Construímos um aparato experimental para medir os instantes de disparo do órgão elétrico de peixes elétricos de campo fraco da espécie Gymnotus carapo, que produz estes pulsos para localizar objetos dentro da água e para se comunicar socialmente. O aparato foi desenvolvido de maneira a iisolar o animal de perturbações externas como vibrações mecânicas, sons, campos elétricos e variações de luminosidade do ambiente. A principal característica de nosso aparato é um conjunto de eletrodos, distribuídos nos vértices do tanque de experimentos, que permitem obter as medidas (longas séries de instantes de disparo) sem restringir os movimentos do peixe e até mesmo inferir a sua posição comparando as amplitudes em diferentes eletrodos, o que possibilita relacionar a posteriori os padrões de disparo ao comportamento do animal. Desenvolvemos um programa de computador em linguagem C que, através de uma interface digital­analógica reproduz a série temporal da voltagem de um pulso de um peixe verdadeiro e utilizamos este sinal elétrico para estimular os animais. Os pulsos artificiais foram aplicados a um dipolo elétrico que imita a geometria do órgão elétrico de um peixe e os intervalos entre pulsos foram produzidos por diferentes distribuições: aleatória, intervalos gravados previamente do próprio ou de outro peixe, sequências manipuladas para repetir determinados trechos reais intercalados com trechos aleatórios, etc. Um segundo computador foi utilizado para detectar os instantes dos pulsos de estímulo e resposta e armazenar estas sequências em arquivos. Posteriormente utilizamos estas sequências para calcular a informação mútua média entre os sinais e verificamos que diferentes peixes reconhecem e reagem (alterando seus disparos elétricos) a determinados trechos da série de estímulo real de maneira bastante reprodutível. Também desenvolvemos outro programa de controle para detectar os pulsos do peixe em um dos aquários e estimular, em tempo real, o peixe de outro aquário e vice­versa. Assim, a única forma de interação entre os peixes é através dos pulsos elétricos e esta interação ocorre de modo bidirecional. Os dados destes experimentos também foram analisados utilizando o cálculo da informação mútua média entre os padrões dos dois peixes e encontramos evidências de que neste caso o fluxo de informação é maior que nos experimentos unidirecionais. Nosso aparato permitiu utilizar com sucesso a teoria da informação para estudar a dinâmica de disparo durante a interação elétrica entre peixes e possibilita diversos experimentos futuros em que pretendemos relacionar os padrões elétricos ao comportamento social dos animais e a sua interação com o meio ambiente. / We built an experimental apparatus to measure the electric organ discharge times from weakly electric fishes of the Gymnotus carapo species. Such fishes use these pulses to actively locate objects in water as well as in social communication. Our apparatus was designed to allow such measures in the absence of some external perturbations the fishes are sensitive to, such as mechanical vibrations, electric fields and changes in the laboratory luminosity. A set of eight electrods were installed in the corners of the experimental tank and allows to obtain the discharge times without need to restrain the movements of the fish. Actually, from the maximal amplitudes of the discharge in different elecrodes we can infer the position and movements of the fish and relate its electrical dynamics to its behavior. A computer program (C language) was written to use a digital to analog interface to reproduce the time series of a discharge pulse from a real fish (recorded previously) and this electrical signal was used to stimulate the animals. The artificial pulses were applied to an electrical dipole built to mimic the geometry of the electrical organ of a living fish. The intervals between discharges were chosen from sequences obtained from different distributions: random, sequencies from real living fishes, handled sequencies where we repeated some real patterns with random patterns in between, etc. The detection of the stimuli and response pulses were done in another computer with the software Dasylab and the discharge times sequencies were recorded in harddisk for further analysis. Both sequencies were used to compute the average mutual information between the signals and we verified that different fishes recognize and react (changing their pulse interval pattterns) to the same regions of the real stimuli sequence. We also developed another control program (C language) to detect the discharges of a fish in one tank and to stimulate, in real time, a fish in another tank with those pulses, and vice­versa, in a bidirectional way. In this way, the only interaction between the fishes is through their electric pulses. The data analysis also consisted in obtaining the average mutual information between the sequencies of the two fishes and we found evidences that the flow of information is higher than that found in unidirectional experiments. Our apparatus allowed us to succesfully apply information theory to study the dynamics of the discharge intervals when the fishes are interacting. In the future we intend to extensivelly use such experiments to relate the electrical patterns to social behavior and to the interaction of these fishes with their environment.

