Finished drafting the introduction.
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164
main.tex
164
main.tex
@@ -198,140 +198,35 @@ process, in which different neuronal populations contribute to varying
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degrees~(\bcite{clemens2010intensity}) and by different
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degrees~(\bcite{clemens2010intensity}) and by different
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mechanisms~(\bcite{hildebrandt2009origin}).
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mechanisms~(\bcite{hildebrandt2009origin}).
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% How can song recognition be modelled functionally (feat. Jan Clemens & Co.)?
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% How did we expand on the previous framework (feat. Clemens et al.)?
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% How did we expand on the previous framework?
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In the current study, we leverage functional modelling to trace the emergence
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In the current study, we leverage functional modelling to trace the emergence
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of intensity invariance through individual processing steps of the grasshopper
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of intensity invariance through individual processing steps of the grasshopper
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song recognition pathway. The model pathway we propose here expands on a
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song recognition pathway. The model pathway we propose here is based on a
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previous functional model framework for song recognition --- including feature
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previous functional model framework for song recognition in both
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extraction, evidence accumulation, and categorical decision making --- in both
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crickets~(\bcite{clemens2013computational}; \bcite{hennig2014time}) and
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crickets~(\bcite{clemens2013computational}; \bcite{hennig2014time}) and
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grasshoppers~(\bcite{clemens2013feature}; review on
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grasshoppers~(\bcite{clemens2013feature}; review on
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both:~\bcite{ronacher2015computational}). The exisiting framework relies on
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both:~\bcite{ronacher2015computational}). The exisiting framework relies on
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pulse trains as input signals, which were designed to capture the essential
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pulse trains as input signals, which were designed to capture the essential
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structural properties of natural song envelopes~(\bcite{clemens2013feature}).
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structural properties of natural song envelopes~(\bcite{clemens2013feature}).
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We extended this framework by a physiologically plausible preprocessing stage
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It includes feature extraction by a bank of linear-nonlinear feature detectors,
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--- including spectral filtering, envelope extraction, logarithmic compression,
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evidence accumulation by temporal averaging of each feature, and categorical
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and intensity adaptation --- which allows the model to operate on unmodified
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decision making by a weighted linear combination of feature values. We adopted
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recordings of natural grasshopper songs. The result is a comprehensible account
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the general structure of the existing framework and extended it by a
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of the known
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physiologically plausible preprocessing stage --- including spectral filtering,
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envelope extraction, logarithmic compression, and intensity adaptation ---
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which allows the model to operate on unmodified recordings of natural
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% Its evolutionary significance makes the grasshopper auditory system ---
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grasshopper songs. The resulting model pathway thus covers the entire auditory
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% specifically, the pathway responsible for species-specific song recognition
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processing stream from the initial reception of airborne sound waves to the
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% --- an intriguing candidate for attempting to construct a functional model
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generation of a high-dimensional feature representation that allows for the
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% framework.
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categorical recognition of conspecific songs. It incorporates anatomical,
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physiological, and ethological evidence from several decades of research on the
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% Another, more substantial modification is that
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grasshopper auditory system. In the following, we provide a side-by-side
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% the feature detector outputs are temporally averaged in a way that does not
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account of the known physiological processing steps along the song recognition
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% condense them into single feature values but retains their time-varying
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pathway and their functional approximations in the model pathway. We then
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% structure. This is in line with the fact that songs are no discrete units but
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elaborate on the computational mechanisms that contribute to the emergence of
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% part of a continuous acoustic stream that the auditory system has to process in
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intensity-invariant song representations, the interaction between these
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% real time. Moreover, a time-varying feature representation only stabilizes
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mechanisms, the overall capacity for intensity invariance in the system, and
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% after a certain delay following the onset of a song, which emphasizes the
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the ethological implications of our findings.
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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 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.
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%% BACKUP OF PREVIOUS ITERATION:
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% % Why functional models of sensory systems?
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% Our scientific understanding of sensory processing systems is based on the
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% distributed accumulation of specific anatomical, physiological, and ethological
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% evidence. This leaves us with the challenge of integrating the available
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% knowledge fragments into a coherent whole in order to address more and more
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% far-reaching questions, from the interaction between individual processing
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% steps to comparisons between similar systems across different species. One way
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% to deal with this challenge is to build a unified framework that captures the
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% essential functional aspects of a sensory system. However, building such a
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% framework is a challenging task in itself. It requires a wealth of existing
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% knowledge of the system and the stimuli it operates on, a clearly defined
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% scope, and careful abstraction of the underlying structures and mechanisms.
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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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% One sensory system that has been extensively studied over the years is the
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% auditory system of grasshoppers~(\textit{Acrididae}). Grasshoppers rely on
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% their sense of hearing for intraspecific communication --- including mate
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% attraction~(\bcite{helversen1972gesang}) and
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% evaluation~(\bcite{stange2012grasshopper}), sender
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% localization~(\bcite{helversen1988interaural}), courtship
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% display~(\bcite{elsner1968neuromuskularen}), and rival
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% deterrence~(\bcite{greenfield1993acoustic}) --- and have evolved a variety of
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% acoustic signals for different behavioral
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% contexts~(\bcite{otte1970comparative}). The most conspicuous acoustic signals
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% of grasshoppers are their species-specific calling songs, which broadcast the
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% presence of the singing individual to potential mates within range. These songs
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% are 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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% \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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% \bcite{sevastianov2023evolution}), with over 6800 recognized species in the
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% \textit{Acrididae} family~(\bcite{cigliano2024orthoptera}).
