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Perception and motor control traditionally are studied separately. However, motor activity can serve as a scaffold to shape the sensory flow. This tight link between motor actions and sensing is particularly evident in active sensory systems. Here, we investigate how the weakly electric mormyrid fishGnathonemus petersiiof undetermined sex structure their sensing and motor behavior while learning a perceptual task. We find systematic adjustments of the motor behavior that correlate with an increased performance. Using a model to compute the electrosensory input, we show that these behavioral adjustments improve the sensory input. As we find low neuronal detection thresholds at the level of medullary electrosensory neurons, it seems that the behavior-driven improvements of the sensory input are highly suitable to overcome the sensory limitations, thereby increasing the sensory range. Our results show that motor control is an active component of sensory learning, demonstrating that a detailed understanding of contribution of motor actions to sensing is needed to understand even seemingly simple behaviors.SIGNIFICANCE STATEMENTMotor-guided sensation and perception are intertwined, with motor behavior serving as a scaffold to shape the sensory input. We characterized how the weakly electric mormyrid fishGnathonemus petersii, as it learns a perceptual task, restructures its sensorimotor behavior. We find that systematic adjustments of the motor behavior correlate with increased performance and a shift of the sensory attention of the animal. Analyzing the afferent electrosensory input shows that a significant gain in information results from these sensorimotor adjustments. Our results show that motor control can be an active component of sensory learning. Researching the sensory corollaries of motor control thus can be crucial to understand sensory sensation and perception under naturalistic conditions.

A Henneguya sp., morphologically resembling Henneguya nyongensis Fomena & Bouix, 1996, was isolated from the gills of Peter’s elephantnose fish, Gnathonemus petersii Gunther, imported from Nigeria. Plasmodia were located between lamellae and within the gill epithelium, often leading to lamellar fusion. Although slightly smaller, the myxospores from these fish were morphologically consistent with H. nyongensis. In valvular view, spores are elongate, pyriform with a rounded posterior and tapering caudal processes. Myxospore bodies are 9.6–12.3 (mean 11.2) µm long and 4.0–4.7 (mean 4.3) µm wide. Polar capsules are pyriform, elongate, 4.5–5.2 (4.7) µm long and 1.3–1.6 (1.4) µm wide, with a characteristic neck-like structure at the apical end. Sequence generated for the 18S small subunit rRNA gene did not directly match any sequences available on GenBank, but demonstrated 91% nucleotide similarity to an unpublished Henneguya sp. infecting Mormyrus kannume Forsskal. Herein, the description of H. nyongensis is supplemented with new data on histopathology, molecular characterisation, and expanded host and geographical range.

The highly mobile chin appendage of Gnathonemus petersii, the Schnauzenorgan, is used to actively probe the environment and is known to be a fovea of the electrosensory system. It receives an important innervation from both the trigeminal sensory and motor systems. However, little is known about the premotor control pathways that coordinate the movements of the Schnauzenorgan, or about central pathways originating from the trigeminal motor nucleus. The present study focuses on the central connections of the trigeminal motor system to elucidate premotor centers controlling Schnauzenorgan movements, with particular interest in the possible connections between the electrosensory and trigeminal systems. Neurotracer injections into the trigeminal motor nucleus revealed bilateral, reciprocal connections between the two trigeminal motor nuclei and between the trigeminal sensory and motor nuclei by bilateral labeling of cells and terminals. Prominent afferent input to the trigeminal motor nucleus originates from the nucleus lateralis valvulae, the nucleus dorsalis mesencephali, the cerebellar corpus C1, the reticular formation, and the Raphe nuclei. Retrogradely labeled cells were also observed in the central pretectal nucleus, the dorsal anterior pretectal nucleus, the tectum, the ventroposterior nucleus of the torus semicircularis, the gustatory sensory and motor nuclei, and in the hypothalamus. Labeled terminals, but not cell bodies, were observed in the nucleus lateralis valvulae and the reticular formation. No direct connections were found between the electrosensory system and the V motor nucleus but the central connections identified would provide several multisynaptic pathways linking these two systems, including possible efference copy and corollary discharge mechanisms.