Animals must sense their surroundings to update their internal representations of the external environment, and exploratory behaviours such as sensory sampling are influenced by past experiences. This thesis investigates how voluntary sensory sampling activities undergo learning-dependent changes. Studies of freely behaving animals impose two major challenges: 1) the accuracy of biological measurements is compromised by movement-induced artifacts, and 2) large degrees of freedom in unrestrained behaviours confound well-controlled studies. Pulse-type weakly electric fish (WEF) are an ideal choice to study adaptive sensory sampling from unrestrained animals, since they generate readily observable and quantifiable sensory capture events expressed by discrete pulses of electric organ discharges (EODs). To study the voluntarily movements and sensory sampling while animals navigated in darkness, we developed three novel experimental techniques to track movements and detect sensory sampling from a freely behaving WEF: 1) an EOD detector to remotely and accurately measure the sensory sampling rate, 2) an electrical tracking method to track multiple WEF using their own EODs, and 3) visual tracking algorithm for robust body tracking through water under infrared illumination. These techniques were successfully applied to reveal novel sensory sampling behaviours in freely exploring Gymnotus sp. Cortical activity precedes self-initiated movements by several seconds in mammals; this observation has led into inquiries on the nature of volition. Here we demonstrate the sensory sampling enhancement also precedes self-initiated movement by a few seconds in Gymnotus sp. Next, we tested whether these animals can be trained to learn a location of food using electrically detectable landmarks and, if so, whether they can use their past experiences to optimize their sensory sampling. We found that animals revisited the missing food location with high spatial accuracy, and they intensified their sensory sampling near the expected food location by increasing the number of EOD pulses per unit distance travelled.

Sensorische Nervenzellen kodieren Informationen über die Umwelt mittels elektrischer Impulse, sogenannte Aktionspotentiale oder Spikes. Diese werden weitergeleitet zu postsynaptischen Neuronen im zentralen Nervensystem, welche unterschiedliche Auslesestrategien verwenden. Integratorzellen summieren alle ankommenden Aktionspotentiale auf, wodurch sie die Gesamtaktivität einer präsynaptischen Population messen. Koinzidenzdetektoren hingegen, werden nur durch das synchrone Feuern der zuführenden Neuronenpopulation aktiviert. Die grundlegende Frage dieser Dissertation lautet: Welche Information eines zeitabhängigen Signals kodieren die synchronen Spikes einer Neuronenpopulation im Vergleich zu der Summe all ihrer Aktionspotentiale? Hierbei verwenden wir die Theorie stochastischer Prozesse: wir berechnen Spektralmaße, die es ermöglichen Aussagen darüber zu treffen welche Frequenzkomponenten eines Signals vorwiegend transmittiert werden. Im Gegensatz zu früheren Studien, verstehen wir unter einem synchronen Ereignis nicht zwangsläufig, dass die gesamte Population simultan feuert, sondern, dass ein minimaler Anteil („Synchronizitätsschranke") gleichzeitig aktiv ist. Unsere Analyse zeigt, dass die synchrone Populationsaktivität als ein Bandpass-Informationsfilter agieren kann: die synchronen Spikes kodieren hauptsächlich schnelle Signalanteile. Damit stellt die Selektion simultaner Neuronenaktivität ein potentielles Mittel dar um gleichzeitig anwesende, konkurrierende Signale voneinander zu trennen. Dabei hängen die genauen Charakteristika der Informationsfilterung ausschlaggebend von der Synchronizitätsschwelle ab. Insbesondere zeigt sich, dass eine Symmetrie in der Schwelle vorliegt,die die Äquivalenz der Kodierungseigenschaften von synchronem Feuern und synchronem Schweigen offenlegt. Unsere analytischen Ergebnisse testen wir mittels numerischer Simulationen und vergleichen sie mit Experimenten am schwach elektrischen Fisch. / Populations of sensory neurons encode information about the environment into electrical pulses, so called action potentials or spikes. Neurons in the brain process these pulses further by using different readout strategies. Integrator cells sum up all incoming action potentials and are thus sensitive to the overall activity of a presynaptic population. Coincidence detectors, on the other hand, are activated by the synchronous firing of the afferent population. The main question of this thesis is: What information about a common time-dependent stimulus is encoded in the synchronous spikes of a neuronal population in comparison to the sum of all spikes? We approach this question within the framework of spectral analysis of stochastic processes, which allows to assess which frequency components of a signal are predominantly encoded. Here, in contrast to earlier studies, a synchronous event does not necessarily mean that all neurons of the population fire simultaneously, but that at least a prescribed fraction ('synchrony threshold') needs to be active within a small time interval. We derive analytical expressions of the correlation statistics which are compared to numerical simulations and experiments on weakly electric fish. We show that the information transmission of the synchronous output depends highly on the synchrony threshold. We uncover a symmetry in the synchrony threshold, unveiling the similarity in the encoding capability of the common firing and the common silence of a population. Our results demonstrate that the synchronous output can act as a band-pass filter of information, i.e. it extracts predominantly fast components of a stimulus. If signals in different frequency regimes are concurrently present, the selection of synchronous firing events can thus be a tool to separate these signals.

