Began pruning the introduction.

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

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