that the strong sensitivity of mormyromast B-cells of EOD waveform distortions cannot be attributed to any of the seven waveform parameters tested. In addition, EOD stimuli must have the correct duration for B-cells to respond to waveform distortions. Thus, waveform tuning appears to be based on the specific combination of several waveform parameters that occur only with natural EODs.

SummaryGoal-directed behavior in most cases is composed of a sequential order of elementary motor patterns shaped by sensorimotor contingencies. The sensory information acquired thus is structured in both space and time. Here we review the role of motion during the generation of sensory flow focusing on how animals actively shape information by behavioral strategies. We use the well-studied examples of vision in insects and echolocation in bats to describe commonalities of sensory-related behavioral strategies across sensory systems, and evaluate what is currently known about comparable active sensing strategies in electroreception of electric fish. In this sensory system the sensors are dispersed across the animal's body and the carrier source emitting energy used for sensing, the electric organ, is moved while the animal moves. Thus ego-motions strongly influence sensory dynamics. We present, for the first time, data of electric flow during natural probing behavior in Gnathonemus petersii (Mormyridae), which provide evidence for this influence. These data reveal a complex interdependency between the physical input to the receptors and the animal's movements, posture and objects in its environment. Although research on spatiotemporal dynamics in electrolocation is still in its infancy, the emerging field of dynamical sensory systems analysis in electric fish is a promising approach to the study of the link between movement and acquisition of sensory information.

Tapirs (Perissodactyla: Tapiridae) are the only living vertebrates, beyond the order Proboscidea, found to possess a true proboscis, defined as a flexible tubular extension of the joint narial and upper labial musculature that serves, at least in part, to grasp food. Tapirs show only partial homology and analogy with elephants in the narial and upper labial structures, as well as in the skull bones and teeth. However, superficially similar extensions in other extant vertebrates differ greatly in anatomy and function. Therefore, they deserve new names: prorhiscis (e.g. Mammalia: Saiga tatarica), prorhinosis (e.g. Chondrichthyes: Callorhinchus spp.), prorhynchis (e.g. Osteichthyes: Campylomormyrus spp.) and progeneiontis (e.g. Osteichthyes: Gnathonemus spp.). Among non-mammalian vertebrates, no bird or reptile is known to possess a proboscis. Among fishes, there are various extensions of the rostrum, jaws, 'nose' and 'chin' that lack the required narial involvement. The skulls of extinct mammals within (e.g. deinotheres) and beyond (e.g. astrapotheres) the Proboscidea confirm that a proboscis evolved independently in several mammalian lineages before the Pliocene. This convergence with tapirs presumably reflects, in part, the advantages of concentrating the olfactory sensor on what is, effectively, the tip of a long mobile upper lip. However, the proboscis does not appear to have arisen de novo in any vertebrate post-Pliocene, and its continued evolution has apparently depended on the further development of its length, flexibility and innervations, as epitomized by elephants.

Vertebrates have repeatedly been noted for having remarkably constant ratios of brain to body O2 consumption, the brain using 2–8 % of resting body O2 consumption, suggesting that evolution has put strict limits on the energetic cost of brain function. Only man, with a value of 20 %, is an exception to this rule. However, the results presented here suggest that, in the electric fish Gnathonemus petersii, the brain is responsible for approximately 60 % of body O2 consumption, a figure three times higher than that for any other vertebrate studied, including man. The exceptionally high energetic cost of the G. petersii brain appears to be a consequence both of the brain being very large and of the fish being ectothermic. It was also found that G. petersii has a high ability to utilise O2 at low levels. Thus, during falling [O2], this species was found to maintain both its O2 uptake and its electric discharge rate down to an ambient O2 level of 0.8 mg l-1 (at 26 °C), although it was unable to tolerate an [O2] below 0.3 mg l-1. During severe hypoxia (

