diff --git a/protocol/chapters/discussion.tex b/protocol/chapters/discussion.tex
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+
+We detected 25766 chirps in this dataset which included 27 valid trials of recordings that lasted 6 hours each. The number of detected chirps is close to the currently largest reported dataset by \textcite{obotiWhyBrownGhost2022} which contains multiple different experiments. This was achieved by extending the chirp detection algorithm by \textcite{henningerTrackingActivityPatterns2020}. We added a dynamic search frequency and combined peaks of the envelope in the EOD amplitude, the envelope of the dynamic search frequency, and the instantaneous frequency for chirp detection. The chirps we detected on a dataset published by \textcite{raabElectrocommunicationSignalsIndicate2021} indicate that individuals that win the competition for a superior shelter chirp less than the losers. Moreover, in some of these pairings, chirps emitted by the loser are temporally correlated with the offset of an agonistic interaction. This indicates, that chirps might be used by losers to terminate chasing events.
+
+\subsection{Assessing detection performance}
+
+While our chirp detector detected many chirps, the performance of the algorithm was not quantified yet. We only used a small trial dataset to visually asses whether chirps are detected or not. Additionally, the EOD$f$ of the fish in the trial data set were rather far apart, making it easy to correctly assign chirps. To assess the performance of our analysis data set, we visually inspected each iteration of the algorithm (in 5 seconds snippets) for one whole recording. However, for future uses of this detector, it is imperative to quantify the detection performance and tune the parameters accordingly. This is especially important considering that the main innovation of this algorithm is not to detect but to correctly assign chirps. We predict that the performance in the correct assignment will most likely drop with decreasing difference frequencies. If this is the case we have to quantify this parameter as well. In conclusion, future work on this algorithm should include a synthetic data set that reflects the natural variability to quantify the detection performances of multiple chirp detection parameters.
+
+\subsection{The problem of normalization}
+
+Currently, a major flaw of the chirp detector is that the peak detection parameters are fixed for each feature, and decided upon by trial and error. We reached the most error-free performance with high thresholds. A side effect is that the great majority of the chirps are only detected on a single electrode. If chirps would be detectable on multiple electrodes, the smallest number of electrodes on which a chirp must be detected could be used as an additional detection parameter. But to do this, ideally, we would need to be able to use the same peak detection parameters across all electrodes and for all features. And to achieve this we should normalize our feature arrays not only across features but also across electrodes. The difficulty with normalization arises with the rolling windows and the changing electrodes. If there is no chirp in a certain window, normalizing across electrodes over just that window would scale up all noise so that the peak detector would find many peaks. Most of the time
+ these peaks do not occur at the same time and thus are not falsely detected as chirps, but sometimes, they are. The current method to fix this is to not normalize at all and hard-code peak detection thresholds. This solution is not flexible and might fail completely for new data sets without manually adjusting the parameters. The seemingly easy approach would be to save the data of all feature arrays for each iteration to disk and normalize across all data before running the chirp detector. But the fact that for each iteration we jump between electrodes makes this step complicated and impractical. Another approach could be, to determine 'chirp-less' windows by a single parameter before normalization. If e.g. there in no peak in the search frequency, we could just skip to the next window instead of running the whole detection routine. Normalization could then be performed only in windows that have a peak in search frequency. 
+
+\subsection{Why can the instantaneous frequency go down during a chirp?} \label{ref:insta}
+
+While computing the instantaneous frequency we often encountered troughs instead of peaks in frequency during chirps. We simply circumvented this issue by taking the absolute of this instantaneous frequency. This phenomenon is critical because chirps are defined by a positive frequency excursion and not by a decrease in frequency. This decrease was only found after we computed the band pass filter which is filtering \SI{5}{\hertz} around the baseline of the EOD$f$ of the individual fish. If we band pass-filtered in a way so that the chirp was still included in the signal, all peaks in the instantaneous frequency were positive. This indicates that the narrow band pass filter around the signal is causing these frequency drops. One explanation for this issue could be the reduced amplitude during a chirp. The band pass filter removes the high frequency components that a chirp introduces. If the amplitude reduction is high enough, the frequency information shifts to noise.  If this noise has only low frequency components the instantaneous frequency drops. If the chirp contrast is low and the amplitude of the baseline does not decrease strongly, the increase in frequency that is common during a chirp might be reflected in a peak. But if the contrast is high and the amplitude in the baseline breaks down this would reflect in a trough in the instantaneous frequency. Against this theory stands the fact that we observed the troughs in frequency in cases where the amplitude of the baseline did not break down strongly. 
+
+\vspace{\baselineskip}
+
+Another explanation are the properties of the band pass filter which can introduce a phase shift while the fish emits chirps. If the phase is shifted backward the frequency temporarily increases, and vice versa for forward phase shifts. We still do not fully understand this change in frequency yet. Further analysis should include plotting the transfer function of the filter since narrow pass bands could introduce anomalies. Additionally, we would like to simulate chirps with different parameters, such as chirp phase, height, width, and contrast to understand which parameters might result in a trough of the instantaneous frequency.
+
+\subsection{Losers chirped more than winners}
+
+The detected chirps in this experiment indicate that winners chirped less than losers in this competition experiment. We can hypothesize that chirps are used as a submissive signal conveying information about the physical condition \parencite{davies1978deep}, and therefore can settle the competition without fights that escalate. 
+This result shows similarities to rises, another signal variation used by \textit{Aperonotus leptorhynchus}. \textcite{raabElectrocommunicationSignalsIndicate2021} showed that the loser of a competition experiment emitted more rises than the winner of the competition. Furthermore, the outcome of the competition is influenced by the body size: A larger body size is a predictor for winning the competition for the shelter \parencite{raabElectrocommunicationSignalsIndicate2021}. The frequencies of the individuals, on the other hand, do not seem to play a role in the outcome of the competition experiment. This needs to be further analyzed because the EOD$f$ is sexually dimorphic \parencite{meyer1987hormone}, and we did not take the sex of the pairings into account for our analysis.  
+
+\subsection{Why chirps might only terminate chasing in some dyads}
+
+Our results indicated that chirps can terminate chasing, hence we computed the chirp rate to the chasing on- or offset and physical events. Only for the chasing offset we could find a connection for some pairings in the competition experiment in a way that the chirp increases right before the offset of the chasing event. \textcite{raabElectrocommunicationSignalsIndicate2021} showed that rises are used by the subordinate to motivate mutual assessment. In the few cases where we observed increasing chirp frequencies at the offset of a chasing event, chirps may have been used by the loser of the competition to signal submission. Still, the mean for all pairings did not show a correlation of chirps and the chasing offsets. This can be explained by not including factors like sex and size difference in our analysis. The sex of the fish has an important role for the communication behavior in the mating season, where, the female emits a big chirp to signalize the spawning \parencite{hagedornCourtSparkElectric1985a, henningerStatisticsNaturalCommunication2018}. The size difference is also important for the outcome of the competition experiment, so there may be a link between the increase of chirping behavior and the offset of the chasing events with the size difference. This should be dissected further in the analysis of the behavioral data set. 
+
+\subsection{Summary}
+
+In conclusion, we were able to build the first chirp detector that might be usable on large data sets including multiple and freely moving individuals. Chrips can be detected by combining features from the changes in amplitude, instantaneous frequency, and a dynamic search window that is above the fish's own EOD$f$. We tested this detector on a data set of a competition experiment. We found that more chirps are produced when the size difference between the individuals was small and that the subordinates sometimes drastically increase their chirp frequency before a chasing event ends. This is the first step towards analyzing chirp-correlated behaviors in complex laboratory or field recordings to gain a clearer picture of the potentially multiple meanings of chirps. These first observations could inspire hypotheses that can then be verified in controlled experiments. 
\ No newline at end of file
diff --git a/protocol/chapters/introduction.tex b/protocol/chapters/introduction.tex
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+Successful communication depends on the transfer of reliable information \parencite{endler1993some, searcy2010evolution, hebets2016systems}. In the animal kingdom, information is commonly conveyed by signals which are produced or displayed by one individual to be received by another \parencite{hebets2016systems, searcy2010evolution, seyfarth2010central}. The main purpose of these signals is the prediction of upcoming events and the behavior of other animals in order to make suitable decisions \parencite{endler1993some, seyfarth2010central}. Thereby, signal reliability is a necessity for communication to occur in the first place, as neither the sender would produce signals nor the receiver would respond to them if it were not beneficial for both \parencite{seyfarth2017origin}. Thus, signals conveying reliable information are evolutionary stable even if a sender's and receiver's relationship may be cooperative or competitive \parencite{seyfarth2017origin}. However, the success of communication does not only depend on the signal and its properties itself. Any signal possesses an inherent vagueness that limits its specificity in any situation \parencite{seyfarth2003signalers, seyfarth2017origin}. Therefore, a second necessity for successful communication is contextual information. The context limits the possible interpretations of the information conveyed by the signal, thus increasing its specificity \parencite{seyfarth2017origin}. Hence, both the signal properties and the contextual information frame the communicative event whose success is further determined by the underlying degree of reliability. As such, successful communication is one of the main factors for an animal's survival and reproduction which is why it is the active focus of a wide field of research today.
+
+\subsection{Communication across modalities}
+
+As diverse as the contexts of animal signaling, as diverse are the modalities of the signals themselves. They range from chemical, tactile, and acoustic to visual and even electric, whereby the majority is effectively multimodal \parencite{bradbury1998principles, seyfarth2010central}. One of the most famous examples is the waggle dance of the honeybee \textit{Apis mellifera}, performed to communicate the location of a potent food source to workers in the hive \parencite{von1965tanze, riley2005flight}. As the performance is mainly a visual display it likewise involves scent \parencite{thom2007scent} as well as sound and air flow \parencite{tsujiuchi2007dynamic}. The former is also used by many ant species to indicate a profitable foraging spot. Thereby, the colony is guided by pheromone trails whose compounds are released via specified glands of individual ants \parencite{sudd1959interaction, david2009trail}. Another type of dance besides the honeybee's can be found in the peacock spider \textit{Maratus volans}. However, instead of dancing their way to a promising food source, male peacock spiders perform to impress females to find a partner to mate with \parencite{girard2011multi}. Not only because of the complex motion patterns and ornament displays but likewise owing to the vibratory signals emitted by the males, the peacock spider dance probably marks one of the most impressing communication signals in the animal kingdom \parencite{girard2011multi}. Considering the above it appears that invertebrates in general exhibit an interesting repertoire of possibilities to communicate, as there are also cases of facial patterns signifying status and rival assessment in paper wasps \parencite{tibbetts2008visual} as well as vibrational signals transmitted through plants by various insects to convey sex and species information together with directional cues for the potential mate \parencite{virant2004vibrational}.
+
+\vspace{\baselineskip}
+
+Striking examples of communication are also found among vertebrates. In this group, one of the most prevalent ways of conveying information is active vocalization. For instance, various monkey species exhibit an impressing repertoire of different calls, whereby its type depends on the context of the situation \parencite{SCHLENKER2016894}. One prominent example are the predator-specific calls in vervet monkeys indicating an attack by either a leopard, snake, or eagle \parencite{seyfarth1980}. Moreover, many species are likewise capable of communicating via facial and bodily gestures, which has for example been found in squirrel monkeys \parencite{anderson2010flexibility}, and chimpanzees \parencite{hobaiter2011gestural}. However, not only our closest ancestors but songbirds also use their voices through inter-individual contact. Especially during courtship, males like to show off their song repertoire to convey their qualities to a potential female mate and to repel competitors \parencite{kroodsma1991function, byers2009female}. Whereas a large fraction of animal communication can hardly be overheard, there is an equally large part escaping from human perception. Rodents, for example, emit calls with frequencies way above \SI{20}{\kilo\hertz}: so-called ultrasonic vocalizations (USV) which exceed the human hearing threshold \parencite{wohr2013affective}. These USVs are situation-dependent and differ in frequency. In rats, for instance, three major call types are known: Aversive and appetitive calls emitted by juveniles and adults, and pup calls indicating social isolation \parencite{wohr2013affective, seffer2014pro}. But this is not the only strategy for transferring information in the non-perceivable realm of animal communication. Some groups developed sensory systems enabling them to act both as the sender and receiver of sensory information by using the very same mechanisms \parencite{jonesCommunicationSelfFriends2021}. Such systems are found in pretty distinct species, namely bats \parencite{simmons1979echolocation} and toothed whales \parencite{kamminga1988echolocation}, which rely on echolocation, and weakly-electric fish using electrolocation \parencite{heiligenberg1973electrolocation}. The peculiar thing about these systems is that the animals possessing them actively generate signals to navigate, communicate, and detect prey \parencite{jonesCommunicationSelfFriends2021}. These signals are reflected by objects or other animals and allow for the detection of their properties, distance, and direction. In echolocation, bats and toothed whales emit high-frequency (\SI{20}{\kilo\hertz}) tonal sounds which are produced in the larynx or nose and whose reflections are then gathered by parts of the acoustic system \parencite{schnitzler2003spatial, park2016ultrasonic}. Electrolocation in weakly-electric fish on the other hand is based on the discharge of their electric organ \parencite{heiligenberg1973electrolocation, meyer1987hormone}. They hunt and navigate by perceiving alterations of their electric field caused by other animals or objects \parencite{von1999active, heiligenberg1973electrolocation}. The ability to sense electric fields is not uncommon in aquatic species. Sharks and rays have likewise developed the ability to perceive astoundingly weak electric fields \parencite{kalmijn1971electric}. They use these inputs for prey detection and electrolocation as well, however, they do not produce an electric signal themselves \parencite{kalmijn1971electric, kalmijn1973electro}. The electric eel, for example, is one the few species that is capable of actively producing electricity \parencite{catania2014shocking}. In contrast to weakly-electric fish, however, they do not constantly surround themselves with an electric field, but rather emit a mixture of low- and high-voltage pulses \parencite{catania2014shocking, catania2015electric}. The low-voltage pulses are used for sensing the environment whereas high-voltage pulses serve as a predatory strike to incapacitate prey \parencite{brown1950electric, catania2014shocking, catania2015electric}. The latter can reach strikingly high voltages of up to 600 V \parencite{brown1950electric, catania2014shocking}.
+
+\subsection{Beyond human perception: Weakly electric fish}
+As stated above, weakly-electric fish sample their environment by surrounding themselves with an electric field produced by discharging their electric organ. Based on the waveform of the electric organ discharge (EOD), weakly-electric fish are classified into two distinct groups: Wave-type and pulse-type electric fish \parencite{bendaPhysicsElectrosensoryWorlds2020}. Whereas wave-type EODs are characterized by a continuous discharge and a periodic waveform, pulse-type EODs show brief pulses interrupted by intervals of varying length \parencite{bendaPhysicsElectrosensoryWorlds2020}. For the wave-type category, the gymnotiform brown ghost knifefish \textit{Apteronotus leptorhynchus} is one of the best and most extensively studied species \parencite{zupanc1993evoked, meyer1987hormone, malerAtlasBrainElectric1991, hupeEffectDifferenceFrequency2008, englerSpontaneousModulationsElectric2000a, hagedornCourtSparkElectric1985a, dunlapHormonalBodySize2002}. Its EOD has a quasi-sinusoidal waveform and typically lies in a frequency range of \SIrange{600}{1000}{\hertz} (Fig. \ref{fig:Electric fields}B) \parencite{zupanc1993evoked, zakonEODModulationsBrown2002a}. Moreover, \textit{A. leptorhynchus} shows a sexual dimorphism in EOD frequency (EOD$f$) with females having a lower EOD$f$ ($<$ \SI{750}{\hertz}) than male individuals ($>$ \SI{750}{\hertz}) \parencite{meyer1987hormone, zakonEODModulationsBrown2002a}. Communication between two individuals occurs first and foremost because of the physical properties of the electric field. When interacting, the electric fields of two fish superimpose and result in amplitude modulation (AM), also called beat (Fig. \ref{fig:Electric fields}A \& D) \parencite{zupanc1993evoked, walzNeuroethologyElectrocommunicationHow2013, benda2005spike, bendaPhysicsElectrosensoryWorlds2020}. That is, by interacting with another individual, one fish perceives its own EOD being modulated by another electric field. The resulting beat has a specific frequency which is given by the frequency difference of the two interacting fish \parencite{benda2005spike, walzNeuroethologyElectrocommunicationHow2013, bendaPhysicsElectrosensoryWorlds2020}. Because of the frequency dimorphism in \textit{A. leptorhynchus}, this allows for a passive exchange of information since the beat frequency concomitantly transfers the gender relation of the fish. Simply speaking, the greater the beat frequency, the more likely it is that the other fish is of the opposite gender. However, \textit{A. leptorhynchus} is likewise capable of producing active communication signals. One of the most extensively studied are chirps \parencite{zupanc1993evoked, zakonEODModulationsBrown2002a, englerSpontaneousModulationsElectric2000a, hupeElectrocommunicationSignalsFree2009, obotiWhyBrownGhost2022, dunlapDiversityStructureElectrocommunication2003}. Chirps are transient frequency modulations that, depending on the type, are characterized by a duration of ten to a few hundred milliseconds and an increase of EOD$f$ up to several hundred Hertz (Fig. \ref{fig:introplot}, red track) \parencite{bendaPhysicsElectrosensoryWorlds2020, zupanc1993evoked, engler2001differential, zakonEODModulationsBrown2002a}. They have been shown to play a role in courtship and the synchronization of spawning \parencite{hagedornCourtSparkElectric1985a, henningerStatisticsNaturalCommunication2018}, but are also associated with aggressive encounters \parencite{triefenbachChangesSignallingAgonistic2008b}, where they are suggested to serve as a submissive signal \parencite{walzNeuroethologyElectrocommunicationHow2013, henningerStatisticsNaturalCommunication2018}. However, because of the various contexts in which chirps are emitted, their function is still an ongoing debate \parencite{obotiWhyBrownGhost2022}.
+
+\begin{figure}[H]
+    \centering
+    \includegraphics[width=0.7\linewidth]{figures/Fishies_cropped.pdf}
+    \mycaption{Properties of the electric field in weakly-electric fish}{\textbf{A:} The fish surround themselves with a bipolar electric field. The superposition of electric fields alters their spatial properties. Thereby, the phase of the EOD influences the extend and direction of the electric field of each fish (red marker in EOD waveform). \textbf{B:} Quasi-sinusoidal waveform of the EOD of a single wave-type electric fish. \textbf{C:} Amplitude modulations (AM) of an individual fish recorded by external electrodes. The AM is caused by the movements of the fish relative to the recording electrodes. \textbf{D:} Amplitude modulation or beat caused by the superposition of two electric fields. The frequency of the beat is given by the difference in frequency between the individual fish (Figure from \cite{raab2022social}).}
+    \label{fig:Electric fields}
+\end{figure}
+
+\subsection{The problem of detection}
+For the exploration of the chirp function, two requirements have to be met. First, chirps need to be recorded to be analyzed in the first place. Secondly, the analysis depends on the ability to detect chirps in the recordings. Optimally, the method fulfilling these two requirements is suited for deployment in laboratory settings as well as in the field. Chirp recording is the least problematic part. As chirps alter the EOD$f$, they can readily be recorded by a pair of electrodes placed in the vicinity of the fish \parencite{bendaPhysicsElectrosensoryWorlds2020}. A common approach in laboratory settings is to simulate a conspecific with a pair of electrodes which should cause the recorded individual to chirp. These can then be recorded with a second pair of electrodes \parencite{zupanc1993evoked, hagedornCourtSparkElectric1985a, dunlapDiversityStructureElectrocommunication2003, englerSpontaneousModulationsElectric2000a}. However, the laboratory does not reflect the natural habitat of the animals since the behavioral context of chirps emitted in the wild is highly variable \parencite{henningerStatisticsNaturalCommunication2018}. Thus, there is a need of recording fish in their natural habitat while freely behaving and interacting with each other. This is why recently, research is more and more shifted to the field \parencite{henningerStatisticsNaturalCommunication2018, fugere2011electrical, zubizarreta2020seasonal}. With this increasing change in research approach, new methods that allow for the recording of multiple fish in the wild were required. One approach was developed by \textcite{henningerStatisticsNaturalCommunication2018} and successfully tested in Panama, where the authors simultaneously recorded multiple individuals of four species of weakly-electric fish. They used a grid-like array of at least 54 electrodes which was submerged at the cut bank side of a stream. Later, a refined version of the grid was applied for further recordings in Colombia \parencite{raab2022AdvancesNoninvasiveTracking}. As such, the recording problem has been solved for the laboratory as well as the natural setting. The detection of chirps, however, turned out to be the bigger problem. As part of successful detection, it is required to correctly identify the chirp in the EOD track as well as assign the individual chirp to the correct animal. This is particularly difficult in recordings of multiple fish because the detection of a chirp implies that the underlying EOD$f$ is tracked for every animal. One approach for the analysis of electric signals in weakly-electric fish are spectrograms (Fig. \ref{fig:introplot}). Since chirps are very fast changes in the EOD$f$, a sufficiently high resolution in the time domain is necessary to resolve them. But, an increase of the resolution in time is accompanied by a resolution decrease in the frequency domain, which renders the distinction of the EOD$f$s of the chirping fish impossible. Given the situation of two fish with a similar EOD$f$, and the individual with the lower frequency emitting a chirp, it is unfeasible to assign the chirp to the correct individual by only using the spectrogram. Thus, chirp detection is surely not a trivial problem. \textcite{raab2022AdvancesNoninvasiveTracking}, based on previous work by \textcite{henningerTrackingActivityPatterns2020}, developed an algorithm that solves one part of the issue. It is capable of extracting EOD tracks by using both the EOD$f$ in the spectrogram as well as the spatial distribution of EOD power across electrodes \parencite{raab2022AdvancesNoninvasiveTracking}. This way, it is possible to obtain the EOD track of individual fish in recordings with multiple fish in space and time (Fig. \ref{fig:introplot}). Only with this groundwork, it is achievable to detect chirps in the first place. 
+
+\vspace{\baselineskip}
+
+To overcome the remaining problem of chirp assignment, we refined previous work on this issue \parencite{henningerStatisticsNaturalCommunication2018}. In the following, we describe and test the first draft of a chirp detection algorithm capable of assigning chirps to individual fish in recordings with multiple animals. Our approach was to include a dynamic filter that searches for a frequency range between the individual EOD$f$'s free of interference. In a second step, we tested the algorithm with a data set obtained by \textcite{raabElectrocommunicationSignalsIndicate2021}. Thereby, individuals of the species \textit{Apteronotus leptorhynchus} were recorded while competing for a superior shelter in dyads. We were able to detect 25766 chirps in this data set alone and analyzed the context in which they were emitted. Thereby, we could not fully replicate suggestions for chirp function from the literature. However, some of our results indicate the association of chirps with a specific type of event during aggressive encounters.
+
+\begin{figure}[H]
+    \centering
+    \includegraphics[width=1\linewidth]{figures/introplot.pdf}
+
+    \mycaption{Spectrogram of signal containing the EODs of two fish.}{The line plots indicate the instantaneous frequency of the respective individual, which we obtained by filtering the signal around its frequency component. \textbf{A:} Fish 1 produces a chirp that is visible by the frequency excursion in the instantaneous frequency and spectrogram if the frequency resolution is sufficient (NFFT=133.3, 20\% overlap). \textbf{B:} If the frequency resolution is lower, fish can be distinguished in the spectrogram, but chirp detection and assignment are not reliable (NFFT=1333.3, 20\% overlap). }
+    \label{fig:introplot}
+\end{figure}
+
+\todo[inline, color=orange]{Bilder von Lepto und seiner EOD waveform?}
+\todo[inline, color=orange]{Messy in-text Quellen}
+\todo[inline, color=orange]{Messy references Quellen}
diff --git a/protocol/chapters/methods.tex b/protocol/chapters/methods.tex
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+\subsection{Chirp detection algorithm}
+
+We developed and tested the improved algorithm on the raw data from a competition experiment (more information in section \ref{Behaviour}, or in the Paper by \cite{raabElectrocommunicationSignalsIndicate2021}) with 15 electrodes arranged in a grid. To be able to analyze the communication signals of the fish, which are most prominently represented as changes in their EOD$f$, we used the frequency tracks that were already computed by \textcite{raab2022AdvancesNoninvasiveTracking}. A frequency track is an approximation of how the fundamental frequency of the EOD of a single fish evolves. But because they are estimated using a spectrogram, they lack the temporal information needed to resolve transient frequency changes of a chirp. For this reason, we use the raw data, which has the appropriate temporal resolution to resolve chirps. To extract the EOD of single individuals, the frequency tracks are used to build specific band pass filters for every fish. Because the baseline EOD$f$ of a single fish changes with temperature and communication signals, filtering had to be performed in time windows. Additionally, to obtain the best signal for a freely moving fish, we choose the electrode with the highest power for every single rolling window. For each time window, we extract the amplitude of the baseline, the instantaneous frequency, and the amplitude of the search frequency, which all change during the production of a chirp. Simultaneously detected peaks on all three features are classified as a chirp. All signal processing steps that the raw data in a single rolling window snippet goes through are summarized in figure \ref{fig:algorithm}. The following paragraphs describe the processing steps in the same order as they are organized in the code of the algorithm. 
+
+\subsubsection{Loading and preparing the data set}
+
+The parameters used by the algorithm are all adjustable using a \codeword{yaml} configuration file, which includes sections for the path to the data set as well as to the output directory. The program loads the raw data as well as the frequency tracks for every fish in the recording and generates time windows, in which it iterates across the raw data set. For each window in time, it iterates through the respective windows on the frequency tracks of all the fish in the recording. This approach is arguably less intuitive compared to iterating through the full time of the recording on the frequency track of a single fish and then skipping to the next individual. However, it reduces the number of times the more memory- and hence computationally demanding raw data set needs to be loaded. 
+
+\subsubsection{Rolling windows and their overlap}
+
+To reduce the edge effects caused by filtering, we overlapped the rolling windows (window duration of \SI{5}{\second}) by one second and discarded the first and last \SI{250}{\milli\second} of a single window. This resulted in a true overlap of \SI{0,5}{\milli\second} in which chirps might be detected twice. To resolve this issue, we grouped all chirps that occurred less than \SI{20}{\milli\second} apart from each other as a single chirp. 
+
+\subsubsection{Following a fish through space}
+
+The raw data set was recorded using an electrode grid. Hence, the EOD amplitudes of single fish varies between electrodes, because the amplitude decreases with the distance between an electrode and a moving fish. These changes in amplitude convey information on where the individual of interest is located in space. For optimal detection of chirps, we should use the strongest electrode for one fish, since it should have the highest signal-to-noise ratio. Additionally, if multiple fish are further apart, it is advantageous to use the electrode that is closest to one fish, but not the other, to increase the odds of correct sender assignment. For each iteration of the algorithm, we start off with a short (\SI{5}{\second}) snippets of the approximated frequency track and respective powers of the frequencies of an individual fish on the same temporal extent. To decide upon which raw data snippet for this fish across the pool of 15 electrodes the algorithm should load for optimal performance, we first had to determine, to which electrode the fish was the closest in the current window in time. To determine the best electrodes, we simply use the ones that had the highest power in the tracked frequencies, since power is proportional to the amplitude squared. In the current implementation, we repeat the following pipeline for the two electrodes with the highest power for the current fish of interest. 
