First methods paragraph (WIP).
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127
main.tex
127
main.tex
@@ -109,11 +109,11 @@ conspicuous acoustic signals of grasshoppers are their species-specific calling
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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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can vary greatly within species~(\bcite{rowell1972variable},
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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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\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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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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species in the \textit{Acrididae} family~(\bcite{cigliano2024orthoptera}). It
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is this diversity of species, and the crucial role of acoustic communication in
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@@ -122,7 +122,7 @@ candidate for attempting to construct a functional model framework. As a
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necessary reduction, the model we propose here focuses on the pathway
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responsible for the recognition of species-specific calling songs, disregarding
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other essential auditory functions such as directional
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hearing~(\bcite{helversen1984parallel}, \bcite{ronacher1986routes},
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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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@@ -133,7 +133,7 @@ system, one has to understand the properties of the songs it is designed to
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recognize. Grasshopper songs are amplitude-modulated broad-band acoustic
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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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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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@@ -141,7 +141,7 @@ 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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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, or envelope, of the
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song is sufficient for recognition~(\bcite{helversen1997recognition}). However,
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the essential recognition cues can vary considerably with external physical
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@@ -150,7 +150,7 @@ in order to reliably recognize songs under different conditions. For instance,
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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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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}). Another, perhaps even
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more fundamental external source of song variability lays in the attenuation of
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@@ -167,13 +167,13 @@ 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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dynamics and simultaneous insensitivity to slower, more sustained amplitude
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dynamics. Intensity invariance in different auditory systems is often
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associated with neuronal adaptation~(\bcite{benda2008spike},
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\bcite{barbour2011intensity}, \bcite{ozeri2018fast}, more
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associated with neuronal adaptation~(\bcite{benda2008spike};
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\bcite{barbour2011intensity}; \bcite{ozeri2018fast}; more
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general:~\bcite{benda2021neural}). In the grasshopper auditory system, a number
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of neuron types along the processing chain exhibit spike-frequency adaptation
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in response to sustained stimulus
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intensities~(\bcite{romer1976informationsverarbeitung},
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\bcite{gollisch2002energy}, \bcite{hildebrandt2009origin},
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intensities~(\bcite{romer1976informationsverarbeitung};
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\bcite{gollisch2002energy}; \bcite{hildebrandt2009origin};
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\bcite{clemens2010intensity}) and thus likely contribute to the emergence of
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intensity-invariant song representations. This means that intensity invariance
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is not the result of a single processing step but rather a gradual process, in
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@@ -196,8 +196,8 @@ informative features of the song pattern and then integrate the gathered
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information into a final categorical percept. Previous authors have proposed a
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functional model framework that describes this process --- feature extraction,
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evidence accumulation, and categorical decision making --- in both
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crickets~(\bcite{clemens2013computational}, \bcite{hennig2014time}) and
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grasshoppers~(\bcite{clemens2013feature}, review on
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crickets~(\bcite{clemens2013computational}; \bcite{hennig2014time}) and
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grasshoppers~(\bcite{clemens2013feature}; review on
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both:~\bcite{ronacher2015computational}). Their framework provides a
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comprehensible and biologically plausible account of the computational
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mechanisms required for species-specific song recognition, which has served as
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@@ -219,7 +219,7 @@ larger, generic set of unfitted Gabor basis functions in order to cover a wide
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range of possible song features across different species. Gabor functions
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approximate the general structure of the filters used in the existing framework
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as well as the filter functions found in various auditory
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neurons~(\bcite{rokem2006spike}, \bcite{clemens2011efficient},
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neurons~(\bcite{rokem2006spike}; \bcite{clemens2011efficient};
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\bcite{clemens2012nonlinear}). The fitted sigmoidal nonlinearities in the
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existing framework consistently exhibited very steep slopes and are therefore
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replaced by shifted Heaviside step-functions, which results in a binarization
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@@ -233,23 +233,23 @@ after a certain delay following the onset of a song, which emphasizes the
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temporal dynamics of evidence accumulation towards a final categorical
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decision. The most notable difference between our model pathway and the
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existing framework, however, lays in the addition of a physiologically inspired
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preprocessing portion, whose starting point corresponds to the initial
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reception of airborne sound waves. This allows the model to operate on
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unmodified recordings of natural grasshopper songs instead of condensed pulse
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train approximations, which widens its scope towards more realistic,
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ecologically relevant scenarios. For instance, we were able to investigate the
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contribution of different processing stages to the emergence of
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intensity-invariant song representations based on actual field recordings of
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songs at different distances from the sender.
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preprocessing stage, whose starting point corresponds to the initial reception
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of airborne sound waves. This allows the model to operate on unmodified
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recordings of natural grasshopper songs instead of condensed pulse train
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approximations, which widens its scope towards more realistic, ecologically
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relevant scenarios. For instance, we were able to investigate the contribution
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of different processing stages to the emergence of intensity-invariant song
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representations based on actual field recordings of songs at different
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distances from the sender.
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% Forgive me, it's friday.
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In the following, we outline the structure of the proposed model of the
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grasshopper auditory pathway, from the initial sound reception at the tympanal
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membrane up to the generation of a high-dimensional, time-varying feature
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representation that is suitable for species-specific song recognition. We
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provide a side-by-side account of the known physiological processing steps and
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their functional approximation by basic mathematical operations. We then
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elaborate on two key mechanisms that drive the emergence of intensity-invariant
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song representations within the auditory pathway.
