Added MA literature selection and grasshopper sketch SVGs.

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14 changed files with 4302 additions and 99 deletions

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cite.bib
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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",
@article{balakrishnan2001song,
title={{Song pattern recognition in the grasshopper Chorthippus biguttulus: The mechanism of syllable onset and offset detection}},
author={Balakrishnan, Rohini and von Helversen, Dagmar and von Helversen, Otto},
journal={J Comp Physiol A},
volume={187},
pages={255--264},
year={2001},
}
@article{bauer1987separate,
title={Separate localization of sound recognizing and sound producing neural mechanisms in a grasshopper},
author={Bauer, Maria and von Helversen, Otto},
journal={J Comp Physiol A},
volume={161},
pages={95--101},
year={1987},
}
@article{benda2008spike,
title={Spike-frequency adaptation generates intensity invariance in a primary auditory interneuron},
author={Benda, Jan and Hennig, R Matthias},
journal={J Comp Neurosci},
volume={24},
pages={113--136},
year={2008},
}
@article{bhavsar2017brain,
title={Brain regions for sound processing and song release in a small grasshopper},
author={Bhavsar, Mit Balvantray and Stumpner, Andreas and Heinrich, Ralf},
journal={J Insect Physiol},
volume={99},
pages={15--24},
year={2017},
}
@article{breckow1985mechanics,
title={Mechanics of the transduction of sound in the tympanal organ of adults and larvae of locusts},
author={Breckow, Joachim and Sippel, Martin},
journal={J Comp Physiol A},
volume={157},
pages={619--629},
year={1985},
}
@misc{cigliano2018orthoptera,
title={{Orthoptera Species File. Version 5.0/5.0}},
author={Cigliano, MM and Braun, H and Eades, DC and Otte, D},
year={2018}
}
@article{clemens2013computational,
title={Computational principles underlying the recognition of acoustic signals in insects},
author={Clemens, Jan and Hennig, R Matthias},
journal={J Comp Neurosci},
volume={35},
pages={75--85},
year={2013},
}
@article{clemens2011efficient,
title={Efficient transformation of an auditory population code in a small sensory system},
author={Clemens, Jan and Kutzki, Olaf and Ronacher, Bernhard and Schreiber, Susanne and Wohlgemuth, Sandra},
journal={PNAS},
volume={108},
pages={13812--13817},
year={2011},
}
@article{clemens2013feature,
title={Feature extraction and integration underlying perceptual decision making during courtship behavior},
author={Clemens, Jan and Ronacher, Bernhard},
journal={J Neurosci},
volume={33},
pages={12136--12145},
year={2013},
}
@article{clemens2012nonlinear,
title={Nonlinear computations underlying temporal and population sparseness in the auditory system of the grasshopper},
author={Clemens, Jan and Wohlgemuth, Sandra and Ronacher, Bernhard},
journal={J Neurosci},
volume={32},
pages={10053--10062},
year={2012},
}
@article{clemens2010intensity,
title={Intensity invariance properties of auditory neurons compared to the statistics of relevant natural signals in grasshoppers},
author={Clemens, Jan and Weschke, Gerroth and Vogel, Astrid and Ronacher, Bernhard},
journal={J Comp Physiol A},
volume={196},
pages={285--297},
year={2010},
}
@article{creutzig2010timescale,
title={Timescale-invariant pattern recognition by feedforward inhibition and parallel signal processing},
author={Creutzig, Felix and Benda, Jan and Wohlgemuth, Sandra and Stumpner, Andreas and Ronacher, Bernhard and Herz, Andreas VM},
journal={Neural Comput},
volume={22},
pages={1493--1510},
year={2010},
}
@article{creutzig2009timescale,
title={Timescale-invariant representation of acoustic communication signals by a bursting neuron},
author={Creutzig, Felix and Wohlgemuth, Sandra and Stumpner, Andreas and Benda, Jan and Ronacher, Bernhard and Herz, Andreas VM},
journal={J Neurosci},
volume={29},
pages={2575--2580},
year={2009},
}
@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},
journal={J Neurosci},
volume={32},
pages={17332--17344},
year={2012},
}
@article{gollisch2002energy,
title={Energy integration describes sound-intensity coding in an insect auditory system},
author={Gollisch, Tim and Sch{\"u}tze, Hartmut and Benda, Jan and Herz, Andreas VM},
journal={J Neurosci},