The “novelty response” of weakly electric mormyrids is a transient acceleration of the rate of electric organ discharges (EOD) elicited by a change in stimulus input. In this study, we used it as a tool to test whether Gnathonemus petersii can perceive minute waveform distortions of its EOD that are caused by capacitive objects, as would occur during electrolocation. Four predictions of a hypothesis concerning the mechanism of capacitance detection were tested and confirmed: (1) G. petersii exhibited a strong novelty response to computer-generated (synthetic) electric stimuli that mimic both the waveform and frequency shifts of the EOD caused by natural capacitive objects (Fig. 3). (2) Similar responses were elicited by synthetic stimuli in which only the waveform distortion due to phase shifting the EOD frequency components was present (Fig. 4). (3) Novelty responses could reliably be evoked by a constant amplitude phase shifted EOD that effects the entire body of the fish evenly, i.e., a phase difference across the body surface was lacking (Figs. 3, 4). (4) Local presentation of a phase-shifted EOD mimic that stimulated only a small number of electroreceptor organs at a single location was also effective in eliciting a behavioral response (Fig. 5). Our results indicate that waveform distortions due to phase shifts alone, i.e. independent of amplitude or frequency cues, are sufficient for the detection of capacitive, animate objects. Mormyrids perceive even minute waveform changes of their own EODs by centrally comparing the input of the two types of receptor cells within a single mormyromast electroreceptor organ. Thus, no comparison of differentially affected body regions is necessary. This shows that G. petersii indeed uses a unique mechanism for signal analysis, which is different from the one employed by gymnotiform wavefish.

Corollary discharge signals associated with the motor command that elicits the electric organ discharge are prominent in the electrosensory lobe of mormyrid fish (Gnathonemus petersii). Central pathways and structures that convey these signals from the motor command nucleus to the electrosensory lobe are known anatomically, but these structures and their contributions to the various corollary discharge phenomena have not been examined physiologically. This study examines one such structure, the mesencephalic command associated nucleus (MCA).

Mormyrid fish (Gnathonemus petersii) can discriminate between ohmic and capacitive electrical objects during active electrolocation. The neural basis of this ability was investigated by recording cells in the dorsolateraland medial zones of the electrosensory lobe. Natural electric organ discharges (EODs) distorted by capacitive objects and EODs with computer generated phase shifts were used as stimuli.

Institute for Zoology, Department of Neuroethology/Sensory Ecology, University of Bonn, 53115 Bonn,North Rhine-Westphalia, GermanyABSTRACT The nocturnally active weakly electric fishGnathonemus petersii is known to employ active electro-location for the detection of objects and for orientation inits environment. The fish emits pulse-type electric signalswith an electric organ and perceives these signals withmore than 3,000 epidermal electroreceptor organs, themormyromasts, which are distributed over the animal’sskin surface. In this study, we measured the metricdimensions of the mormyromasts from different bodyregions to find structural and functional specialization ofthe various body parts. We focused on the two fovealregions of G. petersii, which are located at the elongatedand movable chin (the Schnauzenorgan; SO) and at thenasal region (NR), the skin region between the mouthand the nares. These two foveal regions were compared tothe dorsal part (back) of the fish, which contains typicalnonfoveal mormyromasts. While the gross anatomy of themormyromasts from all skin regions is similar, the metricdimensions of the main substructures differed. The mor-myromasts at the SO are the smallest and contain thesmallest receptor cells. In addition, the number of recep-tor cells per organ is lowest at the SO. In contrast, at theback the biggest receptor organs with the highest amountof receptor cells per organ occur. The mormyromasts atthe NR are in several respects intermediate betweenthose from the back and the SO. However, mormyromastsat the NR are longer than those at all other skin regions,the canal leading from the receptor pore to the innerchambers were the longest and the overlaying epidermallayers are the thickest. These results show that mormyro-masts and the epidermis they are embedded in at bothfoveal regions differ specifically from those found on therest of the body. The morphological specializations lead tofunctional specialization of the two foveae. J. Morphol.273:629–638, 2012.

Abstract. Newly imported, adult weakly discharging electric fish Gnathonemus petersii , were examined for sex differences in the waveform of their electric organ discharge and in external morphology. The discharge waveform of gonadally ripe subjects are quantitatively described at three stages corresponding to the Nigerian pre-rainy, rainy breeding, and post-rainy season in 1988. A sex difference was found in external morphology, with males displaying a distinct indentation at the dorsal margin of the anal fin, regardless of season. Pronounced sex differences in electric organ discharges were found only in fish imported during the Nigerian rainy season, and only in two phase of the tetraphasic discharge, with males exhibiting longer durations of phases 2 and 3. These sex differences were reflected in the Fourier transformation, with male discharges exhibiting lower peak power spectral frequencies than those of females. No sex differences were found in total duration of the electric discharge. Correlations between the electric discharge measures and fish length, as well as between the discharge measures themselves, revealed relationships which varied in both direction and degree as a function of sex and/or season. These results indicate that the duration of individual phases, rather than the entire duration of the electric organ discharge, function in the communication of sex differences in this mormyrid species. Previous work has shown that phase durations of the electric discharge are steroid-dependent and therefore indicative of physiological states of the animal. Thus, the kinds of information conveyed during breeding and non-breeding seasons may be quite different, given great differences in gonadal steroid levels. These findings explain previously unsuccessful attempts to find sex differences as well as a report of a sex difference in a direction opposite to that shown here in G. petersii.

