added Caputis papers
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@ -315,7 +315,7 @@ Respiration was then switched to normal tank water and the fish was transferred
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For the recordings fish were positioned centrally in the experimental tank, with the major parts of their body submerged into the water. Those body parts that were above the water surface were covered with paper tissue to avoid drying of the skin. Local analgesia was refreshed in intervals of two hours by cutaneous application of Lidocaine (2\,\%; bela-pharm, Vechta, Germany) around the surgical wounds. Electrodes (borosilicate; 1.5\,mm outer diameter; GB150F-8P; Science Products, Hofheim, Germany) were pulled to a resistance of 50--100\,\mega\ohm{} (model P-97; Sutter Instrument, Novato, CA) and filled with 1\,M KCl solution. Electrodes were fixed in a microdrive (Luigs-Neumann, Ratingen, Germany) and lowered into the nerve. Recordings of electroreceptor afferents were amplified and lowpass filtered at 10\,kHz (SEC-05, npi-electronics, Tamm, Germany, operated in bridge mode). All signals, neuronal recordings, recorded EOD, and the generated stimulus, were digitized with sampling rates of 20 or 40\,kHz (PCI-6229, National Instruments, Austin, TX). RELACS (\url{https://github.com/relacs/relacs}) running on a Linux computer was used for online spike and EOD detection, stimulus generation, and calibration. Recorded data was then stored on the hard drive for offline analysis.
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\subsection{Identification of P-units and ampullary cells}
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Recordings were classified as P-units if baseline action potentials phase locked to the EOD with vectors strengths between 0.7 and 0.95, a baseline firing rate larger than 30\,Hz, a serial correlation of subsequent interspike intervals below zero, a coefficient of variation of baseline interspike intervals below 1.5 und during stimulation below 2. As ampullary cells we classified recordings with vector strengths below 0.15, baseline firing rate above 10\,Hz, baseline CV below 0.18, CV during stimulation below 1.0, and a response modulation during stimulation below 80\,Hz \citep{Grewe2017}. We here selected only those cells of which the neuron's baseline activity as well as the responses to band-limited white noise stimuli were recorded.
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Recordings were classified as P-units if baseline action potentials phase locked to the EOD with vectors strengths between 0.7 and 0.95, a baseline firing rate larger than 30\,Hz, a serial correlation of subsequent interspike intervals below zero, a coefficient of variation of baseline interspike intervals below 1.5 und during stimulation below 2. P-units are clearly distinguised from T-type electrorecptors, that we did not analyze here, by having firing rates much lower than the EOD frequency of the fish (no 1:1 locking to the EOD) \notejb{CITE TUNIT PAPER}. As ampullary cells we classified recordings with vector strengths below 0.15, baseline firing rate above 10\,Hz, baseline CV below 0.18, CV during stimulation below 1.0, and a response modulation during stimulation below 80\,Hz \citep{Grewe2017}. We here selected only those cells of which the neuron's baseline activity as well as the responses to band-limited white noise stimuli were recorded.
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\subsection{Electric field recordings}
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For monitoring the EOD without the stimulus, two vertical carbon rods ($11\,\centi\meter$ long, 8\,mm diameter) in a head-tail configuration were placed isopotential to the stimulus. Their signal was differentially amplified with a gain factor between 100 and 500 (depending on the recorded animal) and band-pass filtered (3 to 1500\,Hz pass-band, DPA2-FX; npi electronics, Tamm, Germany). For an estimate of the transdermal potential that drives the electroreceptors, two silver wires spaced by 1\,cm were located next to the left gill of the fish and orthogonal to the fish's longitudinal body axis (amplification 100 to 500 times, band-pass filtered with 3 to 1\,500\,Hz pass-band, DPA2-FX; npi-electronics, Tamm, Germany). This local EOD measurement recorded the combination of the fish's own EOD and the applied stimulus.
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@ -434,12 +434,14 @@ First, the input $y(t)$ is thresholded by setting negative values to zero:
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\label{eq:threshold2}
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\lfloor y(t) \rfloor_0 = \left\{ \begin{array}{rcl} y(t) & ; & y(t) \ge 0 \\ 0 & ; & y(t) < 0 \end{array} \right.
