Added some (older) papers on neuronal types, structure, and connectivity to the literature archive.
Expanded cite.bib accordingly.
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cite.bib
@ -100,6 +100,16 @@
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pages={2575--2580},
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year={2009},
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}
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@article{eichendorf1980projections,
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title={Projections of auditory ventral-cord neurons in the supraesophageal ganglion of Locusta migratoria},
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author={Eichendorf, Annette and Kalmring, Klaus},
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journal={Zoomorphologie},
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volume={94},
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number={2},
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pages={133--149},
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year={1980},
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publisher={Springer}
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}
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@article{fisch2012channel,
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title={Channel noise from both slow adaptation currents and fast currents is required to explain spike-response variability in a sensory neuron},
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author={Fisch, Karin and Schwalger, Tilo and Lindner, Benjamin and Herz, Andreas VM and Benda, Jan},
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@ -151,6 +161,37 @@
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pages={433--450},
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year={1996},
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}
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@article{kalmring1975afferent,
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title={The afferent auditory pathway in the ventral cord of Locusta migratoria (Acrididae) I. Synaptic connectivity and information processing among the auditory neurons of the ventral cord},
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author={Kalmring, Klaus},
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journal={Journal of comparative physiology},
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volume={104},
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number={2},
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pages={103--141},
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year={1975},
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publisher={Springer}
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}
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@article{kalmring1972akustische,
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title={Akustische Neuronen im Bauchmark der Wanderheuschrecke Locusta migratoria},
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author={Kalmring, Klaus and Rheinlaender, J{\"u}rgen and Rehbein, Hansgeorg},
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journal={Zeitschrift f{\"u}r vergleichende Physiologie},
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volume={76},
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number={3},
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pages={314--332},
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year={1972},
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publisher={Springer}
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}
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@article{kalmring1978coding,
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title={The coding of sound signals in the ventral-cord auditory system of the migratory locust, Locusta migratoria},
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author={Kalmring, Klaus and K{\"u}hne, Roland and Moysich, Fritz},
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journal={Journal of comparative physiology},
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volume={128},
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number={3},
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pages={213--226},
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year={1978},
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publisher={Springer}
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}
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@article{lang2000acoustic,
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title={Acoustic communication distances of a gomphocerine grasshopper},
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author={Lang, Friederike},
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@ -211,6 +252,17 @@
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pages={1965--1974},
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year={2008},
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}
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@article{rehbein1974structure,
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title={Structure and function of acoustic neurons in the thoracic ventral nerve cord of Locusta migratoria (Acrididae)},
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author={Rehbein, Hansgeorg and Kalmring, Klaus and R{\"o}mer, Heiner},
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journal={Journal of comparative physiology},
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volume={95},
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number={3},
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pages={263--280},
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year={1974},
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publisher={Springer}
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}# Cited
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@article{rehbein1976auditory,
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title={Auditory neurons in the ventral cord of the locust: Morphological and functional properties},
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author={Rehbein, Hansgeorg},
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@ -218,7 +270,8 @@
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volume={110},
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pages={233--250},
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year={1976},
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}
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}# Cited
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@article{romer1985responses,
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title={Responses to model songs of auditory neurons in the thoracic ganglia and brain of the locust},
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author={R{\"o}mer, Heiner and Seikowski, Ulrich},
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@ -265,6 +318,16 @@
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pages={154--175},
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year={2023},
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}
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@article{sippel1983nonlinear,
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title={Non-linear analysis of the transmission of signals in the auditory system of the migratory locust Locusta migratorta},
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author={Sippel, Martin and Breckow, Joachim},
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journal={Biological Cybernetics},
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volume={46},
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number={3},
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pages={197--205},
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year={1983},
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publisher={Springer}
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}
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@article{stumpner1994song,
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title={{Song production and song recognition in a group of sibling grasshopper species (Chorthippus dorsatus, Ch. dichrous and Ch. loratus: Orthoptera, Acrididae)}},
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author={Stumpner, Andreas and von Helversen, Otto},
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literature/Kalmring_Kühne_Moysich_1978.pdf
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literature/Rehbein_Kalmring_Römer_1974.pdf
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@ -8,28 +8,34 @@
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@ -40,32 +46,32 @@
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@ -74,5 +80,5 @@
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72
main.bbl
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main.bbl
@ -513,6 +513,78 @@
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\field{pages}{63\bibrangedash 101}
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\range{pages}{39}
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\endentry
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\entry{rehbein1976auditory}{article}{}
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\name{author}{1}{}{%
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{{un=0,uniquepart=base,hash=34d133bc807dc9040cb5b3db821041f3}{%
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family={Rehbein},
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\field{labeltitlesource}{title}
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\field{journaltitle}{J Comp Physiol A}
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\field{title}{Auditory neurons in the ventral cord of the locust: Morphological and functional properties}
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\field{volume}{110}
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\field{year}{1976}
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\field{pages}{233\bibrangedash 250}
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\range{pages}{18}
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\endentry
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{{un=0,uniquepart=base,hash=96806c4105cbfbf30da29ebb55c2d4e1}{%
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@ -134,6 +134,25 @@ $\rightarrow$ More general, simpler, unfitted formalized Gabor filter bank
|
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\section{Developing a functional model of\\the grasshopper auditory pathway}
|
||||
|
||||
The auditory pathway of grasshoppers comprises a number of neuronal stages and
|
||||
identified cell types~(\bcite{rehbein1974structure}; \bcite{rehbein1976auditory}).