One of the most powerful and important applications that the brain accomplishes is solving the sensory "cocktail party problem:" to adaptively suppress extraneous signals in an environment. Theoretical studies suggest that the solution to the problem involves an adaptive filter, which learns to remove the redundant noise. However, neural learning is also in its infancy and there are still many questions about the stability and application of synaptic learning rules for neural computation. In this thesis, the implementation of an adaptive filter in the brain of a weakly electric fish, A. Leptorhynchus, was studied. It was found to require a cerebellar architecture that could supply independent frequency channels of delayed feedback and multiple burst learning rules that could shape this feedback. This unifies two ideas about the function of the cerebellum that were previously separate: the cerebellum as an adaptive filter and as a generator of precise temporal inputs.

One of the most powerful and important applications that the brain accomplishes is solving the sensory "cocktail party problem:" to adaptively suppress extraneous signals in an environment. Theoretical studies suggest that the solution to the problem involves an adaptive filter, which learns to remove the redundant noise. However, neural learning is also in its infancy and there are still many questions about the stability and application of synaptic learning rules for neural computation. In this thesis, the implementation of an adaptive filter in the brain of a weakly electric fish, A. Leptorhynchus, was studied. It was found to require a cerebellar architecture that could supply independent frequency channels of delayed feedback and multiple burst learning rules that could shape this feedback. This unifies two ideas about the function of the cerebellum that were previously separate: the cerebellum as an adaptive filter and as a generator of precise temporal inputs.

One of the most powerful and important applications that the brain accomplishes is solving the sensory "cocktail party problem:" to adaptively suppress extraneous signals in an environment. Theoretical studies suggest that the solution to the problem involves an adaptive filter, which learns to remove the redundant noise. However, neural learning is also in its infancy and there are still many questions about the stability and application of synaptic learning rules for neural computation. In this thesis, the implementation of an adaptive filter in the brain of a weakly electric fish, A. Leptorhynchus, was studied. It was found to require a cerebellar architecture that could supply independent frequency channels of delayed feedback and multiple burst learning rules that could shape this feedback. This unifies two ideas about the function of the cerebellum that were previously separate: the cerebellum as an adaptive filter and as a generator of precise temporal inputs.

One of the most powerful and important applications that the brain accomplishes is solving the sensory "cocktail party problem:" to adaptively suppress extraneous signals in an environment. Theoretical studies suggest that the solution to the problem involves an adaptive filter, which learns to remove the redundant noise. However, neural learning is also in its infancy and there are still many questions about the stability and application of synaptic learning rules for neural computation. In this thesis, the implementation of an adaptive filter in the brain of a weakly electric fish, A. Leptorhynchus, was studied. It was found to require a cerebellar architecture that could supply independent frequency channels of delayed feedback and multiple burst learning rules that could shape this feedback. This unifies two ideas about the function of the cerebellum that were previously separate: the cerebellum as an adaptive filter and as a generator of precise temporal inputs.