Weakly electric fish can perceive electric properties of objects by monitoring the responses of their epidermal electroreceptors (mormyromasts) to their own electric organ discharges (EOD), a process known as active electrolocation. Mormyrid fish can distinguish capacitative from resistive properties of objects. It is mainly animate objects that possess capacitative properties. Water conductivity is a critical environmental factor that varies widely from season to season and has strong effects on the emitted EOD. The two goals of this study were: (1) to investigate the ability of Gnathonemus petersii to detect the properties of capacitative objects in waters of different ion content and (2) to test a recently formulated hypothesis which states that the detection of the features of a capacitative object depends on a comparison of the inputs from the two types of mormyromast primary afferents. Individuals of G. petersii were tested in a conditioned electrolocation procedure. With increasing water conductivities from 50 to 1100 μS cm−1, EOD amplitude decreased and the detection threshold for small capacitances increased. At 50 μS cm−1, the smallest detectable capacitative value was below 0.5nF; this increased to about 20nF at 800 μS cm−1. When conductivity approached about 1000 μS cm−1, fish were no longer able to electrolocate, probably because of the reduction in EOD amplitude at high conductivities. The fish’s ability to discriminate a capacitative object unequivocally from every resistive object was also tested at different conductivities. Below about 800 μS cm−1, all fish could do so. Above that conductivity, however, fish could no longer discriminate between capacitative and resistive objects of similar impedance, although they could still discriminate between objects of different impedances. The two types of receptor afferents (from the ‘A’ and ‘B’ cells) of mormyromast electroreceptor organs have different thresholds, with the B afferents being more sensitive. I suggest that only the B receptor cells remain active at about 800 μS cm-1, when the EOD amplitude is much reduced. With input from B afferents only, an unambiguous capacitance detection was no longer possible. This supports the hypothesis that capacitance detection is achieved by comparing inputs of A and B electroreceptor cells.

1. Experiment 1 employed a repeated measures design to examine the effects of captivity on sex differences in the electric organ discharge (EOD) of Gnathonemus petersii, newly imported from Africa, and maintained individually or in groups. 2. On the day of import, males exhibited longer durations of phases 2 and 3 of the EOD and lower peak power spectral frequencies (PPSFs) than females. 3. After 14 days in captivity in the laboratory, the sex differences were eliminated. After 37 days of captivity, all sex differences were still abolished, or even reversed depending on housing conditions. Males exhibited the most dramatic changes in EODs and females appeared to have higher testosterone (T) levels than males. 4. Experiment 2 was designed to investigate the effects of captivity on both behavior and endocrine status in 58 newly imported males. In this independent group design, EOD data and blood were collected from subjects over 15 days. 5. Decreases in phase 3 of the EOD and increases in PPSFs progressed over the 15 day experimental period, becoming statistically significant by days 10 and 15, respectively. Regardless of housing conditions, both T and 11-keto T dramatically decreased to near non-detectable levels by Day 5 in the laboratory. 6. Captivity causes rapid and profound changes in the endocrine system which result in dramatic changes in steroid-sensitive EODs. These findings directly link captivity, hormones, and behavior, and show why feral animals brought into captivity usually do not exhibit sexual behavior.

Gnathonemus petersii discriminates between ohmic and capacitive objects. To investigate the sensory basis of this discrimination we recorded from primary afferents that innervate either A or B mormyromast sensory cells. Modified and natural electric organ discharges were used as stimuli. In both A and B fibres frequencies below the peak-power frequency (3.8 to 4.5 kHz) of the electric organ discharge caused minimal first-spike latencies and a maximum number of spikes. A fibres did not discriminate phase-shifted stimuli, whereas B fibres responded significantly with a decrease in first-spike latency if the phase shift was only — 1°. In both A and B fibres an amplitude increase caused a decrease in spike latency and an increase in spike number; an amplitude decrease had the reverse effect. If stimulated with quasi-natural electric organ discharges distorted by capacitive objects, the responses of A fibres decreased with increasing signal distortion. In contrast, the responses of B fibres increased until amplitude effects began to dominate. Gnathonemus may use the physiological differences between A and B fibres to detect and discriminate between capacitive and purely ohmic objects.