+
+\subsubsection{Feature extraction}
+
+After determining the best electrodes, we load the raw data snipped for the respective time window. This results in a data set of the frequency track of a fish (Figure \ref{fig:algorithm}, A, red)  and the raw data (Figure \ref{fig:algorithm}, A, spectrogram), which the algorithm uses to extract more information. For each frequency track of an individual fish in each window, the raw signal \SI{5}{\hertz} is then band pass filtered with cut off frequencies above and below the baseline of the frequency track. This effectively provides an approximation of the recorded signal as if the current fish was the only one in the area. However fast frequency excursions, e.g. during chirps, are lost because they exceed the frequency limits of the cutoff frequencies. In other words, the peak of the chirp is filtered out by the band pass filter and that is why we see a trough in the amplitude. We use this to our advantage because the amplitude of the signal drops at theses points in time. To use the amplitudes of this signal, we extract the envelope using a low pass (cut off at \SI{25}{\hertz}) filter multiplied by the square-root of two (Figure \ref{fig:algorithm}, B, red). In addition to the amplitude trough, a chirp should also change the frequency of the signal, even if the actual peak gets lost due to filtering. To detect such transient frequency changes, we also compute the instantaneous frequency of the filtered baseline, which can be achieved by extracting the inverse of every single period using the zero crossings of the signal (Figure \ref{fig:algorithm}, B, orange).  But using just the envelope and the instantaneous frequency of the EOD baseline of the fish was not sufficient to determine whether the anomaly we detected is a chirp or is caused e.g. by the movement of the fish. To deal with this issue, we used the notion that chirps are always up-modulations of the frequency that reach values of +\SI{50}{\hertz} to +\SI{300}{\hertz} above the baseline EOD$f$ of the emitting fish \parencite{zakonEODModulationsBrown2002a}. The first approach by \textcite{henningerStatisticsNaturalCommunication2018} was to filter a band at approximately +\SI{10}{\hertz} above the baseline and look for peaks in the power of this filtered band. We call this area the 'search frequency'. However, this method comes with the limitation, that if there are multiple fish and one of them has a baseline EOD$f$ that is coincidentally higher than the frequency of the current analyzed fish, the search frequency will become unusable. If the search frequency is close to- or at the baseline frequency of a fish with higher EOD$f$, chirps from the fish with a lower EOD$f$ would not be correctly assigned to the sender (Figure \ref{fig:henninger}). We adopted the method first documented in \textcite{henningerStatisticsNaturalCommunication2018} but introduce a dynamically adjusted search frequency. In our version of the chirp detection algorithm, the search frequency is still confined to a region of +\SI{20}{\hertz} to +\SI{100}{\hertz} above the baseline. In contrast to \textcite{henningerStatisticsNaturalCommunication2018} in this window, the frequency tracks of all other individuals are evaluated to find a region with the largest frequency difference from all other individuals that is still within the usual frequency peaks of chirps. The search frequency is then extracted in this window. This should make chirp detection and, most importantly, correct assignment to the sender, possible with recordings that include multiple individuals (Figure \ref{fig:dynamic}). The search window chosen in the example in Figure \ref{fig:algorithm}, A is indicated by orange dashed lines on the spectrogram. The envelope of the search frequency is visualized by the orange line in panel B. 
+
+\begin{figure}[H]
+    \centering
+    \includegraphics[width=\linewidth]{figures/henninger.pdf}
+    \mycaption{Fixed search frequency}{Schematic sketch of a fixed search frequency used by \textcite{henningerStatisticsNaturalCommunication2018}. The search frequency is placed at a static \SI{10}{\hertz} above the baseline of the lower fish. \textbf{A:} If there is no upper fish with a higher EOD$f$ in the search frequency, a chirp detection can be implemented. The decrease in amplitude and the increase of amplitude in the search frequency can be used for the chirp detection algorithm. \textbf{B:}
+    In the case that there is another fish with an EOD$f$ in the search frequency, chirp detection is impaired. The EOD at the search frequency contains the chirp of the lower fish and the EOD of the upper fish, which is making the detection of the chirp unreliable.}
+    \label{fig:henninger}
+\end{figure}
+
+\begin{figure}[H]
+    \centering
+    \includegraphics[width=\linewidth]{figures/dynamic.pdf}
+    \mycaption{Dynamic search frequency}{Schematic sketch of a dynamic search frequency implemented in the new algorithm. The search frequency is dynamic in a range of \SIrange{20}{100}{\hertz} above the baseline of the lower fish. \textbf{A:} If there is no upper fish with a higher EOD$f$ in the search frequency, chirp detection can be implemented. The decrease in amplitude and the increase of amplitude in the search frequency can be used for the chirp detection algorithm. \textbf{B:}
+    In the case that there is another fish with an EOD$f$ in the search frequency, the search frequency changes dynamically to a space with the largest difference to the upper fish. The chirp assignment is now possible with a correct sender assignment.}
+    \label{fig:dynamic}
+\end{figure}
+
+\subsubsection{Feature processing}
+
+The features we extracted, particularly the filtered baseline EOD, are also subject to change e.g. when the fish moves. But changes due to movements happen on a larger time scale. To reduce the impact of this noise, we additionally band-pass filtered the baseline envelope with cutoff frequencies \SI{2}{\hertz} (low-cutoff) and \SI{100}{\hertz} (high-cutoff). Additionally, we invert the baseline envelope to turn troughs into peaks for detection purposes. For the instantaneous frequency, we took the absolute and shifted it by the median of the EOD$f$ ($|EODf_{inst} - med({EODf_{inst}}|$). The instantaneous frequency during a chirp was negative in some cases. This irregularity is discussed in section \ref{ref:insta}. 
+The search frequency required no additional processing. The processed features are visualized in the panel C of Figure \ref{fig:algorithm}. We then detected the peaks of all three features using a prominence threshold of 0.00005 for the baseline envelope, 0.000004 for the search frequency, and 2 for the instantaneous frequency.
+
+\subsubsection{Peak classification}
+
+Since all three features should coincide temporally with the peak in frequency during a chirp, the first criterion for a chirp was that peaks were detected on all three features simultaneously. More specifically, we chose \SI{20}{\milli\second} as a tolerance window where the peaks must co-occur. Additionally, since we repeated the algorithm on the two electrodes of the highest power for the current time window, we set a threshold of the number of electrodes on which the chirp must appear to be accepted as a chirp. In the current implementation, the chirp must be detected on just a single electrode. If a chirp is detected we compute the mean of the three time points on which the peaks were found and appended the resulting time stamp to the chirp times of the current fish.
+
+\begin{figure}[H]
+    \centering
+    \includegraphics[width=\linewidth]{figures/10.0_11245.5.pdf}
+    \mycaption{Core processing pipeline of the chirp detector.}{The left side shows how the data is modified while it passes the algorithm. The right side is color-coded with respect to the plots and indicates what kind of data is visualized and how it is processed. \textbf{A:} The spectrogram on the top is used to visualize the raw data. The red line on the spectrogram indicates the tracked frequency. \textbf{B:} The subplots show the three features that the algorithm extracts from the raw data using the frequency tracks of the individual fish. Red indicates the envelope of the baseline EOD$f$ for a single fish, which we obtain by filtering the raw signal around the tracked frequency. The envelope of the 'search frequency' we obtain by filtering a narrow band inside a dynamically adjusted window above the baseline EOD$f$ of the respective fish is indicated in orange. The search frequency band is also indicated by dashed lines in the same color on the spectrogram. The yellow line is the instantaneous frequency of the filtered baseline (red), which changes if the signal disappears out of the filtering window during a chirp. \textbf{C} illustrates how the three features appear after they are processed. The dots indicate the detected peaks. The dots on the spectrogram indicate the detected chirps after the peaks are sorted.}
+    \label{fig:algorithm}
+\end{figure}
+
+\subsection{Behavioral data used to test the algorithm } \label{Behaviour}
+
+We tested the algorithm on a data set published by \textcite{raabElectrocommunicationSignalsIndicate2021}. The dataset was recorded using 21 mature \textit{Apteronotus leptorhynchus} from a tropical fish supplier. 9 males and 12 females, all in non-breeding conditions, were used. The experiment was conceptualized to understand the role of rises, another communication signal, during the competition of two fish for a superior shelter. The experiments took place in a \SI{100}{\liter} tank with a superior shelter in the center surrounded by other, less optimal shelters. The bottom of the tank was equipped with 15 mono-polar electrodes with low-noise buffer headstages. A reference electrode was positioned in a corner of the tank. The electric signals were first amplified and then digitized at \SI{20}{\kilo\hertz} per channel. The movement of the fish was recorded by a video camera mounted above the aquarium. Behavior (chasing events and contacts) were annotated manually using the software package BORIS \parencite{https://doi.org/10.1111/2041-210X.12584}.
+
+\subsection{Data analysis}
+
+The chirp detection algorithm as well as our subsequent analysis were written in Python 3.10.9 using the packages numpy, scipy, matplotlib, \href{https://github.com/janscience/thunderfish}{thunderfish} and \href{https://github.com/janscience/audioio}{audioio} \parencite{2020SciPy-NMeth, Hunter:2007, harris2020array}. All scripts as well as the required package versions are publically available in a \href{https://whale.am28.uni-tuebingen.de/git/raab/GP2023_chirp_detection}{git repository} (\url{https://whale.am28.uni-tuebingen.de/git/raab/GP2023_chirp_detection}). The temporal relation between chirps and chasing events was analyzed using a cross-correlation analysis. We computed the temporal differences between agonistic events (chasing onset, -offset and contact) and the chirps up to $\pm$ \SI{90}{\second} before and after the event. We then convolved the temporal differences with a Gaussian kernel with a standard deviation of \SI{2}{\milli\second}. To generate a baseline to compare the convolution with we randomly shuffled the chirp intervals in 50 permutations and recomputed the convolution for each. The baseline distribution was then obtained by the median across the permutations with the 5th and 95th percentile respectively.
\ No newline at end of file
diff --git a/protocol/chapters/results.tex b/protocol/chapters/results.tex
new file mode 100644
index 0000000..8520774
--- /dev/null
+++ b/protocol/chapters/results.tex
@@ -0,0 +1,43 @@
+\subsection{Chirps during dyadic competitions}
+
+We used our algorithm to detect chirps in a dyadic competition experiment where two individuals competed for one superior shelter. In this exemplary recording almost all physical contacts, chasing events, and chirps happened during the dark phase of the experiment (Figure \ref{fig:timeline}, top rows in raster plot). We can estimate that the winner (yellow) did not chirp as much as the loser (red) of the competition. In this example, the winner emitted only 10 chirps whereas the loser produced close to 400 chirps(Figure \ref{fig:timeline}).
+\begin{figure}[H]
+    \centering
+    \includegraphics[width=\linewidth]{figures/timeline.pdf}
+    \mycaption{Overview of a single trial of the competition experiment.}{The raster plots indicate the occurrence of the type of event. Rasterplots of chirps are color-coded with respect to the emitter of the chirp. The line plots indicate the EOD$f$ of the two fish over time as it was tracked on a spectrogram. Red was the winner of the competition, and yellow was the loser. The shaded area indicates the phase where lights were turned off.}
+    \label{fig:timeline}
+\end{figure}
+
+Losers tended to chirp more than winners (Figure \ref{fig:winnerloser} A). This could be observed in 16 of the 22 valid trials (total n = 28) however this difference was not significant (Wilcoxon-test: z-statistic=67.0, pvalue=0.054). The outcome of the competition experiment was influenced by the size of the individuals. The winner of the dyadic interaction tended to have a greater size than the losers (Figure \ref{fig:winnerloser} B). For small size differences, there was an increase in chirp count, indicating that chirps increased in relevance for competition as the size difference decreased. The frequencies of the emitted EOD for winner and loser did not show any relevance for the outcome of the competition because the distribution for winner and loser is evenly spread over the range of EOD$f$ (Figure \ref{fig:winnerloser} C). 
+
+\begin{figure}[H]
+    \centering
+    \includegraphics[width=\linewidth]{figures/chirps_winner_loser.pdf}
+    \mycaption{Competition outcome }{compared to the emitted chirps in 22 valid parings (total n=28). \textbf{A:} Winner and loser of the competition experiment displayed with their quantity of chirps. Connecting lines indicate the pairing in the experiment. \textbf{B:} Winner and Loser compared to their size difference [cm] and the number of chirps they emitted. Losers (orange) are usually smaller than winners ($\Delta$ size $<$ 0) \textbf{C}: EOD$f$s [Hz] of the winner and loser compared to their quantity of chirps.}
+    \label{fig:winnerloser}
+\end{figure}
+
+\subsection{Chirps emitted by loser fish might disrupt chasing events}
+
+\begin{wrapfigure}{R}{0.6\textwidth}
+\vspace{-1cm}
+  \begin{center}
+    \includegraphics[width=\linewidth]{figures/kde.pdf}
+    \end{center}
+    \mycaption{Chasing-triggered chirps}{for two selected dyads. The time axis is centered around the offset of a chasing event. The black line indicates the bootstrapped baseline. The gray area is the bootstrapped confidence interval. The red line indicates the chirp rate estimated by a convolution with a Gaussian kernel. \textbf{A:} In most cases, there was no change in the chirp rate around an offset of a chasing event. \textbf{B:} But in a subset of the dataset chirping increased just before the chasing stopped.}
+    \label{fig:kde}
+\end{wrapfigure}
+
+Losers tending to chirp more often than winners already hints towards a behavioral significance to chirps during competitions. We evaluated the temporal correlation of chirps and agnostic behaviors to further investigate the behavioral significance of chirps in dyadic encounters. For this, we computed the chasing-triggered chirp rate of the on- and offset of chasing events and contacts. The chasing-triggered chirp rate consists of chirps centered around agonistic events. After this sorting, we estimated the distribution of chirps for the events using a convolution with a Gaussian kernel. We focused on the offset of chasing events because the onset and physical contact did not show consistent increases in chirp rate. For approximately a third of all dyadic competitions, there was an increase in chirp rate before the offset occurred (Figure \ref{fig:kde} right). In the other cases, there was no interaction of the offset and chirps (Figure \ref{fig:kde} left). This was compared to a permutated baseline to show a significant increase in chirp rate. The summarized chirp rates across all competitions did not show an increase in chirp rate around the offsets of the chasing events.
+
+\newpage
+\FloatBarrier
+\begin{wrapfigure}[14]{hr}{0.4\textwidth}
+    \centering
+    \includegraphics[width=\linewidth]{figures/chirps_in_chasing.pdf}
+    \mycaption{Proportion of time spent and chirps emitted in chasing events.}{Relative time spent of the dark recording phase (3 h) in chasing events for single trials (\textbf{left}) and the relative quantity of chirps that were emitted during a chasing event for each recording (\textbf{right}). The lines show that the proportion of chirps during chasing events was only elevated for a subset of the competitions.}
+    \label{fig:chasing}
+\end{wrapfigure}
+\FloatBarrier
+
+To summarize this, we compared the percentage of time spent in chasing events with the percentage of chirps emitted in chasing events. This comparison should indicate that when the two fish are in a chasing event, and the chirps are used to carry information specifically related to competition, there should be an increase in the percentage of chirps in these chasing events. In 15 out of 27 pairings the percentage of time spent in chasing events was higher than the percentage of chirps emitted during chasing, showing that chirps may not be used for transferring information about their physical status in those cases. This indicates that chirps are not specifically used for the offset of chasing events because their percentage of occurrences is often lower than the time spent in these chasing events. 
diff --git a/protocol/chapters/titlepage.tex b/protocol/chapters/titlepage.tex
new file mode 100644
index 0000000..8208c95
--- /dev/null
+++ b/protocol/chapters/titlepage.tex
@@ -0,0 +1,42 @@
+\begin{titlepage}
+\begin{center}
+
+% Head block
+\vspace*{0.5cm}
+\Large
+Protokoll\\
+Neurobiologisches Großpraktikum\\
+\vspace{1cm}
+
+% Title and author
+\Huge
+\textbf{Detecting chirps in freely interacting weakly electric fish}\\
+\Large
+\vspace{1cm}
+\textbf{Sina Prause, Alexander Wendt and Patrick Weygoldt}
+\vspace{1cm}
+
+% Add a nice figure below the authors
+% \begin{figure}[ht]
+%     \centering
+%     %\includegraphics[scale=0.6]{images/drawing.pdf}
+%     \mycaption{A short bold}{ and a long regular caption.}
+% \end{figure}
+\vfill
+
+% Bottom block
+\large
+\vspace{0.5cm}
+Supervised by:\\
+\vspace{0.5cm}
+Till Raab \& Jan Benda\\
+Neuroethology\\
+Institute for Neurobiology\\
+Tuebingen University\\
+\vspace{1cm}
+\today \\
+\vspace{0.5cm}
+
+        
+\end{center}
+\end{titlepage}
diff --git a/protocol/chirpdetection.bib b/protocol/chirpdetection.bib
new file mode 100644
index 0000000..ce354d2
--- /dev/null
+++ b/protocol/chirpdetection.bib
@@ -0,0 +1,2001 @@
+@article{davies1978deep,
+  title={Deep croaks and fighting assessment in toads Bufo bufo},
+  author={Davies, Nick B and Halliday, Tim R},
+  journal={Nature},
+  volume={274},
+  pages={683--685},
+  year={1978},
+  publisher={Springer}
+}
+@article{raab2022AdvancesNoninvasiveTracking,
+	abstract = {Recent technological advances greatly improved the possibility to study freely behaving animals in natural conditions. However, many systems still rely on animal-mounted devices, which can already bias behavioral observations. Alternatively, animal behaviors can be detected and tracked in recordings of stationary sensors, e.g., video cameras. While these approaches circumvent the influence of animal-mounted devices, identification of individuals is much more challenging. We take advantage of the individual-specific electric fields electric fish generate by discharging their electric organ (EOD) to record and track their movement and communication behaviors without interfering with the animals themselves. EODs of complete groups of fish can be recorded with electrode arrays submerged in the water and then be tracked for individual fish. Here, we present an improved algorithm for tracking electric signals of wave-type electric fish. Our algorithm benefits from combining and refining previous approaches of tracking individual specific EOD frequencies and spatial electric field properties. In this process, the similarity of signal pairs in extended data windows determines their tracking order, making the algorithm more robust against detection losses and intersections. We quantify the performance of the algorithm and show its application for a data set recorded with an array of 64 electrodes distributed over a 12 m<sup>2</sup> section of a stream in the Llanos, Colombia, where we managed, for the first time, to track Apteronotus leptorhynchus over many days. These technological advances make electric fish a unique model system for a detailed analysis of social and communication behaviors, with strong implications for our research on sensory coding.},
+	author = {Raab, Till and Madhav, Manu S. and Jayakumar, Ravikrishnan P. and Henninger, J{\"o}rg and Cowan, Noah J. and Benda, Jan},
+	doi = {10.3389/fnint.2022.965211},
+	issn = {1662-5145},
+	journal = {Frontiers in Integrative Neuroscience},
+	title = {Advances in non-invasive tracking of wave-type electric fish in natural and laboratory settings},
+	url = {https://www.frontiersin.org/articles/10.3389/fnint.2022.965211},
+	volume = {16},
+	year = {2022},
+	bdsk-url-1 = {https://www.frontiersin.org/articles/10.3389/fnint.2022.965211},
+	bdsk-url-2 = {https://doi.org/10.3389/fnint.2022.965211} %das glaub ich nicht 
+	}
+
+
+@book{alcockAnimalBehavior2019,
+  title = {Animal Behavior},
+  author = {Alcock, John and Rubenstein, Dustin R.},
+  date = {2019},
+  edition = {Eleventh edition},
+  publisher = {{Oxford University Press}},
+  location = {{Sunderland, Massachusetts}},
+  isbn = {978-1-60535-548-1},
+  langid = {english},
+  pagetotal = {548},
+  keywords = {Animal behavior,Evolution},
+  file = {/home/weygoldt/Data/zotero/storage/EFDQJQWC/alcock_2019_animal behavior.pdf}
+}
+
+@article{arnegardSexualSignalEvolution2010,
+  title = {Sexual {{Signal Evolution Outpaces Ecological Divergence}} during {{Electric Fish Species Radiation}}},
+  author = {Arnegard, Matthew~E. and McIntyre, Peter~B. and Harmon, Luke~J. and Zelditch, Miriam~L. and Crampton, William~G.~R. and Davis, Justin~K. and Sullivan, John~P. and Lavoué, Sébastien and Hopkins, Carl~D.},
+  date = {2010-09},
+  journaltitle = {The American Naturalist},
+  shortjournal = {The American Naturalist},
+  volume = {176},
+  number = {3},
+  pages = {335--356},
+  issn = {0003-0147, 1537-5323},
+  doi = {10.1086/655221},
+  url = {https://www.journals.uchicago.edu/doi/10.1086/655221},
+  urldate = {2021-12-06},
+  abstract = {Natural selection arising from resource competition and environmental heterogeneity can drive adaptive radiation. Ecological opportunity facilitates this process, resulting in rapid divergence of ecological traits in many celebrated radiations. In other cases, sexual selection is thought to fuel divergence in mating signals ahead of ecological divergence. Comparing divergence rates between naturally and sexually selected traits can offer insights into processes underlying species radiations, but to date such comparisons have been largely qualitative. Here, we quantitatively compare divergence rates for four traits in African mormyrid fishes, which use an electrical communication system with few extrinsic constraints on divergence. We demonstrate rapid signal evolution in the Paramormyrops species flock compared to divergence in morphology, size, and trophic ecology. This disparity in the tempo of trait evolution suggests that sexual selection is an important early driver of species radiation in these mormyrids. We also found slight divergence in ecological traits among closely related species, consistent with a supporting role for natural selection in Paramormyrops diversification. Our results highlight the potential for sexual selection to drive explosive signal divergence when innovations in communication open new opportunities in signal space, suggesting that opportunity can catalyze species radiations through sexual selection, as well as natural selection.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/N94DR6WJ/Arnegard et al._2010_Sexual Signal Evolution Outpaces Ecological Diverg.pdf}
+}
+
+@article{bastianArginineVasotocinModulation,
+  title = {Arginine Vasotocin Modulation of Chirping},
+  author = {Bastian, J and Schniederjan, S and Nguyenkim, J},
+  pages = {16},
+  abstract = {Summary South American weakly electric fish produce a variety of electric organ discharge (EOD) amplitude and frequency modulations including chirps or rapid increases in EOD frequency that function as agonistic and courtship and mating displays. In Apteronotus leptorhynchus, chirps are readily evoked by the presence of the EOD of a conspecific or a sinusoidal signal designed to mimic another EOD, and we found that the frequency difference between the discharge of a given animal and that of an EOD mimic is important in determining which of two categories of chirp an animal will produce. Type-I chirps (EOD frequency increases averaging 650 Hz and lasting approximately 25 ms) are preferentially produced by males in response to EOD mimics with a frequency of 50–200 Hz higher or lower than that of their own. The EOD frequency of Apteronotus leptorhynchus is sexually dimorphic: female EODs range from 600 to 800 Hz and male EODs range from 800 to 1000 Hz. Hence, EOD frequency differences effective in evoking type-I chirps are most likely to occur during male/female interactions. This result supports previous observations that type-I chirps are emitted most often during courtship and mating. Type-II chirps, which consist of shorter-duration frequency increases of approximately 100 Hz, occur preferentially in response to EOD mimics that differ from the EOD of the animal by 10–15 Hz. Hence these are preferentially evoked when animals of the same sex interact and, as previously suggested, probably represent agonistic displays. Females typically produced only type-II chirps. We also investigated the effects of arginine vasotocin on chirping. This peptide is known to modulate communication and other types of behavior in many species, and we found that arginine vasotocin decreased the production of type-II chirps by males and also increased the production of type-I chirps in a subset of males. The chirping of most females was not significantly affected by arginine vasotocin.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/5TBQKKJP/Bastian et al. - Arginine vasotocin modulation of chirping.pdf}
+}
+
+@article{bastianElectrolocationPresenceJamming1987,
+  title = {Electrolocation in the Presence of Jamming Signals: Behavior},
+  shorttitle = {Electrolocation in the Presence of Jamming Signals},
+  author = {Bastian, Joseph},
+  date = {1987},
+  journaltitle = {Journal of Comparative Physiology A},
+  shortjournal = {J. Comp. Physiol.},
+  volume = {161},
+  number = {6},
+  pages = {811--824},
+  issn = {0340-7594, 1432-1351},
+  doi = {10.1007/BF00610223},
+  url = {http://link.springer.com/10.1007/BF00610223},
+  urldate = {2022-04-22},
+  abstract = {Electrolocation behavior of Apteronotus leptorhynchus was studied by monitoring the animal's ability to maintain orientation to a variety of moving electrolocation targets. The primary goal of this study was to determine the relative effectiveness of various types of electrical 'jamming signals' in disrupting electrolocation performance.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/ZU8HV2RA/Bastian - 1987 - Electrolocation in the presence of jamming signals.pdf}
+}
+
+@incollection{bendaPhysicsElectrosensoryWorlds2020,
+  title = {The {{Physics}} of {{Electrosensory Worlds}}},
+  booktitle = {The {{Senses}}: {{A Comprehensive Reference}}},
+  author = {Benda, Jan},
+  date = {2020},
+  pages = {228--254},
+  publisher = {{Elsevier}},
+  doi = {10.1016/B978-0-12-805408-6.00016-6},
+  url = {https://linkinghub.elsevier.com/retrieve/pii/B9780128054086000166},
+  urldate = {2021-11-29},
