First methods paragraph (WIP).

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2026-02-02 17:13:00 +01:00
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@@ -109,11 +109,11 @@ 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
traits~(\bcite{tishechkin2016acoustic}, \bcite{tarasova2021eurasius}), which
can vary greatly within species~(\bcite{rowell1972variable},
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
species diversification~(\bcite{vedenina2011speciation},
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
@@ -122,7 +122,7 @@ 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},
hearing~(\bcite{helversen1984parallel}; \bcite{ronacher1986routes};
\bcite{helversen1988interaural}).
% What are the signals the auditory system is supposed to recognize?
@@ -133,7 +133,7 @@ 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},
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,
@@ -141,7 +141,7 @@ 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
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, or envelope, of the
song is sufficient for recognition~(\bcite{helversen1997recognition}). However,
the essential recognition cues can vary considerably with external physical
@@ -150,7 +150,7 @@ 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},
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
@@ -167,13 +167,13 @@ 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
dynamics. Intensity invariance in different auditory systems is often
associated with neuronal adaptation~(\bcite{benda2008spike},
\bcite{barbour2011intensity}, \bcite{ozeri2018fast}, more
associated with neuronal adaptation~(\bcite{benda2008spike};
\bcite{barbour2011intensity}; \bcite{ozeri2018fast}; more
general:~\bcite{benda2021neural}). In the grasshopper auditory system, a number
of neuron types along the processing chain exhibit spike-frequency adaptation
in response to sustained stimulus
intensities~(\bcite{romer1976informationsverarbeitung},
\bcite{gollisch2002energy}, \bcite{hildebrandt2009origin},
intensities~(\bcite{romer1976informationsverarbeitung};
\bcite{gollisch2002energy}; \bcite{hildebrandt2009origin};
\bcite{clemens2010intensity}) and thus likely contribute to the emergence of
intensity-invariant song representations. This means that intensity invariance
is not the result of a single processing step but rather a gradual process, in
@@ -196,8 +196,8 @@ informative features of the song pattern and then integrate the gathered
information into a final categorical percept. Previous authors have proposed a
functional model framework that describes this process --- feature extraction,
evidence accumulation, and categorical decision making --- in both
crickets~(\bcite{clemens2013computational}, \bcite{hennig2014time}) and
grasshoppers~(\bcite{clemens2013feature}, review on
crickets~(\bcite{clemens2013computational}; \bcite{hennig2014time}) and
grasshoppers~(\bcite{clemens2013feature}; review on
both:~\bcite{ronacher2015computational}). Their framework provides a
comprehensible and biologically plausible account of the computational
mechanisms required for species-specific song recognition, which has served as
@@ -219,7 +219,7 @@ larger, generic set of unfitted Gabor basis functions in order to cover a wide
range of possible song features across different species. Gabor functions
approximate the general structure of the filters used in the existing framework
as well as the filter functions found in various auditory
neurons~(\bcite{rokem2006spike}, \bcite{clemens2011efficient},
neurons~(\bcite{rokem2006spike}; \bcite{clemens2011efficient};
\bcite{clemens2012nonlinear}). The fitted sigmoidal nonlinearities in the
existing framework consistently exhibited very steep slopes and are therefore
replaced by shifted Heaviside step-functions, which results in a binarization
@@ -233,23 +233,23 @@ after a certain delay following the onset of a song, which emphasizes the
temporal dynamics of evidence accumulation towards a final categorical
decision. The most notable difference between our model pathway and the
existing framework, however, lays in the addition of a physiologically inspired
preprocessing portion, whose starting point corresponds to the initial
reception of airborne sound waves. This allows the model to operate on
unmodified recordings of natural grasshopper songs instead of condensed pulse
train approximations, which widens its scope towards more realistic,
ecologically relevant scenarios. For instance, we were able to investigate the
contribution of different processing stages to the emergence of
intensity-invariant song representations based on actual field recordings of
songs at different distances from the sender.
preprocessing stage, whose starting point corresponds to the initial reception
of airborne sound waves. This allows the model to operate on unmodified
recordings of natural grasshopper songs instead of condensed pulse train
approximations, which widens its scope towards more realistic, ecologically
relevant scenarios. For instance, we were able to investigate the contribution
of different processing stages to the emergence of intensity-invariant song
representations based on actual field recordings of songs at different
distances from the sender.
% Forgive me, it's friday.
In the following, we outline the structure of the proposed model of the
grasshopper auditory pathway, from the initial sound reception at the tympanal
membrane up to the generation of a high-dimensional, time-varying feature
representation that is suitable for species-specific song recognition. We
provide a side-by-side account of the known physiological processing steps and
their functional approximation by basic mathematical operations. We then
elaborate on two key mechanisms that drive the emergence of intensity-invariant
song representations within the auditory pathway.
grasshopper auditory pathway, from the initial reception of sound waves up to
the generation of a high-dimensional, time-varying feature representation that
is suitable for species-specific song recognition. We provide a side-by-side
account of the known physiological processing steps and their functional
approximation by basic mathematical operations. We then elaborate on two key
mechanisms that drive the emergence of intensity-invariant song representations
within the auditory pathway.
