Formalizing stuff (WIP).

This commit is contained in:
j-hartling 2025-11-06 09:27:17 +01:00
parent 48f3c8cd41
commit d9fb7d3e5d
12 changed files with 3195 additions and 80 deletions

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@article{clemens2011,
author = "Jan Clemens, Olaf Kutzki; Bernd Ronacher; Susanne Schreiber; Sandra Wohlgemuth",
journal = "PNAS",
volume = "108",
number = "33",
year = "2011",
DOI = "https://doi.org/10.1073/pnas.1104506108",
}

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\documentclass[a4paper, 12pt]{article} \documentclass[a4paper, 12pt]{article}
\usepackage{parskip} \usepackage{parskip}
\usepackage{amsmath}
\usepackage[
backend=biber,
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\title{Physiologically inspired model of the grasshopper auditory system} \title{Emergent intensity invariance in a physiologically inspired model of the grasshopper auditory system}
\author{Jona Hartling, Jan Benda} \author{Jona Hartling, Jan Benda}
\date{} \date{}
\begin{document} \begin{document}
\maketitle{} \maketitle{}
\newcommand{\bp}{h_{\text{BP}}(t)} % Bandpass filter function
\newcommand{\lp}{h_{\text{LP}}(t)} % Lowpass filter function
\newcommand{\hp}{h_{\text{HP}}(t)} % Highpass filter function
\newcommand{\fc}{f_{\text{cut}}} % Filter cutoff frequency
\newcommand{\infint}{\int_{-\infty}^{\infty}} % Indefinite integral
\newcommand{\bi}{b_\theta}
\newcommand{\feat}{f_\theta}
\section{The sensory world of a grasshopper} \section{The sensory world of a grasshopper}
Strong dependence on acoustic signals for ranged communication\\ Strong dependence on acoustic signals for ranged communication\\
@ -53,39 +67,108 @@ How can a human observer conceive a grasshopper's auditory percepts?\\
- How to integrate the available knowledge on anatomy, physiology, ethology?\\ - How to integrate the available knowledge on anatomy, physiology, ethology?\\
$\rightarrow$ Abstract, simplify, formalize $\rightarrow$ Functional model framework $\rightarrow$ Abstract, simplify, formalize $\rightarrow$ Functional model framework
\section{Pre-split pathway: Population pre-processing}
\section{Developing a functional model of\\the grasshopper auditory pathway}
\subsection{Population-driven signal pre-processing}
"Pre-split portion" of the auditory pathway:\\
Tympanal membrane $\rightarrow$ Receptor neurons $\rightarrow$ Local interneurons
Similar response/filter properties within receptor/interneuron populations (\cite{clemens2011})\\
$\rightarrow$ One population-wide response trace per stage (no "single-cell resolution")
\textbf{Stage-specific processing steps and functional approximations:}
Initial: Continuous acoustic input signal $x(t)$
Filtering of behaviorally relevant frequencies by tympanal membrane\\ Filtering of behaviorally relevant frequencies by tympanal membrane\\
- Bandpass 5-30 kHz $\rightarrow$ Bandpass filter 5-30 kHz
\begin{equation}
x(t)\,*\,\bp; \quad\quad \fc\,=\,5\,\text{kHz},\,30\,\text{kHz}
\end{equation}
Extraction of signal envelope (AM encoding) by receptor population\\ Extraction of signal envelope (AM encoding) by receptor population\\
- Full-wave rectification + lowpass 500 Hz $\rightarrow$ Full-wave rectification, then lowpass filter 500 Hz
\begin{equation}
|x(t)|\,*\,\lp; \quad\quad \fc\,=\,500\,\text{Hz}
\end{equation}
Logarithmically compressed intensity tuning curve of receptors\\ Logarithmically compressed intensity tuning curve of receptors\\
- Decibel transformation $\rightarrow$ Decibel transformation