SUMMARYWeakly electric fish generate electric fields with an electric organ and perceive them with cutaneous electroreceptors. During active electrolocation,nearby objects are detected by the distortions they cause in the electric field. The electrical properties of objects, their form and their distance,can be analysed and distinguished. Here we focus on Gnathonemus petersii (Günther 1862), an African fish of the family Mormyridae with a characteristic chin appendix, the Schnauzenorgan. Behavioural and anatomical results suggest that the mobile Schnauzenorgan and the nasal region serve special functions in electroreception, and can therefore be considered as electric foveae. We investigated passive pre-receptor mechanisms that shape and enhance the signal carrier. These mechanisms allow the fish to focus the electric field at the tip of its Schnauzenorgan where the density of electroreceptors is highest (tip-effect). Currents are funnelled by the open mouth (funnelling-effect), which leads to a homogenous voltage distribution in the nasal region. Field vectors at the trunk, the nasal region and the Schnauzenorgan are collimated but differ in the angle at which they are directed onto the sensory surface. To investigate the role of those pre-receptor effects on electrolocation, we recorded electric images of objects at the foveal regions. Furthermore, we used a behavioural response(novelty response) to assess the sensitivity of different skin areas to electrolocation stimuli and determined the receptor densities of these regions. Our results imply that both regions – the Schnauzenorgan and the nasal region – can be termed electric fovea but they serve separate functions during active electrolocation.

Three weakly electric fish (Gnathonemus petersii) were force-choice trained in a two-alternative procedure to discriminate between objects differing in their electrical characteristics. The objects were carbon dipoles in plexiglass tubing (length 2.5 cm, diameter 0.6 cm). Their electrical characteristics could be changed by varying the impedance of an external circuit to which they were connected (Fig. 1). In one (the ‘capacitance dipole’) the resistance was very low(< 3 Ω) and the capcitance variable. In the other (the ‘resistance dipole’) the resistance was variable and the capacitance low (

The receptive field of a sensory neuron is known as that region in sensory space where a stimulus will alter the response of the neuron. We determined the spatial dimensions and the shape of receptive fields of electrosensitive neurons in the medial zone of the electrosensory lateral line lobe of the African weakly electric fish, Gnathonemus petersii, by using single cell recordings. The medial zone receives input from sensory cells which encode the stimulus amplitude. We analysed the receptive fields of 71 neurons. The size and shape of the receptive fields were determined as a function of spike rate and first spike latency and showed differences for the two analysis methods used. Spatial diameters ranged from 2 to 36 mm (spike rate) and from 2.45 to 14.12 mm (first spike latency). Some of the receptive fields were simple consisting only of one uniform centre, whereas most receptive fields showed a complex and antagonistic centre-surround organisation. Several units had a very complex structure with multiple centres and surrounding-areas. While receptive field size did not correlate with peripheral receptor location, the complexity of the receptive fields increased from rostral to caudal along the fish's body.

International audience; Weakly electric fish react to resistance and capacitance of objects that locally amplify and distort their self-generated Electric Organ Discharge (EOD) received by their skin receptors. The successive-layer structure of tissues gives certain biological materials a kind of electrical anisotropy. A polarized object, for instance, will conduct current differently in one versus the other direction. This diode-like electric anisotropy should make a significant difference to a Mormyrid who emits a directional, biphasic EOD and whose receptors are sensitive to EOD amplitude and distortion changes. The ability of Gnathonemus petersii (Mormyridae) to discriminate polarity was investigated on a virtual object by manipulating changes in a circuit comprised of diodes combined in various ways. The “novelty response,” an increase in the discharge rate in response to perceived changes, was used to assess the fish’s sensitivity. Indeed, G. petersii detects polarized objects and discriminates between polarity directions. However, the diode-like anisotropy entails a voltage threshold. Because voltage decreases with distance, and the EOD comprises opposite phases of different amplitudes, the active spaces of detection and discrimination are different and depend on the object orientation. Electric polarity thus extends the “palette” of dielectric properties used by this fish to evaluate object quality, direction, and distance.