+  abstract = {Electric fish generate electric fields to infer properties of their environment and to communicate with each other. What does an electrosensory world look like? What are the basic physical laws governing electric image formation? This is discussed in four sections: temporal and spatial properties of electric fields generated by electric fish, electrolocation of small objects, electronavigation based on large nonconducting boundaries, and electrocommunication by frequency modulations of beats. Guided by the specific physics of these problems, behavioral, physical, and simulation findings are set into context. The dipole nature of the fish’s electric field and of polarized nearby objects inevitably limit electrolocation to the fish’s near field. Detection of capacitive objects as a different electric color is only possible if capacity, water conductivity and spectral properties of the fish’s electric field match. Water surface and large rocks as large nonconducting boundaries provide cues which are potentially detectable within a range of a meter and usable for electronavigation. Electrocommunication signals cover ranges up to 2 m and in wave fish are frequency modulations of beats on various timescales, ranging from tens of milliseconds to several minutes. We are just starting to understand their variety, their meaning and behavioral significance in courtship, breeding, aggression, group cohesion, etc. In addition to physical constraints, many selection pressures from predators and conspecifics act on electrosensory systems. We need more field data and need to explore the electric properties of ecological niches occupied by electric fish to really understand their adaptations and the costs of the various selection pressures.},
+  isbn = {978-0-12-805409-3},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/L64E9A6F/Benda_2020_The Physics of Electrosensory Worlds.pdf}
+}
+
+@article{bendaSynchronizationDesynchronizationCodeNatural2006,
+  title = {A {{Synchronization-Desynchronization Code}} for {{Natural Communication Signals}}},
+  author = {Benda, Jan and Longtin, André and Maler, Leonard},
+  date = {2006-10},
+  journaltitle = {Neuron},
+  shortjournal = {Neuron},
+  volume = {52},
+  number = {2},
+  pages = {347--358},
+  issn = {08966273},
+  doi = {10.1016/j.neuron.2006.08.008},
+  url = {https://linkinghub.elsevier.com/retrieve/pii/S0896627306006301},
+  urldate = {2022-04-21},
+  abstract = {Synchronous spiking of neural populations is hypothesized to play important computational roles in forming neural assemblies and solving the binding problem. Although the opposite phenomenon of desynchronization is well known from EEG studies, it is largely neglected on the neuronal level. We here provide an example of in vivo recordings from weakly electric fish demonstrating that, depending on the social context, different types of natural communication signals elicit transient desynchronization as well as synchronization of the electroreceptor population without changing the mean firing rate. We conclude that, in general, both positive and negative changes in the degree of synchrony can be the relevant signals for neural information processing.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/IZHUWCGN/Benda et al. - 2006 - A Synchronization-Desynchronization Code for Natur.pdf}
+}
+
+@article{carlsonBehavioralResponsesJamming2007,
+  title = {Behavioral Responses to Jamming and ‘Phantom’ Jamming Stimuli in the Weakly Electric Fish {{Eigenmannia}}},
+  author = {Carlson, Bruce A. and Kawasaki, Masashi},
+  date = {2007-08-24},
+  journaltitle = {Journal of Comparative Physiology A},
+  shortjournal = {J Comp Physiol A},
+  volume = {193},
+  number = {9},
+  pages = {927--941},
+  issn = {0340-7594, 1432-1351},
+  doi = {10.1007/s00359-007-0246-6},
+  url = {http://link.springer.com/10.1007/s00359-007-0246-6},
+  urldate = {2022-04-22},
+  abstract = {The jamming avoidance response (JAR) of the weakly electric fish Eigenmannia is characterized by upward or downward shifts in electric organ discharge (EOD) frequency that are elicited by particular combinations of sinusoidal amplitude modulation (AM) and differential phase modulation (DPM). However, non-jamming stimuli that consist of AM and/or DPM can elicit similar shifts in EOD frequency. We tested the hypothesis that these behavioral responses result from non-jamming stimuli being misperceived as jamming stimuli. Responses to nonjamming stimuli were similar to JARs as measured by modulation rate tuning, sensitivity, and temporal dynamics. There was a smooth transition between the magnitude of JARs and responses to stimuli with variable depths of AM or DPM, suggesting that frequency shifts in response to jamming and non-jamming stimuli represent different points along a continuum rather than categorically distinct behaviors. We also tested the hypothesis that non-jamming stimuli can elicit frequency shifts in natural contexts. Frequency decreases could be elicited by semi-natural AM stimuli, such as random AM, AM presented to a localized portion of the body surface, transient changes in amplitude, and movement of resistive objects through the electric field. We conclude that ‘phantom’ jamming stimuli can induce EOD frequency shifts in natural situations.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/375CPLIU/Carlson and Kawasaki - 2007 - Behavioral responses to jamming and ‘phantom’ jamm.pdf;/home/weygoldt/Data/zotero/storage/7YP9YJ7S/fnint-16-965211.pdf}
+}
+
+@article{carlsonElectricSignalingBehavior2002,
+  title = {Electric Signaling Behavior and the Mechanisms of Electric Organ Discharge Production in Mormyrid Fish},
+  author = {Carlson, Bruce A.},
+  date = {2002-09},
+  journaltitle = {Journal of Physiology-Paris},
+  shortjournal = {Journal of Physiology-Paris},
+  volume = {96},
+  number = {5-6},
+  pages = {405--419},
+  issn = {09284257},
+  doi = {10.1016/S0928-4257(03)00019-6},
+  url = {https://linkinghub.elsevier.com/retrieve/pii/S0928425703000196},
+  urldate = {2022-04-21},
+  abstract = {Mormyrid fish communicate and navigate using electric organ discharges (EODs). The EOD is highly stereotyped and provides information on sender identity, including species, sex, reproductive condition, and possibly relative status and individual identity. By contrast, the sequence of pulse intervals (SPI) is variable and plays more of a role in signaling behavioral states. Various types of SPI displays may be produced, including tonic patterns such as ‘random’ and ‘regularized’, and phasic patterns such as ‘bursts’ and ‘cessations’. Certain displays have been linked to specific behaviors such as aggression, submission, courtship and active exploration. In addition, interacting pairs of fish may produce stereotyped displays involving the relative timing of their EODs. The EOD waveform is controlled by the morphological and physiological properties of cells in the electric organ termed electrocytes. Differences in the innervation, morphology, size and membrane characteristics of electrocytes have been directly linked to species and sex differences in the EOD. The generation of each EOD is initiated in the medullary command nucleus (CN), which thereby determines the timing of EOD output. CN does not have any properties of a pacemaker, but rather appears to integrate descending inputs that affect the probability of EOD production. The precommand nucleus (PCN) provides a major source of excitatory input to CN and is itself inhibited by corollary discharge feedback following the production of each EOD. Changes in the activity of PCN and its inhibitory feedback neurons modify EOD output, and therefore drive the generation of SPI patterns. Current studies are addressing the mechanisms underlying the generation of these patterns and preliminary results suggest that different types of signals may be controlled by distinct components of the electromotor system. This is similar to findings in other electrogenic teleosts, suggesting that it may be a general feature in the motor control of signaling behavior.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/GK32AQ43/Carlson - 2002 - Electric signaling behavior and the mechanisms of .pdf}
+}
+
+@article{cramptonElectroreceptionElectrogenesisElectric2019,
+  title = {Electroreception, Electrogenesis and Electric Signal Evolution},
+  author = {Crampton, William G. R.},
+  date = {2019},
+  journaltitle = {Journal of Fish Biology},
+  volume = {95},
+  number = {1},
+  pages = {92--134},
+  issn = {1095-8649},
+  doi = {10.1111/jfb.13922},
+  url = {https://onlinelibrary.wiley.com/doi/abs/10.1111/jfb.13922},
+  urldate = {2021-12-01},
+  abstract = {Electroreception, the capacity to detect external underwater electric fields with specialised receptors, is a phylogenetically widespread sensory modality in fishes and amphibians. In passive electroreception, a capacity possessed by c. 16\% of fish species, an animal uses low-frequency-tuned ampullary electroreceptors to detect microvolt-range bioelectric fields from prey, without the need to generate its own electric field. In active electroreception (electrolocation), which occurs only in the teleost lineages Mormyroidea and Gymnotiformes, an animal senses its surroundings by generating a weak ({$<$} 1 V) electric-organ discharge (EOD) and detecting distortions in the EOD-associated field using high-frequency-tuned tuberous electroreceptors. Tuberous electroreceptors also detect the EODs of neighbouring fishes, facilitating electrocommunication. Several other groups of elasmobranchs and teleosts generate weak ({$<$} 10 V) or strong ({$>$} 50 V) EODs that facilitate communication or predation, but not electrolocation. Approximately 1.5\% of fish species possess electric organs. This review has two aims. First, to synthesise our knowledge of the functional biology and phylogenetic distribution of electroreception and electrogenesis in fishes, with a focus on freshwater taxa and with emphasis on the proximate (morphological, physiological and genetic) bases of EOD and electroreceptor diversity. Second, to describe the diversity, biogeography, ecology and electric signal diversity of the mormyroids and gymnotiforms and to explore the ultimate (evolutionary) bases of signal and receptor diversity in their convergent electrogenic–electrosensory systems. Four sets of potential drivers or moderators of signal diversity are discussed. First, selective forces of an abiotic (environmental) nature for optimal electrolocation and communication performance of the EOD. Second, selective forces of a biotic nature targeting the communication function of the EOD, including sexual selection, reproductive interference from syntopic heterospecifics and selection from eavesdropping predators. Third, non-adaptive drift and, finally, phylogenetic inertia, which may arise from stabilising selection for optimal signal-receptor matching.},
+  langid = {english},
+  keywords = {ampullary electroreceptor,electric-organ discharge,sensory ecology,tuberous electroreceptor},
+  annotation = {\_eprint: https://onlinelibrary.wiley.com/doi/pdf/10.1111/jfb.13922},
+  file = {/home/weygoldt/Data/zotero/storage/GYAIY79S/crampton_2019_electroreception, electrogenesis and electric signal evolution.pdf;/home/weygoldt/Data/zotero/storage/VWP4VJS5/jfb.html}
+}
+
+@article{desantanaUnexpectedSpeciesDiversity2019,
+  title = {Unexpected Species Diversity in Electric Eels with a Description of the Strongest Living Bioelectricity Generator},
+  author = {de Santana, C. David and Crampton, William G. R. and Dillman, Casey B. and Frederico, Renata G. and Sabaj, Mark H. and Covain, Raphaël and Ready, Jonathan and Zuanon, Jansen and de Oliveira, Renildo R. and Mendes-Júnior, Raimundo N. and Bastos, Douglas A. and Teixeira, Tulio F. and Mol, Jan and Ohara, Willian and e Castro, Natália Castro and Peixoto, Luiz A. and Nagamachi, Cleusa and Sousa, Leandro and Montag, Luciano F. A. and Ribeiro, Frank and Waddell, Joseph C. and Piorsky, Nivaldo M. and Vari, Richard P. and Wosiacki, Wolmar B.},
+  options = {useprefix=true},
+  date = {2019-09-10},
+  journaltitle = {Nature Communications},
+  shortjournal = {Nat Commun},
+  volume = {10},
+  number = {1},
+  pages = {4000},
+  publisher = {{Nature Publishing Group}},
+  issn = {2041-1723},
+  doi = {10.1038/s41467-019-11690-z},
+  url = {https://www.nature.com/articles/s41467-019-11690-z},
+  urldate = {2021-12-01},
+  abstract = {Is there only one electric eel species? For two and a half centuries since its description by Linnaeus, Electrophorus electricus has captivated humankind by its capacity to generate strong electric discharges. Despite the importance of Electrophorus in multiple fields of science, the possibility of additional species-level diversity in the genus, which could also reveal a hidden variety of substances and bioelectrogenic functions, has hitherto not been explored. Here, based on overwhelming patterns of genetic, morphological, and ecological data, we reject the hypothesis of a single species broadly distributed throughout Greater Amazonia. Our analyses readily identify three major lineages that diverged during the Miocene and Pliocene—two of which warrant recognition as new species. For one of the new species, we recorded a discharge of 860\,V, well above 650\,V previously cited for Electrophorus, making it the strongest living bioelectricity generator.},
+  issue = {1},
+  langid = {english},
+  keywords = {Biodiversity,Ichthyology,Taxonomy},
+  annotation = {Bandiera\_abtest: a Cc\_license\_type: cc\_by Cg\_type: Nature Research Journals Primary\_atype: Research Subject\_term: Biodiversity;Ichthyology;Taxonomy Subject\_term\_id: biodiversity;ichthyology;taxonomy},
+  file = {/home/weygoldt/Data/zotero/storage/5ATDSSBH/de santana_2019_unexpected species diversity in electric eels with a description of the strongest living bioelectricity generator.pdf;/home/weygoldt/Data/zotero/storage/QH2WZLB3/s41467-019-11690-z.html}
+}
+
+@book{dugatkinPrinciplesAnimalBehavior,
+  title = {Principles of {{Animal Behavior}} ({{Third Edition}})},
+  author = {Dugatkin, Lee Alan},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/NTKZXXJN/dugatkin_principles of animal behavior (third edition).pdf}
+}
+
+@article{dulkaTestosteroneModulatesFemale1994,
+  title = {Testosterone Modulates Female Chirping Behavior in the Weakly Electric Fish, {{Apteronotus}} Leptorhynchus},
+  author = {Dulka, J.G. and Maler, L.},
+  date = {1994-03},
+  journaltitle = {Journal of Comparative Physiology A},
+  shortjournal = {J Comp Physiol A},
+  volume = {174},
+  number = {3},
+  issn = {0340-7594, 1432-1351},
+  doi = {10.1007/BF00240215},
+  url = {http://link.springer.com/10.1007/BF00240215},
+  urldate = {2022-04-22},
+  abstract = {The weakly electric fish, Apteronotus leptorhynchus, produces a wave-like electric organ discharge (EOD) utilized for electrolocation and communication. Both sexes communicate by emitting "chirps": transient increases in EOD frequency. In males, chirping behavior and the jamming avoidance response (JAR) can be evoked by an artificial EOD stimulus delivered to the water at frequencies 1-10 Hz below the animal's own EOD. In contrast, females rarely chirp in response to this stimulus even though they show consistent JARs. To investigate whether this behavioral difference is hormone dependent, we implanted females with testosterone (T) and monitored their chirping activity over a 5 week period. Our findings indicate that elevations in blood levels of T cause an enhancement of chirping behavior and a lowering of basal EOD frequency in females. Elevated blood levels of T also appear to modulate the quality of chirps produced by hormone treated females. The effects of T on female chirping behavior and basal EOD frequency appear specific, since the magnitude of the JAR was not affected by the hormonal treatment. These findings suggest that seasonal changes in circulating concentrations of T may regulate behavioral changes in female chirping behavior and basal EOD frequency.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/Z9LXY765/Dulka and Maler - 1994 - Testosterone modulates female chirping behavior in.pdf}
+}
+
+@article{10.1242/jeb.038653,
+    author = {Dunlap, K. D. and DiBenedictis, B. T. and Banever, S. R.},
+    title = "{Chirping response of weakly electric knife fish (Apteronotus leptorhynchus) to low-frequency electric signals and to heterospecific electric fish}",
+    journal = {Journal of Experimental Biology},
+    volume = {213},
+    number = {13},
+    pages = {2234-2242},
+    year = {2010},
+    month = {07},
+    abstract = "{Brown ghost knife fish (Apteronotus leptorhynchus) can briefly increase their electric organ discharge (EOD) frequency to produce electrocommunication signals termed chirps. The chirp rate increases when fish are presented with conspecific fish or high-frequency (700–1100 Hz) electric signals that mimic conspecific fish. We examined whether A. leptorhynchus also chirps in response to artificial low-frequency electric signals and to heterospecific electric fish whose EOD contains low-frequency components. Fish chirped at rates above background when presented with low-frequency (10–300 Hz) sine-wave stimuli; at 30 and 150 Hz, the threshold amplitude for response was 1 mV cm–1. Low-frequency (30 Hz) stimuli also potentiated the chirp response to high-frequency (∼900 Hz) stimuli. Fish increased their chirp rate when presented with two heterospecific electric fish, Sternopygus macrurus and Brachyhypopomus gauderio, but did not respond to the presence of the non-electric fish Carassius auratus. Fish chirped to low-frequency (150 Hz) signals that mimic those of S. macrurus and to EOD playbacks of B. gauderio. The response to the B. gauderio playback was reduced when the low-frequency component (\\&lt;150 Hz) was experimentally filtered out. Thus, A. leptorhynchus appears to chirp specifically to the electric signals of heterospecific electric fish, and the low-frequency components of heterospecific EODs significantly influence chirp rate. These results raise the possibility that chirps function to communicate to conspecifics about the presence of a heterospecific fish or to communicate directly to heterospecific fish.}",
+    issn = {0022-0949},
+    doi = {10.1242/jeb.038653},
+    url = {https://doi.org/10.1242/jeb.038653},
+    eprint = {https://journals.biologists.com/jeb/article-pdf/213/13/2234/1425922/2234.pdf},
+}
+
+@article{dunlapDiversityStructureElectrocommunication2003,
+  title = {Diversity in the Structure of Electrocommunication Signals within a Genus of Electric Fish, {{Apteronotus}}},
+  author = {Dunlap, K. D. and Larkins-Ford, J.},
+  date = {2003-02},
+  journaltitle = {Journal of Comparative Physiology A},
+  shortjournal = {J Comp Physiol A},
+  volume = {189},
+  number = {2},
+  pages = {153--161},
+  issn = {0340-7594, 1432-1351},
+  doi = {10.1007/s00359-003-0393-3},
+  url = {http://link.springer.com/10.1007/s00359-003-0393-3},
+  urldate = {2022-04-22},
+  abstract = {Some gymnotiform electric fish modulate their electric organ discharge for intraspecific communication. In Apteronotus leptorhynchus, chirps are usually rapid (10–30 ms) modulations that are activated through nonN-methyl-D-aspartate (non-NMDA) glutamate receptors in the hindbrain pacemaker nucleus. Males produce longer chirp types than females and chirp at higher rates. In Apteronotus albifrons, chirp rate is sexually monomorphic, but chirp structure (change in frequency and amplitude during a chirp) was unknown. To better understand the neural regulation and evolution of chirping behavior, we compared chirp structure in these two species under identical stimulus regimes. A. albifrons, like A. leptorhynchus, produced distinct types of chirps that varied, in part, by frequency excursion. However, unlike in A. leptorhynchus, chirp types in A. albifrons varied little in duration, and chirps were all longer (70–200 ms) than those of A. leptorhynchus. Chirp type production was not sexually dimorphic in A. albifrons, but within two chirp types males produced longer chirps than females. We suggest that species differences in chirp duration might be attributable to differences in the relative proportions of fast-acting (non-NMDA) and slowacting (NMDA) glutamate receptors in the pacemaker. Additionally, we map species difference onto a phylogeny and hypothesize an evolutionary sequence for the diversification of chirp structure.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/UEEVAQ6Z/Dunlap and Larkins-Ford - 2003 - Diversity in the structure of electrocommunication.pdf}
+}
+
+@article{dunlapHormonalBodySize2002,
+  title = {Hormonal and {{Body Size Correlates}} of {{Electrocommunication Behavior}} during {{Dyadic Interactions}} in a {{Weakly Electric Fish}}, {{Apteronotus}} Leptorhynchus},
+  author = {Dunlap, Kent D},
+  date = {2002-03},
+  journaltitle = {Hormones and Behavior},
+  shortjournal = {Hormones and Behavior},
+  volume = {41},
+  number = {2},
+  pages = {187--194},
+  issn = {0018506X},
+  doi = {10.1006/hbeh.2001.1744},
+  url = {https://linkinghub.elsevier.com/retrieve/pii/S0018506X01917441},
+  urldate = {2022-04-22},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/V7DPBXS6/Dunlap - 2002 - Hormonal and Body Size Correlates of Electrocommun.pdf}
+}
+
+@article{dunlapTemperatureDependenceElectrocommunication2000,
+  title = {Temperature {{Dependence}} of {{Electrocommunication Signals}} and {{Their Underlying Neural Rhythms}} in the {{Weakly Electric Fish}}, {{{\emph{Apteronotus}}}}{\emph{ Leptorhynchus}}},
+  author = {Dunlap, K.D. and Smith, G.T. and Yekta, A.},
+  date = {2000},
+  journaltitle = {Brain, Behavior and Evolution},
+  shortjournal = {Brain Behav Evol},
+  volume = {55},
+  number = {3},
+  pages = {152--162},
+  issn = {1421-9743, 0006-8977},
+  doi = {10.1159/000006649},
+  url = {https://www.karger.com/Article/FullText/6649},
+  urldate = {2022-04-22},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/65GL9F82/Dunlap et al. - 2000 - Temperature Dependence of Electrocommunication Sig.pdf}
+}
+
+@incollection{dunlapWeaklyElectricFish2017,
+  title = {Weakly {{Electric Fish}}: {{Behavior}}, {{Neurobiology}}, and {{Neuroendocrinology}}},
+  shorttitle = {Weakly {{Electric Fish}}},
+  booktitle = {Hormones, {{Brain}} and {{Behavior}}},
+  author = {Dunlap, Kent D. and Silva, Ana C. and Smith, G. Troy and Zakon, Harold H.},
+  date = {2017},
+  pages = {69--98},
+  publisher = {{Elsevier}},
+  doi = {10.1016/B978-0-12-803592-4.00019-5},
+  url = {https://linkinghub.elsevier.com/retrieve/pii/B9780128035924000195},
+  urldate = {2021-11-30},
+  isbn = {978-0-12-803608-2},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/RGDVDH5U/Dunlap et al._2017_Weakly Electric Fish Behavior, Neurobiology, and .pdf}
+}
+
+@article{dyeDynamicsStimulusdependencePacemaker1987,
+  title = {Dynamics and Stimulus-Dependence of Pacemaker Control during Behavioral Modulations in the Weakly Electric Fish,{{Apteronotus}}},
+  author = {Dye, John},
+  date = {1987},
+  journaltitle = {Journal of Comparative Physiology A},
+  shortjournal = {J. Comp. Physiol.},
+  volume = {161},
+  number = {2},
+  pages = {175--185},
+  issn = {0340-7594, 1432-1351},
+  doi = {10.1007/BF00615239},
+  url = {http://link.springer.com/10.1007/BF00615239},
+  urldate = {2022-04-22},
+  abstract = {I. Weakly electric fish generate around their bodies low-amplitude, AC electric fields which are used both for the detection of objects and intraspecific communication. The types of modulation in this signal of which the high-frequency wave-type gymnotiform, Apteronotus, is capable are relatively few and stereotyped. Chief among these is the chirp, a signal used in courtship and agonistic displays. Chirps are brief and rapid accelerations in the normally highly regular electric organ discharge (EOD) frequency. 2. Chirping can be elicited artificially in these animals by the use of a stimulus regime identical to that typically used to elicit another behavior, the jamming avoidance response (JAR). The neuronal basis for the JAR, a much slower and lesser alteration in EOD frequency, is well understood. Examination of the stimulus features which induce chirping show that, like the JAR, there is a region of frequency differences between the fish's EOD and the interfering signal that maximally elicits the response. Moreover, the response is sex-specific with regard to the sign of the frequency difference, with females chirping preferentially on the positive and most males on the negative Df. These features imply that the sensory mechanisms involved in the triggering of these communicatory behaviors are fundamentally similar to those explicated for the JAR.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/DW3EUHU9/Dye - 1987 - Dynamics and stimulus-dependence of pacemaker cont.pdf}
+}
+
+@article{englerSpontaneousModulationsElectric2000a,
+  title = {Spontaneous Modulations of the Electric Organ Discharge in the Weakly Electric Fish, {{Apteronotus}} Leptorhynchus: A Biophysical and Behavioral Analysis},
+  shorttitle = {Spontaneous Modulations of the Electric Organ Discharge in the Weakly Electric Fish, {{Apteronotus}} Leptorhynchus},
+  author = {Engler, G. and Fogarty, C.M. and Banks, J.R. and Zupanc, G.K.H.},
+  date = {2000-08},
+  journaltitle = {Journal of Comparative Physiology A},
+  shortjournal = {J Comp Physiol A},
+  volume = {186},
+  number = {7-8},
+  pages = {645--660},
+  issn = {0340-7594, 1432-1351},
+  doi = {10.1007/s003590000118},
+  url = {http://link.springer.com/10.1007/s003590000118},
+  urldate = {2022-04-22},
+  abstract = {Brown ghosts, Apteronotus leptorhynchus, are weakly electric gymnotiform ®sh whose wave-like electric organ discharges are distinguished by their enormous degree of regularity. Despite this constancy, two major types of transient electric organ discharge modulations occur: gradual frequency rises, which are characterized by a relatively fast increase in electric organ discharge frequency and a slow return to baseline frequency; and chirps, brief and complex frequency and amplitude modulations. Although in spontaneously generated gradual frequency rises both duration and amount of the frequency increase are highly variable, no distinct subtypes appear to exist. This contrasts with spontaneously generated chirps which could be divided into four \`natural\` subtypes based on duration, amount of frequency increase and amplitude reduction, and time-course of the frequency change. Under non-evoked conditions, gradual frequency rises and chirps occur rather rarely. External stimulation with an electrical sine wave mimicking the electriceld of a neighboring ®sh leads to a dramatic increase in the rate of chirping not only during the 30 s of stimulation, but also in the period immediately following the stimulation. The rate of occurrence of gradual frequency rises is, however, unaected by such a stimulation regime.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/C5V9NLE2/Engler et al. - 2000 - Spontaneous modulations of the electric organ disc.pdf}
+}
+
+@article{fortuneSpookyInteractionDistance2020a,
+  title = {Spooky {{Interaction}} at a {{Distance}} in {{Cave}} and {{Surface Dwelling Electric Fishes}}},
+  author = {Fortune, Eric S. and Andanar, Nicole and Madhav, Manu and Jayakumar, Ravikrishnan P. and Cowan, Noah J. and Bichuette, Maria Elina and Soares, Daphne},
+  date = {2020-10-22},
+  journaltitle = {Frontiers in Integrative Neuroscience},
+  shortjournal = {Front. Integr. Neurosci.},
+  volume = {14},
+  pages = {561524},
+  issn = {1662-5145},
+  doi = {10.3389/fnint.2020.561524},
+  url = {https://www.frontiersin.org/articles/10.3389/fnint.2020.561524/full},
+  urldate = {2022-04-22},
+  abstract = {Glass knifefish (Eigenmannia) are a group of weakly electric fishes found throughout the Amazon basin. Their electric organ discharges (EODs) are energetically costly adaptations used in social communication and for localizing conspecifics and other objects including prey at night and in turbid water. Interestingly, a troglobitic population of blind cavefish Eigenmannia vicentespelea survives in complete darkness in a cave system in central Brazil. We examined the effects of troglobitic conditions, which includes a complete loss of visual cues and potentially reduced food sources, by comparing the behavior and movement of freely behaving cavefish to a nearby epigean (surface) population (Eigenmannia trilineata). We found that the strengths of electric discharges in cavefish were greater than in surface fish, which may result from increased reliance on electrosensory perception, larger size, and sufficient food resources. Surface fish were recorded while feeding at night and did not show evidence of territoriality, whereas cavefish appeared to maintain territories. Surprisingly, we routinely found both surface and cavefish with sustained differences in EOD frequencies that were below 10 Hz despite being within close proximity of about 50 cm. A half century of analysis of electrosocial interactions in laboratory tanks suggest that these small differences in EOD frequencies should have triggered the “jamming avoidance response,” a behavior in which fish change their EOD frequencies to increase the difference between individuals. Pairs of fish also showed significant interactions between EOD frequencies and relative movements at large distances, over 1.5 m, and at high differences in frequencies, often {$>$}50 Hz. These interactions are likely “envelope” responses in which fish alter their EOD frequency in relation to higher order features, specifically changes in the depth of modulation, of electrosocial signals.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/3CZYRFIN/Fortune et al. - 2020 - Spooky Interaction at a Distance in Cave and Surfa.pdf}
