Began pruning the introduction.
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2
cite.bib
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cite.bib
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volume={154},
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volume={154},
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pages={837--846},
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pages={837--846},
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year={1984},
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year={1984},
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}# Cited
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}
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@article{helversen1988interaural,
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@article{helversen1988interaural,
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title={Interaural intensity and time discrimination in an unrestraint grasshopper: a tentative behavioural approach},
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title={Interaural intensity and time discrimination in an unrestraint grasshopper: a tentative behavioural approach},
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144
main.tex
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main.tex
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\documentclass[a4paper, 12pt]{article}
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\documentclass[a4paper, 12pt]{article}
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\usepackage[left=2cm,right=2cm,top=2cm,bottom=2cm,includeheadfoot]{geometry}
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\usepackage[left=2cm,right=2cm,top=2cm,bottom=2cm,includeheadfoot]{geometry}
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\usepackage[onehalfspacing]{setspace}
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% \usepackage[onehalfspacing]{setspace}
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\usepackage{graphicx}
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\usepackage{graphicx}
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\usepackage{svg}
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\usepackage{svg}
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\usepackage{import}
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\usepackage{import}
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@@ -106,95 +106,71 @@
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\newcommand{\pclp}{p(c,\,\tlp)} % Probability density (lowpass interval)
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\newcommand{\pclp}{p(c,\,\tlp)} % Probability density (lowpass interval)
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\newcommand{\muf}{\mu_{f_i}} % Average feature value
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\newcommand{\muf}{\mu_{f_i}} % Average feature value
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\section{Exploring a grasshopper's sensory world}
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\section{Introduction}
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% Why functional models of sensory systems?
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% Why functional models of sensory systems?
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Our scientific understanding of sensory processing systems results from the
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Our scientific understanding of sensory processing systems is based on the
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distributed accumulation of anatomical, physiological and ethological evidence.
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distributed accumulation of specific anatomical, physiological, and ethological
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This process is undoubtedly without alternative; however, it leaves us with the
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evidence. This leaves us with the challenge of integrating the available
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challenge of integrating the available fragments into a coherent whole in order
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knowledge fragments into a coherent whole in order to address more and more
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to address issues such as the interaction between individual system components,
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far-reaching questions, from the interaction between individual processing
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the functional limitations of the system overall, or taxonomic comparisons
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steps to comparisons between similar systems across different species. One way
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between systems that process the same sensory modality. Any unified framework
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to deal with this challenge is to build a unified framework that captures the
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that captures the essential functional aspects of a given sensory system thus
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essential functional aspects of a sensory system. However, building such a
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has the potential to deepen our current understanding and fasciliate systematic
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framework is a challenging task in itself. It requires a wealth of existing
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investigations. However, building such a framework is a challenging task. It
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knowledge of the system and the stimuli it operates on, a clearly defined
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requires a wealth of existing knowledge of the system and the signals it
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scope, and careful abstraction of the underlying structures and mechanisms.
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operates on, a clearly defined scope, and careful reduction, abstraction, and
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formalization of the underlying structures and mechanisms.
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% Why the grasshopper auditory system?
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% Why the grasshopper auditory system?
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% Why focus on song recognition among other auditory functions?
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% Why focus on song recognition among other auditory functions?
