In some fish lineages, evolution has led to unique sensory adaptations that provide information which is not available to terrestrial animals. These sensory systems include, among others, electroreception, which together with the ability of fish to generate electric discharges plays a role in social communication and object location. Most studies on electric phenomena in aquatic animals are dedicated to selected groups of electric fishes that regularly generate electric signals (Mormyriformes, Gymnotiformes). There exist, however, several species (hitherto described as non-electric) which, though able to perceive electric signals, have now been found to also generate them. In this article, we introduce a tool that we have designed to investigate such electric activity. This required significant adaptations of the equipment used in fish with regular discharge generation. The necessary improvements were realized by using a multielectrode registration setup allowing simultaneous visualization and quantification of behavior and associated electric activity of fish, alone or in groups, with combined electro-video clips. Precise synchronization of locomotor and electric behaviors made it possible to determine the electrically active fish in a group, and also the location of the electrogenic structure inside the fish's body. Our simple registration procedure, together with data presentation, should attract a broad audience of scientists taking up the challenge of uncovering electric phenomena in aquatic animals currently treated as electrically inactive.
Purpose of this study introduces a multielectrode array (MEA) registration system in order to generate electric field images of the episodic discharges generated by weakly electric fish. A multielectrode registration system has several important features: the design of the multielectrode lattice, the amplifier circuit, the choice of reference points for differential measurements, the recovery of the absolute values of the electric field potentials, and the application of principal components analysis. Methods. There are several advantages of our MEA registration as compared with the traditional twoelectrode registration: (a) the signal-to-noise ratio is significantly increased, (b) it is possible to construct the spatial distribution of the electric field for a single electric discharge, (c) the signals’ sources can be easily separated and identified, and (d) quantitative data on the electrical potential distribution can be obtained throughout the entire experimental tank. Results. The results illustrate an example of applied MEA registration. Electric discharges were recorded from a weakly electric catfish, Clarias gariepinus, using an array of 8 x 8 electrodes at a sampling rate of 20 kHz. Data show oscillograms and two-dimensional plots of the spatial distribution of the electrical field.
The Asian broadhead walking catfish (Clarias macrocephalus) generates weak electric monopolar pulses during spawning. Males emit a single pulse when attracting a female, and while in amplexus, only females emit a single burst of similar pulses. This burst is a necessary component in the mating ritual of C. macrocephalus. Release of milt occurs about 5 s prior to the onset of a burst, which is immediately followed by the release of eggs. Following sperm release, the male remains in tight embrace with the female. Though both male and female could perceive each other’s electric pulses via ampullary receptors (communication mode), we postulate that egg release can be facilitated by direct action of the female’s burst on the male’s neuromuscular system (contraction mode). Shedding light on the function of weak episodic electric emission, we propose that the modification of electrogenic structures evolved towards increasing the efficiency of direct bodily impact. As extant clariids exhibit intermediate features between non-electric and strongly electric catfishes, Clarias should be considered a “serviceable transition,” which Charles Darwin deemed a possible intermediate form between these two groups.
This paper explores the plasticity of sexually dimorphic characters in subadult female Brevimyrus niger, an African weakly electric mormyrid species. Thirty-five fish were exposed in a staggered fashion (five fish a week) to aromatizable 17α-methyltestosterone over a period of 7 weeks; 18 fish served as untreated controls. 17α-MT induced precocious vitellogenesis that mirrored the natural maturational process during seasonal ovarian recrudescence. At the same time, 17α-MT exposure resulted in complete masculinization of the females’ anal fin support structure normally observed during rainy season in adult males. We discuss possible hormonal mechanisms acting along the brain-pituitary-gonad axis that would explain the occurrence of precocious vitellogenesis and the male-typical transformation of the female’s anal fin ray bases. Our findings are relevant to commercial aquaculture as the use of 17α-MT in fish hatcheries can pose serious environmental issues.
This osteological survey of 249 specimens of Brevimyrus niger ranging in size from 44 to 137 mm standard length (LS) demonstrated that developmental changes in anal‐fin morphology can serve as a predictor of sexual maturity in this species. Anal‐fin ray bases begin to expand when fish reach c. 90 mm LS at which size and above there were roughly equal numbers of individuals observed with expanded and unmodified anal‐fin bases, reflecting a 1:1 sex ratio.
The aim of this study was to allow nonhuman animals to control their environment using operant conditioning procedures and to assess the effect of control on cognitive tasks. The study tested 4 predictions: (a) rats (Rattus norvegicus) will control a light stimulus; (b) animals will exhibit preferences for particular stimulus strengths; (c) animals who exert control over environmental stimuli will show improved performance on cognitive tasks compared with animals who lack control; and (d) at the end of the operant phase, experimental subjects will have lower corticosterone levels than animals who lack control. Experimental subjects did show control over a light stimulus and performed significantly better over time in a discrimination task compared with subjects who could not control their environment. There was no difference in corticosterone levels between control and experimental subjects. The results will both contribute to our understanding of how control of environmental stimuli affects the welfare of animals in captive environments and aid in designing experimental conditions that will increase validity and reliability in research.