+}
+
+@article{fotowatStatisticsElectrosensoryInput2013,
+  title = {Statistics of the {{Electrosensory Input}} in the {{Freely Swimming Weakly Electric Fish Apteronotus}} Leptorhynchus},
+  author = {Fotowat, Haleh and Harrison, Reid R. and Krahe, Rüdiger},
+  date = {2013-08-21},
+  journaltitle = {Journal of Neuroscience},
+  shortjournal = {J. Neurosci.},
+  volume = {33},
+  number = {34},
+  eprint = {23966697},
+  eprinttype = {pmid},
+  pages = {13758--13772},
+  publisher = {{Society for Neuroscience}},
+  issn = {0270-6474, 1529-2401},
+  doi = {10.1523/JNEUROSCI.0998-13.2013},
+  url = {https://www.jneurosci.org/content/33/34/13758},
+  urldate = {2021-12-01},
+  abstract = {The neural computations underlying sensory-guided behaviors can best be understood in view of the sensory stimuli to be processed under natural conditions. This input is often actively shaped by the movements of the animal and its sensory receptors. Little is known about natural sensory scene statistics taking into account the concomitant movement of sensory receptors in freely moving animals. South American weakly electric fish use a self-generated quasi-sinusoidal electric field for electrolocation and electrocommunication. Thousands of cutaneous electroreceptors detect changes in the transdermal potential (TDP) as the fish interact with conspecifics and the environment. Despite substantial knowledge about the circuitry and physiology of the electrosensory system, the statistical properties of the electrosensory input evoked by natural swimming movements have never been measured directly. Using underwater wireless telemetry, we recorded the TDP of Apteronotus leptorhynchus as they swam freely by themselves and during interaction with a conspecific. Swimming movements caused low-frequency TDP amplitude modulations (AMs). Interacting with a conspecific caused additional AMs around the difference frequency of their electric fields, with the amplitude of the AMs (envelope) varying at low frequencies due to mutual movements. Both AMs and envelopes showed a power-law relationship with frequency, indicating spectral scale invariance. Combining a computational model of the electric field with video tracking of movements, we show that specific swimming patterns cause characteristic spatiotemporal sensory input correlations that contain information that may be used by the brain to guide behavior.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/Y7BB5U4Z/fotowat_2013_statistics of the electrosensory input in the freely swimming weakly electric fish apteronotus leptorhynchus.pdf;/home/weygoldt/Data/zotero/storage/RP5T26ZR/13758.html}
+}
+
+@article{fugereElectricalSignallingDominance2011,
+  title = {Electrical Signalling of Dominance in a Wild Population of Electric Fish},
+  author = {Fugère, Vincent and Ortega, Hernán and Krahe, Rüdiger},
+  date = {2011-04-23},
+  journaltitle = {Biology Letters},
+  shortjournal = {Biol. Lett.},
+  volume = {7},
+  number = {2},
+  pages = {197--200},
+  issn = {1744-9561, 1744-957X},
+  doi = {10.1098/rsbl.2010.0804},
+  url = {https://royalsocietypublishing.org/doi/10.1098/rsbl.2010.0804},
+  urldate = {2022-04-21},
+  abstract = {Animals often use signals to communicate their dominance status and avoid the costs of combat. We investigated whether the frequency of the electric organ discharge (EOD) of the weakly electric fish,               Sternarchorhynchus               sp., signals the dominance status of individuals. We correlated EOD frequency with body size and found a strong positive relationship. We then performed a competition experiment in which we found that higher frequency individuals were dominant over lower frequency ones. Finally, we conducted an electrical playback experiment and found that subjects more readily approached and attacked the stimulus electrodes when they played low-frequency signals than high-frequency ones. We propose that EOD frequency communicates dominance status in this gymnotiform species.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/I36IDJTE/Fugère et al. - 2011 - Electrical signalling of dominance in a wild popul.pdf}
+}
+
+@article{g.DifferentialProductionChirping2001,
+  title = {Differential Production of Chirping Behavior Evoked by Electrical Stimulation of the Weakly Electric Fish, {{Apteronotus}} Leptorhynchus},
+  author = {G., Engler and G., Zupanc},
+  date = {2001-11-01},
+  journaltitle = {Journal of Comparative Physiology A: Sensory, Neural, and Behavioral Physiology},
+  shortjournal = {Journal of Comparative Physiology A: Sensory, Neural, and Behavioral Physiology},
+  volume = {187},
+  number = {9},
+  pages = {747--756},
+  issn = {0340-7594, 1432-1351},
+  doi = {10.1007/s00359-001-0248-8},
+  url = {http://link.springer.com/10.1007/s00359-001-0248-8},
+  urldate = {2022-04-22},
+  abstract = {Apteronotus leptorhynchus (Gymnotiformes) produces wave-like electric organ discharges distinguished by a high degree of constancy. Transient frequency and amplitude modulations of these discharges occur both spontaneously and during social interactions, which can be mimicked by external electrical stimulation. The so-called chirps can be divided into four different types. Independent of the type of chirp produced under spontaneous conditions, the ®sh generate only signi®cant numbers of type-2 chirps under evoked conditions. The rate of production of chirps of this type is largely determined by the frequency relative to the ®sh's frequency and signal intensity. Frequencies of ‹10 Hz of the ®sh's own discharge frequency most e€ectively elicit chirps. Type-2 chirps can also be evoked through stimulation at or near the higher harmonic frequencies of the ®sh's frequency, but the chirp rate decreases with increasing number of the higher harmonic component. Over a certain range, the rate of production of type-2 chirps increases with increasing stimulus intensity. At very high intensities the generation of type-2 chirps is accompanied by the production of a novel type of electrical signal (\`abrupt frequency rise\`) characterized by a frequency increase of approximately 20 Hz and high repetition rates of roughly 10 s±1. We hypothesize that the di€erent types of electric modulations subserve di€erent behavioral functions.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/YUJTTKTQ/G. and G. - 2001 - Differential production of chirping behavior evoke.pdf}
+}
+
+@article{gavassaSignalModulationMechanism2012,
+  title = {Signal Modulation as a Mechanism for Handicap Disposal},
+  author = {Gavassa, Sat and Silva, Ana C and Gonzalez, Emmanuel and Stoddard, Philip K},
+  date = {2012-04-01},
+  journaltitle = {Animal behaviour},
+  shortjournal = {Anim Behav},
+  volume = {83},
+  number = {4},
+  eprint = {22665940},
+  eprinttype = {pmid},
+  pages = {935--944},
+  issn = {0003-3472},
+  doi = {10.1016/j.anbehav.2012.01.012},
+  url = {https://europepmc.org/articles/PMC3365606},
+  urldate = {2021-12-01},
+  abstract = {Signal honesty may be compromised when heightened competition provides incentive for signal exaggeration. Some degree of honesty might be maintained by intrinsic handicap costs on signalling or through imposition of extrinsic costs, such as social punishment of low quality cheaters. Thus, theory predicts a delicate balance between signal enhancement and signal reliability that varies with degree of social competition, handicap cost, and social cost. We investigated whether male sexual signals of the electric fish Brachyhypopomus gauderio would become less reliable predictors of body length when competition provides incentives for males to boost electric signal amplitude. As expected, social competition under natural field conditions and in controlled lab experiments drove males to enhance their signals. However, signal enhancement improved the reliability of the information conveyed by the signal, as revealed in the tightening of the relationship between signal amplitude and body length. Signal augmentation in male B. gauderio was independent of body length, and thus appeared not to be curtailed through punishment of low quality (small) individuals. Rather, all individuals boosted their signals under high competition, but those whose signals were farthest from the predicted value under low competition boosted signal amplitude the most. By elimination, intrinsic handicap cost of signal production, rather than extrinsic social cost, appears to be the basis for the unexpected reinforcement of electric signal honesty under social competition. Signal modulation may provide its greatest advantage to the signaller as a mechanism for handicap disposal under low competition rather than as a mechanism for exaggeration of quality under high competition.},
+  langid = {english},
+  pmcid = {PMC3365606},
+  file = {/home/weygoldt/Data/zotero/storage/F5SH3TI6/gavassa_2012_signal modulation as a mechanism for handicap disposal.pdf}
+}
+
+@article{gupteGuidePreProcessing2022,
+  title = {A Guide to Pre‐processing High‐throughput Animal Tracking Data},
+  author = {Gupte, Pratik Rajan and Beardsworth, Christine E. and Spiegel, Orr and Lourie, Emmanuel and Toledo, Sivan and Nathan, Ran and Bijleveld, Allert I.},
+  date = {2022-02},
+  journaltitle = {Journal of Animal Ecology},
+  shortjournal = {Journal of Animal Ecology},
+  volume = {91},
+  number = {2},
+  pages = {287--307},
+  issn = {0021-8790, 1365-2656},
+  doi = {10.1111/1365-2656.13610},
+  url = {https://onlinelibrary.wiley.com/doi/10.1111/1365-2656.13610},
+  urldate = {2022-07-03},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/6JKARH64/Gupte et al. - 2022 - A guide to pre‐processing high‐throughput animal t.pdf}
+}
+
+@article{hagedornCourtSparkElectric1985a,
+  title = {Court and Spark: Electric Signals in the Courtship and Mating of Gymnotoid Fish},
+  shorttitle = {Court and Spark},
+  author = {Hagedorn, Mary and Heiligenberg, Walter},
+  date = {1985-02},
+  journaltitle = {Animal Behaviour},
+  shortjournal = {Animal Behaviour},
+  volume = {33},
+  number = {1},
+  pages = {254--265},
+  issn = {00033472},
+  doi = {10.1016/S0003-3472(85)80139-1},
+  url = {https://linkinghub.elsevier.com/retrieve/pii/S0003347285801391},
+  urldate = {2022-04-22},
+  abstract = {By mimicking tropical rainy season conditions in aquaria, we stimulated two species of gymnotoid electric fish, Eigenmannia virescens and Apteronotus leptorhynchus, to spawn in captivity. Their courtship activity, breeding behaviour and electric social communication were monitored in several groups over 2 years. Groups of both species established dominance hierarchies correlated with electric organ discharge frequency, aggressiveness and size. Spawning was preceded by several nights of courtship during which the male modulated its electric organ discharge to produce 'chirps'. Continual bouts of chirping lasted for hours on evenings prior to spawning. These electrical signals play a significant role in courtship and spawning, as gravid E. virescens females could be stimulated to spawn by playing back into the tank a tape recording of male courtship chirps. In both species the chirp involves a slight increase in frequency followed by a cessation of the dominant frequency. This suggests a common mode of signal production in these two different genera offish. Chirps are short and abrupt during aggressive encounters, but assume a softer and more raspy quality during courtship.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/AXKBPGUV/Hagedorn and Heiligenberg - 1985 - Court and spark electric signals in the courtship.pdf}
+}
+
+@article{harvey-girardLongTermRecognition2010,
+  title = {Long Term Recognition Memory of Individual Conspecifics Is Associated with Telencephalic Expression of {{Egr-1}} in the Electric Fish, {{{\emph{Apteronotus}}}}{\emph{ Leptorhynchus}}},
+  author = {Harvey-Girard, Erik and Tweedle, Jessica and Ironstone, Joel and Cuddy, Martin and Ellis, William and Maler, Leonard},
+  date = {2010},
+  journaltitle = {The Journal of Comparative Neurology},
+  shortjournal = {J. Comp. Neurol.},
+  pages = {NA-NA},
+  issn = {00219967, 10969861},
+  doi = {10.1002/cne.22358},
+  url = {https://onlinelibrary.wiley.com/doi/10.1002/cne.22358},
+  urldate = {2022-04-21},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/3T8A925Z/Harvey-Girard et al. - 2010 - Long term recognition memory of individual conspec.pdf}
+}
+
+@article{heiligenbergJammingAvoidanceResponse1991,
+  title = {The Jamming Avoidance Response of the Electric Fish, {{Eigenmannia}}: Computational Rules and Their Neuronal Implementation},
+  shorttitle = {The Jamming Avoidance Response of the Electric Fish, {{Eigenmannia}}},
+  author = {Heiligenberg, Walter},
+  date = {1991-02},
+  journaltitle = {Seminars in Neuroscience},
+  shortjournal = {Seminars in Neuroscience},
+  volume = {3},
+  number = {1},
+  pages = {3--18},
+  issn = {10445765},
+  doi = {10.1016/1044-5765(91)90062-S},
+  url = {https://linkinghub.elsevier.com/retrieve/pii/104457659190062S},
+  urldate = {2022-04-22},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/YL7NZ2UB/Heiligenberg - 1991 - The jamming avoidance response of the electric fis.pdf}
+}
+
+@article{heiligenbergMotorControlJamming1996,
+  title = {Motor Control of the Jamming Avoidance Response of {{Apteronotus}} Leptorhynchus: Evolutionary Changes of a Behavior and Its Neuronal Substrates},
+  shorttitle = {Motor Control of the Jamming Avoidance Response of {{Apteronotus}} Leptorhynchus},
+  author = {Heiligenberg, W. and Wong, C.J.H. and Metzner, W. and Keller, C.H.},
+  date = {1996-11},
+  journaltitle = {Journal of Comparative Physiology A},
+  shortjournal = {J Comp Physiol A},
+  volume = {179},
+  number = {5},
+  pages = {653--674},
+  issn = {0340-7594, 1432-1351},
+  doi = {10.1007/BF00216130},
+  url = {http://link.springer.com/10.1007/BF00216130},
+  urldate = {2022-04-22},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/4C6J4G4K/Heiligenberg et al. - 1996 - Motor control of the jamming avoidance response of.pdf}
+}
+
+@book{heiligenbergPrinciplesElectrolocationJamming1977,
+  title = {Principles of {{Electrolocation}} and {{Jamming Avoidance}} in {{Electric Fish}}: A {{Neuroethological Approach}}},
+  shorttitle = {Principles of {{Electrolocation}} and {{Jamming Avoidance}} in {{Electric Fish}}},
+  author = {Heiligenberg, Walter},
+  date = {1977},
+  publisher = {{Springer Berlin Heidelberg}},
+  location = {{Berlin, Heidelberg}},
+  url = {https://doi.org/10.1007/978-3-642-81161-6},
+  urldate = {2022-04-22},
+  abstract = {This booklet, together with the following two, -which are well under way and will succeed it at intervals of, we hope, no more than six months, sets the stage for a new editorial enterprise in the field of brain science. The accent is on the functional aspects of brains rather than on their developU?? ment, hence the title of the series. The central question being how neural activity is related to behavior, there will be, naturally, a wide scatter of subU?? jects, and Heiligenberg's monograph on electric fish may be considered typU?? ical of the expected standard deviation from the mean. Deviations in other directions may go as far as the sensory neuron, or brain theory, or aphasia, or farther. The next contributions planned for the series are: Precht, Neuronal Operations in the Vestibular System, and Movshon, Genes and Environment in the Development of the Visual Cortex. Our aim is to apU?? proach the central area by means of something like an evolving handbook of brain science. The individual monographs should describe promising and successful approaches, even in areas where the last word is far from being said. Besides originaI monographs and compounds of the author's own published papers, reviews are also we1come if they are more than the sum of the parts. The publisher promises speedy publication, and the editors will see that the manuscripts will be readable as well as interesting. T??bingen, Summer 1977 V.},
+  isbn = {978-3-642-81161-6 978-3-540-08367-2},
+  langid = {english},
+  annotation = {OCLC: 858930874},
+  file = {/home/weygoldt/Data/zotero/storage/XPFMH74B/Heiligenberg - 1977 - Principles of Electrolocation and Jamming Avoidanc.pdf}
+}
+
+@article{henningerStatisticsNaturalCommunication2018,
+  title = {Statistics of {{Natural Communication Signals Observed}} in the {{Wild Identify Important Yet Neglected Stimulus Regimes}} in {{Weakly Electric Fish}}},
+  author = {Henninger, Jörg and Krahe, Rüdiger and Kirschbaum, Frank and Grewe, Jan and Benda, Jan},
+  date = {2018-06-13},
+  journaltitle = {The Journal of Neuroscience},
+  shortjournal = {J. Neurosci.},
+  volume = {38},
+  number = {24},
+  pages = {5456--5465},
+  issn = {0270-6474, 1529-2401},
+  doi = {10.1523/JNEUROSCI.0350-18.2018},
+  url = {https://www.jneurosci.org/lookup/doi/10.1523/JNEUROSCI.0350-18.2018},
+  urldate = {2021-11-30},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/UXNHBJ2B/Henninger et al._2018_Statistics of Natural Communication Signals Observ.pdf}
+}
+
+@article{henningerTrackingActivityPatterns2020,
+  title = {Tracking Activity Patterns of a Multispecies Community of Gymnotiform Weakly Electric Fish in Their Neotropical Habitat without Tagging},
+  author = {Henninger, Jörg and Krahe, Rüdiger and Sinz, Fabian and Benda, Jan},
+  date = {2020-01-01},
+  journaltitle = {Journal of Experimental Biology},
+  pages = {jeb.206342},
+  issn = {1477-9145, 0022-0949},
+  doi = {10.1242/jeb.206342},
+  url = {https://journals.biologists.com/jeb/article/doi/10.1242/jeb.206342/267458/Tracking-activity-patterns-of-a-multispecies},
+  urldate = {2022-04-22},
+  abstract = {Field studies on freely behaving animals commonly require tagging and often are focused on single species. Weakly electric fish generate a species- and individual-specific electric organ discharge (EOD) and therefore provide a unique opportunity for individual tracking without tagging. Here, we present and test tracking algorithms based on recordings with submerged electrode arrays. Harmonic structures extracted from power spectra provide fish identity. Localization of fish based on weighted averages of their EOD amplitudes is found to be more robust than fitting a dipole model. We apply these techniques to monitor a community of three species, Apteronotus rostratus, Eigenmannia humboldtii and Sternopygus dariensis, in their natural habitat in Darién, Panama. We found consistent upstream movements after sunset followed by downstream movements in the second half of the night. Extrapolations of these movements and estimates of fish density obtained from additional transect data suggest that some fish cover at least several hundreds of meters of the stream per night. Most fish, including E. humboldtii, were traversing the electrode array solitarily. From in situ measurements of the decay of the EOD amplitude with distance of individual animals, we estimated that fish can detect conspecifics at distances of up to 2 m. Our recordings also emphasize the complexity of natural electrosensory scenes resulting from the interactions of the EODs of different species. Electrode arrays thus provide an unprecedented window into the so-far hidden nocturnal activities of multispecies communities of weakly electric fish at an unmatched level of detail.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/NRM5SUFL/Henninger et al. - 2020 - Tracking activity patterns of a multispecies commu.pdf}
+}
+
+@article{hupeEffectDifferenceFrequency2008,
+  title = {The Effect of Difference Frequency on Electrocommunication: {{Chirp}} Production and Encoding in a Species of Weakly Electric Fish, {{Apteronotus}} Leptorhynchus},
+  shorttitle = {The Effect of Difference Frequency on Electrocommunication},
+  author = {Hupé, Ginette J. and Lewis, John E. and Benda, Jan},
+  date = {2008-07},
+  journaltitle = {Journal of Physiology-Paris},
+  shortjournal = {Journal of Physiology-Paris},
+  volume = {102},
+  number = {4-6},
+  pages = {164--172},
+  issn = {09284257},
+  doi = {10.1016/j.jphysparis.2008.10.013},
+  url = {https://linkinghub.elsevier.com/retrieve/pii/S0928425708000429},
+  urldate = {2022-04-21},
+  abstract = {The brown ghost knifefish, Apteronotus leptorhynchus, is a model wave-type gymnotiform used extensively in neuroethological studies. As all weakly electric fish, they produce an electric field (electric organ discharge, EOD) and can detect electric signals in their environments using electroreceptors. During social interactions, A. leptorhynchus produce communication signals by modulating the frequency and amplitude of their EOD. The Type 2 chirp, a transient increase in EOD frequency, is the most common modulation type. We will first present a description of A. leptorhynchus chirp production from a behavioural perspective, followed by a discussion of the mechanisms by which chirps are encoded by electroreceptor afferents (P-units). Both the production and encoding of chirps are influenced by the difference in EOD frequency between interacting fish, the so-called beat or difference frequency (Df). Chirps are produced most often when the Df is small, whereas attacks are more common when Dfs are large. Correlation analysis has shown that chirp production induces an echo response in interacting conspecifics and that chirps are produced when attack rates are low. Here we show that both of these relationships are strongest when Dfs are large. Electrophysiological recordings from electroreceptor afferents (P-units) have suggested that small, Type 2 chirps are encoded by increases in electroreceptor synchrony at low Dfs only. How Type 2 chirps are encoded at higher Dfs, where the signals seem to exert the greatest behavioural influence, was unknown. Here, we provide evidence that at higher Dfs, chirps could be encoded by a desynchronization of the P-unit population activity.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/MZ94KQXQ/Hupé et al. - 2008 - The effect of difference frequency on electrocommu.pdf}
+}
+
+@article{hupeElectrocommunicationSignalsFree2009,
+  title = {Electrocommunication Signals in Free Swimming Brown Ghost Knifefish, {{{\emph{Apteronotus}}}}{\emph{ Leptorhynchus}}},
+  author = {Hupé, Ginette J. and Lewis, John E.},
+  date = {2009-12-15},
+  journaltitle = {Journal of Experimental Biology},
+  volume = {212},
+  number = {24},
+  pages = {4101--4101},
+  issn = {1477-9145, 0022-0949},
+  doi = {10.1242/jeb.039081},
+  url = {https://journals.biologists.com/jeb/article/212/24/4101/9642/Electrocommunication-signals-in-free-swimming},
+  urldate = {2022-04-22},
+  abstract = {Brown ghost knifefish, Apteronotus leptorhynchus, are a species of weakly electric fish that produce a continuous electric organ discharge (EOD) that is used in navigation, prey capture and communication. Stereotyped modulations of EOD frequency and amplitude are common in social situations and are thought to serve as communication signals. Of these modulations, the most commonly studied is the chirp. This study presents a quantitative analysis of chirp production in pairs of free-swimming, physically interacting male and female A. leptorhynchus. Under these conditions, we found that in addition to chirps, the fish commonly produce a second signal type, a type of frequency rise called abrupt frequency rises, AFRs. By quantifying the behaviours associated with signal production, we find that Type 2 chirps tend to be produced when the fish are apart, following periods of low aggression, whereas AFRs tend to be produced when the fish are aggressively attacking one another in close proximity. This study is the first to our knowledge that quantitatively describes both electrocommunication signalling and behavioural correlates on a subsecond time-scale in a wave-type weakly electric fish.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/KEGA444A/Hupé and Lewis - 2009 - Electrocommunication signals in free swimming brow.pdf}
+}
+
+@article{jonesCommunicationSelfFriends2021,
+  title = {Communication with Self, Friends and Foes in Active-Sensing Animals},
+  author = {Jones, Te K. and Allen, Kathryne M. and Moss, Cynthia F.},
+  date = {2021-11-15},
+  journaltitle = {Journal of Experimental Biology},
+  volume = {224},
+  number = {22},
+  pages = {jeb242637},
+  issn = {0022-0949, 1477-9145},
+  doi = {10.1242/jeb.242637},
+  url = {https://journals.biologists.com/jeb/article/224/22/jeb242637/273391/Communication-with-self-friends-and-foes-in-active},
+  urldate = {2022-04-22},
+  abstract = {Animals that rely on electrolocation and echolocation for navigation and prey detection benefit from sensory systems that can operate in the dark, allowing them to exploit sensory niches with few competitors. Active sensing has been characterized as a highly specialized form of communication, whereby an echolocating or electrolocating animal serves as both the sender and receiver of sensory information. This characterization inspires a framework to explore the functions of sensory channels that communicate information with the self and with others. Overlapping communication functions create challenges for signal privacy and fidelity by leaving active-sensing animals vulnerable to eavesdropping, jamming and masking. Here, we present an overview of active-sensing systems used by weakly electric fish, bats and odontocetes, and consider their susceptibility to heterospecific and conspecific jamming signals and eavesdropping. Susceptibility to interference from signals produced by both conspecifics and prey animals reduces the fidelity of electrolocation and echolocation for prey capture and foraging. Likewise, activesensing signals may be eavesdropped, increasing the risk of alerting prey to the threat of predation or the risk of predation to the sender, or drawing competition to productive foraging sites. The evolutionary success of electrolocating and echolocating animals suggests that they effectively counter the costs of active sensing through rich and diverse adaptive behaviors that allow them to mitigate the effects of competition for signal space and the exploitation of their signals.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/3RBB4EDE/Jones et al. - 2021 - Communication with self, friends and foes in activ.pdf}
+}
+
+@article{k.RetreatSiteSelection2002a,
+  title = {Retreat Site Selection and Social Organization in Captive Electric Fish, {{Apteronotus}} Leptorhynchus},
+  author = {K., Dunlap and L., Oliveri},
+  date = {2002-07-01},
+  journaltitle = {Journal of Comparative Physiology A: Sensory, Neural, and Behavioral Physiology},
+  shortjournal = {Journal of Comparative Physiology A: Sensory, Neural, and Behavioral Physiology},
+  volume = {188},
+  number = {6},
+  pages = {469--477},
+  issn = {0340-7594, 1432-1351},
+  doi = {10.1007/s00359-002-0319-5},
+  url = {http://link.springer.com/10.1007/s00359-002-0319-5},
+  urldate = {2022-04-22},
+  abstract = {Gymnotiform fish use their electric organ discharge for electrolocation and communication. They are active nocturnally and seek retreat sites during the day. We examined retreat site selection in Apteronotus leptorhynchus by assessing their preference for retreat tubes that differed in opacity and dimension. Isolated fish preferred opaque to clear tubes, long and narrow diameter tubes to short, wide diameter tubes, and openended to closed tubes. We also assessed how groups of fish distributed themselves in tubes according to sex and electric organ discharge frequency under four conditions: (1) unlimited tube availability, (2) limited tube availability, (3) variation in tube opacity, and (4) variation in tube dimension. When tube availability was unlimited, fish generally preferred to occupy tubes alone. However, females, but not males, often cohabited tubes with consexuals. When tube availability was limited, females were more often than males found outside of tubes. When tubes varied by opacity and dimension, fish clustered most commonly in preferred tube types (opaque and long tubes). Males with the highest electric organ discharge frequencies usually occupied the most preferred tube type. Thus, fish have clear preferences in selecting retreat sites and groups of fish reveal their dominance relationships when presented with variation in retreat sites.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/HHN4Z9HR/K. and L. - 2002 - Retreat site selection and social organization in .pdf}