\begin{equation}
20\,\cdot\,\log_{10} \frac{x(t)}{x_{\text{max}}}
\end{equation}
Spike-frequency adaptation in receptor and interneuron populations\\ Spike-frequency adaptation in receptor and interneuron populations\\
- Highpass 10 Hz $\rightarrow$ Highpass filter 10 Hz
\begin{equation}
x(t)\,*\,\hp; \quad\quad \fc\,=\,10\,\text{Hz}
\end{equation}
\section{Post-split pathway: Feature extraction}
Template matching by individual ascending neurons\\ \subsection{Feature extraction by individual neurons}
- Separate convolution with each of a set of Gabor kernels\\
- Pathway splitting: Single population response into several separate branches "Post-split portion" of the auditory pathway:\\
- Expansion into a higher-dimensional sound representation Ascending neurons (AN) $\rightarrow$ Central brain neurons
Diverse response/filter properties within AN population (\cite{clemens2011})\\
- Pathway splitting into several parallel branches\\
- Expansion into a decorrelated higher-dimensional sound representation\\
$\rightarrow$ Individual neuron-specific response traces from this stage onwards
\textbf{Stage-specific processing steps and functional approximations:}
Template matching by individual ANs\\
- Filter base (STA approximations): Set of Gabor kernels\\
- Gabor parameters: $\sigma, \phi, f$ $\rightarrow$ Determines kernel sign and lobe number
\begin{equation}
k(t)\,=\,e^{-\frac{t^{2}}{2\sigma^{2}}}\,\cdot\,\sin(2\pi f t\,+\,\phi)
\end{equation}
$\rightarrow$ Separate convolution with each member of the kernel set
\begin{equation}
c_i(t)\,=\,x(t)\,*\,k_i(t)
= \infint x(\tau)\,\cdot\,k_i(t\,-\,\tau)\,d\tau
\end{equation}
Thresholding nonlinearity in ascending neurons (or further downstream)\\ Thresholding nonlinearity in ascending neurons (or further downstream)\\
- Step-function (or sigmoid) threshold\\ - Binarization of AN response traces into "relevant" vs. "irrelevant"\\
- Binarization of response values into "relevant" vs. "irrelevant" $\rightarrow$ Heaviside step-function $H(c\,-\,\theta)$ (or steep sigmoid threshold?)
\begin{equation}
\bi(t)\,=\,\begin{cases}
\;1, \quad c(t)\,\geq\,\theta\\
\;0, \quad c(t)\,<\,\theta
\end{cases}
\end{equation}
Temporal averaging by neurons of the central brain\\ Temporal averaging by neurons of the central brain\\
- Lowpass 1 Hz\\ - Finalized set of slowly changing kernel-specific features (one per AN)\\
- Finalized set of slowly changing kernel-specific features\\ - Different species-specific song patterns are characterized by a distinct combination
- Different (species-specific) songs are characterized by a distinct combination of feature values of feature values $\rightarrow$ Clusters in high-dimensional feature space\\
$\rightarrow$ Lowpass filter 1 Hz
\begin{equation}
\feat(t)\,=\,\bi(t)\,*\,\lp; \quad\quad \fc\,=\,1\,\text{Hz}
\end{equation}
\section{Pre-split intensity invariance:\\Logarithm-highpass mechanism}
\section{Post-split intensity invariance:\\Threshold-lowpass mechanism} \section{Two mechanisms driving the emergence of intensity-invariant song representation}
\section{Conclusion and outlook}
\subsection{Logarithmic scaling \& spike-frequency adaptation}
Song signal $s(t)$ with variable scale $\alpha$ and fixed-scale additive noise $\eta(t)$
\begin{equation}
\alpha\,\cdot\,s(t)\,+\,\eta(t)
\end{equation}
\subsection{Threshold nonlinearity \& temporal averaging}
\section{Discriminating species-specific song\\patterns in feature space}
\section{Conclusions \& outlook}
\end{document} \end{document}