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One sensory system about which extensive information has been gathered over the
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One sensory system that has been extensively studied over the years is the
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years is the auditory system of grasshoppers~(\textit{Acrididae}). Grasshoppers
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auditory system of grasshoppers~(\textit{Acrididae}). Grasshoppers rely on
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rely on their sense of hearing primarily for intraspecific communication, which
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their sense of hearing for intraspecific communication --- including mate
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includes mate attraction~(\bcite{helversen1972gesang}) and
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attraction~(\bcite{helversen1972gesang}) and
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evaluation~(\bcite{stange2012grasshopper}), sender
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evaluation~(\bcite{stange2012grasshopper}), sender
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localization~(\bcite{helversen1988interaural}), courtship
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localization~(\bcite{helversen1988interaural}), courtship
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display~(\bcite{elsner1968neuromuskularen}), rival
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display~(\bcite{elsner1968neuromuskularen}), and rival
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deterrence~(\bcite{greenfield1993acoustic}), and loss-of-signal predator
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deterrence~(\bcite{greenfield1993acoustic}) --- and have evolved a variety of
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alarm~(SOURCE). In accordance with this rich behavioral repertoire,
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acoustic signals for different behavioral
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grasshoppers have evolved a variety of sound production mechanisms to generate
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contexts~(\bcite{otte1970comparative}). The most conspicuous acoustic signals
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acoustic communication signals for different contexts and ranges using their
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of grasshoppers are their species-specific calling songs, which broadcast the
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wings, hindlegs, or mandibles~(\bcite{otte1970comparative}). Among the most
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presence of the singing individual to potential mates within range. These songs
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conspicuous acoustic signals of grasshoppers are their species-specific calling
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are usually more characteristic of a species than morphological
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songs, which broadcast the presence of the singing individual --- mostly the
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males of the species --- to potential mates within range. These songs are
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usually more characteristic of a species than morphological
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traits~(\bcite{tishechkin2016acoustic}; \bcite{tarasova2021eurasius}), which
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traits~(\bcite{tishechkin2016acoustic}; \bcite{tarasova2021eurasius}), which
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can vary greatly within species~(\bcite{rowell1972variable};
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can vary greatly within species~(\bcite{rowell1972variable};
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\bcite{kohler2017morphological}). The reliance on songs to mediate reproduction
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\bcite{kohler2017morphological}). The reliance on songs to mediate reproduction
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represents a strong evolutionary driving force, that resulted in a massive
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represents a strong evolutionary driving force that resulted in a massive
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species diversification~(\bcite{vedenina2011speciation};
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species diversification~(\bcite{vedenina2011speciation};
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\bcite{sevastianov2023evolution}), with over 6800 recognized grasshopper
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\bcite{sevastianov2023evolution}), with over 6800 recognized species in the
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species in the \textit{Acrididae} family~(\bcite{cigliano2024orthoptera}). It
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\textit{Acrididae} family~(\bcite{cigliano2024orthoptera}).
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is this diversity of species, and the crucial role of acoustic communication in
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% Could go lower to concluding part:
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its emergence, that makes the grasshopper auditory system an intriguing
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% Its evolutionary significance makes the grasshopper auditory system ---
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candidate for attempting to construct a functional model framework. As a
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% specifically, the pathway responsible for species-specific song recognition
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necessary reduction, the model we propose here focuses on the pathway
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% --- an intriguing candidate for attempting to construct a functional model
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responsible for the recognition of species-specific calling songs, disregarding
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% framework.
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other essential auditory functions such as directional
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hearing~(\bcite{helversen1984parallel}; \bcite{ronacher1986routes};
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\bcite{helversen1988interaural}).
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% What are the signals the auditory system is supposed to recognize?
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% What are the signals that the auditory system is supposed to recognize?
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% Why is intensity invariance important for song recognition?
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Grasshopper songs are amplitude-modulated broad-band acoustic signals. They
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% (Obviously, split this paragraph)
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consist of a series of noisy syllables and relatively quiet pauses, which form
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To understand the functional challenges faced by the grasshopper auditory
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a characteristic repetitive pattern~(\bcite{helversen1977stridulatory};
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system, one has to understand the properties of the songs it is designed to
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\bcite{stumpner1994song}). Song recognition depends on certain structural
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recognize. Grasshopper songs are amplitude-modulated broad-band acoustic
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parameters of this pattern --- such as the duration of syllables and
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signals. Most songs are produced by stridulation, during which the animal pulls
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the serrated stridulatory file on its hindlegs across a resonating vein on the
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forewings~(\bcite{helversen1977stridulatory}; \bcite{stumpner1994song};
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\bcite{helversen1997recognition}). Every tooth that strikes the vein generates
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a brief pulse of sound. Multiple pulses make up a syllable; and the alternation
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of syllables and relatively quiet pauses forms a characteristic, through noisy,
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waveform pattern. Song recognition depends on certain temporal and structural
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parameters of this pattern, such as the duration of syllables and
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pauses~(\bcite{helversen1972gesang}), the slope of pulse
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pauses~(\bcite{helversen1972gesang}), the slope of pulse
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onsets~(\bcite{helversen1993absolute}), and the accentuation of syllable onsets
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onsets~(\bcite{helversen1993absolute}), and the accentuation of syllable onsets
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relative to the preceeding pause~(\bcite{balakrishnan2001song};
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relative to the preceeding pause~(\bcite{balakrishnan2001song};
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\bcite{helversen2004acoustic}). The amplitude modulation of the song is
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\bcite{helversen2004acoustic}) --- which are sufficiently conveyed by the
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sufficient for recognition~(\bcite{helversen1997recognition}). However, the
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amplitude modulation of the song alone~(\bcite{helversen1997recognition}).