+}
+
+@book{kandelPrinciplesNeuralScience2012,
+  title = {Principles of {{Neural Science}}, {{Fifth Edition}}.},
+  author = {Kandel, Eric and Schwartz, James and Jessell, Thomas and Jessell, Department of Biochemistry {and} Molecular Biophysics Thomas and Siegelbaum, Steven and Hudspeth, A. J},
+  date = {2012},
+  publisher = {{McGraw-Hill Publishing}},
+  location = {{Blacklick}},
+  url = {https://public.ebookcentral.proquest.com/choice/publicfullrecord.aspx?p=4959346},
+  urldate = {2021-11-30},
+  abstract = {The field's definitive work from a Nobel Prize-winning author 900 full-color illustrations Principles of Neural Science, 5e describes our current understanding of how the nerves, brain, and mind function. From molecules to anatomic structures and systems to cognitive function, this comprehensive reference covers all aspects of neuroscience. Widely regarded as the field's cornerstone reference, the fifth edition is highlighted by more than 900 full-color illustrations. The fifth edition has been completely updated to reflect the tremendous amount of new research and development in neuroscien.},
+  isbn = {978-0-07-181001-2},
+  langid = {english},
+  annotation = {OCLC: 1027191624},
+  file = {/home/weygoldt/Data/zotero/storage/KW7MD25A/kandel_2012_principles of neural science, fifth edition.pdf}
+}
+
+@article{kawasakiIndependentlyEvolvedJamming,
+  title = {Independently Evolved Jamming Avoidance Responses Employ Identical Computational Algorithms: A Behavioral Study of the {{African}} Electric Fish, {{Gymnm}}'chusniloticus},
+  author = {Kawasaki, M},
+  pages = {14},
+  abstract = {An African electric fish, Gymnarchus, and a South American electric fish, Eigenmannia, are believed to have evolved their electrosensory systems independently. Both fishes, nevertheless, gradually shift the frequency of electric organ discharge away when they encounter a neighbor of a similar discharge frequency. Computational algorithms employed by Gymnarchusfor this jamming avoidance response have been identified in this study for comparison with those of extensively studied Eigenmannia.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/D3ABU8GJ/Kawasaki - Independently evolved jamming avoidance responses .pdf}
+}
+
+@article{kawasakiSensoryHyperacuityJamming1997,
+  title = {Sensory Hyperacuity in the Jamming Avoidance Response of Weakly Electric Fish},
+  author = {Kawasaki, Masashi},
+  date = {1997-08},
+  journaltitle = {Current Opinion in Neurobiology},
+  shortjournal = {Current Opinion in Neurobiology},
+  volume = {7},
+  number = {4},
+  pages = {473--479},
+  issn = {09594388},
+  doi = {10.1016/S0959-4388(97)80025-6},
+  url = {https://linkinghub.elsevier.com/retrieve/pii/S0959438897800256},
+  urldate = {2022-04-22},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/WJR2CI3N/Kawasaki - 1997 - Sensory hyperacuity in the jamming avoidance respo.pdf}
+}
+
+@article{kolodziejskiSIGNALPRODUCTIONFUNCTION,
+  title = {{{SIGNAL PRODUCTION AND FUNCTION IN WEAKLY ELECTRIC FISH}}: {{A COMPARATIVE INVESTIGATION OF SEXUALLY DIMORPHIC COMMUNICATION BEHAVIOR IN APTERONOTUS}}},
+  author = {Kolodziejski, Johanna A},
+  pages = {24},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/EE8FLWJW/Kolodziejski - SIGNAL PRODUCTION AND FUNCTION IN WEAKLY ELECTRIC .pdf}
+}
+
+@article{kramerSEXUALLYDIMORPHICJAMMING,
+  title = {{{THE SEXUALLY DIMORPHIC JAMMING AVOIDANCE RESPONSE IN THE ELECTRIC FISH EIGENMANNIA}} ({{TELEOSTEI}}, {{GYMNOTIFORMES}})},
+  author = {Kramer, Bernd},
+  pages = {24},
+  abstract = {Eigenmannia's jamming avoidance response (JAR) is a frequency change of its electric organ discharge (EOD) in response to an electric stimulus of similar frequency (small AF; AF = FFlgh — FStlm). It is assumed that the response to an undamped stimulus, AR = FRMponM. — F R M I , is stereotyped and non-habituating, and improves the fish's electrolocation performance in the presence of a jamming stimulus, such as the EOD of a nearby conspecific. Adult females gravid with eggs (N = 3) gave good responses (frequency decrease of at least 3 Hz) to —AF (stimulus frequency higher than fish frequency), but no response or only weak responses ({$<$}0-5 Hz) to + AF (stimulus frequency lower than fish frequency). After 2-75 years, a sexually mature female still showed the same behaviour, whereas an immature female (see below) had changed its behaviour considerably on becoming sexually mature.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/S59VXE38/Kramer - THE SEXUALLY DIMORPHIC JAMMING AVOIDANCE RESPONSE .pdf}
+}
+
+@article{kramerWaveformDiscriminationPhase1999,
+  title = {Waveform Discrimination, Phase Sensitivity and Jamming Avoidance in a Wave-Type Electric Fish},
+  author = {Kramer, B.},
+  date = {1999-05-15},
+  journaltitle = {Journal of Experimental Biology},
+  volume = {202},
+  number = {10},
+  pages = {1387--1398},
+  issn = {1477-9145, 0022-0949},
+  doi = {10.1242/jeb.202.10.1387},
+  url = {https://journals.biologists.com/jeb/article/202/10/1387/8187/Waveform-discrimination-phase-sensitivity-and},
+  urldate = {2022-04-22},
+  abstract = {The electric organ discharge (EOD) of most species of frequency-clamped and phase-locked to the EOD the freshwater knifefishes (Gymnotiformes) of South (frequency difference 0 Hz). Opening the electronic America is of the wave, not the pulse, type. Wave EODs are feedback loop immediately restored discrimination usually of constant frequency and amplitude, and show a performance on an on/off basis, and a strong jamming bewildering multitude of species-characteristic waveforms. avoidance response (JAR; a frequency shift away from the The EOD of Eigenmannia is sexually dimorphic in stimulus) accompanied every behavioural decision (to go waveform and in the intensity of its higher harmonics. In for a food reward). The strong habituation of the JAR that a go/no go paradigm, trained food-rewarded fish occurs in response to stimuli of no behavioural discriminated between these waveforms, and naive consequence (the usual test situation) was not seen in the (untrained) fish showed a significant preference. To present experiments. The proposed sensory model (which determine whether spectral or waveform (time) cues are is based on time-marking T electroreceptors) is supported used by the fish, artificial stimuli of identical amplitude by these experiments, and a biological function for the JAR spectrum were synthesized that differed only in phase – subserving EOD waveform discrimination is shown to be relationship between their harmonics, i.e. waveform, and useful in a social context.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/QVVYBTBK/Kramer - 1999 - Waveform discrimination, phase sensitivity and jam.pdf}
+}
+
+@article{malerAtlasBrainElectric1991,
+  title = {An Atlas of the Brain of the Electric Fish {{Apteronotus}} Leptorhynchus},
+  author = {Maler, L. and Sas, E. and Johnston, S. and Ellis, W.},
+  date = {1991-01},
+  journaltitle = {Journal of Chemical Neuroanatomy},
+  shortjournal = {Journal of Chemical Neuroanatomy},
+  volume = {4},
+  number = {1},
+  pages = {1--38},
+  issn = {08910618},
+  doi = {10.1016/0891-0618(91)90030-G},
+  url = {https://linkinghub.elsevier.com/retrieve/pii/089106189190030G},
+  urldate = {2022-04-22},
+  abstract = {This atlas consists of a set of six macrophotographs illustrating the important external landmarks of the apteronotid brain, as well as 54 transverse levels through the brain stained with cresyl violet. There are 150 lam between levels and the scales have 1mm divisions (100 lam small divisions). In general the neuroanatomy of this brain is similar to that of other teleosts except that all parts known to be concerned with electroreception are greatly hypertrophied (electrosensory lateral line lobe, nucleus praeminentialis, caudal lobe of the cerebellum, torus semicircularis dorsalis, optic tectum and nucleus electrosensorius). There are other regions of this brain which are hypertrophied or which have not been described in other teleosts, but which are not known to be directly linked to the electrosensory/ electromotor system; these regions are mentioned in the accompanying text.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/X5YXPTL5/Maler et al. - 1991 - An atlas of the brain of the electric fish Apteron.pdf}
+}
+
+@article{metznerGymnotiformJARCommunication,
+  title = {Gymnotiform {{JAR}} and Communication},
+  author = {Metzner, W},
+  pages = {11},
+  abstract = {Summary Over the past decade, research on the neural basis of communication and jamming avoidance in gymnotiform electric fish has concentrated on comparative studies of the premotor control of these behaviors, on the sensory processing of communication signals and on their control through the endocrine system, and tackled the question of the degree to which these behaviors share neural elements in the sensory–motor command chain by which they are controlled. From this wealth of investigations, we learned, first, how several segregated premotor pathways controlling a single central pattern generator, the medullary pacemaker nucleus, can provide a large repertoire of behaviorally relevant motor patterns. The results suggest that even small evolutionary modifications in the premotor circuitry can yield extensive changes in the behavioral output. Second, we have gained some insight into the concerted action of the brainstem, the diencephalon and the long-neglected forebrain in sensory processing and premotor control of communication behavior. Finally, these studies shed some light on the behavioral significance of multiple sensory brain maps in the electrosensory lateral line lobe that long have been a mystery. From these latter findings, it is tempting to interpret the information processing in the electrosensory system as a first step in the evolution towards the ‘distributed hierarchical’ organization commonly realized in sensory systems of higher vertebrates.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/UBSYM2JD/Metzner - Gymnotiform JAR and communication.pdf}
+}
+
+@article{middletonCellularBasisParallel2006,
+  title = {The Cellular Basis for Parallel Neural Transmission of a High-Frequency Stimulus and Its Low-Frequency Envelope},
+  author = {Middleton, Jason W. and Longtin, André and Benda, Jan and Maler, Leonard},
+  date = {2006-09-26},
+  journaltitle = {Proceedings of the National Academy of Sciences},
+  shortjournal = {Proc. Natl. Acad. Sci. U.S.A.},
+  volume = {103},
+  number = {39},
+  pages = {14596--14601},
+  issn = {0027-8424, 1091-6490},
+  doi = {10.1073/pnas.0604103103},
+  url = {https://pnas.org/doi/full/10.1073/pnas.0604103103},
+  urldate = {2022-04-21},
+  abstract = {Sensory stimuli often have rich temporal and spatial structure. One class of stimuli that are common to visual and auditory systems and, as we show, the electrosensory system are signals that contain power in a narrow range of temporal (or spatial) frequencies. Characteristic of this class of signals is a slower variation in their amplitude, otherwise known as an envelope. There is evidence suggesting that, in the visual cortex, both narrowband stimuli and their envelopes are coded for in separate and parallel streams. The implementation of this parallel transmission is not well understood at the cellular level. We have identified the cellular basis for the parallel transmission of signal and envelope in the electrosensory system: a two-cell network consisting of an interneuron connected to a pyramidal cell by means of a slow synapse. This circuit could, in principle, be implemented in the auditory or visual cortex by the previously identified biophysics of cortical interneurons.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/8D5PYXVQ/Middleton et al. - 2006 - The cellular basis for parallel neural transmissio.pdf}
+}
+
+@article{nelsonPreyCaptureBlack,
+  title = {Prey Capture in Black Ghost Knifefish},
+  author = {Nelson, M E and MacIver, M A},
+  pages = {9},
+  abstract = {Summary Sensory systems are faced with the task of extracting behaviorally relevant information from complex sensory environments. In general, sensory acquisition involves two aspects: the control of peripheral sensory surfaces to improve signal reception and the subsequent neural filtering of incoming sensory signals to extract and enhance signals of interest. The electrosensory system of weakly electric fish provides a good model system for studying both these aspects of sensory acquisition. On the basis of infrared video recordings of black ghost knifefish (Apteronotus albifrons) feeding on small prey (Daphnia magna) in the dark, we reconstruct three-dimensional movement trajectories of the fish and prey. We combine the reconstructed trajectory information with models of peripheral electric image formation and primary electrosensory afferent response dynamics to estimate the spatiotemporal patterns of transdermal potential change and afferent activation that occur during prey-capture behavior. We characterize the behavioral strategies used by the fish, with emphasis on the functional importance of the dorsal edge in prey capture behavior, and we analyze the electrosensory consequences. In particular, we find that the high-pass filter characteristics of P-type afferent response dynamics can serve as a predictive filter for estimating the future position of the prey as the electrosensory image moves across the receptor array.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/PSCRVCS3/Nelson and MacIver - Prey capture in black ghost knifefish.pdf}
+}
+
+@report{obotiWhyBrownGhost2022,
+  type = {preprint},
+  title = {Why the Brown Ghost Chirps at Night},
+  author = {Oboti, Livio and Pedraja, Federico and Ritter, Marie and Lohse, Marlena and Klette, Lennart and Krahe, Rüdiger},
+  date = {2022-12-29},
+  institution = {{Animal Behavior and Cognition}},
+  doi = {10.1101/2022.12.29.522225},
+  url = {http://biorxiv.org/lookup/doi/10.1101/2022.12.29.522225},
+  urldate = {2023-01-17},
+  abstract = {Since the pioneering work by Moeller, Szabo and Bullock, weakly electric fish have been widely used as a model to approach the study of both spatial and social cognitive abilities in a vertebrate taxon typically less approachable than mammals or other terrestrial vertebrates. Through their electric organ, weakly electric fish generate low-intensity electric fields around their bodies with which they scan the environment and manage to orient themselves and interact with conspecifics even in complete darkness. The brown ghost knifefish is probably the most studied species due to the large repertoire of individually variable and sex-specific electric signals it produces. Their rich electric vocabulary is composed of brief frequency modulations - or chirps - of the oscillating dipole moment constantly emitted by their electric organ. Because chirps come in different types, each carrying very specific and behaviorally salient information, they can be used as references to specific internal states during recordings - of either the brain or the electric organ - or during behavioral observations. Not surprisingly, this made the fortune of this model in neuroethology during the past 7 decades. Yet, truth is, this is not necessarily all true. Although established, to date this view has been supported only by correlative evidence and, to be fair, we still do not have a convincing framework to explain why these freshwater bottom dwellers emit electric chirps. Here we provide evidence for a previously unexplored role of chirps as specialized self-directed signals used to expand conspecific electrolocation ranges during social encounters.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/D5XL8XBF/Oboti et al. - 2022 - Why the brown ghost chirps at night.pdf}
+}
+
+@article{patricelliNewDimensionsAnimal2016,
+  title = {New Dimensions in Animal Communication: The Case for Complexity},
+  shorttitle = {New Dimensions in Animal Communication},
+  author = {Patricelli, Gail L and Hebets, Eileen A},
+  date = {2016-12},
+  journaltitle = {Current Opinion in Behavioral Sciences},
+  shortjournal = {Current Opinion in Behavioral Sciences},
+  volume = {12},
+  pages = {80--89},
+  issn = {23521546},
+  doi = {10.1016/j.cobeha.2016.09.011},
+  url = {https://linkinghub.elsevier.com/retrieve/pii/S2352154616301760},
+  urldate = {2022-04-22},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/A4HISNKA/Patricelli and Hebets - 2016 - New dimensions in animal communication the case f.pdf}
+}
+
+@article{pedrajaElectricImagingEvolution2014,
+  title = {Electric {{Imaging}} through {{Evolution}}, a {{Modeling Study}} of {{Commonalities}} and {{Differences}}},
+  author = {Pedraja, Federico and Aguilera, Pedro and Caputi, Angel A. and Budelli, Ruben},
+  editor = {Bethge, Matthias},
+  date = {2014-07-10},
+  journaltitle = {PLoS Computational Biology},
+  shortjournal = {PLoS Comput Biol},
+  volume = {10},
+  number = {7},
+  pages = {e1003722},
+  issn = {1553-7358},
+  doi = {10.1371/journal.pcbi.1003722},
+  url = {https://dx.plos.org/10.1371/journal.pcbi.1003722},
+  urldate = {2022-04-21},
+  abstract = {Modeling the electric field and images in electric fish contributes to a better understanding of the pre-receptor conditioning of electric images. Although the boundary element method has been very successful for calculating images and fields, complex electric organ discharges pose a challenge for active electroreception modeling. We have previously developed a direct method for calculating electric images which takes into account the structure and physiology of the electric organ as well as the geometry and resistivity of fish tissues. The present article reports a general application of our simulator for studying electric images in electric fish with heterogeneous, extended electric organs. We studied three species of Gymnotiformes, including both wave-type (Apteronotus albifrons) and pulse-type (Gymnotus obscurus and Gymnotus coropinae) fish, with electric organs of different complexity. The results are compared with the African (Gnathonemus petersii) and American (Gymnotus omarorum) electric fish studied previously. We address the following issues: 1) how to calculate equivalent source distributions based on experimental measurements, 2) how the complexity of the electric organ discharge determines the features of the electric field and 3) how the basal field determines the characteristics of electric images. Our findings allow us to generalize the hypothesis (previously posed for G. omarorum) in which the perioral region and the rest of the body play different sensory roles. While the ‘‘electrosensory fovea’’ appears suitable for exploring objects in detail, the rest of the body is likened to a ‘‘peripheral retina’’ for detecting the presence and movement of surrounding objects. We discuss the commonalities and differences between species. Compared to African species, American electric fish show a weaker field. This feature, derived from the complexity of distributed electric organs, may endow Gymnotiformes with the ability to emit site-specific signals to be detected in the short range by a conspecific and the possibility to evolve predator avoidance strategies.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/62N75HD8/Pedraja et al. - 2014 - Electric Imaging through Evolution, a Modeling Stu.pdf}
+}
+
+@article{petzoldCoadaptationElectricOrgan2016,
+  title = {Co-Adaptation of Electric Organ Discharges and Chirps in {{South American}} Ghost Knifefishes ({{Apteronotidae}})},
+  author = {Petzold, Jacquelyn M. and Marsat, Gary and Smith, G. Troy},
+  date = {2016-10},
+  journaltitle = {Journal of Physiology-Paris},
+  shortjournal = {Journal of Physiology-Paris},
+  volume = {110},
+  number = {3},
+  pages = {200--215},
+  issn = {09284257},
+  doi = {10.1016/j.jphysparis.2016.10.005},
+  url = {https://linkinghub.elsevier.com/retrieve/pii/S0928425716300109},
+  urldate = {2022-04-21},
+  abstract = {Animal communication signals that simultaneously share the same sensory channel are likely to coevolve to maximize the transmission of each signal component. Weakly electric fish continuously produce a weak electric field that functions in communication. Fish modulate the electric organ discharge (EOD) on short timescales to produce context-specific signals called chirps. EODs and chirps are simultaneously detected by electroreceptors and processed in the electrosensory system. We analyzed these signals, first to explore whether EOD waveform is encoded in the signal received by electroreceptors and then to examine how EODs and chirps interact to influence conspicuousness. Our findings show that gross discrimination of sinusoidal from complex EOD waveforms is feasible for all species, but fine discrimination of waveform may be possible only for species with waveforms of intermediate complexity. The degree of chirp frequency modulation and chirp relative decay strongly influenced chirp conspicuousness, but other chirp parameters were less influential. The frequency difference between the interacting EODs also strongly impacted chirp conspicuousness. Finally, we developed a method for creating hybrid chirp/EOD combinations to independently analyze the impact of chirp species, EOD species, and EOD difference frequency on chirp conspicuousness. All three components and their interactions strongly influenced chirp conspicuousness, which suggests that evolutionary changes in parameters of either chirps or EODs are likely to influence chirp detection. Examining other environmental factors such as noise created by fish movement and species-typical patterns of sociality may enrich our understanding of how interacting EODs affect the detection and discrimination of chirps across species.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/EB8TNV5R/Petzold et al. - 2016 - Co-adaptation of electric organ discharges and chi.pdf}
+}
+
+
+@article{raabDominanceHabitatPreference2019,
+  title = {Dominance in {{Habitat Preference}} and {{Diurnal Explorative Behavior}} of the {{Weakly Electric Fish Apteronotus}} Leptorhynchus},
+  author = {Raab, Till and Linhart, Laura and Wurm, Anna and Benda, Jan},
+  date = {2019-07-05},
+  journaltitle = {Frontiers in Integrative Neuroscience},
+  shortjournal = {Front. Integr. Neurosci.},
+  volume = {13},
+  pages = {21},
+  issn = {1662-5145},
+  doi = {10.3389/fnint.2019.00021},
+  url = {https://www.frontiersin.org/article/10.3389/fnint.2019.00021/full},
+  urldate = {2021-11-29},
+  abstract = {Electrocommunication and -localization behaviors of weakly electric fish have been studied extensively in the lab, mostly by means of short-term observations on constrained fish. Far less is known about their behaviors in more natural-like settings, where fish are less constrained in space and time. We tracked individual fish in a population of fourteen brown ghost knifefish (Apteronotus leptorhynchus) housed in a large 2 m3 indoor tank based on their electric organ discharges (EOD). The tank contained four different natural-like microhabitats (gravel, plants, isolated stones, stacked stones). In particular during the day individual fish showed preferences for specific habitats which provided appropriate shelter. Male fish with higher EOD frequencies spent more time in their preferred habitat during the day, moved more often between habitats during the night, and less often during the day in comparison to low-frequency males. Our data thus revealed a link between dominance indicated by higher EOD frequency, territoriality, and a more explorative personality in male A. leptorhynchus. In females, movement activity during both day and night correlated positively with EOD frequency. In the night, fish of either sex moved to another habitat after about 6 s on average. During the day, the average transition time was also very short at about 20 s. However, these activity phases were interrupted by phases of inactivity that lasted on average about 20 min during the day, but only 3 min in the night. The individual preference for daytime retreat sites did not reflect the frequent explorative movements at night.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/P4GTJH4A/Raab et al._2019_Dominance in Habitat Preference and Diurnal Explor.pdf}
+}
+
+@article{raabElectrocommunicationSignalsIndicate2021,
+  title = {Electrocommunication Signals Indicate Motivation to Compete during Dyadic Interactions of an Electric Fish},
+  author = {Raab, Till and Bayezit, Sercan and Erdle, Saskia and Benda, Jan},
+  date = {2021-10-01},
+  journaltitle = {Journal of Experimental Biology},
+  volume = {224},
+  number = {19},
+  pages = {jeb242905},
+  issn = {0022-0949, 1477-9145},
+  doi = {10.1242/jeb.242905},
+  url = {https://journals.biologists.com/jeb/article/224/19/jeb242905/272395/Electrocommunication-signals-indicate-motivation},
+  urldate = {2021-11-29},
+  abstract = {ABSTRACT             Animals across species compete for limited resources. Whereas in some species competition behavior is solely based on the individual's own abilities, other species assess their opponents to facilitate these interactions. Using cues and communication signals, contestants gather information about their opponent, adjust their behavior accordingly, and can thereby avoid high costs of escalating fights. We tracked electrocommunication signals known as ‘rises’ and agonistic behaviors of the gymnotiform electric fish Apteronotus leptorhynchus in staged competition experiments. A larger body size relative to the opponent was the sole significant predictor for winners. Sex and the frequency of the continuously emitted electric field only mildly influenced competition outcome. In males, correlations of body size and winning were stronger than in females and, especially when losing against females, communication and agonistic interactions were enhanced, suggesting that males are more motivated to compete. Fish that lost competitions emitted the majority of rises, but their quantity depended on the competitors’ relative size and sex. The emission of a rise could be costly since it provoked ritualized biting or chase behaviors by the other fish. Despite winners being accurately predictable based on the number of rises after the initial 25\hspace{0.25em}min, losers continued to emit rises. The number of rises emitted by losers and the duration of chase behaviors depended in similar ways on physical attributes of contestants. Detailed evaluation of these correlations suggests that A. leptorhynchus adjusts its competition behavior according to mutual assessment, where rises could signal a loser's motivation to continue assessment through ritualized fighting.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/8SDSSL8W/Raab et al_2021_Electrocommunication signals indicate motivation to compete during dyadic.pdf}
+}
+
+@article{roseInsightsNeuralMechanisms2004,
+  title = {Insights into Neural Mechanisms and Evolution of Behaviour from Electric Fish},
+  author = {Rose, Gary J.},
+  date = {2004-12},
+  journaltitle = {Nature Reviews Neuroscience},
+  shortjournal = {Nat Rev Neurosci},
+  volume = {5},
+  number = {12},
+  pages = {943--951},
+  issn = {1471-003X, 1471-0048},
+  doi = {10.1038/nrn1558},
+  url = {http://www.nature.com/articles/nrn1558},
+  urldate = {2021-12-06},
+  abstract = {Both behaviour and its neural control can be studied at two levels. At the proximate level, we aim to identify the neural circuits that control behaviour and to understand how information is represented and processed in these circuits. Ultimately, however, we are faced with questions of why particular neural solutions have arisen, and what factors govern the ways in which neural circuits are modified during the evolution of new behaviours. Only by integrating these levels of analysis can we fully understand the neural control of behaviour. Recent studies of electrosensory systems show how this synthesis can benefit from the use of tractable systems and comparative studies.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/WXE6F24X/Rose_2004_Insights into neural mechanisms and evolution of b.pdf}