% SCRAPPED UNTIL FURTHER NOTICE:
% Multi-species, multi-individual communally inhabited environments\\
@@ -280,30 +280,38 @@ song representations within the auditory pathway.
% - How to integrate the available knowledge on anatomy, physiology, ethology?\\
% $\rightarrow$ Abstract, simplify, formalize $\rightarrow$ Functional model framework
\section{Developing a functional model of\\the grasshopper auditory pathway}
\section{Developing a functional model of the\\grasshopper song recognition pathway}
% Either pick up in intro and/or discussion, or move entirely:
The grasshopper auditory system has been studied extensively over the past
decades; and a corresponding number of involved neuron types has been
described~(\bcite{rehbein1974structure}; \bcite{kalmring1975afferent};
\bcite{rehbein1976auditory}; \bcite{eichendorf1980projections}). The functional
model we propose here focuses on the pathway responsible for song recognition
and assumes a strict feed-forward organization of three consecutive neuronal
populations: Peripheral auditory receptor neurons~\mbox{(1st order)}, local
interneurons of the metathoracic ganglion~\mbox{(2nd order)}, and ascending
neurons~\mbox{(3rd order)} projecting towards the supraesophageal ganglion.
The essence of constructing a functional model of a sensory processing system
is to gain a sufficient understanding of the system's essential structural
components and the functional roles they might fulfill; and to then build a
formal framework of manageable complexity around these two aspects.
Anatomically, the organization of the grasshopper song recognition pathway can
be outlined as a hierarchical feed-forward network of three consecutive
neuronal populations~(Fig.\,\ref{fig:pathway}a-c): Peripheral auditory receptor
neurons, whose axons enter the ventral nerve cord at the level of the
metathoracic ganglion; local interneurons that remain exclusively within the
thoracic region of the ventral nerve cord; and ascending neurons projecting
from the thoracic region towards the supraesophageal
ganglion~(\bcite{rehbein1974structure}; \bcite{rehbein1976auditory};
\bcite{eichendorf1980projections}). The input to the network originates from
Previous authors have reported a marked increase in response heterogenity
within the population of ascending neurons compared to receptors and local
interneurons, which exhibit almost identical filter characteristics,
respectively~(\bcite{clemens2011efficient}). Based on these findings, the model
pathway can be divided into two distinct portions~(Fig.\,\ref{fig:pathway}c+d).
In the preprocessing portion, generated
The preprocessing portion comprises the tympanal membrane, receptors, and
local interneurons. The different signal representations
Due to the similar response properties within the involved
The input to the network originates from
sound-induced vibrations of the tympanal membrane on each side of the thorax,
which are transduced into electro-chemical signals by the receptor neurons. The
output from the network converges somewhere in the supraesophageal ganglion,
where the recognition of conspecific songs is presumed to take
place~(\bcite{romer1985responses}; \bcite{ronacher1986routes};
\bcite{bauer1987separate}; \bcite{bhavsar2017brain}). Functionally, the
ascending neuron population is characterized by a marked increase in response
heterogenity compared to the preceding receptor neurons and local interneurons,
which exhibit relatively homogeneous response properties across their
respective populations~(\bcite{clemens2011efficient}). Based on these
considerations, the organisation of the model
pathway~(Fig.\,\ref{fig:pathway}d) comprises two distinct overall stages:
1) "Pre-split portion" of the auditory pathway:\\
@@ -324,13 +332,26 @@ $\rightarrow$ Individual neuron-specific response traces from this stage onwards
\centering
\def\svgwidth{\textwidth}
\import{figures/}{fig_auditory_pathway.pdf_tex}
\caption[Grasshopper auditory system]{\textbf{The auditory system of
grasshoppers.}}
\caption{\textbf{Schematic organisation of the song recognition pathway in
grasshoppers compared to the structure of the model pathway.}
\textbf{a}:~Course of the pathway in the grasshopper, from
the tympanal membrane over receptor neurons (1st order),
local interneurons (2nd order) of the metathoracic
ganglion, and ascending neurons (3rd order) further
towards the central brain.
\textbf{b}:~Connections between the three neuronal
populations within the metathoracic ganglion.
\textbf{c}:~Network representation of neuronal connectivity.
\textbf{d}:~Flow diagram of the different signal
representations (boxes) and transformations (arrows) along
the model pathway. The pathway consists of a
population-wide preprocessing stream followed by several
parallel feature extraction streams.
}
\label{fig:pathway}
\end{figure}
\FloatBarrier
\subsection{Population-driven signal pre-processing}
\subsection{Population-driven signal preprocessing}
Grasshoppers receive airborne sound waves by a tympanal organ at each side of
the thorax~(Fig.\,\ref{fig:pathway}a). The tympanal membrane acts as a