Mormyrids, a family of weakly electric fish, use electric pulses for communication and for extracting information from the environment (active electroreception). The electromotor system controls the timing of pulse generation. Ethological studies have described several sequences of pulse intervals (SPIs) which are related to distinct behaviors (e.g. mating or exploratory behaviors). Accelerations, scallops, rasps, and cessations are four SPI patterns reported in these fish, each showing characteristic temporal structures and large variability. This paper presents a computational model of the electromotor command circuit that reproduces SPI patterns as a function of the inputs to the model while keeping the same internal network configuration. The topology of the model is based on a simplified representation of the network with four neuron clusters (nuclei). An initial ad hoc tuned configuration (S-T) was built to reproduce nucleus characteristics and network topology as described by detailed morphological and electrophysiological studies. Then, a genetic algorithm (GA) was developed and applied to automatically tune the synaptic parameters of the model connectivity. Two different configurations obtained from the GA are presented here: one optimized to a set of synthetic examples of SPI patterns (S-GA) and another configuration adjusted to patterns recorded from freely-behaving Gnathonemus Petersii specimens (R-GA). Robustness analyses to input variability were performed to discard overfitting and assess validity. Results show that the set of SPI patterns are consistently reproduced, both for synthetic data and for recorded data. This model can be used as a tool to test novel hypotheses regarding temporal structure in electrogeneration.

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.

Karyotyping is a basic method to investigate chromosomal evolution and genomic rearrangements. Sixteen genera within the basal teleost order Osteoglossiformes are currently described cytogenetically. Our study adds information to this chromosomal dataset by determining the karyotype of Campylomormyrus compressirostris, a genus of African weakly electric fish that has not been previously examined. Our results indicate a diploid chromosome number of 2n=48 (4sm+26m+18a) with a fundamental number of FN=72. This chromosome number is identical to the number documented for the sister taxon of the genus Campylomormyrus, i.e., Gnathonemus petersii (2n=48). These results support the close relationship of Campylomormyrus and Gnathonemus. However, the karyotype formula of C. compressirostris is different from Gnathonemus petersii, thereby confirming the high variability of karyotype formulae within the Mormyridae. We infer that the differences in chromosome number and formula of Campylomormyrus relative to other mormyrids may be caused by Robertsonian fusion and pericentric inversion. In addition to the karyotype description and classification of Campylomormyrus, a ChromEvol analysis was used to determine the ancestral haploid chromosome number of osteoglossiform taxa. Our results indicate a relatively conservative haploid chromosome number of n=24 for the most recent common ancestor of Osteoglossiformes and for most of the internal nodes of osteoglossiform phylogeny. Hence, we presume that the high chromosome variability evolved recently on multiple independent occasions. Furthermore, we suggest that the most likely ancestral chromosome number of Mormyridae is either n=24 or n=25. To the best of our knowledge this is the first attempt to determine and classify the karyotype of the weakly electric fish genus Campylomormyrus and to analyze chromosomal evolution within the Osteoglossiformes based on Maximum Likelihood and Bayesian Inference analyses.

The Electrosensory Lateral Line lobe (ELL) is the first central target where the electrosensory information encoded in the spatiotemporal pattern electroreceptor afferent discharges is processed. These afferents encode the minute amplitude changes of the basal electric field through both a change in latency and discharge rate. In the ELL the time and rate-coded input pattern of the sensory periphery goes through the granular cell layer before reaching the main efferent cells of the network: large fusiform (LF) and large ganglion (LG) cells. The evidence until now shows that granular cells are inhibitory. Given that large fusiform cells are excited by the sensory input, it remains a mystery how the afferent input produce excitation through a layer composed by only inhibitory cells. We addressed this problem by modeling how the known circuitry of the ELL could produce excitation in LF cells with only inhibitory granular cells. Alternatively we show that a network composed of a mix of excitatory and inhibitory granular cell not only performs better, as expected, carrying excitation to LF cells but it does so robustly and at higher sensitivity by enhancing the contrast of the electric image between the periphery and the ELLs output. We then show with refined histological methods that a subpopulation of the granular cells indeed are excitatory, providing the necessary input for this contrast enhancing mechanism. Copyright 2016 Elsevier Ltd. All rights reserved.