+}
+
+@article{s.ElectrocommunicationSignalsFemale2002b,
+  title = {Electrocommunication Signals in Female Brown Ghost Electric Knifefish, {{Apteronotus}} Leptorhynchus},
+  author = {S., Tallarovic and H., Zakon},
+  date = {2002-09-01},
+  journaltitle = {Journal of Comparative Physiology A: Sensory, Neural, and Behavioral Physiology},
+  shortjournal = {Journal of Comparative Physiology A: Sensory, Neural, and Behavioral Physiology},
+  volume = {188},
+  number = {8},
+  pages = {649--657},
+  issn = {0340-7594, 1432-1351},
+  doi = {10.1007/s00359-002-0344-4},
+  url = {http://link.springer.com/10.1007/s00359-002-0344-4},
+  urldate = {2022-04-22},
+  abstract = {Female communication behaviors are often overlooked by researchers in favor of male behaviors, which are usually more overt and easier to elicit. Very little is known about female electrocommunication behaviors in brown ghost knifefish, a weakly electric wavetype Gymnotiform fish. Most behavioral studies have focused on males, and fish are usually restrained and played a stimulus near their own electric organ discharge frequency to evoke chirps (abrupt short-term frequency rises) or the jamming avoidance response. Our study focuses on categorizing and describing spontaneous and evoked electric organ discharge modulations in free-swimming female fish that were either electrically coupled to tanks containing a conspecific (male or female), or left isolated. Cluster analysis of signals produced under isolated and social conditions revealed three categories of rises: short rise, medium rise and long rise; and one category of frequency decrease (dip). Females produce significantly more short rises when electrically coupled to tanks containing lower-frequency females, and produce more long rises when electrically coupled to tanks containing males. Short rises may have an intrasexual aggressive function, while long rises may serve as an advertisement of status or reproductive condition in intersexual interactions.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/R3ZH5DJU/S. and H. - 2002 - Electrocommunication signals in female brown ghost.pdf}
+}
+
+@article{scheffelIntraandInterspecificElectrocommunication2006,
+  title = {Intra-and Interspecific Electrocommunication among Sympatric Mormyrids in the {{Upper Zambezi River}}},
+  author = {Scheffel, Andreas and Kramer, Bernd},
+  date = {2006},
+  publisher = {{Science Publishers}},
+  isbn = {1578083281},
+  file = {/home/weygoldt/Data/zotero/storage/87TV78YK/kramer4_.pdf}
+}
+
+@article{schumacherObjectDiscriminationActive2016,
+  title = {Object Discrimination through Active Electrolocation: {{Shape}} Recognition and the Influence of Electrical Noise},
+  shorttitle = {Object Discrimination through Active Electrolocation},
+  author = {Schumacher, Sarah and Burt de Perera, Theresa and von der Emde, Gerhard},
+  options = {useprefix=true},
+  date = {2016-10},
+  journaltitle = {Journal of Physiology-Paris},
+  shortjournal = {Journal of Physiology-Paris},
+  volume = {110},
+  number = {3},
+  pages = {151--163},
+  issn = {09284257},
+  doi = {10.1016/j.jphysparis.2016.11.008},
+  url = {https://linkinghub.elsevier.com/retrieve/pii/S0928425716300304},
+  urldate = {2022-04-22},
+  abstract = {The weakly electric fish Gnathonemus petersii can recognise objects using active electrolocation. Here, we tested two aspects of object recognition; first whether shape recognition might be influenced by movement of the fish, and second whether object discrimination is affected by the presence of electrical noise from conspecifics. (i) Unlike other object features, such as size or volume, no parameter within a single electrical image has been found that encodes object shape. We investigated whether shape recognition might be facilitated by movement-induced modulations (MIM) of the set of electrical images that are created as a fish swims past an object. Fish were trained to discriminate between pairs of objects that either created similar or dissimilar levels of MIM of the electrical images. As predicted, the fish were able to discriminate between objects up to a longer distance if there was a large difference in MIM between the objects than if there was a small difference. This supports an involvement of MIMs in shape recognition but the use of other cues cannot be excluded. (ii) Electrical noise might impair object recognition if the noise signals overlap with the EODs of an electrolocating fish. To avoid jamming, we predicted that fish might employ pulsing strategies to prevent overlaps. To investigate the influence of electrical noise on discrimination performance, two fish were tested either in the presence of a conspecific or of playback signals and the electric signals were recorded during the experiments. The fish were surprisingly immune to jamming by conspecifics: While the discrimination performance of one fish dropped to chance level when more than 22\% of its EODs overlapped with the noise signals, the performance of the other fish was not impaired even when all its EODs overlapped. Neither of the fish changed their pulsing behaviour, suggesting that they did not use any kind of jamming avoidance strategy.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/THIJF8WX/Schumacher et al. - 2016 - Object discrimination through active electrolocati.pdf}
+}
+
+@article{serrano-fernndezGradualFrequencyRises2003,
+  title = {Gradual Frequency Rises in Interacting Black Ghost Knifefish, {{Apteronotus}} Albifrons},
+  author = {Serrano-Fern�ndez, P.},
+  date = {2003-09-01},
+  journaltitle = {Journal of Comparative Physiology A: Sensory, Neural, and Behavioral Physiology},
+  shortjournal = {Journal of Comparative Physiology A: Sensory, Neural, and Behavioral Physiology},
+  volume = {189},
+  number = {9},
+  pages = {685--692},
+  issn = {0340-7594, 1432-1351},
+  doi = {10.1007/s00359-003-0445-8},
+  url = {http://link.springer.com/10.1007/s00359-003-0445-8},
+  urldate = {2022-04-22},
+  abstract = {The present paper highlights the relationship between social status and production of gradual frequency rises in interacting Apteronotus albifrons. The gradual frequency rise production was mathematically inferred and a discrete classification deliberately avoided. The results showed little gradual frequency rise production before the hierarchy settlement. Afterwards, only the dominant fish kept this gradual frequency rise production at low levels, while the subdominant fish drastically increased it in all following interaction contexts. The hypothesis of gradual frequency rises being involved in communication as submissive signals was thus strengthened.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/4F9ICCH3/Serrano-Fern�ndez - 2003 - Gradual frequency rises in interacting black ghost.pdf}
+}
+
+@article{shifmanComplexityHighfrequencyElectric2018,
+  title = {The Complexity of High-Frequency Electric Fields Degrades Electrosensory Inputs: Implications for the Jamming Avoidance Response in Weakly Electric Fish},
+  shorttitle = {The Complexity of High-Frequency Electric Fields Degrades Electrosensory Inputs},
+  author = {Shifman, Aaron R. and Lewis, John E.},
+  date = {2018-01},
+  journaltitle = {Journal of The Royal Society Interface},
+  shortjournal = {J. R. Soc. Interface.},
+  volume = {15},
+  number = {138},
+  pages = {20170633},
+  issn = {1742-5689, 1742-5662},
+  doi = {10.1098/rsif.2017.0633},
+  url = {https://royalsocietypublishing.org/doi/10.1098/rsif.2017.0633},
+  urldate = {2022-04-22},
+  abstract = {Sensory systems encode environmental information that is necessary for adaptive behavioural choices, and thus greatly influence the evolution of animal behaviour and the underlying neural circuits. Here, we evaluate how the quality of sensory information impacts the jamming avoidance response (JAR) in weakly electric fish. To sense their environment, these fish generate an oscillating electric field: the electric organ discharge (EOD). Nearby fish with similar EOD frequencies perform the JAR to increase the difference between their EOD frequencies, i.e. their difference frequency (DF). The fish determines the sign of the DF: when it has a lower frequency (DF {$>$} 0), EOD frequency is decreased and vice versa               .               We study the sensory basis of the JAR in two species:               Apteronotus leptorhynchus               have a high frequency (               ca               1000 Hz), spatio-temporally heterogeneous electric field, whereas               Eigenmannia               sp. have a low frequency (               ca               300 Hz), spatially uniform field. We show that the increased complexity of the               Apteronotus               field decreases the reliability of sensory cues used to determine the DF. Interestingly,               Apteronotus               responds to all JAR stimuli by increasing EOD frequency, having lost the neural pathway that produces JAR-related decreases in EOD frequency. Our results suggest that electric field complexity may have influenced the evolution of the JAR by degrading the related sensory information.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/QZM9PLW2/Shifman and Lewis - 2018 - The complexity of high-frequency electric fields d.pdf}
+}
+
+@book{sillarNeuroethologyPredationEscape,
+  title = {The {{Neuroethology}} of {{Predation}} and {{Escape}}},
+  author = {Sillar, Keith T},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/EUML6JSK/sillar_the neuroethology of predation and escape.pdf}
+}
+
+@article{silvaSexualSeasonalPlasticity2008,
+  title = {Sexual and Seasonal Plasticity in the Emission of Social Electric Signals. {{Behavioral}} Approach and Neural Bases},
+  author = {Silva, Ana and Quintana, Laura and Perrone, Rossana and Sierra, Felipe},
+  date = {2008-07},
+  journaltitle = {Journal of Physiology-Paris},
+  shortjournal = {Journal of Physiology-Paris},
+  volume = {102},
+  number = {4-6},
+  pages = {272--278},
+  issn = {09284257},
+  doi = {10.1016/j.jphysparis.2008.10.016},
+  url = {https://linkinghub.elsevier.com/retrieve/pii/S0928425708000557},
+  urldate = {2022-04-21},
+  abstract = {Behav­ior in elec­tric fish includes modu­ l­a­tions of a ster­ eo­typed elect­ ric organ disc­ harge (EOD) in addit­ ion to locom­ o­tor disp­ lays. Gym­not­i­for­mes can modu­ l­ate the EOD rate to prod­ uce sig­nals that par­tic­i­pate in dif­fer­ent behavi­ors. We studi­ed the reprod­ uc­tive behav­ior of Brachy­hyp­o­po­mus pinni­caud­a­tus both in the wild and lab­o­ra­tory sett­ ings. Duri­ng the breed­ing seas­ on, fish pro­duce sexu­ ­ally dimorp­ hic social elec­tric sig­nals (SES): males emit three types of chirps (dis­tin­guished by their durat­ ion and intern­ al struc­ture), and accel­er­a­tions, whereas females inter­rupt their EOD. Since these SES imply EOD fre­quency mod­u­la­ tions, the pacem­ aker nucleus (PN) is involved in their gene­ r­a­tion and cons­ ti­tutes the main tar­get organ to explore seas­ onal and sexu­ al plas­tic­ity of the CNS. The PN has two types of neur­ ons, pace­mak­ers and relays, which receive modu­ ­lat­ ory inputs from pre-pacem­ aker struct­ ures. These neur­ ons show an aniso­ tropic ro­stro-cau­dal and dorso-ven­tral dis­tri­bu­tion that is par­all­eled by dif­fer­ent field potential wave­ forms in dis­tinct por­tions of the PN. In vivo glu­ta­mate inject­ ions in dif­fer­ent areas of the PN pro­voke dif­fer­ent kinds of EOD rate mod­u­lat­ ions. Vent­ ral inject­ ions pro­duce chirp-like responses in breedi­ng males and EOD inter­rup­tions in breedi­ng females, whereas dors­ al inject­ ions pro­voke EOD freq­ uency rises in both sexes. In the non-breed­ing seas­ on, males and females respond with inter­rup­tions when stim­u­ lated vent­ rally and fre­quency rises when injected dors­ ally. Our results show that changes of glut­ a­mate effects in the PN could explain the sea­sonal and sex­ual dif­fer­ences in the gen­er­a­tion of SES. By means of behav­ioral recordi­ngs both in the wild and in labo­ ­ra­tory sett­ ings, and by elect­ rop­ hysi­­o­logi­­cal and phar­ ma­co­log­i­cal experi­­ments, we have iden­ti­fied sexu­ al and seas­ onal plas­tic­ity of the CNS and explored its under­ly­ing mech­a­nisms.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/TNVJTC38/Silva et al. - 2008 - Sexual and seasonal plasticity in the emission of .pdf}
+}
+
+@article{sinzSimultaneousSpiketimeLocking2020,
+  title = {Simultaneous Spike-Time Locking to Multiple Frequencies},
+  author = {Sinz, Fabian H. and Sachgau, Carolin and Henninger, Jörg and Benda, Jan and Grewe, Jan},
+  date = {2020-06-01},
+  journaltitle = {Journal of Neurophysiology},
+  shortjournal = {Journal of Neurophysiology},
+  volume = {123},
+  number = {6},
+  pages = {2355--2372},
+  issn = {0022-3077, 1522-1598},
+  doi = {10.1152/jn.00615.2019},
+  url = {https://journals.physiology.org/doi/10.1152/jn.00615.2019},
+  urldate = {2022-04-21},
+  abstract = {Locking of neuronal spikes to external and internal signals is a ubiquitous neurophysiological mechanism that has been extensively studied in several brain areas and species. Using experimental data from the electrosensory system and concise mathematical models, we analyze how a single neuron can simultaneously lock to multiple frequencies. Our findings demonstrate how temporal and rate codes can complement each other and lead to rich neuronal representations of sensory signals.           ,              Locking of neural firing is ubiquitously observed in the brain and occurs when neurons fire at a particular phase or in synchronization with an external signal. Here we study in detail the locking of single neurons to multiple distinct frequencies at the example of p-type electroreceptor afferents in the electrosensory system of the weakly electric fish Apteronotus leptorhynchus (brown ghost knifefish). We find that electrosensory afferents and pyramidal cells in the electrosensory lateral line lobe (ELL) lock to multiple frequencies, including the electric organ discharge (EOD) frequency, beat, and stimulus itself. We identify key elements necessary for locking to multiple frequencies, study its limits, and provide concise mathematical models reproducing our main findings. Our findings provide another example of how rate and temporal codes can coexist and complement each other in single neurons and demonstrate that sensory coding in p-type electroreceptor afferents provides a much richer representation of the sensory environment than commonly assumed. Since the underlying mechanisms are not specific to the electrosensory system, our results could provide the basis for studying multiple frequency locking in other systems.             NEW \& NOTEWORTHY Locking of neuronal spikes to external and internal signals is a ubiquitous neurophysiological mechanism that has been extensively studied in several brain areas and species. Using experimental data from the electrosensory system and concise mathematical models, we analyze how a single neuron can simultaneously lock to multiple frequencies. Our findings demonstrate how temporal and rate codes can complement each other and lead to rich neuronal representations of sensory signals.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/SY6ZR3F7/Sinz et al. - 2020 - Simultaneous spike-time locking to multiple freque.pdf}
+}
+
+@article{sohJammingAvoidanceResponse2018,
+  title = {Jamming {{Avoidance Response Inspired}} by {{Wave-type Weakly Electric Fish}}},
+  author = {Soh, Jaehyun and Kim, DaeEun},
+  date = {2018-11},
+  journaltitle = {Journal of Bionic Engineering},
+  shortjournal = {J Bionic Eng},
+  volume = {15},
+  number = {6},
+  pages = {982--991},
+  issn = {1672-6529, 2543-2141},
+  doi = {10.1007/s42235-018-0086-9},
+  url = {http://link.springer.com/10.1007/s42235-018-0086-9},
+  urldate = {2022-04-22},
+  abstract = {Weakly electric fish use the electric field to detect objects in the neighborhood or communicate with conspecifics. They generate electric field with their electric organ and the electroreceptors sense the distortion of electric field caused by nearby objects. They use a modulated frequency signal of the Electric Organ Discharge (EOD), and it can be disturbed by similar frequency signals that neighboring weakly electric fish emit. They have a particular behavior response to change their EOD frequencies to avoid signal jamming. It is called jamming avoidance response. Inspired by the behavior of wave-type weakly electric fish, we propose an engineering perspective of jamming avoidance response model with the amplitude-phase modulation graph. The time course of the amplitude-phase graph of the EOD signal provides a cue to detect the jamming signal. We argue that the temporal integration can determine the shift direction of the EOD as well as the amount of the frequency shift to be moved frequency for the jamming avoidance response. Alternatively, as a fast adapting measure, the cross product of point vectors in the amplitude-phase graph allows early decision for jamming avoidance. We demonstrate the methods with simulations and the real experiments in the underwater environment.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/9SP3FGBL/Soh and Kim - 2018 - Jamming Avoidance Response Inspired by Wave-type W.pdf}
+}
+
+@article{stamperJammingAvoidanceResponse2012,
+  title = {Beyond the {{Jamming Avoidance Response}}: Weakly Electric Fish Respond to the Envelope of Social Electrosensory Signals},
+  shorttitle = {Beyond the {{Jamming Avoidance Response}}},
+  author = {Stamper, Sarah A. and Madhav, Manu S. and Cowan, Noah J. and Fortune, Eric S.},
+  date = {2012-12-01},
+  journaltitle = {Journal of Experimental Biology},
+  volume = {215},
+  number = {23},
+  pages = {4196--4207},
+  issn = {1477-9145, 0022-0949},
+  doi = {10.1242/jeb.076513},
+  url = {https://journals.biologists.com/jeb/article/215/23/4196/11288/Beyond-the-Jamming-Avoidance-Response-weakly},
+  urldate = {2022-04-21},
+  abstract = {Recent studies have shown that central nervous system neurons in weakly electric fish respond to artificially constructed electrosensory envelopes, but the behavioral relevance of such stimuli is unclear. Here we investigate the possibility that social context creates envelopes that drive behavior. When Eigenmannia virescens are in groups of three or more, the interactions between their pseudo-sinusoidal electric fields can generate ʻsocial envelopesʼ. We developed a simple mathematical prediction for how fish might respond to such social envelopes. To test this prediction, we measured the responses of E. virescens to stimuli consisting of two sinusoids, each outside the range of the Jamming Avoidance Response (JAR), that when added to the fishʼs own electric field produced low-frequency (below 10Hz) social envelopes. Fish changed their electric organ discharge (EOD) frequency in response to these envelopes, which we have termed the Social Envelope Response (SER). In 99\% of trials, the direction of the SER was consistent with the mathematical prediction. The SER was strongest in response to the lowest initial envelope frequency tested (2Hz) and depended on stimulus amplitude. The SER generally resulted in an increase of the envelope frequency during the course of a trial, suggesting that this behavior may be a mechanism for avoiding low-frequency social envelopes. Importantly, the direction of the SER was not predicted by the superposition of two JAR responses: the SER was insensitive to the amplitude ratio between the sinusoids used to generate the envelope, but was instead predicted by the sign of the difference of difference frequencies.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/GTDNSJ5A/Stamper et al. - 2012 - Beyond the Jamming Avoidance Response weakly elec.pdf}
+}
+
+@article{stamperSpeciesDifferencesGroup2010a,
+  title = {Species Differences in Group Size and Electrosensory Interference in Weakly Electric Fishes: {{Implications}} for Electrosensory Processing},
+  shorttitle = {Species Differences in Group Size and Electrosensory Interference in Weakly Electric Fishes},
+  author = {Stamper, Sarah A. and Carrera-G, Erika and Tan, Eric W. and Fugère, Vincent and Krahe, Rüdiger and Fortune, Eric S.},
+  date = {2010-03-05},
+  journaltitle = {Behavioural Brain Research},
+  shortjournal = {Behavioural Brain Research},
+  volume = {207},
+  number = {2},
+  pages = {368--376},
+  issn = {01664328},
+  doi = {10.1016/j.bbr.2009.10.023},
+  url = {https://linkinghub.elsevier.com/retrieve/pii/S0166432809006317},
+  urldate = {2022-04-22},
+  abstract = {In animals with active sensory systems, group size can have dramatic effects on the sensory information available to individuals. In “wave-type” weakly electric fishes there is a categorical difference in sensory processing between solitary fish and fish in groups: when conspecifics are within about 1 m of each other, the electric fields mix and produce interference patterns that are detected by electroreceptors on each individual. Neural circuits in these animals must therefore process two streams of information—salient signals from prey items and predators and social signals from nearby conspecifics. We investigated the parameters of social signals in two genera of sympatric weakly electric fishes, Apteronotus and Sternopygus, in natural habitats of the Napo River valley in Ecuador and in laboratory settings. Apteronotus were most commonly found in pairs along the Napo River (47\% of observations; maximum group size 4) and produced electrosensory interference at rates of 20–300 Hz. In contrast, Sternopygus were alone in 80\% of observations (maximum group size 2) in the same region of Ecuador. Similar patterns were observed in laboratory experiments: Apteronotus were in groups and preferentially approached conspecific-like signals in an electrotaxis experiment whereas Sternopygus tended to be solitary and did not approach conspecific-like electrosensory signals. These results demonstrate categorical differences in social electrosensory-related activation of central nervous system circuits that may be related to the evolution of the jamming avoidance response that is used in Apteronotus but not Sternopygus to increase the frequency of electrosensory interference patterns.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/BTUW8VXR/Stamper et al. - 2010 - Species differences in group size and electrosenso.pdf}
+}
+
+@book{stevensSensoryEcologyBehaviour2013,
+  title = {Sensory Ecology, Behaviour, and Evolution},
+  author = {Stevens, Martin},
+  date = {2013},
+  edition = {First edition},
+  publisher = {{Oxford University Press}},
+  location = {{Oxford, United Kingdom}},
+  isbn = {978-0-19-960177-6 978-0-19-960178-3},
+  langid = {english},
+  pagetotal = {247},
+  keywords = {Animal behavior,Animal ecophysiology,Evolution},
+  annotation = {OCLC: ocn812257233},
+  file = {/home/weygoldt/Data/zotero/storage/IJ5VZMKE/stevens_2013_sensory ecology, behaviour, and evolution.pdf}
+}
+
+@article{stoddardPredationCrypsisEvolution2019,
+  title = {Predation and {{Crypsis}} in the {{Evolution}} of {{Electric Signaling}} in {{Weakly Electric Fishes}}},
+  author = {Stoddard, Philip K. and Tran, Alex and Krahe, Rüdiger},
+  date = {2019},
+  journaltitle = {Frontiers in Ecology and Evolution},
+  volume = {7},
+  pages = {264},
+  issn = {2296-701X},
+  doi = {10.3389/fevo.2019.00264},
+  url = {https://www.frontiersin.org/article/10.3389/fevo.2019.00264},
+  urldate = {2021-12-01},
+  abstract = {Eavesdropping by electroreceptive predators poses a conflict for weakly electric fish, which depend on their Electric Organ Discharge (EOD) signals both for navigation and communication in the dark. The EODs that allow weakly electric fish to electrolocate and communicate in the dark may attract electroreceptive predators such as catfishes and Electric Eels. These predators share with their prey the synapomorphy of passive electric sense supported by ampullary electroreceptors that are highly sensitive to low-frequency electric fields. Any low-frequency spectral components of the EOD make weakly electric fish conspicuous and vulnerable to attack from electroreceptive predators. Accordingly, most weakly electric fish shift spectral energy upwards or cloak low-frequency energy with compensatory masking signals. Subadults and females in particular emit virtually no low-frequency energy in their EODs, whereas courting males include a significant low-frequency component, which likely attracts females, but makes the signals conspicuous to predators. Males of species that coexist with the most predators tend to produce the least low-frequency signal energy, expressing sexual dimorphism in their signals in less risky ways. In these respects, electric signals follow the classic responses to opposing forces of natural and sexual selection, as exemplified in the visual signals of guppies and the acoustic signals of Túngara frogs. Unique to electric fish is that the electric signal modifications that help elude detection by electroreceptive predators are additions to the basal signal rather than losses of attractive components. These enhancements that enable crypsis are energetically costly, but have also provided the evolutionary substrates for subsequent sexual selection and species identity characters.},
+  keywords = {important},
+  file = {/home/weygoldt/Data/zotero/storage/W4D4C8DY/stoddard_2019_predation and crypsis in the evolution of electric signaling in weakly electric fishes.pdf}
+}
+
+@article{stoddardPredationEnhancesComplexity1999,
+  title = {Predation Enhances Complexity in the Evolution of Electric ®sh Signals},
+  author = {Stoddard, Philip K},
+  date = {1999},
+  volume = {400},
+  pages = {3},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/SP5R3CNX/Stoddard_1999_Predation enhances complexity in the evolution of .pdf}
+}
+
+@article{sullivanMolecularSystematicsAfrican2000,
+  title = {Molecular Systematics of the African Electric Fishes ({{Mormyroidea}}: Teleostei) and a Model for the Evolution of Their Electric Organs},
+  shorttitle = {Molecular Systematics of the African Electric Fishes ({{Mormyroidea}}},
+  author = {Sullivan, J.P. and Lavoue, S. and Hopkins, C.D.},
+  date = {2000-02-15},
+  journaltitle = {Journal of Experimental Biology},
+  volume = {203},
+  number = {4},
+  pages = {665--683},
+  issn = {1477-9145, 0022-0949},
+  doi = {10.1242/jeb.203.4.665},
+  url = {https://journals.biologists.com/jeb/article/203/4/665/8367/Molecular-systematics-of-the-african-electric},
+  urldate = {2021-12-06},
+  abstract = {We present a new molecular phylogeny for 41 species of monophyletic. Within the Mormyrinae, the genus Myomyrus African mormyroid electric fishes derived from the 12S, 16S is the sister group to all the remaining taxa. Other welland cytochrome b genes and the nuclear RAG2 gene. From supported clades within this group are recovered. A this, we reconstruct the evolution of the complex electric reconstruction of electrocyte evolution on the basis of our organs of these fishes. Phylogenetic results are generally best-supported topology suggests that electrocytes with concordant with earlier preliminary molecular studies of a penetrating stalks evolved once early in the history of the smaller group of species and with the osteology-based mormyrids followed by multiple paedomorphic reversals to classification of Taverne, which divides the group into electrocytes with non-penetrating stalks.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/YWT6TGRM/Sullivan et al._2000_Molecular systematics of the african electric fish.pdf}
+}
+
+@article{takizawaFrequencyDecelerationEigenmannia,
+  title = {Frequency Deceleration in {{Eigenmannia}}},
+  author = {Takizawa, Y and Rose, G J and Kawasaki, M},
+  pages = {10},
