Navigation in Familiar Environments by the Weakly Electric Elephantnose Fish,Gnathonemus petersii L. (Mormyriformes,Teleostei)

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.     

Trained Weakly-electric Fishes Pollimyrus isidori and Gnathonemus petersii (Mormyridae,Teleostei) Discriminate between Waveforms of Electric Pulse Discharges

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.     

Maze Learning and Recall in a Weakly Electric Fish,Mormyrus rume proboscirostris Boulenger (Mormyridae,Teleostei)1

Mormyrus rume proboscirostris, African weakly electric fish, were trained to seek shelter in a meander maze, and following path acquisition released into the empty arena with all maze cues removed, either from the original start box or from a novel site (recall). We demonstrate that fish use their active electrosense, sight, and lateral line synergistically in maze acquisition and recall. In the presence of an electric roadmap consisting of an array of aluminum and Plexiglas objects, fish employed landmark orientation. But fish ignored visual markers and relied on internalized motor routines, which was inconsistent with evidence for cognitive mapping.     

The gross anatomy and microscopic anatomy of the tongue and lower jaw of Gnathonemus petersii (Gthr. 1862) (Mormyridae,Teleostei)

Summary This article deals with the gross and microscopic anatomy of the tongue and tongue region of the lower jaw of Gnathonemus petersii (Mormyriformes, Teleostei). The osteology coincides largely with that of Mormyrus caschive; the cranial muscles differ from those of most other fish through the absence of the m. intermandibularis posterior. The innervation of the tongue appears to be by the ramus palatinus superior n. facialis (VII). Microscopically the overall picture of the tongue is that of a deflated tube which consists at the outer portion of a compact continuous fibrous connective tissue surrounding an inner part which contains mainly loosely arranged collagen and reticular or elastic fibers with a few cells and little recognizable matrix. A comparison is made between the tongue structure of G. petersii and those of three other bony fish, Phoxinus phoxinus (Cyprinoidei), Plecoglossus altivelis (Salmonoidei) and Polypterus (Braehyopterygii).     

Electric Organ Discharge Displays during Social Encounter in the Weakly Electric Fish Brienomyrus niger L. (Mormyridae)

We investigated the electric organ discharge (EOD) activity of the mormyrid fish Brienomyrus niger during social encounters. The fish were contained in porous ceramic shelters and tested alone and in pairs in an experimental tank designed to restrict communication to the electrosensory modality. We moved one fish toward and away from a stationary conspecific, beginning at a distance known to be outside the range of communication (250 cm). Baseline EOD activity was recorded prior to interaction and categorized as ‘variable’, ‘regular’, and ‘scallop’. When moved closer together, the fish modulated this baseline activity in four ways: (1) At 100–130 cm apart, the stationary fish emitted a maximum of sudden EOD rate increases which defined the outer limit of its communication range. (The associated Electric Field Gradient was 1 μV/cm). (2) Long EOD cessations, which we called social silence, lasted from 5–130 s and occurred most frequently when the fish were 36 to 55 cm apart (EFG: 100 μV/cm). The duration of social silence was negatively correlated (r = ? 0.862) with the responding fish's size, and was independent of the partner's sex and size. Fish whose EOD baseline pattern was ‘scallop’ were least likely to fall electrically silent, and those that were categorized as ‘regular’ or ‘variable’ were most likely to cease discharging. (3) Within electrolocation range, fish ‘regularized’ their EOD activity while the partner was ‘silent’ (EFG: 1 mV/cm). (4) Following long EOD cessations the fish resumed discharging with characteristic EOD rebound patterns. The possible ethological significance of these findings is discussed.     

Adenosine triphosphatases in electroreceptor organs (ampullary organs and mormyromasts) of Gnathonemus petersii Mormyridae

Summary Cytochemical techniques were used for the light and electron microscopical localization of ATPase in the ampullary organ and the mormyromast, both cutaneous electroreceptors inGnathonemus petersii (Mormyridae).At the light microscope level, two different techniques gave the same results, namely that high concentrations of the enzyme are present in the mormyromast and certain epidermal cells and weak concentrations in the ampullary organ.The enzyme was localized at the ultrastructural level using the lead capture method. It was found in the cytoplasm of type III accessory cells of the ampullary organ, in the apical cytoplasm of SC1 sensory cells and the accessory cells surrounding them and on the membrane of the SC2 sensory cells of the mormyromast. The ATPase of these various cells was inhibited byp-chloromercuribenzoate.The enzyme in the mormyromast SC1 and their accessory cells was not dependent on Mg²⁺ ions. However, that in the type III accessory cells of the ampullary organ and in the SC2 of the mormyromast was strictly dependent on Mg²⁺. In addition, there was a Ca²⁺-dependent ATPase in the microvilli of the SC2 of certain mormyromasts.     