SUMMARY Spike timing-dependent plasticity that follows anti-Hebbian rules has been demonstrated at synapses between parallel fibers and inhibitory interneurons known as medium ganglionic layer (MG) neurons in the cerebellum-like electrosensory lobe of mormyrid fish. This plasticity is expressed when presynaptic activation is associated with a characteristically broad,postsynaptic action potential, lasting 7-15 ms, occurring within a window of up to 60-80 ms following synaptic activation. Since the site of plastic change is presumably in the apical dendrites, it is important to know where, when and how this broad spike is generated and the manner in which such events propagate within the intrinsic network of the electrosensory lobe. The electrosensory lobe has a strict layered organization that makes the preparation suitable for one dimension current source density analysis. Using this technique in an `in vitro' interface slice preparation, we found that following either parallel fiber stimulation or an orthogonal field stimulus, a sink appeared in the ganglionic layer and propagated into the molecular layer. Intracellular records from MG somata showed these stimuli evoked broad action potentials whose timing corresponds to this sink. TTX application in the deep fiber layer blocked the synaptically evoked ganglionic layer field potential and the `N3' wave of the outer molecular layer field potential simultaneously, while the molecular layer`N1' and `N2' waves corresponding to synaptic activation of the apical dendrites remained intact. These results confirm the hypothesis that the broad spikes of MG cells originate in the soma and propagate through the molecular layer in the apical dendritic tree, and suggest the possibility that this backpropagation may contribute to `boosting' of the synaptic response in distal apical dendrites in certain circumstances.

SUMMARY African weakly discharging electric fish (Mormyridae) use their self-generated electric signals and electroreceptive abilities for orientation and communication in the context of courtship and territorial interactions. This paper documents socially mediated changes in the electric organ discharge(EOD) of subadult Gnathonemus petersii under non-breeding environmental conditions. Increases in EOD duration and changes in the relative phase amplitudes occurred in dominant fish during same-sex(male–male, female–female) and opposite-sex interactions. Similar changes were also observed in fish that were restricted in their physical interactions, suggesting that direct contact is not necessary to induce dominance-typical EOD waveforms. The possible communicative functions of these changes are discussed.

SUMMARYAfrican mormyrids, such as Gnathonemus petersii, migrate:nocturnally, from daytime shelters to find food and return by morning, and seasonally, spawning in swamps flooded during the rainy season. The present study examined whether the fish use landmarks detected viaelectrolocation to locate an aperture, whether they detect changes in landmark size and respond appropriately, whether landmarks or hydrostatic pressure are the primary cues for navigation and whether fish of different developmental stages behave differently with respect to landmarks and navigation. The fish's task was to locate and swim through a circular aperture in a wall dividing an aquarium into two compartments. Two groups of fish were trained to find the aperture with a landmark present, while a control group had no such landmark. The water level remained constant throughout training. At the end of training,the fish's task was to locate the aperture after the landmark size had changed or the water level had increased. The results show that G. petersiiuse landmarks to orient and navigate. They can detect changes in landmark size and will modify their locomotor behaviour to integrate the change into an internal representation. If the water level changes, increasing hydrostatic pressure, the fish orient to a landmark, if present. If no landmark is present, the fish rely on an internal representation oriented to hydrostatic pressure. Larger, early-adult G. petersii located the aperture faster than smaller, sub-adult fish.

SUMMARYSeveral insects use template-matching systems to recognize objects or environmental landmarks by comparing actual and stored retinal images. Such systems are not viewpoint-invariant and are useful only when the locations in which the images have been stored and where they are later retrieved coincide. Here, we describe that a vertebrate, the weakly electric fish Gnathonemus petersii, appears to use template-matching to recognize visual patterns that it had previously viewed from a fixed vantage point. This fish is nocturnal and uses its electrical sense to find its way in the dark, yet it has functional vision that appears to be well adapted to dim light conditions. We were able to train three fish in a two-alternative forced-choice procedure to discriminate a rewarded from an unrewarded visual pattern. From its daytime shelter, each fish viewed two visual patterns placed at a set distance behind a transparent Plexiglas screen that closed the shelter. When the screen was lifted, the fish swam towards one of the patterns to receive a food reward or to be directed back into its shelter. Successful pattern discrimination was limited to low ambient light intensities of approximately 10 lx and to pattern sizes subtending a visual angle greater than 3°. To analyze the characteristics used by the fish to discriminate the visual training patterns, we performed transfer tests in which the training patterns were replaced by other patterns. The results of all such transfer tests can best be explained by a template-matching mechanism in which the fish stores the view of the rewarded training pattern and chooses from two other patterns the one whose retinal appearance best matches the stored view.