+  abstract = {The algorithm for the control of the jamming avoidance modulation appear to be sufficient to explain previous response (JAR) of Eigenmannia has been the subject of experimental results on which the former theory is based. debate for over two decades. Two competing theories have Specifically, fish of the genus Eigenmannia show been proposed to explain how fish determine the correct differential deceleration responses to asymmetric beat direction to shift their pacemaker frequency during envelopes. The deceleration responses do not require jamming. One theory emphasizes the role of time- phase modulation and show a sensitivity for amplitude asymmetric beat envelopes, while the other emphasizes modulation depth and selectivity for amplitude the role of amplitude- and phase-difference computations modulation rate comparable with that of JARs that are that arise from the differences in spatial geometry of the elicited when amplitude- and phase-difference electric fields of neighboring fish. In repeating earlier information is available.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/53JHQH9U/Takizawa et al. - Frequency deceleration in Eigenmannia.pdf}
+}
+
+@article{tallarovicElectricOrganDischarge2005,
+  title = {Electric Organ Discharge Frequency Jamming during Social Interactions in Brown Ghost Knifefish, {{Apteronotus}} Leptorhynchus},
+  author = {Tallarovic, Sara K. and Zakon, Harold H.},
+  date = {2005-12},
+  journaltitle = {Animal Behaviour},
+  shortjournal = {Animal Behaviour},
+  volume = {70},
+  number = {6},
+  pages = {1355--1365},
+  issn = {00033472},
+  doi = {10.1016/j.anbehav.2005.03.020},
+  url = {https://linkinghub.elsevier.com/retrieve/pii/S0003347205003088},
+  urldate = {2022-04-22},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/6LWWAMB9/Tallarovic and Zakon - 2005 - Electric organ discharge frequency jamming during .pdf}
+}
+
+@article{tallarovicElectricOrganDischarge2005b,
+  title = {Electric Organ Discharge Frequency Jamming during Social Interactions in Brown Ghost Knifefish, {{Apteronotus}} Leptorhynchus},
+  author = {Tallarovic, Sara K. and Zakon, Harold H.},
+  date = {2005-12},
+  journaltitle = {Animal Behaviour},
+  shortjournal = {Animal Behaviour},
+  volume = {70},
+  number = {6},
+  pages = {1355--1365},
+  issn = {00033472},
+  doi = {10.1016/j.anbehav.2005.03.020},
+  url = {https://linkinghub.elsevier.com/retrieve/pii/S0003347205003088},
+  urldate = {2022-04-22},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/SJSDLW23/Tallarovic and Zakon - 2005 - Electric organ discharge frequency jamming during .pdf}
+}
+
+@article{triefenbachApteronotusLeptorhynchus,
+  title = {Apteronotus Leptorhynchus},
+  author = {Triefenbach, Frank Alexander},
+  pages = {88},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/IN7BUH9W/Triefenbach - Apteronotus leptorhynchus.pdf}
+}
+
+@article{triefenbachChangesSignallingAgonistic2008b,
+  title = {Changes in Signalling during Agonistic Interactions between Male Weakly Electric Knifefish, {{Apteronotus}} Leptorhynchus},
+  author = {Triefenbach, Frank A. and Zakon, Harold H.},
+  date = {2008-04},
+  journaltitle = {Animal Behaviour},
+  shortjournal = {Animal Behaviour},
+  volume = {75},
+  number = {4},
+  pages = {1263--1272},
+  issn = {00033472},
+  doi = {10.1016/j.anbehav.2007.09.027},
+  url = {https://linkinghub.elsevier.com/retrieve/pii/S0003347208000110},
+  urldate = {2022-04-22},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/BQD2LMTY/Triefenbach and Zakon - 2008 - Changes in signalling during agonistic interaction.pdf}
+}
+
+@article{triefenbachEffectsSexSensitivity2003,
+  title = {Effects of Sex, Sensitivity and Status on Cue Recognition in the Weakly Electric Fish {{Apteronotus}} Leptorhynchus},
+  author = {Triefenbach, Frank and Zakon, Harold},
+  date = {2003-01},
+  journaltitle = {Animal Behaviour},
+  shortjournal = {Animal Behaviour},
+  volume = {65},
+  number = {1},
+  pages = {19--28},
+  issn = {00033472},
+  doi = {10.1006/anbe.2002.2019},
+  url = {https://linkinghub.elsevier.com/retrieve/pii/S0003347202920191},
+  urldate = {2022-04-22},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/T47EX9PE/Triefenbach and Zakon - 2003 - Effects of sex, sensitivity and status on cue reco.pdf}
+}
+
+@book{vonderemdeEcologyAnimalSenses2016,
+  title = {The {{Ecology}} of {{Animal Senses}}},
+  editor = {von der Emde, Gerhard and Warrant, Eric},
+  options = {useprefix=true},
+  date = {2016},
+  publisher = {{Springer International Publishing}},
+  location = {{Cham}},
+  doi = {10.1007/978-3-319-25492-0},
+  url = {http://link.springer.com/10.1007/978-3-319-25492-0},
+  urldate = {2021-12-01},
+  isbn = {978-3-319-25490-6 978-3-319-25492-0},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/P2L46XDG/von der emde_2016_the ecology of animal senses.pdf}
+}
+
+@article{walzNeuroethologyElectrocommunicationHow2013,
+  title = {The Neuroethology of Electrocommunication: {{How}} Signal Background Influences Sensory Encoding and Behaviour in {{Apteronotus}} Leptorhynchus},
+  shorttitle = {The Neuroethology of Electrocommunication},
+  author = {Walz, Henriette and Hupé, Ginette J. and Benda, Jan and Lewis, John E.},
+  date = {2013-01},
+  journaltitle = {Journal of Physiology-Paris},
+  shortjournal = {Journal of Physiology-Paris},
+  volume = {107},
+  number = {1-2},
+  pages = {13--25},
+  issn = {09284257},
+  doi = {10.1016/j.jphysparis.2012.07.001},
+  url = {https://linkinghub.elsevier.com/retrieve/pii/S092842571200040X},
+  urldate = {2022-04-22},
+  abstract = {Weakly-electric fish are a well-established model system for neuroethological studies on communication and aggression. Sensory encoding of their electric communication signals, as well as behavioural responses to these signals, have been investigated in great detail under laboratory conditions. In the wave-type brown ghost knifefish, Apteronotus leptorhynchus, transient increases in the frequency of the generated electric field, called chirps, are particularly well-studied, since they can be readily evoked by stimulating a fish with artificial signals mimicking conspecifics. When two fish interact, both their quasi-sinusoidal electric fields (called electric organ discharge, EOD) superimpose, resulting in a beat, an amplitude modulation at the frequency difference between the two EODs. Although chirps themselves are highly stereotyped signals, the shape of the amplitude modulation resulting from a chirp superimposed on a beat background depends on a number of parameters, such as the beat frequency, modulation depth, and beat phase at which the chirp is emitted. Here we review the influence of these beat parameters on chirp encoding in the three primary stages of the electrosensory pathway: electroreceptor afferents, the hindbrain electrosensory lateral line lobe, and midbrain torus semicircularis. We then examine the role of these parameters, which represent specific features of various social contexts, on the behavioural responses of A. leptorhynchus. Some aspects of the behaviour may be explained by the coding properties of early sensory neurons to chirp stimuli. However, the complexity and diversity of behavioural responses to chirps in the context of different background parameters cannot be explained solely on the basis of the sensory responses and thus suggest that critical roles are played by higher processing stages.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/Y4U2QMH7/Walz et al. - 2013 - The neuroethology of electrocommunication How sig.pdf}
+}
+
+@article{walzStaticFrequencyTuning2014,
+  title = {Static Frequency Tuning Accounts for Changes in Neural Synchrony Evoked by Transient Communication Signals},
+  author = {Walz, Henriette and Grewe, Jan and Benda, Jan},
+  date = {2014-08-15},
+  journaltitle = {Journal of Neurophysiology},
+  shortjournal = {Journal of Neurophysiology},
+  volume = {112},
+  number = {4},
+  pages = {752--765},
+  issn = {0022-3077, 1522-1598},
+  doi = {10.1152/jn.00576.2013},
+  url = {https://www.physiology.org/doi/10.1152/jn.00576.2013},
+  urldate = {2022-04-21},
+  abstract = {Although communication signals often vary continuously on the underlying signal parameter, they are perceived as distinct categories. We here report the opposite case where an electrocommunication signal is encoded in four distinct regimes, although the behavior described to date does not show distinct categories. In particular, we studied the encoding of chirps by P-unit afferents in the weakly electric fish Apteronotus leptorhynchus. These fish generate an electric organ discharge that oscillates at a certain individual-specific frequency. The interaction of two fish in communication contexts leads to the emergence of a beating amplitude modulation (AM) at the frequency difference between the two individual signals. This frequency difference represents the social context of the encounter. Chirps are transient increases of the fish's frequency leading to transient changes in the frequency of the AM. We stimulated the cells with the same chirp on different, naturally occurring backgrounds beats. The P-units responded either by synchronization or desynchronization depending on the background. Although the duration of a chirp is often shorter than a full cycle of the AM it elicits, the distinct responses of the P-units to the chirp can be predicted solely from the frequency of the AM based on the static frequency tuning of the cells.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/TSX64FN3/Walz et al. - 2014 - Static frequency tuning accounts for changes in ne.pdf}
+}
+
+@article{wormSocialInteractionsLive2017,
+  title = {Social Interactions between Live and Artificial Weakly Electric Fish: {{Electrocommunication}} and Locomotor Behavior of {{Mormyrus}} Rume Proboscirostris towards a Mobile Dummy Fish},
+  shorttitle = {Social Interactions between Live and Artificial Weakly Electric Fish},
+  author = {Worm, Martin and Kirschbaum, Frank and von der Emde, Gerhard},
+  editor = {Engelmann, Jacob},
+  options = {useprefix=true},
+  date = {2017-09-13},
+  journaltitle = {PLOS ONE},
+  shortjournal = {PLoS ONE},
+  volume = {12},
+  number = {9},
+  pages = {e0184622},
+  issn = {1932-6203},
+  doi = {10.1371/journal.pone.0184622},
+  url = {https://dx.plos.org/10.1371/journal.pone.0184622},
+  urldate = {2021-11-30},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/PBXKKV2A/Worm et al._2017_Social interactions between live and artificial we.pdf}
+}
+
+@article{yuCodingConspecificIdentity2012,
+  title = {Coding {{Conspecific Identity}} and {{Motion}} in the {{Electric Sense}}},
+  author = {Yu, Na and Hupé, Ginette and Garfinkle, Charles and Lewis, John E. and Longtin, André},
+  editor = {Gabbiani, Fabrizio},
+  date = {2012-07-12},
+  journaltitle = {PLoS Computational Biology},
+  shortjournal = {PLoS Comput Biol},
+  volume = {8},
+  number = {7},
+  pages = {e1002564},
+  issn = {1553-7358},
+  doi = {10.1371/journal.pcbi.1002564},
+  url = {https://dx.plos.org/10.1371/journal.pcbi.1002564},
+  urldate = {2022-04-21},
+  abstract = {Interactions among animals can result in complex sensory signals containing a variety of socially relevant information, including the number, identity, and relative motion of conspecifics. How the spatiotemporal properties of such evolving naturalistic signals are encoded is a key question in sensory neuroscience. Here, we present results from experiments and modeling that address this issue in the context of the electric sense, which combines the spatial aspects of vision and touch, with the temporal aspects of audition. Wave-type electric fish, such as the brown ghost knifefish, Apteronotus leptorhynchus, used in this study, are uniquely identified by the frequency of their electric organ discharge (EOD). Multiple beat frequencies arise from the superposition of the EODs of each fish. We record the natural electrical signals near the skin of a ‘‘receiving’’ fish that are produced by stationary and freely swimming conspecifics. Using spectral analysis, we find that the primary beats, and the secondary beats between them (‘‘beats of beats’’), can be greatly influenced by fish swimming; the resulting motion produces low-frequency envelopes that broaden all the beat peaks and reshape the ‘‘noise floor’’. We assess the consequences of this motion on sensory coding using a model electroreceptor. We show that the primary and secondary beats are encoded in the afferent spike train, but that motion acts to degrade this encoding. We also simulate the response of a realistic population of receptors, and find that it can encode the motion envelope well, primarily due to the receptors with lower firing rates. We discuss the implications of our results for the identification of conspecifics through specific beat frequencies and its possible hindrance by active swimming.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/53FIAG8P/Yu et al. - 2012 - Coding Conspecific Identity and Motion in the Elec.pdf}
+}
+
+@article{zaitounyTrackingSinglePigeon2017,
+  title = {Tracking a Single Pigeon Using a Shadowing Filter Algorithm},
+  author = {Zaitouny, Ayham and Stemler, Thomas and Small, Michael},
+  date = {2017-06},
+  journaltitle = {Ecology and Evolution},
+  shortjournal = {Ecol Evol},
+  volume = {7},
+  number = {12},
+  pages = {4419--4431},
+  issn = {20457758},
+  doi = {10.1002/ece3.2976},
+  url = {https://onlinelibrary.wiley.com/doi/10.1002/ece3.2976},
+  urldate = {2022-07-03},
+  abstract = {Miniature GPS devices now allow for measurement of the movement of animals in real time and provide high- quality and high-resolution data. While these new data sets are a great improvement, one still encounters some measurement errors as well as device failures. Moreover, these devices only measure position and require further reconstruction techniques to extract the full dynamical state space with the velocity and acceleration. Direct differentiation of position is generally not adequate. We report on the successful implementation of a shadowing filter algorithm that (1) minimizes measurement errors and (2) reconstructs at the same time the full phase-space from a position recording of a flying pigeon. This filter is based on a very simple assumption that the pigeon's dynamics are Newtonian. We explore not only how to choose the filter's parameters but also demonstrate its improvements over other techniques and give minimum data requirements. In contrast to competing filters, the shadowing filter's approach has not been widely implemented for practical problems. This article ­addresses these practicalities and provides a prototype for such application.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/ZQFQF52L/Zaitouny et al. - 2017 - Tracking a single pigeon using a shadowing filter .pdf}
+}
+
+@article{zakonEODModulationsBrown2002a,
+  title = {{{EOD}} Modulations of Brown Ghost Electric Fish: {{JARs}}, Chirps, Rises, and Dips},
+  shorttitle = {{{EOD}} Modulations of Brown Ghost Electric Fish},
+  author = {Zakon, Harold and Oestreich, Joerg and Tallarovic, Sara and Triefenbach, Frank},
+  date = {2002-09},
+  journaltitle = {Journal of Physiology-Paris},
+  shortjournal = {Journal of Physiology-Paris},
+  volume = {96},
+  number = {5-6},
+  pages = {451--458},
+  issn = {09284257},
+  doi = {10.1016/S0928-4257(03)00012-3},
+  url = {https://linkinghub.elsevier.com/retrieve/pii/S0928425703000123},
+  urldate = {2022-04-22},
+  abstract = {Weakly electric ‘‘wave’’ fish make highly regular electric organ discharges (EODs) for precise electrolocation. Yet, they modulate the ongoing rhythmicity of their EOD during social interactions. These modulations may last from a few milliseconds to tens of minutes. In this paper we describe the different types of EOD modulations, what they may signal to recipient fish, and how they are generated on a neural level. Our main conclusions, based on a species called the brown ghost (Apteronotus leptorhynchus) are that fish: (1) show sexual dimorphism in the signals that they generate; (2) make different signals depending on whether they are interacting with a fish of the opposite sex or, within their own sex, to a fish of that which is dominant or subordinate to it; (3) are able to assess relative dominance from electrical cues; (4) have a type of plasticity in the pacemaker nucleus, the control center for the EOD, that occurs after stimulation of NMDA receptors that causes a long-lasting (tens of minutes to hours) change in EOD frequency; (5) that this NMDA receptor-dependent change may occur in reflexive responses, like the jamming avoidance response (JAR), as well as after certain long-lasting social signals. We propose that NMDA-receptor dependent increases in EOD frequency during the JAR adaptively shift the EOD frequency to a new value to avoid jamming by another fish and that such increases in EOD frequency during social encounters may be advantageous since social dominance seems to be positively correlated with EOD frequency in both sexes.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/25W4ITMC/Zakon et al. - 2002 - EOD modulations of brown ghost electric fish JARs.pdf}
+}
+
+@article{zakonMolecularEvolutionCommunication2008,
+  title = {Molecular Evolution of Communication Signals in Electric Fish},
+  author = {Zakon, Harold H. and Zwickl, Derrick J. and Lu, Ying and Hillis, David M.},
+  date = {2008-06-01},
+  journaltitle = {Journal of Experimental Biology},
+  volume = {211},
+  number = {11},
+  pages = {1814--1818},
+  issn = {1477-9145, 0022-0949},
+  doi = {10.1242/jeb.015982},
+  url = {https://journals.biologists.com/jeb/article/211/11/1814/9504/Molecular-evolution-of-communication-signals-in},
+  urldate = {2021-12-06},
+  abstract = {Animal communication systems are subject to natural selection so the imprint of selection must reside in the genome of each species. Electric fish generate electric organ discharges (EODs) from a muscle-derived electric organ (EO) and use these fields for electrolocation and communication. Weakly electric teleosts have evolved at least twice (mormyriforms, gymnotiforms) allowing a comparison of the workings of evolution in two independently evolved sensory/motor systems. We focused on the genes for two Na+ channels, Nav1.4a and Nav1.4b, which are orthologs of the mammalian muscle-expressed Na+ channel gene Nav1.4. Both genes are expressed in muscle in non-electric fish. Nav1.4b is expressed in muscle in electric fish, but Nav1.4a expression has been lost from muscle and gained in the evolutionarily novel EO in both groups. We hypothesized that Nav1.4a might be evolving to optimize the EOD for different sensory environments and the generation of species-specific communication signals. We obtained the sequence for Nav1.4a from non-electric, mormyriform and gymnotiform species, estimated a phylogenetic tree, and determined rates of evolution. We observed elevated rates of evolution in this gene in both groups coincident with the loss of Nav1.4a from muscle and its compartmentalization in EO. We found amino acid substitutions at sites known to be critical for channel inactivation; analyses suggest that these changes are likely to be the result of positive selection. We suggest that the diversity of EOD waveforms in both groups of electric fish is correlated with accelerations in the rate of evolution of the Nav1.4a Na+ channel gene due to changes in selection pressure on the gene once it was solely expressed in the EO.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/BDSH8IAI/Zakon et al._2008_Molecular evolution of communication signals in el.pdf}
+}
+
+@article{zakonSodiumChannelGenes2006,
+  title = {Sodium Channel Genes and the Evolution of Diversity in Communication Signals of Electric Fishes: {{Convergent}} Molecular Evolution},
+  shorttitle = {Sodium Channel Genes and the Evolution of Diversity in Communication Signals of Electric Fishes},
+  author = {Zakon, H. H. and Lu, Y. and Zwickl, D. J. and Hillis, D. M.},
+  date = {2006-03-07},
+  journaltitle = {Proceedings of the National Academy of Sciences},
+  shortjournal = {Proceedings of the National Academy of Sciences},
+  volume = {103},
+  number = {10},
+  pages = {3675--3680},
+  issn = {0027-8424, 1091-6490},
+  doi = {10.1073/pnas.0600160103},
+  url = {http://www.pnas.org/cgi/doi/10.1073/pnas.0600160103},
+  urldate = {2021-12-06},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/JQPNV23N/Zakon et al._2006_Sodium channel genes and the evolution of diversit.pdf}
+}
+
+@article{zupancElectricInteractionsChirping2006,
+  title = {Electric Interactions through Chirping Behavior in the Weakly Electric Fish, {{Apteronotus}} Leptorhynchus},
+  author = {Zupanc, G. K. H. and Sîrbulescu, R. F. and Nichols, A. and Ilies, I.},
+  date = {2006-02},
+  journaltitle = {Journal of Comparative Physiology A},
+  shortjournal = {J Comp Physiol A},
+  volume = {192},
+  number = {2},
+  pages = {159--173},
+  issn = {0340-7594, 1432-1351},
+  doi = {10.1007/s00359-005-0058-5},
+  url = {http://link.springer.com/10.1007/s00359-005-0058-5},
+  urldate = {2022-04-22},
+  abstract = {The weakly electric fish Apteronotus leptorhynchus produces wave-like electric organ discharges distinguished by a high degree of regularity. Transient amplitude and frequency modulations (‘‘chirps’’) can be evoked in males by stimulation with the electric field of a conspecific. During these interactions, the males examined in this study produced six types of chirps, including two novel ones. Stimulation of a test fish with a conspecific at various distances showed that two electrically interacting fish must be within 10 cm of each other to evoke chirping behavior in the neighboring fish. The chirp rate of all but one chirp type elicited by the neighboring fish was found to be negatively correlated with the absolute value of the frequency difference between the two interacting fish, but independent of the sign of this difference. Correlation analysis of the instantaneous rates of chirp occurrence revealed two modes of interactions characterized by reciprocal stimulation and reciprocal inhibition. Further analysis of the temporal relationship between the chirps generated by the two fish during electric interactions showed that the chirps generated by one individual follow the chirps of the other with a short latency of approximately 500–1000 ms. We hypothesize that this ‘‘echo response’’ serves a communicatory function.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/Z4NXZVQ9/Zupanc et al. - 2006 - Electric interactions through chirping behavior in.pdf}
+}
+
+@article{zupancOscillatorsModulatorsBehavioral2002,
+  title = {From Oscillators to Modulators: Behavioral and Neural Control of Modulations of the Electric Organ Discharge in the Gymnotiform Fish, {{Apteronotus}} Leptorhynchus},
+  shorttitle = {From Oscillators to Modulators},
+  author = {Zupanc, Günther K.H.},
+  date = {2002-09},
+  journaltitle = {Journal of Physiology-Paris},
+  shortjournal = {Journal of Physiology-Paris},
+  volume = {96},
+  number = {5-6},
+  pages = {459--472},
+  issn = {09284257},
+  doi = {10.1016/S0928-4257(03)00002-0},
+  url = {https://linkinghub.elsevier.com/retrieve/pii/S0928425703000020},
+  urldate = {2022-04-22},
+  abstract = {The brown ghost (Apteronotus leptorhynchus) is a weakly electric gymnotiform fish that produces wave-like electric organ discharges distinguished by their enormous degree of regularity. Transient modulations of these discharges occur both spontaneously and when stimulating the fish with external electric signals that mimic encounters with a neighboring fish. Two prominent forms of modulations are chirps and gradual frequency rises. Chirps are complex frequency and amplitude modulations lasting between 20 ms and more than 200 ms. Based on their biophysical characteristics, they can be divided into four distinct categories. Gradual frequency rises consist of a rise in discharge frequency, followed by a slow return to baseline frequency. Although the modulatory phase may vary considerably between a few 100 ms and almost 100 s, there is no evidence for the existence of distinct categories of this type of modulation signal. Stimulation of the fish with external electric signals results almost exclusively in the generation of type-2 chirps. This effect is independent of the chirp type generated by the respective individual under non-evoked conditions. By contrast, no proper stimulation condition is known to evoke the other three types of chirps or gradual frequency rises in non-breeding fish. In contrast to the type-2 chirps evoked when subjecting the fish to external electric stimulation, the rate of spontaneously produced chirps is quite low. However, their rate appears to be optimized according to the probability of encountering a conspecific. As a result, the rate of non-evoked chirping is increased during the night when the fish exhibit high locomotor activity and in the time period following external electric stimulation. These, as well as other, observations demonstrate that both the type and rate of modulatory behavior are affected by a variety of behavioral conditions. This diversity at the behavioral level correlates with, and is likely to be causally linked to, the diversity of inputs received by the neurons that control chirps and gradual frequency rises, respectively. These neurons form two distinct sub-nuclei within the central posterior/prepacemaker nucleus in the dorsal thalamus. In vitro tract-tracing experiments have elucidated some of the connections of this complex with other brain regions. Direct input is received from the optic tectum. Indirect input arising from telencephalic and hypothalamic regions, as well as from the preoptic area, is relayed to the central posterior/prepacemaker nucleus via the preglomerular nucleus. Feedback loops may be provided by projections of the central posterior/prepacemaker nucleus to the preglomerular nucleus and the nucleus preopticus periventricularis. \# 2003 Elsevier Ltd. All rights reserved.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/PX5Y7FUS/Zupanc - 2002 - From oscillators to modulators behavioral and neu.pdf}
+}
+
+@article{zupancWalterHeiligenbergJamming2006,
+  title = {Walter {{Heiligenberg}}: The Jamming Avoidance Response and Beyond},
+  shorttitle = {Walter {{Heiligenberg}}},
+  author = {Zupanc, G. K. H. and Bullock, T. H.},
+  date = {2006-06},
+  journaltitle = {Journal of Comparative Physiology A},
+  shortjournal = {J Comp Physiol A},
+  volume = {192},
+  number = {6},
+  pages = {561--572},
+  issn = {0340-7594, 1432-1351},
+  doi = {10.1007/s00359-006-0098-5},
+  url = {http://link.springer.com/10.1007/s00359-006-0098-5},
+  urldate = {2022-04-22},
+  abstract = {Walter Heiligenberg (1938–1994) was an exceptionally gifted behavioral physiologist who made enormous contributions to the analysis of behavior and to our understanding of how the brain initiates and controls species-typical behavioral patterns. He was distinguished by his rigorous analytical approach used in both behavioral studies and neuroethological investigations. Among his most significant contributions to neuroethology are a detailed analysis of the computational rules governing the jamming avoidance response in weakly electric fish and the elucidation of the principal neural pathway involved in neural control of this behavior. Based on his work, the jamming avoidance response is perhaps the best-understood vertebrate behavior pattern in terms of the underlying neural substrate. In addition to this pioneering work, Heiligenberg stimulated research in a significant number of other areas of ethology and neuroethology, including: the quantitative assessment of aggressivity in cichlid fish; the ethological analysis of the stimulus–response relationship in the chirping behavior of crickets; the exploration of the neural and endocrine basis of communicatory behavior in weakly electric fish; the study of cellular mechanisms of neuronal plasticity in the adult fish brain; and the phylogenetic analysis of electric fishes using a combination of morphology, electrophysiology, and mitochondrial sequence data.},
+  langid = {english},
+  file = {/home/weygoldt/Data/zotero/storage/VU8DHZ6E/Zupanc and Bullock - 2006 - Walter Heiligenberg the jamming avoidance respons.pdf}
+}
+
+@Article{Hunter:2007,
+  Author    = {Hunter, J. D.},
+  Title     = {Matplotlib: A 2D graphics environment},
+  Journal   = {Computing in Science \& Engineering},
+  Volume    = {9},
+  Number    = {3},
+  Pages     = {90--95},
+  abstract  = {Matplotlib is a 2D graphics package used for Python for
+  application development, interactive scripting, and publication-quality
+  image generation across user interfaces and operating systems.},
+  publisher = {IEEE COMPUTER SOC},
+  doi       = {10.1109/MCSE.2007.55},
+  year      = 2007
+}
+
+@Article{         harris2020array,
+ title         = {Array programming with {NumPy}},
+ author        = {Charles R. Harris and K. Jarrod Millman and St{\'{e}}fan J.