African weakly electric mormyrid fish show a high diversity of their electric organ discharge (EOD) both across and within genera. Thanks to a recently developed technique of artificial reproduction in mormyrid fish, we were able to perform hybridizations between different genera and within one genus (Campylomormyrus). The hybrids of intergenus hybridizations exhibited different degrees of reduced survival related to the phylogenetic distance of the parent species: hybrids of the crosses between C. rhynchophorus and its sister genus Gnathonemus survived and developed normally. Hybrids between C. rhynchophorus and a Mormyrus species (a more basal clade compared to Campylomormyrus s) survived up to 42days and developed many malformations, e.g., at the level of the unpaired fins. Hybrids between C. numenius and Hippopotamyrus pictus (a derived clade, only distantly related to Campylomormyrus) only survived for two days during embryological development. Eight different hybrid combinations among five Campylomormyrus species (C. tamandua, C. compressirostris, C. tshokwe, C. rhynchophorus, C. numenius) were performed. The aim of the hybridizations was to combine species with (1) either caudal or rostral position of the main stalk innervating the electrocytes in the electric organ and (2) short, median or long duration of their EOD. The hybrids, though they are still juveniles, show very interesting features concerning electrocyte geometry as well as EOD form and duration: the caudal position of the stalk is prevailing over the rostral position, and the penetration of the stalk is dominant over the non-penetrating feature (in the Campylomormyrus hybrids); in the hybrid between C. rhynchophorus and Gnathonemus petersii it is the opposite. When crossing species with long and short EODs, it is always the long duration EOD that is expressed in the hybrids. The F1-Hybrids of the cross C. tamandua×C. compressirostris are fertile: viable F2-fish could be obtained with artificial reproduction.

The African weakly electric fish Gnathonemus petersii is capable of cross-modal object recognition using its electric sense or vision. Thus, object features stored in the brain are accessible by multiple senses, either through connections between unisensory brain regions or because of multimodal representations in multisensory areas. Primary electrosensory information is processed in the medullary electrosensory lateral line lobe, which projects topographically to the lateral nucleus of the torus semicircularis (NL). Visual information reaches the optic tectum (TeO), which projects to various other brain regions. We investigated the neuroanatomical connections of these two major midbrain visual and electrosensory brain areas, focusing on the topographical relationship of interconnections between the two structures. Thus, the neural tracer DiI was injected systematically into different tectal quadrants, as well as into the NL. Tectal tracer injections revealed topographically organized retrograde and anterograde label in the NL. Rostral and caudal tectal regions were interconnected with rostral and caudal areas of the NL, respectively. However, dorsal and ventral tectal regions were represented in a roughly inverted fashion in NL, as dorsal tectal injections labeled ventral areas in NL and vice versa. In addition, tracer injections into TeO or NL revealed extensive inputs to both structures from ipsilateral (NL also contralateral) efferent basal cells in the valvula cerebelli; the NL furthermore projected back to the valvula. Additional tectal and NL connections were largely confirmatory to earlier studies. For example, the TeO received ipsilateral inputs from the central zone of the dorsal telencephalon, torus longitudinalis, nucleus isthmi, various tegmental, thalamic and pretectal nuclei, as well as other nuclei of the torus semicircularis. Also, the TeO projected to the dorsal preglomerular and dorsal posterior thalamic nuclei as well as to nuclei in the torus semicircularis and nucleus isthmi. Beyond the clear topographical relationship of NL and TeO interconnections established here, the known neurosensory upstream circuitry was used to suggest a model of how a defined spot in the peripheral sensory world comes to be represented in a common associated neural locus both in the NL and the TeO, thereby providing the neural substrate for cross-modal object recognition.

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.

Animal multisensory systems are able to cope with discrepancies in information provided by individual senses by integrating information using a weighted average of the sensory inputs. Such sensory weighting often leads to a dominance of a certain sense during particular tasks and conditions, also called sensory capture. Here we investigated the interaction of vision and active electrolocation during object discrimination in the weakly electric fish Gnathonemus petersii. Fish were trained to discriminate between two objects using both senses and were subsequently tested using either only vision or only the active electric sense. We found that at short range the electric sense dominates over vision, leading to a decreased ability to discriminate between objects visually when vision and electrolocation provide conflicting information. In line with visual capture in humans, we call this dominance of the electric sense electrosensory capture. Further, our results suggest that the fish are able to exploit the advantages of multiple senses using vision and electrolocation redundantly, synergistically and complementarily. Together our results show that by providing similar information about the environment on different spatial scales, vision and the electric sense of G. petersii are well attuned to each other producing a robust and flexible percept.

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.