The present study describes a measurement-based model of electric image generation in the weakly electric mormyrid fish Gnathonemus petersii. Measurements of skin impedance, internal resistivity and fish body dimensions have been used to generate an electrical-equivalent model of the fish and to calculate electrical images and equivalent dipole sources for elementary resistive objects. These calculations allow us to understand how exafferent and reafferent signals are sensed by electroreceptors. An object's electric image consists of the modulation of the transcutaneous voltage profile generated by the fish's own discharge. The results suggest a set of rules for electrolocation: (1) the side of the fish where modulation is larger indicates the side on which the object is situated; (2) the object's position in the electroreceptive field is indicated by the point of maximum modulation of the transcutaneous voltage; (3) the degree of focus of the image indicates the distance to the object. In addition, center-surround opposition originating at pre-receptor level is proposed. Both experimental measurements and modeling indicate that fish skin impedance is relatively low (400-11 000IMG src="/images/symbols/capomega.gif" WIDTH="13" HEIGHT="13" ALIGN="BOTTOM" NATURALSIZEFLAG="3"cmSUP2/SUP) and mainly resistive. This low skin impedance appears to enhance the local electric organ discharge modulation, the center-surround effect, the signal-to-noise ratio for electrolocation and the active space for electrocommunication.

The weakly electric fish Gnathonemus petersii searches at night for insect larvae in tropical African streams. The aim of this study was to determine the contributions of different sensory modalities to foraging. The time that fish needed to find two randomly placed chironomid larvae was measured. The influence of various senses on search time was investigated by blocking the use of one or more senses. Active electrolocation was used by most fish for prey detection in the dark. In addition, passive electrolocation played a role in some individuals. If light was available, vision could become the dominant sense in some individuals, replacing active electrolocation. The presence of chemical cues decreased prey detection time in most fish. Prey movements also shortened search times when active electrolocation and vision were not possible, indicating that the mechanosensory lateral line also plays a role in the detection of moving prey. The results show that G. petersii uses several senses simultaneously during foraging. Each individual favours a specific combination of the available sensory inputs. If one sensory modality is eliminated, fish can switch to other modalities, indicating that the food detection system is flexible and plastic.

Mormryid electric fish (Gnathonemus petersii) respond to novel stimuli with an increase in the rate of the electric organ discharge (EOD). These novelty responses were used to measure the fish's ability to detect small changes in the amplitude and latency of an electrosensory stimulus. Responses were evoked in curarized fish in which the EOD was blocked but in which the EOD motor command continued to be emitted. An artificial EOD was provided to the fish at latencies of 2.4 to 14.4 ms following the EOD motor command.

Fish of the family Mormyridae emit weak, pulse-like electric organ discharges (EODs). The discharge rhythm is variable, but the waveform of the EOD is constant for each fish, with species- and individual characteristics. The ability of Pollimyrus isidori and Gnathonemus petersii (Mormyridae) to discriminate between different EOD waveforms was tested using a differential conditioning procedure. Fish were first trained to respond to a reference signal in swimming to a dish to receive a bloodworm (food reward). The reference signal consisted of a 10-Hz train of the digitally recorded EOD of a conspecific. Second, an alternative signal (10-Hz train of a different EOD, either from another species, or from a conspecific of the other sex) was associated with air bubbles as punishment. The two signals were played at successive trials in random order. On each trial the latency was measured between the onset of the signal and the response. 7 out of the 8 P. isidori tested and both of the two G. petersii tested associated the reference EOD with food. Among these, five P. isidori and two G. petersii responded differentially (p < 0.01) to EODs of different species. P. isidori similarly discriminated between conspecific EODs of different sexes. The quantity of different alternative EODs which could be tested was limited when fish eventually habituated to the punishment. Even when the amplitude of the EODs was randomly changed at each trial, two out of two G. petersii differentiated between EODs of the two species, and three out of three P. isidori tested differentiated between EODs within their own species. Response latencies to the rewarded signal during the basic training and during discrimination (when it had to be distinguished from the S-) were similar. G. petersii showed differential responses for S+ and S- also in the rhythm of discharge exhibited during playback, after five EOD pulses for one fish, and after a single pulse for the other. Mormyrids may therefore distinguish between conspecifics and members of other species, and even between individual conspecifics, by their EOD waveform.

