diff --git a/cite.bib b/cite.bib index bfc05a6..1763485 100644 --- a/cite.bib +++ b/cite.bib @@ -100,6 +100,16 @@ pages={2575--2580}, year={2009}, } +@article{eichendorf1980projections, + title={Projections of auditory ventral-cord neurons in the supraesophageal ganglion of Locusta migratoria}, + author={Eichendorf, Annette and Kalmring, Klaus}, + journal={Zoomorphologie}, + volume={94}, + number={2}, + pages={133--149}, + year={1980}, + publisher={Springer} +} @article{fisch2012channel, title={Channel noise from both slow adaptation currents and fast currents is required to explain spike-response variability in a sensory neuron}, author={Fisch, Karin and Schwalger, Tilo and Lindner, Benjamin and Herz, Andreas VM and Benda, Jan}, @@ -151,6 +161,37 @@ pages={433--450}, year={1996}, } +@article{kalmring1975afferent, + title={The afferent auditory pathway in the ventral cord of Locusta migratoria (Acrididae) I. Synaptic connectivity and information processing among the auditory neurons of the ventral cord}, + author={Kalmring, Klaus}, + journal={Journal of comparative physiology}, + volume={104}, + number={2}, + pages={103--141}, + year={1975}, + publisher={Springer} +} +@article{kalmring1972akustische, + title={Akustische Neuronen im Bauchmark der Wanderheuschrecke Locusta migratoria}, + author={Kalmring, Klaus and Rheinlaender, J{\"u}rgen and Rehbein, Hansgeorg}, + journal={Zeitschrift f{\"u}r vergleichende Physiologie}, + volume={76}, + number={3}, + pages={314--332}, + year={1972}, + publisher={Springer} +} +@article{kalmring1978coding, + title={The coding of sound signals in the ventral-cord auditory system of the migratory locust, Locusta migratoria}, + author={Kalmring, Klaus and K{\"u}hne, Roland and Moysich, Fritz}, + journal={Journal of comparative physiology}, + volume={128}, + number={3}, + pages={213--226}, + year={1978}, + publisher={Springer} +} + @article{lang2000acoustic, title={Acoustic communication distances of a gomphocerine grasshopper}, author={Lang, Friederike}, @@ -211,6 +252,17 @@ pages={1965--1974}, year={2008}, } +@article{rehbein1974structure, + title={Structure and function of acoustic neurons in the thoracic ventral nerve cord of Locusta migratoria (Acrididae)}, + author={Rehbein, Hansgeorg and Kalmring, Klaus and R{\"o}mer, Heiner}, + journal={Journal of comparative physiology}, + volume={95}, + number={3}, + pages={263--280}, + year={1974}, + publisher={Springer} +}# Cited + @article{rehbein1976auditory, title={Auditory neurons in the ventral cord of the locust: Morphological and functional properties}, author={Rehbein, Hansgeorg}, @@ -218,7 +270,8 @@ volume={110}, pages={233--250}, year={1976}, -} +}# Cited + @article{romer1985responses, title={Responses to model songs of auditory neurons in the thoracic ganglia and brain of the locust}, author={R{\"o}mer, Heiner and Seikowski, Ulrich}, @@ -265,6 +318,16 @@ pages={154--175}, year={2023}, } +@article{sippel1983nonlinear, + title={Non-linear analysis of the transmission of signals in the auditory system of the migratory locust Locusta migratorta}, + author={Sippel, Martin and Breckow, Joachim}, + journal={Biological Cybernetics}, + volume={46}, + number={3}, + pages={197--205}, + year={1983}, + publisher={Springer} +} @article{stumpner1994song, title={{Song production and song recognition in a group of sibling grasshopper species (Chorthippus dorsatus, Ch. dichrous and Ch. loratus: Orthoptera, Acrididae)}}, author={Stumpner, Andreas and von Helversen, Otto}, diff --git a/literature/Eichendorf_Kalmring_1980.pdf b/literature/Eichendorf_Kalmring_1980.pdf new file mode 100644 index 0000000..7d1cbd0 Binary files /dev/null and b/literature/Eichendorf_Kalmring_1980.pdf differ diff --git a/literature/Kalmring_1975.pdf b/literature/Kalmring_1975.pdf new file mode 100644 index 0000000..0028784 Binary files /dev/null and b/literature/Kalmring_1975.pdf differ diff --git a/literature/Kalmring_Kühne_Moysich_1978.pdf b/literature/Kalmring_Kühne_Moysich_1978.pdf new file mode 100644 index 0000000..45dd323 Binary files /dev/null and b/literature/Kalmring_Kühne_Moysich_1978.pdf differ diff --git a/literature/Kalmring_Rheinlaender_Rehbein_1972.pdf b/literature/Kalmring_Rheinlaender_Rehbein_1972.pdf new file mode 100644 index 0000000..768a059 Binary files /dev/null and b/literature/Kalmring_Rheinlaender_Rehbein_1972.pdf differ diff --git a/literature/Rehbein_Kalmring_Römer_1974.pdf b/literature/Rehbein_Kalmring_Römer_1974.pdf new file mode 100644 index 0000000..63bec94 Binary files /dev/null and b/literature/Rehbein_Kalmring_Römer_1974.pdf differ diff --git a/literature/Sippel_Breckow_1983.pdf b/literature/Sippel_Breckow_1983.pdf new file mode 100644 index 0000000..82f4213 