|
||||
|
||||
Along the auditory pathway of grasshoppers (Fig.\,\ref{fig:pathway}a+b),
|
||||
|
||||
1) "Pre-split portion" of the auditory pathway:\\
|
||||
Tympanal membrane $\rightarrow$ Receptor neurons $\rightarrow$ Local interneurons
|
||||
|
||||
Similar response/filter properties within receptor/interneuron populations (\cite{clemens2011efficient})\\
|
||||
$\rightarrow$ One population-wide response trace per stage (no "single-cell resolution")
|
||||
|
||||
2) "Post-split portion" of the auditory pathway:\\
|
||||
Ascending neurons (AN) $\rightarrow$ Central brain neurons
|
||||
|
||||
Diverse response/filter properties within AN population (\cite{clemens2011efficient})\\
|
||||
- Pathway splitting into several parallel branches\\
|
||||
- Expansion into a decorrelated higher-dimensional sound representation\\
|
||||
$\rightarrow$ Individual neuron-specific response traces from this stage onwards
|
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|
||||
\begin{figure}[!ht]
|
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\centering
|
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\def\svgwidth{\textwidth}
|
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@ -148,9 +167,9 @@ $\rightarrow$ More general, simpler, unfitted formalized Gabor filter bank
|
||||
|
||||
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
|
||||
mechanical resonance filter, that focuses vibrations of specific frequencies on
|
||||
different membrane areas while attenuating
|
||||
others~(\bcite{michelsen1971frequency}; \bcite{windmill2008time};
|
||||
mechanical resonance filter: Vibrations that fall within specific frequency
|
||||
bands are focused on different membrane areas, while others are
|
||||
attenuated~(\bcite{michelsen1971frequency}; \bcite{windmill2008time};
|
||||
\bcite{malkin2014energy}). This processing step can be approximated by an
|
||||
initial bandpass filter
|
||||
\begin{equation}
|
||||
@ -158,10 +177,10 @@ initial bandpass filter
|
||||
\label{eq:bandpass}
|
||||
\end{equation}
|
||||
applied to the acoustic input signal $\raw(t)$. The auditory receptor neurons
|
||||
connect directly to the tympanal membrane. Besides performing the
|
||||
mechano-electrical transduction, the receptor population further is substrate
|
||||
to several known processing steps. First, the receptors extract the signal
|
||||
envelope~(\bcite{machens2001discrimination}), which likely involves a
|
||||
connect directly to the tympanal membrane~(Fig.\,\ref{fig:pathway}a). Besides
|
||||
performing the mechano-electrical transduction, the receptor population is
|
||||
substrate to several known processing steps. First, the receptors extract the
|
||||
signal envelope~(\bcite{machens2001discrimination}), which likely involves a
|
||||
rectifying nonlinearity~(\bcite{machens2001representation}). This can be
|
||||
modelled as full-wave rectification followed by lowpass filtering
|
||||
\begin{equation}
|
||||
@ -177,52 +196,25 @@ logarithmic compression is achieved by conversion to decibel scale
|
||||
\label{eq:log}
|
||||
\end{equation}
|
||||
relative to the maximum intensity $\dbref$ of the signal envelope $\env(t)$.
|
||||
The axons of the receptor neurons project into the metathoracic ganglion, where
|
||||
they synapse onto local interneurons~(Fig.\,\ref{fig:pathway}b). Both the local
|
||||
interneurons~(\bcite{hildebrandt2009origin}; \bcite{clemens2010intensity}) and,
|
||||
to a lesser extent, the receptors themselves~(\bcite{fisch2012channel}) display
|
||||
spike-frequency adaptation in response to sustained stimulation.