The weakly electric fish Gnathonemus petersii uses its electric sense to actively probe the environment. Its highly mobile chin appendage, the Schnauzenorgan, is rich in electroreceptors. Physical measurements have demonstrated the importance of the position of the Schnauzenorgan in funneling the fish's self-generated electric field. The present study focuses on the trigeminal motor pathway that controls Schnauzenorgan movement and on its trigeminal sensory innervation and central representation. The nerves entering the Schnauzenorgan are very large and contain both motor and sensory trigeminal components as well as an electrosensory pathway. With the use of neurotracer techniques, labeled Schnauzenorgan motoneurons were found throughout the ventral main body of the trigeminal motor nucleus but not among the population of larger motoneurons in its rostrodorsal region. The Schnauzenorgan receives no motor or sensory innervation from the facial nerve. There are many anastomoses between the peripheral electrosensory and trigeminal nerves, but these senses remain separate in the sensory ganglia and in their first central relays. Schnauzenorgan trigeminal primary afferent projections extend throughout the descending trigeminal sensory nuclei, and a few fibers enter the facial lobe. Although no labeled neurons could be identified in the brain as the trigeminal mesencephalic root, some Schnauzenorgan trigeminal afferents terminated in the trigeminal motor nucleus, suggesting a monosynaptic, possibly proprioceptive, pathway. In this first step toward understanding multimodal central representation of the Schnauzenorgan, no direct interconnections were found between the trigeminal sensory and electromotor command system, or the electrosensory and trigeminal motor command. The pathways linking perception to action remain to be studied.

Weakly electric fish orient, hunt and communicate by emitting electrical pulses, enabling them to discriminate objects, conspecifics and prey. In addition to the electrosensory modality - although dominating in importance in these fishes - other modalities, like vision, play important roles for survival. The visual system of Gnathonemus petersii, a member of the family mormyridae living in West African blackwater streams shows remarkable specializations: Cone photoreceptors are grouped in bundles within a light reflecting tapetum lucidum, while the rods are also bundled but located at the back within a light-scattering guanine layer. Such an organization does not improve light sensitivity nor does it provide high spatial resolution. Thus, the function of the grouped retinal arrangement for the visual performance of the fish remains unclear. Here we investigated the contrast sensitivity of the temporal transfer properties of the visual system of Gnathonemus. To do so, we analyzed visual evoked potentials in the optic tectum and tested the critical flicker fusion frequency in a behavioral paradigm. Results obtained in Gnathonemus are compared to results obtained with goldfish (Carassius auratus), revealing differences in the filter characteristics of their visual systems: While goldfish responds best to low frequencies, Gnathonemus responds best at higher frequencies. The visual system of goldfish shows characteristics of a low-pass filter while the visual system of Gnathonemus has characteristics of a band-pass filter. Furthermore we show that the visual system of Gnathonemus is more robust towards contrast reduction as compared to the goldfish. The grouped retina might enable Gnathonemus to see large, fast moving objects even under low contrast conditions. Due to the fact that the electric sense is a modality of limited range, it is tempting to speculate that the retinal specialization of Gnathonemus petersii might be advantageous for predator avoidance even when brightness differences are small.

Summary Many sighted animals use color as a salient and reliable cue [ 1 ] to identify conspecifics [ 2 , 3 , 4 ], predators, or food [ 5 , 6 , 7 ]. Similarly, nocturnal, weakly electric fish Gnathonemus petersii might rely on “electric colors” [ 8 ] for unambiguous, critical object recognitions. These fish identify nearby targets by emitting electric signals and by sensing the object-evoked signal modulations in amplitude and waveform with two types of epidermal electroreceptors (active electrolocation) [ 9 , 10 , 11 , 12 ]. Electrical capacitive objects (animals, plants) modulate both parameters; resistive targets (e.g., rocks) modulate only the signal’s amplitude [ 11 , 12 ]. Ambiguities of electrosensory inputs arise when object size, distance, or position vary. While previous reports suggest electrosensory disambiguations when both modulations are combined as electric colors [ 8 , 13 , 14 ], this concept has never been demonstrated in a natural, behaviorally relevant context. Here, we assessed electric-color perception (1) by recording object-evoked signal modulations and (2) by testing the fishes’ behavioral responses to these objects during foraging. We found that modulations caused by aquatic animals or plants provided electric colors when combined as a ratio. Individual electric colors designated crucial targets (electric fish, prey insect larvae, or others) irrespective of their size, distance, or position. In behavioral tests, electrolocating fish reliably identified prey insect larvae of varying sizes from different distances and did not differentiate between artificial prey items generating similar electric colors. Our results indicate a color-like perceptual cue during active electrolocation, the computation [ 15 ], reliability, and use of which resemble those of color in vision. This suggests “color” perception as a sensory concept beyond vision and passive sensing.

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.

