j_hartling/paper_2025
1
0
Fork 0
Code Issues Pull Requests Actions Packages Projects Releases Wiki Activity

Began pruning the introduction.

Browse Source
This commit is contained in:
j-hartling
2026-05-18 18:19:48 +02:00
parent a205d70fe4
commit 85dc43fb38
4 changed files with 72 additions and 80 deletions
Download Patch File Download Diff File Expand all files Collapse all files

6
.gitignore vendored
View File

@@ -17,8 +17,8 @@ data/*
*.cb
*.cb2
.*.lb
*.bbl
*.bcf
*.bbl*
*.bcf*
*.blg
*-blx.aux
*-blx.bib
@@ -28,4 +28,4 @@ data/*
*.synctex(busy)
*.synctex.gz
*.synctex.gz(busy)
*.pdfsync
*.pdfsync

2
cite.bib
View File

@@ -253,7 +253,7 @@
volume={154},
pages={837--846},
year={1984},
}# Cited
}
@article{helversen1988interaural,
title={Interaural intensity and time discrimination in an unrestraint grasshopper: a tentative behavioural approach},

BIN
main.pdf
View File

Binary file not shown.

144
main.tex
View File

@@ -1,7 +1,7 @@
\documentclass[a4paper, 12pt]{article}
\usepackage[left=2cm,right=2cm,top=2cm,bottom=2cm,includeheadfoot]{geometry}
\usepackage[onehalfspacing]{setspace}
% \usepackage[onehalfspacing]{setspace}
\usepackage{graphicx}
\usepackage{svg}
\usepackage{import}
@@ -106,95 +106,71 @@
\newcommand{\pclp}{p(c,\,\tlp)} % Probability density (lowpass interval)
\newcommand{\muf}{\mu_{f_i}} % Average feature value
\section{Exploring a grasshopper's sensory world}
\section{Introduction}
% Why functional models of sensory systems?
Our scientific understanding of sensory processing systems results from the
distributed accumulation of anatomical, physiological and ethological evidence.
This process is undoubtedly without alternative; however, it leaves us with the
challenge of integrating the available fragments into a coherent whole in order
to address issues such as the interaction between individual system components,
the functional limitations of the system overall, or taxonomic comparisons
between systems that process the same sensory modality. Any unified framework
that captures the essential functional aspects of a given sensory system thus
has the potential to deepen our current understanding and fasciliate systematic
investigations. However, building such a framework is a challenging task. It
requires a wealth of existing knowledge of the system and the signals it
operates on, a clearly defined scope, and careful reduction, abstraction, and
formalization of the underlying structures and mechanisms.
Our scientific understanding of sensory processing systems is based on the
distributed accumulation of specific anatomical, physiological, and ethological
evidence. This leaves us with the challenge of integrating the available
knowledge fragments into a coherent whole in order to address more and more
far-reaching questions, from the interaction between individual processing
steps to comparisons between similar systems across different species. One way
to deal with this challenge is to build a unified framework that captures the
essential functional aspects of a sensory system. However, building such a
framework is a challenging task in itself. It requires a wealth of existing
knowledge of the system and the stimuli it operates on, a clearly defined
scope, and careful abstraction of the underlying structures and mechanisms.
% Why the grasshopper auditory system?
% Why focus on song recognition among other auditory functions?
One sensory system about which extensive information has been gathered over the
years is the auditory system of grasshoppers~(\textit{Acrididae}). Grasshoppers
rely on their sense of hearing primarily for intraspecific communication, which
includes mate attraction~(\bcite{helversen1972gesang}) and
One sensory system that has been extensively studied over the years is the
auditory system of grasshoppers~(\textit{Acrididae}). Grasshoppers rely on
their sense of hearing for intraspecific communication --- including mate
attraction~(\bcite{helversen1972gesang}) and
evaluation~(\bcite{stange2012grasshopper}), sender
localization~(\bcite{helversen1988interaural}), courtship
display~(\bcite{elsner1968neuromuskularen}), rival
deterrence~(\bcite{greenfield1993acoustic}), and loss-of-signal predator
alarm~(SOURCE). In accordance with this rich behavioral repertoire,
grasshoppers have evolved a variety of sound production mechanisms to generate
acoustic communication signals for different contexts and ranges using their
wings, hindlegs, or mandibles~(\bcite{otte1970comparative}). Among the most
conspicuous acoustic signals of grasshoppers are their species-specific calling
songs, which broadcast the presence of the singing individual --- mostly the
males of the species --- to potential mates within range. These songs are
usually more characteristic of a species than morphological
display~(\bcite{elsner1968neuromuskularen}), and rival
deterrence~(\bcite{greenfield1993acoustic}) --- and have evolved a variety of
acoustic signals for different behavioral
contexts~(\bcite{otte1970comparative}). The most conspicuous acoustic signals
of grasshoppers are their species-specific calling songs, which broadcast the
presence of the singing individual to potential mates within range. These songs
are usually more characteristic of a species than morphological
traits~(\bcite{tishechkin2016acoustic}; \bcite{tarasova2021eurasius}), which
can vary greatly within species~(\bcite{rowell1972variable};
\bcite{kohler2017morphological}). The reliance on songs to mediate reproduction
represents a strong evolutionary driving force, that resulted in a massive
represents a strong evolutionary driving force that resulted in a massive
species diversification~(\bcite{vedenina2011speciation};
\bcite{sevastianov2023evolution}), with over 6800 recognized grasshopper
species in the \textit{Acrididae} family~(\bcite{cigliano2024orthoptera}). It
is this diversity of species, and the crucial role of acoustic communication in
its emergence, that makes the grasshopper auditory system an intriguing
candidate for attempting to construct a functional model framework. As a
necessary reduction, the model we propose here focuses on the pathway
responsible for the recognition of species-specific calling songs, disregarding