volume={22},
pages={10434--10448},
year={2002},
}
@article{hennig2014time,
title={Time and timing in the acoustic recognition system of crickets},
author={Hennig, R Matthias and Heller, Klaus-Gerhard and Clemens, Jan},
journal={Front Physiol},
volume={5},
pages={286},
year={2014},
}
@article{hildebrandt2009origin,
title={The origin of adaptation in the auditory pathway of locusts is specific to cell type and function},
author={Hildebrandt, K Jannis and Benda, Jan and Hennig, R Matthias},
journal={J Neurosci},
volume={29},
pages={2626--2636},
year={2009},
}
@article{hildebrandt2015neural,
title={A neural mechanism for time-window separation resolves ambiguity of adaptive coding},
author={Hildebrandt, K Jannis and Ronacher, Bernhard and Hennig, R Matthias and Benda, Jan},
journal={PLoS Biol},
volume={13},
pages={e1002096},
year={2015},
}
@article{hoy1996tympanal,
title={Tympanal hearing in insects},
author={Hoy, Ronald R and Robert, Daniel},
journal={Annu Rev Entomol},
volume={41},
pages={433--450},
year={1996},
}
@article{lang2000acoustic,
title={Acoustic communication distances of a gomphocerine grasshopper},
author={Lang, Friederike},
journal={Bioacoustics},
volume={10},
pages={233--258},
year={2000},
}
@article{machens2001discrimination,
title={Discrimination of behaviorally relevant signals by auditory receptor neurons},
author={Machens, Christian K and Prinz, P and Stemmler, Martin B and Ronacher, Bernhard and Herz, Andreas VM},
journal={Neurocomputing},
volume={38--40},
pages={263--268},
year={2001},
}
@article{machens2001representation,
title={Representation of acoustic communication signals by insect auditory receptor neurons},
author={Machens, Christian K and Stemmler, Martin B and Prinz, Petra and Krahe, R{\"u}diger and Ronacher, Bernhard and Herz, Andreas VM},
journal={J Neurosci},
volume={21},
pages={3215--3227},
year={2001},
}
@article{malkin2014energy,
title={Energy localization and frequency analysis in the locust ear},
author={Malkin, Robert and McDonagh, Thomas R and Mhatre, Natasha and Scott, Thomas S and Robert, Daniel},
journal={J R Soc Interface},
volume={11},
pages={20130857},
year={2014},
}
@article{michelsen1971physiology,
title={{The physiology of the locust ear: I. Frequency sensitivity of single cells in the isolated ear}},
author={Michelsen, Axel},
journal={Z vergl Physiologie},
volume={71},
pages={49--62},
year={1971},
}
@article{michelsen1971frequency,
title={{The physiology of the locust ear: II. Frequency discrimination based upon resonances in the tympanum}},
author={Michelsen, Axel},
journal={Z vergl Physiologie},
volume={71},
pages={63--101},
year={1971},
}
@article{neuhofer2008evolutionarily,
title={Evolutionarily conserved coding properties of auditory neurons across grasshopper species},
author={Neuhofer, Daniela and Wohlgemuth, Sandra and Stumpner, Andreas and Ronacher, Bernhard},
journal={Proc R Soc B},
volume={275},
pages={1965--1974},
year={2008},
}
@article{rehbein1976auditory,
title={Auditory neurons in the ventral cord of the locust: Morphological and functional properties},
author={Rehbein, Hansgeorg},
journal={J Comp Physiol A},
volume={110},
pages={233--250},
year={1976},
}
@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},
journal={J Comp Physiol A},
volume={156},
pages={845--860},
year={1985},
}
@article{roemschied2014cell,
title={Cell-intrinsic mechanisms of temperature compensation in a grasshopper sensory receptor neuron},
author={R{\"o}mschied, Frederic A and Eberhard, Monika JB and Schleimer, Jan-Hendrik and Ronacher, Bernhard and Schreiber, Susanne},
journal={eLife},
volume={3},
pages={e02078},
year={2014},
}
@phdthesis{romschied2016neural,
title={Neural mechanisms of temperature compensation in an insect auditory system.},
author={R{\"o}mschied, Frederic Alexander},
school={Humboldt-Universit{\"a}t zu Berlin, Lebenswissenschaftliche Fakult{\"a}t},
year={2016},
}
@article{ronacher1986routes,
title={{Routes and stations in the processing of auditory directional information in the CNS of a grasshopper, as revealed by surgical experiments}},
author={Ronacher, Bernhard and Helversen, Dagmar v and Helversen, Otto v},
journal={J Comp Physiol A},
volume={158},
pages={363--374},
year={1986},
}
@article{ronacher2015computational,