Several species of Mormyrid weakly electric fish have a mobile chin protuberance that serves as a mobile antenna during prey detection, tracking behaviors, and foraging for food. It has been proposed that it constitutes a fovea of the electrosensory system. The distribution of the three types of receptor organs involved in active imaging of the local surroundings, prey detection, and passive electroreception, and their central projection to the electrosensory lobe (ELL), have been studied in Gnathonemus petersii. Density distributions were compared for different body regions. Primary afferent projections were labeled with biocytin or biotinylated dextrans. This showed that there is considerable central "over-representation" of the mandibular and nasal regions of the sensory surface involved in electrolocation, at the expense of the other body regions investigated. This over-representation is not a mere effect of the very high density of receptor organs in these areas, but is found to be due to central magnification. This magnification differs between the subclasses of electroreceptors, suggesting a functional segregation in the brain. We conclude that the chin protuberance and the nasal region are the regions of greatest sensitivity for the resistive, capacitive, and low-frequency characteristics of the environment, and are probably most important in prey detection, whereas other regions of the skin with a lesser resolution and sensitivity to phase distortion of the EOD, in particular the trunk, are probably designed for imaging larger, inanimate features of the environment. Our data support the hypothesis that the chin appendage and nasal region are functionally distinct electrosensory foveae.

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.

Mormyrus rume rume, Hyperopisus bebe bebe, Campylomormyrus tamandua and Gnathonemus petersii sampled from the Anambra river with a fleet of gill nets, traps and hook and line were examined for endo-parasitic helminthes from October 2005 to September 2006. Recovered helminthes were Rhadinorhynchus horridus from the intestine of H. bebe bebe and G. petersii; Procamallanus laeviconchus from the stomach of M. rume rume and C. tamandua; Spinitectus mormyri from the stomach of M. rume rume; Contracaecum sp. from the coelom of H. bebe bebe, G. petersii and C. tamandua, whereas an unidentified cestode infected the intestine of all the mormyrids. G. petersii constitutes a new host record for R. horridus; M. rume rume for S. mormyri and M. rume rume and C. tamandua for P. laeviconchus. In cases of mixed infection the parasites occupied their preferred habitats. The overall prevalence of the endo-parasites in the fish hosts was 41.9%, which is within the range (< 50%) typical of southern Nigerian freshwater lotic habitats. Prevalence, mean intensity and mean abundance of R. horridus in its host fishes were higher in the dry (October/November-March) than the rainy season (April-September/October); in the nematodes (P. laeviconchus, S. mormyri and Contracaecum) the reverse was the case, whereas no definite pattern was exhibited by the unidentified cestode. P. laeviconchus appeared reddish from engorgement of blood but no damage was evident at the point of attachment. Local inflammation where R. horridus was attached to the mucosa of the intestine was not considered severe. R. horridus and P. laeviconchus are probably the most important parasites of the mormyrids in terms of fishery management in the Anambra river system.

To elucidate the specific properties of retinae with grouped photoreceptors the neural pathways in the outer and inner plexiform layer were studied. Photoreceptor bundles in this species consist of more than 100 rods and up to 50 cones, and are usually regarded as functional units. Golgi impregnation in thick and thin sections and light microscopy were used to identify bipolar cell types linking photoreceptors to amacrine and/or ganglion cells. Nine different types were distinguished based on their dendritic morphology and the position of the axon terminal in the inner plexiform layer. Small cells have dendritic fields smaller than the diameter of a photoreceptor bundle and are contacted mostly by cones. The dendritic field size of bushy cells matches that of a photoreceptor bundle; they are contacted mainly by rods. Flat cells receive about equal input from rods and cones; their dendritic field size exceeds the bundle diameter. Within the three major classes there are subtypes addressing three sublaminae of the inner plexiform layer, the proximal On-centre region (sl b), the distal Off-centre region (sl a) and a central sublayer (sl c) probably with transient activity. These observations suggest that cone vision has a spatial acuity better than the “bundle grain”. In rod dominated vision the resolution matches that of the bundles; for this pathway, the hypothesis of the bundle as a functional unit is confirmed. The mesopic flat cell pathway has a resolution inferior to the “bundle grain”; it may therefore be dedicated to movement detection.