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% % What are the signals that the auditory system is supposed to recognize?
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% Grasshopper songs are amplitude-modulated broad-band acoustic signals. They
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% consist of a series of noisy syllables and relatively quiet pauses, which form
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% a characteristic repetitive pattern~(\bcite{helversen1977stridulatory};
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% \bcite{stumpner1994song}). Song recognition depends on certain 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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% \bcite{helversen2004acoustic}) --- which are sufficiently conveyed by the
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% amplitude modulation of the song alone~(\bcite{helversen1997recognition}).
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% % Why is intensity invariance important for song recognition?
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% Grasshopper songs, like all acoustic signals, are subject to sound attenuation,
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% which depends on the distance from the sound source, the frequency content of
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% the signal, and the vegetation of the habitat~(\bcite{michelsen1978sound}).
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% This has two major consequences for the receiving auditory system. First, the
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% amplitude dynamics of the song pattern degrade with increasing distance to the
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% sender, which limits the effective communication range of grasshoppers
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% to~\mbox{1\,-\,2\,m} in their typical grassland
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% habitats~(\bcite{lang2000acoustic}). Second, the intensity of a song at the
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% receiver's position varies with the position of the sender, which should
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% ideally not affect song recognition. The auditory system thus needs to achieve
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% a certain degree of intensity invariance --- a time scale-selective sensitivity
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% to faster amplitude dynamics and simultaneous insensitivity to more sustained
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% amplitude dynamics. Intensity invariance is commonly associated with neuronal
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% adaptation~(\bcite{benda2008spike}; \bcite{barbour2011intensity};
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% \bcite{ozeri2018fast}; more general:~\bcite{benda2021neural}). Different neuron
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% types in the grasshopper auditory system exhibit spike-frequency adaptation in
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% response to sustained stimulation~(\bcite{romer1976informationsverarbeitung};
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% \bcite{gollisch2004input}; \bcite{hildebrandt2009origin};
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% \bcite{clemens2010intensity}; \bcite{fisch2012channel}). Accordingly, intensity
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% invariance is not the result of a single processing step but rather a gradual
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% process, in which different neuronal populations contribute to varying
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% degrees~(\bcite{clemens2010intensity}) and by different
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% mechanisms~(\bcite{hildebrandt2009origin}).
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% % How can song recognition be modelled functionally (feat. Jan Clemens & Co.)?
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% % How did we expand on the previous framework?
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% In the current study, we use a physiologically inspired functional model of the
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% grasshopper song recognition pathway to investigate the emergence of intensity
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% invariance along the auditory processing stream. The model pathway expands on a
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% previous functional model framework for song recognition --- including feature
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% extraction, 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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% both:~\bcite{ronacher2015computational}). The exisiting framework relies on
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% pulse trains as input signals, which were designed to capture the essential
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% structural properties of natural song envelopes~(\bcite{clemens2013feature}).
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% The extracted song features are compressed into a single preference value for
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% each presented input signal, which serves as predictor for the behavioral
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% response of the animal. We extended this framework by a physiologically
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% plausible preprocessing stage --- including envelope extraction, logarithmic
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% compression, and intensity adaptation --- that allows the model to operate on
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% recordings of natural grasshopper songs. Furthermore, we modified the feature
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% extraction stage in a way that retains the time-varying structure of individual
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% song features.
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\section{Methods}
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\section{Methods}
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% This maybe does not quite fit here, but it is the most general part of the
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% This maybe does not quite fit here, but it is the most general part of the
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@@ -1632,6 +1527,21 @@ natural song variation.
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\section{Conclusions \& outlook}
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\section{Conclusions \& outlook}
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% RIPPED FROM INTRODUCTION:
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% RIPPED FROM INTRODUCTION:
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|
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% Why functional models of sensory systems?
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% Our scientific understanding of sensory processing systems is based on the
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% distributed accumulation of specific anatomical, physiological, and
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% ethological evidence. This leaves us with the challenge of integrating the
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% available knowledge fragments into a coherent whole in order to address more
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% and more far-reaching questions, from the interaction between individual
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% processing steps to comparisons between similar systems across different
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% species. One way to deal with this challenge is to build a unified framework
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% that captures the essential functional aspects of a sensory system. However,
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% building such a framework is a challenging task in itself. It requires a
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% wealth of existing knowledge of the system and the stimuli it operates on, a
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% clearly defined scope, and careful abstraction of the underlying structures
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% and mechanisms.
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% Multi-species, multi-individual communally inhabited environments\\
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% Multi-species, multi-individual communally inhabited environments\\
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% - Temporal overlap: Simultaneous singing across individuals/species common\\
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% - Temporal overlap: Simultaneous singing across individuals/species common\\
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% - Frequency overlap: Little speciation into frequency bands (likely unused)\\
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% - Frequency overlap: Little speciation into frequency bands (likely unused)\\
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