title={Computational principles underlying recognition of acoustic signals in grasshoppers and crickets},
author={Ronacher, Bernhard and Hennig, R Matthias and Clemens, Jan},
journal={J Comp Physiol A},
volume={201},
pages={61--71},
year={2015},
}
@article{sevastianov2023evolution,
title={{Evolution of calling songs in the grasshopper subfamily Gomphocerinae (Orthoptera, Acrididae)}},
author={Sevastianov, Nikita and Neretina, Tatiana and Vedenina, Varvara},
journal={Zool Scr},
volume={52},
pages={154--175},
year={2023},
}
@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},
journal={Bioacoustics},
volume={6},
pages={1--23},
year={1994},
}
@article{suga1960peripheral,
title={Peripheral mechanism of hearing in locust},
author={Suga, Nobuo},
journal={Jpn J Physiol},
volume={10},
pages={533--546},
year={1960},
}
@article{tarasova2021biguttulus,
title={{Songs and morphology in three species of the Chorthippus biguttulus group (Orthoptera, Acrididae, Gomphocerinae) in Russia and adjacent countries}},
author={Tarasova, Tatiana and Tishechkin, Dmitry and Vedenina, Varvara},
journal={Zookeys},
volume={1073},
pages={21--53},
year={2021},
}
@article{tarasova2021eurasius,
title={{Songs and morphology in grasshoppers of the Stenobothrus eurasius group (Orthoptera: Acrdidae: Gomphocerinae) from Russia and adjacent countries: clarifying of taxonomic status}},
author={Tarasova, Tatiana A and Sevastianov, Nikita S and Vedenina, Varvara Yu},
journal={Zootaxa},
volume={4965},
pages={244--260},
year={2021},
}
@article{tishechkin2016acoustic,
title={Acoustic signals in insects: A reproductive barrier and a taxonomic character},
author={Tishechkin, D Yu and Vedenina, V Yu},
journal={Entomol Rev},
volume={96},
pages={1127--1164},
year={2016},
}
@article{vedenina2014stable,
title={{Stable and variable parameters in courtship songs of grasshoppers of the subfamily Gomphocerinae (Orthoptera, Acrididae)}},
author={Vedenina, V Yu and Shestakov, LS},
journal={Entomol Rev},
volume={94},
pages={1--20},
year={2014},
}
@article{vedenina2003complex,
title={Complex courtship in a bimodal grasshopper hybrid zone},
author={Vedenina, V Yu and von Helversen, O},
journal={Behav Ecol Sociobiol},
volume={54},
pages={44--54},
year={2003},
}
@article{vedenina2011speciation,
title={Speciation in gomphocerine grasshoppers: Molecular phylogeny versus bioacoustics and courtship behavior},
author={Vedenina, Varvara Yu and Mugue, Nikolay},
journal={J Orthoptera Res},
volume={20},
pages={109--125},
year={2011},
}
@article{vedenina2013narrow,
title={{A narrow hybrid zone between the grasshoppers Stenobothrus clavatus and Stenobothrus rubicundus (Orthoptera: Gomphocerinae): Female preferences for courtship songs}},
author={Vedenina, Varvara Yu and F{\"a}hsing, Sylvia and Sradnick, Jan and Kl{\"o}pfel, Anja and Elsner, Norbert},
journal={Biol J Linn Soc Lond},
volume={108},
pages={834--843},
year={2013},
}
@article{von1984parallel,
title={{Parallel processing in auditory pattern recognition and directional analysis by the grasshopper Chorthippus biguttulus L. (Acrididae)}},
author={von Helversen, Dagmar},
journal={J Comp Physiol A},
volume={154},
pages={837--846},
year={1984},
}
@article{von1993absolute,
title={{Absolute steepness of ramps as an essential cue for auditory pattern recognition by a grasshopper (Orthoptera; Acrididae; Chorthippus biguttulus L.)}},
author={von Helversen, Dagmar},
journal={J Comp Physiol A},
volume={172},
pages={633--639},
year={1993},
}
@article{von2004acoustic,
title={{Acoustic communication in a duetting grasshopper: Receiver response variability, male strategies and signal design}},
author={von Helversen, Dagmar and Balakrishnan, Rohini and von Helversen, Otto},
journal={Anim Behav},
volume={68},
pages={131--144},
year={2004},
}
@article{von1977stridulatory,
title={The stridulatory movements of acridid grasshoppers recorded with an opto-electronic device},
author={von Helversen, Otto and Elsner, Norbert},
journal={J Comp Physiol A},
volume={122},
pages={53--64},
year={1977},
}
@article{windmill2008time,
title={Time-resolved tympanal mechanics of the locust},
author={Windmill, JFC and Bockenhauer, S and Robert, D},
journal={J R Soc Interface},
volume={5},
pages={1435--1443},
year={2008},
}