In dieser Arbeit wurden die Morpho-Physiologie und das Verhalten zweier Arten Afrikanischer schwach elektrischer Fische, Marcusenius victoriae und Petrocephalus degeni, im Labor und in einem ihrer natürlichen Habitate im Lwamunda Sumpf in Uganda untersucht. Die zwei Hauptziele dieser Arbeit waren (i) tageszeitabhängige Verhaltensrhythmen (Aktivität, Habitatnutzung) im Labor und im Freiland zu untersuchen und (ii) die Ausprägung und Plastizität der morpho-physiologischen Merkmale von P. degeni zu untersuchen, die ihnen erlauben bei natürlich vorkommender, geringer Sauerstoffverfügbarkeit (Hypoxie) zu überleben. Tageszeitabhängige Verhaltensrhythmen beider Arten wurden im Labor über 42 Stunden und im natürlichen Habitat dieser Fische für sechs Tage erfasst. In den Laborversuchen verbrachten beide Arten tagsüber annähernd 100% der Zeit in einem bereitgestellten Versteck und schwammen nachts heraus um aktiv ihre Umwelt zu erkunden. Im Habitat wurden die meisten Fische in strukturell komplexen Habitaten unter schwimmenden Pflanzen detektiert. Nachts schwammen die Fische aktiv in die offenen und ungeschützten Bereiche der Lagune, vermutlich um nach Futter zu suchen und zu interagieren. Die Begleitende in-situ Messung der Sauerstoffverfügbarkeit zeigte, dass beide Arten präsent und vermutlich sogar am aktivsten waren während Phasen extremer nächtlicher Hypoxie. Zur Untersuchung der respiratorischen Merkmale von P. degeni wurden Respirometrieversuche mit hypoxie-akklimatisierten Tieren durchgeführt, Hämoglobin- und Laktatkonzentration im Blut gemessen, und morphologische Parameter an den ersten beiden Kiemenbögen erfasst. Die Fische zeigten niedrige Sauerstoffverbrauchsraten, welche sie bis zu einem sehr niedrigem äußeren Sauerstoffpartialdruck aufrechterhielten. Zusätzlich zeigten sie hohe Hämoglobin- und Laktatkonzentrationen im Blut. Bis zu 75 Tage Normoxie-Akklimatisierung führte zu reduzierter Hämoglobinkonzentration und kürzeren Kiemenfilamenten. / In this thesis, I investigated the morpho-physiology and behavior of two species of African mormyrid weakly electric fish, Marcusenius victoriae and Petrocephalus degeni, in the laboratory and in one of their natural habitats, the Lwamunda Swamp in Uganda. The two main objectives of this work were to (i) observe behavioral rhythms and habitat use patterns of both species under natural and laboratory conditions, and (ii) assess expression and plasticity of morpho-physiological traits that might enable P. degeni to survive naturally occurring low oxygen conditions (hypoxia). Behavioral rhythms were recorded in the laboratory over 42 hours and in the habitat on six sampling days. In the laboratory, both species spent close to 100% of the time in their shelter during the day and actively explore their environment at night. In the swamp lagoon, fish were most often encountered in structurally complex habitats under floating vegetation and ventured into open and unsheltered areas of the lagoon at night, presumably to forage and interact. Concomitant in-situ oxygen measurements revealed that these fish were present, and presumably most active during periods of extreme nocturnal hypoxia in their swamp habitat. To investigate respiratory traits of swamp-dwelling P. degeni, I conducted respirometry experiments and measured blood lactate and hemoglobin and gill morphometrics on the first two gill arches. Fish showed low routine oxygen consumption rates, which they maintained until a very low ambient oxygen partial pressure was reached. Additionally, they had high concentrations of hemoglobin and lactate in their blood. Up to 75 days of normoxia exposure reduced blood hemoglobin and gill filament length.