1. A repertoire of five probing motor acts (PMAs) is used by Marcusenius cyprinoides in the exploratory behaviour of novel objects, introduced into the fish's surroundings: chin probing, ‘va et vient,’ lateral probing, tangential probing, and stationary probing. 2. Four of these PMAs are displayed at a defined distance from the object: 6.8±0.4 cm for a metallic rod and 3.1±0.5 cm for a porous porcelain tube. 3. The exploratory behaviour is accompanied by the appearance of a characteristic electric organ discharge activity mode in the interpulse interval histogram. This pattern is typically emitted during va et vient, but occurs also during other acts of exploratory behaviour. 4. The results indicate that PMAs are probably used to reproducably determine the conductivity characteristics of an object.

The sequence of spike-like receptor potentials (RPs) of the tuberous organ of a Mormyrid fish (Gnathonemus petersii) has been studied in vivo. In stable conditions, the receptor is submitted only to the natural effector potentials of the fish, that is, the electric organ discharges (EODs). The intervals of time between RPs were analysed in histograms, considering only the cases with bimodal curves. Two modes of discharges were considered among the mean EOD frequencies: the stable and the burst modes. The burst EODs were shown to regulate the mean frequencies of RPs near a receptor equilibrium frequency. Two series of 10 experiments were performed. In the first series, in 8 experiments a regulation of the receptor mean frequency (RRMF) could be found. In the second, when the EODs of the fish were attenuated previously by a bath to which a curarizing agent had been added, only in 4 experiments a RRMF was found. Both modes of EODs are needed during RRMF, suggesting a counting mechanism during the stable mode of the effector. A parallelism between behavior, EODs and RP rates and RRMF is suggested.

Hearing sensitivity and psychophysical tuning curves were determined for the mormyridGnathonemus petersii. Pure tone hearing thresholds were determined from 100 Hz to 2,500 Hz, with best sensitivity being about −31 dB (re: 1 dyne/ cm2) from 300 Hz to 1,000 Hz. In order to determine frequency tuning of the auditory system, psychophysical tuning curves (PTC's) were measured with the masker presented simultaneously with, or just ahead of, the 500 Hz test signal. The sound level for different frequencies needed to just mask the test tone were determined from 100 to 800 Hz. Maximum masking occurred in both forward and simultaneous conditions when the masker and the test tone were at the same frequency. As the masker was moved in frequency from 500 Hz, higher sound levels of maskers were needed to afford masking of the test tone. The data were similar in simultaneous and forward masking, with theQ 10 dB, a measure of sharpness of tuning, being about 5 in both cases. Data were compared for other species for which behavioral thresholds and PTC's are available.Gnathonemus hears about as wide a range of frequencies as the goldfish,Carassius auratus, although the PTC's for the two species are strikingly different. The PTC's forGnathonemus resemble those determined in a forward-masking paradigm for the clown knife fish,Notopterus chitala, even thoughGnathonemus has a wider hearing bandwidth.

1. Gnathonemus petersii respond to each other's electric organ discharge (EOD) with an “echo” discharge of their own at a latency of about 12 msec. This response persists until interfish distances reach about 30 cm (Fig. 4). 2. Artificial electrical stimuli were used to further characterize the response. Response threshold, latency dependence on stimulus intensity, polarity characteristics (Figs. 5–7), and differential regional sensitivity indicate that “medium” electroreceptors in the anterior region of the animal underlie the response. 3. Response probability depends upon the delay of the stimulus after the last EOD and also upon the instantaneous EOD rate (Figs. 8, 9). The echo response in turn resets the EOD rhythm of the responding animal (Fig. 10). These results suggest that the echo input pathway terminates on the presumed mesencephalic command center.

Inter-individual similarities in the electric organ discharge activity of immobile, isolated and undisturbed mormyrid fish were investigated. Two types of analysis were performed on the discharge patterns of 10 Gnathonemus petersii: (1) The Bout Interval Criterion method was used to categorise the intervals between consecutive electric pulses; (2) an analysis of sequences of acts was performed to study the serial ordering of the interpulse intervals. Interpulse intervals were demonstrated to belong to distinct classes, having similar limits for most animals. Most fish show five classes of interpulse intervals (23 to 68 ms; 69 to 108 ms; 109 to 170 ms; 171 to 212 ms; >212 ms), to which a sixth class (

