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\documentclass[11pt]{article}
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\usepackage[utf8]{inputenc}
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\usepackage{textcomp}
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\usepackage{xcolor}
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\usepackage{graphicx}
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\usepackage[ngerman,english]{babel}
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\usepackage[left=25mm, right=25mm, top=20mm, bottom=20mm]{geometry}
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\setlength{\parskip}{2ex}
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\usepackage[mediumqspace,Gray,squaren]{SIunits} % \ohm, \micro
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\usepackage{natbib}
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%\bibliographystyle{jneurosci}
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\usepackage[breaklinks=true,bookmarks=true,bookmarksopen=true,pdfpagemode=UseNone,pdfstartview=FitH,colorlinks=true,citecolor=blue,urlcolor=blue]{hyperref}
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\setlength{\parindent}{0em}
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\begin{document}
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\hspace*{\fill} September 24, 2025
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\noindent Dear Arvind Kumar,
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we would like to submit our manuscript ``Spike generation in
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electroreceptor afferents introduces additional spectral response
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components by weakly nonlinear interactions'' for publication in
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eNeuro. It is the result of a collaborative effort on the theoretical
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side, Benjamin Linder, HU Berlin, and the experimental and numerical
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modeling side, Jan Grewe and myself at the University of Tuebingen,
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within the DFG priority program 2205 ``Evolutionary optimization of
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neuronal processing''.
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Computational neuroscientists have a strong interest in characterizing
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non-linearities and study their functional consequences. However,
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experimental backing of these theoretical findings are scarce. For
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example, encoding of dynamic stimuli in spike trains often can be well
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approximated by linear response theory, in particular when driving the
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neuron in the supra-threshold regime at low signal-to-noise
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ratios. This has been exploited in numerous theoretical studies. At
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less noise or stronger stimulus amplitudes non-linear effects become
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more prominent. In the weakly non-linear regime, the second term of
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the Volterra series becomes relevant. Benjamin Lindner developed
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analytical solutions of this term for the leaky integrate-and-fire
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neuron, which predicts non-linear interactions whenever one or the sum
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of two stimulus frequencies matches the neuron's baseline firing
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rate. Until now, however, these fundamental non-linearities arising
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from the core mechanism of spike generation have not been reported in
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any experimental data.
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With our work we set out to fill this gap. We scan a large set of
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electrophysiological data measured in two types of electrosensory
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neurons of the electric fish \textit{Apteronotus leptorhynchus} for
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signatures of these non-linearities. In ampullary cells, non-linear
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interaction between two stimulus frequencies are prominent, whereas in
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P-units they are harder to find. Estimating the second-order
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susceptibilites from real data turns out to be a hard problem as
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limited data leads to poor estimates. Comparison with models that have
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been fitted to individual P-units, we are able to deduce the presence
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of non-linear interactions also in some of the P-units. Finally, we
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discuss our findings and the relevance of non-linear interactions in
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the neuroethological context.
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We believe that this analysis of electrophysiological data close to
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expectations from theoretical work is of strong interest to many
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readers of eNeuro and can inspire future research in other sensory
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systems.
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Best regards,\\%[-2ex]
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%\hspace*{0.17\textwidth}\includegraphics[width=0.3\textwidth]{JanBenda-Signature2020}\\
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Prof. Dr. Jan Benda, on behalf of all authors
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\end{document}
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