other essential auditory functions such as directional
hearing~(\bcite{helversen1984parallel}; \bcite{ronacher1986routes};
\bcite{helversen1988interaural}).
\bcite{sevastianov2023evolution}), with over 6800 recognized species in the
\textit{Acrididae} family~(\bcite{cigliano2024orthoptera}).
% Could go lower to concluding part:
% Its evolutionary significance makes the grasshopper auditory system ---
% specifically, the pathway responsible for species-specific song recognition
% --- an intriguing candidate for attempting to construct a functional model
% framework.
% What are the signals the auditory system is supposed to recognize?
% Why is intensity invariance important for song recognition?
% (Obviously, split this paragraph)
To understand the functional challenges faced by the grasshopper auditory
system, one has to understand the properties of the songs it is designed to
recognize. Grasshopper songs are amplitude-modulated broad-band acoustic
signals. Most songs are produced by stridulation, during which the animal pulls
the serrated stridulatory file on its hindlegs across a resonating vein on the
forewings~(\bcite{helversen1977stridulatory}; \bcite{stumpner1994song};
\bcite{helversen1997recognition}). Every tooth that strikes the vein generates
a brief pulse of sound. Multiple pulses make up a syllable; and the alternation
of syllables and relatively quiet pauses forms a characteristic, through noisy,
waveform pattern. Song recognition depends on certain temporal and structural
parameters of this pattern, such as the duration of syllables and
% What are the signals that the auditory system is supposed to recognize?
Grasshopper songs are amplitude-modulated broad-band acoustic signals. They
consist of a series of noisy syllables and relatively quiet pauses, which form
a characteristic repetitive pattern~(\bcite{helversen1977stridulatory};
\bcite{stumpner1994song}). Song recognition depends on certain structural
parameters of this pattern --- such as the duration of syllables and
pauses~(\bcite{helversen1972gesang}), the slope of pulse
onsets~(\bcite{helversen1993absolute}), and the accentuation of syllable onsets
relative to the preceeding pause~(\bcite{balakrishnan2001song};
\bcite{helversen2004acoustic}). The amplitude modulation of the song is
sufficient for recognition~(\bcite{helversen1997recognition}). However, the
essential recognition cues can vary considerably with external physical
factors, which requires the auditory system to be invariant to such variations
in order to reliably recognize songs under different conditions. For instance,
the temporal structure of grasshopper songs warps with
temperature~(\bcite{skovmand1983song}). The auditory system can compensate for
this variability by reading out relative temporal relationships rather than
absolute time intervals~(\bcite{creutzig2009timescale};
\bcite{creutzig2010timescale}), as those remain relatively constant across
different temperatures~(\bcite{helversen1972gesang}). Another, perhaps even
more fundamental external source of song variability lays in the attenuation of
sound intensity with increasing distance to the sender. Sound attenuation
depends on both the frequency content of the signal and the vegetation of the
habitat~(\bcite{michelsen1978sound}). For the receiving auditory system, this
has two major implications. First, the amplitude dynamics of the song pattern
are steadily degraded over distance, which limits the effective communication
range of grasshoppers to~\mbox{1\,-\,2\,m} in their typical grassland
habitats~(\bcite{lang2000acoustic}). Second, the overall intensity level of
songs at the receiver's position varies depending on the location of the
sender, which should ideally not affect the recognition of the song pattern.
\bcite{helversen2004acoustic}) --- which are sufficiently conveyed by the
amplitude modulation of the song alone~(\bcite{helversen1997recognition}).
% Why is intensity invariance important for song recognition?
Grasshopper songs, like all acoustic signals, are subject to sound attenuation,
which depends on the distance from the sender, the frequency content of the
signal, and the vegetation of the habitat~(\bcite{michelsen1978sound}). The
amplitude dynamics of the song pattern degrade fairly quickly, which limits the
effective communication range of grasshoppers to~\mbox{1\,-\,2\,m} in their
typical grassland habitats~(\bcite{lang2000acoustic}). Moreover, the intensity
of a song at the receiver's position varies with the location of the sender,
which should ideally not affect the recognition of the song.
This neccessitates that the auditory system achieves a certain degree of
intensity invariance --- a time scale-selective sensitivity to faster amplitude
dynamics and simultaneous insensitivity to slower, more sustained amplitude
@@ -1620,12 +1596,28 @@ natural song variation.
\section{Conclusions \& outlook}
\textbf{The role of repetitive songs for the feature representation:}
Most grasshopper songs are produced by stridulation, which refers to the
pulling of the serrated stridulatory file on the hindlegs across a resonating
vein on the forewings~(\bcite{helversen1977stridulatory};
\bcite{stumpner1994song}; \bcite{helversen1997recognition}). Every "tooth" that
strikes the vein generates a brief sound pulse; multiple pulses make up a
syllable; and the repetition of syllables and pauses results in a
characteristic amplitude-modulated waveform pattern.
\textbf{Excursion into time-warp invariance:}
For instance, the temporal structure of grasshopper songs warps with
temperature~(\bcite{skovmand1983song}). The auditory system can compensate for
this variability by reading out relative temporal relationships rather than
absolute time intervals~(\bcite{creutzig2009timescale};
\bcite{creutzig2010timescale}), as those remain relatively constant across
different temperatures~(\bcite{helversen1972gesang}).
\textbf{Song recognition pathway: Grasshopper vs. model:}\\
The model pathway includes a rather large number of Gabor kernels compared to
the 15 to 20 ascending neurons in the grasshopper auditory
system~(\bcite{stumpner1991auditory}).
\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