The cerebellum of mormyrid fish is of interest for its large size and unusual histology. The mormyrid cerebellum, as in all ray-finned fishes, has three subdivisions--valvula, corpus, and caudal lobe. The structures of the mormyrid valvula and corpus have been examined previously, but the structure of the mormyrid caudal lobe has not been studied. The mormyrid caudal lobe includes a posterior caudal lobe associated with the electrosense and an anterior caudal lobe associated with lateral line and eighth nerve senses. In this article we describe cellular elements of the posterior caudal lobe and of the eminentia granularis posterior (EGp) in the mormyrid fish Gnathonemus petersii. The EGp gives rise to the parallel fibers of the posterior caudal lobe. We used intracellular injection of biocytin, extracellular injection of biotinylated dextran amine, and immunohistochemistry with antibodies to gamma-aminobutyric acid, inositol triphosphate receptor I, calretinin, and Zebrin II. The histological structure of the posterior caudal lobe is markedly irregular in comparison to that of the corpus and the valvula, and a tight modular organization of cerebellar elements is less apparent here. Most Purkinje cell bodies are in the middle of the molecular region. Their dendrites are only roughly oriented in the sagittal plane, extend both ventrally and dorsally, and branch irregularly. Climbing fibers terminate only on smooth dendrites near the soma. Most Purkinje cell axons terminate locally on eurydendroid cells that project outside the cortex. The results provide an additional variant to the already large set of different cerebellar and cerebellum-like structures.

Electroreceptive fish detect nearby objects by processing the information contained in the pattern of electric currents through the skin. The distribution of local transepidermal voltage or current density on the sensory surface of the fish's skin is the electric image of the surrounding environment. This article reports a model study of the quantitative effect of the conductance of the internal tissues and the skin on electric image generation in Gnathonemus petersii (Günther 1862). Using realistic modelling, we calculated the electric image of a metal object on a simulated fish having different combinations of internal tissues and skin conductances. An object perturbs an electric field as if it were a distribution of electric sources. The equivalent distribution of electric sources is referred to as an object's imprimence. The high conductivity of the fish body lowers the load resistance of a given object's imprimence, increasing the electric image. It also funnels the current generated by the electric organ in such a way that the field and the imprimence of objects in the vicinity of the rostral electric fovea are enhanced. Regarding skin conductance, our results show that the actual value is in the optimal range for transcutaneous voltage modulation by nearby objects. This result suggests that “voltage” is the answer to the long-standing question as to whether current or voltage is the effective stimulus for electroreceptors. Our analysis shows that the fish body should be conceived as an object that interacts with nearby objects, conditioning the electric image. The concept of imprimence can be extended to other sensory systems, facilitating the identification of features common to different perceptual systems., Synopsis This paper analyzes the contribution of the body and skin conductance of weakly electric fish in shaping the electric image, using a realistic computational model. Object recognition, a relevant issue in sensory systems, is not yet fully understood. How pre-receptor mechanisms and interactions between objects shape images is of interest in all sensory systems, leading to general concepts and a specialized jargon. The authors rescue the generality of two concepts for understanding sensory systems. These concepts were introduced early in electroreception research: object perturbed field (change in the basal field generated by the presence of an object), and imprimence (equivalent sources produced at the location of the object). The fish body is an object: generating its imprimence, modifying the basal field, and interacting with other objects. Analogously, the human body interferes, reflecting light or sound in the generation of visual and acoustic images. The contribution of internal and skin conductivities in image generation has been controversial since the seminal work of Lissmann and Machin in 1958. We show that the high internal conductance of the fish increases and redirects the currents that illuminate objects, increasing and shaping the electric image. Skin resistance mainly influences image magnitude.

An examination of the permanent bony structures of the anal fin complex in the mormyrid fish, Gnathonemus petersii, revealed two new structural sexual dimorphisms: longer proximal pterygiophores and wider anal fin rays in males than in females. Both structures are thought to facilitate the male’s courtship-associated anal fin reflex. Adult male mormyrid fishes are characterized by a dorsally directed indentation of the posterior body wall (anal fin indentation). The expression of this indentation in males, presumably driven by anal fin musculature, was correlated with the fish’s gonadal state: large indentations were associated with high gonado-somatic indices and small indentations with low indices.

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.

In pairs of interacting Gnathonemus petersii, a mormyrid electric fish, dominant individuals were larger (longer body). Pairs of interacting fish of similar body size emitted more parallel displays at the onset of an interaction. Over the course of 10 min interactions, head butting increased and parallel display decreased. This decrease occurred primarily in pairs that contained a prior resident and intruder, as compared with two intruders. The patterns of electric organ discharge during parallel display, and the possible sensory modalities mediating this behaviour are discussed.