Binary files /dev/null and b/literature/Sippel_Breckow_1983.pdf differ diff --git a/main.aux b/main.aux index 9ab6ae2..4448817 100644 --- a/main.aux +++ b/main.aux @@ -8,28 +8,34 @@ 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comparative physiology} + \field{number}{3} + \field{title}{Structure and function of acoustic neurons in the thoracic ventral nerve cord of Locusta migratoria (Acrididae)} + \field{volume}{95} + \field{year}{1974} + \field{pages}{263\bibrangedash 280} + \range{pages}{18} + \endentry \entry{ronacher2015computational}{article}{} \name{author}{3}{}{% {{un=0,uniquepart=base,hash=4ce9349f95ced7a4831a5efa0fe5b00f}{% diff --git a/main.bcf b/main.bcf index 644ec9c..0ec1a3f 100644 --- a/main.bcf +++ b/main.bcf @@ -2365,18 +2365,20 @@ clemens2013computational clemens2013feature ronacher2015computational - michelsen1971frequency - windmill2008time - malkin2014energy - machens2001discrimination - machens2001representation - suga1960peripheral - gollisch2002energy - hildebrandt2009origin - clemens2010intensity - fisch2012channel - clemens2011efficient - clemens2011efficient + rehbein1974structure + rehbein1976auditory + clemens2011efficient + clemens2011efficient + michelsen1971frequency + windmill2008time + malkin2014energy + machens2001discrimination + machens2001representation + suga1960peripheral + gollisch2002energy + hildebrandt2009origin + clemens2010intensity + fisch2012channel diff --git a/main.blg b/main.blg index 66c8e12..5613353 100644 --- a/main.blg +++ b/main.blg @@ -1,25 +1,27 @@ [0] Config.pm:307> INFO - This is Biber 2.19 [0] Config.pm:310> INFO - Logfile is 'main.blg' -[70] biber:340> INFO - === Di Dez 9, 2025, 15:29:26 -[82] Biber.pm:419> INFO - Reading 'main.bcf' -[134] Biber.pm:979> INFO - Found 14 citekeys in bib section 0 -[145] Biber.pm:4419> INFO - Processing section 0 -[153] Biber.pm:4610> INFO - Looking for bibtex file 'cite.bib' for section 0 -[154] bibtex.pm:1713> INFO - LaTeX decoding ... -[183] bibtex.pm:1519> INFO - Found BibTeX data source 'cite.bib' -[296] UCollate.pm:68> INFO - Overriding locale 'en-US' defaults 'normalization = NFD' with 'normalization = prenormalized' -[296] UCollate.pm:68> INFO - Overriding locale 'en-US' defaults 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PDF statistics: 1083 PDF objects out of 1200 (max. 8388607) 687 compressed objects within 7 object streams diff --git a/main.pdf b/main.pdf index 8407deb..2fc32d5 100644 Binary files a/main.pdf and b/main.pdf differ diff --git a/main.synctex.gz b/main.synctex.gz index c7e2ce5..3050b7d 100644 Binary files a/main.synctex.gz and b/main.synctex.gz differ diff --git a/main.tex b/main.tex index 0f32f7c..0d6fe8f 100644 --- a/main.tex +++ b/main.tex @@ -134,6 +134,25 @@ $\rightarrow$ More general, simpler, unfitted formalized Gabor filter bank \section{Developing a functional model of\\the grasshopper auditory pathway} +The auditory pathway of grasshoppers comprises a number of neuronal stages and +identified cell types~(\bcite{rehbein1974structure}; \bcite{rehbein1976auditory}). + +Along the auditory pathway of grasshoppers (Fig.\,\ref{fig:pathway}a+b), + +1) "Pre-split portion" of the auditory pathway:\\ +Tympanal membrane $\rightarrow$ Receptor neurons $\rightarrow$ Local interneurons + +Similar response/filter properties within receptor/interneuron populations (\cite{clemens2011efficient})\\ +$\rightarrow$ One population-wide response trace per stage (no "single-cell resolution") + +2) "Post-split portion" of the auditory pathway:\\ +Ascending neurons (AN) $\rightarrow$ Central brain neurons + +Diverse response/filter properties within AN population (\cite{clemens2011efficient})\\ +- Pathway splitting into several parallel branches\\ +- Expansion into a decorrelated higher-dimensional sound representation\\ +$\rightarrow$ Individual neuron-specific response traces from this stage onwards + \begin{figure}[!ht] \centering \def\svgwidth{\textwidth} @@ -148,9 +167,9 @@ $\rightarrow$ More general, simpler, unfitted formalized Gabor filter bank Grasshoppers receive airborne sound waves by a tympanal organ at each side of the thorax~(Fig.\,\ref{fig:pathway}a). The tympanal membrane acts as a -mechanical resonance filter, that focuses vibrations of specific frequencies on -different membrane areas while attenuating -others~(\bcite{michelsen1971frequency}; \bcite{windmill2008time}; +mechanical resonance filter: Vibrations that fall within specific frequency +bands are focused on different membrane areas, while others are +attenuated~(\bcite{michelsen1971frequency}; \bcite{windmill2008time}; \bcite{malkin2014energy}). This processing step can be approximated by an initial bandpass filter \begin{equation} @@ -158,10 +177,10 @@ initial bandpass filter \label{eq:bandpass} \end{equation} applied to the acoustic input signal $\raw(t)$. The auditory receptor neurons -connect directly to the tympanal membrane. Besides performing the -mechano-electrical transduction, the receptor population further is substrate -to several known processing steps. First, the receptors extract the signal -envelope~(\bcite{machens2001discrimination}), which likely involves a +connect directly to the tympanal membrane~(Fig.