|
||||
This behavior is crucial to render subsequent signal representations invariant
|
||||
to variations in sound intensity.
|
||||
|
||||
|
||||
"Pre-split portion" of the auditory pathway:\\
|
||||
Tympanal membrane $\rightarrow$ Receptor neurons $\rightarrow$ Local interneurons
|
||||
|
||||
Similar response/filter properties within receptor/interneuron populations (\cite{clemens2011efficient})\\
|
||||
$\rightarrow$ One population-wide response trace per stage (no "single-cell resolution")
|
||||
|
||||
\textbf{Stage-specific processing steps and functional approximations:}
|
||||
|
||||
Initial: Continuous acoustic input signal $x(t)$
|
||||
|
||||
Filtering of behaviorally relevant frequencies by tympanal membrane\\
|
||||
$\rightarrow$ Bandpass filter 5-30 kHz
|
||||
|
||||
Extraction of signal envelope (AM encoding) by receptor population\\
|
||||
$\rightarrow$ Full-wave rectification, then lowpass filter 500 Hz
|
||||
|
||||
Logarithmically compressed intensity tuning curve of receptors\\
|
||||
$\rightarrow$ Decibel transformation
|
||||
|
||||
Spike-frequency adaptation in receptor and interneuron populations\\
|
||||
$\rightarrow$ Highpass filter 10 Hz
|
||||
%
|
||||
Next, the axons of the receptor neurons project into the metathoracic ganglion,
|
||||
where they synapse onto local interneurons~(Fig.\,\ref{fig:pathway}b). Both the
|
||||
local interneurons~(\bcite{hildebrandt2009origin};
|
||||
\bcite{clemens2010intensity}) and, to a lesser extent, the receptors
|
||||
themselves~(\bcite{fisch2012channel}) display spike-frequency adaptation in
|
||||
response to sustained stimulus intensity levels. This mechanism allows for the
|
||||
robust encoding of faster amplitude modulations against a slowly changing
|
||||
overall baseline intensity. Functionally, this processing step resembles a
|
||||
highpass filter
|
||||
\begin{equation}
|
||||
\adapt(t)\,=\,\db(t)\,*\,\hp, \qquad \fc\,=\,10\,\text{Hz}
|
||||
\label{eq:highpass}
|
||||
\end{equation}
|
||||
%
|
||||
over the logarithmically scaled envelope $\db(t)$. The projections of the local
|
||||
interneurons remain within the metathoracic ganglion and synapse onto a small
|
||||
number of ascending neurons~(Fig.\,\ref{fig:pathway}b).
|
||||
|
||||
\subsection{Feature extraction by individual neurons}
|
||||
|
||||
"Post-split portion" of the auditory pathway:\\
|
||||
Ascending neurons (AN) $\rightarrow$ Central brain neurons
|
||||
|
||||
Diverse response/filter properties within AN population (\cite{clemens2011efficient})\\
|
||||
- Pathway splitting into several parallel branches\\
|
||||
- Expansion into a decorrelated higher-dimensional sound representation\\
|
||||
$\rightarrow$ Individual neuron-specific response traces from this stage onwards
|
||||
|
||||
\textbf{Stage-specific processing steps and functional approximations:}
|
||||
|
||||
Template matching by individual ANs\\
|
||||
@ -267,6 +259,20 @@ $\rightarrow$ Lowpass filter 1 Hz
|
||||
%
|
||||
\section{Two mechanisms driving the emergence of intensity-invariant song representation}
|
||||
|
||||
\textbf{Definition of invariance (general, systemic):}\\
|
||||
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
|
||||
of one or more other parameters (variation to be discarded)
|
||||
$\rightarrow$ Selective input-output decorrelation
|
||||
|
||||
\textbf{Definition of intensity invariance (context of neurons and songs):}\\
|
||||
Intensity invariance = Time scale-selective sensitivity to certain faster
|
||||
amplitude dynamics (song waveform, small-scale AM) and simultaneous
|
||||
insensitivity to slower, more sustained amplitude dynamics (transient baseline,
|
||||
large-scale AM, current overall intensity level)\\
|
||||
$\rightarrow$ Without time scale selectivity, any fully intensity-invariant
|
||||
output will be a flat line
|
||||
|
||||
\subsection{Logarithmic scaling \& spike-frequency adaptation}
|
||||
|
||||
Envelope $\env(t)$ $\xrightarrow{\text{dB}}$ Logarithmic $\db(t)$ $\xrightarrow{\hp}$ Adapted $\adapt(t)$
|
||||
|
||||
Loading…
Reference in New Issue
Block a user