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.
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 current study demonstrated that juvenile Mormyrus rume proboscirostris, an African freshwater weakly electric fish, used their active electrosense in group cohesion. Data also indicated that sight and mechano-reception could play a synergistic role in controlling this behaviour. The developmental change from a larval monophasic electric organ discharge to the adult biphasic waveform was accompanied by a reversal of the fish's social spacing. Light was aversive to social spacing in the younger fish (aged 49 and 65 days), but facilitated aggregation in the older fish (245 days).
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.
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.
SUMMARY African weakly discharging electric fish (Mormyridae) use their self-generated electric signals and electroreceptive abilities for orientation and communication in the context of courtship and territorial interactions. This paper documents socially mediated changes in the electric organ discharge(EOD) of subadult Gnathonemus petersii under non-breeding environmental conditions. Increases in EOD duration and changes in the relative phase amplitudes occurred in dominant fish during same-sex(male–male, female–female) and opposite-sex interactions. Similar changes were also observed in fish that were restricted in their physical interactions, suggesting that direct contact is not necessary to induce dominance-typical EOD waveforms. The possible communicative functions of these changes are discussed.
The ability to integrate multisensory information is a fundamental characteristic of the brain serving to enhance the detection and identification of external stimuli. Weakly electric fish employ multiple senses in their interactions with one another and with their inanimate environment (electric, visual, acoustic, mechanical, chemical, thermal, and hydrostatic pressure) and also generate signals using some of the same stimulus energies (electric, acoustic, visual, mechanical). A brief overview provides background on the sensory and motor channels available to the fish followed by an examination of how weakly electric fish 'benefit' from integrating various stimulus modalities that assist in prey detection, schooling, foraging, courtship, and object location. Depending on environmental conditions, multiple sensory inputs can act synergistically and improve the task at hand, can be redundant or contradictory, and can substitute for one another. Over time, in repeated encounters with familiar surrounds, loss of one modality can be compensated for through learning. Studies of neuronal substrates and an understanding of the computational algorithms that underlie multisensory integration ought to expose the physiological corollaries to widely published concepts such as internal representation, sensory expectation, sensory generalization, and sensory transfer.
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.
Adult males of African weakly discharging electric fish (family: Mormyridae) are distinguished from juveniles and adult females by a dorsally directed indentation of the posterior ventral body wall and by massive bone expansion of the bases of a select number of anal-fin rays. These sexually dimorphic structures seem to facilitate the anal-fin reflex that is displayed during courtship when the male envelopes its anal fin around the female's to form a common spawning pouch. Expanded bone could provide additional surface for muscle attachment and thus assist in part with the courtship sequence. Based on the fact that the expression of the male sexually dimorphic electric organ discharge (EOD) is under androgen control, and that the female EOD can be masculinized through testosterone administration, we hypothesized that androgens should also drive anal-fin ray bone expansion in male mormyrids and equally effect male-like changes in treated juveniles and adult females. Exogenous androgen treatment (17α-methyltestosterone) of adult femaleBrienomyrus nigerresulted in a male-like EOD, and male-typical structural transformations (body wall indentation and anal-fin ray bone expansion). Some of these changes were immediate and receded following hormone withdrawal (EOD), while others developed more slowly and were apparently permanent (indentation and bone formation). 17α-Methyltestosterone administration affected only those targets in females that are normally involved in the male's reproductive behavior, i.e., its courtship signal (EOD) and two morphological features (body-wall indentation and bone expansion). Rays of the dorsal or caudal fins were never affected.
The spectral sensitivity of the weakly electric mormyrid fish Gnathonemus petersi was investigated under dark‐ and light‐adapted conditions using a transient change (startle) in its electric organ discharge (EOD) rate as response measure. The startle was resistant to habituation and graded with light intensity. Under both lighting conditions, the fish responded optimally to a monochromatic light of 525 nm. A porphyropsin pigment (520–5402) appears to mediate spectral sensitivity over most of the visible spectrum. However, G. petersi responded more strongly to 625‐ and 675‐nm lights (dark‐ and light‐adapted fish) and a 725‐nm light (light‐adapted fish only) than predicted by the presence of a single rod pigment. These data suggest that at least one additional visual pigment (most likely of cone cells) maximally absorbing long wavelength light (600 nm or longer) is present. The spectral sensitivity data are consistent with the sensitivity hypothesis in that heightened sensitivity to long wavelength light is predicted for fish living in blackwater habitats which are characterized typically by low light levels and transmission of predominantly long wavelengths. Histology of the retina showed photoreceptors grouped into bundles and ensheathed by pigment epithelial cells. Our results demonstrated a functional visual sense in a species of fish much better known and studied for its electrosensory and electromotor abilities.