Permanent and Seasonal Expressions of Sexual Dimorphisms in a Weakly Electric Fish, Mormyrus rume proboscirostris Boulenger 1898 (Mormyridae, Teleostei)

Several sexually dimorphic characters of the anal-fin complex in the mormyrid fish, Mormyrus rume proboscirostris, assist during courtship when the male envelops the female's anal fin with its own to form a common spawning pouch (anal-fin reflex). We found that developmental growth and seasonally cycling gonadal activity selectively affect their expression. The structures defining the anal fin undergo a permanent sexually dimorphic transformation at a time when ripe spermatozoa first appear in the testis of young males. However, the expression of a dorsally directed indentation of the posterior ventral body wall, affecting the dorsal margin of the anal fin, appeared to be more plastic as it correlated with the gonadosomatic index, that is, testis size. We surmised that this indentation is influenced by cyclic anabolic action on muscle involved with the execution of the anal-fin reflex.     

The anal fin complex in a weakly discharging electric fish, Gnathonemus petersii(Mormyridae)

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.     

Zur Funktion des elektrischen Organs von Gnathonemus petersii (Gthr. 1862) (Mormyriformes,Teleostei)

Journal of Comparative Physiology A - In der vorliegenden Arbeit werden Untersuchungen an schwachelektrischen Fischen der Art Gnathonemus petersii (Günther 1862) beschrieben.     

Zur Empfindlichkeit des schwachelektrischen Fisches Gnathonemus petersii (Gthr. 1862) (Mormyriformes,Teleostei) gegenüber elektrischen Feldern

Zusammenfassung An schwachelektrischen Fischen der Art Gnathonemus petersii (Günther, 1862) wird die Empfindlichkeit gegenüber von außen einwirkenden elektrischen Feldern und gegenüber Veränderungen des vom Tier selbst aufgebauten elektrischen Feldes gemessen.Als Reaktion für die Wahrnehmung dieser Reize dienen charakteristische Änderungen in der Pulsfrequenz des elektrischen Organs.Die Reizschwelle gegenüber von außen einwirkenden elektrischen Feldern ist von Tier zu Tier unterschiedlich. Die empfindlichsten Tiere reagieren auf eine Feldstärke von etwa 0,7 mV/cm. Die Reizung erstreckt sich hierbei auf einen kleinen Bereich der Körperoberfläche des Tieres. Am empfindlichsten ist der Bereich zwischen Auge und Kiemenöffnung.Die Verteilungsdichte der epidermalen Sinnesorgane wurde bestimmt; sie ist in den Bereichen hoher Empfindlichkeit besonders groß. Damit wurden frühere Befunde anderer Autoren bestätigt, welche diese Sinnesorgane als die Elektrorezeptoren erkannten.Die Empfindlichkeit gegenüber Veränderungen des vom Fisch selbst aufgebauten elektrischen Feldes deckt sich in ihrem allgemeinen Verlauf mit der Empfindlichkeit gegenüber von außen einwirkenden elektrischen Feldern. Somit kommen für beide Arten von Reizen die gleichen Rezeptoren in Frage.

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Summary Weakly electric fish (Gnathonemus petersii, Mormyridae) have been investigated to determine the sensitivity to externally applied electric fields and to disturbances of the electric field emitted by the fish.Characteristic changes in the pulse frequency of the electric organ were used as criterion for the perception of these stimuli.The threshold value for externally applied fields varies for different specimen. The most sensitive annuals showed a response to a field intensity of about 0,7 mV/cm, the stimulus being applied to a small portion of the surface of the body of the animal. The most sensitive area is located between the eye and the gill slit.The distribution of epidermal sense organs has been determined and compared with the sensitivity to electric fields. Areas of high sensitivity coincide with areas of high density of epidermal sense organs. These findings confirm the results of other investigators who found these sense organs to be electroreceptors.The sensitivity to disturbances of the electric field emitted by the fish itself coincides with the sensitivity to externally applied fields. This result justifies the suggestion that both kinds of stimuli are perceived by the same types of receptors.

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Die Verfasser danken Herrn Prof. Dr.-Ing. habil. H. Tischner für die Anregung zu dieser Untersuchung und dem Bundesministerium der Verteidigung für die Bereitstellung der Mittel

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Jetzt: Zoophysiologisches Institut der Universität Tübingen.     

Enzyme activity during the metabolism of glycogen. II. Cytochemical study of glycogen synthetase in the sensory cells of the tuberous organ of Gnathonemus petersii (Mormyridae).