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@ -43,7 +43,7 @@ style=authoryear,
\newcommand{\pc}{p(c_i,\,T)} % Probability density (general interval)
\newcommand{\pclp}{p(c_i,\,\tlp)} % Probability density (lowpass interval)
\section{The sensory world of a grasshopper}
\section{Exploring a grashopper's sensory world}
Strong dependence on acoustic signals for ranged communication\\
- Diverse species-specific sound repertoires and production mechanisms\\
@ -87,6 +87,19 @@ How can a human observer conceive a grasshopper's auditory percepts?\\
- How to integrate the available knowledge on anatomy, physiology, ethology?\\
$\rightarrow$ Abstract, simplify, formalize $\rightarrow$ Functional model framework
\textbf{Precursor work for model construction (special thanks to authors):}
Linear-nonlinear modelling of behavioral responses to artificial songs\\
- Feature expansion as implemented in our model: Major contribution!\\
- Bank of linear filters, nonlinearity, temporal integration, feature weighting\\
$\rightarrow$ \cite{clemens2013computational} (crickets)\\
$\rightarrow$ \cite{clemens2013feature} (grasshoppers)\\
$\rightarrow$ \cite{ronacher2015computational}\\
\textbf{Own advancements/key differences}:\\
1) Used boxcar functions as artificial "songs" (focus on few key parameters)\\
$\rightarrow$ Now actual, variable songs (as naturalistic as possible)\\
2) Fitted filters to behavioral data\\
$\rightarrow$ More general, simpler, unfitted formalized Gabor filter bank
\section{Developing a functional model of\\the grasshopper auditory pathway}
@ -96,7 +109,7 @@ $\rightarrow$ Abstract, simplify, formalize $\rightarrow$ Functional model frame
"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})\\
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:}
@ -140,7 +153,7 @@ $\rightarrow$ Highpass filter 10 Hz
"Post-split portion" of the auditory pathway:\\
Ascending neurons (AN) $\rightarrow$ Central brain neurons
Diverse response/filter properties within AN population (\cite{clemens2011})\\
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
@ -327,18 +340,29 @@ duty cycle-encoding quantity, mediated by threshold function $\nl$
on the magnitude of the derivative of $c_i(t)$ in temporal proximity to time
points at which $c_i(t)$ crosses threshold value $\thr$\\
$\rightarrow$ The steeper the slope of $c_i(t)$, the less $T_1$ changes with scale variations\\
$\rightarrow$ Extreme amplitudes of $c_i(t)$ (peaks/troughs)
$\rightarrow$ If $T_1$ is invariant to scale variation in $c_i(t)$, then so is $\feat(t)$
$\rightarrow$ Only amplitudes of \\
$\rightarrow$ Absolute amplitudes of peaks/troughs of $c_i(t)$ \\
$\rightarrow$ Acuity of peaks/troughs in $c_i(t)$ matters, not their absolute amplitude
- From graded stimulus to categorical behavioral decision:\\
- Suggests a relatively simple rule for optimal choice of threshold value $\thr$:\\
$\rightarrow$ Find amplitude $c_i$ that maximizes absolute derivative of $c_i(t)$ over time\\
$\rightarrow$ Optimal with respect to intensity invariance of $\feat(t)$, not necessarily for
other criteria such as song-noise separation or diversity between features
- Nonlinear operations can be used to detach representations from graded physical
stimulus (to fasciliate categorical behavioral decision-making?):\\
1) Capture sufficiently precise amplitude information: $\env(t)$, $\adapt(t)$\\
$\rightarrow$ Closely following the AM of the acoustic stimulus\\
2) Quantify relevant stimulus properties on a graded scale: $c_i(t)$\\
$\rightarrow$ More decorrelated representation, compared to prior stages\\
3) Nonlinearity: Distinguish between "relevant vs irrelevant" values: $\bi(t)$\\
$\rightarrow$ Trading a graded scale for two or more categorical states\\
4) Represent stimulus properties under relevance constraint: $\feat(t)$\\
$\rightarrow$ Graded again but highly decorrelated from the acoustic stimulus\\
5) Categorical behavioral decision-making requires further nonlinearities\\
$\rightarrow$ Parameters of a behavioral response may be graded (e.g. approach speed),
initiation of one behavior over another is categorical (e.g. approach/stay)
\section{Discriminating species-specific song\\patterns in feature space}
\section{Conclusions \& outlook}
\end{document}