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\subsection{Invariant processing in the grasshopper auditory system}
\label{sec:general_inv}
% Invariance in the general (systemic) sense:
% Invariance in the general (systemic) sense (could be skipped if too much):
The notion of invariance is fundamental for sensory processing systems.
Invariance, in the general sense, can be described as the property of a
transformation to maintain variation across certain meaningful input parameters
@@ -1785,52 +1785,61 @@ that are robust to noise masking~(\bcite{einhaupl2011attractiveness}).
% Trading SNR for log-HP intensity invariance (+variability, +general principle):
The SNR of each song representation prior to $\adapt(t)$ increases
monotonically with $\sca$~(excluding $0<\sca\ll1$, noise regime). These
representations maintain and improve the initial SNR of $\raw(t)$ and hence
never achieve intensity invariance. In contrast, the SNR of the
intensity-invariant $\adapt(t)$ never exceeds its saturation level even for
arbitrarily high $\sca$. The saturation level of $\adapt(t)$ varies across
species and songs. This variability is likely rooted in the way in which
logarithmic compression acts on the specific distribution of $\env(t)$, which
depends on the $\fc$ of the lowpass filter as well as the temporal structure
and frequency spectrum of the rectified $\filt(t)$. Overall, $\adapt(t)$ has
never been observed to exceed a SNR of around~10 across all songs. The low SNR
of $\adapt(t)$ partially results from the amplification of smaller values of
$\env(t)$ by the logarithm, which raises the noise floor of $\adapt(t)$. Still,
the reduction in SNR is substantial --- considering that the SNR of preceeding
song representations has been orders of magnitude higher --- but is likely a
necessary price to pay for the intensity invariance of $\adapt(t)$. After all,
a transformation cannot compress a range of different input intensities into a
representations maintain the full extent of the initial SNR of $\raw(t)$ and
hence never achieve intensity invariance. In contrast, the SNR of $\adapt(t)$
saturates for sufficiently high $\sca$. Accordingly, $\adapt(t)$ is
intensity-invariant but cannot have a higher SNR than the saturation level,
which indicates a fundamental trade-off. The saturation level of $\adapt(t)$
varies across species and songs. This variability is likely rooted in the way
in which logarithmic compression acts on the specific $\env(t)$, which depends
on the $\fc$ of the lowpass filter as well as the temporal structure and
frequency spectrum of the rectified $\filt(t)$. Across all songs, the
saturation level of $\adapt(t)$ has never been observed to exceed a SNR of
around~10. This is a substantial reduction in SNR, considering that the SNR of
preceeding representations had been orders of magnitude higher. Part of this
reduction stems from the amplification of smaller values of $\env(t)$ by
logarithmic compression, which raises the noise floor of $\adapt(t)$ relative
to the song. Accordingly, the low SNR of $\adapt(t)$ appears to be a necessary
price to pay for its intensity invariance through logarithmic compression and
adaptation. But the trade-off between intensity invariance and SNR likely goes
beyond the particular mechanisms along the pathway. After all, a transformation
is not expected to compress a range of different input intensities into a
constant output intensity without sacrificing some of the corresponding input
SNR. Accordingly, the trade-off between intensity invariance and SNR is not
expected to be specific to the particular mechanisms along the pathway but
presumably applies to any transformation that achieves or improves intensity
invariance.
SNR. This suggests that the trade-off is a more general principle that applies
to any transformation that achieves or improves intensity invariance.
Thresholding and temporal averaging renders feature $f_i(t)$
intensity-invariant for sufficiently large $\sca$. The trade-off between
intensity invariance and SNR is mediated by threshold value $\thr$. A lower
$\thr$ ($\thr\to0$) improves intensity invariance by shifting the saturation
point towards lower $\sca$ but also decreases the SNR of $f_i(t)$. The
saturation level of $f_i(t)$ is independent of $\thr$ as long as the intensity
invariance by the previous mechanism is neglected. The SNR of $f_i(t)$ is
therefore determined solely by the pure-noise response of $f_i(t)$. The
distribution $\pci$ of the pure-noise kernel response $c_i(t)$ is largely a
normal distribution with mean $\mu\approx0$ for all kernels $k_i(t)$. The value
of the pure-noise $f_i(t)$ is hence 0.5 for $\thr=0$ and decreases for higher
$\thr$. If $\thr$ is set above the maximum of $c_i(t)$, the pure-noise feature
value is 0, which results in an "unlimited" SNR of $f_i(t)$. In this case, any
non-zero feature value that is sustained for a sufficient duration could serve
as indicator for the presence of $\soc(t)$, although at the cost of a higher
saturation point. The maximum of the pure-noise $c_i(t)$ is assumed to be very
The second mechanism of intensity invariance consists of thresholding and
temporal averaging of $c_i(t)$ into $f_i(t)$. Here, the trade-off between
intensity invariance and SNR is mediated by the threshold value $\thr$. A lower
$\thr$~($\thr\to0$) improves the intensity invariance of $f_i(t)$ by shifting
the saturation point towards lower $\sca$. However, a lower $\thr$ also raises
the noise floor of $f_i(t)$ by including more of the pure-noise $c_i(t)$, which
decreases the SNR of $f_i(t)$. The distribution $\pci$ of the pure-noise
$c_i(t)$ is very close to a normal distribution with mean $\mu\approx0$ for all
kernels $k_i(t)$. The value of the pure-noise $f_i(t)$ is hence 0.5 for
$\thr=0$ and decreases for higher $\thr$. If $\thr$ is set above the maximum of
$c_i(t)$, the pure-noise feature value is 0, which results in an "unlimited"
SNR of $f_i(t)$. In this case, any non-zero feature value that is sustained for
a sufficient duration could serve as indicator for the presence of $\soc(t)$ in
$\raw(t)$, although at the cost of a higher saturation point. Of course, this
would require a fine evolutionary tuning of $\thr$ to the properties of the
natural noise in a certain habitat to avoid false positives.
The saturation level of $f_i(t)$ is independent of $\thr$ as long as the
intensity invariance by the previous mechanism is neglected.
If $f_i(t)$ can achieve an arbitrarily high SNR, it can counteract the
comparably low SNR of $\adapt(t)$
The maximum of the pure-noise $c_i(t)$ is assumed to be very
small due to the various SNR improvements along the pathway, so that the
required increase in $\thr$ and hence the saturation point of $f_i(t)$ is not
expected to be substantial. However, exploiting the capacity of $f_i(t)$ for
arbitrarily high SNR would certainly require a fine evolutionary tuning of
$\thr$ to the properties of both the species-specific song and the natural
noise in a certain habitat.
expected to be substantial.
\newpage
\subsection{Intensity invariance versus intensity invariance}
% \newpage
% \subsection{Intensity invariance versus intensity invariance}
Two consecutive mechanisms of intensity invariance do not necessarily add up to
a stronger overall intensity invariance. If the first mechanism results in a