The weaklyelectric fish Gnathonemus petersii detects, localizes, and analyzes objects during active electrolocation even in complete darkness. This enables these fish to lead a nocturnal life and find and identifytheir prey(small insect larvae) on the ground of their freshwater habitat. During active electrolocation, fish produce a series of brief electric signals, electric organ discharges (EOD), with an electric organ in their tail. Each EOD builds up a stable electric field around the fish, which is distorted onlyby nearbyobjects. Field distortions lead to changes of the transepidermal electric current flow at a region of the fish’s electroreceptive skin surface called the ‘electric image’. Within the electric image, locallyperceived EODs can be either altered in amplitude or waveform byan object. Fish measure both parameters to assess object properties, such as the capacitive and resistive components of the object’s complex impedance, the object’s size and shape, and its distance from the fish. None of these object properties can be evaluated in isolation, but have to be inferred during parallel processing of electric image spatial and qualitative parameters. Two anterior skin regions of G. petersii appear to possess particular properties for special electrolocation tasks and we therefore refer to them as ‘foveal’ regions. Because of its high electroreceptor density, the electric field geometry around it, and its behavioral use, the ‘nasal region’ between the nares and the mouth at the head of the fish is suggested to be a fovea for long-range guidance and object detection. We propose that the ‘Schnauzenorgan’, a long and flexible chin appendix covered denselywith electroreceptor organs, is a second electroreceptive fovea associated with a short-range (food) identification system. Together, these two electric foveae constitute an effective preydetection and identification sy stem. # 2003 Elsevier Ltd. All rights reserved.

We review modelling and experimental work dealing with the mechanisms of generation of electric image. We discuss: (1) the concept of electric image in the context of the reafference principle; (2) how waveform codes an impedance related qualia of the object image, referred to as “electric colour”; (3) that some characteristics of the spatial profiles generated by pre-receptor mechanisms are suitable for edge detection; (4) which parameters of the spatial profiles provide information for distance discrimination; (5) that electric images are distributed representations of the scene.

This study examined how weakly electric fish, Gnathonemus petersii, integrate multiple sensory modalities (passive and active electrosenses, and vision) to maintain proximity to tubular structures, serving as the fish’s hiding place or shelter during the daytime. By moving the shelter along a linear 2-meter path, causing a mechanical disturbance, we challenged the fish’s shelter-seeking behavior and used the length of travel that shelter proximity was maintained (contact distance) as an indicator of how well the animal maintained its shelter. In order to determine the contribution of vision and electrosense to this behavior, four groups of fish were tested in which: (1) all three modalities were intact; (2) vision alone was eliminated by optic nerve transection; (3) the active electrosense was silenced by spinal cord transection rendering the electric organ inoperative; and (4) both vision and active electrosense were deactivated. Further elimination or minimization of various sensory cues was achieved by testing the fish with optically transparent, acrylic shelters (Plexiglas) that stimulate active, but not passive, electrosense, and aluminum shelters that theoretically stimulate all three modalities. As expected, performance was optimal when all three modalities were operating, but better than expected from quantitative models based on additive processes alone. Although the absence of one sense (vision or active electrosense) caused initial deficits, these were fully compensated for over repeated daily exposure to the task, suggesting that learning might generate sensory substitution and/or the formation of sensory expectation. Finally, environmental conditions, such as shelter opacity, also affected shelter-seeking performance, sometimes in a negative direction. These results demonstrate that: (1) the integration of multiple sensory inputs in G. petersii can be synergistic, additive, redundant, or even inhibitory, and (2) multisensory processes also take into account the respective sensory cues; i.e. (a) the prevailing ambient light intensity and optical qualities of the object; (b) the geometry and strength of the DC potential emanating from the object (‘battery effect’); and (c) the complex perceived impedance differential with the surrounding medium.

The weakly electric fish Gnathonemus petersii uses active electrolocation for non-visual orientation and object detection. Fish emit brief electric signals, which build up an electric field around their body. Nearby objects project electric images onto the fish's skin, which are perceived by epidermal electroreceptors, enabling the animal to discriminate between objects and to detect their electric properties. To test up to what distance object recognition is possible and whether complex backgrounds influence the discrimination performance, animals were trained to detect various objects of different sizes and shapes in front of different stagnant or moving backgrounds. Our results show that G. petersii is able to distinguish between objects up to a distance of about 3 cm even in complete darkness. Moving backgrounds improve the discrimination distance up to 4 cm. Using electric fish as models, underwater robots can be given an electric sense, which will enable them to navigate in dark or turbid environments. Our bionic approach will allow the robots to detect and analyse objects and recognize different object properties such as material composition, size and shape. Object recognition based on active electrolocation will still be possible in opaque surroundings where visual orientation is impossible.