\,\ref{fig:pathway}a). Besides +performing the mechano-electrical transduction, the receptor population is +substrate to several known processing steps. First, the receptors extract the +signal envelope~(\bcite{machens2001discrimination}), which likely involves a rectifying nonlinearity~(\bcite{machens2001representation}). This can be modelled as full-wave rectification followed by lowpass filtering \begin{equation} @@ -177,52 +196,25 @@ logarithmic compression is achieved by conversion to decibel scale \label{eq:log} \end{equation} relative to the maximum intensity $\dbref$ of the signal envelope $\env(t)$. -The axons of the receptor neurons project into the metathoracic ganglion, where -they synapse onto local interneurons~(Fig.\,\ref{fig:pathway}b). Both the local -interneurons~(\bcite{hildebrandt2009origin}; \bcite{clemens2010intensity}) and, -to a lesser extent, the receptors themselves~(\bcite{fisch2012channel}) display -spike-frequency adaptation in response to sustained stimulation. -This behavior is crucial to render subsequent signal representations invariant -to variations in sound intensity. - - -"Pre-split portion" of the auditory pathway:\\ -Tympanal membrane $\rightarrow$ Receptor neurons $\rightarrow$ Local interneurons - -Similar response/filter properties within receptor/interneuron populations (\cite{clemens2011efficient})\\ -$\rightarrow$ 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\\ -$\rightarrow$ Bandpass filter 5-30 kHz - -Extraction of signal envelope (AM encoding) by receptor population\\ -$\rightarrow$ Full-wave rectification, then lowpass filter 500 Hz - -Logarithmically compressed intensity tuning curve of receptors\\ -$\rightarrow$ Decibel transformation - -Spike-frequency adaptation in receptor and interneuron populations\\ -$\rightarrow$ Highpass filter 10 Hz -% +Next, the axons of the receptor neurons project into the metathoracic ganglion, +where they synapse onto local interneurons~(Fig.\,\ref{fig:pathway}b). Both the +local interneurons~(\bcite{hildebrandt2009origin}; +\bcite{clemens2010intensity}) and, to a lesser extent, the receptors +themselves~(\bcite{fisch2012channel}) display spike-frequency adaptation in +response to sustained stimulus intensity levels. This mechanism allows for the +robust encoding of faster amplitude modulations against a slowly changing +overall baseline intensity. Functionally, this processing step resembles a +highpass filter \begin{equation} \adapt(t)\,=\,\db(t)\,*\,\hp, \qquad \fc\,=\,10\,\text{Hz} \label{eq:highpass} \end{equation} -% +over the logarithmically scaled envelope $\db(t)$. The projections of the local +interneurons remain within the metathoracic ganglion and synapse onto a small +number of ascending neurons~(Fig.\,\ref{fig:pathway}b). + \subsection{Feature extraction by individual neurons} -"Post-split portion" of the auditory pathway:\\ -Ascending neurons (AN) $\rightarrow$ Central brain neurons - -Diverse response/filter properties within AN population (\cite{clemens2011efficient})\\ -- 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\\ @@ -267,6 +259,20 @@ $\rightarrow$ Lowpass filter 1 Hz % \section{Two mechanisms driving the emergence of intensity-invariant song representation} +\textbf{Definition of invariance (general, systemic):}\\ +Invariance = Property of a system to maintain a stable output with respect to a +set of relevant input parameters (variation to be represented) but irrespective +of one or more other parameters (variation to be discarded) +$\rightarrow$ Selective input-output decorrelation + +\textbf{Definition of intensity invariance (context of neurons and songs):}\\ +Intensity invariance = Time scale-selective sensitivity to certain faster +amplitude dynamics (song waveform, small-scale AM) and simultaneous +insensitivity to slower, more sustained amplitude dynamics (transient baseline, +large-scale AM, current overall intensity level)\\ +$\rightarrow$ Without time scale selectivity, any fully intensity-invariant +output will be a flat line + \subsection{Logarithmic scaling \& spike-frequency adaptation} Envelope $\env(t)$ $\xrightarrow{\text{dB}}$ Logarithmic $\db(t)$ $\xrightarrow{\hp}$ Adapted $\adapt(t)$