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grasshopper auditory pathway, from the initial reception of sound waves up to
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the generation of a high-dimensional, time-varying feature representation that
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is suitable for species-specific song recognition. We provide a side-by-side
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account of the known physiological processing steps and their functional
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approximation by basic mathematical operations. We then elaborate on two key
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mechanisms that drive the emergence of intensity-invariant song representations
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within the auditory pathway.
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% SCRAPPED UNTIL FURTHER NOTICE:
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% Multi-species, multi-individual communally inhabited environments\\
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@@ -280,30 +280,38 @@ song representations within the auditory pathway.
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% - How to integrate the available knowledge on anatomy, physiology, ethology?\\
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% $\rightarrow$ Abstract, simplify, formalize $\rightarrow$ Functional model framework
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\section{Developing a functional model of\\the grasshopper auditory pathway}
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\section{Developing a functional model of the\\grasshopper song recognition pathway}
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% Either pick up in intro and/or discussion, or move entirely:
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The grasshopper auditory system has been studied extensively over the past
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decades; and a corresponding number of involved neuron types has been
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described~(\bcite{rehbein1974structure}; \bcite{kalmring1975afferent};
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\bcite{rehbein1976auditory}; \bcite{eichendorf1980projections}). The functional
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model we propose here focuses on the pathway responsible for song recognition
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and assumes a strict feed-forward organization of three consecutive neuronal
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populations: Peripheral auditory receptor neurons~\mbox{(1st order)}, local
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interneurons of the metathoracic ganglion~\mbox{(2nd order)}, and ascending
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neurons~\mbox{(3rd order)} projecting towards the supraesophageal ganglion.
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The essence of constructing a functional model of a sensory processing system
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is to gain a sufficient understanding of the system's essential structural
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components and the functional roles they might fulfill; and to then build a
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formal framework of manageable complexity around these two aspects.
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Anatomically, the organization of the grasshopper song recognition pathway can
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be outlined as a hierarchical feed-forward network of three consecutive
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neuronal populations~(Fig.\,\ref{fig:pathway}a-c): Peripheral auditory receptor
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neurons, whose axons enter the ventral nerve cord at the level of the
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metathoracic ganglion; local interneurons that remain exclusively within the
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thoracic region of the ventral nerve cord; and ascending neurons projecting
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from the thoracic region towards the supraesophageal
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ganglion~(\bcite{rehbein1974structure}; \bcite{rehbein1976auditory};
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\bcite{eichendorf1980projections}). The input to the network originates from
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Previous authors have reported a marked increase in response heterogenity
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within the population of ascending neurons compared to receptors and local
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interneurons, which exhibit almost identical filter characteristics,
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respectively~(\bcite{clemens2011efficient}). Based on these findings, the model
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pathway can be divided into two distinct portions~(Fig.\,\ref{fig:pathway}c+d).
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In the preprocessing portion, generated
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The preprocessing portion comprises the tympanal membrane, receptors, and
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local interneurons. The different signal representations
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Due to the similar response properties within the involved
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The input to the network originates from
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sound-induced vibrations of the tympanal membrane on each side of the thorax,
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which are transduced into electro-chemical signals by the receptor neurons. The
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output from the network converges somewhere in the supraesophageal ganglion,
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where the recognition of conspecific songs is presumed to take
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place~(\bcite{romer1985responses}; \bcite{ronacher1986routes};
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\bcite{bauer1987separate}; \bcite{bhavsar2017brain}). Functionally, the
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ascending neuron population is characterized by a marked increase in response
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heterogenity compared to the preceding receptor neurons and local interneurons,
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which exhibit relatively homogeneous response properties across their
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respective populations~(\bcite{clemens2011efficient}). Based on these
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considerations, the organisation of the model
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pathway~(Fig.\,\ref{fig:pathway}d) comprises two distinct overall stages:
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1) "Pre-split portion" of the auditory pathway:\\
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@@ -324,13 +332,26 @@ $\rightarrow$ Individual neuron-specific response traces from this stage onwards
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\centering
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\def\svgwidth{\textwidth}
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\import{figures/}{fig_auditory_pathway.pdf_tex}
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\caption[Grasshopper auditory system]{\textbf{The auditory system of
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grasshoppers.}}
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\caption{\textbf{Schematic organisation of the song recognition pathway in
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grasshoppers compared to the structure of the model pathway.}
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\textbf{a}:~Course of the pathway in the grasshopper, from
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the tympanal membrane over receptor neurons (1st order),
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local interneurons (2nd order) of the metathoracic
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ganglion, and ascending neurons (3rd order) further
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towards the central brain.
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\textbf{b}:~Connections between the three neuronal
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populations within the metathoracic ganglion.
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\textbf{c}:~Network representation of neuronal connectivity.
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\textbf{d}:~Flow diagram of the different signal
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representations (boxes) and transformations (arrows) along
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the model pathway. The pathway consists of a
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population-wide preprocessing stream followed by several
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parallel feature extraction streams.
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}
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\label{fig:pathway}
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\end{figure}
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\FloatBarrier
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\subsection{Population-driven signal pre-processing}
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\subsection{Population-driven signal preprocessing}
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Grasshoppers receive airborne sound waves by a tympanal organ at each side of
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the thorax~(Fig.\,\ref{fig:pathway}a). The tympanal membrane acts as a
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