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\end{equation}
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(\subfigrefb{flowchart}{A}). This thresholds models the transfer function of the synapses between the primary receptor cells and the afferent. Together with a low-pass filter
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(\subfigrefb{flowchart}{A}). This threshold models the transfer function of the synapses between the primary receptor cells and the afferent \notejb{CITE A MOSER PAPER} and may also include nonlinear properties introduced by low-threshold Kv1 channels known to be present in the afference \citep{TroySmith2006,Nogueira2013}. Together with a low-pass filter
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\begin{equation}
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\label{eq:dendrite}
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\tau_{d} \frac{d V_{d}}{d t} = -V_{d}+ \lfloor y(t) \rfloor_{0}
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\end{equation}
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the threshold operation is required for extracting the amplitude modulation from the input \citep{Barayeu2024}. The low-pass filter models passive signal conduction in the afferent's dendrite (\subfigrefb{flowchart}{B}) and $\tau_{d}$ is the membrane time constant of the dendrite. Dendritic low-pass filtering was also necessary to reproduce the loose coupling of P-unit spikes to the EOD while maintaining high sensitivity at small amplitude modulations.
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the threshold operation is required for extracting the amplitude modulation from the input \citep{Barayeu2024}. As detailed in the discussion, the details of the threshold nonlinearity \eqnref{eq:threshold2} and the low-pass filter \eqnref{eq:dendrite} do not matter here, since we compute all cross-spectra between the spiking response and the amplitude-modulation $s(t)$ --- and not the full input signal $y(t)$, \eqnref{eq:ram_equation}.
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The low-pass filter models passive signal conduction in the afferent's dendrite (\subfigrefb{flowchart}{B}) and $\tau_{d}$ is the membrane time constant of the dendrite. Dendritic low-pass filtering was also necessary to reproduce the loose coupling of P-unit spikes to the EOD while maintaining high sensitivity at small amplitude modulations.
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The dendritic voltage $V_d(t)$ is then fed into a stochastic leaky integrate-and-fire (LIF) model with adaptation,
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\begin{equation}
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@ -76,7 +76,16 @@
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strengthen the links between well-written theoretical analysis and
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the practical field of experimental physiology.}
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\response{READ PAPERS AND CITE THEM}
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\response{READ PAPERS AND CITE THEM You are right, there are more
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nonlinear mechanisms potentially contributiong to the threshold
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nonlinearity. We now mention this in the methods when introducing
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the threshold nonlinearity (after eq. 13) and cite the corresponding
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manuscripts. We also added a paragraph to the methods reiterating
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what we have in the discusssion, that the details of this
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nonlinearity are not so important in the context of the present
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manuscript, since we compute all cross-spectra between the resulting
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amplitude modulation and the spikes responses. MAYBE ALSO IN THE
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CONCLUSION.}
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\issue{Second, and along the same lines, the discussion could be
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improved by mentioning the effects and significance of these
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@ -85,7 +94,7 @@
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interference avoidance responses, and b) transient changes, as in
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chirps.}
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\response{THINK ABOUT IT AND ADD TO DISCUSSION}
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\response{HERE: STATIONARY ANALYSIS, CHIRPS ARE TRANSIENT, NEEDS RESEARCH. JAR SIGNALS are slow but change continously so they might hit the baseline frequency Add something somewhere around line 660.}
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\issue{Finally, the precise description of the methods could be
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expanded for reaching a broader biology audience; in particular, the
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@ -109,7 +118,11 @@
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between the extreme profiles of P (signal amplitude) and T (signal
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slope)?}
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\response{SAY SOMETHING ABOUT T-UNITS AND THAT WE DEFINITELY EXCLUDED THEM}
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\response{In \textit{Apteronotus} T-units are characterized by 1:1
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locking to the EOD, i.e. by having a baseline firing rate matching
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the EOD frequency. We definitely have no T-units in our data
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set. This we eplain now in the ``Identification of P-units and
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ampullary cells'' section in the methods.}
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\issue{In line 147, rather than using the term
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"laterally," I believe it would enhance clarity to state "parallel
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@ -124,7 +137,7 @@
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field foveal perioral region where the majority of receptors are
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located.}
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\response{ADD SOMETHING TO STIMULATION SECTION}
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\response{ADD SOMETHING TO STIMULATION SECTION LINE 110}
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\issue{Line 148, the phrase "band limited white noise" lacks
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clarity. Upon my initial reading, I assumed that the cutoff limit
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@ -7132,7 +7132,7 @@ microelectrode recordings from visual cortex and functional implications.},
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year={2014},
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}
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@article{Smith2006,
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@article{TroySmith2006,
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title={Distribution of {Kv1}-like potassium channels in the electromotor and electrosensory systems of the weakly electric fish {\textit{Apteronotus leptorhynchus}}.},
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author={G. Troy Smith and Graciela A. Unguez and Christopher M. Weber},
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journal={J Neurobiol},
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