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essential recognition cues can vary considerably with external physical
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factors, which requires the auditory system to be invariant to such variations
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% Why is intensity invariance important for song recognition?
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in order to reliably recognize songs under different conditions. For instance,
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Grasshopper songs, like all acoustic signals, are subject to sound attenuation,
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the temporal structure of grasshopper songs warps with
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which depends on the distance from the sender, the frequency content of the
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temperature~(\bcite{skovmand1983song}). The auditory system can compensate for
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signal, and the vegetation of the habitat~(\bcite{michelsen1978sound}). The
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this variability by reading out relative temporal relationships rather than
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amplitude dynamics of the song pattern degrade fairly quickly, which limits the
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absolute time intervals~(\bcite{creutzig2009timescale};
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effective communication range of grasshoppers to~\mbox{1\,-\,2\,m} in their
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\bcite{creutzig2010timescale}), as those remain relatively constant across
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typical grassland habitats~(\bcite{lang2000acoustic}). Moreover, the intensity
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different temperatures~(\bcite{helversen1972gesang}). Another, perhaps even
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of a song at the receiver's position varies with the location of the sender,
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more fundamental external source of song variability lays in the attenuation of
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which should ideally not affect the recognition of the song.
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sound intensity with increasing distance to the sender. Sound attenuation
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depends on both the frequency content of the signal and the vegetation of the
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habitat~(\bcite{michelsen1978sound}). For the receiving auditory system, this
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has two major implications. First, the amplitude dynamics of the song pattern
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are steadily degraded over distance, which limits the effective communication
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range of grasshoppers to~\mbox{1\,-\,2\,m} in their typical grassland
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habitats~(\bcite{lang2000acoustic}). Second, the overall intensity level of
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songs at the receiver's position varies depending on the location of the
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sender, which should ideally not affect the recognition of the song pattern.
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This neccessitates that the auditory system achieves a certain degree of
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This neccessitates that the auditory system achieves a certain degree of
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intensity invariance --- a time scale-selective sensitivity to faster amplitude
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intensity invariance --- a time scale-selective sensitivity to faster amplitude
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dynamics and simultaneous insensitivity to slower, more sustained amplitude
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dynamics and simultaneous insensitivity to slower, more sustained amplitude
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@@ -1620,12 +1596,28 @@ natural song variation.
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\section{Conclusions \& outlook}
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\section{Conclusions \& outlook}
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\textbf{The role of repetitive songs for the feature representation:}
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Most grasshopper songs are produced by stridulation, which refers to the
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pulling of the serrated stridulatory file on the hindlegs across a resonating
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vein on the forewings~(\bcite{helversen1977stridulatory};
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\bcite{stumpner1994song}; \bcite{helversen1997recognition}). Every "tooth" that
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strikes the vein generates a brief sound pulse; multiple pulses make up a
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syllable; and the repetition of syllables and pauses results in a
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characteristic amplitude-modulated waveform pattern.
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\textbf{Excursion into time-warp invariance:}
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For instance, the temporal structure of grasshopper songs warps with
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temperature~(\bcite{skovmand1983song}). The auditory system can compensate for
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this variability by reading out relative temporal relationships rather than
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absolute time intervals~(\bcite{creutzig2009timescale};
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\bcite{creutzig2010timescale}), as those remain relatively constant across
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different temperatures~(\bcite{helversen1972gesang}).
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\textbf{Song recognition pathway: Grasshopper vs. model:}\\
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\textbf{Song recognition pathway: Grasshopper vs. model:}\\
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The model pathway includes a rather large number of Gabor kernels compared to
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The model pathway includes a rather large number of Gabor kernels compared to
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the 15 to 20 ascending neurons in the grasshopper auditory
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the 15 to 20 ascending neurons in the grasshopper auditory
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system~(\bcite{stumpner1991auditory}).
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system~(\bcite{stumpner1991auditory}).
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\textbf{Definition of invariance (general, systemic):}\\
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\textbf{Definition of invariance (general, systemic):}\\
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Invariance = Property of a system to maintain a stable output with respect to a
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Invariance = Property of a system to maintain a stable output with respect to a
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set of relevant input parameters (variation to be represented) but irrespective
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set of relevant input parameters (variation to be represented) but irrespective
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