+                 van der Walt and Ralf Gommers and Pauli Virtanen and David
+                 Cournapeau and Eric Wieser and Julian Taylor and Sebastian
+                 Berg and Nathaniel J. Smith and Robert Kern and Matti Picus
+                 and Stephan Hoyer and Marten H. van Kerkwijk and Matthew
+                 Brett and Allan Haldane and Jaime Fern{\'{a}}ndez del
+                 R{\'{i}}o and Mark Wiebe and Pearu Peterson and Pierre
+                 G{\'{e}}rard-Marchant and Kevin Sheppard and Tyler Reddy and
+                 Warren Weckesser and Hameer Abbasi and Christoph Gohlke and
+                 Travis E. Oliphant},
+ year          = {2020},
+ month         = sep,
+ journal       = {Nature},
+ volume        = {585},
+ number        = {7825},
+ pages         = {357--362},
+ doi           = {10.1038/s41586-020-2649-2},
+ publisher     = {Springer Science and Business Media {LLC}},
+ url           = {https://doi.org/10.1038/s41586-020-2649-2}
+}
+
+@ARTICLE{2020SciPy-NMeth,
+  author  = {Virtanen, Pauli and Gommers, Ralf and Oliphant, Travis E. and
+            Haberland, Matt and Reddy, Tyler and Cournapeau, David and
+            Burovski, Evgeni and Peterson, Pearu and Weckesser, Warren and
+            Bright, Jonathan and {van der Walt}, St{\'e}fan J. and
+            Brett, Matthew and Wilson, Joshua and Millman, K. Jarrod and
+            Mayorov, Nikolay and Nelson, Andrew R. J. and Jones, Eric and
+            Kern, Robert and Larson, Eric and Carey, C J and
+            Polat, {\.I}lhan and Feng, Yu and Moore, Eric W. and
+            {VanderPlas}, Jake and Laxalde, Denis and Perktold, Josef and
+            Cimrman, Robert and Henriksen, Ian and Quintero, E. A. and
+            Harris, Charles R. and Archibald, Anne M. and
+            Ribeiro, Ant{\^o}nio H. and Pedregosa, Fabian and
+            {van Mulbregt}, Paul and {SciPy 1.0 Contributors}},
+  title   = {{{SciPy} 1.0: Fundamental Algorithms for Scientific
+            Computing in Python}},
+  journal = {Nature Methods},
+  year    = {2020},
+  volume  = {17},
+  pages   = {261--272},
+  adsurl  = {https://rdcu.be/b08Wh},
+  doi     = {10.1038/s41592-019-0686-2},
+}
+
+@article{LAIDRE2013R829,
+title = {Animal signals},
+journal = {Current Biology},
+volume = {23},
+number = {18},
+pages = {R829-R833},
+year = {2013},
+issn = {0960-9822},
+doi = {https://doi.org/10.1016/j.cub.2013.07.070},
+url = {https://www.sciencedirect.com/science/article/pii/S0960982213009317},
+author = {Mark E. Laidre and Rufus A. Johnstone},
+abstract = {Summary
+The study of animal signals began in earnest with the publication in 1872 of Charles Darwin’s The Expressions of the Emotions in Man and Animals, which laid the basis for a comparative study of signals across all animals, including humans. Yet even before Darwin, the exceptional diversity of animal signals has gripped the attention of natural historians and laymen alike, as these signals represent some of the most striking features of the natural world. Structures such as the long ornamented tail of the peacock, the roaring sounds of howler monkeys, audible kilometers away, and the pheromone trails laid by ants to guide their nestmates to resources are each examples of animal signals (Figure 1). Indeed, because signals evolved for the purpose of communicating (Box 1), their prominence can be hard for even a casual observer to overlook. Animal signals therefore raise many scientific questions: What are their functions? What information do they transmit? How are they produced? And why did they evolve?}
+}
+
+@article{endler1993some,
+  title={Some general comments on the evolution and design of animal communication systems},
+  author={Endler, John A},
+  journal={Philosophical Transactions of the Royal Society of London. Series B: Biological Sciences},
+  volume={340},
+  number={1292},
+  pages={215--225},
+  year={1993},
+  publisher={The Royal Society London}
+}
+
+@article{hebets2016systems,
+  title={A systems approach to animal communication},
+  author={Hebets, Eileen A and Barron, Andrew B and Balakrishnan, Christopher N and Hauber, Mark E and Mason, Paul H and Hoke, Kim L},
+  journal={Proceedings of the Royal Society B: Biological Sciences},
+  volume={283},
+  number={1826},
+  pages={20152889},
+  year={2016},
+  publisher={The Royal Society}
+}
+
+@article{marler1967animal,
+  title={Animal Communication Signals: We are beginning to understand how the structure of animal signals relates to the function they serve.},
+  author={Marler, Peter},
+  journal={Science},
+  volume={157},
+  number={3790},
+  pages={769--774},
+  year={1967},
+  publisher={American Association for the Advancement of Science}
+}
+
+@incollection{searcy2010evolution,
+  title={The evolution of animal communication},
+  author={Searcy, William A and Nowicki, Stephen},
+  booktitle={The Evolution of Animal Communication},
+  year={2010},
+  publisher={Princeton University Press}
+}
+
+@article{seyfarth2010central,
+  title={The central importance of information in studies of animal communication},
+  author={Seyfarth, Robert M and Cheney, Dorothy L and Bergman, Thore and Fischer, Julia and Zuberb{\"u}hler, Klaus and Hammerschmidt, Kurt},
+  journal={Animal Behaviour},
+  volume={80},
+  number={1},
+  pages={3--8},
+  year={2010},
+  publisher={Elsevier}
+}
+
+@article{kalmijn1971electric,
+  title={The electric sense of sharks and rays},
+  author={Kalmijn, Ad J},
+  journal={Journal of Experimental Biology},
+  volume={55},
+  number={2},
+  pages={371--383},
+  year={1971},
+  publisher={Company of Biologists}
+}
+
+@techreport{kalmijn1973electro,
+  title={Electro-orientation in sharks and rays: theory and experimental evidence},
+  author={Kalmijn, Ad J},
+  year={1973},
+  institution={SCRIPPS INSTITUTION OF OCEANOGRAPHY LA JOLLA CA}
+}
+
+@article{catania2014shocking,
+  title={The shocking predatory strike of the electric eel},
+  author={Catania, Kenneth C},
+  journal={Science},
+  volume={346},
+  number={6214},
+  pages={1231--1234},
+  year={2014},
+  publisher={American Association for the Advancement of Science}
+}
+
+@article{catania2015electric,
+  title={Electric eels use high-voltage to track fast-moving prey},
+  author={Catania, Kenneth C},
+  journal={Nature Communications},
+  volume={6},
+  number={1},
+  pages={8638},
+  year={2015},
+  publisher={Nature Publishing Group UK London}
+}
+
+@article{brown1950electric,
+  title={The electric discharge of the electric EEL},
+  author={Brown, MV},
+  journal={Electrical Engineering},
+  volume={69},
+  number={2},
+  pages={145--147},
+  year={1950},
+  publisher={IEEE}
+}
+
+@article{wohr2013affective,
+  title={Affective communication in rodents: ultrasonic vocalizations as a tool for research on emotion and motivation},
+  author={W{\"o}hr, Markus and Schwarting, Rainer KW},
+  journal={Cell and tissue research},
+  volume={354},
+  number={1},
+  pages={81--97},
+  year={2013},
+  publisher={Springer}
+}
+
+@article{seffer2014pro,
+  title={Pro-social ultrasonic communication in rats: insights from playback studies},
+  author={Seffer, Dominik and Schwarting, Rainer KW and W{\"o}hr, Markus},
+  journal={Journal of neuroscience methods},
+  volume={234},
+  pages={73--81},
+  year={2014},
+  publisher={Elsevier}
+}
+
+@article{virant2004vibrational,
+  title={Vibrational communication in insects},
+  author={Virant-Doberlet, Meta and Cokl, Andrej},
+  journal={Neotropical Entomology},
+  volume={33},
+  pages={121--134},
+  year={2004},
+  publisher={SciELO Brasil}
+}
+
+@article{girard2011multi,
+  title={Multi-modal courtship in the peacock spider, Maratus volans (OP-Cambridge, 1874)},
+  author={Girard, Madeline B and Kasumovic, Michael M and Elias, Damian O},
+  journal={PLoS One},
+  volume={6},
+  number={9},
+  pages={e25390},
+  year={2011},
+  publisher={Public Library of Science San Francisco, USA}
+}
+
+@article{tibbetts2008visual,
+  title={Visual signals of status and rival assessment in Polistes dominulus paper wasps},
+  author={Tibbetts, Elizabeth A and Lindsay, Rebecca},
+  journal={Biology Letters},
+  volume={4},
+  number={3},
+  pages={237--239},
+  year={2008},
+  publisher={The Royal Society London}
+}
+
+@article{riley2005flight,
+  title={The flight paths of honeybees recruited by the waggle dance},
+  author={Riley, Joe R and Greggers, Uwe and Smith, Alan D and Reynolds, Don R and Menzel, Randolf},
+  journal={Nature},
+  volume={435},
+  number={7039},
+  pages={205--207},
+  year={2005},
+  publisher={Nature Publishing Group UK London}
+}
+
+@article{sudd1959interaction,
+  title={Interaction between ants on a scent trail},
+  author={Sudd, JH},
+  journal={Nature},
+  volume={183},
+  number={4675},
+  pages={1588--1588},
+  year={1959},
+  publisher={Nature Publishing Group UK London}
+}
+
+@article{schnitzler2003spatial,
+  title={From spatial orientation to food acquisition in echolocating bats},
+  author={Schnitzler, Hans-Ulrich and Moss, Cynthia F and Denzinger, Annette},
+  journal={Trends in Ecology \& Evolution},
+  volume={18},
+  number={8},
+  pages={386--394},
+  year={2003},
+  publisher={Elsevier}
+}
+
+@article{https://doi.org/10.1111/2041-210X.12584,
+author = {Friard, Olivier and Gamba, Marco},
+title = {BORIS: a free, versatile open-source event-logging software for video/audio coding and live observations},
+journal = {Methods in Ecology and Evolution},
+volume = {7},
+number = {11},
+pages = {1325-1330},
+keywords = {behavioural analysis, behaviour coding, coding scheme, ethology, observational data, ttime-budget},
+doi = {https://doi.org/10.1111/2041-210X.12584},
+url = {https://besjournals.onlinelibrary.wiley.com/doi/abs/10.1111/2041-210X.12584},
+eprint = {https://besjournals.onlinelibrary.wiley.com/doi/pdf/10.1111/2041-210X.12584},
+abstract = {Summary Quantitative aspects of the study of animal and human behaviour are increasingly relevant to test hypotheses and find empirical support for them. At the same time, photo and video cameras can store a large number of video recordings and are often used to monitor the subjects remotely. Researchers frequently face the need to code considerable quantities of video recordings with relatively flexible software, often constrained by species-specific options or exact settings. BORIS is a free, open-source and multiplatform standalone program that allows a user-specific coding environment to be set for a computer-based review of previously recorded videos or live observations. Being open to user-specific settings, the program allows a project-based ethogram to be defined that can then be shared with collaborators, or can be imported or modified. Projects created in BORIS can include a list of observations, and each observation may include one or two videos (e.g. simultaneous screening of visual stimuli and the subject being tested; recordings from different sides of an aquarium). Once the user has set an ethogram, including state or point events or both, coding can be performed using previously assigned keys on the computer keyboard. BORIS allows definition of an unlimited number of events (states/point events) and subjects. Once the coding process is completed, the program can extract a time-budget or single or grouped observations automatically and present an at-a-glance summary of the main behavioural features. The observation data and time-budget analysis can be exported in many common formats (TSV, CSV, ODF, XLS, SQL and JSON). The observed events can be plotted and exported in various graphic formats (SVG, PNG, JPG, TIFF, EPS and PDF).},
+year = {2016}
+}
+
+@inbook{von1965tanze,
+	address = {Berlin, Heidelberg},
+	author = {von Frisch, Karl},
+	booktitle = {Tanzsprache und Orientierung der Bienen},
+	doi = {10.1007/978-3-642-94916-6_2},
+	isbn = {978-3-642-94916-6},
+	pages = {3--330},
+	publisher = {Springer Berlin Heidelberg},
+	title = {Die T{\"a}nze der Bienen},
+	url = {https://doi.org/10.1007/978-3-642-94916-6_2},
+	year = {1965},
+	bdsk-url-1 = {https://doi.org/10.1007/978-3-642-94916-6_2}}
+
+@article{bradbury1998principles,
+  title={Principles of animal communication},
+  author={Bradbury, Jack W and Vehrencamp, Sandra Lee and others},
+  year={1998},
+  publisher={Sinauer Associates Sunderland, MA}
+}
+
+@article{thom2007scent,
+  title={The scent of the waggle dance},
+  author={Thom, Corinna and Gilley, David C and Hooper, Judith and Esch, Harald E},
+  journal={PLoS biology},
+  volume={5},
+  number={9},
+  pages={e228},
+  year={2007},
+  publisher={Public Library of Science San Francisco, USA}
+}
+
+@article{tsujiuchi2007dynamic,
+  title={Dynamic range compression in the honey bee auditory system toward waggle dance sounds},
+  author={Tsujiuchi, Seiya and Sivan-Loukianova, Elena and Eberl, Daniel F and Kitagawa, Yasuo and Kadowaki, Tatsuhiko},
+  journal={PLoS One},
+  volume={2},
+  number={2},
+  pages={e234},
+  year={2007},
+  publisher={Public Library of Science San Francisco, USA}
+}
+
+@article{david2009trail,
+  title={Trail pheromones of ants},
+  author={Morgan, David},
+  journal={Physiological entomology},
+  volume={34},
+  number={1},
+  pages={1--17},
+  year={2009},
+  publisher={Wiley Online Library}
+}
+
+@article{SCHLENKER2016894,
+title = {What Do Monkey Calls Mean?},
+journal = {Trends in Cognitive Sciences},
+volume = {20},
+number = {12},
+pages = {894-904},
+year = {2016},
+issn = {1364-6613},
+doi = {https://doi.org/10.1016/j.tics.2016.10.004},
+url = {https://www.sciencedirect.com/science/article/pii/S1364661316301711},
+author = {Philippe Schlenker and Emmanuel Chemla and Klaus Zuberbühler},
+keywords = {primate semantics, primate call evolution, primate implicatures, primate linguistics, evolution of meaning, evolution of communication},
+abstract = {A field of primate linguistics is gradually emerging. It combines general questions and tools from theoretical linguistics with rich data gathered in experimental primatology. Analyses of several monkey systems have uncovered very simple morphological and syntactic rules and have led to the development of a primate semantics that asks new questions about the division of semantic labor between the literal meaning of monkey calls, additional mechanisms of pragmatic enrichment, and the environmental context. We show that comparative studies across species may validate this program and may in some cases help in reconstructing the evolution of monkey communication over millions of years.}
+}
+
+@article{
+seyfarth1980,
+author = {Seyfarth, Robert M.  and Dorothy, L. Cheney  and Marler, Peter },
+title = {Monkey Responses to Three Different Alarm Calls: Evidence of Predator Classification and Semantic Communication},
+journal = {Science},
+volume = {210},
+number = {4471},
+pages = {801-803},
+year = {1980},
+doi = {10.1126/science.7433999},
+URL = {https://www.science.org/doi/abs/10.1126/science.7433999},
+eprint = {https://www.science.org/doi/pdf/10.1126/science.7433999},
+abstract = {Vervet monkeys give different alarm calls to different predators. Recordings of the alarms played back when predators were absent caused the monkeys to run into trees for leopard alarms, look up for eagle alarms, and look down for snake alarms. Adults call primarily to leopards, martial eagles, and pythons, but infants give leopard alarms to various mammals, eagle alarms to many birds, and snake alarms to various snakelike objects. Predator classification improves with age and experience.}}
+
+@article{hobaiter2011gestural,
+  title={The gestural repertoire of the wild chimpanzee},
+  author={Hobaiter, Catherine and Byrne, Richard W},
+  journal={Animal cognition},
+  volume={14},
+  pages={745--767},
+  year={2011},
+  publisher={Springer}
+}
+
+
+@article{anderson2010flexibility,
+  title={Flexibility in the use of requesting gestures in squirrel monkeys (Saimiri sciureus)},
+  author={Anderson, J. R. and Kuroshima, H. and Hattori, Y. and Fujita, K.},
+  journal={American Journal of Primatology},
+  volume={72},
+  number={8},
+  pages={707--714},
+  year={2010},
+  publisher={Wiley Online Library}, 
+  doi={https://doi.org/10.1002/ajp.20827}
+}
+
+@article{kroodsma1991function,
+  title={The function (s) of bird song},
+  author={Kroodsma, Donald E and Byers, Bruce E},
+  journal={American Zoologist},
+  volume={31},
+  number={2},
+  pages={318--328},
+  year={1991},
+  publisher={Oxford University Press UK}
+}
+
+@article{byers2009female,
+  title={Female mate choice and songbird song repertoires},
+  author={Byers, Bruce E and Kroodsma, Donald E},
+  journal={Animal Behaviour},
+  volume={77},
+  number={1},
+  pages={13--22},
+  year={2009},
+  publisher={Elsevier}
+}
+
+@article{heiligenberg1973electrolocation,
+  title={Electrolocation of objects in the electric fish Eigenmannia (Rhamphichthyidae, Gymnotoidei)},
+  author={Heiligenberg, Walter},
+  journal={Journal of comparative physiology},
+  volume={87},
+  number={2},
+  pages={137--164},
+  year={1973},
+  publisher={Springer}
+}
+
+@article{simmons1979echolocation,
+  title={Echolocation and pursuit of prey by bats},
+  author={Simmons, James A and Fenton, M Brock and O'Farrell, Michael J},
+  journal={Science},
+  volume={203},
+  number={4375},
+  pages={16--21},
+  year={1979},
+  publisher={American Association for the Advancement of Science}
+}
+
+@article{kamminga1988echolocation,
+  title={Echolocation signal types of odontocetes},
+  author={Kamminga, Cees},
+  journal={Animal sonar: processes and performance},
+  pages={9--22},
+  year={1988},
+  publisher={Springer}
+}
+
+@article{park2016ultrasonic,
+  title={Ultrasonic hearing and echolocation in the earliest toothed whales},
+  author={Park, Travis and Fitzgerald, Erich MG and Evans, Alistair R},
+  journal={Biology Letters},
+  volume={12},
+  number={4},
+  pages={20160060},
+  year={2016},
+  publisher={The Royal Society}
+}
+
+@article{meyer1987hormone,
+  title={Hormone-induced and maturational changes in electric organ discharges and electroreceptor tuning in the weakly electric fish Apteronotus},
+  author={Meyer, J Harlan and Leong, Margaret and Keller, Clifford H},
+  journal={Journal of Comparative Physiology A},
+  volume={160},
+  pages={385--394},
+  year={1987},
+  publisher={Springer}
+}
+
+@article{von1999active,
+  title={Active electrolocation of objects in weakly electric fish},
+  author={von der Emde, Gerhard},
+  journal={Journal of experimental biology},
+  volume={202},
+  number={10},
+  pages={1205--1215},
+  year={1999},
+  publisher={Company of Biologists The Company of Biologists, Bidder Building, 140 Cowley~…}
+}
+
+
+@article{zupanc1993evoked,
+  title={Evoked chirping in the weakly electric fish Apteronotus leptorhynchus: a quantitative biophysical analysis},
+  author={Zupanc, G{\"u}nther KH and Maler, Leonard},
+  journal={Canadian Journal of Zoology},
+  volume={71},
+  number={11},
+  pages={2301--2310},
+  year={1993},
+  publisher={NRC Research Press Ottawa, Canada}
+}
+
+@article{benda2005spike,
+  title={Spike-frequency adaptation separates transient communication signals from background oscillations},
+  author={Benda, Jan and Longtin, Andr{\'e} and Maler, Len},
+  journal={Journal of Neuroscience},
+  volume={25},
+  number={9},
+  pages={2312--2321},
+  year={2005},
+  publisher={Soc Neuroscience}
+}
+
+@article{engler2001differential,
+  title={Differential production of chirping behavior evoked by electrical stimulation of the weakly electric fish, Apteronotus leptorhynchus.},
+  author={Engler, G and Zupanc, GKH},
+  journal={Journal of Comparative Physiology A: Sensory, Neural \& Behavioral Physiology},
+  volume={187},
+  number={9},
+  year={2001}
+}
+
+@article{triefenbach2008changes,
+  title={Changes in signalling during agonistic interactions between male weakly electric knifefish, Apteronotus leptorhynchus},
+  author={Triefenbach, Frank A and Zakon, Harold H},
+  journal={Animal Behaviour},
+  volume={75},
+  number={4},
+  pages={1263--1272},
+  year={2008},
+  publisher={Elsevier}
+}
+
+@article{fugere2011electrical,
+  title={Electrical signalling of dominance in a wild population of electric fish},
+  author={Fug{\`e}re, Vincent and Ortega, Hern{\'a}n and Krahe, R{\"u}diger},
+  journal={Biology Letters},
+  volume={7},
+  number={2},
+  pages={197--200},
+  year={2011},
+  publisher={The Royal Society}
+}
+
+@article{zubizarreta2020seasonal,
+  title={Seasonal and social factors associated with spacing in a wild territorial electric fish},
+  author={Zubizarreta, Luc{\'i}a and Quintana, Laura and Hern{\'a}ndez, Daniel and Teixeira de Mello, Franco and Meerhoff, Mariana and Massaaki Honji, Renato and Guimar{\~a}es Moreira, Renata and Silva, Ana},
+  journal={Plos one},
+  volume={15},
+  number={6},
+  pages={e0228976},
+  year={2020},
+  publisher={Public Library of Science San Francisco, CA USA}
+}
+
+@phdthesis{raab2022social,
+  title={Social structures and interactions in electric fish explored by large-scale signal tracking},
+  author={Raab, Till},
+  year={2022},
+  school={Universit{\"a}t T{\"u}bingen}
+  }
+
+
+@article{seyfarth2003signalers,
+  title={Signalers and receivers in animal communication},
+  author={Seyfarth, Robert M and Cheney, Dorothy L},
+  journal={Annual review of psychology},
+  volume={54},
+  number={1},
+  pages={145--173},
+  year={2003},
+  publisher={Annual Reviews 4139 El Camino Way, PO Box 10139, Palo Alto, CA 94303-0139, USA}
+}
+
+@article{seyfarth2017origin,
+  title={The origin of meaning in animal signals},
+  author={Seyfarth, Robert M and Cheney, Dorothy L},
+  journal={Animal Behaviour},
+  volume={124},
+  pages={339--346},
+  year={2017},
+  publisher={Elsevier}
+}
\ No newline at end of file
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diff --git a/protocol/figures/timeline.pdf b/protocol/figures/timeline.pdf
new file mode 100644
index 0000000..ff3e792
Binary files /dev/null and b/protocol/figures/timeline.pdf differ
diff --git a/protocol/main.tex b/protocol/main.tex
new file mode 100644
index 0000000..f754341
--- /dev/null
+++ b/protocol/main.tex
@@ -0,0 +1,132 @@
+\documentclass[a4paper]{article}       % Default text size and document class
+
+% Format -----------------------------------------------------------------------------
+\usepackage[                    % To set up the page
+  a4paper,                        % A4 paper
+  width=160mm,                    % Text body width
+  top=25mm,bottom=25mm,           % Top and bottom margins
+  bindingoffset=6mm               % Offset of left and right pages for printing
+  ]
+ {geometry}
+\usepackage{setspace}           % So set spacings (e.g. line height)
+  \onehalfspacing                 % Sets line width to 1.5
+%\usepackage{sectsty}            % To control sectional headers
+%  \chapternumberfont{\huge}       % Set size of chapter number
+%  \chaptertitlefont{\huge}        % Set size of chapter title 
+
+\usepackage{multicol}           % Writes in multiple columns
+\usepackage{wrapfig}
+\usepackage{lipsum}             % Inserts blind text for previewing
+\usepackage{placeins}           % adding float barriers 
+\setlength{\parindent}{0pt}     % remove indentation for new paragraphs
+
+% Language ---------------------------------------------------------------------------
+\usepackage[english]{babel}     % To enable language support other than us english
+\usepackage[babel]{csquotes}    % Omits spell checking in quotations
+
+% Images, long tables & append pdf pages ---------------------------------------------
+\usepackage{float}              % Enables some options for image placement
+\usepackage{graphicx}           % To include graphics
+\usepackage{                    % Long table stuff
+  csvsimple,                      % So a csv can be converted to a table
+  longtable,                      % To make long tables
+  booktabs                        % To page break long tables properly
+  }
+\usepackage{pdfpages}           % Appends pdf pages
+\usepackage{xcolor}
+
+% Custom fancy image caption ---------------------------------------------------------
+\usepackage{caption}
+  \captionsetup[figure]{labelsep=space}
+%  \captionsetup[table]{labelsep=space}
+  \newcommand{\mycaption}[2]{\caption[#1]{\textbar\,\textbf{#1} #2}}
+
+% Math and code support -----------------------------------------------------------------------
+\usepackage{amsmath, amsfonts}  % For fancy math
+\usepackage[separate-uncertainty = true]{siunitx} % for SI units
+\sisetup{detect-all}
+\usepackage{xcolor}
+\NewDocumentCommand{\codeword}{v}{%
+\texttt{\textcolor{black}{#1}}%
+}
+% Citation setup ---------------------------------------------------------------------
+\usepackage[]{hyperref}         % To make clickable links
+  \hypersetup{hidelinks,}         % To draw no boxes around the links
+\usepackage[                    % Citation setup  
+    backend=biber,                % Citation backend
+    style=apa,           % Citation format
+    % minbibnames=1,
+    % maxbibnames=99,
+    %maxnames=2,                 % Max number of names displayed in text
+    %sortlocale=de_DE,             % Sorts by german conventions (ß = ss)
+    %natbib=true,                   % Enables \citep and \citet. Use \parencite and \textcite in future
+    url=false,                    % Disaples url
+    doi=true,                     % To print the doi
+    eprint=false                  % Disables link to eprint (?)
+]{biblatex}
+\DeclareDelimFormat[parencite]{finalnamedelim}{\addspace and\space} % disabels & and uses 'and'
+
+\DeclareDelimFormat[bib,biblist]{finalnamedelim}{\addspace and\space} % disabels & and uses 'and' in the bibfile
+
+\addbibresource{chirpdetection.bib} % Where the bibliography file is
+\renewcommand{\familydefault}{\sfdefault}
+\usepackage{tgheros}
+\usepackage{wrapfig}
+%\setlength{\marginparwidth}{1.8cm}
+\usepackage[disable]{todonotes}
+%\DeclareUnicodeCharacter{0301}{\textcolor{red}{*************************************}} % for finding unicode erros in the pdf 
+
+
+\setlength\bibitemsep{2\itemsep} 
+
+% Abstract title same size as sectional headers --------------------------------------
+% see here: https://tex.stackexchange.com/questions/366169/how-to-change-font-size-for-abstract-title
+\makeatletter
+\renewenvironment{abstract}{%
+    \if@twocolumn
+      \section*{\abstractname}%
+    \else %% <- here I've removed \small
+      \begin{center}%
+        {\bfseries \Large\abstractname\vspace{\z@}}%  %% <- here I've added \Large
+      \end{center}%
+      \quotation
+    \fi}
+    {\if@twocolumn\else\endquotation\fi}
+\makeatother
+
+% Document ---------------------------------------------------------------------------
+\begin{document}
+\input{chapters/titlepage.tex} % Add the path to the titlepage here
+\listoftodos
+\section{Introduction}
+   \label{chap:introduction}
+   \input{chapters/introduction.tex}
+
+\pagebreak
+\section{Methods}
+  \label{chap:methods}
+  \input{chapters/methods.tex}
+
+\pagebreak
+\section{Results}
+  \label{chap:results}
+  \input{chapters/results.tex}
+
+\pagebreak
+\section{Discussion}
+\label{chap:discussion}
+\input{chapters/discussion.tex}
+
+\pagebreak
+\printbibliography 
+%\pagebreak
+%\section{Appendix}
+%  \label{chap:appendix}
+%  \input{chapters/appendix.tex}
+
+% Append pdf outputs from other programs to the document
+% \includepdf[pages=-]{appendix/ab.pdf}
+% \includepdf[pages=-]{appendix/cd.pdf}
+% \includepdf[pages=-]{appendix/ef.pdf}
+
+\end{document}
\ No newline at end of file
diff --git a/requirements.txt b/requirements.txt
index 72cac28..ad47b80 100644
--- a/requirements.txt
+++ b/requirements.txt
@@ -1,7 +1,7 @@
-audioio==0.9.5
+audioio==0.10.0
 cmocean==2.0
 cycler==0.11.0
-ipython==8.10.0
+ipython==8.12.0
 matplotlib==3.7.0
 numpy==1.23.5
 pandas==1.5.3
@@ -9,5 +9,5 @@ paramiko==2.11.1
 PyYAML==6.0
 scipy==1.10.1
 scp==0.14.5
-thunderfish==1.9.9
+thunderfish==1.9.10
 tqdm==4.64.1