1. The electric organ discharge (EOD) patterns seen in pairs of mormyrid fishes (Gnathonemus petersii) during displays related to aggression and establishment of dominance are described. 2. A new method of reliably separating the discharges of the two fish was used. In this method a fine wire was attached to the tail of one animal (Fig. 1). 3. Discharge patterns were examined at different stages during the characteristic sequence of overt behavioral events which usually occurred after an intruder was put into a tank in which another fish of the same species had been resident for 1 hour or more. The resident attacked the intruder immediately, the intruder being initially unresponsive. After a few minutes, however, the two fish entered into intense, mutual, antiparallel displays (Fig. 2). The displays occurred repeatedly during a period of 0.5–30 min. This period ended suddenly with one of the fish clearly dominant as shown by one-sided attacks and avoiding behavior by the submissive fish. 4. All attacks were accompanied by a smooth acceleration to a high discharge rate which was usually terminated abruptly (Fig. 3). Anti-parallel behavior was accompanied by similar accelerations in both fish. Interdischarge intervals during these high rates changed discretely between those of about 15 msec and those of about 9 msec (Figs. 3–9). Initial attacks before the antiparallel period usually produced no effect or a brief acceleration in the discharges of the attacked animal. Similar attacks when dominance was well established caused a slowing of the discharge rate of the attacked fish. 5. The echo response in which one fish responds to the EOD of another with a discharge of its own at a latency between 11 and 14 msec was seen at all stages of the encounter. This latency corresponded rather exactly to the gap in the interval histogram between the shorter intervals around 9 msec and the longer ones around 15 (Figs. 11, 12). This correspondence led to a degree of avoidance of near synchronous discharges during those attacks which did not cause either slowing or accelerations in the attacked animal (Fig. 11). A degree of synchrony avoidance also occurred during the mutually high discharge rates of antiparallel behavior. This resulted from the phase locking of the two discharge trains which was often present at these times and which was probably due to the echo responses (Fig. 13). 6. Several features of the individual discharge trains and of their interaction were examined during the period of antiparallel activity. This was done in order to see if some critical parameter could be detected which would allow one to predict the winner of the encounter and which might be used as a signal by the fish themselves. No single feature among those we examined was clearly and consistently related to the outcome of the encounter.

The sensory basis of group cohesion in the weak-electric fish Gnathonemus petersii was investigated in a circular tank with groups of four fish each, interacting through a wide-meshed plastic screen with intact or operated conspecifics, or with other stimulus objects. We confined these stimuli to one or two peripheral holding compartments. The response measures were obtained from the free swimming fish and included (1) the time the fish spent together as a group, (2) the time they spent in front of the holding compartments, (3) the circular distribution of the fish's positions, and (4) the mean nearest neighbour distances. Under empty compartment conditions, four different groups were tested, consisting of either (1) intact, electrically active fish, or (2) electrically ‘silent’ fish (with their electric organ surgically rendered inoperative), or (3) blind, or (4) ‘silent’ and blind animals. The loss of either sensory modality, vision or feedback from electric organ discharge, led to changes of comparable size, decreasing the time spent as a group and increasing the mean nearest neighbour distance. In fish lacking both modalities, group cohesion was further impaired. With stimuli present in one or both holding compartments, the strength of social attraction depended on the nature of the stimulus: the more intact stimulus conspecifics were present, the more densely did the fish group in front of the stimulus compartment. ‘Wired-in’ electric organ discharges (simulating waveform and intensity) and electrically ‘silent’ fish were equally attractive, but only half as attractive as intact fish. Blind free swimming fish aggregated with intact and also with ‘silent’ conspecifics. Under dim light conditions, group cohesion was predominantly, though not exclusively, affected by electrosensory feedback from the electric organ discharge and visual input. Mechanical and olfactory cues may also be involved.

We investigated group cohesion in four species of African weakly electric fish (Brienomyrus niger, Gnathonemus petersii, Marcusenius cyprinoides and Pollimyrus isidori). The attraction among members of the same and different species served as the criterion for species recognition. To identify possible mechanisms underlying this ability, conspecifics were allowed to interact through a wide-meshed plastic screen with either a pair of confined conspecifics or a pair from a related species. Members of each of the four test species were optimally attracted to their own kind, but also responded selectively to the presence of the other species. These interspecific interactions ranged from attraction to avoidance. The difference in the fish's preference to aggregate in inter- and intraspecific interactions pointed to species-specific social cues that facilitate group cohesion in mormyrid fish. Since all four species respond to each other's electric organ discharge with changes in their own discharge rate, species recognition cannot be merely a function of electroreceptor characteristics but must involve the integration of electroreceptive and other sensory cues.