diff --git a/cite.bib b/cite.bib index 4a1beb9..cf3ad27 100644 --- a/cite.bib +++ b/cite.bib @@ -1,8 +1,371 @@ -@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}, +} \ No newline at end of file diff --git a/figures/grasshopper_sketch.svg b/figures/grasshopper_sketch.svg new file mode 100644 index 0000000..85929bc --- /dev/null +++ b/figures/grasshopper_sketch.svg @@ -0,0 +1,876 @@ + + + + diff --git a/figures/grasshopper_sketch_full.svg b/figures/grasshopper_sketch_full.svg new file mode 100644 index 0000000..5a234ee --- /dev/null +++ b/figures/grasshopper_sketch_full.svg @@ -0,0 +1,1093 @@ + + + + diff --git a/figures/grasshopper_sketch_half.svg b/figures/grasshopper_sketch_half.svg new file mode 100644 index 0000000..dd4eecd --- /dev/null +++ b/figures/grasshopper_sketch_half.svg @@ -0,0 +1,1066 @@ + + + + diff --git a/figures/grasshopper_sketch_half_backup.svg 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grasshoppers and crickets} + \field{volume}{201} + \field{year}{2015} + \field{pages}{61\bibrangedash 71} + \range{pages}{11} \endentry \enddatalist \endrefsection diff --git a/main.bcf b/main.bcf index c3a9289..1ee8e73 100644 --- a/main.bcf +++ b/main.bcf @@ -2362,8 +2362,11 @@ cite.bib - clemens2011 - clemens2011 + clemens2013computational + clemens2013feature + ronacher2015computational + clemens2011efficient + clemens2011efficient diff --git a/main.fdb_latexmk b/main.fdb_latexmk index a27c4ec..570886a 100644 --- a/main.fdb_latexmk +++ b/main.fdb_latexmk @@ -1,14 +1,14 @@ # Fdb version 4 -["biber main"] 0 "main.bcf" "main.bbl" "main" 1763132946.72563 -1 - "cite.bib" 1762424399.63955 252 7fc49109df64576e199ce34111f01817 "" - "main.bcf" 1763132946.69547 107531 bb4289418b1a6604c7c781cbf02f8242 "pdflatex" +["biber main"] 0 "main.bcf" "main.bbl" "main" 1763453762.09677 -1 + "cite.bib" 1763394695.45527 13526 f2f482e82b02ce762bbde9705fe514aa "" + "main.bcf" 1763453762.06487 107781 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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} \ No newline at end of file