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.

In part I (Shuai et al. 1998. Biophys. J . 75:1712–1726), we presented a cellular model of the A- and B-electroreceptors of the weakly electric fish Gnathonemus petersii. The model made clear the cellular origin of the differences in the response functions of A- and B-receptors, which sensitively code the intensity of the fish's own electric organ discharge (EOD) and the variations in the EOD waveform, respectively. The main purpose of the present paper is to clarify the cellular origin of the inverse waveform tuning of the B-receptors by using the receptor model. Inverse waveform tuning means that B-receptors respond more sensitively to the 180° inverted EOD than to undistorted or less distorted EODs. We investigated how the A- and B-receptor models respond to EODs with various waveforms, which are the phase-shifted EODs, whose shift angle is varied from −1° to −180°, and single-period sine wave stimuli of various frequencies. We show that the tuning properties of the B-receptors arise mainly from the combination of two attributes: 1) The waveform of the stimuli ( B stim ) effectively sensed by the B-receptor cells. This consists of a first smaller and a second larger positive peak, even though in the original phase-shifted EOD stimuli, the amplitudes of the two positive peaks are reversed. 2) The effective time constant of dynamical response of the receptor cells. It is on the order of the duration of a single EOD pulse. We also calculated the response properties of the A- and B-receptor models when stimulated with natural EODs distorted by various capacitive and resistive objects. Furthermore, we investigated the effect of EOD amplitude on the receptor responses to capacitive and resistive objects. The models presented can systematically reproduce the experimentally observed response properties of natural A- and B-receptor cells. The mechanism producing these properties can be reasonably explained by the variation in the stimulus waveforms effectively sensed by the A- and B-receptor cells and by time constants.

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.

The paraventricular organ of Gnathonemus petersii was investigated with light and electronmicroscopical techniques. It contains high concentrations of dopamine, noradrenaline and serotonin, but the synthesizing enzymes are not or hardly present. Consequently, the cerebrospinal fluid-contacting neurons might pick up their biogenic amines from the ventricular fluid. Dense subependymal axonal plexuses in the everted telencephalon probably release these substances into the ventricle. However, electronmicroscopical observations suggest release rather than uptake by the paraventricular organ. The possible significance of intraventricular release, transport and uptake of biogenic amines is discussed.

Male Gnathonemus petersii (Mormyridae) exhibit two structural, sexually dimorphic characters: anal-fin ray bone expansion and an indentation of the posterior ventral body wall (formerly described as anal-fin indentation). Females lack this bone expansion, but may show a slight indentation. Morphometric data on both characters were obtained from radiographs of 414 fish (males and females) ranging in size from 60 to 276 mm Ls. Both body wall indentation and bone expansion began to develop in males of about 120 mm Ls. At 160–180 mm, the sex ratio of fish with expansion to fish without expansion was 50 : 50. Androgens seem to affect the expression of both these sexual dimorphisms. Bone expansion may provide increased bone surface for muscle attachment and thus facilitate the anal-fin reflex during courtship behaviour.

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.

The retina of the weakly electric fish Gnathonemus petersii is a so-called grouped retina where photoreceptors are bundled. These bundles are regarded as functional units and this type of retinal specialization is uniquely found in teleosts. To understand how this anatomical organization influences visual information processing we investigated the morphology and distribution of retinal ganglion cells (GCs) and the response properties of retinal afferents terminating in the major retinorecipient area, the optic tectum. GCs were classified based on their dendritic morphology (dendritic field diameters 280 μm: giant GCs). Within these classes subtypes were distinguished based on the ramification patterns of the dendrites in the sublaminae of the inner plexiform layer. Properties of presumed optic nerve terminals were investigated in the optic tectum using extracellular recordings. Physiological classes could be observed based on their response to visual stimuli (on; off; on-off, and fast units). Receptive field sizes and spatiotemporal properties were classified and the topographical representation of the visual space was mapped in the tectum. Gratings of low spatial frequencies were best responded to and followed up to high temporal frequencies (>30 Hz). Most of the recorded units were directionally selective. No evidence of distorted topographies in the tectum was found, i.e., no overrepresentation of the retina was seen in the tectum opticum. The grouped retina of G. petersii seems to be optimized for the detection of large, fast objects in an environment of low optical quality.

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.