Glycogen synthetase (2.4.1.11) forms I (independent or active) and D (dependent or passive) as well as the enzymes active in the transformation of the pathways, protein kinase and phosphatase transferase, were studied in the sensory cells and glycogen rich epidermal cells of the weakly electric fish Gnathonemus petersii (Mormyridae). For light microscopy an indirect cytochemical method which differentiated between glycogen originally present and that produced during incubation in the presence of UDPG was used. This differentiation was obtained by iodine, PAS and alpha and beta amylases. Glycogen synthetase is present in the sensory cells in the I and D forms. The epidermal cells only contain the D form. Protein kinase (active I yields D) has only been found in the sensory cells but phosphatase transferase (active D yields I) has been found in both the epidermal cells and the sensory cells, but only within certain organs. Electron microscopy studies of glycogen synthetase I and D and protein kinase were restricted to the sensory cells only. As with the light microscope it was possible to differentiate between native glycogen and newly formed glycogen. This was done using ultrathin sections and staining with uranyl acetate, lead citrate or by the PATAg reaction. It was possible from these observations to locate precisely the positions of these enzymes. In fact, glycogen synthetase I and D are found both in the sensory cytoplasm and in the sensory cavity with the polysaccharide filaments. Protein kinase is also abundant in the sensory cytoplasm especially in the periphery of the cell near the microvillary border.     

Spatial aspects of electrolocation in the mormyrid fish, Gnathonemus petersii

The range of electrolocation in the weakly electric fish, Gnathonemus petersii, was determined for plastic and aluminium cubes. A characteristic change in the fish's EOD activity, and abrupt change to more uniform EOD intervals (regularization), was used as the criterion for object detection. The average response distances extending laterally from the fish's longitudinal axis were significantly different (p less than 0.05) for the aluminium cube (5.4 cm) and the plastic cube (7.0 cm).     

Electric organ discharge variability of Mormyridae (Teleostei: Osteoglossomorpha) in the Upper Volta system

The electric organ discharges (EODs) of five mormyrid species ( Marcusenius senegalensis , Brevimyrus niger , Petrocephalus bovei , Pollimyrus isidori , Hippopotamyrus pictus ) from different sampling sites from the Upper Volta system in West Africa were investigated. EOD waveforms were recorded at high sampling rates in order to compare signal waveform parameters of the different species from different locations. Except for H. pictus , EODs within a species differed significantly from one another in some parameters and waveform variability at least between some sampling sites. In addition, each species showed a continuous spectrum of waveform variations, all or only parts of which were found at certain localities. Although there was variability and sometimes similarities between species, the EOD waveforms were species specific. Knowing their variation spectrum, they can be used for species determination and are probably used for species recognition by the mormyrids. Similarities or differences in EOD waveform expression within a species were not related to geographical distance. By contrast, we suggest that biotic environmental factors at a given location influence the expression of EOD waveforms. These factors affect absolute measurements such as EOD duration and fast Fourier transformation peak frequency as well as the amount of variation for certain waveform parameters across species in a similar manner for a given site. Although EOD waveform might be important for the establishment of reproductive barriers between species, our results suggest that differences in waveforms may not necessarily reflect different species or speciation processes in progress.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 61–80.     

Enzyme activity during the metabolism of glycogen. I. Demonstration of phosphorylases in the sensory cells of the tuberous organ of Gnathonemus petersii (Mormyridae). Histochemical and ultrastructural methods.

Phosphorylase activities were investigated by histochemical and ultrastructural procedures in the electroreceptive sensory cells of the tuberous organ of Gnathonemus petersii. Ater incubation in G1P, G1P activated by AMP (Takeuchi and Kuriaki medium) or in G1P activated by ATP+MgSO4 (Guha and Wegman medium) newly formed polysaccharides were analysed with the iodine and P.A.S. reactions under light microscopy and, under electron microscopy, with the periodic acid thiocarbohydrazide (TCH) silver proteinate (PATAg reaction, Thiery), The newly formed polysaccharides proved the presence of glycogen phosphorylase (2.4.1.1) activities and of their branching enzymes (2.4.1.18). When G1P was activated by ATP+MgSO4, they appeared as glycogen particles with the same constitution as native glycogen. After incubation in G1P and in G1P activated by AMP they appeared as glycogen and polyglucose filaments too. In the latter case they were high concentrated. The results show that the phosphorylases are principally present in this sensory cell in their inactive form.     

Marcusenius desertus sp. nov. (Teleostei: Mormyridae), a mormyrid fish from the Namib desert

We critically compared Marcusenius specimens from the mouth of the Cunene River on the Namibia/Angola border, a harsh desert environment on the Atlantic Ocean coast virtually devoid of aerial insects with aquatic larvae which are an important food item, with Marcusenius multisquamatus Kramer & Wink, 2013 from the escarpment region of that same river, in a relatively rich and productive subtropical savannah environment. River mouth specimens were differentiated in morphology and electric organ discharges, as determined by ANOVA/MANOVA comparisons, principal component and discriminant analyses on morphological and electrophysiological characters, and genetics, including sequences of the mitochondrial cytochrome b gene, indicating reproductive isolation. Specimens from the river mouth differed from M. multisquamatus, their closest relatives, by having a shorter snout, a smaller eye diameter, and smaller nares separation. River mouth specimens were also differentiated from other, increasingly less-close relatives, such as M. altisambesi Kramer et al., 2007 from the Okavango River, Botswana, and from M. krameri Maake et al., 2014 from the Limpopo System, South Africa. We therefore designate the new species Marcusenius desertus sp. nov. for the Cunene River mouth population.